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ANNALS

of CARNEGIE MUSEUM

THE CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 63 22 FEBRUARY 1 994 NUMBER 1

CONTENTS

ARTICLES

Geographic variation and systematics of the striped whipsnakes
taeniatus complex; Reptilia: Serpentes: Colubridae)

y\'rtS0 Nifijsj

o t m

cophrs

Jeffrey D. Camper and James R. Dixon 1

U-shaped orientation of Hunter-Schreger bands in the enamel of Moropus
(Mammalia: Chalicotheriidae) in comparison to some other
Perissodactyla Wighart v. Koenigswald 49

Five new species of Hydroptila from eastern United States (Insecta: Tri-

choptera: Hydroptilidae) Jan L. Sykora and Steven C. Harris 67

Description of the immature stages of six species of Sphaenognathus, with
comparative notes on phylogeny and natural history (Insecta: Coleop-
tera: Lucanidae) Giovanni Onore 77

REVIEW

The Scent of Orchids: Olfactory and Chemical Investigations [Roman

Kaiser] Fredrick W. Wright, Jr. 101

Erratum 103

Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS:

K. C. Beard

L. Krishtalka

M. A. Schmidt, ELS, Assistant Editor

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THE CARNEGIE

MUSEUM OF
NATURAL HISTORY

@ This paper meets the requirements of ANSI/NISO Z39. 48-1992 (Permanence of Paper).

ANNALS OF CARNEGIE MUSEUM

Vol, 63, Number 1, P. 1-48

22 February 1994

GEOGRAPHIC VARIATION AND SYSTEMATICS OF THE
STRIPED WHIPSNAKES {MASTICOPHIS TAENIATUS COMPLEX;

REPTILIA: SERPENTES: COLUBRIDAE)

Jeffrey D, Camper1’2
James R. Dixon1
Abstract

Geographic variation in morphological and protein characters in Masticophis taeniatus sensu lato
and Masticophis hilineatus were investigated. Populations of striped whipsnakes in southern Texas
and eastern Mexico allocated to M. t. schotti, M. t. ruthveni, and M. t. australis are not conspecific
with those of M. t. taeniatus and M. t. girardi of the western United States and Mexico. The southeastern
populations are elevated to specific status and referred to Masticophis schotti, whereas the western
whipsnake populations remain M. taeniatus. Striped whipsnake populations in south-central Mexico
allocated to M. t. australis are not separable from those making up M. t. ruthveni and the two taxa
are synonymized. Masticophis schotti and M. taeniatus differ from one another in maxillary tooth
number, preanal dorsal scale row frequency, number of apical scale pits, and color pattern. An allelic
difference across the Balcones Escarpment of central Texas at the supernatant amino acid transferase
C S-Aat-A ) locus exists between M. t. girardi and M, s. schotti . Masticophis hilineatus shows little
geographic variation. A phylogenetic analysis of geographic samples supports the monophyly of both
M. taeniatus and M. schotti.

Resumen

Se estudio la variation geografica de caracteres morfoldgicos y proteicos en Masticophis taeniatus
sensu lato y Masticophis hilineatus. Las poblaciones de las culebras rayadas en el sur de Texas y este
de Mexico, ante conocidas como M, t. schotti, M. t. ruthveni , y M. t. australis no son eonespedficas
con M t. taeniatus y M. t. girardi del oeste de Estados Unidos y Mexico. Las poblaciones del sureste
son el v adas a la categoria de especie y re fen das como Masticophis schotti, mientras que las poblaciones
del oeste retienen el nombre de M. taeniatus. Las poblaciones del sur del centre de Mexico, conocidas
como M. t. australis, las consideramos sinonimias con M. t. ruthveni. Estas dos especies difieren en
el numero de dientes maxilares, la frecuencia de Imeas de escamas dorsales preanales, el numero de
foestas apicales y el patron de coloration. Existe una difereneia en el locus del supemadante del
arm noacid o de transferencia ( S-Aat-A ) entre M. t. girardi y M. s. schotti, loeaiizandose sobre el Balcones
Escarpment del centro de Texas. Masticophis hilineatus present© poca variation geografica. Un analisis
filogenetico de las muestras geograficas sostiene la monofiiia de M. taeniatus y M. schotti.

Introduction

The status of populations of the striped whipsnake, Masticophis taeniatus sensu
lato, in Texas and Mexico has long been enigmatic to herpetologists. The first
name applied to snakes of this species-group was Leptophis taeniatus (Hallowell,
1852). Striped whipsnakes from central and western Texas and western Mexico
were described as Masticophis ornatus (Baird and Girard, 1853). This form was
subsequently recognized as a subspecies of M. taeniatus (Cope, 1875). Stejneger

1 Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas 77843-
2258.

2 Present address: Department of Biology, Suite 61588, Austin College, Sherman, Texas 75091-1177.
Submitted 16 August 1991.

Annals of Carnegie Museum

vol. 63

and Barbour (1917) did not recognize the genus Masticophis and relegated all
species of Masticophis to Coluber. Hence they substituted C. t. girardi for Coluber
taeniatus ornatus, which became a secondary junior homonym of Coluber ornatus
Shaw, 1802 ( =Chrysopelea ornata of Asia). Intergradation of color pattern char-
acters between M. t. taeniatus and M. t. girardi was thought to occur in extreme
western Texas (Ortenburger, 1928). Schmidt and Smith (1944) stated that the use
of Masticophis taeniatus ornatus “appears to be in accord with the opinion of the
International Commission on Zoological Nomenclature with regard to secondary
homonyms.” However, Article 59 of the 1985 Code stated that all secondary
junior homonyms replaced before 1961 are permanently invalid; therefore, M. t.
girardi should be used instead of M. t. ornatus.

Baird and Girard (1853) described Masticophis schotti from southern Texas, in
the Tamaulipan Biotic Province. Ortenburger (1928) examined only three spec-
imens of this taxon. Gloyd and Conant (1934) considered M. schotti a subspecies
of M. taeniatus, based on two juvenile specimens that they believed to be inter-
grades from the eastern portion of the Edwards Plateau. A darker form with
reduced striping from the lower Rio Grande valley of Texas and Tamaulipas,
Mexico, described as Masticophis ruthveni (Ortenburger, 1923), was also consid-
ered a subspecies of M. taeniatus by Gloyd and Conant (1934). Based on the
assumption that juvenile M. t. ruthveni do not have light stripes, Smith (1941)
described Masticophis taeniatus australis from Guanajuato, Mexico.

Masticophis bilineatus was described by Jan (1863) from Mexico. Cope (1891)
later applied the name Bascanium semilineatum to the same taxon from Arizona
(Smith, 1941). Subspecies described include Masticophis bilineatus lineolatus
(Hensley, 1950) from the Ajo Mountains of Arizona, based on minor color pattern
differences, and Masticophis bilineatus slevini (Lowe and Norris, 1955), the pu-
tatively dwarf population on Isla San Esteban, Baja California, Mexico. Our
interest in investigating geographic variation in M. bilineatus and the relationship
of this taxon to M. taeniatus arose because of parapatry among similarly-patterned
M. bilineatus, M. t. girardi, and M. t. ruthveni in south-central Mexico, and the
apparent sympatry of M. bilineatus and M. t. taeniatus in central Arizona. The
objectives of this study are to: (1) determine evolutionary relationships within
the striped whipsnake complex, Masticophis taeniatus sensu lato; (2) determine
evolutionary and geographic relationships between Masticophis bilineatus and M.
taeniatus ; and (3) document patterns of geographic variation in morphological
and protein characters of M. taeniatus and M. bilineatus.

Materials and Methods

A total of 1633 Masticophis taeniatus and 335 M. bilineatus from throughout the ranges of these
species was examined (Fig. 1, Appendix 1). The states for 36 characters were recorded for each M.
taeniatus, and 35 for each M. bilineatus. Four measurements (total length, tail length, head length
[snout to posterior margin of the last supralabial], and greatest head width) were recorded to nearest
0. 1 mm. Because the character has been used to discriminate among subspecies of M. bilineatus (Lowe
and Norris, 1955), the length of the right posterior chin shield divided by the distance from the anterior
edge of the mental scute to the posterior edge of the posterior chin shield was calculated for M.
bilineatus. The following data were also recorded: sex; age (adult or juvenile); anal plate condition
(single/divided); number of scale rows at neck, midbody, and ten ventral scutes anterior to the vent
(Dowling, 1951); numbers of supralabials, infralabials, preoculars, postoculars, loreals, primary tem-
porals, secondary temporals, ventrals, subcaudals, and maxillary teeth; and supralabials contacting
the orbit. Head scale variation is presented for the right side of the head only. Fourteen color pattern
characters were also recorded. Presence and width (both dorsally and laterally) of a light nuchal collar

1994

Camper and Dixon— Striped Whipsnakes

Fig. 1.— Map showing location of specimens examined in this study. Closed circles = Masticophis t.
taeniatus, open circles = M. t. girardi, circled stars = M. t. taeniatus x girardi, black squares = M. t.
schotti, open squares = M. t. ruthveni, and triangles = Masticophis bilineatus. Stars indicate type
localities for respective taxa.

Annals of Carnegie Museum

vol. 63

and number of light bands were recorded for M. taeniatus only. Other color pattern characters included:
number of stripes; stripe color (light or dark); stripe position (scale row location); presence of dorsal
speckling; presence of light head scale borders; dorsal color pattern; ventral pattern of neck, belly, and
tail; and length of stripes expressed in numbers of ventral scutes. Scale row reductions were recorded
for each specimen and presented in the formula recommended by Dowling (1951). The length of the
inverted hemipenis was recorded in terms of subcaudals subtended.

Measurements were used to calculate five ratios: tail length/snout-vent length (TLR), head length/
head width (HR), head length/snout-vent length (HLSVL), head width/snout-vent length (HWSVL),
and posterior chin shield/mental-posterior chin shield distance (CSR, M. bilineatus only). Tests of
geographic variation used analysis of variance (ANOVA) and Duncan’s multiple range (univariate)
and canonical discriminant analyses (multivariate) employing the GLM and CANDISC procedures
of the Statistical Analysis System software package, respectively (SAS Institute, 1985). A significance
level of a = 0.05 was used on all statistical tests. Phenograms based on distance and correlation
coefficient matrices were generated with the unweighted pair-group method using arithmetic means
(UPGMA) algorithm of the Numerical Taxonomy System of Multivariate Statistical Programs, NT-
SYS (Rohlf et al., 1980). Version 2.4 of Phylogenetic Analysis Using Parsimony (PAUP) and the
associated CONTREE were used to perform phylogenetic analyses (Swofford, 1985). Genotype data
were analyzed with Biosys-I computer software (Swofford and Selander, 1981).

Sexual, phylogenetic, and geographic variation analyses were conducted on the 22 samples shown
in Fig. 3. Samples 1-17 and 22 represent M. taeniatus sensu lato and 18-21 are M. bilineatus. Samples
included mapped localities grouped in 80.5 km2 unit areas delineated after examining the geographic
distribution of locality data and considering major biogeographic boundaries. Cluster analysis of the
80.5 km2 unit areas, using the UPGMA algorithm on separate sexes, yielded these 22 samples (Fig. 3).

Heart, liver, kidney, and skeletal muscle tissue removed from freshly sacrificed specimens of M.
taeniatus sensu lato (33), and M. bilineatus (3) were used to analyze protein variation by horizontal
starch gel electrophoresis (Harris and Hopkinson, 1976; Selander et al., 1971; Table 1). Data from
M. bilineatus were not included in analyses because too few individuals were available to accurately
estimate allozyme variation. Voucher specimens are deposited in the Texas Cooperative Wildlife
Collections, University of Texas at El Paso, and New Mexico State University (Appendix 2).

Results

Ontogenetic Variation.— Due to insufficient samples of juveniles, the descrip-
tion of ontogenetic variation is limited to color pattern differences between adults
and hatchlings. The hatchling color pattern of M. taeniatus sensu lato is invariable
throughout its range (Fig. 2). The holotype of M. t. australis also exhibited this
pattern and is identical to all other juvenile specimens examined. Adult color
patterns are shown in Fig. 1 3 ( taeniatus ), 1 5 ( girardi ), 1 7 ( schotti ), and 1 9 {ruthveni).
The color pattern of M. bilineatus changes little ontogenetically; therefore, only
adult color patterns are shown (Fig. 20).

Hatchling M. taeniatus have a ground color of dark olive green dorsally, with
two white or cream lateral stripes. One stripe encompasses the lower half of scale
row 1 and the lateral edges of the ventrals. The other stripe is on the upper half
of scale row 3 and the lower half of scale row 4. The dorsal pattern is most similar
to that of adult M. t. schotti or some M. t. ruthveni, differing only in ground color
shade and absence of dorsal speckling. Hatchlings have a cream-colored venter
with a narrow, diffuse dark olive lateral stripe forming the lower border of the
lower light stripe. The medial portion of the venter is immaculate. Some hatchling
M. taeniatus have paired black spots on the first five to ten ventral scutes. Spots
may also be scattered on the chin shields and gular scales. Hatchlings are most
similar to adult M. t. taeniatus in ventral color pattern.

Sexual Variation.— Eleven characters and snout-vent length (SYL) were tested
for sexual variation (Table 2). Samples 4, 11, 15, 16 (M. taeniatus ), and 18 (M.
bilineatus) were chosen because they are the largest (Fig. 3). A lower limit of 500
mm SVL was chosen for adult size in analyses of SVL. Males had significantly
larger body sizes than females in samples 1 1 and 18 (Table 2).

1994

Camper and Dixon— Striped Whipsnakes

Table 1.— Proteins examined, tissues of origin, and electrophoretic conditions yielding best resolution
o/Masticophis taeniatus allozymes. Mitochondrial and supernatant loci are denoted by M- and

S- prefixes, respectively .

Protein

Enzyme Commission
number

Locus

Tissue

Electrophoretic

conditions2

Aminopeptidase

3.4.11.1

AP-A

liver

Aspartate

aminotransferase

2.6.1. 1

M-AAT-A

liver

2.6.1. 1

S-AAT-A

liver

Creatine kinase

2.73.2

CK-A

muscle

2.73.2

CK-C

muscle

Dihydrolipoamide

dehydrogenase

1. 8.1.4

DDH-A

liver

Esterase15

EST-D

liver

Fumarate hydratase

4.23.2

FUMHA

liver

General protein

GPS

muscle

Glucose-6-phosphate

dehydrogenase

1.1.1.49

G6PDH-A

heart

Glutamate dehydrogenase

1,43.2

GTDHP-A

liver

Glyceraldehyde-3-phosphate

dehydrogenase

1.2332

GAPDH-A

muscle

Isocitrate dehydrogenase

1.1.1.42

M-IDH-A

liver

1.1.1.42

S-IDH-A

liver

Lactate dehydrogenase

1.1.1.27

LDH-A

liver

1.1.1.27

LDH-B

liver

Malate dehydrogenase

1.1.1.37

M-MDH-A

liver

1.1.1.37

S-MDIIA

liver

Mannose-6-phosphate isomerase

5.33.8

MPI-A

muscle

Peptidase"

3.4.-V-

PEP-A

liver

PEP-B

liver

PEP-C

liver

PEP-S

liver

Phosphoglucomutase

5. 4. 2. 2

PGM-A

muscle

5. 4.2. 2

PGM-B

muscle

Proline dipeptidase

3.4.13.9

PROS

liver

Pyruvate kinase

2.7.1.40

PK-A

liver

Superoxide dismutase

135.1.1

SSOD-A

liver

Xanthine dehydrogenase

13.1.204

XDH-A

liver

a A: Tris-HCl pH 8.5/borate pH 8.2; B: Discontinuous tris-citrate-borate (Poulik, 1957); C: Tris-citrate
pH 8.0; D: Tris-citrate pH 7.0.
b The substrate used was methyl-umbelliferyl acetate.

c Substrates used were leucyl-alanine for Peptidase A and C, leucyl-glycyl-glycine for Peptidase B, and
leucyl-leucyl-leucine for Peptidase S.

Mean ventral counts were greater for females in samples 1 5 and 1 6, and greater
for males in sample 18 (Table 2). Males had significantly higher mean subcaudal
counts than females (Table 2). Males in samples 11 and 15 had greater mean
maxillary tooth counts than females. Males had relatively longer tails, as measured
by TLR, in sample 1 1 , whereas females had relatively longer tails in sample 1 5
(Table 2). Sexual variation was not found in HR, HLSVL, or HWSVL. Males in
samples 1 1 and 15 had relatively longer heads (Table 2).

Males in samples 4, 11, 15, and 16 had significantly fewer posterior scale rows
than females. Masticophis taeniatus sensu lato always has 1 5 scale rows anteriorly
and at midbody. The first scale row reduction (SRR1) is lateral (scale row 3) and
paired. Therefore, SRR1 reduces scale rows from 15 to 13, and is considered a

Annals of Carnegie Museum

vol. 63

Fig. 2.— Color pattern of juvenile Masticophis taeniatus sensu lato.

single reduction site. The second (SRR2) and third (SRR3) reduction sites are
unpaired, located middorsally, and occur separately. Most males in samples 4
(85%) and 11 (68%) had three scale row reduction sites (15-15-11, Table 2),
whereas most females in these samples (80% and 69%, respectively) had two scale
row reduction sites (15-15-12). Most males in samples 15 (57%) and 16 (48%)
had two (15-15-12) scale row reductions (Table 2), whereas most females in
samples 15 (75%) and 16 (84%) had one scale row reduction site (15-15-13). This
is also illustrated by the small number of females in samples 1 5 and 1 6 with more
than one scale row reduction site (Table 2). Females have a greater relative girth
posteriorly as indicated by the more posteriad location of scale row reduction
sites. The locations of all reduction sites were significantly different between the
sexes in sample 1 1 , but only SRR2 was significantly different in sample 4 (Table 2).

Virtually all M. bilineatus possess 17-17-13 scale rows. Scale row reduction 1
occurs laterally and is paired on scale row 4. The location of SRR2 and SRR3
were significantly different between the sexes in sample 18 (Table 2). Unlike the
other four samples (M. taeniatus ), males in sample 18 had SRR2 and SRR3
significantly more posteriad than did females.

Geographic Variation. — Geographic variation in ventral and subcaudal scutes
was analyzed by plotting sample statistics on modified Dice-Leraas diagrams (Fig.
4-7). Samples with nonoverlapping shaded boxes (±2 SE) have means that are
significantly different. Males in samples 15-17, 20, and 21 had significantly fewer
mean ventrals than most in samples 1-14, 18, and 19 (Fig. 4). In females, samples

1994

Camper and Dixon— Striped Whipsnakes

Fig. 3.— Distribution and location of the 22 samples of Masticophis used in this study. The locality
between samples 1 7 and 21 is in sample 1 3 and is represented by one specimen only.

Table 2.— Statistically significant (a = 0.05) sexual variation in samples 4, 11, 15, 16, and 18.

Annals of Carnegie Museum

vol. 63

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b Wileoxon Rank Sum test on raw data.

1994

Camper and Dixon— Striped Whipsnakes

LOCALITY

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180 185 190 195 200 205 210 215 220 225

230 235

VENTRALS

Fig. 4.— Modified Dice-Leraas diagram of geographic variation in ventrals of males of samples 1-22
shown in Fig. 3. Horizontal line is range, vertical line is mean, clear box is ± 1 SD, and shaded box
is ±2 SE.

Annals of Carnegie Museum

vol. 63

LOCALITY

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VENTRAL5

Fig. 5. —Modified Dice-Leraas diagram of geographic variation in ventrals of females of samples 1-
22 shown in Fig. 3. Explanation as in Fig. 4.

1994

Camper and Dixon— Striped Whipsnakes

1 9
16

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5UBCAUDAL5

Fig. 6.— Modified Dice-Leraas diagram of geographic variation in subcaudals of males of samples 1-
22 shown in Fig. 3. Explanation as in Fig. 4.

Annals of Carnegie Museum

vol, 63

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SUBCAUDALS

Fig. 7.— Modified Dice - Lera as diagram of geographic variation in subcaudals of females of samples
1=22 shown in Fig. 3. Explanation as in Fig. 4.

1994

Camper and Dixon —Striped Whipsnakes

Table 3. — Geographic variation in HLSVL among samples 1-22 shown in Fig. 3. Means of samples
grouped by the same letter are not significantly different.

X (SD)

Range

Sample

Grouping

0.034 (0.002)

0.030-0.037

0.000

0.033 (0.002)

0.029-0.036

0.000

0.033 (0.002)

0.030-0.036

0.001

164

0.033 (0.002)

0.021-0.040

0.000

0.033 (0.003)

0.029-0.048

0.000

0.032 (0.002)

0.030-0.038

0.000

0.031 (0.002)

0.028-0.035

0.000

0.031 (0.002)

0.023-0.041

0.000

0.031 (0.002)

0.027-0.039

0.000

0.030 (0.002)

0.025-0.036

0.000

0.030 (0.003)

0.027-0.049

0.000

0.030 (0.002)

0.018-0.037

0.000

0.030 (0.002)

0.026-0.036

0.000

0.030 (0.002)

0.028-0.034

0.001

0.030 (0.002)

0.027-0.035

0.000

0.030 (0.002)

0.027-0.035

0.001

120

0.030 (0.002)

0.025-0.035

0.000

0.030 (0.002)

0.027-0.035

0.000

145

0.029 (0.002)

0.015-0.035

0.000

0.029 (0.001)

0.028-0.032

0.000

0.029 (0.004)

0.023-0.058

0.000

0.029 (0.003)

0.018-0.034

0.001

16, 17, 20, and 21 had significantly fewer mean ventrals than most other samples
(Fig. 5). However, ranges overlap extensively for both sexes. In both sexes there
were significant differences in mean ventral counts between parapatric samples
14 and 15. The boundary between sample 14 ( girardi ) and sample 15 (schotti)
forms the only contact zone between these taxa that is well-represented by spec-
imens.

An increase in average subcaudal number in southern populations of males was
evident among samples 1=13 and 15-17 (Fig. 6). A significant difference in mean
subcaudal counts between samples 1 4 and 1 5 existed (Fig. 6). Sample 20 had a
significantly lower mean subcaudal count than all other samples except 1,2, and
2 1 . A northwest to southeast clinal increase in subcaudals was present in females
of samples 1-10 (Fig. 3, 7). Females in sample 22 (M. t. taeniatus x M. t. girardi )
were intermediate between sample 10 {taeniatus) and sample 1 1 {girardi). A north
to south clinal decrease in subcaudal number was evident among females in
samples 14-16 and 18-21 (Fig. 3, 7).

ANOYAs were significant for hemipenis length, maxillary tooth counts, TLR,
HR, HLSVL, and HWSVL. However, Duncan’s multiple range tests showed
significant variation in HLSVL only. Sample 20 had a significantly larger mean
HLSVL {P < 0.05, d.f. = 1254, n = 1255). Samples grouped by the same letters
have means that are not significantly different (Table 3). Samples 1-4, 15, and
17 had the largest mean hemipenis lengths; however, no significant groupings
were found {P < 0.05, d.f. = 447, n = 448). Average maxillary tooth counts for
samples 11-14 and 18-21 were greater than those for samples 1-10 and 15-17;
however, no significant groupings were found {P < 0.05, d.f. = 693, n = 694).
Samples 9, 11-13, and 16-17 had proportionately the longest tails, averaging
>47% of SVL, whereas samples 1-4 had the lowest TLR values with averages

Annals of Carnegie Museum

vol. 63

VECTOR I (88.65%)

Fig. 8.— Projections on the first three canonical vectors of sample means of 22 samples of male striped
whipsnakes, Masticophis.

<44% of SYL. However, no significant groupings were found (P < 0.05, d.f. =
877, n = 878). No significant groupings of samples based on mean HR were found
(P < 0.05, d.f. = 1252, n = 1253). Relative head width as measured by HWSVL
was greater in southern samples, but no significant groupings were found (P <

0.05, d.f. = 1250, n = 1251).

Lowe and Norris (1955) implied that a lower CSR was characteristic of the
insular and Ajo Mountains populations of M. bilineatus. Kruskal- Wallis tests
comparing the Ajo Mountains population with four southern Arizona populations
and southwestern New Mexico material were significant (X2 = 11.76, P < 0.05,
d.f. = 3). However, a Duncan’s multiple range test comparing means of these
populations showed no significant differences. A Kruskal- Wallis test showed no
significant variation in CSR among samples 18-21 (X2 = 5.08, P > 0.05, d.f. =
3). The population of M. bilineatus from Isla San Esteban did not differ signifi-
cantly in SVL from three mainland Sonora populations (Kruskal- Wallis test, X2
= 3.02, P > 0.05, d.f. = 3). The insular specimens were not significantly smaller,
V = 0.981, d.f. = 266, P > 0.05, with mean SYLs of 899 mm for 254 mainland
adults and 854 mm for 14 adults from Isla San Esteban.

Multivariate analyses utilized the following characters: scale pit number, preanal
scale rows, ventral counts, TLR, number of transverse light bands, number of
stripes, maxillary tooth counts, HR, HLSVL, HWSVL, SRR1, and SRR2. A
canonical discriminant analysis using Wilk’s Lambda, Pillai’s Trace, Hotelling-
La wley’s Trace, and Roy’s Greatest Root was conducted to test the null hypothesis
of no overall significant variation due to locality. All were significant at P < 0.05.
In the canonical discriminant analysis, canonical vectors I— III explained most
(99.03%) of the variation (Table 4). Samples 1-14 and 22 separated from samples

1994

Camper and Dixon— Striped Whifsnak.es

Table A.— Eigenvalues and percent influence for 12 canonical vectors for the 22 samples of male striped

whipsnakes, Masticophis shown in Fig. 3.

Canonical vector

Eigenvalue

Percent influence

Cumulative

75.5058

88.65

6.3298

7.43

96.08

III

2.5133

2.95

99.03

0.2564

0.30

99.33

0.1852

0.22

99.55

0.1440

0.17

99.72

VII

0.0849

0.10

99.82

VIII

0.0487

0.06

99.88

0.0442

0.05

99.93

0.0252

0.03

99.96

0.0196

0.02

99.98

XII

0.0152

0.02

100.00

15-21 along canonical vector I which summarized 88.65% of the variation (Fig.
8; Table 4). Number of stripes was the most influential character along this axis
(Table 5). Canonical vector II accounted for 7.43% of the variation and separated
samples 1-10 from 1 1-14 and 22. Band number influenced this axis greatly (Table
5). Canonical vector III accounted for 2.95% of the variation and separated sam-
ples 15-17 from 1 8-2 1 (Fig. 8). Preanal dorsal scale row counts and SRR2 position
influenced this vector the most (Table 5).

The patterns of separation seen in the canonical discriminant analysis were
concordant with cluster analysis (Fig, 9). Characters used in cluster analysis in-
cluded: scale pit number, number of scale rows around the anterior body, preanal
scale rows, ventral counts, subcaudal counts, hemipenis length, number of trans-
verse bands, number of stripes, maxillary tooth counts, TLR, HR, HLSVL,
HWSVL, SRR1, SRR2, and SRRS. Three distinct clusters separating at distance
values >1.0 were found (Fig. 9). Samples 1-14 and 22 form the largest cluster,
with samples 15-17 and 1 8-2 1 forming two smaller clusters, respectively. Western
samples 1, 2, and 7 grouped together, sample 22 grouped with the geographically
proximate samples 9-10, but not with 1 1 or 12 (Fig. 3, 9). The significance of the
relative position of sample 6 to samples 1-10 and the split among samples 11-
14 Is unclear, especially since neither were separated in Fig. 8.

Table 5.— Variable coefficients for canonical variates I -HI for 22 samples of male striped whipsnakes ,

Masticophis.

Character

Vector I

Vector II

Vector III

Scale pit

0.1428

0.0817

0.1390

Preanal scale rows

-0.3550

0.1028

1.1623

Ventrals

0.0361

-0.0169

0.4014

TLR

-0.0304

0.0374

-0.2538

Bands

0.0732

2.6390

-0.1375

Stripes

8.2383

-0.9094

0.5162

Maxillary teeth

0.0420

0.2498

0.4193

0.0460

-0.0912

0.0365

SRR1

-0.1404

0.1409

0.2080

SRR2

0.1767

-0.0922

-1.3304

HLSVL

0.0341

-0.0474

0.1031

HWSVL

0.0326

0.0412

-0.0650

Annals of Carnegie Museum

vol. 63

SAMPLE

16
17

20
21

I I _J I I —I — I I

1.98 1.73 1.48 1.23 0.98 0.73 0.48 0.23

DISTANCE

Fig. 9. — Phenogram based on UPGMA clustering of 16 morphological characters among 22 samples
of male striped whipsnakes, Masticophis. Cophenetic correlation = 0.921.

A phylogenetic analysis based on the number of scale rows around the anterior
body, preanal scale rows, number of transverse bands, number of stripes, scale
pit number, dorsal speckling, head scale bordering, maxillary tooth counts, ventral
counts, subcaudal counts, hemipenis length, and stripe color yielded the consensus
tree shown in Fig. 10. Samples 1 8-2 1 were used for outgroup rooting. The mulpars
option of PAUP version 2.4 produced nine most parsimonious trees 19 steps long
with a consistency index of 0.842. Two monophyletic groups are evident, one
consisting of samples 1-14 and 22 and another containing samples 15-17. Place-
ment of samples within these groups and within the outgroup may not represent
an accurate phylogenetic hypothesis due to the high probability of reticulate evo-
lution among samples within groups (Thorpe, 1987). This tree differs from the
phenogram (Fig. 9) in that sample 22 was placed with samples 11-14 and not
samples 6 and 10. Otherwise the phylogenetic tree (Fig. 10) and the phenogram
(Fig. 9) have similar topologies. The presence of 1 5 dorsal scale rows around the
anterior body and at midbody is a synapomorphy uniting samples 1-17 and 22
within the M. taeniatus complex. Samples 1 8-2 1 have the pleisiomorphic con-
dition of 1 7 scale rows around anterior and midbody.

Allozyme Variation. — The protein products of 29 presumptive gene loci were

1994

Camper and Dixon— Striped Whipsnakes

Fig. 10.— Phylogenetic tree of 22 samples of Masticophis shown in Fig. 3. Consistency index = 0.842.

assayed for variation in M. taeniatus sensu lato. Twelve of the 29 loci were
polymorphic (Table 6). Masticophis t. girardi exhibited polymorphism at ten of
the 12 variable loci. Masticophis t. girardi and M. t. schotti exhibited allelic
differences in the S-Aat-A locus across their contact zone at the southern edge of
the Balcones Escarpment (Fig. 11, Table 6). Both M. t. taeniatus and M. t. girardi
are fixed for the A allele, while alleles B and C occur in M. t. schotti and alleles
A and C are present in M. t. ruthveni . Neither an M. t. girardi (TCWC 64840),
collected in Tamaulipan Biotic Province vegetation at the escarpment of the
Edwards Plateau in Uvalde County, Texas, nor an M. t. ruthveni (TCWC 65282)
from Brownsville, Texas, exhibited any protein activity at the S-Aat-A locus. The
A allele was found in only one specimen of M. t. ruthveni from Hidalgo County,
Texas, approximately 350 km south of the contact zone. The M-Aat-A locus also
exhibited variation. The B allele was present at all M. t. girardi localities repre-
sented by more than one specimen, except the Brewster County and El Paso
County, Texas, samples. This allele was also present in the homozygous state in
a specimen of M. t. taeniatus from Socorro County, New Mexico, near the in-
tergradation zone.

Genetic identity values were highest between M. t. ruthveni and both M. t.
taeniatus and M. t. girardi (Table 7). The greatest genetic distances were between
the parapatric M. t. girardi and M. t. schotti for all three measures used (Tables
7, 8). Even though M. t. ruthveni had S-Aat-A allele A, it clustered with M. t.
schotti and not M. t. taeniatus or M. t. girardi (Fig. 12).

In addition to the results of morphological and allozyme analyses, additional
evidence supporting differentiation of samples 15-17 from samples 1-14 and 22
is the absence of a broad intergradation zone between samples 14 and 15, such
as the one seen in sample 22 (see systematic accounts). Only eight specimens
exhibited intermediate morphological character states between samples 11-14

Annals of Carnegie Museum

vol. 63

Fig. 1 1.— Distribution in Texas and Mexico of genotypes of the S-Aat-A locus and morphological
hybrids. Location of hybrids denoted by an asterisk, numbers are sample sizes per locality.

and 15-17 (Fig. 1 1). The Balcones Escarpment contact zone between samples 14
and 1 5 is well-represented by specimens. Only four putative hybrids were found
there (Fig. 1 1). Four specimens having intermediate color patterns were found at
three localities in Mexico. These may represent a contact zone running northwest
to southeast between samples 12 or 13 and 15 or 17 in southeastern Coahuila
and western Nuevo Leon, respectively. They are considered putative hybrids,
even though the nearest localities for samples 1 2 and 1 3 are in central Coahuila
and central Zacatecas, respectively. This gap may be a collecting artifact. In ca-

1994

Camper and Dixon— Striped Whipsnakes

Table 6.— Allele frequencies at 12 polymorphic loci examined in the subspecies of Masticophis taeniatus.

Locus and
allele

M. i. taeniatus

M. t. girardi

M. t. schotti

M. t. ruthveni

S-Aat-A

1.000

0.500

0.800

0.500

0.200

M-Aat-A

0.833

0.632

1.000

0.167

0.316

0.053

Ck-A

1.000

0.937

1.000

0.062

Ck-C

1.000

0.929

1.000

0.071

Ddh-A

1.000

0.899

1.000

0.111

Est-D

1.000

0.967

1.000

0.033

Gapdh-A

0.800

0.937

1.000

0.200

0.062

S-Idh-A

1.000

0.971

1.000

0.029

Mpi-A

1.000

0.937

1.000

0.062

Pep-S

1.000

0.800

1.000

0.200

Pgm-A

1.000

0.969

1.000

0.031

Pk-A

1.000

0.964

1.000

0.036

Annals of Carnegie Museum

vol. 63

Table 7.— Matrix of Nei’s (1972) genetic identity above the diagonal and genetic distance below,
calculated from the 29 loci examined for the subspecies o/ Masticophis taeniatus.

Population

1 taeniatus

*****

0.997

0.967

0.989

2 girardi

0.003

*****

0.963

0.986

3 schotti

0.034

0.038

*****

0.992

4 ruthveni

0.011

0.014

0.008

*****

nonical discriminant analysis, the eight presumed hybrids did not have variable
coefficients either similar to one another or intermediate between parental forms.
Instead, Edwards Plateau hybrids grouped with either samples 1-10 or 16-17.
The Nuevo Leon hybrids grouped with samples 1 1-14 and the Coahuila hybrids
grouped with samples 1-10. These specimens show a disharmonic combination
of character states typical of interspecific hybrids (Lawson and Lieb, 1990). We
consider this strong evidence of severely restricted gene flow between samples 14
and 15 across the Balcones Escarpment, and of reproductive isolation of these
samples. The results of phylogenetic analyses show that two monophyletic groups
exist (Fig. 10), one contains samples 1—14 and 22 and the other contains samples
15-17. Therefore, samples 1-10 are considered to be Masticophis taeniatus tae-
niatus and samples 11-14 Masticophis taeniatus girardi, with sample 22 containing
specimens intermediate between them. These appear to be reproductively isolated
from sample 15, which is Masticophis schotti schotti, and samples 16-17, which
are Masticophis schotti ruthveni . Samples 18-21 correspond to Masticophis bili-
neatus.

Systematics

Masticophis taeniatus (Hallowell)

Leptophis taeniata Hallowell, 1852:181. Holotype: USNM 21 10; adult male (547 mm SVL); collected
by Dr. S. W. Woodhouse, date unknown. Type locality: New Mexico west of the Rio Grande;
restricted to Shiprock, San Juan Co., New Mexico, USA (Smith and Taylor, 1950). See Parker
(1982) for synonymies.

Definition. — A species of whipsnake with 1 5 anterior and midbody dorsal scale
rows; four dark stripes, one in the center of each of the first four dorsal scale rows
(Fig. 13); and dorsal head plates edged in cream or white (Fig. 14). Transverse
light gray bands are present in some specimens (Fig. 1 5). All specimens lack paired
light speckling on the dorsal scales and red or pink pigment immediately posterior
to the angle of the jaw. Two subspecies, M. t. taeniatus and M. t. girardi, are
recognized (see subspecies accounts).

Table 8.— Matrix of Rogers’ (1972) genetic distance, below the diagonal, and Cavalli-Sforza and
Edwards (1967) chord distance, above the diagonal, calculated from the 29 loci examined for the

subspecies o/Masticophis taeniatus.

Population

1 taeniatus

*****

0.090

0.190

0.117

2 girardi

0.026

*****

0.209

0.145

3 schotti

0.051

0.068

*****

0.115

4 ruthveni

0.030

0.046

0.022

*****

1994

Camper and Dixon— Striped Whipsnakes

I I 11 — 1 1 I

0.20 0.17 0.13 0.10 0.07 0.03 0.00

DISTANCE

Fig. 12. — Phenogram showing the relationships of the subspecies in Masticophis taeniatus sensu lato,
based on UPGMA of Cavalli-Sforza and Edwards (1967) chord distance from Table 14. Cophenetic
correlation = 0.714.

Variation.— See Table 9 for meristic variation. Variation in primary and sec-
ondary temporal scales included 39 different arrangements in M. taeniatus. The
lower primary temporal scale was divided, 3-2 pattern, in 531 (43.2%) specimens.
No divisions, 2-2 pattern, were found in 423 (34.4%) specimens. All other tem-
poral scale arrangements occurred at a frequency of <5%. No geographic com-
ponent to temporal scale variation was found.

The number of scale row reduction sites ranged from one (15-15-13) to five
(15-15-9) in both sexes of M. taeniatus. Most males (473, 70.9%) had three (15-

A B

Fig. 13.— Color pattern of adult Masticophis taeniatus taeniatus : A) pattern found throughout sub-
species’ range; B) pattern found in the southern portion of its range and in smaller specimens.

Annals of Carnegie Museum

vol. 63

15-11) reduction sites, whereas most females (446, 75.7%) have only two (15-15-
12). Two reduction sites were present in 168 (25.2%) males and three reduction
sites were present in 122 (20.7%) females. One reduction site (15-15-13) was
present in five (0.8%) males and 17 (2.9%) females. Three (0.5%) females and 19
(2.9%) males had four reduction sites (15-15-10). Five reduction sites (15-15-9),
were present in only two (0.3%) males and one (0.2%) female. Variation in SRR1
for males: X = 121 (91-151, n = 643); females: x - 122 (95-146, n = 576).
Variation in SRR2 for males: x = 132 (109-175, n = 633); females: X = 139
(1 12-180, n = 546). Variation in SRR3 for males: X = 159 (118-206, n = 460);
females: X = 164 (123-190, n = 119).

Distribution. — This species is found in the Great Basin and Chihuahuan deserts
and the Edwards Plateau of Texas (Fig. 1). Masticophis taeniatus occurs north to
the Snake and Columbia river valleys of Idaho and Washington, respectively. It
occurs east of the Cascade Mountains, Oregon, the Sierra Nevadas, California,

1994

Camper and Dixon— Striped Whipsnakes

Fig. 15.— Color patterns of adult Masticophis taeniatus girardi : A) light pattern common throughout
the Chihuahuan Desert; B) dark pattern common on the Edwards Plateau; C) intermediate pattern
found throughout the subspecies’ range.

and the Continental Divide in Mexico. It is absent from the Wasatch and Uinta
mountain ranges in Utah. In Arizona, M. taeniatus is found mainly north of the
Gila River. It ranges east into extreme western Colorado and to the Llano Estacado
in eastern New Mexico, and south to northeastern Jalisco, Mexico. The eastern
and southern range limits of M. taeniatus in Mexico are poorly understood.

The subspecies of M. taeniatus intergrade along the upper Rio Grande valley
in central New Mexico (sample 22). Specimens exhibiting intermediate color
patterns have been found from the vicinity of Albuquerque south to Las Cruces.
This intergradation zone may be widest at the southern end, stretching from
Carlsbad Caverns National Park, Eddy County, west to eastern Grant County.
However, this may be an artifact of collecting because records are lacking from
northern Catron, southern Valencia, and western Socorro counties in west-central
New Mexico. Intergrades have a striping pattern similar to that shown in Fig.
1 3B, with 0-8 (X = 4) faint transverse bands present anteriorly. Remnants of the
nuchal collar, characteristic of M. t. girardi (Fig. 1 5), occur in most intergrades.
The back, head, and stripes are usually dark gray or brown. The venter is more
heavily pigmented than in most M. t. taeniatus, but less so than in M. t. girardi.
Thirty-seven male and 3 1 female intergrades were examined.

Masticophis taeniatus taeniatus (Hallowell)

(Fig. 13, 14)

Definition and Diagnosis.— The desert striped whipsnake, M. t. taeniatus, has
1 5 dorsal scale rows anteriorly and at midbody, light borders on head scales, and
four dark longitudinal stripes. It is the only Masticophis with 15 dorsal scales
rows lacking transverse bands, dark ventral mottling, light dorsal speckling, and
red pigment at the angle of the jaw. Masticophis t. taeniatus has fewer subcaudals
and maxillary teeth than M. t. girardi.

Distribution. — Masticophis t. taeniatus has primarily a Great Basin Desert dis-
tribution (Fig. 1). It occurs south into southwestern New Mexico east of the
Continental Divide. Its range slightly overlaps the range of M. bilineatus across
central and southeastern Arizona. Masticophis t. taeniatus does not occur in Mex-
ico or Texas, contrary to what Ortenburger (1928), Parker (1982), and Conant

Annals of Carnegie Museum

vol= 63

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1994

Camper and Dixon— Striped Whipsnakes

and Collins (1991), report. Only two specimens from Washington were examined
(Appendix 1) and additional localities (Nussbaum et al., 1983) for Washington
were not confirmed. The specimen from near Oakland, Alameda County, Cali-
fornia (CAS 1 0639) is a M. t. taeniatus, but we believe its locality data are incorrect.

Color Pattern Variation. — This subspecies shows little variation in color pattern.
The head, back, and stripes are usually dark olive green, but may be brown to
almost black in some southern specimens. In many smaller, presumably younger
specimens and those from the southern part of its range, lateral stripes 2 and 4
are lighter than stripes 1 and 3, with a slight darkening between stripes 1 and 3
(Fig. 13B), but the pattern depicted in Fig. 13A is most common. The head plates
are edged with light pigment (Fig. 14). In life, the posterior quarter of the venter
and entire ventral surface of the tail are pink. The rest of the venter is usually
cream with a blurry stripe on the lateral portions of the ventrals (Fig. 1 3). More
than 90% of each sample has black spotting on the lower jaw, throat, and neck
regions (Fig. 3). Northern specimens have dark ventral pigment occurring at high
frequencies; samples 1-3 (32%-46%). Dark ventral pigmentation occurs at fre-
quencies of <15% in all other samples except sample 9 (39% with dark ventral
pigment). This may be due to the influence of M. t. girardi.

Masticophis taeniatus girardi (Stejneger and Barbour)

(Fig. 15, 16)

Masticophis ornatus Baird and Girard, 1853:102. Holotype: USNM 1971; skin of adult, sex unknown,
collected by J. D. Graham, date unknown. Type locality: USA, Texas, between San Antonio and
El Paso; restricted to Fort Davis, Jeff Davis Co. (Smith and Taylor, 1950). See Parker (1982) for
synonymies.

Coluber taeniatus girardi Stejneger and Barbour, 1917:89.

Diagnosis. — The central Texas whipsnake, M. t. girardi , differs from M. t.
taeniatus in having transverse light banding, darker dorsal and ventral coloration,
and higher subcaudal and maxillary tooth counts. It lacks the dorsal light flecking
and red pigment at the angle of the jaw that are present in M. schotti.

Distribution. —Masticophis t. girardi has a Chihuahuan Desert-western Mexican
Plateau distribution with an eastward extension onto the Edwards Plateau of Texas
(Fig. 1). The hiatus in the range of M. t. girardi in southern Coahuila, northern
Zacatecas, and northwestern San Luis Potosi, Mexico, may be a sampling artifact.
The western boundary of the range follows the continental divide with a few
localities west of it in Durango and northeastern Jalisco, Mexico. In central Texas,
the range limits of M. t. girardi closely follow the Balcones Escarpment. A Throck-
morton County, Texas, specimen (TNHC 26579) is typical M. t. girardi, but is
so far out of range that its locality data are believed to be incorrect (Dixon, 1987).

Color Pattern Variation. — The dorsal head plates are light edged in all but the
darkest specimens (Fig. 1 6). A light collar is present in all specimens, but is broken
into two light nape blotches by dark pigment in 48.9% of specimens examined
(Fig. 16). The lateral portions of the collar (nape blotches) range from one to five
scales wide. Most specimens (42.3%) have nape blotches three scales wide, whereas
40.9% have them two scales wide. The medial collar width ranges up to five scales,
with 48.9%, 23.3%, 14.0%, and 10.7% having a medial collar width of zero, one,
two, and three scales wide, respectively. Dorsal light bands, posterior to the collar,
range from zero to ten, averaging 5.3 in 479 specimens. No geographic variation
in band number was found.

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vol. 63

The dorsal color patterns illustrated in Fig. 1 5 represent the ends of a continuum
of variation. Fig. 15C represents a common intermediate condition. Pattern 15A
differs from 15C in that some light areas are present between stripes, especially
stripes 3 and 4. Color patterns were scored as belonging to one of these three

Table 10.— Dorsal color pattern frequencies found among samples 11-14 of Masticophis taeniatus
girardi shown in Fig. 3. Pattern types refer to Fig. 15.

Sample

Pattern

Light ( 1 5 A)

Dark (15B)

Intermediate ( 1 5C)

1 1

64.70%

16.81%

18.48%

81.08%

2.70%

5.41%

88.00%

4.00%

8.00%

13.70%

60.27%

15.34%

1994

Camper and Dixon— Striped Whipsnakes

patterns (Fig. 1 5). The light pattern (Fig. 1 5A) is more common throughout the
Chihuahuan Desert (samples 11-13, Table 10). The dark pattern (Fig. 15B) is
dominant on the Edwards Plateau (sample 14), occurring at low frequencies in
samples 11-13. Specimens scored as intermediates (Fig. 15C) were most common
in the northern part of the range (samples 1 1 and 1 4) and occur at low frequencies
in samples 12-13 (Table 10). The anterior ventral pattern of M. t. girardi consists
of black pigment covering >50% of the chin shields, gulars, and anterior ventrals
in >99% of specimens. The belly is mostly black in >92% of specimens. The
subcaudals have dark pigment confined to the lateral edges in >88% in samples
11-14 (Fig. 3). The posterior quarter of the venter and the subcaudals are pink
in life. In dark specimens, the pink may be reduced to only the small areas between
the dark blotches on the ventrals.

The predominance of the dark pattern (Fig. 15B) on the Edwards Plateau may
be associated with thicker vegetation such as Juniperus, Five melanistic specimens
of M. t. girardi were found. One such Edwards Plateau specimen is an adult female
1090 mm SVL (CAS 103476) from Yal Verde County, Texas. Three additional
specimens (CM 48192, LACM 116256, and KU 80828) are from the Cuatro
Cienegas basin in central Coahuila, Mexico (Camper and Dixon, 1990). Another
specimen (KU 39567) is an adult female (938 mm SVL) from western Coahuila,
Mexico.

Masticophis schotti (Baird and Girard)

Masticophis schotti Baird and Girard, 1853:160. Holotype: USNM 1972, adult male (943 mm SVL)
collected by Arthur Schott, date unknown. Type locality: Eagle Pass, Maverick Co., Texas, USA.
See Parker (1982) for synonymies.

Definition. — The only species of Masticophis with 15 anterior and midbody
dorsal scale rows, paired white, cream, or yellow spots on the anterior comers of
the dorsal scales, uniformly dark head plates, and red pigment immediately pos-
terior to the jaw angle. Two subspecies, M. s. schotti and M. s. ruthveni, differ
from one another primarily in color pattern. See subspecies accounts for descrip-
tions of type specimens and color patterns.

Variation.— See Table 9 for meristic variation. Thirty-one temporal scale pat-
terns were found. The nondivided 2-2 pattern was the most frequent, 35.3% (102
specimens). The lower primary was divided, 3-2 pattern, in 69 (23.9%) specimens.
The lower primary and secondary temporals were fused in 33 (1 1.4%) specimens.
Other temporal scale arrangements existed at frequencies < 5% with no geographic
variation discernible.

The number of scale row reduction sites ranged from one (15-15-13) to four
( 1 5 - 1 5 - 1 0) in males and from one to three (15-15-11) in females. Most males (9 6 ,
54.9%) had two (15-15-12) reduction sites, whereas most females (97, 74. 1%) had
only one (15-15-13). Two reduction sites were present in 30 (22.9%) females and
one reduction site was present in 24 (13.7%) males. Three reduction sites (15-15-
1 1) were present in 53 (30.3%) males and four (3.1%) females. Two (1.1%) males
had four reduction sites (15-15-10). Variation in SRR1 in males: x = 119 (88-
139, n = 165); females: x = 1 17 (76—138, n — 118). Variation in SRR2 in males:
x = 151 (112-184, n = 137); females: x = 157 (115-186, n — 30). Variation in
SRR3 in males: x = 159 (136-190, n = 49); females: x = 161 (154-172, n = 4).

Distribution. — This species is found from the Balcones Escarpment of central
Texas south to the Gulf Coastal Plain and inland onto the Mexican Plateau (Fig.
1). The western limits of its range in northern Mexico are unclear.

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vol. 63

Fig. 17.— Color pattern of adult Masticophis schotti schotti.

Masticophis schotti schotti (Baird and Girard),

new combination

(Fig. 17, 18)

Definition and Diagnosis. —Masticophis s. schotti has 15 anterior and midbody
dorsal scale rows, paired cream-colored light spots on the anterior dorsal scale
comers and invariably two prominent light stripes. It differs from M. s. ruthveni
in having lighter ground color, lighter dorsal speckling, and a more prominent
upper light stripe that always extends past ventral 110.

Distribution. — The geographic range of M. s. schotti is apparently restricted to
the Tamaulipan Biotic Province of Blair (1949; Fig. 1). Few specimens from
Mexico exist, and the western range limits in the Coahuila Folded Belt Region
are unclear.

Color Pattern Variation. — Adult M. s. schotti are invariant in color pattern (Fig.
17). The grayish-green ground color does not fade in preservative. In life, the
subcaudals are pink in most specimens, except for their lateral tips which are
grayish-green. Grayish-green speckling is present on the subcaudals in 15.0% of
M. s. schotti examined. The red pigment behind the jaw angle fades in preservative.

Masticophis schotti ruthveni (Ortenburger),

new combination

(Fig. 18, 19)

Masticophis ruthveni Ortenburger, 1923:2. Holotype: UMMZ 57681, adult male (1104 mm SVL),
collected by A. I. Ortenburger, date unknown. Type locality: Brownsville, Cameron Co., Texas,
USA. Paratypes: UMMZ 57682-57684 and 57686-57694, UIMNH 43493, and MCZ 62561.

1994

Camper and Dixon —Striped Whipsnakes

Fig. 18.— Head color pattern of adult Masticophis schotti and Masticophis bilineatus.

Masticophis taeniatus australis Smith, 1941:390. Holotype: USNM 10240, juvenile female, collected
by A. Duges in 1879. Type locality: “Guanajuato,” Guanajuato, Mexico. New synonymy.

Definition and Diagnosis.— Ruthven’s whipsnake, M. s. ruthveni, has 1 5 anterior
and midbody dorsal scale rows, paired dorsal spotting, and a variable color pattern.
Because the holotype of M. t. australis is a typical juvenile M. s. ruthveni, we
herein synonymize the former. It differs from M. s. schotti in having narrower,
fainter stripes that are variable in number, darker dorsal spots, and a more variable
ground color that is usually darker. The upper light stripe does not extend to
ventral 110.

Distribution. —Masticophis s. ruthveni is found on the Gulf Coastal Plain from
the Rio Grand south to central Veracruz, Mexico (Fig. 1). Gaps along the east
coast of Mexico in Tamaulipas and Veracruz are believed to result from collecting
bias. This taxon ranges west into the Sierra Madre Oriental and onto the southern
part of the Mexican Plateau west to northern Michoacan. The western range limits
of M. s. ruthveni are unclear. Two putative M. t. girardi x M. s. ruthveni hybrids

30 Annals of Carnegie Museum vol. 63

A B C

Fig. 19.— Color patterns of adult Masticophis schotti ruthveni : A) two-striped pattern most common
throughout the subspecies’ range; B) four-striped pattern found uncommonly throughout the subspe-
cies’ range; C) rare stripeless pattern.

from central Nuevo Leon indicate that M. s. ruthveni may not occur as far west
in northern Mexico as it does on the southern part of the Mexican Plateau. In
the USA, this subspecies occurs in Cameron, Hidalgo, Starr, Willacy, and south
central Kenedy counties, Texas. The juvenile identified as M. s . schotti by Martin
(1958) from La Joya de Salas, Tamaulipas, Mexico, (UMMZ 1 10818) is a M. s.
ruthveni, as is UIMNH 3836 from 25 miles south of Monterrey, Nuevo Leon,
Mexico (Shannon and Smith, 1949).

Color Pattern Variation. — Specimens exhibiting only the lower light stripe were
the most common, occurring at a frequency of 56.8% (Fig. 19 A; Table 11). Two
light stripes were found in 37.5% of M. s. ruthveni (Fig. 19B). The upper light
stripe, occurring on the upper margin of scale row 3 and lower margin of scale
row 4, is narrow and faint. It extended as far posterior as ventral 1 10 (M. s. schotti
character state) in only four (3.5%) M. s. ruthveni. The 5.8% of M. s. ruthveni that
lacked stripes had a dark olive, almost black, ground color with bright yellow
dorsal speckling (Fig. 19C). Stripeless specimens occur in southern Tamaulipas
and eastern and southwestern San Luis Potosi, Mexico. Little geographic variation
in color pattern was found. The frequencies of the one- and two-striped morphs
are equal in the lowland sample 16 (Table 1 1). The ventral color pattern varies
from dense speckling on the lateral quarter of the ventrals to sparse speckling
toward the ventral midline, except in stripeless specimens, where dense ventral
speckling is present throughout. The venter is the same color as the dorsum in
striped specimens. In stripeless specimens, it is lighter toward the midline. Dorsal

Table 1 1.— Dorsal color pattern frequencies found among samples 16-17 Masticophis schotti ruthveni
shown in Fig. 3. Pattern types refer to Fig. 19.

Pattern

Sample

1 -stripe ( 1 9 A)

42.86%

67.80%

2-stripes (19B)

42.86%

22.03%

0-stripes ( 1 9C)

14.29%

10.17%

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Camper and Dixon— Striped Whipsnakes

pigment extends ventrally to the lateral subcaudal tips. Medially, the subcaudals
are pink in living specimens.

Masticophis hilineatus Jan
(Fig. 18, 20)

Masticophis hilineatus Jan, 1863:65. Holotype: MTKD 15523, female (467 mm SVL), date of collection
and collector unknown. Type locality: Restricted to Guaymas, Sonora, Mexico (Smith and Taylor,
1950). Restriction of type locality to Casas Grandes, Chihuahua by Schmidt (1953) is in error
(see remarks).

Bascanium semilineatum Cope, 1891:626. Holotype: USNM 1981, female, 816 mm SVL, collected
by A. Schott, date unknown. Type locality: Colorado River bottom, Arizona.

Zamenis semilineatus: Gunther, 1894:121.

Bascanion semilineatum: Van Denburgh, 1896:347.

Coluber semilineatus: Stejneger and Barbour, 1917:80.

Masticophis semilineatus: Ortenburger, 1923:2.

Coluber hilineatus: Bogert and Oliver, 1945:362.

Masticophis hilineatus lineolatus Hensley, 1950:272. Holotype: UIMNH 5611, male, 1011 mm SVL,
collected by M. Hensley, 23 May 1949. Type locality: North branch of Alamo Canyon, Ajo
Mountains, 12.9 mi S and 5 mi E of the Ajo-Tucson-Sonoyta junction, Pima Co., Arizona. New
synonymy.

Masticophis hilineatus slevini Lowe and Norris, 1955:93. Holotype: SDNHM 3826, female, 833 mm
SVL, collected by Mrs. G. Bancroft, 18 April 1930. Type locality: Isla San Esteban, Gulf of
California, Sonora, Mexico. New synonymy.

Definition and Diagnosis. — Masticophis hilineatus is the only striped Mastico-
phis with a combination of 1 7 anterior and midbody dorsal scale rows, two dark
lateral stripes on each side of the body, and paired light spots on the anterior
comers of the dorsal scales. Because the previously described subspecies do not
differ from other populations in the characters on which their descriptions were
based, we herein synonymize them with M. hilineatus. It differs from M. taeniatus
and M. schotti in having more dorsal scale rows. It differs from other striped
whipsnakes that have 1 7 dorsal scale rows in dorsal ground color and by possessing
paired light spots on the dorsal scales.

Description of Holotype.— Dorsal scale rows 17-17-13. A portion of the venter
is missing and the tail is incomplete so ventral and subcaudal counts were not
made. A paratype, (MTKD 15068) an adult female (748 mm SVL) has 196
ventrals, 130 subcaudals and 17-17-13 dorsal scale rows. The paratype has a
dorsal scale row reduction formula as follows:

17J^n|15 — 8<i18) 146 + 7(-124) 13(196)

Scale row reduction sites of the holotype involve the same dorsal scale rows as
those of the paratype. The holotype has 20 maxillary teeth and one apical scale
pit. It has a divided anal plate; supralabials 8-9; infralabials 11-10; preoculars
2-2; postoculars 2-2; and loreals 1-1. Supralabials 4 and 5 contact the right orbit
and supralabials 5 and 6 contact the left orbit. Supralabial 3 on the left side is
divided. Both right primary temporals are divided, 4-2 pattern, and all but the
lower left secondary are split, 4-3 pattern.

The color pattern is characteristic for the species (Fig. 20B). Two dark stripes
are present, the lower stripe is located on the lower half of scale row 1 and adjacent
tips of the ventrals, and the upper stripe is located on the upper half of scale row
2 and lower half of scale row 3. The dorsal scales have paired light spots at their
anterior comers. The dorsal coloration is dark olive green fading to light green

Annals of Carnegie Museum

vol. 63

A B

Fig. 20.— Color pattern of adult Masticophis bilineatus : A) lighter pattern more common in larger
specimens; B) darker pattern more common in smaller specimens.

posteriorly. The light stripe between the upper dark stripe and the middorsum
does not fade anterior to ventral 1 10. The head plates are uniformly colored the
same shade as the dorsum (Fig. 18). The venter is immaculate except for the dark
spots present on the chin shields, gulars, and first 10-15 ventrals. The subcaudals
are also immaculate, except for their lateral tips, which are the same color as the
dorsum.

Remarks. — We believe that Schmidt (1953) is in error in restricting the type
locality of M. bilineatus to Casas Grandes, Chihuahua, Mexico. This locality is
east of the continental divide where this species does not occur.

Variation. — See Table 9 for meristic variation. Thirty-one right temporal scale
patterns were recorded for M. bilineatus. A divided lower primary, 3-2 pattern,
occurred in 72 (22.4%), 64 (19.9%) had all but the lower secondary divided, 4-3
pattern. No temporals were divided in 52 (16.2%), both primary temporals were
divided, 4-3 pattern, in 30 (9.4%). Other temporal scale patterns occurred at
frequencies <5%. No geographic component to temporal scale variation was
found.

The number of scale row reduction sites ranged from two to five in males and
from two to six in females. Most males (150, 91.5%) and females (155, 95.1%)
had three reduction sites (17-17-13). Eleven (6.7%) males and five (3.1%) females
had four (17-17-12) scale row reduction sites. One (0.6%) male and two (1.2%)
females had only two (17-17-14) reduction sites. Two (1.2%) males had five (17-
17-11) reduction sites and one (0.6%) female had six (17-17-10) reduction sites.
Variation in SRR1 for males: x = 1 15 (81-136, n = 158); females: x = 1 14 (91-
129, n = 153). Variation in SRR2 in males: x = 123 (91-167, n = 154); females:
x = 121 (93-154, n = 147). Variation in SRR3 in males: x = 130 (108-192, n =
117); females: x- 127 (95-189, n - 121).

1994

Camper and Dixon —Striped Whipsnakes

Distribution. — In the north, M. bilineatus has a Sonoran Desert distribution
that interdigitates with that of M. t. taeniatus in eastern and central Arizona.
Sympatric populations may occur in southwestern Yavapai County, western Gila
County, and southern Graham County, Arizona. Masticophis bilineatus and M.
t. girardi appear to be parapatric in western Durango and northern Jalisco, Mexico.
In New Mexico, USA, and northern Sonora, southern Zacatecas, northern Jalisco,
and Aguascalientes, Mexico, the eastern range limits of M. bilineatus follow the
Continental Divide. From the Sonoran Desert it occurs south along the Gulf of
California lowlands to Colima with an eastward range extension onto the Mexican
Plateau in Jalisco, Aguascalientes, and Zacatecas. The specimen supposedly from
Isla San Pedro Martir (UO 32528) is probably from Isla San Esteban (D. Hews,
personal communication). Two specimens, both labelled MCZ 4582, from San
Pedro, Coahuila, Mexico, are M. taeniatus and M. bilineatus. Since the latter is
so far out of range, the locality data are believed to be incorrect. The same is true
for USNM 46499, purportedly from Cuicatlam, Oaxaca, Mexico.

Color Pattern Variation. — The dorsal color pattern of M. bilineatus varies little,
with both patterns present in specimens of all sizes and from all of its range (Fig.
20). The head plates of M. bilineatus are uniformly dark and identical in shade
to the rest of the dorsum (Fig. 18). Pattern 20B is most common in smaller
individuals, whereas pattern 20 A is more common in larger specimens. An adult
male (TCWC 21903, 733 mm SVL) from Mazatlan, Sinaloa, Mexico, has an
aberrant color pattern. The light line between the upper dark stripe and the back
is absent, forming a uniformly dark dorsum starting at the upper half of scale row
2 on each side. Dark pigment is present on the lateral tips of the ventrals and on
the ventral and posterior edges of each scale in row 1 , forming a narrow diffuse
lower dark stripe. The paired cream-colored spots at the anterior comers of the
dorsal scales are present in this and all adult M. bilineatus.

The ventral color pattern of M. bilineatus varies little, with dark spotting present
on the chin shields, gulars, and first 10-15 ventrals in 131 (45.6%) specimens,
and absent in 168 (54.4%). Ventral spots form a median row that usually extends
10-15 ventrals posterior to the head, but in the Isla San Esteban population it
extends approximately 40 ventrals farther posteriorly. Dark spotting on the belly
was present in only 37 (12.0%) specimens, whereas 272 (88.0%) lacked belly
spotting. Dark spotting was present on the subcaudals of 1 37 (44.3%) M. bilineatus,
175 (55.7%) had dark pigment on the lateral subcaudal tips only. Subcaudal
spotting occurred at a frequency of about 50% in samples 18 and 19, but was
virtually absent from samples 20 and 21 (Fig. 3).

Discussion

Masticophis taeniatus and M. schotti are well-differentiated morphologically.
Many color pattern differences are obvious and were weighted highest by the
canonical discriminant analysis (see species accounts). Additionally, M. taeniatus
and M. schotti differ in a number of meristic and mensural character states. The
differences are greatest between M. t. girardi and M. s. schotti across the southern
Balcones Escarpment. They differ in M. t. girardi having a higher frequency of
three SRR sites, two apical scale pits, higher maxillary tooth counts, significantly
higher mean ventral counts, and significantly smaller mean HWSVL. The Balcones
Escarpment is a well-known geographic boundary for many reptile species, in-
cluding 37 species of snakes (Smith and Buechner, 1947). The occurrence of M.

Annals of Carnegie Museum

vol. 63

t. girardi (TCWC 64840) off the Edwards Plateau in Tamaulipan desert vegetation
establishes the existence of a narrow zone of sympatry between M. t. girardi and
M. s. schotti. The locality is about 38 km southwest of the nearest M. t. girardi
locality and approximately 33 km northwest of the nearest M. s. schotti locality.
A hybrid specimen (TCWC 49204) is known from a locality 20 km to the east.
Increases in ventral counts in M. s. ruthveni and decreases in maxillary tooth and
ventral counts of M. t. taeniatus away from the contact zone may indicate character
displacement (Mayr, 1970). Discordant patterns of character state distributions
are found in intermediate specimens from this area, indicating that they may be
interspecific hybrids (Lawson and Lieb, 1 990). Evidence suggests that the southern
Balcones Escarpment may be a narrow hybrid zone (sensu Moore, 1977).

Two additional contact zones may exist between M. taeniatus and M. schotti.
The first of these is the previously mentioned area of northeastern Mexico, where
four putative hybrids were found. However, this is problematical, because no
specimens of M. t. girardi are known as yet from near this locality. The second
is a M. t. girardi (CAS 165260) from northeastern Jalisco which is sandwiched
in an area of parapatry between the ranges of M. s. ruthveni and M. bilineatus,
suggesting a potential contact zone involving all three species.

Despite character state similarities among these three species, and areas of
sympatry involving M. taeniatus and M. bilineatus, no hybrids involving the latter
species were found. Masticophis bilineatus differs from the other two species in
having the plesiomorphic character state of 17 dorsal scale rows. Masticophis
taeniatus and M. schotti are shown to be sister species on the basis of the synapo-
morphic condition of 1 5 dorsal scale rows. Little intraspecific variation in anterior
and midbody dorsal scale rows are known for any Masticophis. Thus, M. bilineatus
is considered the most divergent of the three species examined.

Southward decreases in ventrals of M. bilineatus (Fig. 4, 5) as found in this
study were also reported by Ortenburger ( 1 928). Ventral variation among samples
18-21 indicates a step cline, whereas subcaudal variation is more gradual (Fig.
4-7). Differences between snake populations inhabiting Sonora and Sinaloa were
discussed for M. flagellum (Wilson, 1970) and Hypsiglena (Dixon and Dean,
1986). The latter authors identified a hybrid zone in the foothills along the Rio
Fuerte in northern Sinaloa that is a transition area from lower Sonoran Desert
vegetation to tropical thorn scrub. Two male M. bilineatus from the lowlands
along the Rio Fuerte agree with northern (Sonoran Desert) specimens in ventral
number. Perhaps the contact zone between the low and high ventral count forms
is farther east in the foothills, as in Hypsiglena, or immediately to the south of
the Rio Fuerte. However, no specimens of M. bilineatus and Hypsiglena are
available from between Los Mochis and Guamuchil, Sinaloa (Dixon and Dean,
1986). The southern low- ventral form of M. bilineatus may constitute a separate
subspecies (C. H. Lowe, personal communication). However, multivariate anal-
yses did not separate these samples (Fig. 8, 9) and Masticophis bilineatus is there-
fore considered monotypic.

Genetic differentiation is low among the taxa examined here. Since other snake
species are known to show low levels of genic differentiation (Gartside et al., 1977;
Murphy and Ottley, 1980; Murphy, 1983; Murphy and Crabtree, 1985; Lawson,
1987; Lawson and Lieb, 1990), this is not indicative of measurable gene flow.
Murphy (1983) reported Nei’s genetic identity values of 0.89 between Masticophis
lateralis and M. aurigulus, and 0.94 between Crotalus ruber and C. catalinensis.
Speciation with little genic differentiation is well-known among many taxa (see

1994

Camper and Dixon— Striped Whipsnakes

citations in Gartside et al., 1977). They reported Rogers’ S values of 0.91-0.94
with little morphological differentiation between the ribbon snakes, Thamnophis
proximus and T. sauritus. Rogers’ S values among the morphologically better-
differentiated M. taeniatus and M. schotti range from 0.93-0.97. Degree of genic
differentiation is more closely associated with time since divergence (Avise and
Ayala, 1976). Differentiation of M. schotti and M. taeniatus may have resulted
from a Recent speciation event, possibly divergence in allopatry during habitat
changes brought about by Wisconsin glaciation, with secondary contact occurring
presently along the Balcones Escarpment. Populations of whipsnakes now known
as M. taeniatus may have been isolated in the Chihuahuan Desert refugia of north-
central Mexico, while the ancestors of M. schotti were restricted to the Tamaulipan
grassland refugium of eastern Mexico (Morafka, 1977). However, because M.
taeniatus and M. schotti are sister species, one cannot distinguish between a
primary and secondary origin for the Balcones Escarpment contact zone (Wiley,
1981).

The existence of the A allele of the S-Aat-A locus in one specimen of M. s.
ruthveni from extreme southern Texas is problematical. Three possibilities exist.
First, the presence of this allele in M. schotti may be the result of introgression
from M. t. girardi in Mexico. This may occur at one of the two potential contact
zones previously mentioned. With the apparent absence of M. t. girardi near the
potential contact zone in northern Mexico, and a lack of intermediate specimens
from the south-central Mexican plateau area, little evidence in support of this
hypothesis exists. Second, the observed pattern of geographic variation in alleles
at this locus may be a function of small sample size, with all alleles being present
in both species. It has been shown that sample sizes as small as those used here
are adequate for estimating genetic divergence (Nei, 1978; Gorman and Renzi,
1979). However, answering this question necessitates a population genetics ap-
proach and not simply a measure of genetic divergence. Even though sample sizes
are insufficient for population genetic studies of these species, the fixed difference
across the Balcones Escarpment contact zone appears real. Evidence for this
hypothesis lies in the step clinal pattern of variation of several morphological
characters and the paucity of specimens with intermediate phenotypes or geno-
types. Finally, the A allele of the S-Aat-A locus may be a residual allele common
to the ancestor of M. schotti and M. taeniatus such as that hypothesized for
Anguilla by Avise et al. (1990). It appears that there is character state divergence
at the S-Aat-A locus with respect to the Balcones Escarpment contact zone, with
the A allele not yet extinct in M. schotti populations located away from the contact
zone. Since M. schotti is variable at this locus, with the A allele still present, Recent
divergence with differential selection pressures within each of these two species
is suspected. The last hypothesis is preferred, since it is most concordant with the
more conclusive morphological data set. However, more specimens from through-
out the ranges of both species must be assayed for genetic variation in order to
fully test these hypotheses.

Although we have not previously discussed the relationship of Masticophis to
other related genera, for example Coluber, Mastigodryas, Dendrophidion, and
possibly Drymobius, at this time we do not propose the synonymy of Masticophis
and Coluber. It is obvious to us that the latter genera may well be congeneric, as
suggested by Schatti (1986, 1987) and Stejneger and Barbour (1917). We suspect
that Masticophis sensu stricto is not a valid genus because of the evidence from
osteology and hemipenis morphology presented by Schatti (1986, 1987).

Annals of Carnegie Museum

vol. 63

Acknowledgments

We would like to thank the following for live specimens or tissue samples: R. Baker, B. Buchanan,

J. Campbell, L. Densmore, C. Farquhar, R. Green, C. Harrison, R. Jennings, J. Johnson, B. Keeley,

K. King, W. Lamar, T. Lee, C. Lieb, R. Macken, R. Martin, H. McCrystal, P. Mckeown, A. Price, A.
Rentfro, N. Scott, R. Wallace, R. Worthington, and the Texas Herpetological Society. Special thanks
go to R. and B. Houston, who graciously gave us access to their land in Llano County, Texas. Marc
Allard, G. Baumgardner, H. Camper, J. Derr, I. Greenbaum, H. Greene, B. Hanks, M. Morris, A.
Price, M. Retzer, K. Reed, M. Sims, S. Smith, and P. Sudman provided help during data collection
or analysis. We thank the following institutions (and curators) for loans of specimens: American
Museum of Natural History (AMNH, C. Myers, R. Zweifel), Academy of Natural Sciences, Phila-
delphia (ANSP, J. Cadle, E. Malnate), Angelo State University (ASC, M. Engstrom), Arizona State
University (ASU, M. Douglas), Baylor U ni versity-Strecker Museum (SM, D. Lintz), Brigham Young
University (BYU, J. Sites), Chicago Academy of Sciences (CA, R. Vasile), California Academy of
Sciences (CAS, R. Drewes, J. Vindum), Carnegie Museum of Natural History, Pittsburgh (CM, E.
Censky, C. J. McCoy), Cornell University (CU, K. Adler, R. Schoknecht), Dallas Museum of Natural
History (DMNH, K. Krumke), Drake University (DURC, J. Christiansen), Ernest A. Liner (EAL),
Field Museum of Natural History, Chicago (FMNH, H. Marx), Fort Worth Museum of Science and
History (FWM, W. Voss), Illinois Natural History Survey (INHS, K. Cummings, L. Page, M. Retzer),
University of Kansas Museum of Natural History (KU, J. Collins, W. Duellman), Los Angeles County
Museum (LACM, R. Bezy, J. Wright), Harvard University-Museum of Comparative Zoology (MCZ,
P. Alberch, J. Rosado), Kerr Wildlife Management Area, Kerrville, Texas (KWMA), Louisiana State
University-Museum of Zoology (LSUMZ, D. Rossman), Midwestern State University (MU, F. Stangl),
National Museum of Natural History (USNM), R. McDiarmid, R. Reynolds, G. Zug), New Mexico
State University (NMSU, J. Lapointe), San Diego Natural History Museum (SDNHM, G. Pregill),
Southern Illinois University (SIUC, R. Brandon, M. Morris), Southwestern Research Station, Portal,
Arizona (SWRS, W. Sherbrooke), Staatliches Museum fur Tierkunde (MTKD, F. Obst), Sul Ross State
University (SRSU, J. Scudday), Texas A&I University (TAIC, A. Chaney, S. Smith), Texas A&M
U ni versity-T exas Cooperative Wildlife Collections (TCWC, M. Retzer, K. Friend), Texas Tech Uni-
versity Museum (TTU, C. Jones), Tulane University (TU, H. Dundee), University of Arizona (UAZ,
G. Bradley, C. Lowe), University of California at Berkeley-Museum of Vertebrate Zoology (MVZ, D.
Good, H. Greene), University of Florida-Florida State Museum (FSM, D. Auth), University ofldaho-
Museum of Natural History (UIM, R. Wallace), University of Illinois-Museum of Natural History
(UIMNH, L. Maxson), University of Michigan Museum of Zoology (UMMZ, A. Kluge), University
of New Mexico-Museum of Southwestern Biology (UNM, H. Snell), University of Northern Arizona-
Museum of Northern Arizona (MNA, D. Howard), University of Oklahoma-Stovall Museum (UO,
J. Krupa), University of Southwestern Louisiana (USL, J. Jackson), University of Texas-Texas Me-
morial Museum (TNHC, R. Martin), University of Texas at Arlington (UTACV, J. Campbell, J.
Darling), University of Texas at El Paso (UTEP, C. Lieb), University of Utah (UU). In particular, we
would like to thank J. V. Vindum (CAS) for calling our attention to four recently accessioned specimens
from Mexico in his care. We thank Jose Luis Villarreal B. for translating the abstract. This research
was funded by the Texas Agricultural Experiment Station and a Texas A&M University Faculty-Staff
minigrant.

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Appendix 1

Specimens Examined

(See Acknowledgments for explanation of
museum abbreviations)

Masticophis taeniatus . -UNITED STATES: ARIZONA: Apache Co., Chinle (MCZ 8972), Concho

(SDNHM 44192), 1 mi S Hat (UNM 11976), Petrified Forest National Park (UTACV 14342), 2 mi

N Springerville (BYU 16673), 4 mi E St. Johns (MVZ 9327), 4 mi S St. Johns (UAZ 39492), 1 1 mi

S jet US 666 & AZ 61 (UAZ 27270); Cochise Co., 8 mi S. Willcox (NMSU 1965); Coconino Co., no

1994

Camper and Dixon— Striped Whipsnakes

locality (FMNH 25721, UAZ 25726), Bright Angel Canyon (UAZ 25716-25717), Buckfarm Canyon
(MNA Z7.2405), Coal Mine Canyon (CA 1 5905), Glen Canyon (UAZ 39719), Grand Canyon (AMNH
63857, 97792, MNA Z7.2827, SDNHM 3158-31 59, USNM 50953, 239259-239260), 22 mi S Cam-
eron (MVZ 25484), Coleman Lake (MVZ 29083), 13 mi N Deadman’s Flat (MZ 8680), 1 1 mi E
Desert View (UAZ 34753), Flagstaff (LACM 27773), 17 mi ENE Flagstaff (MNA Z7.624), 28 mi N
Flagstaff (BYU 22113), 3 mi N Fredonia (UAZ 25723), 5.7 mi S Fredonia (UAZ 25731), 14 mi E
Jacob Lake (CA 10123), 42 mi SW Kayenta (CM 66657), Oak Creek (CAS 35235), Rainbow Lodge
(MVZ 17885), San Francisco Mtns (MNA Z1.44), 15 mi E Seligman (LACM 20592), 10 mi N Sunset
Crater National Monument (UTEP 4123), 15 mi N The Gap (SM 3961), 12 mi SE Tuba City (CAS
156190), 19 mi SE Tuba City (AMNH 114528), 24.7 mi SE Tuba City (UAZ 25720), Two Guns
(SDNHM 32324), Wupatki National Monument (INHS 6500, LACM 20591, MNA Z7.5, Z7.656),
2.6 mi N US 89 on alt US 89 (UAZ 25730); Gila Co., no locality (USL 8946), Crook National Forest
(UIMNH 72093), 1 1 mi N Miami (FMNH 106182-106283), Pine (BYU 1 1291), 3 mi NW Pine (CA
17900), Pioneer Peak S of Globe (UIMNH 72092), Pinal Mtns (UAZ 25722), 8 mi W Roosevelt
(UAZ 25725), Roosevelt Reservoir (USNM 105228), 4.3 mi S Rose Creek Campgrounds (UIMNH
72094), Rose Creek Lodge (MVZ 49918), near San Carlos (CM 26675), Sierra Ancha Experimental
Forest-SAEF (CM 53796), SAEF T4.5N R21E Sec 31 (ASU 10534-10535), 7 mi SE SAEF Head-
quarters (ASU 1 1299), Cams Ranch in SAEF (MVZ 6309), Tick Rat (ASU 10533), 6.8 mi N Young
(UAZ 35973); Graham Co., Graham Mtns (USNM 51764), 7.5 mi E Klondyke (AMNH 111226),
San Carlos Indian Reservation (CAS 158216-158217), Santa Teresa Mtns (SDNHM 62706); Greenlee
Co., Blue Mtns (UNM 22319-22320), 40 mi NW Clifton (UAZ 25724), 7 mi SW New Mexico state
line on AZ 78 (UNM 33141); Mohave Co., Cerbat Mtns (UAZ 40571), 7.6 mi E Hackberry (MVZ
170750), 16.3 mi S & 9.5 mi E Hackberry (UAZ 25734), Hualpai Mtn Park (TCWC 9341), 32 mi E
Kingman (MVZ 175138), 35 mi W Kingman (TCWC 9340), Mt Trumbull (UAZ 25719), 1 1.8 mi E
jet US 40 & AZ 93 on US 40 (LACM 135203), 8.2 mi W jet Main St Valley Rd & Virgin Mtn Rd
(UAZ 257 15), 15.1 mi E jet Main St Valley Rd & Navajo Trail (UAZ 25718), 13.5 mi E Yucca (MNA
Z7.656); Navajo Co., 12 mi W Cibecue (MCZ 101217), New Orabi (AMNH 110454, UAZ 25727),
Paiute Canyon (MVZ 17884), S of Taylor (BYU 13599); Pinal Co., SE Superior (UIMNH 72095);
Yavapai Co., Bradshaw Mtns (AMNH 63978), 9.8 mi NW Congress (UIMNH 71995), Cottonwood
(ASU 208223, CM 66655, FMNH 208223), 4 mi SW Cottonwood (CM 66656), Granite Dells (UAZ
9333-9334), 3 mi W Iron Springs (UAZ 25732), Milk Creek (UAZ 25728), Mingus Mtn (CM 66660),
1 mi N Montezuma’s Castle National Monument (EAL 3084), Peeples Valley (UIMNH 72096),
Prescott (ASU 1614, UAZ 25729, USNM 15704), 1 2 mi SW Prescott (AMNH 126735), 13.6 mi NNE
Prescott (CM 66659), 4 mi W Sedona (UIMNH 43200), 7 mi W Sedona (CM 66661), Whipple (MCZ
5865, USNM 1 1422-1 1423), Yamell (CA 3151, SDNHM 32325), 1 mi S Yamell (SDNHM 5290),
6 mi N Yamell (CA 3468); Yuma Co., 3 mi S & 6.2 mi W Gladden (UAZ 25735), Kofa Mtns (UNM
41621, USNM 221843-221844). CALIFORNIA: Alameda Co., near Oakland (CAS 10639); Inyo Co.,
Argus Mtns (SDNHM 17061), 18 mi N Cowhom Valley (SDNHM 48083), 5 mi W & 1.25 mi S
Independence (MVZ 38410-3841 1), 6 mi W Independence (AMNH 1 10680); Panamint Mtns (MVZ
19256, 64131); Lassen Co., 9 mi E Amedee (MVZ 24579), Bare Creek Ranch (LSUMZ 8802-8803),
Fort Sage Mtn (MVZ 20481), Lassen National Park (MVZ 68313), 5 mi E Litchfield (MVZ 36136),
6 mi E Susanville (UMMZ 91821); Modoc Co., 6 mi N Eagleville (LSUMZ 8804); Mono Co., 2 mi
S Benton (MVZ 3714), between Hot Creek & Mammoth Jet (CAS 8395), near Hot Creek geysers
(SDNHM 19357-19360), Sierra Nevadas (LACM 2270), Topas Lake (SDNHM 28860), Whitmore
Tubs (SDNHM 34240); San Bernardino Co., Chemehuevi Mtns (LACM 2272-2274), Clark Mtn
(MVZ 28564-28566, 41703), 1 1.2 km S Lima (MVZ 100271), Midhills Campground (MVZ 150189);
Siskiyou Co., Ash Creek at Klamath River (UNM 26436), 3 mi S & 1 mi E Capco (MVZ 85073),
Gazelle (CAS 36063), 15 mi NE Weed (MVZ 17259). COLORADO: Garfield Co., 6 mi N Mesa Co
line (UIMNH 80484), 7 mi W Rifle (USNM 40197); Mesa Co., Colorado National Monument-CNM
(CM 42791-42792, 43537, SM 13299), Debeque Canyon (CM 43538), Fruita Entrance to CNM
(UIMNH 53217), Grand Junction (FMNH 920), Palisade (USNM 29610), Plateau Creek (USNM
40196); Moffat Co., Jet Green & Yampa rivers (UIMNH 64659); Montezuma Co., 6 mi S Cortez
(BYU 41686), Mesa Verde National Park (KU 106150-106151, 129712-129714); Montrose Co.,
Paradox Valley (SDNHM 8680). IDAHO: Ada Co., Boise (CAS 41362-41363, KU 8443, UMMZ
63990-63993), Cinder Cone Butte (CAS 64203-64206, FMNH 6963), Snake River SE Melba (SDNHM
1432), Swan Falls near Snake River (TCWC 64836-64837), near Swan Falls (UIM 329 [6]); Bannock
Co., Pocatello (KU 6468, 11803); Bingham Co., no locality (MVZ 28765); Butte Co., Arco (BYU
30771); Cassia Co., Albion (UMMZ 71466), Raft River Valley (BYU 42032); Elmore Co., Cleft (CAS
64185, 64207), 4 mi W Hammett (MVZ 21562), 14 mi S Mtn Home (UIM 196); Fremont Co., Yellow
Stone (USNM 56018); Gooding Co., 2 mi S Hagerman (MVZ 18302), between Snake River & Bliss
(CAS 4065); Jerome Co., Jerome (TCWC 57856); Owyhee Co., 9.8 mi E Bruneau (UMMZ 133006,

Annals of Carnegie Museum

vol. 63

133398), W of Homedale (UMMZ 68299), E of Hot Springs & S of Bruneau (SDNHM 1431), Fossil
Butte (UIM 390 [2]), 18 mi SW Nampa (BYU 40813), 22 mi N Nevada State Line (MVZ 24580-
24581), lower Reynolds (UIM 291 [2]), Squaw Creek (UIM 103), Sucker Creek (CM 91569), Warm
Springs Ferry (CAS 55246); Twin Falls Co., SW 0.25 Sec 28 T17S R13E (UMMZ 125687). NEVADA:
Churchill Co., 90 mi E Fallon (BYU 16650), 8.5 mi SE Hausen (MVZ 21511); Clark Co., 20 mi SW
Indian Springs (UNM 4802), 1 mi N Lee Canyon Rd to Charles Mtns (LACM 59025), 10 mi W Hwy
95 on NV 156 (MVZ 182611), Spring Mtns (CAS 2003); Douglas Co., 3 mi NE Topaz Lake (MVZ
18635); Elko Co., Carlin (UMMZ 43123-43127, 43156, 149867-149869), 15 mi W Wendover, UT
(CAS 10025); Esmeralda Co., 12 mi E Oasis (KU 82339); Eureka Co., Dunphy (SDNHM 27787,
27851); Humboldt Co., 10 mi SW Denio (KU 109967), 6 mi S Golconda (MVZ 20622), Pine River
Mtn (MVZ 1514, 1516), Quinn River Crossing (MVZ 1513, 1515), 5 mi S Quinn River Crossing
(MVZ 21515), 2.5 mi N Ryepatch pump station valley (UMMZ 78026), Valmy (SDNHM 27788), 2
mi W Winnemucca (CAS 120818), 10.2 mi SW Winnemucca (UIMNH 4396); Lander Co., Antelope
Spring (USNM 8120), 3 mi E Austin (TNHC 7441), 16.5 mi S Battle Mtn (USNM 218839), 19.3 mi
SSE Battle Mtn (USNM 218838), Kingston Creek (MVZ 12171), Toquime Range (MVZ 42085),
Toiyabe Range (MVZ 42084); Lincoln Co., Caliente (CAS 36976), 0.5 mi E Panaca (MVZ 56917),
13 mi NNW Pioche (MVZ 61816), N end Timpahute (MVZ 14283); Mineral Co., Excelsior Mtns
(MVZ 10865), 18 mi SE Hawthorne (SDNHM 37702), 15 mi ENE Luning (UMMZ 124602); Nye
Co., Cloverdale (USNM 44525), 2 mi E Garrett Ranch (SDNHM 40654), Mercury (BYU 17409,
18755-18756), Nevada Test Site (MVZ 140912), N Twin River (MVZ 12843, 16180), 0.5 mi S Oak
Spring (MVZ 1 3082), Rainer Mesa (BYU 23858, 30547), Toiyabe Range (UNM 52432), 0.5 E Tonopah
(UNM 463), 18 mi E Tonopah (SM 3962), 24.4 mi ENE Tonopah (CAS 21320), S Twin River (MVZ
12170), White River Valley (UMMZ 86127), Wisconsin Creek (MVZ 12841-12842); Ormsby Co.,
Carson City (CAS 6527, SDNHM 27785, USNM 9520), 4 mi SW Carson City (MVZ 20395); Pershing
Co., Humboldt Range (MVZ 21513-21515), Imlay (MVZ 43150), 5 mi E Lovelock (MVZ 11983),
15.3 mi NW Lovelock (SDNHM 37791); Washoe Co., Little High Rock Canyon (MVZ 7546), 6 mi

5 Pahrum (MVZ 24582), Pyramid Lake (CAS 6526, 40505, 44146-44147, USNM 44536, 50801),
0.5 mi S Pyramid Lake (MVZ 32100), near Reno (KU 6667), 12 mi NNW Reno (CAS 93781-93784),
Smoke Creek 1 mi from CA state line (MVZ 20482), Sutcliffe (CAS 22188); White Pine Co., Baker
(CAS 14372), 1.7 mi E El Dorado (MVZ 70341), Major Woods (CAS 14371), Mt. Moriah (MVZ
24583), Spring Valley (UMMZ 84911), 8 mi N Yelland’s Ranch (SDNHM 19704). NEW MEXICO:
Bernalillo Co., 3 mi S Alameda (UNM 18254), Albuquerque (KU 5536, UNM 460, 4722, 11407,
12254, 22551,22562, 33751,38812), 1 7 mi E Albuquerque (AMNH 115716), Hwy 10 10.6 mi S US
66 (UNM 32384), Sandia Mtns (UNM 11611, 15600, 18253, 25658); Catron Co., 10 mi N Alma
(UNM 33122), 4 mi W Baldy Creek (NMSU 2956-2957), Glenwood Fish Hatchery (NMSU 3734,
4401), 0.9 mi NE Glenwood (AMNH 120697), 1 mi S Glenwood (UNM 4723), 1.3 mi N Glenwood
(CM 48768), 2.5 mi N Glenwood (BYU 13979), 4 mi S Glenwood (UNM 18740), 5 mi S Glenwood
(KU 6477-6478), 6 mi S Glenwood (AMNH 114035), 6.5 mi S & 0.5 mi W Glenwood (TCWC
65001), 13.5 mi E Glenwood (UNM 1 1535), 1 mi W Mogollon (UTEP 171 1), Nichols Canyon (NMSU
5328), Pleasanton (UNM 32169), 4 mi S Pleasanton (UTEP 1710), San Francisco Hot Springs (UAZ
25714, UNM 38853), Whitewater Canyon (UNM 6794); Chaves Co., Bishops Cap (UTEP 2540), Elk
(LACM 20598), Jornada Experimental Range (USNM 102252), 4 mi N Las Cruces (LACM 133881),
8 mi N Las Cruces (LACM 2279), 1 1 mi S Las Cruces (DMNH 1930), 1 1 mi N Las Cruces (LSUMZ
10027), 12 mi N Las Cruces (LACM 2278), 22 mi W & 15 mi N Las Cruces (LACM 133882), 14.9
mi E Mayfield (LACM 20597), 2 mi W & 1 mi S Mesilla (LACM 133883), 5 mi S Mesilla (UTEP
107), Organ Mtns (UTEP 9509), 3.9 mi N Organ (LACM 103360), Picacho Canyon (NMSU 2833),
1 mi S Radium Springs (CU 5056), 1 mi N Radium Springs (NMSU 1984), 1 mi E Rincon (UTEP
7639), 13 mi N & 4.6 mi E University Park (KU 72916), 3.2 mi S US 70 on Aguirre Springs Rd
(NMSU 5837); Eddy Co., no locality (UTEP 7660), 30 mi SW Carlsbad (CM 18297), NM 137 W
Carlsbad (UNM 37796), Carlsbad Caverns National Park (TTU 3581, UMMZ 121794, 121797-
121798, 123477, 125338, UNM 30458, 30926), Dog Canyon Rd 4 mi N TX state line (TTU 9679),
Guadelupe Mtns (UMMZ 123470, UNM 25729), Rattlesnake Canyon (KU 8386), Robinson Draw
(UMMZ 123473), 1 mi N Whites City (UMMZ 121795-121796); Grant Co., Blue Creek (NMSU
5259), 5 mi NW Buckhom (UNM 6425), Gertie Canyon in Big Burro Mtns (MVZ 7119), Burro Mtns
(NMSU 4694, UTEP 1974-1975), 4 mi S Cliff (NMSU 4391), 4 mi N Cliff (NMSU 3733), 5.5 km E

6 6.2 km N Cliff (UNM 37768), 7.5 mi N Cliff (NMSU 4395), 10.5 mi N Cliff (UAZ 35078), 16 mi
SW Cliff (NMSU 4392), 16 mi S Cliff (NMSU 4396), City of Rocks State Park (KU 72917), Dwyer
(KU 109968), Gila National Forest (UNM 38913), Gila River (NMSU 6019, UNM 461), Hachita
(AMNH 85071), 18 mi N Lordsburg (UIMNH 72091), N of Mimbres (CA 5312), 1 mi N Mimbres
(UNM 32862, UTEP 7638), 5 mi N Mimbres (UMMZ 792 1 5), Mimbres River Valley (NMSU 4399),
Pinos Altos (UAZ 25713), 2.75 mi NNW Pinos Altos (AMNH 84991), Redrock (NMSU 3730), 5 mi
NE San Lorenzo (UTEP 1345), Sapillo Creek Bridge (NMSU 4512), 2 mi W Sapillo Creek Bridge

1994

Camper and Dixon— Striped Whipsnakes

(NMSU 4390), Sawmill Creek (UNM 38932), Silver City (MVZ 42593, NMSU 4397, 4513, 4696-
4697), 1 mi S Silver City (UNM 4721), 2.9 mi W Silver City (UTACV 2621), 3 mi S Silver City
(NMSU 4493), 10 mi S & 8 mi W Silver City (UNM 15205), 13 mi W Silver City (UNM 15294), 19
mi W Silver City (NMSU 4695), 20 mi S Silver City (TCWC 63047), 25 mi SW Silver City (NMSU
4400), Slades Canyon (NMSU 4394), 1 mi NNE Vanadium (NMSU 4389), 10.7 mi NNE White Signal
(UTEP 8763) Jet NM 61 & Hwy 90 (LACM 76408), Sec 12 T18S R16W (NMSU 4398), 13.3 mi NE
jet NM 90 & US 70 (UNM 32033); Hidalgo Co., Alamo Hueco Mtns (AMNH 78996, UTEP 1 1355),
4.5 mi NW Antelope Wells (NMSU 1868), 39 mi S Hatchita (ASC 10952), 0.7 mi S Grant Co line
(UNM 6793), Little Hatchet Mtns (AMNH 74725, 75410, UTEP 10570), 1.8 mi W jet NM 530 &
92 (NMSU 5347); Lincoln Co., Corona (LACM 2277), 2 mi NW Carrizozo (UMMZ 67658), 4 mi
NW Carrizozo (AMNH 1 1 1227), 2 mi NW Hondo (UNM 23558), Tularosa Malpaise (UNM 37153),
0.5 mi N White Oaks (UNM 31673); Luna Co., 5.7 mi N Columbus (LACM 103361), 21.5 mi W
Columbus (TCWC 56339), 19 mi NNE Deming (UTEP 5676), 10 mi NW Florida (NMSU 4514),
Florida Mtns (UTEP 200, 1 1261), 2 mi N Nutt (LACM 109526), Tres Hermanas Mtns (UNM 462);
McKinley Co., 5.9 mi N Ya-ta-hey (UTEP 9540); Otero Co., Alamo Mtn NW 0.25 Sec 21 T26S R13E
(NMSU 6329), 1 mi NE Alamogordo (NMSU 4698), Dog Canyon (USL 24962), 7.5 mi E LaLuz
(LACM 20599), SW 0.25 Sec 33 T17S R10E (NMSU 6186-6187); Quay Co., 9 mi W & 1.5 mi S
Tucumari Mtn (UNM 19705); Rio Arriba Co., 14 mi NW Abiqui (UAZ 39721), Nutria (USNM 8432);
Sandoval Co., 4 mi S Algdones (UNM 16697), Bandelier National Monument (UNM 30534), 15.2
mi SE Cuba (UNM 22071), Jemez Mtns (UNM 30879), Jemez Pueblo (UNM 459, 4720), Juan Tabo
Picnic Area (UNM 10902, 13518,37797), Placitas (UNM 458, 18255), 2 mi SW Placitas (CM 58692),
Sante Fe National Forest (UTEP 8926); San Juan Co., Chaco Canyon National Monument (UNM
3395, 21835-21839), 2.6 mi S Newcomb (UAZ 36364), Shiprock (USNM 21 10 holotype, 44503), 4
mi W Shiprock (UTACV 1182), NE 0.25 Sec 24 T32N R7W (UNM 4153), Wool Canyon (UU 3738);
San Miguel Co., 1 mi below Conchas dam (UNM 21909), 3 mi below Conchas dam (UNM 18256),
1 mi N Sena (UNM 32863), 7.3 mi SE Trujillo (LACM 20600); Santa Fe Co., 4 mi N Golden (UNM
31669), Jemez Mtns (UNM 364643), 5 mi NE Los Cerillos (UNM 456), 1.5 mi S Madrid (UNM
23514), 2 mi N Madrid (UNM 10243); Sierra Co., Arrey (NMSU 1477), Caballo Dam (UTEP 152),
Elephant Butte (LACM 2276), 2 mi N Ft McCrae (NMSU uncatalogued), Hillsboro (UAZ 40319),
0.8 mi W Hillsboro (UTEP 10698), 3 mi W Hillsboro (NMSU 2028), 3 mi S Hillsboro (LACM 76409),
16 mi E Hillsboro (LACM 76417), Hospital Canyon (NMSU 5531), 2 mi N Lake Valley (NMSU
4402), 1 mi N Truth or Consequences (LACM 2275), 3 mi W Truth or Consequences (NMSU 2020),
12 mi N Truth or Consequences (UNM 13730), 1.5 mi W jet NM 52 & 135 on 135 (NMSU 5484);
Socorro Co., 2.3 mi E Bingham (UNM 12762), 6 mi E Bingham (LACM 20594), 12 mi SE New
Bingham (LACM 20596), Chupadera Mesa (LACM 63453), 15 mi W Corrizozo (UNM 16745), 3 mi
E & 3 mi S Dusty (TCWC 66165), 13.5 mi NE Magdalena (UNM 30880, 30895), Magdalena Mtns
(UNM 4154-4155, 13855), Mockingbird Gap (LACM 20595), Rio Salado (UNM 14876), 8.9 km W
& 1.4 km N 1-40 bridge over Rio Salado (UNM 39258), 15 mi S & 3 mi E Rosedale (KU 47791), 25
mi SW San Antonio (UNM 455), 8.7 mi W Socorro (LACM 20593), 16.1 mi W Socorro (UNM
1 1 525); Torrance Co., 9 mi W Mountainair (UNM 457); Valencia Co., 9 mi N Acoma (MVZ 12811 2),
5 mi E Grants (CU 3052), 6 mi NE Grants (UNM 22332), 8 mi E Grants (UNM 8843), 10.4 mi S
Grants (CU 5602), Mesa Encantada (UNM 30896). OREGON: Crook Co., no locality (MVZ 26924);
Deschutes Co., 3 mi W Terrebonne (UNM 33169), 4 mi W Terrebonne (UNM 30941); Harney Co.,
Fields (UMMZ 133342, 133379), 3 mi S Fields (UMMZ 133091), 6 mi S Fields (UMMZ 133001,
133092 [3]), Frenchglen (CM 20745); Lake Co., Picture Pass Summit (UMMZ 135986); Malheur Co.,
5 mi S Adrian (UMMZ 124574); Wasco Co., Maupin (MVZ 62055); TEXAS: Bandera Co., 18 mi
NW Medina (TNHC 1904), 6 mi S Tarpley (SM 7491), 13 mi W Tarpley (UNM 37798); Bexar Co.,
Helotes (SM 228-229), 1 mi N Helotes (MVZ 68462), Helotes Creek (CU 609, SM 6384), San Antonio
(UNM 13522, UTEP 9595), 17 mi N San Antonio (AMNH 22743, 74548), 18 mi N San Antonio
(CM 22850), 22 mi N San Antonio (CU 1771-1772); Blanco Co., 4.9 mi W Johnson City (USL 1793),
5.7 mi E Johnson City (USL 1792), 17 mi SE Johnson City (TNHC 28913), 20 mi NW Johnson City
(MCZ 62561), 16.9 mi E Sandy (LSUMZ 14149); Brewster Co., no locality (SRSU 1469, UMMZ
71925), Alpine (CAS 7503, SRSU 1844, 4587), 10 mi S Alpine (SRSU 1694), 22 mi S Alpine (TCWC
28940), 42 km S Alpine (KU 176790), 32.1 mi S Alpine (LSUMZ 23342), 59.5 mi S Alpine (USL
6799), 65 mi S Alpine (SRSU 1845), Big Bend National Park (AMNH 72527, 77318, 111228, CA
7821, CM 5006, 60005, FMNH 26617-26618, 26803-26804, 27707-27708, 27845, LACM 103362-
103363, MVZ 25362, SM 6388-6392, SRSU 3927, 5079, TCWC 16127-16128, 40111, TTU 4, 629,
UAZ 34784, 39228, 40391, UMMZ 66026-66029, 72088, 95435, UNM 9025-9026, 18258, 20801,
20891-20893, 22238, 22381, USNM 103638, UTACV 2619), 1 mi N BBNP on Hwy 118 (UMMZ
32386), 1 3 mi N BBNP on Hwy 1 1 8 (UIMNH 47645), Black Gap Wildlife Management Area (DMNH
296, 302, TCWC 12296, TNHC 12603, 12797, 12984), Boquillas (CA 4860), Lajitas (KU 51945),
Marathon (SM 5926-5927), 12 mi S Marathon (UMMZ 20827), 12 mi S & 2 mi W Marathon (FWM

Annals of Carnegie Museum

vol. 63

6910), 13 mi S Marathon (SRSU 4665), 22 mi E Marathon (MVZ 53916), 31 mi SE Marathon (SRSU
1671), Reed Plateau (TCWC 64999), Rosillos Mtns (FMNH 75480-75482, UMMZ 1 14344), 25 mi
W Sanderson (AMNH 94276), Santiago Mtns (UMMZ 1 14203), Study Butte (NMSU 3210), 7 mi W
Study Butte (NMSU 3211), 10 mi S Study Butte (SRSU 2193), 28.5 km N Study Butte (UTACV
12736), Terlingua Ranch (TCWC 65285), 1.1 mi W jet Hwys 170 & 1 18 on 170 (TCWC 63376), 4
mi N jet Hwys 118 & 170 on 118 (UTEP 10569), 48.9 mi S jet Hwys 118 & 90 on 118 (TCWC
63378); Burnet Co., Clear Creek (CAS 33072, SM 752), 12 mi W Inks Dam (TNHC 2639), 7 mi NW
Marble Falls (TAIC 1414); Coleman Co., 0.25 mi N Santa Anna (UTACV 2620); Comal Co., 1 mi
NW Fischer (TCWC 25397), Hunter (UMMZ 74328), New Braunfels (UMMZ 69662), 3.5 mi NNE
New Braunfels (MVZ 99224), 5 mi NW New Braunfels (UMMZ 74074); Concho Co., 50 mi SE San
Angelo (LACM 74105); Crockett Co., Howard Springs (USNM 1970), 17 mi E Iraan (TCWC 401 12),
15 mi W Ozona (UAZ 25757), jet Pecos River & Independence Creek (TU 14472); Culberson Co.,
Guadelupe Mtns National Park (SM 4881-4882, TCWC 65904, TTU 9675, UMMZ 70134, 123469,
123514, USNM 147878), 6 mi N Kent (MU 4130), 2 mi NE Nickle (KU 72759-72760), Pine Springs
(SDNHM 25483), 2 mi W Van Horn (TTU 4741), 15 mi N Van Horn (UMMZ 91474), 25 mi N
Van Horn (TCWC 18299-18300); Edwards Co., 0.5 mi W Real Co line on TX 41 (TNHC 44638), 4
mi E Rocksprings (TTU 9715), 7 mi E Rocksprings (UMMZ 102443), 20 mi S Rocksprings (TCWC
65923), 21 mi NE Rocksprings (DMNH 5254), 44.2 km S jet Hwys 290 & U10 (UTACV 14737); El
Paso Co., El Paso (DMNH 2341, 41 13, NMSU 5501, UMMZ 74072, UTEP 49, 319-322, 816, 2693,
3368, 3677, 4122, 9073, 10572, 10781, 10804, 11178), Hueco Tanks State Park (UTEP 418, 725,
969, 1043, 1656, 2841, 4369, 10452), 5 mi E Hueco Tanks SP (MVZ 37008); Gillespie Co., 8 mi NE
Fredericksburg (TCWC 27390), 10 mi N Fredericksburg (SM 12621), 18 mi N Fredericksburg (TCWC
5204); Hays Co., no locality (ASU 2413), Dripping Springs (TNHC 29046), 4 mi E Dripping Springs
(TU 18219), Pollard Wildlife Refuge (TCWC 38792), San Marcos (AMNH 32412), 12 mi WNW San
Marcos (SIU-C 226), 15 mi WNW San Marcos (UMMZ 105236), 17 mi NW San Marcos (FSM
56659), Wimberley (FMNH 55049, UTACV 1582), 3 mi S Wimberley (LSUMZ 5852), 7 mi SE
Wimberley (TCWC 19030-19031), Wren Ranch Blanco River (FMNH 38061); Hudspeth Co., Eagle
Mtns (UTACV 2361, UTEP 1937), 4.5 mi S Hilltop (SRSU 1559), 9.6 mi W Indian Hot Springs
(SRSU 4336), Quitman Mtns (UTEP 7455), Sierra Blanca (UTACV 591), 9 mi W Sierra Blanca (KU
40335), 8.2 mi W Van Horn (UTEP 1 1171); Jeff Davis Co., no locality (NMSU 5838), 14 mi NW
Alpine (FSM 56661), 15 mi NW Alpine (SRSU 1846), 29.4 mi NW Alpine (FSM 56660), Davis Mtns
(FWM 2530, SM 11532, SRSU 1506, TAIC 4870, TCWC 612, UMMZ 49975-49976, 52925, 81978,
UTEP 6812, 10497, 10735), Davis Mtns State Park (AMNH 115717, NMSU 3186, SM 13273), Ft
Davis (SRSU 1849-1850, USNM 1971 holotype), 3 mi N Ft Davis (UMMZ 69666), 14 mi N & 8
mi E Ft Davis (KU 56236), 16 mi W Ft Davis (TNHC 7044), 20 mi NW Ft Davis (FSM 56662), 1
mi S Kent (SRSU 1973), 8.5 mi S Kent (UTACV 2701), 3 mi NW McDonald Observatory Rd on
118 (UNM 37799); Kendall Co., Boeme (SM 1901), 1 1 mi N Boeme (TTU 52), Kendalia (LSUMZ
40055); Kerr Co., 3.6 mi W Hunt (UTACV 2618), 5 mi W Hunt (TCWC 199), 12.9 mi from Hunt
(TCWC 47133), 8 mi SW Ingram (TCWC 198), 8.3 mi S & 2.5 mi W Kerrville (TNHC 1 1933), 10
mi W Kerrville (TCWC 197), 13 mi S Kerrville (TAIC 1058, 1330, 1839, 2618), 17 mi NW Kerrville
(TU 1 4496), 25 mi W Kerrville (UTACV 2 1 94), Kerr Wildlife Management Area (KWMA 1 specimen,
TCWC 19033, 25396); Kimble Co., no locality (SRSU 1851), 3.8 mi N Cleo (SM 12620), Crow Ranch
(TCWC 65287), Junction (TCWC 64809, 65284), 6 mi W Junction (KU 61061), 10 mi E Junction
(MU 3040), 10 mi W Junctin (TNHC 29049), 10 mi NNW Junction (TTU 6048), 12 mi NE Junction
(SRSU 1852), 15 mi NW Junction (ASC 8130), US 290 SW comer of county (LACM 66805), 1.5 mi
W Telegraph (TAIC 3050); Llano Co., 12.8 km S Cherokee (TCWC 63772-63773, 63876, 64841,
65000, UTACV 1 1126), Enchanted Rock State Natural Area (AMNH 74547, MVZ 128100, TNHC
4498, 4501, 4503, 29048, TU 13549), S of Llano (TNHC 29045), 2.5 mi W Llano (TNHC 12290),
3 mi W Llano (TCWC 64996), 6 mi SW Llano (UTACV 508), 8 mi E Llano (TCWC 64996), 9 mi
N Llano (TCWC 531), 19.2 km NNE Llano (UTACV 14738), 19 mi SW Llano (TCWC 58488-58489);
Mason Co., 1 mi E Katemcy (ASC 10549), 2 mi WNW Katemcy (ASC 10813), 3.2 mi WNW Katemcy
(ASC 10205), 9 mi SE Mason (ASC 386), 9.1 mi W Mason (TCWC 401 10), 10 mi W Mason (TCWC
8734), 12 mi S Mason (TCWC 33797), 12 mi NE Mason (TCWC 3298), 13 mi W Mason (SM 11531),
20 mi ESE Mason (TCWC 31125), Olga Zesch Ranch (TCWC 31126); McCulloch Co., S Brady at
San Saba River (FWM 3174), FM 734 at San Saba River (TCWC 19032), 2 mi N San Saba River on
Hwy 71 (TCWC 64838); Medina Co., Rio Medina (CM 19919), Hwy 173 at Verde Creek (TCWC
63775); Menard Co., 2.5 E Menard (TCWC 38015), 3 mi E Menard (TCWC 42362), 10 mi W Menard
(SM 8951), 10 mi SW Menard (TNHC 12262); Mills Co., 15 mi SSW Goldthwaite (TNHC 14101);
Pecos Co., ANSP 15621, 15654, 17076; LACM 66802 15 mi E Ft Stockton; TCWC 27383 13 mi E
& 1 mi N Bakersfield; UNM 12079 Huckabee Ranch; Presidio Co., 8 mi NE Candelaria (TCWC
27637, 27639-27643), 9 mi NE Candelaria (TCWC 27639), 24 mi SW Ft Davis (TTU 4290), Kingston
Hot Springs (TCWC 64810-64811), 3 mi W Lajitas (AMNH 112229), 12.1 mi W Lajitas (TCWC

1994

Camper and Dixon —Striped Whipsnakes

65286), 20.2 mi W Lajitas (TCWC 64324), Madera Canyon N Hwy 170 (SRSU 3931), 16 mi S Marfa
(TCWC 30516), 63 mi S Marfa (TNHC 14975-14978), 3 mi E Porvenir (FMNH 46037), 15 mi E
Porvenir (TNHC 3589), 1 mi SE Presidio (TCWC 27892), 3 mi SE Presidio, (TCWC 27893), 7 mi E
Presidio (ASC 10379), 12 mi N Presidio (SRSU 1848), 16.5 mi N Presidio (SRSU 2298), 29.9 mi N
Presidio (CM 60598), 7 mi SE Redferd (TCWC 27894), 7.3 mi SE Redford (USL 6834), 12.8 mi SE
Redford (TCWC 40112), 13 mi SE Redford (SRSU 3561), 13.5 mi SE Redford (TCWC 26171), 17.5
mi SE Redford (LSUMZ 28669), 7 mi N Ruidosa (SRSU 1972), 4.5 mi S Shafter (UTEP 2773), 6.8
mi SW Shafter (LSUMZ 23332), 13 mi N Shafter (DMNH 4227), 19 mi N Shafter (SRSU 4733), 9
mi W jet US 67 & FM 170 on 170 (TCWC 29375), 23.2 mi E jet US 67 & FM 170 on 170 (TCWC
63377), 11 mi W Valentine (TNHC 2851, 3528, 3819, 3837, 3914, 4042, 4262, 15299, 17899);
Reagan Co., 9 mi W & 1 1 mi S Big Lake (ASC 9572), 9 mi S & 12 mi W Big Lake (TCWC 31376);
Real Co., 4.5 mi S Leakey (CM 42793), Prade Ranch (FMNH 55018-55019); Reeves Co., Toyahvale
(TNHC 1927); San Saba Co., 2 mi W Bend (FWM 5219), 6 mi SE Bend (ASC 385, 5195), 1 1 mi
NW Bend (FWM 5460-5461), 17.5 mi W Lampasas (UTACV 5571), near San Saba (TCWC 20170-
20171), 11 mi SW San Saba (FWM 1759-1760), 12.1 mi SE San Saba (FWM 6061), 18 mi NNW
San Saba (TNHC 1 1985-11896); Sutton Co., E of Sonora (LACM 66804), 4 mi S Sonora (KU 82337),
14.9 mi WNW Sonora (UTEP 113); Terrell Co., Chandler Ranch (SRSU 1732, 1971, 1974), 7 mi N
Dryden (MU 3897), 8.8 mi W Dryden (UTEP 9156), 21 mi N Dryden (MU 3042), 30 mi W Dryden
(SRSU 1975), 20 mi W Langtry (KU 82338), 2 mi W Sanderson (SRSU 3016), 4.7 mi W Sanderson
(TAIC 4352), 5.7 mi W Sanderson (UIMNH 34274), 7.9 mi W Sanderson (UTEP 9051), 7 mi SW
Sheffield (TTU 2530), 10 mi S Sheffield (TNHC 7994), 13 mi S Sheffield (TNHC 7681), 21 mi S
Sheffield (TNHC 8225), 25.9 mi S Sheffield (TCWC 28885), 30 mi S Sheffield (TNHC 7838); Throck-
morton Co., 15 mi SW Throckmorton (TNHC 26579); Travis Co., no locality (LACM 103364), Austin
(AMNH 4191, FSM 1 1989, TNHC 35221, TTU 86), NW of Austin (TNHC 26828), 7 mi SW Austin
(FSM 56656-56658), 8 mi NW Austin (TNHC 2069), 10 mi N Austin (AMNH 122738), 12 mi W
Austin (TNHC 19651, 20493), 15 mi SW Austin (TNHC 4497), 17 mi NW Austin (TNHC 4500),
18 mi NNW Austin (TNHC 4499), 20 mi NNW Austin (TNHC 4502), 20 mi W Austin (UIMNH
2652-2654), Gaines Ranch (TNHC 1847,4897,9084, 12295, 1 5295, 20542), Hamilton’s Pool (LACM
66801), near Lake Travis (TCWC 20239), old river above Winfree (TNHC 29050), RR 620 1 mi W
jet with RR 2769 (UAZ 25758), US 290 E of Hays Co line (FSM 7801-7802); Upton Co., 24 km
WNW Rankin (UTACV 14472-14474); Uvalde Co., 3.2 mi S Nueces River on Hwy 55 (TCWC
64840), 8.2 mi N jet Hwys 1275 & 1835 (USL 19742); Val Verde Co., Caulk Ranch (TAIC 3217),
Comstock (TNHC 29047), N of Comstock on Pandale rd (Taic 4780), 1 mi W Comstock (ASC 9570,
UAZ 40392), 1.5 mi SE Comstock (ASC 9568), 6.2 mi W Comstock (LSUMZ 23331), 7.9 mi NW
Comstock (ASC 9565), 8 mi SW Comstock (USNM 32782), 15 mi SE Comstock (ASC 9564), 16.6
mi N Comstock (ASC 9571), 17 mi SE Comstock (ASC 9567), 18 mi SE Comstock (ASC 9569), 19
mi NE Comstock (TNHC 32512), 21.6 mi N Comstock (TNHC 32530), 7 mi NW Del Rio (UTACV
1339), 17 mi W Del Rio (MU 3041), 18 mi N Del Rio (CAS 103476), 18 mi NW Del Rio (SRSU
1843), 25.2 mi N Del Rio (UNM 8848), 36 mi NNW Del Rio (ASC 1524), 36.3 mi N Del Rio (UNM
8325), 40 mi N & 30 mi W Del Rio (ASC 5763), Devils River (USNM 32781), Evans Creek at Lake
Amistad (SRSU 2746), 16.6 mi S Everett’s Crossing (TAIC 4283), Fawcett Ranch (SRSU 3457, 3414,
TTU 4355), 13 mi W Juno (ASC 9566), 2 mi W. Langtry (KU 61062), 7 mi N Langtry (TAIC 3558),
8 mi W Langtry (CM 88738), 20.9 km N Langtry- (UTACV 14613), 25 mi N Langtry (TAIC 4118),
35 mi N Langtry (TAIC 3465 [3], 3476, 3498, 3556-3557, 3560, 3984), 29.8 mi W Loma Alta (TCWC
13837), Pandale hwy jet (TAIC 3540), 7.1 km S Pandale (UTACV 16341), mouth of Pecos River
(UMMZ 74335-74336), Pecos River at Lake Amistad (SRSU 4812), US 90 at Pecos River (TAIC
1915, 2423, TCWC 33796, UMMZ 92798), US 90 2 mi W Amistad Dam (TAIC 2364), 1 mi W
Shumla (TNHC 35223), Hwy 163 29 mi N jet with US 90 (UMMZ 175826), US 90 9 mi ESE Terrel
Co line (UTEP 10576); Williamson Co., 3 mi N Georgetown (UTACV 1340). UTAH: Beaver Co.,
Baker’s Canyon (USNM 36346), Beaver Creek hills (AMNH 29367-29368), Milford Valley, (BYU
564, 1640-1641), 50 mi W Milford BYU 563, 1637, 1809), White Sage Valley (SDNHM 40691); Box
Elder Co., Locomotive Springs (UU 917-918, 2000-2004, 8062), 22 mi N Wendover (BYU 42416);
Carbon Co., 25.3 mi S & 20 mi W Ouray (AMNH 1 12918), Price (BYU 2792, KU 82340, UU 1368,
1409-1410), 3 mi W Price (BYU 22951); Davis Co., Antelope Island (LACM 121003), Farmington
(AMNH 14231); Duchesne Co., Montwell (BYU 16719), 5 mi W Roosevelt (BYU 16718); Emery
Co., Morrison Salt Wash 10 mi W Green River (AMNH 64840), San Rafael (BYU 2906), upper
Temple Mtn (BYU 18997), 6 mi N Temple Mtn (BYU 20732), 2 mi NW Temple Jet (BYU 16614),
Woodside (BYU 258), 2 mi NNE Jet Hwy 24 & road to San Rafael Swell (CAS 92465); Garfield Co.,
Hog Springs (UTACV 12737), 4 mi NW Hog Springs (BYU 12694), 1 mi W Star Springs (BYU
12623); Grand Co., Arches National Park (AMNH 115603, BYU 13036), near Castleton (LACM
103365-103366), 2 mi NW Castleton (UMMZ 68588), 25 mi N Cisco (FMNH 33909), 12 mi SE
Cresent Jet (BYU 30533), near Colorado state line (KU 5533), Green River (BYU 257), 2 mi S Moab

Annals of Carnegie Museum

vol. 63

(UMMZ 68587), 5 mi N Thompson (UMMZ 68586); Iron Co., Antelope Springs (SDNHM 40662),
Cedar City (BYU 392), 1 mi N Cedar City (KU 20906-20907), Sulphur Springs (SDNHM 40661);
Juab Co., 18 mi S & 9 mi W Eureka (CM 91 197), 20 mi W Nephi (UIMNH 2135), Yopaz Mtn (BYU
9072); Kane Co., 10 mi S Alton (BYU 11312), Glen Canyon (UAZ 39720), Kanab (UAZ 25759), 3
mi N Kanab (UMMZ 73390), 7 mi N Kanab (MNA 7.254), 19 mi E Kanab (BYU 8715), Kaiparowits
Plateau (UU 1288, 1606), Orderville (SDNHM 25485), Piaria (BYU 2893); Millard Co., Cove Fort
(CAS 54150), 3 mi N Cove Fort (BYU 568), 20 mi NW Delta (SDNHM 674-675), 30 mi N Delta
(CU 1082, SDNHM 1274), 65 mi W Delta (UIMNH 2132-2134), Fillmore (MCZ 13361-13362), 2
mi N Fillmore (UMMZ 70654), 10 mi NW Fillmore (UMMZ 70652), 10 mi SW Fillmore (UMMZ
70653), 10-11 mi W Fillmore (UMMZ 70655), 20 mi W Hinckley (BYU 16590), 7 mi S Kanosh
(CAS 47756), Maple Grove Forest Canyon (UU 3386), 18.1 mi E NV state line (CM 49031); Piute
Co., 8.4 mi N Circleville (ASU 21 199), 4.4 mi S Marysville (LACM 20590); Salt Lake Co., 0.5 mi
from mouth of Emigration Canyon (UU 14167), Fort Douglas (CAS 14167, 30923-30924, 38761),
Salt Lake City (CU 3500, MCZ 8899, 9051, 9053); San Juan Co., no locality (FMNH 25274, 25450,
25452, 25712, UTEP 5456), Bluff (AMNH 4871), 6 mi SW Bluff (CM 66658), Devils Lane near
Needles (LACM 103367), Indian Creek (BYU 21709-21710), 6 mi W La Sal (BYU 18579), 25 mi
SE La Sal (LACM 29037), 12 mi S Moab (UNM 6695), 36 km S Moab (KU 191935); Sanpete Co.,
Ephraim (UU 89), Maple Canyon (UMMZ 64681); Sevier Co., Monroe (SDNHM 38275); Tooele Co.,
13.8 mi N Bonnville Station (UU 5656-5657), 10 mi W Castle Cliff (CM 53604), Dugway Proving
Ground (BYU 14823), 9 mi N Dugway Proving Ground entrance (LACM 103368), Gold Hill (BYU
2998), near Grantsville (UU 2478, 2482), W of Grantsville (UU 1966-1967), Lone Rock Skull Valley
(LACM 121038-121080, MVZ 197596), between Ophir & Mercur (UU 2032), Stansbury Island
(DURC 379 [2], LACM 121004, UU 1607), Stockton (SDNHM 24679), Tooele (UU 38), W of Vernon
(UU 1219a, 8115), Wendover (LACM 24680), 14 mi N & 9 mi E Wendover (UU 3253); Uintah Co.,
Dinosaur National Monument (BYU 10028), near Jensen (USNM 66196), LaPoint (MVZ 30256), 7
mi E Myton (KU 56097), Oil Town (BYU 22479), 10.6 mi S & 12 mi W Ouray (AMNH 112917),
29 mi S & 19 mi W Ouray (AMNH 112919), near Vernal (CM 7527), 10 mi N Vernal (MVZ 30255),
10 mi W Vernal (MVZ 30254), White River (CM 1430); Utah Co., 3 mi S Allen Ranch (BYU 13042),
Cedar Valley (BYU 216, 110, 1109, 14686-14687), Chimney Rock Pass (BYU 2842, 14685), Dividend
(BYU 247), mtns W of Elberta (BYU 22093), 5 mi SW Goshen Bay (UIMNH 23875), Lake Mtn
(BYU 386), W of Lehi (BYU 2779, 14684), Meseda Bench Prospect (BYU 14986), W of Meseda
(UIMNH 34771-34772), Provo (FMNH 41680, UO 3687, USNM 8122), Rock Pile (BYU 30431),
near Springville (AMNH 122735), Tintic (BYU 2031-2034, 2136, 2714), Utah Lake (LACM 121081,
UU 881); Washington Co., 16 mi NW St. George (USNM 44268), Snow Canyon (BYU 18967), Zion
National Park (AMNH 63965, 64141, BYU 215, 394, 1098, 1399, 8739, CAS 56714, DMNH 4276,
LACM 132466, SDNHM 3160, UU 74, 341, 805; Wayne Co., no locality (FMNH 62895), Hanksville
(BYU 8391-8392); Weber Co., Ogden (MCZ 4595, USNM 10716); County unknown, no locality
(ANSP 5363, MCZ 1943, USNM 14002), near Casisson, Colorado (KU 6668), Rush Lake (USNM
81244); WASHINGTON: Walla Walla Co., 3 mi E Walla Walla (UMMZ 56952-56953). MEXICO:
CHIHUAHUA: no locality (USNM 14272), 18.1 mi E Aldama (UNM 34249), 26.6 mi E Aldama
(UNM 34248), 4.5 mi N Hwy 16 on E Colonias Access rd to Aldama (UNM 34250), 4.3 mi E jet
Hwy 16 & 45 on 16 (UNM 34251), 18.6 mi W El Ancon (UAZ 35018), 13.6 mi N Ascension (UAZ
36562), 5 mi N Camargo (UMMZ 117765), 5 km SW Chihuahua (CM 60003), 9.8 mi NW Chihuahua
(UIMNH 46021), 30 km N Chihuahua (SDNHM 49501), 72 mi N Chihuahua (UTEP 4097), Coyame
(UAZ 350 1 7), 23.2 mi ENE Coyame (EAL 3489), Falomir (UIMNH 52432), 5 mi N Falomir (UIMNH
52433), 30 mi SW Gallego (MVZ 66116), 50 mi E General Trias (UNM 34247), 6 mi NE Janos
(UTEP 4228), 5.2 mi S La Mula (SRSU 2816), Lago Santa Maria (USNM 46594), 30 mi N Las
Delicias (CM 60006), Norogachi (AMNH 73752), 100 mi SSW Ojinaga (DMNH 2273), 13 mi N
Pachera (MVZ 59286), Santa Barbara (AMNH 68 1 93-68 198, 6825 1 , 68947-68950), rd to Santa Clara
12 mi from Hwy 45 (UTACV 4532), Sierra del Nido (MVZ 68873, 71014, 73053, UTEP 2519), 2
mi N Yepomera (UAZ 34783), 1 1 .4 mi N Yepomera (UAZ 34425). COAHUILA: 3 mi E Americanos
(KU 39567), Cuatro Cienegas (USNM 248109), 3 mi S Cuatro Cienegas (AMNH 77315), 3 mi W
Cuatro Cienegas (TNHC 33012), 11 km SW Cuatro Cienegas (LACM 116256), 14 km S Cuatro
Cienegas (CM 48192), del Carmen Mtns (FMNH 25300), 1 mi E tip San Marcos Mtns (KU 80282),
San Pedro (MCZ 4582), 5 mi S San Pedro (USNM 105300), 1 1 mi N Zapata (UIMNH 48148).
DURANGO: 8 mi W Durango (AMNH 102520), 84.8 mi W Durango (TCWC 33798), 41 mi E El
Verfel (UTACV 8361), 27.5 mi S jet hwys 30 & 45 at La Zarca (UNM 33474), 3 mi SW Lerdo
(AMNH 67326-67328), 13.3 mi W Mapuni (UTEP 7749), 4 mi E Metates (UTEP 9374), 3.8 mi NE
Pedricena (TCWC 44007), 6 mi NE Pedricena (UIMNH 17857), 12 mi S Villa Ocampo (AMNH
86000). JALISCO: 8.1 mi ESE Tepetatillo (CAS 165260). ZACATECAS: 4 mi W Colorado (FMNH
106181), Sierra Organos (CAS 165223).

Masticophis schotti. — UNITED STATES: TEXAS: Atascosa Co., Christine (UMMZ 74329), 4 mi

1994

Camper and Dixon— Striped Whipsnakes

E Christine (SM 1760), 10 mi W Jourdanton (SDNHM 30412), 3.1 mi SE Kyote (TAIC 2229), Lytle
(UIMNH 6299), near Pleasanton (TNHC 28758), 6 mi S Pleasanton (UMMZ 71341-71342), 9 mi E
Pleasanton (CM 8457), 7 mi E Poteet (UMMZ 74068-74069); Bee Co., Beeville (CA 10838-10840),
1 mi E Beeville (TNHC 28031), 6 mi S Berclair (TNHC 20403), 3.8 mi N Skidmore (TNHC 24445),
Tulsita (TNHC 1 5294); Bexar Co., San Antonio (ASC 4790, CM 1 676 1 , UMMZ 7407 1 , UTEP 9931),
near San Antonio (SDNHM 20970), 12 mi S San Antonio (UMMZ 74066), 12 mi S San Antonio
(UMMZ 74330-74332), 16 mi E San Antonio (UMMZ 74327), 20 mi SW San Antonio (FMNH
108658), 20 mi S San Antonio (UMMZ 74070), Somerset (CM 19911); Cameron Co., no locality
(FMNH 38037), Brownsville (AMNH 24991, ANSP 22253, CAS 55354-55355, CU 1199, 1261-
1262, FMNH 27170, KU 8393, MCZ 17998-18000, 46506, 15173, 62561 paratype, TCWC 65282,
UIMNH 43493 paratype, UMMZ 57681 holotype, 57682-57684 & 57686-576894 paratypes, 59036,
60873-60876, USNM 75981, 84041, UU 1093-1095), 5 mi E Brownsville (TTU 5231), 10 mi NE
Brownsville (UMMZ 74065), 12 mi NE Brownsville (UMMZ 74062-74064), 20 mi NE Brownsville
(UMMZ 74073), 25-30 mi NNE Brownsville (AMNH 122736-122737), Harlingen (KU 8395), E
Laguna Atascosa National Wildlife Refuge (LACM 59023), S Laguna Atascosa NWR (TCWC 61778),
33 mi NW hdqrs Laguna Atascosa NWR (USNM 238845), Laguna Heights (TAIC 4987), 3 mi NE
San Benito (MVZ 78363), 3 mi S jet FM 106 & 1847 on 1847 (UTACV 8392); Duval Co., 3.9 mi
NE Benavides (LSUMZ 23333, 23337), 8.8 mi SW Benavides (23336), 1 mi W Freer (TAIC 2355),
1 mi N Freer (UMMZ 1 14405), 13.9 mi NE Freer (UMMZ 126173), 1.6 mi NE Realitos (LSUMZ
23339), 1.9 mi NW Realitos (LSUMZ 23334), San Diego (LSUMZ 23338, 23340), 7.9 mi SW San
Diego (LSUMZ 23335), 4.7 mi jet 1329 & 2295 on 2295 (TAIC 3328 [2]); Frio Co., Dilley (UTACV
293), 7 mi NW Dilley (TCWC 63774, 63919), 3.7 mi SW Moore (TNHC 47670), Hwy 57 at Frio
River (USL 15084); Goliad Co., 8 mi E Goliad (TNHC 17900, 17930); Hidalgo Co., Alamo (USNM
1 59384), Edinburg (AMNH 74549-7455 1 , CA 6350-635 1), N of Edinburg (CU 589), 1 mi N Edinburg
(TCWC 18301), 12 mi W Hidalgo (TCWC 65283), 13 mi N Edinburg (TCWC 36389), 19 mi S Encino
(UIMNH 3832-3833), 14.7 mi NW La Joya (SM 8949), 6 mi S Linn (SRSU 2120), 6 mi S McAllen
(TNHC 9059), 1 mi W Red Gate (TAIC 4666), 1 mi W jet FM 2984 & 676 (TAIC 1474); Jim Hogg
Co., 9.3 mi N Guerra (TCWC 60755), 2 mi S Randado (TAIC 4398), 5.8 mi S jet Hwys 3973 & 649
on 649 (TCWC 48658); Jim Wells Co., Alice (TTU 791), 1.5 mi E Alice (UMMZ 69663), 3 mi S Ben
Bolt (LSUMZ 1 3288), 5 mi S Ben Bolt (TNHC 23 1 69), La Copita Experimental Ranch (TCWC 63408,
64389), 3 mi N Premont (UIMNH 16874), 10.9 mi N Premont (SM 8950); Kenedy Co., 9 mi S
Armstrong (TNHC 28003), 15.6 mi Armstrong (TAIC 3037), 12 mi S Norms (KU 61063), 10.2 mi
S Riviera (FSM 43495), 5.7 mi S Sarita (TAIC 3099), 6 mi S Sarita (UTACV 16827), 8 mi S Sarita
(LSUMZ 27724); Kleberg Co., Laureles Division King Ranch (TAIC 5307), Kingsville (CAS 103473-
103474, EAL 5000, TAIC 203, 361, 1707, 1731, 1942, 1976, 2059, 2073, 2382, 3563, 4255, 4772,
4857, 5257, 5307, 5364), 8.5 mi S Kingsville (TAIC 693 [2], 924, 2069), 2.5 mi S Riviera (TNHC
28030), 8 mi E Riviera (TAIC 933), 0.5 mi W Riviera Beach (TAIC 208); Kinney Co., 2 mi W
Bracketville (TNHC 42218), 5 mi WSW Bracketville (TNHC 47577), 5.9 mi E Bracketville (LSUMZ
32643), 9.9 mi W Bracketville (TCWC 63677), 13.6 mi E Spofford (TCWC 63918), 1.8 mi E & 4 mi

5 jet Rio Grande & Sycamore Creek (UTACV 7980); LaSalle Co., Cotulla (CU 1760), between Encinal

6 Atlee on 1-35 (TNHC 41907); Live Oak Co., 13 mi SSE Campbellton (TCWC 14796), George West
(TNHC 24492), near George West (TNHC 4306), 5 mi N George West (SM 8948), 3 mi W Three
Rivers (TCWC 10587), 5 mi SSW Three Rivers (TNHC 25892), 0.3 mi S Whitsett (KU 145894-
145895); Maverick Co., Eagle Pass (USNM 1972 holotype), 1 mi W Quemado (UMMZ 113153);
McMullen Co., N McMullen Co (UMMZ 74333), 2 mi E San Miguel Creek on Hwy 173 (TNHC
28757), Tilden (CU 612), 25 mi S & 12 mi E Tilden (TNHC 47506), jet Hwys 624 & 16 (TAIC 2046);
Refugio Co., 3 mi W Woodsboro (DMNH 3070); San Patricio Co., Aransas Pass (TAIC 3977), 1.5
mi NW Mathis (TNHC 24554), 2 mi N Mathis (TNHC 24567), 2.5 mi SE Mathis (TNHC 20406),

5 mi SW Mathis (TNHC 1 530 1), 7 mi SW Mathis (TNHC 28362), along Nueces Bay (SDNHM 23 1 90-
23191), 1 mi S Sinton (FSM 22104), Welder Wildlife Refuge (MVZ 128099, TCWC 65281); Starr
Co., 10.4 mi S Agua Nueva (TCWC 63112), Falcon Heights (TCWC 51838), 5 mi W McCook (CAS
103475), 5 mi W Roma (TCWC 6311 1); Val Verde Co., 2 mi W Del Rio (TNHC 42219), 2.5 mi W
Del Rio (TNHC 42220), W Sacatosa Creek at Hwy 85 (CAS 9856); Webb Co., 15 mi NNE Laredo
(AMNH 126734), 37.8 mi NW Laredo (TCWC 42361), 4.1 mi W jet Hwys 44 & 83 on 83 (TAIC
733); Willacy Co., 5 mi W Port Mansfield (TAIC 4072); Zapata Co., no locality (TNHC 23182), 1
mi NE Escobas (TCWC 48657), 5.3 mi S Laredo (UMMZ 74067), along Rio Grande (UMMZ 92797),

6 mi N San Ygnacio (TCWC 36388), 8.4 mi NE San Ygnacio (UTACV 10438); Zavala Co., 5.4 mi
S & 6.2 mi E LaPryor (TCWC 64997). MEXICO: COAHUILA: 3 mi E Muzquiz (KU 38331), 20 mi
E Nava (KU 39960), 18.5 mi SE Saltillo (EAL 3377), 3.4 mi S Villa Union (SDNHM 49792).
GUANAJUATO: Guanajuato (USNM 10240), Hwy 57 at San Luis Potosi state line (TCWC 65903).
HIDALGO: Ixmiquilpan (AMNH 72422), 77 km N Ixmiquilpan (ANSP 28689), 8.3 mi N Jacala
(UMMZ 106396), 12 mi S Jacala (UAZ 27006), 5 km S Venados (SDNHM 58399), 9 km S Zacualpan

Annals of Carnegie Museum

vol. 63

(TTU 6800), 1 1 km S & 1 km W Zacualpan (TTU 6799), 2 mi W Hwy 85 at S entrance to Zimipan
(LACM 128447). MICHOACAN: 2 mi W Morelia (MVZ 78364), Tacicuaro (FMNH 100296, USNM
111312). NUEVO LEON: 4 mi W Allende (KU 68117), Aramberri (KU 87748), 10.4 mi W Cerralvo
(EAL 3571), 12 km E & 15 km S Galeana (TU 17551-17552), 1 mi E Iturbida (TCWC 30430), 2 mi
S Linares (AMNH 69938), 19 km N & 9 km E Matehuala (FWM 7255), 4 mi S Monterrey (TU
17547), 13 mi E Monterrey (MCZ 46321), 25 mi S Monterrey (UIMNH 3836), 24.8 km S Sabinas
Hidalgo (TCWC 60760-60763). QUERETERO: 6.3 mi El Madrono (TCWC 29489), 0.2 mi E Jalpan
(TCWC 32928), km 106 on rd from Jalpan to San Juan del Rio (MCZ 157827), 1 mi N Pena Blanca
(TCWC 45658), 8 km N Queretero (SDNHM 49502), 6.4 mi NE San Juan del Rio (TCWC 38459),
1 mi E San Pablo Jet (TCWC 53061), 1 mi NW Toliman (TCWC 41012). SAN LUIS POTOSI:
Alvarez (MCZ 19024, 19028-19031, 46452), Bledos (LSUMZ 4204), between Charcas & Venado
(UMMZ 77245), 7.5 mi N El Valles (AMNH 67160), La Naranja (TU 17562), 8 mi NE Maiz (AMNH
1 10413), 13 mi E Maiz (AMNH 85246), 35 km W Maiz (UIMNH 17858), 30 mi NW Maiz (MVZ
129331), 10 mi E Matehuala (CM 60004), 20 mi S Matehuala (AMNH 93427), 24 mi S Matehuala
(EAL 24), 38 km SW Rio Verde (LSUMZ 5396), 8.5 mi SW San Luis Potosi (UMMZ 120221), 12.7
mi SW San Luis Potosi (FWM 8670), 43.4 km E San Luis Potosi (UTACV 12365), 4.5 mi NNW
Santo Domingo (EAL 4201). TAMAULIPAS: 4 mi E Aldama (UMMZ 101260), 50 mi S Brownsville
(USNM 64681), 25 mi N El Limon (UMMZ 102898), 1 mi S Fortuna (UMMZ 110956), 1.3 mi N
Gomez Farias (SM 11425), 4.3 km S Gomez Farias (UTACV 16135), 3 mi N Gomez Farias (SM
8947), Juamave (UMMZ 95204), La Joya de Salas (UMMZ 110818), 29 km N Mante (UMMZ
143724), 27 mi N Mante (UIMNH 3831), Matamoros (USNM 1974), 25 mi SW Matamoros (KU
61064), Miquihuana (MCZ 19552-19553), 11.7 km NW Morales (TNHC 28899), 8 km S Nuevo
Laredo (UTACV 12364), 0.3 mi SW Rancho Carricitos (TCWC 49933), 12 km SE Reynosa (UMMZ
143723), 20 mi ESE Reynosa (MVZ 36748), Soto La Marina (USNM 37546), Victoria (AMNH 75935),
7 mi N Victoria (MCZ 46322), 15.5 mi SW Victoria (UMMZ 1 14675), 22 km N Victoria (USNM
1 11279), 18 mi SW Victoria (AMNH 104465), 63 km E Victoria (TCWC 26730), 3 mi N Villagran
Monterrey (TNHC 32323), 1 mi S jet Hwy 101 & San Carlos rd (FSM 42308). VERACRUZ: 9 mi
SE Jalapa (UO 33467), 35 mi S Nautla (TAIC 2648).

Masticophis taeniatus x M. Schotti. — UNITED STATES: TEXAS: Uvalde Co., 13 mi N Uvalde
(TCWC 49204); Val Verde Co., 8.5 mi NW Del Rio on US 90 (CAS 9864), mouth Pecos River (UMMZ
74334). MEXICO: COAHUILA: del Carmen Mtns (FMNH 25301), 14.2 mi N Saltillo (TCWC 54321-
54322). NUEVO LEON: 20 mi SSW Galeana (FWM 7931), 5 mi E San Roberto (ANSP 28688).

Masticohpis bilineatus.— UNITED STATES: ARIZONA: no locality, (ANSP 5362); County un-
known, Colorado Bottom (USNM 1981); Cochise Co., Bisbee (BYU 256), 25 mi W Bisbee (CA 7809),
30 mi W Bisbee (CA 7470), 6 mi SSW Carr Canyon (AMNH 84969), Chiricahua Mtns (AMNH
75361, CU 10679, LSUMZ 8799, MVZ 7943, 8193, 44932, 97078, TCWC 63048, UMMZ 1 14130,
UTEP 2824, 9592), 1 mi N Dox Cabezos (MVZ 6942), 1 5 mi S Ft Huachuca (UIMNH 6062), Huachuca
Mtns (ANSP 16488, BYU 30029, 31880, MCZ 11409, MVZ 5406, 44742), Portal (AMNH 80816,
91620-91621, 109438-109439), 0.5 mi E Portal (CU 10660), 0.4 mi S & 0.8 mi W Portal (AMNH
1 1159), 1 mi SW Portal (AMNH 84972, 99333, CM 66092), 1.5 mi E Portal (MVZ 137578), 2 mi
W Portal (CA 5070), 2 mi S Portal (CM 66057), 2 mi E Portal (LACM 2257), 3 mi W Portal (AMNH
91622), 3.5 mi SW Portal (ASC 6597), 4 mi NW Portal (UMMZ 121839, 121917), 5 mi W Portal
(UMMZ 114633), Southwestern Research Station-SWRS (AMNH 80817, 84971, 111193-111194,
123866, FSM 8690, LACM 2255, MVZ 67188, 67204), 0.25 mi W SWRS (AMNH 84970), 2.5 mi
W SWRS (LACM 133880), 3.4 mi S Silverspur (UIMNH 72001), Tombstone (CU 2320), 20 mi W
Willcox (CA 16456); Gila Co., no locality (USL 18840), 2 mi NW Coolidge Dam (UMMZ 1 14634),
20 mi N Globe (TNHC 16793), Roosevelt Lake (USL 8408), 8 mi S Salt River (ASU 1716), 10 mi
N Salt River on Hwy 60 (CA 9520), Salt River Canyon (ASU 10536-10538), Sierra Ancha Experi-
mental Forest (CM 51907-51908); Graham Co., 5 mi SE Bonita (ASU 7030), Camp Grant (USNM
8434), 0.25 mi S Cyclone (UIMNH 24555), Gila River Sec 28 T6S R28E (CM 70672), Marijilda Base
Camp (ASU 7020, 7032-7033), 3 mi E Marijilda Base Camp (ASU 7014), Marijilda Canyon (ASU
7770, CM 4866 1 , 5 1 492, 70891, 70900, 70933, 70944), Pinaleno Mtns (CA 1 7006, CM 698 1 9, 7 1 04 1 ,
71067,71254,71537,71629,71667, 107289), 1 1 mi SW Salford (CA 11083), 1 1 mi S & 4.5 mi W
Safford (ASU 7011, 7017), 25 mi W Safford (CA 12974), 30 NW Safford (UIMNH 71999-72000),
Mt Turnbull (USNM 54719), 5.4 mi SW jet Hwys 366 & 666 on 366 (UTEP 9545); Greenlee Co.,
Eagle Creek (CM 71184, 70746), Gila River (CM 71425); Maricopa Co., 10-20 mi N Ajo (ASU
13846), 1 1 mi N Apache jet (CM 48620), Ashdale Ranger Station (USL 6724), 7.8 mi ENE Carefree
(CM 69160), Cave Creek (ASU 753), E of Mesa (ASU 59), Morristown (SDNHM 8414), 4.8 mi N
Morristown (CA 16158), 1 mi N Sunflower (ASU 3686), Tortilla Flats (ASU 9085, CAS 80670), 5.5
mi SE Wickenburg (UIMNH 71996-71997), 7 mi SE Wickenburg (UIMNH 71998); Mohave Co., 15

1994

Camper and Dixon— Striped Whipsnakes

km W Bagdad (ASU 14488), Hwy 93 at mile post 157.7 (ASU 22134); Pima Co., 0.5 mi S Ajo (UAZ
36767), 2 mi W Ajo (UAZ 40028), 24 mi S Ajo (NMSU 2634), Kitt Pk Observatory (UAZ 25158),
1.3 mi E jet Hwy 86 & Kitt Pk Rd on 86 (UAZ 32787), 2.8 mi from Hwy 86 & Kitt Pk Rd on 86
(UAZ 20794), 3.2 mi E jet Hwy 86 & Kitt Pk Rd on 86 (UAZ 20756), 3.6 mi from jet Hwy 86 &
Kitt Pk Rd on 86 (UAZ 20796), 4.1 mi E jet Hwy 86 & Kitt Pk Rd (UAZ 20795), 7.3 mi from jet
Hwy 86 & Kitt Pk Rd on 86 (UAZ 20786), 3.7 mi below jet Kitt Pk Rd & Picnic Area Rd (UAZ
20783), Organ Pipe Cactus National Monument (CM 48524, UIMNH 5611), 0.5 mi from Pima
Canyon mouth (TCWC 30720), 22.9 mi W Robles Jet (UAZ 20799), Sabino Canyon Rec Area (INHS
7571, 8007), Santa Rita Mtns (MU 2494), Sells (FSM 56664), 8 mi N Sonoita (UAZ 36250), 8.7 mi
N Sonoita (UAZ 20765), 15.2 mi N Sonoita (UAZ 34699), 8 mi N Tanque Verde Rd on Mt Lemon
Rd (MVZ 180258), 14 mi N Tucson (CA 3929), 50 mi S Tucson (EAL 883); Pinal Co., 30 mi S
Phoenix (CA 16157), Picket Post Mtns (CA 9825), Superior (ASU 1622, CA 9525, 9680), 5 mi W
Superior (CA 13579-13580), 6 mi E Winkleman (UIMNH 37712); Santa Cruz Co., 7.7 mi W Ft
Huachuca (UTEP 11162), 8.9 mi WNW Lochiel (LSUMZ 40983), Patagonia (CU 5234), 3 mi S
Patagonia (CU 5492), 2 mi SE Ruby (LSUMZ 32900), 8.8 mi W jet Hwys 89 & 289 (EAL 2770), 9.8
mi S Jet Hwys 82 & 83 on 83 (TCWC 63379); Yavapai Co., Bradshaw Mtns (AMNH 63977), 2.2 mi
NE Hillside (BYU 37042). NEW MEXICO: Hidalgo Co., Adobe Creek center Sec 24 T31S R19W
(AMNH 77453), Animas Mtns (AMNH 84976, 87276, UNM 6979), 8 mi W Animas (AMNH 84974-
84975), 8.5 mi S & 2 mi W Animas (NMSU 6395), 3.5 mi W Cloverdale (UNM 8298), 7.5 mi NW
Cloverdale (LACM 2253), 10.5 mi N & 13.5 mi E Cloverdale (UTEP 1344), 16 mi NE Cloverdale
(CM 18222), Guadalupe Mtns (UNM 3334, 6786, 8297), Indian Creek Canyon (UNM 13820), Pe-
loncillo Mtns (AMNH 75949-75950, 119512, UNM 7687), 7 mi S Rodeo (MVZ 67203), 8 mi SSE
Rodeo (CU 5768), 17 mi N & 3 mi E Rodeo (UNM 39100), San Geronimo Trail 5 mi E AZ state
line (LACM 2254), 2 mi SW Steins (LSUMZ 8806), 8.2 mi S jet Hwys I- 10 & 80 on 80 (LACM
76563), SW 0.25 Sec 15 T34S R21W (UNM 32349). MEXICO: AGUASCALIENTES: 2 km W
Calvillo (UTEP 7747), 4.5 mi E Calvillo (LSUMZ 35033). CHIHUAHUA: near Batopilas (USNM
46382). COLIMA: no locality (MCZ 1 1409), Colima (AMNH 19636). DURANGO: Santa Ana (KU
74567), 2 km E Villa Corona (UTEP 3977). JALISCO: 2 mi E Bolanos (KU 91424), 10.1 km SW
Cocula (TCWC 48059), Guadalajara (USNM 32212), 4.8 km S Huejucar (KU 102966), Hwy 15 S of
Lago de Chapala (TCWC 33793), 5.3 mi S Nayarit state line near Ixtlan del Rio (UIMNH 40361),
13 mi N Sayula (AMNH 93339), 4.8 km E Totatiche (KU 100512), 10 mi NE Union de Tula (MVZ
7220). NAYARIT: 17.9 mi SW Acaponeta (FSM 24761), 1 1.4 mi S Acaponeta turnoff (LACM 6990),
Jesus Maria (AMNH 74953-74956), between Marquezado & Auzata (AMNH 19640), 15 mi E San
Bias (UIMNH 7 1988), Hwy 1 5 near Sinaloa border (LACM 6989), 2.4 mi N Tecuala turnoff (UIMNH
85728), Tepic (USNM 46417), rd from Tepic to Paga (LACM 8739). OAXACA: Cuicatlam (USNM
46499). SINALOA: 4 mi S Casa Blanca (KU 83406), 0.2 mi W Concordia (LACM 6992), 10 mi W
Concordia (UMMZ 102483), 15.6 mi N Culiacan (UIMNH 71992), 37.5 mi N Culiacan (UIMNH
71991), 72 mi S Culiacan (FSM 42096), Elota (UIMNH 83457), 12 mi N Escuinapa (UIMNH 41593),
17.2 mi NW Escuinapa (TNHC 25475), 30 mi S Escuinapa (UIMNH 71994), Isla Palmito del Verde
(KU 73569-73570), 1 5 mi N Los Mochis (CM 54004), Mazatlaan (TCWC 21903), 1.1 mi N Mazatlan
(UIMNH 83456), 15.8 mi N Mazatlan (LACM 103106), 26.8 mi N Mazatlan (LACM 103104), 29
mi N Mazatlan (UIMNH 71989), 52.4 mi N Mazatlan (UIMNH 71900), Hwy 15 5 mi N Nayarit
state line (LACM 6991), Rosario (KU 73568), San Ignacio (LACM 6986), Terreros (LACM 6993), 8
km N Villa Union (KU 80757), 5.8 mi NW Villa Union (CAS 241 14), 18 mi SW Villa Union (CAS
120883), 34 mi NE Villa Union (LACM 103105), 6.2 mi N & 10.6 mi E jet Hwys 40 & 15 (CAS
23954-23955). SONORA: no locality (ASU 8421), 47 km E Agua Prieta (UTACV 17796), Alamos
(MCZ 43251), W of Alamos (CAS 140518), 1 mi W Alamos (LACM 103109), 2 mi SSE Alamos
(LACM 103107), 4 mi W Alamos (LACM 103108), 7 mi W Alamos (ASU 6037-6038, 6454, 6645),
17 mi W Alamos (ASU 6510), 17 mi S Casa Blanca (MVZ 67430), 0.4 mi NE El Coyote (MVZ
136780), S El Novillo 9MVZ 136781), Guaymas (MTKD 15068 paratype, 15523 holotype, USNM
15880), 4 mi SW Guaymas (KU 48919-48920), Guirocoba (AMNH 63722-63723, MVZ 50793-
50796), 5 mi S Hermosillo (UMMZ 72103), 5 mi SW Hermosillo 9FMNH 102669), 32 mi SE
Hermosillo (AMNH 84978), Isla San Esteban (KU 91587, LACM 20601-20604, 74047, MVZ 74954-
74956, 76499, SDNHM 3826, 40855, TTU 8017, UMMZ 128928), Isla San Pedro Martir (UO 32528),
Isla Tiburon (USNM 222054-222058), 5 mi SE La Pintada (AMNH 808 1 5), La Posa (FMNH 102670,
UIMNH 17838), Las Chispas (AMNH 3469), 25 mi W Maxatan (AMNH 84977), San Luis Mtns
(USNM 21053-21054, UTACV 17794-17795), 4.5 mi S Tubutama (CAS 16642), 5 mi S Tubutama
(CAS 17297), between Vicam & Guaymas (LACM 103110). ZACATECAS: 9 mi SSW Jalpa (UTEP
7748), 1.7 mi N Los Bajios (LSUMZ 26233), San Juan Capistrano (USNM 46481), 7 km S Santa
Rosa (UTEP 3978), Zapogui (USL 12283).

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vol. 63

Appendix 2
Electrophoresis Samples

Masticophis bilineatus. - ARIZONA: Cochise Co. (TCWC 63696); Pima Co. (TCWC 64995). NEW
MEXICO: Hidalgo Co. (TCWC 64842).

Masticophis s. schotti. — TEXAS: Frio Co. (TCWC 63774, 63919); Jim Wells Co. (TCWC 64839);
San Patricio Co. (TCWC 65281); Zavala Co. (TCWC 64997).

M. s. ruthveni.- TEXAS: Cameron Co. (TCWC 65282); Hidalgo Co. (TCWC 65283). MEXICO:
GUANAJUATO: (TCWC 65903).

Masticophis t. taeniatus. — IDAHO: Ada Co. (TCWC 64836-64837). NEW MEXICO: Catron Co.
(TCWC 65001); Hidalgo Co. (UTEP 1 1355); Sierra Co. (NMSU uncatalogued); Socorro Co. (TCWC
66165).

M. t. girardi. -TEXAS: Brewster Co. (TCWC 64999, 65285); Edwards Co. (TCWC 65923); El Paso
Co. (TCWC 64998, UTEP 10804); Kimble Co. (TCWC 64809, 65284, 65287); Llano Co. (TCWC
63772-63773, 63876, 64841, 64996, 65000); McCulloch Co. (TCWC 64838); Medina Co. (TCWC
63775); Presidio Co. (TCWC 64810-64811, 65286); Uvalde Co. (TCWC 64840).

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 1, P. 49-65

22 February 1994

U-SHAPED ORIENTATION OF HUNTER-SCHREGER BANDS IN
THE ENAMEL OF MOROPUS (MAMMALIA: CHALICOTHERIIDAE)

IN COMPARISON TO SOME OTHER PERISSODACTYLA

Wighart v. Koenigs wald1
Abstract

In the Chalicotheriidae the main shearing facets of upper and lower molars possess a very significant
modification of the direction of Hunter-Schreger bands (HSB). In contrast to the primitive horizontal
HSB orientation, chalicotheres show distinctly U-shaped HSB that intersect the main cutting edges
at almost right angles. This relationship is maintained throughout the various stages of wear. This
specific characteristic is shared with Brontotheriidae. Certain other Perissodactyla, such as Tapiroidea
and Rhinocerotidae, but definitely not the Equoidea, follow a similar tendency. Functionally, the
reorientation of the HSB is interpreted as an adaptation to reduce abrasion. The reorientation of the
HSB seems to be related to interprismatic matrix (IPM) parallel to the prisms.

ZUSAMMENFASSUNG

Bei den Chalicotheriidae zeigen die Hauptschneidekanten der oberen und unteren Molaren eine
aulfallende Abweichung von der normalerweise horizontalen Orientierung der Hunter-Schreger-Ban-
der (HSB). Die HSB sind U-formig gebogenen und stehen annahemd senkrecht auf den Hauptschnei-
dekanten. Dieser Winkel bleibt durch diese spezielle Anordnung auch wahrend der fortschreitenden
Abkauung erhalten. Dieses abgeleitete Merkmal teilen die Chalicotheriidae mit den Brontotheriidae.
Andere Perissodactyla, wie die Tapiroidea und Rhinocerotidae, nicht aber die Equoidea, verfolgen
eine ahnliche Strategic. Funktionell wird diese Reorientierung der HSB als Anpassung gedeutet, durch
die der Abrieb der Schneidekante reduziert wird. Die Reorientierung der HSB scheint an eine Inter-
prismatische Matrix (IPM) gebunden sein, die parallel zu den Prismen liegt.

Introduction

The prismatic enamel of most large mammals is characterized by Hunter-
Schreger bands (HSB). These are light and dark bands often visible in low-powered
light microscopy. The SEM shows their nature as decussating layers of enamel
prisms (Fig. 1). The most common orientation of these bands is roughly parallel
to the alveolar surface of the jaw, which is here termed the “horizontal orienta-
tion.”

In contrast to most other large mammals, several groups of perissodactyls such
as Chalicotheriidae, Brontotheriidae, Tapiroidea, and Rhinocerotidae (but not
Equoidea) modified the horizontal orientation of HSB. The vertical orientation
of HSB in Rhinocerotidae was described by Rensberger and Koenigswald (1980)
and Boyde and Fortelius (1986). Rensberger and Koenigswald (1980) interpreted
the modified orientations found in Tapiroidea and Brontotheriidae as structural
intermediates in the evolution toward Rhinocerotidae.

This paper describes the modified orientation of HSB in the schmelzmuster of
Chalicotheriidae. The study is based on various jaws (including milk dentitions)
of Moropus elatus and Moropus sp. from the Agate Springs Quarries, lower Har-
rison Formation, Nebraska (Holland and Peterson, 1914). The extremely well-

1 Institut fur Palaontologie der Universitat Bonn, Nussallee 8, D-531 15 Bonn, Germany.
Submitted 3 March 1993.

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vol. 63

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Koenigswald— Enamel Structure of Moropus

preserved enamel and the specific schmelzmuster allows a tracing of the orien-
tation of the HSB over almost the entire surface of the teeth. The material was
generously made available by the Carnegie Museum of Natural History.

The schmelzmuster of Moropus is compared with that of other Chalicotheriidae,
Brontotheriidae, and other Perissodactyla. This survey allows a discussion of
functional aspects of the orientation of HSB.

Hunter- Schreger Bands

Hunter-Schreger bands (HSB) constitute one of the basic enamel types (Ko-
enigswald, 1980, 1982) widely distributed among mammals. This enamel type is
characterized by layers of enamel prisms of regularly alternating direction. Prisms
of adjacent layers often decussate at a high angle of up to 90°. The thickness of
the bands counted in prism diameters is significant for rodent incisors (pauciserial,
multiserial, or uniserial) (Korvenkontio, 1934; Wahlert, 1968), but shows a wide
individual variation in larger mammals (Fig. 1). Average thickness between 5 and
20 prisms have been counted by Kawai (1955) for various species. Mostly com-
monly, thickness tends to be around ten prisms. But thickness is only one of the
characters of the HSB. The bands share regular bifurcation patterns as shown in
Fig. 5C, 6C, and prisms change over from one band to the next in the transitional
zones between bands (Koenigswald and Pfretzschner, 1987). As seen in cross
sections through the enamel, HSB often form only one layer of a given schmelz-
muster and are combined with an inner and/or outer layer of radial enamel, a
different enamel type in which the prisms do not decussate.

As planar elements, the HSB show two distinct directions (Fig. 2). The one
observable in the vertical cross section between the enamel-dentine junction (EDJ)
and the outer surface is defined as inclination (Korvenkontio, 1934). The incli-
nation is 0° when the bands are perpendicular to the EDJ. The second direction
is the lateral orientation which can be studied on the translucent enamel surface
or in tangential sections through the layer with HSB. Based on commonality among
Mammalia, the least derived lateral orientation is horizontal— that is, parallel to
the alveolar margin of the jaws. This paper concentrates on modifications of the
lateral orientation of HSB.

Methods

Enamel prisms function like optic fibers when exposed to a strong source of
light. The individual prisms are too thin to be observed under a low-powered
microscope, but since all prisms within one band have the same orientation, their
optic activity sums up, and HSB become visible even at low magnification. When
the enamel is illuminated parallel to the bands, they become visible as light and
dark areas. The prisms of all dark areas point toward the light source, whereas
the prisms in light areas form a larger angle with the direction of the light (Ko-
enigswald and Pfretzschner, 1987). By handling the tooth in front of a strong light
source like fiber optics under a binocular microscope, the lateral orientation of
the HSB can be studied at 20 to 50 times magnification. At the same time, the

Fig. 1.— Moropus sp., Agate Springs quarries, Nebraska; Harrison Formation, Miocene. Scanning
electron micrograph of longitudinal section through the protoconid of the lower molar, depicting
Hunter-Schreger bands of varying thickness. (EDJ = enamel dentine junction.)

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vol. 63

Fig. 2. — Diagram to illustrate inclination and lateral orientation of Hunter-Schreger bands (HSB).
Inclination is defined as the angle between the HSB and a plane parallel to the enamel dentine junction
(EDJ). It is seen best in a vertical cross section. Lateral orientation is usually horizontal but in some
cases, as in chalicotheres, rotated. The HSB include an angle with the plane parallel to the alveolar
margin of the jaw. In the enamel depicted here, two layers of different enamel types form the schmelz-
muster, as in chalicotheres or brontotheres, having an inner layer with HSB and on outer layer with
radial enamel.

unilateral bifurcation of the bands then becomes visible (Koenigswald and Pfretz-
schner, 1987).

In order to facilitate the mapping of these observations, casts were made from
the teeth studied and the directions of the HSB were drawn on the plaster casts
as observed.

A thick layer of outer radial enamel or a structured enamel surface can often
obscure the visibility of the HSB. Pericymata on the enamel surface, although
occasionally parallelling the HSB, must not be mistaken for HSB, since they are
fundamentally independent structures.

To study the schmelzmuster and the orientation of the HSB from sections of
teeth or tooth fragments, they have to be embedded into epoxy resin, sectioned,
and ground in the desired direction. On a tangential section through the layer
containing HSB, the lateral orientation of the HSB can be studied even when they
are not visible from the outside. The prisms show the same optical properties in
sections as in surface views. These properties become even more visible when
etched (e.g., with 2N HC1 for 2-5 sec). The etched surfaces, coated with a con-

1994

Koenigswald— Enamel Structure of Moropus

mesial

buccal

ial 1

M sup.

Mesostyle

Metacone

Metaloph

Hypocone

M inf.

Hypoconid

Metaloph id
Hypolophid

Entoconid

Fig. 3.— Moropus sp. Tooth cusps in upper and lower molars. Modified after Coombs (1978) and
Thenius (1989).

ducting element (e.g., gold palladium), can be studied for further details of the
complex enamel structure under the SEM.

Morphological terms used for the description of Moropus (Fig. 3) follow Thenius
(1989) and Coombs (1978).

SCHMELZMUSTER OF MOROPUS

The schmelzmuster of Moropus is characterized by HSB that continue from
the EDJ almost to the outer surface (Fig. 1). The angle of decussation is reduced
in an outer zone which continues into a thin radial enamel. The inclination of
the HSB is slight and concave upwards. In a tangential view of the HSB are slightly
wavy and bifurcate frequently. The lateral orientation of the HSB varies: it is
horizontal at the base of the crown but strongly bent to intersect the main cutting
edges almost vertically. A detailed description is given below. The prisms generally

Annals of Carnegie Museum

vol. 63

Fig. A.— M or opus sp., Agate Springs quarries, Nebraska; Harrison Formation, Miocene. A: SEM
micrograph of enamel prism cross sections in a tangential ground and etched section of the lower
molar. Most prisms have an open but somewhat irregular prism sheath. B: Prisms in cross section
and longitudinal aspect at the border between HSB. The IPM is almost parallel to the prisms.

have an incomplete prism sheath (Fig. 4A). The cross section is rounded or angular.
Prisms show a “seam” (Lester and Koenigswald, 1989), especially near the EDJ.
The IPM crystallites run almost parallel to the prisms and surround them.

Orientation of Hunter- Schreger Bands

Upper molar morphology is characterized by a single series of shearing facets
which is formed by the ectoloph (Fig. 3). While parastyle and mesostyle remain
almost vertical, the paracone and metacone are strongly inclined, which results
in a strong flexion of the ectoloph. The enamel on the buccal side is much thicker
than on the lingual side. The protocone and the hypocone are on the lingual side.
The hypocone forms a second shearing facet positioned lingually from the meta-
cone. The protocone is only slightly linked to the ectoloph by the protoloph and
forms an almost isolated rounded cusp.

In the upper molars the HSB are oriented horizontally in the cingulum and at
the base of the crown. The horizontal orientation of the HSB is retained on the
lingual side of the hypocone and on all sides of the protocone to its full height.
On the buccal side of the ectoloph, however, the lateral orientation of the HSB
is strongly modified as follows. Parastyle and mesostyle retain the horizontal HSB,
but on the prominent paracone the ectoloph forms two cutting edges that enclose
an angle of roughly 90°. HSB are bent in a U-shaped fashion to intersect the
cutting edges at an almost right angle (Fig. 5A, C). Below the tip of the paracone

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Koenigswald— Enamel Structure of Moropus

pas

ms hy me pas pa me hy

1cm

^ 1mm

Fig. 5.— Moropus sp., Agate Springs quarries, Nebraska; Harrison Formation, Miocene. Upper M3.
A: diagrammatic HSB orientation in the ectoloph between parastyle and mesostyle. B: diagrammatic
HSB orientation in the ectoloph between mesostyle and metastyle. C: detailed mapping of the U-shaped
HSB orientation in the paracone (detail from A). Abbreviations: if, interface between fields of HSB;
hy, hypocone; me, metacone; ms, mesostyle; pa, paracone, pas, parastyle.

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vol. 63

the HSB pass through a short horizontal section. If the paracone rib is prominent,
the HSB may even be flexed downward forming an acute angle. Between the
horizontal HSB related to the paracone and the inclined HSB related to the
parastyle, there is a distinct interface in which the HSB of both dental elements
meet at an acute angle. This interface is marked by a slight crease in the enamel
surface. But between paracone and mesostyle the fields of HSB grade into each
other without a distinct border. Since HSB bifurcate frequently, such a transition
from strongly inclined to horizontal HSB is made possible.

In the distal part of the ectoloph the metacone is not morphologically distinct.
However, the orientation of the HSB clearly indicates the position of this cusp
within the ectoloph. On the metacone the HSB orientation is similar to that of
the paracone. Whereas HSB are steeply oriented on both flanks they pass through
a narrow horizontal stretch below the cusp. Because of the U-shape, the HSB are
again oriented almost normal to the cutting edge between metacone and mesostyle.
On the other flank of the metacone they intersect the descending ectoloph at a
high angle. The hypocone, which is linked to the ectoloph by the metaloph, shows
a U-shaped orientation of the HSB on the buccal side. On the lingual side HSB
orientation is more or less horizontal.

The upper milk dentition is generally molariform and follows the same basic
pattern. In the premolars the protocone is relatively bigger and somewhat buc-
colingually compressed, thus forming a short cutting edge comparable to the
ectoloph. The HSB are slightly flexed and therefore approach the cutting edges at
almost right angles as well.

In the lower molars trigonid and talonid form similar triangles which are con-
nected at the twin cones of metaconid and metastylid. The W-shaped buccal side
forms the shearing facets working against the ectoloph of the upper molars. The
enamel is again thicker on the buccal side than on the lingual side.

The W-shaped cutting edge is formed by the paralophid, protolophid, metalo-
phid, and hypolophid (Fig. 3). The base of the crown and the lingual sides of the
main cusps show distinct horizontally-oriented HSB (Fig. 6A). In a protoconid
and hypoconid the HSB are similarly U-shaped as in the ectoloph of the upper
molars. While the HSB pass through a narrow section of the horizontal orientation
and the buccal side of the protoconid and hypoconid, they bend upward toward
the cutting edges on both flanks of each cone (Fig. 6A). Distinct interfaces are
found between the cusps situated more to the lingual side. The interfaces are best
seen in the paralophid and hypolophid separating the field of HSB related to the
paraconid and protoconid and the entoconid and hypoconid, respectively (Fig.
6B, C). Distinct interfaces separate the HSB fields of the twin cusp metaconid/
metastylid from that of the protoconid and hypoconid. These interfaces converge
onto the central valley and do not reach the base of the crown.

The molariform P4 and the milk teeth follow a similar pattern as far as they
could be investigated. In the anterior premolars (P2 and P3) the HSB are slightly
flexed towards the shearing facets. In the incisors of the lower jaw HSB are all
horizontally oriented.

The pattern described above was found in all available teeth of Moropus elatus
and Moropus sp. as well as in the European Chalicotherium goldfussi and the
Asian Chalicotherium cf. habereri. The congruence of this character in several
related genera excludes the possibility that the U-shaped HSB orientation is an
individual variation and implies that it is characteristic of the Chalicotheriidae
in general. Individual variation within the group occurs on a much smaller scale,
and concerns the thickness, bifurcation, and exact orientation of HSB.

1994

Koenigswald— Enamel Structure of Moropus

Fig. 6.— Moropus sp., Agate Springs quarries, Nebraska; Harrison Formation, Miocene. Lower M,.
A: diagrammatic HSB orientation on the buccal side. B: schematic HSB orientation on the hypolophid
with interface. C: detailed mapping of the HSB orientation in the hypolophid with an interface between
areas of the hypoconid and entoconid. Abbreviations: end, entoconid; if, interface between fields of
HSB; hyd, hypoconid; med, metaconid; msd, metastylid; pad, paraconid; prd, protoconid.

Annals of Carnegie Museum

vol. 63

Orientation of the HSB in
Other Perissodactyls

Equoidea

In the well-documented evolution of the Equidae, the HSB consistently retain
a horizontal orientation. The basic schmelzmuster, as in Hyracotherium , shows
horizontal HSB penetrating from the EDI almost to the outer surface where a
thin radial enamel is found. The crystallites of the IPM run parallel to the prisms
(Pfretzschner, 1 993, in press). In Mesohippus and Anchitherium this basic schmelz-
muster is retained, but the IPM crystallites form an angle with prisms and are
arranged in inter-row sheets.

When the molars of Equidae become hypsodont as in Merychippus, Hipparion,
or Equus, the schmelzmuster is reorganized. A layer of “modified radial enamel”
characterized by thick, vertical inter-row sheets is introduced between the EDI
and the layer of HSB (Pfretzschner, 1993, in press; Koenigswald et al, 1993).
Nevertheless, the horizontal orientation of the HSB is retained unchanged.

Paiaeotherium sp. (Palaeotheriidae) shows well-developed HSB (Remy, 1976)
which are generally oriented horizontally. Although the paracone and metacone
in the ectoloph of the upper molars are similarly inclined and shaped as in Mom-
pus , the HSB retain their horizontal orientation (Fig. 7 A). This documents that
HSB orientation is independent of the shape of the ectoloph.

Brontotheriidae

In comparison to the Chalicotheriidae studied, the enamel in the larger Bron-
totheriidae, such as Menodus proutii, is fairly thick. The schmelzmuster of Bron-
totheriidae is in most cases two-layered. An inner layer is formed by HSB which
are only very slightly inclined toward the occlusal surface concave up. The thick-
ness of HSB with an oblique lateral orientation is difficult to quantify since the
likelihood of finding these bands at an angle other than 90° is very high. The
thickness counts therefore are commonly too high. The HSB disappear in the
outer layer of radial enamel. This outer layer is almost as thick as the inner layer
and prevents observation of the HSB from the outer surface of the enamel.

The prisms of the radial enamel rise apically at a maximum of about 45°. The
prisms have incomplete prism sheaths and the IPM is mainly parallel to the
prisms.

In areas with thin enamel, the outer radial enamel may be reduced in thickness
or missing. On the occlusal surface, however, the outer radial enamel on the
outside is worn off more rapidly than the inner layer of HSB. Therefore, the
direction of the HSB can be observed near the occlusal surface. In the angled
ectoloph of the upper molars, a U-shaped orientation is found at the paracone
and the metacone as described for M or opus. In lower molars a similar EJ-shaped
orientation of the HSB is found at the hypoconid and the protoconid just as in
Moropus. In the paralophid, protolophid, metalophid, and hypolophid, interfaces
with very sharp turns of the HSB are present as described for the Chalicotheriidae.
These interfaces are visible at the occlusal surface but can be observed better in
a tangential ground section.

Tapir oidea

The schmelzmuster of Tapirus (Tapiridae) is characterized by horizontal HSB
which extend to the outer surface. The prisms show an open prism sheath and

1994

Koenigswald— Enamel Structure of Moropus

pas

1cm

Fig. 7. — Hunter-Schreger band orientation in Equoidea and Rhinocerotidae. A: Palaeotherium sp.
(Equoidea) Frohnstetten, Germany, lower Oligocene. Diagrammatic illustration of the horizontal HSB
orientation in the ectoloph of an upper molar. B: Floridaceras whiiei (Rhinocerotidae). Diagrammatic
illustration of the vertical HSB in the ectoloph of an upper molar. Abbreviations: me, metacone; ms,
mesostyle; pa, paracone; pas, parastyle.

are surrounded by IPM, the crystallites of which are almost parallel to the prisms.
HSB orientation is modified in Tapirus which was discussed for lower molars by
Rensberger and Koenigswald (1980). In the lower molars, the HSB of the lingual
and buccal sides of the main cusps are horizontal. In the transverse lophs, the
HSB bend into an almost vertical direction. In the middle lophs, the fields of
HSB of the two involved cusps meet at a distinct interface which often can be
seen in the worn occlusal surface as the angle between HSB (Fortelius, 1984: fig.
2b) or on the unworn surface as a slight groove (Fig. 8).

In the upper molars, the horizontal HSB dominate. Only in the uppermost parts
of the transverse lophs can similar orientations of the HSB and incipient interfaces
be observed.

In Colodon cingulatus (Helatelidae) from the Oligocene of Montana, a similar
schmelzmuster was found (Rensberger and Koenigswald, 1980), but no material
for sectioning was available.

Lophiodon lautricense (Lophiodontidae), a large tapiroid from the Eocene of
Europe, follows the tapiroid schmelzmuster, even though an outer zone of radial
enamel occurs in some parts of the molars. The prism sheath is generally incom-
plete and the IPM is parallel to the prisms.

In the upper molars the horizontal HSB are widely retained. Only in the trans-
verse lophs do the HSB turn into a vertical position. Interfaces were not seen in
the available material. In the lower molars the protoconid and the hypoconid
show horizontal HSB. Toward the transverse lophs, the HSB turn vertically.
Interfaces are well-marked by a slight groove in the enamel surface and on the
occlusal surface by an irregularity of the HSB. The lower molars have horizontal
FISB which are strongly modified in the lingual walls of metaconid and entoconid.

Annals of Carnegie Museum

vol. 63

Fig. 8 ,—Tapirus prisms , Eppelsheim, Germany,, Upper Miocene. Unretouched photo of the anterior
loph of the lower right M3 with horizontal HSB in the cusps, inclined HSB in the loph and an interface
in the center of the loph where the areas of protoconid (left) and metaconid (right) merge into each
other. Length of bar equals 5 mm.

Heavy undulation of the HSB forms vertical structures in the outer enamel. These
unusual structures resemble the modification of HSB found in hyaenids (personal
observation).

Rhinocerotidae

The schmelzmuster in the molars of most rhinos (e.g., Cododonta antiquitatis )
consists of an inner layer of HSB and an outer layer of radial enamel The outer
layer varies considerably in different genera. It can be very thin or even missing
as in Subhyracodon or Floridaceras whitei.

Rhinocerotid upper and lower molars are characterized by vertical HSB as in
Floridaceras whitei (Fig. 7B). The vertical orientation is not restricted to certain
parts of the occlusal surface as in Chalicotheriidae, Brontotheriidae, or Tapiroidea,
but continues around the entire tooth and does not include any interfaces. These
HSB bifurcate in the same manner as horizontal HSB. These vertical HSB have
been studied by Rensberger and Koenigswald (1980) and Boyde and Fortelius
(1986). The authors were not aware that this peculiar structure in rhino teeth,
which results in a specific roughness of the occlusal surface, was already observed
and carefully figured more than 100 years earlier by Quenstedt (1867: pi. 1:1, pi
2:35).

In incisors the HSB are horizontal but intersect the very steep shearing facets
at nearly right angles (Koenigswald, 1985; Koenigswald and Clemens, 1992). In
Coelodonta, the prisms have incomplete prism sheaths and the IPM is mostly
parallel to the prisms.

Based on the enamel surface investigation of a mandible of Metamynodon sp.

1994

Koenigswald —Enamel Structure of Moropus

Table 1 .—Perissodactyl tooth material investigated in this study. 1, investigated by light microscopy on
natural surfaces only; 2, investigated in sections under light microscope and SEM.

Chalicotheriidae

Moropus elatus, Agate Springs quarries, Nebraska; Harrison Formation, Miocene; Carnegie Muse-
um of Natural History; 1 .

Moropus sp., Agate Springs quarries, Nebraska; Harrison Formation, Miocene; Carnegie Museum
of Natural History; 1+2 [KOE 1436].

Chalicotherium goldfussi, Eppeisheim, Germany; Upper Miocene; Hessisches Landesmuseum,
Darmstadt; 1.

Chalicotherium cf. haberreri, Pavlowelar, Kazahkstan; Turolian; Paleontological Museum, Mos-
cow; 1.

Equoidea

Hyracotherium sp., Bighorn Basin, Wyoming; Will wood Formation, lower Eocene; U.S. Geological
Survey, Denver; 1+2 [KOE 1022].

Equus sp., Heidenheim, Germany; upper Pleistocene; 1+2 [KOE 34].

Palaeotherium sp., Frohnstetten, Germany; Oligocene; 1+2 [KOE 1666].

Brontotheriidae

Menodus proutii, White River, Nebraska; Oligocene; Hessisches Landesmuseum, Darmstadt; 1 .

large brontothere, gen. indet., Nebraska; Chadron Formation, White River Group, Nebraska; Ne-
braska State Museum, Lincoln; 1+2 [KOE 1637],

Tapiroidea

Tapirus priscus, Gau Weinheim, Germany; upper Miocene; Hessisches Landesmuseum, Darm-
stadt; 1.

Tapirus priscus, Esselbom, Germany; upper Miocene; Hessisches Landesmuseum, Darmstadt; 1 .

Tapirus sp., China; Pleistocene; 1+2 [KOE 73].

Colodon cingulatus, Oligocene, Montana; Carnegie Museum of Natural History; 1 .

Lophiodon lautricense, Robiac, France; late Eocene; Hessisches Landesmuseum, Darmstadt; 1.

Rhinocerotidae

Ceratotherium simum, Africa; Recent; 1+2 [KOE 610].

Rhinoceros sp., Sangiran Java; Pleistocene; 1+2 [KOE 71].

Coelodonta antiquitatis, Urspringhohle, Germany; 1+2 [KOE 52].

Subhyracodon occidentale, M3; Museum of Comparative Zoology, Harvard University; 1+2 [KOE
356].

Floridaceras whitei, Hawthorne Formation; Miocene; Thomas Farm, Florida; Museum of Compar-
ative Zoology, Harvard University; 1+2 [KOE 357].

(Amynodontinae), the HSB orientation and the schmelzmuster seem to follow
the same pattern as Rhinocerotidae.

Discussion

Hunter-Schreger bands were evolved several times in parallel among various
mammalian lineages (Koenigswald and Clemens, 1992), for instance in the early
Paleocene by condylarthrans (Koenigswald et al., 1987), in primates, carnivores,
rodents, and a few marsupials. Despite this multiple origin, the usual orientation
of HSB is horizontal. Therefore this orientation is regarded as the most primitive
condition for mammals in general as well as for perissodactyls.

The interpretation of the variability of the schmelzmuster among the Perisso-
dactyla from a phylogenetic perspective allows us to differentiate the derived
genera within the various families and superfamilies, respectively. The Equoidea
retain the horizontal HSB. However, when the molars become hypsodont, a
significant layer of modified radial enamel is introduced between the EDJ and

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vol. 63

the layer of HSB. This schmelzmuster developed convergently in several other
large herbivores with hypsodont molars (Pfretzschner, in press; Koenigswald et
al., 1993). These large herbivores share the characteristic of having the IPM at a
high (close to 90°) angle with the prisms.

Other Perissodactyla, such as Brontotheriidae, Chalicotheriidae, Tapiroidea,
and Rhinocerotidae, follow different pathways of modification in their schmelz-
muster. Brontotheriidae, widely accepted to be closely related to Equoida, keep
the crystallites of the IPM parallel to the prisms and develop the U-shaped ori-
entation of the HSB. The strongly curved HSB intersect the occlusal surface at
almost right angles. They share this derived character with the Chalicotheriidae.
Differences between some Chalicotheriidae and Brontotheriidae are expressed
mainly in the thickness of the outer radial enamel. Whether the very similar
orientation of the HSB in both families is a synapomorphy must be tested by a
larger survey of the schmelzmuster in both groups.

In Tapiroidea the HSB turn in a vertical direction only in the transverse lophs
immediately below the crest. Tapirus and Lophiodon differ in the enamel on the
lingual side of the lower molars.

The Rhinocerotidae have a very derived schmelzmuster with vertical HSB. The
vertical orientation of HSB in Rhinocerotidae is unique among Perissodactyla.
However, a similar orientation evolved convergently in astrapotheres (Fortelius,
1984, 1985; Rensberger and Pfretzschner, 1992), but not in Arsinoitherium as
assumed by Fortelius (1984) and since disproven by Pfretzschner (in press).

Whereas in the Rhinocerotidae the HSB are vertical around the entire molars,
in tapirs, chalicotheres, and brontotheres, the HSB of the various cusps developed
vertical HSB individually. The interfaces, found as distinct lines between the
cusps even when they are united in the same loph, are the most obvious feature
indicating this different strategy.

Hyracoidea, which are regarded by some as a sister group of the Perissodactyla
(Fischer, 1986), have a horizontal HSB orientation in some genera (Fortelius,
1985), but, astonishingly, most genera lack HSB altogether.

A strong functional significance of the modified orientation of the HSB in
perissodactyls is indicated first by the restriction of the modifications to the main
cutting edges, and second by the parallel occurrence of this tendency in several
lineages such as in Chalicotheriidae, Brontotheriidae, Tapiroidea, and Rhinocer-
otidae. The functional significance here is that the HSB intersect the main shearing
facets at almost right angles. The modified orientation of HSB brings a large
number of prisms into a direction favorable for reduced wear (Rensberger and
Koenigswald, 1980; Boyde and Fortelius, 1986). This modification has reached
its greatest perfection in the Rhinocerotidae, but is not found in Equoidea or any
member of the Artiodactyla.

Even if the teeth are to some extent worn, the high angle between the cutting
edge and the HSB is maintained in chalicotheres and brontotheres and especially
in Rhinos.

Three apparently antagonistic hypotheses were formulated to explain the adap-
tive significance of vertical HSB in Rhinocerotidae. Rensberger and Koenigswald
(1980) argued that the vertical HSB allow a maximum number of prisms to
intersect the occlusal surface at almost right angles. Their experimental data dem-
onstrated that there is less abrasion of the enamel when prisms are at right angles
to the occlusal surface compared to prisms parallel to the occlusal surface. Pfretz-
schner (1992, in press) argues that vertical HSB would ideally compensate mas-

1994

Koenigswald— Enamel Structure of Moropus

ticatory stresses in the enamel of hypsodont teeth near the EDJ. Rensberger (1 992),
on the other hand, argues that the maximum tensile stresses in the flat lophs of
rhinos are normal to the cutting edges, causing fracturing parallel to the edges.
Orientation of the HSB perpendicular to these edges resists such fracturing. The
material studied here makes it possible for us to evaluate these hypotheses.

Vertical HSB were achieved in Rhinocerotidae and Astrapotheriidae in low-
crowned molars. Since this character antedates hypsodonty in both lineages, it
cannot be an adaptation specifically related to hypsodonty. Vertical HSB, which
are found to reduce abrasion in the cutting edges, are not unique to Perissodactyla.
A similar orientation of HSB as in Tapiroidea has been found in the camassials
of some Carnivora, in which the horizontal HSB change direction abruptly when
they approach the cutting edge to intersect the blades of the camassial notch
vertically.

There is a general demand to stabilize the cutting edges, especially in low-
crowned teeth, to keep the functional apparatus in shape. Wear of the oblique
cutting edges in low-crowned teeth, in Chalicotheriidae and in Brontotheriidae
but also in Carnivora, changes the morphology of teeth significantly and demands
a readjustment of the antagonist. The specific quality of high-crowned teeth is
that, due to the almost vertical cutting edges, the morphology of the occlusal
surface is maintained during progressive wear for a relatively long time. If wear
of cutting edges is reduced by prisms penetrating at high angles (as in vertical
HSB), a strong selective pressure applies to HSB reorientation from horizontal to
U-shaped or vertical.

However, this explanation does not contradict the hypothesis of Pfretzschner
(in press). The animals with vertical HSB in the entire molars, such as the early,
low-crowned Rhinocerotidae, have an appropriate prerequisite to build hypsodont
molars. This prerequisite is present in the Rhinocerotidae and the Astrapotherii-
dae. According to Pfretzschner’s hypothesis, most other herbivores, like horses
or bovids, that evolved hypsodont molars but retained horizontal HSB introduced
the “modified radial enamel” close to the EDJ. In the modified radial enamel,
thick layers of IPM lie between rows of prisms. These layers therefore were called
“inter row sheets” by Boyde (1964). It is mechanically important that the crys-
tallites of the IPM are at a high angle to the prisms. The thick descending layers
of IPM in the modified radial enamel and the rows of ascending prisms function
very similarly to vertical HSB. The orientation of structural elements is very
important since the enamel is very anisotropic in its physical properties.

Among small mammals, some rodent lineages modify the enamel of their in-
cisors from primarily horizontal uniserial HSB to vertical HSB. Several different
pathways to achieve this character can be distinguished in the Myoxidae, Dip-
didae, and Spalacidae (Koenigswald, 1993). The functional significance of these
modifications in the very thin rodent enamel is not yet fully understood. The
uniserial HSB with an IPM at right angles to the bands seems to be functionally
more isotropic to crack propagation than the thick HSB of the larger mammals.

Modification of the primarily horizontal HSB occurs only occasionally among
large mammals. The occurrence in Perissodactyla is unusually frequent. Obvi-
ously, with exception of Equoidea, most other groups in the Perissodactyla change
HSB orientation instead of introducing a large angle between the IPM and the
prisms. They share this character with astrapotheres. Therefore, the modification
of HSB orientation seems to be linked to IPM parallel to the prisms. In contrast
to the modified radial enamel, which is linked to the ability to reorient the crys-

Annals of Carnegie Museum

vol. 63

tallites of the IPM at an angle to the prisms, the lateral reorientation of the HSB
demonstrates second pathway to strengthen the enamel in cases when IPM crys-
tallite orientation cannot be changed.

This study of HSB orientation in Perissodactyla demonstrates that the quality
of enamel is a source of functional as well as phylogenetic information. Neither
aspect excludes the other since the evolution of characteristics always has to have
some functional significance if the new combination of characteristics is to be
advantageous for the animal. It is especially true in enamel that many character-
istics evolved in parallel in various lineages that indicate such functional aspects.
On the other hand, different groups often solve similar functional problems in
different pathways. This allows us to identify taxonomic groups of higher rank,
e.g., at the genus or family level.

Acknowledgments

I am very indebted to M. R. Dawson, Carnegie Museum of Natural History, for providing the
material of Moropus and for her hospitality during my stay in Pittsburgh. Further material was kindly
provided by R. M. Hunt, Jr., Nebraska State Museum, Lincoln, and F. Schrenk, Darmstadt. For
intensive discussions, I thank M. Sander and H. U. Pfretzschner, Bonn. The latter gave permission
to refer to his unpublished paper. Furthermore, two anonymous reviewers contributed to the clarity
of the discussion. I thank W. Haas who kindly helped photograph the HSB in some critical specimens.
D. Kranz and G. Oleschinski provided the photos and the artwork. Finally, I am very grateful to M.
Sander, C. Gee, and Th. Hansen for bringing the English into readable form.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 1, P. 67-75

22 February 1994

FIVE NEW SPECIES OF HYDROPTILA FROM EASTERN
UNITED STATES (INSECTA: TRICHOPTERA: HYDROPTILIDAE)

Jan L. Sykora

Research Associate, Section of Invertebrate Zoology
Steven C. Harris1

Research Associate, Section of Invertebrate Zoology

Abstract

Five new species of Hydroptila from the eastern United States similar to H. strepha Ross are described
and figured: Hydroptila antennopedia and H. parachelops from Pennsylvania, New Hampshire, and
Maine; H. morsei from South Carolina, Texas, and Florida; H. blicklei from Maine; H. holzenthali
from Mississippi. Male genitalia of H. strepha, determined to occur only in Pennsylvania and West
Virginia, are illustrated.

Introduction

The United States Army Corps of Engineers (Pittsburgh District) has supported
our ongoing study of caddisflies in reservoir tailwaters of the upper Ohio River
drainage basin. As part of this project, ultraviolet light traps were operated month-
ly from May through October at the outflow from Youghiogheny River Lake in
southwestern Pennsylvania (J. L. Sykora, unpubl. reports to U.S. Army Corps of
Engineers). Of the more than 100 species collected, two were tentatively identified
as Hydroptila strepha Ross. Comparisons of the genitalia of the specimens col-
lected from the Youghiogheny River to the figures of H. strepha suggested dif-
ferences, prompting an examination of the holotype of H. strepha collected from
the Susquehanna River in Pennsylvania. Close examination indicated that the
specimens from the Youghiogheny River represented two new species closely
related to H. strepha and other members of the H. consimilis group of Marshall
(1979).

The discovery of two new species similar to H. strepha suggested that reex-
amination of the species was in order. The question of the identity of H. strepha
was reinforced by the unusual distribution pattern for the species, with records
from the northeastern United States, Minnesota, Mississippi, Texas, and South
Carolina. Therefore, an effort was made to secure additional H. strepha material.
Collections of specimens identified as H. strepha and related species were obtained
from the University of New Hampshire, Clemson University, Illinois Natural
History Survey, National Museum of Natural History, University of North Texas,
University of Tennessee, and University of Minnesota. Specimens from these
collections were compared with the holotype of H. strepha resulting in the iden-
tification of three additional new species: one from Mississippi; one from Maine;
and another from South Carolina, Texas, and Florida. The material available to
us indicates H. strepha is known only from Pennsylvania and West Virginia.

The genus Hydroptila is the largest and most successful of the family Hydrop-

' Department of Biology, Clarion University, Clarion, Pennsylvania 16214.

Submitted 9 April 1993.

Annals of Carnegie Museum

vol. 63

Fig. 1-5 .—Hydroptila antennopedia, new species: 1, head of paratype; 2, male genitalia, lateral view;
3, tenth abdominal segment, dorsal view; 4, male genitalia, ventral view; 5, phallus, dorsal view.

tilidae with world- wide, cosmopolitan distribution (excluding polar regions). Mar-
shall (1979) estimated that this genus included over 150 species worldwide. Since
that time, many more species have been described from different parts of the
world. Up to 1979, 60 species were recorded by Blickle (1979) in the continental
United States, a number that has been constantly growing. Of these new species
many were discovered in the southern United States. At present, 95 Hydroptila
species are known from the continental United States, 72 of which are described
from the eastern United States and, according to Masteller and Flint (1992), 27
species of Hydroptila are known from Pennsylvania.

Marshall (1979) recognized several species-groups which are characterized by
male genitalia. Species described in this paper are members of the widely dis-
tributed consimilis group. Terminology utilized in the descriptions follows that
of Marshall (1979).

Specimens are deposited at the Carnegie Museum of Natural History (CMNH),
the Department of Entomology, Clemson University (CU), the Department of
Entomology, University of New Hampshire (NH), the Illinois Natural History
Survey (INKS), the University of Minnesota (UM), the National Museum of
Natural History (NMNH), or in the junior author’s collection (SCH).

Hydroptila antennopedia, new species
(Fig. 1-5)

Diagnosis. — This species is distinguished from any other known species of
Hydroptila by a combination of features in the male genitalia. Segment X is
trilobed; the inferior appendages expand gradually toward a rounded apex with
a dorsoapical point; and the apical section of the phallus is divided posteriorly
into two sections of equal length.

Description.— Male: Length 2.1 mm. Antennae 34-segmented. Color brown in alcohol. Stemite of
abdominal segment VII with short, pointed process; segment VIII annular, quadrate from lateral view;
segment IX with blunt, broad dorsal lobe, and a narrow mediolateral projection. Dorsal aspect of
segment X broad, three-lobed with central lobe rounded at apex, lateral aspect almost rectangular
with oblique apical margin. Subgenital plate shorter than inferior appendages, rounded at apex and

1994

Sykora and Harris— Five New Hydroptila

bearing two short spines close to the apical margin. Inferior appendages in lateral view with slightly
sinuate ventral and dorsal margins, expanding into a ventrally rounded apex with a distinct dorsoapical
point; ventral aspect with a quadrangular broad base and narrow, triangular apical section. Phallus
relatively short, distal section divided into two narrow, long processes, the dorsal one produced from
a broad cup-shaped base, the ventral process a rod-shaped continuation of the ejaculatory duct; short
paramere arising anteriorly of neck making full revolution.

Female. — Unknown.

Type Specimens. — Holotype, male (CMNH): PENNSYLVANIA, Fayette Co.,
Youghiogheny River Lake outflow near Confluence, June 17-18, 1991, light trap.
Paratypes: The specimens made available to us and identified as H. antennopedia
included many examples from Pennsylvania, New Hampshire, Maine, and one
specimen from Minnesota. PENNSYLVANIA, Fayette Co., Youghiogheny River
Lake outflow near Confluence 1 male (SCH); same August 24-25, 1991, 1 male
(CMNH); Forest Co., Otter Run, July 1, 1990, E. C. Masteller, 5 males; Warren
Co., 2 mi S Warren, 25 June 1987, O. S. Hint, Jr., 2 males (NMNH). MAINE,
Allagash, July 5, 1959, 5 males; same July 6, 1959, 4 males; same July 8, 1959,
2 males; same July 10, 1959, 40 males; same July 13, 1959, 8 males; same July
19, 1959, 39 males; same July 22, 1959, 17 males; same July 23, 1959, 9 males;
same July 24, 1959, 4 males; same July 25, 1959, 21 males; same July 26, 1959,
69 males; same July 27, 1959, 22 males; same July 29, 1959, 90 males; same July
30, 189 males; same August 1, 1959, 186 males; same August 2, 1959, 1 male.
Dennistown, July 22, 1959, 1 male; same August 27, 1959, 1 male; Kingfield,
August 13, 1964, 40 males; Round Pond, July 13, 1959, 1 male (NH); Allagash,
July 13, 1959, 1 male; same July 29, 1959, 1 male; same August 1, 1959, 10
males; same August 2, 1959, 1 1 males; Big Black River, July 25, 1961, A. Brower,
120 males; Oxbow, July 22, 1961, A. Brower, 1 male (NMNH). MINNESOTA,
Finland Co., July 25, 1965, D. Etnier, 1 male (UM). NEW HAMPSHIRE, Cole-
brooke, July 17-24, 1957, 2 males; same July 10-17, 1957, 1 male (NH).

Remarks.— One paratype of H. antennopedia bears a pair of thin, leg-like ap-
pendages extending from beneath the “scent caps” (Fig. 1). According to Schmid
(1980) who illustrated similar appendages in H. ampoda, these erectile organs
may carry androconia in the setae or scales with odoriferous function. This new
species is placed near H. strepha Ross (1941) based on the trilobed segment X
and the short, pointed apical sections of the phallus. It differs in the shape of
inferior appendages which are tubular in H. strepha, but club-shaped in H. an-
tennopedia. All the specimens designated as paratypes from Maine were previously
identified as H. strepha (Blickle, 1964). Morse and Blickle (1957) also reported
H. strepha from New Hampshire. Based on our reexamination, it is quite possible
that H. strepha does not occur in New Hampshire and Maine. The material
available to us indicates that H. strepha is known from the original locality in
north-central Pennsylvania (Susquehanna River) and West Virginia (Pendleton
County, Smoke Hole State Park, Briggs Run, April 9, 1977, Don and Mignon
Davis, Det. Hint; Pendleton County, Smoke Hole Camp, May 14, 1963, Field
and Hint, 1 male). As considerable confusion exists in the identity of H. strepha,
the species is redrawn from the type (Fig. 6-9).

Etymology. — Latin, meaning leg-like antennae.

Hydroptila parachelops, new species
(Fig. 10-13)

Diagnosis. — This species is distinguished by a combination of several morpho-
logical characters of male genitalia. These include the trilobed segment X with

Annals of Carnegie Museum

vol. 63

Fig. 6-9.— Male genitalia of Hydroptila strepha Ross: 6, lateral view; 7, tenth abdominal segment,
dorsal view; 8, phallus, dorsal view; 9, inferior appendages, ventral view.

very small central lobe and large lateral lobes, ventrally curved, tubular inferior
appendages and long, biramose apical section of the phallus.

Description.— Male: Length 3.0 mm. Antenna 29-segmented. Brown in alcohol. Stemite of segment
VII with short, pointed apicomesal process; segment VIII tubular with straight apical margins and
covered with sparse setae; segment IX emarginate ventrally and dorsally with relatively short ventro-
lateral extensions; segment X with dorsal aspect wide at the base and slightly narrowed at the excised,
flared apex; central lobe small and membranous, lateral sections slightly sclerotized, wide in the middle,
tapering posteriorly. Inferior appendages from lateral view almost tube-shaped, long and narrow,
curved ventrally, with slightly enlarged distal section, and rounded apex; ventral view triangular with
broad base and straight mesal margins, the distal section rapidly tapering towards obtuse apex with
small, sclerotized lateral point. Subgenital plate round with sclerotized ventral “strip” bearing two
setae and a membranous dorsal section. Phallus with relatively short base, forked distal section; dorsal
portion heavily sclerotized and curved with apex pointed ventrally; ventral branch narrow, parallel-
sided, lightly sclerotized and bent at the base; neck with paramere making at least one full revolution,
ejaculatory duct sinuous in neck, continuing into ventral branch of the phallus.

Female. — Unknown.

Type Specimens. — Holotype, male (CMNH): PENNSYLVANIA, Fayette Co.,
Youghiogheny River Lake outflow near Confluence, Pennsylvania, August 24-
25, 1991, light trap. Paratypes: same, 1 male (SCH); MAINE, Dennistown, July
31, 1959, 5 males (INHS); same, July 19, 1959, 2 males; same July 25, 1959, 13
males; Oquossoc, July 28, 1959, 1 male; same July 30, 1959, 2 males; same July
31, 1959, 1 male (NH).

Remarks.— Hydroptila parachelops is closely related to H. strepha Ross (1941)
and H. chelops Harris (1985), on the basis of trilobed segment X, tube-shaped
inferior appendages, and forked distal section of phallus with curved apex. It
differs from the latter in the shape of forked phallus which has only one sclerotized
apical branch. From the former it could be distinguished by the triangular, ventral
aspect of inferior appendages with straight mesal margins similar to H. arctia

1994

Sykora and Harris— Five New Hydroptila

Fig. 10-13.— Male genitalia of Hydroptila parachelops, new species: 10, lateral view; 11, tenth ab-
dominal segment, dorsal view; 12, ventral view; 13, phallus, dorsal view.

Ross (1938). Those specimens of H. parachelops collected in Maine and obtained
from the Illinois Natural History Survey and the University of New Hampshire
collections were previously identified as H. strepha.

Etymology.— Latin, close to chelops.

Hydroptila morsei , new species
(Fig. 14-17)

Diagnosis. — This species is characterized by male genitalia with segment IX
extended anteriorly into a ligament attached to segment VIII, trilobed segment
X bearing sickle-shaped lateral lobes, rod-shaped inferior appendages and mem-
branous apex of phallus with sinuous tip.

Description. — Male: Length 2.1 mm. Antennae 28-segmented. Color light brown in alcohol. Venter
of abdominal segment VII with short apicomesal projection; segment VIII tubular with rounded
posteroventral edges. Segment IX recessed within segment VIII, in lateral aspect rounded anteriorly
with antero ventral margin extended into sclerotized ligament-like section attached to the posteroventral
margin of segment VIII. Dorsal aspect of segment X trilobed with broad central lobe and sclerotized
lateral lobes flared and sickle-shaped; in lateral view broadly truncate with apex of the lateral lobes
upturned. Subgenital plate membranous and extending over the midlength of inferior appendages,
rounded apically and bearing two short setae. Inferior appendages in lateral aspect almost as long as
segment X, rod shaped, in ventral view each with broad triangular base with short lateral spines.
Phallus with proximal section broad and slightly longer than the distal portion; distal section mem-
branous with a slightly curved tip and bulbous base encircled by a short spiral paramere making one
full revolution.

Female. — Unknown.

Type Specimens. — Holotype, male (CU): SOUTH CAROLINA, Dorchester
Co., Four Holes Swamp, Goodsons Lake, 13 August 1976, J. Morse. Paratypes:
same, 4 males (3 males in CU, 1 male in SCH); Berkeley Co., Four Holes Swamp,
Mims Lake, 7 May 1976, 1. Morse, 2 males (1 male in CU, 1 male in CMNH);
TEXAS, Hardin Co., Cypress Creek, off TX 326, N. Kountze, UV light, October
23, 1992, Moulton and Alexander, 6 males; Hardin Co., Hickory Creek, off US

Annals of Carnegie Museum

vol. 63

Fig. 14-17. —Male genitalia of Hydroptila morsei, new species: 14, lateral view; 15, dorsal view; 16,
ventral view; 17, phallus, dorsal view.

287/69, N. Kountze, October 23, 1992, UV light, Moulton and Alexander, 1 male
(SCH); FLORIDA, Highlands Co., Archbold Biological Station, March 6, 1964,
S. W. Frost, 2 males (NMNH).

Remarks.— Hydroptila morsei is closely related to H. strepha Ross (1941) but
it is distinguished by the shape of segment X and the phallus. In H. morsei, the
distal portion of the tenth segment is characterized by well-developed lateral lobes
separated from a large central section by deep incisions while in H. strepha these
incisions are shallow and the central section is small. In H. strepha the distal
portion of the phallus has a broad, sclerotized base extending into the lateral,
chitinized margin with a membranous lateral part. In H. morsei the section sur-
rounding the ejaculatory duct is entirely membranous. This species was collected
from five sites located in South Carolina, Texas, and Florida. It was not recorded
by Harris et al. (1991) from Alabama and its distribution in the southern United
States remains unclear.

Etymology. — Named in honor of Dr. John Morse who collected this species.

Hydroptila blicklei, new species
(Fig. 18-21)

Diagnosis. — The male of this species is characterized by the combination of
several characters. The inferior appendages are wide at the base extending distally
to pointed, divergent apices; segment X is trilobed with rounded central lobe
slightly protruding dorsad and a pair of pointed, divergent lateral lobes; ventral
aspect of subgenital plate with “X”-shaped sclerotized central section; phallus
membranous with a short spiral paramere.

Description.— Male: Length 2. 1 mm. Antennae broken off. Color yellowish-brown in alcohol. Stemite
of abdominal segment VII with short, pointed apicomesal process; segment VIII generally quadrate
with slightly excised posteroventral margin and rounded ventrolateral aspect; segment IX retracted
into VIII, triangular from lateral view with spoon-shaped ventral extensions and incised posteroventral
margin; apex of segment X divided into three lobes, a membranous apically rounded mesal lobe and
a pair of sharp sickle-shaped lateral lobes; subgenital plate from lateral view partially sclerotized,

1994

Sykora and Harris —Five New Hydroptila 73

Fig. 18-21.— Male genitalia of Hydroptila blicklei, new species: 18, lateral view; 19, tenth abdominal
segment, dorsal view; 20, ventral view; 2 1 , phallus, dorsal view.

almost as long as segment X and extending over midlength of inferior appendages, ventrally with
“X”-shaped, sclerotized section, bearing a pair of setae centrally; inferior appendages longer than
segment X, in lateral view elongated, narrowly triangular and gradually tapering toward pointed tips,
in ventral aspect with a broad base extending into blade-shaped apices diverging distally; phallus
curved ventrally, central portion narrow tapering to a long neck, distal section membranous almost
triangular from dorsal view with slightly curved apex, ejaculatory duct sinuous in the neck and
continuing through the apical, membranous section with a slender, spiral paramere arising anteriorly
of neck making slightly more than one full revolution.

Female . — Unknown.

Type Specimens. — Holotype, male (NH): MAINE, Dennistown, July 29, 1959.
Paratypes: MAINE, Oquossoc, July 31,1959,1 male (NH); Oxbow, July 22, 1961,
A. Brower, 10 males (NMNH).

Remarks. — This species is another member of the H. consimilis group closely
related to H. strepha Ross (1941) and H. roberta Hamilton and Holzenthal (1986).
It differs from the former by the lateral aspect of inferior appendages which are
straight in H. blicklei and curved in H. strepha. From the latter it could be
distinguished by the phallus with spiral paramere, trilobed tenth segment with
membranous mesal section and by segment IX with spoon- shaped antero ventral
section. These specimens collected by Dr. A. E. Brower were made available to
us by Dr. J. S. Weaver, Jr. Ill and Dr. O. S. Flint, Jr. They were originally identified
as H. strepha (Blickle, 1964).

Etymology. --Named for Dr. R. L. Blickle who first studied this species.

Hydroptila holzenthali, new species
(Fig. 22-25)

Diagnosis. — The male of this species is characterized by the combination of
several characters, including bilobed segment X, triangular subgenital plate with
two apical lobes bearing terminal setae, wedge-shaped inferior appendages, and
the very long, thin and tubular phallus.

Annals of Carnegie Museum

vol. 63

Fig. 22-25. —Male genitalia of Hydroptila holzenthali, new species: 22, lateral view; 23, tenth abdom-
inal segment, dorsal view; 24, phallus, dorsal view (40% reduction); 25, ventral view.

Description. — Male: Length 3 mm, antennae broken off, color light brown in alcohol. Postero ventral
margin of segment VII with short apicomesal process. Segment VIII tubular, posteroventral margin
irregular with rounded posteroventral comers. Segment IX heavily sclerotized, in lateral view with
rounded anteroventral section extending deeply into segment VIII; dorsal and ventral aspects widely
incised. Segment X in dorsal view bilobed, lobes flared, in lateral view rectangular and broad with
rounded apex. Subgenital plate membranous, triangular, incised in the middle, almost as long as
inferior appendages, apex divided into two short lobes bearing two setae. Inferior appendages almost
as long as segment X, in lateral view wedge-shaped, with posteroventral comers rounded, dorsal
margin ending in small posterodorsal point; in ventral view widely separated at broad, shouldered
quadrangular bases with a long and prominent seta located at rounded posterolateral comer. Tubular
phallus very long, extending over more than three segments, with funnel-shaped base constricted at
midlength without spiral paramere and with one third of ejaculatory duct protruding freely from
membranous sheath.

Female. — Unknown.

Type Specimen. — Holotype, male (CU): MISSISSIPPI, Stone Co., Flint Creek,
Hwy 26, 7.9 km E Wiggins, 7 June 1979, Col. R. W. Holzenthal.

Remarks. — The holotype of this species was collected in Mississippi and orig-
inally identified as H. strepha (Harris et al., 1982). Hydroptila holzenthali is closely
related to H. quinola Ross (1947). It is distinguished by the absence of defined
thumb-like projections on the base of inferior appendages, and a membranous
and quadrangular subgenital plate. In addition, the apices of inferior appendages
in H. holzenthali are slightly hooked while in H. quinola the apices are rounded.

Etymology. — Named for Dr. Ralph W. Holzenthal, University of Minnesota who collected the
holotype.

Acknowledgments

This project was supported by contract DACW 599 1 M0705 from the U.S. Army Corps of Engineers,
Pittsburgh District, Pittsburgh, Pennsylvania. Special thanks go to J. M. Fowles and M. Koryak from
the U.S. Army Corps of Engineers for their assistance in the selection of sites, logistics of sample
collection, and critical review of the manuscript. We also acknowledge the help of O. S. Flint, Jr.,
National Museum of Natural History, J. Morse, Clemson University, and J. S. Weaver, Jr. Ill, who

1994

Sykora and Harris— Five New Hydroptila

provided us with additional specimens of Hydroptila. The type of H. strepha was loaned for use in

this study by the Illinois Natural History Survey.

Literature Cited

Buckle, R. L. 1964. Hydroptilidae (Trichoptera) of Maine. Entomological News, 75(6): 159-1 62.

. 1979. Hydroptilidae (Trichoptera) of America north of Mexico. New Hampshire Agriculture

Experiment Station, University of New Hampshire, Durham, New Hampshire, Station Bulletin
506, 97 pp.

Hamilton, S. W., and R. W. Holzenthal. 1986. Two new species of caddisflies from Georgia
(Trichoptera: Polycentropodidae, Hydroptilidae). Proceedings of the Entomological Society of
Washington, 88(1): 163-1 66.

Harris, S. C. 1985. New Hydroptilidae (Trichoptera) from Alabama. Journal of the Kansas Ento-
mological Society, 58(2):248-253.

Harris, S. C, P. G. Lago, and R. W. Holzenthal. 1982. An annotated checklist of the caddisflies
(Trichoptera) of Mississippi and southeastern Louisiana. Part II: Rhyacophiloidea. Proceedings
of the Entomological Society of Washington, 84(3): 509 -512.

Harris, S. C., P. E. O’Neil, and P. K. Lago. 1991. Caddisflies of Alabama. Bulletin of the Geological
Survey of Alabama, 142:1-442.

Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the
British Museum of Natural History (Entomology), 39(3): 135-239.

Masteller, E. C, and O. S. Flint, Jr. 1992. Trichoptera Biodiversity of Pennsylvania. 2nd ed.
[Report to Pennsylvania Fish Commission.] 2 volumes, 67 pp.

Morse, W. J., and R. L. Blickle. 1957. Additions and corrections to the New Hampshire list of
Trichoptera. Entomological News, 68:127-131.

Ross, H. H. 1938. Description of Nearctic caddis flies (Trichoptera). Bulletin of the Illinois Natural
History Survey, 21(4):101-183.

. 1941. Description and records of North American Trichoptera. Transactions of the American

Entomological Society, 67:35-126.

— -. 1947. Descriptions and records of North American Trichoptera with synoptic notes. Trans-

actions of the American Entomological Society, 73:125-168.

Schmid, F. 1 980. Genera des Trichopteres du Canada et des Etats adjacent. Les Insectes et Arachnides
du Canada, Partie 7. Agriculture Canada, Ottawa, 296 pp.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 1, P. 77-99

22 February 1994

DESCRIPTION OF THE IMMATURE STAGES OF SIX SPECIES OF
SPHAENOGNATHUS, WITH COMPARATIVE NOTES ON
PHYLOGENY AND NATURAL HISTORY
(INSECTA: COLEOPTERA: LUCANIDAE)

Giovanni Onore1

Research Associate, Section of Invertebrate Zoology

Abstract

Third instar larvae of six Andean species of Sphaenognathus and pupae of Sphaenognathus lindenii
are described and illustrated, with comparative notes on larvae of the related genus Chiasognathus.
Keys are provided to larvae of genera of Chiasognathini and to species of Sphaenognathus for which
larvae are known. A character considered diagnostic for Chiasognathini is to have all setae on the
raster uniformly inclined laterally. The absence of a basal tooth on the mandible in Sphaenognathus
peruvianus supports its placement in the subgenus Chiasognathinus. Derived larval characters are
presented to support three monophyletic taxa: the tribe Chiasognathini, the genus Sphaenognathus,
and a species-group containing S. lindenii, S. oberon, and S. metallifer.

Introduction

The family Lucanidae is of worldwide distribution and includes about a thou-
sand described species (Benesh, 1960). According to Blackwelder (1944), there
are 128 species in the Neotropical region. The larvae of several species of Lu-
canidae have been described. Significant publications on larvae of this group
include Hayes (1929), Van Emden (1935, 1941, 1952), Medvedev (1952), Ritcher
(1966) and Alderson (1975a, 1975Z?). In the Neotropical region, larvae of only
four species have been described: Sclerostomus ruficollis Luederwaldt, Pholidotus
spixi Nagel, Pycnosiphorus femoralis (Guerin), and Chiasognathus grand Stephens
(Nagel, 1934; Cekalovic, 1982; Cekalovic and Castro, 1983; Costa et al, 1988).
Most lucanid larvae live in decaying logs (Arnett, 1973; Borror et al., 1981), but
the larvae of at least one species live in the soil (Milne, 1933).

The larvae described in the present work are in the tribe Chiasognathini, rep-
resented by Chiasognathus and Sphaenognathus. At present, according to Benesh
(1990) and incorporating the synonymies made by Lacroix (1969), Chiasognathus
includes four species, all from Chile. Sphaenognathus includes about 40 Neo-
tropical species (Bartolozzi et al., 1992) and an additional species from Australia
described by Moore (1978), thus implying a Gondwanian distribution for the tribe
with ancient connections across Antarctica.

The only species of the tribe for which the larva has been described is Chia-
sognathus grand Stephens (Cekalovic and Castro, 1983). Notes on adults of Chia-
sognathus were published by Montalegre (1925), Ureta (1934), and Mathis (1981);
information on the adults of Sphaenognathus was provided by Howden and
Campbell (1974), Perrault (1991), Bartolozzi et al. (1992), and Bartolozzi and
Onore (1993). The phylogeny of Lucanidae based on adults was discussed by

1 Pontificia Universidad Catolica del Ecuador, Departamento de Biologia, 1 2 de Octubre y Carrion,
Apartado 2184, Quito, Ecuador.

Submitted 20 May 1993.

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vol. 63

Holloway (I960, 1968, 1969), Lacroix (1969), and Howden and Lawrence (1974).
Phylogenetic hypotheses based on larvae were discussed by Van Emden (1952)
and Lawrence (1981).

Materials and Methods

To collect larvae, soil was sampled in areas where adults had been found. The soil was dug to a
depth of 20 cm, and larvae, pupae, and associated adults were collected. Intact larvae were fixed by
boiling gently in water for 3-5 min, and then preserved in 80% ethanol. The pupae were maintained
alive in moist boxes until they became adults that were then killed and mounted for identification.

To study the larvae, the mandibles were removed using two strong needles, and then successively
the labium and maxillae were removed using scissors. Larval structures submerged in 80% ethanol
were examined at 50 x with a stereomicroscope. The determination of sex in larvae was made by the
presence (male) or absence (female) of Herold’s organs as described by Menees (1957). The shape of
the last two antennomeres and number and position of teeth of the stridulatory organs of the meso-
thoracic and metathoracic legs were used as characters to identify the larvae to genus. To distinguish
species the following characters were used: number of teeth on the left mandible, shape of epipharyngeal
pterotormae, shape of the prothoracic lateral sclerome, number of teeth on plectrum and pars stridens,
shape of the thoracic spiracle and setation of lateral anal lobes and raster. The terminology used follows
that of Edwards (1930), Boving (1936), Ritcher (1966) and Britton (1970).

I borrowed material from the following institutions or persons: CMNH, Carnegie Museum of Natural
History, Pittsburgh, Pennsylvania, USA; CV, Carlota Vergara Collection, Santiago, Chile; FSCA,
Florida State Collection of Arthropods, Gainesville, Florida, USA; IZAV, Instituto de Zoologia Agrico-
la, Universidad Central de Venezuela, Maracay, Venezuela; OSUO, Department of Entomology Col-
lection, Oregon State University, Corvallis, Oregon, USA; QCAZ, Museo del Departamento de Biologia
de la Pontificia Universidad Catolica del Ecuador, Quito, Ecuador; WSUC, James Entomological
Collection, Washington State University, Pullman, Washington, USA.

Natural History

In 1986, L. Coloma found a great number of adults of Sphaenognathus oberon
in Casha Totoras, Bolivar Province, Ecuador. With the intention of finding larvae
of this stag beetle in the vicinity, I searched carefully in rotten logs and old stumps.
No larvae were found in rotting wood, so I extended the search to the soil where
digging and sifting soon revealed several larvae of S. oberon. Investigating the
same habitat at other localities, I found the larvae of S. lindenii , S. metallifer, S.
nobilis, S. peruvianus (Fig. 1), and S. subtilis.

I never found eggs in the natural habitat, so in March, 1988, 1 dissected a dozen
freshly killed females. These contained immature eggs in various stages of de-
velopment. The number of eggs varied from 12-96. First instar larvae were found
in groups up to a maximum of 54 individuals. They were in the first 3-10 cm of
topsoil which was rich in humus and roots. Very often a dead female was found
buried in the soil close to the group of larvae. Species of plants typically found
growing in the topsoil near larvae were: Salpichroa sp. (Solanaceae), Gynoxys
buxifolia (Asteraceae), Orthrosanthus chimboracensis (Iridaceae), Equisetum bo-
gotense (Equisetaceae), Barnadesia spinosa (Asteraceae), Hypericum laricifolium
(Hypericaceae), Lachemilla orbiculata (Rosaceae), Ribes andicola (Saxifragaceae),
Hesperomeles pernettifolia (Rosaceae), Chusquea sp. (Poaceae), and Elaphoglos-
sum sp. (Lomariopsidaceae).

Second instar larvae were less frequently encountered in the soil than third
instar larvae. Third instar larvae were distributed uniformly, roughly equidistant
from one another. They were buried 1 0-20 cm deep, and there was a maximum
of 36 specimens per square meter. In some areas larvae were numerous, while in
other, apparently identical, sites no larvae were found. In uniform habitat, the
females may be arbitrary when choosing oviposition sites. Perhaps they settle

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Onore— Immature Stages of Sphaenognathus

when they encounter some obstruction to flight, such as small bushes, and then
lay their eggs.

Soil and fragments of roots were found in dissections of the gut, thus indicating
that larvae may eat decomposing organic materials mixed with soil and small
roots. When dug from the soil, larvae rolled into the characteristic C-shape of
scarabaeiform larvae and remained inactive. Specimens unearthed at dusk and
left on the surface were found in the same place the following morning. As the
sun came up, they desiccated and died. Unearthed larvae of S. lindenii have the
terga of abdominal segments 3-5 covered with moist soil, suggesting some special
property of the cuticle or some secretion in those specific areas. I observed the
same phenomenon in recently uncovered larvae of Sphaenognathus oberon and
S. metallifer.

When disturbed, the larvae of S. lindenii produced vibrations using a stridu-
latory apparatus. Simultaneously and rhythmically, the metathoracic trochanter
was rubbed against the mesothoracic coxa. No sound audible to human ears was
produced, but vibrations could be felt when the stridulating larvae was held in
hand. The stridulation may be used by larvae for protecting or marking the
territory where they feed, as suggested by the evenly spaced distribution of larvae
in the soil. Mature third instar larvae were 10-20 cm deep in suboval cavities
with internal dimensions about 35 by 70 mm. Sometimes these cavities contained
numerous white Collembola. I found dead larvae in the soil, covered by a white
fungus. Others were full of brown liquid and very soft. One contained a dipteran
puparium. Close to farms, domestic pigs actively dig the soil searching for the
white grubs, which they eat voraciously. Dogs also follow farmers when they are
preparing or plowing the soil, eating the larvae, pupae, and adults as they are
turned up.

Pupae were in cavities as described above, each with its larval exuvium retracted
to the caudal end. The pupae (Fig. 50-52) reposed on their dorsal surfaces and
rotated their abdomens when disturbed. Pupae were found in the soil from No-
vember to February and adults from December to March. First, second, and third
instar larvae, however, were found throughout the year, suggesting a life cycle of
more than a single year.

In my opinion the subterranean habit of the larvae may be an adaptation that
protects them from the rapid temperature changes in the upper layers of the soil
that occur at the high elevations where they live.

Keys and Diagnoses
Chiasognathini Felsche, 1889

Diagnosis. — All setae on the raster inclined laterally, never medially or erect
(Fig. 44-49).

Key to Larvae of Genera of Chiasognathini

1. Antenna with subapical segment swollen distally with an acute projecting
lobe, medial to which is a suboval sensory spot facing medially (Fig. 7);
apical segment pyriform. Mesocoxa with pars stridens consisting of a
single row of teeth expanded proximally into a field of teeth as wide as
several teeth, each tooth subequal in size to those in the main row (Fig.

35). Metatrochanter with plectrum consisting of row of fewer than 48
teeth (Fig. 40) Chiasognathus

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vol. 63

1'. Antenna with subapical segment distally rounded with sensory spot facing
laterally (Fig. 6); apical segment subconical. Mesocoxa with pars stridens
consisting of a single row of teeth, proximally with at most a patch of
very small sparse granules (Fig. 36-39). Plectrum of metatrochanter con-
sisting of row of more than 48 teeth (Fig. 41-43) Sphaenognathus

Genus Chiasognathus Stephens, 1831

I have seen larvae and associated adults of the following species: Chiasognathus
granti, 1 male, Chile, Cayutue, Lago Todos Los Santos, Llanguihue, 14 October
1969, J. Solervinces A.; Chiasognathus sp., 1 female, Chile, Maniuales, L. Pena;
C. granti, 14 larval skins, Chile, Lago Penihueico, Volcan Chosuenco, August
1988, O. Skew, ex boar stomach; C granti, 2 males, XI Region, Coyhaique [no
further data]. All specimens examined with mandibles worn; material deposited
in QCAZ, CMNH, and CV.

Genus Sphaenognathus Buquet, 1838

Larva C-shaped, subcylindrical and thickened posteriorly (Fig. 1); mediolateral
length, including head, 45-86 mm; color whitish blue when teneral and whitish
yellow when fully developed.

Head capsule globose (Fig. 1-4), 8.0-11.6 mm wide, dark to light reddish brown
with anterior pattern of clypeus yellowish brown; epicranial suture impressed,
yellowish, approximately as long as frontal suture (epicranial suture appears short-
er in Fig. 4 due to angle of view); surface of cranium bare to lightly punctate with
2-8 dorsoepicranial setae; stemma circular to ovate, not pigmented, without def-
inite margin, not or slightly raised above head surface; frontal suture distinct,
often asymmetrical; frons lightly punctate, becoming rugosopunctate anteriorly,
with an ovate depression at top of epicranial suture corresponding to ocellar area;
on each side, 1 or 2 setae at anterior angle, 1 or 2 exterior frontal setae, and 0 or
1 anterior frontal setae; clypeofrontal suture distinct, impressed. Clypeus trape-
zoidal, 3. 0-3. 7 times as long as wide, 1 or 2 anterior clypeal setae and 1 or 2
exterior clypeal setae on each side; preclypeus lightly sclerotized, rugosopunctate.
Labrum suboval, rugosopunctate, wider than long, with 2 anterior labral setae,
10-20 posterior labral setae. Apex of epipharynx (Fig. 8, 9, 11-14), in frontal
view, with 13-18 long, thick setae directed anteromedial] y; haptomerum with at
least two truncate coarse heli; chaetopariae consisting of 18-93 long setae and
numerous small setae directed medioapically; tormae united, laeotorma and dex-
iotorma almost symmetrical; pterotormae symmetrical, with apex rounded, acute
or broadly truncate; epitorma with rounded apex and deeply impressed in surface
of pedium; haptolachus with three suboval nesia, the medial nesium sometimes
hardly distinct from sclerotized sensory cone, the right nesium larger than left
nesium; a patch of small setae on each side of sensory area. Mandibles subtrian-
gular (Fig. 2, 15-23), asymmetrical, brownish; dorsa of mandibles (Fig. 16, 17,
19-23) with small dorsomolar setae on base of molar area, one large caudolateral
seta at scissorial notch and one large lateral seta on molar area; ventral surface
of mandibles (Fig. 15, 18) with patch of ventromolar setae and a patch of ven-
trolateral setae; basolateral angle with ventral subtriangular process; basolateral
angle with rounded postartis; scissorial area of left mandible (Fig. 15, 16) with
4-5 teeth (unknown in S. subtilis), inner margin of left mandible between scissorial
and molar areas with or without basal tooth (Fig. 15, 16, 19-23); teeth of molar
area (Fig. 15, 16) not well-defined, with masticatory concave area projecting

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Onore— Immature Stages of Sphaenognathus

Fig. 1-3. —Third instar larva of Sphaenognathus peruvianus. 1, left lateral view; 2, head, frontal view;
3, head and thorax, left lateral view.

anteriorly and retracted posteriorly; aria present (Fig. 1 6); scissorial area of right
mandible with teeth often not well-defined; molar area with masticatory area
convex. Maxillae (Fig. 25, 26) each with galea and lacinia separate; galea with
strong subcorneal uncus and 5-7 large setae lateral to uncus; lacinia with terminal
subconical uncus, 10-17 large setae behind uncus; stipes with large strip of small

Scutum

Scutellum

Prescutum

Pleural Area

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vol. 63

Dorsoepicranial Setae
Frontal Suture
Ocellar Area
Frons

Exterior Frontal Seta
Stemma
Seta of Anterior Angle
Clypeofrontal Suture
Preclypeus
Exterior Clypeal Seta
Anterior Ciypeal Seta
Postclypeus

Labrum

Posterior Labral Setae

Anterior Labral Seta

^ f — r

Sensory Spot

1mm

Fig. 4-1.— Third instar larvae of Chiasognathini. 4, Sphaenognathus nobilis, head; 5, S. subtilis, left
antenna, dorsal view; 6, S. oberon, third and fourth antennomeres of right antenna, lateral view; 7,
Chiasognathus grand, third and fourth antennomeres of right antenna, lateral view.

teeth set in a lightly sclerotized area; maxillary palpus with four palpomeres; basal
palpomere as long as wide, with subcircular unpigmented area; second palpomere
shorter than basal, subconical with subcircular unpigmented area; third palpomere
subcylindrical, as long as second palpomere; apical palpomere subconical, longer
than basal palpomere; palpomere 1-3 setose, apical palpomere glabrous. Labium
(Fig. 24): labial palpi with two palpomeres, apical palpomere subconical, twice
or more length of basal palpomere; hypopharyngeal sclerome heavily sclerotized,
concave medially, asymmetrical; sublateral lobes of sclerome reduced to two
unpigmented spots, the right spot smaller than left; sclerome with left side of
anterior margin with strong conical teeth directed dorsally. Antennae (Fig. 4-6)

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Onore- Immature Stages of Sphaenognathus

Fig. 8—10.— Epipharynges of third instar larvae of Chiasognathini. 8, Sphaenognathus nobilis\ 9, S.

peruvianus; 10, Chiasognathus granti.

geniculate, each with four antennomeres; basal antennomere subcylindrical, about
as long as apical; second antennomere clavate, longer than third; third antenno-
mere clavate with a subapical and externally directed sensory spot; fourth anten-
nomere subconical, externally directed and with a sensory spot; second to fourth
antennomeres setose.

Thorax (Fig. 3, 27) divided into pro thorax, mesothorax, and metathorax; pro-
thoracic dorsum with two narrow transverse rows of long setae, anterior row with
many setae, posterior row with few setae; each side of prothorax with sclerotized
furrow, continuous (Fig. 27) or interrupted below the middle (Fig. 3); mesothorax
divided into two dorsal areas, posterior one with row of long setae; metathorax
with dorsum similar to mesothorax; pleural area of mesothorax and metathorax

Annals of Carnegie Museum

vol, 63

Fig. 1 1-14.— Epipharynges of third instar larvae of Sphaenognathus. 11,5. suhtilis\ 12, S. metallifer ;
13, S. lindenir, 14, S. oberon.

each with a tubercle with a patch of long setae. Prothoracic legs (Fig. 31) slightly
shorter than mesothoracic and metathoracic legs (Fig. 33, 34); metathoracic legs
directed anterolaterally and perpendicular to those of mesothorax and prothorax.
Each leg with setose coxa, trochanter, femur, tibia, and tarsungulus (Fig. 31, 33,
34); tarsungulus abruptly attenuated at apex forming very short median tooth,
flanked by two setae rising from same position on each side (Fig. 32); coxae of
mesothoracic legs (Fig. 36-39) with a stridulatory area or pars stridens, consisting
of a slightly curved, longitudinal row of subconical teeth raised into slight carina,
lateral end of carina diffusing into small patch of granules, medial end with very
small patch or no patch of granules; trochanter of each metathoracic leg (Fig. 41-
43) with a stridulatory area or plectrum consisting of a single longitudinal row of

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Onore— Immature Stages of Sphaenogna thus

Fig. 15-23.— Mandibles of third instar larvae of Sphaenognathus. 15, S. nobilis, left mandible, ventral
view; 16, S. nobilis, left mandible, dorsal view; 17, S. nobilis, right mandible, dorsal view; 18, S.
nobilis, right mandible, dorsal view; 18, S. nobilis , right mandible, ventral view; 19, S. metallifer, left
mandible, dorsal view, 20, S. peruvianus, left mandible, dorsal view; 21, S. lindenii, left mandible,
dorsal view; 22, S. subtilis, left mandible, dorsal view; 23, S. oberon, left mandible, dorsal view.

short rounded, suboval or elongate teeth; teeth decreasing in size or becoming
elongate distally; trochanter swollen apically with a moderate projecting lobe.

Abdominal terga 1-8 becoming longer posteriorly (Fig. 1); tergum 9 about same
length as tergum 6; abdominal segment 1 with two dorsal areas, posterior area

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Fig. 24-26 . — Mouthparts of third instar larvae of Sphaenognathus. 24, S. subtilis, labium, dorsal view;
25, S. metallifer, right maxilla, dorsal view; 26, S. subtilis, right maxilla, dorsal view.

with row of long setae; segments 2-6 with prescutum, scutum, and scutellum;
scutum and scutellum divided by asperities; scutum covered by short thick setae;
scutellum covered by short thick setae and posteriorly with transverse row of long
setae; segments 7-9 with two dorsal areas, each with transverse row of long setae;
segment 10 with one dorsal area with transverse row of long setae; pleural area
of segments 1-9 each with prominent tubercle bearing patch of long setae; anal
opening (Fig. 44-48) Y-shaped, stem of Y much longer than arms; upper anal
lobe very small, insignificant; lateral lobe with kidney-shaped and glabrous pad;
raster (Fig. 48) consisting of mixed short to long stout setae directed laterally;
sternum 10 about two times as long as tergum 10; sternum 9 about half as long
as tergum 9; Herold’s organ on anal border of sternum 9 present in male (Fig.
49) and absent in female; sterna 1-8 bilobed, each with transverse row of long
sparse setae.

Spiracles (Fig. 28-30) yellowish brown with cribriform respiratory plate; spira-
cles present on nine segments; respiratory plate of prothoracic spiracle kidney-
shaped (Fig. 29) or C-shaped (Fig. 28) with concavity directed anteriorly with
subcircular bulla; respiratory plate of abdominal spiracles C-shaped (Fig. 30)
becoming smaller posteriorly; abdominal spiracle 5 subequal in size to respiratory
plate of prothoracic spiracle.

Key to Known Larvae of Sphaenognathus

F. Prothorax with lateral sclerotized furrow clearly interrupted by a non-
sclerotized area (Fig. 3). Raster with more than 900 short to long setae

(Fig. 1) Sphaenognathus peruvianus

F. Pro thorax with lateral sclerotized furrow continuous, not interrupted
by a nonsclerotized area (Fig. 27). Raster with fewer than 900 short to
long setae (Fig. 48) 2

2(F). Anal lobes glabrous (Fig. 46) Sphaenognathus subtilis

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Onore— Immature Stages of Sphaenognathus

Prothorax
Mesothorax
Metathorax

Lateral Furrow

Pleural Area
4,?^ Tu be rcle

Respiratory Plate

Fig. 27-34.— Third instar larvae of Sphaenognathus. 27, S. nobilis, thorax, left lateral view; 28, S.
subtilis, left thoracic spiracle; 29, S. nobilis, left thoracic spiracle; 30, S. nobilis, left spiracle, abdominal
segment 3; 31, S', peruvianas, right prothoracic leg; 32, S. peruvianas, tarsungulus of right prothoracic
leg; 33, S. metallifer, left mesothoracic leg; 34, S. peruvianas, right metathoracic leg.

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Fig. 35-39.— Pars stridens on coxa of left mesothoracic leg, third instar larvae of Chiasognathini. 35,
Chiasognathus granti\ 36, Sphaenognathus nobilis ; 37, S. metallifer ; 38, S. subtilis\ 39, S. peruvianus.

2'. Anal lobes densely tomentose (microsetae 0.02-0.12 mm long) (Fig.
44,45,47) 3

3(2'). Left mandible with five teeth in the scissorial area (Fig. 15, 16). Re-
spiratory plate of thoracic spiracle kidney-shaped (Fig. 29) .........

Sphaenognathus nobilis

3'. Left mandible with four teeth in the scissorial area (Fig. 19, 21, 23).
Respiratory plate of thoracic spiracle C-shaped (Fig. 28) ...........

Sphaenognathus lindenii, S. metallifer, and S. oberon

Descriptions

Sphaenognathus lindenii Murray, 1857
Larva

Mediolateral Length. — 68-7 6 mm.

Head. — Width of cranium 9.3-1 1.6 mm; cranium surface dark reddish brown,
yellowish spotted. Dorsoepicranial setae 2-5 on each side. Frons on each side
with 1 or 2 setae at anterior frontal angle, 1 or 2 exterior frontal setae, and 0-1
anterior frontal setae. Clypeus trapezoidal with length 3. 5-3. 7 times width, with
1 or 2 anterior clypeal setae and 1 or 2 exterior clypeal setae on each side. Labrum
with 10-15 posterior and two anterior labral setae. Epipharynx (Fig. 13): hap-
tomerum with 2-3 heli; pedium with 24-27 sensilla; right chaetoparia with 10-
1 5 long setae and some smaller setae; left chaetoparia with 1 0-24 long setae and
some smaller setae; pterotormae subparallel and rounded apically. Left mandible
with four blade-like scissorial teeth, basal tooth conical and dull (Fig. 21). Maxillae
each with 5-6 large setae lateral to uncus of galea and 13-16 large setae behind
uncus of lacinia.

Thorax. — Respiratory plate of thoracic spiracle C-shaped. Prothorax with scler-
otized lateral area forming a long furrow. Each mesothoracic leg with pars stridens

1994

Onore— Immature Stages of Sphaenognathus

Fig. 40=43.— Plectrum on trochanter of right metathoracic leg, third instar larvae of Chiasognathini.
40, Chiasognathus sp.; 41, Sphaenognathus metallifer ; 42, S. subtilis; 43, S. nobilis.

consisting of 92-108 teeth; metathoracic legs each with plectrum consisting of
83-105 suboval teeth, distal teeth becoming more elongate.

Abdomen. — Upper and lateral anal lobes covered by short thick setae 0.04-
0.12 mm long; anal pad bare (Fig. 45). Raster consisting of 366-386 short to long
setae.

Larval Material. — Description based on the following third instar larvae: five
males and 14 females with associated adults; deposited in QCAZ and CMNH
collections.

Locality. —Ecuador: Pichincha, La Cocha, 3000 m, February to March 1989,
G. Onore, 5-25 cm deep in sod soil.

Pupa

Male Pupa. —(Fig. 50-52). Shape elongate, exarate; length 43-54 mm; color
yellowish brown, becoming greenish brown just prior to adult eclosion.

Head. — Ceratotheca elongate, with few short scattered setae. Ophthalmothecae
distinctly visible with sclerotized lunate border anteriorly. Maxillary palpi distinct,
longer than mandibles.

Thorax. — Podothecae with three segments discernible; metatarsi reaching the
fifth abdominal segment. Pterothecae closely appressed and curving ventrally
around body. Mesonotum and metanotum each with a prominent bump medio-
posteriorly; pronotum subtrapezoidal, 1 .75 times as wide as long, with two prom-
inent bumps medioposteriorly; pronotum, mesonotum, and metanotum with few
scattered, short setae.

Abdomen.— Abdominal sterna with small scattered setae; apical segment with
genitalia distinctly visible between urogomphi; nine abdominal terga visible; terga
1-8 with a transverse row of small setae and at posterior border a transverse row

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Fig. 44-49.— Posterior abdominal structures of Chiasognathini. 44, Sphaenognathus nobilis, caudal
view of last abdominal segment; 45, S. lindenii, caudal view of last abdominal segment; 46, S. subtilis,
caudal view of last abdominal segment; 47, S. metallifer, caudal view of last abdominal segment; 48,
S. subtilis, sternum of last abdominal segment; 49, Chiasognathus grand, sterna 9-10 of male larva
showing Herold’s organ.

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Onore— Immature Stages of Sphaenognathus

Ophthalmotheca

Fig. 50-52.— Pupa of Sphaenognathus lindenii (very fine setae on abdominal terga not shown). 50,
dorsal view; 51, left lateral view; 52, ventral view.

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vol. 63

of long setae (these setae not depicted in Fig. 50, 51). Abdominal spiracles eight
on each side.

Female Pupa. —Length 52 mm. Similar to male except mandible shorter than
in male and external genitalia very reduced.

Pupal Material — Description based on the following pupae: four males and
one female with associated adults; deposited in QCAZ and CMNH collections.

Locality. — Ecuador: Pichincha, La Cocha, January 1988, G. Onore, 15-20 cm
deep; in pupal cells, in sod soil; between roots of Lachemilla orbiculata (Rosaceae).

Sphaenognathus metallifer
Bomans and Lacroix, 1972

Larva

Mediolateral Length.— 68-86 mm.

Head.— Width of cranium 8.8-10.1 mm; cranium surface dark reddish brown,
yellowish spotted. Dorsoepicranial setae 2-4 on each side. Frons on each side
with one seta at anterior frontal angle, one exterior frontal seta, and no anterior
frontal seta. Clypeus trapezoidal with length 3. 1-3.6 times width, with one anterior
clypeal seta and one exterior clypeal seta on each side. Labrum with 1 5-20 pos-
terior and two anterior labral setae. Epipharynx (Fig. 1 2): haptomerum with 2-5
heli; pedium with 28-50 sensilla; right chaetoparia with 17-26 long setae and
some smaller setae; left chaetoparia with 20-27 long setae and some smaller setae;
pterotormae subparallel and rounded apically. Left mandible with four blade-like
scissorial teeth, basal tooth conical and dull (Fig. 19). Maxillae each with 5-7
large setae lateral to uncus of galea and 15-17 large setae behind uncus of lacinia
(Fig. 25).

Thorax. — Respiratory plate of thoracic spiracle C-shaped. Pro thorax with scler-
otized lateral area forming a long furrow. Each mesothoracic leg with pars stridens
(Fig. 37) consisting of 83-88 teeth; metathoracic legs each with plectrum (Fig. 41)
consisting of 73-85 suboval teeth, distal teeth becoming more elongate.

Abdomen.— Upper and lateral anal lobes covered by short thick setae 0.04-
0. 12 mm long; anal pad bare (Fig. 47). Raster consisting of 462-464 short to long
setae.

Material. — Description based on the following third instar larvae: six males
and ten females with associated adults; deposited in QCAZ and CMNH collec-
tions.

Locality. — Ecuador: Canar, El Tambo, 2900 m, February 1990, G. Onore, 5-
20 cm deep in grassy soil.

Sphaenognathus nobilis Parry, 1874
Larva

Mediolateral Length.— 45-72 mm.

Head.— Width of cranium 8.0-9. 1 mm; cranium surface dark reddish brown,
yellowish spotted. Dorsoepicranial setae 2-3 on each side (Fig. 4). Frons on each
side with one seta at anterior frontal angle, one exterior frontal seta, and no anterior
frontal setae. Clypeus trapezoidal with length 3.10-3.14 times width, with one
anterior clypeal seta and one exterior clypeal seta on each side. Labrum with 8-
1 2 posterior labral seta and two anterior labral setae. Epipharynx (Fig. 8): hap-
tomerum with 2-4 heli; pedium with 23-3 1 sensilla; right chaetoparia with 1 5-
1 6 long and some smaller setae; left chaetoparia with 13-18 long and some smaller
setae; pterotormae rounded apically and internally directed. Left mandible with

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Onore— Immature Stages of Sphaenognathus

five blade-like scissorial teeth, basal tooth conical and doll (Fig. 15, 16). Right
mandible in dorsal view (Fig. 17) and in ventral view (Fig. 18) with teeth not
well-defined. Maxillae each with 4-6 large setae lateral to uncus of galea and 1 5=
1 7 large setae behind uncus of lacinia.

Thorax. — Respiratory plate of thoracic spiracle kidney-shaped (Fig. 27, 29).
Prothorax with sclerotized lateral area forming a long furrow (Fig. 27). Each
mesothoracic leg with pars stridens consisting of 83-86 teeth (Fig. 36); metatho-
racic legs each with plectrum consisting of 50-57 suboval teeth (Fig. 43).

Abdomen. — Upper and lateral anal lobes covered by short setae 0.02-0.07 mm
long; anal pad bare (Fig. 44). Raster with 586-730 short to long setae.

Material. — Description based on the following third instar larvae: five males
and 13 females with associated adults; deposited in IZAV, QCAZ, and CMNH
collections.

Locality. — Venezuela: Merida, La Mucuy, 2700 m, 18 Apr 1989, G. Onore and
C. Bordon. In topsoil 3-1 5 cm deep, between roots of cultivated Cupressus macro -
carpa (Cupressaceae) and native Podocarpus rospigliosi (Podocarpaceae).

Sphaenognathus oberon Kriesche, 1922
Larva

Mediolateral Length. — 62-86 mm.

Head. — Width of cranium 10.0-10.6 mm; cranium surface dark reddish brown,
yellowish spotted. Dorsoepicranial setae 2-4 on each side. Frons on each side
with 1 or 2 setae at anterior frontal angle, 1 or 2 exterior frontal setae, and no
anterior frontal setae. Clypeus trapezoidal with length 3. 6-3, 7 times width, with
one anterior clypeal seta and one exterior clypeal seta on each side. Labrum with
10-15 posterior labral setae and two anterior labral setae. Epipharynx (Fig. 14):
haptomerum with 2-4 heli; pedium with 28-64 sensilla; right chaetoparia with
16-25 long setae and some smaller setae; left chaetoparia with 23-30 long setae
and some smaller setae; pterotormae subparallel and rounded apically. Left man-
dible with four scissorial teeth blade-like, basal tooth conical and dull (Fig. 23).
Maxillae each with 5-7 large setae lateral to uncus of galea and 10-17 large setae
behind uncus of lacinia. Apical antennomere subconical (Fig. 6).

Thorax. — Respiratory plate of thoracic spiracle C-shaped. Pro thorax with scler-
otized lateral area forming a long furrow. Each mesothoracic leg with pars stridens
consisting of 85-87 teeth; metathoracic legs each with plectrum consisting of 60-
83 suboval teeth, distal teeth becoming more elongate.

Abdomen. —Upper and lateral anal lobes covered by short thick setae 0.5-0. 12
mm long; anal pad bare. Raster with 412-508 small to long setae.

Material. — Description based on the following third instar larvae: seven males,
three females, and three larvae of undetermined sex with damaged genital regions,
all with associated adults; deposited in QCAZ and CMNH collections.

Locality. — Ecuador: Bolivar, Santiago, Casha Totoras, 2900 m, March 1989,
G. Onore, 10-25 cm deep in the soil covered by cultivated Solanum tuberosum
(Solanaceae).

Sphaenognathus peruvianus (Waterhouse, 1869)

Larva

Mediolateral Length. — 58-76 mm (Fig. 1).

Head. —Width of cranium 8.0-10.0 mm; cranium surface reddish brown, yel-
lowish spotted (Fig. 2, 3), 3-8 dorsoepicranial setae on each side. Frons on each

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side with one or two setae at anterior frontal angle, one or two exterior frontal
setae, and 0-1 anterior frontal setae. Clypeus trapezoidal with length 3.4 times
width, with one anterior clypeal seta and one or two exterior clypeal setae on each
side. Labrum with 19 posterior labral setae and two anterior labral setae. Epi-
pharynx (Fig. 9): haptomerum with two heli; pedium with 21-29 sensilla; right
chaetoparia with 30-41 long and some smaller setae; left chaetoparia with 30-52
long and some smaller setae; pterotormae subtriangular and slightly internally
directed. Left mandible with four blade-like scissorial teeth and without basal
tooth (Fig. 2, 20). Maxillae each with five large setae lateral to uncus of galea and
15-16 large setae behind uncus of lacinia.

Thorax. — Respiratory plate of thoracic spiracle C-shaped (Fig. 3). Prothorax
with sclerotized lateral area consisting of furrow interrupted by a nonsclerotized
area, the part of furrow ventral to interruption shorter than dorsal (Fig. 3). Pro-
thoracic leg (Fig. 31). Each mesothoracic leg with pars stridens (Fig. 39) consisting
of 79-88 teeth; metathoracic legs each with plectrum (Fig. 34) consisting of 66-
82 conical teeth, distal teeth becoming more elongate. Tarsungulus of pro thoracic,
mesothoracic, and metathoracic legs (Fig. 32).

Abdomen . — Upper and lateral anal lobes covered by very sparse, scattered short
setae 0.02-0.06 mm long; anal pad bare. Raster with 940-990 short to long setae.

Material — Description based on the following third instar larvae: seven males
and seven females with associated adults; deposited in QCAZ and CMNH col-
lections.

Locality. — Ecuador: Loja, 4°16'S, 79°20'W, 2800 m, 24 March 1990, G. Onore,
5-15 cm in topsoil.

Sphaenognathus subtilis Lacroix, 1987
Larva

Mediolateral Length. — 8 1 mm.

Head. — Width of cranium 10.8 mm; cranium surface dark reddish brown. One
dorsoepicranial seta on each side. Frons on each side with one seta at anterior
frontal angle, one exterior frontal seta, and no anterior frontal seta. Clypeus
trapezoidal with length 3.2 times width, with one anterior clypeal seta on each
side; one exterior clypeal seta on the left side and two setae on the right side.
Epipharynx (Fig. 1 1): haptomerum with two heli; pedium with 50 sensilla; right
chaetoparia with 1 2 long setae and some smaller setae; left chaetoparia with six
long setae and some smaller setae, pterotormae rounded apically. Left mandible
with scissorial teeth and the inner margin of the scissorial area worn (Fig. 22).
Maxillae (Fig. 26) each with 6-7 large setae lateral to uncus of galea and 12-17
large setae behind uncus of lacinia. Labium (Fig. 24). Antenna (Fig. 5).

Thorax.— Respiratory plate of thoracic spiracle C-shaped (Fig. 28). Prothorax
with sclerotized lateral area forming a long furrow. Each mesothoracic leg with
pars stridens (Fig. 38) consisting of 84 teeth; metathoracic legs each with plectrum
(Fig. 42) consisting of 62 suboval teeth, distal teeth becoming elongate.

Abdomen. — Anal lobes and anal pad bare (Fig. 46). Raster (Fig. 48) with about
324 small to long setae.

Material. — Description based on one female third instar larva, fully developed,
associated with adults; deposited in CMNH collection.

Locality. — Ecuador: Napo, Cosanga, 1500 m, G. Onore. Under rotten log in
grass-covered soil.

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Onore — Immature Stages of Sphaenognathus

Discussion

For this study I examined larvae of the following genera: Ceratognathus, Ceru-
chus, Chiasognathus, Dorcus, Figulus, Lamprima, Lissapterus , Lissotes, Lucanus,
Platycerus, Sinodendron, Sphaenognathus, Syndesus, and Rhyssonotus. I assessed
the characters of Pholidotus from the description and illustrations of Costa et al.
(1988). The characters discussed below indicate that the genera most closely
related to Chiasognathus and Sphaenognathus are Rhyssonotus and Pholidotus,
and only these genera are discussed further. The same relationships were found
by Lacroix (1969) and Holloway (1 960) in their phylogenetic interpretations based
on adults.

The following larval characters have been considered useful in distinguishing
various groups of Lucanidae.

1. Mandible. The number of teeth in the left mandible has been used for
distinguishing higher taxa of Lucanidae (Van Emden, 1935, 1941; Ritcher, 1966;
Lawrence, 1981). However, this character is a problem to assess in larvae that
have not recently molted, due to normal wear of the mandibles. The teeth are
divided into two categories: the teeth of the scissorial area and the teeth basal to
the scissorial area (between the scissorial area and the molar area) which I here
designate “basal teeth” (Fig. 15, 16). The mandibles in S. lindenii, S. metallifer
(Fig. 1 9), and S. oberon have four scissorial teeth; the same condition is found in
Rhyssonotus sp. Costa et al. (1988) illustrated four scissorial teeth in Pholidotus
spixi. Sphaenognathus nobilis has five scissorial teeth (Fig. 15, 16), and the con-
dition is unknown in S. subtilis and Chiasognathus grand. Two basal teeth are
present in Rhyssonotus, and one basal tooth in S. lindenii, S. oberon, S. metallifer,
and S. nobilis (Fig. 15, 16, 19, 21, 23). Basal teeth are absent in S. peruvianus
(Fig. 20). This absence is interpreted as derived, and strengthens the reestablish-
ment by Bartolozzi et al. (1992) of C bias o gnat hinus (Didier and Seguy, 1953) as
a distinctive and perhaps monophyletic subgenus of Sphaenognathus containing
S. peruvianus and S. gaujoni (Oberthiir).

2. Prothorax. A sclerotized, anteriorly projecting lobe on the pro thorax was
used as a character by Ritcher (1966) and Lawrence (1981). This structure is not
present in Chiasognathus and Sphaenognathus. A second character, the lateral
sclerotized furrow (Fig. 1, 27), is continuous in all known larvae of Sphaenog-
nathus except S. peruvianus wherein the furrow is interrupted by a nonsclerotized
area. The latter condition is considered derived, and occurs as well in Rhyssonotus.
The character was not illustrated for Pholidotus by Costa et al. (1988).

3. Mesothoracic legs. The shape, number and position of teeth of the pars
stridens of the mesocoxae have been used by Van Emden (1952), Ritcher (1966),
Klausnitzer (1978), and Lawrence (1981). Larvae of species of Sphaenognathus
have similar structures except for the number of teeth (Fig. 36-39). The same
condition is present in Pholidotus and Rhyssonotus. Chiasognathus larvae differ
in having a distinctive field of teeth at the proximal end of the main row of teeth
on the pars stridens (Fig. 35).

4. Metathoracic legs. The plectrum of the metatrochanter is functionally related
to the pars stridens. In the genera Sphaenognathus (Fig. 41-43), Chiasognathus
(Fig. 40), Pholidotus, and Rhyssonotus, this structure is very similar with variation
only in the number and shape of the teeth.

5. Setation of anal sclerite. The dense vestiture of setae in the area surrounding
the anal pad separates S. lindenii, S. metallifer, S. oberon, and S. nobilis (Fig. 44,
45, 47) from S. peruvianus and S. subtilis (Fig. 46), which lack this vestiture.

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Chiasognathus, Rhyssonotus, and Pholidotus have a reduced number of dense
setae.

6. Epipharynx. The basic anatomy of the epipharynx is similar in Sphaenog-
nathus (Fig. 9, 11-14), Chiasognathus (Fig. 10), Pholidotus, and Rhyssonotus.
Perhaps the most important character common to the above-mentioned genera
is the vestiture of setae lateral to the nesia of the haptolachus. In Chiasognathini,
the sensilla in the pedium and the setae of the paria seem to have the same origins
and they are sometimes represented as “emerging setae”; in the haptomerum
there are two or more coarse nail-like spines which I interpret as heli following
Boving (1936).

7. Tarsungulus. Van Emden (1941), Medvedev (1952), Paulian (1959), and
Ritcher (1966) used the number of setae present on the tarsungulus as a character
separating Lucaninae from Dorcinae and some other subfamilies. Sphaenognathus
(Fig. 32), Chiasognathus, Pholidotus and Rhyssonotus have two setae on the tar-
sungulus. This character indicates that they should be included in Dorcinae.
However, other characters support a relationship with Lucaninae.

8. Raster. The setae of the raster, localized in patches, are all inclined laterally
in species of Chiasognathini (Fig. 1 , 44-49).

A larval diagnosis for the family Lucanidae remains speculative with so few
larvae available for study. Larval features suggest that Chiasognathini is related
to Lucaninae and Dorcinae in agreement with Holloway’s analysis of adults. That
author concludes, “There appears to be no valid reason, at the present time, for
recognizing as distinct groups the . . . Chiasognathinae, Dorcinae . . . ; probably
all . . . belong in Lucaninae” (Holloway, 1960).

The most recent larval keys to lucanid subfamilies, based primarily on Palearctic
and Nearctic genera, will not place all Chiasognathini in Lucaninae because S.
peruvianus lacks the tooth between the inner margin of the scissorial area and the
molar area (Fig. 20), a character considered diagnostic for Lucaninae. An im-
portant character linking Chiasognathini to Lucaninae is the length of the dorsal
surface of abdominal segment 10, being half or less the length of the ventral
surface of that segment (Fig. 1).

Known larvae of Sphaenognathus are very similar in anatomy with the excep-
tion of S. peruvianus, which presents features distinctive enough to justify reval-
idation of the subgenus Chiasognathinus.

Phylogenetic Notes

Although characters of larvae do not permit complete resolution of the phy-
logeny of the species described here, several apomorphic characters support the
monophyly of three taxa as follows.

1 . The monophyly of a lineage consisting of all examined species of Chiasog-
nathini is supported by four characters tentatively considered to be apomorphic.
Chiasognathini is also characterized by the presence of larval stemmata, but this
is plesiomorphic and not phylogenetically informative.

a. All setae on the raster inclined or directed laterally. Other Scarabaeoidea
have the setae uniformly directed medially or in many different directions.

b. Reduction in size of the third anal lobe. The third lobe is not reduced in
Syndesus and other Scarabaeoidea.

c. Presence of a basal tooth on the scissorial margin of the mandible. This is
also present in Lucaninae, although absent in other Scarabaeoidea. Its

1994

Onore- Immature Stages of Sphaenogna thus

absence in Sphaenognathus peruvianus is hypothesized to represent a sec-
ondary loss.

d. Presence of short thick setae on the anal lobes. Such short thick setae are
absent in Syndesus, but their occurrence in other Lucanidae is not known.
They appear to have been secondarily lost in Sphaenognathus subtilis.

2. Chiasognathus shares three characters with Syndesus that may be considered
plesiomorphic for Chiasognathini. The derived condition of each of these char-
acters may represent synapomorphies for Sphaenognathus as here delimited.

a. Teeth on the pars stridens in a single row. The pars stridens consists of a
field of multiple, scattered teeth in Chiasognathus and Syndesus.

b. Apex of penultimate antennomere not extended past the base of apical
antennomere. In Chiasognathus and Syndesus the distal end of the pen-
ultimate antennomere is modified, and the unmodified condition in
Sphaenognathus is interpreted as a secondary apomorphic reduction.

c. Apical antennomere oblate to cylindrical. This antennomere is pyriform
or conical in Chiasognathus and Syndesus.

3. The monophyly of a group consisting of Sphaenognathus lindenii, S. oberon,
and S. metallifer is supported by one synapomorphy.

a. Presence of minute, slender setae on the anal lobe. These tiny setae are
absent in other Chiasognathini, Syndesus, and many other genera of Lu-
canidae.

Alterations in the existing classification of Chiasognathini based on the above

characters would be premature, and must wait until a more extensive analysis

of characters from all development stages has been completed.

Acknowledgments

This research was conducted with support and facilities from the International Visitors Program at
Carnegie Museum of Natural History, for which I thank Director J. E. King. I thank L. Arcos Teran
and T. De Vries for support from the Departamento de Biologia de la Pontificia Universidad Catolica
del Ecuador.

I am grateful to the institutions and persons providing specimens for study: OSUO, A. Asquith;
FSCA, D. H. Habeck; QCAZ, L. Arcos; WSUC, R. S. Zack, Jr.; and C. Vergara. I thank the following
persons for donating or exchanging specimens: J. F. Lawrence (CSIRO, Canberra, Australia); H. F.
and A. T. Howden (Carleton University, Ottawa, Canada); and L. E. Pena G., J. Numhauser T., and
J. Solervinces A. (all of Santiago, Chile). Field assistance was provided by L. A. Coloma (Departamento
de Biologia, Pontificia Universidad Catolica del Ecuador) and C. Bordon (Maracay, Venezuela). I am
grateful to H. F. Howden (Carleton University, Canada), D. W. Roubik (Smithsonian Tropical Re-
search Institute, Panama), and R. E. Woodruff (Bureau of Entomology, Division of Plant Industry,
Gainesville, Florida) for valuable suggestions and encouragement.

Adult Lucanidae associated with larvae were determined by L. Bartolozzi (Universita degli Studi
di Firenze, Firenze, Italy). J. Auber (Universite Pierre et Marie Curie, Paris, France) kindly examined
specimens to confirm the presence of the stemmata. L. Baert (Institut Royal des Sciences Naturelles,
Bruxelles, Belgium) provided some rare publications on lucanid larvae. S. Thompson (Section of
Botany, CMNH) assisted with botanical details.

Assistance with illustration was provided by C. W. Young (CMNH); M. A. Klinger (CMNH) rendered
Fig. 50-52; M. A. Daman assisted with manuscript preparation. Two anonymous reviewers provided
useful and constructive criticism. Finally, my special thanks to R. Davidson and J. E. Rawlins (CMNH)
for friendly help, scientific advice, and endless patience in revising this paper.

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Cekalovic, T. K. 1982. Descripcion de la larva y pupa de Pycnosiphorus femoralis (Guerin, 1839).
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Cekalovic, T. K., and M. M. Castro. 1983. Chiasognathus granti Stephens, 1831 (Coleoptera:
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Costa, C., S. A. Vanin, and S. A. Casari-Chen. 1988. Larvas de Coleoptera do Brasil. Museu de
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Holloway, B. A. 1960. Taxonomy and phylogeny in the Lucanidae (Insecta: Coleoptera). Records
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Paulian, R. 1959. Faune de France. 63. Coleopteres Scarabeides, Lechevalier, Paris.

Perrault, G. G. 1991. A propos d’un peuplement de Sphaenognathus Buquet dans les Andes du
Venezuela. (Coleoptera, Lucanidae). Nouvelle Revue Entomologique (N.S.), 8(3):258.

Rjtcher, P. O. 1966. WMte Grubs and their Allies. Oregon State University Press, Corvalis, 219 pp.
Ureta, E. R. 1934. Sobre algunas costumbres del Chiasognathus grand Steph. Revista Chilena de
Historia Natural, 38:287-292.

Van Emden, F. 1935. Die Gattungsunterschiede der Hirschkafer-larven, ein Beitrag zum naturlichen
System de Familie (Coleoptera: Lucanidae). Stettiner Entomologische Zeitung, 93:178-200.

— -. 1941. Larvae of British beetles. II. A key to the British Lamellicomia larvae. Entomologists

Monthly Magazine, 77:1 17-127, 181-192.

— — — . 1952. The larvae of Dendezia and Figulus, with notes on some other larvae of Lucanidae.
Revue de Zoologie et de la Botanique Africaine, 46(3-4):301-310.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 1, P. 101-102

22 February 1994

REVIEWS

THE SCENT OF ORCHIDS: OLFACTORY AND CHEMICAL INVESTI-
GATIONS. Roman Kaiser. 1993. Elsevier Science Publishers B.V., Amsterdam,
The Netherlands. 259 pp., with charts, tables, line drawings, color photographs
and index. ISBN 0-444-89841-7. $175.00.

The Scent of Orchids is an expensive volume which, according to the publishers,
is intended to convey the enormous variation in scent and appearance of orchid
flowers to people engaged in the fragrance field, professional scientists, naturalists,
and hobbyists. Many writers have reported on these variations, but the detailed
analyses of the aroma chemistry of a large number of orchid species from five
different geographic areas provided in this work are an innovation.

Roman Kaiser, a Swiss research chemist, photographer, and student of botany
has, over the past ten years, documented more than 2000 orchid species employing
olfactory descriptions and photographs. His treatise, divided into three parts and
an appendix, incorporates many of the results of his investigations.

Kaiser’s work provides an insight into the large number of odors produced by
orchids which can be detected by vertebrate and insect pollinators, describes
methods by which these scents may be collected and analyzed, tabulates the
chemical composition of the scents of 155 orchid species, and provides a sub-
stantial reference list of orchid fragrance studies.

Part One, Introduction to the World of Orchid Scents, contains a brief history
of the early study and appreciation of orchids, their habitats and geographical
distribution, and a general description of an orchid’s floral structure. This is
followed by a section discussing the nature of plant scents, their toxicity to and
storage by plants, and the importance of their volatility. Many readers may be
surprised to learn that the scent given off by a plant is controlled by the plant’s
biological clock and is not dependent upon visible external stimuli. In addition
to describing the principal methods of plant pollination in which scent is a factor,
Kaiser provides a brief resume of human efforts to utilize scented plant oils first
for rituals and subsequently for medicinal and cosmetic purposes.

Of particular interest to scientists involved in the investigation of floral fra-
grances is the section devoted to a description of the author’s techniques for
trapping scents without damage to plants or flowers. Kaiser describes procedures
for recovering aroma samples in the range of 1-300 ng and determining their
qualitative and quantitative compositions by gas chromatography and mass spec-
trometry. A section on principles of orchid pollination and the part played by
plant fragrances presents little new information but might be useful to readers
unfamiliar with these topics.

Finally, in a section on the verbal description of scents, Kaiser emphasizes the
importance of discrete recounting of smells to complement the complex analytical
systems or as a substitute when the latter cannot identify all olfactory components
of a fragrance. He borrows from perfumery four floral-scent groups identified both
as to olfactory response and chemical composition and relates these more familiar
flower fragrances to some specific orchid scents. The general orchidist will find
the floral-image scent groups useful to describe the complex olfactory scent of

101

102

Annals of Carnegie Museum

vol. 63

some orchid species. Orchid taxonomists may be disappointed that the author
did not discuss utilization of scent composition patterns for orchid species iden-
tification, but his work demonstrates that both the quantitative and the qualitative
composition of scent within a species can vary from plant to plant, and is also
dependent upon plant maturity and the hour when a sample is collected.

Part Two, Interdisciplinary Discussion of Orchid Scents, describes floral scents
of 146 selected species, representing approximately 10% of all orchid genera,
conveniently arranged according to their country of origin into four geographic
regions (American tropics, African tropics, Indo-Australian tropics and subtrop-
ics, and Europe). This section provides descriptions of habitats, scent chemistry,
and scent release patterns, as well as high-quality, close-up color photographs of
more than 140 orchid species, some seldom illustrated. Although these photo-
graphs will prove attractive to any orchidist, the orchid grower primarily interested
in illustrations of orchid species should consider one of the many less expensive
books of orchid illustrations as an alternative.

Part Three, The Chemistry of Orchid Scents, includes remarks on previous
investigations of orchid fragrances, limitations of the present study, and some
generalizations that can be drawn from the investigations described in earlier parts
of the volume. Detailed specifications of the equipment used in the study are
provided as well as characterizations of the analytical samples for which aroma
chemistry is recorded. This is followed by tables displaying the chemical com-
position of the scents collected from orchid species described in Part Two.

The Appendix includes a list of 74 references worldwide to studies of floral
fragrances of Orchidaceae and other plant families and, following this, a list of
36 books and journals for further reading about orchids.

The care with which the translation of the original text from German into
English has been made provides comfortable reading. Throughout Parts One and
Two, geometric diagrams are included to illustrate the molecular structure of
principal components of fragrances of some orchid species. Unfortunately, there
is neither a guide to the nonchemist for interpreting these diagrams nor an ex-
planation of their significance. Even those comfortable with the diagrams will find
the lack of captions inconvenient; the reference numbers of the diagrams appear
in the text, but not always on the same page with the diagram. This book will be
a useful reference for the investigator of floral fragrances, but probably is not a
useful purchase, given its cost, for the much wider audience of amateur and
professional botanists.

FREDRICK W. WRIGHT, JR., Volunteer, Section of Botany.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 1, P. 103

22 February 1994

Erratum

Volume 62, number 3, page 248, line 4. It should read: Superdivision Neodiapsida

incertae sedis.

103

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ANNALS

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THE CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213
VOLUME 63 31 MAY 1994 NUMBER 2

CONTENTS

ARTICLES

Palaeophichthys parvulus Eastman, 1908, a gnathorhizid dipnoan from the

Middle Pennsylvanian of Illinois, USA Hans-Peter Schultze 105

Exploitation of mammals at the Early Bronze Age site of West Row Fen

(Mildenhall 165), Suffolk, England Sandra L. Olsen 115

Tephrodytes brassicarvalis, new genus and species (Anura: Pelodytidae), from
the Arikareean Cabbage Patch beds of Montana, USA, and pelodytid-
pelobatid relationships Amy C. Henrici 155

Taiwanese species of Neopanorpa (Insecta: Mecoptera: Panorpidae)

George W. Byers 185

JUN 1 4 1994

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PALAEOPHICHTHYS PAR VULUS EASTMAN, 1908,
A GNATHORHIZID DIPNOAN FROM THE MIDDLE
PENNSYLVANIAN OF ILLINOIS, USA

Hans-Peter Schultze1

Abstract

Palaeophichthys Eastman, 1908 is a gnathorhizid dipnoan. Monongahela Lund, 1970, a taxon based
on isolated tooth plates, is its junior synonym. Palaeophichthys has tooth plates and a shoulder girdle
typical of a gnathorhizid. The genus is distinct from Gnathorhiza in the skull-roof pattern (unpaired
median B-, C-, E- and F-bones) and in details of the tooth plates (the fourth tooth ridge originates
lateral to the apex). The genus Palaeophichthys ranges from Middle Pennsylvanian to Early Permian
in east-central North America.

Introduction

In an unlikely place— a volume on the Devonian fishes of Iowa— Eastman ( 1 908)
erected the monotypic genus Palaeophichthys (P. parvulus) based on a single
specimen from the Middle Pennsylvanian of the Mazon Creek area, Grundy
County, Illinois. His holotype, a tiny fish on counterpart halves of a typical nodule
from Mazon Creek, had been part of the S. S. Strong collection, Accession No.
[222] 3 of the Peabody Museum of Natural History. Donated by Yale University
to the Museum of Comparative Zoology at Harvard College, it was catalogued
as MCZ 5090a+b. Working from the unprepared specimen, on which much of
the detail was obscured by an infilling of white kaolin, Eastman provided only a
superficial description and a sketchy line drawing in which the eye and gaping
mouth were wholly imaginary. The magnification of his figure was mis-stated as
“x2-l” instead of the actual x 3 (Schevill, 1932). In default of morphological
detail, Eastman attempted to classify his new genus, Palaeophichthys, mainly on
the basis of a median fin, in which the dorsal is confluent with the caudal and
anal. He mentioned a similar confluent unpaired fin in the dipnoans Phanero-
pleuron and Uronemus, but found other features that suggested affinities with
coelacanths. “On the whole,” he concluded, “the most plausible interpretation of
Palaeophichthys seems to be to regard it as an aberrant and extremely degenerate
offshot of fringe-finned ganoids adapted to a mud-grovelling mode of existence”
(Eastman, 1908:254).

In a subsequent publication, Eastman (1917:272-273) added to the hypodigm
of Palaeophichthys parvulus a second and topotypic specimen, USNM 4433 (sic,
mis-cited as “4453”) in the R. D. Lacoe collection at the National Museum of
Natural History, Smithsonian Institution. From its water-worn condition this
specimen, a half-nodule, appears to have been recovered from the bed of Mazon
Creek itself. Aside from observations on the gross form and squamation of the
fish, his second paper added no further information on its morphology or system-

1 Museum of Natural History and Department of Systematics and Ecology, University of Kansas,
Lawrence, Kansas 66045.

Submitted 23 November 1992.

105

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Annals of Carnegie Museum

vol. 63

atic position. “This peculiar crossopterygian genus,” Eastman observed, “has been
previously referred to the Coelacanthidae, but is distinguished from all other
members of the family by its elongate, anguilliform body and continuous median
fins. In the latter respect an agreement is to be noted with the specialized and
problematical genus Tarrasius, from the Lower Carboniferous of Scotland, and
also with Conchopoma gadiforme Kner, from the Lower Permian of Rhenish
Prussia. Possibly both Tarrasius and Conchopoma should be regarded as aberrant
coelacanths.” The second specimen was illustrated by a retouched and slightly
reduced photograph that reveals little morphological detail.

Lacking a proper description, later authors followed Eastman in assigning Pa-
laeophichthys to either the Actinistia (coelacanths), to primitive actinopterygians
( Tarrasius ), or to the Dipnoi ( Conchopoma ). Jordan (1923) and Hay (1929) placed
the Tarrasiidae with Palaeophichthys within the Actinistia; Zittel (1923, 1932)
cited Palaeophichthys as probably within the Coelacanthidae. After Moy-Thomas
(1934) demonstrated that Tarrasius is, in fact, a primitive actinopterygian, Pa-
laeophichthys was regarded as an actinopterygian by Berg (1936), Romer (1945),
Moy-Thomas and Miles (1971), and Bardack (1979). lessen (1973) did not assign
the genus to any group despite comparison with Tarrasius and other forms with
a continuous fin fold, and Lehman had a similar approach (1966: “not a cros-
sopterygian”). As the third group used for comparison by Eastman (1908, 1917),
the dipnoans were suggested as the proper place for Palaeophichthys by Vorobyeva
and Obruchev (1964: within Ctenodontidae together with Ctenodus, Tranodis,
Sagenodus, Megapleuron, Proceratodus, and Nielsenia ), Romer (1966: Dipnoi
incertae sedis), Baird (personal communication in Bardack [1979:51 1]: “a senior
synonym of the dipnoan Conchopoma edesi . . .”; and in Schultze and Bardack
[1987:1]: “. . . a dipnoan, probably a new genus”) and Carroll (1987:Dipnoi in-
certae sedis).

For most of the past half century, the type specimen has been inaccessible to
researchers, having been borrowed around 1938 by T. Stanley Westoll of the
University of Newcastle-upon-Tyne, England. Professor Westoll retired without
publishing his conclusions on the affinities of Palaeophichthys, and in 1985 the
specimen was returned. With the newly repatriated specimen in hand, the present
restudy was undertaken to establish the true affinities of the fish.

The type and hypotype specimens have been prepared by removing the en-
crusting kaolin from the nodules with porcupine quills, insect pins, and small
bristle brushes. Red latex casts (Baird, 1955) were then made from the natural
molds.

Systematic Paleontology

Subclass Dipnoi Muller, 1845
Family Gnathorhizidae Miles, 1977
Genus Palaeophichthys Eastman, 1908
Synonym: Monongahela Lund, 1970
For synonymy list see Schultze (1992:200-201)

Diagnosis. — Gnathorhizid dipnoan with four unpaired median skull roof bones
(B, C, E, and F). Pterygoid tooth plate with four ridges, the most posterior (fourth)
ridge originating posterolaterally to the apex, whereas the other three diverge

1994

Schultze — Palaeophichthys, A Gnathorhizid Dipnoan

107

anterolaterally. Prearticular tooth plate with three ridges. Elongated body (depth
= 9.5-12.5% of total length; head length = 14-15% of total length).

Range. — Early Westphalian D to Wolfcampian, Permo-Carboniferous.

Type species. —P. parvulus Eastman, 1908.

Palaeophichthys parvulus Eastman, 1 908

Palaeophichthys parvulus, Eastman, 1908:253, fig. 37.

P. parvulus, Eastman, 1917:272, PI. 10, fig. 2.

P. parvulus, Schevill, 1932:85.

P. parvulus, Vorobyeva and Obruchev, 1964:314.

P. parvulus, lessen, 1973:177.

P. parvulus, Schultze, 1992:201.

Diagnosis.— Angle between first and second ridge of pterygoid tooth plate 40°,
and between second and third ridge 28°.

Stratigraphic Position and Locality. — Francis Creek Shale, Carbondale For-
mation, Westphalian D, Middle Pennsylvanian; Mazon Creek area, Grundy Coun-
ty, Illinois, U.S.A.

Holotype.— MCZ 5090 a,b, complete specimen.

Holotype. — USNM 4433, poorly preserved complete specimen.

Description

A small, typical Mazon Creek concretion contains the holotype in part and
counterpart; only one part of the concretion of the hypotype is preserved. As
common for Mazon Creek, only the molds of the bones are preserved. The de-
scription is therefore based on latex casts prepared by Donald Baird, and the
bones are described as if they are preserved three-dimensionally. The holotype is
better preserved than the hypotype; thus the description refers mainly to the part
and counterpart of the holotype.

The specimens are elongate but not eel-like as described by Eastman (1908).
The holotype (Fig. 1) is 3.6 cm long with a depth of 0.45 cm, the hypotype is 5.2
and 0.7 cm, respectively. The head is 5.5 mm long (7.4 mm in hypotype), less
than % of the total length. The part (MCZ 5090a) shows the right side of the
specimen (MCZ 5090 a, Fig. 2), with the skull roof seen in oblique dorsal view;
a smooth ostracod lies at the approximate place of the stomach. The counterpart
(MCZ 5090 b, Fig. 3) shows bones of the left side and internal views of bones of
the right side of the skull roof in oblique ventral view, and bones of the palate,
gill arches, and shoulder girdle. Despite the small size, the skull roof, palate, and
ceratohyal are fully ossified, and the complete body covered by scales.

Skull Roof (Fig. 2).— The skull roof displays three median bones, broad B- and
C-bones, and a narrower E-bone. Only the inner side of the lateral portion of
these bones can be seen in the hypotype. The posterior part of bone B, with its
occipital commissure, is missing. The counterpart (Fig. 3) shows the impression
of two small bones anterior to bone E, a median F-bone and the right lateral
Q-bone with large evaginations for the pores of the supraorbital canal. The three
median bones are displaced laterally over their neighboring bones toward the wide
groove of the lateral line (Fig. 2). Only part of the inside of the left I and J bones,
and part of the right J-bone in the comer between bones B and C are exposed.
The anterior part of bone Z carries the broad lateral line canal, whereas the
branching-off of the occipital commissure is only weakly preserved. In front of

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Annals of Carnegie Museum

vol. 63

Fig. 1 .—Palaeophichthys parvulus Eastman, 1908. Latex cast of holotype MCZ 5090a dusted with
NH4C1.

bone Z lies a small bone Y, followed anteriorly by a longer, only partly preserved
X-bone. The pattern of the skull roof cannot be restored from the two specimens.

Cheek. — Because only a few bones are preserved behind the orbit, the cheek
pattern cannot be established.

Lower jaw. — The right ‘‘angular” is completely preserved on the right side (Fig.
2), whereas only the posterior part of the left “angular” is preserved on the
counterpart (Fig. 3). The position of the mandibular canal is marked as a deep

1mm

Fig. 2.— Palaeophichthys parvulus Eastman, 1908. Head of holotype MCZ 5090a. Abbreviations:
“Ang,” angular; B, C, E, I, J, X, Y, Z, skull-roof bones; Clei, cleithrum; Gu, gular; 1c, main lateral
canal; n.sp, neural spine; occ, occipital commissure; Op, operculum; Po, postorbital bone fragments;
Pra, prearticular; Pt, pterygoid; r, rib; Sop, suboperculum, “Spl,” splenial.

1994

Schultze — Palaeophichth ys, A Gnathorhizid Dipnoan

109

Fig. 3 . — Palaeophichthys parvulus Eastman, 1908. Palate and shoulder girdle regions of holotype MCZ
5090b. Abbreviations: “Ang,” angular; C, E, F, J, KL, Q, X, Y, skull roof bones from the inside; chy,
ceratohyal; Cla, clavicle; Clei, cleithrum; Gu, gular; n.sp, neural spine; Op, operculum; Pra, prearticular;
Pt, pterygoid; Psp, parasphenoid; Vo, vomer.

groove on the “angular.” Anterior to the “angular” the “splenial” reaches an-
terolaterally to the prearticular. The labial side of the right prearticular is exposed
partially on the part and the lingual side on the counterpart. The prearticular
tooth plate is only seen in lateral view; it bears three ridges.

Palate. — Opposed to the tooth plate of the lower jaw, the tooth plate of the
right pterygoid shows indication of four ridges. These ridges are clearly visible on
the left pterygoid (Fig. 3, 4). A long anterior ridge is combined with three short
ridges that are directed anterolaterally and posterolaterally. The first three ridges
diverge from the apex of the tooth plate, whereas the most posterior ridge orig-
inates laterally to and lower than the apex. The ridges show indications of cusps.
The angles between the first and second, second and third, and third and fourth
ridges are 40°, 28°, and 55°, respectively. The posterior flange of the pterygoid has
a steep medial side and a narrow width. The vomerine tooth is located on a
pointed, anteriorly directed base in front of the anterior ridge of the pterygoid
tooth plate. No cusps are visible at the margin of the vomerine tooth plate. Only
the most posterior end of the parasphenoid is visible between the bones of the
shoulder girdle anterior to the ribs.

Opercular and Gular Region. — The counterpart (Fig. 3) shows a large opercu-
lum, which is anteroposteriorly longer than dorsoventrally deep (4:3). The sub-
operculum, preserved on the part below the operculum, is as long as the oper-
culum, but much lower in depth (5:2). The suboperculum is the best preserved
bone in the hypotype and has the same shape and size relations as in the holotype.

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Annals of Carnegie Museum

vol. 63

Fig. 4. — Palaeophichthys parvulus Eastman, 1908. SEM of Latex cast of holotype MCZ 5090b, left
pterygoid tooth plate; x72 (each white line corresponds to 100 ixm).

Part and counterpart show indications of a lateral gular adjacent to clearly pre-
served principal gulars. The principal gular plate shows striations radiating toward
the margin. The ceratohyal is completely visible in the hypotype, but only the
posterior part is in the holotype (Fig. 3). It is an elongated bone with narrow
middle portion and expanded anterior and posterior ends.

Poster anial Skeleton. — The shoulder girdle of the holotype (Fig. 3) and hypotype
consists of a relative short, broad, massive cleithrum and a long, narrow clavicle.
The clavicle widens dorsally, where it overlaps the ventral part of the cleithrum.
The body of the part and counterpart of holotype and hypotype shows a complete
covering of scales. The scales (Fig. 5) are round and elongated, with parallel ridges
on the anterior covered field and converging ridges on the posterior exposed field.
They show the division in fields typical for post-Devonian dipnoans. Ribs extend
posteriorly from the shoulder girdle 40-50% of the body length and indicate the
extent of the abdominal cavity. The weakly ossified series of spines extends pos-
teriorly nearly as far back as the ribs.

Paired fins or their traces are not preserved in either specimen. A continuous
fin fold surrounds the posterior part of the body. Dorsally it occupies 60% of body
length and ventrally 42%. There are no supporting elements visible for the fin
fold. The lepidotrichia are not articulated and also seem not to bifurcate.

Age

Lund (1970, 1973) described age pattern in tooth plates of Palaeophichthys
( Monongahela ) stenodonta and dunkardensis. In both species the number of cusps
increases with size increase of tooth plates; the cusps are worn in larger tooth
plates. The fourth ridge of the pterygoid plate appears late in subadult stage (Lund,
1970:253). This ridge is developed in P. parvulus, which indicates that we are
dealing at least with subadult, if not adult, specimens, despite their small size.

1994

Schultze— Palaeophichthys, A Gnathorhizid Dipnoan

111

Fig. 5 . — Palaeophichthys parvulus Eastman, 1908. Scale of holotype MCZ 5090 from the middle of
the body above the beginning of the anal fin.

Further indications of subadult or adult stage are the full ossification of skull roof,
palate, and ceratohyal; the complete covering of the body by scales; and the lack
of cusps on the vomerine tooth.

Comparison

Palaeophichthys parvulus is a tooth-plated dipnoan (after the terminology of
Campbell and Barwick, 1983), and has no relationship to the denticulated dipnoan
Conchopoma. The tooth plates of Palaeophichthys resemble those of gnathorhi-
zids, and the tooth plate of the pterygoid agrees with the diagnosis of the gna-
thorhizid genus Monongahela (Lund, 1 970) in that the most posterior fourth ridge
originates laterally to the apex of the tooth plate. On the basis of this evidence,
Monongahela is, therefore, considered a junior synonym of Palaeophichthys. The
genus Monongahela was described at a time when the true nature and affinities
of Palaeophichthys were entirely unknown. Palaeophichthys parvulus is distinct
from the two species P. stenodonta and P. dunkardensis described by Lund (1970,
1973): it has a different angle between the ridges of the tooth plates, different
shape of cusps on the ridges (Table 1), and different position of the fourth ridge.
Differences in cusp shape also occur on the vomerine tooth plate. The most
posterior fourth ridge of P. parvulus originates closer to the apex than in the other
two species.

The skull roof pattern of Palaeophichthys is unique within Paleozoic dipnoans
in having three, possibly four, median unpaired bones (B, C, E, and F): in Sa-
genodus and Conchopoma bones B and C are unpaired, but bone E is paired; in
Megapleuron and Gnathorhiza bones B and E are unpaired, but bone C is paired;
whereas Ctenodus and Tranodis have paired C- and E-bones. The composition

Table 1. — Comparison of the pterygoid tooth plates of the three species of Palaeophichthys. * Data

from Lund (1970, 1973).

Angle between

1. a. 2. ridge
(range)

Angle between

2. a. 3. ridge
(range)

Shape of cusps
on ridges

P. dunkardensis*

14.37

15.49

rounded

(7-28)

(9-32)

P. stenodonta*

41.65

48.35

laterally flattened

(12-60)

P. parvulus

distinct

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Annals of Carnegie Museum

vol. 63

of the lateral-line bones differs from Gnathorhiza : the occurrence of Z-, Y-, X-
and elongate KL-bones is more primitive than the reduction to fewer larger bones
in Gnathorhiza (Berman, 1976). The length/depth relations of operculum and
suboperculum also are different in the two genera: the operculum is longer than
deep in Palaeophichthys, whereas it is deeper than long in Gnathorhiza. On the
other hand, Palaeophichthys possesses a broad cleithrum and a very elongated
clavicle, as does Gnathorhiza (Berman, 1976).

In summary, Palaeophichthys (= Monongahela) is distinct in its skull roof
pattern from other Paleozoic dipnoans, but the tooth plates indicate close rela-
tionship to Gnathorhiza.

Conclusions

Palaeophichthys becomes the fourth identifiable dipnoan genus from the Middle
Pennsylvanian of the Mazon Creek area, joining Conchopoma (Denison, 1969;
Schultze, 1975), Ctenodus (Baird, 1978), and Megapleuron (Schultze, 1977). How-
ever, it is now well understood that two ecologically distinct assemblages constitute
the Mazon Creek fauna (Johnson and Richardson, 1966). On present evidence,
Palaeophichthys is restricted to the Braidwood fauna, a nonmarine association
that is found in the more northerly area of deposition, including the beds along
Mazon Creek in Grundy County. Conchopoma and Megapleuron, on the other
hand, are found only in the marginal-marine Essex fauna, which is best known
from Pit Eleven in Will and Kankakee counties, whereas Ctenodus is recorded
from both assemblages. Although Megapleuron is otherwise known only by two
specimens from another Pennsylvanian locality in North America (Linton, Ohio:
specimen 063. 107-2 in David S. Hamilla’s private collection) and from the Lower
Permian of France, Conchopoma and Palaeophichthys occur commonly in other
Paleozoic localities of North America. Conchopoma is known from the Middle
Pennsylvanian of Linton (Schultze, 1977; Hook and Baird, 1986) and Five Points
(Hook and Baird, in press), Ohio, and from the Upper Pennsylvanian of Vermilion
County, Illinois (Schultze, 1977). Palaeophichthys was described as Monongahela
from the Upper Pennsylvanian of Allegheny County and the Lower Permian of
Washington County, Pennsylvania (Lund, 1970, 1973). It is surprising that only
two specimens of Palaeophichthys have been found in the thousands of concretions
from the Mazon Creek area.

Acknowledgments

The author is grateful to the chain of persons who delivered the holotype of Palaeophichthys parvulus
into his hands: Alick D. Walker, Robert W. Hook, Charles R. Schaff, Mary Ann Turner, S. Christopher
Bennett, and Donald Baird. Donald Baird contributed to the introduction and prepared the peels.
Robert W. Hook, Richard Lund, and Jiri Zidek have helpfully criticized the manuscript. John Chom
kindly photographed the holotype. J. Elder and J. Wigles worth, Word Processing Center, Division of
Biological Sciences, The University of Kansas, kindly typed different versions of the manuscript. The
author thanks the National Science Foundation for partial support of the project through grant BSR-
8806856.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 2, Pp. 1 15-153

31 May 1994

EXPLOITATION OF MAMMALS AT THE EARLY BRONZE AGE
SITE OF WEST ROW FEN (MILDENHALL 165),

SUFFOLK, ENGLAND

Sandra L. Olsen

Assistant Curator, Division of Anthropology

Abstract

The large assemblage of faunal material from the Early Bronze Age site of West Row Fen (Mildenhall
165) in Suffolk, England, provides important data on the economy and subsistence during a poorly
known time period in Great Britain. Dominated by gracile, short-homed cattle and small, Soay-sized
two-homed sheep, this primarily domestic fauna demonstrates that by this period there was minimal
dependence on hunting in comparison to stock rearing. Other common domesticates in the assemblage
are goats, pigs, and dogs. Wild species that were hunted include hare, red deer, roe deer, and aurochs,
the latter represented by only one extremely large humems. It is unclear whether horses at West Row
Fen were domestic or wild. Preserved carnivores include the wild cat, red fox, and domestic dog. The
presence of the western hedgehog, the western mole, and the water vole suggests a wet woodland
habitat typical of the fens during this period.

Aging of the domestic livestock by epiphyseal fusion and dental eruption and wear show differential
treatment of cattle, sheep and goats, and pigs. Cattle were probably used more for draft than for their
dairy products, whereas sheep or goats were being milked. Most pigs were slaughtered in their first
year of life to control population growth, although some were allowed to reach reproductive maturity
and were killed as needed.

Major taphonomic processes included carnivore gnawing, sedimentary abrasion probably caused
by trampling, moderately light root-etching, and, more rarely, erosion, weathering, and rodent gnawing.
Butchering patterns, previously poorly known from the Bronze Age of Britain, suggest that cow horns
were heavily utilized, metal axes may have been used on cattle carcasses, and dogs were eaten.
Pathologies are those frequently associated with domestic animals, such as malocclusions, tooth
anomalies, and bone modification caused by heavy draft. A total of 1 20 bone and antler artifacts were
identified, including awls, a pin, a scoop, a spatula, tubular beads, a dagger, pressure-flakers, a handle,
socketed mace-heads, and a wedge.

Introduction

West Row Fen contains a remarkably well-preserved Early Bronze Age village
that reveals much about the lives of its inhabitants. Located in Suffolk County
just west of the modern town of Mildenhall (Fig. 1), the site had been protected
for centuries by an overlying cap of Iron Age peat. The fen was drained in 1759,
and erosion subsequently removed much of the peat (Martin and Murphy, 1988:
353), exposing the richness of Neolithic and Bronze Age settlements in the fenlands
around West Row. Although plowing has damaged and destroyed most of these
prehistoric deposits, one site, Mildenhall 165, remained relatively undisturbed.

Major excavations of the site were first undertaken in 1982 and continued
through 1986. The archaeological work was conducted by the Suffolk Archaeo-
logical Unit and English Heritage, under the supervision of Dr. Edward Martin.
Approximately 4200 m2 were excavated, yielding over 3 1 ,000 pieces of animal
bone. All lithic and bone artifacts were recorded three-dimensionally using a
reference datum, and un worked faunal material was recorded by feature, layer
within a feature, or 1 -m2 unit.

Submitted 9 December 1992.

115

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Annals of Carnegie Museum

vol. 63

Fig. 1.— Map of East Anglia showing the location of the site of West Row Fen.

The site lies primarily on a penannular sand ridge about 2.3 m above an adjacent
hollow. While some plow damage was noted on top of the sand ridge, most of
the deposits were intact at lower elevations. Some of the deepest features lie below
the water table and therefore contain well-preserved organic matter.

A small lithic scatter of Mesolithic age and a few Neolithic pits and lithic scatters
were found, but the village dates predominantly from the latter part of the Early
Bronze Age. Radiocarbon dates for the Early Bronze Age component place it at
ca. 2290-1780 B.C., calibrated.

The environment in the vicinity of the site can be inferred from the preserved
wood, macrobotanical remains, pollen, terrestrial mollusks, and small vertebrates.
The rich woodlands surrounding the site included oaks on the higher ground and
alders on lower elevations. After the village land was cleared, wet tussock con-
ditions persisted. Scrub and wet woodland plants such as elder, hazel, holly, sloe,
and willow were also present around the periphery. Cultivated plants consisted
primarily of emmer wheat, accompanied by spelt wheat, hulled barley, and flax
(Martin and Murphy, 1988:356).

The remains of mollusks, fish, amphibians, reptiles, and birds are still under-
going investigation by other researchers, but do not contradict environmental
indications from plant remains. The microfauna from flotation and fine screening,
containing numerous rodents and insectivores, is also being analyzed separately
and is excluded from this study.

Portions of three round houses fabricated from light timber were found on the
ridge (Fig. 2). The most complete was about 5 m in diameter and had a porch on
the south-east side (Martin and Murphy, 1988:355). A midden area filled one of
the lower parts of the site.

1994

Olsen “Mammal Exploitation at West Row Fen

117

Fig. 2. —Plan of excavated area showing locations of features and houses. Key to features: 1, house;
2, house; 3, house; 0901, water pit; 0912, water pit; 0921, flax retting pit; 0934, charcoal clamp; 4112,
water pit filled with ash and charcoal; 4226, large water pit; 4227, large feature containing water pit;
4284, antler soaking pit.

The most informative features were the numerous pits distributed over the site
(Fig. 2). The dry pits along the ridge contained charcoal, animal bones, potsherds,
flint, and other domestic debris. The wet pits (0901, 0912, 41 12, 4226, 4227, and
4284) in low-lying areas of the site served several functions. Some were lined
with alder logs to support the sidewalls and allow easy access, and probably
functioned as shallow wells. Pit 092 1 contained seeds and fragments of flax (Linum
usitatissimum ) and appears to be the oldest known retting pit in Britain (Martin
and Murphy, 1988:355). Pit 4284 (diameter, 2 m; depth, 1.05 m), contained a
split red deer antler (length, 42 cm) that was apparently placed in water to soak
prior to undergoing further manufacturing.

The stone implements recovered include scrapers, knives, saws, borers, arrow-
heads, mace-heads, hammerstones, querns, and rubbing stones (Martin and Mur-
phy, 1988:355). Seven small jet toggles, ranging from 1.3 to 2.3 cm in length,
may have served as earrings or buttons. Pottery was mostly Grimston-type plain
ware.

The large assemblage of mammalian fauna from West Row Fen, Mildenhall
165, documents a heavy dependence on domestic species in the Early Bronze Age
with a secondary reliance on wild animals to supplement the diet. At present, few
large collections from the British Bronze Age have been studied. The material
from the Middle Bronze Age deposit at Grimes Graves, in Norfolk (Legge, 1981,
1 992), and from the Late Bronze Age layers at Runnymede in Surrey (Done, 1991,
Serjeantson, 1991) are the most notable exceptions.

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vol. 63

Fig. 3.— Distribution of animal bone fragments throughout the excavated area of the site.

Spatial Distribution of Mammalian Fauna in Site

The unmodified faunal material was distributed unevenly across the site (Fig.
3; Table 1). The densest concentrations were just west of the midden area, but
the midden itself and the area around the flax-retting pit, water pits, and a charcoal
clamp in the western part of the site also contained considerable amounts of
animal bone. Another concentration was found during the 1985 excavations in
the southern part of the site around the antler soaking pit. Three of the pits (0901,
0912, and 4226) contain large numbers of bones, suggesting that the pits were
filled in with food refuse when their original function ended. The species repre-

Table 1 .—Number of identifiable bone fragments in features.

Feature

Canis

Sus

Capreolus

Cervus

Ovis/Capra

Bos

House 1

House 2

House 3

Pit 0901

Pit 0912

Pit 0934

Pit 41 12

Pit 4226

Pit 4227

Pit 4284

Neolithic pits

Ditch

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Olsen— Mammal Exploitation at West Row Fen

119

Table 2. —Frequencies of mammalian taxa from West Row Fen.

Taxon

NISP

% of NISP

MNI

%of

combined
MNI (175)

Individual
usable
meat (kg)b

Combined
usable
meat (kg)c

Class Mammalia

23,189

n.a.

Erinaceus europaeus

Talpa europaea

Lepus, cf. L. capensis

2.25

6.75

Order Rodentia

n.a.

—

?Arvicola terrestris

Felis sylvestris

Vulpes vulpes

Canis familiaris

Cervus, Bos, or Equus

979

n.a.

—

Order Artiodactyla

n.a.

—

Sus scrofa

757

315

Suborder Ruminantia

1198

n.a.

—

Family Cervidae

n.a.

—

Capreolus capreolus

10.5

Cervus elaphus

139

475

Family Bovidae

n.a.

—

Capra hirca

Ovis aries

Ovis/Capra

1617

60d

12.5

750

Bos primigenius

450

Bos taurus

3230

350

19,250

Equus caballus

300

Totals

31,451

175

a n = 8262 fragments (excludes those only identifiable to Class Mammalia).
b Usable meat weight for an average adult individual (based on Milisauskas, 1978).
c Usable meat for the species is calculated by multiplying the usable meat weight per individual by
the MNI for that species.

d MNI recalculated to combine Ovis/ Capra (MNI = 59), Ovis aries, and Capra hirca.

sented in the pits were domesticates, red deer, and roe deer, all of which appear
to have been consumed by the inhabitants. The areas in and around the houses
were kept remarkably clean of animal remains. One possibly significant intraspe-
cific concentration of bones was a cluster of eight dog bones in water pit 4226,
which probably represents food refuse. Animal burials and ceremonial treatment
of animal remains were not observed. Bones of wild species, such as hare, red
deer, and roe deer, were scattered and distributed in the same pattern as the
domestic species.

Methodology

In order to determine relative frequencies of mammalian species at West Row
Fen, both the NISP (number of identified specimens) and the MNI (minimum
number of individuals) were calculated (Grayson, 1979). The MNI was calculated
using a zone technique, which records the portions of each bone present in detail
(Dobney and Rielly, 1988). Each bone is divided into zones according to diagnostic
features and the ways in which bones break most frequently. To calculate MNIs
the number of times a zone appears for the left and right bones is counted sep-
arately. The zone with the largest number (for either the left or the right element)
yields the MNI for that bone. The MNI for a species is the count from the bone
with the highest MNI. Table 2 lists the taxa identified, followed by the NISP and

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Element Frequencies for Capreolus

capreolus

Percentage of M N I

Fig. 4. —Element frequencies for Capreolus capreolus.

MNI for each taxon by absolute numbers and percentages of identifiable mam-
malian fauna (excluding those elements only identified to Class Mammalia).

Table 2 also lists the estimated weight of usable meat in each of the food-
producing species. According to this measure, cattle were clearly the most im-
portant food species at West Row Fen, followed by sheep and goats, and then red
deer. The single aurochs would have yielded as much meat as 36 sheep or goats.
Pigs, which are third most abundant, rank fifth in meat yield. However, meat
yield must be considered in a temporal context. For example, despite the enormous
dietary contribution made by a single wild aurochs, it represents only one event
that would provide a windfall for a few weeks or even months if the meat was
smoked or dried. But aurochs and other wild game would not be as consistently
available as domestic pigs. Domestic animals are “walking larders” and can pro-
vide a secure food source in periods of environmental stress when wild species
cannot. Calculating the actual amount of meat consumed at West Row Fen using
estimated weight of usable meat is a technique fraught with problems (Guilday,
1970) and is not attempted here.

Element frequencies (Fig. 4-8) are useful in contrasting butchering patterns and
body part utilization between hunted and domestic species, especially when the
hunted species are field dressed to lessen the burden of hauling home a whole
carcass, and domestic species are killed near the village. This calculation pertains
more to red deer than roe deer, which are easily carried back whole. There was
no clear evidence of field dressing or discarding of bones of low utility in this
assemblage.

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Element Frequencies for Cervus elaphus

Percentage of M N I

Fig. 5.— Element frequencies for Cervus elaphus.

Figures 4-8 imply differential preservation due to various taphonomic factors.
Phalanges are poorly preserved but teeth are in abundance, implying that the low
number of phalanges was caused by natural or cultural processes rather than
inferior recovery techniques. Phalanges have marrow cavities and are easier to
break than teeth. Dogs may have chewed on the phalanges, destroying them or
leaving only undiagnostic fragments. Humans may also have reduced the pha-
langes beyond recognition with hammerstones while retrieving the marrow. Distal
tibiae were very well-represented, probably because they are very dense and resist
destruction. Flat elements constructed of thin cortical bone not bolstered with
considerable amounts of cancellous tissue, like the scapula, ulna, and innominate
tended to be slightly less well-preserved. These elements are susceptible to break-
age during excessive pedoturbation (Olsen, 1989), such as trampling in this case.

Tooth eruption and wear stages were recorded for domestic ungulates with
reference to Silver (1969), Grant (1982), Bull and Payne (1982), and Legge (1992).
The absence of complete tooth rows, however, impeded reconstruction of mor-
tality patterns for all of the species at West Row Fen. In recording the wear patterns
on the teeth of Ovis/ Capra and Bos taurus, Grant’s (1982) technique was used,
but as it requires nearly complete tooth rows, final determinations of ages were
made chiefly on the basis of Silver’s (1969) eruption dates and information com-
piled by Legge (1992). Both Bull and Payne (1982) and Silver (1969) were used
to age pigs. Redding’s (1981) methods for calculating percentages of animals
surviving past the ages of fusion for long bones was applied to the domestic
animals from West Row Fen.

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Element Frequencies for Sus scrofa

Teeth
Mandible
Scapula
Humerus
Radius

^ Ulna

5 Metacarpal

Innominate
Femur
Tibia
Metatarsal
Phalanx 1
Phalanx 2
Phalanx 3

0 20 40 60 80 100

Percentage of M N I

Fig. 6.— Element frequencies for Sus scrofa.

All fragments with butcher marks were drawn so that the orientation of each
mark and the number of bones with marks in those positions could be determined.
This information is summarized on illustrations of articulated skeletons for each
of the three most prevalent species (Fig. 18, 20, 23). Distinctions between cut-
marks, chopmarks, and scraping were noted. The best examples of each type of
butcher mark were replicated by making silicone rubber molds and epoxy resin
casts for examination in a scanning electron microscope.

Taphonomic traces, pathologies, and anomalies were recorded. Specific causes
are difficult to attribute to pathological alterations of archaeological bone, but
comparisons can be made with living individuals with known histories in order
to make tentative diagnoses.

Conditions of Preservation

Soil conditions at the site of West Row Fen are typical of these fenlands and
are significant regarding bone preservation. The site’s location in sandy deposits
resting on top of chalk provides the soil with a pH that is sufficiently alkaline for
bone conservation. Although less bone survived in the overlying peat layer, the
peat’s acidity apparently did not filter down enough to alter the sand’s pH and
dissolve the bone in that layer. Sand is generally not a good medium for bone
preservation because the loose grains allow water to flow through too easily, but

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Element Frequencies for 0 vis/Capra

Percentage of M N I

Fig. 7.— Element frequencies for Ovis/ Capra.

in this case the chalk inhibited drainage and made the ground water alkaline
enough to maintain a suitable environment for preserving bone. Mineral accre-
tions, apparently rich in iron, were found adhering directly to some bone surfaces,
but these could be removed with gentle brushing in most cases. Butcher marks
as well as manufacturing and use-wear traces on bone artifacts were readily visible
on the well-preserved bone surfaces.

Taphonomy

Despite the generally good condition of the faunal material from West Row
Fen, the effects of several deleterious taphonomic agents were observed. The
frequencies of taphonomic effects on identifiable bone fragments are displayed in
Table 3. Of these processes, carnivore gnawing was the most destructive agent.
Whole condyles of cattle bones were occasionally missing as a result of heavy
gnawing (Fig. 9). Since most butcher marks occur in the epicondylar regions of
long bones, many were probably destroyed as a result of carnivore activity. Car-
nivores probably caused impact scars and spiral fractures as often as humans, so
the extent of marrow extraction by humans was difficult to assess. In this case,
the most likely culprits in carnivore gnawing were the village dogs, since they are
the most common carnivores in the assemblage, and foxes and wild cats were the
only other carnivores identified. Because of their specialized dentition, cats rarely

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Element Frequencies for Bos taurus

Teeth
Mandible
Scapula
Humerus
Radius
Una

5 Metacarpal

^ Innominate

Femur
Tibia
Metatarsal
Phalanx 1
Phalanx 2
Phalanx 3

0 20 40 60 80 100

Percentage of M N I

Fig. 8. —Element frequencies for Bos taurus.

chew on bones. Both foxes and cats are too small to have inflicted serious damage
on cattle bones. Given the prevalence of severe gnawing on ungulate long bones,
this taphonomic factor may explain why small toe bones, carpals, and tarsals of
these species were underrepresented in the faunal assemblage (Fig. 4-8). It prob-
ably also contributed to the paucity of remains of smaller taxa such as birds,
insectivores, rodents, and hares. Four small bone splinters that have a sheen
extending over their broken edges and medullary surfaces appear to have been
acid-etched. The best explanation is that these pieces passed through the digestive
system of a carnivore (Payne and Munson, 1985; Horwitz, 1990), most likely the
domestic dog.

Sedimentary abrasion was the second most destructive taphonomic process.
Abrasion striations caused by the great amount of sand in the soil were often
observed on bone surfaces. The cause of this abrasion is some form of pedotur-
bation which, given the environmental and geologic settings, most likely involved
trampling by ungulates and people (Olsen and Shipman, 1988). Much of the
postdepositional breakage of bones may also have been caused by trampling.
Many of the cow scapulae are heavily abraded and exhibit a recurrent breakage
pattern in which a V-shaped notch is broken out of the center of the blade. The
mouth of the V is at the vertebral border. This breakage pattern might be expected
if a hoofed animal stepped on a scapular blade that was lying flat on the ground.
The missing area on these scapulae represents the thinnest, weakest part of the

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blade. The scapular spine is also rarely preserved. Abrasion usually appears as
sets of fine parallel striations that sweep over the bone surface rather than gouging
deeply into it (Olsen and Shipman, 1988). Abrasion can vary from a polish to
coarse striae as great as 1 mm in width. Although it is normally easy to distinguish
between sedimentary abrasion and cutmarks, abrasion can obscure or erase evi-
dence of butchery.

Root-etching was also distributed throughout the assemblage, but was never

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Fig. 10.— Perforations in a lumbar vertebra of cow probably caused by large roots.

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Table 3. — Taphonomic modifications to identifiable bone fragments.

Taphonomic process

NISP

% of NISP

Carnivore gnawing

347

4.2

Sedimentary abrasion

333

4.0

Root etching

270

3.3

Erosion

168

2.0

Weathering

100

1.2

Rodent gnawing

0.1

Burning

0.1

a n = 8262, number of identifiable bone fragments, excluding bones identified only as Class Mammalia.

dense on any given bone fragment. The small dendritic grooves, etched by the
acid produced at the tips of rootlets, indicate that there had been a ground cover
of vegetation shortly after the bones were deposited. This is also evident from
the peat deposit overlying the sandy soil. A few bones are bored with round holes
that show no indication of tool marks (Fig. 10). Some of these holes may have
been made by the roots of larger plants, possibly tree seedlings. Similar perfora-
tions have been observed by the author in human burials in the eastern United
States where the roots were still in place. Baker and Brothwell (1980:38) recorded
comparable perforations thought to be caused by roots pushing through the cra-
nium of a cow. These perforations are often difficult to distinguish from holes
made in the ends of long bones to remove marrow. Surficial destruction from
root damage was minimal, although some cutmarks were probably obliterated as
a result of this process.

Erosion and weathering were the fourth and fifth most common taphonomic
processes witnessed at West Row Fen. Erosion caused deterioration of the outer

Fig. 1 1.— Scanning electron micrograph of a cutmark made with a dull metal knife. Note the ragged
margins and unstriated, flat-bottomed groove.

128 Annals of Carnegie Museum vol. 63

5 CM

Fig. 12.— Homcore of a domestic short-homed bull with axe chopmarks around the base made during
removal of the horn.

cortical bone surfaces and probably erased some evidence of cultural modification.
Weathering generally resulted only in shallow surface cracks with an absence of
surface exfoliation. Rodent gnawing was surprisingly rare at West Row Fen, an
indication that vermin were kept to a minimum during the occupation of the
village. Bones were buried relatively rapidly, and disturbance by burrowing ani-
mals was probably minimal.

Butchering

Detailed descriptions and illustrations of butchering patterns accompany the
discussions of most species. Bronze tools were not abundant at West Row Fen at
this technological stage (E. Martin, personal communication), but there is evidence
of their use in butchery. Certain criteria can reveal whether a stone or a metal
tool was used to cut bone in well-preserved cases (Olsen, 1988). Most of the fine
cutmarks were probably made with stone tools such as unretouched blades or
flakes. These tools leave very narrow V-shaped grooves with microscopic parallel
striations running along the walls of the groove (Olsen and Shipman, 1988). Metal
knives, however, leave wider grooves with smoother walls and fewer parallel
striations. Metal knife marks are less common than those made with stone tools,
but in one example multiple cuts were made with a dull metal knife that left
broad, flat-bottomed grooves with very ragged margins (Fig. 1 1).

Although stone butchering tools were dominant on the basis of cutmark analysis,
metal axes may have been used to chop antlers and bones. The chopmark produced
by a chipped stone axe is a broad, open V-shaped notch with visible macroscopic
striations running down into the groove roughly perpendicular to the long axis of
the mark (Olsen and Shipman, 1988). A chopmark made with a metal axe has
much smoother walls, with only very faint microscopic striations. Distinguishing
between chopmarks made with very thin ground stone axes and those made with
metal axes is difficult, however, since both cuts have relatively smooth walls. A

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Table 4.™ Perforations in the ends of elements for marrow extraction.

Bag number

Taxon

Element

End perforated

0238

Ovis aries

radius

distal

0912

O vis /Capra

radius

both

0912

Ovis/Capra

radius

distal

0912

Ovis/Capra

metacarpal

both

0912

Ovis/Capra

tibia

distal

0912

Ovis/Capra

metatarsal

proximal

0912

Ovis/Capra

first phalanx

proximal

0916

Ovis/Capra

radius

distal

0916

Ovis/Capra

tibia

distal

5311-11

Ovis/Capra

radius

proximal

few thin ground stone axes were found at West Row Fen, so it is possible that
these were used to chop through bones. The West Row people apparently used
either ground stone or metal axes to hack through the bases of hom cores (Fig.
12) and tough joints of cattle, as well as deer antler beams and tines.

The use of unretouched stone blades for cutting meat and metal axes for chop-
ping through hom and bone seems to indicate efficiency. Metal axes produce
greater results with fewer blows than chipped stone axes; unretouched blades are
sharper, more easily obtained, and easier to manufacture than metal blades.

2 CM

Fig. 13.-— Perforations made in the ends of caprine long bones to extract marrow: a, proximal radius;
b, proximal metacarpal; c, proximal metatarsal.

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Fig. 14.— Posterior surface of a caprine radius with a rectangular opening for marrow extraction.

Marrow Extraction

Given the intensive carnivore gnawing of long bones at West Row Fen, it is
difficult to assess the frequency of marrow extraction by humans based on spiral
or “green bone” fractures. There are other types of modifications that were per-
formed by humans for this purpose, however. Long bones of sheep or goats, were
perforated at one or both ends (Table 4) in order to remove the marrow. In these
cases, openings were made with a gouge and then reamed or enlarged by scraping
around the interior margins until they measured from 4 to 14 mm in diameter
(Fig. 1 3). Experimentation on the articular ends of fresh bones showed that this
can be done easily without leaving noticeable tool marks around the rim or wall
of the perforation. In three specimens the epiphysis was removed from the di-
aphysis so that the soft metaphyseal material could be reamed out. In one case,
the intact epiphysis was found near the perforated diaphysis in the deposit. Marrow
was also extracted from a sheep or goat radius by incising a rectangular opening
in the posterior surface of the diaphysis near the proximal end (Fig. 14).

Two proximal cow phalanges were opened to extract marrow from the cavity
by percussion with a chipped stone hammer (Fig. 1 5). The two phalanges were
found together and were probably derived from the same individual.

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1 CM

Fig. 15.— Proximal phalanx of a domestic cow opened with a hammerstone to extract marrow (dorsal
surface).

Burning

Only five identifiable bones were found to have been burned (Table 3). Most
burning does not occur during cooking, because meat is an excellent insulator,
but instead takes place after the meal when the bones are discarded in the fire.
Such disposal reduces the odor and unsanitary conditions caused by rotting meat
and marrow, while providing some fuel for the fire. The prevalence of domestic
dogs in the village (as indicated by their remains and the incidence of carnivore
gnawing) may have led to the practice of throwing the bones to the dogs, rather
than discarding them in the fire. Moreover, the nearby woodlands would have
yielded far superior sources of fuel than that provided by bones.

Pathology

Only a few cases of pathologically altered bones or teeth were observed in this
faunal assemblage. Most were congenital anomalies, healed injuries typically as-
sociated with domestication, or changes probably caused by traction. The scarcity
of pathologies suggests that the domestic animal populations were healthy, but
also that many individuals did not avoid culling long enough to develop serious
diseases or to mend from an injury. There were no fetal and few neonatal remains
recovered, but dogs may have destroyed many of these fragile bones.

The Mammalian Fauna

Erinaceus europaeus, Western Hedgehog. — Three hind limb elements (a femur
and two tibiae) were identified. Hedgehogs, known at least by the early Mesolithic

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in Britain at the site of Thatcham, Berkshire (Davis, 1987:174), occur both in
woodlands and grasslands (Corbet and Ovenden, 1980:120).

Talpa europaea, Northern Mole.— The fused sacrum and innominates of a
northern mole were recovered from the area containing the Neolithic pits. Because
of this species’ habit of burrowing, the mole may have entered the site either
during or after its human occupation. They presently occur in British grasslands
and deciduous woodlands (Corbet and Ovenden, 1980:122).

Lepus, cf L. capensis, Brown Hare. — Nineteen bones of hares were identified
in this assemblage. Rabbits, Oryctolagus cuniculus, apparently did not spread
through Britain until the Norman Invasion (A.D. 1066) (Corbet and Southern,
1977). The remains identified at West Row Fen have proportions comparable to
Lepus rather than to Oryctolagus. Lepus capensis occurs in the area today, but it
is possible that another species could have occupied Britain in prehistoric times
(Grant, 1984). The blue hare, L. timidus was recorded in the early postglacial
from Hartledale (Tinsley, with Grigson, 1981:218). At West Row Fen, no cranial
remains were recovered and the partial mandible did not allow identification to
species level. The MNI for hares is only three, but dogs could have destroyed
many of the scraps left from meals of this small game.

Order Rodentia, Genus and Species Indeterminate. —Only a few rodents were
found in the general faunal assemblage identified here. Most of the rodent elements
were recovered from the sieved material and are currently being studied by T. P.
O’Connor. The paucity of rodent bones may be due in part to the activities of
village dogs.

?Arvicola terrestris, Northern Water Vole. — This species is normally associated
with freshwater marshes, lakes, or slow-moving streams; however, it may live in
grasslands away from water (Corbet and Ovenden, 1980:164). Although water
voles could have been consumed by humans, it is likely that they entered the
archaeological deposits intrusively at the time the peat was forming. Eleven of
the 14 bones, probably all from the same immature individual, came from a single
pit.

Felis sylvestris, European Wild Cat. — One tooth and three limb bones were
recovered. These elements are difficult to distinguish from the domestic cat, F.
domesticus, but there is no evidence that domestic cats entered Britain before the
Iron Age. A cluster of kitten skeletons found at Gussage All Saints, an Iron Age
site in Dorset occupied around 500 b.c. (uncalibrated), was used to establish
domestication (Harcourt, 1979). The relatively large size of the radius and femur
from West Row Fen supports their identification as wild cat rather than a domestic
breed. Wild cats may have lurked around middens at night to catch rats and mice,
although dogs would certainly have helped deter this behavior. Alternatively, wild
cats may have been hunted for their fine fur to make small bags or garments.

Vulpes vulpes, Red Fox.—O ne lower jaw and nine limb bones were found
strewn across a small area and could represent one adult animal. None of the
bones has unfused epiphyses, and the age of fusion for the identified bones ranged
from 19 to 28 weeks (Davis, 1987). Foxes might have been killed because they
were pests and a threat to the young livestock, as foxes scavenge sheep and goat
carcasses (Stallibrass, 1984) and could kill a newborn lamb or kid.

Two cases of butcher marks on fox bones are almost certainly the result of
skinning. The first consists of several short, shallow cuts on the lateral side of the
shaft of a fifth metacarpal (bag 0882) just distal to the proximal articular surface.
These probably occurred when an annular incision was made around the wrist to
remove the hide. The second mark was found on the anterior surface of a distal

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tibia (bag 0933) and could represent an analogous cut around the ankle (Guilday
et al., 1962:71). The removal of hides frequently stops short of including the
strongly adhering skin of the feet and toes since this skin is of little value. Annular
cuts at the wrist and ankle allow the hide to be removed in one piece, exclusive
of the feet.

Canis familiaris, Domestic Dog. — The quantity of gnawed bones in this assem-
blage indicates that dogs were adept at cleaning up the food refuse produced by
the village. Hunting was minimal at West Row Fen in comparison to the rearing
of domestic stock, although dogs could have been trained to assist in periodic
hunts. It is more likely that dogs were used to herd and protect livestock from
predators such as foxes and wild cats. The dogs were medium-sized and the
cranium and mandibles show characteristics of domestication such as foreshort-
ening of the rostrum (Fig. 1 6). Domestic dogs are known in Britain since Mesolithic
times at Star Carr (7538-7607 b.c., uncalibrated) (Degerbol, 1961).

The dentition and epiphyseal fusion data on dogs show that most of the remains
preserved are from adults. The only exceptions are a mandible containing a
deciduous second premolar (aged five weeks to six months; Silver, 1969) and a
radius that was unfused distally (aged less than 47 weeks; Silver, 1969). One
cervical, probably from an old individual, bears traces of possible osteoarthritis
in the form of lipping around the cranial end of the centrum on the ventral surface.

Approximately 63% of the Canis material represents the skull and first two
vertebrae, whereas the remaining 37% is mostly appendicular. Two atlas vertebrae
(bags 5382 and 10498) have cutmarks on the ventral surfaces associated with
decapitation. Diagonal marks immediately adjacent to the right anterior articular
facet of one and transverse marks on the body of the other (Fig. 17) were most
likely made while removing the head from the body. Another cervical (bag 4237)
also bears transverse cutmarks on its dorsal surface. Deer, sheep, pigs, and cattle
were also decapitated as a normal part of the butchering process at West Row
Fen; therefore, no sacrificial or ceremonial inferences need to be made in the case
of dogs.

The anterior surfaces of two dog radii show fine transverse cuts near the proximal
ends, as does the posterior surface of a tibia. Deep cuts like these at the elbow
and behind the knee suggest disarticulation of the carcass as a part of food prep-
aration rather than skinning (Guilday et al., 1962:67). The internal surface of one
rib was also butchered in a way consistent with food preparation or consumption.

Cunliffe (1991) reported a high frequency of butchered dog bones at the Iron
Age site of Highfield. At West Row Fen, the lack of dog burials, the presence of
butchering marks, the scattering of their bones throughout the site, the deposition
of a partial skeleton and isolated bones in refuse pits (Table 1), and the occurrence
of one spirally-fractured and one burnt dog bone suggest that these animals were
probably eaten and were not particularly revered as pets.

Capreolus capreolus, Roe Deer. — Roe deer were present in Britain intermittently
during the Pleistocene interstadials and continuously after the Preboreal (ca. 8300
b.c.), when forests replaced grasslands. They appear in prehistoric sites in or near
forests from the Mesolithic on, but their numbers generally decline in the Neolithic
as domestic stock becomes increasingly more important (Grigson, 1984).

At West Row Fen, roe deer element frequencies do not vary dramatically, which
is compatible with the idea that these small animals were probably carried into
the site whole. The sample is too small to construct a mortality pattern, but
epiphyseal fusion and dental evidence show that adults were taken more frequently
than juveniles. Roe deer would have provided about the same amount of meat

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5 CM

Fig. 16. — Domestic dog cranium from West Row Fen: a, dorsal view; b, basicranial view.

as the small Soay sheep, but, as wild game, would not have yielded milk and
wool. Roe deer rank seventh at West Row in the amount of contributed meat
(Table 2).

Roe deer antlers, because of their small size, spatulate shape, thin cortical layer,
and predominantly spongy interior, were much less useful for manufacturing
artifacts than were the antlers of red deer. As far as can be discerned, all of the

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5 C M

Fig. 17.— Ventral view of a dog atlas, illustrating locations of cutmarks.

worked antlers from West Row Fen are derived from red deer. One of the un-
modified roe deer antlers, however, was collected after it was cast by the deer.

Cutmarks on the medial shaft of one roe deer metacarpal imply skinning. The
distal epicondyle of a humerus bears several annular cutmarks, like those usually
associated with the removal of the forearm flexors and extensors. Two astragali
and one calcaneous exhibit transverse cuts that represent disarticulation of the
ankle. All of these are common butcher marks made in the preparation of a
medium or large mammal for consumption.

Cervus elaphus, Red Deer. — Red deer have a paleontological distribution pat-
tern similar to that of roe deer— flourishing in the Pleistocene interstadials and
returning with the forests of the Holocene. They, too, have been identified in sites
that date back to the Preboreal (Davis, 1987). At West Row Fen, red deer remains
were more plentiful than those of roe deer according to their NISP, but less
common in their MNIs. Red deer yield about nine times as much meat as roe
deer, however, and have antlers that are far more useful for manufacturing arti-
facts. By weight of usable meat, the red deer is the third most important species
consumed at West Row Fen.

The one definite piece of seasonality information comes from an antler still
attached to the frontal bone (bag 5349) of a deer that was killed between September
and March. Four red deer antlers were chopped through the beam just above the
pedicle, suggesting that they were cut from the frontal during butchering. The
most remarkable of these was found in Pit 4284 and is 42 cm long. It had been
split in half longitudinally by chopping with a metal axe in preparation for artifact
manufacture. Three cast antlers demonstrate the apparent need at West Row Fen
to gather this valuable raw material for artifact manufacture. Two of the cast
antlers were made into mace-heads. Additional antler artifacts include four rod-
like flakers, eight modified tines, a wedge, and a hollow handle.

That red deer provided meat and hides as well as antlers for making artifacts
is evident from the butchering patterns. The atlas of one red deer bears several
long cutmarks on the ventral surface reflecting decapitation. The anteromedial
surfaces of two radii exhibit fine cuts near the proximal ends, inflicted when the
elbow joint was disarticulated. Finally, a proximal phalanx was cut on the side
just above the distal condyle during skinning.

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Table 5. —Survival based on epiphyseal fusion for Sus scrofa. Ages of fusion based on Silver, 1969:252-

253.

Age in months

Element

End8

Left

Unfused

Right

Left

Fused

Right

% Surviving11

humerus

radius

tibia

femur

radius

tibia

femur

humerus

a P = proximal, D = distal.
b Surviving beyond age of fusion.

All red deer bones retaining epiphyseal areas are fused. Identified teeth include
one left and right deciduous premolar and 28 permanent cheek teeth. There is
little evidence that juvenile red deer were hunted with any frequency.

Sus scrofa, Pig. — Wild boars, like the roe and red deer, invaded Britain once
the forests began to dominate and were present from the Preboreal onward. The
earliest evidence for small, apparently domestic, pigs is derived from the Neolithic
causewayed camp at Windmill Hill (ca. 2960 b.c., uncalibrated; Davis, 1987:177).

Osteological features alone do not reveal whether wild boars were hunted oc-
casionally by West Row people. Two large adult pig bones— a temporal and an
ulna— were recovered, but since fully adult domestic pig bones are rare in this
collection, the maximum size of a typical domestic male cannot be estimated.
Milisauskas (1978:67) estimates a significant difference between the amount of
usable meat produced by an average wild boar (53.75 kg) compared to an early
domestic pig (15 kg).

Most of the fused epiphyses were those that fuse at birth or within the first two
years of life (Table 5). Those that fuse between 2 and 3.5 years were always unfused.
The quantity of immature bones argues that most, if not all, of the pigs at West
Row Fen were domesticated and that culling of young individuals occurred.

Based on the teeth (Table 6), a slightly different picture of the age structure of
pigs at West Row Fen emerges. Bull and Payne (1982) demonstrated that there
are only minor differences between wild boars and modem domestic pigs in the
timing of tooth eruption and epiphyseal fusion. Therefore, the age categories for
wild boars have been applied to the dentition of Early Bronze Age domestic pigs.
Wild boars mature only slightly quicker than modem domestic pigs, which have
undergone far more artificial selection than the Bronze Age pigs. The dental
evidence from West Row Fen shows that at least some individuals survived long
enough to reach full size. Based on the right mandibles, the most common jaws
in the assemblage, 45% of the pigs were being killed in the first 16 months of life.
Another 36% died between 19 and 35 months, and 18% survived to about four
years or beyond. This pattern suggests that while the mortality rate was heavy in
the first year, at least some pigs were reserved for breeding and were kept to grow
until they were needed for their meat. It was uncommon, however, for a pig to
survive beyond three years.

The average dressed weight of a mature European wild boar is about 36.4 kg,
but it is just 13.6 kg for a wild piglet (Henry, 1969; Biddick, 1984:165). Assuming
comparable ratios for early domestic pigs in Britain, the mortality patterns bear

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Table 6.— Age distribution of Sus scrofa based on mandibles with two or more teeth. Aging of dentition
done by using Silver (1969) for dates of eruption and Bull and Payne (1982) for wear stages.

Age category

Left

Right

1 week to 1 6 months

7 weeks to 1 6 months

4 months to 16 months

7 months to 16 months

8 months to 16 months

19 months to 23 months

19 months to 35 months

31 months to 35 months

Over 35 months

Total

on the relative importance of pork in the diet of the West Row people. Although
some individuals were kept to adulthood, most were slaughtered before full growth
had been achieved. This would considerably lower their relative dietary contri-
bution compared to cattle and red deer, which were primarily slaughtered as
adults. However, pork has a higher caloric value than beef, venison, or mutton.
Per kilogram, mutton provides 1500 calories; beef, 2000 calories; and pork, 3700
calories (Flannery, 1969). If calories are considered over pure units of meat weight,
pigs would again increase in relative value, although their caloric value would
depend in part on whether juveniles have the same proportion of fat as adults.

Butchering marks are not very common on pig bones (Fig. 18; Table 7) except
on the distal epicondyles of humeri. Marks on a lacrimal and on the lingual surface
of a mandible (not figured) illustrate that skin and meat were removed from the
head. One occipital bears traces of decapitation. Cutmarks on a scapula, humeri,
an innominate, and a femur were probably made while removing meat and dis-
articulating the joints. Ribs were cut away from the vertebrae in the process of
disarticulating the carcass. Ribs were also scraped on the internal surfaces probably
during human consumption of the intercostal muscles. Removal of the hock is
indicated by cuts on the astragalus, calcaneus, and navicular bones. One pubis
(not figured) bears a cut probably made while skinning or eviscerating the animal.

Live pigs do not produce useful products such as wool and milk, although their
manure may be used for fuel or fertilizer. One possible benefit of raising pigs is
that they root up the soil thus serving the same function as a plow (Reynolds,
1976).

Although they may have been supplied with grain and legumes occasionally,
pigs could have survived primarily on garbage and pannage provided by nearby
forests. Grass and bracken would have been available locally in the spring and
summer. Stubble left in pastures grazed by sheep and cattle was another possible
food source for swine. If pigs were kept primarily for their meat and lard, their
numbers would probably have been maintained below the level that would require
much provisioning with grain that people could have consumed. Though pigs are
inexpensive to feed, if their numbers get too large, they begin to become a nuisance.
Too many pigs are difficult to manage because they tend to get into gardens and
destroy food intended for people (Rappaport, 1968). Because a female produces
eight to ten piglets at a time (Towne and Wentworth, 1950:253), some culling
would be necessary to prevent overpopulation.

Ovis aries/Capra hirca, Domestic Sheep or Goat. — Sheep and goats were brought

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Fig. 18. —Distribution of butcher marks on pig bones.

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Table 1 .—Key to butchering patterns for Sus scrofa.

Location and type

Number of bones cut

1. Skinning

2. Decapitation

3. Disarticulation of shoulder

4. Muscle removal

5. Disarticulation of elbow joint

6. Muscle removal

7. Segmentation of thorax

8. Disarticulation of hip

9. Muscle removal

1 0. Severing Achilles tendon

1 1 . Disarticulation of ankle

12. Disarticulation of ankle

into Britain across the English Channel during the Neolithic (Ryder, 1983:64-65)
and increased in numbers compared to other livestock in the Bronze Age as the
forests diminished (Trow-Smith, 1957). By NISP, caprines were second in fre-
quency to cattle at West Row Fen, but the MNI for caprines was higher than for
cattle (Table 2). Since it takes the meat of 28 sheep or goats to match that of a
cow, however, it is clear that far more pounds of beef were produced in the village
than mutton or goat meat.

Of the six bones identified as those of domestic goats, three were horn cores.
The breed of these goats is not discemable; they appear to have been small and
gracile like the sheep, but are far less common. Sheep thrive much better in cool,
wet lowland farming communities, whereas goats prefer hotter, more arid climates
(Redding, 1984:237) and rugged terrain. According to Grant (1984), sheep were
also much more plentiful than goats during the Iron Age in Britain. Mixed herds
of sheep and goats are common among livestock herders, partly because goats
are thought to improve herd management and to provide leadership for the sheep,
although this has never been scientifically documented (Redding, 1984:29).

The sheep from West Row Fen were small, slender-limbed, two- horned indi-
viduals that are within the size range of the Soay breed. According to Reynolds
(1987), Soay sheep probably developed as a breed during the Bronze Age and
were common until the Iron Age, when four-homed breeds like the Hebridian
and Manx Loghtan breeds arose.

Because of their fragmentary nature, most of the bones in this size range could
only be identified as caprines (Ovis aries/Capra hired). Although the better pre-
served material hints that sheep considerably outnumbered goats, we cannot make
that assumption for the bulk of the less-identifiable material.

Survival patterns reflected by epiphyseal fusion for Ovis/Capra indicate that
there was a serious drop in the proportion of individuals living beyond the first
year (Table 8). At ten months, over 90% were still alive, but by 13 months only
half to a third remained. By 3 to 3.5 years, only a third to a quarter survived.
This pattern is similar to the one seen in the pig dentition records and suggests
that culling of young individuals took place. In this case, the reason for killing
juveniles is probably more than just a desire for the meat, although lamb is
preferable to mutton. Cultures relying on sheep and goat dairy products need to
kill enough of the young to have a surplus of milk for human consumption (Davis,

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Table 8 .—Survival based on epiphyseal fusion for Ovis aries/Capra hirca. Ages of fusion based on
Silver, 1969:252-253, except for phalanges. At West Row the proximal epiphyses fused after the distal
epiphyses in cattle and caprines, counter to Silver.

Age in months

Element

End*

Left

Unfused

Right

Left

Fused

Right

Surviving1 2 3 * * * * * * * * * * * 15

humerus

radius

13-16?

phalanx 1

13-16?

phalanx 2

18-24

tibia

18-24

metacarpal

20-28

metatarsal

30-36

femur

radius

36-42

tibia

36-42

femur

36^12

humerus

a P = proximal, D = distal.
b Surviving beyond age of fusion.

1987:180; Legge, 1992). Males are usually killed in their first year, leaving only
enough to breed with the females. Many of the females are allowed to mature
beyond the age of two or three years in order to reproduce and yield milk.

Determined age categories based on caprine mandibles (Table 9) are quite broad
and do not often coincide because tooth rows and comparable teeth are poorly
preserved from one mandible to the next. Most mandibles preserve only two or
occasionally three teeth, but molar alveoli provided additional evidence for aging
jaws. Despite these constraints, the data indicate that at least two individuals were
killed between the ages of two and three months. Nineteen died before they were
two years of age; eight survived past the age of three years, seven of those past
5.5 years, and one beyond eight years. This evidence indicates that many indi-

Table 9 .—Age distribution of Ovis aries/Capra hirca based on mandibles with two or more teeth. Age

based on Silver (1969) and Legge (1992).

Age in months Left Right

1- 24 2

2- 3 2

3 1

>3 1

3- 10 1

3-24 2 7

9-12 3

>18 1

18-24 1

<24 4 5

>21 2 4

21-24 2

21-34 1

10-65 1

34-65 1 1

65-100 5 6

>100 1

Total 22 32

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141

1 CM

Fig. 19.— Pathological exostosis on a proximal phalanx of a caprine. Arrows point to abnormal flanges
of bone on both sides.

viduals were being maintained well beyond the age when mature weight is achieved,
which suggests that their primary function was dairy production rather than meat
provisioning. The scant dental data do not reveal whether very young lambs and
kids were regularly culled, as would be expected if people were utilizing dairy
products.

Unfortunately, there is no evidence for the use of sheep wool at West Row Fen,
since no textiles or antler combs suitable for plucking fleece have been found at
the site. Sheep shears are not known until Roman times (Reynolds, 1987). How-
ever, Bronze Age textiles made from Soay-type wool have been found in England
(Ryder, 1983:47). A sample from Rylston, in Yorkshire, consists of a piece of
yarn made of fine fibers associated with pieces of a generalized medium and a
hairy medium wool (Ryder, 1969).

The only pathology among the caprine bones is a proximal phalanx that has
indentations on both sides of the distal epicondyle with lipping above (Fig. 1 9).
The unusual symmetry of these injuries suggests that they may have been induced
by a human-made trap or a hobble.

Butcher marks are fairly common on the bones of caprines at West Row Fen
(Fig. 20; Table 10). In contrast to the West Row cattle, the horn cores and frontals
of sheep and goats do not exhibit chopmarks or cutmarks indicative of the use
of their horns as a raw material. Only one chopmark was observed on a caprine
bone, in contrast to many on bones of cattle. Skinning marks were found on the
maxilla, anterior surfaces of the metapodials, and proximal phalanges of caprines.
One hyoid bore traces of cutmarks made when the tongue was removed. The
heads of sheep were removed by cutting between the occipital and the atlas, leaving
small cutmarks on an occipital and both dorsal and ventral surfaces of an atlas.
Disarticulation cuts were found on caprine cervical vertebrae; at the shoulder on

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Fig. 20. — Distribution of butcher marks on caprine bones.

the scapular neck and humeral head; the elbow joint on the humerus, radius, and
ulna; the distal radius and the scaphoid of the wrist; above the acetabulum of the
innominate; and at the ankle on the astragalus, navicular, and calcaneus. Some
fileting and other types of meat removal are indicated by midshaft cuts or scraping
on the scapula, humerus, radius, femur, and tibia.

To determine how the vertebrae and ribs were butchered, it is necessary to
examine the bones that could only be identified as small ruminants, since it is
difficult to distinguish among fragments from sheep, goats, and roe deer. These
show minor cutmarks on the neural spines of the thoracic vertebrae, a chopmark
completely through a thoracic centrum and neural arch (probably with a metal
axe), disarticulation of the ribs and vertebrae at the rib’s head and tubercle, and
scraping of the internal surface of the rib. All of these cutmarks are commonly
found on carcasses butchered for food consumption.

Bos primigenius, Wild Aurochs. — Most of the Bos remains represent domestic
cattle. Although a few of the elements are large, only one is attributable to the
wild aurochs— a distal humerus with a trochlear breadth of 9.9 cm and proportions
comparable to that of a bison.

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Table 10 .—Key to butchering patterns for Ovis/Capra.

Number of bones

Location and type

Cut Chopped Scraped

Skinning

Muscle removal

Tongue removal

Decapitation

Segmentation of neck

Disarticulating shoulder

10.

Muscle removal

11.

Disarticulating shoulder

12.

Muscle removal

13.

Disarticulating elbow

14.

Disarticulating elbow

15.

Disarticulating elbow

16.

Disarticulating elbow

17.

Disarticulating elbow

18.

Fileting upper forelimb

19.

Fileting upper forelimb

20.

Disarticulating wrist

1 1

21.

Disarticulating wrist

22.

Disarticulating wrist

23.

Skinning, tendon removal, or

disarticulating of joint

24.

Same as 23

25.

Same as 23

26.

Disarticulating hip

27.

Eviscerating or skinning

28.

Muscle removal

29.

Muscle removal

30.

Muscle removal

31.

Muscle removal

32.

Fileting lower leg

33.

Disarticulating ankle

34.

Severing Achilles tendon

35.

Disarticulating ankle

36.

Disarticulating ankle

37.

Skinning

38.

Skinning, tendon removal, or

disarticulation of metatarsal-
phalangeal joint

1 1

The wild aurochs appears to have become extinct in Britain during the Bronze
Age (Tinsley, with Grigson, 1981), but its remains have been identified in mixed
Beaker and Early Bronze Age deposits at Snail Down; Early Bronze Age levels at
Lowes Farm near Littleport, Cambridgeshire; and at nearby County Farm, Mil-

denhall Fen (Tinsley, with Grigson, 1981:219). A whole skeleton was recovered
from Early Bronze Age levels at Charterhouse Warren Farm, Blagdon, in Somerset
(1295 b.c., uncalibrated or 1629 B.C., calibrated — Everton, 1975; Clutton-Brock
and Burleigh, 1983).

Bos taurus, Domestic Cattle. — Cattle are well-suited to the lowlands of the fens,
where water is in good supply. They can be used for draft, and yield dairy products,
meat, large hides, bones, and horn. However, cattle require considerable pasture

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Table 1 1 .—Survival based on epiphyseal fusion for Bos taurus. Ages of fusion based on Silver (1969),
except for phalanges. At West Row the proximal epiphyses fused after the distal epiphyses in cattle and

caprines, counter to Silver.

Age in months

Element

End0

Left

Unfused

Right

Left

Fused

Right

% Surviving*1

12-18

humerus

12-18

radius

18?

phalanx 1

18?

phalanx 2

24-30

tibia

24-30

metacarpal

27-36

metatarsal

femur

42—48

radius

42-48

tibia

42-48

humerus

a P = proximal, D = distal.
b Surviving beyond age of fusion.

and keeping them reduces the mobility of their owners, which means less op-
portunity to move to fresh pastures as needed.

The Bronze Age witnessed the development or arrival of a small, short-homed
variety of cattle in Britain, which was apparently represented at West Row Fen.
The two most-complete horn cores have basal circumferences (180 and 195 mm)
well within the range of either Neolithic domestic males or wild females, but the
reconstructed length of the outer curvature (roughly 200 to 250 mm) is only about
half that for Neolithic domestic males and wild females (Grigson, 1982), suggesting
that they were derived from domestic short-homed males.

At West Row Fen, most of the limb bones of domestic cattle are very slender,
but few could be measured due to their comminuted state and the extent of
carnivore gnawing on condyles. With the exception of the enormous aurochs
humerus, the few large specimens may represent the limited number of intact
bullocks allowed to reach adulthood. The data from the epiphyseal fusion of cattle
long bones (Table 1 1) indicate that most individuals survived past the age when
full growth was obtained. The 90% representation of fused distal tibiae may be
an accident of preservation, but it still appears that over 50% of the cattle survived
beyond 3.5 years of age. Immature bones are less likely to be preserved, given
the activity of dogs and sedimentary abrasion from trampling. However, pro-
portionally far more immature caprine and pig bones are preserved in the assem-
blage, and they are smaller and more fragile than those of immature cattle.

Cattle mandibles are only slightly better preserved than those of caprines. For
the most part, they contain only two to three teeth, and no complete tooth rows
are preserved. The largest age group represented by mandibles (Table 12) is in
the six- to eight-year bracket and there are more minimum numbers of individuals
in the full adult categories than for young juveniles. The age distribution does not
imply culling of individuals in the first year of life, suggesting that cattle were
kept primarily for meat and draft rather than for their dairy products (Legge,
1992). The high number of survivors beyond six years also argues in favor of
their use for pulling plows and carts.

Several pathologies and anomalies were noted on cattle teeth and bones. The
mandible of an individual over 28 months in age (bag 0901) shows the absence
of the second permanent premolar, with no sign of infection or injury. The absence

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Table 12.— Age distribution o/Bos taurus based on mandibles with two or more teeth. Age based on

Legge, 1992.

Age in months

Left

Right

MNI

% of total MNI

1-3

3=6

6=15

15=26

26-36

36=72

72=96

Total

of teeth is an anomaly found in many domestic species, including cattle (Baker
and Broth well, 1 980: 1 37). Another mandible of an individual older than 24 months
(bag 5285-11) exhibits malocclusion and unusual wear patterns. An isolated third
lower molar (bag 0297) is worn in an uncharacteristically jagged pattern, appar-
ently from malocclusion. A smaller than usual postyzygopophysis, apparently a
congenital anomaly, occurs on one thoracic vertebra (bag 04 1 8). Exostosis asso-
ciated with either an injury or osteoarthritis caused lipping of the semilunar notch
on one adult ulna (bag 5279-11). Baker and Broth well (1 980: 1 1 5) reported a similar
pathology from the site of Crandon Bridge.

Importantly, one innominate (bag 5285-11) has a swelling on the ischium in-
volving the acetabular rim and the area just lateral to it (Fig. 21). Baker and
Brothwell (1980) reported osteoarthritis on a bovine acetabulum from the Mote
of Mark, and Baker has observed many examples of ebumation of the acetabulum
of cattle from Roman to Medieval deposits at Winchester (Brothwell, 1981). These
pathologies have been interpreted as representing excessive strain on the hip joints
due to an activity that leads to overrotation of the femoral head, such as pulling
a plow or a heavy cart.

Two adult proximal phalanges of cattle (bags 5 1 0 1 -II and 5 1 1 8 -II) show a heavy
development of the tuberosities on the volar surface (Fig. 22), which may indicate
that these animals were used for draft. The palmar (plantar in the hind foot) anular
ligament spans over the tendons of the digital flexors (Getty, 1975:858-59), bind-
ing them against the bone, and attaches to the tuberosities on the volar surface
of the proximal phalanx. Heavily developed tuberosities could indicate greater
than usual stress on these flexor tendons, which insert on the mesial and distal
phalanges. Exostosis development in third phalanges related to stress at the point
of insertion for the flexor tendons has been associated with plowing in water
buffalo and cattle (Higham et ah, 1981).

The pathologies of cattle bones and teeth from West Row Fen are those often
associated with domestication. Malocclusion and the anomalous absence of teeth
are frequently related to genetic changes brought about by breeding, like the
foreshortening of the rostrum. Exostosis caused by injury, strain, or old age is
more common in domestic animals because they are provisioned and protected
despite their impaired movement. The exostosis on the ulna could have been
caused by the additional stress placed on the elbow joint while plowing or pulling
heavy loads. Pathologies of the bones of the hips and feet in European prehistoric
domestic cattle are more common than those seen in sheep and pig remains, and

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Fig. 21.— Innominate of Bos taurus exhibiting a swelling on the ischium involving the acetabular rim
and area lateral to it: a, anterior view; b, lateral view.

have been attributed to the impact of stress and strain in cattle due to their use
for draft (Baker and Broth well 1980:1 17).

The butchering patterns for cattle (Fig. 23, Table 13) are similar to those for
sheep and goats with two exceptions. Horn cores and the frontal bones around
the cores were often chopped with an axe when removing the horn (Fig. 12).
These numerous, heavy marks demonstrate that cow horn was a valued resource
for the manufacture of artifacts or glue.

The second difference involves the frequency of heavy chopping compared to
fine cutmarks. Chopmarks are rare on the bones of small ungulates, but are much
more common on the elements of cattle, which reflects the thicker ligaments and
tendons as well as the heavier bones found in cattle.

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147

2 CM

Fig. 22.— Proximal phalanges of oxen showing hypertrophy of the tuberosities for the palmar (or
plantar) anular ligament, perhaps resulting from strain inflicted on the digital flexors during plowing
or while pulling a heavy cart.

Possible skinning marks occur on the nasal, maxilla, and mandible, as well as
on a metacarpal, and first and second phalanges. One hyoid bone bears two
cutmarks made during removal of the tongue. Both atlas and axis vertebrae exhibit
signs of butchery associated with decapitation. Disarticulation cuts were made
between cervical vertebrae in the neck, on the mandible and malar to remove the
lower jaw, and at the shoulder, elbow, wrist, hip, and ankle. Meat was removed
from the bone of the mandible, scapula, humerus, radius, innominate, femur, and
tibia. The most thorough fileting was performed on scapulae, which were often
scraped on both surfaces and along the spine. The attention given to this element
suggests that the shoulder roast was one of the most preferred cuts of meat. Ribs
of large ungulates (probably cattle) were disarticulated from the vertebrae, leaving
cutmarks or chopmarks on the rib heads and tubercles.

Cattle at West Row Fen were used for meat and draft, based on evidence from
their butchery, mortality pattern, and pathologies. In terms of the quantity of
identified specimens and meat poundage, cattle outrank all other species of animals
consumed by the people at West Row Fen. The mortality pattern does not suggest
that cattle were kept primarily for their dairy products, although it is unlikely that
this resource was completely overlooked. Cow horn was an important by-product,
judging from the number of chopped horn cores and frontals.

Equus caballus, Horse. —Wild horses inhabited Britain throughout the Pleis-
tocene and into the Holocene, but never crossed over to Ireland. Small numbers

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Fig. 23. — Distribution of butcher marks on cattle bones.

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149

Table 13 .—Key to butchering patterns for Bos tauirus.

Number of bones

Location and type Cut Chopped Scraped

1 . Horn removal

2. Skinning

3. Skinning

4. Skinning

5. Disarticulating mandible

6. Muscle removal

7. Skinning

8. Tongue removal

9. Decapitation

10. Decapitation

1 1 . Segmenting neck

1 2. Fileting shoulder

13. Fileting shoulder

14. Disarticulating shoulder

15. Muscle removal

16. Muscle removal

17. Disarticulating elbow

18. Disarticulating elbow

19. Disarticulating elbow

20. Fileting upper forelimb

2 1 . Disarticulating wrist

22. Disarticulating wrist

23. Skinning

24. Skinning, tendon removal, or

disarticulating joint

25. Same as 24

26. Muscle removal

27. Segmenting pelvis

28. Disarticulating hip

29. Disarticulating hip

30. Disarticulating hip

3 1 . Muscle removal

32. Muscle removal

33. Severing Achilles tendon

34. Disarticulating ankle

35. Disarticulating ankle

36. Disarticulating ankle

Not shown: Marks on subspinous fossa

of scapula from fileting shoulder

of horse remains have been found in Mesolithic, Neolithic, and Early Bronze Age
sites in Britain. It is unclear how early domestic horses arrived in Britain, but
evidence at Newgrange, in Ireland, demonstrates that they were introduced there
by the time of the Beaker Culture (Wijngaarden-Bakker, 1974). Whether these
early domestic horses were ridden is unknown, because bronze mouth bits and
antler cheek pieces from harnesses do not appear until the Late Bronze Age at
sites such as Runnymede in Britain (Longley, 1980) and Newgrange in Ireland
(Burgess, 1974). Based on the scant osteological evidence, it is not possible to say
whether the horses from West Row Fen were wild or domesticated.

Just seven teeth and eight bone fragments were identified as horse. Two of the
teeth are deciduous premolars and one cervical is unfused at the caudal epiphyseal

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plate, representing the remains of at least one immature horse. The other teeth
(a worn lower third incisor and a lower third molar) belong to an adult.

The distal epicondyle of an adult horse metapodial was butchered. Multiple
transverse cuts on either the medial or lateral surface of this bone are probably
indicative of skinning rather than disarticulation, because they are well above the
joint. There is no evidence that horses were eaten at West Row Fen, but the single
butchered bone suggests that their hides may have been utilized.

Bone and Antler Artifacts

Of the collection of 120 bone and antler artifacts recovered from West Row
Fen, 76 are bone awls made from sheep or goat bones. Twelve of these are drilled
near the base for sewing or for suspension. A pin, a scoop, and a spatula have
also been identified. Ornaments are restricted to three simple, undecorated tubular
beads. The most spectacular piece is a well-made and finely polished bone dagger
with a tanged handle that may have been a skeuomorph of a more effective bronze
weapon.

Many of the artifacts were made from red deer antler, an apparently plentiful
and useful raw material. Of the twelve pressure flakers recovered, eight were
modified tines, and four were rods cut from antler beams by grooving-and-snap-
ping. A hollow antler handle with two rivet holes and two socketed mace-heads
provide clues about how pieces of composite tools were attached to one another.
A single antler wedge may have been used in conjunction with an antler mace-
head to split wood.

Eighteen pieces of debitage helped decipher manufacturing techniques. These
include a split antler (42 cm long) from the soaking pit, a severed antler tine, nine
grooved-and-snapped ruminant metapodials, as well as other small indeterminate
fragments bearing manufacturing traces.

Conclusions

West Row Fen fits into and generally supports a trend seen in smaller British
faunal assemblages from the Neolithic through the Iron Age. During the Early
Neolithic (ca. 3500 B.C. to 2750 B.C.), the Windmill Hill culture on the Salisbury
Plain depended primarily on cattle (70%), followed by pigs (17%) and sheep/goats
(13%) (Ryder, 1983:65). In the Late Neolithic, cattle and pigs occur most fre-
quently in the archaeological record. Sheep and goats were still fairly rare. Cattle
began to outnumber pigs in Beaker sites and were prevalent throughout the Bronze
Age. From the Early Bronze Age, sheep and goats increased and pigs declined in
numbers (Tinsley, with Grigson, 1981). Ryder (1983:72-73) attributes this to the
clearing of woodlands.

The Bronze Age of Britain took place during the Subboreal period, which was
marked by a decrease in precipitation that reduced productivity in farming and
regeneration of forests, but improved conditions for pastoralism in low-lying,
poorly drained areas (Ryder, 1 983:72—73) such as the fens of East Anglia. This
increase in pastoralism is probably responsible for the scarcity of Bronze Age
settlement sites in Britain, because nomadism would also have increased. Because
of the paucity of large Bronze Age sites, little is known of the relative frequencies
of domestic animals and wild game during this period. West Row Fen demon-
strates the rise of importance of caprines and the decline of pigs during the Early
Bronze Age.

1994

Olsen— Mammal Exploitation at West Row Fen

151

West Row Fen is dominated by cattle in terms of number of bone fragments
and estimated meat poundage, but the minimum number of individuals for ca-
prines (sheep/goat) is higher than for cattle (Table 2). Pigs rank third in both
numbers of fragments (NISP) and minimum numbers of individuals, but fifth in
terms of usable meat. Red deer were the most important wild species, providing
as much as a third of the meat at West Row Fen. Their antlers were also employed
in the production of a variety of useful artifacts. Roe deer are much smaller and
contributed considerably less meat, and their antlers were of little use in tool
manufacture.

At Grimes Graves (Fig. 1), in neighboring Norfolk County, the Middle Bronze
Age deposits are dominated by cattle, followed closely by caprines, with pigs far
behind in third position (Legge, 1992:16-17). However, the ratio of caprines to
pigs varies from one region to the next. The Late Bronze Age deposits of Run-
nymede (Done, 1991) produced slightly more pig than caprine bones, suggesting
that fewer forests had been cleared in this area during the Bronze Age. A small
collection of bones from Mill Pot Cave, Wetton, Staffordshire, associated with
Bronze Age pottery, yielded 76% Ovis/ Capra, 14% Bos taurus, 4% Sus scrofa, and
4% Equus caballus, based on the minimum numbers of individuals (Ryder et al.,
1971).

Most of the Iron Age sites from which there are large faunal assemblages contain
predominantly sheep, but these are in downlands that are poorly watered and
unsuitable for cattle. There are several Iron Age sites, such as Woodyates, Wood-
cuts, Longbridge Deverill (phase A), Grimthorpe Hill Fort, and Catcote, where
cattle outnumber sheep (Ryder, 1983:79).

West Row Fen, with its three known house structures and large midden, is an
important site because Bronze Age settlements of respectable size are rare in
Britain, as they are on the continent. The economy of the village, reconstructed
from the mammalian remains, implies the degree to which the villagers were
sedentary. The possible use of cattle for plowing as well as pulling carts would
imply the importance of agriculture relative to herding at West Row Fen. Although
pigs rank third in frequency among livestock, their numbers still indicate that the
community as a whole was not nomadic (Towne and Wentworth, 1950:69), al-
though shepherds may have left the village seasonally with the caprine herds. The
presence of swine suggests that some forests were present, but the greater number
of sheep indicates considerable clearing of forests for pasture. Sheep outnumber
goats in the fens at this time because of the cool, moist climate of the region and
the flat terrain. Red and roe deer are primarily forest dwellers, but their small
numbers support the data from sheep that much of the area was cleared for
agriculture and grazing. That dogs were eaten is demonstrated by the number of
butcher marks on their bones and the treatment of their remains.

Acknowledgments

I am grateful to the director of the West Row Fen excavations, Edward Martin,
of the Suffolk Archaeological Unit, and Sebastian Payne, of English Heritage, for
making this research possible. English Heritage funded the research that enabled
this report to be written. Edward Martin furnished the map and data in Fig. 2
and 3. Melinda Zeder kindly provided information for distinguishing between
sheep and goats. All of the faunal material and bone artifacts from West Row
Fen, Mildenhall 165, are the property of English Heritage, Fortress House, 23
Savile Row, London, United Kingdom W1X 2HE.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 2, Pp. 155-183

31 May 1994

TEPHRODYTES BRAS SIC ARVALIS, NEW GENUS AND SPECIES
(ANURA: PELODYTIDAE), FROM THE ARIKAREEAN CABBAGE
PATCH BEDS OF MONTANA, USA, AND
PELODYTID-PELOBATID RELATIONSHIPS

Amy C. Henrici

Scientific Preparator, Section of Vertebrate Paleontology

Abstract

Tephrodytes brassicarvalis, n. gen. and sp., is based on the remains of a nearly complete skeleton,
several partial skeletons, and some isolated bones from the Arikareean Cabbage Patch beds of western
Montana, USA. It possesses the hallmark pelodytid character of a fused tibiale and fibulare and differs
from other known pelodytids in the following unique combination of characters: 1) frontoparietal
fontanelle concealed, 2) posterior tip of frontoparietal present, 3) otic plate of squamosal present, 4)
otic ramus of squamosal reduced, 5) vertebral neural arches elongate, 6) sacral diapophyses widely
expanded, and 7) anterior lamina of scapula absent.

Because the new anuran shares similarities with both pelodytids and pelobatids, and because the
pelodytids have been included in the Pelobatidae, a cladistic analysis was undertaken to examine
pelodytid-pelobatid relationships. This analysis also examines pelobatoid relationships. Results of
this study reveal the following hypotheses of relationships. The Pelobatidae and Pelodytidae are best
considered as separate families. Miopelodytes, Pelodytes, and Tephrodytes, whose relationships are
unresolved, are members of the Pelodytidae. Similarities between Tephrodytes and some of the pelobat-
ids are the result of convergence. Within the Pelobatinae Eopelobates, Macropelobates, and Pelobates
form one clade, in which Eopelobates and Pelobates are sister taxa, and Scaphiopus and Spea form
another. The Eopelobatinae is synonymous with the Pelobatinae. Leptobrachium, chosen as a repre-
sentative taxon for the Megophryinae, is the sister taxon to the Pelobatinae.

Introduction

The Cabbage Patch beds, exposed in the Blackfoot, Flint Creek, Deer Lodge,
and Divide intermontane basins of western Montana (Fig. 1), have yielded a
diverse fauna that includes mammals, birds, reptiles, amphibians, and fish (Ras-
mussen, 1989). This study reports a new anuran genus and species, Tephrodytes
brassicarvalis, in the family Pelodytidae, from these beds. The new taxon is based
on a nearly complete skeleton, two partial skeletons, and isolated bones, all of
which are well-preserved. They were collected by Donald Rasmussen from lo-
calities that he discovered in the Flint Creek Basin in Granite and Powell counties.
These specimens and isolated bones are referred to a single species because com-
parison of them with the holotype, and each other, reveals no significant differences
in their morphology. Other taxa of frogs also have been collected by Rasmussen
from the Cabbage Patch beds, and these are under study by the author.

The Cabbage Patch beds are Arikareean, late Oligocene-early Miocene in age.
During the Arikareean in western Montana a large basin, the Clark Fork Basin,
was being filled predominantly with fluvial and lacustrine fine-grained, tuffaceous
sediments (Rasmussen, 1989). Rasmussen (1977) suggested that the ash com-
ponent in these sediments blew in from the west, possibly originating in the
Cascades or the Columbia Plateau area, and was deposited in the uplands. From

Submitted 20 April 1993.

155

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Fig. 1.— Map of western Montana showing the location of basins in which the Cabbage Patch beds
are exposed (modified from Fields et al., 1985). The basin names are: 1) Blackfoot, 2) Flint Creek, 3)
Deer Lodge, and 4) Divide. The pelodytid fossils came from the Flint Creek Basin.

there, Rasmussen (1977) further speculated, the ash was washed into the depo-
sitional basins, of which the Clark Fork was one, and was extensively reworked
before final deposition. Only remnants of this formerly widespread unit exist
today in the Flint Creek and adjacent basins (Rasmussen, 1989). The depositional
environments of the three localities the fossils came from were determined to be
fluviatile overbank for localities KU-MT-12 and KU-MT-25, and lacustrine for
locality KU-MT-8 (Rasmussen, 1977). The holotype came from KU-MT-25.

The Pelodytidae was named by Bonaparte in 1850 (Frost, 1985), although Cope
(1866) is often given credit for naming the family. Boulenger (1897) included the
pelodytids in the Pelobatidae, and since then the pelodytids have been ranked
either as a separate family (Taylor, 1941; Lynch, 1973; Duellman, 1975; Sanchiz,
1978; Laurent, 1979; Dubois, 1983; Frost, 1985; Cannatella, 1985; Duellman and
Trueb, 1986) or incorporated with the Pelobatidae (Noble, 1924, 1931; Griffiths,
1963; Kluge and Farris, 1969; Vergnaud-Grazzini, 1970; Spinar, 1972; Savage,
1973). More recent classifications consider the pelodytids a separate family (Lynch,
1973; Duellman, 1975; Laurent, 1979; Dubois, 1983; Frost, 1985; Cannatella,

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Henrici - New Pelodytid Frog from Montana

157

1985; Duellman and Trueb, 1986; Ford and Cannatella, 1993). Because Tephrod-
ytes possesses several features (sculpturing on the dermal skull bones, presence
of posterior tip of the frontoparietal, frontoparietals cover frontoparietal fonta-
nelle, squamosal articulates with maxilla, and presence of elongate vertebral neural
arches) that also occur in some pelobatids, but were previously unreported for
the pelodytids, an examination of the pelodytid-pelobatid relationship is war-
ranted. Thus a cladistic analysis was undertaken to see if the new information
from Tephrodytes helps to resolve whether the pelodytids should be placed in the
Pelobatidae or in their own family. This analysis also addresses pelobatoid re-
lationships which were unresolved in the cladistic analysis of anurans undertaken
by Ford and Cannatella (1993).

The holarctic family Felodytidae is not very diverse. Two extant species of
Pelodytes, P. punctatus and P. caucasicus, occur in southwestern Europe and the
Caucasus region of southeastern Asia, respectively (Frost, 1985). The occurrence
of Pelodytes in the middle Pleistocene of France was recorded by Rage (1969)
and he questionably referred some of the fossils to P. punctatus (Rage, 1972).
Pelodytes arevacus and specimens attributed to Pelodytes are known from the
middle Miocene of Spain (Sanchiz, 1978). Miopelodytes was the only pelodytid
previously known from North America, and it is based on a single specimen from
the middle Miocene Elko shales near Elko, Nevada (Taylor, 1941). Additional
material, including tadpoles, has been collected from this locality and are being
described by Dr. Ted Ca vender (personal communication). Propelodytes wagneri,
from the middle Eocene of Messel, Germany, was considered to be a pelodytid
by Weitzel (1938). However, both Hecht (1963) and Sanchiz (1978) cast doubt
on this assignment, and Wuttke (1988) has referred to these specimens as Eopelo-
bates wagneri.

According to the most recent classification that includes fossil taxa (Duellman
and Trueb, 1986) the family Pelobatidae includes 13 genera in three subfamilies.
Seven of these are in the Megophryinae, which has no fossil record, and they
occur in southeastern Asia and from the Indo-Australian Archipelago to the
Philippine Islands (Duellman and Trueb, 1986).

The Eopelobatinae (Spinar, 1972) originally included only Eopelobates, which
has a holarctic distribution and ranges from the early Eocene to the Pliocene.
Nessov (1981) later added Aralobatrachus and Kizylkuma, which are based on
isolated elements from the late Cretaceous of Uzbekistan, but these taxa have
since been reassigned to the Discoglossidae (Rocek and Nessov, 1993). Studies
of the development of the frontoparietal by Rocek (1981, 1988) have revealed
that the frontoparietal in Eopelobates bayeri consists of three ossifications instead
of two. The presence of the median ossification has been inferred for E. anthra-
cinus, the genotype, and this character is included in the revised diagnosis for the
genus (Spinar and Rocek, 1984). Because the frontoparietal is apparently paired
in the North American Eopelobates grandis and E. guthriei, Rocek (1981) sug-
gested that they are probably more closely related to Scaphiopus than to Eopelobat-
es. I agree with his suggestion concerning E. guthriei. However, E. grandis pos-
sesses several features that do not support a close relationship with Scaphiopus.
Both specimens are currently under study by the author. Kuhn (1941) described
seven genera containing eight species that Estes (1970) later synonymized as
Eopelobates hinschei. However, Rocek (1981) questioned the taxonomic place-
ment of these taxa, in part because the frontoparietal is apparently paired. Here-
after, discussion concerning Eopelobates will be limited to the well-described E.
anthracinus and E. bayeri.

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1mm

Fig. 2.— Photographs of the holotype of Tephrodytes brassicarvalis, KU 19928. A, dorsal view; B, left
lateral view; C, right lateral view; and D, counterpart.

The fossorial Pelobatinae consists of the middle Oligocene Macropelobates from
Mongolia, the extant Pelobates from Europe, western Asia, and North Africa, and
the extant Scaphiopus from southern Canada to southern Mexico. The earliest
record for Pelobates is the late Eocene of Belgium (Bohme et al., 1982) and that
for Scaphiopus is the middle Oligocene of North America (Estes, 1970). If E.
guthriei is indeed a species of Scaphiopus , then the record can be extended back
to the early Eocene.

Abbreviations

Anatomical— Al, anterior lamina of scapula; As, angulosplenial; At, anterior tubercle of scapula;
C, columella; Cl, clavicle; D, dentary; Ex, exoccipital-prootic complex; F, femur; Fp, frontoparietal;
H, humerus; II, ilium; Is, ischium; M, metatarsals; Mx, maxilla; N, nasal; Op, otic plate of squamosal;
Or, otic ramus of squamosal; Pal, palatine process of maxilla; Pm, premaxilla; Po, postorbital process
of frontoparietal; Pp, posterior process of maxilla; Qj, quadratojugal; R, radio-ulna; S, sacrum; Sc,
scapula; Sp, sphenethmoid; Sq, squamosal; Tf, tibiofibula; fTF, fused tibiale and fibulare; U, urostyle;
V, vertebra; Zp, zygomatic process of maxilla.

Institutional. — AMNH, American Museum of Natural History; CM, Carnegie Museum of Natural
History; FMNH, Field Museum of Natural History; KU, University of Kansas, Museum of Natural
History; UNSM, University of Nebraska State Museum; and USNM, United States National Museum.

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Fig. 3.— Tephrodytes brassicarvalis, KU 19928. Line drawings of same views as in Fig. 2.

Systematic Paleontology

Class Amphibia
Order Anura
Family Pelodytidae
Tephrodytes, new genus

Type Species. — Tephrodytes brassicarvalis , new species.

Diagnosis. — Tephrodytes differs from all other pelodytids by having frontoparie-
tals that meet medially to conceal the frontoparietal fontanelle. It differs from
Pelodytes in the following unique combination of characters that are not known
for Miopelodytes: 1) posterior tip of frontoparietal present, 2) otic plate of squa-

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mosal present, 3) presacral vertebral neural arches elongate, and 4) anterior lamina
of scapula absent. It is distinguished from Miopelodytes by possession of reduced
otic ramus of squamosal and expanded sacral diapophyses.

Etymology. — From the Greek tephra, ashes, and dytes, diver, enterer; in ref-
erence to the tuffaceous sediments from which this frog was recovered, and in
which it may have estivated.

Tephrodytes brassicarvalis, new species

Holotype.— KU 19928 (Fig. 2, 3): partially complete, loosely articulated skel-
eton. The skull includes both frontoparietals, nasals, premaxillae, maxillae, squa-
mosals, right ?quadratojugal, right exoccipital-prootic complex, right ?columella,
right pterygoid, and both dentaries and angulosplenials. The postcranial skeleton
is preserved in part and counterpart and includes: fourth through eighth vertebrae,
sacral vertebra, urostyle, right humerus, right radio-ulna, both ilia, right tibiofibu-
la, and right fused tibilae and fibulare.

Type Locality and Horizon.— KU-MT-25, Cabbage Patch #10; Flint Creek
Basin, Powell County, Montana. Lower Cabbage Patch beds, Arikareean.

Referred Specimens. — From the type locality: KU 19221, partial left frontoparietal, left exoccipital-
prootic complex, and vertebral column; KU 23489, right scapula and clavicle, sacral vertebra, urostyle,
ilia, and hindlimbs; KU 18191, right maxilla; KU 19940, left maxilla; KU 18195, right ilium; KU
19917, left ilium; KU 19918, right ilium; KU 19919, right ilium.

From locality KU-MT-12, Cabbage Patch #4, Flint Creek Basin, Granite County, Montana, Middle
Cabbage Patch beds, Arikareean: KU 18266, proximal half of right humerus; KU 18270, proximal
end of fused tibiale and fibulare; KU 18273, proximal end of fused tibilae and fibulare.

From locality KU-MT-8, Pikes Peak # 1 , Flint Creek Basin, Powell County, Montana, Upper Cabbage
Patch beds, Arikareean: KU 20654, sacrum; KU 20659, complete right humerus and proximal end
of left humerus from different individuals.

Diagnosis.— As for genus (currently monotypic).

Etymology. — From the Latin brassica, cabbage, and arvalis, of a cultivated field,
in reference to the Cabbage Patch beds.

Description

The holotype (Fig. 2, 3) is a three-dimensionally preserved, nearly complete
skeleton in which the cranial bones are very closely associated. Its length, from
tip of snout to distal end of urostyle, is approximately 4.0 cm. That this specimen
had achieved adulthood before death is suggested by the well-ossified bones of
the skull and postcranial skeleton. Measurements of complete bones appear in
Table 1. The following description is based on the holotype unless otherwise
stated.

Skull. — Both halves of the subrectangular frontoparietals are preserved in KU
19928 (Fig. 2A, 3A). The right half is slightly crushed along a small portion of
the medial edge and its anteriormost end is covered by the right nasal. In the left
frontoparietal the anterior half of the lateral edge is broken away, the postero-
medial end lies underneath the posterior end of the right frontoparietal, and the
posterior tip is covered by matrix and the left transverse process of the third
vertebra.

The frontoparietal is widest near the posterior end (Fig. 2A, 3A, 4A). From the
posterior end it narrows to the midpoint of its length where it flares slightly in a
lateral direction to form the postorbital process (Fig. 4A). From there to the

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Table 1 . —Measurements for ail complete bones of Tephrodytes brassicarvalis. The holotype is KU

19928.

18191

19221

19917

19918

19928

20654

20659

23489

Frontoparietal length

7.3

Frontoparietal width

2.0

Nasal length

3.1

Nasal width

4.5

Maxilla length

12.5

10.4

8 th vertebra length
Sacral diapophyses

2.4

1.9

length

Sacral diapophyses

7.8

9.9

5.0

width

6.8

8.0

Urostyle length

Scapula length

11.2

3.5

Humerus length

Ilium length

20.7

17.5

18.3

9.7

14.0

Femur length

13.1

Tibiofibula length
Tibiale-fibulare

13.9

length

6.8

anterior end the frontoparietal is slightly narrower (Fig. 3A). The medial edge of
the left frontoparietal, where exposed, is straight. In Fig. 2 and 3, however, crushing
along the medial edge of the right frontoparietal causes it to appear as though it
is tapering. The posterolateral comer of the frontoparietal is drawn out to a point,
the posterior tip (^processus paroccipitalis of Rocek, 1981), which is oriented
posteriorly. I prefer to use Estes’ (1970) term posterior tip instead of the processus
paroccipitalis of Rocek ( 1 98 1 ) to avoid confusing this process with the paroccipital
process of the occiput.

Light sculpturing in the form of pits and grooves occurs on the relatively flat
dorsal surfaces of the frontoparietals. The sculpturing is most pronounced on the
posterolateral corner and is absent along the medial edge. In KU 19221 (Fig. 4A),
a larger individual than the holotype, the posterior half of one frontoparietal is
preserved. Sculpturing covers most of its dorsal surface, and the pits and grooves
are larger.

Unlike other pelodytids the frontoparietal fontanelle was not exposed in Te-
phrodytes. In some anurans that have an exposed fontanelle, such as Spea, the
frontoparietals are sutured together only at the posterior end. From there anteriorly
the two halves are free, which allows exposure of the fontanelle. In Pelodytes,
which also has an exposed frontoparietal fontanelle, even the posterior ends of
the frontoparietals are not in contact. Concerning Tephrodytes the preserved por-
tion of frontoparietal in KU 1 922 1 has several small indentations along the medial
edge which are interpreted here as a zigzagging suture scar (Fig. 4A). This provides
evidence that the posterior halves of the frontoparietals were sutured together. In
the holotype the medial edge of the right frontoparietal does not taper, indicating
that the two halves would have been in contact, thus covering the fontanelle.

Nasal bones are preserved only in the holotype. The right is undistorted (Fig.
2A, 3A) but the left is incomplete and its posteromedial comer is covered by the
anterior end of the left frontoparietal. The area where the two nasals meet medially

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Fig. 4 . — Tephrodytes brassicarvalis. A, posterior half of frontoparietal, KU 19921; B, presacrals 1-3,
KU 19921; C, vertebral column and part of sacral diapophysis, KU 19921; and D, right ilium, KU
19917. Scale in mm, in which A and B are drawn to same scale.

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Fig. 5. —Right maxilla of Tephrodytes brassicarvalis, KU 18191.

is relatively long. The width of the right nasal is greater than its length (Table 1).
Its anterior process is a small nubbin, and the concave anterolateral margin forms
the posteromedial border of the external narial opening. Laterally, the nasal is
drawn into a maxillary process which is directed posterolaterally (Fig. 2C, 3C).
There is no sculpturing on the dorsal surface of the nasals.

The right and left premaxillae, which bear teeth, are preserved in the holotype
(Fig. 2B, C; 3B, C). Here, the lateral ends of both premaxillae are overlain by the
medial ends of the maxillae. The medial end of the left premaxilla is covered by
the medial end of the right premaxilla. This telescoping causes the snout to appear
narrower than it actually is. The alary process is elongate in the left premaxilla
and incomplete in the right. Sculpturing is light and occurs as tiny pits but does
not extend onto the alary process.

Both maxillae are preserved in the holotype (Fig. 2A, B, C; 3A, B, C). Two
isolated maxillae (KU 18191, Fig. 5; and KU 19940) have also been recovered.
In the holotype the pars facialis of the maxilla is slightly crushed at the anterior
end where it reaches its greatest height. The pars facialis pinches in at the level
of the orbit and increases in height at the posterior end to form the zygomatic
process. In KU 18191 the dorsal surface of the zygomatic process is exposed and
bears a suture scar, which provides evidence that the zygomatic ramus of the
squamosal articulated with it. The posteroventral end of the maxilla is drawn
into a point, the posterior process, which extends posteriorly beyond the level of
the zygomatic process. Small teeth are present on the pars dentalis for about 3A
the length of the bone from its anterior end. In the holotype the tip of a tooth
that was prepared appears to be fang-like and recurved medially. In medial aspect,
in KU 19940, the palatine process arises off the medial face of the pars facialis
just dorsal to the pars palatina. In this specimen the tip of the palatine process is
broken off at the level of the dorsal edge of the pars facialis. In KU 18191, exposed
in lateral view, the tip of the palatine process extends beyond the dorsal edge of
the pars facialis. It seems likely that the palatine process was long enough to reach,
and articulate with, the lateral wing of the sphenethmoid, as in other pelobatoids.
Sculpturing in the holotype is light and occurs as pits and grooves which are larger
at the posterior ends of the maxillae. In KU 19940, the largest maxilla, only a
few pits are present on the posterior end.

In the holotype there is a sliver of bone located adjacent to the posteroventral
edge of the right maxilla. Based on its position and size it is believed to be the
quadratojugal. Even if this bone is not the quadratojugal its presence in Tephrod-

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ytes is suggested by the presence of a posterior process of the maxilla. Estes (1970)
has implied that anurans possessing a posterior process also have a quadratojugal.
To test Estes’ theory I examined the skull of all anuran taxa in the CM amphibian
osteology collection. With the exception of Scaphiopus and Spea all possessed a
quadratojugal and the maxilla bore a well-developed posterior process. Scaphiopus
and Spea were the only taxa that lacked a quadratojugal, and the posterior process
of the maxilla was reduced or absent in these taxa.

Both squamosals are preserved in the holotype. The left squamosal is complete
(Fig. 2A, B; 3A, B) and the right is missing the otic plate (Fig. 2A, C; 3A, C). The
squamosals are pushed anteriorly out of position so that the ventral ramus of
each is adjacent to the posterior end of the maxilla, causing the skull to appear
shorter than it actually is. The zygomatic ramus of the squamosal is relatively
thin and long, reaching its greatest thickness near the midpoint of its length. The
otic ramus, in lateral aspect, is reduced to a nubbin. Projecting medially from the
otic ramus is the otic plate which would overlap the crista parotica of the exoc-
cipital-prootic complex, if the bones of the skull were articulated. The medial
end of the otic plate is rounded. Sculpturing, consisting of a few large pits and
some tiny pits, is mostly confined to the area where the three rami meet. A few
small pits occur on the zygomatic ramus as well.

Only the right exoccipital-prootic complex is preserved in the holotype (Fig.
2A, 3A). Its medial end is pushed under the right frontoparietal and the lateral
end is missing. There is a left exoccipital-prootic complex in KU 19221 in which
the left frontoparietal covers its dorsal surface and the left parasphenoid alae
covers its ventral surface. In this specimen the medial wall of the auditory capsule
is exposed revealing the anterior and posterior acoustic foramina and the superior
and inferior perilymphatic foramina. The presence of a dorsal endolymphatic
foramen cannot be determined because of damage in the area where it would
occur. The prootic foramen is not completely surrounded by bone.

A small, T-shaped bone lies ventral to the right exoccipital-prootic complex
and medial to the posterior end of the right squamosal in the holotype. Based on
its shape and position it is believed to be the medial end of the right columella.
Note that only the footplate of the columella is visible in Fig. 3 A and C. A small
bone was found ventral to the columella and has been removed. This element is
domed shaped, with a smooth dorsal surface and a rough ventral surface. Possibly
it is a calcified operculum. It is equally possible that it is merely a fragment of
weathered bone. The occurrence of a calcified operculum in a fossil frog is not
unprecedented, as Kluge (1966) documented the presence of one in the lower
Miocene Scaphiopus neuter. Unfortunately, I have been unable to compare the
operculum of S. neuter with the possible operculum of Tephrodytes.

Preparation of the palatal region is hindered by the presence of the humerus,
radio-ulna, and several unidentified bones underneath the skull. The right pter-
ygoid, exposed in ventral view, bears an elongate anterior ramus, the anterior end
of which lies between the lower and upper jaws. Although the exact length of the
anterior ramus cannot be determined, it is unquestionably greater than twice the
length of the medial ramus. The short medial ramus has a blunt medial end. The
triangular posterior ramus is the shortest of the three rami. A portion of the left
vomer is exposed in the holotype. Although several tooth sockets of the vomer
are visible, no other details can be discerned.

Both lower jaws have been pushed dorsally inside the mouth of Tephrodytes.
Not much of either dentary could be exposed by preparation because the maxillary

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teeth overlap them (Fig. 2B, C; 3B, C). Only part of the angulosplenial (Fig. 2C,
3C) could be prepared without causing damage to the rest of the skull. Neither
bone exhibits any noteworthy features.

Postcranial Skeleton. — The vertebral column is preserved as part and counter-
part in the holotype (Fig. 2D, 3D). Presacrals four through eight and the sacrum
are exposed, but there is considerable damage. In KU 19921 the eight presacrals
and part of the left sacral diapophysis are exposed in ventral view (Fig. 4C), the
atlas and first three presacrals are exposed in dorsal view (Fig. 4B), and the bone
is well-preserved in all. The following description is based on KU 19921.

The eight presacral vertebrae have imbricate neural arches, as indicated by the
elongate neural spine of the third presacral, the only one completely preserved
and exposed (Fig. 4B). That the vertebrae are procoelous is best demonstrated in
the seventh and eighth presacral (Fig. 4C). The atlantal cotyles of the atlas are
closely juxtaposed. The dorsal surface of the atlantal neural arch is smooth. Its
posterior end is drawn out to form the neural spine, the posterior end of which
is broken off. Transverse processes of the second through fourth vertebrae are
elongate and directed laterally, except for the fourth which is directed slightly
posteriorly. The transverse processes of the fifth through eighth vertebrae are
thinner and shorter. Those of the fifth are directed laterally, the sixth moderately
anteriorly, and the seventh and eighth strongly anteriorly. Free ribs are not pre-
served and are presumably absent.

The sacral vertebra is preserved in the holotype (Fig. 2D, 3D), KU 23489 (Fig.
6), and KU 20654 (isolated sacrum). The diapophyses are widely expanded and
their length is greater than their width. In the holotype length was determined
from the right diapophysis of the counterpart. The width was determined by
doubling the distance from the lateral edge of the left diapophysis to the middle
of the centrum. In the holotype the length of the sacral diapophyses is roughly
equivalent to the length of the last four presacrals. The length of four presacrals
was determined by measuring the length of the eighth presacral, which has been
exposed in dorsal view by preparation, and multiplied by four. Variation in the
shape of the sacral condyle occurs. It is monocondylar in KU 23489, forming a
dorsoventrally compressed oval and distinctly bicondylar in KU 20654. In the
holotype it is neither distinctly monocondylar nor bicondylar but instead forms
a flattened oval that is slightly pinched in the middle.

The urostyle is preserved as part and counterpart in the holotype (Fig. 2A, D;
3A, D), and in KU 23489 (Fig. 6) the anterior half of the urostyle is exposed. In
the holotype it is at least as long as the last five presacral vertebrae. Transverse
processes on the urostyle are not apparent in either example.

A right scapula and clavicle, exposed in KU 23489 (Fig. 6, 7B), are the only
preserved bones of the shoulder girdle. The short, stocky scapula bears both the
pars acromialis and glenoidalis on the ventral end and they are narrowly separated
at their distal ends. Arising from the anterior edge of the scapula near the pars
acromialis is the anterior tubercle. An embayment lies between it and the pars
acromialis. The anterior tubercle also occurs in Pelodytes, and in that taxon the
clavicle reaches the ventral edge of the anterior tubercle, and thus overlaps the
scapula anteriorly. Because the ventral end of the scapula of Tephrodytes is very
similar in shape to that of Pelodytes , it is inferred that the scapula is anteriorly
overlain by the clavicle in Tephrodytes. In Tephrodytes the bone along the anterior
edge of the scapula is somewhat thick and rounded, and the margin is concave.
Thus, there is no anterior lamina.

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Fig. 6.— Postcranial skeleton of Tephrodytes brassicarvalis as preserved in KU 23489. Scale in mm.

The clavicle is oriented almost vertically in the matrix (Fig. 6). The process on
the lateral end that articulates with the scapula is broken off and the medial end
lies under the scapula. Enough of the clavicle is preserved to show that it is bowed,
which suggests that the shoulder girdle of Tephrodytes is arciferal. The preserved
portion of the clavicle is roughly the same length as that of the scapula. Thus, if
the clavicle was complete it would be longer than the scapula.

The right humerus (Fig. 2C, 3C) and radio-ulna are preserved in the holotype
in articulation. Here the humerus is exposed in dorsal and lateral views and the
proximal third of this bone is missing. Also, there are three isolated humeri (KU
1 8266 and 20659, two humeri). The following description is based on KU 20659,
a complete humerus. The shaft is straight and bears the crista ventralis on its
proximal half. The triangular fossa is not very deep. Well-developed medial and
lateral epicondyles bound the round humeral ball. The medial epicondyle pro-

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Fig. 7.— Comparison of scapula from specimens of similar size. A, Pelodytes punctatus, CM 54754;
B, Tephrodytes brassicarvalis, KU 23489; C, Scaphiopus holbrooki, CM 92261.

trudes medially, is triangular in medial aspect, and is joined to the shaft by a thin
crest. The smaller lateral epicondyle is crest-like. In lateral view a groove, the
surface of which is covered with finished bone, lies between the roughened bone
of the lateral epicondyle and the ball. The triangular olecranon scar is medially
situated.

The radio-ulna is exposed in lateral view, and the anterior half of the bone is
also exposed in dorsal view. The posterior end is poorly preserved, hindering its
preparation. The olecranon process is rounded. A sulcus is not visible on the
anterior half of the shaft. Also, in KU 23489 there is a bone that is possibly a
radio-ulna (Fig. 6).

Both ilia are preserved in the holotype. The left is complete whereas the right
is missing most of its shaft. There are several isolated ilia; the best preserved is
KU 19917 (Fig. 4D). In that specimen, the shaft, which lacks crests, is bowed
ventrally and has an oval cross section. The dorsal acetabular expansion bears an
oval, dorsolaterally-proj ecting dorsal protuberance which has roughened, unfin-
ished bone on its surface. This protuberance is larger in KU 19917 than in the
holotype, which is a smaller individual. A groove runs from the shaft to the dorsal
acetabular expansion. The ventral acetabular expansion and the dorsal acetabular
expansion are subequal in length. The acetabulum is large, roughly bell-shaped,
and somewhat deep. Its dorsal border projects slightly from the lateral surface of

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the dorsal acetabular expansion whereas the ventral border projects strongly from
the lateral surface of the ventral acetabular expansion. The dorsal ends of both
ischia are preserved in KU 23489 (Fig. 6), in which they appear to be medially
fused. The ischia do not extend greatly posteriorly as occurs in Eopelobates and
Megophrys (Zweifel, 1956; Estes, 1970).

Both femora and tibiofibulae are preserved in KU 23489 (Fig. 6) and partial
tibiofibulae occur in the holotype. The femur is slightly S-shaped, and the distal
end is wider than the proximal end. Presence of a femoral crest cannot be deter-
mined because of the orientation of the femora in the matrix. In KU 23489 the
tibiofibula is slightly longer than the femur (Table 1). The longitudinal sulcus of
the tibiofibula is restricted to the proximal and distal ends.

Complete fusion of the tibiale and fibulare occurs in Tephrodytes. The left fused
tibiale and fibulare is preserved in the holotype where it lies underneath the sacrum
(Fig. 2D, 3D). In KU 23489 a left fused tibiale and fibulare is partly overlain by
other bones (Fig. 6). Two isolated fused tibiale and fibulare (KU 1 8270 and 1 8273,
Fig. 8) were recovered as well. In the former two examples the longitudinal sulcus
is restricted to the proximal and distal ends. In the latter two examples, which
are considerably smaller and missing their extreme distal ends, the longitudinal
sulcus extends most of the length of the bone. The latter two bones are probably
from juveniles. In all examples, the fused tibiale and fibulare have an hourglass
shape and a small foramen is located near the midpoint of the shaft.

Only a few metatarsals and phalanges are preserved in KU 23489 and the
holotype making it impossible to determine the phalangeal formula. Metatarsals
1-4 are preserved very close to the distal end of the left fused tibiale and fibulare
(Fig. 6). A small phalanx is associated with metatarsal 1. None of the distal tarsal
bones are preserved; possibly they were cartilaginous. A prehallux modified as a
spade has not been found and presumably was absent, as in Pelodytes.

Family Assignment

Tephrodytes brassicarvalis possesses a fused tibiale and fibulare, which occurs
only in the frog families Pelodytidae and Centrolenidae. Of the numerous char-
acters that distinguish the pelodytids from the centrolenids (Duellman and Trueb,
1986) three are preserved in Tephrodytes, and are: the atlantal cotyles are closely
juxtaposed, the vertebral neural arches are imbricate, and the sacral diapophyses
are widely expanded. Tephrodytes possesses no characters that occur in the cen-
trolenids but not in the pelodytids, thereby making its affinity with the pelodytids
certain. Those characters that would indicate centrolenid affinities are: nonim-
bricate neural arches, dilated sacral diapophyses, and scapula not overlain ante-
riorly by clavicle.

During the course of this study I became aware of some inconsistencies in
several of the characters used in different definitions of the Pelodytidae. Taylor
(1941), Lynch (1973), and Duellman and Trueb (1986) all included in their def-
initions the fusion of presacral vertebrae I and II. None of the eight specimens
of Pelodytes punctatus that I examined had fused presacral vertebrae. They were
also free in the P. punctatus specimens examined by Cannatella (1985). Sanchiz
(1978), however, observed the free state in Pelodytes arevacus, P. caucasicus, and
both the free and fused states in Pelodytes punctatus. He concluded that in P.
punctatus the free state occurred in both subadults and adults. Zweifel (1956),
apparently following Taylor (1941), stated that they are fused in Pelodytes, al-
though neither author mentioned which species were examined. In Miopelodytes

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5mm

Fig. 8.— Fused tibiale and fibulare of Tephrodytes bmssicarvalis. A, KU 18273; B, KU 18270.

Taylor observed that the first two presacrals are fused. However, I found this
impossible to determine in Miopelodytes because of bone damage. Thus, fusion
of presacrals I and II is not diagnostic of the family and appears to be variable
in Pelodytes.

In his definition of the Pelodytidae, Lynch (1973) reported that the articulation
between the sacrum and the urostyle is monocondylar, whereas in their definition
of the family, Duellman and Trueb (1986) stated that it is bicondylar. Further-
more, Boulenger (1896), Taylor (1941), and Rage (1974) observed that the artic-
ulation is bicondylar, and Cannatella (1985) reported it as monocondylar. Zweifel
(1956) suggested that the type of articulation varies intraspecifically. Sanchiz
(1978) contested ZweifePs (1956) conclusion because he observed that the sacral
condyle of Pelodytes arevacus and P. punctatus has a unique condition of being
intermediate between the monocondylar and bicondylar states. In Tephrodytes
the form of the sacral condyle varies, being either monocondylar, bicondylar, or
the intermediate condition described by Sanchiz (1978). The type of articulation
cannot be determined for Miopelodytes. Thus, it appears that the type of articu-
lation between the sacrum and urostyle is a variable feature within the family.

Both Lynch (1973) and Duellman and Trueb (1986) included the presence of
transverse processes of the urostyle in their definitions of the Pelodytidae. How-
ever, of the eight specimens of Pelodytes punctatus that I examined, only one
possessed distinct transverse processes on the urostyle. In this specimen, CM
543 18F, the process on the right side is well-developed whereas that of the left
is a nubbin. The presence or absence of transverse processes does not appear to
be the result of variation between the sexes because they were both present and
absent in the females examined. Age as the cause of variation can be ruled out
because the specimens examined were of similar size. Geographical variation also
is not a factor because the specimens were from a single locality. Neither Mio-
pelodytes nor Tephrodytes bears transverse processes on the urostyle. Thus, the
presence of transverse processes of the urostyle varies in P. punctatus, has not
been reported for either Miopelodytes or Tephrodytes, and should not be regarded
as a diagnostic character of the family.

Evans and Milner (1993) noted that the ilia of Pelobates possess a spiral groove
at the base of the shaft whereas pelodytids lack this groove. All of the pelodytid
and pelobatid specimens that I examined possess this spiral groove, with the
exception of Miopelodytes. The presence of a spiral groove could not be determined

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in Miopelodytes because of damage to the ilium. Also, Sanchiz’s (1978) illustra-
tions of the ilium of Pelodytes arevacus and P. punctatus depict a groove. This
character is possibly diagnostic of the Pelobatoidea, although I have not surveyed
its distribution outside of the primitive anurans.

Pelobatoid Relationships

The monophyly of the Pelobatoidea has been established by Cannatella (1985),
and confirmed by Ford and Cannatella (1993). Supporting synapomorphies for
this hypothesis of relationship are the presence of an ossified sternum, the presence
of a palatine process of the maxilla (Cannatella, 1985; Ford and Cannatella, 1993),
and the presence of the adductor longus muscle (Cannatella, 1985). Cannatella
(1985) deviated from traditional classifications that include the megophryines in
the Pelobatidae by placing the megophryines in their own family, the Megophryi-
dae. Following his taxonomy, the Pelobatoidea thus includes the Megophryidae,
Pelobatidae, and Pelodytidae. He concluded, based on a cladistic analysis of
primitive anurans, that the pelodytids and pelobatids are more closely related to
each other than either is to the megophryids. Fossil taxa were not included in his
study. However, Ford and Cannatella (1993), in a cladistic analysis of anurans,
determined that the Megophryidae, Pelobatidae, and Pelodytidae form an unre-
solved trichotomy. The extinct Eopelobatinae were removed from the Pelobatidae
by Ford and Cannatella (1993) because these workers questioned the monophyly
and relationships of this group. Thus, they considered the Eopelobatinae to be a
metataxon, having either uncertain relationships within the Pelobatoidea or com-
prising a sister group to the Pelobatoidea. This left Pelohates, Scaphiopus, and
Spea as the remaining members of the Pelobatidae. The extinct pelobatine Ma-
cropelobates was not included in their study.

Previous studies have suggested somewhat different hypotheses of relationships.
Lynch (1973) proposed that the pelodytids are the sister group of the pelobatids
and higher anurans, whereas Sokol (1981) suggested that Pelodytes is most closely
related to the megophryine and pelobatine pelobatids. Boulenger (1899) hypoth-
esized that Pelodytes is more closely related to Pelobates than to Scaphiopus.

There has been considerable debate concerning the relationship of the extinct
Eopelobates and Macropelobates to other pelobatids. Zweifel (1956) suggested
that Eopelobates could be a subgenus of the megophryine genus, Megophrys.
Similarly, Kluge (1966) considered Eopelobates to be more closely related to the
megophryines than to the pelobatines. Estes (1970) observed that Eopelobates is
a primitive member of the family that is similar to both the megophryines and
the pelobatines. However, the above conclusions were based, in part, on exam-
ination of E. grandis and E. guthriei, which may not be species of Eopelobates
(Rocek, 1981). Derivation of the pelobatines and the megophryines from the
eopelobatines was suggested by Spinar (1972). Rocek (1981), on the other hand,
observed that both Eopelobates and Pelobates possess a frontoparietal derived
from three ossifications, a right and left ossification and a medial ossification
situated posterior to them. He homologized the medial ossification with the ex-
trascapular of Eusthenopteron. Because Eopelobates and Pelobates are the only
anurans known to share this feature, Rocek (1981) concluded that they evolved
separately from all other anurans. This hypothesis was rejected on the basis of
parsimony by both Cannatella (1985) and Milner (1988), and Milner has further
suggested that the medial ossification is merely a synapomorphy defining a clade
within the Pelobatidae.

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A common ancestor, possibly Macropelobates (Noble, 1924), or a form close
to Macropelobates (Zweifel, 1956; Estes, 1970) has been suggested for Scaphiopus
and Pelobates, although Estes (1970) has cautioned that Macropelobates occurred
too late in time to be the actual ancestor. Rocek (1982) theorized that Macro-
pelobates is more closely related to Scaphiopus than Pelobates based on morpho-
logical similarities and paleogeographical grounds. The Turgai Straits, which sep-
arated Europe and Asia from the Jurassic to the Eocene, would have prevented
dispersal of pelobatines into Europe from an Asian ancestor (Rocek, 1982).

Cladistic Analysis

To obtain a better understanding of the interrelationship of the pelobatoids, a
cladistic analysis was undertaken. Representative pelobatoids employed in the
ensuing character analysis are as follows: Leptobrachium, Eopelobates, Macro-
pelobates, Pelobates, Scaphiopus, Spea, Miopelodytes, Pelodytes, and Tephrodytes.
A list of species examined is presented in Table 2. Leptobrachium was chosen as
a representative megophryine because it is possibly the most primitive member
of that group (Estes, 1 970). Character states for Eopelobates were determined from
the text, figures, and photographs in Estes (1970), Spinar (1972), and Spinar and
Rocek (1984). Only E. anthracinus and E. bayeri were considered because the
status of other species in this genus has been questioned (Rocek, 1981). For
Macropelobates the character states were determined from examination of the
holotype and an undescribed, disarticulated specimen, consisting mostly of post-
cranial bones, that was discovered in the collections of the American Museum of
Natural History by Max K. Hecht. Because this specimen exhibits no differences
from the holotype of Macropelobates, other than being slightly smaller, I concur
with Hecht’s identification of it. The descriptions of Macropelobates by Noble
(1924), Estes (1970), and Rocek (1981, 1982) were also used. For Miopelodytes
character states were ascertained from examination of the holotype. Character
states for the remaining anurans were determined through examination of dry
skeletons and cleared and stained specimens. In this study Spea is treated as a
genus, even though it is commonly considered to be a subgenus of Scaphiopus,
to avoid problems in assigning character states for features in which the two taxa
differ. For each character, polarity assessments were based on the outgroup al-
gorithm (Maddison et al., 1984). Alytes, Bombina, and Discoglossus comprised
the outgroup. The distribution of character states for the outgroup can be found
in Table 3. Some of the characters used in the analysis are new, although the
majority of them come from the studies of Zweifel (1956), Kluge (1966), Estes
(1970), Spinar (1972), Sanchiz (1978), Rocek (1981, 1982), Spinar and Rocek
(1984), and Cannatella (1985).

Character Analysis.— A total of 25 cranial and postcranial characters were an-
alyzed in this study. These characters are discussed below.

1. Sculpturing commonly occurs on the outer surface of pelobatoid dermal
skull bones. Although the style of sculpturing varies (i.e., pits and grooves, pus-
tulose) and the distribution and amount of sculpturing on dermal skull bones
varies, only its presence or absence is noted here. Leptobrachium, Spea, and
Pelodytes lack sculpturing on dermal skull bones (state 0). Sculpturing (state 1)
occurs in Eopelobates, Macropelobates, Pelobates, Scaphiopus, and Tephrodytes.
Taylor (1941) made no mention of whether dermal skull bone sculpturing occurs
in Miopelodytes. However, the squamosal, which was not identified by Taylor
(1941) in his description of the holotype, bears sculpturing on the zygomatic and

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Table 2.— List of Recent and fossil specimens examined.

Taxon

Skeleton

Cleared and stained

Recent

Bombinatoridae

Bombina bombina

Bombina orientalis

Discoglossidae

Alytes obstetricans

Discoglossus jeanneae

Discoglossus pictus

Pelobatidae

Leptobrachium hasselti

Leptobrachium montanum

Megophrys montcola

Pelobates cultripes (adult)

Pelobates cultripes (tadpole series)

Pelobates fuscus

Scaphiopus couchi

Scaphiopus holbrooki

Scaphiopus hurteri

Spea bombifrons

Spea hammondi

Spea intermontanus

Pelodytidae

Pelodytes punctatus

Fossil

Pelobatidae

Macropelobates osborni

Scaphiopus skinneri

Pelodytidae

Miopelodytes gilmorei

otic rami. Poor preservation and their orientation in the matrix prevents the
identification of sculpturing on other dermal skull bones.

2. The frontoparietal of pelobatoids is either paired (state 0), consisting of a
right and left half, or is tripartite (state 1), being composed of a right and left
ossification and a medial ossification situated posterior to them (Rocek, 1981,
1988). The frontoparietal is paired in Leptobrachium, Scaphiopus, Spea, Mio -
pelodytes, Pelodytes, and Tephrodytes.

Rocek (1981) has documented the tripartite condition in Pelobates fuscus, P.
syriacus, and Eopelobates bayeri, and has inferred it for E. anthracinus (Spinar
and Rocek, 1984). I have observed it in cleared and stained tadpoles of P. cultripes .
Rocek (1981) noticed that during the ontogeny of the frontoparietal in Pelobates
the three ossifications fuse together leaving no trace of a suture. In adults of
Eopelobates bayeri the median suture is visible although it does not extend to the
posterior end of the bone because of the medial ossification (Rocek, 1981; Spinar
and Rocek, 1984).

For Macropelobates neither Noble (1924:fig. 1) nor Estes (1970:fig. 27) illus-
trated a median suture of the frontoparietal. However, Rocek (1981) detected a

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Table 3.— Distribution of character states among taxa used in the phylogenetic analysis. 0 indicates
the primitive state, 1 and 2 derived states , and 9 missing or inapplicable data. Usage of 1 and 2 does

not imply consecutively derived states.

Character numbers

1234567891 1

[111

111112

2 2

0 1

12 3 4

5 6 7 8 9 0

1 2

Leptobrachium

Eopelobates

Macropelobates

Pelobates

Scaphiopus

Spea

Miopelodytes

Pelodytes

Tephrodytes

Discoglossus

Alytes

Bombina

small notch on the posterior margin of the frontoparietal in Macropelobates which
is asymmetrical on the dorsal surface and symmetrical on the posterior surface.
Because this notch divides the frontoparietal into two symmetrical halves, he
suggested that it is paired. I found that in the holotype the median suture can be
clearly observed at the posterior end of the frontoparietal, arising from the notch,
thereby providing direct evidence for the paired nature of the frontoparietal in
Macropelobates.

3. In some anurans the two halves of the frontoparietal do not meet along the
midline of the skull, which allows exposure of the frontoparietal fontanelle (state
0). This occurs in Spea, Miopelodytes, and Pelodytes. The frontoparietal bones
are in contact medially and thus cover the frontoparietal fontanelle (state 1) in

Leptobrachium, Eopelobates, Macropelobates , Pelobates, Scaphiopus, and Te-
phrodytes.

4. The posterolateral corner of the frontoparietal of some pelobatoids is drawn
out to a point, the posterior tip (state 1). This occurs in Eopelobates, Macropelobat-
es, Pelobates, Scaphiopus, Spea, and Tephrodytes. Leptobrachium and Pelodytes
lack the posterior tip of the frontoparietal (state 0). The condition in Miopelodytes
cannot be determined.

5. The anterolateral margin of the nasal of pelobatoids varies in shape, being
either concave (state 0), forming the posterior border of the narial opening, or
straight (state 1). The margin is concave in Pelobates, Scaphiopus, Spea, and
Tephrodytes. Leptobrachium and Eopelobates have a straight margin. Cannatella
(1985) considered the anterolateral narial margin in Pelodytes to be straight.
However, because the specimens that I examined all have a concave margin, I
consider the state for Pelodytes to be primitive.

In a reconstruction of the skull of Macropelobates, Estes (1970) suggested that
the anterolateral margin of the nasal is concave. However, this portion of the
bone is not preserved (Noble, 1924; Rocek, 1982), and thus the shape of the
anterolateral margin is unknown. The condition of the nasal in Miopelodytes
cannot be determined.

6. The zygomatic ramus of the squamosal either does not articulate with the
maxilla (state 0), or articulates with the maxilla (state 1). In Leptobrachium, Spea,

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and Pelodytes it does not articulate with the maxilla. The squamosal articulates
with the maxilla in Eopelobates, Pelobates, Scaphiopus, and Tephrodytes. In Mac -
ropelobates the anterior end of the zygomatic ramus of the left squamosal and the
posterior end of the left maxilla are both broken away, thereby eliminating direct
evidence of their articulation, or lack thereof. Rocek (1982) observed that the
dorsal portion of the broken edge of the maxilla is very thick, indicating a strong
zygomatic process (processus zygomaticomaxillaris of Rocek, 1981), with which
the zygomatic ramus of the squamosal articulates. However, the presence of the
palatine process at the anterior end of the maxillary fragment indicates that the
preserved portion of maxilla formed the ventral border of the orbit. Thus, the
thickened bone Rocek (1982) believed to be the zygomatic process is actually the
orbital margin. Therefore, it cannot be determined if the squamosal articulated
with the maxilla in Macropelobates. Although Taylor (1941) made no mention
of it, a right squamosal, bearing both a zygomatic and otic ramus, is preserved
in the holotype of Miopelodytes. Unfortunately, it cannot be determined if the
zygomatic ramus of the squamosal articulated with the zygomatic process of the
maxilla because the bones are not preserved in articulation and the portion of the
maxilla where the zygomatic process would be, if present, is covered by a portion
of the pterygoid.

7. The otic ramus of the squamosal in lateral view is either a tiny nubbin (state
0) or is slightly elongate and somewhat deep (state 1). The otic ramus is reduced
in Leptobrachium, Spea, Pelodytes, and Tephrodytes. It is well-developed in Eopel-
obates, Macropelobates, Pelobates, Scaphiopus, and Miopelodytes.

8. The quadratojugal is present (state 0) in Leptobrachium, Eopelobates, Pelo-
bates, Pelodytes, and Tephrodytes. It is absent (1) in Scaphiopus and Spea. In
Macropelobates its presence or absence is controversial. Noble (1924) did not
mention this bone in his description of Macropelobates. Estes (1970) argued for
the probable presence of the quadratojugal, as he observed that the maxilla bore
a posterior process with which the quadratojugal would articulate. Rocek (1982)
observed that the posterior end of the maxilla is broken away and that the bone
along this break, in the posteroventral comer, is very thin suggesting that the
posterior process was absent, and thus the quadratojugal was absent. However,
the presence of the palatine process and teeth on the dentary process indicates
that the preserved portion of maxilla is from the orbital region and not the
posterior end of the maxilla, as suggested by Rocek (1982). Because there is no
solid evidence for the presence or absence of the quadratojugal, its state is here
considered to be unknown. It cannot be determined if the quadratojugal was
present in Miopelodytes.

9. The postchoanal ramus of the vomer, when present, forms the posterior
border of the internal nares. A short postchoanal ramus (state 0) occurs in Pe-
lodytes. This ramus is absent (state 1) in Pelobates and Leptobrachium. Scaphiopus
and Spea possess an elongate postchoanal ramus (state 2) that almost reaches the
maxilla. The presence or absence of the postchoanal ramus remains unknown in
Eopelobates, Macropelobates, Miopelodytes, and Tephrodytes due to the incom-
pleteness of these specimens.

10. In Pelodytes and Tephrodytes the anterior ramus of the pterygoid is elongate,
being greater than twice the length of the medial ramus (state 1). The anterior
ramus of the pterygoid is less than twice the length of the medial ramus (state 0)
in Leptobrachium, Eopelobates, Pelobates, Scaphiopus, and Spea. The pterygoid

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is incompletely known in Macropelobates and Miopelodytes. In Eopelobates the
length of the rami has not been described.

1 1 . The parahyoid bone is present (state 0) in Pelodytes and is absent (state 1)
in Leptobrachium, Pelobates, Scaphiopus, and Spea. It has not been identified for
Eopelobates, Macropelobates, Miopelodytes, and Tephrodytes. However, this char-
acter is considered to be unknown for these anurans because it is impossible to
determine whether the parahyoid bone does not occur or was not preserved in
these frogs.

1 2. The neural arch of the presacral vertebrae either barely extends beyond the
level of the posterior zygapophyses (state 0) or forms an elongate spine which
extends posteriorly beyond the level of the posterior zygapophyses (state 1). Lep-
tobrachium and Pelodytes have vertebral neural arches that are not elongate. The
vertebral neural arches are elongate in Eopelobates, Macropelobates, Pelobates,
Scaphiopus, Spea, and Tephrodytes. Unfortunately, this character cannot be as-
sessed in Miopelodytes because of damage.

13. The transverse processes of the last two presacral vertebrae of pelobatoids
are directed either moderately anteriorly (state 0), or strongly anteriorly (state 1).
Those directed moderately anteriorly are not situated close alongside the vertebral
centrum and do not reach the level of the anterior zygapophyses of the preceding
vertebra. In contrast, those directed strongly anteriorly tend to lie close alongside
the vertebral centrum and the tips of the transverse processes almost reach the
level of the anterior zygapophyses of the preceding vertebra. Leptobrachium,
Macropelobates, Scaphiopus, and Spea possess transverse processes with a mod-
erately anterior orientation. They are directed strongly anteriorly in Eopelobates,
Pelobates, Miopelodytes, Pelodytes, and Tephrodytes.

14. In some pelobatoids the sacral diapophyses are expanded (state 0) whereas
in others they are widely expanded (state 1). Traditionally, for descriptive and
comparative purposes, the length of the sacral diapophyses has been compared
to the equivalent number of vertebrae (Zweifel, 1956; Estes, 1970; Rocek, 1982).
For example, the length of the sacral diapophyses of Scaphiopus is equal to the
length of two presacral vertebrae whereas those of Pelobates are equal to the length
of four presacral vertebrae (Zweifel, 1956). This method works well with Recent
specimens but is difficult to apply to fossils because the vertebral centra are
commonly not exposed or poorly preserved. Instead, for the purpose of this study,
sacral diapophyses considered to be expanded are those having a width greater
than length. Widely expanded sacral diapophyses are those having a length greater
than width. Expanded sacral diapophyses occur in Leptobrachium, Scaphiopus,
Spea, and Miopelodytes, whereas widely expanded diapophyses occur in Eopel-
obates, Macropelobates, Pelobates, Pelodytes, and Tephrodytes.

15. In pelobatoids the scapula is either short (state 0) or long (state 1). The
scapula is considered to be short if its length is less than that of the clavicle, and
long if its length is greater than that of the clavicle. Pelodytes and Tephrodytes
have a short scapula. It is long in Leptobrachium, Eopelobates, Pelobates, Sca-
phiopus, and Spea. In Miopelodytes the right scapula is preserved but neither
clavicle is preserved. However, when the scapula of Miopelodytes is compared to
the scapula from a Scaphiopus specimen of similar size, it is apparent that the
scapula of Miopelodytes is short. For Macropelobates a similar comparison with
Pelobates reveals that the scapula is long.

16. Some pelobatoid scapulae bear an anterior lamina (state 0), which is a thin

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ridge of bone along the anterior edge of the scapula. An anterior lamina occurs
in Eopelobates, Macropelobates, Pelobates, and Pelodytes. It is absent (state 1) in
Leptobrachium, Scaphiopus, Spea, and Tephrodytes (Fig. 7). The presence or
absence of the anterior lamina cannot be determined with certainty in Miopelod-

ytes.

1 7. Both Tephrodytes and Pelodytes bear a small tubercle, the anterior tubercle,
at the anterior margin of the ventral end of the scapula (state 1, Fig. 7). Lepto-
brachium, Macropelobates, Pelobates, Scaphiopus, and Spea lack the anterior
tubercle (state 0). Spinar (1972) makes no mention of this tubercle in his detailed
description of the scapula of E. bayeri, so it is most likely absent. Unfortunately,
the presence or absence of the anterior tubercle cannot be determined for Mio-
pelodytes because the scapular bone is so badly damaged.

18. The clavicle either overlaps the scapula anteriorly (state 0) or abuts the
scapula’s ventral edge, and thus does not overlap it anteriorly (state 1). In Recent
pelobatids the scapula is not overlain anteriorly by the clavicle (Lynch, 1973;
Trueb, 1973; Duellman, 1975; Duellman and Trueb, 1986). However, some con-
fusion exists as to whether or not the scapula is anteriorly overlain by the clavicle
in the pelodytids. Trueb (1973), and Duellman and Trueb (1986) stated that the
scapula is not anteriorly overlain by the clavicle, whereas Lynch (1973) and
Duellman (1975) pointed out that the scapula is partially overlain by the clavicle.
Examination of specimens of Pelodytes, in which the clavicle and scapula are in
articulation, shows that the scapula is anteriorly overlain by the clavicle. The
scapula is anteriorly overlain by the clavicle in Pelodytes and Tephrodytes, and
is not overlain in Leptobrachium, Eopelobates, Pelobates, Scaphiopus, and Spea.
The state of this character cannot be determined in Macropelobates and Miope-
lodytes.

19. The medial end of the coracoid is considered to be expanded if the width
of the medial end is greater than the width of the lateral end (state 1). It is expanded
in Leptobrachium, Eopelobates, Macropelobates, and Pelobates. The medial end
is not expanded (state 0) in Scaphiopus, Spea, and Pelodytes. The coracoids of
Miopelodytes and Tephrodytes are not known.

20. The sternum is ossified (state 1) in Leptobrachium, Eopelobates, Pelobates,
and Pelodytes. It is cartilaginous in Scaphiopus and Spea (state 0). A sternum has
not been identified in Macropelobates, Miopelodytes, and Tephrodytes. Possibly
an ossified sternum was present but not preserved in these taxa. Therefore the
state of this character is considered to be unknown.

21. The ischium does not extend posteriorly beyond the dorsal acetabular
expansion of the ilium (state 0) in Pelobates, Scaphiopus, Spea, Miopelodytes,
Pelodytes, and Tephrodytes. It does extend posteriorly beyond the dorsal acetab-
ular expansion of the ilium (state 1) in Leptobrachium, and Eopelobates. Both
Zweifel (1956) and Estes (1970) illustrate the ischium of Macropelobates as not
extending greatly posteriorly. My examination of this genus has convinced me
that the ischium does extend greatly beyond the level of the dorsal acetabular
expansion. The difference in our interpretations could be because the specimen
has been extensively prepared in the time between their study and mine. Also,
Estes (1970) reconstructed the pubis as unossified, although Noble (1924) pointed
out that it was either ossified or that there was no room for it. This difference in
interpretation caused Estes (1970) to orient the ischium somewhat incorrectly,
making it appear as though it did not extend posteriorly.

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are discussed in the text.

22. Typically, the tibiale and fibulare of adult anurans are fused at their prox-
imal and distal ends (state 0). However, those of Miopelodytes, Pelodytes, and
Tephrodytes are completely fused to form a single bone (state 1).

23. Pelobatoids have either three free distal tarsal bones (state 0) in which distal
tarsals 1 and 2 are free and distal tarsals 3 and 4 fused, or two free distal tarsal
bones (state 1) in which distal tarsal 1 is free and distal tarsals 2, 3, and 4 are
fused (Lynch, 1973). Eopelobates and Pelodytes possess three distal tarsal bones,
whereas Leptobrachium, Pelobates, Scaphiopus, and Spea possess two. The num-
ber of distal tarsal bones is unknown for Macropelobates, Miopelodytes, and Te-
phrodytes.

24. In some pelobatoids the prehallux is modified as a bony spade which
enhances burrowing ability. A spade is absent (state 0) in Leptobrachium, Eopelo-
bates, Pelodytes, Miopelodytes, and Tephrodytes. It occurs (state 1) in Macropel-
obates, Pelobates, Scaphiopus, and Spea.

25. In Scaphiopus and Spea the gracilis muscle partially overlaps the sartorius
muscle and its tendon, thereby partially concealing these structures (state 1). The
gracilis muscle does not partially conceal the sartorius muscle and its tendon in
the other extant pelobatoids (state 0). Obviously, the state for this character cannot
be determined in the fossil taxa Eopelobates, Macropelobates, Miopelodytes, and
Tephrodytes.

Results. — Two equally parsimonious trees were generated by PAUP (version
3. OS for Macintosh, Swofford, 1991) utilizing the branch and bound method in
which the trees were rooted using the outgroup method. The trees have a length
of 54 steps and the consistency index is 0.519. All characters were treated as
unordered. Although PAUP was run using both the ACCTRAN and DELTRAN
optimizations, the results presented here are based on the trees obtained using
the ACCTRAN optimization because it favors reversals. A consensus of the two
trees is presented in Fig. 9.

The results of this cladistic analysis differ from those of Cannatella (1985) and
Ford and Cannatella (1993). Here, the monophyletic Pelobatoidea is divided into
two, well-supported clades, the Pelodytidae and the Pelobatidae. Tephrodytes is

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a member of the Pelodytidae, but its relationships with Miopelodytes and Pelodytes
are unresolved. The Pelobatidae includes the representative megophryine Lep-
tobrachium, Eopelobates, Pelobates, Macropelobates, Scaphiopus, and Spea. That
Eopelobates is a member of the Pelobatidae is well-supported. However, because
it is most closely related to Pelobates, it should be considered a member of the
Pelobatinae, thereby making the Eopelobatinae a synonym of the Pelobatinae.

Autapomorphies for the Pelobatoidea were not included in this study because
they do not contribute toward resolution of pelobatoid relationships. Thus, the
synapomorphies of the Pelobatoidea in this study all exhibit homoplasy and are:
anterior lamina of scapula absent (character 16, state 1), which reverses to present
(state 0) at Node F and Pelodytes ; and sternum ossified (character 20, state 1),
which reverses to cartilaginous (state 0) in Scaphiopus and Spea.

The Pelodytidae is united at Node C by three nonhomoplasious characters and
one that exhibits homoplasy. These are: anterior ramus of pterygoid elongate
(character 10, state 1); scapula with anterior tubercle (character 17, state 1); pos-
terior presacral vertebrae with transverse processes oriented strongly anteriorly
(character 13, state 1), which also evolves at Node F; and tibiale and fibulare
completely fused (character 22, state 1). There are two equally parsimonious
solutions regarding the relationships of the pelodytids, in which Tephrodytes is
more closely related to either Miopelodytes or Pelodytes. Acceptance of the former
theory stipulates that character 14, sacral diapophyses widely expanded (state 1),
evolves at Node C, and reverses to expanded (state 0) in Miopelodytes, and that
four synapomorphies, all homoplasious, define the Tephrodytes + Miopelodytes
clade. These characters are: sculpturing present on dermal skull bones (character
1, state 1), posterior tips of frontoparietal present (character 4, state 1), zygomatic
ramus of squamosal articulates with maxilla (character 6, state 1), and vertebral
neural arches elongate (character 12, state 1). All four characters also evolve at
Node D, the Pelobatinae, and characters 1 and 6 further undergo reversal in Spea.
Because the state for characters 4,6, and 1 2 cannot be determined in Miopelodytes,
one derived character, presence of dermal skull bone sculpturing, actually supports
the association between Tephrodytes and Miopelodytes. On the other hand, ac-
ceptance of the hypothesis that Tephrodytes and Pelodytes are more closely related
is supported by one character, sacral diapophyses widely expanded (character 14,
state 1). Assuming this relationship, three characters evolve in Tephrodytes that
also evolve in the pelobatines (Node D) and are: posterior tips of frontoparietal
present (character 4, state 1), squamosal articulates with maxilla (character 6, state
1), and vertebral neural arches elongate (character 12, state 1). Furthermore, the
presence of dermal skull bone sculpturing evolves at Node A, the Pelobatoidea,
and reverses to absent (state 0) in Pelodytes. Resolution of this trichotomy prob-
ably cannot be accomplished without further knowledge of Miopelodytes and
Tephrodytes.

The Pelobatidae, united at Node B by four nonhomoplasious and four hom-
oplasious characters, contains two subfamilies: the Pelobatinae (Node D) and the
Megophryinae (Leptobrachium). Although Eopelobates has recently been removed
from the Pelobatoidea (Ford and Cannatella, 1993), been placed in its own sub-
family, the Eopelobatinae (Spinar, 1972), and has been considered to be close to
the megophryines (Zweifel, 1956; Kluge, 1966) or intermediate between the me-
gophyrines and pelobatines (Estes, 1970; Spinar, 1972), this study establishes
Eopelobates as a member of the family Pelobatidae, and the subfamily Peloba-
tinae.

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The Pelobatidae is a monophyletic group that shares four nonhomoplasious
and four homoplasious derived characters. The nonhomoplasious synapomor-
phies are: postchoanal ramus of vomer absent (character 9, state 1), parahyoid
bone absent (character 1 1 , state 1), scapula long (character 1 5, state 1), and scapula
not anteriorly overlain by clavicle (character 18, state 1). Of these, however,
character 15 is the only one that is known for all members of the Pelobatidae.
Other characters supporting this clade are: frontoparietal fontanelle covered by
frontoparietals (character 3, state 1), which also evolves in Tephrodytes ; medial
end of coracoid expanded (character 19, state 1), which reverses to not expanded
at Node C; ischium extends posteriorly (character 21, state 1), which reverses in
Pelobates and Node C; and two free distal tarsal bones (character 23, state 1),
which reverses to three free distal tarsal bones in Eopelobates.

Containing two clades, Scaphiopus and Spea in one, and Eopelobates, Pelobates,
and Macropelobates in the other, the Pelobatinae (Node D) is supported by five
synapomorphies, all of which exhibit some degree of homoplasy. Three of these
evolve convergently in Tephrodytes and are: posterior tips of frontoparietals pres-
ent (character 4, state 1), squamosal articulates with maxilla (character 6, state
1), and vertebral neural arches elongate (character 12, state 1). The remaining
characters are: otic ramus of squamosal present (character 7, state 1), which also
evolves in Miopelodytes and reverses to reduced in Spea\ and spade present
(character 24, state 1), which reverses to absent in Eopelobates.

The clade of Scaphiopus and Spea (Node E) is defined by six synapomorphies,
of which half exhibit no homoplasy. These are: quadratojugal absent (character
8, state 1); postchoanal ramus of the vomer elongate (character 9, state 2); medial
end of coracoid not expanded (character 19, state 0); sternum cartilaginous (char-
acter 20, state 0); ischium not extended posteriorly (character 21, state 0); and
sartorius muscle and associated tendon concealed, in part, by gracilis major (char-
acter 25, state 1). Characters 19 and 21 have undergone reversal from the derived
state at Node B and character 20 reverses from the derived state at Node A.

Eopelobates, Pelobates, and Macropelobates form a clade supported by two
synapomorphies: sacral diapophyses widely expanded (character 14, state 1), which
also evolves in Tephrodytes and Pelodytes; and anterior lamina of scapula present
(character 16, state 0), a reversal from absent at Node A. Pelodytes also possesses
an anterior lamina of the scapula. The association of Macropelobates with Eopelo-
bates and Pelobates counters Rocek’s (1982) hypothesis that Macropelobates is
closest to Scaphiopus. The basis of his conclusion was that Macropelobates shares
13 characters with Scaphiopus and only four with Pelobates. However, he was
unable to determine the primitive state from the derived state for many of his
characters. What Rocek (1982) considered to be advanced specializations shared
between Macropelobates and Scaphiopus are: lack of quadratojugal, and propor-
tions of squamosal. A quadratojugal was not preserved with the holotype of
Macropelobates, which has an incomplete and somewhat disarticulated skull, and
Rocek (1982) inferred that it was absent. However, I concluded that the presence
or absence of a quadratojugal could not be determined for Macropelobates (see
Character Analysis, character 8). This leaves the proportions of the squamosal as
Rocek’s (1982) only synapomorphy between Macropelobates and Scaphiopus. Of
the four characters shared between Macropelobates and Pelobates, Rocek (1982)
was unable to determine whether they were primitive or derived. In this study
only two of the five synapomorphies for Scaphiopus and Spea could be determined
in Macropelobates. These are: medial end of coracoid expanded (character 19,

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state 1), and ischium expanded posteriorly (character 2 1 , state 1). Both Scaphiopus
and Spea possess the primitive state for these characters, which represents a
reversal. Thus, based on the characters analyzed in this study, Macropelohates is
more closely related to Eopelobates and Pelobates than to Scaphiopus and Spea.
However, additional knowledge about Macropelohates may lead to a different
hypothesis of relationship.

Two synapomorphies define the clade of Eopelobates and Pelobates (Node G):
frontoparietal derived from three ossifications (character 2, state 1); and transverse
processes of posterior presacral vertebrae oriented strongly anteriorly (character
13, state 1), which also evolves in the pelodytids. Milner (1988) was correct in
his assumption that a frontoparietal derived from three ossifications was merely
a synapomorphy for a clade within the Pelobatidae and not indicative of a separate
origin for Eopelobates and Pelobates from other anurans as proposed by Rocek
(1981).

Discussion

Examination of Tephrodytes, which has the diagnostic pelodytid character of a
fused tibiale and fibulare, revealed there are several derived characters that also
occur in some pelobatids, but not in Pelodytes. Thus, it seemed possible that these
characters would support inclusion of the pelodytids in the Pelobatidae, and that
Pelodytes possesses the primitive state for these characters as a result of reversal.
However, the phylogeny (Fig. 9) derived from this cladistic analysis suggests that
pelodytids are the sister group to the clade including the megophryines and pelo-
batines. Thus, it is best to retain the pelodytids in their own family. Derived
characters that occur in both Tephrodytes and the pelobatines, including Eopelo-
bates, are the result of convergence. These characters are: posterior tip of fron-
toparietal present (character 4), zygomatic ramus of squamosal articulates with
maxilla (character 6), and presacral vertebral neural arches elongate (character
1 2). Furthermore, a concealed frontoparietal fontanelle evolved convergently in
Tephrodytes and the pelobatids. Note, however, that in Spea the frontoparietal
fontanelle is exposed, a reversal. Ford and Cannatella (1993) listed the fused
tibiale and fibulare as the only unambiguous synapomorphy for the Pelodytidae.
To this I add the following: anterior ramus of pterygoid elongate, and scapula
with anterior tubercle.

The Pelobatidae, used here in its traditional sense to include the megophryines
and pelobatines, is monophyletic. Although numerous characters support this
relationship, none of them are unique to pelobatids. Three of the characters
supporting the Pelobatidae that exhibited no homoplasy in this study also evolved
in the Neobatrachia: parahyoid bone absent, scapula long, and scapula not overlain
anteriorly by clavicle. All of the characters supporting the Pelobatinae exhibit
some degree of homoplasy. This is caused, in part, by several of these features
evolving convergently in Tephrodytes, and also by the loss of the spade in Eopelo-
bates. That Eopelobates, used here to include E. anthracinus and E. bayeri, is a
member of the Pelobatinae is well-supported, thereby making the Eopelobatinae
a synonym of the Pelobatinae. One unique character, frontoparietal derived from
three ossifications, supports the close relationship between Eopelobates and Pelo-
bates. Macropelohates is the sister taxon to this clade, but there are no characters
unique to this clade. Within this study three nonhomoplasious characters support
the clade of Scaphiopus and Spea. Of these only one, sartorius muscle and as-
sociated tendon partially concealed by gracilis major, appears to be unique. The

1994

Henrici— New Pelodytid Frog from Montana

181

elongate postchoanal ramus of the vomer also occurs in the rhinophrynid, Chelo-
mophrynus, and the quadratojugal has also been lost in Ascaphus, Leiopelma, and
Notohatrachus.

In conclusion, a phylogeny for the Pelobatoidea is provided. This phylogeny
follows the more traditional approach in that the pelobatids and pelodytids are
sister clades and the megophryines are included in the Pelobatidae. Relationships
within the Pelodytidae are unresolved, and more information on the extinct mem-
bers is required. Within the Pelobatidae, the relationships of Eopelohates and
Macropelobates as presented here differ from previous phylogenetic hypotheses.
However, additional knowledge of these taxa could result in a different phylogeny.

Acknowledgments

I am grateful to the following for loan of specimens in their care: the late C. J. McCoy, CM; N.
Hotton III, USNM; R. Hunt, UNSM; D. Miao, KU; M. C. McKenna, AMNH; and H. Voris, FMNH.
Illustrations in Figures 4 and 5 were skillfully prepared by A. D. Redline. Thanks are due to D. Baird
for his assistance in coining the specific name, and to M. Hecht who brought the second only known
specimen of Macropelobates to my attention. This investigation benefitted from discussion with F. B.
Sanchiz. K. C. Beard and J. E. Rawlins provided assistance with running PAUP, and A. Campbell
provided technical assistance. The late C. J. McCoy, who read an earlier version of this manuscript,
D. C. Cannatella, and an anonymous reviewer provided helpful comments for which I am grateful.
This research was supported in part by the M. Graham Netting Research Fund.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 2, Pp. 185-192

31 May 1994

TAIWANESE SPECIES OF NEOPANORPA
(INSECTA: MECOPTERA: PANORPIDAE)

George W. Byers1
Abstract

Neopanorpa youngi n. sp. is described and illustrated. It is compared in detail with N. ophthalmica
Navas and N. gradana Cheng. All known Taiwanese species of Neopanorpa are differentiated in separate
keys for males and females, and all are illustrated by taxonomic details.

Introduction

Taiwan (formerly Formosa) has a surprisingly rich fauna of Mecoptera, in-
cluding numerous species of Panorpa and fewer of Neopanorpa and Bittacus. The
island has an area of nearly 36,000 km2 (13,836 sq mi). Except for an alluvial
plain about 30 km wide along its western side, it is largely mountainous, much
of its area above 1500 m and several peaks rising above 3500 m. But Taiwan is
crossed at midlength by the Tropic of Cancer. As a result, it has a diversity of
habitats, ranging from lowland tropical rain forest to high montane forest and
grassland. This has undoubtedly influenced the variety of its insect fauna.

Neopanorpa includes approximately 95 species at present, all in southeastern
Asia. The ranges of species seldom overlap, but I suspect this is due to insufficient
collecting. Species of Neopanorpa occur in India, Nepal, Bhutan, and Sikkim,
eastward through Burma, Thailand, Laos, and Vietnam to southern China, chiefly
south of 35° latitude, and southeastward through peninsular Malaysia to Sumatra,
Java, and Borneo. Much of this composite range overlaps that of the more boreal
genus Panorpa.

In Neopanorpa, the wings are more slender, particularly near their bases, than
in Panorpa. Vein 1A in the fore wing is relatively short (around 25% of wing
length) and ends at the wing margin before the level of the origin of the radial
sector (ORs). In Panorpa, 1A extends well beyond the level of ORs and is often
more than 40% as long as the entire fore wing. There are correlated differences
in both male and female genital structures as well, but these are not easily described
in this limited treatment.

The genus Neopanorpa is represented in Taiwan by at least nine species. Navas
(1911) named two of these as Campodotecnum formosanum and C. ophthalmi-
cum\ but Enderlein’s genus Campodotecnum was subsequently relegated to the
synonymy of Neopanorpa (Esben-Petersen, 1913). In 1912, Esben-Petersen de-
scribed Panorpa sauteri, which was later transferred to Neopanorpa. Issiki (1927)
added N. makii and N. magna ; and Issiki and Cheng (1947) described N. dispar.
Cheng (1952) named N. gradana and N. k-maculata. On a recent collecting trip
to Taiwan, Dr. C. W. Young found the ninth species, described here.

1 Snow Entomological Museum, Department of Entomology, University of Kansas, Lawrence, Kansas
66045-2119.

Submitted 13 July 1993.

185

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Neopanorpa youngi, new species

Description based on four males, three females, pinned.

Head. — Dorsum mostly piceous from area around ocelli backward onto occiput;
dark yellowish brown along margins of eyes. Rostrum amber-brown; mouth parts
yellowish brown except tips of maxillary palpi black. Scape and pedicel yellowish
brown, flagellum brown to dark brown, with 42-44 flagellomeres (number may
vary in one individual). Antennal length about 12-13 mm.

Thorax.— Pronotum unevenly dull yellowish brown, with diffuse black medial
band. Mesonotum and metanotum mostly sordid yellowish brown, with piceous
medial stripe that widens abruptly across anterior mesonotum. Pleural areas,
coxae, and mera dull pale yellowish brown, with small, darkly sclerotized spots
at coxal articulations and below wing bases. Hairs of pleura short and pale, longer
on anterior surfaces of middle and hind coxae. Legs yellowish brown; tarsi dark-
ening gradually to brown on apical tarsomeres.

Wings strongly tinged with yellowish brown, bands and spots brown. Apical
band enclosing one or two small to large pale areas. Pterostigmal band complete,
branched posteriorly near midlength; a slender, irregular, diagonal band con-
necting inner (basal) branch of pterostigmal band to marginal spot. Basal band
entire (holotype) or interrupted near M, connected in some individuals by slender,
diagonal band to marginal spot.

Abdomen of Male. — Terga 2-4 mostly black; posterior prolongation of tergum
3 (part of notal organ) dark yellowish brown, highly arched dorsad (Fig. 4); median
elevation on tergum 4 black, bluntly bilobed, with hairs directed cephalad. Tergum
5 and segment 6 unevenly dark yellowish brown; sterna 2-5 pale yellowish brown.
Segments 7-9 yellowish brown except for brown shading on sides of hypovalves.
Ninth tergum distinctly bilobed at apex (Fig. 7), extending far beyond ends of
basistyles (Fig. 3). Ninth sternum wide in ventral aspect, about 80% of greatest
width of genital bulb measured across basistyles. Hypovalves (Fig. 1) narrowly
rounded apically, appearing nearly acute, mesal edges rolled at base; blunt lobe
on dorsal surface of each (Fig. 2) extending between basistyles, thus not evident
in lateral aspect (Fig. 3). Dististyles slender, not strongly curved toward apex, each
with sharply tipped inner basal lobe (Fig. 5). Aedeagus (Fig. 6) with two strongly
bowed ventral appendages approximately in position of ventral parameres (but
possibly homologous with branches from parameres forming aedeagal hamulus
in Panorpa ; see Byers, 1993), each terminating in pale, wrinkled lobe directed
abruptly laterad at nearly right angle to bowed part of appendage. Ventral valves
strigose; lateral processes continuous with ventral parameres; dorsal parameres
inclined together, with uneven, almost truncate ends.

Abdomen of Female. —Terga 2-5 piceous-black, corresponding sterna pale yel-
lowish brown; tergum 6 dark yellowish brown, following segments yellowish brown.
Well-developed laterotergites on segments 7 and 8. Cerci (Fig. 22) two-segmented,
apical segment about three times as long as basal one. Subgenital plate (Fig. 21)
notched apically, with median basal membranous indentation; most hairs short,
10-12 long marginal and apical setae. Genital plate (Fig. 19, 20) with short oval
axial portion bearing thick, darkened lateral lobes conspicuously expanded dorsad;
spaiulate arms subparallel.

Body length, male, about 13-14.5 mm (holotype 13 mm); female about 12-13
mm (allotype 13 mm). Length of fore wing, male, 13.9-15.3 mm (holotype 13.9
mm); female 13.2-15.0 mm (allotype 15.0 mm).

Holotype, male, “Taiwan: Kaohsiung, Shanping. 640 m., 1 1-20 April 1988, J.

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Byers— Taiwanese Neopanorpa

187

Fig. 1-7 .—Neopanorpa youngi, n. sp. 1, genital bulb, male paratype, ventral aspect. 2, right hypovalve,
male, right lateral aspect. 3, genital bulb, paratype, left lateral aspect. 4, notal organ of abdominal
terga 3 and 4, left lateral aspect. 5, right dististyle, male, ventral (and slightly lateral) aspect. 6, aedeagus,
male paratype, ventral aspect. 7, ninth abdominal tergum, male, dorsal aspect (diagrammatic). Fig.
8-9. Neopanorpa ophthalmica, male syntype (Museum of Zoology, Barcelona). 8, right dististyle,
ventral aspect. 9, genital bulb, ventral aspect. Scale: Fig. 1-3, 5-9.

Rawlins, C. Young, R. Davidson.” Allotype, two female and one male paratypes,
same data as holotype; one male paratype same locality but 21-30 April 1988,
C. Young, R. Davidson, J. Rawlins; one male paratype same locality but 1-10
May 1988, R. Davidson, C. Young, J. Rawlins. Holotype, allotype, one male and
two female paratypes in the Carnegie Museum of Natural History, Pittsburgh,
Pennsylvania; two male paratypes in the Snow Entomological Museum, Univer-

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15. magna

16. gradana

17. gradana

18. k-maculata

Fig. 10-18.— Taxonomic details of males of Taiwanese species of Neopanorpa. 10, genital bulb, N.
sauteri, ventral aspect. 11, aedeagus, N. sauteri, ventral aspect. 12, abdominal segments 2-6, male
holotype of N. sauteri (Deutsches Entomologisches Institut, Berlin-Friedrichshagen), right lateral as-
pect, anterior at right (genital bulb missing). 13, genital bulb, N. formosana, male syntype (Museum
of Zoology, Barcelona), ventral aspect. 14, genital bulb, N. makii, after Issiki 1927. 15, genital bulb,
N. magna, after Issiki 1927. 16, genital bulb, N. gradana, from Cheng 1952. 17, same, with hypovalves
removed to show aedeagus, from Cheng 1952. 18, genital bulb, N. k-maculata, from Cheng 1952.
Scale: Fig. 10-13.

sity of Kansas, Lawrence, Kansas. The holotype is intact; the allotype is missing
the apical one-third of the right antenna, and its terminal abdominal segments
are dissected and mounted in polyvinyl alcohol, on the pin with the specimen.

This species is named for my friend and former student, Dr. Chen-Wen Young
of the Carnegie Museum of Natural History, who collected most of the specimens.
A native of Taiwan, Dr. Young is an outstanding field entomologist and collector
and a very capable taxonomist of Tipulidae.

Neopanorpa youngi superficially resembles N . ophthalmica, particularly in wing

1994

Byers —Taiwanese Neopanorpa

189

19. youngi

20. youngi

o 22. youngi

21 . youngi

23. ophthalmica

25. sauteri

26. k-maculata

27. makii

28. dispar

29. dispar

30. magna

Fig. 19=30.— Taxonomic details of females of Taiwanese species of Neopanorpa. 19, genital plate,
female allotype of iV. youngi, n. sp., ventral aspect. 20, same, right lateral aspect. 21, subgenital plate,
allotype of N. youngi, ventral aspect. 22, terminal abdominal segment and cerci, allotype of N. youngi,
ventral aspect. 23, genital plate, N. opthalmica, ventral aspect. 24, same, N. formosana. 25, same, N.
sauteri. 26, same, N. k-maculata. 27, same, N. makii. 28, same, N. dispar. 29, subgenital plate, N.
dispar, ventral aspect. 30, genital plate, N. magna, ventral aspect. Fig. 23=30 after Issiki and Cheng
1947. Scale a— Fig. 19, 20; scale b— Fig. 21, 22.

pattern and the subtriangular shape of the male’s hypovalves. The hypovalves in
ophthalmica, however, are wider, less acutely tipped, uniformly dark and with
more and longer hairs than those of youngi. A conspicuous difference in the genital
bulb of the males is with the width of the ninth sternum (hypandrium) in ventral

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aspect (Fig. 1, 9). In youngi, the width of the sternum is approximately 80% of
the greatest width of the genital bulb measured across the basistyles, while in
ophthalmica it is only about 50-55% (but nearly 68% in N. gradana). Differences
between youngi and both ophthalmica and gradana include the inner basal lobe
of the dististyle, sharply acute in youngi but relatively blunt in ophthalmica (Fig.
8) and gradana (Fig. 1 7), and the apex of the ninth tergum, with distinct lobes
separated by a U-shaped emargination in youngi but only shallowly indented,
virtually truncate in ophthalmica and gradana, thus without lobes. In aedeagal
structure, youngi resembles sauteri and gradana (compare Fig. 6, 11, and 17).
The most obvious difference is in the up-curved ventral appendages I interpret
as ventral parameres, the ends of which are enlarged, directed laterally, and pale
in youngi but flat and disc-like in sauteri (actually widened slightly ventrad, or
caudad). Cheng (1952) described the distal ends of these appendages in gradana
(Fig. 17) simply as “broad lobes.”

Females of N. youngi differ from those of all other known Taiwanese species
in the genital plate, particularly the large, blackened dorsal swelling at each side,
at the bases of the posterior arms. Cheng (1952) described somewhat similar
structures in N. k-maculata as “black and stout,” but I cannot interpret his
illustration (Fig. 26) in more detail. Such basal enlargements are not mentioned
in the descriptions of other species. The ovoid shape of the axial portion of the
plate is also not seen in other Taiwanese species. The apically notched subgenital
plate (Fig. 21) of N. youngi is scarcely different from its counterpart in several
other species; the membranous medial area at the base has not been mentioned
or illustrated for other species.

Species Identification

Since I have not seen specimens of some species of Neopanorpa from Taiwan,
I have relied on descriptions and illustrations provided by S. Issiki and F. Y.
Cheng. A few figures drawn by Dr. Cheng have in fact been copied directly from
his paper in Psyche, with permission of the current editor, Dr. David Furth.

Key to Males of Taiwanese Neopanorpa

(Note: Male of N. dispar Issiki and Cheng is unknown.)

Hypovalves long-elliptical in ventral aspect, slightly overlapped me-
dially (Fig. 13, 14) 2

Hypovalves subtriangular in ventral aspect, or rounded at apex, over-
lapped medially or apically or not overlapped 3

Apical band on fore wing connected to pterostigmal band along costal

margin .formosana Navas

Apical band on fore wing narrowly connected to pterostigmal band

along vein R3 makii Issiki

Hypovalves not overlapped, either medially or apically 4

Hypovalves overlapped along mesal edges or at apex 5

Hypovalves long, extending far beyond ends of basistyles, bowed later-
ad (Fig. 1 8), with small mesal lobe near base of each; sternum 9 pro-
longed so that hypovalves originate shortly before ends of basistyles;
dististyles only slightly curved, outer margins shallowly concave ....

k-maculata Cheng

Hypovalves short, almost straight, without mesal lobe near base, their
apices slightly narrowed and inclined mesad; sternum 9 not prolonged,

2(1).

2'.

3(F).

3'.

4(3).

4'.

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Byers— Taiwanese Neopanorpa

191

hypovalves terminating near level of ends of basistyles; dististyles (Fig.

1 5) longer than basistyles, strongly bowed outward (laterad)

magna Issiki

5(3'). Ninth abdominal tergum (epiandrium) shallowly emarginate apically,

almost truncate ................................................ 6

5'. Ninth abdominal tergum distinctly bilobed, with lobes separated by

deep U-shaped emargination 7

6(5). Hypovalves nearly acute at apex, subtriangular in ventral aspect, over-
lapped slightly along most (about 70%) of their length

ophthalmica Navas

6'. Hypovalves rounded at apex, narrower at midlength than at apex,
overlapped only apically for about 30% of their length (Fig. 16) ....

gradana Cheng

7(5'). Median projection from third abdominal tergum (notal organ) extend-
ing over terga 4, 5, and most of 6 (Fig. 12); inner basal lobe of dististyle

blunt at apex sauteri Esben-Petersen

7'. Median projection from third abdominal tergum extending only over
two-thirds of tergum 4; apex of inner basal lobe of dististyle acutely
pointed youngi, n. sp.

Key to Females of Taiwanese Neopanorpa

(Note: Female of gradana Cheng is unknown; its
position in this key is based mainly on the
wing pattern of the male.)

1 . Marginal spot in fore wing not extended to join basal (proximal) branch

of pterostigmal band 2

1'. Marginal spot in fore wing extended diagonally across wing to join

basal branch of pterostigmal band 6

2(1). Genital plate with well-developed anterior apodemes (Fig. 27, 28) ... 3

2'. Genital plate without evident anterior apodemes 4

3(2). Length of subgenital plate of eighth sternum approximately equal to

its greatest width (Fig. 29) dispar Issiki and Cheng

3'. Length of subgenital plate about 1.3 times its greatest width makii Issiki
4(2'). Apical band of fore wing including extensive pale spots from cell R2

to cell Mf, wing markings grayish brown gradana Cheng

(see Note above)

4'. Apical band entire or including only small pale spots; wing markings

dark brown to blackish brown 5

5(4'). Genital plate without distinct axial portion, arms abruptly widened

subapically (Fig. 30); body largely black magna Issiki

5'. Genital plate with distinct axial portion (Fig. 24); arms gradually wid-
ened subapically; body color mainly brown formosana Navas

6(1'). Band of pigment from marginal spot to pterostigmal band broad with
rather even edges, narrowest part as wide as distance between M1+2

and M3 where bands join k-maculata Cheng

6'. Connection between marginal spot and pterostigmal band of varying

widths but mostly slender, with irregular edges 7

7(6'). Axial portion of genital plate broadly triangular in ventral aspect (Fig.

23, 25) 8

7'. Axial portion of genital plate ovoid (Fig. 19) youngi, n. sp.

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Annals of Carnegie Museum

vol. 63

8(7). Arms of genital plate (apical plate) slightly divergent (Fig. 25); axial

portion narrower than distance between arms . . . sauteri Esben-Petersen
8'. Arms of genital plate approximately parallel (Fig. 23); axial portion
wider than distance between arms ................. ophthalmica Navas

Acknowledgments

I am indebted to my late friend and colleague Dr. Syuti Issiki and to Dr. Fung Ying Cheng for

providing certain descriptions and illustrations of some species of Neopanorpa.

Literature Cited

Byers, G. W. 1993. Autumnal Mecoptera of southeastern United States. The University of Kansas
Science Bulletin, 5 5 (2): 5 7-9 6.

Cheng, F. Y. 1952. Additions to the mecopterous fauna of Formosa. Psyche, 59:89-94.

Esben-Petersen, P. 1912. H. Sauter’s Formosa-Ausbeute. Neuroptera Planipennia. Entomologische
Mitteilungen, 1:197-198. (A second paper under the same general title was published in the same
journal in 1913.)

. 1913. Mecoptera and Planipennia collected in Java by Edward Jacobson. Notes from the

Leyden Museum, 35:225-236.

Issiki, S. 1927. New and rare species of Mecoptera from Corea, Formosa and Japan. Insecta Mat-
sumurana, 2:1-12.

Issiki, S., and F. Y. Cheng. 1947. Formosan Mecoptera with descriptions of new species. Memoirs
of the College of Agriculture, National Taiwan University, 1(4): 1-17.

Navas, L. 1911. NevroptSres nouveaux de l’extreme Orient. Revue Russe d’Entomologie, 11:11 1—
117.

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ANNALS

of CARNEGIE MUSEUM

THE CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213
VOLUME 63 16 SEPTEMBER 1994 NUMBER 3

CONTENTS

ARTICLES

Biology and immature stages of Chloropidae (Insecta: Diptera) associated

with spike-rushes (Cyperaceae: Eleocharis) I. Stem borers

Teresa K. Wearsch and B. A. Foote 193

Beads and pendants from Trants, Montserrat: Implications for the prehis-
toric lapidary industry of the Caribbean

David R. Watters and Richard Scaglion 215

A primitive cricetid (Mammalia: Rodentia) from the Middle Eocene of

Jiangsu Province, China . Ban-yue Wang and Mary R. Dawson 239

Mediocampus, a new stinkbug genus from the Dominican Republic (Insecta:

Heteroptera: Pentatomidae) Donald Bo Thomas 257

REVIEW

New Jersey Ferns and Fern Allies [James D. Montgomery and David E.

Fairbrothers] Bonnie L. Isaac 263

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 3, Pp. 193-213

16 September 1994

BIOLOGY AND IMMATURE STAGES OF CHLOROPIDAE
(INSECTA: DIPTERA) ASSOCIATED WITH SPIKE-RUSHES
(CYPERACEAE: ELEOCHARIS) L STEM BORERS

Teresa K. Wearsch12
B. A. Foote1 2

Research Associate, Section of Invertebrate Zoology

Abstract

The life cycles and larval feeding habits of one species of Chlorops (C. obscuricornis Loew) and three
species of Diplotoxa [D. inclinata Becker, D. nigripes (Coquillett), and D. sp. near versicolor (Loew)]
associated with spike-rushes of the genus Eleocharis (Cyperaceae) are presented. Larvae of the four
species show resource partitioning by attacking either stems or rhizomes, by feeding at different times,
or by utilizing different species of Eleocharis.

This study compares the morphology of the larvae; describes and illustrates the eggs, three larval
instars, and puparia of three species of Diplotoxa ; and provides a key separating the third-instar larvae
of all four species of Eleocharis stem borers.

Introduction

The sedge genus Eleocharis, containing about 1 50 species in the world, is nearly
cosmopolitan in distribution and is particularly well-represented in warmer regions.
It is easily distinguished from other genera of Cyperaceae by the presence of a
single, terminal, spike-like inflorescence that lacks obvious subtending bracts.
Both annual and perennial species occur. Determining species is difficult, as well-
developed seed heads and achenes are necessary for identification (Gleason, 1963;
Braun, 1967; Scoggan, 1978). Species of Eleocharis frequently form nearly pure
stands in open wetlands, perhaps due to their ability to secrete compounds that
inhibit the growth of other hydrophytic plants (Wooten and Elakovich, 1991).

The importance of spike-rushes to chloropid flies was indicated by Todd and
Foote (1987) who reported that a stand of E. smallii Britton contained ten of the
22 species of Chloropidae collected in eight vegetation types occurring in a fresh-
water marsh near Kent, Ohio.

Chlorops, a member of the subfamily Chloropinae, is the largest chloropid genus
in North America, containing 35 species (Sabrosky, 1987). Most of the available
information on the biology of the genus is based on studies of the gout fly, C.
pumilionis Bjerkander (C. taeniopus Meigen, auct .), an economically important
pest of cereal grains in Europe. The stem-boring larvae of that species were de-
scribed in detail by Frew (1923 a, 1923 b), Balachowsky and Mesnil (1935), Good-
liffe (1939, 1942), Nye (1958), and Dennis (1961). Other European species that
are stem borers of grasses are C. speciosa Meigen, C. brevimana Loew, C. interrupta
Meigen, and C. marchali Mesnil (Ferrar, 1987). A few species ofNearctic Chlorops
have larvae that are primary invaders of sedge stems (Valley et al., 1 969). Recently,

1 Department of Biological Sciences, Kent State University, Kent, Ohio 44242.

2 Present address: 36903 Julian Street, Avon, Ohio 4401 1.

Submitted 19 April 1993.

193

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Rogers et al. (1991) elucidated the life history and described the immature stages
of C. certimus Adams, a species whose larvae attack stems of the sedge genus
Carex.

Diplotoxa, also a member of the subfamily Chloropinae, is a cosmopolitan
genus that has been recorded in nearly all the biogeographic regions. Five of the
seven Nearctic species, D. alternata (Loew), D. inclinata Becker, D. messoria
(Fallen), D. recurva (Adams), and D, versicolor (Loew), are practically transcon-
tinental in distribution; D. unicolor Becker is restricted to the western states and
provinces; and D. nigripes (Coquillett) occurs only in the eastern and midwestem
states and provinces (Sabrosky, 1965).

Few papers have been published on the larval feeding habits and host plants
of Diplotoxa. Wendt (1968) reported that larvae of D. messoria are phytophagous
in European species of Eleocharis, but gave no information concerning host plant
specificity, larval feeding habits, or life cycle parameters. Valley et al. (1969) reared
D. messoria, D. nigripes, and D. sp. (near D. inclinata) from stems of Eleocharis
in North America. Spencer (1977) reared a New Zealand species from inflores-
cences of a grass of the genus Echinochloa. No immature stage of any species has
been described (Ferrar, 1987).

The present paper elucidates the life cycles and larval feeding habits of one
species of Chlorops and three of Diplotoxa that attack the stems and rhizomes of
Eleocharis spp. in northeastern Ohio. The immature stages of three species of
Diplotoxa are described and illustrated, and a key is given to the third instars of
all four species of Eleocharis stem borers. In addition, certain morphological
structures found in the larvae are compared.

Materials and Methods

Collecting Techniques.— Adults were collected weekly by sweeping vegetation of suitable habitats
with a standard aerial insect net. Collecting sites were all located in Portage County in northeastern
Ohio. Eggs were collected in nature by pulling up Eleocharis plants in the field and examining them
in the laboratory. Larvae and puparia were found during the winter and spring by examining developing
rhizomes, the bases of young shoots, and new stems of Eleocharis that were approximately 0. 5-1.0
in in height.

Rearing Techniques.— Adult flies were sexed and paired, and placed in baby food jars (9x6 cm)
which had their bottoms removed. The jars were inverted, and the top pressed into the bottom of a
small plastic Petri dish (5.5 x 1.3 cm) containing moist peat moss. The open bottom of the jar was
covered with 1-4 layers of cheesecloth held in place by a rubber band. A small pellet of honey and
brewer’s yeast pressed to the side of the jar served as adult food. The peat moss was moistened daily,
and the fly food replaced periodically. Small sections of Eleocharis stem, approximately 5-6 cm in
length, were oriented vertically in the peat moss to provide oviposition sites. These were replaced
periodically with fresh sections, as females would not oviposit on decomposed material.

Eggs removed from the rearing jars were placed in small Petri dishes (5.4 x 1.3 cm) containing
discs of moist paper toweling; occasionally whole stem sections containing eggs were transferred.

Newly-hatched larvae were placed on young shoots of Eleocharis and examined daily for information
on larval feeding habits, length of larval stadia, and sites of pupation. Larvae were transferred to fresh
plant material regularly, as they quickly abandoned decaying stems. Puparia were transferred to small
Petri dishes containing moist peat moss. Parasitoid wasps emerging from puparia were killed and
preserved in 70% ETOH.

Preservation and Preparation of Specimens. — Approximately 10-20 eggs of each species were mea-
sured and preserved in KAAD. Larvae were killed in hot (not boiling) water and were either preserved
in 70% alcohol or treated for further study, and drawn using standard light microscopy techniques.

Abbreviations used in figures are: A, antenna; AP, anterior papilla; APB, anterior end of parastomal
bar; ApT, apical tooth; ASL, anal slit; ASp, anterior spiracle; AT, accessory tooth; Cph, cephalo-
pharyngeal skeleton; DC, dorsal cornu; FP, frontal papilla; GP, genal papilla; HS, hypopharyngeal
sclerite; IP, interspiracular process; LP, lateral papilla; LS, ligulate sclerite; m, membrane; MH, mouth-
hooks; MP, maxillary palp; Pa, papilla; PB, parastomal bar; PcS, pseudocephalic segment; PfP, pre-

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Wearsch and Foote— Chloropid Biology and Immature Stages

195

frontal papilla; PhS, tentoropharyngeal sclerite; PP, perianal pad; PSP, posterior spiracular plate; R,
ramus; SB, spinule band; ShS, subhypopharyngeal sclerite; SSI, spiracular slit; StB, stigmatic bulb;
StSc, spiracular scar; StT, stigmatic tube; Tr, trabecula; TP, thoracic papilla; VC, ventral cornu; WP,
wing process of hypostomal sclerite.

Life Histories
Chlorops obscuricornis Loew

This species is strictly Nearctic in distribution, ranging from Virginia and New
York west to Manitoba and Oregon and south to Florida and California (Sabrosky,
1965). Valley et al. (1969) reported that the larvae were primary invaders of stems
of Eleocharis smallii.

Rearings were initiated from numerous adults and immature stages collected
from stands of E. smallii, a perennial species having sizeable reddish or purplish
rhizomes. The fidelity of C. obscuricornis to its host plant was shown by Todd
and Foote (1987) who found adults almost exclusively in a stand of E. smallii.
At another marsh, a few adults were swept from a stand of E. obtusa (Willd.)
Schultes, an annual species. Numerous adults were also swept from another pe-
rennial species, E. rostellata Torrey, in a calcareous fen. The flight period, as
determined by weekly sweep samples of a stand of E. smallii occurring in a
roadside ditch, lasted from early June to mid- July (Fig. 54). Adults were most
abundant in mid- June.

Laboratory-reared females lived 12-19 days ( n = 8); males, 8-1 1 days ( n = 8).
No courtship displays were noted, and it appeared that males merely assaulted
any nearby fly of proper size and configuration. Nonreceptive females dislodged
males by vigorous kicking of their hind legs. During copulation the male situated
himself above the female, facing in the same direction. The fore tarsi rested on
the bases of her folded wings, his middle tarsi were applied to the lateral surface
of her abdomen, and the hind tarsi grasped the female’s genitalic segments.

Both field-collected and reared females laid eggs readily on lengths of Eleocharis
stems in the breeding jars. All field-collected eggs were found near the bases of
stems. The incubation period lasted 5-7 days (n = 12). No larvae were reared to
the pupal stage in the laboratory-initiated rearings. Second-instar larvae were
found during late March feeding as primary invaders in stems of E. smallii. A
few larvae occurred in young shoots, but most were in larger, older stems. Fully
grown larvae were found by the middle of April. Shortly before forming puparia,
larvae retreated to the crown, tunneled into one of the rhizomes, reversed direc-
tion, and inserted their posterior spiracles into the hollowed-out portion of the
rhizome. Field-collected puparia produced adults in 10-15 days under laboratory
conditions ( n = 4).

The abbreviated flight period (Fig. 54) suggests that C. obscuricornis is uni-
voltine in northern Ohio. Overwintering apparently occurred as young larvae in
quiescence. Larvae collected during the winter months became active in the lab-
oratory and began feeding on Eleocharis stem tissue. Larval feeding was completed
in nature in April and early May, pupation occurred in late May, and adults
emerged in June. Eggs were deposited between mid- June and early July. Larvae
fed in the stems until late October.

Diplotoxa inclinata Becker

Diplotoxa inclinata has a transcontinental distribution, ranging from Quebec
to California and south to New Jersey and Texas (Sabrosky, 1 965). It was abundant

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vol. 63

— aao_ 12 T-4o

Fig. 1-12 .—Diplotoxa nigripes. 1. Lateral habitus of larva, third instar. 2. Ventral view of anterior
end, same. 3. Dorsal view of posterior end, same. 4. Lateral view of cephalopharyngeal skeleton, first
instar. 5. Ventral view of cephalopharyngeal skeleton, third instar. 6. Lateral view of cephalopharyngeal
skeleton, second instar. 7. Same, third instar. 8. Mandible, second instar. 9. Same, third instar. 10.
Egg. 1 1. Puparium, dorsal view. 12. Same, lateral view.

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Wearsch and Foote Ch lo rg pi d Biology and Immature Stages

197

Fig. 13-17. — Diplotoxa nigripes. 13. Posterior spiracular plate, first instar. 14. Same, second instar.
15. Same, third instar. 16. Anterior spiracle, second instar. 17. Same, third instar.

in freshwater marshes in northeastern Ohio that contained stands of its host plant,
Eleocharis smallii (Todd and Foote, 1987). Several adults were also swept from
a small stand of E. obtusa. This was one of the first species of Diplotoxa to emerge
in the Kent area, as six adults were collected as early as May 5. The flight period
lasted from early May until early September (Fig. 55). Weekly sweep samples of
adults strongly suggest that there are two generations a year in northeastern Ohio.

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28 — 055 29 — 2J2

Fig. 18-29 .—Diplotoxa sp. near versicolor. 18. Lateral habitus, third instar. 19. Dorsal habitus, same.
20. Lateral view of segment 1, third instar. 21. Ventral view of cephalopharyngeal skeleton, third
instar. 22. Mandible, second instar. 23. Same, third instar. 24. Lateral view of cephalopharyngeal
skeleton, first instar. 25. Same, second instar. 26. Same, third instar. 27. Puparium, lateral view. 28.
Same, dorsal view. 29. Egg.

Adults usually rested head downward on Eleocharis stems. Paired adults placed
in breeding jars with lengths of Eleocharis spent most of their time on the stems
and did not wander freely about the enclosure. Field-collected females lived 7-
21 days in the laboratory (n = 10); males, 12-17 days (n = 10). In contrast, reared
females lived only 4-18 days (n = 5). Reared females usually had a premating
period that lasted less than 24 hr, and one female mated approximately six hours

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Wearsch and Foote— Chloropid Biology and Immature Stages

199

Fig. 30-35. — Dip lot oxa sp. near versicolor. 30. Posterior spiracular plate, first instar. 31. Same, second
instar. 32. Same, third instar. 33. Anterior spiracle, second instar. 34. Same, third instar. 35. Perianal
pad, third instar.

after emerging. No overt courtship behavior was observed. Mating usually oc-
curred during late afternoon. During coitus, the male positioned himself above
the female at about a 45° angle to her body. The head was positioned directly
above the anterior portion of her scutellum, the front tarsi rested slightly in front
of the wing bases of the female, and the mid and hind tarsi clasped the sides of
the female’s abdomen. The wings of both sexes remained folded during mating
which lasted from eight to over 30 minutes (n ----- 1 2).

The preoviposition period from mating to the first deposition of eggs usually

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Fig. 36-49. —Diplotoxa inclinata. 36. Egg. 37. Anterior spiracle, second instar. 38. Same, third instar.
39. Ventral view of segment 1, third instar. 40. Dorsal view of stigmatic tubes, third instar. 41. Lateral
view of cephalopharyngeal skeleton, first instar. 42. Same, second instar. 43. Mandible, second instar.
44. Lateral view of cephalopharyngeal skeleton, third instar. 45. Mandible, third instar. 46. Ventral
view of subhypopharyngeal and ligulate sclerites, third instar. 47. Ventral view of cephalopharyngeal
skeleton, third instar. 48. Puparium, dorsal view. 49. Same, lateral view.

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Wearsch and Foote- Chloropid Biology and Immature Stages

201

Fig. 50-53 .—Diplotoxa inclinata. 50. Posterior spiracular plate, first instar. 51. Same, second instar.
52. Same, third instar. 53. Perianal pad, third instar.

was less than a day, and one female began ovipositing six hours after mating. If
Eleocharis stems were not available, females delayed oviposition for over a week
then laid a few eggs on the peat moss that formed a moist substrate in the breeding
jars. These females died a day or so after oviposition. In contrast, females confined
with Eleocharis readily oviposited on the stems and lived for several days after
ovipositing. Five females laid 86, 89, 72, 62, and 86 eggs, respectively (x = 79),
over a 16-day period. Usually 4-5 eggs were laid by a female each day once
oviposition began. Most of the eggs were deposited at the base of a stem just
above the level of the peat moss substrate. A few eggs were placed higher on the
stems, but none was placed on the inflorescence. The incubation period lasted 4-
5 days (n = 25).

In nature, eggs were attached to stems just above the water level and oriented
parallel to the stem. One or two eggs were usually found on each stem, although
a few stems supported as many as five eggs. Stems occurring in deeper water were
more heavily infested than those in shoreline situations.

Newly-hatched larvae crawled down the stem and entered young shoots at the
base of the plant where they fed as primary invaders. Following the first larval
molt, larvae ate their way downward to the bases of the shoots and penetrated
into the rhizomes attached to the crown. Here, they fed on succulent meristematic

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tissues, leaving a distinct feeding trail through the rhizome. When a larva reached
a tiller arising from the rhizome, it entered the upright shoot and consumed its
contents. It then returned to the rhizome and fed along it until another tiller was
encountered. This process was repeated several times, and the larva usually was
fully grown by the time it reached the tip of the rhizome. Pupation occurred within
the feeding trail near the distal end of a rhizome. Shortly before forming a pu-
parium, the larva inserted its posterior spiracles into the living tissue adjacent to
the feeding trail.

In the laboratory, newly-hatched larvae were placed on the tip of a rhizome,
the succulent tissue of which was surrounded by a protective sheath. Larvae
quickly removed under the sheath and began to feed on the meristem. However,
duration of larval stadia under laboratory conditions could not be determined.
Collections of larvae and puparia in nature allowed for estimations of the third
larval and pupal stadia. The third stadium lasted 5-6 days (n = 1 2); the prepupal
period, 30-48 hr ( n = 5); and the pupal period, 10-13 days ( n = 4).

Overwintering occurred as second instars in quiescence, feeding recommenced
in April, and pupation occurred in the last two weeks of April.

The third instars and pupae of this species were easily distinguished from those
of the other species of Diplotoxa by their greenish color.

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Diplotoxa nigripes (Coquillett)

Diplotoxa nigripes ranges from South Dakota and Kansas east to Quebec, Maine,
and the District of Columbia (Sabrosky, 1965). It was, by far, the most abundant
of the three species of Diplotoxa collected in Ohio.

Adults were abundant between June 13 and October 3 in strands of Eleocharis
obtusa, but were only occasionally collected in stands of E. smallii. No adults
were obtained in the sweep samples taken weekly in a stand of E. smallii growing
in a roadside ditch, nor in weekly net samples from reed canary grass (Phalaris
arundinacea L.) and two species of sedges ( Carex lacustris Willd., C. stricta Lam.)
in a freshwater marsh near Kent during the warm seasons of 1984 (Todd and
Foote, 1987) and 1989. The larval rearings as well as the adult collections indicate
that the primary host of this species is E. obtusa.

Adults usually rested on the stems of the host plant facing downward. Field-
collected males lived 14-37 days in the laboratory ( n = 4); females, 7^45 days ( n
= 8). In contrast, females reared in the laboratory rarely lived more than 30 days.
Mating was observed three times in nature on stems of the host plant during late
afternoon. No overt courtship behavior was observed. During copulation, the
male was positioned above the female and facing in the same direction with his
head at about the level of her scutellum. The wings of both sexes remained folded
during mating. The male’s fore tarsi rested on the bases of the female’s wing, and

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the middle and hind tarsi were pressed against the sides of her abdomen. Mating
lasted an average of ten minutes (n = 3). Males terminated mating by moving
anteriorly off the body of the female.

Field-collected females laid numerous eggs in the laboratory breeding jars,
although no reared females oviposited. Three females held in the laboratory for
11 days laid 56, 61, and 64 eggs, respectively. Laboratory-held females showed
no particular preference for any part of the host plant, although more eggs were
deposited on inflorescences and near the base of the stem. In contrast, host plants
collected in nature never had eggs anywhere except on the stem bases. Eggs were
not inserted into the stem tissue, but were affixed to the upper end of the reddish
brown sheath that encircled the lower six inches of the stem. Only one or two
eggs were found on each stem.

The incubation period lasted 5-7 days under laboratory conditions, although
more than 80% of the eggs laid after the first week of October failed to hatch. The
incubation period of those few October-laid eggs that did hatch was considerably
extended, lasting 13-15 days. These observations suggest that autumn-deposited
eggs enter diapause and overwinter.

The larval stages were very difficult to rear in the laboratory because they would
not stay in stems of the host plant once decay commenced, a process that usually
began less than a day after the plant was removed from nature. They fed as primary
invaders within young shoots, leaving a distinct feeding trail of decaying tissue

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as they progressed. Each larva usually attacked at least two shoots, although
occasionally as many as five were damaged. Usually only one third instar was
found in each plant, although two or more larvae may have initially invaded the
plant.

Fully-grown larvae retreated to the base of the shoot in which they had fed and
ate a hole into the rhizome to which the shoot was attached. They then reversed
position and inserted their relatively long spiracular tubes into the cavity created
in the rhizome. This behavior probably ensured an adequate oxygen supply for
the developing pupa. The prepupal period from formation of the puparium to
appearance of the contained pupa lasted four days; the pupal period, 19-25 days
(n = 5).

With an incubation period of 5-7 days, a larval period of ca 30 days, a prepupal
and pupal period of 23-29 days, and a preoviposition period of at least two or
three days, a life cycle was completed in about two months. Overwintering oc-
curred as eggs or as newly-hatched larvae within shoots of the host plant. Larval
development accelerated in April, and larvae became fully grown in mid- to late
May. Pupation then ensued, and adults emerged during June. Adult numbers
increased again in late August, suggesting that two generations a year are produced
in northern Ohio.

Diplotaxa sp. near versicolor (Loew)

This sibling species of D. versicolor was common in northeastern Ohio in fresh-
water marshes containing its host plants, Eleocharis ohtusa and E. smallii. Nu-
merous adults of D. versicolor were swept from a small (3 x 1 m) stand of the
path rush, Juncus tenuis Willd. (Juncaceae), but were not encountered in stands
of Eleocharis, suggesting that the two species are trophically separated.

The earliest seasonal record for adults in northeastern Ohio was May 19; the
latest, October 8 (Fig. 56). Field-collected females lived 15-47 days in the labo-
ratory (n = 5); males, 9-40 days (n = 7). Mating behavior and copulation position
resembled that of D. nigripes.

Laboratory-reared females showed no preference for any particular species of
Eleocharis and readily oviposited on both E. obtusa and E. smallii as well as on
stems of the rush genus Juncus. However, eggs were found only on stems of E.
obtusa in nature, usually 2. 5-5.0 cm above the crown. The incubation period

lasted 4-6 days (n = 1 5).

Newly-hatched larvae that were placed near the tips of young shoots of Eleo-
charis quickly crawled down the stem until they encountered the leaf sheath that
surrounded the lower half of the stem. They then moved under the edge of the
sheath and continued their downward movement to the crown. At the base of the
shoot each larva paused and broke into the stem, subsequently feeding as stem
borers. Usually only one or two larvae, occasionally three, were found in any one
shoot.

Apparently there was only one generation a year in northern Ohio (Fig. 56).
Larvae that had fed heavily on young shoots during the summer months remained
in the plant tissues as second instars through the ensuing winter. However, over-
wintering larvae did not appear to be in diapause, as they quickly resumed feeding
when brought into the laboratory. Feeding larvae formed a distinct linear trail of
browned, decaying tissues as they moved along the length of the shoot.

Fully grown larvae were found near the bases of shoots during late March and
early April. Before forming puparia in late April, larvae moved to the base of the

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shoots at the crown. Prior to pupating, larvae formed a small cavity in the firm
tissue occurring where the shoots united with the roots. The larvae then reversed
direction and placed their posterior spiracular tubes in the cavity similar to the
behavior observed in D. nigripes. The pupal period for the single pupa that
produced an adult was 2 1 days.

Fifteen of the 1 6 field-collected puparia produced parasitic wasps belonging to
an undetermined species of Chaenusa (Braconidae).

Key to Third-instar Larvae

1 . Posterior spiracles at tips of short stigmatic tubes; spiracular plates lacking

spiracular hairs C. ohscuricornis

Posterior spiracles at tips of very elongate stigmatic tubes (Fig. 3); spirac-
ular plates with spiracular hairs (Fig. 52) 2

2. Spiracular plates appearing trilobed, with unbranched spiracular hairs (Fig.

52); tentoropharyngeal and hypopharyngeal sclerites separate (Fig. 44);

living larva usually greenish D. inclinata

Spiracular plates unlobed, with dichotomously branched spiracular hairs
(Fig. 32); tentoropharyngeal and hypopharyngeal sclerites partially fused;
body of living larva white 3

3. Each spiracular hair with 5-7 branches (Fig. 15) D. nigripes

Each spiracular hair with ten or more branches (Fig. 32)

D. sp. nr. versicolor

Descriptions of Immature Stages
Diplotoxa inclinata

Egg-—( Fig. 36) length 1.0-1.10 mm, greatest width 0.20-0.25 mm. White,
elongate, tapered. Ventral surface very flat, transparent, unridged. Chorion finely
striated, with numerous ridges forming a complex pattern. Micropylar end larger
than posterior end, both ends smooth and nonstriated.

First-instar Larva. — Similar to third-instar larva except in following characters.
Length 1.00-2.80 mm, greatest width 0.20-0.31 mm. White, transparent. Inter-
segmental constrictions conspicuous. Posterior spiracular plates (Fig. 50) circular,
borne at end of elongated stigmatic tubes. Spiracular openings heavily sclerotized,
probably two in number. Four unbranched, spiracular hairs, middle two twice as
long as remaining two. Cephalopharyngeal skeleton (Fig. 41): length 0.30-0.41
mm; heavily pigmented, posterior portion of tentoropharyngeal cornua hyaline;
hypopharyngeal and tentoropharyngeal sclerites fused; no parastomal bars; thin
sclerotized wing projecting dorsoposteriorly from hypopharyngeal sclerite; man-
dibles with two accessory teeth.

Second-instar Larva.— Similar to third-instar larva except in following char-
acters. Length 3.50-4.70 mm, greatest width 0.40-0.60 mm. Posterior spiracular
plate (Fig. 51) circular with three heavily sclerotized and indistinct spiracular
openings. Four unbranched spiracular hairs, all hairs equal in length. Stigmatic
scar and spiracular trabeculae, indistinct. Anterior spiracles (Fig. 37) enclosed in
transparent, ensheathing membrane; spiracles creamy white, fan-shaped, with six
marginal papillae. Cephalopharyngeal skeleton (Fig. 42) more pigmented than
that of first instar. Posterior third of ventral cornua not pigmented. Hypopha-
ryngeal and tentoropharyngeal sclerites almost entirely fused, leaving only faint
tract of fusion line between sclerites. Indication of fusion between parastomal bar

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and hypopharyngeal sclerites. Remnant of sclerotized wing extending dorsopos-
teriorly from anterior ridge of hypopharyngeal sclerite, mandibles (Fig. 43) fused
dorsally, strongly pigmented except for apical and accessory teeth. One large apical
tooth, one and possibly two smaller accessory teeth. Only one window visible.

Third-instar Larva. —Length 5.60-7.50 mm, greatest width 0.60-0.85 mm.
Similar to third-instar larva of D. nigripes except for following characters. First
segment (Fig. 39) bilobed apically, each lobe bearing one short, fleshy two-seg-
mented antenna and one maxillary palp with slightly sclerotized C-shaped ring
basally and containing 11-13 papillae. Each lobe also bearing eight sensory pa-
pillae, one dorsal and one frontal papilla, three submaxillary papillae (two anterior
and one posterior), two lateral papillae, and one genal papilla. Genal rami leading
into mouth cavity. Facial mask with numerous, large, posteriorly pointed, V-shaped
spinules. Segments 2 and 3 with 10-15 irregular rows of fine spinules, only 5-10
of these rows completely encircling segment. Segments 4-1 1 with 15-25 overlap-
ping, irregular rows of large, blunt spinules, only 10-15 of these rows continue
around segment. Perianal pad (Fig. 53) level with ventral surface; surrounded
anteriorly by compact rim of unequal-sized, irregularly-spaced, round spinules
and ventrally by three distinct rows of small rounded spinules.

Anterior spiracles (Fig. 38) large and conspicuous, creamy white, extending
perpendicularly from body, fan-shaped, with six finger-like marginal papillae, each
papilla enclosed within ensheathing membrane. Stigmatic tubes (Fig. 40) elongate,
constricted. Posterior spiracular plates (Fig. 52) posterodorsally at apices of stig-
matic tubes. Spiracular bulb and spiracular trunk creamy white, large, conspic-
uous, and trifurcating into three large lobed structures (Fig. 40), each lobe pos-
sessing dorsal spiracular opening. Spiracular trunk heavily pigmented, spiracular
openings indistinct, appearing as continuous, dark, trilobed structure. Stigmatic
scar somewhat circular, not conspicuous. Four unbranched spiracular hairs. Tra-
beculae indistinguishable.

Cephalopharyngeal skeleton (Fig. 44) length 0.64-0.78 mm. Heavily sclerotized
except for posterior ends of dorsal and ventral cornua. Hypopharyngeal and ten-
toropharyngeal sclerites separate. Parastomal bars present, fused anteriorly to
dorsal surface of hypopharyngeal sclerite. Hypopharyngeal slcerite H-shaped in
ventral view (Fig. 47). Dorsal cornua of tentoropharyngeal sclerite not joined by
bridge; floor of tentoropharyngeal sclerite faintly pigmented. Mandibles (Fig. 45)
heavily pigmented except for accessory tooth, fused dorsally, two circular win-
dows, accessory tooth with protrusion anteriorly.

Diplotoxa nigripes

Egg-—{ Fig. 10) length 1.15-1.80 mm, greatest width 0.20-0.25 mm. White.
Elongate, with micropylar end only slightly larger than posterior end, micropyle
turned upward. Ventral surface somewhat flat, opaque with one or two faint ridges.
Chorion with 11-12 large, prominent reticulations; micropylar and opposite end
without reticulations. Eggs with 4-5 diverging reticulations, one diverging towards
posterior end on dorsal surface, remainder on lateral surface.

First-instar Larva. —Similar to third instar except in following characters. Length
1.20-2.18 mm, greatest width 0.20-0.43 mm. White, integument transparent.
Posterior spiracular plates (Fig. 13) borne at distal end of elongated stigmatic
tubes. Spiracular tubes pigmented, spiracular openings indistinct, but probably
two in number, radiating out from stigmatic scar. Four spiracular hairs, each with
one main trunk bifurcating at distal end. Metapneustic. Cephalopharyngeal skel-

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eton (Fig. 4) pigmented except on posterior portion of ventral cornua and tip of
dorsal cornua, length 0.35-0.50 mm. Hypopharyngeal and tentoropharyngeal
sclerites fused, no parastomal bars. Thin sclerotized wing projecting dorsopos-
teriorly from dorsoanterior ridge of hypopharyngeal sclerite. Mandibles of one
sclerite, one accessory tooth.

Second-instar Larva. —Similar to third instar except in following characters.
Length 2.40-3.81 mm, greatest width 0.55-0.68 mm. Posterior spiracular plate
(Fig. 14) circular to oval, three sclerotized and indistinct spiracular openings
appearing as dark, trilobed structure. Four branched spiracular hairs. Stigmatic
scar not distinct. Trabeculae indistinct. Anterior spiracles (Fig. 16) creamy white,
fan-shaped, with seven marginal papillae. Transparent membrane enclosing pa-
pillae. Cephalopharyngeal skeleton (Fig. 6) pigmented except on posterior end of
ventral cornua. Length 0.58-0.68 mm. Hypopharyngeal and tentoropharyngeal
sclerites showing fusion line. Most of each parastomal bar fused with dorsal surface
of hypopharyngeal sclerite. Anterior end of bar forming knob-like protuberance
anterodorsally from hypopharyngeal sclerite. Mandibles (Fig. 8) well-pigmented
except for apical and accessory teeth, one sclerite; one large apical tooth and two
accessory teeth. Subhypostomal and ligulate sclerites semitransparent.

Third-instar Larva. —(Fig. 1) length 5.50-6.70 mm, greatest width 0.85-1.00
mm. Creamy white, integument transparent to translucent. Body elongate, cylin-
drical, tapering anteriorly from third thoracic segment. Posterior end tapering and
terminating in two elongated stigmatic tubes. Body of stigmatic tube (Fig. 3) thick,
integument extending beyond body, terminating in spiracular plates. First segment
(Fig. 2) bilobed apically, each lobe bearing short, two-segmented antenna, two
maxillary palps, each palp with slightly sclerotized basal C-shaped ring, six max-
illary papillae; six sensory papillae including one prefrontal, three frontal (two
anterior, one posterior), one or two lateral, one genal papillae. Genal rami thin,
not bifurcating, directed toward oral opening. Facial mask with numerous rows
of spinules that completely encircle larva.

Spinule bands with 20-30 short, irregular rows of spinules at anterior end of
all thoracic and abdominal segments. Spinules in abdominal segments reduced
and indistinct, forming a fine, linear fold. Spinule bands more numerous on ventral
surface, only 10-15 rows continuing around segment.

Anterior spiracles (Fig. 1 7) large, conspicuous, perpendicular to body. Creamy
white, fan-shaped with seven finger-like marginal papillae, ensheathing membrane
extending closely over papillae.

Posterior spiracular plates (Fig. 1 5) circular, located distally on stigmatic tubes.
Spiracular bulb and spiracular trunk not conspicuously enlarged or trifurcating
(Fig. 3). Three spiracular openings not distinct, appear as continuous, dark, tri-
lobed structure. Stigmatic scar present but not conspicuous. Four spiracular hairs,
each immediately bifurcating into two main branches and many secondary branch-
es. Trabeculae approximately 18 per spiracular opening.

Cephalopharyngeal skeleton (Fig. 7) length 0.88-0.95 mm. Heavily sclerotized
except transparent posterior ends of dorsal and ventral cornua. Hypopharyngeal
and tentoropharyngeal sclerites not completely fused. Parastomal bar fused with
dorsal surface of hypopharyngeal sclerite, anterior end of parastomal bar extending
anterodorsally from hypopharyngeal sclerite. Dorsal cornua not joined by bridge,
floor of tentoropharyngeal sclerite lightly pigmented. Floor of hypopharyngeal
sclerite (Fig. 5) wide, lightly sclerotized, H-shaped when viewed ventrally. Man-
dibles (Fig. 9) heavily pigmented except for accessory tooth, one posterior window.

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Not connected dorsally, large apical tooth, one blunt accessory tooth. Subhypo-
stomal sclerite beneath anterior edge of hypopharyngeal sclerite, with two large
circular windows, slightly visible when viewed internally. Ligulate sclerite semi-
transparent, Y-shaped, anterior to subhypostomal sclerite, anvil shaped when
viewed laterally.

Puparium.— (Fig. 11, 12) length 5.10 mm, greatest width 1.50 mm. Mostly light
golden-brown, dark brown to black anteriorly, stigmatic tubes black. Cuticle very
thin and transparent, developing pupa clearly visible. Elongated anterior and
posterior ends dorsoventrally flattened to form distinct lateral ridges on segments
1-4 and 9-11. Stigmatic tubes large, heavily sclerotized, extending in various
positions. Anterior spiracles poorly developed, located anteriorly on dorsal ce-
phalic cap, fan-shaped and silver-white, with seven marginal papillae. Posterior
spiracular plate rounded, spiracular openings indistinct. Perianal pad depressed.
Puparium retaining spinule bands of third-instar larva.

Diplotoxa sp. near versicolor

Egg. -“(Fig. 29) length 0.80-1.00 mm, greatest width 0.12-0.15 mm. White,
elongate, and tapered. Ventral surface somewhat flattened, transparent and un-
ridged. Chorion striated, with ridges diverging and converging especially at ends
and on the dorsal surface. Micropylar end larger than posterior end, both ends
smooth and nonstriated.

First- instar Larva.— Similar to third instar except in following characters. Length
0.90-2.20 mm, greatest width 0.15-0.21 mm. White, integument transparent.
Posterior spiracular plates (Fig. 30) at ends of elongated stigmatic tubes. Spiracular
trunks deeply pigmented. Spiracular openings not distinguishable, probably two
in number positioned to form dark, heart-shaped structure. Four spiracular hairs,
each with single long trunk terminating in either three or four shorter branches.
Larva metapneustic. Cephalopharyngeal skeleton (Fig. 24) length 0.25-0.35 mm,
pigmented except for hyaline posterior portion of ventral cornua and tip of dorsal
cornua. Hypopharyngeal and tentoropharyngeal sclerites fused, parastomal bars
lacking. Thin, sclerotized wing projecting dorsoposteriorly from dorsoanterior
ridge of hypopharyngeal sclerite. Mandibles with one accessory tooth.

Second-instar Larva. — Similar to the third instar except in following characters.
Length 3.20-4.00 mm, maximum width 0.45-0.60 mm. Posterior spiracular plate
(Fig. 31), circular to oval, three indistinct spiracular openings on brown, trilobed
structure. Four, multibranched spiracular hairs, each with short trunk that forks
into two main branches that each terminate in 2-6 shorter branches. Stigmatic
scar and trabeculae indistinct. Anterior spiracles (Fig. 33) creamy white, fan-
shaped, with five marginal papillae; with transparent ensheathing membrane around
papillae. Cephalopharyngeal skeleton (Fig. 25) more pigmented than in first instar.
Posterior ends of ventral cornua not pigmented. Hypopharyngeal and tentoro-
pharyngeal sclerites almost entirely fused. No dorsal wing extending from hy-
popharyngeal sclerite. Parastomal bars fused to hypostomal sclerite except for
anterior end which appears as thick, anterodorsal protuberance. Mandibles well-
pigmented except for apical and accessory teeth; one large apical tooth and two
accessory teeth; 4-6 circular windows. Subhypostomal and ligulate sclerites poorly
pigmented.

Third-instar Larva.— { Fig. 18, 19) similar to third-instar larva of D. nigripes
except in following characters. Length 4.50-5.50 mm, maximum width 0.70-0.95
mm. Maxillary palp of segment 1 (Fig. 20) with 7-1 1 papillae. Facial mask with

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numerous large, V-shaped spinules. Segment 1 with 20-25 compact, short, irreg-
ular rows of spinules at anterior end, spinule rows completely encircling segment.
Segments 2 and 3 with 1 5-20 thick, irregular rows of large, blunt spinules that
encircle anterior end of each segment. Spinule bands dark due to slight scleroti-
zation or impregnation by dirt. Segments 4-8 -with 15-20 short, irregular rows of
V-shaped spinules encircling segments. Spinule bands on dorsal surface more
linear, fine, and composed of smaller spinules. Ventral surface of segments 9 and
10 containing approximately 20-30 short, irregular, fine rows of spinules, only
7-1 5 rows encircling segments. Segment 1 1 with reduced spinule bands, individual
spinules indistinct, forming fine lines, no spinule bands on dorsal surface. Perianal
pad (Fig. 35) with three fine rows of spinules anteriorly and 3-5 rows posteriorly,
cluster of irregularly spaced large, U-shaped spinules between anterior and pos-
terior rows.

Anterior spiracles (Fig. 34) large, conspicuous, creamy white, extending per-
pendicularly from body, fan-shaped, with five finger-like marginal papillae, each
papilla enclosed within ensheathing membrane. Stigmatic tubes elongate. Pos-
terior spiracular plates (Fig. 32) posterodorsally at end of stigmatic tubes. Stigmatic
bulbs and spiracular trunks normal. Spiracular trunks heavily pigmented, spirac-
ular openings indistinct, appearing as continuous, dark, trilobed structure. Stig-
matic scar somewhat ovoid. Four, multibranched spiracular hairs, each with main
trunk diverging into three or four branches that terminate in many smaller branch-
es. Trabeculae indistinguishable.

Cephalopharyngeal skeleton (Fig. 26) length 0.70-0.90 mm, greatest width 0.15
mm. Heavily sclerotized except for posterior end of ventral cornua. Hypopha-
ryngeal and tentoropharyngeal sclerites partially fused. Parastomal bars fused with
dorsal surface of hypopharyngeal sclerite, anterior end of parastomal bar extending
out anterodorsally from hypopharyngeal sclerite. Hypopharyngeal sclerite H-shaped
when viewed ventrally (Fig. 21). Floor of tentoropharyngeal sclerite (Fig. 21)
lightly pigmented except for transparent posterior end, dorsal cornua not joined
by bridge. Mandibles (Fig. 23) heavily pigmented except for accessory tooth, one
circular window. Subhypostomal sclerite beneath anterior edge of hypopharyngeal
sclerite, somewhat heart-shaped. Slightly visible when viewed laterally, ligulate
sclerite V-shaped, elongate, anterior of subhypostomal sclerite, semitransparent.
No dentate sclerites.

Puparium. —(Fig. 27, 28) similar to puparium of D. nigripes except in following
characters. Length 2.80-3.30 mm, greatest width 0.75-0.90 mm. Gold to reddish
brown, dark brown cephalic cap. Cuticle slightly transparent. Anterior spiracles
poorly developed, anterior on dorsal cephalic cap, fan-shaped with five marginal
papillae. Spinule bands appearing ridge-like, completely encircling segments, be-
coming coarse and deeply grooved at anterior and posterior ends of puparium.
Intersegmental constrictions distinct. Segment 1 partially invaginated. Posterior
stigmatic tubes flattened dorsoventrally, deeply pigmented posteriorly. Spiracular
plates black, oblong, spiracular openings not visible. Anal slit distinct, slightly
depressed.

Discussion

This study suggests that the larval stages of many, if not all, species of the genus
Diplotoxa are associated with spike-rushes of the genus Eleocharis. Adults of
seven species were collected in Ohio or Montana from stands of Eleocharis
(Wearsch, 1968), eggs of at least five species were discovered on Eleocharis stems,
and larvae of four species were found feeding in stems or rhizomes.

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Eleocharis plants are infrequently attacked by other insect larvae except for a
few species of Lepidoptera and Coleoptera. Eleocharis species grow in marshy
areas, along with other sedges and rushes, frequently form extensive pure stands,
and are relatively undisturbed by human activity. The many sprouts, which are
continually produced at the base of the plant during the warm season, and the
thick, branched rhizome system offer very succulent food sources. Meristematic
tissue in monocots such as Eleocharis is basal in position within the stem, and
typically has a higher energy content than nonmeristematic tissue (Hirose et al.,
1989), which explains the concentration of feeding by chloropid larvae at the base
of the stem or in rhizomes.

Primary invaders (larvae feed on living plant tissue) of Eleocharis stems are
best represented by C. obscuricornis, D. sp. near versicolor, and D. nigripes. In
contrast, the morphology of the larvae of D. inclinata includes some features of
a secondary invader (larva feeds on tissue damaged by a primary invader) and
some of a primary invader. A comparison of the cephalopharyngeal skeleton of
various species of primary and secondary invaders in the family Chloropidae
indicates that secondary invaders possess separate tentoropharyngeal and hypo-
pharyngeal sclerites, whereas these sclerites are fused in primary invaders.

Although the overall appearance of chloropid larvae is similar, species are
distinguishable in size, color, and appearance of the stigmatic tubes and spiracular
hairs. Chlorops larvae differ from those of Diplotoxa in having spiracular openings
at the distal ends of sclerotized extensions that protrude from short stigmatic
tubes, and in lacking spiracular hairs. In contrast, larvae of Diplotoxa possess very
elongate stigmatic tubes and have long spiracular hairs.

Identifying species of Diplotoxa from preserved larval specimens is difficult,
although living larvae of D. inclinata are easily distinguished from those of the
other species we studied by their greenish color. The bulb and internal stigmatic
chamber of the stigmatic tubes of this species are enlarged, have a dark creamy
color, and branch into three prominent lobes. The spiracular plate possesses a
trilobed structure bearing the spiracular slits and has only four nonbranching
spiracular hairs. Diplotoxa sp. near versicolor and D. nigripes have semitransparent
internal stigmatic chambers, with the spiracular slits opening onto the flat spi-
racular plate. Both have a complex network of spiracular hairs.

The anterior spiracles of all three species of Diplotoxa consist of a fan-shaped
structure with apical papillae. The number of papillae is relatively constant and
distinctive for each species. Diplotoxa sp. near versicolor, the smallest of the three
species, possesses five papillae; D. inclinata, the next largest, has six; and D.
nigripes, the largest, has seven.

The cephalopharyngeal skeletons of D. sp. near versicolor and D. nigripes differ
in only a few structures, whereas both differ markedly from that of D. inclinata.
The skeletons of the first-instar larvae of D. nigripes and D. sp. near versicolor
are almost identical, having a wing projecting from the anterior end of the hy-
popharyngeal sclerite, but the skeleton of D. nigripes is much larger. In the second
instar of both species the wing of the hypopharyngeal sclerite is reduced or absent,
and a knob-like structure is present. This knob probably is the anterior portion
of the parastomal bar, the remainder of which has fused with the dorsal surface
of the hypopharyngeal sclerite. The hypopharyngeal and tentoropharyngeal scler-
ites are not completely fused in the third instars of both species, suggesting that
larvae of these species originally fed as secondary invaders of damaged stems.
Fusion of these two sclerites probably gives greater support and rigidity to the
cephalopharyngeal skeleton.

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Diplotoxa inclinata apparently became a primary invader very recently, as the
tentoropharyngeal and hypopharyngeal sclerites in the third instar are still sep-
arate. However, the cephalopharyngeal skeleton of the first instar resembles that
of D. sp. near versicolor and D. nigripes. The projecting wing of the hypostomal
sclerite is reduced in the second instar, and the parastomal bar, although evident,
has fused anteriorly with the base of the projecting wing of the hypopharyngeal
sclerite. In the third instar, this fusion is more nearly complete but not to the
degree seen in third instars of the other two species. The third instar of D. inclinata
has the mandibles fused dorsally, which may give greater rigidity for feeding upon
the hard tissues of the rhizome.

The shape of the posterior spiracles is helpful in distinguishing among larvae
of the four species. Two spiracular openings occur in the first instar, and three in
the second and third instars. The spiracles in D. nigripes and D. sp. near versicolor
are similar, although D. nigripes is larger, and all three Diplotoxa species possess
spiracular hairs. The first instar has four large hairs with one or two terminal
branches. The hairs of the second instars are more complex: each hair has a main
trunk that divides immediately into two large branches, each of which in turn
divides two or more times. The third instar possesses a very fine network of four
sets of spiracular hairs. The basal trunk of each hair radiates immediately into
three or four main branches, each of which branches again to form a fine network.
In D. inclinata, in contrast, the spiracular hairs of all three instars are reduced
and unbranched. The first instar has two very long and two shorter hairs, whereas
in the second and third instars the four hairs are equally long.

In C. obscuricornis the posterior spiracles are reduced, sclerotized structures
with three spiracular slits but no spiracular hairs. Oldroyd (1964) reported that
some stem-boring species of Chlorops have become so well-adapted to their en-
vironment that the posterior spiracles are vestigial. They absorb oxygen through
the cuticle. Larvae of C. obscuricornis show a tendency toward reduction of the
posterior spiracles.

Larvae of all four species, whether feeding on stems or rhizomes, inserted their
stigmatic tubes into living tissue or air spaces before forming puparia. This prob-
ably is a mechanism for ensuring sufficient oxygen for pupal development. Under
natural conditions, pupae in living plants developed normally and produced adults.
When cut plants containing puparia were brought into the laboratory, the pupae
died shortly after the plants began to decay. Pupae that were removed from stems
and placed upon peat moss continued to develop and eventually produced adults.

Literature Cited

Balachowsky, A., and L. Mesnil. 1935. Les Insectes nuisibles aux Plantes cultivees. Leurs Moeurs,
Leur Destruction. Vol. 1. Establissements Busson, Paris, 1137 pp.

Braun, E. L. 1967. The Vascular Flora of Ohio. Vol. 1: The Monocotyledoneae. The Ohio State
University Press, Columbus, viii + 464 pp.

Dennis, E. B. 1961. New or uncommon plant diseases and pests. Gout-fly damage in spring wheat.
Plant Pathology, 19:166.

Ferrar, P. 1987. A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha.
Entomonograph, Volume 8, parts 1 and 2. E. J. Brill/Scandanavian Science Press, Leiden and
Copenhagen, 907 pp.

Frew, J. G. H. 1923a. On the larval anatomy of the gout-fly of barley ( Chlorops taeniopus Meig.)
and two related acalyptrate muscids, with notes on their winter host-plants. Proceedings of the
Zoological Society of London, 192:783-821.

— . 1 923 b. On the morphology of the head capsule and mouth parts of Chlorops taeniopus Meig.

(Diptera). Journal of the Linnaean Society of London, 35:399-403.

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Gleason, H . A. 1963. The New Britton and Brown Illustrated Flora of the U nited States and Adj acent
Canada. Vol. 1. New York Botanical Garden, New York. Ixxv + 482 pp.

Goodliffe, F. D. 1939. A joint infestation of barley by Lasiosina cinctipes Mg., together with Chlorops
taeniopus Mg. (Diptera: Chloropidae) and their hymenopterous parasites. Society of British En-
tomology, 1:248-252.

. 1942. Studies on insects bred from barley, wheat, maize and oats. Bulletin of Entomological

Research, 32:309-325.

Hirose, T., M. J. A. Werger, and J. W. A. Van Rheenen. 1989. Canopy development and leaf
nitrogen distribution in a stand of Carex acutiformis. Ecology, 70:1610-1618.

Nye, I. W. B. 1958. The external morphology of some of the dipterous larvae living in the Gramineae
of Britain. Transactions of the Royal Entomological Society of London, 1 10:41 1-487.

Oldroyd, H. 1964. The Natural History of Flies. W. W. Norton and Company, New York City,
xiv + 324 pp.

Rogers, T. P., B. A. Foote, and J. L. Todd. 1991. Biology and immature stages of Chlorops certimus
and Epichlorops exilis (Diptera: Chloropidae), stem-borers of wetland sedges. Journal of the New
York Entomological Society, 99:664-683.

Sabrosky, C. W. 1965. Chloropidae. Pp. 773-793, in A Catalog of the Diptera of America North
of Mexico (A. Stone, C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson, eds.), United
States Department of Agriculture Agricultural Handbook 276, iv + 1696 pp.

— — . 1987. Chloropidae. Pp. 1049-1067, in Manual of Nearctic Diptera. Vol. 2 (J. F. McAlpine,
B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood, eds.), Research
Branch Agriculture Canada Monograph 28.

Scoggan, H. 1978. The Flora of Canada. Part 2. National Museum of Canada Publications in
Botany, 7:91-545.

Spencer, K. A. 1977. A revision of the New Zealand Chloropidae. Journal of the Royal Society of
New Zealand, 7:433-472.

Todd, J. L., and B. A. Foote. 1987. Resource partitioning in Chloropidae (Diptera) of a freshwater
marsh. Proceedings of the Entomological Society of Washington, 89:803-810.

Valley, K., T. Wearsch, and B. A. Foote. 1969. Larval feeding habits of certain Chloropidae
(Diptera). Proceedings of the Entomological Society of Washington, 71:29-34.

Wearsch, T. 1 968. Biology and immature stages of certain Chloropidae (Diptera). M.S. thesis, Kent
State University, Kent, Ohio, 96 pp.

Wendt, H. 1968. Faunistisch-okologische Untersuchungen an Halmfliegen der Berliner Umgebung
(Dipt. Chloropidae). Deutsche Entomologische Zeitschrift (N.F.), 15:49-105.

Wooten, J. W., and S. D. Elakovich. 1991. Comparisons of potential allelopathy of seven freshwater
species of spikerushes (. Eleocharis ). Journal of Aquatic Plant Management, 29:12-15.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 3, Pp. 215-237

16 September 1994

BEADS AND PENDANTS FROM TRANTS, MONTSERRAT:
IMPLICATIONS FOR THE PREHISTORIC LAPIDARY
INDUSTRY OF THE CARIBBEAN

David R. Watters
Associate Curator, Division of Anthropology

Richard Scaglion1

Research Associate, Division of Anthropology

ABSTRACT

Analysis of a collection of lithic beads and pendants from Montserrat suggests that the Trants site
may have been a prehistoric lithic bead manufacturing center specializing in camelian beads. Camelian
is represented in all stages of the manufacturing process. The absence of known local sources for
camelian, the relatively low proportion of finished camelian beads, the relatively high proportion of
broken to unbroken bead “blanks” (indicative of breakage sustained during the manufacturing process),
and the relative homogeneity of bead blanks compared to finished beads support the hypothesis of a
specialized local manufacturing industry. The presence of a variety of other exotic (nonlocal) materials
in finished beads and pendants, including some whose likely sources are in South America, is further
evidence of long-distance trading by the early Saladoid residents of Trants. Although the S. W. Howes
collection of the Museum of the American Indian, Heye Foundation, was surface-collected, the analysis
presented here is consistent with more recent findings of lithic beads from excavated contexts.

Introduction

The Trants site, further described in Watters (1980) and Petersen and Watters
(1991), is located on the windward (east) coast of Montserrat, one of the smaller
volcanic islands in the northern Lesser Antilles (Fig. 1). Montserrat is a moun-
tainous island with a total land area of about 98 km2. Trants is situated on the
only sizeable stretch of relatively flat land on the windward coast. Diagnostic
artifacts of the Saladoid ceramic series, including white-on-red painted and zone-
incised-crosshatch sherds, are indicative of an early Ceramic Age population
movement onto Montserrat. Two early radiocarbon dates from Trants, 2430 ±
80 and 2390 ± 90 B.P. (ca. 480 and 440 B.C. uncorrected), suggest an onset of
Saladoid migration into the Lesser Antilles earlier than previously assumed (Pe-
tersen and Watters, 1991).

This paper describes a collection of surface-collected lithic beads and pendants
from Trants in the Museum of the American Indian. Analysis of this collection,
coupled with an analysis of excavated specimens (Bartone and Crock, 1991)
suggests that lithic beads may have been manufactured at Trants. Camelian in
particular is amply represented in all stages of bead manufacture. The presence
of finished beads and pendants of other nonlocal materials, including some whose
likely sources are in South America, is suggestive of long-distance trading by the
early Saladoid residents of Trants.

1 Current address: Department of Anthropology, University of Pittsburgh, Pittsburgh, Pennsylvania
15260.

Submitted 1 November 1993.

215

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vol. 63

Greater Antilles

Northern

Lesser

Antilles

1 00 km

Barbuda

Kitts

Antigua

MONTSERRAT

Fig. l.—The Trants prehistoric site is located on the east coast of Montserrat in the northern Lesser
Antilles.

The Howes Collection

In 1924, the Museum of the American Indian (MAI), Heye Foundation, in New
York City, cataloged a collection of some 1073 artifacts from Montserrat. Seymour
Wylde Howes (Fig. 2), a recipient of the O.B.E. (Order of the British Empire) for
his innovations in agricultural practices, assembled this collection while manager
of the Trants Estate (Fig. 3) where he lived until his death in 1 927. Queenie Howes
van der Veer, Howes’ daughter, distinctly remembers her father collecting beads
and pieces of pottery from newly-tilled fields (personal communications, 1 979—
1993). S. W. Howes personally searched whatever field was being worked each
day and also encouraged the workers to bring him artifacts they found, for which
he paid them a nominal amount. Bruce Howes, S. W. Howes’ son, recollects
(personal communication, 1993) that the majority of the beads were recovered
from Pigeon Field, an area that corresponds for the most part to the fields num-
bered 1, 2, 3, 8, and 9 by Petersen and Watters ( 1 99 1 :fig. 1).

Queenie van der Veer and Bruce Howes are confident that all of the objects in
their father’s collection come from the Trants site. Although the possibility exists
that workers may have found beads elsewhere and given them to Howes in ex-
change for the modest reward that he paid them, it is important to note that no
lithic beads have been recovered from any other prehistoric site on Montserrat
(Watters, 1980).

The arrival of the Howes collection at the MAI created considerable interest.
The beads were particularly noteworthy. Harrington (1924) described them in the
first published article dealing with artifacts from the Howes collection. Most of
the beads were stored at the MAI Research Branch in the Bronx, where our research
was conducted in 1985 and 1991. Forty-six beads were on display and were not
available for study.

Table 1 lists the 1053 items currently present in the S. W. Howes collection

1994

Watters and Scaglion — Frants Beads and Pendants

217

Fig. 2.— Seymour Wylde Howes, O.B.E., the person who assembled the Howes collection.

according to four general categories: groundstone, shell, pottery, and beads and
related artifacts. There are 502 beads and related objects in the Howes collection.
Although the MAI has three other collections from Montserrat, the Huckerby
collection contains the only other stone bead (catalog number 8/6068), which we
also examined. Our study thus includes all lithic beads and related materials from
Montserrat available for study at the MAI.

The 503 beads and related artifacts were classed using a scheme based on an
object’s substance, configuration, condition, and perforation. The first classificatory
level, substance, identifies the composition (Table 2). There are 494 lithic (stone),
one ceramic, one shell, and seven glass objects. Only the 494 lithic objects are
treated in this paper. The second classificatory level, configuration, includes form,
shape, and geometry. Artifacts were assigned to one of four basic forms: bead,
pendant, blank, and raw material. A relationship between a fifth basic form,
“roller” (n = 4), and a bead and pendant manufacturing industry could not be
established; therefore, these artifacts are not considered here.

There is some ambiguity in the way that the terms “bead” and “pendant” have
been employed in popular usage. For the purpose of this study, a “bead” is defined
as a single-perforated object that is symmetrical around a longitudinal axis running

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vol. 63

Fig. 3.— Trants Estate House as it appeared in the 1920s when Queenie and Bruce Howes lived there
as children.

through the center of its perforation. In contrast, a '‘pendant” is an object that is
not symmetrical with regard to any of the axes running through any of its per-
forations. Thus, a pendant’s center of gravity is offset from the perforation through
which it presumably was strung, causing it to "hang” or "be suspended” with an
inherent orientation (or orientations) relative to its axis (or axes). Figure 4, which
is adapted from Durand and Petitjean Roget (1991:68), illustrates the distinction
between beads and pendants.

Eleven objects that might be classed as "beads,” in that they are symmetrical
around a longitudinal axis running through the center of a perforation, presented

Table 1 .—Expected and observed artifacts.

Artifact category

Expected quantities2
based on catalog .
cards

Observed quantities

Discrepancy

Research branch

Exhibit

Total

Howes Collection

Groundstone

162

162b

Shell

90b

91b

Pottery

252

246b

252b

Beads

551

502c

548c

Total

1056

1000

1053

Huckerby Collection

Beads

Total beads

503d

549d

8 Excludes objects exchanged with University Museum, University of Pennsylvania.
b Groundstone, shell, and pottery specimens in the Research Branch were not all counted; expected
and observed quantities are presumed to be equal; if they are not, discrepancies could pertain.
c Specimens counted.
d Howes and Huckerby collections.

1994

Watters and Scaglion— Trants Beads and Pendants

219

Table 2.— Levels of classification for the 503 artifacts studied.

Classificatory

Classificatory level 4

level 2 Classi- (perforation)

Classificatory (configuration) ficatory —

(substance)

Form

Shape

Geometery

(condition)

her

Stage

gory

lithic

bead

regular

rounded

intact

finished

303

lithic

bead

regular

faceted

intact

finished

lithic

bead

regular

rounded

broken

finished

lithic

bead

regular

faceted

broken

finished

lithic

bead

regular

rounded

intact

unfinished

lithic

bead

regular

faceted

intact

unfinished

lithic

bead

regular

rounded

broken

unfinished

lithic

bead

regular

faceted

broken

unfinished

lithic

bead

regular

rounded

intact

finished

lithic

pendant

—

intact

—

lithic

bead

regular

rounded

broken

finished

lithic

pendant

—

broken

—

lithic

bead

irregular

—

intact

finished

lithic

other

ir-

—

lithic

blank

regular

rounded

intact

—

lithic

blank

regular

faceted

intact

—

lithic

blank

regular

rounded

broken

—

lithic

blank

regular

faceted

broken

—

lithic

blank

—

intact

—

lithic

raw material

—

ceramic

bead

regular

rounded

intact

finished

shell

bead

regular

rounded

intact

finished

glass

bead

regular

rounded

intact

finished

glass

bead

regular

faceted

intact

finished

lithic

roller

regular

rounded

intact

Total

503

a classificatory problem because they have a second perforation, typically trans-
verse and often nearer to one “end” than the other. It is possible that these objects
were strung as beads, with the additional perforation used for the insertion of
objects such as feathers (cf. Fewkes, 1922:232). These double-perforated artifacts
(Fig. 5) were eliminated from the analysis.

For the most part, the beads and pendants are finished products in that the
stages of manufacture are complete and the object is ready to be worn. In some
cases, however, earlier stages in the manufacturing process are represented by
objects designated as “blanks” and “raw materials.” Blanks (Fig. 6), which range
from those merely “roughed out” to those lacking only perforation, illustrate
numerous stages in the manufacturing process.

Other aspects (shape and geometry) of artifact configuration were adapted from
Horace C. Beck’s article on bead classification and nomenclature. Page and figure
citations in the present paper refer to the 1981 reprint edition of Beck’s 1928
work. Beck’s (1981) figures 1, 2, and 3 and plates I, II, III, IV, were particularly
useful.

Following Beck (1981:4), beads and blanks were classed as regular or irregular.
Those with regular shape were then sorted by geometry, as rounded or faceted,
and by condition, intact or broken. The final classificatory level, perforation , has
been explained above.

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vol. 63

Fig. 4.— A prehistoric necklace from Guadeloupe displaying both beads and pendants (adapted from
Durand and Petitjean Roget, 1 99 1 :plate II).

Descriptive Analysis

For the purposes of this study, a coding scheme of 17 variables was devised,
not all of which pertained to every artifact category. In fact, “material” was the
only variable that could be coded for all artifacts.

Rocks and minerals comprising the beads and pendants were first discussed by
Harrington (1924), who concentrated on the “semi-precious stones,” and sub-
sequently by Ball (1941), in a survey of gem and ornamental stone mining in the
New World. Harrington’s material attributions (Table 3) appear to have been
based on his own knowledge and judgment rather than mineralogical or gemo-
logical tests. Judging by his virtually identical materials list, Ball (1941:17 and
table 1) apparently relied on Harrington’s identifications. The MAI catalog cards
for the Howes collection list a greater variety of materials (Table 3).

Neither catalog numbers nor artifact illustrations are provided by Harrington
and Ball; therefore, it is not possible to assign their attributions to specific artifacts.
Our overall material categories are similar to theirs, but we were unable to verify
examples of two of their identifications: lapis lazuli and cannel coal. In this paper,

1994

Watters and Scaglion— Trants Beads and Pendants

221

Fig. 5.— The 11 double-perforated objects having longitudinal and transverse perforations. A, 12/
7035(001); B, 12/7052(001); C, 12/7033(001); D, 12/7120(078); E, 12/7031(003); F, 12/7031(002);
G, 1 2/7031 (00 1);H, 12/7032(002); 1, 12/7032(001); J, 1 2/7030(00 1);K, 12/7028(001). A-D are diorite
(material 25); E-J are unidentified (material 22), possibly quartzite; K is unidentified (material 16).

our material attributions accompany the MAI catalog numbers (e.g., 12/701 1) in
the captions of artifact photographs.

Table 4 presents a frequencies analysis for 29 categories of lithic materials. Five
materials (Table 5) tested and identified by the GIA Gem Trade Laboratory
include camelian, amethyst, turquoise, nephrite jade, and quartzite (Gemological
Institute of America, 1985). Our use of specific names reflects reasonable confi-
dence in identification. Less secure attributions are qualified as probable or pos-
sible', unidentified materials are described.

In order to present a more consolidated table, the “certainty levels” were col-
lapsed, combining the “reasonably confident” and “probable” attributions under
their appropriate material labels, and combining “possible” and “unidentified”
as simply “unidentified.” Results are presented in Table 6.

Raw Materials

The simplest category of objects, raw materials, represents the “first stage” of
the manufacturing process. These specimens are unformed pieces of materials
showing no clear evidence of shaping (Fig. 7). A breakdown of these objects by
material is presented in Table 7, which shows that 90% of the specimens are
camelian.

Read Blanks

Bead blanks are objects whose shape and geometry resemble those of beads,
but which have no perforations or unfinished perforations (Fig. 7). The materials
represented by these 72 objects are grouped in Table 8. It should be noted that

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Annals of Carnegie Museum

vol. 63

Fig. 6. —Materials represented by bead blanks. A, camelian (material 1; 12/701 1), the most common;
B, unidentified (material 31; 12/7041), possibly feldspar; C, diorite (material 25; 12/7042); D, quartz
(material 55; 12/7051).

1994

Watters and Scaglion— T rants Beads and Pendants

223

Table 3.— Previous material attributions of the Howes collection beads and pendants.

Material attribution

Harrington

(1924)

Ball

(1941)

MAI catalog cards*

Camelian

x 12/7011

Amethyst

x 12/7016

Chalcedony

x 12/7040

Quartz (rock) crystal

x 12/7050

Turquoise

x 12/7026

Cannel coal

x 12/7063

Lapis lazuli

Nephrite

Jade

x 12/7073

Jadeite

Agate

Serpentine

x 12/7023

Feldspar

x 12/7029

Altered granitic rock

x 12/7033

Quartz

x 12/7039

Petrified wood

x 12/7062

Chalcedony/calcite/gypsum

x 12/7051

Marble

x 12/7060

Soap stone

x 12/7069

Green (amazon) stone

x 12/7064

White stone

x 12/7030

Brown stoneb

x 12/7045

Gray stone

x 12/7083

a In some instances, only one of several catalog card numbers is used to exemplify the material
attribution.

b This brown stone bead actually is a ceramic bead.

69.4% of bead blanks are camelian. The relatively high proportion of broken to
unbroken bead blanks (24/48) could be interpreted as breakage sustained during
the manufacturing process.

Beads

Most specimens in the MAI collections are completed beads. Following Beck
(1981), they were divided into regular rounded and regular faceted subtypes.

Regular Rounded Beads. — Regular rounded beads comprise 360 of the 503
specimens or 71.6% of the total objects. At least six materials are represented
among the 354 lithic regular rounded beads (Table 9), but the vast majority of
these are diorite (71.2%). Only 1.1% of the finished regular rounded beads are
camelian. Since 90.0% of raw materials and 69.4% of bead blanks are camelian,
a much greater proportion of camelian in the finished objects might have been
expected.

The lengths of the beads are highly variable (Fig. 8, 9, 10). Interestingly, the
lengths of the blanks ( n = 50; mean = 23.84; SD = 8.20; coefficient of variation
= 0.34) are more homogeneous than the lengths of the finished beads (n = 320;
mean = 13.25; SD = 6.99; coefficient of variation = 0.53). This homogeneity of
bead blanks is suggestive of a specialized local manufacturing industry. Regular
rounded beads should ideally be symmetrical; thus, end diameters should be
virtually identical on individual beads. The fact that the mean end diameters in
this sample do not differ significantly (t = -0.317, P > 0.37) suggests relative
precision in the manufacture of these particular objects.

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vol. 63

Table 4.— Lit hie materials: frequencies.

Material
category no.

Description or identification

Fre-

quency

Per-

centage

diorite

260

53.1

camelian

15.5

translucent, high gloss; probably rock crystal or glass

7.8

amethyst

4.5

green and blue-green with reflective flecks; probably adventurine

2.7

whitish gray (some banded), opaque, some luster

2.4

turquoise

2.2

white, opaque, no luster; possibly feldspar

2.0

white, little luster, no apparent black; probably diorite

1.6

rock crystal (translucent, medium gloss)

1.4

white, opaque, no luster; possibly quartzite

1.2

white, opaque, no luster; possibly limestone

1.0

light green, low gloss

0.6

greenish gray, medium gloss; probably nephrite or jadeite

0.6

white, opaque, no luster

0.4

light green, translucent

0.2

green, brown patina, no luster; possibly turquoise

0.2

black and white with red inclusions; probably diorite

0.2

pink, dark inclusions, high gloss

0.2

black, fine grain, medium luster

0.2

rock crystal (translucent, high gloss)

0.2

rock crystal (transparent, high gloss)

0.2

dark green, striated; probably serpentine

0.2

green, medium luster; probably nephrite or jadeite

0.2

nephrite

0.2

dark gray, slight luster; probably serpentine

0.2

light pink, opaque

0.2

red, opaque; possibly camelian

0.2

greenish blue with brown inclusions, opaque, no luster

0.2

Totals

490

100.0

The general shape categories established by Beck (1981:5-6, plate 1) were used
to describe the transverse sections of beads (Fig. 1 1). Regular rounded beads and
bead blanks are of only two basic types, circular and elliptical, with the over-
whelming majority (97.1% of bead blanks and 99.7% of finished beads) being
circular. The infrequency of the elliptical shapes in both beads and blanks suggests
that the few specimens merely represent errors in manufacture. The profiles or

Table 5 . — Gemologically verified materials in the Howes collection.

Material identification8

Specimen number”

Remarks”

Camelian

1 2/701 2(005)d

material 1

Amethyst

12/7017(001)

material 4

Turquoise

12/7026(004)

material 13

Turquoise

12/7065(001)

material 13

Nephrite jade

12/7084(001)

material 79

Quartzite

12/7031(001)

material 22

a GIA Gem Trade Laboratory, Gem Identification Report #1926912, 19 April 1985.
b MAI catalog number followed by the piece number (in parentheses) assigned by Watters.
c Material numbers assigned by Watters (see Table 4).
d The GIA report erroneously lists this as 12/7022.

1994

Watters and Scaglion — T rants Beads and Pendants

225

Table 6. —Lit hie materials: collapsed frequencies.

Material category nos.

Probable material

Frequency

Percentage

25, 28, 34

diorite

269

54.9

camelian

15.5

64, 55, 52, 61

rock crystal

9.6

amethyst

4.5

adventurine

2.7

turquoise

2.2

70, 76, 79

jade group

1.0

67, 82

serpentine

0.4

89, 31, 22, 46, 49, 73, 16, 19, 37, 43, 85, 95, 98

unidentified

9.2

Totals

490

100.0

Fig. 7.— Camelian (material 1) objects. A, four raw material specimens (12/7013); B-C, eight bead
blanks in various stages of the manufacturing process (12/7011).

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vol. 63

Table 7 .—Raw materials: frequencies.

Material category nos.

Probable material

Frequency

Percentage

camelian

90.0

55, 61

rock crystal

10.0

Totals

100.0

“side views” of beads were also based on Beck (1981:7-9, plates II and III).
Convex (barrel) and straight (cylinder) profiles constitute the overwhelming ma-
jority (97.7%) of regular rounded beads (Table 10, Fig. 12) and constitute exclu-
sively the bead blanks.

Regular Faceted Beads. — Whereas the perimeter of the transverse section of a
regular rounded bead is curvilinear, the perimeter of a regular faceted bead is a
polygon. Most transverse sections of regular faceted beads are hexagonal (Fig. 1 1 ,
Table 1 1). The two regular faceted bead blanks are pentagonal and “polygonal,”
the latter a category used by Beck (1981:6). It is possible that faceted beads
(particularly those whose transverse sections are multisided polygons) are not
really a “type” of bead, but rather a “stage” in the manufacturing process and /
or are poorly-made regular rounded beads. Data on the longitudinal sections or
“side view profiles” of regular faceted beads are compiled in Table 12, and the
material frequencies analysis is presented in Table 13. Two blanks, both broken,
are rock crystal and an unidentified material, although the most common material
for both regular faceted and regular rounded beads is diorite. However, faceted
beads made of camelian are proportionally more common.

Pendants

Pendants typically have distinctive forms, very often being zoomorphic rep-
resentations (Fig. 1 3). A materials breakdown shows that the “jade group” (jadeite,
nephrite) constitutes some 41.6% of the sample (Table 14). Pendants are the only
category of artifacts manufactured from jade group materials. (Pendants on display
at the MAI Manhattan facility, although not part of this study, also had a high
proportion of green-colored stones possibly belonging to the jade group.) Tur-
quoise is also heavily represented in studied pendants (25%) as compared with,
for example, regular rounded beads (2%). Clearly, there is a high proportion of
“exotic” materials represented in pendants. The only pendant blank with a ma-
terial attribution is serpentine. It and a finished pendant are the only represen-
tatives of serpentine among all the lithic objects.

Table %.—Bead blanks: frequencies.

Material category nos.

Probable material

Frequency

Percentage

camelian

69.4

diorite

9.7

unidentified

9.7

64, 55

rock crystal

9.7

unidentified

1.4

Totals

100.0

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Watters and Scaglion— Trants Beads and Pendants

227

Fig. 8.— Diorite (material 25) beads, the most common material for regular rounded beads. A, lon-
gitudinal and transverse views of the same short to disc beads (12/7122); B, longitudinal view of
standard to short beads (12/7121) (see Fig. 1 1).

Fig. 9. —Size range in regular rounded quartz (rock crystal) beads (material 64; 12/7056 and 12/7057).
The large specimen (material 52; 12/7049) in the lower right is a truncated bicone.

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vol. 63

Fig. 10.— Regular rounded beads provisionally attributed to adventurine (material 10; 12/7023 and
12/7024).

Discussion

The only locational information on the MAI catalog cards for the S. W. Howes
collection is “Montserrat, British West Indies.” However, Mrs. van der Veer and
Bruce Howes are confident that their father collected artifacts only on the Trants
Estate. In the tilled fields, S. W. Howes also found historic beads (some of which
are still in Mrs. van der Veer’s possession) but these were of no interest to the
MAI. Nevertheless, seven historic beads (Table 2), all made of glass, were included
in the materials sent to the MAI because superficially they resembled certain
prehistoric lithic beads collected by Howes. It is also possible that a few of the
lithic beads, all of which are assumed to be prehistoric in origin, may date to the
historic period of use of the Trants Estate.

Because the artifacts were surface collected, they lack contextual, stratigraphic,
or other provenience data. Fortunately, beads and related lithics were found in
undisturbed prehistoric contexts during excavations in 1979 and 1990. Materials
represented in the excavated objects include camelian, diorite, amethyst, quartz,
feldspar, and jadeite or nephrite, as well as several unidentified materials (Bartone
and Crock, 1991).

Comparatively high frequencies of exotic (nonlocal) lithic artifacts have been

Table 9 .—Regular rounded beads: frequencies.

Material category nos.

Probable material

Frequency

Percentage

25, 28, 34

diorite

252

71.2

64, 52

rock crystal

10.5

amethyst

5.6

adventurine

3.1

unidentified

2.8

turquoise

2.0

unidentified

1.4

camelian

1.1

unidentified

0.8

unidentified

0.8

unidentified

0.3

unidentified

0.3

Totals

354

100.0

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229

TRANSVERSE SECTIONS LONGITUDINAL SECTIONS

Elliptical

Square

Convex Straight

(barrel) (cylinder)

Fig. 1 1.— Drawings illustrating some of the classifications and terms used in this paper (adapted from
Beck, 1981:plates I, II, and III, and fig. 2, 3). A-B, transverse sections of regular rounded beads; C-
E, transverse sections of regular faceted beads; F, longitudinal sections of regular rounded beads; G,
Beck’s terms for classifying regular rounded beads by length; H, all eight beads illustrated in G have
circular transverse sections.

reported on at least four islands in the region— Vieques Island, near Puerto Rico
(Chanlatte Baik and Narganes Storde, 1983; Chanlatte Baik, 1984), St. Croix,
U.S. Virgin Islands (Vescelius and Robinson, 1979; Faber Morse, 1989:fig. 4e-
I), Montserrat (Harrington, 1924), and Grenada (Cody, 1990, 1991 a, 1991 b).
There are reports of exotic lithics on other islands, among them St. Vincent (Bullen
and Bullen, 1972:plate XlXf), Barbados (Drewett, 1991:132), Martinique (Mat-
tioni, 1979:49), Guadeloupe (Clerc, 1970:fig. 27; Durand and Petitjean Roget,
1991), Nevis (Wilson, 1989:436), St. Kitts (Fewkes, 1922:165, plate 85f), St.
Martin (Haviser, 1993), St. Eustatius (Versteeg and Schinkel, 1992:fig 160), and

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vol. 63

Fig. 12.— Diorite (material 25) beads (12/7 119). A, convex or “barrel” beads; B, straight or “cylinder”
beads.

Puerto Rico (Rouse and Alegria, 1990:50, fig. 12, plate 1 1; Rodriguez, 1991:610-
612; Rodriguez and Rivera, 1 99 1 :fig. 6, 7). Sued Badillo (1978) discusses the
prehistoric lapidary industry in the Caribbean.

Many of the lithic materials in the S. W. Howes collection from Montserrat are
duplicated in the collections from Vieques, St. Croix, and Grenada, but there are
some intriguing differences. Although there is no known source for camelian on
Montserrat, it is represented in all stages of manufacture in the Howes collection,
and is also disproportionately represented in the excavated artifacts from Trants,
where it comprises 65% of bead debitage and 88% of the blanks (Bartone and
Crock, 1991). Furthermore, two of the three Howes collection bead blanks ex-

Table 10.— Regular rounded beads: frequencies.

Longitudinal section
category nos.

Shape

Frequency

Percentage

convex (barrel)

188

53.1

straight (cylinder)

158

44.6

straight (truncated bicone)

1.1

convex (oblate, circular, ellipsoid)

0.3

convex (bicone)

0.3

convex (wedge)3

0.3

straight (truncated cone)

0.3

Totals

354

100.0

3 The convex (wedge) category was added by the authors; it is not used by Beck (1981).

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Watters and Scaglion T rants Beads and Pendants

231

Table 1 1.— Regular faceted beads: frequencies.

Transverse section
category nos.

Shape Frequency

Percentage

hexagonal

76.5

octagonal

11.8

square

11.8

Totals

100.0

Table 12 .—Regular faceted beads: frequencies.

Longitudinal section

category nos.

Shape

Frequency

Percentage

straight (cylinder)

55.6

straight (truncated cone)

11.1

convex (barrel)

11.1

convex (truncated bicone)

5.6

straight (bicone)

5.6

straight (truncated bicone)

5.6

straight (chamfered cylinder)

5.6

Totals

100.0

Table 13 .—Regular faceted beads: frequencies.

Material category nos.

Probable material

Frequency

Percentage

diorite

33.3

camelian

22.2

amethyst

11.1

adventurine

5.6

turquoise

5.6

rock crystal

5.6

89, 37

unidentified

16.7

Totals

100.0

T able 14.— Pendants: frequencies.

Material category nos.

Probable material

Frequency

Percentage

70, 76

jade group

33.3

turquoise

25.0

unidentified

16.7

adventurine

8.3

nephrite

8.3

serpentine

8.3

Totals

100.0

changed with the University Museum (Table 1) are camelian. At Trants, camelian
is found in all stages of lithic reduction, whereas amethyst occurs only as finished
beads (Fig. 14). Cody (1991a) documents just the opposite on Grenada, where
amethyst beads are found in all stages of manufacture, but camelian occurs only
as finished beads.

There very likely are geological sources within the West Indies for some of the
materials, such as serpentine and diorite, recognized both in the Howes collection

Fig. 13. —Pendants in the S. W. Howes collection. Front and back views: A, 12/7084(001), material
79, nephrite (verified gemologically); B, 12/7083(001), material 76, “jade group”; C, 12/7088(001),
material 82, probably serpentine; D, 12/7069(001), material 70, “jade group”; E, 12/7074(001), ma-
terial 70, “jade group”; F, 12/7075(001), material 73, unidentified. Single views: G, 1 2/7078(00 1),
material 13, turquoise; H, 1 2/7066(00 1), material 70, “jade group”; I, 12/7067(001), material 10,
probably adventurine; J, 12/7064(001), material 13, turquoise; K, pendant blank, 12/7068(001),
material 98, unidentified. The best quality pendants, being on exhibit, were not available for study.

and in collections from other sites in the region. However, there are no known
West Indian sources for turquoise (Fig. 15) and the “jade group.” Cody (1990,
1991 b) reports only two sources of turquoise in South America: northern Chile
and eastern Brazil. South American sources for the “jade group” (more generally
termed “greenstone”) have been investigated in considerable detail because of
the importance of greenstone pendants among surviving indigenous groups
(Boomert, 1987). Greenstone artifacts in the Lesser Antilles are most commonly
associated with the Saladoid ceramic series, which is generally interpreted as the
earliest of the Ceramic Age population movements into the region.

The S. W. Howes collection thus contains an unusually rich assemblage of
prehistoric beads and pendants of diverse types manufactured from a wide range
of materials (Fig. 1 6) that adds to existing knowledge of the prehistoric lapidary
industry in the Lesser Antilles.

1994

Watters and Scaglion— Trants Beads and Pendants

233

Fig. 14.— Only finished amethyst beads (material 4; 12/7016) were recovered at Trants.

Conclusions

The evidence provided by the beads in the Howes collection, coupled with
information regarding excavated beads, strongly suggests that Trants was a pre-
historic lithic bead manufacturing site specializing in camelian beads. The rela-
tively high proportion of broken blanks is compatible with this interpretation,
and the homogeneous nature of bead blanks provides additional evidence of a
specialized manufacturing industry.

Camelian seems to have been the most common material used for the local
manufacture of lithic beads, despite an absence of known local sources. Diorite
and rock crystal are also represented in early manufacturing stages. Whereas 90.0%
of raw materials and 69.4% of bead blanks are camelian, only 1.1% of finished
beads were of this material, suggesting probable import of raw materials and off-
island trade of the finished products. It may be that certain islands in the Lesser
Antilles, such as Montserrat (camelian) and Grenada (amethyst) were specializing
in the production of lithic beads primarily for off-island trade. If so, the identi-
fication of other island manufacturing sites would provide important clues for
understanding interisland interaction in Caribbean prehistory.

Fig. 15.— Five turquoise beads (material 13; 12/7026 and 12/7027). The disc head on the left, 12/
7026(004), was the specimen tested and identified as turquoise by the GIA Gem Trade Laboratory.

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vol. 63

Fig. 16. —Pendants and beads in the S. W. Howes collection. A, 12/7084(001), material 79 (nephrite;
specimen verified gemologically); B, 12/7064(001), material 13 (turquoise); C, 12/7065(001), material
13 (turquoise; verified gemologically); D, 12/7078(001), material 13 (turquoise); E, 1 2/7067(00 1),
material 10 (probably adventurine); F, 12/7083(001), material 79 (“jade group”); G, 12/7066(001),
material 70 (“jade group”); H, 12/7068(001), material 98 (unidentified); I, 12/7074(001), material 70
(“jade group”); J, 1 2/7069(00 1), material 70 (“jade group”); K, 12/7076(001), material 73 (uniden-
tified); L, 12/7075(001), material 73 (unidentified); M, 12/7088(001), material 82 (probably serpen-
tine); N, 12/7017(001), material 4 (amethyst quartz; verified gemologically); O, 12/7048(004), material
49 (unidentified); P, 12/7119, material 25 (diorite); Q, 12/7023(003), material 10 (probably adven-
turine); R, 12/7056, material 64 (probably rock crystal quartz); S, 12/7012(006), material 1 (camelian;
possibly a historic bead); T, 1 2/70 1 2(002), material 1 (camelian); U, 12/71 19, material 25 (diorite);
V, 12/7016(012), material 4 (amethyst quartz); W, 12/7032(002), material 22 (unidentified, possibly
quartzite; double-perforated object); X, 12/7046(001), material 43 (unidentified); Y, 12/7044(001),
material 37 (unidentified); Z, 12/7047(002), material 46 (possibly limestone); AA, 1 2/7027(00 1),
material 13 (turquoise); BB, 12/7021(001), material 4 (amethyst quartz); CC, 1 2/7029(00 1), material
19 (possibly turquoise); DD, 12/7049(001), material 52 (rock crystal quartz). Three beads (P, R, U),
all of which are associated with MAI catalog numbers containing numerous specimens, lack recorded
piece numbers (in parentheses).

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Watters and Scaglion— Trants Beads and Pendants

235

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vol. 63

Acknowledgments

Watters’ research at the Museum of the American Indian (MAI), Heye Foundation, in New York
was supported by the Netting Research Fund of Carnegie Museum of Natural History (CMNH). The
authors acknowledge the assistance of James B. Petersen, who studied the Howes collection ceramics;
Stanley W. Lantz for his extensive photographic documentation of the collection; Carrie Pantier for
darkroom assistance; staff members Anna C. Roosevelt, Mary Jane Lenz, Eulie Wierdsma, and Nancy
Rosoff of the Museum of the American Indian for help accessing the collection; Queenie Howes van
der Veer and Bruce Howes for discussing their childhood recollections of Trants Estate and for
permission to reproduce personal photographs of S. W. Howes and his home; Marc L. Wilson,
Collections Manager of the CMNH Section of Minerals, for discussing mineral nomenclature and
classification; Sylvia Keller and Colleen Mylan for assisting with manuscript preparation; and the
three anonymous reviewers for their careful reading and insightful comments. Research in 1985 took
place at the Research Branch in the Bronx when it was a facility of the Museum of the American
Indian, Heye Foundation, and in 1991 when it was part of the National Museum of the American
Indian, Smithsonian Institution. We have opted to use the MAI catalog numbers in this report since
that organization originally acquired and cataloged the S. W. Howes collection.

Literature Cited

Ball, S. W. 1941. The mining of gems and ornamental stones by American Indians. Smithsonian
Institution, Bureau of American Ethnology Bulletin, 128:1-77.

Bartone, R. N., and J. G. Crock. 1991. Flaked stone industries at the early Saladoid Trants site,
Montserrat, West Indies. Pp. 124-146, in Proceedings of the Fourteenth International Congress
for Caribbean Archaeology (A. Cummins and P. King, eds.), Barbados Museum and Historical
Society, 694 pp.

Beck, H. C. 1981. Classification and Nomenclature of Beads and Pendants. George Shumway
Publisher, York, Pennsylvania, 79 pp. [Reprint of 1928 original in Archaeologia 77, Society of
Antiquaries of London.]

Boomert, A. 1 987. Gifts of the Amazon: “Green stone” pendants and beads as items of ceremonial
exchange in Amazonia and the Caribbean. Antropologica, 67:33-54.

Bullen, R. P., and A. K. Bullen. 1972. Archaeological Investigations on St. Vincent and the
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Arqueologicas, Universidad de Puerto Rico, Rio Piedras, 90 pp.

Chanlatte Baik, L. A., and Y. Narganes Storde. 1983. Vieques “-Puerto Rico: Asiento de Una
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Clerc, E. 1970. Recherches archeologiques en Guadeloupe. Parall&les, 36/37:68-97.

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versity Microfilms #1344090.]

— — . 1991a. From the site of Pearls, Grenada: Exotic lithics and radiocarbon dates. Pp. 589-604,
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-. 1991 b. Distribution of exotic stone artifacts through the Lesser Antilles: Their implications

for prehistoric interaction and exchange. Pp. 204-226, in Proceedings of the Fourteenth Inter-
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and Historical Society, 694 pp.

Drewett, P. L. 1991. Prehistoric Barbados. London, Archetype Publications, 196 pp.

Durand, J.-F., and H. Petitjean Roget. 1991. A propos d’un collier funeraire a Morel, Guadeloupe:
Les Huecoide sont-ils un mythe? Pp. 53-72, in Proceedings of the Twelfth Congress of the
International Association for Caribbean Archaeology (L. S. Robinson, ed.), Martinique, Inter-
national Association for Caribbean Archaeology, 40 1 pp.

Faber Morse, B. 1989. Saladoid remains and adaptive strategies in St. Croix, Virgin Islands. Pp.
29-42, in Early Ceramic Population Lifeways and Adaptive Strategies in the Caribbean (P. E.
Siegel, ed.), BAR International Series 506, 418 pp.

Fewkes, J. W. 1922. A prehistoric island culture area of America. Pp. 35-281, in Thirty-fourth
Annual Report, Bureau of American Ethnography, Smithsonian Institution.

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Gemological Institute of America, Inc. 1985. Gem Identification Report #1926912, April 19,
1985. GIA Gem Trade Laboratory, New York, 2 pp.

Harrington, M. R. 1924. A West Indian gem center. Indian Notes, 1:184-189.

Haviser, J. B. 1993. A brief review of stone material. Association Archeologique Hope Estate
Bulletin, 2:30-31.

Mattioni, M. 1979. Salvage excavations at the Vive site, Martinique: Final report (L. Allaire, ed.
and trans.), University of Manitoba Anthropology Papers, 23, 56 pp.

Petersen, J. B., and D. R. Watters. 1991. Archaeological testing at the early Saladoid Trants site,
Montserrat, West Indies. Pp. 286-305, in Proceedings of the Fourteenth International Congress
for Caribbean Archaeology (A. Cummins and P. King, eds.), Barbados Museum and Historical
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Rodriguez, M. 1991. Arqueologia de Punta Candelero, Puerto Rico. Pp. 605-627, in Proceedings
of the Thirteenth International Congress for Caribbean Archaeology (E. N. Ayubi and J. B. Haviser,
eds.), Reports of the Archaeological-Anthropological Institute of the Netherlands Antilles, 9(2),
Willemstad, Curasao, 982 pp.

Rodriguez, M., and V. Rivera. 1991. Puerto Rico and the Caribbean Pre-Saladoid “Crosshatch
Connection.” Pp. 45-51, in Proceedings of the Twelfth Congress of the International Association
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Rouse, I., and R. E. AlegrIa. 1990. Excavations at Maria de la Cruz Cave and Hacienda Grande
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Watters, D. R. 1 980. Transect surveying and prehistoric site locations on Barbuda and Montserrat,
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Archaeology, 16:427-450.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 3, Pp. 239-256

16 September 1994

A PRIMITIVE CRICETID (MAMMALIA: RODENTIA) FROM THE
MIDDLE EOCENE OF JIANGSU PROVINCE, CHINA

Ban-yue Wang1 * *
Mary R. Dawson

Curator, Section of Vertebrate Paleontology

Abstract

A new species of the cricetid rodent Pappocricetodon, P. antiquus, from Middle Eocene fissure
deposits in southeastern China combines such derived features as uniserial incisor enamel and a
hystricomorphous zygomasseteric structure with primitive characters including retention of P4 and
less expansion of the anterocone/id of MVM, than in other species of the genus. This addition to the
Eocene record of myomorph rodents lends support to the hypothesis that the most likely sister group
of the Myomorpha is the family Sciuravidae.

Introduction

The Cricetidae, one of the most flourishing families of living rodents in the
world, are known to have diversified and dispersed in the Holarctic since latest
Eocene times. Due to their abundance in the fossil record, cricetids have been
used in the subdivision and correlation of Cenozoic Holarctic continental deposits
(Fahlbusch, 1964; Mein, 1975; Engesser, 1985). This abundance has also led to
numerous investigations of evolution within the family (Mein and Freudenthal,
1971a; Lindsay, 1968, 1977; Martin, 1980). Although the origin and interrela-
tionships of the cricetids have been discussed frequently, the source of the family
has been unclear. Suggested relationships have been with the Sciuravidae (Schaub,
1925; Wilson, 19496; Wood, 1959; Martin, 1980; Walton, 1993) or with the
ctenodactyloids (Lindsay, 1977; Flynn et al., 1985; Vianey-Liaud, 1985; Wang
and Meng, 1986).

Part of the difficulty of studying the origin of the family has been the lack of
good middle and early late Eocene records for the family. For many years this
record consisted of two lower molars referred to Cricetodon schaubi from the Late
Eocene River Section of Shanxi Province, China (Zdansky, 1930). Recently, the
new genus Pappocricetodon was described from China, with two species: P. ren-
cunensis from the late middle Eocene (Rencun Member, Hedi Formation) of
Henan; and P. schaubi , for Zdansky’ s species, from the late Eocene (Zhaili Mem-
ber, Hedi Formation) of Shanxi (Tong, 1992).

From 1987 to 1992 a field party from the Institute of Vertebrate Paleontology
and Paleoanthropology (IVPP), Academia Sinica (Qi et al., 1991), and, from 1 993,
a cooperative field team from the IVPP and the Carnegie Museum of Natural
History (CM) collected a variety of fossil mammals from fissure fillings in the
Triassic Shangqinglong Limestone in the Shanghuang Quarry of Liyang County,
Jiangsu Province, China (Fig. 1). To date five fissures (IVPP Locs. 93006.A-E)

1 Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, P.O. Box 643, Beijing

100044, China.

Submitted 18 February 1994.

239

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Annals of Carnegie Museum

vol. 63

Fig. 1 . — Map of China with enlargement showing parts of Jiangsu and adjoining provinces and location
(*) of Shanghuang Quarry, Liyang County.

have been found to contain Paleogene deposits. In Fissure D (IVPP Loc. 93006. D)
the deposits, composed mostly of red clay with lesser amounts of yellow clay, are
rich in fossil mammals. Cricetid fossils dominate in Fissure D. In addition to the
cricetids, the fossil mammals include varied insectivores, omomyids, leporids,
Yuomys sp., ctenodactyloids, creodonts, Archaeomeryx sp., Anthracotheriidae,
Homacodontidae, Eoentelodon, cf. Helohyus, brontotheres, palaeotheres, and ta-
piroids. All the specimens were collected by quarrying followed by screen washing.
The total mammal fauna from Fissure D suggests a middle Eocene age. The
Cricetidae is represented by the most primitive known and probably oldest mem-
ber of the family, described here as a new species of Pappocricetodon.

The discovery of this primitive cricetid from the middle Eocene both reveals
some primitive features and provides some suggestions on the origin of the Cri-
cetidae and on evolutionary trends within the family.

Methods

Figure 2 illustrates basic terminology used here for cricetid molars, combining conventional rodent
dental terminology (e.g., Wood and Wilson, 1936; Mein and Freudenthal, 19716) with nomenclature
necessitated by the primitive features of these rodents and the opportunity to employ terms suggesting
dental homology. For example, on M1 the crest that extends anterobuccally from the protocone is
termed protoloph, instead of “anterior arm of protocone” (Fahlbusch, 1964; Vianey-Liaud, 1979)
because it appears homologous to the protoloph of more primitive rodents such as sciuravids. The
cusp on the lingual part of the anterior cingulum, the anterocone of Lillegraven and Wilson (1975),
is named the anterostyle, and their parastyle is named the anterocone. “Mure” is used for the anterior
arm of the hypocone whether or not it joins the protocone. The “sinus” of the upper molar (Mein
and Freudenthal, 19716; Vianey-Liaud, 1979) is called “lingual valley.” On the lower molars the
anterior arm of the protoconid is termed the protolophid and the buccal crest of the metaconid, the
metalophid.

Biostratigraphic terminology for Oligocene localities follows Wang (1992) as the most recent treat-
ment of this interval in Asia.

Measurements were all taken by Wang using a micrometer in a Wild binocular microscope. For
consistency only the left side was measured for well-represented teeth; the scarcer M3/3 were all
measured.

In the tooth descriptions, frequency of a character variant is expressed as a ratio, with the upper
number indicating presence of the variant and the lower number, total number of specimens showing
the character. Thus, 101/143 indicates that out of 143 specimens, 101 have the variant described.

Abbreviations for repositories are: AMNH, American Museum of Natural History; CM, Carnegie
Museum of Natural History; IVPP, Institute of Vertebrate Paleontology and Paleoanthropology. Tables
of measurements (in millimeters, mm) of the dentition utilize the positional abbreviations: L, length;

1994

Wang and Dawson —Middle Eocene Cricetids from China

241

paracone
protoloph I
anterior lobe
anterocone
protoconule
anterostyle
protoloph
protocone

mesostyle

metacone

metaloph

posteroloph

mesoloph

mesocone

mure

hypocone

entostyle

mesostyle

metacone

metaloph

posteroloph

mesoloph

mesocone

mure

hypocone

ectolophid
mesoconid
protocol! id
anteroconid
metalophid I
in eta con id
metalophid II

hypoconid

hypoconuhd

posteroloph id

hypolophid

entoconid

mesolophid

mesostylid

Fig. 2.— Terminology used for cricetid molars, showing simplified left M1 and M2, and right Mr.

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W, width; b, buccal; 1, lingual. For statistical treatment, abbreviations used are: n, number of specimens;
O.R., observed range; X, mean; s, standard deviation; C.V., coefficient of variation.

Systematics

Order Rodentia Bowdich, 1821
Superfamily Muroidea Miller and Gidley, 1918
Family Cricetidae Rochebrune, 1883
Pappocricetodon Tong, 1992
Pappocricetodon antiquus, new species
(Table 1)

Holotype. — LM1 (IVPP V 11018.1).

Referred Specimens.- 142 M1 (IVPP V 11018.2-V 11018.143), 144 M2 (IVPP V 11018.144-V
11018.287), 21 M3 (IVPP V 11018.288-V 11018.308), one left lower jaw with I, and M, (IVPP V
1 1018.309), one left lower jaw with M, (IVPP V 1 1018.310), one right lower jaw with M1=2 (IVPP V

11018.311) , 111 Ms (IVPP V 1 1018.312-V 11018.422), 151 M2 (IVPP V 11018.423-V 11018.573),
30 M3 (IVPP V 11018.574-V 11018.603), 4 maxillae (IVPP V 1 1018.604-V 1 1018.607), 3 lower jaws
(IVPPV 11018.608-V 11018.610) and 2 I, (IVPP V 1 1018.61 1-V 11018.612).

Diagnosis. — Cricetid smaller than P. rencunensis, with hystricomorphous skull
and sciurognathous lower jaw; dental formula: 1/1, 0/0, 1/0, 3/3; cheek teeth
brachydont; upper molars have protocone bulbous and mesoloph short; M1 less
enlarged, having anterior lobe weak and anterocone small, protoloph I weak and
usually connecting to protoloph, protoloph II absent; on M1-2 mure usually long,
lingual valley long and oblique; M3 less reduced than in other species of genus,
with distinct metacone and somewhat reduced but less buccad hypocone; lower
molars usually lacking ectomesolophid; less enlarged with anteroconid small
and usually isolated; on M,_2 hypolophid transverse, connecting to hypoconid or
its posterior arm; lower incisor with uniserial enamel.

Etymology. — From Latin, antiquus, ancient, old.

Description. — Four edentulous maxillae of Pappocricetodon antiquus show that the infraorbital
foramen is relatively large and rounded with no ventral constriction. This is the hystricomorphous
condition. There is no bony septum isolating a neurovascular canal such as occurs in dipodoids. The
ventral root of the zygomatic process extends laterally well anterior to P4 and is compressed dorso-
ventrally. The zygomatic plate is flattened and nearly horizontal. It is bordered anteriorly and laterally
by a crescentic ridge extending from anterior to P4 to the lateral edge of the zygoma. This configuration
resembles that of Cricetops, an Asian cricetid, and a dipodoid such as Zapus more closely than that
of a North American muroid such as Eumys. This suggests that in Pappocricetodon the superficial
and lateral branches of the masseter muscle were incompletely differentiated as in dipodoids (Klingener,
1964). The posterior part of the incisive foramen, preserved in three of the four specimens, terminates
anterior to P4 and posterior to the anterior side of the ventral zygomatic root. The premaxillary-
maxillary suture extends obliquely back and down to intersect the lateral side of the incisive foramen.
The foramen appears to be relatively wider posteriorly than in Eumys. The anterior edge of the
maxillary-palatine suture is approximately in line with the juncture of M1 and M2.

One of the interesting features of the four maxillary fragments is evidence on the dental formula of
Pappocricetodon, unknown in the previously described species of the genus (Fig. 3). M1 and M2 each
have three alveoli, one for a large lingual root and two for smaller buccal roots. These alveoli show
that the anterobuccal root of M1 protrudes anteriorly more than does the corresponding root of M2.
On all maxillae there is a small circular alveolus just anterior to the anterobuccal alveolus of M1,
indicating that a small tooth, P4 or dP4, occurred in front of M1.

The anterior alveolar foramen (terminology follows Wahlert, 1974) pierces the maxilla in line with
the posterior margin of the ventral zygomatic root. Behind it are nutrient foramina. The larger spheno-
palatine foramen is at the anterior edge of the maxillary-palatine suture and above the intersection of
M1 and M2.

Six incomplete lower jaws are known, of which three have teeth (V 11018.309; V 11018.310; V

1 1018.31 1) . The horizontal ramus of the mandible is sturdy and the diastema is short (Fig. 4). The

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243

Fig. 3.— Ventral view of edentulous maxillae of Pappocricetodon antiquus, stereo views. Above, left
maxilla, IVPP V 11018.606, showing alveoli of P4 (or dP4) and M1. Below, right maxilla, IVPP V
1 1018.604, showing alveoli of P4 (or dP4), M1, and M2. Scale bar equals 1 mm.

mental foramen is anterior to Mi and slightly above middepth of the jaw. The masseteric fossa extends
forward to below M2. Distinct upper and lower ridges form boundaries of the fossa, intersect anteriorly,
and extend as a distinct ridge to below M,. The jaw is sciurognathous.

A complete cheek tooth series is not yet known. In addition to indications from the alveoli for
presence of P4 (or dP4), further evidence for the presence of a tooth anterior to M1 comes from V
1 1018.90, M1 which has a small wear facet on its anterior side. Thus, the dental formula of Pappo-
cricetodon antiquus was j^f, occurring only in this genus among the Cricetidae.

The lower incisor, compressed transversely and long oval-shaped in cross section, extends into the
ascending ramus. Enamel extends up to about one third of the lateral side but barely curves onto the
medial side. The enamel surface has fine longitudinal ridges as in P. rencunensis and P. schaubi.
Scanning electron microscope examination of the enamel of the lower incisor by Prof. Wighart von
Koenigswald (letter, 17 December 1993) produced the following results (Fig. 5): “The enamel is fairly
thin and has some external ridges. As normal in rodents the Portio interna (PI) and the Portio externa
(PE) are well distinguishable. The PE holds only 1% of the enamel thickness. It is formed by radial
enamel and the outer ridges are exclusively formed by a thickening of the PE. The interprismatic
matrix is at an angle toward the prisms and anastomoses between them. The PI is built of transversely
oriented uniserial Hunter-Schreger-Bands (HSB), which are vertical to the enamel dentine junction
(EDJ) in a longitudinal section. In the transverse section the prisms decussate at almost right angles.
The interprismatic matrix (IPM) is parallel to the prisms.”

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Fig. 4.— Left mandible of Pappocricetodon antiquus, with M,, IVPP V 11018.310, stereo views. Above,
occlusal view showing M,; below, lateral view. Scale bar equals 1 mm.

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Wang and Dawson— Middle Eocene Cricetxds from China

245

Fig. 5.— Detail of uniserial incisor enamel of Pappocricetodon antiquus, IVPP V 1 1018.61 1. Above,
transverse section; below, longitudinal section.

The cheek teeth are brachydont and have slightly elongate cusps and weak lophs (Fig. 6). M1 is
trapezoidal in occlusal view, the buccal wall slightly longer than the lingual. Its anterior lobe is small
with an anterocone that is usually distinct but small. An anterior cingulum extends from the anterocone
to in front of the protocone. The protocone is bulbous, and rarely has a weak crest on the posterobuccal
side (1 1/143). As in P. rencunensis and P. schaubi it extends slightly more buccad than the hypocone.
Usually the protoloph is long (128/141) and reaches the anterocone (121/141), but in few specimens
it is short and free (13/141). The protoconule on the protoloph ranges from moderate (87/143) to
indistinct (56/143). Paracone and metacone are elongate transversely and subequal. The lingual crest
from the paracone is weak (1 19/143) or absent (24/143); if present it intersects the protoloph near the
protoconule (90/143) to form a complete protoloph I, or the anterior end of the mure (29/143). There
is no trace of a protoloph II. The metaloph is complete and joins the mure near the hypocone. The
hypocone is crescentic. Its anterior arm tapers to form the mure, which is usually long and extends
far forward, variably meeting the protoloph (39/143), the lingual crest from the paracone (38/143),
or the buccal side of the protocone (21/143); it may also end freely (35/143). Occasionally (6/143) it
is short and joins the posterior side of the protocone. In V 1 1018.38, V 1 1018.40, ¥ 1 1018.72, and
V 1 1 0 1 8. 1 27 the mure forks into two branches: one reaches to the protocone and the other to protoloph
I to enclose a small basin. The mesoloph is less well-developed than in P. rencunensis and P. schaubi ,
is usually short (107/143), or even absent (1 1/143). It is usually transverse and from medial in position
to closer to the metaloph. In V 1 1 0 1 8.38 it is oblique and extends from the intersection of the metaloph
and the mure. The mesostyle is usually distinct. In some M1 (56/135) the mesostyle has a lingual
crest. The long mesoloph may be formed by a crest from the mesostyle and the short mesoloph. The
mesocone is rarely present (7/143). The anterostyle may be distinct (65/139) or indistinct (74/139)
and in some M1 (25/65) it joins the protoloph. The posteroloph reaches the metacone.

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Fig. 6. —Cheek teeth of Pappocricetodon antiquus. A. Left M1, holotype, IVPP V 1 1018.1; B. left M2,
IVPP V 11018.147; C. left M3, IVPP V 11018.296; D. right M„ IVPP V 11018.419; E. right M2,
IVPPV 11018.555; F. right M3, IVPP V 11018.590.

M2 is rectangular in occlusal view, slightly longer than wide. The anterior cingulum crosses the
anterior wall and joins the paracone buccally. It has a thickened anterostyle anterior to the protocone
that becomes a small bridge following wear. As in M1 the protocone is usually bulbous and rarely (23/
143) has a weak crest on its posterobuccal comer. Protoloph I is complete (137/141), and appears to
be formed by contributions from the protocone and the paracone. In about half of the specimens of
M2 (60/141), the intersection of the two crests can be seen. In some M2 the crest from the protocone
meets the paracone and leaves a free crest from the paracone anterior (4/141) or posterior (9/141) to
it. In V 11018.189, V 11018.240, V 11018.273, and V 11018.277, for example, protoloph I is
incomplete because the two crests do not intersect. The mesostyle is usually present (105/140) and
has a lingual crest extending toward the mesoloph. The mesoloph may be absent (16/140), short (33/
140), or of midlength (69/140). It may be forked (3/102) or double (26/102), with the anterior arm
shorter than the posterior. The long mesoloph of some specimens (22/140) appears to be formed by
the lingual mesostyle crest joining the short mesoloph. The mure is long, its anterior end usually
turning lingually to the buccal side of the protocone (78/143); it may extend to protoloph I (16/143)
or end in the valley (25/143). Occasionally it is short, reaching the posterior side of the protocone (3/
143). A weak entostyle or crest is sometimes present (5 1/143), very rarely forming a lingual cingulum
that joins the anterior cingulum (7/143). In some M2 (21/143) the posterobuccal crest of the protocone
may meet the mure to separate the lingual valley into open lingual and closed buccal parts.

M3, the smallest of the upper molars, varies considerably, especially in talon length and cusp
development. The anterior cingulum, paracone, protoloph I, and protocone are well-developed. In
some M3 (6/2 1 ), the anterior cingulum wraps around the lingual side of the protocone. The talon is
not as reduced as in P. rencunensis and P. schaubi. The mesostyle is usually distinct but weak (16/
21) and the mesoloph varies from weak (9/21) to absent (12/21). In the trigon basin there may be a
longitudinal crest (9/21). The metacone varies from cuspate to crescentic. The hypocone is less buccad
than in P. rencunensis and P. schaubi. The metaloph is shorter than the protoloph I and usually
complete (17/21), enclosing a small basin with the posterior cingulum. The lingual valley is shallower
than in P. rencunensis and P. schaubi.

Of the three lower molars, M2 is the largest. M, has a narrow trigonid as in P. rencunensis. Protoconid,

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247

metaconid, hypoconid, and entoconid are similar in size and height, with buccal and lingual cusps
aligned transversely. The metaconids are cuspate. Most protoconids (89/113) are cuspate, some are
crescentic (24/113). Metalophid II is usually complete (94/113), formed by the posterior arm of the
protoconid (81/113) or joined by a weak crest from the metaconid (13/113). Rarely it is incomplete
(17/113) or absent (2/ 1 1 3). The anteroconid is a distinct cuspule that varies from rounded to elongated
transversely (105/1 10). It is more weakly developed than in P. rencunensis and P. schaubi, usually is
low and isolated (89/105) and lacks a distinct cingulum (61/105), has a weak cingulum (one lingual
or buccal in 22/105), or has lingual and buccal cingula (18/105). It may be connected with the
protoconid (12/105) or the metaconid (4/105). The mesostylid is distinct; it may be isolated (75/1 10),
join the metaconid (31/1 10), or join both metaconid and entoconid (4/1 10). The mesolophid is less
well-developed and may be very short (49/113), of medium length (31/113), or even absent (29/113).
Rarely is it long (4/113). The mesoconid is more weakly developed than in P. rencunensis and P.
schaubi, and either distinct (73/113) or absent (40/113). The ectolophid is usually complete (95/113)
and straight (85/95). A few are curved (10/95), in which case the mesoconid is oblique, the anterior
arm of the ectolophid is not in line with the posterior arm and is usually more weakly developed. In
a few M, the ectolophid is incomplete and interrupted in front of the mesoconid (11/113), behind it
(4/113), or is completely absent (3/113). In the latter case, the mesoconid is isolated. The hypolophid
is usually complete (1 10/113). It differs from that in P. rencunensis and P. schaubi in being transverse
and joining the hypoconid (93/113), or ending freely (3/113). In some M, it extends posteriorly to the
posterolophid (6/113) or meets the anterior arm of the hypoconid (11/113). The posterolophid rarely
has a buccal part (1 1/113). Although the hypoconulid is usually absent (96/1 1 3), in some it is prominent
(17/113) and ends freely (3/113). In V 1 1018.396 the hypoconid also has a free posterior arm.

The M2 is nearly rectangular in occlusal outline. As in M, the four main cusps are subequal. The
lingual cusps are slightly elongate transversely and aligned with the buccal cusps. The protoconid is
usually crescentic, as in P. schaubi but unlike P. rencunensis. The anteroconid is a swelling lateral to
the midline on the anterior cingulum. After wear it forms a bridge joining the protolophid near its
intersection with the metalophid (137/149). The metalophid connects with the protolophid (120/150),
the protoconid (3/150), or the anteroconid (2/150) to form a complete metalophid I (125/150). Oc-
casionally it is incomplete and has a free buccal end (25/150) and leaves the trigonid basin open
anteriorly. Metalophid II, formed by the posterior arm of the protoconid, meets the metaconid to
close the trigonid basin (89/150), or is free, leaving the trigonid basin incompletely open (61/150).
Metalophid II is no greater than medium length and never reaches the lingual margin of M2. The
mesostylid is distinct (121/149) and variable, being isolated (98/149), connecting with the metaconid
(25/149), the entoconid (1/149), or both (6/149). The mesolophid is either shorter than metalophid
II (93/151) or absent (57/151). In one specimen, V 1 1018.461, it is double. The mesoconid is oblique
or transverse in orientation (70/150). Although the ectolophid is usually complete (1 19/150), it varies
from straight (69/150) to Z-shaped (50/150). In the latter case, the ectolophid is separated by the
oblique mesoconid into an anterior part that is slightly more external and lower than the posterior
part. In the few specimens in which the ectolophid is incomplete, the anterior part is very low or
absent (24/150) or the posterior part is very low (2/150). In five M2 (V 1 1018.449, V 1 1018.453, V
1 1018.491, V 1 1018.498, and especially V 1 1018.560) anterior and posterior parts of the ectolophid
are almost absent and the mesoconid is isolated and elongate transversely. The ectomesolophid is
absent (122/150) or weak (28/150). As in M, the hypolophid is usually complete (147/150) and
transverse, joining the hypoconid (95/150) or the posterior arm of the hypoconid (4/150). In some
M2 (48/150), the hypolophid meets the anterior arm of the hypoconid. The hypoconulid is usually
absent (136/150). In one specimen (V 1 1018.467), the hypoconulid ends freely.

M3 is a little smaller than M, and M2. The trigonid is similar to that of M2. The posterior arm of
the protoconid is usually free (21/30). It is variable in length, midlength (13/30), short (12/30), or
long (2/30). Rarely it is absent (3/30). The mesostylid is either present (14/30) or absent (16/30). The
mesolophid is absent (27/30) or very weak and short (3/30). The mesoconid is absent (10/30) or
variably present (20/30) and shaped as in M2. The ectolophid is variable as in M2. Unlike P. rencunensis
and P. schaubi, the ectomesolophid is absent (25/30) or very weak (5/30). The hypolophid is shorter
than in M2 and usually meets the anterior arm of the hypoconid (17/30). However, in some M3 the
hypolophid joins the hypoconid (5/30), the posterolophid (5/30), or is free (3/30).

Comparisons: Pappocricetodon antiquus resembles the two other described species of the genus, P.
rencunensis and P. schaubi, and differs from other cricetids in having relatively less enlarged M1 and
M,; M1 with a smaller anterior lobe and anterocone, protoloph I present but protoloph II absent, long
protoloph usually joining anterocone; M, with short and narrow trigonid and small anteroconid;
M§ less reduced.

Pappocricetodon antiquus is smaller than P. rencunensis and P. schaubi (Table 1). It also differs from
these species in having upper molars with a bulbous protocone and a less well-developed mesoloph;

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Table 1 . —Measurements (in mm) of Pappocricetodon antiquus.

O.R.

c.v.

1.2-1.55

1.38

0.08

5.8

1.05-1.48

1.24

0.08

6.45

0.95-1.25

1.10

0.07

6.36

1.15-1.55

1.31

0.07

5.42

1.0-1.35

1.13

0.06

5.13

0.92-1.15

1.09

0.08

7.61

0.87-1.1

1.03

0.06

6.21

1. 1-1.4

1.26

0.06

4.76

0.8-1.05

0.94

0.05

5.32

1.15-1.45

1.34

0.07

5.22

0.9-1. 2

1.05

0.07

6.67

1.05-1.4

1.24

0.09

7.26

0.87-1.15

1.02

0.06

5.88

M1 less enlarged (M1 length/M2 length in P. antiquus, 1.05; in P. rencunensis, 1.12; in P. schaubi,
1.18), with a smaller anterior lobe and anterocone, resulting in the relatively less elongated buccal
side; M1-2 with a long mure and a long, oblique lingual valley; M3 less reduced with a more distinct
metacone, an indistinct, less buccad hypocone, and a shallower lingual valley; lower molars lacking
ectomesolophid, having a rather transverse hypolophid that joins the hypoconid or posterolophid; M,
less enlarged and having a less developed anteroconid (M, length/M2 length in P. antiquus, 0.94; in
P. rencunensis, 0.95; in P. schaubi, 0.98); M3 less reduced. Except for the less distinct hypocone of
M3, the features mentioned above are primitive.

The incisor enamel of P. antiquus demonstrates an early stage in formation of the uniserial type,
which is characteristic of myomorphs (W. von Koenigswald, letter, 17 December 1993): “Compared
to modem cricetids two primitive characters are found in Pappocricetodon. The orientation of the
Hunter-Schreger-Bands (HSB) is vertical toward the enamel dentine junction (EDJ), and the inter-
prismatic matrix (IPM) is parallel to the prisms of the HSB. In modem cricetids the HSB are inclined
toward the occlusal surface and the IPM is (in lower incisors) oriented at right angles to the prisms
in the third direction. The schmelzmuster of Pappocricetodon is more derived than in any paramyid
or ischyromyid because of the uniserial HSB but primitive compared to modem cricetids.”

Relationships: Pappocricetodon was originally described as a new genus of cricetid based on isolated
teeth (Tong, 1992). No mention was made by Tong of the dental formula of this cricetid. As described
above, Pappocricetodon antiquus is known to have a tiny P4 (or dP4). Reexamination of P. rencunensis
and P. schaubi has shown that two M1 of P. rencunensis (IVPP V 8928.6 and V 8928.20) have an
anterior wear facet similar to that found in P. antiquus, indicating presence of a tooth anterior to M1.
Thus, not only P. antiquus but also P. rencunensis had P4 (or dP4) and the dental formula of Pappo-
cricetodon appears to have been The rudimentary P4 (or dP4) might have been shed during life.

One of the characters differentiating muroids from dipodoids is absence of P4 in the former. Because
Pappocricetodon has P4 (or dP4), it could be asked whether or not Pappocricetodon is correctly assigned
to the Muroidea. A basically similar molar pattern in primitive muroids (Cricetidae) and dipodoids
supports the concept of a monophyletic origin of the Myomorpha (Wilson, 1949a, 19496; Lindsay,
1968, 1977; Martin, 1980; Vianey-Liaud, 1 985). The similarity is such that at least one genus, Simimys,
has been transferred back and forth several times between these two myomorph superfamilies. Clearly,
the primitive condition for the myomorphs must include P4. P4 of Pappocricetodon is very reduced.
More important than its presence as a primitive feature are derived dental characters shared by
Pappocricetodon and other cricetids but absent in dipodoids. These include: 1) M1 has a distinct
anterior lobe and anterocone; and 2) the anteroconid of M, is transversely elongate and has a cingulum.
Pappocricetodon has a large, rounded infraorbital foramen, as in dipodoids and some other cricetids
such as Cricetops, but lacks a neurovascular canal, which is a derived feature of dipodoids. Thus,
derived dental features of Pappocricetodon, especially in the structure of the first molars, support its
reference to the Cricetidae and the most primitive known genus of the family, close to the separation
of muroids and dipodoids. The dipodoids are differently derived, retaining the primitive P4 and
rounded infraorbital foramen, but developing a protective bony septum to form a neurovascular canal.

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249

Origin of the Cricetidae

Now that Pappocricetodon has been recognized as the earliest and most prim-
itive known member of the family Cricetidae, hypotheses on the paleogeographic
and phylogenetic roots of this family, long discussed (numerous papers from
Schaub, 1925, through Tong, 1992), should be further tested. Characters of Pap-
pocricetodon important in this connection are: 1. hystricomorphous-sciurogna-
thous zygomasseteric structure; 2. anterior part of zygomatic plate nearly hori-
zontal; 3. P4 (or dP4) present; 4. upper molars with bulbous protocone; 5.
mesoloph(id) short; 6. metaloph contacts anterior arm of hypocone; 7. M1 and
Mj not enlarged; 8. M1 with small anterior lobe and anterocone; 9. protoloph I
weak on M1; 10. protoloph II absent; 1 1. protoloph short and free buccally on
M1; 12. mure free on M1-2; 13. lingual valley long and oblique on M1-2; 14. M3
and M3 slightly reduced; 15. M3 with distinct metacone; 16. hypocone distinct,
lingually located on M3; 17. posteroloph complete on M3; 18. with narrow
trigonid and small anteroconid; 19. ectolophid absent or incomplete; 20. meso-
conid oblique or transverse; 21. transverse hypolophid joins posterior arm of
hypoconid; 22. incisor enamel uniserial.

Paleogeographically, the origins of the cricetids and other myomorphs can surely
be sought in the Holarctic. In support of this premise is, of course, the occurrence
of Pappocricetodon in the middle to late Eocene of Asia. To this can be added
presence of a myomorph (Dawson et al., 1990) and a ?zapodid (Emry and Korth,
1989) in the early middle Eocene of North America, of Simimys, certainly a
myomorph and probably a dipodoid, in the later Eocene of North America (Wil-
son, 1949^; Lindsay, 1977; Emry, 1981), and of some diversity of cricetids in the
Chadronian and equivalents of North America and Eurasia. The family is not
known from Africa until the Miocene and from South America until the Pliocene.

The Asian rodent record prior to the appearance of Pappocricetodon includes
the families Alagomyidae in the earliest Eocene of Mongolia and Paramyidae in
the middle Eocene of Nei Mongol. (Two species of the Eocene genus Zelomys
were described as members of the family Sciuravidae by Wang and Li, 1990, but
this assignment is still equivocal). Alagomyids and paramyids are generally prim-
itive rodents. Alagomyids lack a hypocone on the upper molars. Paramyids have
upper molars with usually well-developed conules, weak lophs from paracone and
metacone converging toward the protocone to form a fairly well-marked trigon,
and usually only a trace of a hypocone. In both families the lower molars have a
pronounced basin. Neither has derived characters suggestive of close relationships
with myomorphs. Also present in Asia were several families of the superfamily
Ctenodactyloidea, first known from the early Eocene, which have been proposed
as myomorph relatives (Lindsay, 1977; Flynn et al., 1985; Vianey-Liaud, 1985;
Wang and Meng, 1986). Ctenodactyloids include the protrogomorphous early
Eocene Cocomys (Cocomyidae) and hystricomorphous middle Eocene Birbalomys
(Chapattimyidae), Yuomys (Yuomyidae), and Tamquammys (Tamquammyidae).
Among them Cocomys and Tamquammys are similar to the cricetids in some
primitive features such as having upper molars with a free buccal end of the
protoloph and some development of a hypocone. Differences between them and
cricetids are, however, more prominent: in the ctenodactyloids the cheek teeth
increase in size from first to third molars; the upper molars are wider than long;
the metaloph extends toward the protocone, forming a trigon similar to that of
paramyids; a metaconule is present; the lower molars have a large, distinct hy-

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poconulid; and the incisor enamel is pauciserial-multiserial or multiserial. To date
no evidence has been found in derived characters to connect Cocomys or other
ctenodactyloids to even the primitive cricetid Pappocricetodon antiquus.

European Eocene rodents are paramyids, chapattimyids (Pelaez-Campomanes
and Lopez Martinez, 1993), glirids, and theridomorphs. Characters separating
paramyids and chapattimyids from cricetids are mentioned above. Glirids are
clearly related to the paramyid microparamyines (Hartenberger, 1971). The most
primitive theridomorphs, members of the family Pseudosciuridae, occur first in
the middle Eocene. These rodents are hystricomorphous and develop uniserial
enamel. P3 is absent, the upper molars have a well-developed hypocone, the mure
is oblique and separated into anterior and posterior parts. A prominent, feature
is large P4 and P4, which are never reduced. Even primitive theridomorphs differ
from cricetids in having very well-developed conules, transversely-oriented main
cusps on upper and lower molars, and only a rudimentary lingual valley. These
differences from cricetids indicate no close affinity between these two groups.

North American early and middle Eocene rodents include paramyids, cylin-
drodontids, sciuravids, and some probable myomorphs. As mentioned above,
paramyids show no tendency toward developing lophate teeth with well-developed
hypocone and hypolophid that would be expected in relatives of the cricetids.
Cylindrodontids do have a well-developed hypolophid, but their upper molars
remain on a basic paramyid level of development throughout the Eocene.

The North American rodent family that stands out in dental morphology to
have important features shared with the cricetids remains the Sciuravidae (Wilson,
1949Z?). Sciuravids are basically primitive rodents, combining a dental formula
of protrogomorphous/sciurognathous zygomasseteric structure, pauciserial
incisor enamel, and brachydont cheek teeth. In molar morphology, however, some
sciuravids possess derived characters shared with early myomorphs including the
primitive cricetid Pappocricetodon and the primitive ?zapodid Elymys. In the
upper molars conules are reduced; protocone and hypocone are subequal and
separated lingually by a pronounced valley; protoloph and metaloph are oblique,
extending into the valley anterior to the corresponding para- and metacone. This
last character is especially strong in Sciuravus nitidus, S. altidens, and S. poway-
ensis. In S. bridgeri the valley between protocone and hypocone is crossed by a
low crest from each cusp (Dawson, 1968:350-351). In the lower molars of Sci-
uravus the cusps are set obliquely, with lingual cusps anterior to the corresponding
buccal cusps, the ectolophid is complete, the hypolophid is well-developed and
the entoconid is separate from the posterolophid. There is a tendency among
sciuravids toward reduction of the premolars, seen most markedly in Sciuravus ?
rarus (Wilson, 1938), AMNH 12118 (Dawson, 1962), and Pauromys. The Sci-
uravidae are here considered to be the sister group of the myomorphs (Fig. 7).

Two early middle Eocene North American genera are probably myomorphs.
Armintomys tullbergi, the oldest known rodent with a hystricomorphous zygo-
masseteric structure, has incisor enamel that is transitional from pauciserial to
uniserial. It seems to be a primitive myomorph but is probably not a member of
the Dipodoidea (Dawson et al, 1990) because it lacks the derived neurovascular
canal characteristic of that superfamily. It shares with P. antiquus some features,
such as M!-M2 longer than wide, mesostyle present, low lophs, hypocone subequal
to protocone, and bipartite metaloph. It is more primitive than P. antiquus in
some other features including the presence of P3, probably well-developed P4, and

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251

Sciuravidae

Armintomys

Elymys

Zapodidae

Nonomys

Simimys

Pappocrice todon
other cricetids

Fig. 7.— Analysis of relationships between the Sciuravidae and several myomorphs, including Pap-
pocricetodon, based on characters considered to be shared-derived (synapomorphous) at the branching
points: 1, upper molars with hypocone subequal to protocone; 2, upper molars with buccal cusps
anterior to corresponding lingual cusps; 3, conules weak or absent; 4, well-developed hypolophid; 5,
complete ectolophid; 6, hystricomorphy; 7, pauciserial toward uniserial enamel; 8, P3 absent, P4 small;
9, uniserial incisor enamel; 10, zygoma slightly inclined; 11, neurovascular canal separate from in-
fraorbital foramen; 12, P4 absent; 13, M1 with anterocone and anterior lobe; 14, Mj with anteroconid
and anterior cingulum; 15, P4 absent; 16, zygomatic plate developed.

lower crowned teeth. Its sciuravid-like molar pattern, hystricomorphy, and incisor
enamel indicate that Armintomys tullbergi may represent the closest known sister
group to a unit including dipodoids and cricetids.

Elymys (Emry and Korth, 1989) from the early Bridgerian of North America
resembles Pappocricetodon in dental formula and tooth morphology having P4
reduced, upper molars longer than wide, oblique compressed protocone and large
hypocone, protoconule present in M1 but lacking in M2-3, M1 with protoloph
extending anterobuccally and a protoloph I, M2 protoloph I complete, formed by
crests from paracone and protocone. Elymys is more primitive than P. antiquus
in having lower-crowned teeth and lacking the mesoloph(id), anterocone and
anterior lobe on M1, and anteroconid on Because the zygomasseteric structure
and incisor enamel of Elymys are unknown, its relationships remain uncertain.
The suggestion proposed by Emry and Korth (1989:9) is that Elymys may be
ancestral to both Simimys and later zapodids. Elymys and Pappocricetodon appear
to converge toward a common origin.

Simimys, from the Uintan and Duchesnean of North America, combines a
hystricomorphous skull with a separate neurovascular canal as in dipodoids but
the muroid dental formula of M3/M3. It has been regarded as a “more or less
primitive survivor into the late Eocene of a stalk which was ancestral to both
cricetids and the Dipodoidea” (Wilson, 1949(3:23). Recently it has been included
in the Dipodoidea (Emry and Korth, 1989). Nonomys, from the Chadronian, is
another rodent having the derived feature of a separate neurovascular canal. Like
Simimys, it has the muroid dental formula, but its teeth appear to have a highly
derived pattern, cuspate and with well-developed cingula, very different from the
more dipodoid-like pattern in Simimys. Although dental features have been used

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to support the affinities of Nonomys with the muroids (Emry, 1981), the phylo-
genetic importance of the neurovascular canal is here emphasized and this genus
is regarded as a dipodoid.

Paleogene Cricetids in Asia

In the Paleogene of Asia cricetids are represented by eighteen species distributed
among four genera: Pappocricetodon Tong, 1 992 (Middle to Late Eocene of Henan,
Shanxi, and Jiangsu provinces, China); Eucricetodon Thaler, 1966 (Early to Late
Oligocene of Nei Mongol, Gansu, and Yunnan of China, Mongolia and Kazakh-
stan); Cricetops Matthew and Granger, 1923 (Middle Oligocene of Nei Mongol
of China, Mongolia and Kazakhstan); and Selenomys Matthew and Granger, 1 923
(Middle Oligocene of Nei Mongol of China and Mongolia). (Note: Aksyiromys
and Ulkenulastomys, known from one tooth each, were cited as “Myomorpha,
family indet.” by Shevyreva, 1984. Whatever their affinities, they are apparently
not cricetids and accordingly are not considered here.)

Pappocricetodon, the most primitive cricetid, includes three species: P. rencu-
nensis Tong, 1992, P. schaubi (Zdansky, 1930), and P. antiquus, described above,
now the earliest and most primitive member of this genus. In most features P.
antiquus is more primitive than the other two species, but it has a more reduced
hypocone on M3 than P. rencunensis and P. schaubi, and thus appears to be a
sister group to the two latter.

Eucricetodon is represented by ten taxa in Asia: E. asiaticus (Matthew and
Granger, 1923; Lindsay, 1978) from the Middle Oligocene of Mongolia and Ka-
zakhstan; E. near E. asiaticus from the Middle Oligocene of Mongolia; E. affi E.
asiaticus from the Late Oligocene of Kazakhstan; E. caducus (Shevyreva, 1967)
from the Middle Oligocene of Nei Mongol of China and Kazakhstan; E. aff. E.
caducus from the Late Oligocene of Kazakhstan; E. meridionalis Wang and Meng,
1986, and E. leptaleos Wang and Meng, 1986, from the Early Oligocene of Yunnan,
China; E. youngi Li and Qiu, 1980, from the Early Miocene of Qinghai, China;
Eucricetodon sp. from the Late Oligocene of Taben Buluk, Gansu, China (Bohlin,
1946), and Eucricetodon sp. from the Early Oligocene of Mongolia (Janovskaja
etal., 1977).

Tong (1992:6-7) considered Eucricetodon meridionalis to be more similar to
Pappocricetodon than to Eucricetodon on the basis of the following shared features
of tooth morphology: M1 with a very small anterior lobe, a well-developed pro-
toloph joining anterocone with protocone, and a single anterocone; small Mt about
same length as M2 and with small anteroconid and trigonid; no free arm of
hypoconulid. Tong concluded that E. meridionalis was more closely related to
Pappocricetodon than to other cricetids and might be in the same evolutionary
lineage as P. rencunensis and P. schaubi. However, E. meridionalis is here con-
sidered to be a species of Eucricetodon that has more primitive features than other
species of Eucricetodon. All the similarities mentioned by Tong are primitive
features. In comparison with Pappocricetodon and Eucricetodon, E. meridionalis
shares with the latter such derived features as M1 having an enlarged anterior
lobe, a large anterocone, lacking protoloph I but having a complete protoloph II,
a large anteroconid, and Mj longer than M3. E. meridionalis is here retained in
Eucricetodon.

Tong (1992:7) also doubted the position of E. leptaleos in Eucricetodon because
he thought it had the following special characters: 1) short protoloph (^anterior
arm of protocone in Tong, 1992) extending towards paracone or joining crest

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253

from paracone (=protoloph in Tong, 1992); 2) anterocone(id) small and with a
smooth posterior surface; 3) absence of a bridge from anterostyle (=caudaler
anteroconusspom of Dienemann, 1987, and Tong, 1992); 4) M3 having simple
structure; 5) M, equal to M2 in length; 6) weak lophs on lower molars; and 7) M3
with a reduced talonid. Of these, 1, 2, 3, 5, and 6 are primitive features of cricetids
and similar to those of Pappocricetodon, and 4 is variable in cricetids and probably
not phylogenetically significant. Only 7 is a derived feature but it is shared by
Eucricetodon. In fact, in E. leptaleos the anterior lobe of M1 is enlarged and forms
a distinct lobe, the anterocone is large, and protoloph II has begun to form, as in
Eucricetodon . Thus, E. leptaleos appears to be a primitive species of Eucricetodon.

Pappocricetodon rencunensis and P. schaubi share some derived features: a long
protoloph joining the anterocone on M1, a well-developed mesoloph, and a buccad
hypocone on M3. These features differ from those of Eucricetodon, including E.
meridionalis and E. leptaleos , and other later cricetids. However, among the
cricetids Eucricetodon is closest to Pappocricetodon. Possibly primitive cricetids
in Asia may have divided into two lineages, with Pappocricetodon representing a
primitive group, and another lineage represented by Eucricetodon and other Asian
cricetids, in which M1 and M, are enlarged and have a large anterocone(id) and
Mj has a well-developed anterior cingulum. There remain many gaps in the history
of cricetids in Asia, however, with such problems as the origin of Cricetops and
the peculiar Selenomys still to be resolved.

Paleogene Cricetids in the Holarctic

Now that the geologically oldest and morphologically most primitive cricetid
rodents are known to occur in the Eocene of Asia, implications of the entirely
Holarctic Paleogene stratigraphic and biogeographic record of the family should
be reconsidered. Pappocricetodon is important in providing evidence on primitive
characters of cricetids that allows a clearer understanding of character polarity
among Paleogene forms. No attempt is made here to revise the family at the
subfamily or any other level. Currently the infrafamilial arrangement of cricetids
can conservatively be characterized as in a state of flux (contrast, for example,
the familial and subfamilial associations of Martin, 1980, Unay-Bayraktar, 1989,
and Freudenthal et al., 1992).

Cricetids appeared in Europe after the Grande Coupure. Since that event cri-
cetids (sensu lato) diversified and dispersed rapidly in Europe. Thirteen genera
are recognized (Freudenthal et al., 1992) in the European Paleogene record. The
earliest cricetid is Eucricetodon atavus from Hoogbutsel (the Eucricetodon atavus
tooth from the pre-Grande Coupure locality Mohren 6 should be dismissed due
to faunal mixing within the fissure, following the discussion in Dienemann, 1987).
It resembles the Asian E. meridionalis in size and some aspects of tooth mor-
phology, but is more derived in having first molars with a larger anterior lobe
and anterocone(id) and more reduced third molars. The record supports the hy-
pothesis that Eucricetodon migrated from eastern Asia into Europe after the Grande
Coupure.

In North America, as in Europe, a major cricetid radiation occurred in the
Oligocene ( Eumys sp. described by Storer, 1988, from the Duchesnean of Sas-
katchewan has been referred to the Sciuravidae by Dawson, in press), from which
eight cricetid genera have been reported (Martin, 1 980). They have some common
features, such as more or less swollen main cusps, short posteroloph and well-
developed metacone on M3, which differ from those of Asian Paleogene cricetids,

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including Pappocricetodon and Eucricetodon. The North American cricetids Eoeu-
mys, Scottimus, Leidymys, Geringia, and Paciculus are not referable to the Eur-
asian Eucricetodontinae (Tong, 1992, contra Martin, 1980). The Oligocene genera
Geringia and Paciculus have a small anterocone-anteroloph that differentiates
them from contemporary North American cricetids. This state has been regarded
as a reduction from the Eurnys condition. The morphology of the primitive Asian
Pappocricetodon opens the possibility that these genera represent another im-
migration from Asia rather than a reduction from North American endemic
rodents.

Acknowledgments

This report would not have been possible without the encouragement and support of and discussions
with our colleagues Chris Beard and Qi Tao, whom we thank most sincerely. We are also grateful to
Wighart von Koenigswald and Jin Meng for their work on the incisor enamel of this early cricetid.
Illustrations were skillfully done by Andrew D. Redline, photographs by Chris Beard and Wighart
von Koenigswald. We appreciate reviews of this manuscript by Robert Emry and Everett Lindsay,
whose comments helped in clarifying our results. The larger project on Paleogene fissure fillings from
China is supported by grants from the National Science Foundation (DBS 9221231), the Chinese
National Science Foundation (grant no. 49372070), and the M. Graham Netting and Edward O’Neil
funds of Carnegie Museum of Natural History.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 3, Pp. 257-261

16 September 1994

MEDIO CAMP US, A NEW STINKBUG
GENUS FROM THE DOMINICAN REPUBLIC
(INSECTA: HETEROPTERA: PENTATOMIDAE)

Donald B. Thomas1

Research Associate, Section of Invertebrate Zoology

Abstract

A new genus, new species, in the stinkbug family Pentatomidae, Mediocampus dominicanus is
described from the Dominican Republic. The genus is apparently related to the Antillean genera of
the Pentatomini with a xiphoid metastemum, Pharnus, Neopharnus, and Praepharnus. It differs in
that the metastemum is deeply sulcate for reception of the rostrum and the abdominal tubercle is
bicarinate. These genera, especially the new genus, occupies a phylogenetic position somewhat inter-
mediate between the Pentatomini and the Edessini.

Introduction

As a result of intensive collecting in the Dominican Republic by staff members
of the Carnegie Museum, a new genus of stinkbug, herein described, was discov-
ered. The new genus is represented by two female specimens from two localities
separated by a distance of approximately 65 km. Both localities were low elevation
(230 m), riparian woodland habitats. The specimens were collected at lights. The
new genus appears to be related to a small group of genera known to occur primarily
on the island of Cuba.

Mediocampus, new genus
(Fig. 1-3)

Type Species. —Mediocampus dominicanus, n. sp.

Diagnosis.— Metastemum bilaterally elevated, longitudinally, mesially sulcate
for reception of rostrum; broadly notched posteriorly in apposition to basal ab-
dominal production; projecting anteriorly as narrow elongate arm on each side
of mesostemum. Meso sternum flat, narrow, with low bicarinate apical process
projecting anteriorly between procoxae. Basal production of abdomen strongly
sulcately excavated mesially, lateral margins of sulcus obtusely carinate; each
carina projecting anteriorly into metastemal notch as subspinous production (Fig.
2). Bucculae prominently arcuate anteriorly, evanescently contiguous posteriorly.
Apex of scutellum acuminate.

All femora unarmed. Tarsi three-segmented. Antennae five-segmented. Trich-
obothria on abdominal stemites II-VII in line with spiracles. Base of rostrum
originating anterior to middle of head.

Etymology. —Latinized from Spanish Mediocampo, literally medio meaning
“middle,” and ocampo meaning “field,” in reference to the intermediate position
of the genus between the typical pentatomines and the plesiomorphic edessines.

1 USDA-ARS Subtropical Agriculture Research Laboratory, Weslaco, Texas 78596.
Submitted 1 1 March 1 994.

257

Fig. 1 .—Mediocampus dominicanus, dorsal aspect. Bar = 2 mm.

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Thomas “New Dominican Republic Heteropteran

259

Mediocampus dominicanus, new species

Description. — Elongate, ovate; length excluding membrane 1 1 mm, width across
humeri 7 mm. Stramineous dorsally with fuscous yellow hemelytral coria and
scutellum; dorsal and ventral surface with reddish-brown punctures irregularly
distributed. Connexival segments with dark brown anterior and posterior infra-
margins alternating with pale yellow meson.

Head. Length from apices of juga to line of ocelli slightly greater than anteocular
width. Juga broadly contiguous anteriorly, lateral margins strongly sinuate. An-
tennal segment I shortest; II slightly longer than III; III about two-thirds length
of IV; IV and V longest, subequal. Rostrum long, extending to third abdominal
segment (second visible); segment II longest, slightly longer than III; segments I
and IV subequal to one another and about two-thirds length of II.

Thorax. Anterolateral pronotal margin smooth, without marginal bead, sub-
rectilinear in dorsal view. Humeri angular, not produced. Basal margin of pro-
notum arcuately concave; posterior angles not acutely produced. Scutellum pro-
portionately long, apex extending to posterior limit of coria. Posterior margin of
corium evenly arcuate; membrane infuscated. Metastemal scent gland evapora-
torium maculate with spots equal in size and density to the dark punctures
scattered over rest of venter. Ruga of scent gland orifice elongate, extending two-
thirds distance to metapleural margin. Protibia subprismatic, meso- and meta-
tibiae cylindrical with elongate sulcus.

Abdomen. Apices of connexiva acuminate. Lateral angles of stemite VII strongly
spinosely produced. Margin separating segments III and IV effaced mesially. Spira-
cles large, oval.

Genitalia. First gonocoxites briefly contiguous at base, triangulum exposed
medially; second gonocoxites narrow and emarginate posteriorly; eighth parater-
gites strongly angularly produced posteriorly; spiracles present; ninth paratergites
narrowly acuminate (Fig. 3). Males unknown.

Type Specimens. — Holotype, female. Verbatim label data: “DOMINICAN RE-
PUBLIC: Barahona. 9.2 Km NW Paraiso, confluence of Rio Nizao and Rio Coltico
18-03N 71-12W 230m 9-10 Aug 1990 J. Rawlins, S. Thompson.” Deposited
Carnegie Museum Natural History.

Paratype, female, (a) “DOMINICAN REPUBLIC: Pedemales. Along Rio Mu-
lito, 1 3 Km N. Pedemales, 1 8-09N 7 1 -46 W.” (b) “230m, 1 7 July 1 992, J. Rawlins,
S. Thompson, C. Young, R. Davidson. Riparian Woodland.” Deposited Carnegie
Museum Natural History.

Discussion

The character states separating the neotropical subfamily Edessinae from the
cosmopolitan Pentatominae are somewhat ambiguous. Four edessine genera, all
restricted to the New World, are recognized: Edessa Fabricius, Peromatus Amyot
& Serville, Olhia St&l, and Pantochlora St&l. Leston (1955) recognized the tribes
Edessini and Pantochlorini within the subfamily Pentatominae. Rolston and
McDonald (1979) did not acknowledge the tribal separation, but elevated the four
genera to their own subfamily, Edessinae. All have a strongly-elevated metaster-
num which projects over the mesostemum; all have a short rostrum which extends
only to the mesosternum in repose; all have the bucculae of the oral groove united
behind the rostrum, and all have parandria or dorsal genital plates on the interior
surface of the male pygophore. However, none of these character states is unique
to the edessines, but occur singly or in combination in many pentatomines. The

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vol. 63

Fig. 2-3.— 2. Metastemal and abdominal armature, ventral view. Cx = coxa, Mst = metastemum,
Ab2 = second abdominal segment. 3. Female terminalia. Pt8 = eighth paratergite, Gel = first gono-
coxite.

genus Piezosternum Amyot & Serville has all of these characters but is placed in
the Tessaratomidae, subfamily Oncomerinae, by Kumar (1968) because it has the
additional character of the pronotum overlying the base of the scutellum. For this
reason the Edessinae may be considered to be the stock derived most closely to
the sister family Tessaratomidae.

Mediocampus is very “edessoid” in aspect and my first impression on seeing

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Thomas - New Dominican Republic Heteropteran

261

these specimens was that they were a species of the large genus Edessa. On
examining the ventral armature for the characteristic xiphoid metastemum found
in all members of the latter genus, I found the xiphoid metastemum with typical
long anterior arms present, but with the unusual character of a groove or sulcus
mesially for the reception of the long rostrum. A similar, but even more exag-
gerated condition occurs in some South Pacific asopine genera, namely Ealda
Walker and Cantheconidea Bergroth, but that is clearly a case of convergence and
not relation. Another distinctive character is the bifid abdominal tubercle. A
similarly bifid abdominal tubercle occurs in the African genera Glypsus Dallas
and Bathycoelia Amyot & Serville, which otherwise bear little resemblance to the
new Antillean genus.

Rolston et al. (1980) divide the Pentatomini into three sections based on the
ventral armature. Mediocampus belongs to section 3, those having an abdominal
tubercle in apposition to an elevated metastemum. In the key to this section
Mediocampus falls out with a small group of Antillean genera which, like Edessa,
have a xiphoid metastemum: Pharnus St&l, Neopharnus Van Duzee, and Prae-
pharnus Barber and Bruner. In their description of the latter genus, Barber and
Bruner (1932) considered Praepharnus “more closely related to Pharnus than to
Edessa. . . Mediocampus appears to lie between Praepharnus and Edessa. It is
similar to Praepharnus in size, elongate depressed form, straight anterolateral
pronotal margins, and long slender rostrum. It shares with Edessa the long anterior
arms of the metasternum and the posteriorly-united bucculae. It is unlike both
genera in having the bicarinate abdominal tubercle and the sulcate metastemum.
This sulcus is continuous with a sulcus on the abdominal midline, in contrast to
the keeled abdominal midline in Praepharnus.

It is interesting to speculate that these three endemic Cuban genera, plus the
new Hispaniolan genus, might be offshoots from a lineage close to the base of the
edessines and pentatomines that differentiated and persisted in isolation in the
Greater Antilles.

Acknowledgments

I am grateful to John Rawlins for making these specimens available to me for study, and to L. H.

Rolston and Pieter van Doesburg for advice on character states in the Edessinae. The habitus drawing

of Mediocampus dominicanus was executed by Chris Van Dyke.

Literature Cited

Barber, H. G., and S. C. Bruner. 1932. The Cydnidae and Pentatomidae of Cuba. Journal of the
Department of Agriculture of Puerto Rico, 16:231-284.

Kumar, R. 1968. Morphology and relationships of the Pentatomoidea (Heteroptera) IV. Oncome-
rinae (Tessaratomidae). Australian Journal of Zoology, 17:553-606.

Leston, D. 1955. A key to the genera of Oncomerinae St&l (Hemiptera, Pentatomidae). Proceedings
of the Royal Entomological Society of London (B), 24:62-68.

Rolston, L. H., and F. J. D. McDonald. 1979. Keys and diagnoses for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemip-
tera). Journal of the New York Entomological Society, 87:189-207.

Rolston, L. H., F. J. D. McDonald, and D. B. Thomas. 1980. A conspectus of Pentatomini genera
of the Western Hemisphere. Part I (Hemiptera: Pentatomidae). Journal of the New York Ento-
mological Society, 88:120-132.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 3, P. 263

16 September 1994

REVIEW

NEW JERSEY FERNS AND FERN ALLIES. James D. Montgomery and David
E. Fairbrothers. 1992. Rutgers University Press, New Brunswick, New Jersey.
293 pp. with line drawings. ISBN 0-8135-1817-2. $45.00.

New Jersey Ferns and Fern Allies begins with a historical introduction of bo-
tanical field studies in New Jersey, especially those on pteridophytes. This subject,
vast enough to warrant its own book, is substantially condensed in a one-chapter
treatment.

Other prefatory chapters include “Fern Structure and Classification,” “Hy-
brids,” “Keys,” and “Ecology and Distribution.” These chapters discuss the terms
used in the keys, how hybrids arise and some characteristics for hybrids identi-
fication, and the ecology of New Jersey, including how pteridophyte distribution
is related to ecological regions in the state. The authors make these subjects
understandable even for novice botanists.

This fern flora is more than just a description of all 83 species of ferns and fern
allies found in the state. Specimen occurrences for each species are plotted on
maps using three different-sized dots to indicate records from pre-1900, 1900-
1950, and post- 1950. This system reveals chronological trends for each species—
whether its populations are declining, expanding, or remaining static. Although
each dot represents a herbarium specimen, the authors fail to list which herbaria
were consulted. This information would be useful for future floristic projects in
New Jersey. Taxa apparently absent from New Jersey but found in bordering
regions are mentioned in the accounts of closely related species.

Each taxonomic chapter treats a single genus, beginning with a description of
the genus and a key to the species (and hybrids) found in New Jersey. Each species
account provides scientific name, synonyms, common name, and a brief descrip-
tion of the plant, including its habitat, range, and distribution in New Jersey.
Comments include chromosome number and endangered status in the state. Each
distribution map is paired with an excellent line drawing by Kathleen L. John-
Alder, often with a smaller closeup drawing of diagnostic characteristics. The
illustrations in the genus Ophioglossum are reversed. For ease of reference, text
and figures for most species accounts are on facing pages, which also results in a
good deal of blank space.

The keys in the manual, including a vegetative key to ferns of New Jersey, are
easy to use by anyone with a basic knowledge of pteridophyte terminology. It is
refreshing that the authors include hybrids in the keys, instead of mentioning
them only in passing as is too often the case. Thirty-two hybrids are discussed,
with descriptions, illustrations, and maps of the 1 7 that occur most frequently.

The included checklist of all pteridophyte species and hybrids found in New
Jersey will be useful to those involved in floristic studies of New Jersey and nearby
states. The bibliography is excellent for anyone interested in delving deeper into
the pteridophytes of eastern North America.

BONNIE L. ISAAC, Curatorial Assistant, Section of Botany.

263

INSTRUCTIONS FOR AUTHORS

Manuscript style.— Articles should include the follow-
ing items in this order: title page, abstract, text (with
desired headings), acknowledgments, literature cited, ta-
bles, figure captions, and copies of illustrations. All
manuscripts must be typed double-spaced on standard
8‘/2 by 1 1 inch white bond paper, with at least one inch
margins all around, and submitted in triplicate— an orig-
inal for the editors, and two review copies. All pages
should be numbered, including tables, literature cited,
and the list of figure captions. Only correspondence qual-
ity or better dot matrix printouts will be accepted; draft
mode is unacceptable. A fixed-space font (e.g., Courier)
is preferred.

Abstracts. Abstracts should be short, double-spaced
and substantive and included at the head of the first page
of text.

Text. Do not right justify text or break (hyphenate) a
word at the end of a line. Footnotes and acknowledgments
as footnotes are unacceptable. All text is double-spaced.

Measurements. Metric units should be used, if possi-
ble.

1) Two authors in an institutional series:

Matthew, W. D., and W. Granger. 1923. The
fauna of the Houldjin Gravels. American Mu-
seum of Natural History Novitates, no. 97:
1-6.

2) Same authors repeated— use three-em dash:

. 1923. The fauna of the Ardyn Obo For-
mation. American Museum of Natural

History Novitates, no. 98:1-5.

3) Same authors plus a third author— repeat all authors:

Matthew, W. D., W. Granger, and G. G. Simpson.
1928. Paleocene multituberculates from Mon-
golia. American Museum of Natural History
Novitates, no. 331:1-4.

4) Chapter in an edited volume:

Rausch, R. L. 1 963. A review of the distribution
of Holarctic mammals. Pp. 29^43, in Pacific
Basin Biography (J. L. Gressitt, ed.), Bishop
Museum Press, Honolulu, Hawaii, xx + 450

pp.

5) Unpublished dissertation:

Smith, J. P. 1976. Review of Eocene mammals.
Unpublished Ph.D. dissert., University of Cal-
ifornia, Berkeley, 302 pp.

6) Book:

White, M. J. D. 1961. The Chromosomes. Me-
thuen and Co., Ltd., London, 120 pp.

7) Journal articles with usual volume and issue number:

Anderson, W. I. 1 969. Lower Mississippian con-
odonts from northern Iowa. Journal of Pale-
ontology, 43(4):9 16-928.

Proof— The author should answer all queried proof
marks and check the entire proof copy. Return corrected
page proof with the edited manuscript promptly to the
editors.

issn 0097-4463

‘ AN NALS

of CARNEGIE MUSEUM

THE CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 63 30 NOVEMBER 1994 NUMBER 4

CONTENTS

ARTICLES

Archaeology of Trants, Montserrat. Part 1 . Field methods and artifact density

distributions David R» Watters 265

Archaeology of Trants, Montserrat. Part 2. Vertebrate fauna

Elizabeth J. Reitz 297

A new species of Antocha (Subgenus Orimargula) from Sulawesi (Diptera:

Tipulidae) and its mate-clasping behavior Chen W* Young 319

A revised classification of the spiriferid brachiopods John L. Carter,

Je Go Johnson, Remy Gourveneec, and Hou Hong-ffei 327

Index to Volume 63 375

Editors, ANNALS, BULLETIN and SPECIAL PUBLICATIONS:

K. C. Beard

L. Krishtalka
B. C. Livezey
D. R. Watters

M. A. Schmidt, ELS, Assistant Editor

Manuscripts, subscriptions, orders for individual numbers, and changes of address
should be sent to:

Office of Scientific Publications

The Carnegie Museum of Natural History

4400 Forbes Avenue

Pittsburgh, PA 15213-4080

Phone (412) 622-3287

Fax (412) 622-8837

E-mail: schmidtm@clp2.clpgh.org

THE CARNEGIE

MUSEUM OF
NATURAL HISTORY

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 4, Pp. 265-295

30 November 1994

ARCHAEOLOGY OF TRANTS, MONTSERRAT. PART 1.

FIELD METHODS AND ARTIFACT DENSITY DISTRIBUTIONS

David R. Watters

Associate Curator, Division of Anthropology

Abstract

Different research strategies used during archaeological fieldwork in 1978-79 and 1990 at the early
Ceramic Age site of Trants are compared and evaluated. Site size and the area of “major concentration”
of artifacts were both underestimated by the field methods employed during initial research in 1978-
79. The “collection corridor” strategy used in 1990 was an efficient and effective technique for sys-
tematic surface collection and, when combined with subsurface data from spaced test pits, it proved
to be a reliable method for distinguishing artifact densities and determining site parameters. Low-
density artifact distributions existing in peripheral fields were distinguished from a “most dense” area
(ca 90,000 m2) demarcated by surface and subsurface artifacts, in which a “core area” (ca 50,000 m2)
delineated the site’s oval-shaped configuration. A ca 500 B.C. initial occupation at Trants is substan-
tiated by the two oldest radiocarbon dates and by diagnostic pottery ascribed to the Saladoid ceramic
series. Terminal radiocarbon dates (ca A.D. 330) suggest that Trants was inhabited, either continuously
or intermittently, for perhaps 800 years. Trants is one of three early Ceramic Age sites on Montserrat
that have yielded Saladoid ceramics. This article is the first in a series of individually numbered parts
dealing with the results of archaeological field research at the Trants site.

Introduction

Trants is the largest and oldest of three confirmed early Ceramic Age sites on
Montserrat. At Trants, site disturbance by historic and recent cultivation is min-
imal, the stratigraphic integrity of its archaeological deposits is excellent, and the
preservation of its artifacts and ecofacts is superb.

The island of Montserrat is located in the Lesser Antilles, the archipelago
forming the eastern border of the Caribbean Sea and demarcating the western
edge of the Atlantic Ocean. These islands form an 850-km long arc from Sombrero
south to Grenada (Fig. 1). North of Guadeloupe, the Lesser Antilles islands form
a double arc, the Outer or Limestone Arc and Inner or Volcanic Arc. Montserrat
is one of the smaller (98 km2 volcanic islands on the Inner Arc of the northern
Lesser Antilles. Three major mountain masses, trending north to south down the
center of the island, dominate Montserrat’s landscape (Fig. 2). The Trants site is
situated east of the Centre Hills on the only sizeable stretch of relatively flat terrain
near sea level along the windward coast (Fig. 3). The rest of this coast is marked
by precipitous cliffs.

The Trants Estate, comprising the generally level terrain surrounding the Trants
site, encompasses a much larger area (ca 600,000 m2) and is bounded by mountains
to the west and north, the coast to the east, and a river valley to the south. Trants
Estate is named for Dominick Trant, a plantation owner in the 1720s (Marion
Wheeler, personal communication, 1990). Since the 1950s, this estate has been
the property of the Government of Montserrat. Trants Estate no longer is a working
agricultural plantation; instead it is used for pastureland and small garden plots.

Montserratians have collected Amerindian artifacts at Trants for many years.

Submitted: 1 November 1993.

265

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Fig. 1.— The arcuate Lesser Antilles stretch from Sombrero to Grenada. Sombrero lies about 100 km
east of the Virgin Islands, the easternmost islands of the Greater Antilles (see Fig. 2); Grenada lies
about 1 50 km north of the South American continent.

1994

Watters— Trants Archaeology, 1

267

Greater Antilles

Fig. 2.— The Trants prehistoric site is located midway on the east coast of Montserrat in the northern
Lesser Antilles.

Prehistoric beads were the focus of Harrington’s (1924) brief article, the first
publication to discuss artifacts from the site. The beads formed one part of a large
and diverse collection of artifacts from Trants acquired by the Museum of the
American Indian (MAI), Heye Foundation, from Seymour Wylde Howes, the
manager of Trants Estate (Watters and Scaglion, 1994). The lithic beads and
ceramics in the Howes collection were studied respectively by Watters and James
B. Petersen in 1985; they re-examined the MAI materials in 1991; and in 1994
Watters examined objects from the Howes collection that the MAI had exchanged
with the University Museum, University of Pennsylvania in 1932.

The first archaeological survey and excavation at Trants occurred during 1 978—
79, as part of Watters’ (1980) dissertation research on the islands of Montserrat
and Barbuda. Trants, the first site recorded in St. George’s Parish, was assigned
site number MS-G1. The second field season, which took place in 1990, involved
additional survey and testing at Trants, included a study of Trants artifacts housed
at the Montserrat National Trust Museum, and was a collaboration of the Carnegie
Museum of Natural History (CMNH) and the University of Maine at Farmington
(UMF). The CMNH-UMF project was jointly directed by Watters and Petersen.
The Trants site was observed briefly in 1983, 1984, and 1992, during short
stopovers on Montserrat made in the course of other research projects on nearby
islands.

Preliminary information about the 1990 field season has been published already
(Bartone and Crock, 1991; Petersen and Watters, 1991). Data from the 1978—79
project (Watters, 1980) are used in two regional studies: a report on the thin
section petrography of ceramics from four northern Lesser Antilles islands (Don-
ahue et al., 1990), and a compilation of Late Quaternary vertebrate faunas from
archaeological and paleontological contexts throughout the Lesser Antilles islands
(Pregill et al, 1994).

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Fig. 3.— A view (looking west) from an airport tower across the Trants site toward the Centre Hills.
Field 10 is the pasture in the foreground; Field 8 lies just beyond the treeline, where the cement block
building is visible (indicated by the arrow).

This article is Part 1 in a series of separately numbered reports dealing with
the results of the archaeological research at the Trants site. These reports focus
on the 1990 fieldwork although selected articles will deal with aspects of the 1 978—
79 research as well. Part 2 of this series reports the vertebrate fauna recovered
from three 1990 excavation units (Reitz, 1994) and combines those data with the
previously published vertebrate fauna from the 1978-79 research (Steadman et
al., 1984). Other parts will discuss sediment data and settlement patterns, lithic
technology, paleoethnobotanical remains, and invertebrate faunas. Data derived
from museum research on the S. W. Howes collection are being published as
separate articles (Watters and Scaglion, 1994), to divorce those reports from the
individually numbered parts of the Trants field research series.

The purposes of Part 1 in the Trants archaeology series are to: (1) compare and
evaluate field methods used in 1978-79 and 1990; (2) provide data on derived
artifact densities and spatial distributions; (3) interpret these data with respect to
site demarcation and configuration; (4) discuss the chronological placement of the
Trants site and its occupational span; (5) review the relationship of Trants to
Montserrat’s other six prehistoric sites; (6) and furnish the background infor-
mation, about the surface samples, test pits, and excavation units, that is integral
to the presentations in the forthcoming parts of this series.

Field Methods
Archaeological Surveys

Field research on Montserrat in 1978-79 was conducted by the author and one
field assistant. The strategy used in the survey to locate prehistoric sites involved

1994

Watters —Trants Archaeology, 1

269

Fig. 4. — Locations of the “major” and “minor” concentrations of surface artifacts identified during
initial fieldwork in 1978-79. Northing and easting based on the British West Indies (BWI) grid, which
differs from the site grid (see Fig. 1 7).

six randomly selected, 250 m-wide cross-island transects (Watters and Scaglion,
1980). The northern part of the Trants site was discovered near the eastern ter-
minus of Transect 28 at Trants Bay (Watters, 1 980:2 1 4-2 1 5). This area, designated
the “minor concentration,” was identified through surface artifacts found due
west of the north end of the Blackbume Airport runway (Fig. 4). A “major
concentration” of artifacts was detected subsequently in an area outside of the
Transect 28 boundaries, exposed on the surface of a field planted in sea island
cotton immediately north of the road to Blackbume Airport. Few surface artifacts
were observed in the other fields north of the airport road, most of which were
in grass and used as pasture; fields south of that road, being covered by thickets
of thorny “cassie” bushes, were unobservable for the most part. At the end of the
1979 survey, the minor and major concentrations were regarded as spatially
distinct sectors, perhaps representing two site components, that were separated
by an intervening area of about 350 m of grassy pasture having few artifacts.

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Fig. 5.— Systematic collection strategies used to obtain data on surface and subsurface artifact densities
at Trants during the 1990 fieldwork. Note locations of numbered fields (BWI grid).

Based on observed surface artifacts, the total area of the Trants site, combining
the major and minor concentrations, was estimated at about 42,500 m2 (Watters,
1980:table 8), or 4.25 hectares.

Brief visits to Montserrat in 1983 and 1984 reconfirmed the locations of the
major and minor concentrations and provided the opportunity to examine, for
the first time, the fields south of the airport road from which the cassie thickets
had been removed in the interim. Few artifacts were observed in these newly
cleared fields.

Different strategies were used during the CMNH-UMF project in 1990. The
ten-week field season involved seven people although not everyone participated
for the entire project (Petersen and Watters, 1991). The primary objectives were
to: (1) examine the relatively flat terrain (60 hectares) surrounding the Trants site,
from the coast to the mountains, for evidence of cultural materials; (2) define
more accurately the extent and boundaries of the site; and (3) determine the depths

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Watters - T rants Archaeology, 1

271

Fig. 6. —Crew members pin flagging artifacts in a 10 x 10 m surface collection unit in Corridor 1.

of its cultural deposits. These goals were paramount because the Trants site was
potentially threatened by two projects proposed for the adjacent Blackbume Air-
port: realignment of its runway, and construction of new airport buildings.

The archaeological survey area included the terrain east of the mountains to
the coastline between Trants Bay and Farm Bay (Fig. 5). The southern border
was Farm River Ghaut (ghaut is a term used locally to signify a steep-sided valley).
To manage the archaeological survey, the terrain was designated as Fields 1
through 15, using already existing roads, fencelines, earthen ridges, and wind-
breaks as the boundaries of the subdivisions.

Surface collections were made during the very dry month of September and in
early October when the seasonal rains were just starting. The exposure of the
ground surface by the die-back of vegetation enabled excellent observation of
surface artifacts.

Three systematic surface collection strategies were used in Fields 1, 2, and 3
south of the airport road (Fig. 5). Surface artifacts in all three fields were pin
flagged and counted, but the percentages shot-in by transit and collected varied
among the three fields. In general, artifacts were sparsely distributed in these
fields.

A similar systematic surface collection strategy initially was used in the southeast
sector of Field 8, immediately west of the cement block building, in an area
coinciding with part of the previously defined major concentration. The high
density of artifacts in Field 8, in comparison to their scarcity in Fields 1 through
3, required a change in strategy for surface collecting. A more efficient sampling
strategy was needed to maximize the systematic survey coverage and to facilitate
the comparison of spatial distributions of artifacts.

A “collection corridor” technique was devised to sample the surface artifacts.
It involved a linear arrangement of adjacent 10 x 10 m squares in which surface
artifacts were flagged (Fig. 6) and collected but not shot-in by transit. Corridor 1 ,
the corridor spanning the terrain (Fig. 5) from the mountains to the present airport
boundary, traversed Fields 13, 8, and 10 from west to east (Fig. 7). Corridor 2
traversed Fields 8 and 9 from south to north between the airport road and the
estate road (Fig. 5).

Data for two categories of artifacts, pottery and lithics, from the 56 collection

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vol. 63

Fig. 7.— A fully flagged 10 x 10 m surface collection unit in Corridor 1 in Field 8.

squares in Corridor 1 are included in this paper. When plotted by 10 x 10 m
surface squares, counts for both artifact categories have somewhat narrow dis-
tributions, with a decided peak consistently occurring toward the east side of Field
8 (Fig. 8). Field 13 and much of Field 10 yielded many fewer artifacts. When
Corridor 1 pottery and lithics are plotted by weight (Fig. 9), a similar overall trend
is evident (in spite of some minor perturbations for lithics in Field 10). These
surface findings corroborated the observations made in 1978-79, whereby the
major concentration of artifacts occurred mainly in Field 8 (Fig. 4).

Data from 21 collection squares crossing Fields 8 and 9 in Corridor 2 are
presented. Plots of surface pottery and lithic counts conform to one another (Fig.
10), but their distributions are distinctly different from those in Corridor 1. In
Corridor 2 plots, there are two distinct peaks, one in northern Field 8 and a second
in northern Field 9, while the intervening area has many fewer surface artifacts
(Fig. 1 1). A peak in Field 8 was anticipated from the previous fieldwork; the peak
in Field 9 was unexpected since that area fell outside of the boundary of the
previously defined major concentration.

Excavations

In February 1979, a single 2 x 2 m excavation unit was dug on the eastern
edge of the major concentration, in a pasture due east of the cement block building
(Fig. 4). Deposits were dug in 1 0-cm levels within the three strata (I— III) identified.
The upper 20 cm of Stratum I was a mixed deposit of historic and prehistoric
artifacts. Below this disturbed level, prehistoric deposits were intact to 80 cm
(Stratum III) although cultural materials were most abundant to 60 cm, in Stratum
II (Watters, 1980:table 17).

In 1990, the first ten test pits (0.5 x 0.5 m) were excavated south of the airport
road in Fields 1, 2, and 3. Test pits were aligned within transects and were widely

LITHICS COUNT POTTERY COUNT

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Watters- Trants Archaeology, 1

273

TRANTS SURFACE COLLECTION (10X10 m UNITS)

West/East Corridor N420-430

TRANTS SURFACE COLLECTION (10X10 m UNITS)

West/East Corridor N420-430

Fig. 8. —Surface pottery and lithic distributions by count in Corridor 1 (west to east) through Fields
13, 8, and 10.

LITHICS WEIGHT (GRAMS) POTTERY WEIGHT (GRAMS)

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TRANTS SURFACE COLLECTION (10X10 m UNITS)

West/East Corridor N420-430

Fig. 9. —Surface pottery and lithic distributions by weight in Corridor 1. Minor lithic “peaks” in Field
1 0 are the result of weighty cores found on the surface of those collection units.

E790-800

POTTERY COUNT

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TRANTS SURFACE COLLECTION

South/North corridor E570-580

Fig. 10. —Surface pottery and lithic distributions by count in Corridor 2 (south to north through Fields
8 and 9) showing two peaks with a drop in artifact frequency in the intervening area.

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Fig. 1 1. —Corridor 2 flags in Field 9 (looking north). Note the high density of flags (indicated by the
arrow), correlating to the artifact peak in the north part of Field 9, compared to the sparsely distributed
flags in that field’s central and southern parts (foreground).

dispersed in the fields. The basal depths of prehistoric materials ranged from as
shallow as 20 cm to as deep as 70 cm; cultural deposits extended deeper than 30
cm in only three test pits (Table 1). The quantities of artifacts within these test
pits were correspondingly limited.

The collection corridors revealed differentially distributed surface artifact den-
sities in Fields 8, 9, 10, and 1 3. However, one concern was that these distributions
might have been biased by surficial disturbance caused by cultivation, including
sugar cane grown during the historic period and sea island cotton more recently.
To investigate the possibility of biased surface distributions, a systematic sub-
surface testing program was used in the same two collection corridors. A test pit
(0.5 x 0.5 m) was dug every 50 m along the entire length of each corridor. This
testing program made it possible to compare the subsurface distributions in test
pits across the site; it also provided a means to compare surface and subsurface
artifact distributions within each collection corridor.

In the five test pits dug in Corridor 2, one in Field 8 and four in Field 9, the
basal depths of prehistoric artifacts ranged from 50 to 100 cm (Table 1). Pottery
and lithic plots (Fig. 1 2) reveal a definite frequency decrease that correlates very
well with the decrease seen in the Corridor 2 surface data (Fig. 10). The low
density area in each data set corresponds to the southcentral part of Field 9, where
the shallowest test pit also is located (Table 1).

Twelve test pits were dug in Corridor 1, four each in Fields 13, 8, and 10.
Shallower test pits generally occurred in Field 13 and the eastern part of Field 10
(Table 1). These test pits also yielded lesser quantities of pottery and lithic artifacts
(Fig. 13). The peak in the eastern part of Field 8, which was anticipated based on
the surface collected data, was present. However, a second peak for subsurface

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Table 1.— Excavations at the Trants site in 1979 and 1990.

Field

Location

Pre-
historic
depth (cm)

Remarks

1979 excavation unit (2.0 x ;

2.0 m)

east edge of major

prior to 1990 site grid

concentration

1990 test pits (0.5 x 0.5 m)

a. South of airport road in

= 10)

TIPI

T = Transect; P = Test Pit

T1P2

T2P1

T3P1

T3P2

T3P3

T4P1

T4P2

T4P3

T5P1

b. Corridor 1 test pits (n =

1 2; west to east)

N421 E245 SE

N421 E295 SE

N421 E345 SE

N421 E395 SE

N421 E445 SE

N421 E495 SE

N421 E545 SE

N421 E595 SE

N421 E645 SE

110

adjacent to unit N42 1/422 E645

N421 E695 SE

N421 E745 SE

N421 E795 SE

c. Corridor 2 test pits (n =

5; south to

north)

N395 E571 NW

100

adjacent to unit N396 E571

N445 E571 NW

N495 E571 NW

N545 E571 NW

N595 E571 NW

adjacent to unit N596 E571

d. Discretionary test pits (n = 2)

N407 E634 SW

near 1979 excavation unit

N521 E646 SW

100

1990 excavation units (1.0 x

1.0 m)

a. Corridor 1 excavation unit

N42 1/422 E645

110

adjacent to test pit N421 E645 SE; the unit quadrants ex-

cavated are N421 NE and NW and N422 SE and SW

b. Corridor 2 excavation units

N396 E571

100

adjacent to test pit N395 E571 NW

N596 E571

100

adjacent to test pit N595 E571 NW

POTTERY COUNT

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TRANTS TEST PITS (0.5 X 0.5m)

N-

til

South/North Corridor

Fig. 12. — Subsurface pottery and lithic distributions by count for the five test pits in Corridor 2, once
again displaying the marked decrease in artifact frequency (compare with Fig. 10).

LITHICS COUNT POTTERY COUNT

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West/East Corridor

Fig. 13.— Subsurface pottery and lithic distributions by count for the 12 test pits in Corridor 1. The
abundance of subsurface artifacts denoted by the peak in the west side of Field 10 is not duplicated
in the surface density plots (compare with Fig. 8).

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Fig. 14.— View looking west of the 1.0 x 1.0 m excavation unit (N42 1/422 E645) laid out beside its
adjacent (already dug) test pit (N421 E645 SE) in Corridor 1 in Field 10 (see also Fig. 21).

pottery and lithics is evident in the western part of Field 1 0 (Fig. 1 3), an area
where no peak was expected based on the surface data. Lithic frequency actually
is slightly greater for the Field 10 peak. Equally intriguing was an unexpected
frequency decrease, for both pottery and lithics, in the easternmost test pit of
Field 8, between the two peaks. For Corridor 1, the overall distribution patterns
for surface and subsurface artifacts across Fields 13, 8, and 10 were analogous
(Fig. 8, 13). However, it was the test pit plots alone that pointed out the significance
of the subsurface remains in the western edge of Field 10 for defining the extent
of the site.

Two “discretionary” test pits were excavated in Field 10. One was positioned
in the southern part of that field near the 2.0 x 2.0 excavation unit dug in 1979,
in order to compare their stratigraphies. The maximum depth of cultural materials
in this test pit was 80 cm, the same as the nearby 1979 unit (Table 1). A second
discretionary test pit was dug in the northern part of Field 1 0, an area where no
subsurface data and no systematically collected surface data were available. This
test pit contained prehistoric artifacts to about 100 cm; it also yielded the only
complete ceramic vessel excavated at the Trants site.

Three excavation units (each 1.0 x 1.0 m) were dug during the final phase of
the 1990 fieldwork. They were dug adjacent to the three test pits providing the
best stratigraphic information at key areas of the site (Fig. 14). Excavation units
were placed in Field 8, northern Field 9, and southern Field 10. Prehistoric
artifacts, which were recovered from three field strata (A-C) in the excavation
units (Petersen and Watters, 1991), reached depths of 100 to 1 10 cm (Table 1).
Because these excavation units are so widely dispersed, a site-wide correlation is
equivocal for the three field strata identified in the three units in 1990. Also
equivocal is an association between the three strata (I— III) identified in the 1979
excavation unit and the three field strata (A-C) from the 1990 units. Although
stratigraphic correlations across the site cannot be made at this time, the recurrence
of three strata having cultural materials in all four excavation units is noteworthy.

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Fig. 1 5. — Panoramic view (looking SSE) of the Trants site. A, south part of airport runaway; B, cement
block building in the treeline; C, Farm River, the southern border of the site; D, lower slope of the
Centre Hills. Fields 1-3 and 8-13 are the primary fields mentioned in the text (compare with Fig. 5).

Mixed deposits containing historic and prehistoric artifacts were, for the most
part, stratigraphically restricted to field stratum A (and Stratum I from 1979),
especially to its uppermost levels (ca 0-25 cm). Beneath this mixed layer, the
undisturbed prehistoric deposits yielded the kinds of artifacts and ecofacts (ce-
ramics, lithics, bone, crab, mollusks, and coral) that routinely recur in Ceramic
Age middens in the West Indies.

A total of 29 test pits, having a surface area of 7.25 m2 were dug in Fields 1,
2, 3, 8, 9, 10, and 13. When the 3-m2 area of the three excavations units is added,
a total of 10.25 m2 of surface area was excavated in 1990. Combining that total
with the 4-m2 surface area dug in 1979, a cumulative total of 14.25 m2 of surface
area has been excavated at Trants.

Field Methods Summary

In one sense, the Trants site comprises virtually all of the 60 hectares of relatively
flat land stretching eastward from the mountains to the coast (Fig. 15). Surface
artifacts were found in every field examined but their densities varied greatly
across the site. Fields nearest to the mountains, south of the airport road, and
north of the estate road had fewer artifacts. Within the broader landscape, the
surface and subsurface distributions reveal a “most dense” area of artifacts, the
extent of which is still not fully determined but likely encompasses major portions
of Fields 8, 9, 10, and 1 1 (Fig. 16). This ‘‘most dense” area covers about 90,000
m2 (9 hectares) and incorporates all of the “major concentration” but not the
“minor concentration” defined in 1978-79. Interviews in 1990 with Montser-
ratians involved in constructing Blackbume Airport, which opened in 1957

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Fig. 16.— An overall “most dense” area of artifacts, within the broader terrain, covered most of Fields

8, 9, 10, and 11 (BWI grid).

(Wheeler, 1988:56), revealed that the minor concentration really was an artifact
scatter exposed by bulldozers scraping the adjacent land for fill to extend the
northern end of the original runway. The minor concentration, even though it is
situated well away from the “most dense” area, deserves further study since the
depth and extent of its cultural deposits have yet to be determined.

Site Configuration

After the 1978-79 fieldwork, the Trants site was deemed to be a prehistoric
village, the extent of which was approximately co-terminus with the major con-
centration (Watters, 1980:249-252). Some aspects of this view changed dramat-
ically following the 1990 fieldwork.

The surface collection squares and test pits in Corridor 1 confirmed dense
artifact distributions in Fields 8 and 10. In Corridor 2, the collection squares
revealed a pattern previously unrecognized, with a significant decrease in artifact

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density in the middle of the corridor and a marked increase in northern Field 9.
Five test pits and two excavation units in Corridor 2 corroborated the variable
densities initially observed on the surface.

The pattern that was beginning to emerge altered previous perceptions of the
Trants site because, for the first time, a “core area” was recognized (Fig. 17). The
highest artifact densities were obtained in two sectors, the northern sector (north-
ern part of Field 9) and the southern sector (Field 8 and the southern part of Field
10), and these sectors are separated by 100 m of sparsely distributed artifacts
(central-southern part of Field 9). The southern sector maximally extends east-
west about 250 m based on Corridor 1 data. The northern sector’s length is less
well-determined because this part of the site had no corresponding east-west
corridor, and therefore no equivalent systematically collected, quantifiable data.
It has a length of at least 150 m, based on observed surface artifacts, and likely
extends further eastward judging from the discretionary test pit in northern Field
10 (Fig. 17). The core area’s northern and southern sectors consist of opposing
but seemingly complementary curvilinear segments demarcated by dense artifact
distributions.

Although the northern and southern sectors were fairly well-defined during the
1 990 fieldwork, little was known about the eastern and western margins, the areas
that potentially linked the two sectors. Surface artifact scatters indicated that these
connections likely existed. The presence of the connecting link on the eastern
margin was strongly supported by data from the discretionary test pit in northern
Field 10 (Fig. 17). This isolated test pit, the sole source of information for that
part of the site, attained a depth of 1 00 cm with clearly defined stratigraphic layers
and it yielded abundant artifacts.

The western margin, in the more westerly part of Field 9, was the least known
area of the Trants site at the close of the 1990 fieldwork. Corridor 2 was situated
in the eastern part of Field 9 (Fig. 5) and did not traverse this western margin.
In 1992, eight backhoe “trial” or “borrow” pits, dug in connection with the
planned airport development, provided subsurface data about the western margin.
During their brief visit to Montserrat in May 1992, Watters and Petersen were
able to examine the open trial pits, which the contractor had left unfilled as
requested. Four pits each had been dug in Fields 8 and 9. Two Field 9 pits (B6
and B7 on Fig. 1 7), located midway between the northern and southern sectors,
were especially important with respect to the western margin. These pits (Fig. 18)
contained midden material, including abundant artifacts and faunal remains, in
stratified cultural levels that paralleled what had been observed in other excavated
areas of Trants. Even more strongly than was the case for the one discretionary
test pit on the eastern margin, the two borrow pits in Field 9 support the existence
of the western link between the northern and southern sectors. Another trial pit,
on the eastern edge of Field 9, also proved to be significant since it was located
in the area of low artifact density identified in the Corridor 2 research. This pit
had few cultural remains, an observation that matched the findings for the nearby
shallow test pit excavated in Corridor 2 (Fig. 1 7).

Chronology and Occupation

Eight radiocarbon dates, all obtained from charcoal samples in excavated con-
texts, are available for the Trants site. Two are from the 1979 excavation unit
and six from the 1990 fieldwork (Table 2). Samples from Trants provide two of
the oldest dates from the Ceramic Age in the Lesser Antilles, at 2430 ± 80 B.P.

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Fig. 1 7. —The oval-shaped “core area” of the Trants site, as demarcated by the northern and southern
sectors and the less well-defined eastern and western margins (site grid).

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Fig. 18.— Backhoe “trial” pit B7 dug in connection with future development of the new airport.

(Beta-44828) and 2390 ± 90 B.P. (Beta-41682), or respectively ca 480 and 440
B.C. (uncorrected), both of which were obtained from field stratum C, the deepest
cultural stratum in their respective excavation units. The oldest date is from the
excavation unit (N396E571) in Field 8 (Fig. 17, Unit D) of the southern sector
of the site; the next oldest date is from the unit (N596E571) in Field 9 (Fig. 17,
Unit C) of the northern sector; the samples are separated by 200 m. These two
dates are from midden deposits in excavation units located on their respective
outer edges of the opposing curvilinear sectors and midway on the circumference
of the oval (Fig. 17). Another radiocarbon date for the northern sector excavation
unit (Fig. 17, Unit C), 1890 ± 70 B.P. (Beta-41678), or ca A.D. 60, is from field
stratum B and in correct stratigraphic relationship (Table 2).

Five radiocarbon samples from the southeast portion of the Trants site in Field

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Table 2.— Chronological ordering of radiocarbon dated samples from various contexts at Trants.

Radiocarbon

years B.P.

Conversion

(uncorrected)

Lab sample

Unit

Contexts

Depth*

Field

Corridor

2430 ± 80

480 B.C.

Beta-44828

N396E571

50-70

2390 ± 90

440 B.C.

Beta-41682

N596E571

70-80

2140 ± 80

190 B.C.

Beta- 18489

1979 imitb

40-50c

n/a

1960 ± 90

10 B.C.

Beta-41680

N422E645

70-80

1890 ± 70

A.D. 60

Beta-41678

N596E571

30-40

1750 ± 80

A.D. 200

Beta-41679

N407E634d

40-60

n/a

1740 ± 90

A.D. 210

Beta-41681

N422E645

1620 ± 90

A.D. 330

Beta- 18582

1979 unitb

40-50c

n/a

a Depth in cm below ground surface.

b This unit was dug in 1979, prior to laying out the site grid.
c Same stratigraphic level for these two dates.
d This discretionary test pit was not within a corridor.

10 are dated. Two dates are from the same level (40-50 cm in Stratum II) in the
1979 excavation unit (Fig. 17, Unit A), 2140 ± 80 B.P. (Beta- 18489) and 1620
± 90 (Beta- 18582), or respectively ca 190 B.C. and A.D. 330 (Table 2). These
widely divergent dates, separated by 520 rcy, may result from the small sample
sizes. To try to resolve the problem of the discordant dates, a charcoal sample
was submitted from field stratum B (40-60 cm) of the discretionary test pit in
southern Field 10, the test pit nearest the 1979 unit. This procedure assumes
stratigraphic correlation between Stratum II from 1979 and field stratum B from
1990. The derived date, 1750 ± 80 B.P. (Beta-41679) or ca A.D. 200, accords
more closely with the younger date (A.D. 330) of the 1979 unit. The final two
dates, 1960 ± 90 B.P. (Beta-41680) and 1740 ± 90 B.P. (Beta-41681), ca 10 B.C.
and A.D. 210, are from field stratum C in the single Corridor 2 excavation unit
(N422E645) in Field 10 (Fig. 17, Unit B). These two dates are in proper strati-
graphic context.

Temporal and spatial aspects of the radiocarbon dates merit discussion. Samples
separated stratigraphically have the correct temporal relationship in two exca-
vation units. Two samples from one level in another excavation unit produced
discordant dates, and it is the older (190 B.C.) of the two dates that is suspect,
based on crossdating of a stratum in a nearby test pit. These eight dates (Table
2) span either 810 rcy or 690 rcy depending on the terminal date used (A.D. 330
or A.D. 210, respectively). If the suspect 190 B.C. date in the 1979 unit is ignored,
the four remaining dates from the southeast part of the oval (Field 10) span 340
rcy (10 B.C.-A.D 330). Three of those four dates fall within a 130 rcy span (A.D.
200-330). The units producing the Trants site’s two oldest dates are located further
west, midway across the northern and southern sectors of the oval in Fields 8
and 9 (Fig. 1 7). The northern unit also has one date (A.D. 60) contemporary with
the earlier part of the 340 rcy span from Field 10. There are no dates from the
western part of the Trants site.

Ceramics recovered during the 1978-79 and 1990 field seasons include white-
on-red (WOR) painted and zoned-incised-crosshatched (ZIC) sherds (Fig. 19, 20).
Such decoration is regarded as diagnostic of the Saladoid ceramic series, and as
evidence for the earliest Ceramic Age population movement into the Caribbean
(Rouse, 1992; Siegel, 1991). Although the decorated sherds are useful diagnos-
tically, they constitute only 7.5% of the 1990 ceramic sample from Trants studied
so far; even when slipped sherds (that otherwise are undecorated) are added to

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Fig. 19.— Six white-on-red painted sherds recovered from the 1979 excavation unit. Artifact FSNs
(Field Specimen Numbers): A, MS-G1-955; B, MS-G1-510; C, MS-G1-953; D, MS-G1-788; E, MS-
Gl-572; F, MS-G1-966. (Photograph courtesy of James B. Petersen.)

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Fig. 20.— Three renderings of zoned-incised-crosshatched sherds recovered from the 1979 excavation unit. Artifact FSNs: A, MS-G1-806, B, MS-G 1-476
C, MS-G 1-292 and MS-G 1-293.

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decorated sherds, the cumulative total is only 14.4% (Petersen and Watters, in
press). At Trants, WOR and ZIC sherds commingle in excavation units and test
pits rather than being spatially segregated.

The radiocarbon dates and diagnostic artifacts confirm an early Ceramic Age
occupation at Trants, generally corresponding (but beginning earlier) to Period
Ha (300 B.C.-A.D.400) in the chronological framework established by Rouse
(1992:fig. 26). The field research carried out to date cannot answer, with certainty,
whether the Trants site was inhabited continuously or occupied intermittently
during the 810 (or 690) rcy span. Also uncertain is whether the oval was formed
simultaneously, resulting from a single habitation event that was spatially con-
tiguous, or was formed episodically, resulting from multiple habitation events
that were spatially discontinuous but which eventually coalesced to create the
oval. The configuration of the oval (assuming full closure on the eastern and
western margins) is certain; the site formation processes that created the oval are
uncertain. Early Ceramic Age occupation at Trants is certain; the continuity of
that occupation is not.

Discussion
Field Methods

The collection corridor technique was the most efficient (in terms of time ex-
pended) and effective (in terms of quantifiable data secured) of the systematic
surface collection strategies used at the Trants site. It provided the broadest
coverage of the terrain and distinguished the “most dense” and “core area” from
that terrain. The 10 x 10 m collection squares were laid out rapidly using tapes
and chaining pins; artifacts in each square were quickly pin flagged, counted, and
collected; and photographs taken of the individual squares and the corridors
provided permanent visual records of the relative densities of flagged artifacts
(Fig. 7, 11). The corridor technique did result in the forfeiture of certain prove-
nience data, notably the loss of precise locations for the surface artifacts within
each square. Loss of these data was deemed acceptable since the site surface already
had been disturbed by cultivation, thereby making that level of precision some-
what inconsequential and superfluous.

The collection corridor technique proved to be effective but it could have been
even more efficient. Surface artifact density plots (Fig. 8, 9) for Corridor 1 show
that two patterns, one of artifact scarcity in Fields 1 3 and 1 0 and one of abundance
in the east part of Field 8, would have been detectable even when the total number
of collected squares was reduced considerably. In the CMNH-UMF project, every
square (n = 56) was collected for the entire length of Corridor 1. Yet, the overall
distribution would still have been evident even if only every second square ( n =
28) or perhaps every third square (n = 18) had been collected instead. The same
rationale applies to the 21 collection squares in Corridor 2 (Fig. 10). Had the
conformity of the patterns been recognized during the fieldwork at the Trants site
in 1990, the “skipping” squares rationale could have been used in Corridors 1
and 2, thereby freeing up time for doing additional east-west and north-south
collection corridors and thus providing even more extensive spatial coverage of
the terrain and site.

Although the same rationale theoretically could be applied to corridor test pits,
an examination of the plots indicates a decision to skip certain test pits would
result in different distribution patterns. Skipping any test pit would have been a
precarious exercise in Corridor 2 since it contained only five test pits (Fig. 12).

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Increasing the interval between the 12 test pits in Corridor 1 would modify its
overall pattern less drastically than Corridor 2, and skipping one or two test pits
in Field 1 3 would not have greatly affected the low-density pattern observable in
that particular field (Fig. 13). However, had the single test pit in westernmost
Field 1 0 been skipped over, its elimination would have obscured completely the
“unanticipated” subsurface peak which originally called attention to the eastward
extension of the site. For the “most dense” and “core area” at Trants, a 50-m
space between test pits appears to be the minimum interval needed to maintain
the integrity of the subsurface density pattern. The low-density peripheral fields
are suitable for more widely spaced test pits.

The collection corridor strategy corroborated some previous findings such as
the major concentration existing in Field 8, but it also revealed new information
on artifact distributions, such as the frequency decreases in surface and subsurface
artifacts in Corridor 2 and the eastward extension of the site into Field 10. This
technique certainly fulfilled the objectives of demarcating the extent and defining
the boundaries of the Trants site, but it could have been made even more efficient,
by reducing the number of surface collected squares, without compromising the
integrity of the overall surface distribution patterns in each corridor.

Spatial Parameters

At the close of the 1990 CMNH-UMF project, the investigated terrain was
interpreted as three spatially distinct areas. The first, the peripheral area , was
demarcated by the modem fields having sparsely distributed surface artifacts. The
low-density determinations resulted either from systematic sampling or from less
reliable superficial observations made while walking over certain fields. The pe-
ripheral area encompassed Fields 1-6 south of the airport road, Field 12 north
of the estate road, and Fields 13-15 at the base of the mountains (Fig. 5). Field
7, the Blackbume Airport, was the only part not observed at all in the entire 60-
hectare terrain.

The second distinctive area, the most dense area, comprises the higher-density
distributions determined primarily from the surface artifact plots in Corridors 1
and 2. The most dense area incorporates almost all of Fields 8, 9, 10, and 1 1 (Fig.
1 6). Its maximum extent likely is somewhat biased along the northern, western,
and southern boundaries by artifacts that were brought to the surface through
construction of the roads that correspond to those borders.

The third distinctive area is the core area which lies wholly within the most
dense area. It certainly incorporates parts of Fields 8, 9, and 10 and probably part
of Field 1 1 (Fig. 1 7). The circumference of the core area includes the fairly well-
defined, curvilinear-shaped northern and southern sectors and the suggestive, but
less firmly established, linking sectors on its eastern and western margins. The
one discretionary test pit in northern Field 10 and the two trial pits in Field 9
substantiate the existence of these connections. The boundaries of the core area
delineate an oval configuration having an east-west length of as much as 250 m
and north-south extent of 200 m. The middle of the oval is the low-density area
identified in the Corridor 2 plots. The size of the core area alone (50,000 m2, or
5 hectares) is larger than the total site area (42,500 m2) estimated after the 1 978—
79 fieldwork.

In overview, the peripheral area comprises the relatively flat 600,000-m2 terrain,
bounded by the mountains, coast, and river valley, in the midst of which is the

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most dense area of 90,000 m2 which in turn encompasses the 50,000-m2 core area
of the Trants site itself.

Oval Configured Sites

Analysis of spatial patterning in this article relies primarily on artifact distri-
butions and secondarily on radiocarbon date distributions. Another data set,
sediment analyses and distributions, provides independent support for the exis-
tence of the peripheral area, in which low artifact densities and low sediment
values correlate, and the core area, in which high artifact densities, high sediment
values, and the occurrence of anthrosols also correlate. Petersen and Watters
(1991) presented the sediment data in a preliminary manner; a detailed analysis
of grain size characterization, pH, organic matter, and phosphate fractionation
will be provided by Petersen in a separate part of the Trants field research series.

Within the core area of the Trants site are two well-defined areas, the northern
and southern sectors (apparently extending to the eastern and western margins)
consisting of middens with high densities of artifacts and ecofacts and high sed-
iment values, and a site center comprising a 100-m -diameter area with low artifact
density. Although the site center is thought to represent an area of “common
ground” or “community space,” it is premature to label this area a “plaza” or
“dance-court” (cf. Alegria, 1983).

The Trants site’s oval configuration is demarcated by the encircling middens.
The two oldest dates (480 and 440 B.C.), from excavation units opposite one
another on the oval, indicate contemporaneous occupation for those particular
points along the northern and southern sectors. Four other dates indicate more
recent habitation of the oval’s southeast segment. Presently undated are other
midden segments on the oval’s circumference and the site center.

Circular, ring, or oval configured sites occur elsewhere in the Caribbean. Siegel
(1992:372) uses a concentric “ring model” to characterize the Maisabel site on
the north coast of Puerto Rico (Fig. 2). The site plan (Siegel, 1992:fig. 6.2) depicts
a “cleared central portion of the village,” encircled by a “dense midden accu-
mulation” (incorporating five mounded middens), in turn surrounded by a “low
dense midden accumulation defining site periphery.” Low densities of artifacts
in the central portion and the periphery contrast with the high density midden
accumulation. A cemetery was discovered in the central portion of the Maisabel
site. Siegel (1992:126-132, 188-191) contends that: (1) the overall settlement
groundplan was established during initial occupation by Saladoid colonists; (2)
the four areas of the site he studied were occupied contemporaneously; and (3)
the site was continuously occupied. Saladoid and Ostionoid occupation (ca 100
B.C.-A.D. 1200) at Maisabel spans part of Period Ha and all of Periods lib, Ilia,
and IHb in Rouse’s (1992) scheme. The Puerto Rican site of Punta Candelero,
which has an earlier Huecan component with linearly arranged mounds, attains
a semicircular configuration during its later Cuevas component (Rodriguez, 1991:
fig. 6).

The Golden Rock site on St. Eustatius (Fig. 2) apparently had a circular layout
(Versteeg and Schinkel, 1992:212). The report summarizes the spatial data derived
from research by De Josselin De Jong in 1923, during which he mapped five
distinct midden clusters. Three clusters subsequently were damaged by airport
development. One extant cluster (GR-1), the focus of excavations in the 1980s,
contained a midden, house structures, and burials; a presumably complementary

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midden cluster (GR-2) is situated opposite GR-1 (Versteeg and Schinkel, 1992:
fig. 178). Occupation (ca A.D. 600-800) at Golden Rock is primarily Period Ilia
but may extend back into Period lib.

Rouse (1974:167, 1978:703) characterizes the Indian Creek site on Antigua
(Fig. 2) as an oval ring or a circle, with a concentration of shell refuse in a series
of five middens around the periphery of an oval area. Radiocarbon dates indicate
that Indian Creek encompasses the latter part of Period Ila and all of Periods lib.
Ilia, and Illb. Rouse (1974:1 68) mentions two other oval-shaped sites surrounded
by middens, Ostiones on Puerto Rico and Carrier in Haiti.

The oval configured core area at the Trants site most closely parallels the “ring
model” at Maisabel (Siegel, 1992:fig. 6.2), with respect to their comparable low-
density site centers, high-density midden areas, and low-density peripheries. Spa-
tially distinct mounded middens at Maisabel, midden clusters at Golden Rock,
and middens at Indian Creek and Punta Candelero have no counterparts yet
identified at Trants, where the midden accumulation is continuous around the
oval rather than existing as discrete entities. Structural remains have yet to be
detected at Trants but the area where they most likely would occur, at the interface
of the site center and midden areas based on findings at Maisabel and Golden
Rock, has received minimal subsurface testing. A counterpart at Trants for the
Maisabel cemetery within the site center is also unknown. Although comparisons
between specific intrasite components at Trants and Maisabel are not warranted
at this time, their strikingly similar overall configurations suggest future compar-
isons, following larger-scale excavations at Trants, will be fruitful.

Montserrat ’s Ceramic Age Sites

Trants is one of the oldest Ceramic Age sites in the Lesser Antilles. Only two
sites, Hope Estate on St. Martin (560 B.C.) and Fond Brule on Martinique (530
B.C.), report older Ceramic Age radiocarbon dates. The dates from Trants and
from these islands support a migration by ceramic-producing peoples into the
Lesser Antilles at a much earlier date (by at least 500 years) than was previously
accepted (Rouse, 1989; Siegel, 1991). Trants is an important component of this
500 B.C. population movement.

Equally important for establishing this population movement are the other early
Ceramic Age sites on Montserrat. The Radio Antilles site (MS-A1) on the island’s
south coast (Fig. 2) has a single radiocarbon date, 2390 ± 60 B.P. (Beta- 18491),
or 440 B.C. (uncorrected), equivalent to the second oldest date from Trants. There
are two other early dates from Radio Antilles, 2210 ± 70 B.P. (Beta- 18490) and
2120 ± 60 B.P. (Beta- 10582), or respectively 260 B.C. and 170 B.C. Excavations
at Trants and Radio Antilles have produced numerous examples of commingled
WOR and ZIC sherds as well as vessel forms associated with the Saladoid ceramic
series.

More recently, a third Montserrat site containing Saladoid ceramics, the Belham
Valley site on the island’s west coast (Fig. 2), has been identified from artifacts
surface collected in 1964 by Walter Kenyon and curated at the Royal Ontario
Museum (ROM). During Kenyon’s visit, the Belham Valley site was in the process
of being covered over (perhaps destroyed) by grading for a golf course. In 1979,
informants notified Watters (1980:237) of the existence of the Belham Valley site
but his reconnaissance of the golf course fairways failed to detect any surface
artifacts. The materials in the ROM collection are the only artifacts from this site
available for study. The Belham Valley collection, kindly loaned by ROM to

1994

Watters— Trants Archaeology, 1

293

Fig. 21.— Stratigraphy of the east wall (1.5 m length) of the excavation unit and its adjacent test pit
in Field 10 (see also Fig. 14).

CMNH for the past two years, has been studied by Petersen and Watters. It
includes 300 individual vessels, defined primarily by rim sherds, and contains
painted, incised, and undecorated sherds ascribed to the Saladoid ceramic series.
Diagnostic WOR and ZIC artifacts confirm that Belham Valley is Montserrat’s
third early Ceramic Age site. The absence of materials suitable for radiocarbon
dating in the ROM collection means that more precise information on the duration
of occupation at Belham Valley is unavailable.

Early Ceramic Age colonization of Montserrat is documented by Saladoid ce-
ramics in three very widely dispersed sites on relatively flat land occurring near
sea level on the east, south, and west coasts (Fig. 2). Radiocarbon dates at the
sizeable sites of Trants and Radio Antilles document early settlement and suggest
these occupations were sustained. The area and duration of occupation of the
Belham Valley site are unknown, but the site’s location in the largest valley on
the island and its proximity to the largest permanent flowing river suggest it also
was a major habitation site. The other prehistoric sites on Montserrat, Windward
Bluff, Farnsworth, Dagenham Beach, and Little Bay, are more recent post-Saladoid
sites containing no Saladoid ceramics.

Conclusions

The Trants site presently is interpreted as a manifestation of the initial Ceramic
Age population movement into the Lesser Antilles at about 500 B.C. Small scale
excavations (Fig. 21) within the core area recovered common midden materials
such as pottery, lithic tools and debitage, shell and stone beads, and a variety of
vertebrate, molluscan, crab, and coral remains. The pottery is attributed primarily

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to the early part of the Saladoid ceramic series (Petersen and Watters, in press).
Late Saladoid and post-Saladoid ceramics are absent in the Trants excavations.

Spatial distribution of the two oldest radiocarbon dates, one each from the
northern and southern midden sectors, indicates occupation was contemporary
in these areas of the Trants site. The site’s terminal date (A.D. 330) provides a
timespan of some 810 rcy, which incorporates almost all of Period Ha and extends
back into latest Period lb. Dates ranging from 10 B.C.-A.D. 330 on the oval’s
southeast part support the idea of a sustained occupation in that sector. Whether
the site was inhabited continuously or occupied intermittently during that 810
rcy span is uncertain.

Surface and subsurface artifact distributions and sediment distributions delin-
eate a “core area” having an oval-shaped configuration (assuming full closure on
the eastern and western margins) with maximum dimensions of about 250 x 200
m. The intervening area (ca 100 m between the northern and southern sectors)
exhibits definite decreases in surface and subsurface artifact densities as well as
lower sediment values. Whether the oval configuration was laid down simulta-
neously or episodically at Trants is uncertain. Younger oval- or circular-shaped
sites are reported from Golden Rock, Indian Creek, Maisabel, and Punta Can-
delero. These sites for the most part are more recent than the earliest dates at
Trants although the Indian Creek, Maisabel, and Punta Candelero (Huecan com-
ponent) sites include occupations in the later part of Period Ha.

The postulated overall site structure at Trants includes a centrally located “com-
mon ground” (the site center having sparse artifacts) that is surrounded by struc-
tures (yet to be detected) behind which are refuse areas (represented by the mid-
dens), all being contained in the core area of the site. Beyond the midden
accumulation is the site periphery, which is indicated by the modem fields having
low-density artifact distributions.

Field methods used in the 1990 CMNH-UMF project, notably the collection
corridor strategy for surface and subsurface sampling, fulfilled the objectives of
delineating the extent of the Trants site and determining its overall configuration
and boundaries. This field research has verified that Trants is one of the oldest
early Ceramic Age sites in the Lesser Antilles, and forthcoming parts of the Trants
series will show that Trants also is one of the best-preserved sites in the region.

Acknowledgments

Research in 1978-79 was supported by a Fulbright-Hays Doctoral Dissertation Research Abroad
Fellowship (DHEW:OE) and an Andrew Mellon Pre-doctoral Fellowship from the University of
Pittsburgh. The 1990 research was funded by the Netting and O’Neil Research Funds of Carnegie
Museum of Natural History (CMNH), the Center for Latin American Studies (University of Pittsburgh),
and the University of Maine at Farmington (UMF). The project was co-directed by James B. Petersen;
the field crew included Cathy Watters, Robert Bartone, Elizabeth Subin, John Crock, and Ellie Cowie,
and, for frequent on-site consultations, Jah Rebel. James B. Richardson III assisted with the 1992
trial pit study. The author acknowledges the Government of Montserrat for authorizing the fieldwork,
and Ken Sparkes, then Director of Public Works, for providing plans of the proposed airport devel-
opment; the Montserrat National Trust, especially its President Franklin Margetson, for serving as
liaison with Government and issuing the export permit for artifacts analyzed in the U.S.; Erica Gibbs
for providing the crew with delightful field accommodations; Cedric and Carol Osborne, Bert and
Marion Wheeler, and Parker and Meg McChesney, for assistance and advice on various matters; Mrs.
Sarita Weekes-Francis and Grace White for arranging site visits by students from Salem, Harris, and
Bethel schools; Elizabeth Subin, Robert Bartone, and John Crock (UMF) and Tara Reilly, Andrew
Fisher, and Ann Fleming (CMNH) for laboratory assistance; Belinda Cox (UMF), Carrie Pantier,
Nancy Perkins, and Gail Richards (CMNH) for graphics; and Sylvia Keller (CMNH) for preparation
of tables. The author acknowledges the careful reading and constructive criticism of the reviewers.

1994

Watters— Trants Archaeology, 1

295

Earlier versions of this paper were presented in 1 993 at a Trants symposium at the Society for American
Archaeology annual meeting in St. Louis and at the 1 5th International Congress for Caribbean Ar-
chaeology, San Juan, Puerto Rico.

Literature Cited

Alegria, R. A. 1983. Ball courts and ceremonial plazas in the West Indies. Yale University Pub-
lications in Anthropology, 79, 185 pp.

Donahue, J., D. R. Watters, and S. Millspaugh. 1990. Thin section petrography of northern
Lesser Antilles ceramics. Geoarchaeology, 5:229-254.

Harrington, M. R. 1924. A West Indian gem center. Indian Notes, 1:184-189.

— . In press. A preliminary analysis of Amerindian ceramics from the Trants site, Montserrat.

Proceedings of the Fifteenth International Congress for Caribbean Archaeology (M. Rodriguez,
ed.), Centro de Estudios Avanzados de Puerto Rico y el Caribe.

Pregill, G. K., D. W. Steadman, and D. R. Watters. 1994. Late Quaternary vertebrate faunas
of the Lesser Antilles: Historical components of Caribbean biogeography. Bulletin of Carnegie
Museum of Natural History, 30, 51 pp.

Reitz, E. J. 1994. Archaeology of Trants, Montserrat. Part 2. Vertebrate fauna. Annals of Carnegie
Museum, 63:297-317.

Rouse, I. 1974. The Indian Creek excavations. Pp. 1 66-176, in Proceedings of the Fifth International
Congress for the Study of Pre-Columbian Cultures of the Lesser Antilles (R. P. Bullen, ed.),
Antigua Archaeological Society, 178 pp.

. 1978. Cultural development on Antigua, West Indies: A progress report. Actas del XLI

Congreso Intemacional de Americanistas, 3:701-709.

. 1989. Peoples and cultures of the Saladoid frontier in the Greater Antilles. Pp. 383-403, in

Early Ceramic Population Lifeways and Adaptive Strategies in the Caribbean (P. E. Siegel, ed.),
BAR International Series, 506, 418 pp.

— -. 1992. The Tainos: Rise and Decline of the People Who Greeted Columbus. Yale University

Press, New Haven, 232 pp.

Siegel, P. E. 1991. Migration research in Saladoid archaeology: A review. Florida Anthropologist
44:79-91.

— . 1 992. Ideology, power, and social complexity in prehistoric Puerto Rico. Unpublished Ph.D.

dissert. State University of New York at Binghamton, Binghamton, 806 pp. [University Microfilms
#9219121].

Steadman, D. W., D. R. Watters, E. J. Reitz, and G. K. Pregill. 1984. Vertebrates from ar-
chaeological sites on Montserrat, West Indies. Annals of Carnegie Museum, 53:1-29.

Versteeg, A. H., and K. Schinkel (eds.) 1992. The archaeology of St. Eustatius: The Golden Rock
site. St. Eustatius Historical Foundation Publication 2, 284 pp.

Watters, David R. 1980. Transect surveying and prehistoric site locations on Barbuda and Mont-
serrat, Leeward Islands, West Indies. Unpublished Ph.D. dissert. University of Pittsburgh, Pitts-
burgh, 416 pp. [University Microfilms #81 12643].

Watters, D. R., and R. Scaglion. 1980. Utility of a transect survey technique in Caribbean
prehistoric studies: Applications on Barbuda and Montserrat. Pp. 338-347, in Proceedings of the
Eighth International Congress for the Study of Pre-Columbian Cultures of the Lesser Antilles (S.
Lewenstein, ed.), Arizona State University Anthropological Research Paper, 22, 623 pp.

— 1 994. Beads and pendants from Trants, Montserrat: Implications for the prehistoric lapidary
industry of the Caribbean. Annals of Carnegie Museum, 63:215-237.

Wheeler, M. M. 1988. Montserrat, West Indies: A Chronological History. Montserrat National
Trust, Plymouth, 71 pp.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 4, Pp. 297-3 1 7

30 November 1994

ARCHAEOLOGY OF TRANTS, MONTSERRAT. PART 2.

VERTEBRATE FAUNA

Elizabeth J. Reitz1
Research Associate, Division of Anthropology

Abstract

Trants (MS-G1) is a Saladoid site located on the island of Montserrat in the Lesser Antilles.
Vertebrate remains provide evidence for the use of both terrestrial and marine resources. Fish were
obtained primarily from banks or reefs rather than inshore or offshore waters. The terrestrial resources
include animals introduced by humans and indigenous animals eventually driven to extinction as a
result of human activities. The ratio of marine to terrestrial resources is similar to that reported for
other Saladoid sites in the Lesser Antilles. Differences among Saladoid faunal collections from the
Lesser Antilles suggest that people living on each island knew well how to make use of local animals
and were not transient South Americans unfamiliar with the resources offered by Caribbean islands.

Introduction

In 1984 David W. Steadman, David R. Watters, Gregory K. Pregill, and Eliz-
abeth J. Reitz (Steadman et al., 1984 b) argued that analysis of archaeological
faunal remains from Montserrat was important for several reasons. First, it could
identify the indigenous fauna used by Saladoid colonizers during what appears
to be the first human occupation of the island. Second, faunal identification could
establish the contemporaneity of Caribbean peoples with species now extinct on
Montserrat. Third, vertebrate remains could be compared to those found in pre-
Columbian sites elsewhere in the West Indies, thereby providing information
useful on a regional scale.

These contributions are related to several broad Caribbean issues. One of the
most important of these is the character of the adaptation made by Saladoid
peoples when they left the South American mainland to colonize islands whose
vertebrate resources were unfamiliar. Within a broadly similar pattern, it appears
likely that Saladoid immigrants developed strategies appropriate to the exploi-
tation of those vertebrate resources found on each island colonized.

An important aspect of this adaptation is the role of terrestrial animals in
subsistence efforts of Saladoid peoples as they colonized the island chain. Elizabeth
Wing (1989) observed that remains of terrestrial mammals, pigeons, lizards, and
crabs are most abundant in either deposits from the Greater Antilles or early
deposits in the Lesser Antilles. Wing (1989) found that an average of 34% of the
individuals in Greater Antilles faunal assemblages were terrestrial animals while
38% of the individuals in early Lesser Antilles faunal collections were terrestrial
(Table 1). The two most common terrestrial animals were rice rats and hutias. In
contrast, 1 9% of the individuals in faunal assemblages from late deposits in the
Lesser Antilles and 1 7% of the individuals in deposits from the Bahamas were
terrestrial. Ranges for the four categories overlap; and variations in sample size

1 Museum of Natural History, Natural History Building, University of Georgia, Athens, Georgia

30602-1882.

Submitted 1 November 1993.

297

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Table \.— Terrestrial animals in the Lesser Antilles. Data from the Bahamas, Greater Antilles, and
Early and Late sites from the Lesser Antilles from Wing (1989). Trants data from this paper, excluding

Stratum A.

Average %MNI

Lesser Antilles

Early sites

38%

Late sites

19%

Trants

xk” samples

57%

Vs" samples

44%

and recovery techniques limited the conclusions that could be drawn from these
observations (Wing, 1989).

Extensive use of indigenous terrestrial animals at early sites is found in con-
junction with evidence that colonists introduced South American animals such
as opossum, guinea pig, agouti, and dog into the West Indies (Wing, 1989). This
use of indigenous terrestrial fauna and transportation of exotic animals throughout
the Caribbean system extended into the post-Columbian period. Its impact on
the zoogeography of the Caribbean caused the extinction of many indigenous
forms (Olson, 1978, 1982; Pregill and Olson, 1981; Olson and Pregill, 1982;
Steadman et ah, 1984a; Morgan and Woods, 1986; Woods, 1989, 1990). Extinc-
tions were also probably an important factor in early human colonization of the
Lesser Antilles (Keegan and Diamond, 1987).

This issue will be addressed using data from a Saladoid occupation at the Trants
site (MS-G1) on Montserrat. The Trants vertebrate collection is relevant because
the sample size is large and the recovery technique used during excavation was
good. The faunal data provide evidence for the use of indigenous and exotic
terrestrial animals at an early Ceramic Age site in the Lesser Antilles. The Trants
data indicate that Saladoid colonists made use of the resources found on Mont-
serrat in a way not suggested by faunal assemblages from elsewhere in the Lesser
Antilles, but nonetheless consistent with the general Caribbean pattern described
by Wing (1989).

Materials and Methods

Trants is a large, pre-Columbian site located on the eastern, or windward, side of Montserrat, one
of the northern Lesser Antilles (Fig. 1). Montserrat is a small island of volcanic origin and Trants is
located on one of the largest sections of relatively flat land found on the island (Steadman et al.,
19846). Sandy beaches and shallow inshore waters are limited around Montserrat. The east coast of
the island is characterized by rugged cliffs. These are found north of Trants Bay as well as south of
Farm Bay (Fig. 2). The beach at Trants Bay, just north of the site, is composed of cobble but Farm
Bay has a sandy beach. The site is roughly 400 m inland from a rocky portion of the coast between
the two bays, about 300 m from Trants Bay and 600 m from Farm Bay (Fig. 2). A small water course,
Farm River Ghaut, runs south of the site. Today the stream is interrupted by a dam, but in the past
it probably was a permanent water course. The mouth of Farm River Ghaut may have formed one
of the few estuaries on the island, and a mangrove swamp may also have been present in Farm Bay
(Newsom, 1994). The waters of Montserrat contain more patch reefs than fringe reefs and they are
mainly found on the north, west, and south of the island rather than near Trants. However, there are
some small patch reefs and seagrass beds in the limited inshore area adjacent to Trants.

The specimens reported here were excavated from the Trants site (MS-G1) in 1979 by Watters and
in 1990 by Watters and James B. Petersen (Petersen and Watters, 1991; Watters, 1994). In the
discussion that follows, reference will be made to materials recovered from a 2 x 2 m excavation

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Reitz— Trants Archaeology, 2

299

Greater Antilles

Fig. 1.— The Trants prehistoric site is located on the east coast of Montserrat in the northern Lesser
Antilles.

unit (Test Pit 1) dug in the core area in 1979 (Steadman et al., 19846). The 1979 excavation was in
1 0 cm increments and sediments were dry-screened through Vs" mesh. These increments were combined
into three analytical strata: Strata 1(D), I, and II. Strata I and II were interpreted as Saladoid occupations.
Vertebrates from Stratum 1(D), the uppermost level, were not included in the calculations because of
historic period disturbance.

The 1990 vertebrate remains are from three 1 x 1 m excavation units: N396E571, N421/22E645,
and N596E571, all within the core area (Fig. 2). Each of the 1990 excavation units was dug in four
quadrants. Faunal remains in three quadrants were collected in Va" mesh screens and in the fourth
quadrant using Vs" mesh screen. Some 1 mm window-screened samples were also collected, but time
did not permit their study. In order to explore the impact of recovery techniques, vertebrate materials
from the Va" quadrants are compared to those from the Vs" quadrants. These are referred to as Va"
samples and Vs" samples.

The 10 cm-levels of the 1990 excavation units were combined into three analytical strata, two of
which are associated with the Saladoid occupation. Stratum A is the historic hoe zone, Stratum B was
deposited around A.D. 60-200, and Strata C and D were deposited around 480-10 B.C. (Petersen
and Watters, 1991). Data from Stratum A are excluded from this analysis because historic and
prehistoric artifacts were mixed in Stratum A. An Old World rat (Rattus sp.) was found in Feature 3,
Unit 421/22E645 [Provenience Number (PN) 2027] which underlies Stratum A. Since Rattus clearly
is a post-Columbian animal, it was necessary to exclude Feature 3 and all deposits above it to insure
that only pre-Columbian vertebrate remains were studied. The Stratum D assemblage was so small
that it was combined with samples from Stratum C. Although each of the strata from the three units
may be from different time periods within the Saladoid occupation, or represent distinct activity areas
within a large settlement, their temporal or social relationships have not been clarified. Data for each
of the strata and units are presented separately, but are combined in the discussion. Other details of
excavation are provided elsewhere (Petersen and Watters, 1991; Watters, 1994).

Faunal materials from the 1979 excavation were studied by several people. Birds and mammals
were examined by Steadman, reptiles by Pregill, and fish by Reitz. All vertebrate materials recovered
during the 1 990 excavation were studied by Reitz using the comparative skeletal collections at the
Florida Museum of Natural History and the University of Georgia Museum of Natural History. During
the 1990 study, a record was made of the Number of Identified Specimens (NISP), the portion of each
bone recovered, the bone’s symmetry, and an estimate of age at death. Modifications to the bones
(primarily burning) and weights were also recorded but are not presented here; these data are on file
at the Georgia Museum of Natural History and are available upon request.

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Fig. 2. —The Trants site with the excavation units noted using British West Indies grid (Watters, 1 994).
Unit A is the 1979 Test Pit 1, Unit B is N396E571, Unit C is N421/422E645, Unit D is N596E571.
Excavation units are not to scale.

In order to estimate the Minimum Number of Individuals (MNI), samples from the three excavation
units (N396E571, N421/22E645, and N596E571) were kept separate, as were samples from each of
the three strata, creating nine analytical units. Because only a few elements could be identified to
genus, a higher MNI estimate was sometimes obtained by family or tribe, than at the genus or species
level. For example, more individuals might be estimated when Oryzomyini, Oryzomyini A, and
Oryzomyini B were combined than when bones identified as Oryzomyini A or Oryzomyini B were
counted independently. When that was the case, the estimates of MNI for lower taxonomic levels are
included in the species lists in parentheses. Estimates included in parentheses are not included in the
total for each list or in subsequent calculations.

Relative age of oryzomyines was estimated based on the degree of epiphyseal fusion for diagnostic
elements. When animals are young their bones are not fully formed. Along the area of growth the
shaft and the end of the bone, the epiphysis, are not fused. When growth is complete the shaft and
epiphysis fuse. While environmental factors influence the actual age at which fusion is complete
(Watson, 1 978), elements fuse in a regular temporal sequence (e.g., Silver, 1 963; Schmid, 1972; Gilbert,
1980). In most cases, one end of the bone fuses before the other. Bones were recorded as complete,
proximal (p), or distal (d), and either fused (f) or unfused (unf). Hence a humerus recorded as “punfdf ’
would be unfused at the proximal end and fused at the distal end.

Age was estimated for oryzomyines based on toothwear. The criteria defined by Wing (19936) were
applied to those teeth still in the maxilla or mandible. Teeth in Stage 1 are unworn molars, stage 2 is

1994

Reitz— Trants Archaeology, 2

301

Table 2.— Bone count (NISP) and Minimum Number of Individuals (MNI) from N396E571, Trants,
Montserrat, by stratum (A, B or C/D) and recovery technique.

N396E571 lU" samples

N396E571 Vs"

samples

Stratum

C/D

Taxon

NISP

MNI

NISP

MNI

NISP MNI

NISP

MNI

NISP

MNI

NISP MNI

UID fish

274

Serranidae

Epinephelus sp.

Mycteroperca sp.

Carangidae

Caranx sp.

Selene sp.

Lutjanidae

Haemulidae

Sparidae

Bodianus rufus

Halichoeres sp.

Scaridae

Sparisoma sp.

Scombridae

Balistidae

UID turtle
Cheloniidae

UID lizard

Iguana sp.

Colubridae

UID bird

Columbidae

Passeriformes

UID mammal

cf. Oryzomyini

Oryzomyini

Oryzomyini A

(3)

(1)

Oryzomyini B

(1)

(2)

Dasyprocta aguti
Canis familiaris

Totals

135

500

characterized by slight wear on the tips of the cusps, stage 3 teeth are substantially worn, and teeth
classified as Stage 4 were worn to the point that the tooth surface was flat and the dentine entirely
exposed. Animals with toothwear classified as Stages 1 or 2 are considered juveniles and those with
toothwear in Stages 3 or 4 are referred to as adults. As with fusion, environmental variables, especially
type of forage, are known to influence the rate of wear (Grant, 1978).

Measurements of bones were taken where preservation allowed. In the case of birds and mammals,
these measurements followed the guidelines established by Driesch (1976). Additionally, the anterior
width of the centrum of the fish atlas and the alveolar length (AL) of oryzomyine mandibles and
maxilla were recorded. Ideally only completely fused bones would be measured; however, oryzomyines
typically do not live long enough for many bones to fuse. Consequently, greatest length (GL) was taken
of unfused but otherwise complete humerus, femur, tibia, and calcaneus diaphyses. These exceptions
are noted with the measurements. These measurements represent animals that had not attained full
maturity and represent the size/age of individuals in that portion of the oryzomyine population
exploited by people at Trants. This procedure permits comparison of the Trants oryzomyine data with
those reported by Wing (19936).

The species identified are summarized by four faunal categories based on vertebrate class and habitat
preferences: Terrestrial, Inshore, Reef, and Pelagic. All mammals, birds, and reptiles were classified

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Table 3. — Bone count (NISP) and Minimum Number of Individuals (MNI) from N421/22E645, Trants,
Montserrat, by stratum (A, B, or C/D) and recovery technique.

Taxon

N421/22E645 W samples

N421/22E645 Vs" samples

Stratum

C/D

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

UID fish

146

189

535

272

Tylosaurus sp.

Serranidae

Epinephelus sp.

Mycteroperca sp.

Carangidae

Trachinotus goodei

Lutjanidae

Lutjanus sp.

Haemulidae

Anisotremus sp.

Haemulon sp.

Labridae

Bodianus rufus

Halichoeres sp.

Scaridae

Scarus sp.

Sparisoma sp.

Balistidae

UID reptile

UID turtle

Cheloniidae

UID lizard

Iguana sp.

Teiidae

Ameiva sp.

Colubridae

UID bird

Columbidae

Passeriformes

UID mammal

UID large mammal

cf. Oryzomyini

112

Oryzomyini

Oryzomyini A

(1)

(2)

Oryzomyini B

(2)

(1)

Rattus sp.

Dasyprocta aguti

Canis familiaris

Totals

378

425

913

442

as Terrestrial, including ducks, rails, and sea turtles. These animals are closely associated with inshore
waters and their capture might have been from either land or sea. In placing them in the Terrestrial
category preference in capture technique is given to capture of nesting animals. Inshore taxa include
sharks, palometa, porgies, drums, and gobies. The only pelagic fishes were mackerels. All other fishes
were classified as reef inhabitants. Some, if not all, of these animals might be found in other habitats,
either occasionally or because one or two members of the family occupy another habitat routinely. In
these cases, classification reflects the location most members of each family commonly frequent and
hence the most likely habitat in which they would have been captured, although this is not the only
habitat that might have been exploited.

1994

Reitz— Trants Archaeology, 2

303

Table A. —Bone count (NISP) and Minimum Number of Individuals (MNI) from N596E571, Trants,
Montserrat, by stratum (A, B or C/D) and recovery technique.

N596E571 'A" samples

N596E571 W samples

Stratum

C/D

Taxon

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

NISP

MNI

Ginglymostoma

cirratum

Carcharhinidae

UID fish

315

251

802

Serranidae

Diplectrum sp.

Epinephelus sp.

Mycteroperca sp.

Carangidae

Caranx sp.

Trachinotus goodei

Lutjanidae

Lutjanus sp.

Ocyurus chrysurus

Haem ul idae

Anisotremus sp.

Conodon nobilis

Haemulon sp.

Sciaenidae

Bodianus rufus

Halichoeres sp.

Sparisoma sp.

Gobiidae

Scombridae

Balistidae

UID turtle

Cheloniidae

UID lizard

Iguana sp.

Teiidae

Ameiva sp.

(1)

Colubridae

UID bird

Anatidae

Rallidae

Columbidae

Passeriformes

UID mammal

cf. Oryzomyini

Oryzomyini

Oryzomyini A

(1)

Oryzomyini B

Dasyprocta aguti

Totals

118

575

315

1045

Results

Vertebrate remains in the samples from the three Trants excavation units in-
dicate that both terrestrial and marine resources were used by residents (Tables
2-4). In general, similar resources were present in all three units. The highest
percentages were from the Terrestrial and Reef categories. Animals from inshore

304

Annals of Carnegie Museum

vol. 63

waters were very rare and offshore conditions were represented by a single indi-
vidual.

In each unit, Terrestrial vertebrates were more common in Stratum C/D than
in Stratum B. The frequency of Terrestrial vertebrates ranged from 50% to 71%
in Stratum C/D and from 39% to 57% in Stratum B. Terrestrial individuals were
least common in Unit N396E57 1 , Stratum B (Table 2) with 39% of the individuals;
12% of the individuals were rice rats and 9% were pigeons (Columbidae) and
passerine birds (Passeriformes). Terrestrial individuals were most common in
Unit N421/22E645, Stratum C/D (Table 3), representing 71% of the individuals;
13% of the individuals were rice rats (Oryzomyini) and 36% were pigeons and
passerine birds. Oryzomyines occurred in all strata of all units, with the exception
of Stratum C/D in Unit N596E571 (Table 4), which contained no mammals at
all. Although terrestrial resources contributed 50% of the individuals in the sample
from that stratum, these were exclusively lizards, a snake, pigeons, and passerine
birds.

The samples from the quadrants recovered using Va" and Vs" mesh were similar,
at least in terms of MNI (Tables 5, 6). The Va" component contained 32 identified
taxa, and the V s" component contained 39 taxa, although the latter component
contained almost twice as many bones (Table 5). Terrestrial habitats contributed
57% of the individuals in the Va" fraction and 44% in the Vs" samples (Table 6).
Oryzomyine rodents were present in both components in roughly equal numbers,
as were reptiles. Pigeons and passerines were more common in the Va" samples,
whereas fish individuals were more common in the Vs" samples. The difference
between Va" and Vs" recovery techniques is more clear when the measurements
are examined (Table 7). Although grouper individuals of the genus Epinephelus
were more common than those in the genus Mycteroperca in the Va" samples, the
reverse was true in the Vs" samples (Table 5). Significantly, Mycteroperca indi-
viduals are much smaller than the Epinephelus individuals (Table 7), which is
why they were recovered primarily with the smaller-meshed screen. These two
distinct sizes suggest that distinct fishing strategies were used in the capture of
these two grouper genera, an observation that would not have been known if only
the Va" samples had been studied. Since the materials recovered in the Vs" mesh
appear to be more representative of subsistence strategies at Trants, the following
comments will be based on percentages from the Vs" component only.

Fifty-six percent of the Trants individuals in the Vs" component are sharks and
bony fishes (Table 6). Most of the identifications could not be made to the specific
level, hence the exact nature of the habitats from which these fish were taken is
uncertain. Some species, such as the nurse shark ( Ginglymostoma cirratum) are
very common inshore, especially over rocky reefs and sand flats (Randall, 1968:
9). The palometa ( Trachinotus goodei ), porgy (Sparidae), drum (Sciaenidae), and
goby (Gobiidae) are also generally inshore fishes (Randall, 1968:114, 141, 149,
247). Other fishes are generally associated with coral reefs or rocky bottoms
(Randall, 1968:57, 102, 121, 128, 199, 217). These include groupers (Serranidae,
Epinephelus sp., Mycteroperca sp., jacks (Carangidae, Caranx sp., Selene sp.),
snappers (Lutjanidae, Lutjanus sp., Ocyurus chrysurus), grunts (Haemulidae, An-
isotremus sp., Haemulon sp.), wrasses (Labridae, Bodianus rufus, Halichoeres
sp.), and parrotfishes (Scaridae, Sparisoma sp.). Groupers are the most abundant
fish family in the Trants collection, constituting 28% of the individuals in the Vs"
component, and indicating that rocky outcrops were commonly fished. In contrast,
wrasses and parrotfishes, characteristic of reefs, constituted only 4% of the indi-

1994

Reitz— Trants Archaeology, 2

305

victuals in the Vs" samples. The only evidence that offshore waters were exploited
is a single mackerel (Scombridae), which might have been taken as it swam over
a bank or reef. This pelagic species contributed 1% of the individuals in the V*"
fraction.

All reptiles were classified as Terrestrial vertebrates. Sea turtles (Cheloniidae)
are not abundant in the Trants collection, constituting only 2% of the individuals
in the V&" component. The individuals appear to be adults. Their scarcity probably
reflects the rarity of sandy beaches for nesting on Montserrat. The east coast of
the island is primarily composed of cobbles; however, Farm Bay is one of the
east coast’s few sandy beaches (Fig. 2) and one of the island’s few seagrass beds
(Eastern Caribbean Natural Area Management Program, 1980). While the clas-
sification of these turtles as Terrestrial suggests they were taken while nesting, it
is also possible they were taken from the seagrass bed. The other reptiles were
more clearly Terrestrial. These included iguanas (Iguana cf. iguana ) and ameiva
lizards (Ameiva cf. pluvianotata). Although not common on the island today,
iguanas contribute 6% of the individuals in the V\ s" component. Colubrid snakes
(Colubridae) were almost as common (4% of the MNI).

Among birds, ducks and rails might be considered Inshore rather than Terres-
trial resources since they could have been taken from a bay or the Farm River
Ghaut. The two largest groups of birds, however, were clearly Terrestrial. These
included pigeons (Columbidae) and small passerine birds (Passeriformes). Pigeons
contributed 8% of the individuals in the V&" component and native pigeons still
survive on the island in spite of recent habitat destruction. The passerine order,
which includes thrashers and finches, constituted 9% of the Trants individuals.

Eleven of the rodents in the pre-Columbian assemblage were rice rats (Ory-
zomyini), and an additional four were probably Oryzomyini (Table 5), Rice rats
constitute 9% of the individuals in the samples recovered using *// mesh (Table
6). Two species of rice rats were present on the island. Remains were found in
all three excavation units and in most of the strata, but they were most common
in Stratum A, particularly in Unit N421/22E645 (Table 3), suggesting that both
large and small rice rats may have survived into the post-Columbian period. They
were probably driven to extinction by humans in combination with cats, dogs,
and Old World rodents introduced in the recent past.

Although most of the specimens could only be identified as unidentified (UID)
rodent or oryzomyine, the measurements (Table 7) suggest that there were at least
two species (Fig, 3). Steadman found two species of oryzomyines in his study of
the 1979 Trants vertebrate remains (Steadman et at., 1 984/A He classified these
as a small oryzomyine, Undescribed Species A, and a large oryzomyine, Unde-
scribed Species B, and this distinction is followed here. At least four of the Ory-
zomyini individuals were the small rice rat, (Oryzomyini A), and at least five of
the individuals were the large rice rat (Oryzomyini B) (Table 5). The size differ-
ences probably do not reflect simply younger versus older individuals since the
alveolar lengths (Table 7) support the presence of two distinct sizes of rice rats
rather than differences in maturation. The average alveolar length of the small
oryzomyine mandible is 7.08 mm (n = 3) and the average alveolar length of the
large oryzomyine mandible is 10.09 mm in = 4) (Table 7). Allometric correlations
between the width of the femur head and body weight (Wing and Brown, 1979:
127-129) predict an average weight of 181 g for the smaller West Indian rice rats
and up to 300 g for the larger ones (Wing, 1993a). Degree of fusion (Table 7) and
tooth wear (Table 8) indicate that most of the rice rats In Strata B and C/D were

306

Annals of Carnegie Museum

vol. 63

Table 5. —Bone count (NISP) and Minimum Number of Individuals (MNI) for VP samples and Vs"
samples with Stratum A excluded and Strata B and C/D combined, Trants, Montserrat. All three units

are combined.

xk" Samples

Vi" Samples

Taxon

NISP

MNI

NISP

MNI

Sharks

Ginglymostoma cirratum

Nurse shark

Carcharhinidae

Requiem sharks

Fishes

UID fish

617

1477

Serranidae

Sea basses

Epinephelus sp.

Grouper

Mycteroperca sp.

Grouper

Carangidae

Jacks

Caranx sp.

Jack

Selene sp.

Lookdown

Trachinotus goodei

Palometa

Lutjanidae

Snappers

Lutjanus sp.

Snapper

Ocyurus chrysurus

Yellowtail snapper
Haemulidae

Grunts

Anisotremus sp.

Margate

Haemulon sp.

Grunt

Sparidae

Porgies

Sciaenidae

Drums

Labridae

Wrasses

Bodianus rufus

Spanish hogfish

Halichoeres sp.

Wrasse

Scaridae

Parrotfishes

Sparisoma sp.

Parrotfish

Gobiidae

Gobies

Scombridae

Mackerels

Balistidae

Leatherjackets

1994

Reitz ' —Thants Archaeology, 2

307

Table 5.— Continued.

■A" Samples

Vs" Samples

Taxon

NISP

MNI

NISP

MNI

Reptiles

UID turtle

Cheloniidae

Sea turtles

UID lizard

Iguana sp.

122

Iguana

Teiidae

Whiptails

Ameiva sp.

(2)

Araeiva lizard

Colubridae

Non-poisonous snakes

Birds

UID bird

Anatldae

Ducks

Rallidae

Rails

Columbidae

Pigeons and doves
Passeriformes

Song birds

Mammals

UID mammal

UID large mammal

UID rodent

Oryzomyini

Rice rat

Oryzomyini A

(2)

Small rice rat

Oryzomyini B

(2)

(3)

Large rice rat

Cams familiaris

Dog

UID vertebrate

Totals

1188

2154

Table 6.=- Summary table of vertebrate fauna from Trants, Montserrat.

•A" Samples

Vs" Samples

MNI

%MNI

MNI

%MNI

Sharks/fishes

42.9

55.6

Reptiles

17.9

15.6

Pigeons/passerines

26.2

16.7

Other birds

2.4

1.1

Oryzomyine rodents

8.3

8.9

Dog

2.4

2.2

Totals

308

Annals of Carnegie Museum

vol. 63

Table 1 .—Measurements of vertebrate bones from Trants, Montserrat, in mm. Measurement dimensions
follow Driesh (1976). PN refers to provenience number and stratum to level.

Taxon

Element

Dimension

Measure-

ment

Fusion

Stratum

Ginglymostoma cirratum

vertebra

width

9.80

2214

Tylosaurus sp.

atlas

width

4.50

2020

Epinephelus sp.

atlas

width

2.35

2033

Epinephelus sp.

atlas

width

2.80

1913

Epinephelus sp.

atlas

width

2.80

1926

Epinephelus sp.

atlas

width

3.80

2024

Epinephelus sp.

atlas

width

5.96

2019

Epinephelus sp.

atlas

width

6.73

1959

Epinephelus sp.

atlas

width

6.81

1939

Epinephelus sp.

atlas

width

7.05

2050

Epinephelus sp.

atlas

width

7.29

2215

Mycteroperca sp.

atlas

width

1.70

2220

Mycteroperca sp.

atlas

width

1.80

2209

Mycteroperca sp.

atlas

width

1.90

1926

Mycteroperca sp.

atlas

width

1.90

1926

Mycteroperca sp.

atlas

width

1.95

2100

Mycteroperca sp.

atlas

width

2.00

1913

Mycteroperca sp.

atlas

width

2.02

2032

Mycteroperca sp.

atlas

width

2.10

1964

Mycteroperca sp.

atlas

width

2.10

1980

Mycteroperca sp.

atlas

width

2.10

1980

Mycteroperca sp.

atlas

width

2.10

1980

Mycteroperca sp.

atlas

width

2.10

2020

Mycteroperca sp.

atlas

width

2.10

2020

Mycteroperca sp.

atlas

width

2.20

2020

Mycteroperca sp.

atlas

width

2.20

2027

Mycteroperca sp.

atlas

width

2.30

1921

Mycteroperca sp.

atlas

width

2.30

2024

Mycteroperca sp.

atlas

width

2.40

1937

Mycteroperca sp.

atlas

width

2.40

2209

Mycteroperca sp.

atlas

width

2.50

1926

Mycteroperca sp.

atlas

width

2.56

1934

Mycteroperca sp.

atlas

width

2.60

1980

Mycteroperca sp.

atlas

width

2.60

1980

Mycteroperca sp.

atlas

width

2.67

2206

Mycteroperca sp.

atlas

width

2.90

2020

Mycteroperca sp.

atlas

width

3.20

2027

Carangidae

atlas

width

2.80

1909

Caranx sp.

atlas

width

2.00

1947

Selene sp.

atlas

width

3.60

1917

Trachinotus goodei

atlas

width

2.20

2209

Trachinotus goodei

atlas

width

3.40

2209

Lutjanus sp.

atlas

width

5.69

1928

Ocyurus chrysurus

atlas

width

1.40

2220

Sparisoma sp.

atlas

width

5.41

1966

Gobiidae

atlas

width

1.12

2209

Anatidae

carpometacarpus

11.80

1984

Anatidae

carpometacarpus

Did

5.80

1984

Anatidae

carpometacarpus

52.20

1984

Rallidae

humerus

7.30

1993

Rallidae

humerus

10.00

1993

Rallidae

humerus

46.50

1993

Rallidae

tibiotarsus

5.30

1993

Rallidae

tibiotarsus

5.60

1993

Columbidae

carpometacarpus

7.00

1984

Columbidae

carpometacarpus

7.20

1980

1994

Reitz— Trants Archaeology, 2

309

Table 7 Continued.

Taxon

Element

Dimension

Measure-

ment

Fusion

Stratum

Columbidae

carpometacarpus

7.40

2072

Columbidae

carpometacarpus

7.90

1974

Columbidae

carpometacarpus

8.00

2055

Columbidae

carpometacarpus

8.20

1976

Columbidae

carpometacarpus

9.90

2072

Columbidae

carpometacarpus

Did

4.20

2020

Columbidae

carpometacarpus

Did

4.30

1984

Columbidae

carpometacarpus

Did

4.80

1974

Columbidae

carpometacarpus

Did

4.80

1980

Columbidae

carpometacarpus

Did

5.20

2072

Columbidae

carpometacarpus

Did

5.40

2029

Columbidae

carpometacarpus

Did

6.90

2066

Columbidae

carpometacarpus

24.60

1984

Columbidae

carpometacarpus

24.90

1980

Columbidae

carpometacarpus

26.50

1974

Columbidae

coracoid

27.00

1972

Columbidae

coracoid

31.40

1971

Columbidae

coracoid

31.50

2066

Columbidae

coracoid

32.70

2042

Columbidae

coracoid

34.90

2072

Columbidae

coracoid

35.80

2066

Columbidae

coracoid

36.80

2072

Columbidae

femur

7.20

2064

Columbidae

femur

7.30

2072

Columbidae

femur

7.80

2041

Columbidae

femur

7.10

1984

Columbidae

femur

7.20

2223

Columbidae

femur

7.40

2020

Columbidae

femur

7.50

2072

Columbidae

femur

8.50

1926

Columbidae

femur

5.50

2064

Columbidae

femur

6.00

2072

Columbidae

femur

6.50

2041

Columbidae

femur

4.10

1984

Columbidae

femur

4.40

2020

Columbidae

femur

4.70

2072

Columbidae

humerus

7.90

1941

Columbidae

humerus

7.90

1972

Columbidae

humerus

8.10

2022

Columbidae

humerus

8.50

2066

Columbidae

humerus

9.20

1984

Columbidae

humerus

9.20

2203

Columbidae

humerus

9.50

1961

Columbidae

humerus

9.50

1972

Columbidae

humerus

9.50

2024

Columbidae

humerus

12.50

2085

Columbidae

humerus

13.10

2041

Columbidae

humerus

Dip

13.10

1972

Columbidae

humerus

39.50

1972

Columbidae

radius

5.00

2066

Columbidae

radius

5.50

2072

Columbidae

radius

53.90

2066

Columbidae

scapula

Die

6.40

1980

Columbidae

scapula

Die

6.90

1980

Columbidae

scapula

Die

7.00

1980

Columbidae

scapula

Die

7.40

1980

Columbidae

scapula

Die

7.50

1980

310

Annals of Carnegie Museum

vol. 63

Table 1 .—Continued.

Taxon

Element

Dimension

Measure-

ment

Fusion

Stratum

Columbidae

scapula

Die

7.70

2203

Columbidae

scapula

Die

7.80

2220

Columbidae

scapula

Die

7.90

2201

Columbidae

scapula

Die

8.10

2042

Columbidae

scapula

Die

9.50

2042

Columbidae

scapula

Die

9.70

2072

Columbidae

tarsometatarsus

6.80

2061

Columbidae

tarsometatarsus

7.00

2231

Columbidae

tarsometatarsus

7.40

2041

Columbidae

tarsometatarsus

6.50

2061

Columbidae

tarsometatarsus

33.50

2061

Columbidae

tibiotarsus

5.10

1971

Columbidae

tibiotarsus

5.70

2020

Columbidae

tibiotarsus

5.70

2024

Columbidae

tibiotarsus

5.80

2017

Columbidae

tibiotarsus

6.80

2066

Columbidae

tibiotarsus

7.40

2066

Columbidae

tibiotarsus

4.80

1971

Columbidae

tibiotarsus

5.50

2017

Columbidae

tibiotarsus

5.50

2024

Columbidae

tibiotarsus

6.00

2020

Columbidae

tibiotarsus

6.50

2066

Columbidae

tibiotarsus

6.70

2066

Columbidae

tibiotarsus

Dip

8.90

2072

Columbidae

tibiotarsus

Dip

9.30

2041

Columbidae

ulna

5.50

2042

Columbidae

ulna

7.20

2041

Columbidae

ulna

Did

5.10

2029

Columbidae

ulna

Did

5.30

2021

Columbidae

ulna

Did

6.40

2102

Columbidae

ulna

Did

7.20

2072

Columbidae

ulna

Dip

7.20

2042

Columbidae

ulna

Dip

9.80

2041

Columbidae

ulna

57.00

2041

Oryzomyini

femur

4.10

pfdunf

2201

Oryzomyini

femur

4.50

1926

Oryzomyini

femur

4.60

1917

Oryzomyini

femur

4.90

pfdunf

2114

Oryzomyini

femur

34.80

pfdunf

2201

Oryzomyini

femur

44.20

pfdunf

2114

Oryzomyini

humerus

28.80

punfdf

2072

Oryzomyini

humerus

29.40

punfdf

2020

Oryzomyini

humerus

31.20

punfdf

2020

Oryzomyini

humerus

31.70

punfdf

2020

Oryzomyini

mandible

8.54

1974

Oryzomyini

mandible

9.27

2042

Oryzomyini

mandible

9.55

2041

Oryzomyini

mandible

9.60

2042

Oryzomyini

mandible

9.70

2041

Oryzomyini

mandible

10.87

2051

Oryzomyini

maxilla

8.78

2052

Oryzomyini

maxilla

9.29

2053

Oryzomyini

maxilla

9.53

2041

Oryzomyini

maxilla

9.53

2041

Oryzomyini

maxilla

9.53

2041

Oryzomyini

maxilla

9.70

2029

1994

Reitz— Tkants Archaeology, 2

311

Table 1 .—Continued.

Taxon

Element

Dimension

Measure-

ment

Fusion

Stratum

Oryzomyini

maxilla

9.80

2027

Oryzomyini

maxilla

9.90

2020

Oryzomyini

maxilla

9.92

2041

Oryzomyini A

astragalus

4.00

1917

Oryzomyini A

astragalus

4.20

2027

Oryzomyini A

astragalus

4.50

1913

Oryzomyini A

astragalus

4.80

1909

Oryzomyini A

astragalus

4.80

1909

Oryzomyini A

astragalus

5.81

2032

Oryzomyini A

calcaneus

6.37

punf

2029

Oryzomyini A

femur

3.20

2066

Oryzomyini A

humerus

21.70

punfdf

2066

Oryzomyini A

mandible

7.00

2022

Oryzomyini A

mandible

7.20

2209

Oryzomyini A

mandible

7.50

1984

Oryzomyini A

maxilla

6.75

2025

Oryzomyini B

astragalus

6.30

1909

Oryzomyini B

astragalus

6.59

1926

Oryzomyini B

astragalus

6.70

2018

Oryzomyini B

calcaneus

8.03

1980

Oryzomyini B

calcaneus

11.49

2029

Oryzomyini B

femur

5.30

pfdunf

2066

Oryzomyini B

femur

5.40

pfdunf

2087

Oryzomyini B

femur

46.60

pfdunf

2066

Oryzomyini B

femur

46.70

pfdunf

2087

Oryzomyini B

mandible

9.77

2066

Oryzomyini B

mandible

10.10

2019

Oryzomyini B

mandible

10.10

2022

Oryzomyini B

mandible

10.40

2021

Oryzomyini B

tibia

45.10

punfdf

2066

Oryzomyini B

tibia

46.70

punfdf

2066

Dasyprocta aguti

humerus

11.90

2053

Dasypmcta aguti

tibia

9.90

2018

Cams familiaris

maxilla

12.62

2052

Cams familiaris

maxilla

13.90

2052

juveniles. Before extinction, both species were endemic on Montserret and else-
where in the Lesser Antilles. The presence of both rice rats on the same island,
however, is unusual. Although the habitats preferred by the rice rats are not known,
at least some members of this group tolerate disturbed, brushy, or cleared habitats
such as those around human habitations, from which they raid fields and stored
foods (Wolfe, 1982; Nowak and Paradise, 1983:572; Eisenberg, 1989:351).

None of the rice rat materials were found in combinations that suggested animals
that died in situ with little post-mortem disturbance. It has been demonstrated
in other contexts that rodents even smaller than these were consumed (Szuter,
1988, 1991). Element distributions are confused by the use of two different screen
sizes and identifiability. Over a third (36%) of the rice rat bones were cranial
fragments identified in the lkn meshed screen. Only 1 1 carpals or tarsals could be
identified as rice rats, and all were from the lk,f meshed fraction. The Vs" mesh
fraction itself, however, comprised only a quarter of the excavated area of each

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Fig. 3. —Astragali and mandible of oryzomyine A and oryzomyine B. The small astragalus and man-
dible are of oryzomyine A and the large elements are of oryzomyine B.

unit. No butchering marks were found on any of the Trants assemblage other
than burning; 3% of the rice rat bones had been burned. There is no reason to
assume these rice rats were not consumed, and the abundance of these animals
in this assemblage and the absence of intact skeletons seems solid evidence that
these fairly large rodents were eaten.

The land clearing and food storage associated with Saladoid peoples probably
enhanced the resource base for rice rats on Montserrat. Wing (19936) found that
rice rats were extremely abundant in the Hope Estate faunal assemblage associated
with a Saladoid occupation on the island of St. Martin. She anticipated that this
level of use over a long period of time would result in overexploitation of rice
rats. Based on data for a closely related rice rat ( Oryzomys palustris) (Negus et
al., 1961), she proposed that heavy human predation over time would result in
a relative decrease in West Indian rice rat populations and a corresponding de-
crease of rice rats in human deposits. This might also be associated with a relatively
heavy use of young animals as litter size increased in response to low population
density (Negus et al., 1961). Wing (19936) also predicted a decline in size as a
consequence of selective predation upon larger rice rats.

The Trants oryzomyine materials were examined for evidence of overuse as
proposed for Hope Estate. In the first place, rice rats constituted a much lower
percentage of the individuals in the Trants collection than in that from Hope
Estate. However, there was a decline in the percentage of oryzomyines from 1 1%
of the individuals in Stratum C/D to 9% of the individuals in Stratum B. This
decline in the usage of rice rats needs to be tested over a larger portion of the
Trants site. If it is supported by additional stratigraphic analysis, then this may
represent either a change in prey preference on the part of Saladoid residents or
a decline in the island’s rice rat population, or both.

In order to explore whether younger animals were exploited during more recent
time periods, tooth wear was compared among the three strata (Table 8). The

1994

Reitz— Trants Archaeology, 2

313

Table 8. — Toothwear observed for oryzomyines, Trants, Montserrat. Wear stages as defined by Wing

(1993b).

Taxon

Element

Wear

Stratum

Oryzomyini

mandible

Oryzomyini

mandible

Oryzomyini

mandible

2.5

Oryzomyini B

mandible

2.5

Oryzomyini A

mandible

Oryzomyini B

mandible

Oryzomyini

mandible

3.5

Oryzomyini B

mandible

Oryzomyini A

mandible

Oryzomyini A

mandible

Oryzomyini

mandible

Oryzomyini

mandible

Oryzomyini

mandible

Oryzomyini

mandible

Oryzomyini B

mandible

Oryzomyini

maxilla

Oryzomyini

maxilla

Oryzomyini A

maxilla

Oryzomyini

maxilla

Oryzomyini

maxilla

Oryzomyini

maxilla

Oryzomyini B

maxilla

Oryzomyini

maxilla

Oryzomyini

maxilla

Oryzomyini

maxilla

Oryzomyini

maxilla

degree of toothwear in Strata C and B suggests that all individuals were juveniles.
However, the oryzomyine mandibles and maxilla both indicate that those de-
posited in the lower strata were generally younger than those deposited in Stratum
A. This suggests that rice rat population density during the Saladoid occupation
was lower, perhaps due to greater predation compared to the post-Columbian
period. Unfortunately, there were not sufficient measurements in each of the three
strata to assess changes in body size through time. These data do suggest that rice
rats were subject to overpredation during the Saladoid occupation of Trants com-
pared to the post-Columbian occupation of the island, although data from contexts
with better temporal definition are needed to explore this possibility further. The
large number of older rice rats in Stratum A probably indicates that, at least during
part of the time represented by the hoe zone, these were natural deaths in an
unstressed population.

Terrestrial vertebrate resources also included two exotic forms introduced by
humans. One of these is the agouti (Dasyprocta aguti). Steadman identified an
agouti from Stratum 1(D) in the 1979 Trants fauna, a disturbed context. The
1990 sample contained 12 additional agouti bones, unfortunately also from dis-
turbed contexts (Stratum A). The agouti was introduced from the South American
mainland sometime in the pre-Columbian period. They are found in a number
of archaeological deposits throughout the Lesser Antilles and until recently still
lived on some of the islands (Nowak and Paradiso, 1983:816; Wing, 1989). They
are still found on Montserrat and the presence of agouti bones in disturbed contexts

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vol. 63

Table 9. —Comparison of resource use among several Saladoid vertebrate collections . Cay on data are
from Wing (1989); Pearls data are from Stokes (1993); Trants VV data are from this paper; and Trants
Vs" data are from this paper (Trants Vs ”) and from Test Pit 1 (Steadman et ai, 1984b).

Cayon

Pearls

Trants W'

Trants Vi"

Test Pit 1

MNI

%MNI

MNI

%MNI

MNI

%MNI

MNI

%MNI

MNI

%MNI

Terrestrial

58.6

32.8

57.1

44.4

69.6

Inshore

21.9

1.2

6.7

Reef

24.2

42.2

41.7

47.8

26.8

Pelagic

17.2

3.1

1.1

3.6

Totals

may mean that they burrowed into the site or were buried in it quite recently.
Therefore, it is not possible to confirm that these agouti remains are recent or
Saladoid.

The remains of two dogs were identified in the Vs" samples and two additional
dogs were identified in the 14 " samples (Tables 2, 3). A subadult was found in
Feature 2, Unit N396E571 (PN 1934, 1937) and the other three were individuals
represented by bones too fragmentary to estimate age (PN 1923, 1929, 1935,
1943, 2087). Dogs are not endemic to the West Indies and probably accompanied
humans during their migrations into the region. Since most dogs identified in the
Caribbean have been recovered from burials rather than from middens (Wing,
1989), it is possible that they were valued companions rather than sources of
food. However, the Trants dogs were not associated with burials nor were they
articulated burials themselves.

Discussion

The percentage of Terrestrial individuals puts the Trants assemblage above the
average for early sites in the Lesser Antilles as calculated by Wing (1989; Table
1). Although sharks and bony fishes were the most common vertebrate group,
birds and reptiles constituted a significant portion of the assemblage, with mam-
mals somewhat less abundant.

Data from Trants indicate that people living on Montserrat made use of in-
digenous terrestrial fauna, exotic mammals, and marine animals. In this respect,
Trants conforms to the general Caribbean pattern, in which most of the resources
are from the nearest habitats (Wing, 1989). In the case of Trants, it appears that
terrestrial and marine vertebrates were used in about equal numbers. Indigenous
terrestrial fauna included rice rats, pigeons, passerine birds, and iguanas. Since
this part of the Trants project focused on the vertebrate component, land crabs
and mollusks are not included in this calculation; however, these were also very
common in the collection (David Watters, personal communication, 1993). Some
of the terrestrial resources were exotic animals introduced to the island from
South America. As expected, marine resources in the Trants collection included
sea turtles and fishes from banks, reefs, and offshore areas, with emphasis on
those most accessible from the site.

Although these data support a general Saladoid pattern, they also indicate that
a variety of subsistence strategies were practiced in the Lesser Antilles by Saladoid
peoples, each one reflecting local conditions. Comparing the Trants materials to
those from early sites on Grenada (Pearls) and St. Kitts (Cayon) (Wing, 1989;
Stokes, 1993), the significance of adaptations to local conditions and the diversity

1994

Reitz— Trants Archaeology, 2

315

of subsistence strategies practiced in the Lesser Antilles by Saladoid peoples is
clear (Table 9). The only mammal identified in all three collections was the dog.
Both 1 rants and Pearls contained high percentages of oryzomyine rodents— 17%
of the individuals in the Pearls collection and 9% of the Trants lk’e collection. By
contrast, pigeons, so common in the Trants collection, were not identified in the
Pearls assemblage. Iguana were found in all three collections; sea turtles, rare in
the Trants collection, were not identified in the Cay on or Pearls collections.

Although the percentages of fish individuals from reefs and banks was similar
between Pearls and Trants, there were differences among the kinds of fishes iden-
tified. For example, 28% of the individuals in the Trants collection (the lk" sam-
ples) are groupers, more typical of deeper waters, in contrast to 12% of the in-
dividuals in the Pearls collection. Wrasses and parrotfishes, typical of reefs,
contributed 4% of the Trants lh" and 16% of the Pearls assemblages. Shallow
water, inshore species, rare in the Trants collection, contributed almost a quarter
of the Pearls Individuals.

These differences probably reflect the types of marine habitats associated with
each island. Shallow water areas are limited around Montserrat, which has more
patch reefs than fringe reefs compared to islands such as Grenada and Barbuda.
This leads to the conclusion that the subsistence strategy practiced at Trants is a
local adaptation. It also suggests that it is probably not appropriate to develop
models for the colonization of the Caribbean based on the assumption that Sa-
ladoid peoples everywhere made use of marine and terrestrial resources in the
same way (Watters and Rouse, 1989).

One characteristic all three assemblages share Is a high percentage of oryzomyine
Individuals, raising the question of whether rodents could support exploitation at
these levels for a long period of time. (The hutia was introduced to Montserrat.)
Consumption of rice rats increased between pre-horticultural and horticultural
strata and then declined in later ones (Wing, 1993a). It may be that rice rats were
consumed in limited amounts as a supplement to marine resources, iguanas, and
land crabs before gardening began on those islands which were occupied during
pre-horticultural periods. Use of rice rats and other garden pests may then have
increased in levels associated with horticulture. Human horticultural activities
could have contributed to an increase in rice rat population size because these
plots provided optimum habitat for them. Rice rats could have been both a
terrestrial mammalian meat source for Saladoid peoples as well as pests attracted
by gardens and stored foods. Saladoid colonists may not have intentionally sought
out rice rats, but rather taken advantage of a resource that essentially came to
them, much as the garden hunting model suggested by Linares (1976).

The declining representation of rice rats in later horticultural deposits might
indicate that the “vermin” problem either was under control or had declined in
the face of considerable pressure both from humans and their dogs. Ultimately,
habitat destruction and the introduction of predators/competitors such as cats,
mongoose, and Old World rats during the post-Columbian era drove rice rats to
extinction.

Such a pattern would imply a relationship between gardening, food storage,
predation, and the numbers of rice rats in archaeological deposits that has little
to do with preference for or prejudice against marine resources. It is important,
however, to emphasize that without a better pre-Saladoid, Saladoid, and post-
Saladoid stratigraphic sequence, change through time in the use of rice rats on
Montserrat or an association of their use with gardening cannot be proven.

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Annals of Carnegie Museum

vol. 63

Conclusion

An important aspect of Caribbean cultural history is the role played by terrestrial
animals in Saladoid subsistence efforts as they colonized the island chain. The
data from Trants indicate that people living there used both terrestrial and marine
resources. Use of indigenous terrestrial fauna and transportation of exotic animals
throughout the island system were important characteristics of colonization in
the Lesser Antilles. Perhaps some terrestrial resources could not support the
combination of long-term exploitation and predation by introduced carnivores
and eventually became extinct, leaving only their skeletal remains to remind us
of their existence.

On a regional scale, Trants provides additional evidence that extensive use of
terrestrial resources was a consistent Saladoid feature, but that there was much
variation among Saladoid occupations in the Lesser Antilles. Many different sub-
sistence strategies were practiced in the Lesser Antilles by Saladoid peoples, each
one reflecting local conditions. Future work in the Caribbean should attempt to
explore temporal variation and activity areas within Saladoid settlements such
as Trants.

Acknowledgments

I would like to thank David R. Watters and James B. Petersen for the opportunity to study the
Trants collection. I appreciate the assistance of Joel A. Dukes with the identifications and permission
from Elizabeth S. Wing to use the comparative collection at the Florida Museum of Natural History.
I am also grateful to Elizabeth Wing and Anne Stokes for access to their manuscripts while in un-
published form. Funding was provided by the Carnegie Museum of Natural History, the University
of Maine at Farmington, and the Charles Cohn Foundation. Earlier versions of this paper were
presented at the 58th Ann ual Meeting of the Society for American Archaeology in St. Louis, Missouri,
and at the 1 5th International Congress for Caribbean Archaeology, San Juan, Puerto Rico.

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ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 4, Pp. 319-325

30 November 1 994

A NEW SPECIES OF ANTOCHA (SUBGENUS ORIMARGULA)
FROM SULAWESI (DIPTERA: TIPULIDAE) AND
ITS MATE-CLASPING BEHAVIOR

Chen W. Young

Assistant Curator, Section of Invertebrate Zoology

Abstract

Antocha ( Orimargula ) possessiva n. sp., the first species in the subgenus Orimargula from Sulawesi,
and its unusual behavior involving precopulatory mate-clasping are described. Males clasp females
from below over the pleural region of the fifth abdominal segment, and then ride on them dorsally.
Expansion and folding of the pleural region of the fifth abdominal segment of females, and enlarged
basistyles of males, facilitate nongenitalic coupling, and are hypothesized as resulting from sexual
selection favoring precopulatory mate clasping.

Introduction

The genus Antocha Osten Sacken (1860) is a cosmopolitan genus of small-sized
crane flies within the tribe Limoniini (Tipulidae: Limoniinae). The genus is char-
acterized by a prominent anal angle on the wing. Three subgenera are recognized:
Antocha Osten Sacken (1860), Orimargula Mik (1 883), and Proantocha Alexander
(1919). The subgenus Orimargula can be separated morphologically from the
other two subgenera of Antocha by wing venation that lacks a discal cell (Mik,
1883). Antocha ( Orimargula ) is an Old World group, represented by 21 species
in the Ethiopian region, three in the Australasian region, three in the Palearctic
region, and 1 5 in the Oriental region. In Southeast Asia, species of this subgenus
have been recorded only from Borneo and Sumatra. In 1985 an undescribed
species of Antocha ( Orimargula ) was collected on Sulawesi, the first record of the
subgenus from that island. This new species and its unusual mate-clasping be-
havior are described in this paper.

Systematics

Antocha ( Orimargula ) possessiva Young, new species
(Fig. 1-5)

Diagnosis. — This Indomalayan species of Limoniini is placed in the subgenus
Orimargula because the wing venation lacks a discal cell (Fig. 1). It differs from
all other described species of Orimargula by features of the male genitalia, es-
pecially the elongated basistyles (Fig. 3), and the modification of the pleural region

of the fifth abdominal segment of females (Fig. 5).

Description. — Morphological terminology follows Byers (1961) and Young (1987). Description is
based on seven males and four females preserved in 80% ethanol. Body length: male, 4.0-4. 5 mm;
female, 4. 5-5.0 mm. Wing length: male, 4.0 mm; female, 4.5 mm.

Head: Occiput brown, with black setae. Rostrum pale yellow. Eyes large, separated at vertex by
about two-thirds width of eye in frontal view, narrowly separated ventrally by width of two rows of
ommatidia. Antenna 1 6 -segmented, filiform; scape cylindrical, two times as long as wide; pedicel

Submitted 8 April 1994.

319

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Annals of Carnegie Museum

vol. 63

• 5mm

Fig. 1-2 .—Antocha ( Orimargula ) possessiva. 1, wing; 2, head and thorax, right lateral view.

globular; flagellomeres subcylindrical, verticillate; basal flagellomeres distinctly elongated; apical fla-
gellomere 1.5 times as long as penultimate.

Thorax: Grayish brown to dark brown, with velvety-black patches. Cervical sclerites greatly elon-
gated, attached to thorax ventrally (Fig. 2). Pronotum extremely reduced, with setae near anterior
margin. Prescutum enlarged and projecting anteriorly, dark brown anteromedially, fading to grayish
brown at midlength; lateral border of prescutum with a large velvety-black patch. Scutum, scutellum,
and postnotum brown. Pleura grayish brown; anepistemum and pre-epistemum dark brown; latero-
tergite with large velvety-black patch. Legs yellowish brown; claws dark brown, each claw with single
tooth at basal third. Wings (Fig. 1) subhyaline, faintly tinged with grey; stigma indistinct; Rs long,
gently convex, slightly shorter than R3; R2 and R2+3 subequal. Halteres pale brown.

Abdomen: Pale brown. Pleural region modified on fifth segment of females (Fig. 5). Fifth tergum
with lateral area greatly expanded (especially anterolateral comers), which overlap pleural regions;
pleural membrane compressed, forming narrow thickened band, invaginated at basal third. Postero-
lateral comers of terga with 10-16 setae; posterolateral edge of fifth tergum below setae expanded
ventrolaterally into pleural region.

Male Genitalia: Hypopygium as in Fig. 3. Ninth tergum narrow, posterior margin straight. Basistyle
relatively long, slender; ratio of length of ninth tergum to length of basistyle is 0.3. Ninth sternum
simple, undivided, rounded at posterior apex. Dististyles elongate, fused basally for two-thirds of
length. Dorsal dististyle glabrous, subarcuate, acute apically. Ventral dististyle simple, setiferous,
deflected medially at right angle to basistyle; apex recurved anteromedially. Lateral process simple,

.1 mm

Fig. 3-4 .—Antocha ( Orimargula ) possessiva. 3, male hypopygium, dorsal view; 4, female ovipositor,
lateral view.

1994

Young— New Antocha from Sulawesi

321

■ 5 mm

Fig. 5-6.— Antocha ( Orimargula ) possess iva. 5, female abdominal segments III— VI, right lateral view;
6, female abdomen showing male basistyles and dististyle grasping lateral part of fifth abdominal

segment, ventrolateral view.

extending to near midlength of basistyle, apex slightly expanded. Aedeagal sheath well-developed,
curved ventrally. Aedeagal process slender, apically acute, reaching to near midlength of aedeagal
sheath.

Ovipositor (Fig. 4): Eighth tergum reduced. Ninth tergum much reduced, well-separated from tenth
tergum. Cerci slightly shorter than tenth tergum, curved dorsally; dorsal surface flat, broader at base,
narrowed to apex. Eighth sternum expanded laterally, wider than seventh sternum. Tenth sternum
with setae near posterior margin. Hypovalves broad basally, tapered distally, extending to one-third
length of cerci.

Type Material. —Holotype: male, Carnegie Museum of Natural History (CMNH).
Verbatim text of three pin labels: INDONESIA: Sulawesi Utara. Dumoga-Bone
N. P. 22 Aug 1985. Coll. Chen Young / Base Camp 21 1m. PROJECT WALLACE
/ HOLOTYPE Antocha ( Orimargula ) possessiva Young [red paper]. Paratypes
with same data as holotype: ten males, ten females. All specimens were collected
at light. Six pair of paratypes are mounted in clasping position as retained at
death. Two paratypes (one male, one female) are deposited in the collections of
the National Biological Institute, Bogor, Indonesia; other paratypes deposited in
CMNH.

Other Material. — Eighteen specimens (ten males, eight females) with same data
as holotype preserved in 80% ethanol and deposited in CMNH.

Etymology. — The name of this species is the Latin adjective possessivus (-a, -um), meaning to possess,
and refers to the clasping and holding of females by males.

Habitat. — Specimens of A. possessiva were collected at a rain forest site within
the Dumoga-Bone National Park on the northern peninsula of Sulawesi. The
National Park is located just north of the equator (0°38'N, 124°06'E). The site is
at the eastern end of the park, on the west bank of the Tumpah River, a tributary
of the Toraut River. The vegetation is primary lowland forest at approximately
200 m elevation. Specimens were collected at a white cloth sheet illuminated by
a mercury vapor lamp near the edge of a pond along the Tumpah River. The
pond was surrounded by rice fields with irrigation channels on three sides, and
by lowland forest on one side. The river, the pond, and the larger irrigation
channels are suitable and likely habitats for the immature stages of species of
Antocha.

Remarks. — The general appearance of A. possessiva is similar to that of A.
praescutalis Alexander from Moeara Tenam, southern Sumatra, and to A. ma-

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.1mm

Fig. 1 .—Antocha ( Orimargula ) possessiva. Nongenital coupling of the adult, right lateral view.

culipleura Edwards from Mount Kinabalu, northern Borneo. All three species
have similar velvety-black spots on the thoracic pleura. Antocha possessiva differs
from A. maculipleura mainly by its smaller size and indistinct stigma. It differs
from A. praescutalis by the absence of a third velvety-black spot on the anepister-
num.

Antocha ( Orimargula ) possessiva is very distinct from all other species of the
subgenus. Two unique characters occur in male genitalia of this species. The

1994

Young— New Antocha from Sulawesi

323

basistyles are unusually long and the dorsal and ventral dististyles are almost
completely fused (Fig. 3). Both features enhance the structural integrity of the
male genitalia, and may increase the firmness with which the female abdomen is
clasped (Fig. 6) as described below.

Mate-Clasping Behavior
(Fig. 6-7)

Unusual mate-clasping behavior was observed in this species at an illuminated
cloth sheet on a clear night between 2000-2300 hr. Neither sex was observed to
congregate on the sheet, and 20 males and 1 8 females were collected on the sheet.
All flies arrived at the sheet separately and made initial contact with each other
while fluttering or walking about on its surface. When a male contacted a female,
he would mount her dorsally on the thorax between the wings, their bodies oriented
in the same direction (Fig. 7). The male would bend his abdomen ventrally
alongside the female’s abdomen, and then curl it dorsally so as to clasp the pleural
region of the fifth abdominal segment of the female from the ventral side (Fig.
6). This behavior was observed in nine pairs, with the position of the male’s
abdomen occurring (apparently randomly) on either side of the female.

After a male clasped a female, the male remained mounted on the female and
tenaciously retained possession of her through firm nongenital contact. Coupled
pairs were unable to fly, and if disturbed would drop to the ground or crawl across
the surface of the sheet. The male’s grasp was firm, and union with females
persisted even after a pair was killed in cyanide, or dropped into 80% ethanol.
Coupled pairs were easily mounted in conjoined position.

Male-male contact was also observed on the brightly-lit sheet. Such contact
would result in brief coupling attempts by both males, and would quickly end
with one or both males dislodging the other. In a few cases a solitary male contacted
a male that was already clasping a female. The rival male would attempt to dislodge
the guarding male and to gain access to the fifth abdominal segment of the female.
Rival males were never observed to attempt genitalic copulation with the female.

Actual copulation in this species was not observed, either on the sheet or on
the ground. Copulation was observed at the same time in other crane flies (species
of Nephrotoma and Trentepohlia). These species assumed an end-to-end position
in copula on the sheet.

Discussion

The riding of females by males has not been observed previously in the genus
Antocha, nor has it been reported in other species of crane flies. It is difficult to
assess the significance of this behavior because of the artificial conditions of light
and substrate at the time of observation. Several facts needed to interpret the
nature of this behavior remain unknown, including the time when copulation
occurs, the oviposition site, the influence of clasping on mating success of com-
petitive males, and details of the life history. However, the presence of a pleural
modification of the fifth abdominal segment of females to maintain the secure
grip of elongate male basistyles suggests mutually compatible evolutionary nov-
elties in both sexes that may have resulted from sexual selection. The persistent
clasp of the females by males possibly constitutes mate-guarding behavior. Such
guarding behavior can occur either before (precopulatory) or after (postcopulatory)
copulation and insemination, and both conditions have been reported for other
species of insects.

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Precopulatory mate-guarding has been described in the hippoboscid fly, Lynchia
hirsuta (Thornhill and Alcock, 1983). Females of this species lay eggs individually
over a considerable period of time. Males can identify females that are about to
become receptive to mating, and females mate more than once at intervals
throughout their lives. These factors favor selection of precopulatory female-riding
by males because such behavior increases the likelihood that the sperm of the
riding male will fertilize the eggs as they are produced and laid over time.

In A. possessiva and tipulid species in general, adult longevity is relatively short.
Females generally carry a full complement of mature eggs at emergence and mate
at or very soon after emergence (Pritchard, 1983); females usually lay eggs rapidly
over a relatively short period of time. These conditions decrease the time over
which reproductive advantage might be gained by males via precopulatory riding.
In addition, because females in two out of the six coupled pairs of A. possessiva
observed in this study were without eggs, it is assumed that males of this species
are unable to determine the gravid condition in females, or are indifferent to it.
Neither of these circumstances favor selection for male behavior involving pre-
copulatory riding.

Postcopulatory mate-guarding behavior was first reported in three species of
Limoniinae by Adler and Adler (1991). The three species involved are Antocha
saxicola Osten Sacken, Dactylolabis montana (Osten Sacken), and Limonia simu-
lans (Walker). Adler and Adler observed the males of these species guarding their
mates at oviposition sites after uncoupling. They concluded that all three species
were polygamous and sex ratios at oviposition sites were male-biased. Sperm
competition appears to be intense and to favor the last male to mate.

The sex ratio of A. possessiva at the sheet was not male-biased, although the
sex ratio at natural oviposition sites remains unknown. It is possible that the
observed sex ratios and behavior observed here are artifacts of unnatural en-
counters at the mercury lamp, and reflect behavior normally associated with
postcopulatory coupling at oviposition sites nearby. This would explain the te-
nacity of the coupling, and the apparent indifference of males to gravid and
nongravid females. However, abdominal clasping of females by males, and not
copulation, was observed between male and female flies during their initial en-
counters at the sheet. In addition, rival males appear to compete for possession
of the fifth abdominal segment of females, rather than for direct genitalic contact.
Neither of these factors supports the hypothesis that clasping behavior is postcopu-
latory. In fact, postcopulatory guarding of a female that has been previously mated
by another male would not increase reproductive fitness of the guarding male.
For the above reasons, in addition to those presented by Thornhill and Alcock
(1983) in Lynchia, the observed behavior of A. possessiva is interpreted as pre-
copulatory mate guarding.

Pritchard (1983) indicated that some Limoniinae display true mating swarms.
I observed a swarm of Antocha saxicola (Pennsylvania, Monroe County, Pocono
Lake Reservoir, 20 May 1985) and captured both males and females from it. It
is possible that A. possessiva also forms swarms in which males and females make
contact and form riding pairs. The absence of sexual dimorphism in the antennae
suggests that pheromones are probably not involved in this species (Pritchard,
1983). If mating occurs within such swarms in A. possessiva then coupled pairs
might drop from the mating swarm and crawl to a suitable oviposition site where
copulation would take place. Because riding by the male on the female prevents
the female from flying, the mating swarm would most likely occur very close to

1994

Young —New Antocha from Sulawesi

325

or directly over the habitat where oviposition occurs and where the immature
stages develop.

It is also possible that precopulatory contact between males and females is
mandatory in order to stimulate females in the right microhabitat to permit
copulation or to release oviposition. Aspects of female choice may be involved,
although variation in male reproductive success was not observed in this study.
Alternative interpretations are also possible with male-clasping inhibiting females
from mating, or with males being required at oviposition sites in order to me-
chanically assist with oviposition. Antocha is one of the very few genera of crane
flies in which the larvae have lost all functional spiracles and are truly aquatic in
habit (Byers, 1 978). The immature stages and life cycle of only on z Antocha species,
A. saxicola, has been studied (Alexander, 1919; Fuller and Hynes, 1987).

In summary, the female-riding behavior observed in this study is hypothesized
as precopulatory mate-guarding behavior. Because this unusual behavior was
observed under artificial conditions at a collecting light and sheet, the specific
nature of the behavior as presented above is speculative. Further observations on
life history are needed, especially of mating behavior and oviposition under natural
conditions. Of particular value will be observations that will determine the timing
and occurrence of copulation in relation to precopulatory clasping.

Acknowledgments

This paper is Project Wallace Paper No. 143, and is based on material collected by the author on
Project Wallace, sponsored by the Indonesian Institute of Sciences and the Royal Entomological Society
of London. The research was supported by a grant from the M. Graham Netting Field Research Fund,
The Carnegie Museum of Natural History, Pittsburgh. I would like to thank J. E. Rawlins for comments
and suggestions on the manuscript, R. L. Davidson for literature translation, M. A. Klingler for
preparing Fig. 5-7, Mary Ann Daman for manuscript preparation, and three anonymous reviewers
for their comments on the manuscript.

Literature Cited

Adler, P. H., and C. R. L. Adler. 1991. Mating behavior and the evolutionary significance of mate
guarding in three species of crane flies (Diptera: Tipulidae). Journal of Insect Behavior, 4(5):6 1 9—
632.

Alexander, C. P. 1919. The crane-flies of New York, part II. Biology and phylogeny. Cornell
University Agricultural Experiment Station Memoir, 38:691-1 133.

Byers, G. W. 1961. The crane fly genus Dolichopeza in North America. The University of Kansas
Science Bulletin, 42(6):665— 924.

. 1978. Tipulidae. Pp. 285-310, in An Introduction to the Aquatic Insects of North America

(R. W. Merritt and K. W. Cummins, eds.), Kendall/Hunt Publication Co., Dubuque, Iowa, xiii
+ 441 pp.

Fuller, R. L., and H. B. N. Hynes. 1987. The life cycle, food habits and production of Antocha
saxicola Osten Sacken (Diptera: Tipulidae). Aquatic Insects, 9:129-135.

Mik, J. 1883. Zur Kenntniss der ‘Limnobina anomala’ Osten Sacken. Wiener Entomologische Zei-
tung, 2:198-202.

Pritchard, G. 1983. Biology of Tipulidae. Annual Review of Entomology, 28:1-22.

Thornhill, R., and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard University
Press, Massachusetts, ix + 547 pp.

Young, C. W. 1987. A revision of the crane fly genus Dicranoptycha in North America. The
University of Kansas Science Bulletin, 53(5):2 15-274.

ANNALS OF CARNEGIE MUSEUM

Vol. 63, Number 4, Pp. 327-374

30 November 1 994

A REVISED CLASSIFICATION OF THE
SPIRIFERID BRACHIOPODS

John L. Carter

Curator, Section of Invertebrate Paleontology

J. G. Johnson1
Remy Gourvennec2

Hou Hong-fei3
Abstract

A new classification of the impunctate and punctate spiriferid brachiopods is presented. Where
morphological evidence permits, ancestor-descendant relationships are discussed and indicated.

New taxa include the suborder Cyrtinidina Carter and Johnson; new families are Ulbospiriferidae
Johnson and Carter, Lazutkinidae Johnson and Hou, Elymospiriferidae Johnson and Hou, Tenelloder-
midae Carter, Johnson, and Gourvennec, Perissothyrididae Carter, Palaeochoristitidae Carter, Ske-
lidorygmidae Carter, Rastelligeridae Carter, Dimegelasmidae Carter, Spiropunctiferidae Carter; new
subfamilies are Callispiriferinae Johnson, Eurekaspiriferinae Johnson, Branikiinae Johnson and Hou,
Ulbospiriferinae Johnson and Carter, Palaeospiriferinae Carter, Johnson, and Hou, Howellellinae
Johnson and Hou, Araspiriferinae Johnson, Eomartiniopsinae Carter, Elivellinae Carter, Sergospiri-
ferinae Carter, Tangshanellinae Carter, Reticulariopsinae Gourvennec, Rhenothyridinae Gourvennec,
Obesariinae Gourvennec, Eoreticulariinae Gourvennec, Quadrithyridinae Gourvennec, Martinothyri-
dinae Carter, Toryniferinae Carter, Spinolepismatininae Carter, Paralepismatininae Carter, Rastelli-
gerinae Carter, Dentospiriferinae Carter, Yalongiinae Carter, Pseudocyrtininae Carter, Dispiriferininae
Carter, Tethyspirinae Carter, Paralaballinae Carter, and Jiangdospiriferinae Carter.

Introduction

In 1 985 the organizers of the First International Congress on Brachiopods (Brest)
sponsored a revision of the out-of-date brachiopod Treatise volumes. We were
given the task of revising the spiriferids, impunctate and punctate. In the absence
of any recent comprehensive revision and classification for this large group we
submit the following preliminary classification of the spiriferid brachiopods for
the examination and criticism of our colleagues. We recognize that no endeavor
of this magnitude can be perfect or complete and beg the indulgence of our
colleagues for any egregious errors or omissions. Now is the time to bring errors
or omissions to our attention, before the Treatise goes to the press.

Since the first publication of the brachiopod volumes of the Treatise on In-
vertebrate Paleontology (Williams et al., 1965) a great deal of new information
about spiriferid brachiopods has accrued concerning their diversity, morphology,

1 Department of Geosciences, Oregon State University, Corvallis, Oregon 97331-5506 (deceased 5
July 1994).

2 Laboratoire de Paleontologie et de Stratigraphie du Paleozoique, Universite de Bretagne Occidental,
U.F.R. Sciences & Techniques, 6 Avenue le Gorgeu, B.P. 452, F-29275 Brest Cedex, France.

3 30th International Geological Congress, P.O. Box 823, Beijing 100037, People’s Republic of China.
Submitted 31 May 1994.

327

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distribution, and phylogenetic relationships. This is especially true for the spiri-
ferid brachiopods. For example, in the first edition of the Treatise Pitrat (1965)
recognized 187 spiriferid genera. Over 340 additional genera have been described
in the intervening years, almost tripling the number of genus-level taxa. Also,
numerous family-level taxon proposals have accompanied this descriptive explo-
sion, often without discussion or written justification for the new taxa. Thus, a
new evaluation of genera and their assignment to higher taxa is clearly in order.

Waterhouse (1968), Ivanova (1972), and Carter (1974) offered revised classi-
fications for part or all of the spiriferid brachiopods soon after publication of the
original Treatise volumes. These classifications are long outdated and have been
emended numerous times or ignored. None has been accepted as a standard for
the group.

In the following classification considerable emphasis is placed on shell structure
(orders), growth form (suborders or superfamilies), internal structures (families
or subfamilies), and micro-ornament (all family levels). Several possible synapo-
morphies are identified: taleola-like denticles in the Spiriferoidea and Paeckel-
manelloidea, biramous spines in the Elythidae, the tichorhinum in the Cyrtinidae,
development of a syrinx in the Syringothyridoidea, crenulation of the hingeline
in the Superfamily Spondylospiroidea.

Although cladistic analysis was not performed in constructing this classification,
it is hoped that it reflects the phylogeny of the spiriferids to a large degree. Carlson
(1991), in a phylogenetic study of the brachiopod superfamilies, pointed out that
the spiriferids, as classified in the 1965 Treatise, were not monophyletic and did
not comprise a clade. She is presently reanalyzing this group using some of the
information provided here.

Order Spiriferida Waagen, 1883
[nom. correct. Moore, Lalicker, and Fischer, 1952, p. 221
( pro order Spiriferacea Kuhn, 1949, p. 104;
nom. transl. ex suborder Spiriferacea Waagen, 1883, p. 447),
emend. Carter, Johnson, Gourvennec, and Hou, herein]

Diagnosis. — Generally biconvex; generally transverse with moderately wide to
extended straight hingeline; ribbing very fine to coarse; small dorsal and larger
ventral interareas always developed; spiralia directed laterally or posterolaterally
with primary lamellae parallel and close to sagittal plane; jugum absent; shell
substance impunctate.

Discussion. — The Order Spiriferida comprises the impunctate strophic bra-
chiopods with a spiral brachidium and ranges from the Upper Ordovician to the
Upper Permian. The oldest suborder, Spiriferidina Waagen, 1883, is represented
in the Lower Paleozoic by eospiriferid genera such as Eospirifer and Macropleura
that bear a capillate shell fabric and a simple interior with dental and crural plates
and lack a ctenophoridium. Several authors (Rudwick, 1970; Ivanova, 1972) have
suggested derivation of the order from an impunctate strophic orthid based on
general morphology and age of first appearance. We tend to agree with this sug-
gestion but cannot with confidence propose a likely intermediate ancestor. Gour-
vennec and Melou (1990) recently demonstrated the similarity between the micro-
ornament of the Upper Ordovician orthid genus Ptychopleurella and that of the
spiriferid genus Hysterolites. However, this similarity could be accounted for by
convergent or parallel evolution.

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Carter et al.— Spiriferid Brachiopod Revision

329

Rong et al. (1994) recently described a Late Ordovician (mid-Ashgill) species
of Eospirifer from eastern China. This primitive species lacks crural plates and
has very simple posterior cardinalia, quite unlike any known orthid. Furthermore,
“spiriferoid” orthids such as Platystrophia or Mcewanella are not known from
any of the areas where primitive eospiriferids have been recovered. Although the
brachidium is not known for these early Chinese eospiriferids the simple cardinalia
are reminiscent of some early atrypids and the possibility remains that the im-
punctate spiriferids may have been derived from the atrypids, not the orthids. It
can be seen from these discoveries that more evidence is needed to ascertain the
true ancestors of the spiriferids.

The Devonian history of the Spiriferidina is one of proliferation and diversity
exemplified by the superfamilies Spinelloidea and Theodossioidea, lacking a del-
thyrial plate, and the Cyrtospiriferoidea, which have that structure. Late in the
Famennian the Theodossioidea gave rise to several genera assigned to the Ul-
bospiriferidae, with an inner prismatic shell layer such as is found commonly in
various Carboniferous and Permian spiriferids. Late the in Devonian the theo-
dossioids, probably an ulbospiriferid, gave rise to several superfamilies that be-
came dominant elements of the Late Paleozoic brachiopod faunas, namely the
Spiriferoidea, Paeckelmanelloidea, and Brachythyridoidea.

Soon after the proliferation of the capillate Spiriferidina the Delthyridina ap-
peared as a second major branch of impunctate genera, including Howellella and
Delthyris , and characterized by a fimbriate shell fabric. Remnants of a transitional
shell fabric are present in the Silurian and Lower Devonian Cyrtinopsididae and
Acrospiriferidae of the Superfamily Delthyridoidea, but by the Middle Devonian
the two suborders were well represented and distinct. The Devonian history of
the Delthyridina is also one of proliferation and diversity, represented by nu-
merous families and subfamilies. By the Carboniferous most of the delthyridoid
families had disappeared, but the reticularioids, which arose from the Delthyridina
in the Silurian, flourished with great diversity until the end of the era.

The origin of the Delthyridina is not known. Gourvennec (1989, 1991) has
amply demonstrated that the micro-ornament of this group is substantially dif-
ferent from that of the eospiriferids. It is possible that the Delthyridina were
derived from a group other than the eospiriferids or from a common ancestor, in
which case the Spiriferida, as presented here, is diphyletic.

Two additional superfamilies, the Ambocoelioidea and the Martinioidea, split
off from the Spiriferidina in the Silurian. There has been uncertainty about the
derivation of these two superfamilies in the past. The ambocoeliids were judged
by Pitrat (1965) to have been derived from an eospiriferid ancestor, but this was
disputed by Ivanova (1972) who regarded both the Ambocoelioidea and Marti-
nioidea as having separated from the Delthyridina. Here, we follow Pitrat and
also Johnson and Lenz (1992) in regarding the ambocoelioids as having originated
from an eospiriferoid stock. We argue here for a similar derivation of the mar-
tinioids on the basis of shell fabric and internal morphology of early genera.
Ambocoelioid genera are common and diverse in the Devonian, but were reduced
to less than a dozen genera in the Late Paleozoic. The martinioids are much less
diverse and sporadically represented in the Devonian, but became numerous and
diversified in the Carboniferous and Permian, just before their extinction in the
Late Permian.

Stratigraphic Range.— Upper Ordovician-Upper Permian.

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Suborder Spiriferidina Waagen, 1883
[nom. correct. Pitrat, 1965, p. H668
( pro suborder Spiriferacea Waagen, 1883, p. 447)]

[emend. Carter, Johnson, and Gourvennec, herein]

Diagnosis. — Lateral slopes plicate or costate; fold and sulcus commonly well
developed; fine ornament, if present, capillate, pustulose, or imbricate; spinose
ornament absent; ctenophoridium absent in early forms.

Stratigraphic Range. — Upper Ordovician-Upper Permian.

Superfamily Cyrtioidea Frederiks, 1924
[nom. correct. Johnson, Gourvennec, and Hou, herein
(pro superfamily Cyrtiacea Pitrat, 1965, p. H668);
nom. transl Pitrat, 1965, (ex Cyrtiinae Frederiks, 1924, p. 312)]

Diagnosis. — Biconvex, commonly with fold and sulcus, smooth or plicate; or-
nament of nonspinose capillae crossed by variably nodose growth lines; deltidium,
delthyrial plate, or stegidial plates may be present; dental plates present; crural
plates well developed, rudimentary, or lacking.

Stratigraphic Range.— Upper Ordovician (Middle Ashgill)-Lower Devonian
(Emsian).

Family Cyrtiidae Frederiks, 1924

[nom. transl. Ivanova, 1959, p. 55 (ex Cyrtiinae Frederiks, 1924, p. 312)]
[^Eospiriferinae Schuchert, 1929, p. 20]

Diagnosis. — Delthyrial plate and stegidial plates lacking; crural plates com-
monly well developed.

Stratigraphic Range.— Upper Ordovician (Middle AshgilD-Lower Devonian
(Emsian).

Subfamily Cyrtiinae Frederiks, 1924
[Cyrtiinae Frederiks, 1924, p. 312]

Diagnosis. — Ventribiconvex, ventral valve hemipyramidal, with catacline or
procline interarea, narrowly elongate delthyrium; deltidium with medial foramen;
ctenophoridium absent.

Discussion. — The species Cyrtia approximans (Barrande, 1879) was listed as
Devonian Ff2 by Barrande (1879) and from the Acanthopyge Limestone by Hav-
licek, (1959:75). This led Boucot (1963) to list the upper range of Cyrtia as lower
Devonian, but other Devonian occurrences of Cyrtia have not been verified. A
query to Havlicek brought the following response (written communication, July
11, 1991): 44 Cyrtia approximans (Barrande) is a true Cyrtial The data about its
age and type locality, however, are not correct in Barrande’s paper. After revising
this species, I came to the conclusion that Cyrtia approximans is conspecific with
the Silurian Cyrtia spiriferoides Boucek, 1941 (Wenlock, Motol Formation).”

Stratigraphic Range.— Upper Silurian (Upper Wenlockian-Ludlovian).

Genera Included. — Cyrtia Dalman, 1828; Plicocyrtia Boucot, 1863; Dongbeiis-
pirifer Liu, 1977.

Subfamily Eospiriferinae Schuchert, 1929
[Eospiriferinae Schuchert in Schuchert and Levene, 1929, p. 20]

Diagnosis. —Biconvex with curved, commonly apsacline ventral interarea and
open delthyrium; deltidium present; ctenophoridium commonly absent.

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Carter et al.— Spiriferid Brachiopod Revision

331

Discussion. — In the list below of assigned genera we retain Macropleura although
it was regarded as a junior subjective synonym of Hedeina after the type species
of Hedeina was made known by Brunton et al. (1967). Boucot (1975:362) suggested
retaining Macropleura for large-sized Lower Devonian species, but we propose
to include both large and small species based on the amplitude of plications and
interspaces rather than on shell size. Macropleura has a few prominent plications
and deep, U-shaped interspaces. Hedeina is characterized by broad, low plications
and shallow interspaces. Probably, Hedeina was derived from Eospirifer and
belongs to a different branch of eospiriferid phylogeny than Macropleura. Our
definition would include Hedeina ananias Johnson, Boucot, and Murphy (1976)
in Macropleura. We also synonymize Ejnespirifer Fu, 1982 with Macropleura on
the same basis. According to Rong Jia-yu (written communication, March 6,
1994) the type species of Ejnespirifer does not have strut-like accessory plates in
the dorsal valve as were illustrated by Fu (1982:177, fig. 92B).

Stratigraphic Range.— Upper Ordovician (Middle Ashgill)-Lower Devonian
(Emsian).

Genera Included . — Eospirifer Schuchert, 1913; Badainjarania Zhang, 1981; En~
dospirifer Tachibana, 1981; Espella Nilova, 1965 [=Laevispirifer Ushatinskaia,
1977]; Havlicekia Boucot, 1963; Hedeina Boucot, 1957; Janius Havlicek, 1957;
Lobvia Breivel and Breivel, 1977; Macropleura Boucot, 1963 [= Ejnespirifer Fu,
1982]; Mictospirifer Johnson, 1995; Myriospirifer Havlicek, 1978 [=Acutilineolus
Amsden, 1978]; Nurataella Larin, 1973 [=Baterospirifer Rong, Su, and Li, 1984];
Striispirifer Cooper and Muir- Wood, 1951; Yingwuspirifer Rong, Xu, and Yang,
1974; Xinanospirifer Rong, Xu, and Yang, 1974.

Family Hedeinopsidae Gourvennec, 1990
[nom. transl. Johnson, herein
(ex Hedeinopsinae Gourvennec, 1990, p. 142)]

Diagnosis. — Plicate, ventribiconvex, with stegidial plates and delthyrial plate;
crural plates rudimentary or absent; ctenophoridium present.

Stratigraphic Range. — Upper Silurian (Wenlockian-Pridolian).

Subfamily Hedeinopsinae Gourvennec, 1990
[Hedeinopsinae Gourvennec, 1990, p. 142]

Diagnosis. — As for the family.

Discussion.— Examination of the holotype of Tannuspirifer dixoni Jones (1980:
pi. 1, fig. 1), on loan from Jones, reveals that it has stegidial plates.

Stratigraphic Range. — Upper Silurian (Wenlockian-Pridolian).

Genera Included. —Hedeinopsis Gourvennec, 1990; Tannuspirifer Ivanova, 1960.

Superfamily Spinelloidea Johnson, 1970
[nom. transl. Johnson and Hou, herein
(ex Spinnellinae Johnson, 1970, p. 205)]

Diagnosis. — Biconvex, with fold and sulcus; plicate; ornament nonfrilly, cap-
illate; delthyrial plate lacking; dental plates, without median septum.

Stratigraphic Range.— Upper Silurian (Wenlockian)-Upper Devonian (Upper
Famennian).

Family Spinellidae Johnson, 1970
[nom. transl. Wang and Rong, 1986, p. 200
(ex Spinellinae Johnson, 1970, p. 205)]

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vol. 63

Diagnosis . — Multiplicate, with smooth or medially grooved fold and smooth
or medially plicate sulcus.

Stratigraphic Range. — Upper Silurian ( W enlockian)-U pper Devonian (Upper
Famennian).

Subfamily Spinellinae Johnson, 1970
[Spinellinae Johnson, 1970, p. 205]

[=Guerichellinae Paeckelmann, 1931, p. 24]

Diagnosis. — Multiplicate, with smooth fold and sulcus, or with medial plication
in sulcus; crural plates short or lacking.

Stratigraphic Range. — Lower Devonian (Pragian)-Upper Devonian (Upper Fa-
mennian).

Genera Included. —Spinella Talent, 1956; Acutatheca Stainbrook, 1 945; Adolfia
Giirich, 1909 [=Guerichella Paeckelmann, 1913; =C hangs haispirifer Zhao, 1977];
Allanella Crickmay, 1953 [= Minutilla Crickmay, 1967]; Chimaerothyris Paulus,
Struve, and Wolfart, 1963; Eospiriferina Grabau, 1931; Fidespirifer Liashenko,
1973; Guicyrtia Wang and Zhu, 1979; Volgospirifer Shevchenko, 1970.

Subfamily Pinguispiriferinae Havlicek, 1971
[Pinguispiriferinae Havlicek, 1971, p. 27]

Diagnosis. — Plicate, with weak to obscure capillae, crossed by fila; crural plates
short or lacking.

Discussion. — Several genera of the Pinguispiriferinae have been classified with
the eospiriferids in the past, but although capillate they lack the well-developed
capillae typical of eospiriferids. Others differ also in the construction of the car-
dinalia. Nikiforovaena Boucot, 1963, previously unknown in the dorsal interior,
was examined by preparing a dorsal internal mold of the type species; it revealed
the presence of a ctenophoridium and short, divergent crural plates.

Stratigraphic Range.— Upper Silurian (Wenlockian)-Middle Devonian (Fife-
ban).

Genera Included. —Pinguispirifer Havlicek, 1957; Amoenospirifer Havlicek,
1957; Brevispirifer Cooper, 1942; Ljudmilispirifer Cherkesova, 1976; Nikiforo-
vaena Boucot, 1963; Spurispirifer Havlicek, 1971.

Subfamily Callispiriferinae Johnson, new subfamily

Diagnosis. — Non-capillate, with high flat ventral interarea.

Stratigraphic Range. — Lower Devonian (Emsian).

Genera Included. — Callispirifer Perry, 1984; Rochtex Havlicek, 1990.

Subfamily Eurekaspiriferinae Johnson, new subfamily

Diagnosis. — Capillate, with dorsal adminicula.

Discussion. — The presence of dorsal adminicula in Eurekaspirifer suggests an

even greater separate ranking than subfamily, but as no closely related genera are
now known proposal of a more substantial taxonomy seems inappropriate.

Stratigraphic Range. — Lower Devonian (Lower Emsian).

Genera Included.— Eurekaspirifer Johnson, 1966.

Family Echinospiriferidae Liashenko, 1973
[Echinospiriferidae Liashenko, 1973, p. 109]

[=Rigauxidae Brice, 1988, p. 371]

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Diagnosis. — Multiplicate, with plicate fold and sulcus.

Discussion.— A specimen of the type species of Arctospirifer Stainbrook, 1950
was sectioned confirming the absence of a delthyrial plate and the absence of
crural plates. Regauxia Brice, 1988 includes the widely represented North Amer-
ican species Spirifer orestes Hall and Whitfield and Spirifer strigosus Meek, both
of which have been assigned mistakenly to Indospirifer Grabau, 1931 by various
authors.

Stratigraphic Range. — Lower Devonian (Pragian)-Upper Devonian (Upper Fa-
mennian).

Genera Included. —Echinospirifer Liashenko, 1973; Adolfispirifer Krylova, 1962;
Arctospirifer Stainbrook, 1950; Enchondrospirifer Brice, 1971; Hispidaria Cooper
and Dutro, 1982; Howittia Talent, 1956 [=Glyptospirifer Hou and Xian, 1975];
Indospirifer Grabau, 1931 [=Schizospirifer Grabau, 1931]; Regauxia Brice, 1988;
Sergunkovia Nalivkin, 1979.

Superfamily Theodossioidea Ivanova, 1959
[nom. transl. Johnson, Carter, and Hou, herein
(ex Theodossiinae Ivanova, 1959, p. 61)]

Diagnosis. — Uniformly plicate or costate, with fold and sulcus indistinct or
lacking; delthyrial plate lacking.

Discussion. — The superfamily Theodossioidea, with its one new family and
three new subfamilies, unites genera that are plicate or costate over the entire
exterior and which lack a well-defined fold and sulcus. An implication in this
classification is that there are genetic links from the earliest genera of the subfamily
Branikiinae, of Emsian age, to the Middle Devonian genera of the Theodossiinae.
However, the origin of the Branikiinae is uncertain. The genus Theodossia is the
supposed link to Famennian and younger genera of the Ulbospiriferidae, which
are characterized by the presence of an inner prismatic shell layer. The Early and
Middle Devonian genera are uncommon taxa and this together with their unusual
morphology has resulted in their previously being outside of generally accepted
spiriferid systematics.

Stratigraphic Range. — Lower Devonian (Pragian)-Upper Devonian (Upper Fa-
mennian).

Family Theodossiidae Ivanova, 1959
[nom. transl. Johnson, Carter, and Hou, herein
(ex Theodossiinae Ivanova, 1959, p. 61)]

Diagnosis. — Without inner prismatic shell layer.

Stratigraphic Range. — Lower Devonian (Pragian)-Upper Devonian (Frasnian).

Subfamily Theodossiinae Ivanova, 1959
(Theodossiinae Ivanova, 1959, p. 61)

Diagnosis. — Capillate.

Stratigraphic Range.— Middle Devonian (Eifelian)-Upper Devonian (Frasni-
an).

Genera Included. — Theodossia Nalivkin, 1925 [= Vandergrachtella Crickmay,
1953]; Paralazutkinia Jiang, 1978 [=Pinghuangella Jiang, 1978]; Urella Rzhon-
snitskaia, 1960 [=Retzispirifer Kulkov, I960].

Subfamily Branikiinae Johnson and Hou, new subfamily

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Diagnosis. — Moncapillate.

Stratigraphic Range. — Lower Devonian (Pragian)-Middle Devonian (Eifelian).

Genera Included.— Branikia Havlicek, 1957 [=Bidentatus Khodalevich and
Breivel , 1972]; Jilinospirifer Su, 1980; Lenzia Perry, Boucot, and Gabrielse, 1981,

Family Ulbospiriferidae Johnson and Carter, new family

Diagnosis. — With inner prismatic shell layer.

Stratigraphic Range. — Upper* Devonian (Upper Famennian),

Subfamily Ulbospiriferinae Johnson and Carter, new subfamily

Diagnosis. ™ Uniformly costate.

Stratigraphic Range. — Upper Devonian (Upper Famennian).

Genera Included. — Ulbospirifer Grechishnikova, 1965; Cyrtiorina Cooper and
Dut.ro, 1982; Temsm Martynova, 1970 [= Omolonospirifer Simakov, 1970].

Subfamily Palaeospiriferinae Carter, Johnson, and Hou, new subfamily

Diagnosis, — With low, rounded plications.

Stratigraphic Range. — Upper Devonian (Upper Famennian).

Genera Included.— Palaeospirifer Martynova and Sverbilova, 1968 [=Goung-
junspirifer Zhang, 1983].

Family Palaeochoristitidae Carter, new family

Diagnosis. — Cardinal extremities rounded in juveniles, outline variable in adults;
well-developed dental adminicula and dorsal adminicula present; micro-omament
absent.

Discussion. —The origins of this family are obscure because dorsal adminicula
are rare in the Theodossioidea. The only known Devonian theodossioid genus
with dorsal adminicula is the Russian Middle Devonian genus Ur el la Rzhon-
snitskaya, 1960, which is presumed to be in the line of ancestry of the palaeo-
choristitids.

Stratigraphic Range.— Lower Carboniferous (Toumaisian).

Genera Included . — Palaeochoristites Sokolskaya, 1941; Eochoristites Chu, 1933
[=Centrospirifer Tien, 1938].

Superfamily Cyrtospiriferoidea Termier and Termier, 1949
[nom. transl Ivanova, 1972, p. 31
(ex Cyrtospiriferinae Termier and Termier, 1949, p. 99)]

Diagnosis. — Nonfrilly, capillate or pustulose; ddthyrial plate present; cteno-
phoridium without crural plates.

Stratigraphic Range.— Lower Devonian (Emsian)-Upper Devonian (Upper Fa-
mennian).

Family Spinocyrtiidae Ivanova, 1959
[nom. transl. Pitrat, 1965, p. H688
(ex Spinocyrtiinae Struve, 1963, p. 462,
nom . correct, pro Spinocyrtinae Ivanova, 1959, p. 59)]

Diagnosis. — Multiplicate, with smooth or medially grooved fold and smooth
or medially plicate sulcus.

Stratigraphic Range. L ower Devonian (Emsian)-Upper Devonian (Frasnian).

Genera Included. — Acutoria Cooper and Dutro, 1 982; Alaiiformia Struve, 1 963;

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Carter et al.™ Spiriferid Brachiopod Revision

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Eosyringothyris Stainbrook, 1943; Mediospirifer Bublitchenko, 1956 [=Carpi-
naria Struve, 1982]; Duryeella Boucot, 1975; Orthospirifer Pitrat, 1975; Platyr-
achella Fenton and Fenton, 1924; Spinocyrtia Frederiks, 1916; Subcuspidella
Mittmeyer, 1965 [=Tenuicostella Mittmeyer and Geib, 1967].

Family Cyrtospiriferidae Termier and Termier, 1949
[nom. transl. Beznosova, 1958, p. 17
(ex Cyrtospiriferinae Termier and Termier, 1949, p. 99)]

Diagnosis. — Nonfrilly, flanks costate; fold and sulcus finely costate.

Discussion. — The assignments of at least 19 genera to one of two subfamilies
on the basis of shell shape is inadequate. A re-evaluation of the genera of the
Family Cyrtospiriferidae on the basis of internal structures is obviously needed.
An initial appraisal might usefully involve a determination of dental plates (in-
trasinal or extrasinal).

Stratigraphic Range.— Middle Devonian (Upper Gi vetian)-U pper Devonian
(Upper Famennian).

Subfamily Cyrtospiriferinae Termier and Termier, 1949
[Cyrtospiriferinae Termier and Termier, 1949, p. 99]
[=Hunanospiriferinae Beznosova, 1958, p. 17]

Diagnosis. —Wide interarea with acute cardinal angles.

Stratigraphic Range.— Upper Devonian (Frasnian-Upper Famennian).

Genera Included. — Cyrtospirifer Nalivkin in Frederiks, 1924 [=Hunanospirifer
Tien, 1938; Grabauispirifer Gatinaud, 1949; Eurytatospirifer Gatinaud, 1949;
Deothossia Gatinaud, 1949; Lamarckispirifer G atinaud, 1949; Subquadriangulis-
pirifer Sartenaer, 1982]; Austrospirifer Glenister, 1956; Geminisulcispirifer Sar-
tenaer, 1982; Liraspirifer Stainbrook, 1950; Petshorospirifer Fotieva, 1985; Re-
gelia Crickmay, 1952; Sinospirifer Grabau, 1931; Sphenospira Cooper, 1954;
Syringospira Kindle, 1909; Tarandrospirifer Simakov, 1970; Tenticospirifer Tien,
1938 [=Conispirifer Liashenko, 1985].

Subfamily Cyrtiopsinae Ivanova, 1972
[Cyrtiopsinae Ivanova, 1972, p. 33]

[=?Uchtospiriferidae Liashenko, 1973, p. 87]

Diagnosis. — Narrow interarea with rounded cardinal angles.

Discussion. — Cyrtiopsis Grabau has been widely reported in Famennian rocks,
but lack of knowledge of the Cyrtiopsis interior, as well as lack of critical attention
to shell form makes many citations of the genus dubious. Examples are in the
influential paper by Crickmay (1952), including taxa he named Cyrtiopsis na-
hanniensis and C. normandvillana. We cite these two taxa because Crickmay
provided drawings intended to represent the disposition of the dental plates as
intrasinal and recurved distally (1952:pl. 72, fig. 12; pi. 74; fig. 4). We have
examined specimens of these two species in the Crickmay collections, on loan
from the Paleontological Research Institution, Ithaca, New York. Dental lamellae
observed in internal molds of both species are intrasinal, but long and straight
rather than being recurved distally. In this they resemble a prepared internal mold
in our possession of the type species C. davidsoni, but they differ from it in shell
form. Cyrtiopsis davidsoni has a deep, wide, and sharply bounded ventral sulcus
that gives it a distinctive shell shape. We know of no North American species
like it.

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Stratigraphic Range.— Middle Devonian (Upper Gi vetian)-U pper Devonian
(Famennian).

Genera Included. — Cyrtiopsis Grabau, 1923 [=Grabauicyrtiopsis Gatinaud, 1949;
Sinocyrtiopsis Gatinaud, 1949]; Dichospirifer Brice, 1971; Dmitria Sidiachenko,
1961; Eodmitria Brice, 1982; Platyspirifer Grabau, 1931; Uchtospirifer Liashenko,
1957 [=Timanospirifer, Nordispirifer, and Clivospirifer Liashenko, 1973]; Men-
nespirifer Liashenko, 1973 [=Komispirifer Liashenko, 1973]; Acutella Liashenko,
1973/

Superfamily Ambocoelioidea George, 1931
[nom. transl. Johnson and Carter, herein
(ex Ambocoeliinae George, 1931, p. 42]

Diagnosis. — Commonly small, lacking well-developed fold and sulcus; cardinal
process commonly simple, knoblike; outer hinge plates broad, well developed;
cruralium variably developed.

Stratigraphic Range. —Upper Silurian (Upper Wenlockian)-Permian.

Family Ambocoeliidae George, 1931
[nom. transl Ivanova, 1959, p. 56
(ex Ambocoeliinae George, 1931, p. 42)]

Diagnosis. — Ventribiconvex, smooth shells, rarely pauciplicate; commonly
lacking dental plates.

Stratigraphic Range. —Upper Silurian (Upper WenlockianJ-Permian.

Subfamily Ambocoeliinae George, 1931
[Ambocoeliinae George, 1931, p. 42]

Diagnosis. — Crural plates vestigial or lacking; commonly with fine concentric
growth lamellae, and fine capillae on older genera; dental plates lacking in younger
genera.

Discussion. — Pitrat (1965:H667) suggested that the ambocoeliid brachiopods
were derived from the Eospiriferinae, based on age of appearance, nonstriate site
of dorsal diductor attachment, crural plates, and permissive similarities of fine
ornament. Although Pitrat erred in attributing crural plates to the earliest am-
bocoeliids, the derivation seems plausible, beginning with the oldest ambocoeliid
Eoplicoplasia Johnson and Lenz, 1992. Eoplicoplasia morphology suggests der-
ivation from a plicate eospiriferid genus such as Macropleura Boucot. This is
based on the presence of Macropleura in older Silurian beds (Boucot, 1963:706;
Havlicek, 1980:1 1), similar overall shell shape, nonstriate site of dorsal diductor
attachment, dental plates, and the presence on Eoplicoplasia of a relict eospiriferid
radial ornament of fine capillae (Lenz, 1972:pl. 2, fig. 2, 21). Eoplicoplasia and
Plicoplasia both lack crural plates, consistent with their assignment to the Sub-
family Ambocoeliinae (Johnson and Trojan, 1982:128-129).

Stratigraphic Range. —Upper Silurian (Upper Wenlockian)-Permian.

Genera Included. —Ambocoelia Hall, 1 860; Attenuatella Stehli, 1954; Aviformia
Xian, 1988; Biconvexiella Waterhouse, 1983; Bisinocoelia Havlicek, 1953; Cruri-
cella Grant, 1976; Crurithyris George, 1931; Cyrtinoides Iudina and Rzhonsnit-
skaia, 1985 [=Mucroclipeus Goldman and Mitchell, 1990]; Dicoelospirifer Zhang,
1989; Echinocoelia Cooper and Williams, 1935 [= Pyramina Liashenko, 1969];
Eoplicoplasia Johnson and Lenz, 1992; Guangxiispirifer Xian, 1983; Orbicoelia
Waterhouse and Piyasin, 1970; Paracrurithyris Liao, 1979; Plicoplasia Boucot,

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1959; Metaplasia Hall and Clarke, 1893; Spinoplasia Boucot, 1959; Swaicoelia
Hamada, 1968.

Subfamily Rhynchospiriferinae Paulus, 1957
[Rhynchospiriferinae Paulus, 1957, p. 51]

[=Ilmeniinae Durkoop, 1970, p. 195]

Diagnosis.— Crural plates well developed, either discrete or joined to form a
cruralium; commonly with fine capillae; dental plates present or absent.

Stratigraphic Range. — Lower Devonian (Emsian)-Upper Devonian (Frasnian).

Genera Included. —Amboglossa Wang and Zhu, 1 979; Ambothyris George, 1931;
Changtangella Xian, 1982; Choperella Liashenko, 1969; Crurispina Goldman and
Mitchell, 1990; Diazoma Durkoop, 1970 [=Kelusia Mamedov, 1978]; Emanuella
Grabau, 1923; [=Paraemanuella Yang, 1977]; Ilmenia Nalivkin, 1941; Ilmen-
iopsis Xian, 1982; Ilmenispina Havlicek, 1959; Ilmospirifer Liashenko, 1969;
Ladjia Veevers, 1959; Levibiseptum Xian, 1975; Moravilla Havlicek, 1953; Rhyn-
chospirifer Paulus, 1957 [= Kosirium Ficner and Havlicek, 1975; Biarella Mar-
kovski, 1988]; Zhonghuacoelia Chen, 1978.

Family Vemeuiliidae Schuchert, 1929
[nom. transl. Brunton, 1984, p. 101
(ex Vemeuiliinae Schuchert, 1929, p. 21)]

Diagnosis. — Transverse to subcircular, biconvex, with opposite folding forming
ligate to metacarinate anterior margins; ventral interarea narrow to full width of
valves, with open delthyrium or restricted by apical deltidium; growth lines with-
out capillae.

Stratigraphic Range.— Middle Devonian (Eifelian)-Lower Carboniferous.

Genera Included. — Verneuilia Hall and Clarke, 1 893; Minythyra Brunton, 1984;
Nuguschella Tiazheva, 1960.

Family Lazutkiniidae Johnson and Hou, new family

Diagnosis. — Wholly plicate, lacking capillae; with dental plates.

Stratigraphic Range. —Lower Devonian (Emsian)-Middle Devonian (Give-
tian).

Genera Included.— Lazutkinia Rzhonsnitskaia, 1952 [= Yavorskiella Lazutkin
in Yavorsky, 1940]; Prolazutkinia Hou and Xian, 1983.

Family Eudoxinidae Nalivkin, 1979
[nom. correct, et transl. Carter, herein
( pro Subfamily Eudoxininae Nalivkin, 1979, p. 145)]

Diagnosis. — Small to very large; biconvex; costate or costellate; micro-ornament
papillose in some, absent or unknown in most genera; dental adminicula and
protuberant ctenophoridium absent.

Discussion. — The lack of a convex ctenophoridium in this group convincingly
points to its derivation from the eospiriferid-ambocoeliid lineage. There is a great
difference in size between the Early Carboniferous Eudoxina and Paulonia and
the Permian Costicrura and Wilberrya. In the absence of intermediate forms of
Visean or Upper Carboniferous age the homogeneity of this group cannot be
tested easily.

Stratigraphic Range.— Upper Devonian (Famennian)~Upper Permian.

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Genera Included. — Eudoxina Frederiks and Kruglov, 1928; Costicrura Hoover,
1981 ; Paulonia Nalivkin, 1925; Wilberrya Yancey, 1978.

Superfamily Martinioidea Waagen, 1883
[nom. correct. Carter, Johnson, and Gourvennec, herein
(pro Martiniacea Waagen, 1883;
nom. transl. Ivanova, 1972, p. 41,
ex Subfamily Martiniinae Waagen, 1883, p. 524)]

Diagnosis.— Biconvex, with broadly rounded lateral extremities and short
hingeline; lateral slopes smooth or with subdued ribbing; micro-ornament com-
monly capillate, smooth, or very finely spinulose, but not strongly lamellose or
concentrically disposed; ventral median septum absent; dorsal interior with cteno-
phoridium in all but the earliest genera, commonly with crural plates or dorsal
adminicula; surface commonly pitted.

Discussion. — This superfamily is now believed to have been derived from the
ambocoeliids in Late Silurian time through the acquisition of consistently inflated
dorsal valves and pitted micro-ornament. The earliest family, the Tenelloder-
midae, lack a ctenophoridium, attesting to their derivation from the ambocoe-
lioids. Internally, the appearance of crural plates or strong dorsal adminicula is
manifested in several lineages throughout most of the history of the superfamily.

Stratigraphic Range. —Silurian (Upper Wenlock)-Upper Permian.

Family Tenellodermidae Carter, Johnson, and Gourvennec, new family

Diagnosis. — Ctenophoridium absent; surface pitted or simple; dental plates
present.

Discussion. — In the absence of a ctenophoridium these two genera provide a
convincing transition from the ambocoelioids to ctenophoridium-bearing Late
Paleozoic martinioids. The earliest species of both genera occur in the Upper
Wenlockian of Bohemia, but we suggest that Tenellodermis most closely resembles
an ambocoeliid ancester in its simple cardinalia having neither striate cardinal
process nor crural plates.

Stratigraphic Range. — Silurian (Upper Wenlockian)-Middle Devonian (Eife-
lian).

Genera Included. — Cinguloder mis Havlicek, 1971; Tenellodermis Havlicek,
1971.

Family Elythynidae Gourvennec, new family

Diagnosis. — Flanks plicate; ornament of fine pits or spines and pits; cteno-
phoridium, dental plates, and crural plates or dorsal adminicula present.

Discussion. — These genera are unified by their similar internal morphology and
represent the earliest martinioids with a ctenophoridium. Externally, they vary
in their micro-ornament.

Elythyna has fine elongate pits or grooves arranged in quincunx similar to the
Ingelarellidae. Examination of topotype material of the type species, E. salarica,
shows that there are short subparallel crural plates (or dorsal adminicula?) and a
ctenophoridium. Najadospirifer Havlicek possesses discontinuous capillae with
fine ovate pits radially arranged in the interspaces (Havlicek, 197 1 :pl. 1, 2) and
internally the crural plates are very short (Boucot, 1962:pl. 49). Tatjanaspirifer
is assigned here despite its ornament of radial marginal spinules of the reticulari-
oid type. This genus also shows randomly distributed pits which were originally

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Carter et al. — Spiriferid Brachiopod Revision

339

diagnosed as coarse spine bases by Cherkesova. This pitted ornament is unknown
in the reticularioids and strongly suggests affinity with the martinioids. These pits
are sometimes best seen in the fibrous layer and may not be apparent unless the
outer layer is exfoliated, a feature common in the martinioids.

Stratigraphic Range. — Lower Devonian (Pragian)-Middle Devonian (Give-
tian).

Genera Included.— Ely thy na Rzhonsnitskaya, 1952; Najadospirifer Havlicek,
1957; Tatjanaspirifer Cherkesova, 1991.

Family Martiniidae Waagen, 1883
[nom. transl. Ivanova, 1959, p. 56,
ex Subfamily Martiniinae Waagen, 1883, p. 524]

Diagnosis. — Ctenophoridium present; subequally biconvex; lateral slopes smooth
or weakly plicate; crural plates or dorsal adminicula absent; micro-ornament of
scattered surficial pits, and/or capillae, or absent.

Stratigraphic Range.— Upper Devonian (Famennian)-Upper Permian.

Subfamily Martiniinae Waagen, 1883
[Subfamily Martiniinae Waagen, 1883, p. 524]

Diagnosis. — Lacking plates or septa in either valve; micro-ornament of fine pits
only.

Stratigraphic Range. — Lower Carboniferous-Upper Permian.

Genera Included.— Martinia McCoy, 1844 [= Jilinmartinia Lee and Gu, 1980;
Paramartinia Reed, 1949; Pseudomartinia Leidhold, 1928]; Beschevella Poletaev,
1975; Implexina Poletaev, 1971; Kalitvella Lazarev and Poletaev, 1982; Posta-
martinia Want and Yang, 1993; Spinomartinia Waterhouse, 1968; Tiramnia
Grunt, 1977; Weiningia Ching and Liao, 1974 [=Elenchus Aleksandrov, 1973].

Subfamily Eomartiniopsinae Carter, new subfamily

Diagnosis. — Dental adminicula present; lateral slopes commonly weakly ribbed;
micro-ornament of fine pits or absent.

Discussion. — The genus Eomartiniopsis Sokolskaya has been repeatedly re-
ported in Devonian strata but we have doubts about the validity of these reports.
A detailed investigation of the dorsal interiors and micro-ornament of these
species is necessary to resolve this problem. The type species lacks crural plates
or dorsal adminicula. If we assume that all of the species, including Devonian
ones, assigned here by Sokolskaia (1941) also lack crural plates, the number of
Devonian species in need of future investigation is much reduced.

Five species were assigned to this genus by Rzhonsnitskaia (1952), but she did
not provide morphological evidence for her assignment. One of them, Spirifer
kirki, undoubtedly belongs to Warrenella. The same might be said for Spirifer
laevis judging from the presence of a pseudodeltidium, strong apical callosity,
deep muscle scars, etc. Spirifer maia was probably assigned here based on external
aspect, for no information is available concerning its internal structure. Apparently
S. maia possesses a pseudodeltidium which would suggest relationship to War-
renella, but more information is needed to confirm this assignment. The dorsal

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interiors of Eomartiniopsis lazutkini and Spirifer linguifer have never been de-
scribed.

HavHcek (1 959) assigned six more Devonian species and subspecies to Eomarti-
niopsis. He later 1971) restricted his concept of the genus to include only Car-
boniferous species and assigned his previous species to his genera Tenellodermis
and Cingulodermis. However, several Devonian species cannot be assigned to
these genera. Among them, E. sellata [Spirifer (. Martinia ) inflata sellata Paeckel-
mann, 1 9 1 3] is assigned here, but its dorsal interior is poorly known. The ornament
of Eomartiniopsis lazutkini and Spirifer linguifer, both Eifelian and mentioned
above, consists of quincuntially arranged pits, an ornament unknown in Cingu-
lodermis, but which evokes the Xngellarellidae. On the other hand the described
ornament of Cingulodermis may be an artifact of preservation and incompletely
known, for it is seemingly within the lineage of Tenellodermis (its ancestor) and
Eomartiniopsis (its presumed descendant), both of which possess pits.

The last Devonian species assigned to Eomartiniopsis is E. zeravschanica, a
nomen nudum , and is very poorly known. Thus, the presence of Eomartiniopsis
in the Devonian cannot be ascertained. It has been treated as a “grab-bag” genus
for smooth, poorly described reticularioids occurring in Devonian strata. Further
investigation will probably show that most Devonian species of “ Eomartinio -
posis ” would be better assigned to Cingulodermis, Warrenella, or similar genera.

Stratigraphic Range.— 1 Devonian, Lower Carboniferous-Upper Permian.

Genera Included.— Eomartiniopsis Sokolskaya, 1941; Globispirifer Tachibana,
1964; Heteraria Cooper and Grant, 1976; Kisilia Nalivkin, 1979; Merospirifer
Reed, 1949; Rallacosta Cooper and Grant, 1976.

Subfamily Elivellinae Carter, new subfamily

Diagnosis. — Micro-ornament of capillae and fine pits; dental plates becoming
progressively shorter in younger genera.

Stratigraphic Range.— Upper Devonian (Upper Famennian>-Lower Permian.

Genera Included. —Elivella Frederiks, 1924; Martiniella Grabau and Tien, 1931;
Ushkolia Martynova and Sverbilova, 1969; Moumina Frederiks, 1924.

Family Martiniopsidae Kotljar and Popeko, 1967
[nom. transl. Carter, herein,
ex Martiniopsinae Kotljar and Popeko, 1967, p. 182]

Diagnosis. — Dental and crural plates or dorsal adminicula present; lateral slopes
commonly with weak plications; micro-ornament usually weakly to strongly cap-
illate.

Stratigraphic Range.— Lower Carboniferous (To u m a i sia n)~ U pper Permian.

Genera Included.— Martiniop sis Waagen, 1883 [=Rorespirifer Waterhouse and
Piasin, 1970]; Arktikina Grunt, 1977; Crassumbo Carter, 1967; Nodaea Tachi-
bana, 1981.

Family Ingelarellidae Campbell, 1959
[nom. transl. Archbold and Thomas, 1986, p. 582,
ex Ingelarellinae Campbell, 1959, p. 333]

Diagnosis.— Micro-ornament of quincuncially arranged elongate grooves and
ridges; ventral interior with dental adminicula.

Discussion. — The genera in this family are differentiated and assigned to sub-
families on the basis of micro-ornament, the nature of which is poorly known for

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Carter et al.~ Spiriferid Brachiopod Revision

341

several genera. Reassignment or synonymizing of one or more genera is likely to
be necessary.

Stratigraphic Range. —Lower Carboniferous (Visean)-Upper Permian.

Subfamily Ingelarellinae Campbell, 1959
[Subfamily Ingelarellinae Campbell, 1959, p. 333]

Diagnosis. — -Spinules absent; dorsal adminicula well developed.

Discussion. —Ambikella Sahni and Srivastava is based on a single, poorly pre-
served specimen and its validity has not been confirmed.

Stratigraphic Range. — Lower Carboniferous (Visean)-Upper Permian.

Genera Included. —Ingelarella Campbell, 1959; ? Ambikella Sahni and Srivas-
tava, 1956; Fredericksia Paeckelmann, 1931; Tabellina Waterhouse, 1986; To-
miopsis Benediktova, 1956 [=Danzania Pavlova, 1989].

Subfamily Notospiriferinae Archbold and Thomas, 1986
[Subfamily Notospiriferinae Archbold and Thomas, 1986, p. 584]

Diagnosis. — Fold and sulcus well developed; micro-ornament of quincuncially
arranged grooves and ridges, with anteriorly directed spinules at posterior ends
of short grooves, and with deep elongate globose pits extending into secondary
layer under spinules; dorsal adminicula short or absent.

Stratigraphic Range.— Upper Carboniferous or Lower Permian-Upper Per-
mian.

Genera Included.— Not ospirifer Harrington, 1955; Farmerella Clarke, 1992.

Subfamily Glendoniinae Clarke, 1992
[Subfamily Glendoniinae Clarke, 1992, p. 75]

Diagnosis. - Micro-ornament of quincuncially arranged shallow elongate grooves
terminated anteriorly by low elongate spinules and shallow elongate pits.

Stratigraphic Range.— Permian (Asselian-Kazanian).

Genera Included.— Glendonia McClung and Armstrong, 1978; Birchsella Clarke,
1987; Homevalaria Waterhouse, 1986; Kelsovia Clarke, 1990.

Family Gerkispiridae Carter, 1985
[Family Gerkispiridae Carter, 1985, p. 376]

Diagnosis. — Ovate to transverse; lateral slopes costate; hingeline short; fold-
sulcus moderately to well developed; delthyrium with low thin flaring stegidial
plates; ventral interior with short thin dental adminicula, commonly with low
apical myophragm; dorsal interior with short converging crural plates and small
ctenophoridium; micro-ornament of quincuncially arranged very fine uniramous
hollow spinules or papillae that originate in fibrous layer.

Discussion.— The genus Spinospirifer Martynova differs from the other genera
assigned here in having a widely hinged growth form, a single row of spine bases
along each plica, and a weakly capillate micro-ornament. It is tentatively placed
here due to its spinose ornament and impunctate shell substance.

Stratigraphic Range. — Upper Devonian (Lower Famennian)-Lower Carbon-
iferous (Toumaisian).

Genera Included. — Gerkispir a Carter, 1983; Acanthospirina Schuchert and
LeVene, 1929; Oiosia Cooper and Dutro, 1982; Punctothyris Hyde, 1953; ?Spi-
nospirifer Martynova, 1961.

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Family Perissothyrididae Carter, new family

Diagnosis.— Subequally biconvex; transversely subovate; low concave interar-
eas present in both valves; fold moderately developed; sulcus weak, shallow,
poorly delimited; lateral slopes with weak simple ribbing; micro-ornament absent;
delthyrium very wide, partially occluded apically by “pseudodelthyrial plate”
(fused dental flanges?); dental adminicula absent; wide dorsally reflexed dental
flanges directed medially; large ctenophoridium supported by short median ridge;
crural bases medially directed, with ventrally reflexed medial flanges.

Stratigraphic Range.— Lower Carboniferous.

Genera Included. —Perissothyris Carter, 1967; Mongoliopsis Grunt, 1977.

Superfamily Spiriferoidea King, 1 846
[nom. correct. Carter, herein
( pro Superfamily Spiriferacea King, 1846, p. 28;
nom. transl. Schuchert, 1896, p. 333, ex Spiriferidae King, 1846, p. 28)]

Diagnosis. — Ventral beak incurved, ventral interarea low, usually apsacline;
hingeline denticulate except in genera with very narrow hingeline; fold and sulcus
usually ribbed; ctenophoridium present; prismatic shell layer present in some
genera.

Discussion. — Following Ivanova (1981), the impunctate paeckelmanellids are
placed in their own superfamily, the Paeckelmanelloidea. In the previous Treatise
classification, Pitrat (1965) placed the genera in this superfamily in the punctate
superfamilies Syringothrydoidea and Spiriferinoidea. Also, all punctate genera
have been removed from the Spiriferoidea and are now included in the Order
Spiriferinida.

This superfamily is severely restricted herein to include only families believed
to have been derived from a common ancestor in the Late Devonian. Besides the
paeckelmanellids the brachythyridids have been removed from the Spiriferoidea.
The brachythyridids are believed to have been derived from a Late Devonian
theodossiid ancestral stock that diverged from the line that gave rise to the Spi-
riferoidea and Paeckelmanelloidea. The Paeckelmanelloidea and Spiriferoidea are
judged to have arisen from the same indeterminate theodossiid lineage. This
hypothetical theodosiid ancestor may have been strophic, at least partially den-
ticulate, capillate or modified capillate, bearing dental adminicula, and carrying
the tendency to produce a prismatic shell layer.

Stratigraphic Range.— Upper Devonian (Famennian)-U pper Permian.

Family Spiriferidae King, 1 846
[Family Spiriferidae King, 1846, p. 28]

Diagnosis. — Outline variable; hingeline coarsely denticulate except for narrow
areas near delthyrium; delthyrial cover weakly developed or absent; lateral slopes
and fold-sulcus costate to costellate; micro-ornament capillate; short delthyrial
plate variably developed; dorsal adminicula or crural plates not developed; vas-
cular impressions absent or simple.

Discussion. —This family contains two of the earliest appearing members of the
Spiriferoidea, the late Famennian genera Paralellora Carter, 1974, of the Subfam-
ily Prospirinae, and E ohr achy thy ris Brice 1 97 1 , of the Subfamily Sergospiriferinae.
Eobrachythyris is most similar to the theodossioids in growth form and macro-
ornamentation and could be considered to be closest to a hypothetical ancestor
for this superfamily. Such a likely theodossioid ancestor might be Palaeospirifer

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Martynova and Sverlibova, 1968, except that it lacks any indication of a dentic-
ulate or partially denticulate hingeline.

Stratigraphic Range.— Upper Devonian (Famennian)-Upper Permian.

Subfamily Spiriferinae King, 1 846
[nom. transl. Waterhouse, 1968, p. 9,
ex family Spiriferidae King, 1846, p. 28]

Diagnosis. — Cardinal extremities extended in juveniles; outline variable in adults;
flanks and fold-sulcus with numerous costae that commonly bifurcate; fold usually
poorly delimited; dental adminicula usually developed, and when present, mod-
erately divergent.

Discussion. — The oldest representatives of this subfamily appear in the late
Middle Toumaisian. They were probably derived from a prospirin with a tendency
for bifurcation of the ribs such as Paralellora or Unispirifer. The earliest species
of the genus Spirifer, Spirifer gregeri Weller, 1914, first appears in the Middle
Toumaisian of North America. However, wide and narrow hinged species of the
genus Spirifer appear virtually simultaneously only slightly later in Western Al-
berta.

Stratigraphic Range. — Lower Carboniferous (Middle T ournaisian)-U pper
Permian.

Genera Included.— Spirifer {Spirifer) Sowerby, 1818; Spirifer ( Grandispirifer )
Yang, 1959; Spirifer {Mesochorispira) Carter, 1992; Ectochoristites Campbell,
1957; Larispirifer Enokjan and Poletaev, 1 986; Latispirifer Archbold and Thomas,
1985; Warsawia Carter, 1974.

Subfamily Prospirinae Carter, 1974
[Subfamily Prospirinae Carter, 1974, p. 680]

Diagnosis. — Cardinal extremities extended in juveniles; growth form variable
in adults; lateral slopes with relatively few, mostly simple costae; median sulcal
costa usually simple or bifurcating once; lateral sulcal costae usually simple, not
numerous, derived from sulcus-bounding costae; fold clearly delimited from lat-
eral slopes by bounding interspaces; dental adminicula usually well developed;
ventral umbonal callus commonly present.

Discussion. — The early growth stages of Andreaspira Abramov and Grigor’eva,
Austrochoristites Roberts, and Kasakhstania Beznosova are poorly known. Large
juveniles or young adults of these genera are strongly transverse as seen from
illustrations of the growth lines of mature adult specimens. In the absence of
definitive information concerning their early development, their assignment here
is tentative.

Stratigraphic Range.— Upper Devonian (Upper Famennian)-Upper Carbon-
iferous (Upper Serpukhovian, ?Moskovian).

Genera Included.— Prospira Maxwell, 1954; I Andreaspira Abramov and Gri-
gor’eva, 1986; ?A ustrochoristites Roberts, 1971; Finospirifer Ying, 1979; IKa-
sakhstania Beznosova, 1968; Kinghiria Litvinovich, 1966; Paralellora Carter,
1974; Subspirifer Shan and Zhao, 1980; Unispirifer Campbell, 1957 [= ILytha
Frederiks, 1924].

Subfamily Sergospiriferinae Carter, new subfamily

Diagnosis. — Cardinal extremities rounded in juveniles; outline variable in adults;
lateral slopes with few, simple or bifurcating costae; lateral sulcal costae usually

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simple, derived from bounding costae; fold usually well delimited by bounding
interspaces; dental adminicula well developed.

Discussion. — The nature of the hingeline and micro-ornament of Afghanospi-
rifer Plodowski are poorly known. If it should prove to be nondenticulate, then
it might be related to the Palaeochoristididae. The early growth stages of the type
species of E obr achy thy ris Brice are too poorly known to be certain that they are
rounded in all stages. If the cardinal extremities of juveniles of this species prove
to be extended, this genus should be reassigned to the Prospirinae.

Stratigraphic Range.— 1 Upper Devonian (Upper Famennian); Lower Carbon-
iferous (Lower T oumaisian)-U pper Carboniferous; ?Permian.

Genera Included. —lAfghanospirifer Plodowski, 1968; Anthracospirifer Lane,
1963 [=f?Plicatocyrtia Gauri, 1965]; Cancellospirifer Campbell, 1953; lEobrachy-
thyris Brice, 1970; Eochoristitella Qi, 1983; Sergospirifer Ivanova, 1952.

Subfamily Purdonellinae Poletaev, 1986
[nom. nov. Poletaev, 1986, p. 65
{pro Subfamily Munellinae Frederiks, 1924, p. 313)]

Diagnosis. —Outline brachythyridid; hingeline narrow; fold and sulcus mod-
erately to well developed, multicostate, not delineated by discontinuity in ribbing;
costae on lateral slopes often flattened, simple or subfasciculate; ventral interior
with delthyrial plate and diverging dental adminicula; vascular impressions in-
distinct.

Discussion. — The multicostate freely bifurcating ornament of this group suggests
that it is probably derived from the Spiriferinae by neotenous retention of an
ovate growth form throughout ontogeny. Mirifusella Carter is placed here with
considerable doubt. Its simple ribbing and sharply delimited fold are not usual
for this subfamily.

Stratigraphic Range. — Lower Carboniferous (Toumaisian)-Lower Permian.

Genera Included.— Purdonella Reed, 1944; Ala Nalivkin, 1979; Domokhotia
Abramov and Grigorieva, 1983; Eliva Frederiks, 1924; IMirifusella Carter, 1971;
Neomunella Ozaki, 1931; Podtsheremia Kalashnikov, 1966.

Family Choristitidae Waterhouse, 1968
[nom. transl. Ivanova, 1972, p. 40,
ex Subfamily Choristitidinae Waterhouse, 1968, p. 9]

Diagnosis. — Cardinal extremities rounded in juveniles, variable in large adults;
denticulation usually well developed; numerous simple or bifurcating costae usu-
ally present on entire surface; micro-ornament capillate; delthyrial plate absent;
distinctive vascular impressions commonly present, especially in younger genera.

Stratigraphic Range. — Lower Carboniferous-Permian .

Subfamily Angiospiriferinae Legrand-Blain, 1985
[Subfamily Angiospiriferinae Legrand-Blain, 1985, p. 574]

Diagnosis. — Dental adminicula short and close-set or absent; vascular impres-
sions ramiform, or weakly to moderately reticulate; micro-ornament finely can-
cel late or radially granulose, formed by weak capillae and fine, slightly lamellose
growth lines.

Discussion. — This subfamily first appears in the Visean of Eurasia and North
Africa and was undoubtedly derived from some member of the Sergospiriferinae
by the acquisition of short dental adminicula, distinct vascular impressions, and

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345

weakly capillate micro-ornament. The assignment of Quizhouspirifer Xian is ten-
tative because the vascular impressions are unknown.

Stratigraphic Range. — Lower Carboniferous (Visean)-Lower Permian.

Genera Included.— Angiospirifer Legrand- Plain, 1985; Anthracothyrina Le-
grand-Blain, 1984; Brachythyrina Frederiks, 1929 [=Anelasmina Semikhatova,
1939]; Elinoria Cooper and Muir-Wood, 1951; Eobrachythyrina Lazarev and
Poletaev, 1982; Prochoristitella Legrand-Blain, 1969; ? Quizhouspirifer Xian, 1982.

Subfamily Choristitinae Waterhouse, 1968
[Subfamily Choristitidinae Waterhouse, 1968, p. 9]

Diagnosis. — Dental adminicula close-set, subparallel; vascular impressions re-
ticulate.

Discussion.— Settedabania Abramov has an unusual muscle platform in the
ventral valve and lacks any indication of vascular markings.

Stratigraphic Range.— 1 Lower Carboniferous; Upper Carboniferous; ?Lower
Permian.

Genera Included. — Choristites Fischer de Waldheim, 1825 [= Alphachoristites
Gatinaud, 1949; IBetachoristites Gatinaud, 1949; Yatsengina Semikhatova, 1936];
Choristitella Ivanov and Ivanova, 1937; Parachoristites Barkhatova, 1968; ISet-
tedabania Abramov, 1970; Trautscholdia Ustritsky, 1967.

Subfamily Tangshanellinae Carter, new subfamily

Diagnosis. — Outline subovate with rounded cardinal extremities; dental and
dorsal adminicula absent; vascular impressions poorly developed or unknown.

Discussion. — The coarse simple lateral plicae of Capillispirifer Zhang are not
characteristic of this subfamily.

Stratigraphic Range.— Upper Carboniferous-Upper Permian.

Genera Included. — Tangshanella Chao, 1 929; Alphaneospirifer Gatinaud, 1 949
[=Semibrachythyrina Yang, 19 62]; ? Capillispirifer Zhang, 1983; Zhejiangospirifer
Liang, 1982.

Family Imbrexiidae Carter, 1992
[Family Imbrexiidae Carter, 1992, p. 327]

Diagnosis. — Outline transversely subquadrate; cardinal extremities extended in
juveniles, variable in adults; fold and sulcus well developed, often medially sub-
angular, flaring anteriorly in some genera; ventral interior with dental adminicula
and short delthyrial plate; ornament of moderately numerous simple or bifurcating
costae on both flanks and fold-sulcus; micro-ornament of weak capillae and reg-
ularly imbricate growth lamellae.

Discussion. — Carter (1992:328) speculated that this family was derived from
the Prospirinae because of similarities in macro- and micro-ornament between
prospirins and the earliest imbrexiid genus Fernglenia Carter.

Stratigraphic Range. — Lower Carboniferous (?Middle, Upper Toumaisian).

Genera Included. — Imbrexia Nalivkin, 1937; Fernglenia Carter, 1992; Tegu-
locrea Carter, 1992.

Family Trigonotretidae Schuchert, 1893
[nom. transl. Carter, herein,
ex Subfamily Trigonotretinae Schuchert, 1893, p. 156]

Diagnosis. —Usually strongly transverse in adult growth stage; fold and sulcus

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well developed; denticulate hingeline well developed; entire surface ribbed, often
plicate; lateral slopes commonly fasciculate.

Stratigraphic Range . — Lower Carboniferous-Upper Permian.

Subfamily Meospiriferinae Waterhouse, 1968
[Subfamily Neospiriferinae Waterhouse, 1968, p. 9]

Diagnosis. Weakly to strongly fasciculate; costae medium to fine, nearly uni-

form in size anteriorly; micro-omament weakly capillate, often lame Hose or im-
bricate; ventral interior with delthyrial plate.

Discussion. — The genus Rlasispirifer Kulikov is placed here tentatively on the
basis of having a few fasciculate plicae. In addition, its growth form and lack of
dental adminicula are not usual for this subfamily. The neospiriferins are likely
to have been derived from the Imbrexiidae by acquisition of fasciculate plicae on
the flanks.

Stratigraphic Range.— Upper Carboniferous-Upper Permian.

Genera Included. —Neospirifer Frederiks, 1924; Betaneospirifer G atinaud, 1949;
1 Rlasispirifer Kulikov, 1950; Cartorkiurn Cooper and Grant, 1976; Costatispirifer
Archbold and Thomas, 1985; Crassispirifer Archbold and Thomas, 1985; Cra-
tispirifer Archbold and Thomas, 1985; Fusispirifer Waterhouse, 1966 [= Trans-
vers aria Waterhouse and Gupta, 1983]; Gibhospirifer Waterhouse, 1971; Gypospi-
rifer Cooper and Grant, 1976; Imperiospira Archbold and Thomas, 1993;
Kaninospirifer Kulikov and Stepanov, 1975; Lepidospirifer Cooper and Grant,
1969; Pondospirifer Waterhouse, 1 9 7 8 ; Septospirifer W aterlio use, 1971; Tibetospi -
rifer Liu and Wang, 1990.

Subfamily Trigonotretinae Schuchert, 1893
[Subfamily Trigonotretinae Schuchert, 1893, p. 156]

Diagnosis. — Transverse; often thick shelled; fold and sulcus moderately devel-
oped; lateral slopes weakly to strongly plicate, variable fasciculate; finer ribbing
medium to coarse, often forming unequal ribs anteriorly; micro-omament cap-
illate and variably cancellate; ventral interior with dental adminicula, obscured
by thick apical, commonly bulbous, delthyrial callus.

Discussion.— Brachythyrinella Waterhouse and Gupta is tentatively assigned
here on the basis of its macro-omament, which is well illustrated in Thomas
(1971 :pk 1 9). Internally, it differs greatly from the other genera placed here, lacking
a delthyrial plate or umbonal callus. Sulciplica Waterhouse does not have sec-
ondary costae or costellae imposed on the plicae and thus differs from the other
genera in this subfamily.

Stratigraphic Range.— Lower Carboniferous (Yisean)-Upper Permian.

Genera Included. — Trigonotreta Koenig, 1825 [ - Grantonia Brown, 1953];
Aperispirifer Wat.erh.ouse, 1968; IBrachythyrinella Waterhouse and Gupta, 1978;
Frechella Legrand-Blain, 1986; ISulciplica Waterhouse, 1968.

Family Spiriferellidae Waterhouse, 1968
[nom. transl. Carter, herein,
ex Subfamily Spiriferellinae Waterhouse, 1968, p. 9]

[=Family Spiriferellidae Termier, Termier, Lapparant, and Martin, 1974, p. 136]

Diagnosis. — Lateral slopes plicate, coarsely costate, or fasciculate; fold usually
with median furrow; delthyrium commonly partially or completely occluded by
convex stegidial plates; denticulation poorly developed in narrow hinged genera;

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Carter et al. — Spiriferid Brachiopod Revision

347

micro-ornament pustulose, commonly capillate; ventral interior with strong short
dental adminicula; ventral umbonal region often greatly thickened by callus with

deeply impressed ventral muscle field.

Discussion. — Spiriferellaoides Lee, Gu, and Li is based on a single poorly pre-
served specimen. Although it is very difficult to evaluate, it may prove to be a
junior synonym of Spiriferella and for this reason its assignment here is questioned.
Tipispirifer Grant is truly enigmatic. Its growth form and macro-ornament of fine
costae are unusual for this family. It is assigned here on the basis of its pustulose
micro-ornament and stegidial cover of the delthyrium.

The other genera in this family form a close-knit group characterized by thick
ventral umbones and a pustulose micro-ornament. They were probably derived
from the Trigonotretinae by the acquisition of pustulose micro-ornament.

Stratigraphic Range. — Upper Carboniferous (Bashkirianj-Upper Permian (Ve-
dian).

Genera Included.— Spiriferella Chernyshev, 1902; Alispiriferella Waterhouse
and Waddington, 1982; Arcullina Waterhouse, 1986; Elivina Frederiks, 1924;
Eridmatus Branson, 1966; Plicatospiriferella Waterhouse and Waddington, 1982;
Rhombospirifer Duan and Li, 1985; ? Spiriferellaoides Lee, Gu, and Li, 1980;
Timaniella Barkhatova, 1968; ? Tipispirifer Grant, 1976.

Superfamily Paeckelmanelloidea Ivanova, 1972

[nom. correct. Carter, herein

( pro Paeckelmanellacea Ivanova, 1981, p. 22) nom. trans. Ivanova, 1981, p. 22,
ex Paeckelmanellidae Ivanova, 1972, p. 40]

Diagnosis. — Moderately to strongly transverse with maximum width at hinge-
line; ribs usually simple on flanks, but geologically younger forms sometimes with
bifurcations; fold and sulcus smooth or with median rib in sulcus, or more rarely,
multicostate; ventral interarea usually moderately to very high, often catacline to
slightly procline or weakly apsacline; hingeline denticulate; micro-ornament con-
sisting of capillae and anteriorly free growth lamellae; dental adminicula usually
present; vascular markings simple, radial; ctenophoridium present.

Discussion. — The origin of this superfamily is not certain, but it clearly began
in the Famennian. The earliest paeckelmanelloid, Strophopleura Stainbrook, ap-
peared in the early Upper Famennian, slightly before the earliest true spiriferoids
Parallelora or Eobr achy thy ris. The denticulate hingeline and capillate micro-
ornament point to a close relationship with the Spiriferoidea, probably a shared
ulbospiriferid ancestor.

Stratigraphic Range.— Upper Devonian-Upper Permian.

Family Strophopleuridae Carter, 1974
[nom. trans. Carter, herein, ex Strophopleurinae Carter, 1974, p. 677]

Diagnosis. — Ventral septum absent.

Stratigraphic Range.— Upper Devonian-Upper Permian.

Subfamily Strophopleurinae Carter, 1974
[Subfamily Strophopleurinae Carter, 1974, p. 677]

Diagnosis. — Small, with simple lateral ribs; sulcus smooth or with median rib.

Discussion. — Calvustrigis Carter is tentatively placed here because it lacks an
imbricate ornament, differing from all other genera assigned here. Although Iwai-
spirifer Tachibana is poorly known and a denticulate hingeline has not been

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established, its growth form, close-set dental adminicula, and imbricate-capillate
micro-ornament strongly suggest placement here.

Stratigraphic Range. — Upper Devonian (Fan lennian)- U pper Carboniferous
(Moscovian).

Genera Included.— Strophopleura Stainbrook, 1947; Acuminothyris Roberts,
1963; Avisyrinx Martinez Chacon, 1975; ICalvustrigis Carter, 1987; Cantabriella
Martinez Chacon and Rio Garcia, 1987; llwaispirifer T achibana, 1963; Voiseyella
Roberts, 1964 [=Amesopleura Carter, 1967].

Subfamily Bashkiriinae Nalivkin, 1979
[nom. trans. Carter, herein, ex Bashkiriidae Nalivkin, 1979, p. 143]

Diagnosis. ~ Small to medium size; lateral ribs simple; fold and sulcus ribbed;
ventral interarea moderately high to very high, often flattened, catacline to pro-
dine.

Discussion. — Denticulation in the genus Adminiculoria Waterhouse and Gupta
has not been established; otherwise, it could be assigned here confidently.

Stratigraphic Range. — Lower Carboniferous.

Genera Included.— Bashkiria Nalivkin, 1979; ? Adminiculoria Waterhouse and
Gupta, 1978; Celsifornix Carter, 1974; Fusella McCoy, 1844.

Subfamily Pterospiriferinae Waterhouse, 1975
[Subfamily Pterospiriferinae Waterhouse, 1975, p. 15]

Diagnosis. — Med ium sized to large; ribs on flanks simple or bifurcating, rarely
fasciculate; fold and sulcus smooth or with median rib, rarely costate; dental
adminicula short and divergent.

Stratigraphic Range.— Upper Carboniferous-Upper Permian.

Genera Included.— P ter ospirifer Dunbar, 1955; Alispirifer Campbell, 1961;
Haplospirifer Lee and Gu, 1976; Pteroplecta Waterhouse, 1978; Spiriferinaella
Frederiks, 1926; Xizispirifer Liang, 1990.

Family Paeckelmanellidae Ivanova, 1972
[Family Paeckelmanellidae Ivanova, 1972, p. 40]

Diagnosis. —With ventral medium septum; lateral ribs simple.

Stratigraphic Range. — Lower Permian-Upper Permian.

Subfamily Paeckelmanellinae Ivanova, 1972
[nom. trans. Waterhouse, 1975, p. 15,
ex Paeckelmanellidae Ivanova, 1972, p. 40]

Diagnosis. — With short divergent dental adminicula.

Stratigraphic Range.— Lower Permian-Upper Permian.

Genera Included.— Paeckelmanella Likharev, 1934; Darvasia Likharev, 1934;
Odontospirifer Dunbar, 1955.

Subfamily Scenesiinae Carter, new subfamily

Diagnosis. — With short apical delthyrial plate; dental adminicula absent.

Stratigraphic Range.— Fowtt Permian.

Genus Included.— Scenesia Cooper and Grant, 1976.

Superfamily Brachythyridoidea Frederiks, 1 924
[nom. trans). Carter, herein
{ex Brachythyrinae Frederiks, 1924, p. 316)]

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349

Diagnosis. — Outline ovate; hingeline narrow; cardinal extremities rounded in
all growth stages; lateral slopes with coarse, usually simple ribbing; fold-sulcus
weakly to moderately developed; denticulation not present in younger genera;
micro-ornament absent or weakly developed; dental adminicula, subdelthyrial
plate, and dorsal adminicula absent; delthyrium partially covered by thin deltidi-
um or stegidial plates.

Discussion. — The suppressed denticulation and lack of capillate micro-orna-
ment suggest that this group may have been derived independently from the
theodossioids or an intermediate ancestor by neoteny, with concommitant loss
of dental adminicula.

The origin of this superfamily is obscured by the paedomorphically simple
internal morphology of the genera included here. Also, the characteristic rounded
outline and general absence of micro-ornament of these genera could be construed
as either primitive or derived. Assuming that at least some of these characters
give an indication of ancestry, it seems possible that this group was derived from
a subovate, well-inflated form with a few coarse simple ribs on each lateral slope.
Because there is weak denticulation of the hingeline and obscurely capillate micro-
ornament in at least one of the geologically early genera, it seems most likely that
this group was derived from an offshoot of the same stock that produced the other
Late Paleozoic superfamilies of this order, namely, the Theodossioidea.

Stratigraphic Range. — Upper Devonian (Famennian)-Permian.

Family Brachythyrididae Frederiks, 1924
[nom. transl. et correct. Pitrat, 1965, p. H706,
ex Subfamily Brachithyrinae Frederiks, 1924, p. 316]

Diagnosis. — Primary median costa in sulcus absent.

Discussion.— Dalaia Plodowski was originally assigned by Plodowski (1968:
256) to the Reticulariidae, but it lacks spinose or pitted ornament and cannot be
assigned to either the Reticularioidea or Martinioidea. The micro-ornament of
Pustulosplica Waterhouse consists of discontinuous capillae and elongated fine
nodes or pustules, which is unknown in other members of this superfamily. This
micro-ornament is reminiscent of several martiniid genera and assignment of
Pustuloplica here is uncertain.

Stratigraphic Range.— Upper Devonian (Famennian)~Permian (?Kazanian).

Genera Included.— Brachythyris McCoy, 1844 [=Ovalia Nalivkin, 1937]; Ca-
thayspirina Liang, 1990; IDalaia Plodowski, 1968; Ella Frederiks, 1918; Meristo-
rygma Carter, 1974; ? Pustuloplica Waterhouse, 1968.

Family Skelidorygmidae Carter, new family

Diagnosis. — Sulcus with primary median costa, often with additional sulcal
costae that bifurcate from sulcus-bounding ribs.

Stratigraphic Range. — Upper Devonian (Famennian)-U pper Carboniferous
(Bashkirian).

Genera Included.— Skelidorygma Carter, 1974; Litothyris Roberts, 1971.

Suborder Delthyridina Ivanova, 1972
[Suborder Delthyridina Ivanova, 1972, p. 41]

Diagnosis. — Lateral slopes plicate, rarely costate; concentric lamellae, if present,
fimbriate, spinulose, or exopunctate; early forms with internal plates and septa,
tending to become pedomorphically simple in some geologically younger forms;
commonly with ctenophoridium.

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Stratigraphic Range.— Lower Silurian-Upper Permian.

Superfamily Delthyridoidea Phillips, 1841
[nom. transl. Ivanova, 1959, p. 56, nom. correct. Johnson, 1970, p. 184
(ex Delthyridae Phillips, 1841, p. 54)]

Diagnosis. — Biconvex, with fold and sulcus.

Stratigraphic Range. — Lower Silurian (Upper Llandoverian)-Middle Devonian
(Eifelian).

Family Delthyrididae Phillips, 1841
[nom. correct. Johnson, 1970, p. 184
(pro Delthyridae Phillips, 1841, p. 54)]

Diagnosis. — Small to medium size, pauciplicate; commonly with crural plates.
Stratigraphic Range. — Lower Silurian (Upper Llandoverian)-Middle Devonian
(Eifelian).

Subfamily Delthyridinae Phillips, 1 84 1

[nom. transl, nom. correct. Johnson, 1970, p. 184

(ex Delthyridae Phillips, 1841, p. 54)]

Diagnosis. — Ventral median septum present.

Stratigraphic Range.— Upper Silurian (Wenlockian)-Middle Devonian (Eife-
lian).

Genera Included. —Delthyris (Delthyris) Dalman, 1828; Delthyris ( Quadrifarius )
Fuchs, 1929; Ivanothyris Havlicek, 1957.

Subfamily Howellellinae Johnson and Hou, new subfamily

Diagnosis. — Ventral median septum lacking.

Stratigraphic Range. — Lower Silurian (Upper Llandoverian)-Middle Devonian
(Eifelian).

Genera Included. —Howellella (Howellella) Kozlowski, 1946; Howellella (Hys-
terohowellelld) Carls, 1985; Howellella (Iberohowellella) Carls, Meyn, and Ves-
permann, 1993; Acanthospirifer Menakova, 1 964; Aldanispirifer Alekseeva, 1967
[=Holcospirifer Bassett, Cocks, and Holland, 1976]; Howelloidea Su, 1980; Orien-
tospirifer Hou and Xian, 1975; Pseudokymatothyris Chen, 1979; Qiansispirifer
Yang, 1977; Rufispirifer Havlicek, 1987; Xenospirifer Hou and Xian, 1975.

Family Hysterolitidae Termier and Termier, 1949
[nom. transl. Johnson and Hou herein
(ex Hysterolitinae Termier and Termier, 1949, p. 95)]

Diagnosis. — Medium to large size; fold and sulcus smooth or plicate; flanks
commonly multiplicate; crural plates present or absent.

Stratigraphic Range. —Lower Devonian (Upper Lochkovian)-Middle Devo-
nian (Lower Givetian).

Subfamily Hysterolitinae Termier and Termier, 1949

[Hysterolitinae Termier and Termier, 1949, p. 95]
[=Paraspiriferinae Pitrat, 1965, p. H684]

Diagnosis. — Fold and sulcus smooth; mostly lacking crural plates.

Stratigraphic Range. — Lower Devonian (Upper Lochkovian)-Middle Devo-
nian (Eifelian).

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Genera Included. —Hyster elites Schlotheim, 1820; Antispirifer Williams and
Breger, 1916; Arduspirifer, Mittmeyer, 1972; Australospirifer Caster, 1939;
Brachyspirifer (Brachyspirifer) Wedekind in Salomon, 1926; Brachyspirifer (Toro-
sospirifer ) Gourvennec, 1989; Costellispirifer Boucot, 1973 [=Concinnispirifer
Boucot, 1975]; Dixonella Gourvennec, 1989; Dyticospirifer Johnson, 1966; Eu-
ryspirifer Wedekind in Salomon, 1926 [=Rhenospirifer Mittmeyer, 1972]; Oto-
spirifer Hou and Xian, 1975; Paraspirifer Wedekind, 1 926; Patriaspirifer Johnson,
1995; Rostrospirifer Grabau, 1931 [= Neodelthyris Hou, 1963]; Trigonospirifer
Wang, Rong, and Chen, 1987; Xinjiangospirifer Hou and Zhang, 1983.

Subfamily Fimbrispiriferinae Pitrat, 1965
[nom. transl Wang, 1979, p. 77
(ex Fimbrispiriferidae Pitrat, 1965, p. H687)]

Diagnosis. — Fold, sulcus, and flanks plicate; crural plates present or absent.

Discussion. —Boucot (1975) proposed Vandercammenina and Struveina as Old
World-Realm genera of the Acrospiriferinae, but they are here excluded from that
subfamily because of their fimbriate ornament. We regard Fimbrispirifer, a Middle
Devonian genus from eastern North America, as a descendant of Struveina by
late migration of that stock, following a pattern demonstrated for the genus Para-
spirifer (Johnson, 1979).

Stratigraphic Range. — Lower Devonian (Pragian)-Middle Devonian (Lower
Givetian).

Genera Included.— Fimbrispirifer Cooper, 1942; Struveina Boucot, 1975; Van-
dercammenina Boucot, 1975.

Family Acrospiriferidae Termier and Termier, 1949
[nom. transl. Johnson and Hou herein
(ex Acrospiriferinae Termier and Termier, 1949, p. 96)]

Diagnosis.— Medium to large size, transverse, capillate with fila; crural plates
present or absent.

Discussion.— A peculiarity of genera in this family is the presence of fine or
prominent capillae together with a concentric ornament of fila. Examples are
Acrospirifer primaevus as illustrated by Vandercammen (1963:pl. 1, fig. 19, pi. 2,
fig. 1 1), Acrospirifer fallax as illustrated by Jahnke ( 1 97 1 :pl. 6, fig. 10), and Maui-
spirifer hectori as illustrated by Allan (1947:pl. 62, fig. 6). Capillae are so well
developed on Multispirifer that it was originally presumed to be an eospiriferid
(Kaplun, 1 96 1 :88). The capillae of Costispirifer caused Ivanova (1972) to suppose
that it was derived from the eospiriferids. Of particular interest is the ornament
of Euryspirifer Wedekind, 1926. Vandercammen (1963:pl. 8, fig. 6) and Jahnke
( 1 97 1 :pl. 7, fig. 4c) both show an ornament of capillae interrupted by fila. This
type of ornament is especially well represented by Euryspirifer atlanticus, as il-
lustrated by Boucot (1973:pl. 17, fig. 9).

Stratigraphic Range. — Lower Devonian (Upper Lochkovian)-Middle Devo-
nian (Eifelian).

Subfamily Acrospiriferinae Termier and Termier, 1949
[Acrospiriferinae Termier and Termier, 1949, p. 96]

Diagnosis. — Flanks strongly plicate, delthyrium lacking apical deltidium; dental
plates strong or partly buried in umbonal callus; ctenophoridium on floor of
notothyrial chamber attached to myophragm, or elevated.

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Stratigraphic Range.— Lower Devonian (Upper Lochkovian-Emsian).

Genera Included. —Acrospirifer Helmbrecht and Wedekind, 1923; Mauispirifer
Allan, 1947; Xerospirifer Havlicek, 1978.

Subfamily Costispiriferinae Termier and Termier, 1949
[Costispiriferinae Termier and Termier, 1949, p. 98]

Diagnosis. — Multiplicate, with narrow interspaces; ventral interarea low, trap-
ezoidal; broad delthyrium with apical deltidium; fold and sulcus smooth or plicate;
flanks with simple plications; short, widely spaced dental plates, partly buried in
umbonal callus; sockets strong, simple, curved; ctenophoridium not built up.

Discussion. — Although Cumberlandina was originally assigned to the Mucrospi-
riferinae by Boucot (1975:371), that assignment cannot be sustained because
Cumberlandina lacks lamellose concentric ornament. Except for the lack of pli-
cations on the fold and sulcus, Cumberlandina closely resembles Costispirifer.

Stratigraphic Range. — Lower Devonian (Pragian).

Genera Included. — Costispirifer Cooper, 1942; Cumberlandina Boucot, 1975.

Family Elymospiriferinae Johnson and Hou, new subfamily

Diagnosis. — Fold and sulcus plicate; flanks with bifurcating plications.

Stratigraphic Range. — Lower Devonian (Pragian)-Middle Devonian (Eifelian).

Genera Included. —Elymospirifer Wang, 1974; Borealispirifer Hou and Su, 1993;
Multispirifer Kaplun, 1961; Perry spirifer Jones and Boucot, 1983.

Family Cyrtinopsidae Wedekind, 1926
[nom. transl. Boucot, 1957, p. 38
(ex Cyrtinopsinae Wedekind in Salomon, 1926, p. 198)]

Diagnosis. — Frilly growth lamellae, with interrupted capillae; ctenophoridium
lacking or, in younger genera, rudimentary.

Stratigraphic Range.— Upper Silurian (Wenlockian)=Middle Devonian (Give-
tian).

Subfamily Cyrtinopsinae Wedekind, 1926
[Cyrtinopsinae Wedekind in Salomon, 1926, p. 198]
[=Kozlowskiellininae Boucot, 1958, p. 1031
(nom. correct. Kozlowskiellinae Boucot, 1957, p. 317)]

Diagnosis. — With ventral median septum.

Stratigraphic Range.— Upper Silurian (Wenlockian)-Middle Devonian (Give-
tian).

Genera Included. — Cyrtinop sis Scupin, 1896; Kozlowskiellina Boucot, 1958;
Megakozlowskiella Boucot, 1957; Plicocyrtina Havlicek, 1956.

Subfamily Araspiriferinae Johnson, new subfamily

Diagnosis. — Without ventral median septum.

Stratigraphic Range. — Upper Silurian ( W enlockian-Ludlo vian) .

Genera Included. —Araspirifer Havlicek, 1987; Boucotinskia Brunton and Cocks,
1967.

Family Mucrospiriferidae Boucot, 1959
[nom. transl. Johnson and Carter herein
(ex Mucrospiriferinae Boucot, 1959. p. 745)]

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Diagnosis. — Lamellose growth lines; ctenophoridium without crural plates.
Stratigraphic Range.— Middle Devonian (Givetian)~Lower Carboniferous (Vi-
sean).

Subfamily Mucrospiriferinae Boucot, 1959
[Mucrospiriferinae Boucot, 1959, p. 745]

Diagnosis. — Transverse, multiplicate; dental plates present or absent; without
median septum.

Stratigraphic Range. — Middle Devonian (Gi vetian)-U pper Devonian (Famen-
nian).

Genera Included. —Mucrospirifer Grabau, 1931 [=Lamellispirifer Nalivkin, 1937;
Khinganospirifer Su, 1976]; Apousiella Carter, 1972 [=Bouchardopsis Mailleux,
1933]; Eleutherokomma Crickmay, 1950 [=Dmitrispirifer Liashenko, 1973;
Sculptospirifer Su, 1980]; Sulcatospirifer Maxwell, 1954.

Subfamily Tylothyridinae Carter, 1972
[Tylothyridinae Carter, 1972, p. 730]

Diagnosis. — Variable outline and moderately numerous lateral costae; radial
ornament apparently lacking; with dental plates and median septum.

Stratigraphic Range. — Middle Devonian (Upper Givetian)-Lower Carbonif-
erous (Visean).

Genera Included. — Tylothyris North, 1920 [= Welleria Mailleux, 1931]; Tex-
athyris Carter, 1972.

Superfamily Reticularioidea Waagen, 1883
[nom. correct. Gourvennec and Carter, herein
( pro Reticulariacea Waagen, 1883, nom. transl. Pitrat, 1965, p. H717,
ex Subfamily Reticulariinae Waagen, 1883, p. 538)]

Diagnosis. — Subequally biconvex; outline generally transversely subovate to
equidimensional; hinge line short, lateral extremities rounded; fold and sulcus
generally present, commonly weakly developed; ribbing absent or weak, rarely
multicostate; micro-ornament often lamellose, commonly with fine spines or gran-
ules.

Stratigraphic Range.— Silurian (Upper Llando very)-U pper Permian.

Family Reticulariidae Waagen, 1883
[nom. transl. Ivanova, 1959, p. 56,
ex Subfamily Reticulariinae Waagen, 1883, p. 538]

Diagnosis. — Lateral slopes commonly smooth, rarely plicate; fold and sulcus,
if present, smooth; micro-ornament concentrically arranged, consisting of growth
lamellae and generally uniramous spinules or fine tubercles; dental adminicula
generally present, true ventral septum absent, median ridge or myophragm present
in some genera.

Discussion. — This diagnosis is based on the assumption that Reticularia lacks
a true ventral septum but, rather, possesses a median ridge or euseptoid, as
described and illustrated by George (1932:552). If this is actually not the case,
then the following classification is incorrect and we would be compelled to reassign
all of the nonseptate genera to a different family and revise the composition of
the Xenomartiniidae.

Stratigraphic Range. — Silurian (Upper Llando very )-Upper Permian.

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Subfamily Reticulariinae Waagen, 1883
[Subfamily Reticulariinae Waagen, 1883, p. 538]

Diagnosis. — Generally pauciplicate; lacking delthyrial plates, delthyrial ridges
or any kind of apical thickening; commonly lacking crural plates.

Discussion. —Havlicek erected the genus Undispirifer in 1957 and designated
S. undiferus Roemer 1 844 as type species. He pointed out the presence of a well-
developed fold and sulcus and low lateral plications, which are commonly only
perceptible anteriorly. In 1959, he added “basal plates do not meet the floor of
the valve” and “without subdelthyrial plate” (in the comparison with Eoreticu-
laria). In 1967, Vandercammen gave illustrations of the original material of Roe-
mer, considering undiferus and gerolsteinites Steininger, 1 853, as synonyms (form
I and II of the species undiferus, p. 6). His figure shows a specimen labeled
“Original Sp. undiferus F. Roemer” but Vandercammen indicated that this spec-
imen probably is not the type, which was apparently lost (there are effectively
some differences with the original illustration by Roemer). The origin of the label
is unknown. This specimen seems closely related to the type of S. curvata undulata
Roemer, 1844 (pi. 4, fig. 5a, b) which was later designated as the type of S.
gerolsteinensis by Steininger (1853). Recently, Struve (1990) erected the genus
Gerolsteinites, with Spirifera gerolsteinensis Steininger as the type species, and
which is distinguished from Undispirifer by: 1) a well-marked fold and sulcus, 2)
plicae expressed from apex to commissure, and 3) a high ventral interarea. These
arguments do not justify the erection of a new genus because the original diagnosis
of Undispirifer states that there is a well-developed sulcus/fold and lateral plica-
tions are often (but not necessarily) only perceptible anteriorly. In addition, the
original illustration by Roemer (1844:pl. 4, fig. 6a-c) shows undiferus with a well-
developed, if not high, ventral interarea, not far in its form from that of S.
gerolsteinensis (=S. curvatus undulata). It is clear that the characters of the genus
Undispirifer show some variability, but the erection of a new genus does not
appear as a necessity ( Gerolsteinites at most should have subgeneric rank). In any
case the problem of the type is not solved. Some other features illustrate the
variability in this genus. According to Havlicek (1959:254), Undispirifer is devoid
of a delthyrial plate. However, some authors have mentioned such a plate, fol-
lowing Vandercammen (1957:6), who indicated that a delthyrial plate is present
but difficult to identify. There is a possibility that the specimen described by
Vandercammen belongs to another genus, e.g., Rhenothyris, because his descrip-
tion of the species states that some large mature specimens are smooth (1957:5).
Otherwise, although Undispirifer generally lacks crural plates and possesses only
plate-like crural bases (Havlicek, 1 959:254), it seems that in some specimens these
plates join the floor of the valve (Johnson, 1974:pl. 10, fig. 4). The preceding
discussion shows that a revision of the species undiferus and the designation of
a neotype are desirable. Because there is still doubt about the presence or absence
of a delthyrial plate, the genus is assigned here with a question mark.

The subgenus Nakazatothyris Minato and Kato, 1977 is also placed in syn-
onymy of the genus Undispirifer for reasons comparable to the ones mentioned
above: its characters (few plicae better expressed in the vicinity of the fold/sulcus
and near the apex) do not justify the erection of a (sub-)genus because they fall
within a reasonable variation of the genus Undispirifer. Undispirifer aides Xian is
poorly known and its status is not clear.

Stratigraphic Range. — Lower Devonian (Upper Pragian)-Upper Permian.

Genera Included.— Reticular ia McCoy, 1844; IGeorgethyris Minato, 1953;

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Parareticularia Lee and Gu, 1976; Squamularia Gemmellaro, 1899; lUndispirifer
Havlicek, 1957 [=Nakazatothyris Minato and Kato, 1977] [=Gerolsteinites Struve,

1990]; ? Undispirifer aides Xian, 1978.

Subfamily Reticulariopsinae Gourvennec, new subfamily

Diagnosis.— Generally pauciplicate; lacking delthyrial plates, dethyrial ridges
or any kind of apical thickening; dental plates, crural plates and ctenophoridium
present.

Discussion. — The serial sections accompanying the original diagnosis of Pam -
quadrithyris are aberrant and probably erroneous. No comments are given by the
author about the cardinal process, but the genus is compared with Tinge l la (= Re-
ticulariopsis), which possesses a ctenophoridium. After Hou, the two genera are
probably synonyms.

Vandercammen (1958) does not recognize the genus Reticulariopsis, which he
considers as insufficiently described, but he accepts Tingella Grabau, in which he
places R. dereimsi, the type species of Reticulariopsis. Vandercammen assigned
two other species to Tingellas ( reticularioides and concentrica). The material de-
scribed under the name of concentrica by Vandercammen has since been recog-
nized as Gerothyris eifliana, while T. dereimsi (sensu Vandercammen) has been
found only in Belgium and differs from the Spanish species described by Oehlert
and Oehlert (1901), which lacks a delthyrial plate. Furthermore, the type species
of Tingella , T. reticularioides, apparently also lacks a delthyrial plate. Thus Tin-
gella is a junior synonym of Reticulariopsis and the interpretation of Vandercam-
men (delthyrial plate present) is to be rejected (this latter material probably belongs
to Rhenothyris).

Stratigraphic Range. — Silurian (Wenlock)-Middle Devonian (Givetian).

Genera Included. — Corylispirifer Gourvennec, 1989; Kymatothyris Struve, 1970
[=Fallaxispirifer, Su 1976]; Mariaspirifer Cherkesova, 1991; IParaquadrithyris
Yang, 1983; Padvenia Breivel and Breivel, 1988; Proreticularia Su, 1990; Prosse-
rella Grabau, 1910; Reticulariopsis Frederiks, 1916 [=Tingella Grabau, 1931];
Yeothyris Struve, 1992.

Subfamily Rhenothyridinae Gourvennec, new subfamily

Diagnosis. — With delthyrial plate, ridge or ventral apical thickening; dental
plates and ctenophoridium present.

Discussion. — Rhenothyris does not possess a true delthyrial plate but only clearly
expressed subdelthyrial plates or ridges on internal molds. The definition of the
genus Eohowellella is imprecise and not in accordance with the insufficient illus-
trations. For example, the “ventral septum” recorded in the diagnosis is absent
on the serial sections; if such a septum is really present, Eohowellella would better
be assigned to the Quadrithyridinae. The comparison with Howellella suggests
that a ctenophoridium is present although it is not reported in the diagnosis. The
mention of a “finely punctate shell” in the description is probably erroneous(?);
such a character would remove Eohowellella from the Reticularioidea. For all
these reasons, Eohowellella is assigned here with doubt.

The ctenophoridium is not reported in the diagnosis of Nordella and does not
appear in serial sections, but the comparison with Elita suggests close analogies
in the internal structures, and we admit to the presence of a ctenophoridium.
Should this not be the case, Nordella would be better placed in the Eoreticulariinae.

Puanospirifer is poorly known; many characters have been “estimated” or ex-

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trapolated from the author’s comparison with Reticulariopsis (particularly for
internal structures).

The holotype, paratypes, and figured specimens of Spirinella were destroyed
by fire in 1953. Strusz (1984) has given a new diagnosis and good illustrations of
topotype material, but failed to designate a neotype. We propose to designate the
specimen “SUP 55596” (Yass Formation, New South Wales), figured in Strusz
( 1 984:fig. 1 8 A-D) as the lectotype for Spirinella caecistriata Johnston (type species
of the genus Spirinella). Apparently, following the serial sections given by Johnston
(1941, fig. 2), Spirinella possesses incipient delthyrial plates or ridges. Neverthe-
less, these structures are poorly expressed and they are not reported by Strusz
(1984), who revised the species. Consequently, this genus is assigned here with
some doubt. If delthyrial plates and an apical thickening are lacking in Spirinella,
the Reticulariopsinae would be a better receptacle for this genus.

Stratigraphic Range.— Silurian (?Upper Llandovery, Lower Ludlow)-Middle
Devonian (Eifelian, ?Lower Frasnian).

Genera Included. —Rhenothyris Struve, 1970; Deltospirifer Wang and Rong,
1986; lEohowellella Lopushinskaja, 1976; Gerothyris Struve, 1970; Grebenella
Modzalevskaia and Besnosova, 1992; INordella Liashenko, 1973; Pseudoundispi-
rifer Zhang, 1987; IPuanospirifer Jiang, 1978; ISpirinella Johnston, 1941 [=Ec-
tatoglossa Chu, 1974]; Warrenella (Warrenella) Crickmay, 1953 [=Minatothyris
Vandercammen, 1957]; Warrenella (Warrenellina) Brice, 1982.

Subfamily Obesariinae Gourvennec, new subfamily

Diagnosis.— Smooth; lacking dental plates and ctenophoridium; apical thick-
ening in the ventral valve.

Discussion. — Quasimartinia occasionally possesses a sinus on the dorsal valve,
which is quite uncommon in the Reticularioidea.

In the original diagnosis of Uexothyris, no mention is made of a ctenophoridium,
so we consider it to be lacking. Nevertheless, Struve (1992:574) placed this genus
in the Tribe Gerothyridini with Gerothyris and “ Minatothyris both of which
possess a ctenophoridium. In the description of the species assigned to Uexothyris,
the cardinal process is not taken into consideration and no illustration is given.
If Uexothyris should prove to possess a ctenophoridium, it should be rejected
from the Obesariinae and attached to the Rhenothyridinae, hence the question
mark in our assignment.

Echinocoeliopsis is assigned here because of its ornament and lack of dental
plates, but it is apparently devoid of an apical thickening. Another peculiar feature
of this genus is the presence of a bifid shaft supporting the cardinal process. In
some respects (size, form, mode of folding, bifid cardinal process) this genus evokes
the Ambocoeliidae, particularly Metaplasia and allied genera. Otherwise, Echino-
coeliopsis is only known by a few very small specimens.

Stratigraphic Range.— Silurian (Upper Wenlock)~Middle Devonian (Eifelian).

Genera Included. — Obesaria Havlicek, 1957; Alaskospira Kirk and Amsden,
1952 [=Proreticularia Havlicek, 1957]; Quasimartinia Havlicek, 1959 [= Candi -
spirifer Havlicek, 1971]; ? Uexothyris Struve, 1992; ? Echinocoeliopsis Hamada,
1968.

Subfamily Eoreticulariinae Gourvennec, new subfamily

Diagnosis. — Smooth; with cardinal platform or septalium; ctenophoridium ab-
sent.

Discussion.— A mistake occurs in the translation to English of the work of

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357

Nalivkin (1930: 198) concerning Eoreticularia : “spines sometimes absent” should
read “spines always absent” as is evident from the rest of the description. Eoreticu-
laria is distinguished from both Reticular iop sis and Reticularia by the lack of
spines. Havlicek (1971) notes the presence of radially aligned granules and the
absence of marginal spines. The sections given by this author (1959:160, 163“
164) show a septalium and delthyrial plate, but a ctenophoridium is absent.

Stratigraphic Range. — Silurian (Wenlock)-Upper Devonian (Frasnian).

Genera Included. —Eoreticularia Nalivkin in Frederiks, 1924; Chnaurocoelia
Johnson, Boucot, and Murphy, 1976; Vadum Strusz, 1982.

Family Xenomartiniidae Havlicek, 1971

[Xenomartiniidae Havlicek, 1971, p. 24]

Diagnosis.— With ventral median septum.

Discussion. —In his conception of the family, Havlicek emphasized the lack of
dental plates (1971 :23). Here we would rather emphasize the presence of a ventral
septum, which leads us to include the subfamilies Bojothyridinae and Quadri-
thyridinae and reject the Ohesaria group. Close affinities exist between the Obesari-
inae and Xenomartiniidae, so that the latter probably were derived from the
Obesariinae by acquisition of a ventral median septum.

Stratigraphic Range.— Upper Silurian-Middle Devonian.

Subfamily Xenomartiniinae Havlicek, 1971
[nom. transl. Gourvennec herein (ex Xeonmartiniidae Havlicek, 1971, p. 24)]

Diagnosis.— Smooth; lacking dental plates and ctenophoridium.

Discussion. — In 1953, Havlicek described the genus Xenomartinia (type species
X. monosepta) including smooth forms with reticulate ornament and a ventral
septum but lacking dental plates. In the same year Minato proposed the genus
Sinothyris (type species R. maureri) with the following definition: “without dental
plates but with median septum in the ventral valve.” This diagnosis corresponds
to that of Xenomartinia, and thus the two genera would appear to be synonyms.
On the other hand, Minato’s interpretation of the species maureri is clearly er-
roneous (Pitrat, 1965:H717), and this leads to some confusion in the definition
of the genus Sinothyris. Pitrat placed Sinothyris in the synonymy of Reticularia
with some doubt, but we now know that Reticularia possesses a myophragm or
euseptoid rather than a true septum. The real question is one of priority. After
inquiry, we have established that the work by Minato was published on September
30, 1953, while that of Havlicek was published in February 1953. Thus, Sinothyris
is a junior synonym.

Stratigraphic Range. — Lower (Pragian)-Middle (Eifelian) Devonian.

Genera Included. —Xenomartinia Havlicek, 1953 [^Sinothyris Minato, 1953].

Subfamily Bojothyridinae Havlicek, 1990
[nom. transl. Gourvennec herein
(ex Bojothyrididae Havlicek, 1990, p. 186)]

Diagnosis. — With dental plates converging to median septum, commonly pro-
ducing spondylium or spondylium-like structure.

Discussion. — Although it was first described as lacking dental plates (Havlicek,
1 959:244), serial sections in Quadrithyrina apparently show tracks of rudimentary
dental plates (or ridges) converging towards a septum. Thus, it is assigned here
to the Bojothyridinae. On the other hand, the reduced size of these plates and the

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apparently smooth cardinal process suggest strong affinities with the Xenomar-
tiniinae. The same applies for Uralospirifer in which the dental plates are also
reduced but seem somewhat more prominent than in Quadrithyrina. The two
genera have an almost identical internal structure as pointed out by Havlicek
(1959:1 42). Uralospirifer and Altajella are also very close externally, but the latter
possesses a spondylium and cranial plates and lacks an apical thickening of the
shell. The genus Altajella can be seen as a possible ancestor of both the Bojothyridi-
nae and Xenomartiniinae.

Stratigraphic Range.— Upper Silurian (Upper Ludlow)-Middle (Eifelian) De-
vonian.

Genera Included.— Bojothyris Havlicek, 1959; Altajella Kulkov, 1962; IQuad -
rithyrina Havlicek, 1959; Spondylothyris Su, 1980.

Subfamily Quadrithyridinae Gourvennec, new subfamily

Diagnosis. — With divergent or subparallel dental plates.

Discussion. — The commissure of Quadrithyris is sometimes strongly uniplicate.
The crural plates are apparently lacking, but in some species (e.g., Q. rohusta, Q.
orba) small incipient crural plates are discernible on the bottom of the valve in
serial sections. The redefinition of the genus by Boucot (1962) differs somewhat
from the original definition. Some elements of this redefinition (e.g., the presence
of crural plates) have been accepted in our diagnosis.

Stratigraphic Range.— Upper Silurian (Upper Wenlock)-Middle Devonian
(Givetian).

Genus Included. — Quadrithyris Havlicek, 1957.

Family Thomasariidae Cooper and Dutro, 1982
[Thomasariidae Cooper and Dutro, 1982, p. 102]

Diagnosis. — Small, hemipyramidal shells, with strong, long dental plates and a
pair of conjunct apical plates. Dorsal interior with ctenophoridium, lacking crural
plates. Surface with growth lines and fine spines.

Stratigraphic Range.— Upper Devonian (Frasnian).

Genus Included. — Thomasaria Stainbrook, 1945.

Family Elythidae Frederiks, 1924
[nom. transl. Pitrat, 1965, p. H721,
ex Subfamily Elythinae Frederiks, 1924, p. 304]

Diagnosis.— Lateral slopes smooth or with low plications; fold and sulcus, if
present, usually weakly developed; micro-ornament of fine biramous spines.

Discussion. — The development of biramous spines in this family is unique or
synapomorphous and appears early in the history of the superfamily.

Stratigraphic Range. —Middle Devonian-Upper Permian.

Subfamily Elythinae Frederiks, 1924
[Subfamily Elythinae Frederiks, 1 924, p. 304]

Diagnosis. — Spines biramous but not elaborate; ventral interior with dental
adminicula and median ridge; dorsal interior with ctenophoridium.

Stratigraphic Range.— Lower Devonian-Lower Carboniferous.

Genera Included.— E lit a Frederiks, 1918; Kitakamithyris Minato, 1951.

Subfamily Martinothyridinae Carter, new subfamily

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Diagnosis. — Ventral interior with dental adminicula; ventral median ridge ab-
sent; spines elaborate.

Stratigraphic Range. —Lower Carboniferous-Lower Permian.

Genera Included. — Martinothyris Minato, 1953; Latiplecus Lee and Gu, 1976;
Orenburgella Pavlova, 1969.

Subfamily Phricodothyridinae Caster, 1939
[nom. correct. Carter, herein,
pro Subfamily Phricodothyriinae Caster, 1939, p. 145]

Diagnosis. — Lacking dental adminicula and ventral median ridge; spines elab-
orate.

Stratigraphic Range. — Lower Carboniferous-Upper Permian.

Genera Included.— Phricodothyris George, 1932 [=Wajkuria Ustritsky, 1963;
Condrathyris Minato, 1953; Neophricodothyris Likharev, 1934]; Astegosia Cooper
and Grant, 1969; Budarina Jing and Sun, 1981; Nehenothyris Minato, 1953;
Permophricodothyris Pavlova, 1965.

Subfamily Toryniferinae Carter, new subfamily

Diagnosis. — Dental adminicula and low median ridge present; ctenophoridium
absent; spines elaborate.

Stratigraphic Range. — Lower Carboniferous (Toumaisian)-Permian.

Genera Included. — Torynifer Hall and Clarke, 1893; Plicotorynifer Abramov
and Solomina, 1970; Spirelytha Frederiks, 1924; Stepanoviina Zavadovsky, 1968;
Taimyrella Ustritsky, 1963; Toryniferella Weyer, 1967.

Subfamily Anomaloriinae Cooper and Grant, 1976
[nom. transl. Carter, herein

{ex Family Anomaloriidae Cooper and Grant, 1976, p. 2260)]

Diagnosis. — Delthyrium covered with convex psuedodeltidium; ventral interior
with converging thickened dental flanges that fuse apically; dorsal interior with
notothyrial flanges fused to inner socket ridges that partially obscure sockets
medially.

Stratigraphic Range. — Permian.

Genera Included. —Anomaloria Cooper and Grant, 1 969; Zhinania Liang, 1 990.

Order Spiriferinida Ivanova, 1972
[nom. trans. Carter and Johnson, herein
{ex Suborder Spiriferinidina Ivanova, 1972, p. 41)]

[= Order Spiriferinida Cooper and Grant, 1976, p. 2666]

Diagnosis. —Hingeline strophic; commonly transverse and biconvex; flanks
ribbed, rarely smooth; ventral valve inflated and thicker than dorsal valve; ventral
interarea commonly well developed; brachidium, where present, spiraliform, spi-
ralia directed laterally or posterolaterally; jugum generally present; shell punctate.

Discussion. — Punctate spiriferids first appear near the Silurian-Devonian
boundary as the distinctive small hemipyramidal genus Cyrtina Davidson. The
growth form and complex internal structures of this genus indicate that it is
morphologically highly specialized and may not reflect the ancestral stock from
which this large group is derived. The immediate ancestors of the punctate spi-
riferids must have been strophic biconvex punctate orthids or impunctate spirifer-

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ids with ventral sulcus and dorsal fold. Lacking recognizable antecedents, whether
or not these close ancestors were punctate or impunctate cannot be demonstrated.

The Suborder Cyrtinidina comprises three superfamilies, the Cyrtinoidea, Sues-
sioidea, and Spondylospiroidea, all characterized by having spondylia or related
adductor-raising platforms. The Suessioidea were derived from the cyrtinoids in
the Early Carboniferous, but the Carboniferous and Permian record is poor. The
suessioids survived the Permian extinction event and flourished during the Middle
and Late Triassic, when they became extinct. Sometime in the Early or very early
Middle Triassic a stock of suessioids, the Spondylospiroidea, developed a unique
crenulated hingeline. This short-lived group also disappeared in the Late Triassic.

The other punctate suborder, the Spiriferinidina, appeared in the Late Devonian
as two distinct lineages, the superfamilies Pennospiriferinoidea and Syringothyrid-
oidea, both derived from the cyrtinids at about the same time. These groups
secondarily evolved a spiriferid growth form and discrete dental adminicula in
the ventral valve. The syringothyridoids attained large size and survived until the
Permian extinction event. The Pennospirinoidea developed a strong ventral me-
dian septum and became highly diverse throughout the Permian, but disappeared
in the early Triassic, giving rise in the Middle Triassic to the last of the punctate
superfamilies, the Spiriferinoidea. The latter group acquired its rounded growth
form and reduced radial ornament, reminiscent of the reticularioids, by neoteny.

Stratigraphic Range. — Lower Devonian (Lower Lochkovian)-Lower Jurassic.

Suborder Cyrtinidina Carter and Johnson, new suborder

Diagnosis. — Lateral slopes plicate or smooth; ventral valve high, subconical or
hemipyramidal in early forms, variable in later ones; delthyrium commonly cov-
ered in early genera, variably covered in late genera; ventral interior commonly
with elevated adductor attachment structures.

Stratigraphic Range.— Lower Devonian-Lower Jurassic.

Superfamily Cyrtinoidea Frederiks, 1911
[nom. transl. Johnson, 1966, p. 177
(ex Cyrtininae Frederiks, 1911)]

Diagnosis. — Ventral valve hemipyramidal, with high interarea; dorsal valve flat
or weakly convex; nonfimbriate and noncapillate; ventral median septum.

Stratigraphic Range.— Lower Devonian (basal Lochkovian)-Lower Carbonif-
erous (Visean).

Family Cyrtinidae Frederiks, 1911
[nom. transl. Stehli, 1954, p. 350
(ex Cyrtininae Frederiks, 1911, p. 5)]

Diagnosis. — Stout, apically perforated deltidium; spondylium and divided
trichorhinum; bilobed, commonly nonstriate cardinal process, but ctenophori-
dium may form in older growth stages.

Discussion. — We hypothesize that Cyrtina, the oldest genus of the Superfamily
Cyrtinoidea, if not derived from an unknown punctate orthid or intermediate,
was derived from one of the small impunctate Silurian genera of the Cyrtinopsinae
in the Superfamily Delthyridoidea, possibly Kozlowskiellina Boucot, 1958. How-
ever, demonstrating homology between Cyrtina and impunctate taxa is not easy
because general morphological similarity can indicate convergent or parallel evo-
lution, as well as a genetic relationship.

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Cyrtina became abundant at the beginning of the Devonian, already fully de-
veloped with endopunctate shell and tichorhinum. Kozlowskiellina is a small,
lamellose, cyrtinaform genus with conjunct delthyrial plates pierced by an apical
foramen. It has a ventral median septum and dental plates, and a bifid cardinal
process in the dorsal valve. We assume that Kozlowskiellina could have given
rise to Cyrtinopsis at about the same time that Cyrtina appeared, at the beginning
of the Devonian. Cyrtinopsis differs from Kozlowskiellina by the presence of a
spondylium pierced by a median septum and details of micro-ornament and is
closer to Cyrtina in this way (Havlicek, 1959:144-145; Johnson, 1970:pl. 72, fig.
21-29). Cyrtina could have split off from the Kozlowskiellina-Cyrtinopsis phy-
logeny by the acquisition of an endopunctate shell structure and a tichorhinum.

Stratigraphic Range. — Lower Devonian (basal Lochkovian)-Lower Carbonif-
erous (Visean).

Genera Included. — Cyrtina Davidson, 1858 [=Spinocyrtina Frederiks, 1916;
Cyrtinaellina Frederiks, 1926; Trochalocyrtina Wright, 1975]; Cyrtinaella Fred-
eriks, 1916; Squamulariina Frederiks, 1916 [=Pyramidalia Nalivkin, 1947]; Tec-
nocyrtina Johnson and Norris, 1972.

Family Komiellidae Johnson and Blodgett, 1993
[Family Komiellidae Johnson and Blodgett, 1993, p. 952]

Diagnosis. — Spondylium, or dental flanges and median septum not connected,
lacking a tichorhinum; simple, nonstriate cardinal process; jugum sessile.

Discussion. —The name Komiella Barkhatova, 1970 (p. 62, footnote) is a nomen
nudum.

Stratigraphic Range. — Lower Devonian (Emsian)-Upper Devonian (Frasnian).

Genus Included.— Komiella Liashenko, 1985.

Superfamily Suessioidea Waagen, 1883
[nom. correct. Carter, herein

( pro Superfamily Suessiacea Waagen, 1883) nom. trans. Pitrat, 1965, p. H675,
ex Subfamily Suessiinae Waagen, 1883, p. 498]

Diagnosis. — Usually cyrtiniform; ventral valve usually hemipyramidal to sub-
conical with high interarea; ventral median septum present, discrete dental ad-
minicula absent; endopunctae or hemipunctae obscure in some genera.

Discussion. — In this superfamily there is an obvious relationship between the
hemipyramidal or subconical growth form and internal structures of the ventral
valve, namely the adductor-shortening structures such as the spondylium. The
poor Paleozoic record of this superfamily hinders interpretation of relationships
with the Cyrtinoidea from which the suessioids were derived, probably in the
Lower Carboniferous. The Davidsoninidae are unique in this superfamily in being
very large, in probably lacking crural processes and a brachidium, and in having
hemipunctae. The Davidsoninidae are the only Carboniferous representatives of
the Suessioidea, but if one invokes Dollo’s Law they were probably not in the
direct line of ancestry of the Laballidae, which first appear in the Upper Permian
of China with normal brachidial structures intact.

Stratigraphic Range. — Lower Carboniferous (Visean)-Lower Jurassic.

Family Davidsoninidae Ivanova, 1972
[Family Davidsoninidae Ivanova, 1972, p. 41]

Diagnosis. — Lateral slopes ribbed; ventral interior with false spondylium com-

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posed of median septum and high dental flanges; ventral adminicula absent; shell
substance punctate, but punctae possibly not penetrating primary layer (hemi-
punctate); jugum and jugal processes absent; brachidium unknown, possibly ab-
sent.

Stratigraphic Range. — Lower Carboniferous (Visean-Namurian).

Genera Included. —Davidsonina Schuchert and LeVene, 1929; Carbocyrtina
Ivanova, 1975.

Family Laballidae Dagis, 1962
[nom. trans. Dagis, 1965, p. 91,
ex Subfamily Laballinae Dagis, 1962, p. 49]

Diagnosis. — Cyrtiniform; spondylium bisected by high median septum.
Discussion. — This family appears cryptogenically with the genus Eolaballa Liao
and Meng in the Upper Permian of China.

Stratigraphic Range. — Upper Permian-Upper Triassic.

Subfamily Laballinae Dagis, 1 962
[Subfamily Laballinae Dagis, 1962, p. 49]

Diagnosis. — Fold and sulcus well defined; lateral slopes smooth or with one or
two faint plicae; delthyrium open.

Stratigraphic Range.— Upper Permian-Upper Triassic.

Genera Included. — Laballa Moisseiev, 1962; Eolaballa Liao and Meng, 1986;
Pseudolaballa Dagis, 1974.

Subfamily Spinolepismatininae Carter, new subfamily

Diagnosis. — Lateral slopes distinctly plicate; fold and sulcus smooth; micro-
ornament densely spinulose.

Stratigraphic Range. — Upper Triassic.

Genera Included.— Spinolepismatina Dagis, 1974; Klipsteinella Dagis, 1974;
Klipsteinelloidea Sun, 1981.

Subfamily Paralepismatininae Carter, new subfamily

Diagnosis. — Fold and sulcus poorly developed; entirely ribbed; micro-ornament
absent.

Stratigraphic Range. —Middle Triassic (Anisian).

Genus Included.— Par alepismatina Yang and Xu, 1966.

Family Bittnerulidae Schuchert, 1929
[nom. trans. Carter, herein
(ex Subfamily Bittnerulinae Schuchert, 1929, p. 21)]

[=Subfamily Thecocyrtellinae Dagis, 1965, p. 105]

Diagnosis. — Cyrtiniform; dental adminicula and spondylium absent; ventral
septum and dental flanges fused by apical callus or short transverse plate.
Stratigraphic Range. — Middle-Upper Triassic.

Subfamily Bittnerulinae Schuchert, 1929
[Subfamily Bittnerulinae Schuchert, 1929, p. 21]

Diagnosis. — Smooth or obscurely ribbed; fold and sulcus absent or very weak;
delthyrium closed by convex deltidium.

Stratigraphic Range. — Middle-Upper Triassic.

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Genera Included. —Bittnerula Hall and Clarke, 1895; Leiolepismatina Yang and
Xu, 1966; Thecocyrtella Bittner, 1892; Thecocyrtelloidea Yang and Xu, 1966.

Subfamily Hirsutellinae Xu and Liu, 1983
[Subfamily Hirsutellinae Xu and Liu, 1983, p. 82]

Diagnosis.— Lateral slopes ribbed; fold and sulcus weakly to moderately de-
veloped; delthyrium open or partially occluded by various plates.

Stratigraphic R <2 - Middle-Upper Triassic.

Genera Included. —Hirsutella Cooper and Muir-Wood, 1951; Flabellocyrtia
Chorowicz and Termier, 1975; Neocyrtina Yang and Xu, 1966; Spiriferinoides
Tokuyama, 1957.

Family Suessiidae Waagen, 1883
[nom. trans. Pitrat, 1965, p. H679
(ex Subfamily Suessiinae Waagen, 1883, p. 498)]

Diagnosis. — Entirely plicate; delthyrium open; dental adminicula and spon-
dylium absent; large elongate hingeplate bearing adductors present.

Stratigraphic Range. — Lower Jurassic.

Genus Included. —Suessia Deslongchamps, 1855.

Superfamily Spondylospiroidea Hoover, 1991
[nom. trans. Carter, herein
(ex Family Spondylospiridae Hoover, 1991, p, 75)]

Diagnosis. — Spiriferiform to cyrtiniform; hingeline partially to completely cren-
ulate; dental adminicula converging or forming spondylium; punctation well de-
veloped.

Discussion. —This compact group of 14 genera appeared cryptogenically in the
Middle Triassic. Their unifying morphological character is the partially or com-
pletely crenulate hingeline. This articulatory structure has been termed “dentic-
ulate” by numerous authors but must be differentiated from the denticulate hinge-
line of the Spiriferoidea and Paleckelmanelloidea. In these groups the denticles
consist of taleola-like rods embedded within secondary fibrous shell matter of the
interareas. The crenulate teeth of the Spondylospioidea lack “taleolae” and are
similar in each valve and interlocking. This is a unique or possibly synapomor-
phous articulatory structure among the spiriferinids.

This group includes both wide-hinged “spiriferoid” genera and hemipyramidal
“cyrtinoid” genera. Internally, there is a good spondylium in the “cyrtinoid”
genera or converging dental adminicula in the “spiriferoid” genera. The cyrtini-
form spondylium-bearing genera are considered here to be most primitive and to
indicate derivation from the laballids. The spiriferiform wide-hinged genera seem
likely to have been derived from a cyrtiniform ancestor with neotenous loss of
the complete spondylium and concomitant extension of the hingeline.

Stratigraphic Range. — Upper Triassic.

Family Spondylospiridae Hoover, 1991
[Family Spondylospiridae Hoover, 1991, p. 75]

Diagnosis. —Cyrtiniform to globose; lateral slopes ribbed; spondylium bisected
by high median septum; jugum complete, supported by sessile jugal net.

Stratigraphic Range. — Middle (Ladinian)-U pper Triassic (Rhaetian).

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Subfamily Spondylospirinae Hoover, 1991
[Subfamily Spondylospirinae Hoover, 1991, p. 80]

Diagnosis. — Dental adminicula and ventral interarea complete, not pierced by
paired pedicle foramina.

Stratigraphic Range.— Upper Triassic (Kamian-Rhaetian).

Genera Included. — Spondylospira Cooper, 1942; Phenacozugmayerella Hoo-
ver, 1991; Vitimetula Hoover, 1991; Yanospira Dagis, 1977; Zugmayerella Dagis,
1963.

Subfamily Dagyspiriferinae Hoover, 1991
[Subfamily Dagyspiriferinae Hoover, 1991, p. 77]

Diagnosis. — Globose to cyrtiniform; entirely ribbed; apex of spondylium and
ventral interarea pierced by paired elongate foramina.

Stratigraphic Range.— Upper Triassic (Kamian-N orian) .

Genera Included. — Dagyspirifer Hoover, 1991; Pseudospondylospira Hoover,
1991.

Family Rastelligeridae Carter, new family

Diagnosis. — Spiriferiform; usually transverse; fold and sulcus smooth; lateral
slopes usually plicate; dental adminicula subparallel or convergent.

Stratigraphic Range. - Middle-Upper Triassic.

Subfamily Rastelligerinae Carter, new subfamily

Diagnosis.— Strongly transverse; cardinal extremities extended; lateral slopes
strongly plicate; dental adminicula convergent or forming sessile spondylium.

Stratigraphic Range.— Middle (Ladinian)-U pper Triassic (Rhaetian).

Genera Included. —Rastelligera Hector, 1879; Boreiospira Dagis, 1974; Psioi-
diella Campbell, 1968.

Subfamily Dentospiriferininae Carter, new subfamily

Diagnosis. — Cardinal extremities subangular to rounded; lateral slopes smooth
or with few weak plicae; dental adminicula subparallel to convergent.

Stratigraphic Range. — Upper Triassic (Kamian-Norian).

Genera Included. — Dentospiriferina Dagis, 1965; Canadospira Dagis, 1972; Ori-
entospira Dagis, 1965; Psioidea Hector, 1879.

Suborder Spiriferinidina Ivanova, 1972
[Suborder Spiriferinidina Ivanova, 1972, p. 41]

[= Suborder Spiriferinidina Cooper and Grant, 1976, p. 2666]

Diagnosis. — Subequally biconvex; ventral beak usually incurved; strong ventral
median septum invariably present.

Discussion. — This suborder is presumed to have been derived from the cyrtinoid
Family Komiellidae which had discrete dental adminicula in the Upper Devonian.
Two distinct lineages appeared at this time. One formed the Pennospiriferinoidea
with strong ventral median septum and leading to the highly successful radiation
of similar genera in the Late Paleozoic and Triassic. The other lineage included
large transverse genera bearing a strong delthyrial plate, the Syringothyridoidea.
Although radically transformed internally, some of the advantages of the adductor-
shortening structures of the cyrtinoid spondylium are retained in genera with a
conical or hemipyramidal ventral valve, by the development of a simple high

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365

median septum or large transverse delthyrial plate, respectively. In some mod-
erately inflated, subequally biconvex syringothryridoid genera with low ventral
interareas the delthyrial plate is lost, reflecting the lesser need for shortening the
length of the adductors.

Stratigraphic Range.— XJppQt Devonian (Upper Famennian)-Lower Jurassic.

Superfamily Syringothyridoidea Frederiks, 1926
[nom. correct. Carter, herein (pro Syringothyridacea Frederiks, 1926
(nom. imperf.) nom. trans. Ivanova, 1972, p. 319,
ex Syringothyrinae Frederiks, 1926, p. 41 1)]

Diagnosis. — Outline usually spiriferiform; moderately to strongly transverse;
often strongly inequivalved; fold and sulcus invariably developed; lateral slopes
with simple ribbing; micro-ornament consisting of fine short radial striae with
fine elongate pustules or spinules arranged in quincunx between striae, producing
textile-like appearance; dental adminicula present; cardinalia usually stout and
wide; punctae highly variable in size, commonly sparsely and/or irregularly dis-
tributed.

Stratigraphic Range. — Upper Devonian (Upper Famennian)-U pper Permian.

Family Syringothrididae Frederiks, 1926
[nom. correct. Pitrat, 1965, p. H691 (pro Syringothyridae Frederiks, 1926
(nom. imperf.) nom. transl. Ivanova, 1959, p. 55,
ex Syringothyrinae Frederiks, 1926, p. 41 1)]

Diagnosis.— Cardinal extremities subangular to slightly rounded; lateral slopes
with moderately numerous simple ribs; interspaces narrow and subangular to
moderately broad; fold and sulcus smooth medially; ventral interarea high to very
high; perideltidial areas present.

Stratigraphic Range.— Upper Devonian (Upper Famennian)-Lower Permian.

Subfamily Syringothyridinae Frederiks, 1926
[nom. correct. Pitrat, 1965, p. H692
(pro Syringothyrinae Frederiks, 1926, p. 411)]

Diagnosis.— Delthyrial plate and syrinx present.

Stratigraphic Range.— Upper Devonian (Upper Famennian)-Lower Permian.

Genera Included.— Syringothyris Winchell, 1863 [=Syringopleura Schuchert,
1910]; Subansiria Sahni and Srivastava, 1956.

Subfamily Septosyringothyridinae Massa, Termier, and Termier, 1974
[nom. correct. Legrand-Blain, 1974, p. 120,
pro Septosyringothyridae Massa, Termier, and Termier, 1974
(nom. imperf.) nom. transl. Carter, herein
(ex Septosyringothyridae Massa, Termier, and Termier, 1974, p. 168)]

Diagnosis. — Median septum and syrinx present in ventral valve.

Stratigraphic Range. — Lower Carboniferous.

Genera Included. —Septosyringothyris Vandercammen, 1955; Histosyrinx Mas-
sa, Termier, and Termier, 1974.

Subfamily Permasyrinxinae Waterhouse, 1986
[Subfamily Permasyrinxinae Waterhouse, 1986, p. 3]

Diagnosis.— Syrinx absent.

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Discussion. — The validity of Pseudosyringothyris Frederiks is in doubt. Gri-
gor’eva (1977) states that intensive collecting for topotypes of P. karpinskii has
failed to produce specimens internally similar to Frederiks’s description of the
type specimens. The presence of perideltidial areas in Primorewia Likharev and
KotJjar has not been confirmed. If they are truly lacking, then Primorewia should
be reassigned to the Licharewiidae.

Stratigraphic Range. — Lower Carboniferous (Lower Tournaisian)-Upper
Permian.

Genera Included. —Permasyrinx Waterhouse, 1983; Asyrinx Hudson and Sud-
bury, 1959 [=Kungaella Solomina, 1988]; Asyrinxia Campbell, 1957; Cyrtella
Frederiks, 1924 [=Punctocyrtella Plodowski, 1 968]; Myodelthyrium Thomas, 1985;
1 Primorewia Likharev and Kotljar, 1978; ? Pseudosyringothyris Frederiks, 1916;
Pseudosyrinx Weller, 1914; Sulcicosta Waterhouse, 1983; Verkhotomia Sokol-
skaya, 1963.

Family Dimegelasmidae Carter, new family

Diagnosis.— Cardinal extremities well rounded; ventral interarea low to mod-
erately high; lateral slopes with few plications separated by broad rounded inter-
spaces; sulcus sparsely plicate or costate; perideltidial areas present; syrinx absent;
delthyrial plate small or absent; shell substance thin.

Stratigraphic Range.— Upper Devonian (Famennian)-Lower Carboniferous
(Visean).

Genera Included.— Dimegelasma Cooper, 1942 [=Doescherella Abramov and
Grigor’eva, 1987]; Guilinospirifer Xu and Yao, 1988; Zeugopleura Carter, 1988.

Family Licharewiidae Slyusareva, 1958
[nom. trans. Solomina, 1988, p. 44
(ex Licharewiinae Slyusareva, 1958, p. 582)]

Diagnosis. —Perideltidial areas absent; syrinx absent; moderate to thick callus
deposits in ventral umbonal region commonly present.

Discussion. — The presence or absence of perideltidial areas in Pyramidal hyris
Hu, Tumarinia Solomina and Grigor’eva, and Tuotalania Hu has not been dem-
onstrated. The presence of such areas in any of these genera would require transfer
to the Family Syringothyrididae.

Stratigraphic Range.— Upper Carboniferous-Upper Permian.

Genera Included. —Licharewia Einor, 1939 [=Rugulatia Sokolskaya, 1952]; Ol-
gerdia Grigor’eva, 1977; Orulgania Solomina and Cherniak, 1961; Penzhinella
Solomina, 1985; Permospirifer Kulikov, 1950; I Pyramidal hyris Hu, 1983; ITu-
marinia Solomina and Grigor’eva, 1973; 1 Tuotalania Hu, 1983.

Superfamily Pennospiriferinoidea Dagis, 1972
[nom. trans . Carter, herein

(ex Subfamily Pennospiriferininae Dagis, 1972, p. 36)]

Diagnosis. — Spiriferiform to cyrtiniform; ventral interarea usually low or only
moderately high; lateral slopes ribbed; dental adminicula and median septum
discrete; punctae well developed, usually densely spaced.

Stratigraphic Range.— Upper Devonian (Upper Famennian)-Lower Jurassic.

Family Punctospiriferidae Waterhouse, 1975
[nom. correct. Carter, herein,

(pro Family Punctospiriferinidae Waterhouse, 1987, p. 44;

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Carter et al. — Spiriferid Brachiopod Revision

367

nom. trans. Waterhouse 1983 or 1987, ex Subfamily
Punctospiriferinae Waterhouse, 1975, p. 17)]

Diagnosis. — Usually transverse; fold and sulcus narrow, weakly to moderately
developed; dental adminicula short, subparallel to slightly divergent; micro-or-
nament capillate and usually regularly lamellose or subimbricate.

Discussion. — The faintly capillate micro-ornament in Yangkongia Xu and Liu
is very poorly preserved and may in fact be an artifact of preservation. If so,
reassignment would be necessary.

The earliest genus in this superfamily, Ziganella Nalivkin, has a low ventral
median septum or ridge and normal fold-sulcus. This punctospiriferid genus is
believed to have been derived from the Komiellidae or some intermediate ancestor
within this lineage.

Stratigraphic Range.— Upper Devonian (Upper Famennian)-Middle Triassic.

Genera Included. — Punctospirifer North, 1 920; Alipunctifera Waterhouse, 1975;
Lamnaespina Waterhouse, 1976; Liriplica Campbell, 1961; Pustulospiriferina
Waterhouse, 1983; 1 Yangkongia Xu and Liu, 1983; Ziganella Nalivkin, 1960.

Family Spiropunctiferidae Carter, new family

Diagnosis. — Cardinal extremities well rounded; fold and sulcus plicate; micro-
ornament absent.

Stratigraphic Range. — Lower Carboniferous (Upper Visean)-Upper Permian.

Genera Included.— Spiropunctifera Ivanova, 1971; Genuspirifer Liang, 1990.

Family Reticulariinidae Waterhouse, 1975
[nom. trans. Waterhouse, 1983, p. 138,
ex Subfamily Reticulariininae Waterhouse, 1975, p. 15]

Diagnosis. —Outline tran verse; cardinal extremities usually extended, rarely
rounded; fold and sulcus narrow, usually well delimited; sulcus smooth or with
weak median rib; dental adminicula divergent; micro-ornament of coarse hollow
spines.

Stratigraphic Range. — Lower Carboniferous (Upper Visean)-Upper Permian.

Genera Included. — Reticulariina Frederiks, 1916; Altiplecus Stehli, 1954; Gjeli -
spinifera Ivanova, 1975; Spinuliplica Campbell, 1961.

Family Paraspiriferinidae Cooper and Grant, 1976
[Family Paraspiriferinidae Cooper and Grant, 1976, p. 2729]

Diagnosis. — Outline transversely subelliptical; cardinal extremities well round-
ed; fold and sulcus usually well delimited; dental adminicula short, usually di-
vergent; micro-omament regularly and finely lamellose, with lamellae bearing fine
hair-like spinules in some genera.

Stratigraphic Range.— 1 Lower Carboniferous (Upper Visean), Upper Carbon-
iferous-Upper Permian.

Genera Included.— Par aspiriferina Reed, 1944; Callispirina Cooper and Muir-
Wood, 1951; Lamniplica Waterhouse and Rao, 1989; Polystylus Klets, 1993;
Yaoniella Waterhouse, 1983; Zaissania Sokoskaya, 1968.

Family Crenispiriferidae Cooper and Grant, 1976
[Family Crenispiriferidae Cooper and Grant, 1976, p. 2709]

Diagnosis. — Outline usually transversely subelliptical to subtrigonal; fold and
sulcus usually narrow and poorly to moderately delimited; lateral slopes with few

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strong lateral plicae and subimbricate growth varices; dental adminicula very
short; micro-ornament finely pustulose.

Discussion. — Tulungospirifer Ching and Sun is tentatively assigned here on the
basis of its growth form. Its micro-ornament differs from other genera in this
family in having radial grooves and spinules.

Stratigraphic Range. — Lower Carboniferous (Upper Chesterian)-U pper Tri-
assic.

Genera Included. — Crenispirifer Stehli, 1954; Lancangjiangia Jin and Fang,
1977; Metriolepis Cooper and Grant, 1976; Pseudospiriferina Young and Xu,
1966; Spiriferellina Frederiks, 1924 [=Tylotoma Grabau, 1934]; Sulcispiriferina
Waterhouse and Gupta, 1981; I Tulungospirifer Ching and Sun, 1976.

Family Sarganostegidae Cooper and Grant, 1976
[Family Sarganostegidae Cooper and Grant, 1976, p. 2743]

Diagnosis. — External surface with quincuntially arranged very coarse punctae;
micro-ornament absent.

Stratigraphic Range. - Middle-Upper Permian.

Genus Included. — Sarganostega Cooper and Grant, 1969.

Family Balatonospiridae Dagis, 1974
[nom. trans. Carter, herein,
ex Subfamily Baiatonospirinae Dagis, 1974, p. 137]

[=Subfamily Nudispiriferininae Xu and Liu, 1983]

Diagnosis. — Outline transversely subelliptical to subquadrate or subpentagonal;
fold and sulcus poorly developed; entire surface usually ribbed.

Stratigraphic Range. —Middle-Upper Triassic.

Subfamily Yalongiinae Carter, new subfamily

Diagnosis. — Dental adminicula discrete.

Stratigraphic Range.— Middle Triassic.

Genera Included. — Yalongia Xu and Liu, 1983; Aequispiriferina Yang and Yin,
1962; Sinucostella Xu and Liu, 1983.

Subfamily Baiatonospirinae Dagis, 1974
[Subfamily Baiatonospirinae Dagis, 1974, p. 137]

Diagnosis. — Dental adminicula reduced or absent; dental flanges and median
septum fused by transverse plate or callus.

Stratigraphic Range.— Middle (Anisian)-U pper (Kamian) Triassic.

Genera Included. —Balatonospira Dagis, 1974; Dinarispira Dagis, 1974; Koeve -
skallina Dagis, 1965; Nudispiriferina Yang and Xu, 1966; Tylospiriferina Xu,
1978.

Family Pennospiriferinidae Dagis, 1972
[nom. trans. Carter, herein,
ex Subfamily Pennospiriferininae Dagis, 1972, p. 36]

Diagnosis.— Transverse, usually with angular or extended lateral extremities;
dental adminicula subparallel or converging; micro-ornament absent.

Stratigraphic Range. —Upper Carboniferous-Upper Triassic (Kamian), ?Lower
Jurassic.

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Carter et al.™ Spiriferid Brachiopod Revision

369

Subfamily Pennospiriferininae Dagis, 1972
[Subfamily Pennospiriferininae Dagis, 1972, p. 36]

Diagnosis. — Fold and sulcus smooth; dental adminicula converging, fused with
median septum by thick callus.

Discussion. — The micro-ornament of Callospiriferina Rousselle, if any, is poorly
known. Furthermore, the growth form of this genus differs substantially from that
of the type genus.

Stratigraphic Range. — Middle Triassic (Ladinian)-?Lower Jurassic.

Genera Included.— Pennospiriferina Dagis, 1965; ? Callospiriferina Rousselle,
1977.

Subfamily Punctospirellinae Dagis, 1974
[Subfamily Punctospirellinae Dagis, 1974, p. 135]

[=Family Xestotrematidae Cooper and Grant, 1976, p. 2748]

Diagnosis. — Fold and sulcus smooth or weakly ribbed; dental adminicula dis-
crete, usually short, not converging.

Discussion. — This long-ranging subfamily, if it proves to be monophyletic, is
probably the ancestral stock for the other Triassic families in this superfamily.

Stratigraphic Range. — Upper Carboniferous-Middle Triassic.

Genera Included. —Punctospirella Dagis, 1974; Arionthia Cooper and Grant,
1976; Laioporella Ivanova, 1975; Xestotrema Cooper and Grant, 1969 [=Mu-
crospiriferinella Waterhouse, 1982].

Family Lepismatinidae Xu and Liu, 1983
[Family Lepismatinidae Xu and Liu, 1983, p. 82]

Diagnosis. — Cyrtiniform or globose; lateral slopes ribbed; fold and sulcus well
developed; delthyrium open; dental adminicula discrete.

Stratigraphic Range. — Middle Triassic-Lower Jurassic.

Subfamily Lepismatininae Xu and Liu, 1983
[nom. trans. Carter, herein,
ex Family Lepismatinidae Xu and Liu, 1983, p. 82]

Diagnosis.— Fold and sulcus smooth; micro-ornament of dense spinules; car-
dinalia sessile.

Stratigraphic Range. - Middle-Upper Triassic.

Genera Included. —Lepismatina Wang, 1955 [= Costispiriferina Dagis, 1974];
Altoplicatella Xu and Liu, 1983; Pseudolepismatina Ching and Sun, 1976.

Subfamily Pseudocyrtininae Carter, new subfamily

Diagnosis. — Fold and sulcus smooth; cardinalia supported by short median
septum; micro-ornament absent.

Stratigraphic Range. — Upper Triassic.

Genera Included.— Pseudocyrtina Dagis, 1962; Bolilaspirifer Sun, 1981.

Subfamily Dispiriferininae Carter, new subfamily

Diagnosis. — Entirely ribbed; no dorsal septum.

Stratigraphic Range. — Middle Triassic-Lower Jurassic.

Genera Included. —Dispiriferina Siblik, 1965; Qingyenia Yang and Xu, 1966.

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Superfamily Spiriferinoidea Davidson, 1884
[nom. correct. Carter, herein (pro Superfamily Spiriferinacea Davidson, 1884
(nom. imperfect.) nom. trans. Ivanova, 1959, p. 57,
ex Subfamily Spiriferininae Davidson, 1884, p. 354)]

Diagnosis. — Reticulariiform, subequally biconvex; cardinal extremities well
rounded; ventral beak incurved; fold and sulcus usually weakly developed; lateral
slopes smooth or obscurely ribbed, rarely with moderately developed ribbing.

Discussion. —The growth form, internal morphology, and micro-ornament of
this superfamily strongly suggest that it was derived from some member of the
Family Paraspiriferinidae.

Stratigraphic Range. — Middle Triassic-Lower Jurassic.

Family Spiriferinidae Davidson, 1884
[nom. trans. Ivanova, 1959, p. 57
(ex Subfamily Spiriferinidae Davidson, 1884, p. 354)]

Diagnosis. — Lateral slopes smooth or obscurely ribbed.

Stratigraphic Range. —Middle Triassic-Lower Jurassic.

Subfamily Spiriferininae Davidson, 1884
[Subfamily Spiriferinidae Davidson, 1884, p. 354]

[=Subfamily Spiriferellinae Paeckelmann, 1932, p. 25,
pro Spiriferininae Schuchert, 1929, p. 21]

Diagnosis. — Reticulariiform; dental adminicula discrete.

Stratigraphic Range.— Middle Triassic-Lower Jurassic.

Genera Included. — Spiriferina Orbigny, 1847 [=Liospiriferi na Rouselle, 1977];
Calyptoria Cooper, 1989 [=Cingolospiriferina Pozza, 1992]; Mentzelioides Dagis,
1974; Qinghaispiriferina Sun and Ye, 1982; Triadispira Dagis, 1961; Viligella
Dagis, 1965.

Subfamily Paralaballinae Carter, new subfamily

Diagnosis. — Cyrtiniform; fold and sulcus weakly developed; ornament absent;
dental adminicula discrete, thin, short, subparallel.

Stratigraphic Range. — Upper Triassic.

Genus Included. —Paralaballa Sun, 1981.

Subfamily Mentzeliinae Dagis, 1974
[Subfamily Mentzeliinae Dagis, 1974, p. 138]

Diagnosis. — Dental adminicula absent.

Stratigraphic Range.— Middle-Upper Triassic.

Genera Included.— Mentzelia Quenstedt, 1871; Madoia Sun and Ye, 1982;
Paramentzelia Xu, 1978.

Subfamily Tethyspirinae Carter, new subfamily
Diagnosis. — Spondylium present.

Stratigraphic Range. — Middle (Ladinian)-U pper (Kamian?) Triassic.

Genera Included. — Tethyspira Siblik, 1991; Spondylospiriferina Dagis, 1972.

Family Sinucostidae Xu and Liu, 1983
[nom. trans. Carter, herein,
ex Subfamily Sinucostinae Xu and Liu, 1983, p. 112]

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Carter et al.-Spiriferid Brachiopod Revision

371

Diagnosis. — Lateral slopes ribbed; dental adminicula discrete.

Stratigraphic Range.— Middle-Upper Triassic, ?Lower Jurassic.

Subfamily Sinucostinae Xu and Liu, 1983
[Subfamily Sinucostinae Xu and Liu, 1983, p. 112]

Diagnosis. — Dorsal septum absent.

Stratigraphic Range. -Middle-Upper Triassic, ?Lower Jurassic.

Genera Included. —Sinucosta Dagis, 1963 [=Guseriplica Dagis, 1963]; Mentze-
liopsis Trechmann, 1918; Qispiriferina Xu and Liu, 1983.

Subfamily Jiangdaspiriferinae Carter, new subfamily

Diagnosis. — With dorsal median septum.

Stratigraphic Range.— Upper Triassic.

Genus Included. —Jiangdaspirifer Chen, Rao, Zhou, and Pan, 1986.

Suborder uncertain
Superfamily uncertain
Family uncertain

Genus Included.— Plicatosyrinx Minato, 1952.

Rejected from the Spirifers

Genera Included. — Guangshunia Xian 1978 [possible stringocephalid]; Iliella
Rukavischnikova 1980 [Ashgill, not an Eospirifer]; Plectospirifer Grabau, 1931
[its type species is a synonym of Athyrisina squamosa Hayasaka, type species of
Athyrisina ]; Pustulatia Cooper 1956 [possible anoplothecid]; Xerxespirifer Cocks
1979 [rhynchonellid or leptocoeliid].

Acknowledgments

Johnson’s research was supported by National Science Foundation Grant EAR-9204772.

Appendix

Boucot (1973:39-61; 1975:369-370) used the term “ctenophoridium’’ for the
striate site of dorsal diductor attachment in some spiriferid brachiopod genera,
but did not define the term. We remedy that oversight here and call attention that
“ctenophoridium” will appear as an accepted usage in the glossary of the forth-
coming revision of the brachiopod Treatise. Krans (1965:pl. 16) illustrated thin
sections of the ctenophoridium, which he refered to as a cardinal process of
Cyrtospirifer and an unidentified genus.

ctenophoridium.— Striated or comb-like site of dorsal diductor muscle attach-
ment situated on floor of notothyrial cavity or bifid on hinge plates, but not
elevated on a shaft-like cardinal process.

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To include here every dated citation given above would require the editors to print many pages of
text in addition to an already long paper. Therefore, we exclude references for the taxonomic units
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taxon may call, write, or e-mail Carter. Carter’s e-mail address is: JLC4@VMS.CIS.PITT.EDU; tele-
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Devonian of Bohemia. Sbomik Geologickych Ved (Paleontologie), 14:7-34.

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Williams, A., A. J. Rowell, H. M. Muir- Wood, C. W. Pitrat, H. Schmidt, F. G. Stehli, D. V.
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927 pp.

INDEX TO VOLUME 63
CONTENTS

ARTICLES

Geographic variation and systematics of the striped whipsnakes ( Masticophis taeniatus complex;
Reptilia: Serpentes: Colubridae) Jeffrey D. Camper and James R. Dixon

U-shaped orientation of Hunter-Schreger bands in the enamel of Moropus (Mammalia: Chal-
icotheriidae) in comparison to some other Perissodactyla ........ Wighart v. Koenigswald

Five new species of Hydroptila from eastern United States (Insecta: Trichoptera:
Hydroptilidae) Jan L. Sykora and Steven C. Harris

Description of the immature stages of six species of Sphaenognathus, with comparative notes
on phylogeny and natural history (Insecta: Coleoptera: Lucanidae) ....... Giovanni Onore

Palaeophichthys parvulus Eastman, 1908, a gnathorhizid dipnoan from the Middle Pennsyl-
vanian of Illinois, USA Hans-Peter Schultze

Exploitation of mammals at the Early Bronze Age site of West Row Fen (Mildenhall 165),
Suffolk, England Sandra L. Olsen

Tephrodytes brassicarvalis, new genus and species (Anura: Pelodytidae), from the Arikareean

Cabbage Patch beds of Montana, USA, and pelodytid-pelobatid relationships

Amy C. Henrici

Taiwanese species of Neopanorpa (Insecta: Mecoptera: Panorpidae) George W. Byers

Biology and immature stages of Chloropidae (Insecta: Diptera) associated with spike-rushes
(Cyperaceae: Eleocharis ) I. Stem borers Teresa K. Wearsch and B. A. Foote

Beads and pendants from Trants, Montserrat: Implications for the prehistoric lapidary industry
of the Caribbean David R. Watters and Richard Scaglion

A primitive cricetid (Mammalia: Rodentia) from the Middle Eocene of Jiangsu Province,
China Ban-yue Wang and Mary R. Dawson

Mediocampus, a new stinkbug genus from the Dominican Republic (Insecta: Heteroptera:
Pentatomidae) Donald B. Thomas

Archaeology of Trants, Montserrat. Part 1. Field methods and artifact density distributions. .
David R. Watters

Archaeology of Trants, Montserrat. Part 2. Vertebrate fauna. Elizabeth J. Reitz

A new species of Antocha (Subgenus Orimargula) from Sulawesi (Diptera: Tipulidae) and its
mate-clasping behavior. Chen W. Young

A revised classification of the spiriferid brachiopods

John L. Carter, J. G. Johnson, Remy Gourvennec, and Hou Hong-fei

105

115

155

185

193

215

239

257

265

297

319

327

375

376

Annals of Carnegie Museum

vol. 63

REVIEWS

The Scent of Orchids: Olfactory and Chemical Investigations [Roman Kaiser] ..............

Fredrick W. Wright, Jr. 101

New Jersey Ferns and Fern Allies [James D. Montgomery and David E. Fairbrothers]

Bonnie L. Isaac 263

NEW TAXA

NEW GENERA, SPECIES, AND SUBSPECIES

Antocha ( Orimargula ) possess iv a, new species 319

t Dimegelasmidae, new family 366

t Elythynidae, new family 338

Hydroptila antennopedia, new species 68

Hydroptila hlicklei, new species . . 72

Hydroptila holzenthali, new species 73

Hydroptila morsei, new species 71

Hydroptila parachelops, new species 69

Mediocampus, new genus 257

Mediocampus dominicanus, new species 259

Neopanorpa youngi, new species 186

^Palaeochoristitidae, new family 334

t Pappocricetodon antiquus, new species 242

t Rastelligeridae, new family 364

t Skelidorygmidae , new family 349

t Spiropunctiferidae , new family 367

t Tephrodytes, new genus 159

t Tephrodytes brassicarvalis, new species 160

t Ulbospiriferidae, new family 334

AUTHOR INDEX

Byers, George W 185

Camper, Jeffrey D. . . . . 1

Carter, John L. 327

Dawson, Mary R. 239

Dixon, James R. 1

Foote, B. A. 193

Gourvennec, Remy 327

Harris, Steven C 67

t Fossil taxa

1994

Index to Volume 63—1994

377

Henrici, Amy C 155

Hong-fei, Hou 327

Isaac, Bonnie L. 263

Johnson, J. G 327

Koenigswald, Wighart v. 49

Olsen, Sandra L. 115

Onore, Giovanni 77

Reitz, Elizabeth J 297

Scaglion, Richard 215

Schultze, Hans-Peter 105

Sykora, Jan L. 67

Thomas, Donald B. 257

Wang, Ban-yue 239

Watters, David R. 215, 265

Wearsch, Teresa K. 193

Wright, Fredrick W., Jr. 101

Young, Chen W. . . 319

INSTRUCTIONS FOR AUTHORS

Manuscript style.— Articles should include the follow-
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Abstracts. Abstracts should be short, double-spaced
and substantive and included at the head of the first page
of text.

Text. Do not right justify text or break (hyphenate) a
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Measurements. Metric units should be used, if possi-
ble.

1) Two authors in an institutional series:

Matthew, W. D., and W. Granger. 1923. The
fauna of the Houldjin Gravels. American Mu-
seum of Natural History Novitates, no. 97:

1t6.

2) Same authors repeated— use three-em dash:

. 1923. The fauna of the Ardyn Obo For-
mation. American Museum of Natural

History Novitates, no. 98:1-5.

3) Same authors plus a third author— repeat all authors:

Matthew, W. D., W. Granger, andG. G. Simpson.
1928. Paleocene multituberculates from Mon-
golia. American Museum of Natural History
Novitates, no. 331:1-4.

4) Chapter in an edited volume:

Rausch, R. L. 1 963. A review of the distribution
of Holarctic mammals. Pp. 29-43, in Pacific
Basin Biography (J. L. Gressitt, ed.), Bishop
Museum Press, Honolulu, Hawaii, xx + 450

pp.

5) Unpublished dissertation:

Smith, J. P. 1976. Review of Eocene mammals.
Unpublished Ph.D. dissert.. University of Cal-
ifornia, Berkeley, 302 pp.

6) Book:

White, M. J. D. 1961. The Chromosomes. Me-
thuen and Co., Ltd., London, 120 pp.

7) Journal articles with usual volume and issue number:

Anderson, W. 1 . 1969. Lower Mississippian con-
odonts from northern Iowa. Journal of Pale-
ontology, 43(4):9 16-928.

Illustrations. All illustrations will be called figures, and
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illustrations are required, one (original artwork) for re-
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