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ISSN 0097-4463
AN NALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
ARTICLE 1
23 MAY 1986
VOLUME 55
CAPTORHINID REPTILES FROM THE EARLY PERMIAN
OF NEW MEXICO, WITH DESCRIPTION OF A NEW
GENUS AND SPECIES
David S Berman
Associate Curator, Section of Vertebrate Fossils
Robert R. Reisz^
Research Associate, Section of Vertebrate Fossils
Abstract
A new genus and species of single-tooth-rowed captorhinid, Rhiodenticulatus heatoni,
is based on two skulls and partial postcranial skeletons collected from the Lower Permian
Cutler Formation near Arroyo de Agua, north-central New Mexico. A cladistic analysis
of its relationships to other single-tooth-rowed captorhinids suggests that it is a primitive
sister taxon to Labidosaurus and Eocaptorhinus. The dentition of R. heatoni, however,
exhibits several unique derived features which are interpreted as representing an adap-
tation to a specialized diet.
Puercosaurus obtiisidens Williston, 1916, the only previously described captorhinid
from New Mexico, is declared a nomen dubium because the holotypic left dentary is
indeterminate, and there is no basis for accepting that it and the two poorly preserved
captorhinid skulls found at a different locality and referred to the species by Williston
(1916) are conspecific. Additional captorhinid remains have been collected recently from
the Lower Permian Cutler, Abo, and Sangre de Cristo formations at widely scattered
localities in central and northern New Mexico. Though these specimens, as well as the
skulls referred to ^‘‘Puercosaurus obtusidens,"’’ are too poorly preserved to be assigned to
existing or new taxa, they do indicate that the Captorhinidae was diverse and widely
distributed in the Lower Permian of New Mexico.
‘ Address: Department of Biology, Erindale Campus, Univer
Ontario L5L 1C6, Canada.
Submitted 2 August 1985.
! i I M
V
2
Annals of Carnegie Museum
VOL. 55
Introduction
Published accounts of captorhinid reptiles from the late Paleozoic
of New Mexico have been limited to two reports (Williston, 1916;
Langston, 1953). Williston (1916) described a small captorhinid, Puer~
cosaurus obtusidens, on the basis of three poorly preserved and incom-
plete specimens, a left dentary and two skulls, collected from the Cutler
Formation in the Rio Puerco drainage in the north-central part of the
state. Further discoveries of captorhinids were not made until 1934-
1935, when collecting was resumed by field parties from the University
of California, Berkeley. While conducting extensive field work in the
Lower Permian Cutler Formation of the same area, three moderately
well preserved specimens, including two skulls with jaws and articu-
lated postcranial materials were found at the well known Camp quarry
near the small village of Arroyo de Agua (see Langston, 1953, for
histories and vertebrate assemblages of well known localities of the
area). The only published report of these specimens was a brief reference
to them by Langston (1953) in a discussion of the age of the late
Paleozoic vertebrate-bearing strata of New Mexico. Here he notes (1953:
410) “a small romeriid cotylosaur possibly referable to Puercosaurus
obtusidens is more primitive than Romeria texana of the middle Wich-
ita (Putnam)” of the Lower Permian of Texas. Extensive collecting by
the authors during the past several years in the Lower Permian deposits
throughout New Mexico has resulted in the discovery of additional
captorhinid remains from the Cutler, Abo, and Sangre de Cristo for-
mations.
Taxonomic evaluation of the undescribed captorhinid materials of
New Mexico has necessitated a reexamination of the type specimens
of Puercosaurus obtusidens Williston (1916). The partial left dentary,
designated by Williston as the holotype, is not only indeterminate, but
also provides no basis for considering it conspecific with the two partial,
crushed skulls referred by him to the species. Even though the two
referred skulls are undoubtedly captorhinids, they are too poorly pre-
served to be assigned to an established or new taxon. Under these
circumstances P. obtusidens is judged a nomen dubium. On the other
hand, the specimens collected by the University of California, Berkeley,
are sufficiently well preserved and unique to be referred to a new genus
and species, Rhiodenticulatus heatoni. With the exception of the types
of this species, all other Lower Permian captorhinid specimens from
New Mexico are too incomplete to recommend assignment to existing
or new taxa. Yet, they exhibit sufficient variation to indicate that the
group was probably quite diverse and widely distributed in New Mexico
during the Early Permian.
1986
Berman and Reisz— Permian Captorhinid Reptiles
3
Throughout the text the abbreviations CM, FMNH, and UCMP are used to refer to
collections of the Carnegie Museum of Natural History, Field Museum, Chicago, and
the Museum of Paleontology, University of California, Berkeley, respectively.
Systematic Paleontology
Class Reptilia
Order Cotylosauria
Suborder Captorhinomorpha
Family Captorhinidae
Genus Puercosaurus Williston, 1916
Puercosaurus obtusidens Williston, 1916, nomen dubium
Puercosaurus obtusidens Williston, 1916:189-192, fig. 37A~D.
Remarks. -—The original description of Puercosaurus obtusidens Wil-
liston (1916) was based on poorly preserved and incomplete speci-
mens—an incomplete dentigerous left dentary, FMNH 743, designated
as the holotype and two severely crushed skulls, FMNH 745, referred
to the species (Fig. 6; only one of the skulls is figured). Williston (1916)
illustrated the mandible and one of the two skulls, but a partial recon-
struction of the skull was based on both skulls. Although the specimens
were collected from the Lower Permian Cutler Formation near Arroyo
de Agua in the Rio Puerco drainage area, north-central New Mexico,
the holotypic dentary is from the well known Miller bonebed (see
Langston, 1953, for description of locality), whereas the referred skulls
were apparently found at least several kilometers away along the Rio
Puerco (Williston, 1916). The holotypic dentary is too poorly preserved
and incomplete to be reasonably certain that it belongs to that family.
Further, the holotypic dentary and referred skulls do not exhibit any
unique features in common which would demonstrate that they are
conspecific. In view of these circumstances P. obtusidens is declared
here a nomen dubium. Though the skulls FMNH 745 are sufficiently
complete to recognize their captorhinid affinities, assignment to either
a known or new species is not possible.
Genus Rhiodenticulatus^ new genus
Type species. — Rhiodenticulatus heatoni, new species.
Etymology.— Prom the Greek rhio, nose, and denticulatus, with small teeth, referring
to the relatively small teeth of the premaxilla.
Diagnosis. —Small captorhinid that differs from all other single-tooth-
rowed captorhinids in the following features: 1) premaxillary dentition
reduced to three teeth which are subequal in size and equal to or smaller
than precanine maxillary teeth; 2) reduction of maxillary dentition to
1 1 teeth; 3) number of precanines reduced to two; 4) extremely large
4
Annals of Carnegie Museum
VOL. 55
A
Fig. \.—Rhiodenticulatus heatoni, holotype, UCMP 35757. Skull in A, lateral, B, dorsal,
and C, ventral views. Abbreviations: a, angular; art, articular; bo, basioccipital; d, den-
tary; f, frontal; j, jugal; 1, lacrimal; m, maxilla; n, nasal; p, parietal; pa, prearticular; pf,
postfrontal; po, postorbital; pp, postparietal; prf, prefrontal; pt, pterygoid; q, quadrate;
qj, quadratojugal; s, stapes; sp, splenial; sq, squamosal. Scale = 1 cm.
1986
Berman and Reisz— Permian Captorhinid Reptiles
5
single canine with basal diameter as much as twice that of any post-
canine; 5) very broad lacrimal with a height (measured at the highest
level of the dorsal expansion of the maxilla) to length (shortest distance
between orbit and naris) ratio of .65 to .73; and 6) prefrontal extends
far anteriorly to a level about 84 to 90% of the distance from the orbit
to the naris. Distinguished from Labidosaurus and Eocaptorhinus by
its proportionately narrower skull postorbitally. Straight occipital mar-
gin of skull table separates it from Romeria which has a bilateral parietal
embayment and from Labidosaurus and Eocaptorhinus which have a
median embayment. Differs from Romeria and Protocaptorhinus in
having a long, low rectangular quadratojugal with a longitudinal length
that is approximately four times the height. Pointed postcanine teeth
of Rhiodenticulatus heatoni are distinguishable from the blunt teeth of
Labidosaurus and Eocaptorhinus.
Rhiodenticulatus heatoni, new species
Etymology.— ISlsLmQd. in honor of the late Malcolm J. Heaton in recognition of his
significant contributions to our understanding of the morphology and phylogenetics of
the Captorhinidae.
Holotype. — UCMP 35757: partial, articulated skeleton that includes
skull with closely joined lower jaw, vertebral series with ribs from the
axis to the seventh caudal, pectoral and pelvic girdles, right humerus
and proximal ends of ulna and radius, femora, left tibia, fibulae, and
tarsi; skull not attached to postcranial skeleton.
Paratypes. —UCMP 40209: skull with closely joined lower jaw, miss-
ing left postorbital cheek region and posterior half of left mandible.
UCMP 40210: partial, articulated postcranial skeleton preserved in
three small segments: 1) a series of seven postaxial cervical and dorsal
vertebrae with ribs, essentially complete pectoral girdle, and proximal
ends of humeri; 2) series of six vertebrae that includes the last two
presacrals, two sacrals with ribs, and the first two caudals, and pelvis;
and 3) portion of the left hindlimb, including proximal two thirds of
femur and nearly complete tibia. It is quite likely that UCMP 40209
and UCMP 40210 belong to the same individual.
Horizon and locality. — All specimens are from the Cutler Formation
exposures of the Rio Puerco drainage, Rio Arriba County, north-central
New Mexico. An Early Permian Wolfcampian age is generally accepted
for these exposures. Although the holotype and paratypes are listed as
coming from UCMP Camp quarry locality V-2814, Langston (1952:
98) notes that they were probably not found in the main bone level of
the quarry, but rather as float on the slope of Loma Salazar a few feet
away and presumably at or just above the quarry bone level. The Camp
quarry is located in SW1/4NE1/4NE1/4 sec. 8, T. 22 N,, R. 3 E., about
6
Annals of Carnegie Museum
VOL. 55
Fig. 2,—Rhiodenticulatus heatoni, paratype, UCMP 40209. Skull in A, lateral, B, dorsal,
and C, ventral views. Scale = 1 cm.
1 . 1 km southeast of Arroyo de Agua. All three specimens are preserved
in red, indurated concretionary nodules.
Description
Skw//. —Specimens of Rhiodenticulatus heatoni exhibit the general
structural pattern seen in all captorhinids and, therefore, aside from a
1986
Berman and Reisz— Permian Captorhinid Reptiles
7
few structures, Figs. 1 , 2 eliminate the need for a detailed description
of its anatomy. The skulls of the holotype UCMP 35757 and paratype
UCMP 40209 have suffered little distortion, but most of the superficial
features of the skulls, such as sculpturing, have been lost due to weath-
ering and excessive preparation performed prior to this study. In UCMP
40209 the left postorbital region was removed in the late 1930s in an
attempt to study the braincase in thin section. The extent of ossification
of the appendicular and axial portions of the holotypic skeleton suggests
that it is a mature individual. The skulls are triangular, with the post-
orbital width being only about 80 to 82% of the midline length. The
occipital margin of the skull stable is straight.
The downtumed premaxilla possesses three teeth. In the paratype
UCMP 40209 (Fig. 2A) the anterior end of the right maxilla greatly
overlaps the lateral surface of the maxillary process of the premaxilla,
making it appear as though the third premaxillary tooth originates from
the anterior end of the maxilla. Although imperfectly preserved, the
premaxillary teeth obviously had the shape of sharply pointed pegs,
were subequal in size, and were approximately the same size as, or
even possibly slightly smaller than, the precanine maxillary teeth. An-
teriorly the maxilla forms the ventral rim of the naris, gradually expands
to a moderate midlength dorsal swelling, and then tapers to a posterior
terminus at, or just short of, the level of the posterior margin of the
orbit. The right and left maxillae of the holotype possess 1 0 and 1 1
teeth, respectively, whereas both maxillae of UCMP 40209 possess 1 1.
In both skulls the third tooth forms an extremely large canine relative
to any of the other marginal teeth, with a basal diameter equal to, or
greater than, twice that of any of the postcanines. In the holotype the
precanines are slightly larger than the largest postcanines. The post-
canines exhibit a steady decrease in size posteriorly. As in the pre-
maxilla, the maxillary teeth have the form of sharply pointed pegs. In
neither skull is it possible to observe directly that only a single row of
marginal maxillary teeth is present. Indirect evidence for a single row
is present, however, in that the teeth form a straight row along the
outermost margin of the jaw, the postcanines exhibit a steady decrease
in size, and there does not appear to be sufficient space for an additional
tooth row on the alveolar shelf of the maxilla.
The lacrimal is unusual in being very broad. The ratio of its height
(measured at the level of the dorsalmost expansion of the maxilla) to
length (measured as the shortest distance between the orbit and naris)
is about .65 in the holotype and about .73 in UCMP 40209. There is
a correspondingly narrower lateral exposure of the prefrontal as a result
of the expanded height of the lacrimal. The prefrontal is also very long
and extends anteriorly along the dorsal margin of the lacrimal to a level
that is about 90 and 84% of the distance between the orbit and the
naris in the holotype and UCMP 40209, respectively. A long ventral
8
Annals of Carnegie Museum
VOL, 55
process of the prefrontal can be seen in the holotype extending along
the medial margin of the lacrimal on the anterior orbital rim. The
prefrontal and postfrontal are separated by only a small lateral process
of the frontal on the dorsal rim of the orbit. The frontals have a long,
narrow rectangular outline. Measured from the level of their orbital
contribution, the length of the anterior portion of the frontal is almost
one and one half times that of the posterior portion. The pineal opening
in both skulls is large and positioned anterior of the midlength of the
union of the parietals. The supratemporals are not preserved in either
skull. The presence of the postparietal is indicated only in the holotype
and then only as an impression of its ventral surface; its suture with
the parietal is therefore uncertain. The anterior ends of the right jugals
of both skulls appear to wedge between the lacrimal and maxilla, rather
than forming the step-like sutural encroachment onto the lateral surface
of the dorsal margin of the maxilla seen in other captorhinids (Heaton,
1979). This is undoubtedly due to imperfect preservation, however,
inasmuch as the standard condition is present on the nonfigured left
side of the holotypic skull. The quadratojugal has the outline of a long,
low rectangle, with the length exceeding the height by about four times.
Description of the palate is limited by the attached jaws. As in all
captorhinids there is no ectopterygoid, and the rectangular palatine
probably extends posteriorly to the subtemporal fossa. The presence
of a medial jugal process cannot be determined. The denticle fields of
the palate are preserved only in the paratype UCMP 40209. There is
a scattering of denticles along the posterior border of the transverse
flange of the pterygoid. There are also two faint, denticle bearing ridges;
one extends along the medial border of the palatal ramus of the pter-
ygoid, and a second extends obliquely anterolaterally across the palatal
ramus of the pterygoid and onto the palatine. The three columns of
irregularly arranged denticles converge toward the basicranial articu-
lation. Denticles also appear to be present on the parasphenoid.
The braincases of the holotype and UCMP 40209 are exposed in
ventral and occipital views and, though poorly preserved for the most
part, do not appear to exhibit any noteworthy differences from the
standard captorhinid construction. Both stapes of the holotype and the
right of UCMP 40209 are exposed in ventral view and are well enough
preserved to deserve comment. Though the footplates are not fully
exposed, they appear to conform closely to those of Ecocaptorhinus
(Heaton, 1979) and Captorhinus (Fox and Bowman, 1966). It has the
form of a broadly oval disk that thins toward its periphery. The disk
is drawn out posterolaterally into a cone-like structure, with the apex
being smoothly continuous with the columella. The cross-sectional
shape of columella, which remains unchanged throughout its short
length, is that of a mediolaterally flattened blade having a vertical height
1986
Berman and Reisz— Permian Captorhinid Reptiles
9
about three times its horizontal width. A large stapedial foramen pierces
the proximal end of the columella at a slightly anteromedial angle from
the vertical. Occipital view of the holotypic skull (not drawn) clearly
reveals the dorsal process of the left stapes just distal to the stapedial
foramen. It is very narrow, tapers to a point distally, and curves slightly
medially.
The mandibles of both skulls are visible in partial lateral view and
in ventral view; their sutural pattern and shape show no deviation from
those of other captorhinids. The posterior ends of the mandibles are
too damaged to determine whether or not a retroarticular process was
present. Dentary teeth are visible only in the holotype, but unfortu-
nately only the anterior half of the series is visible, and these are only
partially exposed. The first three teeth exhibit a marked increase in
size posteriorly, with the third tooth probably being the largest of the
entire series. On the basis of basal diameter, the fourth and fifth teeth
are slightly smaller than the third, whereas the sixth appears to be equal
to the third in size. The seventh and eighth decrease further in size, as
undoubtedly does the remaining unexposed portion of the series. It is
estimated that the dentary of the holotype held 14 or 15 teeth.
Postcranial skeleton. — Whereas the skull of Rhiodenticulatus heatoni
exhibits notable differences from those of other captorhinids, the op-
posite appears to be true of the postcranial skeleton; this is not unex-
pected inasmuch as this characterizes the history of captorhinids (Hea-
ton and Reisz, 1980).
The holotype appears to possess a complete, articulated vertebral
column from the axis to the sixth caudal vertebra (Fig. 3). Unfortu-
nately, the column is exposed only in ventral view, and small segments
of the series are hidden by the pectoral and pelvic girdles. Despite this,
it can be safely estimated that the entire presacral column consisted of
25 vertebrae. The centra are slightly pinched laterally, and except for
what is believed to be the axial centrum the ventral midlines are still
broadly rounded in transverse section; the axial centrum has a distinct
keel-like ventral midline. The wing-like transverse processes exhibit a
gradual reduction in their lateral extent posteriorly in the column. The
ventral surface of the processes slope anteroventrally, and the lateral
width narrows as the processes extend to the anterior rim of the cen-
trum. Both ends of the centra are slightly beveled to accommodate the
intercentra, giving them a slightly keystone appearance in lateral view.
The intercentra are variably displaced dorsally into the notochordal
canals of the centra, where attempts to fully expose them would result
in damage to the centra. As a result, many of the intercentra appear
to be absent, whereas those that are partially exposed vary in size and
have a lozenge-shaped outline. The first chevron occurs between cau-
dals three and four.
10
Annals of Carnegie Museum
VOL. 55
1986
Berman and Reisz— Permian Captorhinid Reptiles
11
The string of seven postaxial cervical and dorsal vertebrae of UCMP
40210 are exposed in dorsal view only (Fig. 4 A) and undoubtedly
include postaxial cervicals. Although poorly preserved, the neural arch-
es exhibit the swollen appearance so typical of captorhinids. The neural
spines are barely developed and appear as mere nubbins. The zyg-
apophyses are widely spaced from the midline, giving the neural arches
the typical lateral expansion of captorhinids. The transverse processes
extend laterally beyond the zygapophyses. Only the badly weathered
neural arches are exposed in the UCMP 40210 vertebral series which
includes the second to last presacral to the second caudal (not figured),
and they reveal no important differences from the far anterior presacral
of the same specimen.
The ribs of the holotype and paratype UCMP 40210 are moderately
well preserved, but the expansion of the heads is rarely visible, and
the shafts frequently appear as narrow rods. The heads of the postaxial
cervical ribs appear to be holocephalous and articulate in part with the
intercentra. The rib shafts of the cervicals of UCMP 40210 are ex-
panded into blade-like structures, whereas the more posterior rib shafts
of the holotype are subcircular in cross-section. The ribs of the anterior
half of the presacral column are more strongly curved posteroventrally
than those of the posterior half The sacral ribs are straight, thick, and
greatly expanded distally. The anterior caudal ribs of the holotype are
fused to the centra, curve strongly posteriorly, are thicker than the
presacral ribs, and quickly decrease in length more posteriorly in the
column.
The greater portions of the pectoral girdles are preserved in both the
holotype and paratype UCMP 40210, and together they exhibit most
of the important features of this structure (Figs. 3A, B, 4B). The head
of the interclavicle is roughly diamond-shaped, and the long, thin stem
is nearly complete in UCMP 40210, missing only a small part of the
distal end. The ventral plates of the clavicles are not complete, but
impressions on the interclavicles indicate that they were broad and
met medially; there is also no indication of a prominent, thumb-like
posterior process diverging from the main body of the ventral plate as
has been described in Labidosaurus (Williston, 1917) and Captorhinus
(Holmes, 1977). The narrow dorsal stem is directed abruptly dorsally
Fig. 3.~Rhiodenticulatus heatoni, holotype, UCMP 35757. A, ventral view of postcranial
skeleton, B, right lateral view of shoulder region, and C, lateral view of left hindlimb.
Abbreviations: as, astragalus; ax, axis; cal, calcaneum; cl, clavicle; cor, coracoid; cr, caudal
rib; cth, cleithrum; f, femur; fi, fibula; h, humerus; id, interclavicle; Ic, lateral centrale;
of, obturator foramen; r, radius; sc, scapula; sr, sacral rib; t, tibia; u, ulna; 2, 4, 5, distal
tarsals; iii, iv, v, metatarsals. Scales = 1 cm.
12
Annals of Carnegie Museum
VOL. 55
Fig. A.—Rhiodenticulatus heatoni, paratypc, UCMP 40210. A, dorsal view of series of
seven far anterior presacral vertebrae with ribs, and B, ventral view of pectoral girdle
with proximal ends of humeri preserved in a single nodule.
at nearly a right angle to the ventral plate. As in Captorhinus (Holmes,
1977), a distinct, posteriorly directed flange-like expansion of the ven-
tral half of the dorsal stem for the clavicular deltoid muscle is clearly
seen in the holotype. What may be a portion of the cleithrum is present
on the distal end of the dorsal stem of the right clavicle of the holotype.
There are no visible sutural divisions of the endochondral portion of
the pectoral girdle. The scapular blade curves dorsally rather abruptly
from the essentially horizontal coracoid plate. The anterior and pos-
terior margins of the scapular blade are essentially straight and parallel
to each other except for the anterodorsal comer being broadly curved.
The anterior coracoid portion expands a short distance anteriorly be-
yond the scapular blade as a smoothly rounded plate. A coracoid fo-
ramen located ventromedially to the anterior buttress of the glenoid
and a supraglenoid foramen on the posterior margin of the lateral
surface of the scapular blade just above the supraglenoid buttress are
clearly visible in the holotype and UCMP 40210.
Essentially all that is visible of the pelves of the holotype and UCMP
40210 is the worn ventral surface of the puboischiadic plate (Fig. 3 A);
the less complete pelvis of UCMP 40210 is not figured. In both spec-
imens osssification along the puboischiadic suture appears to be com-
plete in that there are no open spaces. The sutural division between
1986
Berman and Reisz— Permian Captorhinid Reptiles
13
the pubis and ischium is barely discemable in the holotype. The anterior
border of the puboischiadic plate is moderately concave. The ischium
is slightly longer and narrower than the pubis. A short distance from
the ventral rim of the acetabulum the pubis is perforated by the ob-
turator foramen.
The humerus is best represented in the holotype (Fig. 3A, B). It is
poorly preserved, but exhibits the same general configuration as those
of Captorhinus and Eocaptorhinus except that the shaft and distal head
have a more slender appearance. Its length, about 1.8 cm, is approx-
imately 90% of that of the femur. All other forelimb elements are either
too incomplete to comment on or are absent. The hindlimb and pes
are preserved only in the holotype (Fig. 3A, C). The preservation of
the femora allows recognition of only some of the major features of
this element. Except in being considerably more slender, particularly
the shaft, the femur is very similar to that of Captorhinus. It is about
2.0 cm long, has a minimum shaft diameter of about 1.3 mm, and a
maximum width of the distal head of 4.3 mm. The head appears rather
massive, with a well developed intertrochanteric fossa. The popliteal
area is a smooth, broadly concave depression. Though the internal
trochanter is well developed, there appears to no distinct step or notch
between it and the head. The tibiae and fibulae of the holotype are
present, but only those of the left limb are well preserved. The tibia
and fibula appear to be identical to those of Captorhinus except in
being noticeably more slender. In typical primitive reptilian fashion
the tibia is much shorter, 1 1.0 mm, than the femur, roughly 55% of
its length. The mediolateral width of the massive proximal end is about
45% of the length, whereas anteroposterior width of the distal end is
about 36% of the length; the narrowest mediolateral width of the shaft
is about 0.8 mm. A deep groove divides the anterior face and articular
surface of the expanded proximal end; the groove is bounded medially
by a prominent cnemial crest. The lateral margin of the tibia is bowed
slightly medially away from the fibula. The left fibula is about 12.3
mm long and the mediolaterally expanded proximal and distal ends
are about 2.5 and 3.5 mm wide, respectively; the narrowest mediola-
teral width of the shaft is about 1.1 mm. The medial margin of the
fibula is strongly concave and the lateral margin only very slightly
convex, giving it the appearance of being bowed laterally away from
the tibia.
The tarsi of the holotype are well ossified. The right, exposed in
ventral view (Fig. 3A), is nearly complete, missing only the first distal
tarsal, whereas the left is represented only by the dorsally exposed
calcaneum and astragalus (Fig. 3C). The tarsal elements conform closely
to the pattern seen in Captorhinus (Peabody, 1951) except for two
apparent deviations; the fourth distal tarsal is relatively smaller and
14
Annals of Carnegie Museum
VOL. 55
the fifth which is relatively larger than in Captorhinus. The typical
pattern in primitive reptiles is for the fourth distal to be considerably
larger than the other distal tarsals. In Rhiodenticulatus, however, the
fourth distal is roughly equal in size to the fifth. The extreme proximal
ends of the third, fourth, and fifth metatarsals are all that remains of
the rest of the right pes.
Discussion
Placement of Rhiodenticulatus heatoni within the Captorhinidae of
the suborder Captorhinomorpha is unquestionable. It should be made
clear, however, that we follow Heaton (1979), Gaffney and McKenna
(1979), Reisz (1980), and Heaton and Reisz (in press) in the assignment
of genera in the two recognized captorhinomorph families, the Early
Pennsylvanian to Early Permian Protorothyrididae (=Romeriidae of
many authors) and Early to Late Permian Captorhinidae. The capto-
rhinids are differentiated from the protorothyridids by their low, wide,
massive skull, hooked premaxillae, loss of tabulars and ectopterygoids,
fully ossified paroccipital processes, stoutly built postcranial skeleton,
25 presacral vertebrae with swollen neural arches and low neural spines,
absence of cleithra, thumb-like process on the ventral plate of clavicle,
short stoutly built limbs, absence of a supinator process of humerus,
and wide manus and pes. Presently, about 1 4 genera of captorhinids
are recognized. Among these, however, only four genera, Romeria,
Protocaptorhinus, Eocaptorhinus, and Labidosaurus, could conceiv-
ably be confused with Rhiodenticulatus, because they possess single-
rowed, marginal dentitions.
Clark and Carroll (1973) and Heaton (1979) presented nearly iden-
tical phylogenies in which the series of successively later occurring,
single-tooth-rowed captorhinids Romeria, Protocaptorhinus, and
Eocaptorhinus forms a single, continuous, phylogenetic lineage de-
picting transitional morphological stages that links the protorothyridids
with the later occurring, multiple-tooth-rowed Captorhinus. Among
the captorhinids with multiple-rowed marginal dentitions, Captorhinus
is the only genus known in great detail and is also generally accepted
as the most primitive. Labidosaurus, the least understood of the single-
tooth-rowed captorhinids, is not included in Heaton’s (1979) phylo-
genetic scheme, but is depicted in Clark and Caroll’s (1973) phyloge-
netic tree as the end member of an offshoot from Protocaptorhinus.
Gaffney and McKenna (1979:7) criticized the systematic methodology
used by Clark and Carroll, and Heaton as being “stratophenetic” {sensu
Gingerich, 1976) in which “similar morphologies are arranged strati-
graphically and connected using usually implicit rather than explicit
criteria, to form what are interpreted as ancestor-descendant lineages.”
1986
Berman and Reisz— Permian Captorhinid Reptiles
15
Gaffney and McKenna, without altering the basic phylogenies of Clark
and Carroll, and Heaton, reexpressed them in the form of a cladogram
and, thus, as a testable hypothesis. Our only serious reservation of their
cladogram of the Captorhinidae is the position of Labidosaurus as a
member of the clade containing Protocaptorhinus. As brought out be-
low, Labidosaurus shares with Eocaptorhinus and Captorhinus several
derived features of the skull not seen in Protocaptorhinus. Further,
restudy of Labidosaurus is greatly needed before its phylogenetic rel-
tionships can be accurately evaluated. Despite this, the cladogram of
Gaffney and McKenna presents a reasonable understanding of the evo-
lutionary relationships of the captorhinids and, along with the detailed
morphological studies of the known captorhinomorphs by Carroll and
Baird (1972), Clark and Carroll (1973), Heaton (1979), and Olson
(1984), provides a basis for assessing the polarity of several character
states of Rhiodenticulatus heatoni.
The maxillary dentitions of the holotype and paratype UCMP 40209
of Rhiodenticulatus are unique among the single-tooth-rowed capto-
rhinids in possessing: 1) a single, extremely large canine whose basal
diameter is equal to, or greater than, twice that of any of the postcanines;
2) 1 1 teeth; and 3) two precanines. It can also be noted that among the
protorothyridids only the Pennsylvanian Cephalerpeton exhibits a sim-
ilar specialization toward a greatly reduced number (16) of maxillary
teeth that includes a low number (four) of precanines (Reisz and Baird,
1983). Protorothyridids typically possess, as does Romeria, a pair of
prominent, subequal canines, yet their basal diameters are far less than
twice that of the largest postcanines. Although a single tooth may be
designated as a canine in Protocaptorhinus and Labidosaurus, it is not
as prominent as either of the paired canines of Romeria. Eocaptorhinus
also exhibits a single, prominent canine, and although the first through
third postcanines may be noticeably shorter, their basal diameters are
only slightly smaller than that of the canine. In the holotypic skull of
Rhiodenticulatus, having a midline length of about 38 mm, the basal
diameter of the canine is about 2.2 mm. This is larger in both absolute
and relative measurements than the canines of Romeria and Proto-
captorhinus, in which the basal diameters range from roughly 1.2 to
1.7 mm for skulls 50 to 53 mm in midline length. On the other hand,
though the maximum basal diameter of the canines in Eocaptorhinus
and Labidosaurus may be as much as 2.6 and 3.0 mm, respectively,
their midline skull lengths are as much as two and four times greater
than that of Rhiodenticulatus.
Previous authors (Clark and Carroll, 1973; Heaton, 1979) have noted
that there is a general reduction in the number of maxillary teeth in
successively later occurring, single-tooth-rowed captorhinids. Approx-
16
Annals of Carnegie Museum
VOL. 55
imate maxillary tooth counts for Protorothyris, Romeria, Protocapto-
rhinus, Labidosaurus, mid Eocaptorhinus are 24-30, 22-23, 18-22, 14-
18, and 17-22, respectively. The maximum of 11 maxillary teeth in
Rhiodenticulatus can only be interpreted as a unique derived character.
Probably related to this trend is the unique occurrence in Rhiodentic-
ulatus of only two precanines. Protorothyridids typically possess five
precanines, but as many as seven or eight have been described in
Paleothyris (Carroll, 1969). A further slight reduction in the number
of precanines occurs in the successively later occurring captorhinids;
Romeria prima possesses six precanines, R. texana, Protocaptorhinus,
and Labidosaurus four or five, Eocaptorhinus three or occasionally
four, and Captorhinus three or rarely four.
The lacrimal of Rhiodenticulatus may be unique among all capto-
rhinomorphs in having an unusually large height to length ratio. The
height was measured at the level of the dorsalmost expansion of the
maxilla, whereas the length was taken as the shortest distance between
the orbit and naris. Despite the small errors expected in making these
sorts of measurements, the height to length ratios of .65 and .73 for
the holotype UCMP 35757 and paratype UCMP 40209, respectively,
are considerably greater than those of other single-tooth-rowed cap-
torhinids, which range from about .25 to .40. In the protorothyridids
Paleothyris and Protorothyris, the lacrimals are very long and narrow,
and have a height to length ratio of about .17. The fact that in Rhio-
denticulatus the ratio is smaller for the larger holotype than for the
paratype, suggests that the ratio decreases somewhat with growth or
increase in size. This notion is reinforced in Romeria texana, where
the ratios for an adult and juvenile described by Clark and Carroll
(1973) are .27 and .40, respectively.
The extreme anterior extent of the prefrontal along the dorsal margin
of the lacrimal in Rhiodenticulatus also sets it apart from all other
captorhinids. In the holotype UCMP 35757 and paratype UCMP 40209
the prefrontal extends anteriorly to a level that is 90 and 84% of the
distance from the orbit to the naris, respectively, whereas in other
captorhinids and in Protorothyris this measurement ranges from ap-
proximately 43 to 58%. It might be suspected that the greater anterior
extension of the prefrontal in Rhiodenticulatus is due to removal, either
as a result of weathering or mechanical preparation, of that portion of
the nasal overlying its anterior end. In Eocaptorhinus, for example,
where additional exposure of the prefrontal could conceivably increase
its preorbital length by as much as 28% (Heaton, 1979), the anterior
extension of the prefrontal would increase from about 44 to 56% of
the distance between the orbit and naris. As pointed out by Heaton
(1979), in Clark and Carroll’s (1973) illustration and reconstruction of
1986
Berman and Reisz— Permian Captorhinid Reptiles
17
the holotype of Romeria texana the prefrontals appear unusually long
due to the exposure of that portion of their anterior ends normally
overlapped by the nasals. For this reason we used Heaton’s (1979)
reconstruction of Romeria texana in calculating the relative anterior
extension of the prefrontal. In the holotype and paratype UCMP 40209
of Rhiodenticulatus both pairs of prefrontals have identical lateral ex-
posure configurations, strongly suggesting that they have not been sig-
nificantly distorted in this way by weathering or excessive preparation.
Rhiodenticulatus exhibits several shared derived features with other
advanced single-tooth-rowed captorhinids. Its possession of only three
premaxillary teeth is considered advanced among the captorhinids in
view of the general trend within the captorhinomorphs toward reduc-
tion in the number of premaxillary teeth. Protorothyridids typically
have five or six premaxillary teeth, although Protorothyris archeri ap-
pears to have four and Cephalerpeton only three (Reisz and Baird,
1983). Premaxillary tooth counts for Romeria prima, R. texana, Pro-
tocaptorhinus, Labidosaurus, and Eocaptorhinus are 4, 5, 4 or 5, 3,
and 4 or 5, respectively. The premaxilla of Captorhinus typically pos-
sesses four teeth and rarely three or five. Rhiodenticulatus is also similar
to the more derived captorhinids Labidosaurus, Eocaptorhinus, and
Captorhinus in having a long, narrow, rectangular quadratojugal in
which the longitudinal length exceeds by almost four times the height,
and the dorsal margin tends to be straight. As pointed out by Heaton
(1979), in the more primitive Romeria and Protocaptorhinus the dorsal
margin of the quadratojugal tends to be more convex. Heaton also
noted that in the reconstruction of Romeria prima by Clark and Carroll
(1973) this feature is erroneously exaggerated and is actually not sig-
nificantly different from that of R. texana and Protocaptorhinus. More
notable, however, is the shorter length of the quadratojugals of Romeria
and Protocaptorhinus, so that the length exceeds the height by no more
than two and one half times. The quadratojugals of the protorothyridids
tend to be more like those of the more primitive captorhinids. The
straight occipital margin of the skull table of Rhiodenticulatus, seen
also in Protocaptorhinus, is a derived feature with respect to the bi-
lateral parietal embayment of the occipital margin of Romeria and the
protorothyridids. On the other hand, Rhiodenticulatus is viewed as
primitive with respect to the median embayment of the occipital mar-
gins of Labidosaurus, Eocaptorhinus, and Captorhinus.
Rhiodenticulatus exhibits at least two characters that link it with the
more primitive captorhinids Romeria and Protocaptorhinus, and ex-
clude it from the more advanced Labidosaurus, Eocaptorhinus, and
Captorhinus. It has been noted by several authors (Clark and Carroll,
1973; Heaton, 1979) that in the evolution of the captorhinids there is
18
Annals of Carnegie Museum
VOL. 55
a marked trend toward relative widening of the postorbital region of
the skull. In Labidosaurus, Eocaptorhinus, and Captorhinus, the post-
orbital lateral expansion of the skull becomes so pronounced that the
lateral margin of the skull in dorsal view is noticeably concave, whereas
in Romeria, Protocaptorhinus, and Rhiodenticulatus it is essentially
straight. Labidosaurus, Eocaptorhinus, and Captorhinus are advanced
over Romeria, Protocaptorhinus, and Rhiodenticulatus in exhibiting
the shared derived feature of blunt (rather than sharply pointed) post-
canine maxillary teeth (Olson, 1984).
Finally, there is one unique feature of Rhiodenticulatus with respect
to all other single-tooth-rowed captorhinids which on first consider-
ation seems unquestionably primitive, its possession of small premax-
illary teeth of subequal size. In all captorhinids the premaxillary teeth
exhibit a steady but dramatic increase in size anteriorly, with the an-
terior teeth reaching sizes equal to, or greater than, the maxillary canine.
Though Rhiodenticulatus is like its protorothyridid predecessors in this
character, implying a primitive state, the alternative interpretation that
it represents an evolutionary reversal is argued below.
On the basis of the above character state analysis we conclude that
the most plausible relationship of Rhiodenticulatus heatoni to other
captorhinids is that depicted by the cladogram of Fig. 5 in which it is
the primitive sister taxon to Labidosaurus, Eocaptorhinus, and Cap-
torhinus (plus all other multiple-tooth-rowed forms). We recognize,
however, that the cladogram possesses a few weaknesses. First, several
of the nodes are defined by only a single character. Second, there are
at least two notable contradictions between the cladogram and the
character state analysis presented. Perhaps the most obvious is the
possession by Rhiodenticulatus of small, subequal premaxillary teeth.
The cladogram requires that this character be interpreted as the result
of a secondary reduction in tooth size, or an evolutionary reversal,
rather than more simply, as our character analysis implies, a primitive
character. The likelihood that such an event occurred, however, seems
very reasonable in light of the several derived modifications of the
dentition of Rhiodenticulatus noted: 1) a single, extremely large canine,
2) reduction of the maxillary dentition to 1 1 teeth, 3) reduction in the
number of precanines to two, and 4) reduction of the premaxillary
dentition to three teeth. Of these, the first three are judged unique to
Rhiodenticulatus among the single-tooth-rowed captorhinids, whereas
the last also occurs in Labidosaurus. It should be noted here, however,
that in our opinion it seems quite likely that the reduction in the number
of premaxillary teeth to three in Rhiodenticulatus and Labidosaurus
was achieved independently given the otherwise marked differences
between their dentitions. A second possible inconsistency between the
Romeria Protocaptorhinus Rhiodenticulatus Labidosaurus Eocaptorhinus Captorhinus
19
1986
Berman and Reisz— Permian Captorhinid Reptiles
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Annals of Carnegie Museum
VOL. 55
placement of Rhiodenticulatus in the cladogram and our character state
analysis concerns the unique derived features of its dentition. If, as
suggested above, the extremely large, single canine, 1 1 maxillary teeth,
and two precanines of Rhiodenticulatus represent the most advanced
stages of general trends within the single-tooth-rowed captorhinids,
then it could be argued that these features indicate an advanced sister
taxon relationship with Labidosaurus and Eocaptorhinus as well. This
interpretation is rejected, however, in favor of the alternative argument
that these unique features of the dentition of Rhiodenticulatus, as well
as its relatively small, few premaxillary teeth, probably reflect an ad-
aptation to a specialized diet not present in the other single-tooth-
rowed captorhinids.
Other New Mexico Captorhinids
In recent years the authors have collected additional captorhinid
remains from the Lower Permian Cutler, Abo, and Sangre de Cristo
formations at widely scattered localities in northern and central New
Mexico. Although these specimens, as well as the two crushed and
incomplete skulls referred to ''Puercosaurus obtusidens^^ by Williston
(1916), are too poorly preserved to be assigned safely to an existing
taxon or made the basis of a new one, they permit the recognition of
at least three possible morphotypes, one each from the Cutler, Abo,
and Sangre de Cristo formations. These specmiens are, therefore, im-
portant as indicators of the diversity and spatial range of the capto-
rhinids in the Lower Permian of New Mexico.
Indeterminate Cutler Captorhinid
All the indeterminate captorhinid specimens from Cutler Formation
of the Rio Puerco drainage, Rio Arriba County, in the north-central
part of the state are considered together as though pertaining to a single
form distinct from Rhiodenticulatus heatoni of the same area. This is
done despite the fact that the indeterminate specimens exhibit some
differences from each other. It is realized that future discoveries may
indicate that the differences between them may be due to either the
presence of more than one undescribed species, or distinct growth stages
of the same species, or both. If conspecificity is being masked by onto-
genetic growth stages, then it is also conceivable that one or more of
the indeterminate Cutler specimens may prove to be conspecific or
congeneric with R. heatoni. This possibility is given some support by
the presence in a few of the unassigned Cutler specimens of at least
one feature considered derived in R. heatoni, the single, greatly enlarged
canine. The unassigned Cutler specimens include:
FMNH 745, two crushed and very incomplete skulls referred to '' Puercosaurus ob-
tusidens” by Williston (1916), who illustrated only one, the same skull shown here in
1986
Berman and Reisz— Permian Captorhinid Reptiles
21
Fig. 6.—'‘'Puercosaurus obtusidens'' Williston (1916). A, dorsal, and B, ventral views of
referred skull FMNH 745. C, lateral view of holotypic dentary FMNH 743. Abbrevia-
tions: d, dentary; f, frontal; j, jugal; m, maxilla; pf, postfrontal; po, postorbital; prf,
prefrontal; pt, pterygoid; qj, quadratojugal; sq, squamosal. Scale = 1 cm.
Fig. 6A, B. Their exact locality is unknown, and according to Williston (1916) they were
found by Mr. Miller in 1911 on the Rio Puerco a few miles below Arroyo de Agua. The
holotypic left dentary of “P. obtusidens'" (Fig. 6C) is too incomplete to assign to the
Captorhinidae with reasonable certainty.
22
Annals of Carnegie Museum
VOL. 55
Fig. 7.— Indeterminate captorhinid from the Cutler Formation. A, lateral view of partial
left maxilla, B, lateral view of posterior portion of left dentary, and C, lateral and dorsal
views of anterior portion of right dentary of CM 28592. D, partial skull CM 28591
showing mainly paired frontals in dorsal view, dentaries in ventral view, and small
portion of left maxilla in both medial and lateral views. Abbreviations: d, dentary; f,
frontal; m, maxilla. Scale = 1 cm.
1986
Berman and Reisz— Permian Captorhinid Reptiles
23
CM 28591, a partial skull (Fig. 7C, D); CM 28589, fourteen dorsal vertebrae, most of
which are articulated in strings of two or three, and associated fragments of ribs and
appendicular elements (Fig. 8A). These vertebrae are indistinguishable from those of R.
heatoni, as are those of most captorhinids, but are included here because they were found
in very close proximity to CM 28591 in NEV4SWV4NEy4 sec. 5, T. 22 N., R. 3 E. about
1.5 km northeast of Arroyo de Agua.
CM 28592, partial left maxilla (Fig. 7A), small portion of both dentaries (Fig. 7B),
presacral vertebra, and left humerus (Fig. 8C). These elements undoubtedly belong to a
single individual and were collected in N^ASW'ASE^A sec. 8, T. 22 N., R. 3 E. about 1.6
km southeast of Arroyo de Agua.
The left premaxilla of the figured skull of FMNH 745 (Fig. 6) appears
to have held four teeth as Williston (1916) described; this estimate
takes into account an unoccupied space. The premaxillary teeth, as in
Rhiodenticulatus, are very small relative to the pre- and postcanines
of the maxilla. Accounting for spaces, the maxilla of FMNH 745 held
approximately 13 to 15 teeth, including two or possibly three preca-
nines, one extremely large canine, and 10 or 11 postcanines that de-
crease gradually in size posteriorly. As in Rhiodenticulatus, the basal
diameter of the canine is about twice that of any of the postcanines.
The dentition of the partial left maxilla of CM 28592 (Fig. 7 A) is
considerably different, however, in that the canine is relatively smaller
when compared to the postcanines, and the third or posteriormost
precanine is nearly as large as the canine, producing a double canine
appearance. A segment of the right maxilla of CM 28591 (Fig. 7D)
shows the canine as dominating the postcanines in size, though not as
greatly as in Rhiodenticulatus. The maxillary dentitions of FMNH 745,
CM 28591, and CM 28592 are single rowed, and the teeth appear as
simple, sharply pointed pegs except for a slight, posterior curvature of
the tips. The frontals of FMNH 745 and CM 28591 (Figs. 6B, 7D) are
complete, and their very narrow contribution to the orbital rim is
clearly discemable. As in Rhiodenticulatus, the portion of frontal an-
terior to its contribution to the orbital rim is considerably larger than
that which is posterior. In FMNH 745 the pineal foramen appears to
be more centrally positioned along the median parietal suture than in
Rhiodenticulatus. The dentary dentition is well preserved in CM 2859 1
except for most of the teeth lacking their tips; the more complete right
dentary is estimated to have held about 18 teeth. The first tooth is
extremely small in typical captorhinid fashion, the second and third
are subequal in size and much larger than the others of the series, and
the following teeth do not exhibit an obvious size pattern except for
the last three being greatly reduced. The anterior seven teeth preserved
on the fragment of right dentary of CM 28592 (Fig. 1C) exhibit the
same size relationships as in CM 28591. In contrast, the first five teeth
of the left dentary of the FMNH 745 are of subequal, moderate size.
The dentary teeth also have the form of simple, sharply pointed pegs.
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Annals of Carnegie Museum
VOL, 55
Fig. 8,— Indeterminate captorhinid from Cutler Formation, Presacral vertebrae of A,
CM 28589, and B, CM 28592, C, dorsal proximal and distal ventral views of left humerus
CM 28592. Scale = 1 cm.
The presacral vertebrae of CM 28589 and CM 28592 (Fig. 8 A, B)
are alike and as far as comparisons will allow like those of Rhioden-
ticulatus. The neural spine is small, triangular in lateral view, and
distinctly set off from the neural arch, which has the expected swollen
appearance. The zygapophyses extend slightly beyond the lateral mar-
gins of the centra, and their articular facets are essentially horizontal.
There is no evidence of a suture between the neural arch and centrum.
The transverse process is positioned on the anterodorsal quadrant of
the lateral surface of the centrum. In lateral view the process is a thin,
ridge-like structure whose base extends anteroventrally to the centrum
1986
Berman and Reisz— Permian Captorhinid Reptiles
25
Fig. 9, —Indeterminate captorhinid CM 41707 from the Abo formation. A, medial view
of left maxilla, B, lateral view of left jugal, C, dorsal view of distal half of right femur,
and D, distal ventral view of left humerus. Scale = 1 cm.
rim. In anterior view its lateral projection diminishes as it extends to
the centrum rim, giving it a wing-like appearance. The ends of the
centra are beveled slightly so as to give them a slightly keystone ap-
pearance in lateral view. The lateral surfaces of the centra are mod-
erately concave in horizontal section, producing a spool-shaped ap-
pearance. The only clearly visible intercentrum is seen in the vertebra
of CM 28592 (Fig. 8B); it has a low, narrowly triangular outline in
lateral view and a crescent- shaped outline in anterior view.
The only appendicular element of the indeterminate specimens from
the Cutler Formation worthy of description is the well preserved left
humerus of CM 28592 (Fig. 8C). It differs from those of Eocaptorhinus
and Captorhinus (Holmes, 1977) mainly in having a more gracile form,
but in this feature is also like that of Rhiodenticulatus. The proximal
and distal ends are relatively narrower, and the entepicondyle extends
far more distally beyond the radial condyle than in Eocaptorhinus or
Captorhinus.
Indeterminate Abo Captorhinid
A second possible New Mexico captorhinid form for which there is
insufficient morphological information to assign to either an existing
or a new taxon is based on a single specimen, CM 41707, collected
from the Abo Formation about 20 km northeast of Socorro in the
central part of the state in SEV4NEV4Wy4 of sec. 14, T. 2 S., R. 3 E. CM
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Annals of Carnegie Museum
VOL. 55
41707 consists of disarticulated elements of the skull and postcranial
skeleton of an individual that are randomly associated and densely
concentrated in a small, strongly indurated, red concretion. Only those
elements close to the outer surface of the concretion were prepared and
include: a left maxilla, left jugal, anterior half of the right mandible, a
presacral vertebra, ribs, greater part of the left humerus, distal half of
right femur, and several unidentified fragments. Only a few of the above
elements are figured here (Fig. 9). The left maxilla (Fig. 9A), although
poorly preserved, retains an accurate outline of its dentition, which
consists of 14 teeth and at least one unoccupied space; in this feature
it is like the maxilla of the indeterminate Cutler specimen FMNH 745.
As in Rhiodenticulatus, there are two moderate sized precanines. Al-
though the third tooth is the largest of the series and should be con-
sidered a canine, the fourth tooth is nearly as large, giving CM 41707
a distinctly double canine appearance like that in the partial left maxilla
of the indeterminate Cutler specimen CM 28592. The basal diameter
of neither canine of CM 41707, however, exceeds that of the largest
postcanine as greatly as does the single, enlarged canine of Rhioden-
ticulatus. The teeth gradually decrease in size from the first canine to
the seventh tooth; this is followed first by four somewhat larger, sub-
equal teeth and then by the last three and smallest teeth of the series.
The jugal (Fig. 9B) is like that of other captorhinids. A smooth flange
on the dorsal margin of the posterior plate clearly indicates the position
of the overlaping postorbital, and the spike-like projection on the pos-
terior margin marks the point of separation between the jugal-squa-
mosal and jugal-quadratojugal contacts.
The anterior half of the right mandible (not figured) is exposed in
lateral and dorsal view, and the first 1 6 teeth are present, though many
are represented by only their bases. As in the captorhinids Eocapto-
rhinus and Captorhinus, the first tooth is extremely small, the second
moderate sized, and the third is greatly enlarged and dominates the
entire series, having a basal diameter of about 2 mm and a height of
about 5 mm. The fourth tooth is the second largest of the series, with
a basal diameter of about 1.5 mm and an estimated height of 2.5 mm,
whereas the fifth is greatly reduced and about equal to the second in
size. Teeth 6, 7, and 8 are of subequal, moderate size, the larger ninth
tooth appears to have been about the size of the third tooth, and the
remaining seven teeth steadily decrease in size posteriorly. All the
dentary teeth appear to have the form of simple pointed pegs and are
aligned in a single row. As in Eocaptorhinus and Captorhinus, the first
three teeth lean obliquely forward and the fourth is nearly vertical. The
one partial vertebra appears to be typical of captorhinids. The left
humerus of CM 41707 (Fig. 9D) is nearly complete, missing only a
1986
Berman and Reisz— Permian Captorhinid Reptiles
27
B
Fig. 10. —Indeterminate captorhinid from the Sangre de Cristo Formation. Lateral and
ventral views of maxillary fragments A, CM 28594, and B, CM 28595. Scale = 1 cm.
portion of its proximal end. The shaft is more slender and the entepi-
condyle possibly less expanded than those of Eocaptorhinus or Cap-
torhinus. The entepicondyle extends distally only slightly beyond the
radial condyle. In contrast, the distal half of the right femur (Fig. 9C)
is very stoutly constructed, especially in comparison with Eocaptor-
hinus and Captorhinus.
Indeterminate Sangre de Cristo Captorhinid
Numerous fragments of captorhind maxillae and dentaries have been
collected from the Lower Permian Sangre de Cristo Formation ap-
proximately 50 km southeast of Santa Fe in the northeastern part of
the state in NEV4 sec. 36, T. 14 N., R. 13 E. Two of the maxillary
fragments are figured (Fig. 10), and they clearly indicate that the cap-
torhinid from this locality had at least two rows of teeth. In one (CM
28594, Fig. lOA) the broken edges of a second row of teeth can be seen
lateral to the posterior end of the main row. Medial wear facets of the
teeth, which are more evident on the other fragment (CM 28595, Fig.
lOB), give them the same blunt, peg-like outlines seen in Eocaptorhinus
and multiple-tooth-rowed forms such as Captorhinus. These two fea-
tures of the dentition indicate clearly that the Sangre de Cristo cap-
torhinid remains are of a distinct and more advanced taxon than the
other representatives of the family in New Mexico.
Acknowledgments
Support for this research was provided by grants from the New Mexico Bureau of
Mines and Mineral Resources (to D.S B. and R.R.R.), the M, Graham Netting Research
Fund through a grant from the Cordelia Scaife May Charitable Trust (to D.S B.), and
the Natural Sciences and Engineering Research Council of Canada (to R.R.R.). We are
grateful to Ms. Diane Scott, Erindale Campus, University of Toronto, for the preparation
of the specimens and the drawing of the illustrations. We are obliged to the University
of California, Berkeley, and the Field Museum of Natural History for the loan of spec-
imens. Special thanks are extended to Dr. Wann Langston, Jr., of the University of Texas,
Austin, for bringing these specimens to our attention and expediting their loan to us.
Thanks are due Drs. Robert L. Carroll of McGill University, Montreal, and E. C. Olson
of the University of California, Berkeley, for critical reading of the manuscript.
28
Annals of Carnegie Museum
VOL, 55
Literature Cited
Carroll, R. L, 1969. A Middle Pennsylvanian captorhinomorph, and the interrela-
tionships of primitive reptiles. J. Paleont., 43:151-170.
Carroll, R. L., and D. Baird. 1972. Carboniferous stem-reptiles of the Family Ro-
meriidae. Bull. Mus. Comp. ZooL, Harvard Univ., 143:321-363.
Clark, J., and R. L. Carroll. 1973. Romeriid reptiles from the Lower Permian. Bull.
Mus. Comp. Zool., Harvard Univ., 144:353-407.
Fox, R. C., AND M. C. Bowman. 1966. Osteology and relationships of Captorhinus
aquti (Cope) (Reptilia: Captorhinomorpha). Univ. Kansas Paleont. Contrib., Ver-
tebrata, 11:1-79.
Gaffney, E. S., AND M. C. McKenna. 1979. A Late Permian captorhinid from Rho-
desia, American Mus, Novit, 2688:1-15.
Gingerich, P, D. 1976. Cranial anatomy and evolution of early Tertiary Plesiadapidae
(Mammalia, Primates). Univ. Michigan, Papers Paleont., 15:1-141.
Heaton, M. J. 1979. Cranial anatomy of primitive captorhinid reptiles from the Later
Pennsylvanian and Early Permian OWahoma and Texas. Bull. Oklahoma Geol.
Survey, 127:1-84.
Heaton, M. J., and R. R. Reisz, 1980. A skeletal reconstruction of the Early Permian
captorhinid reptile Eocatorhinus laticeps (Williston). J. Paleont., 54:136-143.
. In press. The interrelationships of captorhinomorph reptiles. Canadian J. Earth
Sci.
Holmes, R. 1977. The osteology and musculature of the pectoral limb of small cap-
torhinids. J. Morph., 152:101-140.
Langston, W., Jr. 1952. Permian vertebrates of New Mexico. Unpublished Ph.D.
dissert., Univ. California, Berkeley, 212 pp.
. 1953. Permian amphibians of New Mexico, Univ, California Publ., Geol. Sci.,
Bull., 29:349-416.
Olson, E. C. 1 984. The taxonomic status and morphology of Pleuristion brachyocoelus
Case; referred to Protocaptorhinus pricei Clark and Carroll (Reptilia: Captorhino-
morpha). J. Paleont., 58:1282-1295.
Peabody, F. E. 1951. The origin of the astragalus of reptiles. Evolution, 5:339-344.
Reisz, R. R. 1 980. A protorothyridid captorhinomorph reptile from the Lower Permian
of Oklahoma. Royal Ontario Mus., Life Sciences Contrib., 121:1-16.
Reisz, R. R., and D. Baird. 1983. Captorhinomorph “stem” reptiles from the Penn-
sylvanian coal-swamp deposit of Linton, Ohio. Ann, Carnegie Mus., 52:393-411.
Williston, S. W. 1916. The osteology of some American Permian vertebrates, 11.
Univ. Chicago, Walker Mus. Contrib., 1:165-192.
. 1917. Labidosaurus Cope, a lower Permian cotylosaur reptile from Texas. J.
Geol., 25:309-321.
NPi
ISSN 0097-4463
ANNALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 23 MAY 1986 ARTICLE 2
DESCRIPTION OF THE LOWER JAW OF STEGOSAURUS
(REPTILIA, ORNITHISCHIA)
David S Berman
Associate Curator, Section of Vertebrate Fossils
John S. McIntosh^
Research Associate, Section of Vertebrate Fossils
Abstract
Description of a well preserved lower jaw of Stegosaurus reveals previously unknown
structural details. Although comparisons between the lower jaw of Stegosaurus and those
of other stegosaurs are greatly limited, they strongly suggest a close comformity. The
lower Jaw of Stegosaurus exhibits the most primitive grade of organization among the
omithischians with one noticeable exception, the retention of an intercoronoid in cer-
atopsians.
Introduction
During the past several years there has been a renewed effort by the
Carnegie Museum of Natural History to prepare the remaining dinosaur
materials collected by that institution over 60 years ago (1909-1922)
at what is now Dinosaur National Monument. During the routine
preparation of one of the blocks, field no. 83, an almost complete,
excellently preserved left mandible of Stegosaurus was unexpectedly
discovered. Earl Douglass, who was in charge of the quarrying oper-
ations, had identified the contents of field no. 83 as ""Dinosaur, vertebra,
rib, etc.,” and preparation of the block was, therefore, subsequently
30
Annals of Carnegie Museum
VOL. 55
postponed because it was thought to contain little of importance. Al-
though Stegosaurus is one of the most common of the Morrison di-
nosaurs, skull material is extremely rare. To date, only one complete
skull and lower jaw have been found, that of the type specimen of
Stegosaurus stenops (USNM 4934) from the Felch quarry (YPM quarry
1; see Ostrom and McIntosh, 1966) at the entrance of Garden Park,
about 13 km north of Canyon City, Colorado. It was originally de-
scribed briefly by Marsh (1887) and later in much greater detail by
Gilmore (1914). Several braincases of Stegosaurus have been found,
but the anterior half of a right dentary (USNM 4935) from the “stego-
saur” quarry (YPM quarry 13; see Ostrom and McIntosh, 1966), about
6.5 km east of Como Bluff, Wyoming, is otherwise the only other
mandible known. The mandible described herein lacks only the pre-
dentary and teeth, and is noteworthy in exhibiting structural details
heretofore unknown in other specimens.
The following abbreviations are used to refer to repositories of specimens: CM, Car-
negie Museum of Natural History; USNM, National Museum of Natural History; and
YPM, Yale Peabody Museum.
Systematic Paleontology
Class Reptilia
Order Omithischia Seeley, 1888
Suborder Stegosauria Marsh, 1877
Family Stegosauridae Marsh, 1880
Genus Stegosaurus Marsh, 1877
Stegosaurus sp.
Specimen.— CM 41681, left mandible lacking predentary and den-
tition.
Horizon.— Morrison Formation, Late Jurassic.
Locality.— 'Dinosnnr National Monument (originally called the Car-
negie quarry), about 10 km north of Jensen, Uintah County, Utah. CM
41681 was found under the fourteeth cervical vertebra of the type
skeleton of Apatosaurus louisae Holland, CM 30 1 8, at the co-ordinates
F to G- 5 4 on the quarry map published by Gilmore (1936).
Collector.— EdcrX Douglass and party, 1910.
Assignment.— T>oscrrpXions and illustrations by Marsh (1887) and
Gilmore ( 1 9 1 4) of the holotypic lower jaw of Stegosaurus stenops USNM
4934, although containing several errors that will be brought out in
the text below, and by Gilmore (1914) of the right dentary of Stego-
saurus sp. USNM 4935 leave little or no doubt about the assignation
of CM 41681 to this genus. It can also be noted that there exists no
known dinosaur from the Late Jurassic whose lower jaw could be
confused with that of Stegosaurus.
1986
Berman and McIntosh— ^'recFOS'^c/Rt/^ Jaw
31
Description
As Figs. 1-3 indicate, the left mandible CM 4 1 68 1 is complete except
for the absence of the one unpaired element of the lower jaw, the median
predentary, and the dentition. The mandible is well preserved and the
sutures delineating the seven elements that comprise it, except those
of the articular, are clearly discemable. The mandible is narrowly com-
pressed in transverse section. In lateral or medial view the mandible
is shallow, exhibiting a gradual dorsal and ventral expansion posteriorly
from the symphysis to a maximum depth at about three-fifths its length,
then gradually narrows at about the same rate posteriorly to the end
of the retroarticular process. Whereas the expansion of the dorsal mar-
gin is somewhat angular in outline, that of the ventral margin is smooth-
ly convex. There is no development of a distinct coronoid process, but
rather there is a low coronoid eminence whose apex is formed by the
surangular. The plate-like anterior end of the mandible turns abruptly
medially and slightly ventrally, and with the symphysis inclined pos-
teroventrally the joined mandibles would have a scoop-like appearance.
It has not been possible to identify the small rectangular plate of bone
adhering to the medial surface of the anterior end of the dentary; it
might be part of the adjoining left dentary.
The sutures of the lateral surface of the mandible (Fig. 1) are, for the
most part, clearly defined and need little comment. The articular-sur-
angular suture is not evident, and a portion of the angular-surangular
suture is restored on the basis of impression. The dentary, surangular,
and angular border a very large, horizontally elongate, oval mandibular
foramen. A very thin, short strip of the splenial is visible on the mid-
ventral margin of the mandible. Three shallow, parallel grooves occur
shortly posterior of the point where the anterior end of the dentary
curves medially. Each deepens slightly as it extends 1 to 2 cm antero-
dorsally from the inferior jaw margin and ends at a small foramen that
pierces the dentary at a very low angle. Three relatively small foramina
are located near the dorsal border of the surangular; two closely spaced
smaller ones are located a considerable distance posterior to a larger
one. They undoubtedly penetrate to the medial side of the surangular.
Gallon (1974) has described similar foramina in the omithischian di-
nosaur Hypsilophodon and suggested that they may have transmitted
cutaneous branches of the inferior alveolar nerve as in modem lizards
(Oelrich, 1956).
In medial view (Fig. 2) all seven elements of the mandible are visible.
The dentary is the largest of the jaw elements. Its upper half, above
the Meckelian canal, is remarkably thick, having a triangular cross-
sectional outline with the apex directed medially to form a dorsal,
shelf-like surface (Fig. 3). Posteriorly much of the medial edge of the
32
Annals of Carnegie Museum
VOL. 55
1986
Berman and McIntosh— (75 Jaw
33
dorsal shelf is occupied by a series of 20 closely spaced alveoli. The
remaining anterior portion of the medial edge of the shelf takes the
form of a thin, knife-like ridge that continues to the symphysis. The
slightly concave dorsal shelf of the dentary faces dorsomedially at its
posterior end, but because of a gradual twisting as it is followed an-
teriorly, the shelf faces directly dorsally just beyond its midlength and
then anteriorly at the symphysis. Along most of the length of the dorsal
shelf, mainly at the level of the alveoli, is a series of about nine foramina
of varied sizes, which might be interpreted as mental foramina. The
posteriormost pair of foramina are located at the anterior end of a
prominent, 3 cm long groove that deepens anteriorly. The orientation
of the alveolar series suggests that the teeth were directed mainly me-
dially and only slightly dorsally. Directly below the series of alveoli is
a series of foramina; they are apparently arranged so that each foramen
lies opposite an alveolus, but they do not occur the entire length of the
alveolar series.
The Meckelian canal of the dentary is exposed medially as it emerges
from beneath the anterior margin of the splenial, becoming progres-
sively shallower as it extends nearly to the symphysis. The large, sub-
rectangular, plate-like splenial has a sinuous, overlapping suture with
the dentary anteriorly. There is a deep, narrow emargination of the
anterior border of the splenial at the level of the Meckelian canal. The
splenial-prearticular contact is also very sinuous; a moderate sized
inframandibular foramen is located on this suture. The posterodorsal
comer of the splenial appears to enter narrowly the rim of the adductor
fossa. Its ventral margin contacts the dentary along the ventral edge of
the mandible, whereas more posteriorly it wraps a short distance around
the ventral edge to contact the angular on the lateral surface of the
mandible. A large foramen of unknown function penetrates the splenial
in a posterodorsal direction near its anteroventral border. What may
be a large foramen is located near the middorsal margin of the splenial.
A small portion of the angular wraps around the ventral edge of the
mandible and is visible in medial view. Although a sutural contact
between the prearticular and articular cannot be discerned, the prear-
ticular undoubtedly formed the greater part of the ventromedial border
and the articular the posterior border of the large, oval adductor fossa.
The coronoid is roughly triangular in medial view, with its longer side
forming a substantial portion of the dorsal margin of the mandible. It
also contributes greatly to the anteromedial border of the adductor
Fig. 1.— Photograph and illustration of left mandible Stegosaurus CM 41681 in lateral
view. Abbreviations: A, angular; C, coronoid; D, dentary; SA, surangular; SP, splenial.
34
Annals of Carnegie Museum
VOL. 55
1986
Berman and McIntosh— ^rE^GO^^c/i? c/5 Jaw
35
fossa. There is no indication that an intercoronoid was present. Just
posterior to the coronoid there is a small, triangular, thickened area
on the medial surface of the surangular apex of the coronoid eminence.
This thickened area undoubtedly marks the site of concentrated at-
tachment of jaw adductor musculature. The slightly concave articular
surface of the articular is subcircular in outline. At its posteromedial
border is a stout, dorsally directed triangular process. The retroarticular
process is short and bluntly rounded.
Comparison and Discussion
The Stegosaurus mandible CM 41681 exhibits three features of par-
ticular interest that are not noted or shown in earlier descriptions or
illustrations of the genus (Marsh, 1887; Gilmore, 1 9 14): a large, external
mandibular foramen; presence of a large coronoid bone; and a very
weakly developed coronoid process with the apex formed by the sur-
angular.
Several orders of reptiles have small fenestrae on the external surface
of the mandible, but the presence of a large external mandibular fo-
ramen located at the intersection of the dentary, angular, and suran-
gular, as in Stegosaurus, is rare. Such a foramen is found in crocodiles,
prosauropods, theropods, and some thecodonts, but among omithis-
chians it has been reported only in the primitive omithopod Fabro-
saurus (Thulbom, 1970) and the stegosaur Huayangosaurus (Dong et
al., 1982; Zhou, 1983). The external mandibular foramen of Fabro-
saurus and CM 41681 are very similar except that in the former the
dentary forms the anterior half or more of its margin. Huayangosaurus,
from the Middle Jurassic of China, is the only stegosaur in which an
external mandibular foramen has previously been described (Dong et
al., 1982; Zhou, 1983). Although the formaen in Huayangosaurus is
relatively smaller than that in CM 4 1 68 1 , it has the same general outline
shape and precisely the same position and relationships to the sur-
rounding dentary, surangular, and angular. Owen (1863) did not find
evidence of the mandibular foramen in the primitive armored omith-
ischian Scelidosaurus, and recent preparation using modem chemical
techniques has clearly indicated the absence of the foramen (Charig,
1979). Scelidosaurus has often been considered a stegosaur, but Charig
(1979:126), who is currently restudying this animal, has remarked that
Fig. 2.— Photograph and illustration of left mandible of Stegosaurus CM 4168 1 in dorsal
view. Abbreviations: AL, alveoli; AR, articular; C, coronoid; D, dentary; PA, preartic-
ular; SA, surangular.
1986
Berman and McIntosh— Jaw
37
“It has often been regarded as an ancestral stegosaur; it might be an
ancestral ankylosaur; it could be neither.”
Gilmore (1914) presumed that a separate coronoid bone was present
in Stegosaurus, but was unable to verify this on the basis of the then
only known complete jaw of the genus, that of the holotype of S. stenops
(USNM 4934). It is, therefore, gratifying that relatively large coronoid
is clearly evident in CM 41681. Owen (1863) stated that the lower jaw
of SceUdosaurus includes a coronoid, but did not describe it or indicate
it in his figures (pis. 46, 47). Although the presence of a coronoid has
not been demonstrated in other stegosaurs, there is no basis for sug-
gesting its absence. On the contrary, there is increasing evidence that
a coronoid element was present in all omithischians except the had-
rosaurs. It has been explicitly reported in the omithopods Hypsilopho-
don (Gallon, 1974), Camptosaurus (Gilmore, 1909), Iguanodon (Dol-
lo, 1883), and Ouranosaurus (Taquet, 1976), in the ceratopsians
Protoceratops, Montanoceratops, Centrosaurus, and Triceratops (Brown
and Schlaikjer, 1940), in the Edmontonia (Gilmore, 1930;
Russell, 1940) (=Panoplosaurus of Coombs, 1978), and in the pachy-
cephalosaur Stegoceras (Gilmore, 1924). The presence of a second
coronoid element, the intercoronoid, has also been found by Brown
and Schlaikjer (1940) in the Protoceratopsidae and Ceratopsidae. The
intercoronoid, which apparently occurs in all saurischians (for example,
Plateosaurus, Brachiosaurus, Camarasaurus, AUosaurus, Tyrannosau-
rus, and others), has not been reported in any other omithischian group,
and we are unable to find any evidence for its presence in CM 41681.
Among the omithischians the coronoid of the ankylosaur Edmon-
tonia (Gilmore, 1930; Russell, 1940) resembles most closely that of
Stegosaurus in shape, size, and position, but important differences are
evident. The anterior extension of the coronoid in Edmontonia lies
medial to the posterior end of the tooth row, whereas in Stegosaurus
it lies lateral to the tooth row. In Edmontonia, as well as in other
ankylosaurs, and the pachycephalosaur Stegoceras (Gilmore, 1 924) the
surangular is expanded dorsally into a broadly rounded but prominent
coronoid process; only in Edmontonia, however, is the coronoid clearly
shown to curve upward onto the leading edge of the process. In Stego-
saurus, on the other hand, the dorsal margin of the mandible rises
gradually and evenly to the apex of the very low coronoid eminence
whose apex is formed by the surangular. With the exception of Fabro-
Fig. 3. —Photograph and illustration of left mandible of Stegosaurus CM 4 1 68 1 in medial
view. Abbreviations: A, angular; C, coronoid; D, dentary; PA, prearticular; SA, suran-
gular; SP, spleniaL
38
Annals of Carnegie Museum
VOL. 55
saurus (Thulbom, 1970), omithopods possess a large, shaft-like cor-
onoid process formed mainly by the dentary, that extends dorsally
from the main body of the bone; the summit of the process may even
curve anteriorly. There is a suboval coronoid on the medial surface of
the process except in hadrosaurs, which appear to lack this element
(Lull and Wright, 1942; Ostrom, 1961). In Fabrosaurus the dentary
forms the apex of a low coronoid eminence; there is no distinct coronoid
process. Within the omithopods there is an apparent trend toward
reduction and loss of the coronoid. The coronoid is relatively large in
the primitive Fabrosaurus and Hypsilophodon, but in advanced forms
it is greatly reduced, as in Ouranosaurus, and apparently lost in had-
rosaurs.
Only one difference between CM 41681 and previous descriptions
of the lower jaw of Stegosaurus is possibly noteworthy. The Stegosaurus
right dentary USNM 4935 possesses 23 alveoli (Gilmore, 1914), where-
as that of CM 41681 has 20. Whether this should be regarded as an
individual variation or as having taxonomic significance is not known.
It can be noted, however, that the USNM 4935 right dentary is ap-
proximately 20% larger than that of CM 41681, and the difference in
their tooth counts may be related to size. What little is known about
the jaws of other stegosaurs suggests that they compare closely with
that of Stegosaurus. Though the descriptions of the jaw of Huayang-
osaurus (Dong et al., 1982; Zhou, 1983) are very brief, the accompa-
nying figures of its lateral surface suggest that it probably differs in only
minor ways from that of Stegosaurus— i\ may have a slightly more
prominent coronoid process, but with the apex still apparently formed
by the surangular; the teeth appear to be vertically oriented; a lateral
exposure of the coronoid is not indicated; and the anterior end of the
mandible curves more strongly ventrally. The stegosaurs Kentrurosau-
rus (Hennig, 1936), Tuojiangosaurus (Dong et al., 1983), and Chung-
kingosaurus (Dong et al., 1983) are known only from incomplete den-
taries. The dentary of the Kentrurosaurus resembles very closely that
of Stegosaurus in the position, shape, and development of the Meck-
elian canal, but the slightly larger alveoli suggest that it may have held
fewer teeth. All that can be said of the dentaries of the latter two genera
is that they were long and low, as in Stegosaurus.
The possession of a large external mandibular fenestra, large coro-
noid, and very weakly developed coronoid process in which the apex
is formed by the surangular can be justifiably interpreted as primitive
omithischian features, inasmuch as they occur in a wide range of the-
codonts. The above comparisons, therefore, indicate that the lower jaw
of Stegosaurus, as is probably true of other stegosaurs, exhibits the
most primitive grade of organization among the omithischians with
1986
Berman and McIntosh -■5'r£'GD5’^i7R (75 Jaw
39
one conspicuous exception, the retention of the intercoronoid in the
ceratopsians.
Acknowledgments
Support for this project was provided by a grant from the M. Graham Netting Research
Fund through a grant from the Cordelia Scaife May Charitable Trust (to D.S B.). We
are especially grateful to Ms. Amy Henrici for the difficult task of preparing the specimen
and to Ms. Nancy Perkins for drawing the illustrations; both are of Carnegie Museum
of Natural History. We also wish to thank Drs. Peter M. Galton of the University of
Bridgeport, Connecticut, John H. Ostrom of the Yale Peabody Museum of Natural
History, and Mary R. Dawson of the Carnegie Museum of Natural History for critically
reviewing the manuscript.
Literature Cited
Brown, B., and E. M. Schlaikjer. 1940. A new element in the ceratopsian jaw with
additional notes on the mandible. Amer. Mus. Novit., 1092: l-l 3.
Charig, a. J. 1979. A new look at dinosaurs. Mayflower Books, New York, 160 pp.
Coombs, W. P., Jr. 1978. The families of the omithischian dinosaur order Ankylo-
sauria. Palaeontology, 21:143-170.
Dollo, L. 1883. Quatrieme note sur les dinosauriens de Bemissart. Bull. Mus. Hist.
Nat. Belgique, 11:223-252.
Dong Zm, Tang Zl, and Zhou Sw. 1982. Note on the new mid-Jurassic stegosaur
from Sichuan Basin. Vertebrata Palasiatica, 20:85-87.
Dong Zm, Zhou Sw, AND Zhang Yh. 1983. The dinosaurian remains from Sichuan
Basin, China. Palaeont. Sin., New C Ser., 162:1-145.
Galton, P. M. 1974. The omithischian dinosaur Hypsilophodon from the Wealden
of the Isle of Wight. Bull. British Mus. (Nat. Hist.), 25:1-152.
Gilmore, C. W. 1909. Osteology of the Jurassic reptile Camptosaurus, with a revision
of the species of the genus and description of two new species. Proc. U.S. Nat. Mus.,
36:197-302.
— — 1914. Osteology of the armored dinosauria in the U.S. National Museum with
special reference to the genus Stegosaurus. Bull. U.S. Nat. Mus., 89:1-140.
■ 1924. On Troodon validus, an omithopod dinosaur from the Belly River Cre-
taceous of Alberta, Canada. Bull. Univ. Alberta, 1:1-43.
. 1930. On dinosaurian reptiles from the Two Medicine Formation of Montana.
Proc. U.S. Nat. Mus., 77:1-39.
1936. Osteology of Apatosaurus with special reference to specimens in the
Carnegie Museum. Mem. Carnegie Mus., 1 1:175-300.
Hennig, E. 1936. Ein dentale von Kentrurosaurus aethiopicus Hennig. Palaeontogr.,
Suppl. 7, 1:309-312.
Lull, R. S., and N. E. Wright. 1942. Hadrosaurian dinosaurs of North America.
Spec. Pap. Geol. Soc. Amer., 40:1-242.
Marsh, O. C. 1887. Principal characters of American Jursassic dinosaurs. Part IX, the
skull and dermal armor of Stegosaurus. Amer, J. Sci., 43:413-417.
Oelrich, T. M, 1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae).
Misc. Pubis. Mus. Zook, Univ. Michigan, 94:1-122,
Ostrom, J, H. 1961. Cranial morphology of the hadrosaurian dinosaurs of North
America. Bull. Amer. Mus. Nat. Hist., 122:1-186.
Ostrom, J. H,, and J. S. McIntosh. 1 966. Marsh’s dinosaurs. Yale Univ. Press, 388 pp.
Owen, R. 1863. A monograph of the fossil Reptilia of the Liassic formations. Part II.
Palaeontogr. Soc. Monogr., 13: 1-26.
40
Annals of Carnegie Museum
VOL. 55
Russell, L. S. 1940. Edmontonia rugosidens (Gilmore), an armoured dinosaur from
the Belly River series of Alberta. Univ. Toronto Stud. Geol. Ser., 43:1-28.
Taquet, P. 1976. Geologie et Paleontologie du Gisemement de gadoufaoua (Aptien
du Niger). Cahiers de Paleontologie, 191 pp.
Thulborn, R. a. 1970. The skull of Fabrosaurus australis, a Triassic omithischian
dinosaur. Palaeontology, 13:416-432.
Zhou Sw. 1983. A nearly complete skeleton of stegosaur from Middle Jurassic of
Dashanpu, Zigong, Sichuan. J. Chengdu College Geol., SuppL 1:15-26.
^ ISSN 0097-4463
ANNALS
of CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE » PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 23 MAY 1986 ARTICLE 3
BIOCHEMICAL AND MORPHOLOGICAL DIFFERENTIATION
IN SPANISH AND MOROCCAN POPULATIONS OF
DISCOGLOSSUS AND THE DESCRIPTION OF A
NEW SPECIES FROM SOUTHERN SPAIN
(AMPHIBIA, ANURA, DISCOGLOSSIDAE)
Stephen D. Busack^’^
Research Associate, Section of Amphibians and Reptiles
Abstract
Biochemical and morphological divergence among Moroccan and Iberian populations
suggests that populations of Discoglossus inhabiting these regions are not conspecific.
Northern Moroccan Discoglossus are assigned to D. pictus’, Discoglossus galganoi Capula
et al, 1985 inhabits the Iberian peninsula to the north of the Guadalquivir River basin,
and the Discoglossus population residing to the south of the Guadalquivir River basin
on Iberia is described as a new species. An evolutionary scenario for Iberian and Mo-
roccan populations, consistent with the biochemical and morphological data, is inferred
from the physiogeographic history of the region.
Introduction
Until recently only two species of Discoglossus were thought to in-
habit Europe. Spanish, French, and Sicilian populations were called D.
pictus, and populations inhabiting Corsica, Sardinia, Elba, and Monte
Argentario, Italy, were called D. sardus (Knoepffler, 1961a, 1961Z?,
1 962). Recent electrophoretic examinations of Discoglossus have, how-
^ Address: Fellow in Herpetology, California
San Francisco, California 94118.
^ Present address: Department of Genetics j
Urbana, Illinois 61801.
Submitted 1 May 1985.
Academy of Sciences, Golden Gate Park,
41
42
Annals of Carnegie Museum
VOL. 55
Fig. I.™ Localities of examined specimens of Discoglossus galganoi (squares), D. pictus
(triangles), and D. jeanneae (circle). Dotted line traces the presumed northern limit of
D. jeanneae', see Discussion for further explanation.
ever, revealed two additional species, D. montalentii from Corsica
(Lanza et al., 1984) and D. galganoi from north of the Guadalquivir
River basin in Spain and Portugal (Capula et aL, 1985).
My interest in Discoglossus was initially focused on the amount of
genetic differentiation that accumulated between European and African
populations after the formation, 5-7 million years ago, of the Strait of
Gibraltar (Busack, 1986). Because my electrophoretic study revealed
substantial genetic differentiation between Spanish and Moroccan sam-
pies, the morphologies of these populations were also compared. Bio-
chemical and morphological data indicate that Iberian populations to
the north and to the south of the Guadalquivir River basin are not
conspecific, and that neither is conspecific with populations inhabiting
northern Morocco. The data supporting this conclusion are presented
in the following pages, together with a description of a second species
from Spain, and a hypothetical reconstruction of the evolutionary his-
tory of Iberian and Moroccan forms of Discoglossus.
1986
Busack— Spanish and Moroccan Discoglossus
43
Table l.~~ Protein systems" examined by electrophoresis; enzymes are arranged by En-
zyme Commission number.
Protein (abbreviation)
Enzyme Commission
number
Electro-
phoretic
conditions
Albumin (Ab)
B
(Oxidoreductases)
Alcohol dehydrogenase (Adh)
1.1. 1.1
A
Glycerol- 3 -phosphate dehydrogenase (Gpd)
1.1. 1.8
D
L-Lactate dehydrogenase (Ldh)
1.1.1.27
F
Malate dehydrogenase (Mdh)
1.1.1.37
F
Malate dehydrogenase (Me)
1.1.1.40
F
Isocitrate dehydrogenase (led)
1.1.1.42
E
Phosphogluconate dehydrogenase (Pgd)
1.1.1.44
E
Glutamate dehydrogenase (Glud)
L4.1.3
D
Superoxide dismutase (Sod)
1.15.1.1
D
(Transferases)
Aspartate aminotransferase (Aat)
2.6.1. 1
D
Hexokinase (Hk)
2.7.1. 1
G
Creatine kinase (Ck)
2.7.3.2
G
(Hydrolases)
Esterase (Est)
3.1. 1.1
B
Esterase-D (Est-D)
3.1. 1.1
B
Acid phosphatase (Aep)
3.1.3.2
G
Fmetose-bisphosphatase (Hdp)
3.1.3.11
D
N - Acetyl-Beta-glucosaminidase (Hex)
3.2.1.30
G
Dipeptidase I, L-Leucyl-L-Alanine (La)
3.4.11
B
Dipeptidase III, L-Leucylglycyl-glycine (Lgg)
3.4.11
C
Dipeptidase IV, L-Phenylalanyl-L-Proline (Pap)
3.4.11
B
Adenosine deaminase (Ada)
3.5.4.4
A
(Lyases)
Fmctose-bisphosphate-aldolase (Aid)
4.1.2.13
H
Aconitate hydratase (Aeon)
4.2.1.3
E
(Isomerases)
Mannose-6-phosphate isomerase (Mpi)
5.3.1.8
E
Glucose-6-phosphate isomerase (Gpi)
5.3.1.9
F
Phosphoglucomutase (Pgm)
5.4.2.2
E
A = Histidine, pH 7.8 gel and electrode buffer (Harris and Hopkinson, 1976), 150v/3h.
B = LiOH A + B, pH 8.2 gel and LiOH A, pH 8.1 electrode buffer (Selander et al.,
1971), 300v/3h.
C = Poulik, pH 8.7 gel and borate, pH 8.2 electrode buffer (Selander et al., 1971),
250v/3h.
D = Tris citrate II, pH 8.0 gel and electrode buffer (Selander et al., 1971), 130v/4h.
E = Tris citrate II, pH 8.0 + NADP gel and tris citrate II, pH 8.0 electrode buffer
(Selander et al., 1971), 130v/4h.
F = Tris citrate III, pH 7.0 gel and electrode buffer (Ayala et al., 1972), 180v/3h.
G == Tris citrate III, pH 7.0 + 1 5% glycerine gel and tris citrate III, pH 7.0 electrode
buffer (Ayala et al., 1972), 180v/3h.
H ^ Tris citrate III, pH 7.0 + NAD + 2-mercaptoethanol gel and tiis citrate III, pH
7.0 electrode buffer (Ayala et al., 1972), 180v/3h.
44
Annals of Carnegie Museum
VOL. 55
Materials and Methods
Electrophoresis. —OnQ individual was collected from north of the Guadalquivir River
basin in Avila Province, Spain, at San Martin del Pimpollar (40®22'N, 5®03'W; Fig. 1,
square C). Seven specimens were collected south of the Quadalquivir River basin in
Cadiz Province, Spain (Fig. 1, circle), near the towns of Facinas (36®08'N, 5®42'W, 5
specimens) and Benalup de Sidonia (36®20'N, 5M9'W, 2 specimens). Specimens collected
in Tetouan Prefecture at Chechaouene (35®10'N, 5®16'W; Fig. 1, open triangle; 5 spec-
imens) and in the vicinity of Tleta Tarhremt (35®47'N, 5®28'W; Fig. 1, closed triangle;
5 specimens) represent northern Moroccan populations.
Specimens were sacrificed in the field and samples of heart and liver were removed,
frozen, and stored in liquid nitrogen (- 196®C). In the laboratory, tissues were transferred
to a freezer (-76®C) until used in electrophoresis two to 12 months later. Tissue samples
were pooled for each animal. Proteins were separated electrophoretically in horizontal
starch gels (11.5% hydrolyzed starch, Sigma Chemical Co.) and localized by standard
histochemical staining procedures (Ayala et al., 1972; Harris and Hopkinson, 1976;
Selander et al., 1971; Table 1). Genetic interpretations of allozymic data were based on
criteria developed by Selander et al. (1971). Multiple loci within a protein system were
numbered with “ 1 ” designating the most anodally migrating set of allelic products. Alleles
of a locus were lettered, with “a” representing the most anodally migrating product. Data
resulting from the electrophoretic analysis are summarized in Table 2.
Two methods were used to analyze genetic relationships among populations of Dis-
coglossus. The first distinguishes patterns of allele distribution among populations that
occur as a result of chance association from those that occur too frequently to be chance
phenomena. If all patterns of allele distribution are equally probable, the probability, P,
that a particular pattern will occur is 1/S„ where S„ is the sum of the number of possible
patterns of allele distribution. In the case of the three populations of Discoglossus being
compared, S„ = 7 for three items combined three at a time, two at a time, and one at a
time. The probability of observing a specific pattern of allele distribution, r, two or more
times is given by summing the terms of the binomial expansion:
b(r) = (7)/>'(l - Pr-,
where r is the number of replications seen for a given pattern of allele distribution, m
is the total number of alleles in the data set, and is the number of possible combi-
nations of m alleles taken r at a time (Straney, 1980; Patton and Smith, 1981).
The second method is the computation of estimates of, and standard errors for, the
unbiased minimum genetic distance {D\ Nei, 1978, 1971, respectively) between Disco-
glossus populations. Allele frequency data (Table 2) were used directly for the compu-
tation of genetic distances and their standard errors.
Morphology.— CB.puhL et al. (1985:tables 2, 4) published comprehensive tables of mor-
phological measurements for D. galganoi. I have used the data in these tables to elucidate
the morphological features of D. galganoi and to make direct assessments of morpho-
logical differentiation between D. galganoi and other taxa.
Spanish specimens I personally examined were representative of the same Cadiz Prov-
ince populations as my electrophoretic samples, but collected earlier (between 1969 and
1972). Moroccan specimens included those used for electrophoresis as well as additional
material from the same sites. Straight-line measurements of snout-urostyle (SUL), snout
(anterior comer of the eye to the tip of the snout), head (posterior angle of the jaw to
the tip of the snout), eye (horizontal diameter from posterior comer to anterior comer),
tibia, femur, hand (proximal aspect of the central metacarpal tubercle to the tip of the
third digit), and foot (proximal aspect of the metatarsal tubercle to the tip of the third
digit) lengths were taken to 0.1 mm with dial calipers. Head width (angle of jaws).
1986
Busack— Spanish and Moroccan Discoglossus
45
interorbital (between the anterior comer of the eyes), and intemarial (center to center)
distances were also recorded.
Frogs are sometimes sexually dimorphic in many characters and sexes were analyzed
separately. The effect of having limited numbers of individuals of each size representing
each sex of each population was minimized by converting each character measurement
to natural logarithms; the variance of In-transformed data estimates intrinsic variability
and is unaffected by size (Lewontin, 1966; Moriarty, 1977). Transformed data repre-
senting each character were subjected to covariance analysis in which snout-urostyle
length was selected as the independent variable. Although allometry is correctly assessed
only from the study of growth of an individual, I used individuals of different sizes from
a population to obtain estimates of allometric coefficients. Identification of dissimilarities
in the allometric growth influence by using transformed data is acceptable for comparing
populations (Thorpe, 1976).
Linear regression analysis, in which the measurement data were left untransformed,
was then performed for variables demonstrating significant differences in allometric
growth. For ease of presentation and interpretation, only the resulting slope and intercept
values are reported in Table 4. Significance levels for all statistical tests were set (a priori)
at 0.05 and probabilities are those for committing a Type I error in a two-tailed test.
Results
Biochemical Comparisons
Aatl, Aat2, Acp2, Ada, Est-D, Hdp, Me, Pgm, and Sod were mon-
omorphically expressed among all three populations I examined. Table
2 summarizes the distribution of allozymes at the 25 polymorphic loci
I was able to score unambiguously.
Populations of Discoglossus residing to the north and to the south
of the Guadalquivir River basin in Spain and those inhabiting northern
Morocco are genetically well differentiated. Fifteen of 87 alleles iden-
tified among these Discoglossus samples are shared among all popu-
lations. Spanish Discoglossus residing north of the Quadalquivir River
basin share 17 alleles with individuals from northern Morocco and 23
alleles with Discoglossus residing south of the Guadalquivir River basin
in Spain. Populations inhabiting the area south of the Guadalquivir
River basin share 29 alleles with Discoglossus inhabiting northern Mo-
rocco (Table 3).
Seventy alleles, however, differentiate between individuals of Dis-
coglossus from north of the Guadalquivir River basin in Spain and
those from northern Morocco, 64 alleles differentiate individuals from
north of the Guadalquivir River basin and those from south of the
Guadalquivir River basin in Spain, and 58 alleles differentiate indi-
viduals living to the south of the Guadalquivir River basin in Spain
from those inhabiting northern Morocco (Tables 2 and 3). The distri-
bution of these alleles contributes to a genetic distance {D) of 0.74 ±
0.18 between the Spanish sample from north of the Guadalquivir River
basin and the northern Moroccan samples, 0.39 ± 0.12 between Span-
ish samples residing to the north and to the south of the Guadalquivir
46
Annals of Carnegie Museum
VOL. 55
Table 2.— Genic variation within and among samples o/Discoglossus.
D. jeanneae
D. galganoi
D. pictiis
Number of specimens
1
1
10
Mean heterozygosity per lo-
cus
0.16
0.03
0.16
Percentage of loci polymor-
phic
38.2
2.9
44.1
Locus and alleles
Abl
a
0.86
1.00
0.95
b
0.14
0.00
0.00
c
0.00
0.00
0.05
Ab2
a
1.00
1.00
0.00
b
0.00
0.00
1.00
Aeon
a
0.57
0.00
1.00
b
0.29
0.00
0.00
c
0.14
0.00
0.00
d
0.00
1.00
0.00
Acpl
a
0.00
0.00
0.10
b
1.00
1.00
0.90
Adh
a
1.00
1.00
0.80
b
0.00
0.00
0.20
Aid
a
0.00
1.00
0.00
b
0.86
0,00
1.00
c
0.14
0.00
0.00
Ck
a
0.57
0.00
0.10
b
0.00
0.00
0.10
c
0.43
0.00
0.75
d
0.00
0.00
0.05
e
0.00
1.00
0.00
Est
a
0.00
0.00
0.20
b
0.00
0.00
0.40
c
0.29
0.00
0,40
d
0.14
0.00
0.00
e
0.57
1.00
0.00
Glud
a
0.00
1.00
0.20
b
1.00
0.00
0.80
Gpd
a
0.00
LOO
0.00
b
0.07
0.00
0.00
c
0.93
0.00
0.50
d
0.00
0.00
0.50
Gpi
a
1.00
1.00
0.90
b
0.00
0,00
0.10
Hex
a
0.00
0.00
0.10
b
1.00
1.00
0.00
c
0.00
0.00
0.90
Hkl
a
1.00
1.00
0.00
b
0.00
0.00
1,00
Hk2
a
0.00
1.00
0.00
1986
Bus ACK— Spanish and Moroccan Discoglossus
47
Table 2.— Continued.
D. jeanneae
D. galganoi
D. pictus
b
1.00
0.00
0.00
c
0.00
0.00
1.00
Icdl
a
0.00
0.00
0.10
b
0.86
0.00
0.85
c
0.14
0.00
0.05
d
0.00
1.00
0.00
Icd2
a
0.00
0.00
1.00
b
0.00
0.50
0.00
c
1.00
0.50
0.00
La
a
0.00
0.00
0.10
b
0.57
1.00
0.90
c
0.43
0.00
0.00
Ldhl
a
1.00
1.00
0.00
b
0.00
0.00
1.00
Ldh2
a
1.00
0.00
1.00
b
0.00
1.00
0.00
Lgg
a
0.00
0.00
1.00
b
1.00
1.00
0.00
Mdhl
a
0.14
0.00
0.00
b
0.29
0.00
0.00
c
0.43
0.00
0.00
d
0.14
0.00
0.00
e
0.00
1.00
0.60
f
0.00
0.00
0.30
g
0.00
0.00
0.10
Mdh2
a
0.71
0.00
0.90
b
0.29
0.00
0.10
c
0.00
1.00
0.00
Mpi
a
0.00
0.00
0.50
b
0.71
1.00
0.50
c
0.29
0.00
0.00
Pap
a
0.29
0.00
0.00
b
0.71
1.00
0.00
c
0.00
0.00
1.00
Pgd
a
0.86
0.00
0.50
b
0.00
0.00
0.40
c
0.14
0.00
0.10
d
0.00
1.00
0.00
River basin, and 0.39 ± 0.12 between Spanish samples from south of
the Guadalquivir River basin and those from northern Morocco.
Morphological Comparisons
Between sexes. — The 19 male and 8 female specimens in the sample
of D. galganoi examined by Capula et al. (1985) do not demonstrate
48
Annals of Carnegie Museum
VOL. 55
Table 3.— Number of replications of the seven possible patterns of allele distribution
present among the three populations o/Discoglossus. Patterns replicated eight times or
more do not occur as a result of chance association (b < 0.05, see Materials and Methods)
but only alleles shared between two or more populations are informative.
D. galganoi D. jeanneae D. pictus Number
X
X
X
15
X
14
X
10
X
24
X
X
2
X
X
8
X
X
14
sexual dimorphism » There were no significant differences in allometry
identified by the covariance analysis.
While no significant difference was found in the distribution of SUL
between 14 male and 16 female Discoglossus from south of the Gua-
dalquivir River basin in Spain, the allometric relationship between
SUL and foot length is significantly different between the sexes of this
population (F ^ 6.61, R < 0.05). The foot length of a female with an
SUL of 45 mm (an intermediate size in the sample of adults of either
sex) is approximately 95% the length of the foot of an equivalent-sized
male.
Five male and six female Discoglossus from northern Morocco dem-
onstrated sexual dimorphism in the allometric relationship between
SUL and snout length. While no significant differences were identified
between male and female SUL in these samples, the relationship be-
tween SUL and snout length is significantly different between sexes
(F = 16.39, P ^ 0.05). At an SUL of 45 mm, the snout length of a
female specimen is approximately 86% of the snout length of an equiv-
alent-sized male.
Between populations.— M.^\q specimens of D. galganoi differ from
male Discoglossus from northern Morocco in the allometric growth
relationship between SUL and snout (F = 68.01, P c 0.05), head (F =
63,57, P ^ 0.05), tibia (F = 10.66, P < 0.05), femur (F = 16.90, P <c
0.05) and hand lengths (F = 55.07, P ^ 0.05) and in the growth re-
lationships of SUL and head width (F = 5,84, P < 0.05) and SUL and
intemarial distance (F = 8.05, P < 0.05). Female specimens repre-
senting these populations differ in the allometric relationship between
SUL and head (F = 39.56, P <c 0.05) and eye (F = 28,73, P c 0,05)
lengths,
Male specimens of D. galganoi differ from males from south of the
Guadalquivir River basin in Spain in the allometric growth relationship
1986
Busack— Spanish and Moroccan Discoglossus
49
Fig. 2.— Spotted (upper) and striped (lower) color phases of Discoglossus jeanneae.
50
Annals of Carnegie Museum
VOL. 55
of SUL and snout (F 224.33, P c 0.05), head (F = 54.76, P c
0.05), tibia (F = 1 1.29, P c 0.05), femur (F - 7.08, P < 0.05), and
hand lengths (F ^ 45.82, P <c; 0.05), and in the allometric relationship
between SUL and intemarial distance (F ^ 5.63, P < 0.05). Female
specimens representing these populations differ in the allometry be-
tween SUL and head length (F = 72.34, P ^ 0.05), SUL and hand
length (F = 77,57, P c 0.05), and between SUL and intemarial dis-
tance (F= 12.62, P <c 0.05).
Male specimens from south of the Guadalquivir River basin in Spain
and those from northern Morocco differ only in the allometric growth
relationship between SUL and snout length (F = 16.47, P ^ 0.05).
The allometric growth relationship between SUL and all examined
morphological characters is not different in female specimens drawn
from these populations.
Systematic Considerations
Northern Moroccan populations. — Lanza et al. (1984) demonstrated
that specimens of Discoglossus from near Barika, Algeria, are geneti-
cally very similar to those from Sicily, the type locality of D. pictus
(Nei’s average genetic identity = 0.93, Nei’s D = 0.07). Although Ca-
pula et al. (1985:table 7) do not provide a table of allele frequencies
with which I might directly compare results, they do list 12 loci that
distinguished Algerian and Tunisian D. pictus from Iberian D. galga-
noi. Three of these 12 loci (Ada, Aatl, and Acp2) were found to be
monomorphic among populations I compared, and seven were not
considered in my study. Only two of these 12 loci (Icd2 and Ldhl)
distinguished between Moroccan and Iberian populations in my study.
While it is possible that not all Moroccan, Algerian, Tunisian, and
Sicilian populations are conspecific, at this time it is zoogeographically
and systematically conservative to consider populations of Discoglossus
inhabiting North Africa and Sicily D. pictus Otth, 1837.
Iberian populations residing north of the Guadalquivir Basin. —Tht
albumin immunological distance obtained when specimens from Vi-
lla viciosa and Arenas de San Pedro, Spain (Fig. lA, B, respectively),
were compared to those representing D. pictus from Tleta Tarhremt,
Morocco (Fig. 1 , closed triangle), was 1 7 units (Maxson and Szymura,
1984:249). The unbiased genetic distance (Nei’s D) between one in-
dividual I collected from San Martin del Pimpollar, Spain (Fig. 1C),
and D. pictus from northern Morocco was 0.74 ± 0.18. Capula et al.
(1985) reported a genetic distance (Nei’s D) of 0.58 between their
pooled samples representing Portugal and central Spain and those from
Algeria and Tunisia.
It is apparent from genetic (Capula et al., 1985:table 9) and mor-
1986
Busack— Spanish and Moroccan Discoglossus
51
Fig. 3.— Snout length regressed on snout-urostyle length in male Discoglossus galganoi
(squares), D. pictus (triangles), and D. jeanneae (circles). See Table 4 for regression
coefficients.
phological comparisons (this study, Results) that Iberian populations
of Discoglossus residing north of the Guadalquivir River basin are
dearly differentiated from those inhabiting northern Morocco and are
deserving of the species status recently ascribed to them by Capula et
al. (1985). Whether or not, however, the specimens Maxson and Szy-
mura (1984) and I examined biochemically (Fig. 1A-~C) actually rep-
resent D. galganoi is unclear. Until data with which to further assess
the taxonomic status of Iberian populations residing to the north of
the Guadalquivir River basin become available, these populations are
best referred to D. galganoi Capula et aL, 1985.
Iberian populations residing south of the Guadalquivir Basin.— Dis-
coglossus from south of the Guadalquivir River basin in Spain are
morphologically and genetically different from both Moroccan D. pictus
and. D. galganoi (Tables 2, 3, and Results). I consider the extent of
these differences to be representative of species level differentiation and
designate the new species herewith.
Table A.— Estimates of slope (b) and intercept (a) obtained when measurements derived from various morphological features (y) were
regressed againts smui-urostyie length (x) in Discoglossus galganoi, D. jeanneae, and D. pictus. Linear regression results are reported
52
Annals of Carnegie Museum
VOL. 55
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F„ P 17.46, cO.05 20.43, <0.05 158.89, c0.05 36.11, c0.05 26.70, <0.05
Table A. — Continued.
1986
Busack— Spanish and Moroccan Discoglossus
53
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54
Annals of Carnegie Museum
VOL. 55
Discoglossus jeanneae, new species
(Fig. 2)
Holotype.— Carnegie Museum of Natural History (CM) 546575 an
adult female, along highway C-440, 1 5 km ESE Alcala de los Gazules,
CMiz Province, Spain, 6 April 1971, Stephen D. Busack.
Paratypes (21, all from Cadiz Province, Spain).— CM 52126, female, along highway
C-440, 12.9 km ESE Alcala de los Gazules, 18 October 1969; CM 52128--52129, two
females, along highway C-440, 13.8 km WNW Los Barrios, 18 October 1969; CM 52475-
52476, two males, along highway C-440, 6.6 km NW Algeciras, 10 January 1970; CM
53087, male, along highway C-440, between 7.6 and 7.7 km NW Los Barrios, 12 March
1970; CM 53119, male, along highway CA-221, 21.2 km ENE Facinas, 10 April 1970;
CM 53324, male, along highway C-440, 9.01 km NW Los Barrios, 18 June 1970; CM
53884a--d, 4 females, along highway C-440, between 4.0 km WNW Casas del Castano
and 5.0 km WNW Los Barrios, 8 October 1970; CM 54244, female, along highway
C-440, between 1 9 km ESE Alcala de los Gazules and 1 km WNW Casas del Castano,
18 November 1970; CM 54581, female, and CM 54582, male, along highway C-440,
between 3.7 and 1 1.4 km WNW Los Barrios, 19 March 1971; CM 54608-54610, three
females, along highway CA-P-2 1 1 2, between 12.6 and 1 4.2 km NNE Benalup de Sidonia,
2 April 1971; CM 54704, male, along highway C-440, 0.3 km WNW Casas del Castano,
16 May 1971; CM 55742-55743, two males, and CM 55744, female, along highway
C-440, between 18.4 and 22.7 km WNW Los Barrios, 15 January 1972.
Diagnosis.— Discoglossus jeanneae is similar in coloration and pat-
tern to D. galganoi and D. pictus, but is distinguished from them by
biochemical and morphological characters. Discoglossus jeanneae and
D. pictus share no alleles at six electrophoretic loci (Ab2, Hkl, Hk2,
Icd2, Ldhl, and Lgg), D. jeanneae and D. galganoi share no alleles at
three loci (Glud, Hk2, and Ldh2).
Male D. jeanneae have a shorter snout than males of either D. pictus
or D. galganoi of similar SUL (Fig. 3, Table 4). Male D. jeanneae also
have a greater head length, a shorter intemarial distance, and shorter
tibia, femur, and hand lengths than male D. galganoi of comparable
SUL (Table 4). Female D. jeanneae have a shorter head and hand
lengths, and shorter intemarial distances, than D. galganoi of similar
SUL (Table 4).
Description of holotype. — An adult female with the following mea-
surements (mm): SUL 47.9, snout length 5.1, head length 10.8, head
width 15.1, horizontal diameter of eye 4.1, interorbital distance 7.5,
intemarial distance 3.0, tibia length 23.4, femur length 21.7, hand
length 5.9, foot length 15.1. Choanae oblong and relatively large; pre-
vomerine dentigerous processes in two nearly straight series, each with
1 1 teeth, located just behind the choanae and separated by a distance
less than half that of the diameter of a choana. Tongue roundish, thick,
scarcely free behind. Nostrils dorsal, much closer to the tip of the snout
than to the eye, horizontal diameter of the eye slightly greater than the
distance from the naris to the eye. Tympanum indistinct. Fingers rel-
1986
Busack— Spanish and Moroccan Discoglossus
55
Fig. 4.-- A Lower Miocene reconstruction of the Iberian peninsula and North Africa;
heavily blackened areas indicate marine incursions (after lilies, 1975),
atively short, unwebbed; III, IV, II, I in order of decreasing length.
Three prominent palmar tubercles, the largest at the base of finger IV,
the next largest at the base of finger I, and the smallest in the center.
Toes slender, very slightly webbed, IV, III, V, II, I in order of decreasing
length. No subarticular tubercles, no tarsal fold; small ellipsoidal meta-
tarsal tubercle. Heels overlap slightly when femora are held at right
angles to the body axis. Skin of dorsum with several irregularly posi-
tioned diminutive, pustules, skin of venter smooth.
Coloration ofholotype (in alcohol).— The ground color of the top of
the head is citrine drab from mid-eye to snout. Beginning at mid-eye,
and continuing to just below the area of front limb insertion, a dark
olive patch reminiscent of a large italicized “X” appears on a ground
color of a lighter shade of dark olive. A prominent (6.5 mm in length)
and elongated teardrop-shaped patch of chaetura drab angles ventro-
posteriorly from the posterior comer of the eye. This patch, and the
ground color of the “X,” are edged in citrine drab. Small faint patches
shaped as triangles oriented with the base down appear along the upper
lip; a light stripe extends from the front comer of the eye through the
nostril.
Colored as the ‘"X,” a lightly-homed, heart-shaped, shield extends
over the central dorsum. The upper half of the shield is edged in citrine
drab on a ground color of lighter dark olive; intense asymmetrical
patches of dark olive line the lower “V” and the upper and outer curves
of this shield. A broken dorsolateral line of citrine drab is present.
56
Annals of Carnegie Museum
VOL. 55
In lateral view, an ovoid dark olive blotch appears below the dorso-
lateral line and just above the area of insertion of the front limb. The
interaxillary region is light deep olive blending into dark olive buff
towards the venter; the dark olive buff blends into buffy brown in the
region of the groin. The venter is deep olive buff and unpattemed.
The upper surfaces of the hind legs are clove brown with irregular
transverse blotches of fuscous black; upper surfaces of the front legs
are citrine drab with three deep olive transverse blotches.
Pattern polymorphism {in alcohol). —Spoiled individuals are most
common, striped specimens are less common (Fig. 2), and occasionally
an unpattemed D. jeanneae is found. Unstriped individuals exhibit
either a complete or broken anteriodorsal ‘"X.” All have a teardrop
eye patch, triangular blotches on the upper labium, and, in varying
degrees of distinction, a stripe extending from the front comer of the
eye through the nostril. A broken dorsolateral line of citrine drab also
appears, at times faint, at times pronounced, in all unstriped individ-
uals.
Unstriped individuals have a ground color which varies from citrine
drab to light dark olive to fuscous. Coloration on the posterior portion
of the dorsum varies from a pattern in which prominent spots are
connected to suggest a shield-like shape to a simple pattern of widely
spaced and barely discemable spots. Ventral coloration is generally
unpattemed deep olive buff; some specimens, however, demonstrate
olive or clove brown pigmentation of varying intensity on the lower
jaw and upper pectoral region.
In the striped paratype (CM 53119), the “X” and shield symmetry
are broken by a medial stripe of citrine drab. The resultant halves of
the “X” and shield merge on either side of the medial stripe to form
two solid regions of dark olive, one on the right, one on the left, and
each between the medial stripe and the dorsolateral region. The small
ovoid blotch found below the dorsolateral ridge and above the area of
limb insertion in spotted individuals is continuous with the teardrop
eye patch and forms a single blotch in this specimen. The eye-nostril
stripes and upper labial triangles are pronounced and, along the dor-
solateral region, there is a prominent citrine drab stripe extending from
the eye to the groin. The outer edge of this stripe exhibits the darker
coloration reminiscent of the dark olive upper and outer curves of the
heart-shaped shield described for the holotype.
Etymology. — Jeanne A. Visnaw accompanied me during four months of field work in
Spain and Morocco during 1982; in spite of what she learned during her first trip abroad,
Jeanne again accompanied me in 1983. While the husbands and wives of graduate
students often contribute substantially to the success of their spouses, few routinely
sacrifice as much or give as unselfishly as my wife has. I dedicate this new Spanish frog
to her.
1986
Bus ACK— Spanish and Moroccan Discoglossus
57
Discussion
Physiogeographic changes have been extensive in this area of the
western Mediterranean region in the last few million years. The Betic
Cordillera (an alpine ridge located on the Spanish Meseta between
CMiz in the west and Alicante in the east) was subjected to folding
and extensively restructured during Miocene-Pliocene. Lower Miocene
Atlantic waters flowed to the Mediterranean through what is now the
Guadalquivir River basin (Figs. 4-5; Berggren and Van Couvering,
1974; lilies, 1975; Le Pichon et al., 1972), Neogene sedimentation filled
the younger, western portion (Cordoba-Sevilla) of the Guadalquivir
Basin and the northern portion began to emerge from considerable
depth at the end of the Miocene (Tjalsma, 1971:1 20-125), and Pliocene
events allowed Atlantic waters to form the Strait of Gibraltar (Femix
et al, 1967; Hsu, 1983; and Mantura, 1977).
Geologic events such as these have a direct effect on sexually repro-
ducing organisms. The rate at which genetic divergence accumulates
between populations is believed to be a function of the time those
populations have been separated (Zuckerkandl and Pauling, 1965). If
this is true, the more genetically differentiated two populations of ter-
restrial anurans are, the longer they have been separated. Of the three
patterns of allele distribution that are phylogenetically informative
among these three populations of Discoglossus, D. galganoi and Mo-
roccan D. pictus share only two, D. galganoi and D. jeanneae share 8,
and D. jeanneae and Moroccan D. pictus share 14 (Table 3). If unbiased
genetic distance units {D) are considered, Moroccan D. pictus are sep-
arated from D. galganoi by 0.74 ± 0.18 units and D. jeanneae is
separated from both D. galganoi and Moroccan D. pictus by 0.39 ±
0.12 units (Fig. 5). Discoglossus galganoi and Moroccan D. pictus are
probably not sister species.
Males of D. galganoi are more morphologically differentiated from
males of D. jeanneae and males of D. pictus than are female D. galganoi
from female D. jeanneae or D. pictus. The allometric growth relation-
ship between SUL and 10 morphological characters were compared
among males and females of these three species and male D. galganoi
differ from Moroccan D. pictus in seven such relationships, females
differ in only one. Male D. galganoi differ from D. jeanneae in six
allometric relationships, females differ in three. Male D. jeanneae ap-
pear to be little changed from Moroccan D. pictus, however, as only
one allometric relationship is clearly different; female D. jeanneae, on
the other hand, demonstrate no differences among any of the 10 al-
lometric growth relationships when compared with female D. pictus.
The fossil record of Discoglossus is limited. Middle Miocene remains
from Beni-Mellal, Morocco, referred to the genus Discoglossus by Ver-
gnaud-Grazzini (1966), have been reassigned to the extinct discoglossid
58
Annals of Carnegie Museum
VOL, 55
Fig. 5.— A mid-Miocene reconstruction of the Gibraltar area (after Femix et al., 1967),
Shaded areas represent emergent land; geographic and unbiased genetic distances between
sampled populations of Discoglossus are indicated.
genus Latonia by Sanchiz and Alcover {in litt.); as a result, fossil rep-
resentatives of Discoglossus are unknown in North Africa. The Lower
Miocene Discoglossus troschelii from Rott, Germany, was considered
conspecific with D. pictus (=? galganoi) from Spain (Boulenger, 1891).
Neogene specimens from Escobosa de Calatahazor (Soria), Venta del
Moro (Valencia), and Alcoy (Alicante), Pliocene samples from El Ar-
1986
Busack— Spanish and Moroccan Discoglossus
59
quillo II (Teruel), and mid-Pleistocene specimens from Arganda (Ma-
drid), attest to the age, persistence without morphological change, and
widespread nature of D. pictus (==? galganoi) throughout prehistoric
Iberia (Sanchiz, 1977a, 1911 b).
The electrophoretic and morphological data coincide well with one
evolutionary scenario that may be inferred from the geographic history
of the region. The ancestral stock of D. galganoi, which once populated
the Spanish Meseta, and that of D. pictus-D. jeanneae, which once
populated an area now known as southern Spain and northern Morocco,
suffered temporal and climatic separation dating from the Lower Mio-
cene. The Pliocene formation of the Strait of Gibraltar then divided
ancestral D. pictus- D. jeanneae populations into two populations which
evolved to become D. jeanneae and D. pictus.
Additional research is clearly necessary before we can fully under-
stand the evolutionary history, distributional limits, and taxonomy of
Iberian and North African Discoglossus. Until additional data becomes
available, however, conservative limits for the distribution of D. jean-
neae may be drawn. These would include the northern edge of the
Guadalquivir River basin (Fig. 1 , dotted line), regions inundated during
Miocene flooding that currently lay to the east of the headwaters of the
Guadalquivir River (Fig. 1 , dotted line with question marks), and the
shores of the Atlantic Ocean and the Strait of Gibraltar.
Acknowledgments
I wish to extend my gratitude to Benedetto Lanza, who provided me with a manuscript
copy of the description of D. galganoi, and to D. B. Wake and M. M. Frelow of the
Museum of Vertebrate Zoology, University of California, Berkeley, who provided fa-
cilities, financial support, and assistance during the electrophoretic analysis. L. R. Maxson
graciously provided immunological data and a copy of her and Szymura’s unpublished
manuscript, C. J. McCoy and E. J. Censky generously provided sustenance and lodging
during a visit to Pittsburgh, D. B, Wake, L. R, Maxson, and T. Uzzell reviewed and
improved earlier versions of the manuscript, Lezlie Skeetz assisted in translating the
patterns and colors of D. jeanneae into words, G. M. Christman prepared Figure 5, and
Alfredo Salvador assisted with a portion of the field work in Spain. Travel funds were
provided by a National Science Foundation dissertation improvement grant (DEB 81-
20868) and a National Geographic Society research grant (2600-83). Specimens in Spain
were collected under authority of permits 888 (1982) and 22061 (1983) issued by the
Institute Nacional para la Conservacion de la Naturaleza, Madrid. Collecting in Morocco
was authorized by letter from the Embassy of Morocco to the United States, Mohamed
Benjelloun, economic counsellor.
Specimens Examined
D. (Electrophoretic Analysis): SDB 1556, 1691, 1905, 1906, 1930, 1949,
1954, and 1989 at the Universidad de Leon, Spain. (Morphological Analysis): Carnegie
Museum of Natural History (CM) 52125-52129, 52475-52477, 52537, 52626, 53087-
53088, 53119-53120, 53324, 53884(4 specimens), 54244, 54581-54582, 54608-54610,
54657, 54704, and 55742-55743.
D. galganoi.— {p\QcXTOp\ioreX\c Analysis): SDB 1691 at the Universidad de Leon.
60
Annals of Carnegie Museum
VOL. 55
D. (Electrophoretic Analysis): SDB 1773 (2 specimens) and SDB 1774 (3
specimens) at the Museum of Vertebrate Zoology, University of California, Berkeley
(MVZ); MVZ 186124-186125 and 186132-186134. (Morphological Analysis): MVZ
186124-186134.
Literature Cited
Ayala, F, J,, J. R. Powell, M. L. Tracey, C. A, MourAo, and S. Perez-Salas. 1972.
Enzyme variability in the Drosophila wiilistoni group. IV. Genic variation in natural
populations of Drosophila wiilistoni. Genetics, 70:113-139.
Berggren, W. a., and j. A. Van Couvering. 1974. Developments in palaeontology
and stratigraphy, 2. The Late Neogene. Elsevier Sci. Publ. Co., New York, 216 pp.
Boulenger, G. a. 1891. On the occurrence of Discoglossus in the Lower Miocene of
Germany. Ann. Mag. Nat. Hist., ser. 6, 8:83-85.
Busack, S. D. 1986. Biogeographic analysis of the herpetofauna separated by the
formation of the Strait of Gibraltar. National Geographic Research 2(1): 17-36.
Capula, M., G. Nascetti, B. Lanza, L. Bullini, and E. G. Crespo. 1985. Morpho-
logical and genetic differentiation between the Iberian and the other west Mediter-
ranean Discoglossus species (Amphibia Salientia Discoglossoidae). Monitore ZooL
Ital. (N.S.), 19:69-90.
Fernix, F., j. Magne, and D. Mongin. 1 967, The paleogeography of the eastern Betic
ranges of southern Spain during the Caenozoic. Pp. 239-246, in Aspects of Tethyan
biogeography (C. G. Adams and D. V. Ager, eds.), Systematics Assn., London,
336 pp.
Harris, H., and D. A. Hopkinson. 1976. Handbook of enzyme electrophoresis in
human genetics. North-Holland Publ. Co., Amsterdam, Holland.
Hsu, K. J. 1983. The Mediterranean was a desert. Princeton Univ. Press, Princeton,
New Jersey, 197 pp.
Illies, j. H. 1975. Intraplate tectonics in stable Europe as related to plate tectonics in
the Alpine system. Geolog. Rundschau, 64:677-699.
Knoepffler, L.~Ph. \96\a. Contribution a Fetude du genre Discoglossus (Amphibiens,
Anoures). Unpublished thesis, Univ. Paris, 96 pp.
— — . 1961Z?. Les batraciens et principalement le genre Discoglossus dans les lies
Mediterraneennes. Colloques int. Cent. natn. Rech. scient. 94:159-161,
. 1962. Contribution a Fetude du genre Discoglossus (Amphibiens, Anoures).
Vie Milieu, 13:1-94.
Lanza, B., G. Nascetti, M. Capula, and L. Bullini. 1984. Genetic relationships
among west Mediterranean Discoglossus with the description of a new species (Am-
phibia Salientia Discoglossidae). Monitore Zool. Ital. (N. S.), 18:133"“152.
Le Pichon, X., G. Pautot, and J. P. Weill, 1972. Opening of the Alboran Sea. Nature,
236:83-85.
Lewontin, R. C. 1966. On the measurement of relative variability. Syst. ZooL, 15:
141-142.
Mantura, a. j. 1977. The Mediterranean enigma. Oceans, 10:24-27.
Maxson, L. R., and j. M. Szymura. 1984. Relationships among discoglossid frogs:
an albumin perspective. Amphibia-Reptilia, 5:245-252.
Moriarty, D. j. 1977. On the use of variance of logarithms. Syst. ZooL, 26:92-93.
Nei, M. 1971. Interspecific gene differences and evolutionary time estimated from
electrophoretic data on protein identity. Amer. Nat,, 105:385-398.
. 1978. Estimation of average heterozygosity and genetic distance from a small
number of individuals. Genetics, 89:583-590.
Patton, J. L., and M. F. Smith. 1981. Molecular evolution in Thomomys: Phyletic
systematics, paraphyly, and rates of evolution. J, Mamm., 62:493-500.
1986
Busack— Spanish and Moroccan Discoglossus
61
Sanchiz, F. B. 1 911a. CatMogo de los anfibios fosiles de Espana (Noviembre de 1 977).
Acta Geol. Hisp., 12:103-107.
— 19776. Nuevos anfibios del Neogeno y Cuatemario de Europa. Origen, desa-
rrollo y relaciones de la batracofauna espanola. Unpublished Ph.D. thesis, Univ.
Complutense, Madrid, 863 pp.
Selander, R. K., M. H. Smith, S. Y. Yang, W. E. Johnson, and J. B. Gentry. 1971.
Biochemical polymorphism and systematics in the genus Peromyscus. I. Variation
in the old-field mouse {Peromyscus poUonotus). Studies in Genetics VI, University
of Texas PubL, 7103:49-90.
SoKAL, R. R., AND F. J. Rohlf. 1981. Biometry. W. H. Freeman and Co., New York,
859 pp.
Straney, D. O. 1 980. Relationships of phyllostomatine bats: evaluation of phylogenetic
hypotheses. Unpublished Ph.D. dissert., Univ. California, Berkeley, 269 pp.
Thorpe, R. S. 1976. Biometric analysis of geographic variation and racial affinities.
Biol. Rev., 51:407-452.
Tjalsma, R. C. 1971. Stratigraphy and Foraminifera of the Neogene of the eastern
Guadalquivir Basin (southern Spain). Micropaleont. Bull., Utrecht, 4:1-161.
Vergnaud-Grazzini, C. 1966. Les amphibiens du Miocene de Beni-Mellal. Notes
Mem. Serv. Mines Carte Geol. Maroc, 27(198):43-75.
Zuckerkandl, E., and L. Pauling. 1965. Evolutionary divergence and convergence
in proteins. Pp. 97-166, in Evolving genes and proteins (V. Bryson and H. J. Vogel,
eds.), Academic Press, London, 629 pp.
Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.
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ANNALS
of CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 23 MAY 1986 ARTICLE 4
YANOMAMA MATERIAL CULTURE IN THE CARNEGIE
MUSEUM OF NATURAL HISTORY. PART IL
WEARING APPAREL AND FESTIVAL
ARTIFACTS
Lorraine Couture-Brunette
Collections Manager, Section of Anthropology
Abstract
The Carnegie Museum of Natural History has one of the largest, most inclusive, and
best documented collections of Yanomama material culture in the world. The collection,
consisting of two accessions comprising 572 specimens, spans a 5 year period from 1979
to 1984. This corpus of material documents not only traditional Yanomama material
culture, but also shows the changes it has undergone due to the introduction of Western
goods and materials. Part I dealt with material culture categories related to Food Pro-
curement and Household Articles (Couture-Brunette, 1985). Part II deals with Wearing
Apparel and Festival Artifacts; and Part III will deal with Foreign Influence and Mis-
cellaneous Constructions.
Introduction
The Yanomama Indians inhabit an area of approximately 30,000
(Smole, 1976:3) to 100,000 (Migliazza, 1972:20) square miles in north-
ern Brazil and southern Venezuela. They are one of the largest Indian
populations in the Amazonian rain forest. In spite of their large pop-
ulation, they have been able to remain isolated and unacculturated
until the present due to their settlement locations olf major waterways
in the Guyana Shield area. There is no
of Yanomama. While Chagnon (1974:
Submitted 2 April 1985,
63
64
Annals of Carnegie Museum
VOL. 55
of 10,000, Saffirio (1985:36) feels that 16,400 in 320 villages is closer
to the true number.
All Yanomama speak four mutually intelligible languages and several
dialects. The group has been called by several names; terms such as
“Waica,” “Xiriana,” “Shamatari,” and “Yanoama,” have all been used
to designate this family of languages. “Yanomamo,” “Yanam,” “San-
uma,” and “Yanomam” actually denote only one dialect or language
within the larger group. Thus “Yanomama” is used to refer to the
entire family of languages (Migliazza, 1972:33). The Catrimani River
Yanomama of Brazil, from whom the Carnegie Museum of Natural
History collections have come, belong to the linguistic group “Yano-
mam.” They inhabit areas of the Federal Territory of Roraima and
the State of Amazonas.
Since the construction in the mid 1970s of Brazilian Highway BR
210, the Perimetral Norte, deep into Yanomama territory in Brazil,
acculturation has been proceeding at a rapid rate. The introduction of
Western goods, beliefs, and diseases has brought about profound and
permanent changes in the Yanomama lifestyle (Saffirio, 1980; Saffirio
et al., 1983). Like many South American groups before them the Yan-
omama are substituting Western material culture in replacement of
their traditional technology. This loss of native crafts is documented
in the Carnegie Museum of Natural History collections.
Giovanni Saffirio, a member of the Consolata Society for Foreign
Missions, arrived at a mission post on the Catrimani River in 1968.
He was the collector of Yanomama material culture for Carnegie Mu-
seum of Natural History. He has recently been awarded his Ph.D. in
the Department of Anthropology at the University of Pittsburgh, and
has since returned to the Catrimani Mission.
Collection I, accession number 32703, was made during 1979-80,
while collection II, accession number 32735, took place in 1984. Every
effort was made to secure as complete a representation of Yanomama
material culture as possible, with a wide range in variation among
specimens. As a result, the Carnegie Museum of Natural History has
one of the largest, best documented, and most inclusive collections in
the world.
The Catrimani River Yanomama Indians of South America, and the
Carnegie Museum of Natural History collections of Yanomama ma-
terial culture, have been presented in more detail in the Part I article
(Couture-Brunette, 1985). Part II covers the Wearing Apparel and Fes-
tival Artifacts segment of the collection.
The classification of an artifact as either an item of wearing apparel
or a festival artifact is based on the function of the artifact, regardless
of its method of construction. Some specimens which served the same
1986
Couture-Brunette— Yanomama Material Culture, II
65
Fig. L— Women’s aprons, an item of everyday apparel. Accession 32735.
purpose were constructed differently based on their use as an item of
everyday wear or as a festival artifact. In these cases the entire group
of specimens was placed in the category where it was most often used.
In this manner, separation of like artifacts was avoided. Two examples,
one from each of the Festival Artifacts and Wearing Apparel categories,
will illustrate this placement. Women’s aprons are worn every day, but
certain aprons in the Carnegie Museum of Natural History collection
are made specifically for festivals. All aprons were placed into the
Wearing Apparel category in order to discuss the “apron” group as a
whole. Highly decorated aprons which were intended for festival usage
are separately indicated.
It proved impossible to explain fully all manufacturing terminology and methods of
the many different materials (basketry, cordage, knots) used in construction of the spec-
66
Annals of Carnegie Museum
VOL. 55
imens. Standard references were used-Emery (1980) when discussing the cordage, Ado-
vasio (1977) for the basketry, and Shaw (1972) for knot descriptions.
Numbers for the types of cordage and letters for the knots are used throughout the
analysis (Couture-Brunette, 1985:tables 1 and 2). All knots are illustrated (Couture-
Brunette, 1985:fig. 4). “S” and “Z” twist in cordage is also illustrated (Couture-Brunette,
1985:%. 5).
Some of the cordage consists of a number of spun and plied yams which were combined
by twisting. Rather than list these varying numbers of yams as separate constmctions,
they were combined in one cordage type (see Couture-Bmnette, 1985:table 1), with the
number of plies referred to in parentheses after the cordage type number. For example,
cordage type #12(4) consists of four pieces of 2 ply s spin Z twist yams, all combined in
an S twist. Similarly, cordage type #13(4) is composed of four pieces of 3 ply s spin Z
twist yams, combined in an S twist. Cordage types #5 and #6, as well as types #11
through #15, all consist of varying numbers of plies, and will be used with parenthetical
numbers indicative of these.
All Yanomama words in the text are set in italics. Spelling follows SafRrio (1980,
1985). Due to the great number of Yanomama languages and dialects, these words are
specific to the Catrimani River villages only.
Wearing Apparel
Men and women do not wear an abundance of clothing. The only
item which men wear every day is a string around the waist, to which
they tie their penis foreskin. Occasionally men will also wear a “belt”
of native cotton. Women wear cotton “aprons” which consist of a thick
belt in the back with a short fringe in the front.
Both men and women pierce their ears. Women also pierce the center
and comers of their lower lip, and the nasal septum. Everyday “jewelry”
consists of straws worn in the lip holes, and straws or bunches of grass
in the nose and ears. Feather earrings are worn by both men and women,
usually during festivals.
Traditionally, necklaces are made from seeds, shells, feathers, and
other animal parts stmng on yamaasik fiber cordage or cotton string.
Only men wear necklaces made from animal parts, although women
utilize feathers and shells in earrings, necklaces, and aprons.
Many men and women have begun to wear Western shorts and T
shirts since the arrival of Westerners on a frequent basis on Brazilian
Highway BR 210. Young men no longer pierce their ears, and women
near the highway do not pierce their lips or nose (Saffirio, 1980; Saffirio
et aL, 1983). Increasing numbers of necklaces and earrings are made
from trade beads and scrap metal acquired from Westerners.
Women's Clothing
“Aprons” (Figs. 1-2), which are worn every day, are a sign of a
woman’s modesty. They are made by women from beginning to end;
the cotton is spun and the cordage is plied and twisted by them. Women
also wear armbands made from flowers, grasses, or cotton.
Aprons (pesimak). — For analytical purposes, the elements of the apron
1986
Couture-Brunette— Yanom AM A Material Culture, II
67
Fig, 2.— Beaded woman’s apron, CMNH number 32703-12. This would be worn during
a festival.
were divided into three separate constructions —belt, fringe, and cord.
The “belt” is the back portion of the apron. It is constructed in the
same manner as the cotton hammock (Couture-Brunette 1985: figs. 28,
29). Stakes are placed in the ground the necessary distance apart, and
one continuous strand of cotton yam is wrapped back and forth. The
ends are then tied with a cotton cord, and the stakes are removed. Belts
contain approximately 150 warp strands. They average twice the length
of the front portion of the apron: the “fringe.” This is constmcted by
folding short pieces of cordage in half over a length of cotton cord. The
pieces are then twined tightly with cotton or yamaasik string just below
the cord. One end of the fringe cord is tied permanently to the belt.
The term “cord” denotes both the cord over which the fringe is folded
and twined, and the cord which ties the two ends of the belt together.
In most cases these two pieces of cordage are not the same, differing
usually in the number of plies contained within the twisted rope.
Decorative seeds, beads, shells, or feathers are occasionally tied to
each end of the fringe. The entire apron is sometimes dyed with nara
(a reddish dye) in shades of brown, yellow, red, or orange.
Table l.— Yanomama women’s aprons in the collections of Carnegie Museum of Natural History.
68
Annals of Carnegie Museum
VOL. 55
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-44P #13(6) #9 #13(2) A(7), C Z slant S slant 51.2 21.6
Table \. — Continued.
1986
Couture-Brunette— Yanom AM A Material Culture, II
69
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t Two halves of fringe made by two people (?); cordage and decoration are both dilferent.
70
Annals of Carnegie Museum
VOL. 55
The aprons are fairly standardized in construction, although some
cordage variation is apparent in Table 1. The belt cordage is the most
consistent element in all aprons. Of the 27 specimens in collection
32703, 24 of them (89%) used cordage type #9 for the belt, either alone
or in combination with another type. Cordage types #10, #1 1(3), and
#14(4) were each used on one belt, and type #7 (in combination with
type #9) was used on two. Fourteen of the 16 aprons from the 32735
collection (87.5%) used cordage type #9 on the belt. Two belts used a
combination of cordage types #9 and #7, and one belt used type #8.
Twelve of the collection I belts have occasional rows of twining to
hold the warp yams together. Of these belts, 75% of the twining is S
slant and 25% is Z slant (Couture-Bmnette, 1985:fig. 15). Ten of the
collection II belts have rows of twining, 90% S slant and 10% Z slant.
This mixture of S and Z slant twining is interesting in view of the fact
that 100% of the basketry twining is S slant. Stitch slant in twining is
a recognizable attribute, which is standardized over a cultural area
(Adovasio, 1977:30). It is unusual to find this variation in the aprons,
which utilize twining only incidentally, and not in the twined basketry.
There, the greater number of specimens and the greater amounts of
twining would make some variation more understandable.
The cordage used on the fringe is generally the same cordage present
on the belt, but is doubled back on itself and re-twisted. For example,
2 ply s spin Z twist cotton cordage (type #9), the most common belt
cordage, is doubled and S twisted to produce the 2 ply s spin z plied
S twisted cordage of type #13(2), the most common fringe cordage.
This cordage is present on 24 of the 27 collection I aprons and 14 of
the 16 collection II aprons.
The twining which secured the pieces of fringe to the cord is S slant
on 78% of the 32703 aprons and Z slant on 22%. Of the 32735 fringes,
87.5% of the aprons have S slant twining, and one each are Z slant and
stitched.
The most interesting fact about the twining on both belts and fringes
is the fact that S slant twining on one is often paired with Z slant
twining on the other. This can also be seen on Table 1, where eight of
the 32703 aprons, representing 67% of the belts which have twining,
pair S slant twining on the belt with S slant twining on the fringe. One
apron exhibits Z slant twining on both belt and fringe, while two of
the aprons, 17% of the belts which have twining, combine Z slant
twining on the belt with S slant twining on the fringe. The same is tme
of the 32735 aprons; of the 10 belts with twining, 70% of them have
S slant twining on both belt and fringe, 10% pair S slant on the belt
with Z slant on the fringe, and 1 0% pair Z slant on the belt with S slant
on the fringe. One of the aprons utilizes S slant twining on the belt and
stitching on the fringe. These facts suggest that belts and fringes are
1986
Couture-Brunette-— Yanom AM A Material Culture, II
71
made by different women, given the standardization of stitch slant in
twining which was discussed earlier. This deduction was later confirmed
by the collector (Saffirio, personal communication).
The cords, which tie the ends of the belt and serve as the ‘‘backbone”
to the fringe, are different from one another on 16 of the collection I
aprons (59%) and 1 3 of the collection II aprons (81%). These differences
are generally exhibited in the number of plies contained in the cord; if
this variance is regarded as a function of cord diameter they are as
standardized as the other apron elements. Cordage type #13(variable
number of plies) is used on 25 of the 27 collection I aprons in at least
one of the two possible places (belt or fringe) and on all of the collection
II aprons on either the belt or fringe. The number of s spin Z twist
plies used in cordage type #13 was widely variable, with four to eight
the most common amount. Of the 32703 aprons, cordage type #13
with six to eight plies was the most popular, whereas on the 32735
aprons four and five plies were more often used. One apron in the
32703 accession used a five strand sennit cord and another exhibited
a commercially braided cord. One of the 32735 aprons used cord made
from yamaasik fiber.
Knot type “A” was the most popular knot present on the aprons.
Nearly all of the cords were finished off with this knot on each string
end. Knot type “C,” the second most popular knot, was used to attach
the fringe to the belt on 23 of the 27 collection I aprons and 12 of the
16 collection II aprons. The other knots present on the aprons were
generally used to attach decorative elements to the fringe. Occasionally
the fringe had additional pieces of cordage attached to the main cord
with knot type “H,” the lark’s head.
Taking all these facts into consideration, the “standard” apron con-
sists of cordage type #9 on the belt, cordage type #13(2) on the fringe,
and cordage type #13(variable number of plies) making up the cord.
Twenty-two of the 27 aprons in collection I, 81%, are standard. The
percentage was even higher for the 32735 aprons, where 14 of the 16
aprons, 87.5%, are standard.
The two accessions differ more in their mode of decoration than in
their method of construction. These decorative differences between
collections I and II are a reflection of collection bias; the earlier 32703
accession contains more aprons which were worn every day, while
many festival aprons were collected for the later 32735 accession.
Only four specimens from collection I have decorative accessories
added to the apron. Two aprons (32703-44E and -440) have shells
strung on cotton or yamaasik string and tied to each fringe end, whereas
one apron (- 1 4C) has patches of curassow skin and feathers tied to each
fringe end. The fourth apron, - 1 2, is an entirely different construction
(Fig. 2). The belt is standard cordage type #9 and the cord is type
72
Annals of Carnegie Museum
VOL. 55
Fig. 3.— Two traditional women’s necklaces of tirimoku seeds, CMNH numbers 32703-7A
and B (left). Right, two aroariki, men’s necklaces worn for protection against evil spirits
and spells, CMNH numbers 32703-8A and B.
#12(5). The fringe, however, consists of a 27.4 by 13.2 cm beaded
rectangle in red, white, and light and dark blue. There are red (cordage
type #10) and black (cordage type #9) tassels at each comer, and a
small red fringe along the bottom of the apron (cordage type #10). The
beaded design is a geometric motif with opposing triangles within bands
of contrasting color. The beadwork on an apron such as this is very
intricate and time consuming, and the apron was very expensive to
purchase from the maker. It was brought from Themaim, a Maxiko-
piutheri woman from the Mucajai River, in December 1979.
Most of the collection II aprons are decorated with beads, tirimoku
seeds, shells, and feathers tied to each fringe end. Feathers are the most
popular decorative .elements; specimens 32735-291, -296, and -300 use
toucan, curassow, and lovely cotinga feathers to decorate each fringe
end, and the fringe on apron -293 is constmcted entirely from curassow
feathers stitched to the cord. Aprons -297, -298, and -299 used curas-
sow and toucan feathers in combination with beads, shells, and seeds.
Apron -294 utilizes strings of beads and seeds at each fringe end, while
-288 makes use of strings of seeds.
1986 Couture-Brunette— -Yanom AM A Material Culture, II 73
Fig. 4.— Three kotho, belts for boys and men. From top: 32703- 16B, -16C, and -16A.
The fringe on apron -292 appears to have been made by different
women. While the cordage, type #13(2), is the same throughout the
fringe, the diameter of both the spun and plied yams is quite different.
Likewise, the decorations at each end of the fringe are different; one
side has shells stmng on cordage types #9 and #1 3(5), whereas the other
side decoration is beads and seeds. This suggests that decorative ele-
ments on aprons may be specific to individual women, another de-
duction later confirmed by the collector (Saffirio, personal communi-
cation).
Armbands. -—Most of the decorative arm ornaments worn by women
are composed of highly perishable flowers or grasses; therefore, no
specimens of these are present in the Carnegie Museum of Natural
History collections. The only women’s armbands in the collection are
a pair of cotton armbands for a young girl. Although at first glance they
appear to be loom woven cloth, the fabric stmcture is actually plain
interlinking crossed right over left (Emery, 1980:61). This circular
method of construction produces a seamless, stretch fabric.
32735-21 A&B, wao kik, owned by Yaikom. Pink cotton girl’s armband, seamless,
cordage type #9, D— 10,1 cm.
Necklaces. Carnegie Museum of Natural History collection
74
Annals of Carnegie Museum
VOL. 55
contains three traditional seed necklaces (Fig. 3). These are rarely made
by women today because of the availability of trade beads.
32703-7A, bought from a Hewenahipiutheri woman, December 1979. Tirimoku seeds
on cordage type #12(3) tied with three knot “A”s. L— 40.6 cm.
32703-7B, same as 32703-7A except cordage type #3 tied in knots “A” and “G.” L—
55.9 cm.
32735-35, yanuak, woman’s necklace. Fourteen loops of small, round black seeds strung
on cordage type #3 tied in knot “F.” Two knot “A”s one string end. L— 43.8 cm.
Men’s clothing
The penis string is the only item of clothing considered essential for
everyday men’s wear (Saffirio, personal communication). Occasionally
men will also wear a “belt” of native cotton (Fig. 4). Initially this is
made by women in the same fashion as the belt portion of the women’s
apron. But after the ends have been tied and the stakes removed, the
entire length of the belt is very tightly wrapped with yanaaasik or cotton
string every 2 to 5 cm. The belt is often dyed red or brown with nara.
Belts (kothoj.
32703- 16A, man’s belt. Cordage type #9 belt; cordage #13(5,6) cord. Loose warps, no
wrapping. L— -56.6 cm,
32703-16B, man’s belt, from Mahuku of Hewenahipiutheri village, December 1979.
Cordage type #7 belt with cordage type #9 tie every 2 cm. Cordage type #3 cord
tied at each end with knots “E,” “G,” and “F.” Yams dyed red with nara; three
purple serpentine lines painted down belt length. L~73.2 cm.
32703- 16C, boy’s belt. Cordage type #9 belt; cordage type #13(4) cord. Belt wrapped
with green commercial string every 3 cm. Tied each end with knots “A,” “F,” and
“H.” L~38.6 cm.
32703-301, boy’s belt. Cordage type #7 belt; cordage type #13(6) cord. Tightly wrapped
with cordage type #1 every 2 cm; tied with knots “A”(4) and “E”(2). Dyed red with
nara, L— 36.2 cm.
Necklaces.— ThQSQ necklaces (Fig. 5) are worn only by men, because
they are made of animal parts from game the men have hunted and
killed.
32703-9, thihi nak, jaguar tooth necklace from Paxeko, a 30 year old good hunter in
Hewenahipiutheri village. Cordage type #3, knot “E”(?). L— 36.9 cm,
32703-10, opo nak, from Iropitheri village November 1979. Armadillo teeth tied to
cordage type #1 with knot “E.” L— 39.4 cm.
32735-43, opo sina. Cordage type #3 necklace, with pendant of two armadillo tails tied
to untwisted yamaasik fiber with knots “A” and “E.” L— 37.9 cm.
32735-84, necklace of cordage type #3 with pendant of two cock of the rock bird upper
beaks and crests stitched together with commercial pink thread. Tied together with
knots “G” and “F.” L— 36.5 cm.
Festival Artifacts
This category includes all items used at feasts and for ritual or spir-
itual purposes. Items of clothing (feather armbands, headbands, and
1986
Couture-Brunette— Yanom AM A Material Culture, II
75
Fig. 5. —Two necklaces made from animal parts, and thus worn only by men. CMNH
numbers 32703-9 (left) and 32703-10 (right).
earrings) worn only during festivals are included, although highly dec-
orated women’s aprons, usually worn during festivals, are grouped into
the “Wearing Apparel” category with the rest of the aprons as discussed
previously.
Feasts, which can last up to a week, are primarily political events
among the Yanomama. Although each village is self-sufficient in terms
of daily life and food procurement, alliances with other villages are
promoted for marriage alliances and cooperative warfare. The Catri-
mani River Yanomama follow a prescriptive bilateral cross-cousin
marriage rule and use an ‘Troquois-Dra vidian” kinship system (Chag-
non, 1977:56; Saffirio, 1985:1 18). However, one village often does not
provide enough cross-cousins to satisfy this rule. A man must fre-
quently seek a wife outside of his own village as a result (Smole, 1976:
76, 94). When he finds an eligible female, he is obliged to reciprocate
to her father or brothers with his own sister. Alliances between villages
are strengthened by the double tie (Chagnon, 1977:55).
Warfare is another situation calling for village alliances. Several vil-
lages may band together to fight a common enemy, or one village may
flee to an ally when warfare drives them out of their own territory.
76
Annals of Carnegie Museum
VOL. 55
They are occasionally forced to live in the allied village for a lengthy
period of time —up to several years —until their new gardens begin to
produce (Chagnon, 1977:97ff).
Both the Brazilian Indian Agency (FUNAI) and missionary groups
have discouraged warfare among the Yanomama in recent years. As a
result, ritual chest pounding duels during festivals are currently re-
placing fatal raiding parties (Saffirio, personal communication). The
duels allow the Yanomama to settle grievances among themselves or
with allied villages. These duels are usually performed during the fes-
tivals.
Body Decoration
Festival wear consists of feather or animal skin armbands and head-
bands for men and feather earrings for men and women. In addition,
both sexes paint themselves elaborately and cover their hair with bird
down.
Yanomama men and women paint their bodies frequently
during the dry season and almost daily during the rainy season with
black {nara uxirim) and red {nara wakirim) paint. Body painting is
especially colorful and elaborate during feasts, so the paints and storage
equipment were placed into the Festival category.
Red body paint is made from seeds {nara moko) of the Bixa orellana
tree. These have a greasy red outer covering, which is rubbed off and
molded into a ball (Fig. 14) or made into a thick liquid to be stored
in gourds.
32735-93, black paint gourd, owned by Carrera. End sealed with wax. String through
hole below gourd neck: cordage type #3 tied in two knot “A”s. L— 10.7 cm W-8.1
cm.
32735-342, black paint gourd? sealed with leaf plug. Neck wrapped with scotch tape, no
string. L— 11.2 cm W—7.7 cm.
32703-29, lump of red paint. L— 5.9 cm.
32735-310, same as 32703-29. L-=7.0 cm.
32735-311, same as 32703-29. L^7.8 cm.
32703-23, red paint gourd sealed with wax. Cordage type #3 through hole in gourd neck,
knot “A” on visible end. L— 18.8 cm W— 1 1.6 cm.
Apparel. —Festival armbands for women are made of leaves and
flowers; none of these are present in the Carnegie Museum of Natural
History collection. Festival armbands for men are made from bird
feathers and body parts.
White bird down feathers (horomaep) are also used by both Yano-
mama men and women. They coat the hair with resin, and then cover
their heads with the white down. The down on women occupies only
a fringe around the head, but men apply it to completely cover the
hair. The down is stored in gourds {horokoto), and retrieved by poking
a stick into the cut-oflf top.
1986
Couture-Brunette— Yanom AM A Material Culture, II
77
Fig. 6.— Feather earrings {kurakaas sina) worn by men and women during festivals. All
are accession 32735.
32703-20A, horokoto with horomaep, from Mahuku of Hewenahipiutheri village De-
cember 1979. Top cut off; hole for tie string through neck. L—15.4 cm.
32703-20B, same as 32703-20A. L— 16.5 cm.
32703-20C, same as 32703-20A. L-19.3 cm.
32703-2 1C, corked with wad of leaves. Cordage type #1 1(3) tied through hole with knot
‘T”(?). L-12.4 cm.
32703-52A, collected 1979-”80. Cordage type #3 tied in neck hole with knot “A.” L—
15,1 cm.
32703-52B, same as 32703-52A. L-15.4 cm.
32735-106, top cut off; no hole in neck for cordage. L— 12,9 cm.
32735-107, corked with leaf wad; cordage type #4 tied through hole with knots “E” and
“A.” L-12.1 cm.
32735-108B, two holes drilled through neck; cordage type #3 threaded through both
holes and tied inside gourd with unidentifiable knot. L— 16.6 cm.
32735-108B, no stopper; cordage type #9(?) through hole. L— 25.1 cm.
As has been discussed previously, men wear items made from animal
body parts such as wings, skins, teeth, and tails. Women, however, are
permitted to wear feather earrings. These earrings, also worn by men,
consist primarily of single feathers inserted into pieces of straw (Fig.
6). Sometimes the feathered end of the straw is smeared with resin.
78
Annals of Carnegie Museum
VOL. 55
Table 2. — Yanomama feather earrings in the collections of Carnegie Museum of Natural
History.
Type
Accession number
Feathers (number)
Cordage
Length
(cm)
seisi sina
32735-46 A&B
lovely cotinga
#2
9.5
-47 A&B
lovely cotinga
yamaasik
11.2
-53 A&B
lovely cotinga
yamaasik
11.0
-54 A&B
lovely cotinga
yamaasik
17.0
kurakaas sina
-55 A&B
parrot (single)
none
16.5
-56 A&B
parrot (single)
none
17.0
-57 A&B
parrot (single)
none
16.6
-58 A&B
parrot (single)
resin
19.4
-59 A&B
parrot (single)
resin
19.7
-60 A&B
parrot (single)
yamaasik
22.5
-61 A&B
parrot (single)
yamaasik
21.4
-62 A&B
parrot, curassow
yamaasik
15.6
-63 A&B
parrot, curassow
yamaasik
17.7
-64 A&B
parrot, curassow
yamaasik
17.3
-65 A&B
curassow (two)
none
13.4
-66 A&B
curassow (two)
none
9.9
-67
curassow (two)
none
12.8
-68 A&B
curassow (two)
none
11.3
-69 A&B
curassow (two)
none
11.3
-70 A&B
curassow (two)
none
11.5
-71 A&B
toucan, parrot,
curassow
#2 (?)
18.7
-72 A&B
parrot, toucan
none
14.3
-73 A&B
parrot, curassow
#1
15.0
-74 A&B
guan, curassow
#1
12.2
-75 A&B
parrot (single)
none
11.9
-76 A&B
parrot (single)
none
11.6
-77 A&B
guan (single)
none
7.2
-78 A&B
guan (single)
none
10.0
-79 A&B
guan (single)
none
11.7
-80 A&B
parrot (single)
none
11.3
-81 A&B
guan (single)
none
11.6
-82 A&B
curassow (two)
none
12.4
-83 A&B
guan, curassow
yamaasik
13.0
either to protect underlying fiber wrapping or to hold the unwrapped
feather in place.
Table 2 presents the data for the two types of feather earrings. The
first, seisi sina, consists of lovely cotinga bird skin and feather patches
wrapped to pieces of wood, which is used in place of straw because of
the weight of the earring. One pair of earrings is wrapped with the rare
Z twist yamaasik fiber cordage, whereas the others are secured with
unplied yamaasik.
The second type of earring, kurakaas sina, utilizes a variety of feath-
1986
Couture-Brunette— Yanom AM A Material Culture, II
79
ers. Parrot and curassow are the most popular types, whereas toucan
and guan are occasionally present. The commonest earring is the single
or paired feather inserted into straw, seen in 2 1 of the 29 pairs. Eight
pairs have a single earring in the center surrounded by a “bouquet” of
curassow or toucan feathers.
Nine pairs use fiber wrapping to secure the feathers, and two pairs
have resin coating the inserted end. The unplied yamaasik is again the
most popular wrapping, but two pairs use S twist yamaasik, and one
pair utilizes the rare Z twist yamaasik fiber.
Men’s clothing. — Men wear both arm- and head-bands during festi-
vals. These are made by the men from bird or other animal parts. There
are two types of armbands. The first, ara sina, consists of one to four
long red tail feathers from the macaw {Ara macao) tied to pieces of
80
Annals of Carnegie Museum
VOL. 55
Table 2>. — Yanomama ara sina armbands in the collections of Carnegie Museum of
Natural History.
Accession number
Macaw
feathers
Bunched feathers
Cordage
type
Length
(cm)
32703-6A
four
parrot & macaw
#1
66.1
-6B
four
white-winged guan
#1
65.6
-6C
four
macaw, parrot, and white-
winged guan
#1
64.2
-35A
three
parrot
#1
67.5
-35B
three
white-winged guan
#1
59.4
~35C
four
white-winged guan
#1
67.7
-35D
three
white-winged guan
#1
65.6
-35E
three
white-winged guan
#1
63.0
-35F
four
macaw, parrot and white-
winged guan
#1
64.0
-35 G
three
macaw
#1
68.7
32735-121A
four
white-winged guan
#1
60.9
-121B
four
white-winged guan
#1
66.7
-122A
four
white-winged guan
#1
51.8
-122B
four
white-winged guan
#1
64.1
-123
one
white-winged guan
#1
60.4
-124
one
parrot & white-winged guan
#9
61.6
-125
three
white-winged guan
#1
69.5
-126
three
white-winged guan
#1
68.0
-127
four
white-winged guan
#1
52.1
-128A
three
white-winged guan
#1
62.9
-128B
three
white-winged guan
#1
66.4
-129 A
three
white-winged guan
#1
68.4
-129B
three
white-winged guan
#1
66.1
-130A
four
white-winged guan
#1
66.2
-130B
four
white-winged guan
#1
62.0
-131A
three
white-winged guan
#1
64.7
-131B
three
white-winged guan
#1
64.6
-132A
four
macaw & parrot
#1
54.3
-132B
four
parrot
#1
68.6
-133A
four
macaw
#1
68.4
-133B
four
macaw
#1
69.6
-134A
four
parrot
#1
58.9
-134B
four
parrot
#1
72.2
-135A
four
parrot
#1
76.7
-135B
four
parrot
#1
60.0
-136A
four
macaw & parrot
#1
61.8
-136B
four
macaw & parrot
#1
59.0
-137A
four
parrot
#1
65.2
-137B
four
parrot
#1
53.9
-138
three
parrot
#1
69.7
-139
three
macaw
#1
62.0
-140
three
parrot
#2
63.5
1986
Couture-Brunette— Yanom AM A Material Culture, II
81
Table 3. — Continued.
Accession number
Macaw
feathers
Bunched feathers
Cordage
type
Length
(cm)
-141
three
parrot
#1
65.6
-142A
four
parrot
#1
63.1
-142B
three
parrot
#1
57.3
-143 A
three
parrot
#1
65.9
-143B
three
parrot
#1
72.0
-144 A
three
parrot
#1
67.4
-144B
three
parrot
#1
63.3
-MSA
four
parrot
#1
65.5
-MSB
four
parrot
#1
61.4
-146 A
four
parrot
#1
57.9
-M6B
four
parrot
#1
51.2
-147
two
parrot
#2
65.8
-148
four
parrot
#1
59.2
-149 A
four
parrot
#1
59.2
-M9B
four
parrot
#1
58.5
-ISOA
four
parrot
#1 and #2
51.4
-ISOB
four
parrot
#1
59.2
-ISIA
three
parrot
#1
61.9
-ISIB
three
parrot
#1
58.9
-152A
three
parrot
#1
71.1
-152B
three
parrot
#1
64.2
-153A
three
parrot
#1
69.9
-153B
three
parrot
#1
59.4
-1S4A
three
parrot
#1
66.5
-1S4B
three
parrot
#1
67.4
-15SA
three
parrot
#1 and #2
70.8
-ISSB
three
parrot
#1
67.3
bamboo or palm wood (Fig. 7). Bunches of parrot {Amazona sp.),
curassow {Crax alector), or white-winged or piping guan {Aburria pip-
He) feathers are tied in a “bouquet” at the base of the longer macaw
feathers. The armband is then tied around the upper arm with the long
red feathers pointing upwards. They are worn in pairs, with usually
one on each arm.
Table 3 presents the data for the ara sina armbands. Length, cor-
responding to the height of the long red tail feathers when worn, is not
a critical factor in matching pairs of armbands; six pairs differ in size
between pieces by 10 cm or more. It is more important to match the
feathers, both in number of long red macaw feathers and in color and
species of the “bouquet” feathers. All but two sets of armbands matched
both the number of long red macaw feathers and the “bouquet” feathers
82
Annals of Carnegie Museum
VOL. 55
Fig. S. — Yaro sina, man’s festival armband made from bird tails.
on both armbands. One pair, 32735- 142A & B, matched four long red
macaw feathers on one armband with three on the other. Only 32735-
132A & B did not match the “bouquet” feathers on the two armbands.
An interesting application of these facts can be tested with accession
32703 armbands, which were not delivered in pairs. Utilizing the cri-
teria discussed above for matching pairs, it can be suggested that pairs
are composed of specimens -6B and -35C, and -6C and -35F. Both of
these pairs then have matched macaw and “bouquet” feathers. Both
the macaw and the “bouquet” feathers match on three specimens (-35B,
-35D, and “35E); any two of these three could form a pair of armbands.
The “bouquet” feathers of -6 A and -3 5 A or -35G are somewhat matched.
Whatever the pairs of the last two groupings, it is clear that the arm-
bands from accession 32703 are not completely paired; the final set of,
for instance, -35E and -35G is composed of completely unmatched
“bouquet” feathers.
Cordage type #1 is clearly the choice to wrap the feathers to the
wood. Cordage type #2, the rare Z twist yamaasik fiber, is present on
four armbands, and s spin Z twist cotton cordage, type #9, on one.
Interestingly, pairs 32735- 150A & B and -155A & B utilize both S
1986
Couture-Brunette —Yanom AM A Material Culture, II
83
o
2
s
§
U W
S C O W' ^
U
m
^RX
u
fS|
m vn ^ fNi «M fNi
?<<<<<<
m ^
^ fNj
«N CN
R" ri
^ ^ r- ^ '^ '»-«'
w w ’— ' rn m m
^ fN| ^
*“''-^^•^^0X0^
0\ 0\ •>—1 0\ 0\ ^
i g
p p
o o
o
p
o
S g g *-* fl «
S g g g g g
p p p ^ p p
^ s ^
g g g
{« p ^
ess
a S fi
s I §
m Q4 m
g 2
p 5
o g
^ S
g S
ca so
e
^ VI ^o O VI
»n in in \o VO so
•
cn
O
r--
oj
cn
84
Annals of Carnegie Museum
VOL, 55
Fig. 9. — Yaw sina, man’s festival headband made from bird skins, wings, and tails.
and Z twist yamaasik fiber on one armband of the pair, whereas the
other exhibits standard S twist yamaasik fiber. It is possible that these
armbands which utilize mixed cordages were made by two individuals,
one of whom wrapped the long red macaw feathers to the wood while
the other attached the ‘‘bouquet” feathers. Accession 32703 armbands
use S twist yamaasik fiber exclusively.
The second type of men’s armband, the yaw sina (Fig. 8), is com-
posed of bird wings, tails, and skin. Aracari {Ptewglossus pluricinctus),
parrot, toucan (Ramphastos rucanus), and macaw wings, tails, and
skins are tied together to lengths of string. These lengths are tied around
the upper arm, leaving the wings dangling.
Men usually wear the yaw sina armbands as a pair, one on each
arm. Thus the armbands discussed in Table 4 are actually pairs of
1986
Couture-Brunette —Yanom AM A Material Culture, II
85
Table S. — Yanomama yaro sina headbands in the collections of Carnegie Museum of
Natural History.
Accession
number
Wings and tails
Headpiece
Cordage
Knots
Length*
(cm)
32703-57
parrot
curassow
#13(3)
A(3), C, F
32.4
-58
parrot
curassow
#13(3)
A(3), C, F, J
24.2
-59
none
curassow (2)
#12(6)
F
34.8
-61
toucan
none
#10
H, K
-62
none
curassow
#11 (3)
25.7
-63
none
curassow
#11 (3)
K
25.1
-66
parrot, curassow
curassow
#13(3)
A(3), C, F
38.0
-67
toucan, guan
curassow
#13(3)
A(2), C, J
23.5
-68
toucan, guan, ma-
caw, parrot
curassow
#3, 5 (3)
E, F, H, C(2)
30.4
* “Length” refers to the length of the headpiece.
armbands, which have been tied together. Some effort was made to
balance the armbands; a pair with a toucan tail on one armband will
be partnered with a toucan tail on the other. Only the smaller guan or
parrot wings are occasionally present on one member of the pair but
not the other.
Table 4 indicates that toucan tails are the parts used most consistently
by all who make the yaro sina armbands. Only one specimen uses
another bird tail in addition to the toucan tails, and all armbands but
32703-93 have toucan tails on them. Macaw wings are the second most
popular bird part, present on four of the seven armbands. When the
other parts are considered, primarily skins with feathers from the breast
or head, macaw parts are present on five of the seven armbands.
Cordage type #9, the 2 ply s spin Z twist cotton, is used on six of
the seven armbands. Cordage type #1 1(3) is present on four specimens,
and type #13(2) on three. Only two armbands exhibit yamaasik fiber
cordage, and only in minor amounts.
An interesting fact about the yaro sina armbands stems from the
variation possible in their mode of construction. There are many types
of cordage to choose from, and even more knots which can be used.
As with stitch slant in twining, discussed previously, knots tend to
reflect individual choice. Taking these facts into consideration, arm-
band pairs 32703-54 and -65 were probably made by the same man.
Both armbands used knot “H” exclusively to the the string to the bird
parts. None of the other specimens uses knot “H” in this fashion; in
fact, none of the other specimens uses knot '‘H” in any fashion. A
lark’s head is not a “fixed” knot, but rather a suspended knot capable
of movement along another piece of string. Although it can be rendered
relatively immobile by the tightness of the tie, it is unusual to find it
used as a method of permanent attachment. The specimens are not
86
Annals of Carnegie Museum
VOL. 55
Fig. 1 0. —Incomplete yaw sina headbands, consisting of just the curassow skin headband.
overly similar in any other attribute; they both make heavy use of
toucan parts, as do many of the other yaro sina armbands; they both
use cordage types #9 and #13(2), which are the first and third most
popular cordages; and one uses yamaasik fiber cordage, found on one
other specimen. Nevertheless, the attachment of the parts to the string
is unusual enough to merit special attention, and use of the rare lark’s
head knot is a distinguishing characteristic which links these two pairs
of armbands and separates them from the others.
The second item of men’s festival apparel is the headband made of
bird parts or monkey skin. While the monkey headbands are simply
made of skinned monkey tails, the feather yaro sina headbands, like
the yaro sina armbands, are complex constructions of bird wings and
tails tied to cotton or fiber cordage (Fig. 9). The band which passes
across the forehead is made of curassow skin and feathers. A string is
tied at each end of the band to make the head size adjustable, and the
bird wings and tails dangle at each temple.
The yaro sina headbands are not as balanced as the yaro sina arm-
bands; little effort was made to place similar wings and tails at each
temple. As can be seen on Table 5, many of the headbands (32703-
1986
Couture-Brunette— Yanomam A Material Culture, II
87
Fig. 11.— Wisa sina, man’s festival headband made from a skinned black saki monkey
tail. Accession 32735.
58, "59, -62, and -63) consist of just the curassow headband with no
dangling parts (Fig. 10). Although bands are occasionally worn this
way, it is usual to complete them with other bird parts at the temples.
The reverse situation is found on 32703-61, which consists only of the
earpieces to a headband; it has not yet been added to, or has been
detached from, the curassow band.
Cordage type #13(3) is the most popular cordage used on the yaw
sina headbands, present on four of the nine specimens. The rest of the
cordage consists of two pieces of cordage type #1 1(3), and one each of
types #12(6), #10, #3, and #5(3). In this respect the yaw sina headbands
differ from the yaw sina armbands; cordage usage on the armbands
was very consistent, with cordage type #9 present on six of the seven
armbands, type #11(3) on four, and type #13(2) on three. Yamaasik
fiber was used on one headband. Knot “A” was the most popular knot
in total number of ties, whereas knots ‘‘C” and “F” were used on the
greatest number of headbands.
A second type of headband, also utilizing feathers, is woven of masik
vine. Curassow and toucan feathers are inserted into the weave in
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Annals of Carnegie Museum
VOL. 55
Fig. \l.~Horokoto with yakoana, hallucinogenic drug storage gourd. CMNH number
32703-22B.
patches of red, yellow, and black. This headband is not adjustable to
different sizes.
32735-20, mayop ahuk, owned by Tixo. Band is tied at four points with cordage types
#1 and #3 in knots “F”(2) and “A.” A 12 cm length in the back of the headband
has no feathers. D™ 18.0 cm.
Other festival headbands consist of skinned monkey tails (Fig. 1 1).
The fur piece is flattened, and an adjustable string is tied at each end.
32735-194, wixa sina. Black saki monkey skin tied with cordage type #3 in knot “H”
at one end. L--(tail only) 42.7 cm.
32735-195, same as -194 but string tied with knot “E.” L— 41.2 cm.
32735-196, wixa sina tied with cordage type #19 in knot “C.” L— 36.9 cm.
Drug Containers
Most men use hallucinogenic drugs during festivals, al-
though shamans make frequent use of them when performing sha-
manistic rituals. The drug ( Virola sp.), also known as ebene, is made
from inner tree bark (Chagnon et al., 1971:72-74). The latex from the
tree is used as the poison on monkey arrows (Couture-Brunette, 1985:
501).
1986
Couture-Brunette —Yanom AM A Material Culture, II
89
Fig. 13. —Chest pounding tools. From top: paruk xayau, CMNH number 32703-25 A;
paruk xayau, CMNH number 32703-25B; suhumuk, CMNH number 32703-25C.
The moist scrapings are mixed with ashes and kneaded into a ball,
with saliva providing additional moisture if necessary. It is then placed
over the fire on a container to dry (Chagnon, 1977:23). The residue is
ground into a powder, and stored in gourd containers until needed (Fig.
12).
32703-22A, horokoto with yakoana, from Mahuku of Hewenahipiutheri village, Decem-
ber 1979. Corked with ball of wax; cordage type #9 through hole. Knot “A” inside
gourd; two knot “A”s other end. L— 14.2 cm.
32703-22B, same as 32703-22A but corked with roll of animal hide. Cordage type #3
through hole in neck; tied with knot “E.” Outside of gourd covered with resin. L—
15.9 cm.
32735-102, sealed with wax. Cordage type #3 through neck hole; two knot “A”s one
end and three on the other. L— 1 1 .4 cm.
32735-103, brown wood cork. Hole drilled in neck; no string. L— 15.1 cm.
32735-104, owned by Mahuku. Top sealed with wax, no hole or string. Split down side.
L-12.4 cm.
32735-105, top sealed with wax. Cordage type #3, no visible knots. L— 15.3 cm.
32735-341, sealed with wax. L— 12.2 cm.
The drug is blown into the nostrils with a mokamosi, a hollow cane
tube. One end of the tube is lined with wax while the other is unmod-
ified. The tube is loaded with a dose of yakoana, and the receiver places
90
Annals of Carnegie Museum
VOL. 55
CAre^MftO*
CS4 6«
VAOOmA ry^O
Fig. 14.— Haya maro, deer bone flute (top) and nara wakirim, lump of red body paint
(bottom). Accession 'ill 35.
the waxed end inside one nostril. Another man expels a powerful gust
of air into the cane. A man will take a few doses of the drug at a time
in each nostril.
32703-41, collected in 1979--80. Lengthwise cracks and splits along the sides of the tube.
L— 8^4 cm.
32735-89, four parrot feathers in the waxed end. L— 69.8 cm.
32735-90, same as 32735-89. L^69.0 cm.
Chest Pounders
Ritual chest pounding duels during festivals have replaced much of
the revenge warfare among the Catrimani Yanomama. Blows are de-
livered with the aid of chest pounders (paruk xayau), wood branches
which are smoothed and polished (Fig. 1 3). No effort is made to trim
off protruding stubs, and the ends may be pointed or flat.
32703-25A, from Akasi and Paruk of Uxiutheri village, November 1979. Ends unpoint-
ed. L— 35.9 cm.
32703-25B, same as 32703-25A but both ends pointed. L— 40.4 cm.
32735-109, owned by Carlos. Both ends pointed; approximately 5 cm from each end
red and yellow toucan feathers are wrapped onto the pounder with cordage type #1.
L— 22.1 cm.
1986
Couture-Brunette —Yanom AM A Material Culture, II
91
32735-263, one end pointed and other rounded. Cordage type #9 wrapped 10 times
around one end. L— 28.9 cm.
32735-264, ovoid, both ends pointed. Small hole partially drilled into one side. “ME”
carved on other side. L— 18.2 cm.
32735-265, forked. Two top ends pointed, other (bottom) end flat. L— 20.2 cm.
A second type of ritual dueling tool, suhumuk, is used in a different
fashion. This object, also made of wood, is blade shaped with pointed
ends and serrated teeth in the center (Fig. 1 3). Both ends are grasped
by one man, who then hugs his opponent and digs the teeth into his
back while the opponent does the same.
32735-25C, from Akasi and Paruk of Uxiutheri village, November 1979. Four teeth;
one end pointed, other flat. L— 38.5 cm.
32735-191, four teeth; both ends pointed. L— 77.6 cm.
32735-31, maama suhumuk, stone found by Honi on the path near the Arapari River
(KM 135 of Highway BR 210). Honi said these were not longer used in dueling, but
were made and used by ancestors. L— 23.4 cm.
Miscellaneous Festival Accessories
The Yanomama use two kinds of musical instruments, both of them
flutes. The first is a deer bone flute (Fig. 14). Its use is restricted to
festivals, when it is played by the guests as they draw near the host
village. The leg bone is hollowed and holes are drilled into one side.
The joint forms a natural closure for one end, while the other end of
the flute is open. The flute is usually reddened with nara. The second
flute, which is a child’s toy imitation of the ritual flute, will be discussed
in Part III as a miscellaneous construction.
32703-49A, haya maro, collected 1979-80, 2 holes drilled in top. L---12.4 cm.
32703-49B, same as 32703-49A but three holes drilled in top. L— 15.1 cm.
32703-49C, same as 32703-49B. L-14.6 cm.
32735-50, two drilled holes and one natural hole, faint traces of nara. L— 18.2 cm.
32735-51, same as 32735-50. L-=17.8 cm.
32735-52, three drilled holes, reddened with nara. L— 18.9 cm.
32735-303, three drilled holes alternate with three “X”s scratched onto the surface.
Reddened with nara. L— 19.0 cm.
32735-304, same as 32735-303. L--19.4 cm.
Two necklaces, although occasionally worn outside of festivals, are
intimately associated with the spirit world and were thus grouped in
the Festival category with the drug equipment. The first, aroariki (Fig.
3), is worn for protection against evil spells and spirits. It is composed
of cut pieces of tubers which have been dyed brown. The pieces are
threaded on brown dyed string.
32703-8 A, from Puuxim of Wakathautheri village. Cordage type #10, eight knot “A”s.
L— 35.4 cm.
32703-8B, same as 32703-8A. Cordage type #3, with knots “A”(5), “E”(2), and “J”(2).
L--38.2 cm.
92
Annals of Carnegie Museum
VOL. 55
The other type of necklace, marasik, is worn by men. It acts as an
aphrodisiac, attracting partners of the opposite sex. It consists of cut
pieces of tubers, which are dyed red and tied to reddened cotton string.
32735-44A, cordage type #11(3) tied around center of each tuber in knots “E,” “G,”
and “F.” Ends tied together with two knot “A”s. L--70.0 cm.
32735-44B, same as 32735-44A but string tied around tubers with knot “E.” Ends tied
together with knot “E.” L--69.1 cm.
Additional Comments
The two categories of Wearing Apparel and Festival Artifacts are
intimately related, inasmuch as most festival artifacts are items to be
worn. The division of specimens into selected categories was intended
to reflect the function of the artifact. However, in two cases the con-
struction of the specimen outweighed the functional aspects. Specifi-
cally, the women’s aprons and the body paint groups were discussed
together under one category, although in function each group should
have been separated. In the case of the aprons, this combined discussion
followed a constructional basis. Everyday and festival aprons are made
the same way, and differ only in the added decorative elements. As
this was the case, the “apron” group was combined into one discussion
section under “Wearing Apparel” rather than presenting separate but
identical constructional data in two categories. The “body paint” group
was similarly combined for the same reasons.
Acknowledgments
I wish to thank Dr. Giovanni Saffirio for his invaluable advice and explanations on
particularly puzzling specimens. Drs. James B. Richardson and David R. Watters kindly
read through initial drafts and provided helpful comments. Dr. Hugh H. Genoways and
the staff of the Section of Mammals, Drs. Kenneth Parkes and D. Scott Woods of the
Section of Birds, and Dr, Juan Parodiz and Ms. Jay Tripp of the Section of Malacology
assisted in the identification of materials used in manufacture of many of the artifacts.
Ms. Nancy Perkins of the Division of Exhibits drafted the knots as well as the cordage/
basketry twist examples. Specimen photographs were taken by Ms. Barbara Gundy; all
other photographs are used with the kind permission of Giovanni Saffirio.
Literature Cited
Adovasio, J. M. 1977. Basketry technology— a guide to identification and analysis.
Aldine Publ. Co., Chicago, Illinois, 182 pp.
Chagnon, N. a. 1974, Studying the Yanomamo. Holt, Rinehart, and Winston, New
York, New York, 270 pp.
— . 1977. Yanomamo: the fierce people. Holt, Rinehart, and Winston, New York,
New York, 2nd ed., 174 pp.
Chagnon, N. A., P. Le Quesne, and J. M. Cook. 1971. Yanomamo hallucinogens:
anthropological, botanical, and chemical findings. Current Anthropology, 12:
72-74.
Couture-Brunette, L. 1985. Yanomama material culture in the Carnegie Museum
1986
Couture-Brunette— Yanom AM A Material Culture, II
93
of Natural History. Part I. Food procurement and household articles. Ann. Carnegie
Mus. 54:487-532.
Emery, I. 1980. The primary structure of fabrics. The Textile Mus. Press, Washington,
D.C., 2nd ed., 339 pp.
Migliazza, E. 1972. Yanomama grammer and intelligibility. Ph.D. dissertation, In-
diana Univ., University Microfilms #72-30,432, Ann Arbor, Michigan, 457 pp.
Saffirio, G. 1980. Some social and economic changes among the Yanomama of
northern Brazil (Roraima): a comparison of “forest” and “highway” villages. Un-
published MS thesis, Univ. Pittsburgh, Pennsylvania, 1 1 9 pp.
— 1985. Ideal and actual kinship terminology among two Yanomama villages of
the Catrimani River basin (Brazil). Unpublished Ph.D. dissertation, Univ. Pitts-
burgh, Pittsburgh, Pennsylvania, 244 pp.
Saffirio, G., R. Hames, N. Chagnon, and T. Melancon. 1983. The impact of contact:
two Yanomamo case studies. Pp. 1--52, in Working papers on South American
Indians, Bennington College Press, Bennington, Vermont, 6.
Shaw, G. R. 1972. Knots, useful and ornamental. First Collier Books ed., Macmillan
Publ. Co. Inc., New York, New York, 194 pp.
Smole, W. 1976. The Yanoama Indians: a cultural geography. Univ. Texas Press,
Austin, 272 pp.
Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.
ISSN 0097-4463
ANNALS
ofCAf!jNECIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 23 MAY 1986 ARTICLE 5
STANDARD KARYOLOGY OF NINE SPECIES OF
VESPERTILIONID BATS
(CHIROPTERA: VESPERTILIONIDAE)
FROM THAILAND
Karen McBee
Rea Postdoctoral Fellow, Section of Mammals
John W. Bickham^
SONGSAKDI YeNBUTRA^
Jarujin Nabhitabhata^
Duane A. Schlitter
Curator, Section of Mammals
Abstract
Karyotypes of nine species of vespertilionid bats from Thailand are described. Pip-
istrellus mimus (2n = 34, FN = 46), Tylonycteris robustula (2n = 32, FN = 50), Murina
leucogaster (2n = 44, FN = 50), and Miniopterus schreibersi (2n = 46, FN = 52) have
karyotypes essentially identical to ones previously reported from other regions. Pipis-
trellus pulveratus (2n = 32, FN = 50) is reported for the first time and differs by six
Robertsonian fission/fusion events from the primitive MyotisA.ik.Q karyotype. Karyotypes
95
96
Annals of Carnegie Museum
VOL. 55
for Hesperoptenus tickelli (2n = 32, FN = 50) and H. blanfordi (2n = 34, FN = 60) are
reported for the first time and parallel the extreme morphological differences between
the two species. Harpiocephalus mordax (2n = 40, FN = 62) is very distinct from other
members of the subfamily Murininae but is apparently derived from a Murina-\ik&
ancestor. Kerivoula papillosa (2n = 38, FN = 52) though considered little differentiated
from primitive vespertilionines has a relatively highly derived karyotype similar to
Vespertilio.
Introduction
The family Vespertilionidae is distributed worldwide in temperate
and tropical regions. It is the largest family in the order Chiroptera
including approximately 33 genera and 3 1 3 Recent species (Koopman,
1984). Thirteen genera and 34 species are known to occur in Thailand
(Lekagul and McNeely, 1977).
Previous karyotypic studies led Pathak and Sharma (1969) to suggest
that the family has two very different patterns of chromosomal vari-
ability. Some genera such as Myotis exhibit remarkable homogeneity
with all examined species having 2n = 44, FN = 50 or 52. Others such
as Pipistrellus (2n = 26, 28, 30, 32, 34, 36, 38, 42, 44 and FN = 44,
46, 48, 50, 52) are much more heterogeneous. These studies, however,
mostly have been restricted to New World (Baker and Patton, 1967;
Bickham, 1979<2, 1979Z?) and European (Bovey, 1949; Capanna and
Civitelli, 1970; Fedyk and Fedyk, 1970; Zima, 1978) species. Karyo-
typic data for African, Australian, and Asian vespertilionids are sparse.
For example, karyotypes have been reported for only one species of
vespertilionids from Thailand (Harada et al., 1982Z?).
This study presents standard karyotypes of nine species in seven
genera and four subfamilies from Thailand. Karyotypes of five of these
species have been reported from other regions (Pathak and Sharma,
1969; Manna and Talukdar, 1965; Yong et al., 1971; Bickham and
Hafner, 1978; Harada and Kobayashi, 1980; Harada, 1973; Ando et
al., 1977). New data are presented for four species and one subfamily.
Materials and Methods
All animals were collected in Thailand using mist nets. Upon capture, all animals were
subcutaneously injected with a weak solution of baker’s yeast, sugar, and water (Lee and
Elder, 1980) to stimulate bone marrow mitosis. Twenty-four hours later, animals were
sacrificed and humeri removed. Karyotypes were prepared in the field from bone marrow
cells suspended in a hypotonic solution (0.075 M KCl) for approximately 25 min and
then fixed in a 3:1 solution of methanol : glacial acetic acid (Baker et al., 1982). Three
to four drops of the fixed cell suspension were dropped onto clean, dry microscope slides
and ignited with a match. After the flaming suspension extinguished itself, any remaining
liquid was carefully drained away and slides were stained in a 2% solution of Geimsa
in 0.01 M phosphate buffer. Diploid (2n) and fundamental (FN) numbers were deter-
mined from counts of a minimum of 10 mitotic spreads. Description of chromosome
morphology follows the nomenclature of Patton (1967). All specimens were prepared as
1986
McBee et al.™ Thailand Vespertilionid Karyology
97
museum skins and skulls or alcoholics and are housed in the Carnegie Museum of Natural
History (CM), the Texas Cooperative Wildlife Collection, Texas A&M University (TCWC),
or The Museum, Texas Tech University (TTU).
Specimens Examined
Pipistrellus mimus.—UTnM Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife
Sanctuary, 15°29'N, 99®18'E (CM 88129 M, 88130 F); Huai Kha Khang Wildlife Sanc-
tuary, 2.7 km S Khao Nang Rum Wildlife Research Station, 15®27'N, 99®18'E (CM
88132 M); Huai Kha Khang Wildlife Sanctuary, 1.5 km W Khao Nang Rum Wildlife
Research Station, 15°29'N, 99n7'E (CM 88131 M).
Pipistrellus pulveratus. — Uthai Thani Prov,; Lan Saka Dist., Huai Kha Khang Wild-
life Sanctuary, 3.7 km S, 1 km E Khao Nang Rum Wildlife Research Station, 15®27'N,
99°18'E (CM 88134 F, CM 88136 F).
Tylonycteris robustula. — SuKAT Thani Prov.; Tha Chang Dist., 15 km N, 23 km W
Ban Maruan, 09°18'N, 98®58'E (CM 88149 F, CM 88151 F, CM 88152 F, CM 88140
F, TTU 41257 F, TK 21416 F).
Hesperoptenus blanfordi.—UTHM. Thani Prov.; Lan Saka Dist., Huai Kha Khang
Wildlife Sanctuary, 15°19'N, 99°18'E (CM 881 14 M, TTU 41255 F); Huai Kha Khang
Wildlife Sanctuary, 1.5 km W Khao Nang Rum Wildlife Research Station (TK 21279 F).
Hesperoptenus tickelli. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wild-
life Sanctuary, 3.7 km S, 1 km E Khao Nang Rum Wildlife Research Station, 15®27'N,
99®18'E (CM 881 19 M, TK 21 193 M); Huai Kha Khang Wildlife Sanctuary, 2.0 km S
Khao Nang Rum Wildlife Research Station, 15°30'N, 99°16'E (CM 88117 M).
Kerivoula papillosa. —Surat Thani Prov.; Tha Chang Dist., 15 km N, 23 km W Ban
Maruan, 09°18'N, 90®58'E (CM 88164 F).
Miniopterus schreibersi haradai. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha
Khang Wildlife Sanctuary, 2.7 km S Khao Nang Rum Wildlife Research Station, 1 5°30'N,
99®16'E (CM 88156 M); Huai Kha Khang Wildlife Sanctuary, 3.7 km S, 1 km E Khao
Nang Rum Wildlife Research Station, 15°27'N, 99“18T (CM 88157 M).
Murina leucogaster.— Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang Wildlife
Sanctuary, Khao Nang Rum Wildlife Research Station, 15®29'N, 99®18'E (CM 88162 F,
CM 88163 F).
Harpiocephalus mordax. —Uthai Thani Prov.; Lan Saka Dist., Huai Kha Khang
Wildlife Sanctuary, Khao Nang Rum Wildlife Research Station, 15®29'N, 99®18'E (CM
88159 F).
Results
Table 1 is a summary of the known standard karyotypic data for the
family Vespertilionidae including those reported here. The standard
karyotypes of eight species representing seven genera and four sub-
families are presented in Figs. 1--3. A brief description of these karyo-
types follows.
Subfamily Vespertilioninae
Pipistrellus mimus (2n 34, FN = 46; Fig. la). —Four animals
examined have a karyotype that consists of six pairs of large metacentric
to submetacentric chromosomes and one large subtelocentric pair. There
are nine pairs of acrocentric chromosomes ranging in size from medium
98
Annals of Carnegie Museum
VOL. 55
Table l.—A summary of known standard karyotypic data for the family Vespertilionidae.
SM—submetacentric, M—metacentric, ST—subtelocentric, and A— acrocentric.
Taxon
Subfamily Vespertilioninae
Myotis auriculas
Myotis austroriparius
Myotis bechsteini
Myotis blythi
Myotis brandti
Myotis californicus
Myotis capaccinii
Myotis dasycneme
Myotis daubentonii
Myotis elegans
Myotis emarginatus
Myotis evotis
Myotis fortidens
Myotis frater
Myotis grisescens
Myotis horsfieldi
Myotis hosonoi
Myotis keaysi
Myotis keenii
Myotis leibii
Myotis lucifugus
Myotis macrodactylus
2n
FN
X
Y
44
52
SM
A
44
50
SM
SM
44
50
SM
A
44
52
M
A
44
52
SM
A
44
50
SM
A
44
50
SM
A
44
50
SM
SM
44
50
_
_
44
52
M
A
44
50-52
SM
A
44
54
SM
44
52
M
A
44
50
SM
SM
44
50
SM
A
44
56
M
A
44
52
M
A
44
50
SM
SM
44
52
SM
A
44
50
SM
A
44
50
SM
A
44
50
SM
A
44
50
SM
A
44
50
SM
A
44
52
SM
A
44
50
SM
A
44
50
SM
A
44
50
—
44
50
SM
SM
44
50
SM
A
44
50
SM
SM
44
50
SM
SM
44
52
SM
A
44
52*
SM
A
44
50
SM
A
Authority
Bickham, \919b
Baker and Patton, 1967
Bickham, \919b
Zima, 1978
Baker, 1970
Baker et aL, 1974
Zima, 1982
Baker and Patton, 1967
Manfreddi Romanini et
aL, 1975
Zima, 1978
Bovey, 1949
Fedyk and Fedyk, 1970
Zima, 1984
Baker and Patton, 1967
Bovey, 1949
Radjhabli et al., 1969,
1970
Zima, 1978
Baker and Patton, 1967
Bickham, \919b
Osborne, 1965
Harada and Yoshida,
1978
Baker and Patton, 1967
Bickham, 1979^
Harada and Kobayashi,
1980
Harada, 1973
Harada and Yoshida,
1978
Bickham, 1979^
Baker and Bickham, 1980
Baker and Patton, 1967
Bickham, \919b
Baker and Patton, 1967
Baker and Patton, 1967
Harada, 1973
Obaraetal., 1976a
Harada and Yoshida,
1978
1986
McBee et al.— Thailand Vespertilionid Karyology
99
Taxon
Myotis milieri
Myotis myotis
Myotis mystacinus
Myotis nattered
Myotis nigricans
Myotis oxygnathus
Myotis pruinosus
Myotis sodalis
Myotis thysanodes
Myotis velifer
Myotis {Pizonyx) vivesi
Myotis volans
Myotis yumanensis
Lasionycteris noctivagans
Pipistrellus abramus
Pipistrellus affinm
Pipistrellus babu
Pipistrellus endoi
Pipistrellus hesperus
Pipistrellus kuhli
Pipistrellus mimus
Table 1. — Continued.
2N
FN
X
Y
44
52
SM
__
44
50
M
A
44
50
SM
A
44
50
SM
A
44
56
M
A
44
50
SM
__
44
50
SM
__
44
52
SM
A
44
50
SM
SM
44
50
SM
A
44
50
SM
A
44
56
M
A
44
50
SM
A
44
52
SM
ST
44
50
SM
—
44
50
SM
A
44
50
SM
SM
44
52
SM
A
44
50
SM
SM
44
50
SM
A
44
50
SM
SM
44
50
SM
SM
44
50
SM
__
44
50
SM
A
20
28
SM
A
20
28
SM
A
26
44
ST
A
26
44
A
A
26
44
A
A
36
50
SM
A
36
50
M
A
38
50
A
__
38
50
A
28
46
SM
A
44
50
SM
A
44
50
SM
__
44
50
SM
A
34
__
__
__
Authority
Reduker et al., 1983
Bovey, 1949
Bickham and Hafner,
1978
Iliopoulou-Georgudaki
and Giagia, 1984
Radjhabli et al., 1969,
1970
Ando et al., 1977
Haradaand Yoshida, 1978
Zima, 1978
Baker and Patton, 1967
Bickham, \919b
Baker and Bickham, 1980
Radjhabli et al., 1969,
1970
Bickham and Hafner,
1978
Harada and Uchida, 1982
Baker and Patton, 1967
Bickham, \919b
Baker and Patton, 1967
Bickham, \919b
Baker and Patton, 1967
Bickham, \919b
Baker and Patton, 1967
Baker and Patton, 1967
Baker and Patton, 1967
Bickham, 1979^
Baker and Patton, 1967
Bickham, 1979<2
Takayama, 1959
Harada, 1973
Obara et al., \916b,
1976c
Pathak and Sharma, 1969
Dulic, 1981
Ando et al., 1977
Ando et al., 1980
Baker and Patton, 1967
Capanna, 1968
Baker et al., 1974
Zima, 1982
Manna and Talukdar,
1965
100
Annals of Carnegie Museum
VOL. 55
Table Continued.
Taxon
Pipistrellus mordax
Pipistrellus nanus
Pipistrellus nathusii
Pipistrellus pipistrellus
Pipistrellus savii
Pipistrellus pulveratus
Pipistrellus subjlavus
Nyctalus furvus
Nyctalus lasiopterus
Nyctalus leisleri
Nyctalus noctula
Eptesicus andinus
Eptesicus brasiliensis
Eptesicus capensis
Eptesicus circumdatus
Eptesicus diminutus
Eptesicus furinalis
Eptesicus fuscus
Eptesicus guadeloupensis
2N
FN
X
Y
38
48
M
A
34
46
SM
A
34
46-
M
A
36
50
M
A
44
51
SM
A
42
50
M
42
50
M
A
42
51
M
A
44
50
M
—
44
50
M
A
44
50
SM
A
44
52
M
A
44
50
SM
A
44
50
__
—
44
50
SM
A
32
50
SM
__
30
56
SM
A
30
50
SM
44
50
SM
A
44
50
SM
A
42
50
SM
A
42
50
SM
A
42
50
SM
A
42
50
SM
A
46
54
SM
42
50
SM
A
42
50
SM
A
42
50
M
A
42
50
M
M
50
48
SM
A
50
48
SM
A
50
48
SM
A
32
50
SM
A
50
48
SM
__
50
48
SM
A
50
48
SM
A
50
48
SM
A
50
48
SM
A
50
48
SM
A
50
48
SM
A
Authority
Pathak and Sharma, 1969
This study
Pathak and Sharma, 1969
Peterson and Nagorsen,
1975
Bovey, 1949
Fedyk and Ruprecht,
1976
Zima, 1978
Bovey, 1949
Fedyk and Ruprecht,
1976
Zima, 1978
Zima, 1982
Zima, 1984
Capanna, 1968
Park and Won, 1978
Zima, 1982
This study
Baker and Patton, 1967
Bickham, \919a
Ando et al., 1977
Harada et al., 1982<2
Tsuchiya et al,, 1972
Harada, 1973
Ando et al., 1977
Harada et al., 1982t2
Fedyk and Fedyk, 1970
Dulic et al., 1967
Vorontsov, 1969
Zima, 1978
Zima, 1984
Baker and Patton, 1967
Baker and Patton, 1967
Baker et al., 1982
Peterson and Nagorsen,
1975
Heller and Volleth, 1984
Williams, 1978
Baker and Patton, 1967
Williams, 1978
Baker and Patton, 1967
Bickham, 1979a
Genoways and Baker,
1975
1986
McBee et al.— Thailand Vespertilionid Karyology
101
Table \.— Continued.
Taxon
2N
FN
X
Y
Authority
Eptesicus hottentotus
50
48
SM
—
Peterson and Nagorsen,
1975
Eptesicus japonensis
50
48
SM
SM
Ando et al., 1977
Eptesicus lynni
50
48
SM
A
Bickham, 1979^?
Eptesicus nilssoni
50
48
__
—
Ando et al., 1977
50
50
M
A
Zima, 1978
50
48
M
__
Zima, 1982
Eptesicus serontinus
50
48
SM
A
Baker and Patton, 1967
50
48
SM
A
Vorontsov, 1969
50
52
SM
SM
Fedyk and Fedyk, 1970
50
48
SM
—
Baker etal., 1974
50
48
SM
A
Bickham, 1979^2
50
48
SM
_
Baker and Bickham, 1980
Vespertilio murinus
38
50
M
A
Vorontsov, 1969
38
54
M
A
Zima, 1978
38
50
—
—
Obara and Saitoh, 1977
Vespertilio orientalis
38
50
M
A
Ando et al., 1977
38
50
SM
A
Obara and Saitoh, 1977
Vespertilio superans
38
50
M
A
Vorontsov, 1969
38
50
M
A
Zima, 1978
Histiotus montanus
50
48
SM
A
Williams and Mares,
1978
Tylonycteris pachypus
46
52
A
M
Yonget al., 1971
Tylonycteris robustula
32
52
A
M
Yonget al., 1971
32
52
A
M
This study
Hesperoptenus blanfordi
34
60
A
-
This study
Hesperoptenus tickelli
32
46
ST
M
This study
Nycticeius humeralis
46
48
SM
A
Baker and Patton, 1967
46
48
SM
A
Bickham, 1979^?
Scotoecus hindei
30
50
ST
SM
Nagorsen et al., 1976
Rhogeessa genowaysi
42
50
SM
SM
Baker, 1984
Rhogeessa parvula
44
50
SM
SM
Baker and Patton, 1967
44
50
SM
Bickham and Baker, 1977
Rhogeessa tumida
42
50
SM
SM
Baker and Patton, 1967
30
50
__
_
Baker, 1970
42
50
SM
SM
Bickham and Baker, 1977
34
50
SM
SM
Bickham and Baker, 1977
32
50
SM
SM
Bickham and Baker, 1977
30
50
SM
ST
Bickham and Baker, 1977
34
50
_
__
Baker and Bickham, 1980
30
50
__
Baker and Bickham, 1980
52
52
_
Honeycutt et al., 1980
34
50
SM
__
Baker et al., 1985
32N
50
SM
_
Baker etal, 1985
32B
50
SM
__
Baker et al, 1985
30
50
SM
A
Baker et al, 1985
102
Annals of Carnegie Museum
VOL. 55
Table \. — Continued.
Taxon
2N
FN
X
Y
Authority
Scotophilus dinganii
36
52
A
M
Schlitter et al., 1980
36
62*
“
_
Peterson and Nagorsen,
1975
Scotophilus heathi
36
52
M
A
Sharma et al., 1974
Scotophilus kuhlii
36
52
M
A
Pathak and Sharma, 1969
36
48
M
A
Harada et al., 1982^?
Scotophilus temminckii
36
52
SM
A
Pathak and Sharma, 1969
36
48
SM
A
Harada and Kobayashi,
1980
Scotophilus viridis
36
54
A
M
Schlitter et al., 1980
Lasiurus borealis
28
46
SM
A
Baker and Patton, 1967
28
48
SM
A
Baker and Mascarello,
1969
28
48
SM
A
Bickham, 1979<2
Lasiurus cinereus
28
46
SM
A
Baker and Patton, 1967
28
48
SM
A
Bickham, \919a
Lasiurus ega
28
48
SM
A
Bickham, \919a
Lasiurus ega panamensis
28
46
A
A
Baker and Patton, 1967
Lasiurus ega xanthinus
28
46
SM
A
Baker and Patton, 1967
Lasiurus inter medius
26
40
SM
A
Baker and Patton, 1967
26
42
A
A
Baker, 1970
Lasiurus seminolus
28
48
SM
A
Baker and Mascarello,
1969
28
48
SM
A
Bickham, 1979<2
Barbastella barbastellus
32
52
__
—
Matthey and Bovey, 1948
32
50
M
A
Bovey, 1949
32
50
SM
A
Capanna et al., 1968
32
52
SM
A
Zima, 1978
Barbastella leucomelas
32
50
SM
A
Ando et al., 1977
Plecotus auritus
32
52
__
Matthey and Bovey, 1948
32
50
M
A
Bovey, 1949
32
54
SM
A
Fedyk and Fedyk, 1970
Plecotus auritus auritus
32
54
M
A
Ando et al., 1977
32
52
M
A
Zima, 1978
Plecotus auritus sacrimontis
32
50
M
A
Harada, 1973
Plecotus austriacus
32
50
SM
A
Baker, 1970
32
54
SM
A
Fedyk and Fedyk, 1970
32
50
SM
A
Baker et al., 1974
32
52
M
A
Zima, 1978
Plecotus phyllotis
30
50
_
__
Baker and Patton, 1967
30
50
SM
A
Baker and Mascarello,
1969
Idionycteris phyllotis
30
50
SM
_
Bickham, 1979a
30
50
SM
““
Stock, 1983
1986
McBee et al.— Thailand Vespertilionid Karyology
103
Table \. — Continued.
Taxon
2N
FN
X
Y
Authority
Plecotus rafinesquii
32
50
A
A
Baker and Mascarello,
1969
Plecotus townsendi
32
48
__
—
Baker and Patton, 1967
32
50
A
A
Baker and Mascarello,
1969
32
50
A
A
Bickham, 1979a
32
50
A
A
Stock, 1983
Euderma maculatum
30
50
SM
A
Williams et al., 1970
30
50
SM
-
Stock, 1983
Subfamily Miniopterinae
Miniopterus australis
46
50
SM
A
Harada and Kobayashi,
1980
Miniopterus magnater
46
50
SM
A
Harada and Kobayashi,
1980
Miniopterus schreibersi
46
50
_
__
Matthey and Bovey, 1 948
46
50
SM
A
Baker et al., 1974
46
50
SM
A
Bickham and Hafner,
1978
46
50
SM
A
Bickham, 1979a
46
50
SM
A
Harada and Kobayashi,
1980
Miniopterus schreibersi ha-
46
52
SM
A
This study
radai
Miniopterus schreibersi fuli-
46
52
SM
A
Harada, 1973
ginosus
Subfamily Murininae
Murina aurata
44
60
SM
A
Ando et al., 1977
Murina leucogaster
44
50
—
Harada, 1973
44
58
SM
A
Ando et al., 1977
44
50
SM
A
This study
Harpiocephalus mordax
40
62*
-
This study
Subfamily Kerivoulinae
Kerivoula papillosa
38
52**
This study
Subfamily Nyctophilinae
Antrozous pallidus
56
50
SM
A
Bickham, 1979a
Bauerus dubiaquercus
44
52
SM
A
Engstrom and Wilson,
1981
* Obara et al., 1976^, report on inversion polymorphism in chromosome 5.
** Includes sex chromosomes in FN.
* Examination of the figure in Peterson and Nagorsen (1975) gives a FN = 52. This
probably represents a typographical error.
104
Annals of Carnegie Museum
VOL. 55
114 4# 4A 4A #•
la
II!
Fig. 1 . — The standard karyotypes of: a) Pipistrellus mimus F (CM 88 1 35), 2n = 34, FN =
46, inset F. mimus M (CM 88131); b) Pipistrellus pulveratus F (CM 88136), 2n = 32,
FN = 50.
to minute. The X is medium-sized and submetacentric and the Y is
small and acrocentric.
Pipistrellus pulveratus (2n = 32, FN = 50; Fig. lb). —The autosomal
complement includes eight pairs of metacentric or submetacentric chro-
mosomes ranging in size from large to medium. There is one large pair
and one small pair of subtelocentric chromosomes, and five pairs of
acrocentric chromosomes ranging from medium-sized to small. The X
is medium-sized and submetacentric.
Tylonycteris robustula (2n = 32, FN = 50; Fig. 2a).— The karyotype
shown here is similar to that reported by Yong et al. (1971), but there
are slight differences. Both studies report 2n = 32 with nine pairs of
metacentric to submetacentric chromosomes. However, our specimens
had one pair of medium-sized subtelocentric chromosomes and five
pairs of acrocentric chromosomes ranging from medium-sized to mi-
nute, whereas Yong et al. (1971) reported two pairs of subacrocentric
1986
McBee et al.— Thailand Vespertilionid Karyology
105
ll U M M *kH )<«
0#' A5 1
XX 1
2a
• <*
T
XY
HI )) >1 >1 >11 II II
18 M H ••
2b
1.
XY
lir V VI n IX 1
KR w *• Ift IM 0» (
2c
XY
Fig. 2.— The standard karyotypes of: a) Tylonycteris robustula F (TK 21416), 2n = 32,
FN = 50; inset T. robustula M. (CM 88 1 52); b) Hesperoptenus tickelli M (TCWC 47481),
2n = 32, FN = 46; c) Hesperoptenus blanfordi F (CM 88114), 2n = 34, FN = 60.
and two pairs of acrocentric chromosomes. The X is large and acro-
centric and the Y is a small metacentric chromosome.
Hesperoptenus tickelli (2n = 32, FN = 46; Fig. 2b).— The autosomal
complement contains eight pairs of metacentric to submetacentric
chromosomes ranging from large to medium-sized. There also are seven
pairs of medium-sized to minute acrocentric chromosomes. The X is
a large subtelocentric and the Y is a small metacentric chromosome.
Hesperoptenus blanfordi (2n ^ 34, FN = 60; Fig. 2c).— The auto-
somal complement includes 1 3 pairs of metacentric to submetacentric
chromosomes gradually decreasing in size from large to small. There
106
Annals of Carnegie Museum
VOL. 55
is one pair of medium-sized subtelocentric chromosomes and two pairs
of medium-sized acrocentric chromosomes. The X is a medium-sized
subtelocentric chromosome, and the Y is a small subtelocentric chro-
mosome.
Subfamily Kerivoulinae
Kerivoula papillosa (2n = 38, FN = 52; Fig. 3a).— Metacentric to
submetacentric chromosomes include three large pairs, one medium-
sized pair and one small pair. There is one medium-sized pair and one
large pair of subtelocentric chromosomes, and 1 2 pairs of acrocentric
chromosomes grading from large to small. The sex chromosomes were
not identified in this species, but are probably the medium-sized pair
of submetacentric chromosomes. The fundamental number includes
the presumed sex chromosomes.
Subfamily Miniopterinae
Miniopterus schreibersi haradai (2n = 46, FN = 52; Fig. 3b).— The
autosomal complement includes two large and one medium-sized pairs
of metacentric chromosomes, and one medium-sized pair of subtelo-
centric chromosomes. There are 1 8 acrocentric pairs ranging from large
to small. One medium-sized acrocentric pair has a secondary constric-
tion near the centromere. The X is a medium-sized submetacentric
chromosome. Other reports of karyotypes from Miniopterus schreibersi
(Harada, 1973; Ando et al., 1977; Bickham and Hafner, 1978) iden-
tified the smallest pair of chromosomes as biarmed. This was not
apparent from our preparations; however, the medium-sized subtelo-
centric pair of chromosomes we observed is evidently unique to Thai
Miniopterus schreibersi and is not seen in European (Bickham and
Hafner, 1978) or Japanese (Harada, 1973) populations.
Subfamily Murininae
Murina leucogaster (2n = 44, FN = 50; Fig. 3c).— There are two
pairs of large metacentric, and two pairs of medium-sized to small
submetacentric autosomes. The autosomal complement is completed
by 1 7 acrocentric pairs ranging from large to small. The X is medium-
sized and submetacentric. This karyotype is similar to that of Murina
leucogaster from Atesu, Japan (Harada, 1973), but the third largest
Fig. 3.— The standard karyotypes of: a) Kerivoula papillosa F (CM 88164), 2n = 38,
FN =52 (FN includes sex chromosomes); b) Miniopterus schreibersi haradai F (CM
88157), 2n = 46, FN = 52; c) Murina leucogaster F (CM 88163), 2n = 44, FN = 50.
1986
McBee et al,— Thailand Vespertilionid Karyology
107
108
Annals of Carnegie Museum
VOL. 55
chromosome is considerably smaller in our material than in the Jap-
anese bats.
Harpiocephalus mordax (2n ^ 40, FN = 62).— Poor field preparation
of Harpiocephalus mordax made chromosome analysis difficult, but
counts of metaphase spreads consistently gave a diploid number of 40.
The chromosomal complement includes three large pairs, one medium-
sized pair and two small pairs of metacentric to submetacentric chro-
mosomes and five pairs of subtelocentric chromosomes. There are nine
pairs of acrocentric chromosomes gradually decreasing from large to
small. The single individual examined was a female, so sex chromo-
somes could not be identified but are probably the medium-sized pair
of submetacentric chromosomes. The fundamental number includes
the presumed sex chromosomes.
Discussion
This karyological study is consistent with earlier studies (Capanna
and Civitelli, 1 970; Baker and Patton, 1967; Pathak and Sharma, 1 969;
Bickham, 1979Z?), which have indicated extensive chromosomal vari-
ability between genera in vespertilionid bats. Within genera for which
karyotypic data have been obtained on more than one species, three
different patterns of chromosomal variability are apparent. Of the 1 5
genera (Table 1) for which more than one species have been karyotyped,
1 1 , including members of three subfamilies, can be characterized as
conservative genera. These are genera in which all species have the
same or nearly the same standard karyotype. Myotis (2n = 44, FN =
50, 52) and Eptesicus (2n = 50, FN = 48, with the exception of E.
capensis) are typical representatives of this pattern. Scotophilus, Ves~
pertilio, Barbestella, Lasiurus, Plecotus, Miniopterus, and Murina also
exhibit this pattern. Genera exhibiting the second pattern of variation
are interspecifically variable. Five genera currently fill this group (Table
1). Pipistrellus has nine different diploid numbers among the 1 5 species
that have been karyotyped. Nyctalus, with data from four species,
shows three different karyotypes and Tylonycteris and Hesperoptenus
each has two different karyotypes for two species. The third pattern is
one of intraspecific variability and is best documented in the genus
Rhogeessa (Table 1). Three species possess at least nine different karyo-
types. Rhogeessa parvula has a 2n = 44, FN = 50 karyotype, and R.
genowaysi has 2n = 42, FN = 50. R. tumida, however, has 2n = 30,
32a, 32b, 34, 42, 44, 52 and FN = 50, 52).
Pipistrellus exhibits such wide variability that even with karyotypes
for approximately one third of the recognized species no real patterns
of karyotypic relationships are evident within the genus. Several species
share the Myotis-MkQ 2n = 44, FN = 50 karyotype considered primitive
for the family (Bickham, \919a, 1 9 79Z?; Baker and Patton, 1967). Many
1986
McBee et al.— Thailand Vespertilionid Karyology
109
of the other karyotypes can be related to each other and to the primitive
Myotis-likQ karyotype on the basis of Robertsonian fusions and fissions.
Pipistrellus pulveratus has a karyotype of 2n = 32, FN 50 in two
individuals. The karyotypes appear to differ from the primitive Myotis-
like karyotype by six Robertsonian fission-fusion events.
The karyotype of P. mimus cannot be so directly derived from the
Myotis-\ik£ primitive. Manna and Talukdar (1965) reported a karyo-
type of 2n ^ 34 from southwestern India and Pathak and Sharma
(1969) found 2n = 38, FN = 48 for the same species in northeastern
India. We found 2n == 34, FN = 46 for four individuals from north-
western Thailand. This karyotype can be derived from the 2n == 38
karyotype through one centric fusion and the loss, possibly through
tandem fusion, of one small pair of acrocentric chromosomes. The
2n = 38 karyotype has 6 pairs of large metacentric to submetacentric
chromosomes and 12 pairs of acrocentric chromosomes. The Thai
karyotypes have an additional biarmed chromosome that is subtelo-
centric and three fewer pairs of acrocentrics. Neither karyotype can be
derived from the Myotis-likQ primitive karyotype without the loss or
tandem fusion of at least one pair of acrocentric chromosomes. These
data support the suggestion of Pathak and Sharma (1969) that trans-
locations other than Robertsonian fusions may play an important role
in chromosomal evolution in some groups of Pipistrellus, These authors
also suggest that the two karyotypic forms of P. mimus may represent
cryptic species. Cryptic species differentiated by karyotypes have been
discovered for the family among the many karyotypic forms of Rho-
geessa (Baker, 1984), so this explanation of karyotypic variation in P,
mimus is not unreasonable. An alternative explanation of the chro-
mosomal variability observed in P. mimus may involve intraspecific
variability. This phenomenon is very rare among vespertilionids but
is well documented within the genus Rhogeessa. In either case, P.
mimus merits comprehensive cytogenetic study throughout its range
in southern Asia.
Our karyotype of Tylonycteris robustula is slightly different from that
reported by Yong et al. (1971). The extremely small chromosomes they
consider biarmed are probably acrocentric. The T. robustula autosomal
karyotype readily can be derived from the Myotis-\ikQ primitive con-
dition by a series of six centric fusions. The X chromosome, however,
has experienced a pericentric inversion to an acrocentric condition and
the Y, either a pericentric inversion or the addition of a heterochromatic
short arm making it biarmed. An acrocentric or subtelocentric X chro-
mosome is rare among the Vespertilionidae occurring in only two
species of Pipistrellus, two species of Scotophilus, Scotoecus hindei,
two species of Lasiurus, two species of Plecotus, and two species of
Hesperoptenus (Table 1). Tylonycteris robustula also has diploid and
110
Annals of Carnegie Museum
VOL. 55
fundamental numbers in common with the two Plecotus. The karyo-
types are at least superficially the same, except both Plecotus have
acrocentric rather than biarmed Y chromosomes. Assumptions of ho-
mology on the basis of standard karyotypes must be made with caution,
however (Bickham and Baker, 1977; Baker et al., 1985; Haiduk and
Baker, 1982). The similarities between the plecotine genus, Plecotus,
and the vespertilionine genus Tylonycteris are likely the result of con-
vergence. Tate (1942) considered Tylonycteris and Philetor as derived
from an ancestor similar to the Pipistrellus jojfrei group. Neither Phile-
tor nor any members of the P. joffrei group have been karyotyped for
comparison, however.
The genus Hesperoptenus is poorly understood systematically. Four
species are currently recognized, two of which are known from only a
few specimens. The more common forms, H. tickelli and H. blanfordi,
are very different from one another morphologically. Tate (1942) com-
mented that if the genus was not polyphyletic it at least contained
strongly differentiated species. Hesperoptenus blanfordi and H. tickelli
also are karyologically distinct. Whereas the two species have similar
diploid numbers, H. blanfordi has one of the highest FNs reported for
the family, and H. tickelli has an FN of 52, among the most commonly
found in the family. To derive one karyotype from the other would
require one fission/fusion event and five pericentric inversions or het-
erochromatic additions. The H. tickelli karyotype can be derived from
the Myotis-likt primitive karyotype through four Robertsonian fusions
and the loss or tandem fusion of two pairs of acrocentrics. There also
has been a pericentric inversion changing the primitive biarmed X to
a derived subtelocentric configuration. The Y is a derived small meta-
centric chromosome, either through pericentric inversion or hetero-
chromatic addition. The H. blanfordi karyotype is more difficult to
derive from the Myotis-M^iQ primitive requiring at least five Robert-
sonian fusions and five pericentric inversions in the autosomal com-
plement. The X chromosome also is inverted to an acrocentric con-
dition. The more parsimonious scenario might consist of H. tickelli
diverging from the Myotis-\i\iQ ancestor with H. blanfordi being a high-
ly divergent offshoot of H. tickelli. Chromosomal data support the
conclusion that H. tickelli and H. blanfordi, at best, are only distantly
related. Ryan (1966) and Koopman (1971) thought Hesperoptenus was
closely related to Glauconycteris and Chalinolobus. Hill (1976) con-
sidered dental differences between the three genera to be too great and
considered Hesperoptenus more closely aligned with the genus Scoto-
philus. Hesperoptenus tickelli and some Scotophilus have FN and uni-
armed X chromosomes in common. H. tickelli has one fewer pair of
biarmed chromosomes and one fewer pair of acrocentric chromosomes
1986
McBee et al.— Thailand Vespertilionid Karyology
1
than Scotophilus, however. No species of Glauconycteris or Chalinolobus
have been karyotyped for comparison.
Our karyotype of Miniopterus schreibersi haradai agrees well with
previous reports. Miniopterinae is considered the most derived
subfamily of the Vespertilionidae, even being accorded familial status
by some authors (Mein and Tupinier, 1977) yet the karyotype found
throughout this subfamily differs from the primitive Myotis-li^Q karyo-
type by a single Robertsonian fission and two pericentric inversions
(Bickham and Hafner, 1978). Harada (1973) found in M. s. fuliginosus,
and we found in M. s. haradai, a medium-sized subtelocentric chro-
mosome apparently unique to Thai members of the species.
Members of the subfamily Murininae have been regarded as a spe-
cialized offshoot of an early Myotis-VikQ ancestor (Miller, 1907). The
subfamily contains two genera, Murina and Harpiocephalus. All mem-
bers of Murina karo typed so far have had a standard karyotype essen-
tially identical to the 2n = 44 MyotisAi\.Q primitive, agreeing with the
early divergence of Murininae from the vespertilionine line. Tate ( 1 94 1 )
considered the second genus, Harpiocephalus, as a very specialized
offshoot of the line leading to Murina, and Miller (1907) termed Har-
piocephalus as one of the most aberrant genera of the family. The
Harpiocephalus karyotype is derived from the primitive Myotis-\ik.Q
karyotype and the Murina karyotype by two possible pericentric in-
versions indicating that Harpiocephalus probably evolved from a Mu-
rina-likQ ancestor rather than diverging earlier from the line leading to
Murina.
The subfamily Kerivoulinae has been considered the least specialized
of the vespertilionid subfamilies being closely related to the “least
progressive” genera of the subfamily Vespertilioninae (Tate, 1941).
The karyotype of Kerivoula papillosa can be derived from the primitive
Myotis-\ik& karyotype through two Robertsonian fusions and the loss
or tandem fusion of one pair of acrocentric chromosomes.
Within the Vespertilionidae, Kerivoula shares a similar standard
karyotype with the Japanese Pipistrellus endoi and members of the
genus Vespertilio. In the past, the entire genus Pipistrellus has been
considered a part of Vespertilio, but Zima (1978) considers the dis-
tinctive karyotype of Vespertilio as justification for separate generic
status. Ando et al. (1980) suggest P. endoi may be a link between the
genus Vespertilio and its Pipistrellus -likQ ancestor. The subfamily Ker-
ivoulinae may have a similar link to its Pipistrellus -likQ ancestor here.
There are no other data to link the three, however, and postulation of
a common origin is only speculative.
Unquestionably, Robertsonian fusions and fissions have played a
major role in chromosomal evolution of the family Vespertilionidae
112
Annals of Carnegie Museum
VOL. 55
(Bickham, 1979^). However, the standard karyotypes reported here
indicate a greater importance for non-Robertsonian rearrangements
such as inversions and translocations as evolutionary mechanisms than
was previously thought (Bickham, 1979a; Bickham and Baker, 1977).
Pericentric inversions, tandem fusions, or heterochromatic additions
apparently have occurred in Tylonycteris robustula, Miniopterus schrei-
bersi haradai, Kerivoula papillosa, and Harpiocephalus mordax in their
evolution from the 2n = 44 ancestral karyotype. Hesperoptenus tickelli
and H. blanfordi may show an especially high incidence of pericentric
inversions, requiring up to five possible inversion events to be derived
from the ancestral karyotype. Examination of these standard karyo-
types emphasizes how poorly understood are vespertilionid karyolog-
ical relationships. Speculations about relationships based on standard
karyotypes can be misleading, however. G-band analysis has indicated
that constant genera such as Myotis are indeed as constant as was
assumed from standard karyotypes (Bickham, 1 919b). It also has point-
ed out extreme chromosomal differences where standard karyotypes
indicated homology (Baker et al., 1985). Chromosomal banding anal-
ysis should allow a more accurate assessment of the mechanisms of
chromosomal evolution seen in the family Vespertilionidae. G-banding
also will provide a means to test apparent homologies between groups
such as Kerivoula and Vespertilio. The extensive variability between,
and possibly within, species of Pipistrellus also will be much better
characterized by G-banding. Comparison of conserved and derived
chromosome sequences revealed by G-banding is imperative to an
understanding of systematic relationships among the Vespertilionidae.
Acknowledgments
We especially thank Craig S. Hood and Stephen L. Williams for their unflagging
assistance in the field. Keith Studholm, Dorothy Pumo, Bill Frucht, and Carleton J.
Phillips also provided assistance in the field. Dr. Niphan Ratanaworabhan acted as our
hostess and provided invaluable assistance in Thailand. Tweewat Polpakdee, Somechai
Kwanchareon, Supachai Sittilert, Monthida Sitathani, Puangtong Boonsong, and Precha
Luecha acted as our guides and translators, enthusiastically helping us in every possible
way. Gerard McKieman and Karen Muller aided in our literature search and Brian G.
Hanks, Priscilla K. Tucker, Michael S. Smolen and Thomas E. Lee provided aid in the
laboratory, and Teresa A. Heiner assisted in preparation of earlier manuscripts. Don E.
Wilson and Rodney L. Honeycutt provided constructive reviews that enhanced the final
manuscript. McBee received partial funding from a Tom Slick Graduate Research Fel-
lowship at Texas A&M University. This study was supported in part by NSF Grant
PCM-8202794, NIH Grant AIO 4242, and the Office of University Research Services,
Texas A&M University.
Literature Cited
Ando, K., T. Tagawa, and T. A. Uchida. 1977. Considerations of karyotypic evo-
lution within Vespertilionidae. Experientia, 33:877-879.
1986
McBee et al.~ Thailand Vespertilionid Karyology
113
, 1980. The C-banding pattern of 6 Japanese species of Vespertilionid bats.
Experientia, 36:65 3--6 54.
Baker, R. J. 1970. Kaiyotypic trends in bats. Pp, 65-96, in Biology of bats (W. A.
Wimsatt, ed.), Academic Press, New York, lixxii + 1-406.
— — . 1984. A sympatric cryptic species of mammal: a new species of Rhogeessa
(Chiroptera:Vespertilionidae). Syst. Zool., 33:178-183.
Baker, R. J., and J. W. Bickham. 1980. Karyotypic evolution in bats: evidence of
extensive and conservative chromosomal evolution in closely related taxa. Syst.
Zool., 29:239-253.
Baker, R. J., and J. T. Mascarello. 1969. Chromosomes of some vesperilionid bats
of the genera Lasiurus and Plecotus. Southwestern Nat., 14:249-251.
Baker, R. J., and J. L. Patton. 1967. Karyotypes and karyotypic variation of North
American vespertilionid bats. J. Mamm., 48:270-286.
Baker, R. J., J. W. Bickham, and M. L. Arnold. 1985. Speciation in the Rhogeessa
tumida-parvula complex (Chiroptera: Vespertilionidae). Evolution, 39:233-243.
Baker, R. J., B. L. Davis, R. G. Jordan, and A. Binous. 1974. Karyotypic and
morphometric studies of Tunisian mammals: bats. Mammalia, 38:695-710.
Baker, R. J., M. W. Haiduk, L. W. Robbins, A. Cadena, and B. F. Koop. 1982.
Chromosomal studies of South American bats and their systematic implications.
Spec. Publ., Pymatuning Lab. EcoL, 6:303-327.
Bickham, J. W. \919a. Chromosomal variation and evolutionary relationships of
vespertilionid bats. J. Mamm., 60:350-363.
. \919b. Banded karyotypes of 11 species of American bats (Genus Myotis).
Cytologia, 44:789-797.
Bickham, J. W., and R. J. Baker. 1977. Implications of chromosomal variation in
Rhogeessa (Chiroptera:Vespertilionidae). J. Mamm,, 58:448-453.
Bickham, J. W., and J. C. Hafner. 1978. A chromosomal study of three species of
vespertilionid bats from Yugoslavia. Genetica, 48:1-3.
Bovey, R. 1949. Les chromosomes des Chiropteres et des Insectivores. Rev. Suisse
Zool., 56:371-460.
Capanna, E. 1968. Some considerations on the evolution of the karyotype of Micro-
chiroptera. Experientia, 24:624-626.
Capanna, E., and M. V. Civitelli. 1970. Chromosomal mechanism in the evolution
of Chiropteran karyotype. Chromosomal tables of Chiroptera. Caryologia, 23:79-
111.
Capanna, E., L. Conti, and G. de Renzis. 1968. I cromosomi di Barbastella barbas-
tellus. Caryologia, 21:137-145.
Dulic, B. 1981. Chromosomes of three species of Indian microchiroptera. Myotis, 1 8-
19:76-82.
Dulic, B., B. Soldatovic, and D. Rimsa. 1967. La formule chromosomique de la
Noctule, Nyctalus noctula Schreiber (Mammalia, Chiroptera). Experientia, 23:1-4.
Engstrom, M. D., and D. E. Wilson. 1981. Systematics of Antrozous dubiaquercus
(Chiroptera:Vespertilionidae), with comments on the status of Bauerus Van Gelder.
Ann. Carnegie Mus., 50:371-383.
Fedyk, a., and D. Fedyk. 1970. Karyotypes of some species of vespertilionid bats
from Poland. Acta TherioL, 20:295-302.
Fedyk, S., and A. L. Ruprecht. 1976. Karyotypes of Pipistrellus pipistrellus (Schreiber,
1774) and Pipistrellus nathusii (Keyserling and Blasius, 1889) (Chiroptera: Vesper-
tilionidae). Caryologia, 29:283-289.
Genoways, H. H., and R. J. Baker. 1975. A new species of Eptesicus from Guadeloupe,
Lesser Antilles (Chiroptera:Vespertilionidae). Occas. Papers Mus., Texas Tech Univ.,
34:1-7.
Haiduk, M. W., and R. J. Baker. 1982. Cladistic analyis of G-banded chromosomes
of nectar-feeding bats (Glossophaginae:Phyllostomidae). Syst. Zool., 31:252-265.
114
Annals of Carnegie Museum
VOL. 55
Harada, M. 1 973. Chromosomes of nine Chiropteran species in Japan (Chiroptera). La
Kromosomo, 91:2885-2895.
Harada, M., and T. Kobayashi. 1 980. Studies on the small mammal fauna of Sabah,
East Malaysia II. Karyological analysis of some Sabahan mammals (Primates, Ro-
dentia, Chiroptera). Contrib. Biol. Lab., Kyoto Univ., 26:83-95.
Harada, M., and T. A. Uchida. 1982. Karyotype of a rare species, Myotis pruinosus,
involving pericentric inversion and duplicated translocation. Cytologia, 47:539-543.
Harada, M., and T. H. Yoshida. 1978. Karyological studies of four Japanese Myotis
bats (Chiroptera, Mammalia). Chromosoma (Berl.). 65:283-291.
Harada, M., T. A. Uchida, T. H. Yosida, and S. Takada. 1 9^2a. Karyological studies
of two Japanese noctule bats (Chiroptera). Caryologia, 35:1-9.
Harada, M., M. Minezawa, S. Takada, S. Yenbutra, P. Nunpakdee, and S. Ohtani.
1982Z?. Karyological analysis of 12 species of bats from Thailand. Caryologia, 35:
269-278.
Heller, K.-G., and M. Volleth. 1 984, Taxonomic position of Pipistrellus societatis''
Hill, 1972 and the karyological characteristics of the genus Eptesicus (Chiroptera:
Vespertilionidae). Z. Zool. Syst. Evolut.-forsch., 22:65-77.
Hill, J. E. 1976. Bats referred to Hesperoptenus Peters (1869) (Chiroptera: Vespertil-
ionidae) with the description of a new subgenus. Bull, British Mus. (Nat. Hist.) Zool.,
30:1-28.
Honeycutt, R. L., R. J. Baker, and H. H. Genoways. 1980. Results of the Alcoa
Foundation-Suriname Expeditions. III. Chromosomal data for bats (Mammalia:
Chiroptera) from Suriname. Ann. Carnegie Mus., 49:237-250.
Iliopoulou-Georgudaki, j. and E. B. Giagia. 1984. Distribution notes on Myotis
myotis (Borkhausen, 1797) (Chiroptera: Vespertilionidae) from Greece, including the
karyotype of specimens from Lesvos Island. Saugertier. Mitteil., 31:135-139.
Koopman, K. 1971. Taxonomic notes on Chalinolobus and Glauconycteris (Chiroptera,
Vespertilionidae). Amer, Mus. Novit., 2451:1-10.
—. 1984. Bats. Pp. 145-186, in Orders and families of Recent mammals of the
world (S. Anderson and J. K. Jones, Jr,, eds.), John Wiley and Sons, New York,
686 pp.
Lee, M. R., and F. F. B. Elder. 1980. Yeast stimulation of bone marrow mitosis for
cytogenic investigations. Cytogenet. Cell Genet., 26:36-40.
Lekagul, B,, and j. a. McNeely. 1977. Mammals of Thailand. Assoc. Conservation
of Wildlife. Kurusapha Ladprao Press, Bangkok, 758 pp.
Manfreddi Romanini, M, G., C. Pellicciari, F. Bolchi, and E. Capanna. 1975.
Donnees nouvelles sur le contenu en ADN des noyaux postkinetiques chez les
chiroptdres. Mammalia, 39:675-683.
Manna, G. K., and M. Talukdar. 1965. Somatic chromosome number in twenty
species of mammals from India. Mamm. Chromo. Newsletter, 17:77-78.
Matthey, R., and R. Bovey. 1948. La formule chromosomique chez cinq especies de
chiroptero. Experientia, 4:26.
Mein, P., and Y. Tupinier, 1 977. Formule dentaire et position systematique du Mini-
optere (Mammalia, Chiroptera). Mammalia, 41:207-21 1.
Miller, G. S. 1907. The families and genera of bats. Bull. U.S. Natl, Mus., 57:1-282.
Nagorsen, D. W., j. L. Eger, and R. L. Peterson. 1976. Somatic chromosomes of
three African species of bats (Chiroptera) Scotoecus hindei, Tadarida aegyptiaca,
and Tadarida bemmeleni. Mamm. Chromo. Newsletter, 17:9-14.
Obara, Y., and K, Saitoh, 1977. Chromosome studies in Japanese vespertilionid bats.
IV. Karyotypes and C-banding pattern of Vespertilio orientalis. Japan J. Genetics,
52:159-161.
Obara, Y., T. Tomiyasu, and K. Saitoh. 1 916a. Chromosome studies in the Japanese
vespertilionid bats, 1. Karyotypic variations in Myotis macrodactylus Temminck.
Japan J. Genet, 51:201-206.
1986
McBee et al.-~ Thailand Vespertilionid Karyology
115
— . 1 916b. Chromosome studies in the Japanese vespertilionid bats: IL G-banding
pattern of Pipistrellus abramus Temminck. Proc. Japan. Acad., 52:383-386.
— — — ", 1976c, Chromosome studies in the Japanese vespertilionid bats. III. Prelimi-
nary observation of C-bands in the chromosomes of Pipistrellus abramus. Sci. Rep.
Hirosaki Univ., 23:39-42.
Osborne, J. L. 1965. Karyotypes of selected bats (Order Chiroptera). Unpublished
M.S. thesis, Univ. Arizona, Tucson,
Park, S. R., and P. O. Won. 1978. Chromosomes of the Korean bats. J. Mammal.
Soc. Japan, 7:199-203.
Pathak, S., and T. Sharma. 1969. Chromosomes of five species of Indian vespertil-
ionid bats. Caryologia, 22:35-46.
Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus
(Rodentia:Heteromyidae). J. Mamm., 48:27-37.
Peterson, R, L., and D. W. Nagorsen. 1975. Chromosomes of fifteen species of bats
(Chiroptera) from Kenya and Rhodesia. Life Sci. Occas. Papers, Royal Ontario Mus.,
27:1-14.
Radjhabli, S. L, N. N. Vorontsov, and V. T, Volotbuev. 1969. Chromosomes of
three species of the Myotis (Chiroptera- Vespertilionidae) Proc. 2nd Conf Union of
Mammalogy (Moscow, 1969), pp, 12-13,
— . 1970. The karyological analysis of three Myotis species (Chiroptera: Vesper-
tilionidae). Citologia (Moscow), 12:767-773.
Reduker, D. W., T. L. Yates, and I. F. Greenbaum. 1983. Evolutionary affinities
among southwestern long-eared Myotis (Chiroptera: Vespertilionidae). J. Mamm.,
64:666-677.
Ryan, A. M, 1966. A new and some imperfectly known Australian Chalinolobus and
the taxonomic status of African Glauconycteris. J. Mamm., 47:86-91.
Schlitter, D. a., I. L. Rautenbach, and D. A. Wolhuter. 1980, Karyotypes and
morphometries of two species of Scotophilus in South Africa (Mammalia: Vesper-
tilionidae), Ann. Transvaal Museum, 32:231-240.
Sharma, G. P., S. M. Handa, and K. K. Mehta, 1974. Chromosome analysis in a
bat, Scotophilus heathi heathi (Horsfield). Mamm. Chromo. Newletter, 15:12.
Stock, A. D. 1983. Chromosomal homologies and phylogenetic relationships of the
vespertilionid bat genera Euderma, Idionycteris, and Plecotus. Cytogenet. Cell Ge-
net, 35:136-140.
Takayama, S. 1959. The chromosomes of a bat, Pipistrellus tratalitus abramus. Japan
J. Genet, 34:107-110.
Tate, G. H. H. 1941. Results of the Archbold Expeditions. No. 40. Notes on vesper-
tilionid bats. Bull. Amer. Mus. Nat. Hist., 78:567-597.
. 1942. Results of the Archbold Expeditions. No. 47, Review of the vesper-
tilionine bats, with special attention to genera and species of the Archbold Collec-
tions. Bull. Amer. Mus. Nat. Hist., 80:221-297.
Tsuchiya, K., M. Harada, and T. H. Yosida. 1972. Karyotypes of four species of
bats collected in Japan. Ann. Rep. Nat. Institute Genet. Japan, 22:50-51.
Vorontsov, N. N. 1969. The comparative karyology of the vespertilionid bats, Ves-
pertilionidae (Chiroptera). Pp. 16-21, in The mammals (N. N. Vorontsov, ed.) 2nd
All Union Mammal. Conference, Moscow.
Williams, D. F. 1978. Taxonomic and karyologic comments on small brown bats.
Genus Eptesicus, from South America, Ann. Carnegie Mus. 47:361-383.
Williams, D. F., and M. A. Mares. 1 978. Karyological affinities of the South American
big-eared bat, Histiotus montanus (Chiroptera, Vespertilionidae). J. Mamm., 59:
844-846.
Williams, D. F., J. D. Druecker, and H. L, Black. 1970. The karyotype of Euderma
maculatum and comments on the evolution of the Plecotine bats. J. Mamm., 51:
602-606.
116
Annals of Carnegie Museum
VOL. 55
Yong, H. S., S. S. Dhaliwal, and K. L, Teh. 1971. Somatic chromosomes of the flat-
headed bats {Tylonycteris spp.). Experientia, 27; 1353-1 355.
ZiMA, J. 1978. Chromosome characteristics of Vespertilionidae from Czeckoslovakia.
Acta. Sci. Nat Bmo, 12:1-38.
. 1982, Chromosomal homology in the complements of bats of the family Ves-
pertilionidae. II. G-banded karyotypes of some Myotis, Eptesicus and Pipistrellus
species. Folia Zook, 31:31--36.
. 1984. Chromosomes of certain small mammals from southern Bohemia and
the Sumava Mts. (C.S.S.R.). Folia Zook, 33:133-141.
ISSN 0097-4463
ANNALS
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE * PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 23 MAY 1986 ARTICLE 6
RESULTS OF THE CARNEGIE MUSEUM OF NATURAL
HISTORY EXPEDITIONS TO BELIZE. L
SYSTEMATIC STATUS AND GEOGRAPHIC
DISTRIBUTION OF SIBON NEILLI
(REPTILIA, SERPENTES)
C. J. McCoy
Curator, Section of Amphibians and Reptiles
Abstract
The nominal species Sibon neilli Henderson, Hoevers and Wilson (type-locality, “vi-
cinity of Belize City, Belize”) is shown to be a southern subspecies of Sibon sanniola
Cope (type-locality, “Chichen Itza, Yucatan, Mexico”), confirming the arrangement pro-
posed by Kofron (1985). Sibon sanniola neilli differs from Sibon s. sanniola in having
a banded, rather than spotted, color pattern, and higher ventral and subcaudal scale
counts.
Introduction
Sibon neilli Henderson, Hoevers, and Wilson (1977) is an enigmatic
member of the snake fauna of Caribbean Central America. The species
was described from a unique type-specimen, collected in “the vicinity
of Belize City, Belize District, Belize.” Henderson et al. (1977) rec-
ognized that Sibon neilli is very similar to 5. sanniola, a species en-
demic to the northern part of the Yucatan Peninsula, but diagnosed S.
neilli on the basis of higher ventral and subcaudal scale counts, a
different pattern of supralabial, postocular, and temporal scale contacts,
and a banded color pattern. Kofron (1985),
Submitted 24 September 1985.
117
118
Annals of Carnegie Museum
VOL. 55
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1986
McCoy— Status of Sibon neilli
119
tional specimens of S. neilli and without comment, considered S. neilli
a subspecies of Sibon sanniola.
During field work in Belize in 1984 we collected a specimen of Sibon
at Chaa Creek, Cayo District, that has the characteristic banded dorsal
pattern of S. neilli. In addition, there is a specimen in Carnegie Museum
of Natural History from El Peten, Guatemala, that also fits the de-
scription of Sibon neilli. I have compared these specimens with the
holotype and previously only known specimen of S. neilli, with all
other Sibon of this group available from Belize, and with an extensive
series of Sibon sanniola from Yucatan and Quintana Roo, Mexico (see
Specimens Examined).
Results
Ventral and subcaudal scale — Data summaries for ventral
and subcaudal scale counts for Sibon sanniola and Sibon neilli are
presented in Table 1. Both the ventral and subcaudal counts for the
holotype of Sibon neilli exceed the known ranges for these counts in
S. sanniola, as pointed out by Henderson et al. (1977). The range of
ventral counts for males in the “southern” population overlaps the
range for the “northern” population, as do the ranges of ventral counts
for females in the two populations. Subcaudal counts in males barely
overlap in the two populations, and overlap slightly in females. Al-
though the sample size for the southern population is small, it appears
that the differences in segmental counts are consistent. A specimen
from “British Honduras” (FMNH 4247, male) has 153 ventrals and
67 caudals, both counts within the ranges for the northern population.
SupralabiaVpostocular-temporal contact.— The holotype of Sibon
neilli has 9-9 supralabials, with the 4th, 5th, and 6th entering the orbit.
The 7th supralabial is in contact with the lower postocular and the
primary temporal. Henderson et al. (1977) regarded the 7th supralabial-
postocular-temporal contact as a distinctive character of S. neilli, stat-
ing that it “occurs occasionally in sanniola, but in no other species
of Sibon."" In reality, this is the contact pattern that is most common
in Sibon sanniola. In a series of 51 S. sanniola from Yucatan and
Quintana Roo the 7th supralabial contacts only the lower postocular
and primary temporal 88% of the time (right and left sides of the head
scored separately). In the remaining 12% the upper tip of the 7th
supralabial, posterior angle of the lower postocular, and anterior tips
of the primary and secondary temporals make a single point contact.
In specimens with the more common pattern, the anterior tip of the
primary temporal makes a broad contact with the postocular, pre-
venting contact of the 7th supralabial with the secondary temporal.
120
Annals of Carnegie Museum
VOL. 55
Fig. \.~Sibon sanniola neilli (CM 105981), Chaa Creek, Cayo Dist., Belize; from a
Kodachrome photo of the living specimen by Paul S, Freed.
Color pattern. — The holotype of Sibon neilli has a series of 34 dark
dorsal crossbands on a lighter ground color, faded almost to white in
the specimen. The widest crossbands are IVi scales long at the middorsal
line, and taper to about 1 Vi scales at the lowermost dorsal scale rows.
The edges of the bands are jagged, not straight. The bands extend
ventrally only to the outer ends of the ventral scales. The center of the
belly is marked with a series of roughly paired, longitudinal dark “dash-
es,” each usually two ventral scales long. There are about 21 dark
crossbands on the tail, which become increasingly crowded toward the
tail tip. The dark nuchal band extends ventrally to the edges of the
ventrals, and anteriorly as a middorsal dark bar that reaches the frontal
scale. The holotype is much faded, resulting in enhanced contrast be-
tween the dorsal dark bands and the interspaces (Henderson et al.,
1977:fig. 1).
The specimen from Chaa Creek, Belize (CM 105981), has 42 dorsal
body bands that taper ventrally from IVi to IV2 scales wide, and 22
bands on the tail. The nuchal band has a broad anterior extension that
reaches the parietals, and the top of the head is lighter brown. The
sides of the head are much lighter. The belly is lightly marked with
indistinct brown smudges. The only significant difference between the
1986
McCoy— Status of Sibon neilli
121
pattern of this specimen and that of the holotype is the amount of
contrast between the dorsal bands and the interspaces. In life, the dorsal
bands of CM 105981 were medium brown, and the interspaces dark
tan, providing minimal contrast (Fig. 1).
The specimen from Las Cahas, Guatemala (CM 58282), has 38 barely
discernible dark bands on the body, and a banded tail. The bands are
only slightly darker than the medium brown ground color. The head
and nuchal pattern are typical, and the belly is moderately well-marked
with longitudinal dashes.
The typical color pattern of Sibon sanniola is a series of dark, light-
edged middorsal spots, frequently fused to form an irregular “zig-zag”
line, on a light brown to ash gray background. The lateral and ventro-
lateral spot series may either persist, be reduced, or fade completely.
The tail is spotted along the middorsal line. The belly typically is
marked with pairs of dark longitudinal dashes. The nuchal band, with
anterior extension onto the head, is as described for Sibon neilli.
A juvenile specimen from Xunantunich, Belize (MCZ 56994, 180
mm total length), has a pattern of about 47 middorsal spots (many
fused), and two series of lateral dark spots on each side. A very similar
pattern occurs in juvenile S. sanniola (CM 49739, 140 mm; CM 49754,
156 mm). This suggests that the banded adult pattern of S. neilli may
result from ontogenetic fusion of the dorsal, lateral, and ventrolateral
spot series, which remain discrete in 5. sanniola.
Conclusions
In size, habitus, and most details of scutellation Sibon neilli and
Sibon sanniola are identical. The supposedly diagnostic arrangement
of supralabial, postocular, and temporal scales of S. neilli actually is
consistent with the pattern usually found in S. sanniola. Only the color
pattern and numbers of ventral and subcaudal scales are distinctive
characters of the .S', neilli population. Although Kofron (1983) shows
the range of S. sanniola (including S. neilli) as being continuous from
northern Yucatan and Quintana Roo southward into Belize and El
Peten, no specimens are available from the critical areas where inter-
gradation would be expected (Lee, 1980 and personal communication).
The southernmost precise locality for Sibon sanniola is Felipe Carrillo
Puerto, Quintana Roo (Peters, 1953), although FMNH 4247 from
“British Honduras” has both scale counts and color pattern typical of
S. sanniola. The range of S. neilli extends from coastal central Belize
(Belize City), southwestward into El Peten, Guatemala. Despite the
apparent hiatus, I assume that the range of the species is continuous,
as there is no ecological discontinuity between southern Quintana Roo
and central Belize, and I interpret the morphological differences be-
122
Annals of Carnegie Museum
VOL. 55
tween the southern and northern populations as clinal variation within
a single species.
Clinal variation is common in colubrid snake species that have ex-
tensive latitudinal ranges on the Yucatan Peninsula. Such clinal vari-
ation typically involves either the color pattern or segmental counts,
or both. Examples are Conophis lineatus (Wellman, 1963), Leptodeira
frenata (Duellman, 1958), Ninia sebae (Schmidt and Rand, 1957), and
Leptophis mexicanus (Oliver, 1 948), to cite a few of many species that
illustrate this variational pattern. I conclude that Kofron (1985) was
correct in allocating the name Sibon neilli to the southern population
of Sibon sanniola that is characterized by a banded color pattern and
higher ventral and subcaudal scale counts.
Specimens Examined
Sibon sanniola neilli. — Belize: Belize Dist., vicinity of Belize City (MPM 8929, ho-
lotype). Cayo Dist., Chaa Creek, 5 mi. S San Ignacio (CM 105981); vicinity of Augustine
(MPM 8208); Xunantunich (MCZ 56994). Guatemala: El Peten, Las Canas (Munici-
pality San Luis) (CM 58282). Total 5 specimens.
Sibon sanniola sanniola.— Belize: “British Honduras” (FMNH 4247). Mexico: Quin-
tana Roo, Pueblo Nuevo X-Can (CM 45778-45785, 46844-46845, 46881-46883, 49056,
49062, 49136, 49154, 49159, 49163). Yucatan, Chichen Itza (FMNH 20609, 20613,
26988, 36257-36258, 36268, 36272, 36276, 36285, 36287, 36289, 36296); Kantunil
(FMNH 36264, 36286, 36288, 36270); Libre Union (FMNH 36259, 36262, 36266,
36273, 36278, 36280-36283, 36290, 36294-36295); Piste (CM 46955-46958, 47004,
47142-47148, 49734-49741, 49742-49748, 49749-49756); Progreso (FMNH 40734-
40735); Yokdzonot (FMNH 36261, 36263, 36265, 36267, 36269, 36271, 36274-36275,
36277, 36279, 36284, 36290-36293, 36297). Total 101 specimens.
Acknowledgments
Field work in Belize was supported by a grant from the O’Neil Museum Trust, Carnegie
Museum of Natural History. For assistance in the field I thank D. Scott Wood, Robert
C. Leberman, Paul and Mara Freed, and a column of anonymous army ants that enabled
us to capture the Chaa Creek Sibon. I am indebted to the Flemings of Chaa Creek for
their hospitality, to Ray E. Ashton, Jr., of International Expeditions, Inc., for logistical
aid, and to Dora Weyer for steadfast support of our work in Belize. Collecting permits
were granted by Mr. O. Rosado, Department of Forestry, Belize Ministry of Natural
Resources. For loans of specimens I thank R. F. Inger and Hymen Marx, Field Museum
of Natural History (FMNH), Robert W. Henderson, Milwaukee Public Museum (MPM),
and Pere Alberch, Museum of Comparative Zoology, Harvard University (MCZ). I also
thank Julian C. Lee for information on localities, and Ellen J. Censky for technical help.
Literature Cited
Duellman, W. E. 1958. A monographic study of the colubrid snake genus Leptodeira.
Bull. Amer. Mus. Nat. Hist., 114:1-152.
Henderson, R. W., L. G. Hoevers, and L. D. Wilson. 1977. A new species of Sibon
(Reptilia, Serpentes, Colubridae) from Belize, Central America. J. HerpetoL, 11:
77-79.
Kofron, C. P. 1983. Female reproductive cycle of the Neotropical snail-eating snake
Sibon sanniola in northern Yucatan, Mexico. Copeia, 1983:963-969.
1986
McCoy— Status of Sibon neilli
123
1985. Systematics of the Neotropical gastropod-eating snake genera, Tropi-
dodipsas and Sibon. ]. HerpetoL, 19:84-92.
Lee, J. C. 1980. An ecogeographic analysis of the herpetofauna of the Yucatan Pen-
insula. Univ. Kansas Mus. Nat. Hist., Misc. Publ., 67:1-75.
Oliver, J. A. 1948. The relationships and zoogeography of the genus Thalerophis
Oliver. Bull. Amer. Mus. Nat. Hist., 92:157-280.
Peters, J. A. 1953. Snakes and lizards from Quintana Roo, Mexico. Lloydia, 16:227-
232.
Schmidt, K, P., and A. S. Rand. 1957. Geographic variation in the Central American
colubrine snake, Ninia sebae. Fieldiana: ZooL, 39:73-84.
Wellman, J. 1963. A revision of snakes of the genus Conophis (Family Colubridae,
from Middle America). Univ. Kansas Publ., Mus. Nat. Hist., 15:251-295.
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VOLUME 55 23 MAY 1986 ARTICLE 7
KARYOTYPES OF ELEVEN SPECIES OF MOLOSSID
BATS FROM AFRICA (MAMMALIA: CHIROPTERA)
Steven A. Smith ‘
John W. Bickham^
Duane A. Schlitter
Curator, Section of Mammals
Abstract
Standard karyotypic data are reported for 1 1 species of molossid bats collected from
Somalia and Cameroon, Africa. Chromosomal data are reported for the first time for
Chaerephon ansorgei, C. aloysiisabaudiae, Mops midas, M. spurrelli, M. thersites, M.
brachypterus, M. petersoni, M. demonstrator, and M. nanulus (all were formerly mem-
bers of the genus Tadarida). Karyotypes for two of the species we examined have been
reported previously. Although our data corroborate the karyotype of C. pumila described
by Dulic and Mutere (1973), our karyotypic analysis of M. condylurus differs substantially
from that presented by these authors. In addition to these data, we provide a summary
of the available karyotypic data for molossid bats studied to date.
Introduction
The Molossidae is a group of insectivorous, swift-flying bats that live
in tropical and temperate parts of the world. More than half of the 9 1
or so extant species have been regarded as members of the genus Tad-
arida (Corbet and Hill, 1980); the remaining species are spread among
* Address: Department of Wildlife and Fisheries Sciences, Texas A&M University, Col-
lege Station, Texas, 77843.
Submitted 10 June 1985.
125
126
Annals of Carnegie Museum
VOL. 55
Table Summary of karyotype morphology for 11 species of African molossid bats.
Letter designations are: M—metacentric, SM—submetacentric, ST-subtelocentric, A —
acrocentric.
Species
Large
M
Medium
M
Medium
ST
Small
ST
Medium-
small
A
X
Y
FN
Chaerephon ansorgei
Chaerephon aloysiisa-
1
3
4
2
13
ST
A
66
baudiae
1
3
4
2
13
SM
A
66
Chaerephon pumila
1
2
3
0
17
SM
A
58
Mops midas
1
3
4
2
13
SM
A
66
Mops condylurus
1
3
4
2
13
SM
A
66
Mops spurrelli
1
3
4
1
14
SM
_
64
Mops thersites
1
3
3
1
15
SM
ST
62
Mops brachypterus
1
3
0
0
19
SM
_
54
Mops petersoni
1
3
0
0
19
SM
A
54
Mops demonstrator
1
2
1
0
19
SM
A
54
Mops nanulus
1
2
1
0
19
SM
A
54
1 1 Other genera. Until recently, taxonomic assignments and systematic
relationships among the family members had not been examined world-
wide. Freeman (1981), based upon a phenetic study of morphological
traits, provided the first major review of the family. She restricted the
genus Tadarida to include only nine species and assigned the remainder
to Chaerephon, Mops, Mormopterus and Nyctinomops (all former sub-
genera of Tadarida).
Karyotypic data for the Molossidae are available for 25 species, only
six of which are inhabitants of the Old World. In this paper we analyze
the karyotypes of 1 1 African molossid species belonging to the genera
Chaerephon and Mops, and summarize the chromosomal data (Tables
1 and 2) now available for 35 species representing 10 of the 12 genera
recognized by Freeman (1981).
Methods and Materials
Standard karyotypes were obtained in the field from bone marrow preparations (Patton,
1967) of live caught animals. A minimum of five representative chromosome spreads
were examined from each individual to determine diploid (2n) and fundamental numbers
(FN). Photomicrographic enlargements of suitable spreads were used in the final analyses.
Chromosomes were divided into large and medium-sized metacentric, medium and
small subtelocentric, and medium to small acrocentric morphological classes. Deter-
mination of centromere position was difficult because differential contraction of nearly
acrocentric chromosomes caused variation in the number of countable arms. We follow
Warner et al. (1974) in being conservative in the determination of biarmed versus
acrocentric conditions and reiterate their warning that FN values are somewhat arbitrary
and subjective.
Taxonomic designations follow Honacki et al., 1982 (see Freeman, 1981).
1986
Smith et al.— Molossid Karyotypes
127
Xir XX XK XX M MtK
Ad
do dd dl dh All
XY
fkA M
A
X* tn A XX M lA M M M
A.-.
V V
do ftD ot If« An
A T
B
^ 0 XX M /M M M A* AM
l/t
X Y
c
W X« xr> ft firt lifi
An
OH nn n/% rfikM aa «a
X Y
D
Fig. L— Representative karyotypes of A) Chaerephon ansorgei from Cameroun, B) Chae-
rephon aloysiisabaudiae from Cameroun, C) Mops midas from Somalia, and D) Mops
condylurus from Somalia.
Species Accounts
A summary of the chromosomal morphology for the species ex-
amined in this study is presented in Table 1. Representative karyotypes
are presented in Figs, 1-3,
All species examined in this study were characterized by a diploid
128
Annals of Carnegie Museum
VOL. 55
number of 48. Fundamental numbers ranged from 54 to 66. All of
these species had, minimally, four biarmed autosomal elements in-
cluding one large pair of metacentric and at least one medium-sized
pair of metacentric |Chromosomes. In all cases the large metacentric
pair was twice the size of the next largest chromosome pair. The X
chromosomes were medium-sized and submetacentric or subtelocen-
tric in all species; the Y chromosome was medium-sized and acrocentric
in all but one species.
A brief description of the karyotypes for each species reported herein
follows.
Chaerephon ansorgei (Thomas, 1913)
Fig. lA, 2n = 48; FN = 66; U
The autosomal complement includes one pair of large metacentric,
three pairs of medium metacentric, four pairs of medium subtelocen-
tric, and 1 3 medium to small acrocentric chromosomes. The X chro-
mosome is medium-sized and subtelocentric, and the Y is medium-
sized and acrocentric.
Chaerephon aloysiisabaudiae (Festa, 1907)
Fig. IB, 2n = 48; FN = 66; IS
The karyotype of this species is identical to C ansorgei except the
X chromosome in C aloysiisabaudiae appears submetacentric rather
than subtelocentric.
Chaerephon pumila (Cretzschmar, 1826)
2n = 48; FN = 58; SSS, 6$9
The karyotype of our specimens is identical to that reported for this
species by Dulic and Mutere (1973).
Mops midas (Sundevall, 1843)
Fig. 1C, 2n = 48; FN = 66; 3SS, 399
This species is karyotypically identical to the above-mentioned
Chaerephon species and shares the submetacentric condition of the X
chromosome observed in C aloysiisabaudiae.
Mops condylurus (A. Smith, 1833)
Fig. ID, 2n = 48; FN = 66; 4SS, 599
The karyotype of M. condylurus is identical to both M. midas and
C aloysiisabaudiae.
1986
Smith et al.— Molossid Karyotypes
129
Mops spurrelU (Dollman, 1911)
Fig. 2A, 2n - 48; FN - 64; 355
The chromosomal complement from female specimens of M. spur-
relU differ from M. condyiurus in the absence of one less small sub-
telocentric pair and the presence of an extra acrocentric pair. The X
chromosome is submetacentric.
Mops thersites (Thomas, 1903)
Fig. 2B, 2n = 48; FN ^ 62; 355, 499
The autosomes are nearly identical to M. spurrelU but there is one
less medium-sized subtelocentric and one additional acrocentric pair
present. The X chromosome is submetacentric but the Y appears to
be subtelocentric instead of the more commonly observed acrocentric
condition.
Mops bmchypterus (Peters, 1852)
Fig. 3 A, 2n - 48; FN - 54; 19
The autosomes of the female specimen examined consist of one large
pair of metacentric, three pairs of medium-sized metacentric and 1 9
medium to small acrocentric pairs. Although morphologically similar
to M. thersites, it differs chromosomally by lacking subtelocentric pairs
and having additional acrocentric pairs. The X chromosome presum-
ably is submetacentric.
Mops petersoni (El Rayah, 1981)
Fig. 3B, 2n - 48; FN - 54; 15, 19
In addition to being morphologically similar, this species is karyo-
typically identical to M. brachypterus.
Mops nanuius J. A. Allen, 1917
Fig. 3C, 2n - 48; FN - 54; 255, 299
M. nanuius differs from M. petersoni and M. brachypterus by having
one less medium-sized metacentric pair and the presence of a medium-
sized subtelocentric pair. The sex pair is identical to M. petersoni.
Mops demonstrator 1903)
Fig. 3D, 2n = 48; FN - 54; 15
The karyotype of this species is identical to M. nanuius.
Discussion
Until Freeman’s (1981) recent revision of the Molossidae, phylo-
genetic relationships and taxonomic assignments within the family
130
Annals of Carnegie Museum
VOL. 55
M SK AA Ad lA Ad AA
AK
M tt 40 M «• A« It*
X X
4A §• • ««
A
XS Kt n M AA A .
Y V
f/A (^6 Ofl A0 AO 04
A T
B
Fig. 2.— Representative karyotypes of A) Mops spurrelli from Cameroun, B) Mops ther-
sites from Cameroun.
were largely unexplored. Warner et al. (1974) suggested that chromo-
somal studies might be beneficial in evaluating these relationships.
Chromosomal data now available for 36 molossid species representing
10 of 12 genera recognized by Freeman (1981) are summarized in
Table 2.
We detected no intraspecific chromosomal variation within any of
the species examined in this study. This is noteworthy for two reasons.
First, our karyotypes of M. condylurus (FN = 66) from Afgoi, Somalia,
differ substantially from the karyotype of this species (FN = 56) re-
ported from Kisubi, Uganda, by Dulic and Mutere (1973). These lo-
calities are several hundred kilometers apart and this suggests either
that considerable geographic variation in the karyotype occurs within
this species or there are two species currently recognized as M. con-
dylurus. Secondly, our data support the specific distinctiveness of M.
spurrelli and M. nanulus. Freeman (1981) recognized the morpholog-
ical similarity between these two taxa and noted Koopman’s (1975)
suggestion that they might be conspecific. Our data indicate that M.
nanulus (FN = 54) and M. spurrelli (FN = 66) differ by five pairs of
biarmed chromosomes, and considering the scarcity of intraspecific
karyotypic variation within this family, it would seem likely that the
two taxa are specifically distinct.
Variation in FN for the species we examined ranged from 54 to 66
(Table 1). These karyotypes can be conveniently divided into three
1986
Smith et al.— Molossid Karyotypes
131
CC n IK
lA M fn OA Ifi lA M
AR RR (ft (# (ft %( ••
^A^XX' ««
00 AA AA A« M NO Ml M ••
#11 ^41 HA AA A# A# lO* *>« *•>
ITtlUil
il AOONNAI'^ MM M
ii tH »• »l» W ti oo
n/lf Mn
AA tfl HA Ail at «# at Ai ft
U
A
(#
B
X Y
c
h
XY
D
Fig. 3.— Representative karyotypes of A) Mops brachyptems from Cameroon, B) Mops
petersoni from Cameroun, C) Mops nanulus from Cameroun, and D) Mops demonstrator
from Cameroun.
groups. The high FN group (FN 62-66) includes both species of
Chaerephon and four of eight Mops species. Within this group, differ-
ences between the FN = 62-66 karyotypes apparently involve the
absence of medium and small subtelocentric autosomes. Our exami-
nation of Chaerephon pumila (FN ^58) agrees with the karyotype of
this species reported by Dulic and Mutere (1973), and forms an inter-
mediate FN group. Again, differences between the intermediate and
132
Annals of Carnegie Museum
VOL. 55
Table 2.— Summary of molossid karyotype data.
Species
In
FN
Reference
Chaerephon aloysiisabaudiae'^
48
66
This study
Chaerephon ansorgeP
48
66
This study
Chaerephon bivittata^
48
54
Peterson and Nagorsen, 1975
Chaerephon plicata^
48
54
Harada and Kobayashi, 1980; Harada
et al., 1982
Chaerephon pumila^
48
58
Dulic and Mutere, 1973; this study
Eumops auripendulus
42
62
Warner et al., 1974
Eumops glaucinus
38
64
Warner et al., 1974
40
64
Warner et al., 1974
Eumops perotis
48
56
Baker, 1970; Warner et al, 1974
48
58
Wainberg et al., 1974
Eumops underwoodi
48
56
Warner et al., 1974
Molossops abrasus
34
60
Warner et al., 1974; Gardner, 1977
Molossops greenhalli
34
_
Linares and Kiblisky, 1969
34
60
Baker, 1970; Warner et al., 1974
Molossops temminckii
42
56
Gardner, 1977
Molossus ater
48
58
Warner et al., 1974
Molossus molossus
48
56
Baker and Lopez, 1970
48
58
Warner et al., 1974
Molossus rufus
48
58
Wainberg et al., 1974
Molossus Sinaloa
48
58
Warner et al., 1974
Mops brachypterus'
48
54
This study
Mops condylurus^
48
66
This study
48
56
Dulic and Mutere, 1973
Mops demonstrator^
48
54
This study
Mops midas^
48
66
This study
Mops nanulus^
48
54
This study
Mops petersonP
48
54
This study
Mops spurrellP
48
64
This study
Mops thersites^
48
62
This study
Mormopterus kalinowskii’^
48
56
Warner et al., 1974
Mormopterus setiger'^
48
54
Warner et al., 1974
Nyctinomops aurispinosus^
48
58
Warner et al., 1974
Nyctinomops femorosacus^
48
58
Warner et al., 1974
Nyctinomops laticaudatus^
48
58
Warner et al., 1974
Nyctinomops macrotis^
48
58
Warner et al., 1974
48
56
Baker, 1970
Otomops martiensseni
48
56
Dulic and Mutere, 1973
Promops centralis
48
58
Warner et al., 1974
Promops nasutus
40
54
Wainberg, 1966
Tadarida brasiliensis
48
__
Painter, 1925
48
54
KniazefF et al., 1967
48
56
Warner et al., 1974; Baker et al, 1982
Tadarida fulminans
48
54
Peterson and Nagorsen, 1975
* Indicates species formerly recognized as Tadarida, see Freeman (1981).
1986
Smith et al.— Molossid Karyotypes
133
high FN forms appear to be in the absence of small subtelocentric pairs
plus the absence of one pair of medium-sized metacentrics. The low
FN group (FN =54) includes M. brachypterus, M. peter soni, M. dem-
onstrator, and M. nanuius. Karyotypic morphologies of the latter two
are identical and differ from the former pair in having one fewer me-
dium-sized metacentric pair and an additional pair of medium-sized
subtelocentric chromosomes.
Whether or not these karyotype associations reflect phylogenetic re-
lationships within genera is difficult to assess from standard karyotypic
data. Phylogenetic interpretations based on chromosomal data nec-
essarily require identification of homologous pairs using differential
staining techniques (see Haiduk et al., 1981). There is, however, little
concordance between Freeman’s (1981) phenetic classification and the
patterns of karyotypic morphology for these species. Within Chaere-
phon, Freeman’s (1981) analysis clusters C. ansorgei (FN = 66) with
C. bivittata (FN = 54, Peterson and Nagorsen, 1975). C aloysiisabau-
diae (FN = 66) then joins the cluster followed by C. pumila (FN = 58,
Dulic and Mutere, 1973; this study) several junctures later, and further
still, by C plicata (FN = 54, Harada and Kobayashi, 1980; Harada et
al., 1982). Similarly, within the genus Mops, M. demonstrator (FN =
54) clusters phenetically with M. condylurus (FN = 66); M. brachyp-
terus (FN = 54) and M. ther sites (FN = 62) pair together. These dis-
parities suggest the possibility that either morphological and chro-
mosomal characters are evolving at different rates, or that the taxonomic
relationships of these taxa need to be reexamined.
Karyotypic stability for bats, in general, has been recognized by
several authors (Peterson and Nagorsen, 1975; Gardner, 1977; Baker,
1978; Baker et al., 1982; Baker and Bickham, 1980; Bickham, \919a,
\919b\ Bickham and Baker, 1979) and has been suggested for the
Molossidae, specifically, by Warner et al. (1974) and Dulic and Mutere
(1973). Of the 36 molossid species for which chromosomal data are
available only seven species have diploid numbers other than 2n = 48
(Table 1). The modal occurrence of 2n = 48 chromosomes in both Old
and New World genera plus the similarity between this number and
the proposed primitive diploid number for the Vespertilionidae (Baker,
1970) led Warner et al. (1974) to propose 2n = 48 as primitive for the
Molossidae. Our documentation of the 2n = 48 karyotype in 1 1 Old
World molossid species further supports this diploid value as primitive
for the family.
Specimens Examined
Chaerephon aloysiisabaudiae. — Cameroun: 1 6 km S, 2 km E Yaounde (3®43'N, 1 1°32'E),
(16 CM 58678).
134
Annals of Carnegie Museum
VOL. 55
Chaerephon ansorgei. —Cameroun: 25 km S, 13 km E Garoua, (9°05'N, 13®30'E), (13
CM 58679).
Chaerephon Cameroon: 24 km S, 13 km E Garoua (9®05'N, 13®30'E), (13
CM 58724); Somalia: Libsoma Farm 6 km S, 17 km W Afgoi (2®05'N, 44®58'E), (333
CM 85438, 85439-85440; 299 CM 85441-85442); Somalia: Bulo Burti (3®51'N, 45®34'E),
(433 CM 85455-85456, CM 85459-85460; 499 CM 85457-85458, 85461-85462).
Mops brachypterus.—CAMEROvn: 25 km S, 3 km E Yaounde (3®38'N, ir33'E), (19
CM 58687).
Mops condylurus.—^OMAEw: Libsoma Farm, 6 km S, 17 km W Afgoi (2®05'N, 44®58'E),
(433 CM 85423, 85408-85409, 85424; 599 CM 85410-85411, 85412, 85425-85426).
Mops demonstrator.— CAMEROvn: 2 km W Ngaoundere (7®20'N, 13®34'E), (13 CM
58681).
Mops midas. —Somaeia: Libsoma Farm, 6 km S, 17 km W Afgoi (2°05'N, 44®58'E),
(333 CM 85429, 85436, 85428; 399 CM 85427, 85430).
Mops nanulus.—CAMEROuw. 25 km S, 13 km E Garoua (9®05'N, 13®30'E), (13 CM
58694); Cameroon: 24 km S, 13 km E Garoua (9®05'N, 13®30'E), (13 CM 58692; 299
CM 58693, CM 58695).
Mops petersoni. —Cameroun: 25 km S, 3 km E Yaounde (3®38'N, 11°33'E), (13 CM
58688; 19 CM 58691).
Mops spurrel/i.—CAMEROUN: 30 km N, 40 km E Obala (4®22'N, 11°58'E), (13 CM
58730); Cameroon: 25 km S, 3 km E Yaounde (3®38'N, 1 1®33'E), (233 CM 58731, CM
58786).
Mops thersites.—CAMEROUN: 30 km N, 40 km E Obala (4°22'N, 1 1®58'E), (13 CM
58737); Cameroon: 25 km S, 3 km E Yaounde (3°38'N, 1 1°33'E), (299 CM 58743, CM
58745); Cameroon: 7 km S, 8 km W Yaounde (3°48'N, 1 r27'E), (233 CM 58739, CM
58741; 299 CM 58740, CM 58742).
Acknowledgments
Field work in Somalia was supported by a grant from the M. Graham Netting Research
Fund, Cordelia Scaife May Charitable Trust and the Hays Fund of the American Philo-
sophical Society. Assistance in the field was provided by M. J. Smolen, R. Ruiz, Omar
Hagi, Abdulwahab Josuf, and Mohamed Ali. Facilities and support in Somalia was
graciously supplied by Mohamet Abdi Nur, Minister of Agriculture; Mohamed Abikar,
Director, General Ministry of Agriculture; and Abdulcadir Nur, Director, Department
of Plant Protection and Locust Control in the Ministry of Agriculture. Abdullahi Ahmed
Karani, General Manager, National Range Agency, issued the necessary permits to work
in Somalia. John and Jonquil Ash; Bill and Sally Smythe; Tony and Lynette Johnston,
United Nations Development Program in Somalia, graciously assisted in numerous ways.
Our special thanks to all these individuals.
Field work in Cameroun was supported by grants from the M. Graham Netting Re-
search Fund, Cordelia Scaife May Charitable Trust; the Loyalhanna Foundation; and
the National Geographic Society. Assistance in the field was provided by L. W. Robbins,
R. L. Robbins, and S. L. Williams. Permission to conduct field work and collecting
permits were received from the Ministry of Agriculture. We are indebted to Mr. V.
Belinga, Director of Forestry Services in Cameroun and Mr. C. Njiti for assistance in
obtaining the necessary permits. Laboratory work was supported by NSF grant No. PCM-
8202794 to J. W. Bickham. Partial financial support for field work was received from
NIH Grant AIO 4242 to R. Traub.
Literature Cited
Baker, R. J. 1970. Karyotypic trends in bats. Pp. 65-96, in Biology of bats (W. A.
Wimsatt, ed.). Academic Press, New York, 406 pp.
1986
Smith et al.—Molossid Karyotypes
135
■ . 1978. Karyology. Pp. 107-155, in Biology of bats of the New World family
Phyllostomatidae, Part III (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.).
Spec. Publ. Mus., Texas Tech Univ., 16:1-441.
Baker, R. J., and J. W. Bickham. 1980. Karyotypic evolution in bats: evidence of
extensive and conservative chromosomal evolution in closely related taxa. Syst.
ZooL, 29:239-253.
Baker, R. J., M. W. Haiduk, L. W. Robbins, A. Cadena, and B. F. Koop. 1982.
Chromosomal studies of South American bats and their systematic implications.
Spec. Publ. Ser. Pymatuning Lab. EcoL, 3:303-327.
Baker, R. J., and G. Lopez. 1970. Karyotypic studies of the insular populations of
bats of Puerto Rico. Caryologia, 23:465-472.
Bickham, J. W. 1979a. Chromosomal variation and evolutionary relationships of
vespertilionid bats. J. Mamm., 60:350-363.
Bickham, J. W. 1979^. Banded karyotypes of 11 species of American bats (genus
Myotis). Cytologia, 44:789-797.
Bickham, J. W., and R. J. Baker. 1 979. Canalization model of chromosomal evolution.
Bull. Camagie Mus. Nat. Hist., 13:70-84.
Corbet, G. B., and J. E. Hill. 1980. A world list of mammalian species. British Mus.
(Nat. Hist.), London, viii + 226 pp.
Dulic, B., and F. a. Mutere. 1973. Comparative study of the chromosomes of some
molossid bats from eastern Africa. Period. Biol., 75:61-65.
Freeman, P. W. 1981. A multivariate study of the family Molossidae (Mammalia,
Chiroptera): morphology, ecology, evolution. Fieldiana ZooL, Chicago Mus. Nat.
Hist. 1316 n.s. 7.
Gardner, A. L. 1977. Taxonomic implications of the karyotypes of Molossups and
Cynomops (Mammalia:Chiroptera). Proc. Biol. Soc. Washington, 89:545-550.
Haiduk, M. W., R. J. Baker, L. W. Robbins, and D. A. Schlitter. 1981. Chromo-
somal evolution in African Megachiroptera: G- and C-band assessment of the mag-
nitude of change in similar standard karyotypes. Cytogenet. Cell Genet., 29:221-
232.
Harada, M., and T. Kobayashi. 1980. Studies on the small mammal fauna of Sabah,
East Malaysia II. Karyological analysis of some Sabahan mammals (Primates, Ro-
dentia, Chiroptera). Contrib. Biol. Lab. Kyoto. Univ., 26:83-95.
Harada, M., M. Minezawa, S. Takada, S. Yenbutra, P. Nunpakdee, and S. Ohtani.
1982. Karyological analysis of 12 species of bats from Thailand. Caryologia, 35:
269-278.
Honacki, j. H., K. E. Kinman, and J. W. Koeppl (eds.). 1982. Mammal species of
the world. Allen Press, Inc., and the Association of Systematics Collections, Law-
rence, Kansas, 694 pp.
Kniazeff, a. j., D. Constantine, W. A. Nelson-Rees, D. Schmidt, and R. Owens.
1967. Studies in chiropteran cell lines. 41st. Tech. Progr. Rept. Naval Biol. Lab.
Suppl. Rept, CC-8:97-105.
Koopman, K. F. 1975. Bats of the Sudan. Bull. Amer. Mus. Nat. Hist., 154:354-444.
Linares, O. J., and P. Kibilisky. 1969. The karyotype of a new record of Molossups
greenhalli from Venezuela. J. Mamm., 50:831-832.
Painter, T. S. 1925. A comparative study of the chromosomes of mammals. Amer.
Nat, 59:385-408.
Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus
(Rodentia: Heteromyidae). J. Mamm., 48:27-37.
Peterson, R. L., and D. W. Nagorsen. 1975. Chromosomes of fifteen species of bats
(Chiroptera) from Kenya and Rhodesia. Occas. Papers, Royal Ontario Mus. Life
ScL, 27:1-14.
Wainberg, R. L. 1966. Cytotaxonomy of South American Chiroptera. Arch. Biol.
(Liege), 77:411-423.
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Wainberg, R. L., L. H. Delupi de Bianchini, J. J. Bianchini, and G. E. Pollero de
Actis Dato. 1974. Uniformidad cariotipica y radiacion adaptativa en “Eumops”
y “Molossus” (Chiroptera, Molossidae). Physis Secc. C Cont. Org. Terr., 33:249--
254.
Warner, J. W., J. L. Patton, A. L. Gardner, and R. J, Baker. 1974. Karyotypic
analyses of twenty-one species of molossid bats (MolossidaeiChiroptera). Canadian
J. Genet. Cytol., 16:165-176.
^ ISSN 0097-4463
ANNALS
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CARNEGIE MUSEUM OF NATURAL HISTORY
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VOLUME 55 23 MAY 1986 ARTICLE 8
TRILOBITES FROM THE KEOKUK LIMESTONE
(MISSISSIPPIAN) OF MISSOURI
David K. Brezinski^
Research Associate, Section of Invertebrate Fossils
Abstract
A new species of the trilobite genus Griffithides Portlock, G. salinensis, new species,
is described. G. salinensis is a rare component of the fauna of the Keokuk Limestone
(Mississippian) of St Genevieve County, Missouri. It can be distinguished from other
described species assigned to this genus by the posterior termination of the pygidial axis,
which is upturned into a large node-like projection. The type specimens of this species
were collected from just below the Marginarugus magnus bed of the Keokuk Limestone.
In addition to G. salinensis, two specimens also recovered from the same bed are de-
scribed and tentatively assigned to the genus Waribole Richter and Richter.
Introduction
The trilobite genus Griffithides Portlock is relatively poorly known
from Carboniferous rocks of North America as compared to the many
species recognized from correlative rocks of Europe. Of the three North
American species recognized, only one, G. bufo Meek and Worthen, is
known from more than just the type material. In contrast, specific
diversity exhibited by this genus in Europe has prompted some authors
(Hahn and Hahn, 1 970, 1971; Hahn et al., 1983) to subdivide the genus
* Present address: The Maryland Geological Survey, The Rotunda —Suite 440, 71 1 W.
40th Street, Baltimore, MD 21211.
Submitted 21 August 1985.
137
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VOL. 55
into a number of subgenera. The recent recovery from the Keokuk
Limestone of Missouri of a small number of specimens of Griffithides,
which are notably dissimilar to previously recognized species, warrants
the erection of a new species. The specimens of Griffithides, and two
pygidia tentatively assigned to the genus Waribole were recovered from
a light "gray lime grainstone in the Keokuk Limestone along Interstate
55 at Little Saline Creek, St. Genevieve County, Missouri. The spec-
imens were recovered as accessories during field collections made for
brachiopods by J. L. Carter and A. D. Kollar (Carnegie Museum of
Natural History) from a bed just below the Marginarugus magnus bed
or about in the middle of the Keokuk. The brachiopod fauna associated
with these trilobites is dominated by Productus crawfordsvillensis Well-
er, Imbrexia montonana Miller, Rhynchopora beecheri Greger, and
Torynifer pseudolineatus (Hall). The lithologic character of the strata,
which yielded the trilobites, is similar to that in which other North
American species of Griffithides have been found. Moreover, the spec-
imens of Waribole? recovered from the Keokuk are similar to pygidia
I have recovered from the lime grainstones of the Salem Limestone of
southern Indiana. These specimens from the Salem are also found in
association with a species of Griffithides. The consistent occurrence of
these two trilobite genera together in particular lithologies may suggests
a strong ecologic control on their distribution.
Terminology employed in this study follows that utilized by Har-
rington (1959).
Systematic Paleontology
Family Proetidae Salter
Subfamily Griffithidinae Hupe
Genus Griffithides Portlock
Distribution of North American Present in the Keokuk
Limestone (Osagean) of Missouri and Illinois, the Salem Limestone
(Meramecian) of Indiana, and the Pitkin Limestone (Chesterian) of
Oklahoma.
Diagnosis of North American representatives. —Ctphdilon parabolic
in outline, moderately vaulted. Glabella pyriform with frontal lobe
moderately to greatly expanded laterally and reaching the anterior mar-
gin of the cranidium. Lateral preoccipital lobes well-defined, subtrian-
Fig. \. — Griffithides salinensis, new species, A, C, E, holotype pygidium in dorsal, lateral,
and posterior views, CMNH 34553, x2; B, D, paratype pygidium in dorsal and lateral
views, CMNH 34498, x2; F, paratype pygidium, dorsal view, CMNH 34499, x2.5; G,
H, I, paratype cranidium in dorsal, lateral and anterior views, CMNH 34500, x2.5.
1986
Brezinski— Keokuk Trilobites
139
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Annals of Carnegie Museum
VOL. 55
Fig. 2.— Bivariate plot of maximum pygidial width (W) vs maximum pygidial length (Z)
for specimens of Griffithides salinensis, new species, and G. bufo Meek and Worthen.
gular, Ip furrow narrow, deeply incised, widened anterolaterally. Pal-
pebral lobes crescentic to semicircular in outline. Eyes of medium size,
hemispherical in shape. Lateral border furrow well-defined and narrow,
margin sharply rounded. Genal spines short, generally reaching to the
third thoracic segment.
Thorax of nine segments, axial rings semicircular in transverse pro-
file, ornamented by a row of small granules along the posterior margin.
Pleurae sharply rounded at fulcrum, subangular at tip.
Pygidium semicircular to parabolic in outline, strongly vaulted, con-
sisting of an axis composed of 1 1-15 rings and pleural fields of 9-14
ribs. Axis tapers posteriorly, does not reach posterior margin, strongly
convex and steeply downsloping posteriorly. Pleural ribs extend nearly
to the margin, with no well-defined border.
Discussion. — Although Hahn et al. (1983) were able to subdivide the
Eurasian representatives of the genus Griffithides into three subgenera,
no such division of North American species is possible. The main
distinguishing features among North American species of Griffithides
lie mainly in the shape (suboval versus parabolic) and structure (num-
ber of ribs and rings) of the pygidium. Griffithides can be distinguished
from the contemporary trilobite genus Paladin by the presence of a
well-developed pygidial border on the latter.
1986
Brezinski— Keokuk Trilobites
141
Table \. — Univariate measurements of select morphological characters of Griffithides
salinensis, new species. For discussion and illustration of character definition see Shaw
(1957).
Character
N
Mean*
Range*
(W) pygidial width
7
16.2
12.7^19.6
(Z) pygidial length
7
13.4
10.1-16.8
(X) axial width (max.)
7
6.3
4.9-7.4
(Y) axial length
6
12.2
8.4-14.7
number of axial rings
6
15
14-15
number of axial ribs
5
14
13-14
* Measurements in millimeters.
Griffithides salinensis, new species
Figs. 1A~1I
Holotype. -CMNn 34553.
Paratypes.~CMN¥L 34498-34500.
Material ~\0 incomplete pygidia and 1 partial cranidium.
Description.— pyriform with moderate lateral expansion to the frontal lobe.
Frontal lobe strongly convex in transverse profile and extends to the anterior margin. In
longitudinal profile glabella is nearly flat to mildly convex at the posterior terminus,
becoming increasingly convex anteriorly, meeting the anterior margin vertically. Glabella
covered by fine granules. Lateral preoccipital lobes subtriangular with granular ornament;
Ip furrow well-defined and of medium width, becoming broader toward the dorsal furrow.
Palpebral lobes of medium size, semicircular in outline, inclined into the dorsal furrow
at about 45® Facial sutures mildly divergent from a to 0, rounded at wider at co than
a. Occipital lobe not preserved.
Thorax is unknown.
Pygidium parabolic in outline, moderately vaulted, .83 times as long as wide. Axis
tapers posteriorly, .91 the total pygidial length, ,39 the total pygidial (anterior) width,
composed of 14 to 15 rings which are semicircular in transverse profile. Posteriormost
axial ring is enlarged into a large node or nub that overhangs a slightly concave axial
terminus. Each ring is slightly sinuous, being posteriorly bent across axis. A row of 1 2
fine granules ornament the posterior edge of each ring. Pleural fields strongly convex,
made up of 13--14 posteriorly recurved ribs that extend nearly to the margin. The
anteriormost three or four ribs exhibit a well-defined pleural furrow. A row of fine granules
ornament each rib.
Discussion. — G. salinensis, new species, can readily be distinguished
from other North American species of the genus by the upturned node-
like termination of the pygidial axis, by the greater length to width
ratio to the pygidium, and by the greater number of axial rings and
pleural ribs. Only G. bufo Meek and Worthen has been recovered in
sufficient number to allow any close comparison. Fig. 2 is a bivariate
plot comparing the maximum pygidial widths with the maximum py-
gidial lengths for specimens of G. salinensis and G. bufo. There is a
noticeable difference in the rectilinear trends exhibited by each species.
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Annals of Carnegie Museum
VOL. 55
Fig. 3.— Waribolel sp. A, B, dorsal and posterior views of complete pygidium, CMNH
34501, x.2.5.
The plot illustrates that G. salinensis possesses a greater length to width
ratio than does G. bufo and is larger on the average. Unfortunately,
insufficient numbers of G. salinensis are available at present to produce
a reliable regression equation. G. salinensis differs from the poorly
known G. meramecensis Shumard in that the latter exhibits fewer
pygidial ribs and rings (12 and 13, respectively) and lacks the node-
like termination of the axis. Both G. meramecensis and G. salinensis
were recovered from the Keokuk Limestone of Missouri. The com-
parison presented above is based upon the drawing and description of
G. meramecensis presented by Shumard (1855). Inasmuch as the lo-
cation of the holotype is unknown, comparison must be based upon
Shumard’s description and drawing alone. G. salinensis differs from
G. pustulosus Snider in that the pygidium of the former exhibits a
parabolic outline and the prominent terminal node on the pygidial axis.
All Eurasian species of Griffit hides can be distinguished from G. sali-
nensis by the terminal axial node on the pygidium.
Subfamily Cyrtosymbolinae Hupe
Genus Waribole Richter and Richter
Waribolel sp.
Fig. 3A, 3B
Material. — 1 complete and 1 fragmented pygidium from the Keokuk Limestone. IL
lustrated specimen CMNH 34501.
Description.— with low vaulting and relief, semicircular in outline with a
length/ width ratio of .65. Axis tapers posteriorly, is .85 the total pygidial length and .37
the maximum (anterior) pygidium width, composed of 1 1 rings and terminates, poste-
riorly, at inside margin of border. Pleural areas composed of six or perhaps seven ribs
which become increasingly obsolete posteriorly. Each rib composed of two bands of
approximately equal width. Border well-developed, smooth, and slightly concave to the
margins, of nearly equal width all along pygidium.
Discussion. — The genus Waribole is most common in Late Devonian
rocks and has a documented range into the earliest Carboniferous. The
genus, to the best of my knowledge, has not been definitely documented
1986
Brezinski— Keokuk Trilobites
143
from the stratigraphic interval of the Keokuk Limestone (that is, Vi-
sean). Moreover, Waribole has not previously been reported from North
America. If these specimens are in fact not specimens of Waribole,
then they belong to some very similar genus. At present insufficient
specimens are available to make any further inference. The only other
North American trilobite genus with pygidial characteristics somewhat
similar to these two pygidia is Richterelia; Richter ella, however, ex-
hibits a much greater vaulting to the pygidium and the pleural ribs
lack the subdivision into anterior and posterior bands.
Acknowledgments
All specimens utilized in this study are reposited in collections in the Section of
Invertebrate Fossils, Carnegie Museum of Natural History (CMNH). Thanks are owed
to Dr. John L, Carter and Albert Kollar who collected the specimens and made them
available to me. Drafts of the manuscript were read by J. L. Carter and C. A. Kertis,
and critically reviewed by J. H. Stitt and J. F. Taylor.
Literature Cited
Hahn, G., and R. Hahn. 1970. Trilobitae carbonici et permici 11. (Proetidae: Griffi-
thidinae). Pp. 162-331, in Fossilium Catalogus 1. Animalia (F, Westphal, ed.), ’s
Gravenhage (Dr. W. Junk N. V.), 1 19.
1971. Revision von Griffithides {Bollandia) (TriL: UnterKarbon). Palaeonto-
graph., 137:109-154.
Hahn, G. R. Hahn, and C. Brauckmann. 1983. Die Trilobitien des belgischen Koh-
lenkalkes (Unter-Karbon) 5. Griffithides and Cyphinoides. Geol. et Paleont., 1 7: 109-
135.
Harrington, H. J. 1959. General description of trilobita. Pp. 38-1 17, Treatise on
Inverterbrate Paleontology, Part O, Arthropoda 1 (R. C. Moore, ed.), Univ. Kansas
Press, Lawrence, Kansas, 506 pp.
Shaw, A. B. 1957. Quantitative trilobite studies II. Measurement of the dorsal shell
of non-agnostidean trilobites. J. Paleont., 31:193-207.
Shumard, B. F. 1855. Description of a geological section on the Mississippi River
from St. Louis to Commerce. 1st and 2nd Annual Rept., Geol Surv. Missouri, 2:
85-208.
Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.
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CARNEGIE MUSEUM OF NATURAL HISTORY
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VOLUME 55 15 SEPTEMBER 1986 ARTICLE 10
RESULTS OF THE ALCOA FOUNDATION SURINAME
EXPEDITIONS. X. PATTERNS OF CELLULAR
DIVERGENCE AND EVOLUTION IN THE GASTRIC
MUCOSA OF TWO GENERA OF PHYLLOSTOMID
BATS, TRACHOPS AND CHIRODERMA
Keith M. Studholme*’^
Carleton J. Phillips^
Research Associate, Section of Mammals
G, Lawrence Forman^
Abstract
The fundic mucosa in Trachops cirrhosus, Chiroderma trinitatum, and C villosum
(Suborder Microchiroptera) was studied comparatively by transmission electron mi-
croscopy. Trachops is an animalivorous species that mainly feeds on Neotropical frogs,
whereas both species of Chiroderma are frugivorous. In Trachops, two types of entero-
endocrine cells (A and D cells) that possibly produce glucagon and somatostatin, re-
spectively, were identified ultrastructurally. In Chiroderma examples of possible A, EC^,
Di(H), and G cells were identified. The product in possible G-cells in Chiroderma ultra-
structurally matched that found in the pylorus of another stenodermatine bat, Ariteus
flavescens, which has been shown to exhibit gastrin-like immunoreactivity. In Trachops
' Address: Department of Biology, Hofstra University, Hempstead, New York 11550.
^ Present address: Department of Neurobiology and Behavior, SUNY Stony Brook, Stony
Brook, New York 11794.
^ Address: Department of Biology, Rockford College, Rockford, Illinois 61101.
Submitted 7 October 1985.
207
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Annals of Carnegie Museum
VOL. 55
the pepsin-producing chief cells are restricted to the basal-most portion of the gastric
gland and produce an electron-dense product, whereas in Chiroderma the chief cells
occupy up to 75% of the gland and produce a pale product. In Trachops the parietal cells
are less abundant and less active than in Chiroderma but mucous neck cells are far more
abundant. The gastric pits are shallow in Chiroderma, whereas Trachops has deep gastric
pits. The mucous secretory granules in the surface mucous cells in Chiroderma differ
ultrastructurally from those in Trachops. Overall, the ultrastructure of gastric mucosa in
Trachops resembles that found in Phyllostomus (animalivorous/omnivorous) and Pteron-
otus (insectivorous), whereas these features in Chiroderma resemble those of Artibeus
(frugivorous). The general histology of the stomach of Chiroderma is similar to that of
a megachiropteran, Eidolon helvum, suggesting that histological convergence has occurred
in the evolution of the stomach of microchiropteran and megachiropteran fruit bats.
Introduction
Patterns of morphological evolution in bats are complex and un-
doubtedly constrained by many factors (Hill and Smith, 1984). Among
these, dietary specialization appears to explain many chiropteran mor-
phological features. Dental morphology, salivary gland structure, kid-
neys, gastric anatomy, relative brain size, jaws, and even length of
forearm can be correlated with diet (Phillips, 1971; Phillips et al., 1977;
Studier et al., 1983<2; Eisenberg and Wilson, 1978; Findley and Wilson,
1982; Freeman, 1979, 1981). Consequently, bats represent an outstand-
ing mammalian model against which hypotheses about the relation-
ships among structure, function, and biological role of various ana-
tomical features can be tested.
The digestive tract, and the stomach in particular, is especially in-
teresting in bats because diet can be correlated with gross anatomy,
musculature, histochemistry, and histology (for example, Park and Hall,
1951; Kolb, 1954; Schultz, 1965, 1970; Rouk and Glass, 1970; Hart,
1971; Forman, 1971, 1972; Forman et al., 1979; Kamiya and Pirlot,
1975; Ogunbiyi and Okon, 1976; Okon, 1977; Bhide, 1980; Yamada
et al., 1984; Ishikawa et al., 1985). These correlations have been ex-
tended through recent studies in which we have demonstrated differ-
ences in 1) aspects of the ultrastructure of cell types, 2) possible presence
or absence of certain entero-endocrine (endocrine-paracrine) cells, and
3) relative numbers of particular cell types (Phillips and Studholme,
1982; Phillips et al., 1984; Mennone et al., 1 986). A variety of questions
has been left unanswered by previous investigations. For example, we
do not yet know exactly to what extent cellular and subcellular mor-
phology differs in conjunction with diet or to what extent species with
similar diets have similar cellular patterns, regardless of systematic
associations.
For the present investigation we compared two genera of phyllos-
tomid bats, Trachops (a phyllostomine) and Chiroderma (a stenoder-
matine), that represent probable extremes in feeding specialization. We
1986
Studholme et al.— Bat Gastric Mucosa
209
chose Trachops cirrhosus because it has been characterized as an ani-
malivorous species that feeds on small vertebrates, especially Neo-
tropical frogs (Gardner, 1977; Tuttle and Ryan, 1981). As further evi-
dence of feeding specialization, it has been shown that Trachops responds
differentially to the vocal advertisements of edible and presumably
non-edible (poisonous) anurans (Tuttle and Ryan, 1981) and has his-
tologically unique accessory submandibular salivary glands that also
might relate to its dietary habits (Phillips and Tandler, 1985; Phillips
et al., in press). Chiroderma trinitatum and C viliosum are frugivores
(Gardner, 1977), and Forman (1973) and Forman et al. (1979) have
reported that the stomach of Chiroderma viliosum has an unusually
large fundic caecum making it . perhaps the most extreme spe-
cialization for plant feeding ...” among the phyllostomids. Insofar as
previous cellular comparisons are concerned, Phillips and Studholme
(1982) demonstrated a significant ultrastructural difference in chief cells
in the fundic mucosa of representative frugivorous and carnivorous
(insectivorous and animalivorous) bats, including Trachops and Chi-
roderma. This ultrastructural difference was interpreted to mean that
pepsinogen secretion is greatest in the frugivores.
The present investigation addressed the following questions. 1) How
are dietary and structural specializations expressed at the cellular level
in the gastric mucosa of Trachops and Chiroderma? 2) How do spe-
cializations in Trachops and Chiroderma compare with other micro-
chiropterans for which data are available? 3) How do ultrastructural
features of the gastric mucosa in Chiroderma and other frugivorous
microchiropterans compare with those found in Old World frugivores
of the Suborder Megachiroptera?
Methods and Materials
Specimens of Chiroderma trinitatum (2 females; CM 77599, 77600), Chiroderma
viliosum (3 females; CM 76796, 76798, 77601), and Trachops cirrhosus (1 male, CM
63688; 1 female, CM 64048) were collected in the vicinity of Rudi Kappelvliegveld,
Brokopondo, Suriname. All specimens have been deposited in the collections of the
Section of Mammals, Carnegie Museum of Natural History (CM). The bats were captured
in nets at night (1700--2400 h) while feeding. Subsequently, they were kept overnight
without food until between 0900 and 1 200 h when they were anesthetized with an intra-
peritoneal injection of 0.25 cc of sodium pentabarbitol (50 mg/ml). A polyethylene tube
next was inserted into the stomach via the mouth and esophagus and approximately 1
cc of trialdehyde fixative (at ambient temperature) was injected into the digestive tract.
After approximately eight minutes, an incision was made into the abdomen and the
stomach removed and cut into 2 by 2 mm samples that included portions of the fundus.
Only samples from the fundus (Fig. 1) were used for the present report.
The fixation protocol was developed specifically for field projects involving transmis-
sion electron microscopy (Forman and Phillips, in press; Phillips, 1985). The primary
fixative (based on Kalt and Tandler, 1971) consisted of 3% glutaraldehyde, 1% parafor-
maldehyde, 0.5% acrolein, 2,5% dimethyl sulfoxide (DMSO), and 1 mM CaCf in 0.05
M cacodylate buffer at pH 7.2 with 0. 1 M sucrose. All tissues were stored in this primary
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Annals of Carnegie Museum
VOL. 55
fixative for approximately 20 h at ambient temperature (30--40°C), The fixative then was
replaced with fresh buffer (0.05 M cacodylate buffer, pH 7.2, with 0.1 M sucrose); tissues
were left in this solution, unrefrigerated, for ten days. When refrigeration was available,
the tissues were placed in fresh buffer with 3% glutaraldehyde and stored at 4®C. For
processing, the tissues were washed for one hour in 0.05 M cacodylate buffer (pH 7.2)
with 0.1 M sucrose and post-fixed for one hour in 1% OSO4 with cacodylate buffer and
sucrose. Tissues then were dehydrated in an alcohol series and embedded in Epon 812.
Thin sections were post-stained with uranyl acetate (saturated solution in 50% EtOH)
and lead citrate (Reynolds, 1963) and examined and micrographed with a Philips 201
transmission electron microscope (TEM) operated at 60 Kv. Semithin (0.5 nm) sections
were stained with toluidine blue for light microscopy.
Entero-endocrine cells in both genera were identified solely on an ultrastructural basis
using Grube and Forssmann (1979) and Solcia et al. (1981) as guides. The ultrastructure
of the acid-producing cells in our specimens was analyzed comparatively by using avail-
able experimental data as a guide to recognition of secretory state (Ito and Schofield,
1978; Schofield et al., 1979). Our descriptions thus are based on cells judged to be in
the same state of activity in both species.
Results
At the light microscopic (LM) level the fundic mucosa of Trachops
and Chiroderma differed greatly (Fig. 1). In Trachops both surface
mucous cells and chief cells were conspicuous because they contain
toluidine blue stained secretory granules. In Chiroderma gastric pits
either were very shallow or lacking and the basal half of each gastric
gland was composed mostly of chief cells containing unstained granules.
Transmission electron microscopic analysis revealed details of the var-
ious cell types, as described in the following paragraphs. At both the
TEM and LM levels of comparison no noteworthy individual variation,
other than the usual localized differences in fixation, was found among
specimens of the same species.
Entero-endocrine (endocrine-paracrine) cells.— In Trachops cirrho-
sus only two types of entero-endocrine cells were distinguished. The
most common was identified as an A-cell, present in nearly all gastric
glands, positioned among the chief and parietal cells in the lower one-
half of the gland (Fig. 1). These cells contained abundant spherical,
electron-dense secretory granules (averaging 285 nm in diameter) with
a narrow “halo” caused by an apparent space between the electron-
dense material and the granule membrane (Fig. 2a). The cytoplasm
contained scattered lamellar granular endoplasmic reticulum (GER),
relatively few mitochondrial profiles, and lipid-like droplets (Fig. 2a).
The A-cells frequently were juxtaposed to chief cells. The second cell
type, identified as a D-cell, also was found within the most basal portion
of the gastric gland, apparently often juxtaposed to A-cells. The D-cells
were characterized by spherical secretory product (330 nm in diameter)
with a finely granular appearance, exiguous GER, and few mitochon-
drial profiles (Fig. 2b)
In Chiroderma, four distinctive types of entero-endocrine cells were
1986
Studholme et al. — Bat Gastric Mucosa
21
Fig. L— Diagrams of the gross anatomy of the stomachs and histology of the mucosa in
Trachops cirrhosus and a representative of Chiroderma (C. villosum). Tissue samples
used for electron microscopy were taken from areas enclosed by circles in the stomach
diagrams. The mucosa diagrams illustrate the histology as well as the relative height of
the glands and relative numbers of particular cell types. Examples of the ultrastructure
of particular cell types can be found by using the “F” labels as a guide to illustrations.
Abbreviations are: py, pylorus; duo, duodenum; eso, esophagus.
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VOL. 55
found in the lowermost portion of the gastric gland amongst the chief
cells. One type contained highly irregular, somewhat granular secretory
product (Fig. 3b) and corresponded morphologically to an EQ cell.
These cells tended to have a pale, leached appearance with abundant
tubular smooth endoplasmic reticulum (SER) and little or no GER.
The second cell type most closely resembled the Di(H) cell (Grube and
Forssmann, 1979); its cytoplasm contained small, spherical, electron-
dense granules (approximately 1 60 nm in diameter), strands of GER,
lipid-like droplets, and prominent Golgi complexes. The third type,
which was by far the least common, corresponded to an A-cell in having
abundant spherical electron-dense granules (Fig, 3a). The fourth type,
which was common (possibly associated with each gastric gland), con-
tained a mixture of granules most of which were small and contained
a coarsely granular substance equivalent to that characteristic of gastrin-
secreting G-cells (Fig. 4a, b). These cells were distinctive in that the
cell body had elongate processes that extended among the chief cells
so that cross-sectioned profiles were seen commonly (Fig. 4b). These
elongate processes and the cell body itself were generally wedged in
among the chief cells to the extent that chief cell secretory granules
often intruded into the cytoplasm of the G-cell (Fig. 4a, b). This image
was common and probably not an artifact because the granule mem-
brane and cell membranes typically were intact (Fig. 4b). In the pro-
Fig. 2.— a: An A-cell positioned at the base of the fundic gland in Trachops. Note the
spherical, electron-dense product (arrow) and lipid droplets (L) typical of these cells. T.
cirrhosus, CM 63688. b: An example of a D-cell positioned among chief cells in Trachops.
Note the typical secretory product (arrow) of these cells, which are thought to produce
somatostatin. T. cirrhosus, CM 63688.
Fig. 3.— a: An example of an A-cell in Chiroderma. The mature secretory product (arrow)
can be compared to an immature, electron-dense granule (img) associated with the Golgi
complex. C villosum, CM 77601. b: An ECn-cell in Chiroderma; the granules (arrows)
typically are elongate rather than spherical and possibly contain serotonin (5-HT). The
cytoplasm of these cells often is pale and contains abundant tubular smooth endoplasmic
reticulum (SER). A chief cell secretory granule (SG) also can be seen on the left. C
villosum, CM 76798.
Fig. 4.— a: A possible G-cell at the base of a fundic gland in Chiroderma. Note how the
body of the cell is wedged among adjacent chief cells, whose secretory product (SG)
intrudes into the G-cell cytoplasm. The cell and granule membranes (arrow) are intact,
suggesting that this close relationship is not an artifact, bl, basal lamina. C. villosum,
CM 76798. b: A cross-section through a G-cell process that extends between chief cells
filled with product (SG). An arrow denotes an image suggestive of exocytosis; also note
the tubular smooth endoplasmic reticulum (SER) and intact chief cell and granule mem-
brane (MB). C villosum, CM 76798.
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cesses we found tubular smooth endoplasmic reticulum and images
suggestive of exocytosis, suggesting that the G-cell granules might be
released here (Fig. 4b).
Chief cells. — The gastric chief cells in Trachops were clustered at the
base of the gastric gland (Fig. 1 ). The cells were pyramidal and contained
spherical, electron-dense secretory product; the nucleus was irregular
or ovoid and basally positioned; the GER profiles usually were swollen
and spherical in appearance and the cistemae were filled with pale
granular material (Fig. 5 a). The Golgi complex was small and incon-
spicuous.
The chief cells in Chiroderma were extremely abundant, occupying
at least half of each gland (Fig. 1). These cells differed dramatically
from those in Trachops in that the abundant product did not stain with
toluidine blue. Additionally, with the TEM the product was swollen,
pale granules. Although many of the granules were in the form of
individual spheres, others often had a coalesced appearance (Fig. 5b).
The nucleus was basally positioned and usually irregular; lamellar GER
and large Golgi complexes filled most of the remaining cytoplasm.
Parietal (oxyntic) cells. — The cytoplasm of parietal cells in Trachops
was characterized by a modest number of profiles of intracellular can-
aliculi. The microvilli of these canaliculi were closely appressed and
the intercellular spaces between them were filled with electron-dense
product matching that found in the gland lumen (Fig. 6a, b). The
cytoplasm of most parietal cells also contained large numbers of mi-
tochondrial profiles and tubular and spherical vesicles (Fig. 6b). The
parietal cells in our specimens of Trachops nearly always corresponded
to an intermediate activity state (following Ito and Schofield, 1978).
In Chiroderma, the parietal cells could be placed into two categories:
1) an intermediate secretory stage, which was most common; and 2)
an actively secreting stage, which was far less common (Fig. 7a, b).
The ultrastructure of parietal cells in the intermediate stage differed
from those in Trachops in the same stage in that these cells invariably
had extensive, swollen-appearing intracellular canaliculi with thick,
elongate microvilli (Fig. 7b). The cytoplasm of all parietal cells had
abundant mitochondrial profiles and numerous vesicles, scattered GER
profiles, and concentrations of lipofuscin granules (Fig. 7a, b). In active
parietal cells, the intracellular canaliculi were greatly expanded and the
Fig. 5.— a; Chief cell in Trachops; note the spherical electron-dense product (SG) and
example of exocytosis (arrow) into the lumen. NU, nucleus. T. cirrhosus, CM 63688. b:
Chief cells in Chiroderma; note the pale product (SG) and example of coalescing among
secretory granules (CO-SG). NU, nucleus. C. villosum, CM 76798.
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microvilli lining their borders were less compact (Fig. 7a). The cyto-
plasm of these cells contained few vesicles; instead, the Golgi complexes
were the most prominent feature.
Mucous cells.— In Trachops, the mucous neck cells were scattered
among the chief and parietal cells in the lower one-half of the gastric
gland. The apex of each mucous neck cell bordered on the gland
lumen. The nuclei were basally positioned and the apical cytoplasm
contained secretory granules with pale, finely granular material and a
distinctive electron-dense droplet occupying about one-fourth of the
granule diameter (Fig. 8a).
The surface mucous cells and cells lining the gastric pits differed
considerably from the mucous neck cells. The secretory product in
these surface cells was more clearly defined (coalescing granules were
not observed) and consisted of an electron-dense droplet set against a
reticulated, dense substrate (Fig. 9). Some granule polymorphism was
apparent; some of these cells also contained spherical or irregular,
completely electron-dense granules located in the vicinity of the Golgi
complex (Fig. 9).
The mucous neck cells in Chiroderma were relatively small and
sparsely distributed and they primarily were positioned among parietal
cells (Fig. 8b). The secretory product differed considerably from that
in homologous cells in Trachops. The granules in Chiroderma consisted
of a mixture of pale, flocculent material along with a coarser, fibrillar
material and a small, peripheral electron-dense component (Fig. 8b).
Gastric pits were nearly lacking in Chiroderma. The gastric surface
consisted of mucous cells and exfoliating parietal cells with dark, dense
Fig. 6, —a: Parietal cell in Trachops; note the typical image of intracellular canaliculi in
these cells in this species (arrow). NU, nucleus; Mi, mitochondria; M, mucous neck cell.
T. cirrhosus, CM 63688. b: Parietal cell in Trachops; note the intracellular canaliculi
(arrow) and tubular and spherical membrane profiles (V) in the cytoplasm. Abbreviations
as above. T. cirrhosus, CM 63688.
Fig. 7.— a: Actively secreting parietal cell in Chiroderma; note the expanded intracellular
canaliculus and cluster of lipofuscin granules (arrows). NU, nucleus; Mi, mitochondria;
L, lumen. C. villosum, CM 76796. b: Inactive parietal cell in Chiroderma; note the
lipofuscin granules (arrows) and compare the intracellular canaliculi (C) to those seen in
Trachops parietal cells (Fig. 6). Abbreviations as above. C. villosum, CM 76796.
Fig. 8. —a: Mucous neck cells in Trachops. An immature granule (IMG) in association
with the Golgi complex can be compared to mature mucous granules (MSG), which have
an electron-dense droplet (arrow). T. cirrhosus, CM 63688. b: Mucous neck cell in
Chiroderma. This cell is positioned near parietal cells (P) and is packed with mucous
granules (MSG), which have an electron-dense droplet (arrows) but otherwise differ from
those seen in Trachops. C. villosum, CM 76798.
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cytoplasm. The surface mucous cells differed considerably from those
in Trachops because the secretory product was pale and contained
fibrillar material as well as a small, denser sphere at the margin of each
granule (Fig. 10). Unlike the situation in Trachops, in Chiroderma the
surface mucous cells and mucous neck cells appeared to produce only
a single type of granule (Figs. 8b, 10).
Discussion
Comparative transmission electron microscopy has been shown to
be a useful addition to histological and histochemical techniques in the
study of systematics and evolutionary biology of mammals (for ex-
ample, Phillips et al., 1984; Feldman and Phillips, 1984; Tandler et
al., 1986; Phillips et al., in press; Phillips and Tandler, in press). An
analysis by TEM allows for interspecific comparisons of structure
among cells that are thought to be homologous (following criteria of
Rieger and Tyler, 1979) and in the same stage of differentiation. In
making our comparisons of ultrastructure we recognized the fact that,
in effect, we were comparing cells “fixed” at a moment in time and,
thus, to some extent the molecular events and organization that we
were viewing and comparing were aspects of functional state rather
than genic differences that directly determine cell structure (Phillips et
al., 1984). On the other hand, in making comparisons between regu-
lated, polarized secretory cells in the two genera of bats, we were able
to compare directly the ultrastructure of mature secretory granules
containing gene products such as mucus, pepsinogen, and a variety of
peptide hormones (Phillips and Tandler, in press). In our present in-
vestigation we found noteworthy ultrastructural differences among all
cell types and many of their products as well as in the relative numbers
of particular cell types in the fundic glands of two ecologically divergent
species of bats (Table 1).
Entero-endocrine cells are important because they synthesize and
secrete a wide variety of peptides as well as 5-hydroxytryptamine (sero-
tonin) and these products can have a complex, and as yet not fully
understood, controlling or modulating influence on the digestive tract
(Pearse, 1969; Grube and Forssmann, 1979; Solciaetal., 1981). Indeed,
although we use the terms “entero-endocrine” or “endocrine,” at least
some of these cells actually might be regarded as “endocrine-paracrine”
cells. Our identification of entero-endocrine cells was based solely on
Fig. 9. ““Surface mucous cells that line the surface of the stomach and the gastric pits in
Trachops. Note the variety of product images in the cytoplasm (an ows). Those nearest
to Golgi complexes (G) often are the most electron dense. NU, nucleus. T. cirrhosus,
CM 63688.
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Fig. 10.— Surface mucous cells in Chiroderma. These cells are packed with secretory
granules (MSG) that have an electron-dense droplet (arrow) and resemble those in mucous
neck cells (Fig. 8b). Note the sparse microvilli (MV) on the apical surface of these mucous
cells. C. villosum, CM 76798.
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Studholme et al. — Bat Gastric Mucosa
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Table L—A comparative summary of features of the fundic glands in Trachops cirrhosus
and Chiroderma trinitatum and C. villosum.
Feature
Trachops
Chiroderma
Comments
Entero-endo-
A, glucagon
A, glucagon
See Figs. 2, 3, 4
crine (=en-
D, somatostatin
EC„, serotonin
docrine™
D„?
paracrine)
cells
G, gastrin
Chief cells
Moderate num-
Extremely abundant
Ultrastructural differ-
bers
ences in secretory
product; Figs. 1, 5
Parietal cells
Small in size,
Extremely abundant.
Ultrastructural differ-
low average
large in size; very
ences allow for
activity level
high average ac-
comparisons of ac-
in fasted ani-
tivity level in fast-
tivity level; Figs.
mal
ed animal
6, 7
Mucous neck
Common
Scarce
Ultrastructural differ-
cells
ences in secretory
product; Fig. 8
Gastric pit
Abundant, with
Very sparse, gastric
Ultrastructural differ-
and surface
deep gastric
pits nearly non-ex-
ences in secretory
mucous cells
pits
istent
product; Figs.
1, 9, 10
ultrastmcture; thus, we could only infer products from experimental
literature. The following discussion of entero-endocrine cell products
is therefore largely reliant on the accuracy of ultrastructural identifi-
cation of particular cells; immunohistochemistry will be necessary for
more precise identification.
In Trachops we identified A-cells (Fig. 2), which are thought to secrete
glucagon (Moody et al., 1978; Unger et al., 1978) and D-cells, which
secrete somatostatin (Hokfelt et al., 1975; Grube and Forssmann, 1979).
Cells with essentially the same ultrastructure have been found in other
species; A-cells have been described in Pteronotus, Phyllostomus, Car-
ollia, and Artibeus, whereas D-cells have been found previously only
in Pteronotus and Phyllostomus (Phillips et al., 1984). Yamada et al.
(1984) have demonstrated both glucagon- and somatostatin-like im-
munoreactivity in endocrine cells in the fundic region of the stomach
of the common vampire bat, Desmodus rotundus, so although ultra-
structural data are lacking, this species also has both A- and D-cells.
Chiroderma differed from Trachops in that A-cells were rare and in
having ECn-cells, G-cells, and Di(H) cells, none of which was found in
the Trachops material examined by us. However, it must be remem-
bered that cell types that are scarce, or common but localized, could
easily be overlooked in a TEM survey. Light microscopy of semithin
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(0.5 fjLm) sections does not solve this problem because many endocrine
cells cannot be detected and identified with light microscopy and rou-
tine staining.
The apparent presence of abundant glucagon- and somatostatin-pro-
ducing entero-endocrine cells in Trachops is significant because these
cells 1) also are found in other insectivorous, animalivorous, and san-
givorous species (Phillips et aL, 1984; Yamada et aL, 1984), 2) are
abundant, 3) are positioned among chief and parietal cells, and 4)
because each of these products is known to inhibit gastric acid secretion
(Konturek et aL, 1975; Kusomoto et aL, 1979; Stachura et aL, 1981).
Additionally, the physical proximity of A- and D-cells in Trachops
resembles the situation in dogs, in which somatostatin apparently also
inhibits, or limits, glucagon production (Kusomoto et aL, 1979). In any
case, the abundant presence of A- and D-cells in Trachops might cor-
relate with the moderate number of generally inactive parietal cells,
just as we found previously in both Pteronotus and Phyliostomus (Phil-
lips et aL, 1984). Likewise, the relative rarity of glucagon-producing
cells and the possible absence or scarcity of somatostatin-producing
cells in Chiroderma correlates with the extreme activity of the large
and abundant parietal cells (Figs. 1 , 7).
The presence of and Di-cells in Chiroderma (Fig. 3) cannot
readily be related to any particular feature of the fundic gland. The
EQ-cells are thought to have endogenous serotonin ( 5 -hydroxy tryp-
tamine), which also is found in enteric neurons (Grube and Forssmann,
1979; Gershon, 1981). The physiological roles of entero-endocrine cell
serotonin possibly include inhibition of acid release along with stim-
ulation of mucus. Some investigators also regard serotonin as an in-
trinsic neurotransmitter that modulates smooth muscle and affects blood
flow (Gershon, 1981; Ormsbee and Fondacaro, 1985), Examples of an
EQ-type of cell also have been found in Phyliostomus hastatus so
although we failed to find examples of EQ-cells in our specimens of
Trachops, their occurrence in Chiroderma suggests that they might be
widespread in phyllostomid bats. This is further supported by Yamada
et aL (1984) who reported that they found moderate numbers of sero-
tonin-immunoreactive cells in the fundus of vampire bats.
The Di-cells in Chiroderma have an ultrastructure that is indistin-
guishable from that found in D, -cells in Artibeus (Phillips et aL, 1984).
Some authors have linked D, -cells to production of vasoactive intes-
tinal polypeptide (VIP) in the mammalian gut (for example, Grube and
Forssmann, 1979), but others (for example, Larsson et aL, 1979) have
argued that the peptide produced by D, cells is not VIP but a similar
molecule. Some workers now think that VIP in the digestive tract is
found only in nerve fibers and nerve cell bodies (Baecker et aL, 1983).
Another possibility is that the D, cells in Chiroderma produce motilin
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Studholme et al.—Bat Gastric Mucosa
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or bombesin, or both, because these peptides also have been associated
with cells that ultrastructurally fall into the Di cell group (Capella et
aL, 1978; Solcia et al., 1981). The product and physiological role of Di
cells in Chiroderma and Artibeus thus is unknown at the present time.
The possible G-cells found by us at the base of fundic glands in
Chiroderma are noteworthy for several reasons. Heretofore, gastrin-
producing cells have not been identified in the fundic region of the
mammalian stomach (Solcia et al., 1981) and Yamada et al. (1984)
did not find any gastrin-like immunoreactivity in the fundus of vampire
bats. Secondly, the gastrin peptide is known to stimulate gastric acid
secretion (for example. Chew and Hersey, 1982), which is considerable
in Chiroderma, if abundance, large size, and ultrastructure of parietal
cells are reliable indicators of the level of acid secretion in the absence
of quantitative data on gastric acid. Thirdly, the ultrastructure of the
G-cells and their cytoplasmic granules is consistent with published
descriptions of known G-cells in other mammals (Grube and Forss-
mann, 1979; Solcia et al., 1981) and with G-cells in the pylorus of
Ariteus flavescens, which were identified by demonstration of gastrin-
like immunoreactivity (Mennone et al., 1986). The apparent physical
relationship between G-cells and chief cells in Chiroderma is another
matter. It is not known with certainty that gastrin stimulates secretion
of pepsinogen; instead, chief cells in mammals have been shown to
have receptors to the peptides secretin and cholecystokinin (CCK) (Her-
sey et al., 1983, 1984). However, the ultrastructure of possible G-cells
in Chiroderma differs substantially from the ultrastructure of either
secretin or CCK-producing endocrine cells (Solcia et al., 1981) and thus
would not be easily confused with either of these. Gastrin and CCK
have been shown to share a common C-terminal portion of the mol-
ecule (Larsson and Rehfeld, 1977; Solcia et al., 1981) but in mammals
they are produced by separate cells and appear to have separate func-
tions. Indeed, it seems to be typical of entero-endocrine cells that their
various products are stored in membrane-bound granules that have
specific and consistent ultrastructural morphology and gastrin and CCK,
regardless of their molecular similarities, are found in very different-
looking granules (Phillips and Tandler, in press).
Available data on various types of entero-endocrine cells in Tra-
chops, Chiroderma, Phyllostomus, Carollia, Glossophaga, Artibeus, Ar-
iteus, Erophylla, Desmodus, and Pteronotus (Phillips et al., 1984; Ya-
mada et al., 1984; Mennone et al., 1986) raise far more questions than
answers. Some peptides (for example, glucagon, somatostatin, and gas-
trin) have been investigated to the extent that we can begin to relate
physiological data from laboratory studies of other species to our find-
ings in bats. However, most other peptides possibly produced in species-
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Specific patterns of differential abundance in bats are far more difficult
to integrate with other data because less is known about possible func-
tions. The problem is compounded by the fact that some of these
molecules are found in both enteric nerves and endocrine cells and
may be functionally different in each (Hokfelt et al., 1975; Grossman,
1976; Schultzberg et af, 1980; Gershon, 1981; Miller, 1984). Never-
theless, these regulatory molecules probably have played a key role in
the evolution of dietary diversity as seen in bats, and differential pro-
duction of peptides or differential location of cells that produce some
peptides might even contribute to or account for interspecific differ-
ences in histology of the chiropteran digestive tract (Mennone et aL,
1986). Various peptides, and 5-HT as well, ultimately might provide
a key to our understanding of the evolution of histological and func-
tional diversity.
Parietal cell ultrastructure in Chiroderma closely resembles that of
Artibeus and Ametrida (Phillips et al., 1984; Phillips, unpublished data)
and, judging from experimental studies, is indicative of a high rate of
HCl secretion, even in fasted animals (Ito and Schofield, 1978; Scho-
field et al., 1979; Black et al., 1980). This high rate of secretory activity
may be further substantiated by the abundance of lipofuscin granules
(Fig, 7) in Chiroderma parietal cells observed in all of our specimens.
This type of lysosome has been correlated with degradation of by-
products of cellular metabolism and its presence in parietal cells sug-
gests that cellular metabolism is occurring at a high rate (Toth, 1968;
Fawcett, 1981).
Chief cells (which produce pepsinogen and rely on gastric acid for
conversion to pepsin) also are extremely active in phyllostomid fruit
bats, judging from the unusual ultrastructural appearance of the product
(Fig. 5) in Chiroderma, Artibeus, and Ametrida (Phillips and Stud-
holme, 1982; Phillips et al., 1984). This pattern may be a widespread
occurrence in fruit bats because in Eidolon helvum, an African mega-
chiropteran, an analysis of digestive enzymes demonstrated high pepsin
content in both the stomach and lower esophagus (Ogunbiyi and Okon,
1976) and histological study has revealed abundant chief and parietal
cells (Okon, 1977).
Differences among the mucus-producing cells in Trachops and Chi-
roderma were significant, but in keeping with previously reported pat-
terns (Forman, 1972; Phillips et al., 1984). Trachops produces abun-
dant mucus, whereas Chiroderma produces little mucus due to the
limited number of mucous neck cells, the virtual absence of gastric pits
lined with mucous cells, and the relative scarcity of typical mucous
surface epithelial cells. In Trachops, the mucous cells ultrastructurally
resembled those found in Pteronotus and Phyllostomus (particularly in
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Studholme et al.—Bat Gastric Mucosa
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the latter species) and differed from those in Carollia and Artibeus
(Phillips et al., 1984) and Chiroderma. However, Trachops differed
from its nearest studied relative {Phyllostomus, cf. Smith, 1976; Hood
and Smith, 1 982) in having relatively shallower gastric pits (hence fewer
surface mucous cells) and in having a chemically different mucus in
the surface cells. In Phyllostomus, the surface mucous cells exhibit
toluidine blue metachromasia (Forman, 1972), suggesting the presence
of a sulfonated compound (Spicer, 1963), whereas in Trachops the
product is negative to toluidine blue staining. The significance of this
chemical difference is unknown. It might represent a real interspecific
difference but also could be the consequence of physiological state of
the animals at time of sacrifice because Ohara et al. (1984) have dem-
onstrated that histochemical changes in mucous glycoproteins can oc-
cur within hours in fasted laboratory rats. Although our bats all were
handled the same way after capture, we have no way of knowing for
certain whether they fed before being collected. On the other hand,
intraspecific consistency among our specimens does seem indicative
of a real interspecific difference.
The greatly reduced mucus-production in Chiroderma seems to be
characteristic of Neotropical fruit bats (Forman, 1972, 1973). Although
mucus often has been regarded as a major factor in the protection of
the gastric epithelium, its relative scarcity in fruit bats with highly
active parietal cells suggests otherwise. A recent study (Robert et al.,
1984) that demonstrated a lack of correlation between the thickness of
the mucus coat and protection of the stomach lining in laboratory rats
helps to explain the fruit bat data. But how is the stomach protected
in bats such as Chiroderma and Artibeusl One possibility is that the
lining of the stomach is protected by salivary gland secretions (Studier
et aL, 1983Z?; Phillips et al., 1984). Other possibilities include surface-
active phospholipids (Lichtenberger et al., 1983) and H+ disposal by
the surface c6lls through a Na+/H+ exchange system (Olender et al.,
1984).
In summary, it seems reasonably clear that mucous production
(amount and chemistry) in the fundus of the stomach in stenodermatine
fruit bats differs substantially from that in phyllostomine animalivo-
rous species. The mucus produced in fruit bats is less complex in the
sense that mucous neck and surface cells are ultrastructurally the same,
whereas in animalivorous species such as Trachops notable differences
can be found when one compares the secretory granules in these cells.
The biological significance of this intrafamilial divergence in mucous
cells and in ultrastructure of mature product is unknown but the con-
sistency of the pattern in the genera examined to date (Forman, 1972;
Phillips et al., 1984) is noteworthy because when differences in the
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ultrastructure of an exocrine cell product can be correlated with sys-
tematics, a foundation is set for studying evolution at the cellular level
(Tandler et aL, 1986; Phillips and Tandler, in press).
To what extent do our ultrastructural data provide answers for our
original question about the relationship among dietary and structural
specialization and cellular features of the gastric mucosa? Although
Trachops cirrhosus clearly represents an instance of behavioral evo-
lution that includes specialized feeding on Neotropical frogs (Tuttle
and Ryan, 1981), the gastric fundic mucosa at the cellular level is similar
to that of Phyllostomus hastatus and P. elongatus (Phillips et aL, 1 984),
which generally are regarded as “animalivorous” or “insectivorous”
(Gardner, 1977). The only differences of note are the possible absence
of ECn-cells, the slightly shallower gastric pits, and the absence of
toluidine blue positive components in the mucus in Trachops. The
fundic mucosa of Trachops thus seems to be somewhat generalized
even though these bats have adopted a specialized behavior and feeding
strategy.
Chiroderma is very similar to Artibeus at the cellular level (Phillips
et aL, 1984). It differs significantly, however, in having a relatively
greater abundance of chief cells (45“-75% of each fundic gastric gland
as compared to about 25%). Cellular differences in the gastric mucosae
of different Neotropical fruit bats are interesting because they suggest
the possibility of subtle differences in diet or in the assimilation of
nutrients from a shared diet.
Lastly, how do Chiroderma and other previously studied phyllos-
tomid fruit bats compare to the megachiropteran fruit bats? This ques-
tion is significant because evolutionary convergence is virtually un-
studied at the histological, histochemical, and ultrastructural levels.
Available data support the theory that frugivory evolved independently
in these bats (Kamiya and Pirlot, 1975; Smith, 1976) and some phys-
iological studies suggest that the two groups might be different in the
ways that they actually regulate their diets or dietary intake (Thomas,
1984). Furthermore, some data can be interpreted to show that mega-
chiropteran and microchiropteran bats might have had separate origins
altogether (Smith and Madkour, 1980). Given this, the general histo-
logical similarities in gastric mucosa of microchiropteran fruit bats of
the genera Chiroderma, Artibeus, and Ametrida on the one hand (pres-
ent study; Phillips and Studholme, 1982; Phillips et aL, 1984) and the
African megachiropteran. Eidolon helvum, on the other, are indeed
remarkable. Judging from the published data of Ogunbiyi and Okon
(1976) and Okon (1977), the microchiropteran fruit bats and E. helvum
have the following features in common: 1) very shallow, almost non-
existent, gastric pits; 2) a scarcity of mucous neck cells; and 3) abundant.
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231
extremely active parietal and chief cells. The extent of histological
convergence between the stenodermatines and megachiropterans as a
group is less certain because the latter seem to be quite variable and
published descriptions are not always adequate for comparisons. How-
ever, the stenodermatines do appear to share a variety of histological
features with Rousettus, Pteropus, Eonycteris, and Penthetor (Kamiya
and Pirlot, 1975; Bhide, 1980) and future ultrastructural comparisons
will be of interest.
Acknowledgments
Our research and field work were supported by the Research Corporation (grants
C-1251 and C-1855 to Phillips), Hofstra University (HCLAS Executive Committee
grants to Phillips), the Alcoa Foundation, and the M. Graham Netting Research Fund
(to Dr. H. H. Genoways, Carnegie Museum of Natural History). We are particularly
indebted to Hugh H. Genoways for his support and assistance to our research. Stephen
L. Williams, Jane Groen, Nadine M. Sposito, Rodney L. Honeycutt, Ben Koop, and
Mike Arnold all assisted with the field work. In Suriname, Drs. J. P. Schultz and H. A,
Reichart of STINASU were especially helpful and they are gratefully recognized. Lastly,
we greatly appreciate the work of our patient typist, Linda Cossen, of the Hofstra Uni-
versity Special Secretarial Services.
Literature Cited
Baecker, B,, N. Yanaihara, and W. G. Forssmann. 1983. VIP innervation of the
tongue in vertebrates. Anat. Embryol., 167:173“-189.
Bhide, S. A. 1980. Observations on the stomach of the Indian fruit bat, Rousettus
leschenaulti (Desmarest). Mammalia, 44:571-579.
Black, J. A., T. M. Forte, and J. G. Forte. 1 980. Structure of oxyntic cell membranes
during conditions of rest and secretion of HCl as revealed by freeze-fracture. Anat.
Rec., 196:163-172.
Capella, C., E. Hage, E. Solcia, and L. Usellini. 1978. Ultrastructural similarity of
endocrine-like cells of the human lung and some related cells of the gut. Cell Tissue
Res., 186:25-37.
Chew, C. S., and S. J. Hersey. 1982. Gastrin stimulation of isolated gastric glands.
Amer. J. Physiol., 5:G504-G512.
Eisenberg, j. F., and D. E. Wilson. 1978. Relative brain size and feeding strategies
in the chiroptera. Evolution, 32:740-751.
Fawcett, D. W. 1981. The cell. W. B. Saunders Co., Philadelphia, Second ed., 862
pp.
Feldman, J. L., and C. J. Phillips. 1984. Comparative retinal pigment epithelium and
photoreceptor ultrastructure in nocturnal and fossorial rodents: the eastern woodrat,
Neotoma jloridana, and the plains pocket gopher, Geomys bursarius. J. Mamm., 65:
231-245.
Findley, J. S., and D. E. Wilson. 1982. Ecological significance of chiropteran mor-
phology. Pp. 243-260, in Ecology of bats (T. H. Kunz, ed.). Plenum Press, New
York, 444 pp.
Forman, G. L. 1971. Histochemical differences in gastric mucus of bats. J. Mamm.,
52:191-193.
. 1972. Comparative morphological and histochemical studies of stomachs of
selected American bats. Univ. Kansas Sci. Bull., Lawrence, 59:591-729.
232
Annals of Carnegie Museum
VOL. 55
. 1973. Studies of gastric morphology in North American chiroptera (Embal-
lonuridae, Noctilionidae, and Phyllostomatidae). J. Mamm., 54:909=923.
Forman, G. L., and C. J. Phillips. In press. Preparation and fixation of tissues for
histological, histochemical, immunohistochemical, and electron microscopic stud-
ies. In Ecological and behavioral methods for the study of bats (T. H. Kunz, ed.),
Smithsonian Inst, Press, Washington, D.C.
Forman, G. L., C. J. Phillips, and C. S. Rouk. 1979. Alimentary tract, Pp. 205=227,
in Biology of bats of the New World family Phyllostomatidae. Part III (R. J. Baker,
J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., Lubbock,
441 pp.
Freeman, P. 1979. Specialized insectivory: beetle-eating and moth-eating molossid
bats. J. Mamm., 60:467-479.
. 1981. Correspondence of food habits and morphology in insectivorous bats.
J. Mamm., 62:166=173.
Gardner, A. L. 1977. Feeding habits. Pp. 293-350, in Biology of bats of the New
World family Phyllostomatidae. Part II (R. J. Baker, J. K. Jones, Jr., and D. C.
Carter, eds.), Spec. Publ. Mus., Texas Tech Univ., Lubbock, 364 pp.
Gershon, M. D. 1981. The enteric nervous system. Ann. Rev. Neurosci., 4:227-272.
Grossman, M. I. 1976. The gut as an endocrine organ— introductory review. J. En-
docrinol., 70:1-2.
Grube, D., and W. G. Forssmann. 1979. Morphology and function of entero-endo-
crine cells. Horm. Metab. Res., 1:589-606.
Hart, L. A. 1971. Structure of the gastric mucosa as related to feeding habits in selected
species of New World bats. Unpubl. Ph.D. dissert., Virginia Polytechnic Inst, and
State Univ., Blackburg, 65 pp.
Hersey, S. j., D. May, and D. Schyberg. 1983. Stimulation of pepsinogen release
from isolated gastric glands by cholecystokinin-like peptides. Amer. J. Physiol., 7:
G192-G197.
Hersey, S. J., S. H. Norris, and A. J. Gilbert. 1984. Celllular control of pepsinogen
secretion. Ann. Rev. Physiol., 46:393-402.
Hill, J. E., and J. D. Smith. 1984. Bats: a natural history. Univ. Texas Press, Austin,
243 pp.
Hood, C. S., and J. D. Smith. 1 982. Cladistic analysis of female reproductive histology
in phyllostomid bats. Syst. ZooL, 31:241=251.
Hokfelt, T., S. Efendic, C. Hellerstrom, O. Johansson, R. Luft, and A. Arimura.
1975. Cellular localization of somatostatin in endocrine-like cells and neurons of
the rat with special references to the A-cell of the pancreatic islets and to the hy-
pothalamus. Acta Endocrinol., Suppl., 200:5-41.
IsHiKAWA, K., M. Matoba, H. Tanaka, and H. Ono. 1985. Anatomical study of the
intestine of the insect-feeder bat, Myotis frater kaguae. J. Anat., 142:141-150.
Ito, S., and G. C. Schofield. 1978. Ultrastructural changes in mouse parietal cells
after high H+ secretion. Proc. Symp. Gastric Transport, Uppsala, 1977. Acta Physiol.
Scand., Special Suppl., pp. 25-34.
Kalt, M. R., and B, Tandler. 1971. A study of fixation of early amphibian embryos
for electron microscopy, J. Ultrastruct. Res., 36:633-645.
Kamiya, T., and P. Pirlot. 1975. Comparative gastric morphology of Old World bats.
1. In light microscopy. J. Mamm. Soc. Japan, 6:145-154.
Kolb, A. 1954. Biological Beobachtungen an Redermausen. Saugetiekundl. Mitt., 2:
15-26.
Konturek, S. j., j. Biernat, and N. Kweicien. 1975. Eflfect of glucagon on meal-
induced gastric secretion in man. Gastroenterol., 68:448-454.
Kusumoto, Y., T. Iwanaga, S. Ito, and T. Fusita. 1979. Juxtaposition of somatostatin
cell and parietal cell in the dog stomach. Arch. Histol. Japan, 42:459-465.
1986
Studholme et al.— Bat Gastric Mucosa
233
Larsson, L,-L, J. M. Polak, R. Buffa, F. Sundler, and E. Solcia. 1979. On the
immunocytochemical localization of the vasoactive intestinal polypeptide. J. His-
tochem. Cytochem., 27:936-938.
Larsson, L.-L, and J. F. Rehfeld. 1977. Evidence for a common evolutionary origin
of gastrin and cholecystokinin. Nature, 269:335-338.
Lichtenberger, L. M., L. A. Graziani, E. J. Dial, B. D. Butler, and B, A. Hills.
1983. Role of surface-active phospholipids in gastric cytoprotection. Science, 219:
1327^1329.
Mennone, a., C. j. Phillips, and D. E. Pumo. 1986. Evolutionary significance of
interspecific differences in gastrin-like immunoreactivity in the pylorus of phyllos-
tomid bats. J. Mamm., 67:373-384.
Miller, R. J. 1984. The mucosa as a target tissue for gut neuropeptides. Trends
Neurosciences, 7:2-3.
Moody, A. J., H. Jacobsen, and F. Sundby. 1978. Gastric glucagon and gut glucagon-
like immunoreactants. Pp. 369-378, in Gut hormones (S. R. Bloom, ed.), Churchhill
Livingstone, New York, 664 pp.
Ogunbiyi, O. a., and E. E. Okon. 1976. Studies on the digestive enzymes of the
African fruit bat, Eidolon helvum (Kerr). Comp. Biochem. Physiol., 55:359-361.
Ohara, S., M. Kakei, K. Ishihara, T. Katsuyama, and K. Hotta. 1984. Effects of
fasting on mucus glycoproteins in rat stomach. Comp. Biochem. Physiol., 79B:325-
329.
Okon, E. E. 1977. Functional anatomy of the alimentary canal in the fruit bat, Eidelon
helvum, and the insect bat, Tadarida nigeriae. Acta Zool. (Stockh.), 58:83--93.
Olender, E. j., D. Fromm, T. Furukawa, and M. Kolis. 1984. H+ disposal by rabbit
gastric mucosal surface cells. Gastroenterol., 86:698-705.
Ormsbee, H. S., Ill, AND J. D. Fondacaro. 1985. Action of serotonin on the gastroin-
testinal tract. Proc. Soc. Exper. Biol Med., 178:333-338.
Park, H., and E. R. Hall. 1951. The gross anatomy of the tongues and stomachs of
eight New World bats. Trans. Kansas Acad. Sci., 54:64-72.
Pearse, a. G. E. 1969. The cytochemistry and ultrastructure of polypeptide hormone-
producing cells of the APUD-series and the embryologic, physiologic and patho-
physiologic implication of the concept. J. Histochem. Cytochem., 17:303-313.
Phillips, C. J. 1971. The dentition of glossophagine bats: development, morphological
characteristics, variation, pathology, and evolution. Misc. Publ. Mus. Nat. Hist.,
Univ. Kansas, Lawrence, 54:1-138.
— — — . 1985. Field fixation and storage of museum tissue collections suitable for
electron microscopy. Acta Zool. Fenn., 170:87-90.
Phillips, C. J., and K. Studholme. 1982. Comparative ultrastructure of gastric chief
cells in carnivorous and frugivorous bats. IRCS Med. Sci., Cell Membrane Biol.,
10:573-574.
Phillips, C. J., and B. Tandler. 1985. Unique salivary gland structure in two genera
of tropical bats. Anat. Rec., 21 1:152A.
— — -. In press. Mammalian evolution at the cellular level. In Current mammalogy
(H. H. Genoways, ed.), Plenum Press, New York.
Phillips, C. J., G. W. Grimes, and G. L. Forman. 1977. Oral biology. Pp. 121-246,
in Biology of bats of the New World family Phyllostomatidae. Part II (R. J. Baker,
J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ., Mus., Texas Tech Univ.,
Lubbock, 364 pp.
Phillips, C. J., T. Nagato, and B. Tandler. In press. Comparative ultrastructure and
evolutionary patterns of acinar secretory product of parotid salivary glands in Neo-
tropical bats. In Studies in neotropical mammalogy: essays in honor of Philip Hersh-
kovitz (B. D. Patterson and R. M. Timm, eds.), Fieldiana: Zoology, new ser.
234
Annals of Carnegie Museum
VOL. 55
Phillips, C. J., K. M. Studholme, and G. L. Forman. 1984. Results of the Alcoa
Foundation Suriname expeditions. VIII. Comparative ultrastructure of gastric mu-
cosae in four genera of bats (Mammalia: Chiroptera), with comments on gastric
evolution. Ann. Carnegie Mus., 53:71-117.
Phillips, C. J., B. Tandler, and C. A. Pinkstaff. In press. Unique salivary glands in
two genera of tropical microchiropteran bats: an example of evolutionary conver-
gence in histology and histochemistry. J. Mamm.
Reynolds, E. S. 1963. The use of lead citrate at high pH as an electron opaque stain
in electron microscopy. J. Cell. Biol., 17:208-212.
Rieger, R., and S. Tyler. 1979. The homology theorem in ultrastructural research.
Amer. Zool., 19:655-664.
Robert, A., W. Bottcher, E. Golanska, and G. L. Kauffman, Jr. 1984. Lack of
correlation between mucus gel thickness and gastric cytoprotection in rats. Gas-
troenterol., 86:670-674.
Rouk, C. S., and B. P. Glass. 1970. Comparative gastric histology of American bats.
J. Mamm., 51:455-472.
Schofield, G. C., S. Jto, and R. P. Bolander. 1979. Changes in membrane surface
area in mouse parietal cells in relation to high levels of acid secretion. J. Anat., 128:
609-692.
Schultz, W. 1965. Studien iiber den Magen-Darm-Kanal der Chiropteren. Ein Beitrag
zum Problem der Homolgisierung von Abschnitten des Saugetierdams. Zeit. Wis-
senschaft. Zook, 171:241-391.
— . 1970. Einige Bemerkungen zum Bau des Verdauungstraktes und der sys-
tematischen Stellung des Spitzzahnflughundes, Harpionycteris whiteheadi Thomas,
1896 (Megachiroptera). Zeit. Saugetierkunde, 35:81-89.
ScHULTZBERG, M., T. HoKFELT, G. Nilsson, L. Terenius, J. F. Rehfeld, M. Brown,
R. Elde, M. Goldstein, and S. Said. 1980. Distribution of peptide- and cate-
cholamine-containing neurons in the gastro-intestinal tract of rat and guinea pig:
immunohistochemical studies with antisera to substance P, vasoactive intestinal
peptide, enkephalins, somatostatin, gastrin/cholecystokinin, neurotensin and do-
pamine /3-hydroxylase. Neurosci., 5:689-744.
Smith, J. D. 1976. Chiropteran evolution. Pp. 49-69, in Biology of bats of the New
World family Phyllostomatidae. Part I (R. J. Baker, J. K. Jones, Jr., and D. C. Carter,
eds.). Spec. Publ. Mus., Texas Tech Univ., Lubbock, 218 pp.
Smith, J. D. and G. Madkour. 1980. Penial morphology and the question of chirop-
teran phylogeny. Pp. 233-244, in Proceedings of the fifth international bat research
conference (D. E. Wilson and A. L. Gardner, eds.), Texas Tech Univ. Press, Lubbock,
434 pp.
SoLCiA, E., C. Capella, R. Buffa, L. Usellini, R. Fiocca, and F. Sessa. 1981. En-
docrine cells of the digestive system. Pp. 39-58, in Physiology of the gastrointestinal
tract (L. R. Johnson, ed.). Raven Press, New York, 1492 pp.
Spicer, S. S. 1963. Histochemical differentiation of mammalian mucopolysaccharides.
Ann. New York Acad. Sci., 106:379-388.
Stachura, j., a. Tarnawski, J. Bogdal, and K, Ivey. 1981. Effect of glucagon on
human gastric mucosa: histochemical studies. Gastroenterol., 80:474-481.
Studier, E. H., B. C. Boyd, A. T. Feldman, R. W. Dapson, and D. E. Wilson. 1983fl.
Kidney structure in neotropical bats. J. Mamm., 64:445-452.
— . 1983^. Renal function in the Neotropical bat, Artibeus jamaicensis. Comp.
Biochem. Physiol., 74A: 199-209.
Tandler, B., T. Nagato, and C. J. Phillips. 1986. Systematic implications of com-
parative ultrastructure of secretory acini in the submandibular salivary gland in
Artibeus (Chiroptera: Phyllostomidae). J. Mamm., 67:81-90.
Thomas, D. W. 1984. Fruit intake and energy budgets of frugivorous bats. Physiol.
ZooL, 57:457-467.
1986
Studholme et al.— Bat Gastric Mucosa
235
Toth, S. E. 1968. The origin of lipofuscin age pigments. Exper. Genontol., 3:19-30.
Tuttle, M. D., and M. J. Ryan. 1981. Bat predation and the evolution of frog
vocalizations in the neotropics. Science, 214:677-678.
Unger, R. H., R. E. Dobbs, and L. Orcl 1978. Insulin, glucagon, and somatostatin
secretion in the regulation of metabolism. Ann. Rev. Physiol., 40:307-343.
Yamada, J., V, J. M. Campos, N. Kitamura, A, C. Pacheco, T. Yamashita, and V.
Caramaschl 1984. Immunocytochemical study of gastro-entero-pancreatic (GEP)
endocrine cells in the vampire bat (Desmodus rotundus). Gegenbaurs morph. Jahrb.,
Ixipzig, 130:845--856.
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VOLUME 55 15 SEPTEMBER 1986 ARTICLE 9
AN INTERNATIONAL SURVEY OF THE POPULAR AND
TECHNICAL LITERATURE OF MAMMALOGY
Michael A. Mares ^
Research Associate, Section of Mammals
Janet K. Braun*
Abstract
A questionnaire designed to assess the diversity and abundance of the technical, semi-
technical, and popular literature of mammalogy was sent to curators in 1 6 1 mammal
collections and museums in 74 countries; 104 responses were received from 55 nations.
Results show a great disparity in the availability of the different types of literature among
countries. Some nations have a very strong literature at all levels, while many are in
great need of all types of literature. Countries that have a rich literature at the semi-
technical and popular levels also have a strong foundation in the basic technical literature
of mammalogy. A statistical analysis of various socioeconomic factors shows that per
capita income (PCI) is strongly positively related to the availability of literature at all
levels: increasing the PCI by $ 1 000 per year results in a doubling of the available literature
on mammalogy. The analysis suggests that only through international cooperative re-
search on basic mammalogy can the PCI block be bypassed and the stage set for an
increase in the popular literature of mammalogy.
An overview of the mammalogical literature is presented for each country queried.
Responses of the curators are also given. Almost all respondents indicated a willingness
to work in a cooperative manner with foreign scientists in producing semitechnical and
popular literature in mammalogy.
' Address: Stovall Museum and Department of Zoology, University of Oklahoma, Nor-
man, Oklahoma 73019.
Submitted 26 July 1984.
145
146
Annals of Carnegie Museum
VOL. 55
Introduction
Mammalogy is a diverse discipline, encompassing such areas as sys-
tematics, ecology, behavior, physiology, anatomy, and other fields of
specialization. The trend in recent years has been away from basic
systematic and survey research and toward more scientifically elegant
and popular experimental and laboratory-oriented work. We believe
that this is a very healthy and exciting pattern, particularly in the
developed nations of the world. As Mares (1982, 1985, in press) has
pointed out, one very important reason that modem theoretical/em-
pirical research on mammals can be performed in developed countries
is because the foundational research on mammal species was done
decades ago by scores of mammalogists who constructed a framework
upon which current research can be supported. It is no accident that
most of the important research in state-of-the-art mammalogy is being
performed by investigators living in countries that have a long and
noble history of basic faunal research. In many cases, the organisms
being examined by the current crop of new mammalogists have been
studied in one form or another for more than a century.
An unfortunate correlate of the trend toward experimental research
is the tendency by some investigators to view foundational systematic
and survey research as being somehow less scientific and less inherently
valuable than investigations that are dernier cri. This view obtains
because in developed countries there is little work left to be done at
the foundational level. Moreover, younger biologists, trained in the
United States for example, are often quite parochial in their views of
nature. This is not said in a condemnatory fashion, but is merely
observational. Most young mammalogists in developed countries have
only had dealings with very well-studied aspects of their natural en-
vironment. They have seldom been at a real loss for either taxonomic
or ecological information on any vertebrate species with which they
are familiar, and their view of nature reflects this familiarity. They
tend to think of nature as a well-studied entity, believing that most of
the foundational work was completed early in this century and viewing
the research problems that remain to be done as exciting tests of the-
oretical questions. We believe that these views are consistent with an
educational background in a country that is rich in the foundational
literature of mammalogy. We suggest that these sanguine views of basic
field biology are in error.
The literature of mammalogy is extremely diverse. Almost all coun-
tries of the world support one or more mammal collections or natural
history museums that often help to contribute to the basic literature
of the science of mammalogy (for example, Genoways and Schlitter,
1981; Hickman, 1981). Hickman (1981) surveyed the field guides that
1986
Mares and Braun —Mammalogy Literature
147
had been published to the mammal faunas of most countries of the
world. However, in many cases the guides or papers cited by Hickman
are quite old (that is, published in the mid-1800’s or early 1900’s),
and/or are quite difficult to obtain. More important, as Mares (1985,
in press) has argued, field guides and other popular literature owe their
existence to the foundational literature of mammalogy, the taxonomic
and geographic surveys that first clarify the faunal makeup of a country.
It was clear from Hickman (1981), and from our own work in various
countries, that there are pronounced differences in the availability of
the mammal literature of the world. Some countries seem to have a
particularly abundant literature, whereas others have, at best, a very
scant literature on mammals. Moreover, we had not seen any reports
examining the availability of both the popular literature on mammals
and the technical literature that provides its foundational material. If,
indeed, these two types of literature bear some relationship to each
other, we might expect that few countries lacking a strong basic liter-
ature will show a pronounced level of development of popular literature
on mammals.
Just exactly how extensive the foundational literature of mammalogy
might be is not clear. No one, to our knowledge, has attempted a
comprehensive assessment of the actual status of the world’s mam-
malogical literature. We feel that information on the availability of
such literature on a global scale will help point out potential areas
requiring increased research efforts. Such a survey might also indicate
that the world we inhabit is less biologically explored than many sur-
mise.
In this paper, we present results of a survey of the literature of
mammalogy that were obtained through the use of a questionnaire that
was sent to most of the mammal collections of the world. A preliminary
report on these data was given in Mares (1985); however, we herein
present a much more complete literature survey based on many more
responses than were included in the earlier report. These data present
an overview of the popular and technical literature of mammalogy for
many countries of the world. They include data from most countries
that have a mammal collection of greater than 50 specimens, and offer
some information on the availability of mammal literature for almost
all countries of the world that have a mammal collection of even modest
size. Taken together, these data give some indication of the current
status of the world’s mammal literature, including which countries have
reached a high level of literature availability. They also show which
countries need a cohesive plan of work to provide either the technical
or popular literature, or both, that are required if the country’s literature
is to be brought up to acceptable levels.
148
Annals of Carnegie Museum
VOL. 55
Materials and Methods
A questionnaire designed to assess the availability of various types of natural history
literature (primarily that dealing with mammals) in different countries of the world was
sent out in 1981. Respondents were asked to list available publications and indicate if
these publications were available in the common language of the country. The following
questions were used: 1) Is there a publication (in the style of a field guide) available that
deals with the identification of the mammals of your region? 2) Is there a publication
(technical survey) available to the museum specialist which is concerned with the mam-
mals of your region? 3) Are there taxonomic keys available to the mammals of your
region? 4) Are there any works available that deal with the collecting or preparation
techniques of mammals? 5) Are there any publications available that deal with the
operation or importance of natural history museums? 6) Would you judge the number
of available children’s books dealing with natural history topics to be many, few, or rare?
7) Please list three natural history or museum science subjects that you feel need attention
in your particular geographic area. 8) Please list any projects underway that deal with
any of the topics listed in the first seven questions. 9) Would you be interested in
cooperative research with other museum specialists in preparing publications such as
field guides, textbooks, etc., that deal with the fauna of your area? Names of museums,
research organizations, collections, and personnel (curators and directors) were obtained
from Genoways and Schlitter (1981). Museums or research organizations were queried
if: (1) the collections were of a substantial size, or (2) they were the only major museum
or collection in a particular country or region.
The countries queried are listed in alphabetical order and questionnaire responses are
indicated where applicable. Countries which were not surveyed or which did not respond
to the questionnaire are indicated by ‘Wo response received"" before the list of literature
citations. Responses given by the respondents are italicized. Additional literature cita-
tions, not given by respondents, were added to present a more complete overview of the
literature availability for the countries listed and are not italicized. Topics were eliminated
if the respondent(s) did not answer a question. Responses to research needed or projects
underway which did not directly pertain to mammals or museum science were omitted.
Under “Comments” we have either summarized as succinctly as possible, or reproduced
verbatim, informal comments of respondents. In some cases we have added additional
observations that we feel are particularly important to an understanding of the literature
situation in any given country. Willingness of the respondent(s) to work in cooperation
with specialists from other museums and institutions are denoted by WTC at the end
of the comments if more than one-half of the respondents of a particular country answered
positively. The absence of this designation indicates that a negative response was received
or that this question was not answered. A concerted effort was made to verify all references
given by the respondents. Those references which could not be verified and/or are partial
references are denoted by an asterisk following the reference number.
Hickman’s (1981) broad overview of field guides to national mammal faunas includes
many old (pre-1940) natural history publications. We have tried not to duplicate works
cited in Hickman (1981). Moreover, we have attempted to list only the more readily
available recent literature.
Countries responding to the questionnaire were placed into one of six groups based
upon the major geographic regions: Latin American, European, Middle Eastern, African,
Asian, and Australian. Summaries of responses to each question are presented separately
in tabular form (Tables 1-3).
Data on gross national product (GNP), per capita income estimates, population growth
rates, population density, literacy, and the percent of the population living in urban areas
were obtained from The Global 2000 Report to the President (Barney, 1982) and The
World Almanac (Lane, 1983).
1986
Mares and Braun —Mammalogy Literature
149
Results
Survey of Responses
Afghanistan: No response received; but see 26 347, 409, 429,
430.
Algeria: No response received; but see 344, 401.
Angola: No response received; but see 279, 401.
Argentina: Museo Municipal de Ciencias Naturales “Lorenzo Scag-
lia.” Guides: None; but see 447. Surveys: 496; see also 113, 114, 115,
189, 273, 372, 387, 391, 475. Keys: None; but see 272, Technical
manuals: 130*; see also 102. Museum science literature: None. Chil-
dren’s literature: Rare. Research needed: Paleontology. Projects un-
derway: None; also, Mares et aL, Mammals of Tucuman, Los Mami-
feros de Salta. Comments: The recent field guide by Olrog and Lucero
is already out of print, and was difficult to obtain even when it was
still in print. Greenhall has written a key to the bats of Argentina and
Pearson has written a key to the small mammals of Nahuel Huapi and
Lanin National Parks. Both of these are published or in press. Most
of the surveys published in Spanish are difficult to obtain, while those
published in English are either poorly disseminated among the scien-
tists in Argentina or are themselves difficult to obtain. Argentina il-
lustrates a problem that will be encountered time and again during this
survey, and that is the difficulty in obtaining publications that are
published in obscure journals or materials that are unpublished and
circulated among a few specialists. The literature of mammalogy in
Argentina is quite poor, particularly for native students of the Argentine
fauna.
Australia: Arthur Rylah Institute for Environmental Research;
Australian National Wildlife Collection; Central Australian Museum;
Monash University; National Museum of Victoria; Queensland Mu-
seum; Queen Victoria Museum and Art Gallery; South Australian Mu-
seum; Taronga Zoo; Tasmanian Museum and Art Gallery; The Aus-
tralian Museum. Guides: 29*, 218, 234, 300, 312, 313, 314, 389, 500,
663; see also 85, 662. Surveys: 13, 40, 83, 84*, 234, 300, 389, 459,
500, 662, 663; see also 41, 188, 215, 219, 238, 375, 576, 654, 676.
Keys: 12, 39, 144, 300, 306, 337, 663. TecMical manuals: There is no
publication which is specifically oriented in this direction. The Tas-
manian Museum and Art Gallery has a leaflet on collecting animals
for the museum. Works published in North America and Europe are
used. Museum science literature: 483* gives a historical account of the
beginnings of the National Museum of Victoria but does not cover the
last 25 years. Several journals are used including: Kalori, of the Mu-
seums Association of Australia, Museum, Museum News, Curator, and
Museum Journal; see also 533. Children’s literature: Few to many.
150
Annals of Carnegie Museum
VOL. 55
Research needed: Studies on marine mammals; ecology, ethological
and conservation studies on native vertebrates; animal population dy-
namics; handbooks and field keys; collection and preparation technique
manuals; publications dealing with ecological and behavioral tech-
niques for naturalists; museum science publications; field guide to ce-
taceans recorded in Australian waters; detailed surveys of the mammals
of particular geographic regions. Projects underway: The zoology de-
partment of the University of Tasmania is publishing a Fauna of Tas-
mania Series; two field guides to Australian mammals are in prepa-
ration, one by J. W. Calaby and one by M. Archer; B. Marlow is
preparing an identification manual (at a technical level) to all the ter-
restrial mammals of Australia. Comments: Several respondents indi-
cated that a large number of works are currently in preparation. Many
indicated that no comprehensive keys are available to Australian mam-
mals. Many referred to preparation/collection techniques manuals of
the British Museum (Natural History). No publications are available
which deal specifically with museum science and literature; some are
included as part of a treatment of other topics, however. WTC.
Austria: Naturhistorishes Museum Wien; Oberdsterreichisches
Landesmuseum. Guides: JO*, 75*, 90. Surveys: 65*, 411, 432, 670.
Keys: JO*, 75*, 432. Technical manuals: 471 and later editions, 161.
Museum science literature: None. Children’s literature: Few. Research
needed: A modern handbook on European mammals; a survey of Aus-
trian mammals; a semipopular identification manual. Projects under-
way: All of the above are in preparation. Comments: WTC.
Belgium: Institut Royal des Sciences Naturelles de Beligique; La-
boratorium voor Algemende Dierkunde. Guides: 94. Surveys: 192, this
publication is not up to date, and is without notes concerning distri-
butions’, see also 285. Keys: 575* only for Rodentia, Lagomorpha, and
Insectivora. Technical manuals: 285. Museum science literature: None.
Children’s literature: Many, but no original work— most are transla-
tions. Research needed: Mammal distributions; ecology of the Carni-
vora; studies on mammal protection. Projects underway: Mammal dis-
tributions. Comments: WTC.
Belize: No response received’, but see 231, 233, 273, 309, 336.
Bolivia: No response received’, but see 27, 28, 113, 114, 115, 273,
666, 695. Comments: Research is currently underway by Sydney An-
derson and various collaborators on the mammals of Bolivia. Literature
on the mammals of Bolivia is particularly depauperate.
Botswana: No response received’, but see 401, 557, 558.
Brazil: Museu de Ciencias Naturais; Museu Nacional; Museu Par-
aense “Emho Goeldi”; Universidade Estadual Paulista. Guides: 65i.
Surveys: 113, 114, 420, 641, 642’, see also 45, 46, 47, 115, 126, 273,
1986
Mares and Braun— Mammalogy Literature
151
388, 419, 466, 476, 550, 584, 640, 643. Keys: 420, 641, 642, 651; see
also 472. Technical manuals: A collection and preparation manual
written by the Departamento de Zoologia was published by the Mus.
Zool, Univ. Sdo Paulo in 1967 (152). Museum science literature: None.
Children’s literature: None. Research needed: Keys to the mammals of
southeastern Brazil. Projects underway: Chiroptera of southeastern
Brazil; Chiroptera of Mato Grosso, Brasil; chave para determinaqao de
quiropteros brasileiros (reformulation and actualization); guia para
identificaqdo de quiropteros do sul do Brasil (ndo existe verbas para
impressdo) (F. Silva). Comments: Research is currently underway on
mammals of the Cerrado by C. Alho and M. Mares. Brazil has an
extremely complex fauna of mammals and has only been cursorily
examined (474). One respondent, in referring to the availability of field
guides for Brazil, noted that a field guide was published long ago but
is not available today. This same respondent noted that many of the
publications on Brazilian mammals are no longer available, even to
specialists. WTC.
Burundi: No response received; but see 401, 639.
Cameroon: No response received; but see 164, 165, 401.
Central African Republic: No response received; but see 40 1 , 528.
Chile: Coleccion Particular de Fabian Jaksic y Jose Yafiez; Instituto
de Ecologia y Evolucion; Laboratorio de Citogenetica; Museo de Zoo-
logia de la Universidad de Concepcidn. Guides: 412, 454. Surveys:
383, 454; see also 113, 114, 115, 154, 220, 273, 475, 477, 588. Keys:
382, 497; see also 383, 454. Technical manuals: None, although 230
has been used in the past. Museum science literature: None; one re-
spondent reported There is something published by Museo Nacional de
Historia Natural, Casilla 787, Santiago, Chile. Another respondent
suggested 1 54 as a source of information on museum science literature;
see also 694. Children’s literature: None or rare. Research needed:
Ecology; behavior; distribution and zoogeography; taxonomic studies;
keys to Chilean mammals; field guide to Chilean mammals for country
and particular regions; literature for the general public; preparation/
collection technique manuals for the specialist and the general public.
Projects underway: Miscellaneous projects dealing with particular
species. Comments: A recent bibliography of references on terrestrial
Chilean mammals is given in 464. Additional information is given in
495. The research on Chilean mammals has increased significantly in
recent years, although it has not been reflected in available publications
on the topics of interest in this report. Mann’s guide to mammals of
Chile is difficult to obtain. WTC.
Colombia: Instituto de Ciencias Naturales; Museo del Instituto de
la Salle. Guides: None. Surveys: 62, 63, 268, 269, 270, 282*; see also
152
Annals of Carnegie Museum
VOL. 55
78, 113, 114, 115, 267, 273, 363. Keys: 62, 63. Technical manuals:
202*. Museum science literature: 159. Children’s literature: Few or
rare. Research needed: Natural history studies of mammals; systematic
studies on any of the various mammal groups (Rodentia, Ursidae, Cer-
copithecidae). Projects underway: None. Comments: Cabrera and Yepes,
as well as Cabrera’s systematic manual, are either extremely difficult
to obtain and very expensive (the former) or out of print entirely (the
latter). Many of the works cited for Colombia are old or are published
in English. WTC.
Congo: No response received; but see 401, 489.
Costa Rica: No response received; but see 21 1, 231, 233, 309, 310,
323, 626. Comments: The basic information to form a field guide to
the mammals of Costa Rica and, indeed, to produce many other types
of basic mammalogical materials is available in diverse publications
in English. Costa Rica’s fauna is well-studied when compared with the
fauna of most of Latin America. A key to the mammals of Costa Rica
was written by Hooper but is generally unavailable. A key to the bats
of Costa Rica (572) is also generally unavailable.
Czechoslovakia: Insititute of Systematic Zoology; Institute of Ver-
tebrate Zoology; National Museum; Zapadoceske Muzeum v Plzni.
Guides: 24*, 185, 196*, 245, 465. Surveys: 185, 239, 244, 245, 465.
Keys: 24*, 185, 245, 465. Technical manuals: 583*. Museum science
literature: None. Children’s literature: None to many. Research needed:
A handbook and key to Czechoslovakian mammals. Projects underway:
State research programs on nature conservation and on the fauna of
Czechoslovakia; handbook of Czechoslovakian mammals; key to
Czechoslovakian mammals. Comments: One response indicated that
a specialized museum science journal was available, ‘‘Musejni Prace.”
WTC.
Denmark: Natural History Museum; Zoological Museum of the
University. Guides: 373, 548. Surveys: 139, 432; see also 140. Keys:
Keys from a number of different sources and in a variety of languages
are available to the specialist. Non- specialists can use handbooks and
field guides for identification. Technical manuals: No special publica-
tion; techniques are given in a number of field guides for the general
public (youth, teachers, sportsmen). Museum science literature: None.
Children’s literature: Many. Projects underway: None. Comments: You
always want an up-to-date, comprehensive, and confident treatment of
the mammal region, but I think we have a fair picture of the Danish
mammal fauna and new information is added every year. The same
can be said for most countries in N. W. Europe. In referring to the
question on the availability of publications dealing with the operation
and importance of natural history museums the following comments
1986
Mares and Braun— Mammalogy Literature
153
were made: I can think of no special publications. Museums —including
natural history museums— are part of the cultural tradition and de-
pending on the actual financial situation in the country it may go up
and down for the museum. In one period you may pay special attention
to research, in the next to exhibition and other public relation (^sicj e.g.,
in relation to nature conservation and management. In reference to the
question dealing with collecting or preparation techniques manuals,
the following comments were made: No special publication but tech-
niques are mentioned in a number of field guides for youth, guides for
teachers in biology, guides for sportsmen, ‘do it yourself books, ' etc.
Generally, collecting and preparing a mammal should not be encour-
aged in countries in Western Europe and in many countries here mam-
mals not considered game or pest species are totally protected.
Ecuador: No response received', but see 14, 51, 113, 114, 115, 273,
667.
Egypt: Wassif s Collection. Guides: 453. Surveys: None', see also
229, 287, 288, 289, 290, 453, 657, 658, 660. Keys: 525, 536, 537, 538,
539, 540, 541, 542, 659', see also 401. Technical manuals: None. Mu-
seum science literature: None. Children’s literature: Few. Research
needed: [Mammals of] the eastern and western deserts of Egypt; the
Sinai. Projects underway: A study of the Western Desert of Egypt is
currently underway and is scheduled to be published by the Desert Re-
search Institute, Mataria, Cairo (in Arabic). Comments: WTC.
El Salvador: Museo de Historia Natural de El Salvador. Guides:
None. Surveys: 106 (currently unavailable), 181', see also 177, 178, 179,
180, 182, 231, 233, 309. Keys: None', see also 231, 233. Technical
manuals: 25. Museum science literature: None. Children’s literature:
None. Projects underway: Mammals of the Monte Cristo National Park
by Jim Hartman (in preparation); New Bats for El Salvador by V.
Hellebuyck (in preparation). Comments: There was also a comment
indicating that publications dealing with preparation techniques of
mammals are available at the museum library for these specialists. It
was noted that these were not available for the general public. WTC.
England: Booth Museum of Natural History; British Museum (Nat-
ural History). Guides: 93, 136, 137, 139, 141, 142, 559; see also 392.
Surveys: 136, 137, 139, 140, 141, 142, 359', see also 42. Keys: 136,
137, 139, 359; a specialist key is available from Mrs. J. Coy, Depart-
ment of Environment, Faunal Remains Project, 63 University Road,
Southampton S09 5NH England. Technical manuals: 31, 379, 618*,
652’, see also 138. Museum science literature: Various British Museum
(Natural History) publications. Children’s literature: Many. Research
needed: A list of named collections, their content and location; cura-
torial codes and practices; a national inventory of collections. Projects
154
Annals of Carnegie Museum
VOL. 55
underway: A Manual of Curatorship and a Code of Ethics are being
developed by Museums Association, a museums professionals group;
one or two keys to skeletal material are available from specialist mu-
seums groups in England.
Equatorial Guinea: No response received', but see 380, 40 L
Ethiopia: No response received', but see 143, 280, 355, 401, 690,
691, 692, 693.
Federal Republic of Germany: Coll. Pieper/Kiel; Landessamm-
lungen fur Naturkunde; Staatliches Museum fur Naturkunde Stuttgart;
Zoologische Staatssammlung Munchen; Zoologisches Forschungsinsti-
tut und Museum Alexander Koenig. Guides: iO*, 75*, 92, 195, 325*,
341. Surveys: 325* (for Bavaria), 432. Keys: 30*, 75*, 432. Technical
manuals: 148, 149, 471 (and later editions), some information given in
field guides. Museum science literature: 527 and several papers in Natur
und Museum. Children’s literature: Many. Research needed: Detailed
distribution maps; studies of post-cranial osteology; survey of the mam-
mals of Germany; a history of museum collections; type catalogues.
Projects underway: None. Comments: One respondent noted that there
are several small papers dealing with the collection and preparation
techniques of mammals that were issued for the public but are of
questionable value. WTC.
Finland: Zoological Museum, University of Helsinki; Zoological
Museum, University of Oulu. Guides: 547, 549. Surveys: 547, 549,
552. Keys: 549. Technical manuals: 549, 552. Museum science liter-
ature: None. Children’s literature: Few to many. Research needed: Zoo-
geography of the northern parts of Norway, Sweden, Finland, and So-
viet Union; history of the distribution of mammals after the Ice Age;
microtaxonomy of mammals today. Projects underway: None. Com-
ments: One respondent noted that most children’s literature is trans-
lated from other languages. WTC.
France: Laboratoire de Zoologie, Mammiferes et Oiseaux; Musee
Zoologique. Guides: 89, 519. Surveys: 519', see also 362, 506. Keys:
Response affirmative, but none were listed. Technical manuals: 153,
356*. Museum science literature: 80*, 214. Children’s literature: Many.
Research needed: Importance of natural history museums; an atlas of
the fauna of France; inventory of the fauna of parks and reserves; an
altitudinal distribution of the micromammals in the Vosges Mountain
area; a precise list of distributions of the Chiroptera; status of a number
of “pest” (hamster) and introduced species (raccoon, raccoon dog, nu-
tria). Projects underway: Atlas of the distribution of mammals in France
(by the Societe Francaise pour IFtude et la Protection des Mammiferes,
S.F.E.P.M.). Comments: WTC.
French Guiana: No response received', but see 99, 113, 114, 115,
273, 668.
1986
Mares and Braun —Mammalogy Literature
155
German Democratic Republic: Staatliches Museum fiir Tier-
kunde Dresden; Zoologie der Sektion Biowissenschaften der Martin-
Luther-Universitat. Guides: JO*, 75*, 194, 195, 707. Surveys: 92, 664,
665; see also 432. Keys: 30*, 75*, 92, 194, 195, 707, Technical man-
uals: 471 (and later editions). Museum science literature: None. Chil-
dren’s literature: Many. Research needed: Operation of natural history
museums. Projects underway: None.
Ghana: University of Ghana. Guides: 77, 155, 247. Surveys: Nu-
merous publications but no single one; see also 401. Keys: 401. Tech-
nical manuals: Not generally available. Museum science literature: None.
Children’s literature: Few. Research needed: Guide to Ghanaian mam-
mals; books on ecology and scientific method. Projects underway: None.
Comments: There is no lingua franca in Ghana except perhaps English
which is the language of literate people; few books are published in
Ghanaian languages. Books are not generally available in Ghana an-
yway, especially under the present foreign exchange crisis. The respon-
dent noted further that some books that could be used as children’s
books become available periodically. However, the supply of these ma-
terials is extremely erratic. The respondent noted that there is only a
single copy of Meester and Setzer (197 1-1977) available in the entire
country and this new copy is incomplete. WTC.
Greece: No response received; but see 431, 449, 450, 706.
Guatemala: No response received; but see 210, 231, 233, 273, 309,
317, 585.
Guyana: No response received; but see 113, 114, 115, 243, 273.
Honduras: No response received; but see 68, 197, 273.
Hong Kong: No response received; but see 509.
Iceland: Nattwrufraedistofun islands. Guides: None. Surveys: 162*,
223*, 517, 518, 612, 627*. Keys: None. Technical manuals: None.
Museum science literature: None. Children’s literature: Rare. Research
needed: Natural history of whales and seals; natural history museum
publications. Projects underway: None.
India: Central Arid Zone Research Institute; National Zoological
Collection of India. Guides: None. Surveys: 60, 72, 167, 187, 381, 478,
479; see also 96, 97, 98, 169, 328, 351, 482, 573, 574. Keys: 72, 166,
168, 478, 479, 681, 682, 683, 684, 685, 686, 687; see also 163. Tech-
nical manuals: 71, 187. Museum science literature: None. Children’s
literature: None to rare. Research needed: A checklist of mammals; a
field guide to the mammals of India; faunal works. Projects underway:
Faunal work on the groups not covered by Pocock and Ellerman; an
up-to-date checklist of Indian mammals is being prepared by the Zoo-
logical Survey of India. Comments: 128 gives a review of research
literature. First International Workshop on Management of Zoological
Collections: Recent mammal collections in tropical environments
156
Annals of Carnegie Museum
VOL. 55
sponsored by the Zoologcal Survey of India, Calcutta, and the Carnegie
Museum of Natural History, Pittsburgh, USA, was held in Calcutta
from 19-25 January 1984. The results of this workshop, which dealt
almost entirely with museum topics, will be published in the near
future. Two recent publications (423, 6 1 3) are available, which concern
endangered and threatened animals of India. WTC.
Indonesia: Museum Zoologicum Bogoriense. Guides: 399 (for Bor-
neo). Surveys: 131, 357. Keys: 399. Technical manuals: Use primarily
publications of the British Museum (Natural History), American Mu-
seum of Natural History, and the Museum of Berlin. Museum science
literature: None. Children’s literature: Few. Research needed: Taxon-
omy and other information on the biology of mammals; research on
methods of small pest control; research on zoonosis. Projects underway:
Inventory studies of mammals in Indonesia; studies on the biology of
murid rodents as pests on agricultural crops. Comments: The respon-
dent noted that there is no formal publication on mammal surveys for
the Indonesian Archipelago other than for the island of Borneo. A
bibliography of the mammals of Southeast Asia is given in 3 1 5. WTC.
Iran: No response received', but see 100, 147, 360, 414.
Iraq: Natural History Research Centre and Museum. Field guides:
None. Surveys: 139, 169, 253, 254, 255, 262', see also 19, 140, 251,
426, 522. Keys: 253, 254, 255. Technical manuals: 255. Museum sci-
ence literature: None. Children’s literature: Few. Research needed: Field
guide to Iraq's mammals; wildlife of Iraq including modern techniques.
Projects underway: None.
Ireland: National Museum of Ireland, Dublin; Trinity College,
Dublin. Guides: 88, 136, 435*. Surveys: 175, 176, 418; see also 139,
140. Keys: 136, 422, 568. Technical manuals: 31, 618*, 652. Museum
science literature: None, but a publication by the Irish Museums Trust
was due to be released in 1982. Children’s literature: Few to many.
Research needed: Museum maintenance; special techniques manuals;
design and presentation of museum material; simple illustrated litera-
ture dealing with the common mammals; guides to specific areas of
interest in the Natural History Museum. Projects underway: A book on
extant Irish animals has recently been published by the Natural History
Museum; a book on the history of the Natural History Museum is
expected in the near future. Comments: A book on Irish wild mammals
was published as part of the Folens Irish Environmental Library Series.
WTC.
Israel: No response received; but see 10, 11, 43, 44, 74.
Italy: Collezione Microteriologica di Longino Cantoli; Museo Civi-
co di Storia Naturale; Museo Zoologico de “La Specola.” Guides: 90,
699. Surveys: 619, 620; see also 117, 120. Keys: 619, 620, 699. Tech-
nical manuals: 619, 620, 698*. Museum science literature: None; al-
1986
Mares and Braun— Mammalogy Literature
157
though some visitors guides are available for a number of museums;
see also 118, 119. Children’s literature: None to many. Research need-
ed: Literature dealing with training people for collecting, preparing, and
maintenance of natural history collections; publications concerned with
the importance of natural history museums. Projects underway: Soric-
Mae deir ambiente bioclimatico mediterraneo (in press, Contoli); Glir-
Mae, Arvicolidae, Muridae delL ambiente bioclimatico mediterraneo (in
press, Amori, Contoli, and Cristaldi), Comments: One respondent not-
ed that some Italian specialists have field guides currently in press to
Italy’s most important mammals. Comments from another respondent
indicated that survey books having beautiful illustrations that can be
used to explain mammalogy to local and national authorities as well
as to common people are vitally needed in Italy, as are books that
relate nature to the cultural and moral formation of children and to
the general public. Moreover, the respondent felt that books dealing
with collecting and preparation techniques are especially vital, as are
materials that deal with the exhibition of mammalogical material.
Jamaica: Institute of Jamaica, Guides: None. Surveys: None; see
also 231, 233, 27 L Keys: 473; see also 231, 233. Technical manuals:
None. Children’s literature: Few. Research needed: Study on the Ja-
maican cony, Geocapromys brownii; study of the manatee; natural
history study of the mongoose. Projects underway: Studies on the man-
atee and cony (Natural Resources and Conservation Division, Dr. Pat-
rick Fairbairn).
Japan: Department of Oral Anatomy; Laboratory of Wildlife Re-
source Ecology; Natural History Museum; National Science Museum.
Guides: 303, 304. Surveys: 1, 303, 304, 349, 616*; see also 2, 448,
656, 696. Keys: 1, 303, 350*, 616*. Technical manuals: 4*, 303. Mu-
seum science literature: None. Children’s literature: Few to many. Re-
search needed: Complete collections of representative animal faunas;
zoogeographic studies; field guides and taxonomic keys to the mammal
fauna of the Japanese islands. Projects underway: None. Comments:
WTC.
Kenya: National Museum of Kenya. Guides: 151, 155; see also 135.
Surveys: 17, 329, 330, 331, 332, 333; see also 8, 252, 334, 335. Keys:
148, 401. Technical manuals: Field manual for museums (UNESCO).
Museum science literature: None. Children’s literature: Few. Research
needed: Conservation of flora and fauna; ecology and population dy-
namics studies of mammals; general natural history studies of native
species. Projects underway: None. Comments: WTC.
Korea: Natural History Museum. Guides: 680. Surveys: 679*; see
also 318, 319, 425. Keys: 679*. Technical manuals; None. Museum
science literature: None. Children’s literature: Rare. Research needed:
Field guide geared toward the general public and for education; a
158
Annals of Carnegie Museum
VOL. 55
monograph on the mammals of Korea; guide books in English, Projects
underway: None.
Lebanon: No response received; but see 43, 44, 366, 367.
Liberia: No response received; but see 346, 401.
Libya: No response received; but see 240, 293*, 401, 415, 486, 494,
535.
Madagascar: No response received; but see 236, 469, 570*, 607.
Malaysia: Sarawak Museum. Guides: 54, 397; see also 399. Sur-
veys: 54, 397; see also 55, 56, 146, 260, 396*, 398, 399, 400, 521, 624.
Keys: 396*. Technical manuals: None. Museum science literature: None.
Children’s literature: Few. Research needed: Field guide to the mam-
mals of Malaysia; collection/ preparation technique manuals; museum
science literature. Projects underway: None. Comments: An annotated
bibliography of the mammals of Malaysia (116) and a bibliography of
the land mammals of Southeast Asia (315) are available. WTC.
Mexico: Escuela Nacional de Ciencias Biologicas; Universidad Na-
cional Autdnoma de Mexico. Guides: 365, 645, 647. Surveys: 21, 26,
145, 212, 231, 232, 233, 292, 644; see also 22, 23, 49, 52, 105, 125,
127, 201, 203, 273, 320, 321, 322, 323, 326, 364, 438, 491, 492, 529,
625, 661, 678. Keys: 231, 233, 648. Technical manuals: 202*, 646,
Folleto de Divulgacidn, Instituto de Biologia, varias ediciones en Es-
panol. Museum science literature; None; but see 50, 493. Children’s
literature: Few to many. Research needed: Publications explaining the
importance of scientific systematic collections; a brief history of the
“Coleccion de mastozoologia del Instituto de Biologia, U.N.A.M. ”; lo-
cations of natural history specimens from Mexico that are housed out-
side of the country. Projects underway: Mamiferos de Mexico, in prep-
aration by Villa- R. Comments: WTC.
Mozambique: No response received; but see 191, 563.
Morocco: Museum de I’lnstitut Scientifique. Guides: 456, 457, 520.
Surveys: None; but see 95, 401. Keys: None. Technical manuals: None.
Museum science literature: None. Children’s literature: Rare. Projects
underway: None.
Namibia: State Museum. Guides: None. Surveys: 401, 544; see also
133, 286, 324, 507, 581. Keys: 170, 401, 503; see also 134. Technical
manuals: None. Museum science literature: None. Children’s literature:
None to few. Research needed: Semi- popular literature on natural his-
tory and conservation of mammals; comprehensive study on the mam-
mals of Namibia; field guides to small mammals in national parks;
semi-popular literature in non-English languages. Projects underway:
The mammals of Namibia (by C. G. Coetzee); check-list of mammals
of Etosha National Park (by J. E. W. Dixon); taxonomic study of small
mammals of southwest Africa/ Namibia (by the State Museum). Com-
ments: One respondent pointed out the need for children’s books and
1986
Mares and Braun— -Mammalogy Literature
159
Other popular material in the Ovambo language and other Bantu lan-
guages that are commonly used in Namibia. WTC.
Nepal: No response received', but see 3, 217, 311, 401, 416, 417,
545, 577, 669.
New Zealand: Auckland Institute and Museum; Canterbury Mu-
seum; National Museum of New Zealand. Guides: 48, 199, 200, 480,
504*, 677. Surveys: 48, 199, 200, 208, 677. Keys: 200 (key to marine
mammals). Technical manuals: None; only general works on taxider-
my; nothing specializing in mammals. Museum science literature: 610*;
we subscribe to English and American museum journals such as Cu-
rator and Natural History. Children’s literature: Many. Research need-
ed: Cetaceans; a guide to the fauna and flora of marine reserves.
Nicaragua: No response received', but see 53, 231, 233, 273, 309.
Niger: No response received', but see 340, 401.
Nigeria: D.C.D. Happold Collection of Mammals. Guides: None.
Surveys: 510, 511, 512, 513', see also 69, 246, 248, 249, 250, 299, 401.
Keys: 511, 512, 513 (for selected species). Technical manuals: None.
Museum science literature: None. Children’s literature: None to rare.
Research needed: Field guides at all levels for the vertebrates of Nigeria;
conservation literature; general ecological principles in the tropics. Proj-
ects underway: Mammals of Nigeria by D.C.D. Happold (in prepara-
tion, about 400 pp., Oxford University Press).
Northern Ireland: Ulster Museum and Botanic Gardens. Guides:
88, 142. Surveys: 139', see also 140. Keys: 136, 137. Technical manuals:
379, 480*, 652. Museum science literature: None specifically for North-
ern Ireland, although three were listed for England. Children’s litera-
ture: Many. Research needed: Archaeozoology and the history of the
Irish fauna; osteology in natural history museums; a catalogue of the
mammal collections in Britain and Ireland. Projects underway: None.
Comments: WTC.
Norway: No response received', but see 283, 284,
Oman: No response received; but see 257,
Pakistan: No response received; but see 505, 546.
Panama: Museo de Ciencias Naturales. Guides: 171, 209, 233, 402.
Surveys: None; but see 231, 233, 241, 273, 309. Keys: 402; see also
231, 233. Technical manuals: None. Museum science literature: None.
Children’s literature: None. Research needed: Guides to preparation
techniques; children’s natural history books; keys to the mammals of
Panama. Projects underway: None. Comments: WTC.
Papua New Guinea: Papua New Guinea National Museum and Art
Gallery; University of Papua New Guinea. Guides: 103, 104, 275, 357,
406, 500, 590, 598, 601, 634, 655, 701, 703; see also 702. Surveys:
121, 275, 276, 277, 294, 357, 368, 369, 378, 393, 394, 395, 403, 404*,
405, 407, 499, 521, 590, 591, 592, 593, 594, 595, 596, 597, 598, 599,
160
Annals of Carnegie Museum
VOL. 55
600, 601, 602, 603, 604, 605, 606, 628, 629, 630, 632, 633, 634, 703,
704; see also 160, 207, 631, 653. Keys: 295, 370, 404*, 702, 705.
Technical manuals: 652. Museum science literature: None. Children’s
literature: Few to none. Research needed: Handbook to New Guinea
marsupials and monotremes; a guide to the collection and preservation
of vertebrates: conservation of natural history specimens in the tropics;
the role of museums in community education; the role of museum col-
lections in scientific research. Projects underway: None. Comments:
WTC.
Paraguay: No response received; but see 70, 113, 114, 115, 273,
424, 671.
People’s Republic of China: Department of Vertebrate Taxonomy
and Faunology. Field guides: Surveys: 132; see also 16, 18, 183, 498,
689, 700, 708. Keys: None. Technical manuals: None. Museum science
literature: None. Children’s literature: Few. Research needed: An up-
dated survey of the fauna of China. Projects underway: A synopsis of
Chinese mammals with distributional maps. Comments: WTC.
Peru: No response received; but see 113, 114, 115, 273, 343, 460,
461, 462, 463, 565, 566, 567, 609, 621, 623.
Philippines: No response received; but see 15, 59, 263, 315, 358,
487, 488, 523, 530, 608, 614.
Poland: Mammals Research Institute. Guides: 484. Surveys: 348*,
434*, 55i*. Keys: 348*, 434*, 553*. Technical manuals: 433. Museum
science literature: Przeglad Zoologiczny, the journal of the Polish Zoo-
logical Society contains articles on this subject. Children’s literature:
Many. Research needed: None. Projects underway: None. Comments:
WTC.
Portugal: Museu Bocage; Museu e Laboratorio Zoologico. Guides:
20, 150, 455. Surveys: 139; see also 140, 172, 377. Keys: 112, 150.
Technical manuals: 652. Museum science literature: Two publications
by Almaca and one by Sacarrdo are available; the journal Museum
published by UNESCO is available. Children’s literature: Rare to few.
Research needed: Evolution; mammalogy; zoogeography. Projects un-
derway: None. Comments: One respondent noted that the Museum of
Natural History was completely destroyed by fire in March 1978 with
no specimens surviving the conflagration, WTC.
Republica Dominicana: Museo Nacional de Historia Natural.
Guides: None. Surveys: 233, 636; see also 231. Keys: 231, 473, 551,
577; see also 233. Technical manuals: None. Museum science literature:
None. Children’s literature: Few. Research needed: Ecology of the ver-
tebrate fauna of the West Indies and the Caribbean; natural history,
status and evolution of West Indian mammals. Projects underway:
MurciNagos de la Republica Dominicana (J. A. Ottenwalder); status y
explotacion del manati en la Republica Dominicana; habitat preference
1986
Mares and Braun— Mammalogy Literature
161
of the hutia (Plagiodontia aedium); natural history and systematics of
Solenodon; marine mammals of the coast of the Dominican Republic;
evolution and natural history of Capromyidae and fossil mammals (by
C. A. Woods).
.Republic of South Africa: Albany Museum; Department of Zo-
ology ^ University of Cape Town; Jonkershoek Nature Conservation
Station; Kaffrarian Museum; Transvaal Museum. Guides: 35, 155,
216, 503, 558. Surveys: 170, 401, 503, 544, 562; see also 307, 514,
515. Keys: 170, 401, 503, 562. Technical manuals: Minor publications
are available. Museum science literature: Publications available, es-
pecially those published in SAMAB; see also 675. Children’s literature:
Rare to many. Research needed: Systematic and zoogeographic studies;
multi-lingual childrens literature on conservation and general natural
history; techniques books on collection and preservation; a field guide
to mammals of South Africa; general natural history publications aimed
at the general public. Projects underway: Mammals of the Cape Prov-
ince by Swanepoel; Mammals of Transvaal by Rautenbach was pub-
lished in 1982; The Wildlife Society of Southern Africa is preparing a
field guide to the eastern Cape Coast; Mammals of the Orange Free
State by Lynch was published in 1983; Mammals of Natal by Pringle.
Comments: WTC.
Romania: “Grigore Antipa” Museum of Natural History. Guides:
88. Surveys: 122, 481. Keys: 237*, 305. Technical manuals: 428*, 458*.
Museum science literature: 157, 467*; three pre- 193 5 publications were
written by Antipa on the importance of museums. Children’s literature:
Few; for a summary of available literature see 157. Research needed:
Guide to the small mammals of Europe; establishment of general reg-
ulations for organizing and maintaining collections of small mammals;
a manual which centralizes many of the methods (morphological, cy-
togenetic, etc.) used in the systematics of small mammals. Projects
underway: An illustrated guide to the fauna of Romania. Comments:
WTC.
Saudi Arabia: No response received; but see 111, 253, 254, 255,
256, 427.
Scotland: Aberdeen University; The Royal Scottish Museum.
Guides: 108, 138, 141, 142, 359. Surveys: 142, 501. Keys: 108, 138,
139, 141, 142, 359. Technical manuals: 31, 64, 568, 652. Children’s
literature: Many. Research needed: A guide to the Wildlife Protection
Act of 1981; guides to the vertebrates of the eastern Mediterranean
region. Projects underway: Conference is being held to discuss the im-
plications of the Act of Parliament; the lack of field guides is known.
Comments: WTC.
Senegal: No response received; but see 76, 158, 291, 401.
Singapore: Zoological Reference Collection. Guides: 258, 400, 624.
162
Annals of Carnegie Museum
VOL. 55
Surveys: iii; see also 398. Keys: 258, 259*, 400. Technical manuals:
None. Museum science literature: None. Children’s literature: None.
Projects underway: None. Comments: WTC.
Somali Republic: No response received', but see 193, 401.
Spain: Museu de Zoologia de Barcelona. Guides: 91*, 109*, 519.
Surveys: 432', see also 638. Keys: 129, 436*. Technical manuals: 9.
Museum science literature: None. Children’s literature. Few. Projects
underway: None. Comments: WTC.
Sri Lanka: No response received', but see 163, 166, 167, 278, 470.
Sudan: No response received', but see 339, 342, 376, 401, 534.
Suriname: No response received', but see 113, 114, 115, 205, 206,
273, 296, 297, 298, 672, 673.
Switzerland: Naturhistorisches Museum Basel; Zoologisches Mu-
seum des Universitat Zurich. Guides: 66, 92. Surveys: 66, 432, 490.
Keys: 30*, 66, 75*, 92. Technical manuals: None printed in Switzer-
land, but several are available from Germany. Museum science liter-
ature: Birkhdusar- Verlag published the “Raritdten and curiositdten der
natur, sammlungen der naturhistorischen museum Basel” in 1980.
Children’s literature: Few to many. Research needed: Natural history
of Swiss mammals; importance of natural history museums; survey on
distribution of mammals in Switzerland; a museum guide for children.
Projects underway: A children's guide is in preparation by the Natur-
historisches Museum. Comments: WTC.
Taiwan: No response received', but see 308*, 316.
Tanzania: College of African Wildlife Management; The Serengeti
Research Institute. Guides: 155, 401. Surveys: None', but see 235, 582.
Keys: None; keys are available to skulls of some selected mammal
groups; some materials available on identification on mammalian teeth.
Technical manuals: None. Museum science literature: None. Children’s
literature: None to few. Research needed: Keys to the mammals of the
region; preparation manuals; guides to the importance of natural his-
tory museum. Projects underway: Studies on the ecology and behavior
of the lion, cheetah, elephant, and mongoose. Comments: WTC.
Thailand: Division of Environmental Biology. Guides: 361. Sur-
veys: None', but see 101, 281. Keys: 361. Technical manuals: A prep-
aration manual in Thai was published by the TISTR. Museum science
literature: None', see also 611. Children’s literature: Rare. Research
needed: Mammals of southeast Asia; revisions of many southeast Asian
bats and rodents. Projects underway: None. Comments: A bibliography
of the land mammals of Southeast Asia (3 1 5) is available. WTC.
The Netherlands: Rijkamuseum van Natuurlijke Histoire; Zoo-
logical Museum. Guides: 87, 107*, 302*, 374*. Surveys: 301, 432,
519', see also 86, 285, 485. Keys: 137, 139, 301, 432, 519. Technical
manuals: 285. Museum science literature: None. Children’s literature:
1986
Mares and Braun —Mammalogy Literature
163
Many. Research needed: Key to the identification of mammals in owl
pellets; a distribution atlas of mammals; a modern and popular hand-
book on mammals. Projects underway: An owl pellet key is in prepa-
ration at the Rijksmuseum van Natuurlijke Historic, Leiden; an atlas
project has just begun and will result from the collaboration of a large
number of institutes, scientists, and amateurs. Comments: WTC.
Trinidad and Tobago: No response received; but see 124, 213, 231,
233, 502.
Turkey: No response received; but see 173*, 174*, 421, 451, 452.
Uruguay: Museo Nacional de Historia Natural de Montevideo.
Guides: None; see also 186, 586. Surveys: 353, 688; see also 113, 114,
115, 273. Keys: None; but see 354. Technical manuals: None. Museum
science literature: A few articles have been published in the Boletin del
Museo Nacional de Historia Natural since 1973. Children’s literature:
None. Research needed: Ecology; ethology; biogeography. Projects un-
derway: Catdlogo sistemdtico de los vertebrados fdsiles sudamericanos
by A. Mones. Comments: WTC.
USSR: Zoologica Museum of Moscow University. Guides: Publi-
cations available, but none listed. Surveys: Publications available, but
none listed; see 5, 6, 7, 58, 190, 221, 222, 224, 225, 226, 227, 228,
264, 265, 266, 338, 410, 437, 440, 441, 442, 443, 444, 445, 446, 554,
555, 579, 580, 589, 615, 617, 637, 650. Keys: Publications available,
but none listed; see 57, 73, 543, 578, 649, 697. Technical manuals:
None. Museum science literature: None. Children’s literature: Many.
Venezuela: No response received; but see 110, 113, 114, 115, 242,
273, 371, 439, 508, 526, 587. Comments: Although we received no
response to the questionnaire, there are several types of ecological and
systematic projects in mammalogy that are currently underway in the
country. A key to the mammals of Venezuela was written by Handley
but is generally unavailable.
Vietnam: Laboratory of Zoology. Guides: 635. Surveys: 81, 82; see
also 123, 516. Keys: 635. Technical manuals: 153. Museum science
literature: None. Children’s literature: Few. Projects underway: The
rodents of North Vietnam (in French) is in preparation; Key to the
mammals of Vietnam by Tien is in preparation and will be published
in Vietnamese. Comments: A bibliography of the land mammals of
Southeast Asia (315) is available. WTC.
Yemen: No response given; but see 524.
Yugoslavia: Zoology Department Collection and Dr. Dulic’s Col-
lection. Guides: 198*. Surveys: 156, 413*; see also 345, 468. Keys:
390, 413*. Technical manuals: Only booklets and papers dealing with
preparation of animals in general including also the mammals. One
very old one published in 1948 is a small introduction in preparation
techniques in general. Author of this booklet is P. Alinger. Museum
164
Annals of Carnegie Museum
VOL. 55
Table I. — The availability of field guides (fg), surveys (s), keys (k), and museum science
literature (msl) within each geographic region. Percentages are given for each geographic
region and are based on the total number of responses received from each region.
Response
Latin
Ameri-
can
Euro-
pean
African
Middle
East
Asian
Aus-
tralian
Available in the common
fg
17%
90%
50%
62%
87%
language
s
50%
86%
78%
62%
69%
k
39%
83%
64%
69%
81%
msl
17%
31%
7%
8%
25%
Available, but not in the
fg
11%
5%
21%
50%
8%
common language
s
28%
7%
15%
50%
23%
k
11%
2%
7%
100%
23%
msl
5%
3%
7%
8%
Unavailable
fg
72%
5%
29%
50%
30%
13%
s
22%
7%
7%
50%
15%
25%
k
59%
14%
29%
8%
19%
msl
78%
64%
86%
100%
84%
75%
No response
fg
s
k
6%
msl
2%
science literature: None. Children’s literature: Few. Research needed:
A good and comprehensive book on the mammalian fauna; compiled
lists with distributional data; small monographs for different species or
groups of species. Projects underway: Investigation of specific Yugo-
slavian mammal species. Comments: The need for a manual on mam-
mal preparation techniques was emphasized. WTC.
Zaire: Koninklijk Museum voor Midden- Afrika, Musee Royal de
I’Afrique Centrale, Belgium. Guides: 155, 236. Surveys: 531, 532; see
also 67, 79*. Keys: 401. Technical manuals: None. Museum science
literature: 204; a list of West European museums with important col-
lections of African small mammals was given in the supplement to the
African Small Mammal Newsletter in July 1981. Children’s literature:
Rare. Research needed: Importance of conservation of tropical forests;
field guides and general natural history information of small mammals;
conservation of mammals. Projects underway: None.
Zambia; No response received; but see 37, 38, 401.
Zimbabwe: Museum of Zoology. Guides: 327 , 556, 559, 674. Sur-
veys: None; but see 35, 36, 401, 561*, 563, 564, 674. Keys: 556, 559;
see also 184. Technical manuals: None; but see 33, 34, 560. Museum
science literature: None; but see 32. Children’s literature: Few. Research
needed: Translation of available literature into non-English languages;
educational materials. Projects underway: None.
1986
Mares and Braun— Mammalogy Literature
165
Table 2. — The availability of mammal collecting and preparation techniques manuals
within each major geographic region. Percentages are given for each geographic region
and are based on the total number of responses received from each region.
Latin
Ameri-
Euro-
Middle
Aus-
Response
can
pean
African
East
Asian
tralian
Available to the specialist in the
common language
Available to the general public in
25%
50%
27%
29%
40%
the common language
Available to the specialist, but not
25%
37%
13%
14%
20%
in the common language
Available to the general public,
12%
4%
13%
50%
21%
but not in the common language
1%
Generally unavailable
38%
8%
47%
50%
36%
40%
Literature Availability by Topic
An overview of the responses to each of the questions asked is given
in Tables 1-3 for each geographic region. It must be kept in mind that
there was often only a small number of responses for any particular
country and these may not have been filled out with as much care as
might have been desired. Nevertheless, additional library research on
the availability of the literature of mammalogy for a country frequently
was in accord with the impressions of the in-country specialists as
indicated on the returned questionnaires.
The availability of field guides is shown to be quite spotty (Table 1),
with Europe and the Australian region having such publications readi-
Table 3. — The availability of natural history literature for children within each of the
major geographic regions. Percentages are given for each geographic region and are based
on the total number of responses received from each region.
Latin
Ameri-
Euro-
Middle
Aus-
Response
can
pean
African
East
Asian
tralian
Many in the common language
60/0
62%
7%
8%
38%
Few in the common language
22%
24%
36%
100%
46%
56%
Rare in the common language
Many, but not in the common
language
Few, but not in the common lan-
11%
2%
29%
15%
guage
Rare, but not in the common
7%
8%
language
7%
8%
None
61%
7%
14%
15%
6%
No response
5%
166
Annals of Carnegie Museum
VOL. 55
ly available in the common language of the countries queried. In the
Australian area, of course, this is English, but in Europe we found that
most countries responding had access to field guides that were published
in the common language of the country in question. Asia and Africa
had a moderate rate of field guide availability and such books were
primarily available in the common language of the country. Because
of the frequency of having an official language (such as English, for
example) listed for a country, however, there arises the problem of
availability of such guides to the majority of people who may neither
read nor speak the official language. More will be said about this prob-
lem when geographic regions are reviewed individually (below). The
Middle East and especially Latin America are geographic regions that
show a decided scarcity of field guides. Those few that are available
are usually not published in the common language. Fully 72% of the
respondents in Latin America reported that field guides to mammals
are unavailable.
Mares (1982, 1985, in press) has discussed the importance of the
technical literature of mammalogy as a basis for the popular literature.
A review of the responses concerning the availability of technical lit-
erature (Table 1) indicates that such literature is generally more widely
available (to the specialist) than is semi-technical literature on mam-
mals. All geographic regions reported a fairly broad availability of
technical literature on mammals, with Latin American responses re-
porting the lowest values. Generally, such literature is available in the
common language of the country (except in the Middle East, where our
sample size was quite small).
Taxonomic keys are vitally important to the diffusion of mammalogi-
cal information beyond a small coterie of specialists. It is keys that
allow other biologists to identify their research materials with minimal
effort. Such keys are also useful in instructional purposes and to people
in government or other fields that need to have a correct determination
made for a particular specimen. Most respondents (except for Latin
Americans) indicated a fairly good availability of taxonomic keys for
use by specialists and generally these were available in the common
language of the country (Table 1).
One group of publications that proved to be quite scarce in all geo-
graphic regions is collecting and preparation manuals (Table 2). Even
in such well-studied areas as Europe and Australia, only 50% or fewer
of the respondents indicated the availabilty of such works to the spe-
cialist. These types of publications were even less readily available to
the general public.
The types of mammalogical publications that turned out to be the
least available on a worldwide scale were those that are concerned with
1986
Mares and Braun —Mammalogy Literature
167
the operation and importance of natural history museums (Table 1).
The very low values for availability in any language indicate the general
lack of literature in this area.
Children’s literature on natural histoiy was shown to be readily avail-
able only in Europe (Table 3). In most regions, fewer than 10% of the
respondents reported that children’s literature was readily available,
while from 20-50% of them said such books were few.
Literature Availability by Region
Middle East. --Only two of five questionnaires sent to countries of
the Middle East were returned, so it is not possible to speak in any
detail about the status of the literature of this region. Each of the four
countries surveyed has had foundational literature published on mam-
mals, although most of this literature is available only in European
languages. Nevertheless, although there are few guides to the mammals
of these countries and apparently almost no museum literature or pop-
ular literature published in the native languages, a rather firm foun-
dation for future workers has been established.
Latin Twerira.— Eighteen of 32 questionnaires were returned from
10 countries. Unfortunately, this is not a large sample for such a vast
region, but responses were received from Mexico, Central America,
the Caribbean, and South America, giving a good geographical repre-
sentation. In almost all literature areas surveyed, Latin America was
ranked at the lowest levels. Most countries lack even the foundational
literature, regardless of its language of publication. Few faunal surveys
have been published for most countries, taxonomic keys and field
guides are generally unavailable, museum literature is practically non-
existent, and children’s literature is rare to unavailable. These trends
are particularly pronounced for South America, as opposed to Central
America, the Caribbean, and Mexico. Because of their proximity to
North America, these regions have been fairly well-studied, especially
Mexico, the Caribbean, Panama, and Costa Rica. Unfortunately, de-
spite the availability of foundational literature (largely published in
English), there are as yet relatively few publications available of the
nature of field guides, children’s books, and general museum publi-
cations. Indeed, even in Mexico, perhaps the best-studied of the Latin
nations, semitechnical and popular publications are either uncommon
or hard to obtain. For South America, the panorama is much less
positive. There, even the foundational literature is lacking and, as has
been argued by Mares (in press), without this most basic literature of
mammalogy, the preparation of the literature needed to educate the
general population is impossible. Among the South American coun-
tries, only Suriname, Venezuela, Uruguay, Chile, and Argentina have
168
Annals of Carnegie Museum
VOL. 55
had an important portion of their foundational literature completed.
Even in these few countries, however, the semitechnical and popular
literature on mammals is largely unavailable.
Europe. —YovXy -two of 66 questionnaires were returned from Eu-
ropean collections, for a 64% response rate. Of the regions surveyed,
Europe led in all categories of literature availability, reflecting a long
history of nature study, museum collection formation, resident spe-
cialists on mammals, and so on. Generally, basic literature as well as
the semitechnical and popular literature were available in the common
language of each country.
4/nca.— Fourteen of 23 (61%) African collections queried returned
the questionnaires, with six responses coming from two countries, Na-
mibia and the Republic of South Africa. Many African countries were
thus not sampled due either to the fact that questionnaires were not
returned or that a particular country had no collection-oriented mam-
malogist to be queried. There is little doubt that the inclusion of four
responses from the Republic of South Africa skews the African data.
Still, it is evident that the African area (excluding the Middle East) has
a very poor literature base, particularly for works published in non-
European languages. Although 50% of the respondents indicated the
availability of field guides to the mammals of their particular country,
published in the common language, this number is inflated. Many cited
Dorst and Dandelot (1970), for example, which is limited to the larger
mammals of Africa. Fully 86% of the respondents reported that tech-
nical publications were available in the common language of the coun-
try, and 83% said that taxonomic keys were readily available (Table
1), but again these are probably overestimations due to the fact that
some well-surveyed countries responded (for example, Kenya, Repub-
lic of South Africa).
Australian region. — By and large, the Australian area supports a rich
and varied literature at the technical, semitechnical, and popular levels.
Field guides, technical publications, taxonomic keys, and collecting
and preparation manuals were listed as readily available, as was chil-
dren’s literature.
In answer to the question requesting a listing of subject areas needing
research, responses were understandably mixed. Topics suggested with
regularity were surveys and technical publications on a country’s fauna;
taxonomic revisions; the production of both field guides and taxonomic
keys; educational materials aimed at the general public and children
that deal with museum/collection-related topics and natural history;
theoretical and applied ecological research (including conservation);
collecting and preparation technique manuals; and behavioral research.
1986
Mares and Braun— Mammalogy Literature
169
Socioeconomic Factors
In any broad examination dealing with some facet of life in the
diverse nations of the world, it is instructive to examine what country-
wide phenomena might be related to the patterns that have been iden-
tified. We have shown that the world’s literature on mammalogical
subjects has an extremely spotty distribution. We might ask if there
are any obvious measures of a country’s socioeconomic profile that
might be related to the availability of this type of literature. It takes
no special understanding of economics or politics, for example, to see
that those countries generally classified among the economically de-
veloped nations of the world (DC’s) have a higher rate of literature
availability than do those nations generally considered as being under-
developed (UDC’s).
We have examined some basic socioeconomic factors of each nation
that responded to our questionnaire (Table 4). These data include in-
formation on Gross National Product (GNP), per capita income (PCI),
the natural rate of population increase (r), percent literacy, population
density, and the percent of the population that is urban. Utilizing the
SPSS statistical package (SPSS Inc., 1983), we compared these data
to an index of literature availability via multiple regression analysis
and stepwise multiple regression analysis. In order to do this, it was
necessary to score the responses from each country to each of the first
six questions on the questionnaire (mean values were used in the case
of multiple responses). Scores on each question varied from zero for
an unavailability of literature to five for an abundance of literature.
Scores for each question were summed to obtain the country’s total
score (scores may thus vary from zero to 30). Literature availability
scores for those nations that responded fully to a questionnaire are
given in Table 4.
When multiple regression analysis was run so that all independent
variables (the socioeconomic data of Table 4) were examined for their
effect on explaining the variance in the dependent variable (the liter-
ature availability scores of Table 4), two factors were shown to have
statistical importance in explaining variance in literature availability:
the rate of population increase, r, with P = 0.04; and the per capita
income, with P = 0.05. The sign of the beta value for r was negative,
whereas it was positive for the beta values of PCI (Table 5). This means
that the greater the value for r, the lower the availability of literature,
and the greater the PCI, the more literature that is available in a country.
In this analysis, each variable was examined for its effect on the de-
pendent variable, while all other variables were held constant.
In a separate stepwise multiple regression analysis, the single most
Table A. — Gross national product (GNP), per capita income, natural increase in population, percent literacy, population density, percent
urban population, and the literature availability score given for most countries that responded to questionnaires. Countries are listed
alphabetically within each region. The year in which the data were obtained is given in parentheses. Unnumbered countries lack sufficient
VOL. 55
170
Annals of Carnegie Museum
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X X
en os
so o r- so
r- — i F' so
g| S’
ill
fi d
— leNen'^insor^xosO
ed
cei
>
Son ^
O
’S -2 ^ S
d 13 N w
< 03 U Q
— 1 eN rn
in o
eN — I eN
15. England 445.9 (80) 7216 (79) 0.01 (77) 99 (81) 592(81) 78 (73)
16. Federal Republic of
Germany 824.6 (80) 9278 (78) -0.3 (78) 99 (78) 642 (81)
17. Finland 41.3 (79) 6090 (78) 0.4 (78) 99 (78) 37 (81) 60(80)
18. France 585 (80) 8980 (80) 0.4 (77) 99 (78) 252 (81) 73 (75)
Litera-
ture
GNP Natural increase availa-
(in billions of Per capita income in population Population density % Pop. biiity
Country (by region) U.S. dollars) (in U.S. dollars) % % Literacy (per sq. mi.) urban score
1986
Mares and Braun— Mammalogy Literature
171
fN| o b- oo Os
r--
r- r- t--
o\ o
00 oo oo 00
oo 00 oo
m NO (N o
-H CM On
^
so — 1 m
os 00
eq rq rq
^ ^ S' OO
oo 00 b- b-
00 00 00
^ w ^
On O o© in
On O On On
0© rq 00
On b- ON
OO oo'^^
b' b- b- b'
S'bT'^
b- b- b-
o cq ’-I ^
O On b-
d -4 d
d 00
oo 00 oo oo oo
(N OQ ^ VO
On On O m C')
— < cn o rq
00 ON ON On in
On On On On OO
r-- 00 00 NO 00
b- b- b- b- b-
O eN On On
-i' o d d d
oo 00 0© 00 00
On b- —I m
-I o -1 r- o
lO (N 0© ON ’-H
NO ON fN O
VO ON ON NO
oo oo b~ oo 00
b' b^ b- b'
On I O NO lO
r4
ON ON o O
b- b- oo oo
o o o ^
^ o o —<
m o O On
in On lO NO
NO On 00
o
o o o
o o o
m o
CN fs m
o in ON o ON
o in o O
in b- NO
m ^•'ON rq en
b- O
b- oo oo
O m o
«n — < NO
00
ON o
O
o
On
On 00
ON
On
—1
o
On
ON
o o o o
b~ oo
0©
b'
b^ b-
b^
b-
00
00
b»
b-
oo 0© 00 oo
' — '
^ b-
rq
rq
^ in
'd-
NO
oo
ON -4
b-”
-d-
in
d b4
—I
b4
ON
in
ON
NO
NO o ON -4
0©
ON
in
rq NO
O
ON
cn
NO
o
NO ON in rq
S3
cd
g
^ 'O S) fi S
cd
««
3 8 8
ns
w .2
.2 o § g iS
xJ xJ a o
On O’— 'bt fn-vd-in NOb-oooNO
—I rqbtbi rqrqrq rqrqrqrqm
People’s Republic of China 540 (80) 566 (80) 1.6 (78) 70 (78) 278 (81)
Table A. — Continued.
172
Annals of Carnegie Museum
VOL. 55
O
D. C
O c«
CU ^
•se
I ^
o
Bh
la
w C
Z'
.2o
= "O
•Sb
OO OO
(N fN VO
VO O OO
fM
ro O Os
—1 os
(N
(N fS -H
o
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^00
oT
VO VO ^
1^ VO r-
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w
V...^
w w
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00 m
o
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^ VO
_
<n
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00 00 —I
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/~V
^ /—s /-N
/— N
00 OO
OO OO 0©
OO OO 00
00
OO o© OO
OO OO
m m o
VO (N OO
Os
lo — 1 ^
VO VO
fN — H
fsi — 1
—i VO
m —i
^ r--
fN
.-H .=)
(N
<N rn (N
-h’ (N fN
O o
O in
5^-
O O ^
m ^
rn o^
cn <N
Q os OO
oo
•r^ wn o
(N iri VO
(N OS
ro rsi <N
Os os (N
Os Os
(N Os
O <N
%
o o o
oo cn o
m cn oo
m so m
(N os m
lO (N (N
O O
O t--
o tn
00
O ro O
(N VO 00
CO
r-
o c e
0< c3
c'S C
.s5.|
QT) H >
o
frt O rO .S 3
1*1 s
x>
cd
N o
3 5 '3 .3
4)
gN
'K ^
3 o
<
-oO
o
^ o
m
(N rn
OO «— I
VO
in
O O OO
OO 0© r'
os os ^
r- OO
0© o os
^ 00 r-
OS
O —I os
WWW
OO r-
OO vq
q
q q
q
(N CO
q
CTs -h' r'’
d vd VO
cn d
^ 00
fd
CO fN
fN CN
0© os
os O
m
CN m
^ ^
VO 00
^ ''t
^ S'
W
OS d
m
* White Zimbabweans.
^ Black Zimbabweans.
1986
Mares and Braun— Mammalogy Literature
173
Table 5.— Multiple regression analysis of the socioeconomic data versus the index of
literature availability from Table 4 for countries that responded fully to the questionnaire.
B = slope; SE B = standard error of B; P = level of significance; Beta is a standardized B.
Variable
B
SE B
Beta
T
p
Urban population
-0.014
0.031
-0.063
-0.446
0.66
GNP
-0.004
0.003
0.187
1.529
0.13
Population density
0.49 xlO
0.49 X 10
-0.121
-1.017
0.32
PCI
0.55 X 10
0.27 X 10
0.349
1.975
0.05
Literacy
-0.054
0.048
-0.223
-1.128
0.26
r
-2.560
1.233
-0.475
-2.077
0.04
(Constant)
20.421
5.497
3.715
important independent variable in explaining variance in the depen-
dent variable was per capita income (P < 0.000 1): all other independent
variables were non-significant. In this analysis, = 0.33 and the beta
value was 0.91 x 10“^. Since only one independent variable was used,
beta values can be used to show the effect of PCI on literature avail-
ability. We can see that if per capita income in a country were increased
by $ 100/year, the availability of mammal literature would increase by
almost 10%. Increasing PCI by $ 1000/year would essentially double
the availability of literature in a particular society.
Discussion
Clearly, there is a great need to increase the availability of all types
of mammalogical literature throughout the world. In many places (for
example, Latin America), even the foundational literature is lacking,
thus making the development of a more broad-based and diverse lit-
erature more difficult. Many countries lack active mammalogists and
do not support a working collection of mammals. In other countries,
such collections, if they exist at all, are very poorly supported, ill-
housed, and understaffed. Research, such as coordinated faunal surveys
and basic taxonomic studies, is not encouraged. Given these conditions,
the general mammalogical panorama is bleak and the foundational
literature of mammalogy, that most basic literature that is the basis
for more advanced topics such as ecology, physiology, behavior, and
conservation, is unavailable. The questions that we asked of each cu-
rator covered both the basic literature as well as the more advanced
and popular literature. It is consistent with the views of Mares (1985,
in press) and with the points made earlier in this paper, that every
country that reported a rich popular literature also reported a very
broad basic literature. No country having a weak foundational literature
reported an abundance of field guides, children’s books, or other semi-
174
Annals of Carnegie Museum
VOL. 55
technical or popular works. Very few countries reporting a strong foun-
dational literature had a weak popular literature (exceptions are Ghana,
Papua New Guinea, Portugal, Republic of South Africa, and Singa-
pore). The close relationship between the foundational literature and
the extremely important literature of the general public appears ines-
capable. It seems unlikely that a country could begin to publish popular
faunal works on a large scale without first developing the infrastructure
of mammalogy “the collections, the faunal surveys, and the taxonomic
research.
We have shown that there is a great disparity in the way in which
the literature of mammalogy is dispersed throughout the world. Lit-
erature availability does not appear to follow political ideologies, al-
though there is a strong geographic component to the data. Holarctic
countries generally have a strong literature in mammalogy —this lit-
erature is widely available in the common language of the country and
includes both technical and popular literature. If a country is located
north of 35®N latitude, it shows a strong literature at almost all levels.
Countries south of this line, with notable exceptions (for example, the
Australian region and some few African, Asian, and Latin American
countries), lack a strong literature.
The response to the question dealing with suggested research topics
supports our analysis as to the importance of the foundational litera-
ture. In countries having a rich literature, suggested research topics
were often well-formulated and specific. Often they involved fine tuning
of ongoing research efforts, or studies of sweeping scope that were
synthetic in outlook. In countries having a poor literature, suggested
topics involved the most basic types of research, taxonomic surveys,
the construction of taxonomic keys, and so forth.
The analysis of the socioeconomic factors that might affect literature
availability points out some interesting patterns. First, although some
of the data items utilized may be intercorrelated, and even though all
of these factors are the result of complex sociopolitical components,
there are only two items of statistical significance that seem to affect
literature availability, and these two items are themselves related. Per
capita income and the rate of population increase would seem to be,
a priori, inversely related. The faster a population is growing, the lower
the rate of income per person. It is this latter statistic, PCI, that has
such an important effect on literature availability. As per capita income
increases, so does the literature of mammalogy become more diverse.
We suspect that although we limited our analysis to literature on mam-
mals, the same trends will be shown for all natural history literature.
The higher the PCI, the greater the availability of literature on nature.
It is instructive to plot literature availability (as shown by the index)
and PCI of countries (Fig. 1). There is a cluster of countries that have
1986
Mares and Braun— Mammalogy Literature
175
PER CAPITA INCOME ($1000)
Fig. L— A plot of per capita income (PCI) versus an index of literature availability for
tbe countries of the world that responded to the questionnaire. Numbers of countries
refer to Table 4. The asymptotic curve (fitted by eye) shows that after a PCI of about
$4000 per annum, literature availability reaches a relatively stable high level.
both a low per capita income and a low level of literature availability.
With few exceptions, most countries that have a diverse literature on
mammals have PCFs above $5000 US per year. Only about 1 1 coun-
tries show an anomalous relationship of PCI and literature availability.
Several socialist nations (that is, Czechoslovakia, USSR, Poland, Ro-
mania, and Yugoslavia) show a rather abundant literature with rela-
tively low PCI. Given the fact that PCI in a socialist nation is low due
to the many social services that are covered through governmental
programs, and because scientific education is strongly supported in
these countries, it is not surprising that a higher level of literature
availability is maintained with a lower PCI.
Singapore, a city/nation only since 1965, might be expected to sup-
port a poor literature, because few studies concern themselves with
mammals of a densely populated urban area. Another country that
deviates from the expected relationship, Iceland, is also easily ex-
plained. Although the country has a high PCI, it has a depauperate
mammal fauna, thus detailed works on mammals are rare.
176
Annals of Carnegie Museum
VOL. 55
Several other nations that do not fit the observed pattern are the
Republic of South Africa, where the PCI is quite low due to the inclu-
sion of Black incomes in the overall PCI, while White incomes are
quite high. The literature is primarily produced by and for Whites,
thus the unusual location of the PCI versus literature value is easily
explained.
Finally, Mexico, Portugal, and Papua New Guinea also seem to
deviate from the general pattern. Mexico is well-studied due to its
proximity to the United States, whose mammalogists have worked in
Mexico for almost a century. Similarly, Papua New Guinea has had
much of its fauna studied by colonial powers, such as England and
Germany, as well as by Australians. Portugal, although having a low
PCI at present (partially due to the fact that almost one-third of the
labor force is composed of agricultural workers), has had a history of
European research efforts on mammals in the recent past.
In general then, it appears that without a marked increase in per
capita income, mammal literature will not increase. Obviously, this
does not reflect a 1:1 relationship between PCI and literature avail-
ability. Apparently, scientific and educational resources in any country
can only be allocated after basic societal needs are being met. PCI is
a measure of how well such needs are being met, especially in a non-
socialist society. If PCI is too low, few people are able to learn to read,
to complete school, or to go into a complex field of study such as
science. In countries with low PCI’s, higher education will be a luxury,
and research on mammals or other animal groups will be uncommon,
because such studies are not usually viewed as being vital to the national
interest. With few students, fewer professors, and a lack of support for
natural history studies, literature availability will perforce be at min-
imal levels in these countries.
Can a country decide to reverse this pattern and produce the popular
literature that is needed to educate the masses about the importance
of their natural environment without increasing PCI? Our results sug-
gest that this is not a simple task. The only exceptions to the perceived
trend were in countries where PCI is a poor indicator of living standard
(socialist countries) or where foundational research had been completed
in the past due to historical accident (for example, Mexico). The PCI/
literature relationship seems to be a strong one. Elevating the PCI
clearly influences a multiplicity of societal factors in education, com-
merce, science, government, and so on. But there appears to be no
ready mechanism whereby PCI can be bypassed with a resultant flow-
ering of literature. The gradual elevation of the PCI (insofar as it reflects
economic health) very likely leads to a society that has taken care of
its basic economic needs and has developed the time and inclination
1986
Mares and Braun— Mammalogy Literature
177
to dedicate resources to what are, from the standpoint of the basic
requirements of life, rather esoteric pursuits.
An important point in this analysis is that even though the best
m,echanism for increasing the availability of natural history literature
is to raise the PCI, there is another way to elevate the overall level of
literature in any country. Supporting basic research by scientists of any
nation in a particular countiy will, ultimately, yield a significant amount
of foundational literature. This literature will later be used by nationals
of the country as building blocks for a more advanced and compre-
hensive national literature. Mexico is an excellent example of how a
country with a long history of basic research by scientists from another
nation, in this case the United States, is able to utilize the work of
these foreign scientists to develop its own cadre of biologists and the
diversity of its particular literature on nature, despite its low PCI. Papua
New Guinea is another example of this same occurrence.
Our results lead us to be cautiously optimistic. If international co-
operative research efforts on basic systematics, natural historical, and
biogeographical research can be significantly increased, the founda-
tional literature of each country can be established. The very positive
response by almost all curators to the possibility of cooperative re-
search/publication efforts indicates that the door to scientific interac-
tion is open. Colleagues throughout the world are eager to participate
with foreign scientists who might have expertise working with the fauna
of a particular country. Together, these biologists can act to produce
the keystone of the science of natural history, the foundational literature
upon which all more advanced studies are based and upon which the
popular literature of a country rests.
Acknowledgments
We thank Dr. Thomas E. Lacher, Jr., for statistical advice and Mrs. Sonya Johnson
for typing the manuscript. We also express our gratitude to the many curators who took
the time to respond to our questionnaire.
Literature Cited
L Abe, H. 1967. Classification and biology of Japanese Insectivora (Mammalia). I.
Studies on variation and classification. J. Fac. Agric. Hokkaido Univ., 55:191-
265.
2. 1968. Classification and biology of Japanese Insectivora. 11. Biological
aspects. J, Fac. Agric. Hokkaido Univ., 55:429“”458,
3. ——. 1971. Small mammals of central Nepal. J. Fac. Agric. Hokkaido Univ.,
56:367-423.
4. Abe, H., etal. 1968. Survey methods of mammals. Pp. 1-49, Survey methods
of terrestrial fauna (M. Kato, ed.), JIBP/CT-S.
5. Abelentsev, V. L 1968. [Fauna of Ukraine, VoL 1, pt. 3: Weasel family.] Kiev.
6. Abelentsev, V. L, and L G. Pidoplichko. 1956. [Fauna of Ukraine, Vol. 1, pt.
1 : Insectivores and bats.] Kiev.
178
Annals of Carnegie Museum
VOL. 55
7. Afanasev, a. V., U. S. Bazhanov, M. N. Korelov, A, A. Sludskii, and E. L
Strautman. 1953. [The animals of Kazakhstan.] Alma-Ata, 536 pp.
8. Aggundey, I. R., AND D. A. ScHLiTTER. 1984. Annotated checklist of the mam-
mals of Kenya. 1. Chiroptera. Ann. Carnegie Mus., 53:1 19-161.
9. Aguilar-Amat, J. B. d. 191 7. Instruccions per la preparacio i envio de mamifers
amb desti al Museu. Bare. Mus. Barcin. Sci. Nat. Op. Ser. Zook, 1:1-16.
10. Aharoni, B, 1930. Die Saugetiere Palastinas. Z, Saugetierk., 5:327-343.
11 — . — _ 1932. Die Muriden von Palastina und Syrien. Z. Saugetierk., 7:166-240.
12. Aitken, P. F. 1975. Two new bat records from South Australia with a field key
and checklist to the bats of the state. S. Australian Nat., 50:9-15.
13. Aitken, P. (u.d.) Supplement to: Bats of eastern Australia. Checklist and distri-
butions of South Australian bats. S. Australian Mus.
14. Albuja, L. 1982(1983). Murcielagos del Ecuador. Escuela Politecnica Nacional,
Ecuador, 285 pp.
15. Alcasid, G. L. 1969. Checklist of Philippine mammals. Natl. Mus. Philippines,
Manila, 51 pp.
16. Allen, G. M. 1938. The mammals of China and Mongolia (Natural History of
Central Asia, vol. 9, Part 1). Amer. Mus. Nat. Hist., New York, 1-620 pp.
17. . 1939. A checklist of African mammals. Bull. Mus. Comp. Zook, 83:1-
703.
18. . 1 940. The mammals of China and Mongolia (Natural History of Central
Asia, vol. 1 1, Part 2). Amer. Mus. Nat. Hist., New York, 621-1350 pp.
19. Allouse, B. E. 1968. A bibliography of the vertebrate fauna of Iraq and neigh-
boring countries. Mammals. Iraq Nat. Hist. Mus. Pubk, 25:1-36.
20. Alma^a, C. 1968. La faune mammalogique du Portugal dans la ‘Checklist of
Palaearctic and Indian mammals,' 1951, par Ellerman et Morrison-Scott. Arq.
Mus. Bocage, II (12, Suppk):6-9.
21. Alvarez, T. 1963. The recent mammals of Tamaulipas, Mexico. Univ. Kansas
Pubk, Mus. Nat. Hist., 14:363-473.
22. . 1977. Fauna terrestre. Pp. 409-425, in Recursos naturales de la Cuenca
del Papaloapan (J. L. Tamayo and E. Beltran, eds.), Vol. 1. Inst. Mexican© Recurs.
Nat. Renov. Mexico, D.F., 464 pp.
23. Alvarez del Toro, M. 1952. Los animales silvestres de Chiapas. Ed. Gob.
Estado, Tuxtla Gutierrez, Mexico, 247 pp.
24. Andera, M., and I. Horacek. 1982. Poznavame nase savee. Praha.
25. Anderson, R. M. 1965. Methods of collecting and preserving vertebrate animals.
Natl. Mus. Canada Bulk, 69:1-99.
26. Anderson, S. 1972. Mammals of Chihuahua, taxonomy and distribution. Bulk
Amer. Mus. Nat. Hist., 148:149-410.
27. Anderson, S., and W. D. Webster. 1983. Notes on Bolivian mammals. I. Ad-
ditional records of bats. Amer. Mus. Novitates, 2766:1-3.
28. Anderson, S., K. F. Koopman, and G. K. Creighton. 1982. Bats of Bolivia:
An annotated checklist. Amer. Mus. Novitates, 2750:1-24.
29. Andrews and Burrell. 1981. Tasmania’s native mammals. Tasmanian
Museum and Art Gallery.
30. Angermann, R., and H. Hackethall. 1974. Saugetiere. In Exkursions fauna
fur die Gebiete der DDR und BRD. Wirbeltiere (E. Stresemann, Hrgb.), Vol und
wissen VV, Berlin.
31. Anon. 1968. Instructions to curators. No. 1. Mammals (non-marine). British
Mus. (Nat. Hist.) Pubk, 665:1-55.
32. Anon. 1969. National museums of Rhodesia. Mardon Printers, Salisbury.
33. Anon. 1981. Mammal localities. The quarter degree grid system as applied to
Zimbabwe. Trustees of the Nat. Mus. and Monuments of Zimbabwe, Salisbury.
1986
Mares and Braun —Mammalogy Literature
179
34. Anon. 1982. Bats. Data, measurements and preservation. Trustees of the Nat.
Mus. and Monuments of Zimbabwe, Salisbury.
35. Ansell, W. F. H. 1960. Mammals of northern Rhodesia. Government Printer,
Lusaka, 179 pp.
36^ 1973. Addenda and corrigenda to mammals of northern Rhodesia, No.
4. Puku, 7:N19.
37^ — 1978. The mammals ofZambia. Natl. Parks Wildl. Sen, Chilanga, Zambia,
126 pp.
38. Ansell, W. F. H., and B. D. H. Ansell. 1973. Mammals of the north-eastern
montane areas of Zambia. Puku, 7:21-69.
39. Archer, M. 1981. Results of the Archbold Expedition. No. 104. Systematic re-
vision of the marsupial dasyurid genus Sminthopsis Thomas. Bull. Amer. Mus.
Nat Hist, 168:61-224.
40. Archer. M. (ed.). 1982. Carnivorous marsupials. 2 vols. Royal Zoological Society
of New South Wales, Sydney, 804 pp.
41. Archer, M., and G. Clayton (eds.). 1984. Vertebrate zoogeography and evo-
lution in Australasia. Hesperian Press, Western Australia, 1 203 pp.
42. Arnold, H. R. 1978. Provisional atlas of the mammals of the British Isles.
Natural Environment Research Council, The Institute of Terrestrial Ecology, Monks
Wood Experimental Station, Abbots Ripton, Huntington, England, 67 pp.
43. At ALLAH, S. I. 1977. The mammals of the eastern Mediterranean region: Their
ecology, systematics and zoogeographical relationship. Part L Saugetierk, Mitt., 25:
241-320.
44^ _____ 1978. The mammals of the eastern Mediterranean region: Their ecology,
systematics and zoogeographical relationship. Part 2, Saugetierk. Mitt., 26:1-50.
45. Avila-Pires, F. D. de. 1958. Mamiferos colecionados nos arredores de Belem
do Par£ Bol. Mus. Para. Emilio Goeldi, Nova Ser,, ZooL, 19:1-9.
46_ ■ — - — -- 1964. Mamiferos colecionados na regiao do Rio Negro (Amazonas, Brazil).
Bol. Mus. Para. Emilio Goeldi, Nova Sen, ZooL, 42:1-23.
47. Avila-Pires, F. D. de, and E. Gouvea. 1977. Mamiferos do Parque Nacional
do Itatiaia. Bol. Mus. Nac. Rio de Janeiro (ZooL), 291:1-29.
48. Baker, A. N. 1972. New Zealand whales and dolphins. Tuatara, 20:1-49.
49. Baker, R. H. 1956. Mammals of Coahuila, Mexico. Univ. Kansas PubL, Mus.
Nat Hist, 9:125-335.
50. — . 1958. The future of wildlife in northern Mexico— a problem in conser-
vation education. Trans. 23rd N. Amer. WildL Nat, Resour. Conf., pp. 568-575.
5P . 1974. Records of mammals from Ecuador. PubL Mus. Michigan State
Univ. Biol. Sen, 5:129-146.
52. Baker, R. H., and J. K. Greer. 1 962. Mammals of the Mexican state of Durango.
PubL Mus. Michigan State Univ. Biol. Ser., 2:25-154.
53. Baker, R. J., and J. K. Jones, Jr. 1975. Additional records of bats from Nic-
aragua, with a revised checklist of Chiroptera. Occas. Papers Mus., Texas Tech
Univ., 32:1-13.
54. Banks, E. 1949. Bornean mammals, Kuching Press, Sarawak, 83 pp.
55. 1978. Mammals from Borneo. Brunei Mus. J., 4:165-242.
56_ ____ 1980. More mammals from Borneo. Brunei Mus. J., 4:262-273.
57. Bannikov, A. G. 1953. [Identification of mammals of the Mongolian People’s
Republic.] Moscow.
5g__ . 1954. [Mammals of the Mongolian People’s Republic.] Moscow.
59. Barbehenn, K. R., J. P. Sumangil, and J. L. Libay. 1972-1973. Rodents of the
Philippine croplands. Philipp. Agric., 56:217-242.
60. Barnett, S. A., and L Prakash. 1975. Rodents of economic importance in
India. Amold-Heinemann Pubi., New Delhi, 175 pp.
180
Annals of Carnegie Museum
VOL. 55
61. Barney, G. O. 1982. The global 2000 report to the President. Penguin Books,
New York, 766 pp.
62. Barriga-B., E. 1965. Estudios mastozoologicos Colombianos. I. Chiroptera. Cal-
dasia, 9:241-269.
63. . 1966. Estudios mastozoologicos Colombianos. II. Caldasia, 9:491-502.
64. Bateman, J. A. 1971. Animal traps and trapping. Stackpole Books, Harrisburg,
Pennsylvania, 286 pp.
65. Bauer, K., and O. Wettstein-Westerseimb. Mammalia, I. Nachtrag. In Catalog-
us Faunae Austriae, Springer- Verlag, Wien.
66. Baumann, F. 1949. Die freilebenden Saugetiere der Schweiz. Hans Huber Publ.,
Bern, 192 pp.
67. Becke, J, P. van der. 1976. List of the main larger mammals of the Viraenga
National Park. Leopard (Zaire), 4:32-35.
68. Benshoof, L., T. L. Yates, and J. W, Froehlich. 1984. Noteworthy records of
mammals from eastern Honduras. Southwestern Nat., 29:51 1-514.
69. Bergmans, W. 1977. An annotated inventory of three small collections of Ni-
gerian microchiroptera (Mammalia, Chiroptera). Z. Saugetierk., 42:279-289.
70. Bertoni, a. de W. 1 939. Catalogos sistematicos de los vertebrados del Paraguay.
Rev. Soc. Cient. Paraguay, 4:3-37.
7 1 . Biswas, B. 1 968. Mammals, their collection and preservation for study. Pp. 139-
152, in A handbook for zoological collectors (Director, Zoological Survey of India,
ed.), Zoological Survey of India, Calcutta, 1 52 pp.
72. Blanford, W. T. 1888-1891. The fauna of British India, including Ceylon and
Burma. Part 1. Mammalia. Taylor and Francis, London, 250 pp.
73. Bobrinski, N. A., B. A. Kuznetsov, and A. P. Kuzyakin. 1965. Opredelitel’
mlekopitayushchikh SSSR. [Key to the mammals of the USSR.] IzdateFstvo.
Prosveshchenie, Moscow, 382 pp.
74. Bodenheimer, F. S. 1958. The present taxonomic status of the terrestrial mam-
mals of Palestine. Bull. Res. Counc. Isr. Zook, 7B: 165-1 90.
75. Bohlken, H., and H. Reichstein. 1974. Mammalia. In Paul Brohmer Fauna
von Deutschland (W. Tischler, Hrgb.), Quelle and Meyer, Heidelberg.
76. Bohme, W., and R. Hutterer. 1978. Kommentierte Liste einer Saugetier-Auf-
sammlung aus dem Senegal. Bonn. Zool. Beitr., 29: 303-322.
77. Booth, A. H. 1960. Small mammals of West Africa. Longman, London, 68 pp.
78. Borrero-H., j. I. 1967. Mamiferos Neotropicales. Univ. del Valle, Dept, de
Biologia, Cali, Colombia, 110 pp.
79. Bouillan, A., andA.-F. deBont. 1971. Faune du Zaire 1580. Zooleo (Kinshasa),
4:259-284.
80. Bounoure, L. 1956. Le Musee Zoologique de Strasbourg: son activite scientifique
et ses besoius. Terre et Vie, 1956(2):94-100.
8 1 . Bourret, R. 1 942. Les mammiferes de la collection du Laboratoire de Zoologie
de I’Ecole Superieure des Sciences. Notes et Travaux de TEcole Superieure, Univ.
Indochinoise, 1:1-44.
82. - — — . 1944. Mammiferes recemment entres dan les collections du Laboratoire
de Zoologie de TEcole Superieure des Sciences. Notes et Travaux de I’Ecole Su-
perieure des Sciences, Univer. Indochinoise, 3:1-43.
83. Brazenor, W. 1950. The mammals of Victoria and the dental characteristics of
monotremes and Australian marsupials. Nat. Mus. Viet. Australia Handbook No.
1, Brown, Prior and Anderson, Melbourne, 125 pp.
84. Brazenor, C. W. 1954. The mammals of Tasmania. National Museum of Vic-
toria.
85. Breeden, S., and K. Breeden. 1973. Animals of eastern Australia. Collins, Syd-
ney and London, 128 pp.
1986
Mares and Braun --Mammalogy Literature
181
86. Bremer, P. 1977. The mammals of the Noordoostpolder in the former Zuiderzee.
Lutra, 19:49-^61.
87. Brink, F. H. van den. 1955. Zoogdierengids. Van Europa ten westen van 30
oosterlengte. Elsevier, Amsterdam/Brassels, 23 1 pp,
______ 1967£I. a field guide to the mammals of Britain and Europe (Trans, by
H. Kruuk and H. N. Southern). Collins, London, 22 1 pp.
g9_ . 19676. Guide des mammiferes sauvages d’Europe (Trans, by de Bemand
Heuvelmaris). Editions Delachaux et Niestle, Switzerland, 263 pp.
90_ ______ 1969. Guida del mammiferi d’Europa (Trans, by L. Cagnolaro). Labor,
Milano, 243 pp.
91_ ______ 1971. Guia de campo de los mamiferos sal vajes de Europa Occidental.
Ed. Omega, Barcelona.
92_ 1975. Die Saugetiere Europas. Verlag Paul Parey, Hamburg/Berlin,
217 pp.
93_ __ 1976. A field guide to the mammals of Britain and Europe. Collins,
London, 221 pp.
94_ _ — _ 1978. Zoogdierengids. Elsevier, Amsterdam/Brasels, 232 pp.
95. Brosset, a. 1960. Les mammiferes du Maroc oriental. Leur repartition-Leur
statut actuel. Bull. Soc. Sci. Maroc, 40:243-263.
96. — — =. 1962a. The bats of Central and Western India. Part I. J. Bombay Nat.
Hist. Soc., 59:1-57.
97_ _____ 19626. The bats of Central and Western India. Part 11. J. Bombay Nat.
Hist Soc., 59:583-624.
98_ ____ 1962c. The bats of Central and Western India. Part III. J. Bombay Nat.
Hist Soc., 59:707-748.
99. Brosset, A., AND G. DuBost. 1967. Chiropteres de la Guyane frangaise. Mam-
malia, 31:583-594.
100. Brown, R. E. 1980. Rodents of the Kavir National Park, Iran. Mammalia, 44:
89-96.
101. Bruver, W. 1973. Fauna of north Thailand. Nat Hist Bull Siam Soc., 24:463-
466.
102. Budin, O. a. 1982. Taxidermia y captura de mamiferos. Fund. Miguel Lillo
Misc. No. 73, 35 pp.
103. Bulmer, R. N. H., and J. L Menzies. 1972. Karam classification of marsupials
and rodents. Part L J. Polynesian Soc., 81:472-499.
104_ . 1973. Karam classification of marsupials and rodents. Part IT J. Polyne-
sian Soc., 82:86-107.
1 05. Burt, W. H, 1938. Faunal relationships and geographic distribution of mammals
in Sonora, Mexico. Misc. Publ. Mus. ZooL, Univ. Michigan, 39:1-77.
106. Burt, W. H., and R. A. Stirton. 1961. The mammals of El Salvador. Misc.
Publ. Mus. ZooL, Univ. Michigan, 117:1-69.
107. Burton, M. 1976a. Elseviers zoogdierengids. Elsevier, Amsterdam/Brassels.
108. ■ — — 19766. Guide to the mammals of Britain and Europe. Elsevier Phaidon,
Oxford, 256 pp.
109. . 1978. Guia de campo de los mamiferos espanoles y de Europa. Ed. Omega,
Barcelona.
110. Butter worth, B. B., and A. Starrett. 1964. Mammals collected by the Los
Angeles County Museum Expedition to northeastern Venezuela 1958. Contrib.
Sci., Los Angeles Co. Mus. 85:1-8.
111. Buttiker, W., and D. L. Harrison. 1982, On a collection of Rodentia from
Saudi Arabia. Fauna Saudi Arabia, 4:488-502.
112. Cabrera, A. 1914. Fauna Iberica. Mamiferos. Mus. Nac. Cienc. Nat., Madrid,
441 pp.
182
Annals of Carnegie Museum
VOL. 55
113. 1957. Catalogo de los mamiferos de America del Sur. I. Metatheria-
Unguiculata-Camivora. Rev, Mus. Argent. Cienc. Nat. “Bernardino Rivadavia”
Inst. Nac, Invest. Cienc. Nat. ZooL, 4: 1--307.
1 14. 1961. Catalogo de los mamiferos de America del Sur, II, (Sirenia-Peris-
sodactyla-Artiodactyla-Lagomorpha-Rodentia-Cetacea). Rev. Mus. Argent. Cienc.
Nat, “Bernardino Rivadavia” Inst. Nac. Inst. Cienc. Nat. Zool., 4:308-732.
115. Cabrera, A., and J. Yepes. 1960. Mamiferos SudAmericanos. Vols. I and II.
Ediar, Buenos Aires, 187 + 160 pp.
116. Cadigan, F. C., Jr. 1973. Mammals of Malaysia: An annotated bibliography of
volumes 1--24 of the Malayan Nature Journal, Malayan Nat. J., 25:97-1 13.
117. Cagnolaro, L. 1969^2. I Mammiferi del Parco Nazionale dello Stelvio. Azienda
di stato foreste demaniali, ufficio amministrazione del Parco Nazionale dello Stel-
vio, Sondrio, 47 pp.
118. . \969b. I Musei di Storia Naturale e Feducazione naturalistica. Atti L
Riunione S.I.P.S., Roma, pp. 589-597.
119. . 1971. La nuova esposizione generale dei mammiferi nel Museo Civico
di Storia Naturale di Milano. Natura (Milan), 62:421-452.
120. . 1976. Cataloga dei tipi del Museo Civico di Storia Naturale di Milano.
III. I tipi dei mammiferei, con un profilo storico sulla collezione mammalogica.
Atti Soc. Italiano Sci. Nat. Hist. Mus. Civ. Stor. Nat. Milano, 1 17:85-108.
121. Calaby, j. H., and j. M. Taylor. 1980. Re-evaluation of the holotype of Mus
Jentink, 1880 (Rodentia: Muridae) from western New Guinea (Irian Jaya).
Zool. Meded. (Leiden), 55:215-219.
122. Calinescu, R. 1931. Mamiferele Romaniei. Bulet. Min. Agric. Domen 251, 1.
Bucharest.
123. Cao, V. S. 1984. Inventaire des rongeurs du Vietnam. Mammalia, 48:391-395.
124. Carter, C. H., H. H. Genoways, R. S. Loregnard, and R. J. Baker. 1981.
Observations on bats from Trinidad, with a checklist of species occurring on the
island. Occas. Papers Mus., Texas Tech. Univ., 72:1-27.
125. Carter, D. C., and J. K. Jones, Jr. 1978. Bats from the Mexican state of Hidalgo.
Occas. Papers Mus., Texas Tech Univ., 54:1-12.
126. Carvalho, C. T. de, and A. J. Toccheton. 1969. Mamiferos do nordeste do
Para, Brasil. Rev. Biol. Trop., 15:215-226.
127. Ceballos G., G., and C. Galindo L. 1984. Mamiferos silvestres de la cuenca de
Mexico. Publ. Inst. Ecol. Mexico, 12:1-299.
128. Chakraborty, S. 1984. Research on mammals of the Indian subcontinent by
the scientists of the Zoological Survey of India. Zoological Survey of India, Calcutta,
35 pp.
1 29. Chaline, j., H. Baudvin, D. Jammot, and M. C. Saint-Girons. 1 974. Les proies
des Rapaces (petits mammiferes et leur environnement), Ed. Doin, Paris.
1 30. Chani, j. M. 1 980. Guia de metodos de captura para el estudio de los vertebrados.
Univ. Nac. Mar del Plata, 43 pp.
131. Chasen, F, N. 1940. A hand list of Malaysian mammals. (A systematic list of
the mammals of the Malay Peninsula, Sumatra, Borneo and Java, including the
adjacent small islands). Bull. Raffles Mus., 15:1-209.
132. Chen-Huang, S. (ed.X 1962. [The economic fauna of China. Vol. Mammalia.]
Scientific Publ. Co., Peking.
133. CoETZEE, C. G. 1969. The distribution of mammals in the Namib Desert and
adjoining inland escarpment. Sci. Papers Namib Desert Res. Stat., 40:23-36.
134. . 1972. The identification of southern African small mammal remains in
owl pellets. Cimbebasia Ser. A, 2:53-64.
135. Copley, H. 1950. Small mammals of Kenya. Highway Nature Guides, Nairobi,
96 pp.
1986
Mares and Braun— Mammalogy Literature
183
136. Corbet, G. B. 1964. The identification of British mammals. British Mus. (Nat.
Hist.) PubL, 46 pp.
1966. The terrestrial mammals of Western Europe. G. T. Foulis and Co.,
Ltd., London, 264 pp.
138. — — . 1975. Finding and identifying mammals in Britain. British Mus. (Nat.
Hist) PubL, 775:i--56.
139. — , 1978. The mammals of the Palaearctic region: A taxonomic review.
British Mm. (Nat Hist) PubL, 788:1-314.
140. . 1984. The mammals of the Palaearctic region: A taxonomic review.
Supplement. British Mus. (Nat. Hist.), 45 pp.
141. Corbet, G., and D. Ovenden. 1980. The mammals of Britain and Europe.
Collins, London, 253 pp.
142. Corbet, G. B,, AND H. N. Southern. 1977. The handbook of British mammals.
Blackwell Scientific Publications, Oxford, 520 pp.
143. Corbet, G. B., and D. W. Yalden. 1972. Recent records of mammals (other
than bats) from Ethiopia. Bull. British Mus., 22:213-252.
144. CovACEviCH, J., AND A. Easton. 1974. Rats and mice in Queensland. Queensland
Mus. Booklet No. 9, 72 pp.
145. Dalquest, W. W. 1953. Mammals of the Mexican state of San Luis Potosi. La.
State Univ. Stud. Biol. Sci. Sen, 1:1-229.
146. Davis, D. D. 1962. Mammals of the lowland rain forest of North Borneo. Bull.
Natl. Mus. Singapore, 31:1-129,
147. DeBlase, A. F. 1980. The bats of Iran: Systematics, distribution, ecology. Field-
iana-ZooL, Chicago Mus. Nat. Hist,, 4:1-424.
148. DeBlase, A. F., and R, E, Martin. 1974. A manual of mammalogy with keys
to families of the world. Wm. C. Brown Co., Dubuque, Iowa, 329 pp.
1 49^ . 1981. A manual of mammalogy with keys to families of the world. Wm.
C. Brown Co,, Dubuque, Iowa, 436 pp.
1 50. DA Gama, M. M. 1957. Mamiferos de Portugal (chaves para a sua determinagao).
Mem. Mus. ZooL Univ. Coimbra, 246:1-246.
151. Delany, M. J. 1975. The rodents of Uganda. British Mus. (Nat Hist.) PubL,
764:1-165.
152. Departamento DE Zoologia. 1967. Manual de coleta e prepara^ao de animais
terrestres e de Agua Doce. Pubi. Mus. Zool. Univ. Sao Paulo, 223 pp.
153. Didier, R., AND A. Boudarel. 1948. L’art de la taxidermi du XX siecle. EncycL
Biol. (Paris), 30:1-77.
154. Donoso-Barros, R. 1975. Contribucion al conodmiento de los cetaceos vi-
vientes y fosiles del territorio de Chile. Gayana (Zoologia), 36:1-127.
155. Dorst, j., and P. Dandelot. 1 970. A field guide to the larger mammals of Africa.
Collins, London, 287 pp,
156. Dulic, B., and D. Miric. 1967. Catalogus faunae Jugosloviae. IV. Mammalia,
Consiluim Academiamm Scientiaram Rei Publicae Socialisticae Foederativae Ju-
goslaviae, 4(4): 1-46.
157. Dumitrescu, D. 1979. Brochures et livres a caractere de vulgarisation et scien-
tifique elabores por les specialistes du Museum d’Histoire naturelle “Grigore An-
tipa” pendant la periode 1946-1978. Trav. Mus, Hist. Nat. “Grigore Antipa,” XX:
749-758.
158. Dupuy, A, R., AND J. Verschuren, 1977. Wildlife and parks in Senegal. Oryx,
14:36-46,
159. Dusson de Reichel, A. 1973, Guia de los museos de Colombia. Instit. Colombi-
aeo Cult., Div Museos Restauracion, Bogota, Colombia, 141 pp,
160. Dwyer, P. D, 1983. An annotated list of mammals from Mt. Erimbari, Eastern
Highlands Province, Papua New Guinea. Sci. New Guinea, 10:28-38.
184
Annals of Carnegie Museum
VOL. 55
161. Echsel, H., and M. Racek. 1976. Biologische praparation: Arbeitsbuch f. In-
teressierte an Instituten u. Schulen. Jugend and Volk, Wien, Munchen, 248 pp.
162. Einarsson, a. (ed.). 1980. [Wild mammals.] Rit Landvemdar 7, Reykjavik.
163. Eisenberg, J. F, and G. M. McKay. 1970. An annotated checklist of the mam-
mals of Ceylon with keys to the species. Ceylon J. Sci. (Sec. B Zook), 8:69-99.
164. Eisentraut, M. 1963. Die wirbeltiere des Kamerungebirges. Verlag Paul Parey,
Hamburg, 353 pp.
165. , 1973. Die Wirbeltierfauna von Fernando Poo und Westkamerun. Unter
besonderer Beriiksichtigung der Bedentung der pleistozanen Klimaschwankungen
fur die hentige Faunenverteilung. Bonn. Zool. Monogr., 3:1-428.
1 66. Ellerman, j. R. 1 947. A key to the Rodentia inhabiting India, Ceylon and Burma,
based on collections in the British Museum. J. Mamm., 28(3-4):249-278; 357-
386.
167. . 1961. The fauna of India, including Pakistan, Burma and Ceylon. (2nd
Ed.) Vol. 3. Mammalia. Parts 1, 2. Zool. Surv. India, Calcutta, 884 pp.
168. Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic
and Indian mammals 1758-1946. British Mus. (Nat. Hist.), 810 pp.
169. . 1966. Checklist ofPalaearctic and Indian mammals 1758 to 1946. British
Mus. (Nat. Hist.), 810 pp.
170. Ellerman, J. R., T. C. S. Morrison-Scott, AND R. W. Hayman. 1953. Southern
African mammals 1758-1951: A reclassification. British Mus. (Nat. Hist.), 363 pp.
171. Enders, R. K. 1935. Mammalian life histories from Barro Colorado Island,
Panama. Bull. Mus. Comp. Zool, 78:385-502.
172. Engles, H. 1972. Kleinsauger aus Portugal. Bonn. Zool. Beitr., 23:79-86.
173. Faglar, M. 1968. Bats of Turkey. I. Acta Biol. Turcica, 18:5-18.
174. 1969. Bats of Turkey. II. Acta Biol. Turcica, 19:88-106.
175. Fairley, J. S. 1974. Irish wild mammals: A guide to the literature. Corrib. Print-
ers, Galway, 127 pp.
176. . 1975. An Irish beast book. A natural history of Ireland’s furred wildlife.
Blackstaff Press, Belfast, 201 pp.
177. Felten, H. 1955. Fledermause (Mammalia, Chiroptera) aus El Salvador. Teil 1.
Senckenb. Biol., 36:271-285.
178. — -. 1956(2. Fledermause (Mammalia, Chiroptera) aus El Salvador. Teil 2.
Senckenb. Biol., 37:69-86.
179. — 1956^. Fledermause (Mammalia, Chiroptera) aus El Salvador. Teil 3,
Senckenb. Biol. 37:179-212.
180. 1956c. Fledermause (Mammalia, Chiroptera) aus El Salvador. Teil 4.
Senckenb. Biol., 37:341-367.
181. \956d. Quiropteros (Mammalia, Chiroptera) en El Salvador. Comun.
Inst. Trop. Invest. Cient., 5:153-170.
182. -. 1957. Fledermause (Mammalia, Chiroptera) aus El Salvador. Teil 5.
Senckenb. Biol., 38:1-22.
183. Feng, Z., Z. Changlin, and C. Guiquan. 1980. On mammals from southeastern
Xizang (Tibet). Acta Zool. Sinica, 26:91-97.
184. Fenton, M. B. 1975. Observations on the biology of some Rhodesian bats,
including a key to the Chiroptera of Rhodesia. R. Ont. Mus. Life Sci. Contrib.,
104:1-27.
185. FeriancovA-Masarova, Z., and V. Hanak. 1965. Stavovc Slovenska. Vol. 4.
Pp. 301-323, in Vydavatel’-Stvo Slovenskej Akademie vied 19, Bratislavia.
186. Ferreira, R. V. 1970. Fauna: Conservacidn y recursos. Nuestra Tierra, Mon-
tevideo, 60 pp.
187. Fitzwater, W. D., and I. Prakash. 1973. Handbook of vertebrate-pest control.
Indian Council of Agricultural Research, New Delhi, 92 pp.
1986
Mares and Braun— Mammalogy Literature
185
188. Flannery, T. 5 and C Groves. 1983. The mammals of West Irian. Hemisphere
(Woden, Australia), 27:378=^384.
1 89. Fonollat, a. M. P. de. 1 984, Cricetidos de la provincia de Tucuman (Argentina).
Acta ZooL Lilloana, 37:219-225.
190. Foromozov, a. N. (ed.). 1965. [Mammals of the USSR.] Moscow.
191. Frade, F., and J. a. Silva. 1981-1982. Mamiferos de Mozambique (coleczao
do Centro de Zoologia). Garcia de Orta, ZooL, 10:1-12.
192. Frechkop, S. 1958. Faune de Belgique. Mammiferes. Inst. R. Sci. Nat. Belg.
Mem., 545 pp.
1 93. Funaioli, U., and A. M. Simonetta. 1 966. The mammalian fauna of the Somali
Republic: Status and conservation problems. Monit. ZooL Italiano, 74(SuppL):
285-347.
194. Gaffrey, G. 1953. Die Schadel der mitteleuropaishen Saugetiere, Abh. Mus.
Tierk. Dresden, 21:5-123.
I95_ _____ 196L Merkmale der wildlebenden Saugetiere Mitteleuropas. Akad. Ver-
lagsges. Geest and Portig K.-G,, Leipzig, 284 pp.
196. Gaisler, J., V, HolisovA, J. Pelikan, and J. Zejda. 1962. Klk k urcovani
drobnych savcu podle vnejsich znaku. Bmo.
197. GameroL, 1. 1978. Mamiferos de mi tierra. 2 vols. Tegucigalpa, D.C., Honduras.
198. Garms, H., and L, Borm. 1981. [Fauna Europe. Manual for identification of
animals.] Mladinska knjiga, Yugoslavia.
199. Gaskin, D. E. 1968. The New Zealand Cetacea. Fish. Res. Bull, N.Z., 1:1-92.
200. . 1972. Whales, dolphins and seals. Heinemann Educational Books, Auck-
land, 200 pp.
201. Gaumer, G. F. 1917, Monografia de los mamiferos de Yucatan. Departamento
de Talleres Graficos de la Secretaria de Fomento, Mexico, 332 pp.
202. Gavino, Gonzalo, de la Torre, et al. 1974. Tecnicas biologicas selectas de
laboratorio y de campo. Ed. Limusa Mexico.
203. Genoways, H. H., and J. K. Jones, Jr. 1975. Annotated checklist of mammals
of the Yucatan Peninsula, Mexico. IV. Carnivora, Sirenia, Perissodactyla, Artiodac-
tyla. Occas. Papers Mus,, Texas Tech Univ., 26:1-22.
204. Genoways, H. H., and D. A. Schlitter. 1981. Collections of recent mammals
of the world, exclusive of Canada and the United States. Ann. Carnegie Mus., 50:
47-80.
205. Genoways, H. H., and S. L. Williams. 1979. Records of bats (Mammalia:
Chiroptera) from Suriname. Ann. Carnegie Mus., 48:323-335.
206. Genoways, H. H., S. L. Williams, and J. A. Groen. 1981. Results of the Alcoa
Foundation-Suriname expeditions, V. Noteworthy records of Surinamese mam-
mals. Ann. Carnegie Mus., 50:319-332.
207. George, G. 1984, The present day land mammal fauna of Papua New Guinea.
Pp, 1 1 37-1 14 1 , m Vertebrate zoogeography and evolution in Australasia (M. Arch-
er and G. Clayton, eds.), Hesparian Press, Carlisle, Western Australia, 1 203 pp.
208. Gibb, J. H., and J. E. C. Flux. 1973. Mammals. Pp. 334-371, in The natural
history of New Zealand: An ecological survey (G. R. Williams, ed.), A. H. and A.
W, Reed, Wellington, Sydney and London, 437 pp.
209. Goi.dman, E, a. 1920. Mammals of Panama. Smithson. Misc. ColL, 69:1-309.
210. Goodwin, G. G. 1934. Mammals collected by A. W. Anthony in Guatemala
1924-1928. Bull. Amer. Mus, Nat. Hist., 68:1-60.
211. . 1946. Mammals of Costa Rica. Bull. Amer. Mus. Nat Hist., 87:271-
473.
212. . 1969. Mammals from the state of Oaxaca, Mexico, in the American
Museum of Natural History. Bull. Amer. Mus. Nat. Hist., 141:1-270.
186
Annals of Carnegie Museum
VOL. 55
213. Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad
and Tobago. Bull. Amer. Mus. Nat. Hist., 122:187-302.
2 1 4. Gouin, F. 1950. Les mammiferes d’ Alsace. Introduction gen. mamm., Mus. Zool.
Strasburg Livret-guide des collections, Fasc. 1 , 29 pp.
215. Gould, J. 1983. The mammals of Australia (with modem notes by J. M. Dixon).
Macmillan Co. Australia, South Melbourne, 416 pp.
216. Graaff, G. de. 1981. The rodents of southern Africa. Notes on their identifi-
cation, distribution, ecology and taxonomy. Butterworths, Durban and Pretoria,
267 pp.
217. Green, M. J. B. 1981, A checklist and some notes concerning the mammals of
the Langtang National Park, Nepal. J. Bombay Nat. Hist. Soc., 78:77-87.
218. Green, R. H. 1973a. The mammals of Tasmania. Wildl. Aust., 1 1(1):8-1 1.
219. . 1973Z?. Biogeography and ecology of Tasmania. Mammals. Monogr. Biol.,
25:367-396.
220. Greer, J. K. 1965. Mammals of Malleco Province Chile. Publ. Mus. Michigan
State Univ. Biol. Ser., 3:49-152.
221. Gromov, I. M., D. I. Bibikov, N. I. Kalabukhov, and M. N. Meier. 1965.
[Fauna of the USSR Vol. 3, pt. 2: Ground squirrels (Marmotinae)]. Moscow.
222. Gromov, I. M., A. A. Gureev, G. A. Novikov, I. I. Sokolov, P. P. Strelkov,
AND K. K. Chaiskii. 1963. [Mammalian fauna of the USSR.] USSR Acad, of
Sci., Moscow, 2 vols.
223. Gunnlaugsson, T. 1955. [In arctic fox country.] Reykjavik.
224. Gureev, A. A. 1964, [Fauna of the USSR, Mammals, Vol. 3, pt. 10, Lagomorpha.]
Moscow.
225. . 1979. Mammalia, Insectivora: Hedgehogs, moles and shrews (Erinacei-
dae, Talpidae, Soricidae). Fauna SSSR. Akad. Nauk., Leningrad, 120:1-502.
226. Gvozdev, E. V., and V. I. Kapitov (eds.). 1983. [Mammals of Kazakhstan. III.]
Nauka, Alma-Ata, 241 pp.
227. Gvozdev, E. V., AND E. I. Strautman (eds.). 1981a. Mammals of Kazakhstan.
3 Part 1. Carnivora (Canidae, Ursidae, Procyonidae) and Pinnipeda (Phocidae).
Nauka, Alma-Ata, 241 pp.
228. . 1981Z?. Mammals of Kazakhstan. 3 Part 2. Carnivora (Mustelidae, Feli-
dae). Nauka Publ., Alma-Ata, 263 pp.
229. Haim, A., and E. Tchernov. 1974. The distribution of myomorph rodents in
the Sinai Peninsula, Mammalia, 38:201-223.
230. Hall, E. R. 1962. Collecting and preparing study specimens of vertebrates. Misc.
Publ. Mus. Nat. Hist., Univ. Kansas, 30:1-46.
231. — . 1981. Mammals of North America. John Wiley and Sons, New York, 2
vols., 1181 + 90 pp.
232. Hall, E. R., and W. W. Dalquest. 1963. The mammals of Veracruz. Univ.
Kansas Publ, Mus. Nat. Hist., 14:165-362.
233. Hall, E. R., and K. R. Kelson. 1959. Mammals of North America. Ronald
Press, New York, 2 vols., 1083 pp.
234. Hall, L. S., and G. C. Richards. 1979. Bats of eastern Australia. Queensland
Mus. Booklet, 12:1-66.
235. Halsted, D. C. 1979. Birds and larger mammals of Zanzibar. J. East African
Nat. Hist. Soc. Nat. Mus., 1979:41-45.
236. Haltenorth, T., and H. Diller. 1977. Saugetiere Afrikas und Madagaskars.
BLV, Munich, 403 pp.
237. Hamar, V. 1967. Din viata rozatoarelor. Ed. St. Bucuresti.
238. Hampton, J. W. F., ETAL. 1982. Records of the mammal survey group of Victoria,
1966-80, on the distribution of terrestrial mammals in Victoria. Australian Wildl.
Res., 9:177-201.
1986
Mares and Braun— Mammalogy Literature
187
239. HanAk, V. 1967. Verzeichnis der Saugetiere der Tschechoslowakei, Saugetierk.
Mitt., 15:193-221.
240. Hanak, V., AND A. Elgadi. 1984. The bat fauna (Chiroptera) of Libya. Vestn.
Ceskoslovenske Spolec. Zool., 48:165-187.
241. Handley, C. O., Jr. 1966. Checklist of the mammals of Panama. Pp. 753-795,
in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.). Field Museum
of Natural History, Chicago.
242. 1976. Mammals of the Smithsonian Venezuelan Project. Brigham Young
Univ. Sci. Bull. Biol. Ser., 20:1-89.
243. Hanif, M. 1968. Fauna of Guyana. Wild. Cons. Guyana Mem. Nat. Ser., 8:
25-40.
244. HanzAk, J,, and Z. Veselovsky. 1965. Svetem zvirat.-Savci. Statni nakl. detske
knihy, Praha.
245. HanzAk, J., D. CernA, and V. MazAk. 1970. Nasi savci. Albatros, Praha, 347
pp.
246. Happold, D. 1972. Mammals in Nigeria. Oryx, 1 1:469.
247. Happold, D. C. D. 1973. The distribution of large mammals in West Africa.
Mammalia, 37:88-93.
248. Happold, D. C. D,, and M. Happold. 1978<2. The fruit bats of western Nigeria.
Nigerian Field, 43:30-37.
249. 1978^ The fruit bats of western Nigeria. Part 11. Nigerian Field, 43:
72-77.
250. = — — 1978c. The fruit bats of western Nigeria. Part III. Nigerian Field, 43:
121-127.
251. Harrison, D. L. 1956. Mammals from Kurdistan, Iraq, with description of a
new bat. J. Mamm., 37:257-263.
252. — — 1961. A checklist of the bats (Chiroptera) of Kenya Colony. J. East African
Nat. Hist. Soc. Nat Mus., 23:286-295.
253. -- — 1964. The mammals of Arabia. Vol. 1. Insectivora, Chiroptera, Primates.
Ernest Benn, London, 1 92 pp.
254. . 1968. The mammals of Arabia. Vol. 2. Carnivora, Hyracoidea, Artiodac-
tyla. Ernest Benn, London, 193-380 pp.
255. — . 1972. The mammals of Arabia. Vol. 3. Lagomorpha, Rodentia. Ernest
Benn, London, 381-670 pp.
256. . 1973. Checklist: Saudi Arabian mammals. J. Saudi Arab. Nat. Hist Soc.,
1:22-25.
257. . 1977. Mammals obtained by the expedition with a checklist of the mam-
mals of the Sultanate of Oman, Pp. 1 3-26, in The scientific results of the Oman
flora and fauna survey 1975, J. Oman Stud. Spec. Rept.
258. Harrison, J. L. 1966^. An introduction to mammals of Singapore and Malaya.
Malayan Nature Soc., Singapore Branch, Singapore, 340 pp.
259. — . 1966Z?. A guide to mammals of the Malay Peninsula. Mimeographed,
Dept, of Zoology, Univ. of Singapore.
260. Harrison, J. L., and R. Traub. 1950. Rodents and insectivores from Selangor,
Malaya. J. Mamm., 31:337-346.
261. Hassinger, j. D. 1973. A survey of the mammals of Afghanistan. Fieldiana-
Zool., Chicago Mus. Nat. Hist, 60:1=195.
262. Hatt, R. T. 1959. The mammals of Iraq. Misc. Publ. Mus. Zool., Univ. Michigan,
106:1-113.
263. Heaney, L. R. 1984, Mammals of Camiguin Island, Philippines. Proc. Biol. Soc.
Washington, 97:119-125.
264. Heptner, V. G., and N. P, Naumov. 1961. [Mammals of the Soviet Union, Vol.
1: Even-toed and odd-toed mammals.] Moscow. German translation: Jena, 1966.
188
Annals of Carnegie Museum
VOL. 55
265. — — 1967. [Mammals ofthe Soviet Union, VoL 2, pt. l:Sirenia and Carnivora].
Moscow. German translation: Jena, 1974.
266. — 1972. [Mammals of the Soviet Union, Vol. 2, pt. 2: Carnivores (hyaenas
and cats).] Moscow.
267. Hernandez-C., J. 1957. Mammalia. Pp. 213-230, in Informa preliminar sobre
aves y mamiferos de Santander, Colombia (J. A. Borrero-H. and J. Hemandez-C.,
eds.), Anal. Soc. Biol. Bogota, 7:197-230.
268. Hershkovitz, P. 1 947. Mammals of northern Colombia. No. 1 : Squirrels (Sciuri-
dae). Proc. U.S. Nat. Mus., 97(3208): 1-46.
269. . 1948. Mammals of northern Colombia. No. 2: Spiny rats (Echimyidae)
supplemental notes on related forms. Proc. U.S, Nat. Mus., 97(32 14): 125-1 40.
270. . 1949. Mammals of northern Colombia. No. 5: Bats (Chiroptera). Proc.
U.S. Nat. Mus., 99(3246):429-454.
271. — 1951. Mammals from British Honduras, Mexico, Jamaica, and Haiti.
Fieldiana-Zool., Chicago Mus. Nat. Hist., 31:547-569.
272. 1962. Evolution of Neotropical cricetine rodents (Muridae) with special
reference to the phyllotine group. Fieldiana-Zool., Chicago Mus. Nat. Hist,, 46:1-
524.
273. - — . 1977. Living New World monkeys (Platyrrhini). Vol. 1. Chicago Univ.
Press, Chicago.
274. Hickman, G. C. 1981. National mammal guides: A review of references to recent
faunas. Mamm. Rev., 11:53-85.
275. Hill, J. E. 1961. Indo- Australian bats of the genus Tadarida. Mammalia, 25:
29-56.
276. -. 1963. A revision of the genus Hipposideros. Bull. British Mus., 11:1-
129.
277. 1965. Asiatic bats of the genera and (Vespertilion-
idae) with a note on Kerivoula aerosa Tomes. Mammalia, 29:524-556.
278. . 1980. The mammals of Sri Lanka. Spol. Zeylanica, 35:203-211.
279. Hill, J. E., and T. D. Carter. 1941. The mammals of Angola, Africa. Bull.
Amer. Mus. Nat. Hist., 78:1-211.
280. Hill, J. E., and P. Morris. 1971. Bats from Ethiopia collected by the Great
Abbai Expedition, 1968. Bull. British Mus., 21:25-49.
281. Hill, J. E., and K. Thonglonya. 1972. Bats from Thailand and Cambodia. Bull.
British Mus., 22:173-196.
282. Hill, M, G. A. 1970. Parque Regional “Piedras Blancas-Las Palmas.” Bases de
Planeacion, Medellin, Colombia.
283. Hindrum, R. 1981. The bird and mammal faunas of the Julelva River system,
Varanger Peninsula, northern Norway. Tromura (Tromo), 24:1-54.
284. 1982, The bird and mammal faunas of the Reisa River system, Troms
and Finnmark, northern Norway. Tromura (Tromo), 37:1-64,
285. Hoekstra, B. E., E. Van der Straeten, and V. van Laar. 1977. Handleiding
ten behoeve van het inventariseren van landzoogdieren in de Benelux. Wet. Meded.
K.N.N.V. (K. Ned. Natuurhist. Ver.), 119:1-47.
286. Hoesch, W., and E. von Lehmann. 1956. Zur Saugetier-fauna Sudwestafrikas.
Bonn, Zool. Beitr., 7:8-57.
287. Hoogstraal, H. 1962. A brief review of the contemporary land mammals of
Egypt (including Sinai). 1. Insectivora and Chiroptera. J. Egypt. Public Health
Assoc., 37:143-162.
288. -. 1963. A brief review of the contemporary land mammals of Egypt (in-
cluding Sinai). 2. Lagomorpha and Rodentia. J. Egypt. Public Health Assoc., 38:
1-35.
289.
. 1964, A brief review of the contemporary land mammals of Egypt (in-
1986
Mares and Braun— Mammalogy Literature
189
eluding Sinai), 3. Carnivora, Hyracoidea, Perissodactyla and Artiodactyla. J. Egypt.
Public Health Assoc., 39:205”239.
290. Hoogstraal, H., K. Wassif, and M. N. Kaiser. 1956--1957. Results of the
Namru-3 Southeastern Egypt Expedition, 1954. 6. Observations on non-domes-
ticated mammals and their ectoparasites. Bull. Zool. Soc. Egypt, 13:52-75.
291. Hubert, B., F. Adam, and A. Poulet. 1973. Liste preliminaire des rongeurs de
Senegal. Mammalia, 37:76-87.
292. Huey, L. M, 1964, Mammals of Baja California, Mexico. Trans. San Diego Soc,
Nat. Hist, 13:85-168.
293. Hufnagel, E, 1972, Libyan mammals. The Oleander Press, New York, 85 pp.
294. Hughes, R. L, 1965. Comparative morphology of spermatozoa from five mar-
supial families. Australian J. ZooL, 13:533-543.
295. Husson, a. M, 1955. Tabel voor het determineren van de landzoogdieren van
Nederlands Nieuw-Guinea. Zool. Bijdr., 1:1-35. (English translation by E. J. Brill,
Leiden, 1955.)
296. — 1962. The bats of Suriname. Zool. Verb. (Leiden), 58:l"-282.
297. — — — . 1973. Voorlopige lijst van de zoogdieren van Suriname. Zool. Bijdr., 14:
1-15.
298. — — — . 1978. The mammals of Suriname. Zool. Monogr. Rijksmuseum Natuur.
Hist., No. 2. E. J. Brill, Leiden, 569 pp.
299. Hutterer, R., AND D. C. D. Happold. 1983, The shrews of Nigeria (Mammalia:
Soricidae). Bonn. Zool. Monogr., 18:5-79.
300. Hyett, J. 1980. Australian mammals. A field guide for New South Wales, Vic-
toria, South Australia and Tasmania. Nelson, Melbourne, 270 pp.
301. IjssELiNG, M. A., AND A. ScHEYGROND. 1943. Dc zoogdieren van Nederland.
Vols. I and 11. W. J. Theime and Cie, Zutphen, 531 pp.
302. . 1948. Wat is dat voor een dier? W. J. Thieme and Cie, Zutphen.
303. Imaizumi, Y. 1960. (Reprinted in 1975.) Coloured illustrations of the mammals
of Japan. Hoikusha Publ. Co., Osaka, 916 pp.
304. . 1970. The handbook of Japanese land mammals. Shin-Scichoa-Sha, To-
kyo, 350 pp.
305. loNESCu, V. 1968. Vertebratele din Romania. Ed. Acad. R.S.R., Bucuresti,
496 pp.
306. Iredale, T., and E. le G. Troughton. 1934. A check-list of the mammals
recorded from Australia. Australian Mus. Syd. Mem., 6:1-122.
307. Jacobsen, N. H. G. 1977. An annotated checklist of the amphibians, reptiles,
and mammals of the Nylsvley Nature Reserve. South African Nat. Sci. Prog. Rept.,
21:1-65.
308. Jameson, E. W., Jr., and G. S. Jones. The Soricidae of Taiwan, Proc. Biol. Soc.
Washington, 90:459-482.
309. Janson, T, 1981. Animales de Centroamerica en peligro. Ed. Piedra Santa, Gua-
temala City, 122 pp.
310. Janzen, D. H,, AND D. E. Wilson. 1983. Mammals. Pp. 426-501, m Costa Rican
natural history (D. H. Janzen, ed.), Univ. Chicago Press, Chicago.
311. Johnson, D. H., S. D. Ripley, and K. Thonglongya. 1980. Mammals from
Nepal. J. Bombay Nat. Hist Soc., 77:56-63.
312. Jones, F. W. 1923. The mammals of South Australia. Vol. 1. The Monotremes
and carnivorous marsupials. Government Printer, Adelaide, 131 pp.
313. 1924. The mammals of South Australia. Vol. 2. The bandicoots and
herbivorous marsupials. Government Printer, Adelaide, 132-270 pp.
314. , 1925. The mammals of South Australia. Vol. 3. Conclusion, Monodel-
phia. Government Printer, Adelaide, 271-458 pp.
315. Jones, G., AND D. Jones. 1976. A bibliography ofthe land mammals of Southeast
190
Annals of Carnegie Museum
VOL. 55
Asia, 1699“1969. Spec. Publ., Dept. Entomology, Bernice P. Bishop Museum,
Honolulu, Hawaii, 238 pp.
316. Jones, G. S., L. B. Liat, AND J. H. Cross. 1971. A key to the mammals of Taiwan.
Chin. J. Microbiol., 4:267-278.
317. Jones, J. K., Jr. 1966. Bats from Guatemala. Univ. Kansas Publ., Mus. Nat.
Hist., 16:439-472.
318. Jones, J. K., Jr., and D. H. Johnson. 1960. Review of the insectivores of Korea.
Univ. Kansas Publ., Mus. Nat. Hist., 9:549-578.
319. . 1965. Synopsis of the lagomorphs and rodents of Korea. Univ. Kansas
Publ., Mus. Nat. Hist., 16:357-407.
320. Jones, J. K., Jr., J. D. Smith, and H. H. Genoways. 1973. Annotated checklist
of mammals of the Yucatan Peninsula, Mexico. 1. Chiroptera. Occas. Papers Mus.,
Texas Tech Univ., 13:1-31.
321. Jones, J. K., Jr., H. H. Genoways, AND T. E. La WLOR. 1974. Annotated checklist
of mammals of the Yucatan Peninsula, Mexico. 2. Rodentia. Occas. Papers Mus.,
Texas Tech Univ., 22:1-24.
322. Jones, J. K., Jr., H. H. Genoways, and J. D. Smith. 1974. Annotated checklist
of mammals of the Yucatan Peninsula, Mexico. 3. Marsupialia, Insectivora, Pri-
mates, Edentata, Lagomorpha. Occas. Papers Mus., Texas Tech Univ., 23:1-12.
323. Jones, J. K., Jr., P. Swanepoel, and D. C. Carter. 1977. Annotated checklist
of the bats of Mexico and Central America. Occas. Papers Mus., Texas Tech Univ.,
47:1-35.
324. JouBERT, E., AND P. K. N. Mostert. 1975. Distribution patterns and status of
some mammals in South West Africa. Madoqua, 9:5-44.
325. Kahmann, H. 1952. BeitragezurkenntnisderSaugetierfaunainBayem. 5. Bericht
der Naturforsch. Ges. Augsburg, 147-170 pp.
326. Kennedy, M. L., T. L. Best, AND M. J. Harvey. 1984. Bats of Colima, Mexico.
Mammalia, 48:397-408.
327. Kenmuir, D., and R. Williams. 1975. Wild animals—a field guide and intro-
duction to the mammals of Rhodesia. Longman Rhodesia (Pvt.) Ltd., Salisbury,
136 pp.
328. Khajuria, H. 1983. Taxonomic research on Indian mammals. Rec. Zool. Surv.
India, 80:355-374.
329. Kingdon, J. 1971. East African mammals. An atlas of evolution in Africa. Vol.
1 . Primates, Hyraxes, Pangolins, Protoungulates, Sirenians. Acad. Press, New York,
446 pp.
330. . 1974a. East African mammals. An atlas of evolution in Africa. Vol. 2,
Part A. Insectivores and bats. Acad. Press, New York, 341 pp.
331. . 1974Z?. East African mammals. An atlas of evolution in Africa. Vol. 2,
Part B. Hares and rodents. Acad. Press, New York, 703 pp.
332. . 1977. East African mammals. An atlas of evolution in Africa. Vol. 3,
Part A. Carnivores. Acad. Press, New York, 476 pp.
333. . 1979. East African mammals. An atlas of evolution in Africa. Vol. 3,
Part B. Large mammals. Acad. Press, New York, 436 pp.
334. . 1982a. East African mammals. An atlas of evolution in Africa. Vol. 3,
Part C. Bovids. Acad. Press, New York, 393 pp.
335. — . 1982Z?. East African mammals. An atlas of evolution in Africa. Vol. 3,
Part D. Bovids. Acad. Press, New York, 394-746 pp.
336. Kirkpatrick, R. D., and A. M. Cartwright. 1975. List of mammals known to
occur in Belize. Biotropica, 7:136-140.
337. Kirsch, J. A. W., AND J. H. Calaby. 1977. The species of living marsupials: An
annotated list. Pp. 9-26, in The biology of marsupials (B. Stonehouse and D.
Gilmore, eds.), Macmillan, London, 486 pp.
1986
Mares and Braun— Mammalogy Literature
191
338. Kistchinski, A. A. 1973. Birds and land mammals in the USSR. lUCN {Int.
Union Comerv. Nat. Nat. Resow.) Publ. New Ser., 25:177-180.
339. Kock, D. 1 969. Die Fledermaus-Fauna des Sudan (Mammalia^ Chiroptera). Abh.
Senckenb. Naturforsch. Ges., 521:1-238.
340. . 1978. Vergleichende Untersuchung einiger Saugetiere in sudlichen Niger
(Mammalia: Insectivora, Chiroptera, Lagomorpha, Rodentia). Senckenb. Biol., 58:
113-136.
341. Konig, C. 1969. Wildlebende Saugetiere Europas. Belser, Stuttgart, 255 pp.
342. Koopman, K. F. 1975. Bats of the Sudan. Bull. Amen Mus. Nat. Hist., 154:353-
444.
343. . 1978. Zoogeography of Peruvian bats with special emphasis on the role
of the Andes. Amen Mus. Novitates, 2651:1-33,
344. Kowalski, K. 1 979. Terrestrial mammals of Algeria in the collections of the City
Museum in Oran. African Small Mammal News!., 4:8-10.
345. Krystufek, B. 1982. The mammals of Ljubljansko Baije, Yugoslavia. Biol Vestn.
(Ljubljana), 30:33-56.
346. Kuhn, H. J. 1965. A provisional checklist of the mammals of Liberia. Senckenb.
BioL, 46:321-340.
347. Kullmann, E. 1965. Die Saugetiere Afghanistans. (Teil I): Carnivora, Artiodac-
tyla, Primates. Sci. Sp. Ed. Inst. ZooL Parasit., Kabul Univ., Afghanistan, 1965:
1-17.
348. Kuntze, R. 1935. Ssaki. [Mammalia.] Zeszyt 2. Fauna stodkowodna Polski.
Wydawnictwo Kasy im, Mianowskiego, Warszawa, 72 pp.
349. Kuroda, N. 1940. A monograph of the Japanese mammals. Sansiedo Co., Ltd.,
Tokyo and Osaka, 311 pp.
350. . 1963. Systematictaxonomy of animals. 10. Mammals. Nakayame-shoten
Publ. Co., Ltd.
351. Kurup, G. V. 1968. Mammals of Assam and adjoining areas. Proc. ZooL Soc,
(Calcutta), 21:79-99.
352. Lane, H. U. (ed.). 1983. The world almanac and book of facts 1983. Newspaper
Enterprise Assoc., Inc., New York, 976 pp.
353. Langguth, A. 1976. Mamiferos. Pp. 1-6, in Lista de las especies de vertebrados
del Uruguay (A. Langguth, ed.). Mus. Nac. Hist. Nat. Montev., 53 pp.
354. Langguth, A., and S, Anderson. 1980. Manual de identificacion de los mamif-
eros del Uruguay. Div. Publ. Edidones Univ. de la Republica, Montevideo, 65 pp.
355. Largen, M. J., D. Kock, AND D. W. Yalden. 1974. Catalogue of the mammals
of Ethiopia. 1. Chiroptera. Monit. ZooL Italiano, 5:221-298.
356. Larsen, H. 1945. La taxiderime modeme. Ed. de la Frigate, Geneve.
357. Laurie, E. M. O., AND J. E. Hill. 1954. List of land mammals of New Guinea,
Celebes, and adjacent islands, 1758-1952. British Mus. (Nat. Hist.) Publ., London,
175 pp.
358. Lawrence, B. 1939. Collections from the Philippine Islands. Mammals. Bull.
Mus. Comp. ZooL, 86:28-73.
359. Lawrence, M. J., AND R. W. Brown. 1973. Mammals of Britain. Their tracks,
trails and signs. Blandford Press, London, 297 pp.
360. Lay, D, M, 1967. A study of the mammals of Iran, resulting from the Street
Expedition of 1962-63. Fieldiana-ZooL, Chicago Mus. Nat. Hist., 54:1-282.
361. Lekagul, B., and J. A. McNeely. 1977. Mammals of Thailand. Assoc. Conserv.
Wild., Bangkok, 758 pp.
362. leLourarn, H., andM.-C. Saint-Girons. 1977. Les rongeurs de France. Faunis-
tique et biologic. Ann, ZooL Ecol. Anim., 1977:1-159.
363. Lemke, T. O., et al. 1982. Notes on opossums, bats, and rodents new to the
fauna of Colombia, Mammalia, 46:225-234,
192
Annals of Carnegie Museum
VOL. 55
364. Leopold, A. S. 1959. Mammals. Pp. 319-=528, in Wildlife of Mexico. The game
birds and mammals. Univ. Calif Press, Berkeley, 568 pp.
365. . 1977. Fauna silvestre de Mexico. Aves y Mamiferos de caza. Inst. Mex.
Recur. Nat. Renov., Mexico, 655 pp.
366. Lewis, R. E., and J. H. Lewis. 1968, A review of Lebanese mammals. Carnivora,
Pinnipedia, Hyracoidea, and Artiodactyla. J. Zool. (London), 154:517=531.
367. Lewis, R. E., J. H. Lewis, and S. I. Atallah. 1967. A review of Lebanese
mammals. Lagomorpha and Rodentia. J. Zool. (London), 153:45=70.
368. Lidicker, W. Z., Jr. 1978. A phylogeny of New Guinea rodent genera based on
phallic morphology. J. Mamm., 49:609=643.
369. Lidicker, W. Z., Jr., and W. I. Follet. 1968. Isoodon Desmarest, 1817, rather
than Thy lads Illiger 1 8 1 1 , as the valid generic name of the short-nosed bandicoots
(Marsupialia: Peramelidae). Proc. Biol. Soc. Washington, 81:251-256.
370. Lidicker, W, Z., Jr., AND A, C. Ziegler. 1968. Report on a collection of mammals
from eastern New Guinea, including species keys for fourteen genera. Univ. Cal-
ifornia Publ. Zool., 87:1-64.
371. Lopez DE Ceballos, E. 1974. Fauna de Venezuela y su conservacion. Ed. Arte,
Caracas, 170 pp.
372. Lucero, M. M. 1983. Lista y distribucion de aves y mamiferos de la provincia
de Tucuman, Fund. Miguel Lillo Misc., 75:1-61.
373. Lyneborg, L. 1970. Pattedyr i farver. Jorgen Lundo, Politiken, Kobenhavn,
239 pp.
374. . 1972. Wilde zoogdieren in Europa. Moussault, Amsterdam/Standaard,
Antwerpen. Dutch ed. by G. den Hoed.
375. Mack, C. 1961. Mammals from south-western Queensland. Mem. Queensland
Mus., 13:213-229.
376. Mackenzie, P. Z. 1954. Catalogue of wild mammals of the Sudan occurring in
the natural orders. Artiodactyla and Perissodactyla. Publ. Sudan Mus. (Nat. Hist.),
4:1-21.
377. Madureira, M, L,, and C. M. Paixao de Magalhaes. 1980. Small mammals
of Portugal. Arq. Mus. Bocage, 7:179-214.
378. Mahoney, J. A. 1968. Baiyankamys Hinton, 1943 (Muridae: Hydrominae), a
New Guinea rodent genus named for an incorrectly associated skin and skull
(Hydominae, Hydromys) and mandible (Murinae, Rattus). Mammalia, 32:64-71.
379. Mahoney, R. 1973. Laboratory techniques in zoology. Wiley, New York,
518 pp.
380. Malbrant, R., AND A. Maclatchy. 1949. Faune de I’Equateur africain frangais.
Mammiferes. Encycl. Biol., 36:1-323.
381. Mani, M. S. 1974. Biogeography and ecology in India. Dr. W. Junk b.v. Publ.,
The Hague, 773 pp.
382. Mann Fischer, G. 1957. Clave de determinacidn para las especies de mamiferos
silvestres de Chile. Invest. Zool. Chil., 4:89-126.
383. - . 1978. Los pequenos mamiferos de Chile. Gayana Zool., 40:1-342.
384. Mares, M. A. 1982. The scope of South American mammalian biology: Per-
spectives on a decade of research. Pp. 1-26, in Mammalian biology in South
America (M. A. Mares and H. H. Genoways, ed.), Pymatuning Laboratory of
Ecology, Spec. Publ. No. 6, Linesville, Pennsylvania.
385. — . 1985. Mammal and museum literature: An international survey. Acta
Zool. Fennica, 170:57-60.
386. In press. The need for popular natural history publications concerning
mammals. Occas. Papers, Zool, Survey of India.
387. Mares, M. A., R. A. Ojeda, and M. P. Kosco. 1981. Observations on the
distribution and ecology of the mammals of Salta Province, Argentina. Ann. Car-
negie Mus., 50:151-206.
1986
Mares and Braun —Mammalogy Literature
193
388. Mares, M. A., M. R, WiLLiG, K. E. Streilein, and T. E. Lacher, Jr. 1981. The
mammals of northeastern Brazil: A preliminary assessment. Ann. Carnegie Mus.,
50:81-=-137.
389. Marlow, B. J. 1962. Marsupials of Australia. Jacaranda Press, Brisbane,
141 pp.
390. Martino, V. 1930. Kljuc za odredivanje glodara [Keys for identification of ro-
dents]. Glasn. Min. poljoprivrede, Beograd, 8:113-136.
39 1 . Massoia, E. 1 976. Mammalia. Pp. 1-128, in Fauna de agua dulce de la Republica
Argentina, Vol. 44, Fundacion Educ. Cienc. Cult., Buenos Aires.
392. Mathews, L. H. 1982. Mammals in the British Isles. Collins, London, 207 pp.
393. McKean, J.L. 1972, Notes on some collections of bats (Order: Chiroptera) from
Papua-New Guinea and Bougainville Island. CSIRO Wildl. Res., 26:1-35.
394. McKean, J. L., and J. H. Calaby. 1968. A new genus and two new species of
bats from New Guinea. Mammalia, 32:372-378.
395. McKenna, Me, C. 1961. A note on the origin of rodents. Amer. Mus. Novitates,
2037:1-5.
396. Medway, Lord. 1964. Mammals of Borneo: An annotated checklist, Sarawak
Mus. J., 11:602.
397. — . 1965. Mammals of Borneo: Field keys and an annotated checklist. Sin-
gapore, Malaysian Branch of the Royal Asiatic Soc., Malaysia Printers, 193 pp.
398. -. 1969. The wild mammals of Malaya and offshore islands including Sin-
gapore. Oxford Univ. Press, London, 127 pp.
399 1977^ Mammals of Borneo: Field keys and an annotated checklist. Mon-
ogr. Malay. British R. As. Soc., 7:1-172.
400. — — , 1978. The wild animals of Malaya (peninsular Malaysia) and Singapore.
Oxford Univ, Press, Kuala Lumpur, 128 pp.
401. Meester, j., and H. W. Setzer (eds.). 1971-1977. The mammals of Africa: An
identification manual. Parts 1-5. Smithson, Inst. Press, Washington, D.C.
402. Mendez, E. 1970. Los principales mamiferos silvestres de Panama. Privately
printed in Panama, 283 pp.
403. Menzies, j. I. 1971. The lobe-lipped bat {Chalinolobus nigrogriseus Gould) in
New Guinea. Rec. Papua New Guinea Mus., vol, 1, part 2.
404. . 1973. A key to the rats of the genera Rattus, Melomys and Pogonome-
lomys on the New Guinea mainland. Occas. Papers Dept. Biol., Univ. Papua New
Guinea, 2:1-16.
405. — —. 1974. The status of Melomys platyops arfakiensis (Ruemmler) and the
races of Melomys rubex Thomas. Mammalia, 38:647-655.
406. Menzies, J. I., and E. Dennis. 1979. Handbook of New Guinea rodents. Hand-
book Wau Ecol. Instil, Wau, 6:1-68.
407. Menzies, J. L, and A, L. Rand. 1973. Results of the Archbold Expeditions. No.
17. Some original observations on the habits of Dactylopsila trivigata Gray. Amer.
Mus. Novitates, 957:1-7.
408. Metcalf, J. C. 1981. Taxidermy. Duckworth, London.
409. Meyer-Ohme, D. 1965. Die Saugetiere Afghanistans (Teil III): Chiroptera. Sci.,
Quart. J. Fac. Sci., Kabal Univ., (SuppL), August 1965:42-58.
410. Migula, P, 1979. [Mammals of the district Chrzanow. The insectivores and the
rodents.] Stud. Osrodka Dok. Fizjograficznej, 7:257-283.
411. Miller, G. S. 1912. Catalogue of the mammals of western Europe (Europe ex-
clusive of Russia) in collection of the British Museum. British Mus. (Nat. Hist.),
1019 pp.
412. Miller, S. D., and J. Rottmann. 1976. Guia para el reconocimiento de los
mamiferos chilenos. “Expedicion a Chile.” Ed, Gabriela Mistral, Santiago, Chile,
200 pp.
413. Miric, Dj. 1 970. Kljuci za dolocevanje zivali. Mammalia. [Keys for identification
194
Annals of Carnegie Museum
VOL. 55
of animals. Mammals.] Institu za biologijo Univerze v Ljubljani and Dmstvo
biologov Slovenije, Ed. Ljubljana.
414. Misonne, X. 1959. Analyse zoogeographique des mammiferes de I’lran. Mem.
Inst. R. Sci. Nat. Belg., 59(2): 1-1 57.
415. . 1977. Mammiferes du Jebel Uweinat, desert de Libye. Ann. Mus. R. Afr.
Cent. Ser. Quarto Zool., 217:1-44.
416. Mitchell, R. M. 1975. A checklist of Nepalese mammals (excluding bats). Sau-
getierk. Mitt., 23:152-157.
417. . 1978. A checklist of Nepalese bats. Saugetierk. Mitt., 26:75-78.
418. Moffat, C. B. 1938. The mammals of Ireland. Proc. R. Ir. Acad. Sect. B Biol.
Geol. Chem. Sci., 44B, 6:1-128.
419. Mok, W. Y., ET AL. 1982. Lista atualizada de quiropteros de Amazonia Brasileira.
Acta Amazonica, 12:817-823.
420. Moojen, J. 1952. Os roedores do Brasil. Minist. Educ. Cult., Inst. Nac. Livro,
Bibl. Cienc. Bras., Ser. A-II, Rio de Janeiro, 214 pp.
421. Morlok, W. F. 1978. Nagetiere ausderTurkei (Mammalia: Rodentia). Senckenb.
Biol., 59:155-162.
422. Morrison-Scott, T. C. S. 1952. A list of British mammals. British Mus. (Nat.
Hist.), 24 pp.
423. Mukherjee, A. K. 1982. Endangered animals of India. Zool. Survey India, Cal-
cutta, 122 pp.
424. Myers, P., and R. M. Wetzel. 1983. Systematics and zoogeography of the bats
of the Chaco Boreal. Misc. Publ. Mus. Zool., Univ, Michigan, 165:1-59.
425. Nadachowski, A. 1984. On a collection of small mammals from the People’s
Democratic Republic of Korea. Acta Zool. Cracov., 27:47-60.
426. Nadachowski, A., B. Rzebik-Kowalska, and A.-H. H. Kadhim. 1977. The
mammals of Iraq. Przegl. Zool., 21:170-181.
427. Nader, I. A. 1982. New distributional records of bats from the Kingdom of Saudi
Arabia (Mammalia: Chiroptera). J. Zool., London, 198:69-82,
428. Nadra, E. 1955. Colectarea conservarea si naturalizarea vertebratelos pentur
muczee. Ed. de Stat, Bucuresti.
429. Naumann, C., and G. Nogge. 1973. Die Grosssauger Afghanistans. Z. Kolner
Zoo, 16:79-93.
430. Niethammer, J. 1965. Die Saugetiere Afghanistans (Teil II): Insectivora, Lago-
morpha, Rodentia. Science (Fac. Sci., Kabul Univ.), August 1965:18-42.
431. -. 1974. Zur Verbrietung und Taxonomic griechischer Saugetiere. Bonn.
Zool. Beitr., 25:28-55.
432. Niethammer, J., and F. Krapp (eds.), 1978. Handbuch der Saugetiere Europas.
Band 1, Rodentia 1 (Sciuridae, Castoridae, Gliridae, Muridae). Akad. Verlag.,
Wiesbaden, 476 pp.
433. Niezabitowski, L. E. 1924. Mammalia-Ssaki. Pp. 77-99, in Podr§cznik do zbei-
rania i konserwowania zwierz^t nalez^cych do fauny polskeij. Pahstwowe Muzeum
Zoologiczne, zeszyt. Vol. 7.
434. -. 1933. lOucz do oznaczania zwierz^t ss^cych Polski. Koto Przyrodnikow
Uczniow UJ, Krakow, 1 24 pp.
435. NiLamhna, E. (ed.). 1979. Provisional distribution atlas of amphibians, reptiles
and mammals in Ireland. Foras Forbartha, Dublin.
436. Nores, C. 1978. Clave para la identificacion de craneos de los mamiferos ibericos
(excluidos los marinos). Publ. Univ. de Oviedo.
437. Novikov, G. A. 1956. [Carnivorous mammals of the fauna of the USSR.] Mos-
cow. English translation: Jerusalem, 1962, 284 pp.
438. Nunez Garduno, A., et al. 1981. Mamiferos silvestres de la region de El Tuito,
Jalisco, Mexico. An. Inst. Biol. (Mexico), Sec. Zool., 51:647-668.
1986
Mares and Braun— Mammalogy Literature
195
439. Ochoa G,, J., and C. Ibanez. 1985. Distributional status of some bats from
Venezuela. Mammalia, 49:65-73.
440. Ognev, S. I. 1928. Mammals of Eastern Europe and Northern Asia, Vol. 1,
Insectivora and Chiroptera, Moscow. English translation: Jerusalem, 1962.
44 1 ^ . 1931. Mammals of Eastern Europe and Northern Asia, Vol. 2, Carnivora
(Fissipedia). Moscow. English translation: Jerusalem, 1962.
442. — . 1935. Mammals of the USSR and adjacent countries, Vol. 3, Carnivora.
Moscow. English translation: Jerusalem, 1962.
443^ 1940. Mammals of the USSR and adjacent countries, Vol. 4, Rodents.
Moscow. English translation: Jerusalem, 1960.
444__ . 1947. Mammals of the USSR and adjacent countries, VoL 5, Rodents.
Moscow. English translation: Jerusalem, 1963.
445_ 1948. Mammals of the USSR and adjacent countries, Vol. 6, Rodents.
Moscow. English translation: Jerusalem, 1963.
446. . 1950. Mammals of the USSR and adjacent countries, Vol, 7, Rodents.
Moscow. English translation: Jerusalem, 1964.
447. Olrog, C. C., and M, M. Lucero. 1981. Guia de los mamiferos argentinos.
Minist. Cult. Educ. Fund. Miguel Lillo, Tucuman, 151 pp.
448. Omura, H. 1950. Whales in the adjacent waters of Japan. Sci. Rep. Whales Res.
Inst Tokyo, 4:27-113.
449. Ondrias, J. C. 1965. Die Saugetiere Griechenlands. Saugetierk. Mitt., 13:109-
127.
450. . 1966. The taxonomy and geographic distribution of the rodents of Greece.
Saugetierk. Mitt, 14(suppl.):l-136.
451. Osborn, D. I. 1964. The hare, porcupine, beaver, squirrel, jerboas, and dormice
from Turkey. Mammalia, 28:573-592.
452. — — . 1966. Rodents of subfamilies Murinae, Gerbillinae, and Cricetinae from
Turkey. J. Egypt. Public Health Assoc., 40:401-424.
453. Osborn, D. J., AND 1. Helm Y. 1980. The contemporary land mammals of Egypt
(including Sinai). Fieldiana-ZooL, Chicago Mus. Nat. Hist., 5(1 309): 1-579.
454. Osgood, W. H. 1943. The mammals of Chile. Field Mus. Nat His,, (ZooL Ser.),
30:1-268.
455. Palmeirim, J., M, J. Ramos, and D. Dias. 1979. Bats from Portugal in the
collection of Museu Bocage. Arq. Mus. Bocage, 7:53-66.
456. Panouse, j. B. 1951. Les chauve-souris du Maroc. Trav. Inst. Sci, Cherifien Ser.
ZooL, 1:1-120.
457. — . 1957. Les mammiferes du Maroc. Primates, Carnivores, Pinnipedes,
Artiodactyles. Trav. Inst. Sci. Cherifien. Zool., 5:1-210.
458. Papadopol,A. 1964. Confectionarea materioluiui didactic pentru stiinte naturaie.
Ed. Did si Fed., Bucuresti,
459. Parker, S. A. 1973. An annotated checklist of the native land mammals of the
Northern Territory. Rec. S. Australian Mus. (Adelaide), 16:1-57.
460. Patton, J. L., B, Berlin, and E. A. Berlin. 1982. Aboriginal perspectives of a
mammal community in Amazonian Peru: Knowledge and utilization patterns among
the Aguamna Jivaro. Pp. 1 1 1-128, in Mammalian biology in South America (M.
A. Mares and H. H. Genoways, eds.), Pymatuning Laboratory of Ecology, Special
Publ. Series Vol. 6, Linesville, Pennsylvania.
461. Pearson, O. P. 1951. Mammals in the highlands of southern Peru, Bull. Mus.
Comp. ZooL, 106:117-174.
462. — - — . 1957. Additions to the mammalian fauna of Peru and notes on some
other Peruvian mammals. Breviora, 73:1-7.
463. Pearson, O. P., and C. P. Ralph. 1978. The diversity and abundance of ver-
196
Annals of Carnegie Museum
VOL. 55
tebrates along an altitudinal gradient in Pern. Mem. Mus. Hist. Nat. “Javier Prado,”
18:1-97.
464. Pefaur, J. E., AND B. Caceres-Pefaur. 1976. Bibliografia sobre mamiferos te-
rrestres Chilenos: 1942-1972. Bol. Soc. Biol, de Concepcion, Tomo L: 157-1 72.
465. PelikAn, J., J. Gaisler, and P. Rodl. 1979. Nasi savci. [Our mammals.] Akad.
Nakhladatelstvi Ceskoslovenske Akad. Ved., Prague, 163 pp.
466. Peracchi, A. L., and S. T. de Albuquerque. 1971. Lista provisdria dos qui-
ropteros dos estados do Rio de Janeiro e Guanabara, Brasil (Mammalia, Chirop-
tera). Rev. Bras. Biol., 31:405-413.
467. Petranu, C. 1912. Muczelle din Transilvania, Banat, Crisana, si Maranures. Ed.
Cartea Romaneasca, Bucuresti.
468. Petrov, B. 1977. Review of the mammals of Serbia and major tasks of its
research. Arh. Biol. Nauka, 29:1 13-129.
469. Better, F. 1972. The rodents of Madagascar: The seven genera of Malagasy
rodents. Pp. 661-665, in Biogeography and ecology in Madagascar (R. Battistina
and G. Richard- Vindard, eds.), Dr. W. Junk b.v. Publ., The Hague, The Nether-
lands.
470. Phillips, W. W. A. 1980. Manual of the mammals of Sri Lanka. Part I. Wildlife
and Nature Protection Society of Sri Lanka, 116 pp.
471. PiECHOCKi, R. 1967. Saugetiere (Mammalia). Pp. 29-88, in Makroskopische Pra-
parations-technik Leitfaden fur das sammeln, preparieren und konservieren. Teil
I. Wirbeltiere (R. Piechocki, ed.), Akad. Verlag. Geest and Portig K.-G,, Leipzig.
472. Pine, R. H. 1973. Mammals (exclusive of bats) of Belem, Para, Brazil. Acta
Amazonica, 3:47-79.
473. . 1980. Key to the bats of Jamaica, Hispaniola and Puerto Rico on gross
and external characters. Caribbean J. Sci., 15:9-1 1.
474. . 1982. Current status of South American mammalogy. Pp. 27-38, in
Mammalian biology in South America (M. A. Mares and H. H. Genoways, eds.),
Pymatuning Laboratory of Ecology, Spec. Publ. No, 6, Linesville, Pennsylvania.
475. Pine, R. H., J. P. Angle, and D. Bridge. 1 978. Mammals from the sea, mainland
and islands at the southern tip of South America. Mammalia, 42:105-1 14.
476. Pine, R. H., I. R. Bishop, and R. L. Jackson. 1970. Preliminary list of mammals
of the Xavantina/Cachimbo Expedition (central Brazil). Trans. R. Soc. Trop. Med.
Hyg., 64:668-670.
477. Pine, R. H., S. D. Miller, and M. L. Schamberger. 1979. Contributions to the
mammalogy of Chile. Mammalia, 43:339-376.
478. PococK, R. I. 1939. The fauna of British India, including Ceylon and Burma.
Mammalia. Vol. I. Primates and Carnivora (in part). Families Felidae and Viverri-
dae. Taylor and Francis, Ltd., London, 463 pp.
479. . 1941. The fauna of British India, including Ceylon and Burma. Vol. 11.
Carnivora, suborders Aeluroidea (part) and Arctoidea. Taylor and Francis Ltd.,
London, 503 pp.
480. Poole, A. L. (ed.). 1973. Wild animals in New Zealand. Reed, Wellington,
151 pp.
481. Pop, I. andV. Homei. 1973. Mamifere din Romania. 2 Vols. Ed. Stunpfica-
Bucuresti.
482. Prater, S. H. 1948. The book of Indian mammals. Indian Nat. Hist. Ser. Vol.
II, Bombay, 263 pp.
483. Prescott, R. T. M. 1954. Collections of a century. The history of the first hundred
years of the National Museum of Victoria. National Museum, Melbourne.
484. PucEK, Z. (ed.) 1981. Klucze do oznaczania kr§gowc6w Polski. Ssaki-Mammalia.
[Keys to vertebrates of Poland. Mammals.] PWN-Polish Scientific Publishers, War-
szawa, 367 pp.
1986
Mares and Braun— Mammalogy Literature
197
485. Punt, A., P. J. H. VON Bree, J. DE Vlas, AND G. J. WiERSEMA. 1974. De Neder-
landse vieermuizen, [The bats in the Netherlands.] Wet. Meded. K.N.N. V, (K. Ned.
Natuurhist. Ver.), 104:1-48.
486. Qumsiyeh, M. B., and D. A, Schlitter, 1 982. The bat fauna of Jabal Al Akhdar,
northeast Libya. Ann. Carnegie Mus., 51:377-389.
487. Rabor, D. S. 1977. Philippine birds and mammals. Sci. Educ. Center, Univ.
Philipp. Press, Quezon City, 284 pp.
488. Rabor, D. S., A. C. Alcala, and R. B. Gonzales. 1970. A list of the land
vertebrates of Negros Island, Philippines. Silliman J., 17:297-316.
489. Rahm, U. 1966. Les mammiferes de la foret equatoriale de Test du Congo. Ann.
Mus. R. Afr. Centr. Ser. Quarto ZooL, 149:38-121.
490. =•. 1976. Die Saugetiere der Schweiz. Veroff. Naturhist. Mus. Basel, 9:1-87.
491. Ramirez-Pulido, J., et al. 1982. Catalogo de los mamiferos terrestres nativos
de Mexico. Ed. Trillas, Mexico City, 126 pp.
492. . 1983. Lista y bibliografias reciente de los mamiferos de Mexico. Univ.
Auto. Metro. Unidad Iztapalapa, Itzapalapa, 363 pp.
493. Ramirez-Pulido, J., AND M, C. Britton. 1981. An historical synthesis of Mex-
ican mammalian taxonomy. Proc. Biol. Soc. Washington, 94:1-17.
494. Ranck, G. L. 1968. The rodents of Libya: Taxonomy, ecology, and zoogeograph-
ical relationships. Bull. U.S. Nat. Mus., 275:1-264.
495. Rau, j. R., and M. E. Kennedy. 1981. Adenda y actualizacion a la bibliografia
sobre mamiferos Chilenos de Pefaur y Caceres, 1976. Bol. Soc. Biol. Concepcion,
52:135-143.
496. Reig, O. A. 1964. Roedores y marsupiales del partido de General Pueyrredon y
regiones adyacentes (Provincia de Buenos Aires, Argentina). Publ. Mus. Munic.
Cienc. Nat. Mar del Plata, 1:203-224.
497. Reise, D. 1973. Clave para la determinacidn de los craneos de marsupiales y
roedores chilenos. Gayana Zool., 27:1-20.
498. Renji, L., andD. Yongwen. 1984. [Bats from South Anhui]. Acta Theriol. Sinica,
4:321-328.
499. Ride, W, D. L, 1962. On the use of generic names for kangaroos and wallabies
(subfamily: Macropodinae). Australian J. Sci., 24:367-372,
500. Ride, W. D. L. 1970. A guide to the native mammals of Australia. Oxford Univ.
Press, London, 249 pp.
501. Ritchie, J. 1920. The influence of man on animal life in Scotland: A study of
faunal evolution. Cambridge Univ. Press, 550 pp.
502. Robbins, C. B. 1980. Small mammals of Togo and Benin. 1. Chiroptera. Mam-
malia, 44:83-88.
503. Roberts, A. 1951. The mammals of South Africa. Central News Agency, South
Africa, 700 pp.
504. Roberts, G., and G. Tunnicliffe. 1975. Wild animals of New Zealand. A
Microlone Colour Book, Bascands Ltd., Christchurch.
505. Roberts, T. J. 1977. The mammals of Pakistan. Ernest Benn Ltd., London,
361 pp.
506. Rode, P., and R. Didier. 1946. Atlas des mammiferes de France. N. Boubee
and Co., Paris, 219 pp.
507. Roer, H. 1978, Die Fledermaus-fauna von Sudwestafrika. 1. Teil: Microchirop-
tera, J. S. West African Sci. Soc., 32:131-179.
508. Rohl, E. 1949. Mammals. Pp. 4-140, in Fauna descriptiva de Venezuela. Bol
Acad. Cienc. Caracas, 12(36-37): 1-495.
509. Romer, j. D. 1974. Annotated checklist with keys to the bats of Hong Kong.
Mem. Hong Kong Nat. Hist. Soc., 9:1-6.
198
Annals of Carnegie Museum
VOL. 55
5 1 0. Rosevear, D. R. 1953. Checklist and atlas of Nigerian mammals with a foreword
on vegetation. Nigerian Government Printer, Lagos, 133 pp.
51 P _____ 1965. Bats of West Africa. British Mus. (Nat. Hist,), 418 pp.
512. — — . 1969. Rodents of West Africa. British Mus. (Nat. Hist), PubL, 677:1-
604.
513_ _____ 1974, The carnivores of West Africa. British Mus. (Nat. Hist), 548 pp.
514. Ross, G. J. B. 1984. The smaller cetaceans of the southeast coast of southern
Africa. Ann. Cape Prov. Mus. Nat Hist, 15:173-410.
515. Rowe-Rowe, D. T. 1978. The small carnivores of Natal. Lammergeyer,
25:1-48.
516. Ryan, P. F. 1968. A guide to Insectivora and Dermoptera of South Vietnam.
Formosan Sci., 22:79-92.
517. Saemundsson, B. 1932. Spendyrin (Mammalia Islandiae). [Icelandic animals. IT
Mammals.] Sigfus Eymundsson, Reykjavik.
518. Saemundsson, B., and M. Degerbol. 1 939. Mammals. Pp. 1-52, in The zoology
of Iceland. 4. E. Munksgaard, Copenhagen and Reykjavik, 76 pp.
519. Saint-Girons, M.-C. 1973. Les mammiferes de France et du Benelux (faune
marine exceptee). Doin, Paris, 48 1 pp.
520. Saint-Girons, M.-C., and F. Fetter. 1965. Les rongeurs du Maroc. Trav. Inst.
Sci. Cherifien Ser. ZooL, 31:1-55.
521. Sanborn, C. C. 1931. Bats from Polynesia, Melanesia and Malaysia. Field Mus.
Nat. Hist Publ, Zool. Ser., 18:1-29.
522. — — — . 1940, Mammals from Iraq. Appendix F. Pp. 156-162, in The anthro-
pology of Iraq (H, Field, ed.), The upper Euphrates. Field Mus. Nat. Hist. Anthro,
Ser. 30 Pt. 1 No. 1.
523. — . 1952. Philippine Zoological Expedition, 1947-48. Mammals. Fieldiana-
ZooL, Chicago Mus. Nat, Hist., 33:87-158.
524. Sanborn, C. C., and H. Hoogstraal. 1953, Some mammals of Yemen and
their ectoparasites. Fieldiana-ZooL, 34:229.
525. 1955. The identification of Egyptian bats. J. Egypt. Public Health Assoc.,
30:103-121.
526. Schafer, E. 1955. Mamiferos venezolanos. Rev. Shell (Caracas), 4(17):36-40.
527. Schafer, W. 1972. Leme in Museum. Frankfurt.
528. Schlitter, D, A., L. W. Robbins, and S, A. Buchanan. 1 982. Bats of the Central
African Republic (Mammalia: Chiroptera). Ann. Carnegie Mus., 51:133-155.
529. ScHMiDLY, D. J., and F. S. Hendricks. 1 984. Mammals of the San Carlos Moun-
tains of Tamaulipas, Mexico. Spec. Publ. Mus., Texas Tech Univ., 22:15-69.
530. Schneider, R. 1977. A mammalian inventory of St Paul Subterranean National
Park, Phillipines. Tigerpaper, 4:29.
531. Schouteden, H. 1 944. De zoogdieren van Belgisch Congo en van Ruanda-Uran-
di. 1. Primates, Chiroptera, Insectivora, Pholidota. Ann. Mus. Congo Beige Ser.
ZooL, 3:1-168.
532. — _____ 1945. De zoogdieren van Belgisch Congo en van Ruanda-Umndi. 2.
Carnivora (2), Ungulata (1). Ann, Mus. Congo Beige Ser, Zool., 3:169-332.
533. Serventy, D. L. 1979. History of zoology in western Australia. J. R. Soc. West.
Australia, 62:33-43.
534. Setzer, H. W. 1956. Mammals of the Anglo-Egyptian Sudan. Proc. U.S. Natl.
Mus., 106:447-587.
535. — — — . 1957. A review of Libyan mammals. J. Egypt Public Health Assoc., 32:
41-82.
536. — — . 1958a. The jerboas of Egypt J. Egypt Public Health Assoc., 33:87-94.
537_ _____ i95g^. The hares of Egypt J. Egypt Public Health Assoc., 33:145-151,
538. — — , 1958c. The mustelids of Egypt. J. Egypt. Public Health Assoc., 33:199-
204.
1986
Mares and Braun —Mammalogy Literature
199
539 — \95M. The gerbils of Egypt. J. Egypt. Public Health Assoc., 33:205--227.
540, , 1959. The spiny mice {Acomys) of Egypt. J. Egypt. Public Health Assoc.,
34:93-101.
541 — 1961^. The jirds (Mammalia: Rodentia) of Egypt. J. Egypt, Public Health
Assoc., 36:8 1“92.
542. — ^ — . 1961^. The canids (Mammalia) of Egypt. J. Egypt. Public Health Assoc.,
36:113-118.
543. Shidlovskyi, M. V. 1962, [Key to the rodents of Zacaucasia.] Tbilisi.
544. Shortridge, G. C. 1934. The mammals of South West Africa. A biological
account of the forms occurring in that region. Heinemann, London, 2 vols.,
779 pp.
545. Shrestha, T. K. 1983. Wildlife of Nepal: A study of renewable resource of Nepal
Himalaya. Curriculum Development Centre, Tribhuvan Univ,, Kathmandu, Ne-
pal, 734 pp.
546. SiDDiQi, M. S. 1961. Checklist of mammals of Pakistan with particular reference
to the mammalian collection in the British Museum (Natural History), London.
Biologia, 7:93-225.
547. SiivoNEN, L. 1972. Suomen nisakkaat. [Mammals of Finland.] Vol. I and 11.
Kustannusosakeyhtio Otava, Helsinkii, 474 pp.
54g. — \916a. Nordeuropas daggdjur. P, A. Norstedt and Soners Forlag, Stock-
holm, 194 pp.
549 — __ — . 1976^. Pohjolan nisakkaat. Otava, Keumu, Finland, 194 pp.
550. Silva, F. 1 984. Mamiferos silvestres Rio Grande do Sul. Publ. Fund. Zoobotanica
do Rio Grande do Sul, Porto Alegre, 245 pp.
551. Silva, T., G. 1979, Los murcielagos de Cuba: Taxonomia, morphologia, distri-
bucion, biogeografia, ecologia, importancia economica. Ed. Academia, La Habana,
Cuba, 423 pp.
552. Skaren, U., and A. Kaikusalo. 1966. Suomen pikkunisakkaat [Small mammals
of Finland]. Otava, 227 pp.
553. Skuratowicz, W. 1947. Klucz do oznaczania krajowych zweirz§t ss^cych. Ksi?-
gamia Akademicka, Poznan, 67 pp.
554. Sludskii, A. A., A. Bekenov, ET AL. 1977. [Mammals of Kazakhstan. Rodents
(except the marmots, susliks, long-clawed ground squirrels, gerbils, and voles), Vol.
1, pt. 2.] Nauka, Alma-Ata, 536 pp.
555. Sludskii, A. A., S. N. Varshavskii, M. I. Ismagilov, V. 1. Kapdonov, and I. G.
Shubin. 1969. Mammals of Kazakhstan. Vol. 1, pt. 1: Rodents (marmots and
ground squirrels). Nauka, Alma-Ata, 453 pp.
556. Smithers, R. H. N, 1966. The mammals of Rhodesia, Zambia and Malawi.
Collins, London, 159 pp.
557 1968. A check-list and atlas of the mammals of Botswana (Africa). Trust-
ees of the Nat. Mus. of Rhodesia, Salisbury, 169 pp.
558. — . 1971. Mammals of Botswana. Mus, Mem. No. 4. Trustees of the Nac.
Museums of Rhodesia, Salisbury, 340 pp.
559 _ — — ^ 1973<2. Mammals: List of species known to occur in Rhodesia. Nat. Mus,
and Monuments of Rhodesia.
560. — — — . 1973^. Mammals: The preparation of museum study skins. Trustees of
the Nat, Mus. and Monuments of Rhodesia, Causeway, Rhodesia.
561. — 1975. Guide to the rats and mice of Rhodesia. Nat. Mus. and Monuments
of Rhodesia, Salisbury, Rhodesia.
562. — . 1983. The mammals of the southern African subregion. Univ. Pretoria,
Pretoria, South Africa, 736 pp.
563. Smithers, R. H. N., and J. L. P. LobAo Tello. 1976. Checklist and atlas of the
mammals of Mozambique, Mus. Mem. Trust Nat. Mus. Monum. Rhodesia, 8: 1-
184.
200
Annals of Carnegie Museum
VOL, 55
564. Smithers, R. H. N., and V. J. Wilson, 1979. Checklist and atlas of the mammals
of Zimbabwe Rhodesia. Mus. Mem. Trust Nat. Mus. Monum, Rhodesia, 9:1“-193,
565. SouKUP, J. 1961 a. Materiales para el catalogo de los mamiferos Peruanos. Biota,
3:240-276.
566. . 19611?. Materiales para el catalogo de los mamiferos Peruanos, Biota, 3:
277-324.
567. . 1961c. Materiales para el catalogo de los mamiferos Peruanos, Biota, 3:
325-331.
568. Southern, H. N. (ed,), 1964. The handbook of British mammals. Blackwell,
Oxford, 465 pp.
569. SPSS Inc. 1983. SPSS X useTs guide. McGraw Hill, New York.
570. Starck, D. 1974. Die Saugetiere Madagaskars, ihrs Lebenstraume und ihre Ge-
schichte. Sitzungber. Ges. J. G. Goethe-Univ., Frankfort am Main, 1 1:67-124.
571. Starrett, a. 1962. The bats of Puerto Rico and the Virgin Islands, with a
checklist and keys for identification. Caribbean J. Sci., 2:1-7.
572. — — — . unpubl. ms. Keys to the bats of Costa Rica, based primarily on external
characters. Xeroxed.
573. Sterndale, R. A. 1884. Natural history of the mammalia of India and Ceylon.
Thacker, Spink and Co., Calcutta, 540 pp.
574 _= — . 1982. Natural history of the mammalia of India and Ceylon. Himalayan
Books, New Delhi, 540 pp.
575. Straeten, E. van der, 1979. Tabel voor broohbollenandersoek. Tumhout,
57 pp.
576. Strahan, R. (ed.). 1983. Complete book of Australian mammals. Angus and
Robertson PubL, London, Sydney, and Melbourne, 530 pp.
577. Strestha, T. K. 1983. Wildlife of Nepal. Curriculum Development Centre, Trib-
huvan Univ., Kathmandu, Nepal, 734 pp.
578. Stroganov, S. U. 1949. [Keys to the mammals of Karelia.] Petrozavdsk (NV).
579_ __ — __ 1957, [Animals of Siberia: Insectivores.] Moscow.
580. . 1962. [Animals of Siberia: Carnivora.] Moscow. English translation: Je-
rusalem, 1969.
581. Stuart, C. T. 1975. Preliminary notes on the mammals of the Namib Desert
Park. Madoqua (Ser. 2), 4(74-80):5-68.
582. Swynnerton, G. H., and R. W. Hayman. 1951. A checklist of the land mammals
of the Tanganyika Territory and the Zanzibar Protectorate. J. East African Nat.
Hist. Soc. Nat. Mus., 20:274-392.
583. Taborsky, K. 1961. Metodika zoologickych praci v muzekh. 2 dily. Praha.
584. Taddei, V. A. 1974. Phyllostomidae (Chiroptera) do Norte-Ocidental do Estado
de Sao Paulo. IT Glossophaginae, Carolliinae, Stumirinae. Cienc. Cult. (Sao Paulo),
27(7):723-734.
585. Taibel,A. M. 1977. Mammiferi del Guatemala, con speciale riquando alia regione
del Peten, raccolti del maggio al settembre 1932. [Mammals of Guatemala, with
particular regard to the Peten region, collected from May to September 1932,] Atti
Soc. Itai. Sci. Nat Mus. Civ. Stor. Nat. Milano, 118:379“"40L
586. Talice, R, V. 1969. Mamiferos autoctonos. Ed. Nuestra Tierra, Montevideo,
68 pp.
587. Tamayo, F. 1961. Mamiferos de los llanos de Venezuela, El FaroL (Caracas),
23(196):12-19.
588. Tamayo H,, M., and D. Frassinetti C. 1980. Catalogos de los mamiferos fosiles
y vivientes de chile. Bol. Mus. Nac. Hist. Nat. Chile, 37:323-399.
589. Tatarinov, K. A. 1956. [Wild animals of the western regions of the Ukraine].
Akad. Sci. Ukr. RSR, Kiev, 187 pp.
590. Tate, G. H. H. 1936. Some Muridae of the Indo-Australian region. Bull. Amer.
Mus. Nat Hist, 80:221-297.
1986
Mares and Braun —Mammalogy Literature
201
59 1 . 1 939. Results of the Archbold Expeditions No. 24. Oriental Rhinolophus,
with special reference to materials from Archbold Collections. Amer. Mus. Novi-
tates, 1035:1==-12.
592. — — — . 1941^. Results of the Archbold Expeditions No. 35. A review of the
genus Hipposideros with special reference to Indo-Australian species. Bull. Amer.
Mus. Nat. Hist, 78:353-393.
593, ■ — — . 1941^. Results of the Archbold Expeditions No. 36. Remarks on some
old-world leaf-nosed bats. Amer. Mus. Novitates, 1 140:1-1 1.
594 ____ — 1941c. Results of the Archbold Expeditions No. 39. Review of Myotis
of Eurasia. Bull. Amer. Mus. Nat Hist., 78:537-565.
595 ____ — \94\d. Results of the Archbold Expeditions No. 40. Notes on Vesper-
tilionid bats. Bull. Amer. Mus. Nat. Hist., 78:567-597.
595 — . \942a. Results of the Archbold Expeditions No. 47. Review of the Ves-
pertilionine bats, with special attention to genera and species of the Archbold
Collections. Bull. Amer. Mus. Nat. Hist., 80:221-297.
597. . \9A2b. Results of the Archbold Expeditions No. 48. Pteropodidae (Chi-
roptera) of Archbold Collections. Bull. Amer. Mus. Nat. Hist., 80:331-347.
598. 1947. Results of the Archbold Expeditions No. 56. On the anatomy and
classification of the Dasyuridae (Marsupialia). Bull. Amer. Mus. Nat. Hist., 88:1-
155.
599 1 94g Results of the Archbold Expeditions No. 59. Studies on the anatomy
and phylogeny of the Macropodidae (Marsupialia). Bull. Amer. Mus. Nat. Hist.,
91:233-352.
500. — — 1949. Results of the Archbold Expeditions No. 61. A new species of
Rattus from the Snow Mountains of Netherlands New Guinea. Amer. Mus. Nov-
itates, 1421:1-3.
601. . 1951. Results of the Archbold Expeditions No. 65, The rodents of Aus-
tralia and New Guinea. Bull. Amer. Mus. Nat. Hist., 97:183-430.
602. Tate, G. H. H., and R. Archbold. 1936. Results of the Archbold Expedition
No, 8. Four apparently new polyprotodont marsupials from New Guinea. Amer.
Mus. Novitates, 823:1-4.
503. — .. 1938. Results of the Archbold Expeditions No. 18. Two new Muridae
from the western division of Papua. Amer. Mus. Novitates, 982:1-2.
504. — . 1939. Results of the Archbold Expeditions No. 23. A revision of the
genus Emballonura (Chiroptera). Amer. Mus. Novitates, 1035:1-14.
605. 1941a. Results of the Archbold Expeditions No. 31. New rodents and
marsupials from New Guinea. Amer, Mus. Novitates, 1101:1-9.
606. . I94lb. Results of the Archbold Expeditions No. 37. Notes on oriental
Taphozous and allies. Amer. Mus. Novitates, 1 141:1-5.
607. Tattersall, 1. 1982. The primates of Madagascar. Columbia Univ. Press, New
York, 382 pp.
608. Taylor, E. H. 1934. Philippine land mammals. Phillip. Bur. Sci. Monogr., 30:
1-548.
609. Terborgh, J. W., J. W. Fitzpatrick, and L. Emmons. 1 984. Annotated checklist
of bird and mammals species of Cocha Cashu Biological Station, Manu National
Park, Peru. Fieldiana, Zoology, 21:1-29.
610. Thomson, K. W. 1981. Art galleries and museums in New Zealand. A. H, Reed,
Wellington.
611. Thonglongya, K. 1 974. The history of mammalogy in Thailand. Nat. Hist. Bull.
Siam Soc., 25(3-4):53-67.
612. Thorvaldsson, O. 1960. Hreindyr a islandi, 1771-1960. [Reindeer in Iceland,
1771-1960.] Bokautgafa Menningarsjods, Reykjavik, 111 pp.
613. Tikader, B. K. 1983. Threatened animals of India. Zool. Survey India, Calcutta,
307 pp.
202
Annals of Carnegie Museum
VOL. 55
614. Timm, R. M., and E. C. Birney. 1980. Mammals collected by the Menage Sci-
entific Expedition to the Philippine Islands and Borneo. J. Mamm., 61:566-571.
615. Toktosunov, A. 1958. [Rodents of the Kirghiz.] Akad. Nauk Kirghizian SSR
Frunze, 172 pp.
616. Tokuda, M. 1941. A revised monograph of the Japanese and Manchou-Korean
Muridae. Biogeographica, 4:1-155.
617. Tomilin, a. G. 1957. Mammals of the USSR and adjacent countries, Vol. 9,
Cetacea. Moscow. English translation: Jerusalem, 1967.
618. Tompsett, D. H. 1970. Anatomical techniques. E. and S. Livingstone, London.
619. Toschi, a. 1965. Faune dTtalia, Mammalia: Lagomorpha, Rodentia, Carnivora,
Artiodactyla, Cetacea. Vol. VII. Edizioni Calderini, Bologna, 648 pp.
620. Toschi, A., AND B. Lanza. 1959. Fauna dTtalia. Vol. IV. Mammalia: Generalita,
Insectivora, Chiroptera. Accad. Naz. ent:Unione Zool. Ital., Bologna, 488 pp.
621. Tovar, S. A. 1971. Catalogo de mamiferos Peruanos. An. Cient. (La Molina),
9(1-2): 18-37.
622. Troughton, E. 1967. The furred animals of Australia. Angus and Robertson,
Sydney.
623. Tuttle, M. D. 1970. Distribution and zoogeography of Peruvian bats with com-
ments on natural history. Univ. Kansas Sci. Bull., 41:45-86.
624. Tweedie, M. W. F. 1978. Mammals of Malaysia. Longman, Malaysia, 87 pp.
625. Urbano ViDALES, G., andO. Sanchez-Herrera. 1983. Type specimens of mam-
mals in the collection of the Institute of Biology, National University of Mexico.
Occas. Papers Mus., Texas Tech Univ., 87:1-7.
626. Valerio, C. E. 1980. Anotaciones sobre la historia natural de Costa Rica. Ed.
Univ. Estatal a Distancia, San Jose, Costa Rica, 152 pp.
627. Valtysson, H. 1945. [In reindeer country.] Akureyri.
628. Van Deusen, H. M. 1957, Results of the Archbold Expeditions. No. 76. A new
species of wallaby (Genus Dorcopsis) from Goodenough Island, Papua. Amer. Mus.
Novitates, 1826:1-25.
629. . 1960. Notes on the marsupial feather-tail glider of Australia. J. Mamm.,
41:263-264.
630. . 1969. Results of the 1958-1959 Gilliard New Britain Expedition. 5. A
new species of Pteropus (Mammalia: Pteropodidae) from New Britain, Bismarck
Archipelago. Amer. Mus. Novitates, 2371:1-16.
631. . 1978. Results of the Archbold Expeditions. No. 101, Summary of the
seventh Archbold expedition to New Guinea (1964). Amer. Mus. Novitates, 2660:
1-21.
632. Van Deusen, H. M., and G. G. George. 1969. Results of the Archbold Expe-
ditions, No, 90. Notes on the echidnas (Mammalia: Tachyglossidae) of New Guinea.
Amer. Mus. Novitates, 2383:1-23.
633. Van Deusen, H. M., and K. F. Koopman. 1971. Results of the Archbold Ex-
peditions No. 95. The genus Chalinolobus (Chiroptera Vespertilionidae). Amer.
Mus. Novitates, 2468:1-30.
634. Van Deusen, H. M,, and R. F. Peterson, 1958. Chiroptera of New Guinea.
Nat. Hist, New York, 67:452-459.
635. Van Peenen, P. F. D., P. F. Ryan, and R. H. Light. 1969. Preliminary iden-
tification manual for mammals of South Vietnam. Smithson. Inst., Washington,
310 pp.
636. Varona, L. S. 1974. Catalogo de los mamiferos vivientes y extinguidos de las
Antillas. Acad. Cienc. Cuba Ser. Biol., 139 pp.
637. Vereshchagin, N. K. 1959. [The mammals of the Caucasus. History of the
formation of the fauna]. Akad. Nauk SSSR. English translation: Jerusalem, 1967,
816 pp.
1986
Mares and Braun— Mammalogy Literature
203
638. Vericad, J. R. 1972. Estudio faunistico y biologico de los mamiferos montaraces
del Pirineo. Publ. Cent. Pir. Biol. Exp., 4:7-229.
639. Verschuren, J. 1978. Les grandes mammiferes du Burundi. Mammalia, 42:209-
224.
640. Vieira, C. 1942. Ensaio monografico sobre os quiropteros do Brasil. Arq. ZooL
(Sao Paulo), 3:219-471.
541 1950. Xenartros e marsupials do Estado de Sao Paulo. Arq. Zool. (Sao
Paulo), 7:325-362.
542. — — 1953. Roedores e Lagomorfos do Estado de Sao Paulo. Arq. Zool. (Sao
Paulo), 8:129-168.
643. 1955. Lista remissiva dos mamiferos do Brasil. Arq. Zool. (Sao Paulo),
8:341-474.
644. Villa-R., B. 1949. Mamiferos del Soconusco, Chiapas. An. Inst. Biol. Univ. Nac.
Auton. Mex. Ser. Zool., 19:485-528.
645. -. 1952. Los mamiferos silvestres del valle de Mexico. An. Inst. Biol. Univ.
Nac. Auton. Mex. Ser. Zool., 23(l-2):269-492.
646. — . 1963. Sugerencias para preparar en el campo ejemplares de pequenos
mamiferos. Esc. Normal Sup. Mexico, 44 pp.
547^ — 1966. Los murcielagos de Mexico: Su importancia en la economia y la
salubridad— su clasificacion sistematica. An. Inst. Biol. Univ. Nac. Auton. Mex.
Ser. Zool., 491 pp.
648. Villa-R., B., and F. F. Biagi. 1959. Clave para familias de mamiferos Mexicanos.
Acta Zool. Mex., 3:1-8.
649. Vinogradov, B. S., AND A. 1. Argiropulo. 1941. [Fauna of the USSR. Mammals
key to rodents.] Moscow-Leningrad Pub., 241 pp.
650. Vinogradov, B. S., and I. M. Gromov. 1952. [Rodents of the USSR fauna.]
Tabl. Analyt. Fauna USSR, No. 48:1-296.
651. VizoTTO, L. D., andV. A. Taddel 1973. Chave para determinacao de quiropteros
brasileiros. Bol. Cienc. (Sao Jose do Rio Preto), 1:1-72.
652. Wagstaff, R., and J. H. Fidler (eds.). 1 968. The preservation of natural history
specimens, Vol. 11. Zoology-Vertebrata. H. F. and G. Witherby Ltd., London,
404 pp.
653. Waithman, J. 1979. A report on a collection of mammals from Southwest Papua,
1972-1973. Aust. Zool, 20:313-326.
654. Wakefield, N. A. 1974. Mammals of western Victoria. Pp. 35-50, in Proceedings
of the symposium, the natural history of western Victoria (M.H. Douglas and L.
O’Brien, eds.). Australian Inst. Agric. Sci., Horsham, 1 16 pp.
655. Walker, E. P., ET AL. 1964. Mammals of the world. Johns Hopkins, Baltimore,
1500 pp.
656. Wallin, L. 1969. The Japanese bat fauna. Zool. Bidr. Uppsala, 37:223-440.
657. Wassif, K. 1953. On a collection of mammals from northern Sinai. J. Instit.
Egypt Desert, 2:107-1 19.
658. — . 1954. On a collection of mammals from northern Sinai. Bull. Ins. Desert
d’Egypte, 3:107-118.
559^ _____ 1956. Studies on gerbils of the subgenus Dipodillus recorded from Egypt.
Ain Shams Sci. Bull., 1:173-199.
660. Wassif, K., and H. Hoogstraal. 1954. The mammals of south Sinai, Egypt.
Proc. Egypt. Acad. Sci., 9:63-79.
661. Watkins, L. C., J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco,
Mexico. Spec. Publ. Mus., Texas Tech Univ., 1:1-44.
662. Watts, C. H. S ., and H. J. Aslin. 1980. Australian mammals: A field guide for
New South Wales, Victoria, South Australia and Tasmania. Nelson, Melbourne,
321 pp.
204
Annals of Carnegie Museum
VOL. 55
663. 1981. The rodents of Australia. Angus and Robertson, Sydney, 321 pp,
664. Weber, M. 1927. Die Saugetiere: Einfuhrung in die Anatomie und systematik
der recenten und fossilen Mammalia. Zweite Aufl. Bd i Anatomischer Teil unter
Mitwirkung von Dr. H. M. deBurlet. Jena (Fischer), 444 pp.
665. . 1928. Die Saugetiere: Einfuhrung in die Anatomie und systematik der
recenten und fossilen Mammalia. Zweite Aufl. Bd i Anatomischer Teil unter Mit-
wirkung von Dr. O. A’bel. Jena (Fischer), 898 pp.
666. Webster, W. D., AND C. M. Fugler. 1984. Lista de quiropteros de las regiones
nortenas de Bolivia, Mus. Natl. Hist. Nat. La Paz Comunic., 3:13-19.
667. Webster, W. D., and J. K. Jones, Jr. 1984. Notes on a collection of bats from
Amazonian Ecuador. Mammalia, 48;247-252.
668. Webster, W. D., and W. B. McGillivray. 1984. Additional records of bats
from French Guiana, Mammalia, 48:463-465.
669. Weigel, I. 1969. Systematische Ubersicht uber die Insektenfresser und Nager
Nepals nebst Bemerkungen zur Tiergeographie. Ergeb. Forsch Untemehmens Nepal
Himalaya, 3:149-196.
670. Wettstein-Westersheimb, O. 1955. Mammalia. Pp. 1-16, in Catalogus Faunae
Austriae, Teil 21c, Strouhal.
671. Wetzel, R. M., and J. W. Lovett. 1974. A collection of mammals from the
Chaco of Paraguay. Occas. Papers, Univ. Connecticut Biol. Sci. Ser., 2:203-216.
672. Williams, S. L., and H. H, Genoways. 1980. Results of the Alcoa Foundation-
Suriname Expeditions. VI. Additional records of bats (Mammalia: Chiroptera) from
Suriname. Ann. Carnegie Mus., 49:213-236.
673. Williams, S. L., H. H. Genoways, and J. A. Groen. 1983. Results of the Alcoa
Foundation-Suriname Expeditions. VII. Records of mammals from central and
southern Suriname. Ann. Carnegie Mus., 52:329-336.
674. Wilson, V. J. 1975. Mammals of the Wankie National Park, Rhodesia. Natl.
Mus. Monum. Mem. Bulawayo, Zimbabwe, 15:1-147.
675. Wingate, L. R. and P. Swanepoel. 1981. Collections of recent mammals in
southern Africa. Ann. Cape Prov. Mus., 13:169-185.
676. Winter, J. 1984. Mammals of the wet tropics of Australia. Wild. Australia, 2 1 :
24-28.
677. WoDziCKi, K. A. 1950. Introduced mammals of New Zealand: An ecological and
economic study. Bull. Dep. Sci. Ind. Res., Wellington, 98:1-255.
678. WoLoszYN, D., AND B. W. WoLOSZYN. 1982. Los mamiferos de la Sierra de la
Laguna, Baja California Sur. Consejo Nacional de Ciencia y Tecnologia, Circuito
Cultural, Centro Cultural Universitario, 04515 Mexico, D.F., Mexico, 168 pp.
679. Won, P.-H. 1967. Illustrated encyclopedia of fauna and flora of Korea. Vol. 7.
Mammals. Minist. Educ., Seoul.
680. Won, P.-O. 1976. Checklist of the mammals of Korea. Inst. Omithol., Kyung
Hee Univ,, Seoul.
681. Wroughton, R. C. 1918. Summary of the results from the mammal survey. J.
Bombay Nat. Hist, Soc., 25:547-598.
682. — — . \9\9a. Summary of the results from the mammal survey. J. Bombay
Nat. Hist. Soc., 26:19-58.
683. — — ■ — . 1919/?. Summary of the results from the mammal survey. J. Bombay
Nat. Hist. Soc., 26:338-379.
684. - — — . 1919c. Summary of the results from the mammal survey. J. Bombay Nat.
Hist. Soc., 26:776-802.
685. . 1919(7. Summary of the results from the mammal survey. J, Bombay
Nat. Hist. Soc., 26:895-967.
686. — . 1920(2. Summary of the results form the mammal survey. J. Bombay
Nat. Hist. Soc., 27:57-85.
1986
Mares and Braun —Mammalogy Literature
205
19206. Summary of the results from the mammal survey. J. Bombay
Nat Hist Soc., 27:30 N3 13.
688. XiMENEZ, A., A. Langguth, and R. Praderi. 1972. Lista sistemMca de los
mamlferos del Uruguay. An. Mus. Nac. Hist. Nat. Montevideo, 7:l--49.
689. Xu, L., Z. Liu, and X. Yi. 1983. Birds and mammals of Hainan Island. Scientific
Publishing Agency, Beijing, China (for Academia Sinica), 426 pp.
690. Yalden, D. W,, M. J. Largen, AND D. Kock. 1976. Catalogue of the mammals
of Ethiopia. 2. Insectivora and Rodentia. Monit. Zool. Italiano, 8(SuppL):l-l 18.
59 p 1977. Catalogue of the mammals of Ethiopia. 3. Primates. Monit. Zool.
Italiano, 9(Suppl.):l"52.
692. . 1980. Catalogue of the mammals of Ethiopia. 4. Carnivora. Monit. Zool.
Italiano, 13(SuppL):169--272.
693. Yalden, D. W., ET AL. 1984. Catalogue of the mammals of Ethiopia. 5. Artiodac-
tyla. Monit. Zool. Italiano, 19(SuppL):67"-22L
694. YaRez, j. 1982. Estado de las colecciones mastoecologicas nacionales. PubL Ocas.
Mus. Nac. Hist. Natur. Santiago, 38:13-28.
695. Yoneda, M. 1984. Los roedores del Alto Rio La Paz. Mus. Natl. Hist. Nat. La
Paz Comunic., 3:5-12.
696. Yoshikura, M. 1956. Insectivores and bats of South Sakhalin. Kumamoto J.
Sci., 2:259-280.
697. Yudin, B. S, 1971 [Insectivorous mammals of Siberia (Key).] Novosibirsk.
698. Zangheri, P. 1969. II naturalista esploratore, raccogiitore, preparatore. Ed. Hoe-
pli, Milano.
699. Zangheri, P., and A. Pasa. 1969. Piccola fauna Italiana: Uccelli e mammiferi.
Aldo Martello Editore, Milano, 203 pp.
700. Zhuge, Y. 1982. [On the geographical distribution and the mammalian fauna of
Zhejiang Province.] Acta Theriol. Sinica, 2:157-166.
701. Ziegler, A. C. 1971. Suggested vernacular names for New Guinea recent land
mammals. Occas. Papers Bernice P. Bishop Mus., 24:1 39-1 53,
702. . 1972. A guide to the native land mammals of north east New Guinea.
Prelim, version for use of Wau Ecology Institute.
703. 1981. Petaurus abidi, a new species of glider (Marsupialia: Petauridae)
from Papua New Guinea. Australian Mamm., 4:81-88.
704_ ______ 1982. An ecological check-list of New Guinea recent mammals. Pp. 863-
894, in Biogeography and ecology of New Guinea, Vol. 2 Monographiae Biologicae,
VoL 42 (J. L. Gressitt, ed.). Dr. W. Junk, The Hague, 983 pp.
705. Ziegler, A. C., and W. J. Lidicker, Jr. 1 968. Keys to the genera of New Guinea
Recent land mammals. Proc. California Acad. Sci., 36:33“7L
706. ZiMMERMANN, K. 1953. Die Wildsauger von Kreta. Z. Saugetierk., 17:1-72.
707_ _____ 1966. Taschenbuch unserer wildlebenden Saugetiere, mit Bildem von
Lieselotte Finke-Poser, Michael Lissmann und Gerhard Richter. Urania, Leipzig,
169 pp.
708. ZuojiAN, F. 1984. [A checklist of the mammals of Xizang (Tibet).] Acta Theriol.
Sinica, 4:341-358.
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CARNEGIE MUSEUM OF NATURAE HISTORY
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VOLUME 55 7 November 1986 ARTICLE 1 1
CATALOG OF THE RECENT MARINE MAMMALS IN THE
CARNEGIE MUSEUM OF NATURAL HISTORY
Suzanne B. McLaren
Collection Manager, Section of Mammals
Duane A. Schlitter
Curator, Section of Mammals
Hugh H. Genoways
Research Associate, Section of Mammals
Abstract
The Section of Mammals, Carnegie Museum of Museum of Natural History, houses
6 1 2 specimens of Recent marine mammals, including 34 cetaceans, 44 polar bears, six
sea otters, 499 pinnipeds, and 29 manatees. Families represented in the Order Cetacea
include Platanistidae, Physeteridae, Monodontidae, Delphinidae, Phocaenidae, and Bal-
aenopteridae. Families represented in the Order Pinnipedia include Otariidae, Odobeni-
dae, and Phocidae. The single family Trichechidae represents the Order Sirenia in the
collection. For each specimen the following data are recorded: date collected, catalog
number, sex, age or condylobasal length, nature of specimen, condition of specimen, and
comments. The latter category gives information on the condition of the skin and skeletal
material, and also explains the availability of the specimen.
INTRODUCTION
The Section of Mammals, Carnegie Museum of Natural History,
currently holds 6 1 2 specimens of Recent marine mammals, including
34 cetaceans, 44 polar bears, six sea otters, 499 pinnipeds, and 29
manatees. The majority of the collection of marine mammals consists
of specimens preserved as skulls or disarticulated skeletons. However,
Submitted 9 June 1986.
237
238
Annals of Carnegie Museum
VOL. 55
the collection also contains 151 skins and five fluid-preserved speci-
mens as well as body mounts and articulated skeletons. The specimens
are housed primarily in the fur vault and large osteological storage area
of the Section of Mammals, but all mounted material is on display at
this time in the public galleries of the Museum.
The primary purpose of this catalog is to make the availability of
this material known to the scientific community. It is extremely difficult
to obtain new specimens of any marine mammal because most species
are considered to be threatened with extinction (Berger et al., 1979).
All species are placed in at least Appendix II (Threatened) under the
Convention on International Trade in Endangered Species and are
protected under the various Marine Mammal Acts beginning as early
as 1967. Therefore, it is necessary for researchers to make maximum
use of the materials already stored in museum collections. Because
specimens stored in this collection are essentially irreplaceable, none
is available for study on loan. However, all qualified researchers are
welcome to study any of the specimens at the collection facilities of
the Section of Mammals.
All collection data files of the Section of Mammals have been com-
puterized. This catalog presents selected data categories of which some
are available on the computer file, but others have been specifically
generated for this publication. Other categories of data are also available
upon request in printout or machine-readable form. The format of this
catalog is patterned after Napier’s (1976, 1981, 1985) Catalogue of the
Primates in the British Museum (Natural History) and the McLaren
et al. (1984) Catalog of the Recent Scandentia and Primates in the
Carnegie Museum of Natural History.
Classification
The collection of the Section of Mammals and this catalog follow
G. G. Simpson’s (1945) classification in the Principles of Classification
and the Classification of Mammals through the level of subfamily.
Below subfamily, specimens are arranged alphabetically by genus. Spec-
imens stored in the osteological collection and the fur vault are then
arranged numerically within the genus. This catalog is arranged alpha-
betically by genus, species, and in a few cases subspecies, so that re-
searchers may easily see what is available for each taxon. Beyond this
taxonomic arrangement all specimens are presented in numerical order.
It is hoped that by arranging the catalog in this way that work by
researchers in the collection will be facilitated.
Cetaceans in the collection have been examined and identified by C.
W. Potter and G. S. Morgan. The Odobenus have been identified, and
sexed in some cases, by Francis Fay. Hall (1980) has been used for
1986
McLaren et al.™ Catalog of Marine Mammals
239
identification of the Pinnipedia, but scientific names follow Honacki
et al. (1982) except in the case of Physeter catodon (Schevill, 1986).
Explanation of Headings
Locality. —Collection localities appear as listed on the original skin and skull tags.
Latitude and longitude were added to numerous localities as listed in the most recent
edition of the Standard Names Gazetteer for localities outside of Canada (printed by the
United States Board of Geographic Names). Gazetteer of Canada (printed by the Ca-
nadian Permanent Committee on Geographical Names) was used for referencing Ca-
nadian localities.
Of the 6 1 2 marine mammals, 3 1 were zoo specimens. In many cases, original localities
are not known for these animals. To facilitate recognition of locality information per-
taining to captive specimens, these data are followed by an asterisk.
Date of collection.— ¥ or wild taken specimens, this date refers to the day on which
the animal was obtained. In nearly all cases for zoo specimens, the only date which is
available is the day on which the animal died in captivity. In rare cases when more
information is known about a particular captive specimen, those data will be shown
under the “Comments” category.
Three symbols are used for denoting sex: F, M, ? (undetermined by original
collector). Where not denoted on the original tag, sex has sometimes been inferred based
on the examination of adult skulls. This practice has not been used for the preparation
of the current catalog, but in instances where such conclusions have been drawn pre-
viously, the appropriate symbol will be enclosed in brackets.
Age or condylobasal length.— Dming the preparation of this catalog, we were advised
by R, L. Brownell to replace age with condylobasal length. Whenever possible this
measurement was taken. If minor damage made precise measurement impossible, a
rough measurement is shown in brackets. If skull damage was too severe to make this
practical, or when only skins were available, a judgement based on tooth eruption, fusion
of sutures, or pelage was sometimes possible. Additionally, original collectors’ notes on
specimen tags were sometimes utilized in an effort to provide potential researchers with
an idea of whether a specimen was immature [Imm] or adult [Ad]. Immature specimens
were recognized with most certainty. When satisfactory judgement was not possible, a
question mark (?) appears in this category. In one series of Phoca hispida hispida spec-
imens, collector J. K. Doutt noted age approximations (in years) as told to him by local
Inuit hunters. These estimates have been included in brackets under this category, if
measurements cannot be taken. When condylobasal length can be measured, these es-
timates appear in brackets under the “Comments” category.
Nature of specimen. —This category describes the type of preservation used for each
specimen and corresponds to the two letter abbreviation system used on our computer
file. The following list describes each type of preservation used in the marine mammal
collection: SK = skull only; SO = skin only; SS = skin and skull; SB = skin, skull, and
body skeleton; SN = complete skeleton; CO = cranium only; AL = alcoholic (preserved
in 70% ethyl alcohol); BM = body mount; PS = partial skeleton. If peculiarities exist
regarding availability of a specimen these are noted by an asterisk following the two
letter code. There are two types of specimens that carry the asterisk: 1) specimens for
which some of the parts are missing; 2) specimens on exhibit. These latter specimens are
available for examination but work must be planned with the understanding that the
specimens are in exhibit areas and are not housed with the remainder of the collection.
For further explanations regarding the availability status of a specimen refer to the
“Comments” category.
Condition of specimen and comments. — This category is designed to inform the reader
of the usefulness of a specimen for systematics research. The term “Good” is used for
240
Annals of Carnegie Museum
VOL. 55
all specimens and specimen parts, which are in useable condition for study. Damage to
the skull is described so that measurability can be evaluated. All skins are either tanned
or body mounts with the exception of a few newborn polar bears which have been
prepared as conventional study skins. Captives are noted; missing parts and exhibit
specimens are noted. Inuit age estimates of Phoca hispida hispida specimens are included
here.
Taxonomic List
Cetacea
Platanistidae
Iniinae
Inia geoffrensis
Physeteridae
Physeterinae
Physeter catodon
Monodontidae
Delphinapterus leucas
Monodon monoceros
Delphinidae
Delphinus delphis
Sotalia fluviatilis
Tursiops truncatus
Phocaenidae
Phocaena phocaena
Balaenopteridae
Balaenoptera musculus
Carnivora
Ursidae
Ursus maritimus
Mustelidae
Lutrinae
Enhydra lutris lutris
Pinnipedia
Otariidae
Callorhinus ursinus
Eumetopias jubatus
Otaria byronia
Zalophus californicus
Odobenidae
Odobenus rosmarus diver-
gens
Odobenus rosmarus ros-
marus
Phocidae
Phocinae
Erignathus barbatus
Halichoerus grypus
Phoca fasciata
Phoca groenlandica groen-
landica
Phoca hispida hispida
Phoca sibirica
Phoca vitulina concolor
Phoca vitulina mellonae
Phoca vitulina richardii
Phoca vitulina vitulina
Phoca vitulina
Phoca species
Cystophorinae
Cystophora cristata
Sirenia
Trichechidae
Trichechus inunguis
Trichechus manatus lati-
rostris
Trichechus senegalensis
Acknowledgments
During the curation of the marine mammal collections and the subsequent updating
of the computer file, the position held by Suzanne McLaren was supported by National
Science Foundation Grant BSR-81 1 1553. We would like to express our thanks to Dr.
Darryl Domning, Dr. Francis Fay, Gary S. Morgan, and Charles W. Potter for their
assistance in identification of portions of the marine mammal collection. We also ap-
1986
McLaren et al.— Catalog of Marine Mammals
241
predate the assistance of Joseph Bissonnette in helping to pinpoint original collecting
localities for several zoo specimens.
Literature Cited
Berger, T, J., A. M. Neuner, and S. R. Edwards. 1979. Directory of federally con-
trolled species. Assoc. Syst. Collections, Lawrence, Kansas, iii + 1--6 + MA 1--MA
115.
Doutt, J. K. 1942. A review of the genus Phoca. Ann. Carnegie Mus., 29:61-125.
Hall, E. R. 1980. The mammals of North America. John Wiley and Sons, New York,
2nd ed., 2:601-1181.
Honacki, j. H., K. E. Kinman, and J. W. Koeppl (eds.). 1982. Mammal species of
the world. Allen Press, Inc., and Assoc. Syst. Collections, Lawrence, Kansas, ix +
694 pp.
McLaren, S. B., D. A. Schlitter, and H. H. Genoways. 1984. Catalog of the Recent
Scandentia and Primates in the Carnegie Museum of Natural History. Ann. Carnegie
Mus., 53:463-525.
Napier, P. H. 1976. Catalogue of Primates in the British Museum (Natural History).
Part I: Families Callitrichidae and Cebidae. British Museum (Natural History),
London, x + 1 2 1 pp.
— — — . 1981. Catalogue of Primates in the British Museum (Natural History) and
elsewhere in the British Isles. Part II: Family Cercopithecidae, Subfamily Cerco-
pithecinae. British Museum (Natural History), London, vii + 203 pp.
— — . 1985. Catalogue of Primates in the British Museum (Natural History) and
elsewhere in the British Isles. Part III: Family Cercopithecidae, Subfamily Colobinae.
British Museum (Natural History), London, x + 1 1 1 pp.
ScHEViLL, W. E. 1986. The International Code of Zoological Nomenclature and a
paradigm: The name Physeter catodon Linnaeus 1785. Marine Mammal Science, 2:
153-157.
Simpson, G. G. 1945. The principles of classification and a classification of mammals.
Bull. American Mus. Nat. Hist., 85:i + 1-350.
Williams, S. H. 1928. A river dolphin from Kartabo, Bartica District, British Guiana.
Zoologica, 7(4): 105-1 28.
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Belcher Islands, northwest of gions damaged
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CBL Nature of
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252
Annals of Carnegie Museum
VOL. 55
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CBL Nature of
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254
Annals of Carnegie Museum
VOL. 55
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Great Whale River
Catalog of Marine Mammais— Continued.
262
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Catalog of Marine Mammais— Continued.
272
Annals of Carnegie Museum
VOL. 55
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Post 59°40'N; 77°30'W
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282
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Post 59°40'N; 77°30'W numerous loose teeth
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284
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1986
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1986
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ANNALS"*-"
0/ CARNEGIE MUSEUM
CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE » PITTSBURGH, PENNSYLVANIA 15213
VOLUME 55 7 November 1986 ARTICLE 12
TAXONOMIC AND DISTRIBUTIONAL NOTES ON
BATS FROM KENYA
Duane A. Schlitter
Curator, Section of Mammals
ISSA R. Aggundey*
Mazin B. Qumsiyeh^
Rea Postdoctoral Fellow, Section of Mammals
Kimberlyn Nelson^
Rodney L. Honeycutt^
Abstract
Taxonomic and distributional comments are given for six species in three families of
bats from Kenya. Two species {Hipposideros camerunensis and Tadarida russata) are
reported from Kenya for the first time.
Introduction
During field work on the systematics of small mammals in Kenya
from September to November 1985, a number of bats were collected
‘ Curator of Mammals, National Museums of Kenya, P.O. Box 40658, Nairobi, Kenya.
^ Current address: Department of Biological Sciences, Texas Tech University, Lubbock,
TX 79409.
^ Address: Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138.
Submitted 5 February 1986.
298
Annals of Carnegie Museum
VOL. 55
in various parts of Kenya, Some of these specimens add substantially
to the knowledge of the distribution of bats in Kenya and East Africa
while others were of taxonomic interest. This paper summarizes these
data on six species of bats in three families.
Materials and Methods
All specimens reported herein were captured with mist nets set in the normal manner.
Cranial and forearm measurements were taken by means of dial calipers and are given
in millimeters. All specimens are deposited in the Carnegie Museum of Natural History,
Pittsburgh (CM); Museum of Comparative Zoology, Harvard University, Cambridge
(MCZ); and National Museums of Kenya, Nairobi (NMK). Coordinates for localities are
listed only for the first reference to the locality.
Account of Species
Family Hipposideridae
Hipposideros camerunensis Eisentraut, 1956
Hipposideros camerunensis Eisentraut, 1956. Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere,
84:526. Near Buea, Cameroun.
Records (1). — Kenya: Western Prov., Kakamega Dist., Kakamega Forest Station, 3-
1/2 km E Kakamega (0°14'N, 34°52'E) (1 CM).
Measurements. ~Sq\qqXq6. measurements of an adult female, length
of forearm, 74.3; condylocanine length, 24.9; zygomatic breadth, 15.7;
alveolar length of maxillary toothrow, 9.3; and greatest crown breadth
of upper molar toothrows, 11.1.
Remarks. —Hipposideros camerunensis is a rare, large forest species
of Hipposideros resembling closely the more common H. cyclops. It
has been reported from the type locality, Buea, Cameroun, and else-
where only from Shabunda, eastern Zaire (2°42'S, 2r20'E) (Hill, 1963:
81). On 5 November a single adult female was taken in the bottom
shelf of a mist net placed along a cleared trail in the Intermediate
Evergreen Forest near the Forest Guest House at the Kakamega Forest
Station. This specimen from the Kakamega Forest extends the range
of this species about 900 kilometers to the northeast and is the first
record of occurrence for Kenya. For a description of the capture area,
see Zimmerman (1972).
In size, the Kenyan female agrees well with a series of Cameroun
specimens in Carnegie Museum and with the measurements given by
Hill (1963:80).
Hipposideros cyclops (Temminck, 1853)
Phyllorhina cyclops Temminck, 1853. Esquisses Zool, sur la Cote de Guine, p. 75. Boutry
River, Ghana.
Records (2).— Kenya: Coastal Prov. Kwale Dist., Shimba Hills Nature Reserve, Ma-
kandara Picnic Site, 7 km S, 8 km W Kwale (4®15'S, 39®23'E) (2 CM).
1986
SCHLITTER ET AL. — NOTES ON KENYAN BaTS
299
Measurements.— measurements of a male and female, re-
spectively, length of forearm, 66.8, 63.7; condylocanine length, 27.1,
27.2; zygomatic breadth, 14.8, 1 5.2; alveolar length of maxillary tooth-
row, 8.4, 8.7; and greatest crown breadth of upper molar toothrows,
10.4, 10.6.
Remarks.— Hipposider os cyclop s is known currently from two lo-
calities in western Kenya. Hayman (1 935:48) reported this species from
the Yala River and later Harrison (1961:290) listed Kaimosi. The
species is not known from Tanzania.
On 14 October, two adult individuals of H. cyclops were captured
in a mist net set in the cleared Makandara Picnic Site in the Shimba
Hills Nature Reserve. The net was set parallel and adjacent to the forest
edge. These two specimens extend the range of this species to extreme
southeastern Kenya. The geographic range of this species probably
reaches this region of Kenya along a corridor of forest blocks extending
from southwestern Tanzania to northeastern Kenya. This species should
be captured in appropriate forests in this Tanzanian corridor after
careful and concentrated mist netting.
Family Vespertilionidae
Scotophilus nux Thomas, 1 904
Scotophilus Thomas, 1904. Ann. Mag. Nat. Hist., (7) 4:355. Gambage, Ghana.
Records (4).— Kenya: Western Prov., Kakamega Dist., Kakamega Forest Station,
V/i km S, 12 km E Kakamega (2 CM, 1 MCZ, 1 NMK).
Measurements. SQlQcXtd external measurements of four females,
length of forearm, 57.8, 57.6, 54.5, 55.3.
Remarks.— Scotophilus nux has been reported in Kenya only from
8 km SE Kakamega (Aggundey and Schlitter, 19^4:138). The eight
specimens from Kenya referred to by Robbins (1983:23) are from this
locality. Both localities refer to the Kakamega Forest Station. Four
additional specimens of this dark-brown colored forest species of Sco-
tophilus were netted in the forest adjacent to the station and in the
cleared areas around the station complex on 5 and 7 November. These
specimens further verify the occurrence of this species in the forests of
western Kenya. Specimens of Scotophilus dinganii were not taken to-
gether with S. nux at this locality, but the former species was taken in
more heavily disturbed areas near the station complex and seems to
be the most common species of Scotophilus taken in Kenya.
Kerivoula argentata argentata Tomes, 1861
Kerivoula argentata Tomes, 1861. Proc. ZooL Soc. London, 1861:32. Otjoro, Namibia.
Records (I).— Kenya: Coastal Prov., Kwale Dist., Shimba Hills Nature Reserve, 5
km S, 1 km W Kwale (4°13'S, 39°27'E) (1 CM).
300
Annals of Carnegie Museum
VOL. 55
Measurements. — Selected measurements of an adult male, length of
forearm, 35.9; condylobasal length, 13.9; and alveolar length of max-
illary toothrow, 5.9.
Remarks.— Kerivoula argentata occurs primarily in southern Africa
and ranges northward as far as Kenya. Specimens of Kerivoula are
seldom captured and there is a paucity of records for this species in
eastern Africa. It has been reported from Li wale, in southern Tanzania
(Harrison, 1 958:95), an unspecified locality in Uganda (Kingdon, 1974:
304), and from Kibwezi, southeastern Kenya (Aggundey and Schlitter,
1984:139). A single adult male was taken on 12 October in a mist net
set under the trees in the central clearing surrounding the bandas at
the overnight facilities in Shimba Hills Nature Reserve. This specimen
constitutes the second locality record of the species for Kenya and only
the fourth for this region encompassing the northern part of the range
of the species.
The Shimba Hills specimen exhibits the normal white-tipped orange-
rufous dorsal coloration and the somewhat lighter colored ventral fur.
Mensurally it fits within the size range of southern African examples
of the species.
Family Molossidae
Tadarida (Chaerephon) mssata (J. Allen, 1917)
Chaerephon mssata J. Allen, 1917. Bull. Am. Mus. Nat. Hist., 37:458, Medje, Zaire.
Records (3). —Kenya: Rift Valley Prov., Naivasha Dist., Hell’s Gate Canyon, 20 km
S, 14 km W Naivasha (0°54'S, 36°19'E) (1 CM, 1 MCZ, 1 NMK).
Measurements.— SeXQcXQd measurements of two males and one fe-
male, respectively, length of forearm, 45.4, 46.2, 46.2; condylobasal
length, 18,0, 18.0, 17.1; zygomatic breadth, 11.2, 11.9, 11.2; alveolar
length of maxillary toothrow, 7.2, 7.1, 6.7; and greatest crown breadth
of upper molar toothrows, 9.0, 8.7, 8.3.
Remarks.— ThQ taxonomic and distributional status of Tadarida
russata was reviewed by Peterson (1971) and Fenton and Peterson
(1972). This species is presently known from Ghana, Cameroun, and
the type locality in northeastern Zaire (Fenton and Peterson, 1972:20).
On 30 September, two adult males were captured in a mist net set
across the floor of the northern end of Hell’s Gate Canyon. On 3
October a single adult female was taken in a net set in the same area.
These three specimens comprise the first record of the species for Kenya
and extend the range of the species more than 1000 kilometers to the
southeast.
The Kenyan specimens of this species agree in color of pelage with
three Cameroun examples in Carnegie Museum. However, mensurally
they appear to be appreciably larger than these examples and the mea-
1986
SCHLITTER ET AL. — NOTES ON KENYAN BaTS
301
surements given in Fenton and Peterson (1972:21, 22). When more
specimens from throughout the geographic range of this species become
available, the Kenyan population may prove to be significantly larger
and worthy of subspecific recognition.
Tadarida (Tadarida) fulminans (Thomas, 1903)
Nyctinomus fulminans Thomas, 1903. Ann. Mag. Nat. Hist, (7)12:501.
Fianarantsoa, eastern Betsileo, Madagascar.
Records (1).— Kenya: Rift Valley Prov., Nakuru Dist., Njoro River, 3 km S, 3 km W
Nakuru (0°19'S, 36°03'E) (1 CM).
Measurements. SQlQCiQd measurements of an adult male, length of
forearm, 56.7; condylobasal length, 21.2; zygomatic breadth, 13.4; al-
veolar length of maxillary toothrow, 8.3; and greatest breadth of upper
molar toothrow, 9.3.
Remarks. — The first record of occurrence for Tadarida fulminans in
Kenya was a specimen from Nairobi reported by Harrison (1960:74).
This species occurs primarily in southern Africa (Smithers, 1983) and
reaches the northern extent of its range in Kenya. On 8 October a single
adult male was captured in the gorge of the Njoro River southwest of
Nakuru. The specimen was taken in a mist net set across shallow water
and among trees on the steep bank of the watercourse.
Acknowledgments
A research permit (CAB 13/001/1 1C38/17) allowing us to do field studies in Kenya
was received from the Office of the President, We thank Mrs, C. A. Mwango and Miss
L, A, Gwiyo and their staff for their efforts and courtesy in processing this permit on
our behalf and the Secretary, National Council for Science and Technology, and his
colleagues for approving our research project.
Many individuals in the Ministry of Tourism and Wildlife supported the research
project in many ways. We are especially grateful to Mr. D. M. Mbuvi, Assistant Director
(Research), Wildlife Conservation and Management Department and his staff for support,
Mr. J. P. Oriero, Assistant Director (Administration), for issuing permit No. WCMD/
7/9/ Vol. VI, and the numerous Wardens and their staff in the Parks and Reserves in
Kenya who made our stay comfortable and work easier. Mr. F. N. Pertet, Principal,
Wildlife Training College, Naivasha, and his staff, especially Mr. Okumu Mbanda and
Mr. Osbourne Mivelo, were gracious hosts and gave invaluable assistance.
We thank Mr. John S. Karmali and the staff at Nakuru Wildlife Trust House, Lake
Nakuru National Park and Dr. Stephen G. Njuguna and his staff at the Moana Marine
Station, Diani Beach, for logistical assistance.
Finally we are especially indebted to Dr. Richard Leakey and his staff of the National
Museums of Kenya, including Mr. James N. Maikweki, Curator in Charge, Kisumu
Museum, for their overwhelming support. We are truly appreciative of the assistance of
Caroline Plazek and Paul Gathinji who assisted with the field work.
Financial assistance for field work in Kenya was received from the M. Graham Netting
Research Fund, Cordelia Scaife May Charitable Trust, Carnegie Museum of Natural
History; from Barbour and Richmond funds. Museum of Comparative Zoology; from
NIH grant AIO 4242 to Dr. Robert Traub; and the Rea Postdoctoral Fellowship program
at Carnegie Museum of Natural History.
302
Annals of Carnegie Museum
VOL. 55
Literature Cited
Aggundey, L R., and D. A. Schlitter. 1984. Annotated checklist of the mammals
of Kenya. I. Chiroptera. Ann. Carnegie Mus., 53:1 19-161.
Fenton, M. B., and R. L. Peterson. 1972. Further notes on Tadarida aloysiisabaudiae
and Tadarida russata (Chiroptera: Molossidae— Africa). Canadian J. ZooL, 50:19-24.
Harrison, D. L. 1958. Two bats (Microchiroptera) new to the fauna of Tanganyika
Territory. Durban Mus. Novit., 5:95-98.
. 1960. Notes on some Central and East African bats. 3. The free-tailed bat
Tadarida Thomas, 1903, in Kenya Colony. Durban Mus. Novit., 6:74-78.
. 1961. A checklist of the bats (Chiroptera) of Kenya Colony. J. East African
Nat. Hist. Soc., 23:286-295 (for 1960).
Hayman, R. W. 1935. A note on Hipposideros cyclops Temminck and its synonym
Hipposideros langi Allen. Ann. Mag. Nat. Hist., (10)15:47-50.
Hill, J. E. 1963. A revision of the genus Hipposideros. Bull. British Mus. (Nat. Hist.),
Zool., 2(1):1-129.
Kingdon, j, 1974. East African mammals. An atlas of evolution in Africa. Volume
II, Part A (Insectivores and Bats). Academic Press, London, xii + 392 pp.
Peterson, R. L. 1971. The African molossid bat Tadarida russata. Canadian J. Zool.,
49:297-301.
Robbins, C. 1983. A new high forest species in the African bat genus Scotophilus
(Vespertilionidae). Ann. Mus. Roy. Afr. Centr., Sc. Zool., 237:19-24.
Smithers, R. H. N. 1983. The mammals of the southern African subregion. University
of Pretoria, Pretoria, xxii + 734 pp.
Zimmerman, D. A. 1972. The avifauna of the Kakamega Forest, western Kenya, in-
cluding a bird population study. Bull. American Mus. Nat. Hist., 149:257-339.
r'
r
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V.
J c*»j #- *%t,
* 'C^a 5*.#:*,. asiks;-
.>
Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.
o/ CARNEGIE MUSEUM
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VOLUME 55 7 NOVEMBER 1986 ARTICLE 13
RESULTS OF THE ALCOA FOUNDATION-SURINAME
EXPEDITIONS. XL BATS OF THE GENUS
MICRONYCTERIS (MAMMALIA: CHIROPTERA)
IN SURINAME
Hugh H. Genoways
Research Associate, Section of Mammals
Stephen L. Williams
Collection Manager, Section of Mammals
Abstract
Of the 10 recognized species in the genus Micronycteris, seven species {brachyotis,
daviesi, hirsuta, megalotis, minuta, nicefori, and sylvestris) have been reported to occur
in Suriname. Micronycteris daviesi is easily distinguished from the other species by its
large size (forearm over 50 mm) and massive dentition. The next largest species (forearm
over 43 mm)--M hirsuta— is distinguished by having the upper inner incisors separated
at the base but in contact at the tip and having lower incisors that are high and wedged
between canines so that the canines are in contact behind the incisors.
Micronycteris sylvestris has dorsal pelage that is tricolored. The upper incisors of this
species are similar in length to the canines and the first upper premolar possesses accessory
cusps.
The other four species form two species pairs. Micronycteris megalotis and minuta
are the smallest members of the genus in Suriname. They can be distinguished from each
other by the more deeply notched interauricular band in minuta and by the first upper
premolar being smaller than the second premolar in minuta but of about equal size in
megalotis.
Micronycteris brachyotis, which was not encountered during our work in Suriname,
has short ears (less than 1 6 mm from notch) and lacks the faint gray line usually present
on the lower back of specimens of M. nicefori. These species are also distinguished by
Submitted 24 April 1986.
303
VOL. 55
304
Annals of Carnegie Museum
Fig. 1. — Stylistic representation of variation in the interauricular band of members of
the genus Micronycteris. Upper, interauricular band complete (exemplified by M. hir-
suta). Middle, interauricular band deeply notched in middle {M. minuta). Lower, inter-
auricular band absent (M. nicefori). Not to scale.
their upper incisors, which are chisel-shaped and in line with the canines in brachyotis,
but project forward and, therefore, are out of line with the canines in nicefori.
Introduction
The genus Micronycteris is a member of the subfamily Phyllostom-
inae of the New World leaf-nosed bat family Phyllostomidae. Micro-
nycteris is currently recognized as containing 10 small- to medium-
1986
Genoways and Williams —Suriname Micronycteris
305
sized species of bats. The genus was reviewed by Andersen (1906) and
Sanborn (1949).
The genus is divided into six subgenera based primarily upon the
work of Sanborn (1949). We suggest that this level of classification be
carefully examined in the future because the genus seems oversplit at
the subgeneric level. Andersen (1906) recognized seven species divided
into two genera {Micronycteris and Gly phony cteris), whereas Sanborn
(1949) recognized 10 species in one genus. Subsequent to Sanborn’s
work, one species (platyceps) was placed as a junior synonym of brachy-
Otis (Goodwin and Greenhall, 1961) and another species, daviesi, was
described in a separate genus {Barticonycteris; Hill, 1 964), but was later
placed into Micronycteris (Koopman and Cockrum, 1967).
During our field studies in Suriname, six species of the genus Mi-
cronycteris {daviesi, hirsuta, megalotis, minuta, nicefori, and sylvestris)
were captured. Husson (1978) reported a seventh species {brachyotis)
from the country. This means that only three of the currently recognized
species of Micronycteris {behni, pusilla, and schmidtorum) have not
been taken in Suriname. With this large number of species from this
complex genus present in such a small geographic area, we have taken
the opportunity to gain a better insight into the morphological and
ecological relationships of species within the genus.
Methods and Materials
Specimens were taken with mist nets and preserved as skins and skulls or in fluid.
Field weights were taken with Pesola scales, accurate to 1 g. Measurements of forearm
and cranial dimensions were taken with dial calipers accurate to 0. 1 mm. Only adult
specimens (phalangeal epiphyses completely fused) were measured in this study. Mea-
surements were taken as described by Genoways and Williams (1984). Reproductive
condition of the skin and skull specimens was determined by gross dissection in the
field, whereas fluid preserved specimens were dissected in the laboratory. Specimens
listed in each account were deposited in the Section of Mammals, Carnegie Museum of
Natural History.
Acknowledgments
We gratefully acknowledge the logistical support, assistance in acquiring permits, and
field assistance of Dr, loop Schulz, Mr. Henry Reichart, and Mr. Ferdinand Baal. Mr.
Kris Mohadin, Ms. Muriel Hand, and other staff members of STINASU contributed to
the success of field work in Suriname. We are particularly thankful to Mr. Leo Roberts
for accompanying and assisting us with most of the field work. Ms. Marga Werkhoven
and Mr. 1. Douglas provided housing and lab facilities in Paramaribo. Other individuals
who helped collect specimens used in this study include Ms. Jane Casne, Mr. Michael
Arnold, Dr. Rodney Honeycutt, Mr. Ben Koop, Ms. Paisley Seyfarth, Mr. Murray de la
Fuente, Dr. Carleton Phillips, Dr. Robert Baker, and Mr. Keith Studholme.
Financial support for field work in Suriname was received from the Alcoa Foundation
(Charles L. Griswold, President) and the M. Graham Netting Research Fund established
by a grant from the Cordelia S. May Charitable Trust.
306
Annals of Carnegie Museum
vou 55
Fig, 2. — Dental characteristics of members of the genus Micronycteris. A, lower incisors
bifid (exemplified by M. megalotis); B, lower incisors trifid (exemplified by M. sylvestris).
C, upper and lower incisors of M. hirsuta, showing long, narrow lower incisors with
unexpanded crowns and awl-shaped upper incisors; D, upper and lower incisors of M.
megalot is showing short lower incisors with expanded crowns. E, upper premolars (P3,
1986
Genoways and Williams —-Suriname Micronycteris
307
Key to the Species of Micronycteris in Suriname
(see also Medellin et al., 1985)
1 . Interauricular band present (possibly notched or not as broad
in the middle (Fig. lA, B); lower incisors bifid (Fig. 2 A) ..... 2
1'. Interauricular band not present (Fig. 1C); lower incisors trifid
(Fig. 2B) 4
2. Forearm less than 40; greatest length of skull less than 22;
lower incisors with expanded crowns (Fig. 2D) 3
2'. Forearm greater than 40; skull more than 22; upper incisors
awl-shaped; lower incisors long, narrow, and lacking expanded
crown (Fig. 2C); 2n = 30, FN = 32 ....... .Micronycteris hirsuta
3. Calcar longer than foot (claws included); length of interfemoral
membrane more than twice the length of tail; band of skin
between ears with shallow notch in middle; upper premolars
(P3, P4) about the same height (Fig. 2E); 2« = 40, FN =68
Micronycteris megalotis
3 ' . Calcar shorter than foot (claws included); length of interfemoral
membrane less than twice the length of tail; band of skin be-
tween ears deeply notched in middle (Fig. IB); first upper
premolar (P3) distinctly shorter than second upper premolar
(P4) (Fig. 2F); 2n = 28, FN = 52 Micronycteris minuta
4. First upper incisors similar to canines in length; first upper
premolar (P3) having accessory cusps on lingual and posterior
margins 5
4'. First upper incisors distinctly shorter and narrower than ca-
nines; first upper premolar (P3) lacking accessory cusps, only
the main cusps present 6
5. Forearm less than 50; greatest length of skull less than 25;
dorsal hair tricolored; two pairs of upper incisors; 2n = 22,
FN = (40) Micronycteris sylvestris
5'. Forearm greater than 50; greatest length of skull more than 25;
dorsal hair brownish throughout; sagittal crest straight; one
pair of upper incisors; 2n = 28, FN = 52 . . Micronycteris daviesi
6. Length of ear (to notch) less than 16; calcar about the same
length as foot; first pair of upper incisors chisel-shaped (Fig.
2G) and in line with canines; second pair of upper incisors
P4) of M. megalotis (anterior is to the right), note that premolars of about equal size; F,
upper premolars of M. minuta (anterior is to the right), note that P3 is distinctly shorter
than P4. G, chisel-shaped upper incisors of M. brachyotis; H, upper incisors of M. nicefori
which are nearly as broad as they are tall.
308
Annals of Carnegie Museum
VOL. 55
bifid with elongated inner cusp; 2n = 32, FN = 60 .........
Micronycteris brachyotis
6'. Length of ear (to notch) greater than 16; calcar shorter than
length of foot; faint gray line often present on lower back; first
pair of upper incisors not chisel-shaped (Fig. 2H); upper in-
cisors projected forward and out of line with canines; In = 28,
FN = 52 Micronycteris nicefori
Generic Account
Micronycteris Gvdiy , 1866
1866. Micronycteris Gray, Proc. Zool. Soc. London, p. 113, May.
1856. Schizostoma Gervais, Mammiferes in Castelnau Exped. dans les parties centrales
de I’Amer. du Sud . . . pt. 7, p. 38. Type species, Schizostoma minuta Gervais.
Preoccupied by Schizostoma Bronn, 1835, a genus of Mollusca.
1896. Glyphonycteris Thomas, Ann. Mag. Nat. Hist., ser. 6, 18:302, October. Type
species, Glyphonycteris sylvestris Thomas.
1907. Xenoctenes Miller, Bull. U.S. Nat. Mus., 57:124, 29 June. Type species, Schizosto-
ma hirsutum Peters.
1964. Barticonycteris Hill, Mammalia, 28:556, December. Type species, Barticonycteris
daviesi Hill.
Type species.— Phyllophora megalotis Ovdcy , 1842.
Diagnosis.— K genus of small- to medium-sized bats with a well-
developed noseleaf and a tail extending only to the middle of the
interfemoral membrane. In the subfamily Phyllostominae, the dental
formula of i 2(1 )/2, c 1/1, p 2/3, m 3/3 is shared only with the genera
Macrotus and Vampyrum. The one species with only one pair of upper
incisors is M. daviesi. Rostrum not as long as braincase; auditory bullae
small; middle lower premolar approximately same size as last lower
premolar.
Micronycteris {Glyphonycteris) daviesi (Hill, 1964)
Specimen examined (1).— Saramacca: Raleigh Falls, 1.
The species M. daviesi is easily distinguished from other members
of the genus Micronycteris in Suriname by its large size (Table 1; Figs.
3A, 4A) and massive dentition. This species was originally described
as the sole representative of the genus Barticonycteris by Hill (1964).
Shortly thereafter, Koopman and Cockrum (1967) treated Bartico-
nycteris as a synonym of Micronycteris. Most recent authors have fol-
lowed this arrangement (see for example Jones and Carter, 1976), al-
though LaVal (1977) and Hall (1981) are exceptions. Koopman (1978)
treated Barticonycteris as a subgenus of Micronycteris, citing as his
reasons that the characteristics of Barticonycteris “are simply those of
M. {Glyphonycteris), the subgenus including sylvestris and behni carried
one step further.” Hill (1964) had earlier recognized that the closest
1986
Genoways and Williams —Suriname Micronycteris
309
relatives of Barticonycteris were members of Gly phony cteris. We agree
with these assessments of the relationship of daviesi based upon our
own studies, but we believe that the relationships of the taxon are
represented best by placing it as a member of the subgenus Glyphonyc-
teris.
Only one specimen of this rare species was taken during our work
in Suriname (Fig. 5). It was an adult male taken on an island in the
Coppemame River that serves as the headquarters of the Raleigh Falls
Nature Reserve. The bat was netted along a trail on the western side
of the island, about 200 m northeast of the park headquarters and
about 50 m from the river. Vegetation in the area consisted of near-
mature lowland rainforest. Our specimen, weighing 1 8 and with testes
measuring 3, was captured on 24 August at about 2000 hours following
a short rainstorm. Sixteen other species of bats were taken in this area
(Table 2).
Our specimen was found to have a In = 28 and FN = 52. The
X-chromosome was submetacentric, whereas the Y-chromosome was
acrocentric (Honeycutt et al., 1980).
Micronycteris (Glyphonycteris) sylvestris (Thomas, 1896)
Specimens examined (14). — Brokopondo: Brownsberg Nature Park, 8 km S, 2 km W
Brownsweg, 14.
Our specimens were the first of this species reported from Suriname
(Williams and Genoways, 1980). This taxon can be recognized exter-
nally by having tricolored dorsal hair and ears that are about as broad
as they are high. Cranially this species resembles M. daviesi with upper
incisors about the same length as the canines (Figs. 3, 4). Currently M.
sylvestris is considered to be monotypic (Jones and Carter, 1976), al-
though not enough specimens have been available for a proper analysis
of infraspecific variation.
Our 14 specimens were collected from a hollow tree in a mature
tropical hardwood forest on the Brownsberg highlands (Fig. 5). The
opening to the hollow was located about 3 m above the ground. Eight
of the specimens were taken on 24 September and the other six on the
following day. Four males weighed 6, 6, 7, and 7; each had testes that
measured 3. Eight females had weights ranging from 7.5 to 11 with a
mean of 9.3. None of these females evinced gross reproductive activity.
Only 10 other species of bats were captured in nets set along trails near
the hollow tree (Table 2).
The specimens of M. sylvestris from Suriname had a diploid number
of 22 and a probable fundamental number of 36. The fundamental
number could not be determined with certainty because only females
were available for chromosomal analysis. It was supposed that the
X-chromosome was biarmed (Honeycutt et al., 1980).
310
Annals of Carnegie Museum
VOL. 55
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of length of lobasal matic Mastoid bital iOary upper
Catalog No. Locality Sex forearm skull length breadth breadth breadth toothrow molars
1986
Genoways and Williams —Suriname Micronycteris
311
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8 km S, 2 km W Browns weg
CM 63594 Brokopondo: Brownsberg Nature Park F 40.0 20.0 17.9 10.2
8 km S, 2 km W Browns weg
CM 63598 Brokopondo: Brownsberg Nature Park F 39.4 19.7 17.2 10.0
8 km S, 2 km W Brownsweg
1cm
VOL, 55
312
Annals of Carnegie Museum
Fig. 3.— Dorsal view of the crania of the seven species of Mkronycteris occurring in Suriname. A, M. daviesi (CM 63573); B, M. syivestris
(CM 63597); C, M. brachyotis (from Trinidad); D, M. megalotis (CM 68390); E, M. minuta (CM 63584); F, M nicefori (CM 76771);
G, M. hirsuta (CM 68388).
1986
Genoways and Williams "“Suriname Micronycteris
313
Micronycteris (Lampronycteris) brachyotis (Dobson, 1879)
Previous Brokopondo: Gros (about 100 km S Paramaribo on railroad from
Paramaribo to the interior) [5°06'N, 55°15'W] (Husson, 1978).
Husson (1978) first reported this species from Suriname based upon
six males from Gros (Fig. 5). The specimens were taken from an old
goldmine in a savannah area. We did not encounter this species during
our work in Suriname.
Most of the characteristics of the specimens listed by Husson —fore-
arm 40.2 to 42.9, no interauricular band, and second phalanx of third
digit much longer than first— seem to match M. brachyotis closely;
however, the Suriname specimens had the fourth metacarpal the short-
est, whereas in brachyotis the fifth metacarpal is the shortest. The exact
meaning of this difference must await further examination of these
specimens.
Goodwin and Greenhall (1961) were the first to recognize that M.
platyceps, described by Sanborn in 1949, was a junior synonym of the
long described, but poorly known, M. brachyotis.
Micronycteris {Micronycteris) megalotis megalotis (Gray, 1842)
Specimens examined (12).— Brokopondo: 1 km N Rudi Kappelvliegveld, 1; P/2 km
W Rudi Kappelvliegveld, 1; 3 km SW Rudi Kappelvliegveld, 2; Brownsberg Nature
Park, 3 km S, 20 km W Afobakka, 1 . Commewune: Nieuwe Grond Plantation, 1 . Maro-
wune: Oelemarie, 1; Perica, 2. Nickerie: Kayserberg airstrip, 2. Para: Zanderij, 1.
Previous records (Husson, 1978).— Suriname: Plantation Kwatta, near Rijweg; Para-
maribo. No specific district or locality.
Among Surinamese representatives of the genus Micronycteris, M.
megalotis is distinguished by the presence of an interauricular band
which is only slightly notched, a broad interfemoral membrane, upper
premolars (P3, P4) about the same size (Fig. 2E), and bifid lower incisors
(Fig. 2A). The species is polytypic with the nominate subspecies oc-
curring in Suriname and surrounding areas (Jones and Carter, 1976).
M. megalotis has been reported from localities in Suriname previ-
ously and these together with our records indicate that the species may
be expected in most forested situations in the country (Fig. 6). Many
of the capture sites were described by the collectors as being in mature
tropical forest or lowland tropical rainforest. However, at the Nieuwe
Grond Plantation a specimen was taken in a net set along an orchard
path that was bordered on one side by a canal and on the other by
alternating rows of citrus trees and secondary tropical vegetation. At
Perica, the specimens were netted on the edge of secondary growth
forest. M. megalotis was taken at more than half of its nine capture
sites with six species (Table 2)— Saccopteryx bilineata, S. leptura, To-
natia bidens, Carollia perspicillata, Rhinophylla pumilio, and the larger
314
Annals of Carnegie Museum
VOL. 55
1986
Genoways and Williams —Suriname Micronycteris
315
58 57 56 55 54
Fig. 5,— Map of the geographic distribution of four species of Micronycteris in Suriname.
Closed circle, M. daviesi; open circle, M. sylvestris; closed triangle, M. brachyotis; open
triangle, M. hirsuta.
species of Artibeus. It was also taken together with two other species
of the gQmis—minuta and although only at one and two sites,
respectively.
Our 12 specimens consist of 10 males and two females. Testes mea-
surements for the males are as follows (date of capture in parentheses):
Fig. 4.— Lateral view of the crania of the seven species of Micronycteris occurring in
Suriname. A, M. daviesi (CM 63573); B, M. sylvestris (CM 63597); C, M. brachyotis
(from Trinidad); D, M. megalotis (CM 68380); E, M. minuta (CM 63584); F, M. nicefori
(CM 76771); G, M. hirsuta (CM 68388).
316
Annals of Carnegie Museum
VOL. 55
58 57 56 55 54
Fig. 6.— Map of the geographic distribution of Micronycteris megalotis in Suriname.
Closed circles, specimens examined; open circles, previous records.
3, 3 (4 May); 3 (7 July); 2 (13 September); 2 (30 September); 2 (1
October); 2 (3 October); 3, 4.5 (24 October); 2.5 (23 November). A
female taken on 1 October evinced no reproductive activity, whereas
no data are available for the other female. Seven of the males had an
average weight of 5.7 (range, 5—7) and the one female for which data
are available weighed 6,
The karyotype of a male from Suriname had a 2« = 40 and FN ==
68. The X-chromosome was subtelocentric and the Y-chromosome
was acrocentric (Honeycutt et al., 1980).
Micronycteris (Micronycteris) minuta (Gervais, 1856)
Specimens examined (15).-— Brokopondo: 1 km N Rudi Kappelvliegveld, 1; Browns-
berg Nature Park, 8 km S, 2 km W Brownsweg, 2. Commewune: Nieuwe Grond Plan-
1986
Genoways and Williams —Suriname Micronycteris
317
58 57 56 55 54
Fig. 7.— Map of the geographic distribution of Micronycteris minuta in Suriname.
tation, 1. Marowune: 3 km SW Albina, 1; 10 km N, 24 km W Moengo, 1. Nickerie:
Avanavero, 1; Sipaliwini airstrip, 4. Saramacca: Voltzberg, 3. Suriname; Powaka, 1.
Prior to our work in Suriname (Genoways and Williams, 1979; Wil-
liams and Genoways, 1 980) M. minuta was unknown from the country;
however, we took the species in all major regions of the country (Fig,
7). In Suriname, M. minuta would most likely be confused with M.
megalotis; however, minuta can be distinguished by the deeply notched
interauricular band (Fig. 1) and a first upper premolar (P3) that is
distinctly smaller than the second premolar (P4) (Fig. 2F). M. minuta
has some individuals smaller than any individuals of M. megalotis,
but the two species cannot be separated consistently on size alone (Table
1). M. minuta currently is considered to be monotypic (Jones and
Carter, 1976). A comparison of our material with specimens from
318
Annals of Carnegie Museum
VOL. 55
Trinidad and Brazil deposited in the American Museum of Natural
History revealed no notable differences; however, there may be slight
karyotypic differences between specimens from Trinidad and Suriname
(see discussion below).
Micronycteris minuta usually was netted in association with lowland
rainforest. However, at several places such as 1 km N Rudi Kappel-
vliegveld, 10 km N, 24 km W Moengo, and Powaka, this species was
taken in nets set in open areas along or near the forest. At the Nieuwe
Grond Plantation, M. minuta was captured in a series of nets set through
the lawns and gardens surrounding the plantation buildings. Nine species
of bats were collected in association with M. minuta at more than half
of the nine capture sites (Table 2). An interesting association is that
between M. nicefori and M. minuta, which were taken together at five
capture sites. M. minuta was also captured together with two other
members of the genus (megalot is and sylvestris) at single localities.
Our 15 specimens consist of six males and nine females. Testes
measurements for males were as follows (date of capture in paren-
theses): 3 (12 May); 2 (24 September); 2 (28 September); 3 (23 October);
2 (16 November). A female taken on 30 July was carrying a 13 mm
fetus, and another was lactating on 24 September. Females netted on
the following dates evinced no gross reproductive activity: 26 May; 6
August; 28 August; 12 September; 16 November (3 individuals). Four
of the males had weights of 5, 5, 6, and 6 and three females weighed
5, 5, and 8.
The diploid number for this species in Suriname was 28 and the
fundamental number was 52 (Baker et al., 1981). This karyotype dif-
fered slightly from that reported for the species from Trinidad (Baker,
1979). In the Suriname specimen, the smallest pair of autosomes was
biarmed, whereas in the material from Trinidad this pair was acro-
centric.
Micronycteris (Trinycteris) nicefori Sanborn, 1 949
Specimens examined (31). — Brokopondo: 1 km N Rudi Kappelvliegveld, 1. Comme-
wune: Nieuwe Grond Plantation, 1. Marowune: 3 km SW Albina, 1; 10 km N, 24 km
W Moengo, 1; Perica, 2. Nickerie: Avanavero, 3; Kabalebo, 20. Saramacca: Tafelberg,
SE side of Arrowhead Basin (3®54'N, 56°10'W), 600 m, 1. Suriname: Powaka, 1.
This species had not been reported in Suriname before our work
(Genoways and Williams, 1979; Williams and Genoways, 1980), but
we found it to be widespread in the country (Fig. 8). Micronycteris
nicefori is a medium-sized member of the genus (Table 1) that is most
likely to be confused with M. brachyotis. M. nicefori can be distin-
guished by its upper incisors, which are shorter and narrower than the
canines and project forward out of line with the canines (Fig. 2H), long
1986
Genoways and Williams —Suriname Micronycteris
319
58 57 56 55 54
Fig. 8.— Map of the geographic distribution of Micronycteris nicefori in Suriname,
ears, and a faint gray line often present on the lower back. We compared
our specimens with material from Trinidad and Colombia deposited
in the American Museum of Natural History. We could detect no
consistent differences in size or morphology, which supports the idea
that this species is monotypic (Jones and Carter, 1976).
The large sample from Kabalebo is composed of all males. These
individuals were captured in nets set along a newly cut trail, which
passed through the moderate undergrowth of a secondary forest along
a road, which eventually led into the larger trees of a mature rainforest.
Elsewhere the species usually was collected in either secondary or pri-
mary lowland rainforest. The exceptions to this were in the highlands
of Tafelberg where the typical vegetation was lower montane forest
320
Annals of Carnegie Museum
VOL. 55
and at Nieuwe Grond Plantation where a single specimen was taken
in nets set over the lawns and gardens surrounding the plantation
headquarters. M. nicefori shows a high correlation with the distribu-
tions of only seven other species; five of these— Lonchophylla thomasi,
Carollia perspicillata, Rhinophylla pumilio, Sturnim lilium, and Arti-
beus (large species)— were common, widespread species (Table 2). Glos-
sophaga soricina was not a particularly common species in Suriname.
The most interesting distributional correlation was with M. minuta,
which was taken at five of the nine localities where M. nicefori was
captured.
Length of testes of the 20 males from Kabalebo taken on 28 May
averaged 3.5 (range, 3-4). Other males had the following testes lengths
(date of capture in parentheses): 3, 4, 5 (26 May); 5 (6 August); 4 (23
October); 3 (24 October). None of the three females for which data are
available evinced reproductive activity (netted on 30 July, 12 Septem-
ber, and 30 September). Two of the males weighed 7.3 and 8 and three
females weighed 7, 8, and 8.5.
Surinamese specimens of M. nicefori had a 2/t = 28 and FTSF = 52.
The X-chromosome was submetacentric and the Y-chromosome was
acrocentric (Honeycutt et al., 1980; Baker et al., 1981).
Micronycteris (Xenoctenes) hirsuta (Peters, 1869)
Specimens examined (3).— Nickerie: Kabalebo, 1. Para: Zanderij, 2.
Micronycteris hirsuta is the sole member of the subgenus Xenoctenes
(Figs. 3, 4). Miller (1907) originally gave this taxon generic status, but
Sanborn (1949) reduced it to subgeneric level. The species M. hirsuta
can be distinguished from other members of the genus in Suriname by
the following characteristics: ears connected across the forehead by a
low unnotched band (Fig. 1); upper inner incisors separated at base,
but in contact near tip (Fig. 2C); upper outer incisors small; lower
incisors high and wedged tightly between canines (Fig. 2C); lower ca-
nines in contact, or nearly so, behind incisors; and lower incisors bifid
(Fig. 2C). The species is considered to be monotypic (Jones and Carter,
1976).
Our specimens were the first members of the species (Fig. 5) to be
reported from Suriname (Genoways et al., 1981). The specimens from
Zanderij, a reproductively inactive adult female and an immature fe-
male, were netted on 18 May. The specimen from Kabalebo was a
reproductively inactive adult female. The area at Kabalebo was covered
by mixed primary and secondary lowland rainforest, whereas the vi-
cinity of Zanderij was secondary lowland forest associated with a rub-
ber plantation. M. hirsuta was taken in association with 24 other species
of bats (Table 2); however, only four species— Phyllostomus elongatus,
1986
Genoways and Williams “Suriname Micronycteris
321
P. hastatus, Carollia perspicillata, and large Artibeus species— were
captured at both localities. All of these are common, widespread species
in Suriname.
The specimens from Suriname had a 2« = 30 and FN = 32 (Baker
et al,, 1981). This karyotype appears to be identical to the one found
in Middle American populations of M. hirsuta but differs from that
found on Trinidad where 2n = 28 (Baker et al., 1973; Baker, 1979).
Discussion
Seven of the 1 0 recognized species of Micronycteris are known from
the small country of Suriname. As a general rule, specimens were taken
in association with forested habitats, particularly mature lowland rain-
forest. However, there was a relatively low correlation between the
occurrence of any one species of Micronycteris and any of the other
species of the genus (Table 2). Only M. minuta and M. nicefori were
taken together at more than half of their collecting sites. There are two
possible explanations for this fact. First, there can be, and probably
are, subtle differences in the microhabitat required by each of the species
within the forest. The other factor affecting this correlation may be that
the distribution of each species may be clumped around available roost-
ing sites. We saw this phenomenon in at least two places in Suriname.
At Brownsberg Nature Park, specimens of M. sylvestris were taken only
from a hollow tree. At Kabalebo, a large number of male M. nicefori
was captured only in one set of nets. If the occurrence of species of
Micronycteris is clumped around available roost sites and individuals
have relatively small home ranges, then there definitely should be a
reduction in the places that species co-occur.
The seven species of Micronycteris from Suriname form a gradient
in size starting with the small M. minuta and M. megalotis and pro-
gressing through the large M. daviesi at the opposite end of the scale.
The small species really form a species pair based upon size, although
M. minuta probably averages slightly smaller than M. megalotis for
most characters. Between these extremes fall (beginning with the small-
est) M. nicefori, M. sylvestris, and M. brachyotis, and finally M. hirsuta.
Micronycteris sylvestris and M. brachyotis cannot be distinguished on
size alone but there are numerous other useful characters to separate
them. Size is not the only character needed to separate several of the
species of Micronycteris in Suriname, but it is useful in narrowing the
number of comparisons that need to be made.
We stated earlier that the genus Micronycteris seemed to be oversplit
at the subgeneric level. The current arrangement was proposed by
Sanborn (1949) based primarily upon characters of the wings and ears.
Deviations from this arrangement have been suggested by Arnold et
322
Annals of Carnegie Museum
VOL. 55
Table 2.— Species of bats taken in association with specimens o/Micronycteris in Suri-
name. The numbers in each column represent numbers of localities.
Species of bats
M.
daviesi
M.
hirsuta
M.
megalotis
M.
minuta
M.
nicefori
M.
sylvestris
No. of localities for which
data are available
1
2
9
9
9
1
Saccopteryx bilineata
0
1
5
1
1
0
Saccopteryx canescens
0
0
2
0
0
0
Saccopteryx leptura
1
1
5
0
1
0
Cormura brevirostris
0
1
0
1
1
0
Pteronotus parnellii
1
1
3
4
3
1
Noctilio leporinus
0
0
0
1
1
0
Chrotopterus auritus
1
0
2
0
0
0
Micronycteris megalotis
0
0
—
1
2
0
Micronycteris minuta
0
0
1
—
5
1
Micronycteris nicefori
0
0
2
5
~
0
Micronycteris sylvestris
0
0
0
1
0
-=
Tonatia bidens
0
0
5
1
2
0
Tonatia brasiliense
0
0
1
1
0
0
Tonatia carrikeri
0
1
0
2
0
0
Tonatia schulzi
0
0
2
0
1
0
Tonatia silvicola
0
1
1
3
2
1
Mimon crenulatum
0
0
1
1
1
0
Lonchorhina aurita
0
0
0
1
1
0
Phyllostomus discolor
1
1
2
1
0
1
Phyllostomus elongatus
1
2
4
6
3
1
Phyllostomus hastatus
1
2
3
5
3
0
Phyllostomus latifolius
0
0
1
2
0
1
Trachops cirrhosus
1
1
2
0
0
0
Anoura caudifer
0
0
3
0
2
0
Lonchophylla thomasi
1
1
4
7
6
1
Glossophaga soricina
0
0
2
4
5
0
Carollia brevicauda
1
0
0
1
0
0
Carollia perspicillata
1
2
7
5
6
0
Rhinophylla pumilio
1
1
5
6
5
1
Ametrida centurio
0
0
0
1
0
0
Sturnira lilium
1
1
4
6
6
0
Sturnira tildae
0
0
1
2
2
1
Artibeus cinereus
1
0
1
5
4
0
Artibeus concolor
0
0
1
1
1
0
Artibeus (large species)
1
2
7
9
8
1
Uroderma bilobatum
1
1
1
3
3
0
Chiroderma trinitatum
0
1
0
1
1
0
Vampyressa bidens
0
0
2
0
0
0
Vampyressa brocki
0
0
0
1
1
0
Vampyrops aurarius
0
0
0
0
1
0
Vampyrops brachycephalus
0
0
2
1
2
0
Vampyrops helleri
1
1
2
4
3
0
Vampyrodes caraccioli
0
0
0
1
1
0
Mesophylla macconnelli
0
1
1
2
3
0
Desmodus rotundus
0
1
0
0
0
1986
Genoways and Williams —Suriname Micronycteris
323
Table 2. — Continued.
Species of bats
M.
daviesi
M.
hirsuta
M.
megalotis
M.
minuta
M.
nicefori
M.
sylvestris
Thyroptera tricolor
0
0
1
2
1
0
Natalus tumidirostris
0
0
0
1
0
0
Myotis nigricans
0
1
1
3
3
0
Eptesicus brasiliensis
0
1
4
1
3
0
Molossus ater
0
1
0
0
0
0
Molossus molossus
0
1
1
1
1
0
Total species (51)
16
24
36
41
35
10
al. (1983) based on electrophoretic studies. For phenotypic criteria a
useful classification might be devised by using characters of the ears
and the teeth at the front part of the dental arcade (canines and incisors).
The genus can be split into two major groups— in the first the ears are
connected by an interauricular band and the lower incisors are bifid,
and in the second the ears are not connected by an interauricular band
and the lower incisors are trifid. We suggest that future investigation
of subgeneric classifications of this genus examine these groupings and
any subgroupings within them.
Literature Cited
Andersen, K. 1906. On the bats of the genera Micronycteris and Gly phony cteris. Ann.
Mag. Nat. Hist., ser. 7, 18:50~-63.
Arnold, M. L., R. J. Baker, and R. L. Honeycutt. 1983. Genic differentiation and
phylogenetic relationships within two New World bat genera. Biochem. Sys. Ecol.,
11:295=303.
Baker, R. J. 1979. Karyology. Pp. 107=155, in Biology of bats of the New World
family Phyllostomatidae, Part III (R. J. Baker, J. K. Jones, Jr., D. C. Carter, eds.).
Spec. Publ. Mus., Texas Tech Univ., 16:1=441.
Baker, R. J., H. H. Genoways, W. J. Bleier, and J. W. Warner. 1973. Cytotypes
and morphometries of two phyllostomatid bats, Micronycteris hirsuta and Vam-
pyressa pusilla. Occas. Papers Mus., Texas Tech Univ., 17:1-10.
Baker, R. J., H. H. Genoways, and P. A. Seyfarth. 1981. Results of the Alcoa
Foundation-Suriname Expeditions. VI. Additional chromosomal data for bats
(Mammalia: Chiroptera) from Suriname. Ann. Carnegie Mus., 50:333-344.
Genoways, H. H., and S. L. Williams. 1979. Records of bats (Mammalia: Chiroptera)
from Suriname. Ann. Carnegie Mus., 48:323-335.
— . 1984. Results of the Alcoa Foundation-Suriname Expeditions. IX. Bats of the
genus Tonatia (Mammalia: Chiroptera) in Suriname. Ann. Carnegie Mus., 53:327-
346.
Genoways, H. H., S. L. Williams, and J. A. Groen. 1981. Results of the Alcoa
Foundation-Suriname Expeditions. V. Noteworthy records of Surinamese mam-
mals. Ann. Carnegie Mus., 50:319-332.
Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and
Tobago. Amer. Mus. Nat. Hist, 122:187=302.
324
Annals of Carnegie Museum
VOL. 55
Hall, E. R. 1981. The mammals of North America. John Wiley & Sons, New York,
l:xv + 1-600 + 90.
Hill, J. E. 1964. Notes on bats from British Guiana, with the description of a new
genus and species of Phyllostomatidae. Mammalia, 28:553-572.
Honeycutt, R. L., R. J. Baker, and H. H. Genoways. 1980. Results of the Alcoa
Foundation-Suriname Expeditions. III. Chromosomal data for bats from Suriname.
Ann. Carnegie Mus., 49:237-250.
Husson, a. M. 1978. The mammals of Suriname. Zool. Monog., Rijksmuseum Nat.
Hist., 2:xxiv + 1-569.
Jones, J. K., Jr., and D. C. Carter. 1 976. Annotated checklist, with keys to subfamilies
and genera. Pp. 7-38, in Biology of bats of the New World family Phyllostomatidae,
Part I (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas
Tech Univ., 10:1-218.
Koopman, K. F. 1978. Zoography of Peruvian bats with special emphasis on the role
of the Andes. Amer. Mus. Novitates, 2651:1-33.
Koopman, K. F., and E. L. Cockrum. 1967. Bats. Pp. 109-150, in Recent mammals
of the World (S. Anderson and J. K. Jones, Jr., eds.), The Ronald Press Co., New
York, viii + 453 pp.
LaVal, R. K. 1977. Notes on some Costa Rican bats. Brenesia (Museo Nacional de
Costa Rica), 10/11:77-83.
Medellin, R, A., D. E. Wilson, and D. Navarro L. 1985. Micronycteris brachyotis.
Mamm. Species, 251:1-4.
Miller, G. S., Jr. 1907. The families and genera of bats. Bull. U.S. Nat. Mus., 57:
xvii + 1-282.
Sanborn, C. C. 1949. Bats of the genus Micronycteris and its subgenera. Fieldiana:
Zook, 31:215-233.
Williams, S. L., and H. H. Genoways. 1 980. Results of the Alcoa Foundation-Surina-
me Expeditions. II. Additional records of bats (Mammalia: Chiroptera) from Su-
riname. Ann. Carnegie Mus., 49:213-236.
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VOLUME 55 31 DECEMBER 1986 ARTICLE 14
ANNOTATED CHECKLIST OF THE MAMMALS OF KENYA.
II. INSECTIVORA AND MACROSCELIDEA
ISSA R. Agoundey'
Resident Museum Specialist, Section of Mammals
Duane A. Schlitter
Curator, Section of Mammals
Abstract
Four families of Insectivora (Tenrecidae, Chrysochloridae, Erinaceidae, Soricidae) and
one of Macroscelidea (Macroscelididae) are known from Kenya. Of these families, only
the Soricidae, with 31 species, and the Macroscelididae, with five species, are known
from more than a single species. Records of occurrence are annotated by museum spec-
imens or with references to the literature. Suncus murinus seems to be the only introduced
species in the insectivore fauna of Kenya. Distributional records of insect! vores in Kenya
are poor as 1 3 species of the 39 reported are known from only a single locality in Kenya.
A gazetteer of collecting localities is included.
Introduction
This is the second in a series of annotated checklists on Kenya mam-
mals, The first checklist (Aggundey and Schlitter, 1984) covered the
bats. This checklist includes the insectivoran orders Insectivora and
Macroscelidea. Our treatment of this group follows Yates (1984). In
this checklist we cover four Lipotyphlan families, namely the Tenreci-
dae, Chrysochloridae, Erinaceidae, and Soricidae, and a single Men-
otyphlan family, the Macroscelididae, of Kenya.
‘ Address: Curator of Mammals, National Museums of Kenya, P.O. Box 40658, Nairobi,
Kenya.
Submitted 27 January 1986.
325
326
Annals of Carnegie Museum
VOL. 55
In a series of papers, Dollman (1915<2, 1915^?, 1915c, \9\5d, 1915c,
1915/ 1916) reviewed the African species of Crocidura. This review
was followed by Hollister’s (1918) review of Kenyan insectivores in
the United States National Museum. In his massive checklist, Allen
(1939) covered all of the species of Kenyan insectivores recognized at
that time. This work is still important as a starting point in under-
standing the taxonomic relationships of these species. Since these early
papers, there have been no subsequent reviews in a wider context of
Kenyan insectivores except elephant shrews. Corbet and Neal (1965)
and Corbet and Hanks (1968) reviewed the taxonomy and distribution
of Kenyan species in their broader studies of elephant shrews. This
second annotated checklist attempts to synthesize the pertinent liter-
ature on all of the insectivores known from Kenya. The taxonomy of
many groups of insectivores, and particularly the family Soricidae in
Africa, is still in a state of confusion. Undoubtedly many taxonomic
changes and additional distributional records will be required before
a satisfactory arrangement is possible.
We have followed the format from the first part of the series (Ag-
gundey and Schlitter, 1984). Specimen records are included from the
following museums with their accepted acronyms in parentheses.
Carnegie Museum of Natural History, Pittsburgh (CM)
National Museums of Kenya, Nairobi (NMK)
National Museum of Natural History, Smithsonian Institution,
Washington, D.C. (USNM)
As with the checklist on bats, we point out that the distributional
records of this group are also undoubtedly incomplete. Nevertheless,
it is our sincere hope that the information given will stimulate addi-
tional interest and research on the insectivores of Kenya.
Acknowledgments
We were unable to spend as much time gathering distributional records from museum
collections for this checklist as we were for the checklist on bats. Nevertheless, we would
like to thank Charles O. Handley, Jr., for allowing us to record data at the Smithsonian
Institution and for sending photocopies of numerous old and difficult to find articles. In
Kenya, Mrs. Gooderis helped compile the list of the specimens of insectivores in the
collection in Nairobi. R. Hutterer critically reviewed the manuscript. Winnie Woodland
and Mary Ann Schmidt patiently and expertly typed numerous drafts of the manuscript.
We thank them all for their important part in completing this checklist.
This checklist resulted from the first author’s tenure as a Resident Museum Specialist
at Carnegie Museum of Natural History. This visit was made possible by the International
Program, endowed by the Richard King Mellon Foundation. Lastly, the senior author
wishes especially to thank Richard E. Leakey, Director of the National Museums of
Kenya, for making it possible for him to take six months study leave in the United
States.
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 327
Checklist
Order Insectivora
Family Tenrecidae
Genus Potamogale Du Chaillu, 1860
Potamogale velox Du Chaillu, 1860
Potamogale velox Du Chaillu, 1860. Proc. Boston Soc. Nat. Hist., 7:361. Gabon.
Records.— A km S Kakamega (Aggundey, 1977:368, NMK).
Family Chrysochloridae
Genus Lacepede, 1799
Chrysochloris stuhlmanni fasten (St. Leger, 1931)
Chlorotalpa fosteri St. Leger, 1931. Ann. Mag. Nat. Hist., (10)8:605, Dec. Mt. Elgon,
9300 ft, Uganda.
Records.— MX. Elgon (NMK); Cherangani Hills (Duncan and Wrangham, 1971:149;
Meester, 1974:3); southwest side Cherangani Hills, 6500 ft (Duncan and Wrangham,
1971:157); Labot, Cherangani Hills (Duncan and Wrangham, 1971:157).
Family Erinaceidae
Genus Atelerix Pomel, 1848
Atelerix albiventris (Wagner, 1841)
E{rinaceus) albiventris Wagner, 1841. Schreber’s Saugethiere, Suppl., 2:22. Type locality
unknown.
Erinaceus albiventris atratus Rhoads, 1896. Proc. Acad. Nat. Sci., Philadelphia, p. 544,
8 Dec. Lake Rudolf, Ngare Nocbor, Marsabit Dist., Kenya (ca. 2°45'N, 36°45'E).
Erinaceus hindei Thomas, 1910. Ann. Mag. Nat. Hist., (8)5:193, Feb. Kitui, 3500 ft,
Kenya.
Erinaceus sotikae Heller, 1910. Smithsonian Misc. Coll., 56(15):!, 23 Dec. Southern
Guaso Nyiro, Sotik Dist., Kenya.
Ngare Nocbor (Rhoads, 1896:544); Kitui (Peters, 1878:198; Anderson, 1895:
420; Thomas, 1910<3:193); Southern Guaso Nyiro (Heller, 1910^:1; Hollister, 1918:26;
J. Allen, 1922:16); Kapiti Plains (Hollister, 1918:26; J. Allen, 1922:16); Loita Plains
(Hollister, 1 9 1 8:26); Mt. Lololokwi (Hollister, 1 9 1 8:26; J. Allen, 1 922: 1 6); Taveta (True,
1892:469, 480; Hollister, 1918:26); Ulukenia Hills (Hollister, 1918:26; J. Allen, 1922:
16); Upper Ura River (J. Allen, 1922:16); Lokori, Southern Turkana (NMK); Naivasha
(NMK); Nairobi (Harmsen and Jabbal, 1968:158; Kingdon, 1974:32; Kock, 1978:116,
NMK); Busia (CM); Voi (Allen and Lawrence, 1936:39); Lodwar (St. Leger, 1937:526);
Olorgasailie (Toschi, 1949:28); River Kerio Suk (Ruxton, 1926:29); Narro Surra River
(Kollmann, 1914:319); Machakos (Lonnberg, 1912Z?:48).
Family Soricidae
Genus CwciWwm Wagier, 1832
Crocidum allex Osgood, 1910
Crocidura allex Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3):20, 7 Apr.
Naivasha, Kenya.
Crocidura alpina Heller, 1910. Smithsonian Misc. Coll., 56(9):5, 22 July. West slope of
Mt. Kenya, 10,000 ft, Kenya.
328
Annals of Carnegie Museum
VOL. 55
^FCor^/5.~Naivasha Station (Osgood, 1910^7:20; Hollister, 1918:67, USNM); West
slope Mt. Kenya (Heller, 1910^2:5; Hollister, 1918:68, USNM); Aberdare Range (Hol-
lister, 1918:67, USNM); Oljoro O Nyon River (Hollister, 1918:67, USNM); Man Forest
(Toschi, 1947:12, NMK); Ngong (NMK); Loita (NMK); Amala River (NMK); Selangai
(NMK); near Lake Olbollossat (Dollman, 1915/:513); Solai, 8000 ft (Dollman, 1915/:
514); Nanyuki (Southern and Hook, 1963:511); Mt. Kenya, 12,500 ft (Harmsen and
Jabbal, 1968:160); north slope Mt. Kenya (Coe and Foster, 1972:8).
Remarks. —WQim de Balsac and Meester (1977:9) regard both C a.
allex and C a. alpina as valid subspecies in Kenya.
Crocidura bottegi Thomas, 1898
Cwcidura bottegi Thomas, 1898. Ann. Mus. Civ. Stor. Nat. Genova, (2): 18, 677, 21
Feb. Between Badditu and Dime, northeast of Lake Rudolf, Ethiopia.
Records. — MditsdihiX (Heim de Balsac and Meester, 1977:10).
Crocidura elgonius Osgood, 1910
Crocidura bicolor elgonius Osgood, 1910. Ann. Mag. Nat. Hist., (8)5:369. Kirui’s, south-
ern foothills of Mount Elgon, 6000 ft, Kenya (restricted by Moreau et al., 1946:394).
RFCor<7s'. — Kirui’s (Osgood, 1910c:369; Allen and Lawrence, 1936:44; Loveridge, 1937:
519); Kisumu (Hollister, 1918:65); Lukosa River (Hollister, 1918:65, USNM); Kapiti
Plains (Hollister, 1918:65, USNM); Kaimosi (Hollister, 1918:65; Allen and Lawrence,
1936:44; Allen and Loveridge, 1942:159, NMK, USNM); Muguga (NMK); Lake Nakuru
(Hutterer, 1983:225); Nairobi (Hutterer, 1983:225); Cherangani Hills (Ruxton, 1926:
29); Eldoret (Loveridge, 1923:698); Mt. Elgon (Granvik, 1924:9).
Remarks. — Both Heim de Balsac and Meester (1977) and Hutterer
(1984) regard C elgonius as a distinct species.
Crocidura fischeri Pagenstecher, 1885
Crocidura fischeri Pagenstecher, 1885. Jahrb. Wiss. Anst., Hamburg, 2:34, pi. 1, f 1--3.
Nguruman, Kenya.
JRFCorflf5.““Nguruman (Pagenstecher, 1885:34; Hutterer, 1986:26).
Remarks. —ThQ type locality for the species was given by Pagen-
stecher (1885:34) as Nguruman with no other qualifier than Massai-
land. Neumann (1900) also visited Nguruman from 22-27 December
1893 and plots the locality on his map as slightly on the Tanzanian
side of the border (which seems to be the same as the present border).
He labels the locality on his map as “Nguruman (Bagasse)” which leads
us to believe he was referring to a village near Lake Natron or “Ngu-
ruman Salz-See” from his map. Lake Natron seems to have been at a
low level in 1893. Swynnerton and Hayman (1951:284) list the type
locality of Crocidura fischeri as Nguruman, west of Lake Magadi, Kenya
Colony [between L50' and 2®S, 36°5'E; 2500 to 6000 feet]. This locality
reaches from the Rift Valley floor to nearly up to the top of the eastern
face of the Nguruman Escarpment. Hutterer (personal communication)
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 329
has rechecked this type locality as part of a study (Hutterer, 1986) of
C fischeri and concluded that it is in Kenya based on Fischer’s (1884,
1885) accounts of his explorations and especially the photographs and
maps in these accounts. Fischer camped at the foot of a mountain range
northwest of “Natron Swamp” or Lake Natron at 650 meters, near the
foot of Mt. Sambo, where acacia steppe occurs and a flowing stream
came down from the mountains. Mt. Sambo, although on the Tan-
zanian side of the border, is visible far in the background of a pho-
tograph of the campsite. It seems best to follow Swynnerton and Hay-
man (1951) and Hutterer (1986) in placing the type locality on the
Kenyan side of the present border.
Crocidura flavescens nyansae Neumann, 1900
Crocidura flavescens nyansae Neumann, 1900. Zool. Jahrb., Syst., Geog., Biol., 13:544,
10 Oct. Fort Lubwa’s in Ussoga, outlet of Victoria Nyanza, Uganda (restricted to
Fort Thruston, 10 mi E Jinja, Busoga Dist., Uganda, by Moreau et al., 1946:396).
Crocidura kijabaej. A. Allen, 1909. Bull. Amer. Mus. Nat. Hist, 26:173. Kijabe, Kenya.
Records. — KAjabe (J. Allen, 1909:173); Elgeyo Forest (J. Allen, 1914:343); Aberdare
Mountains, 11,000 ft (Dollman, 1915^:568; Hollister, 1918:43); Laikipia (Hollister,
1918:43); Mt. Kenya (Dollman, 19156:568; Hollister, 1918:43); Mt. Umengo (Hollister,
1 9 1 8:43); Naivasha Station (Hollister, 1 9 1 8:43); Nakutishu River, Naivasha Plains (Hol-
lister, 1918:43); Nyeri (Dollman, 19156:568; Hollister, 1918:43); Kaimosi (Hollister,
1918:42; Allen and Lawrence, 1936:41); Kakamega (Hollister, 1918:42); Kisumu (Hol-
lister, 1918:42); Sergoit Lake (Hollister, 1918:42); Jombeni (Dollman, 19156:568); Mweru
(Dollman, 19156:568); Lake Olbollosat (Dollman, 19156:568); Kirui’s (Dollman, 19156:
567); Londiani (Lonnberg, 1918:175); Lake Elmenteita (Osgood, 1936:221); Molo (Os-
good, 1936:221); Mianzini (Thomas, 1891:182); Meru Country (Lonnberg, 19126:52);
Mt. Elgon (Granvik, 1924:8).
Crocidura fulvastra (Sundevall, 1 843)
Sorex fulvaster SnndQvaW, 1843. Kongl. Svenska Vet.-Akad. Handl., Stockholm, p. 172,
for 1842. Bahr-el-Abiad, Sudan.
North of Lokichokio (Hutterer, 1984:215).
Remarks. species includes C. sericea (Sundevall, 1843) as a
synonym according to Hutterer (1984:21 1, 215).
Crocidura fumosa fumosa Thomas, 1 904
Crocidura fumosa Thomas, 1904. Ann. Mag. Nat, Hist., (7)14:238, Sept. Western slope
of Mt. Kenya, 2600 m, Kenya.
Crocidura alchemillae Heller, 1910. Roosevelt’s African Game Trails, American ed., p.
480, London ed., p. 491. Summit of Aberdare range, Kenya.
Records.— WQsXQvn slope ofMt. Kenya, 2600 m (Thomas, 1904:238; Dollman, 191 5£’:
369, 370); summit of Aberdare range (Heller, 1910c:480; Hollister, 1918:55); Kinangop
(Kollmann, 1913:1 39); west side Mt. Kenya (Hollister, 1918:55, NMK); Fort Hall (Thomas,
1904:238; Kollmann, 1913:139; Hollister, 1918:55, NMK); Chyulu Hills (Osgood, 19106:
21, NMK); Nairobi (NMK); Ngong (NMK); Chania River (NMK); Naro Mom (NMK);
330
Annals of Carnegie Museum
VOL. 55
01 Arabel (NMK); Kikuyu (NMK); Thika (NMK); Nyeri (Hollister, 1918:55); Upper
Nzoia River (Hollister, 1918:55); Mt. Elgon (Dollman, 1915^:369, 370); Jombeni Range
(Dollman, 1915e:369, 370); Aberdare Mountains (Dollman, 1915^:369, 370); Machakos
(Thomas, 1904:238; Kollmann, 1913:139); Nandi (Thomas, 1904:238; Kollmann, 1913:
139); Kakamega (Thomas, 1904:238; Kollmann, 1913:139); Donya Sabuk (Lonnberg,
1916:5); Juja Farm (Lonnberg, 1916:5); Nanyuki (Southern and Hook, 1963:512); Mt.
Kenya (Southern and Hook, 1 963:5 1 2; Duncan and Wrangham, 1 97 1 : 1 6 1); 2^bra Farm,
Athi Plains (J. Allen, 1909:173); Lagari (Thomas, 1904:238); Blue Post (Lonnberg, 1912Z?:
53); Kagio (Lonnberg, 1912Z):53); Embu (Lonnberg, 1912Z?:53); Kanyakeni (Lonnberg,
1912Z?:53); Meru (Lonnberg, 1912Z):53); Mt. Kenya, 3800 m (Coe and Foster, 1972:8).
Remarks. — Y^onnhQrg (1 9 1 6:5) considered the series of 10 specimens
from Donya Sabuk to approach C / schist acea, now considered a
synonym of C luna. It is doubtful that all of these listed records rep-
resent C. fumosa as the differences between C fumosa, C. luna, and
C. zaodon are subtle ones. Many of these records could prove to be C.
luna upon reexamination.
Crocidura fuscomurina (Heuglin, 1865)
S.{orex) fusco murinus Heuglin, 1865. Leopoldina, 5:36, in Nova Acta Acad. Caes.
Leop.-Carol., Dresden, June. Meshra-el-Req, Bahr-el-Ghazal Prov., Sudan.
Crocidura ^/co/or Bocage, 1889. Jom. Sci. Math., Phys. e Nat., Lisboa, (2)1:29, March.
“Gambos, dans I’interieur de Mossamedes,” Angola.
Crocidura bicolor cuninghamei Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:240, Sept.
Vumba Island, 1 mi N Sagitu Island, Lake Victoria, Uganda.
Nairobi (Osgood, 1936:230).
Remarks. Hutterer (1 983) reviewed the taxonomy and distribution
of C fuscomurina but did not report any records from Kenya. His
preliminary distribution map (Hutterer, 1983:224) included only rec-
ords for which he had some degree of certainty of identification.
Crocidura hildegardeae Thomas, 1 904
Crocidura hildegardeae Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:240, Sept. Fort Hall,
Kenya.
Crocidura lutreola Heller, 1912. Smithsonian Misc. Coll., 60(1 2):8, 4 Nov. Mt. Mbololo,
Taita Hills, 5000 ft, Kenya.
Crocidura hildegardeae procera Heller, 1912. Smithsonian Misc. Coll., 60( 1 2): 1 0, 4 Nov.
Mt. Lololokwi, 6000 ft, northern Guaso Nyiro, Kenya.
Crocidura ibeana Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:514, May. Ann. Mag.
Nat. Hist., (8)16:362, Oct. Olgerei River, Kenya.
Crocidura hildegardeae altae Heller, 1912. Smithsonian Misc. Coll., 60(1 2):9, 4 Nov.
Mt. Garguez, Mathews Range, 6000 ft, Kenya.
Records. — Fort Hall (Thomas, 1904:240; Dollman, 1915/:508; Hollister, 1918:64,
USNM); Mt. Garguez (Heller, 1912:9, 10; Hollister, 1918:65, USNM); Mt. Lololokwi
(Heller, 1912:10; Hollister, 1918:64); Mt. Mbololo (Heller, 1912:8, 9; Hollister, 1918:
64; Allen and Lawrence, 1936:43, USNM); Olgerei River (Dollman, 1915<2:514); Voi
(Hollister, 1 9 1 8:64, NMK); Ngong (NMK); Narosura River (Kollmann, 1 9 1 4:3 1 9, NMK);
Lemek Valley (NMK); Amala River (NMK); Nyeri (Hollister, 1918:64, NMK); Meru
(Hollister, 1 9 1 8:64, NMK); Amboseli (NMK); Engare Narok (Hollister, 1 9 1 8:64, NMK);
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 331
Isiola River (Hollister, 1 9 1 8:64); Kapiti Plains (Hollister, 191 8:64); Mayo River, Laikipia
(Hollister, 1 9 1 8:64, USNM); Mt. Kenya (Hollister, 1 9 1 8:64); Mt. Sagalla (Hollister, 1918:
64, USNM); Mt. Umengo (Heller, 1912:9; Hollister, 1918:64, USNM); Naivasha Station
(Hollister, 1918:64, NMK); Ndi (Hollister, 1918:64, USNM); Oljoro O Nyon River
(Hollister, 1918:64); Wambugu (Hollister, 1918:64); Taveta (Dollman, 1915^:379); Tsa-
vo River (Dollman, 1915^:380); Mt. Elgon (Dollman, 1915/:508); Baringo (Dollman,
1915/:509); Kaimosi (Allen and Lawrence, 1936:43); Peccatoni (Allen and Lawrence,
1936:43); Wema (Allen and Lawrence, 1936:43); Kazere (Lonnberg, 1912Z):54); Blue
Post (Lonnberg, 1912^:54); Kutu (Lonnberg, 1912Z?:54).
Remarks. —We follow Demeter and Hutterer (1986) and Hutterer
(personal communication) in recognizing C hildegardeae for these
Kenyan records rather than C gracilipes Peters, 1870, an apparently
distinct species.
Crocidura hirta Peters, 1852
Crocidura hirta Peters, 1852. Reise nach Mossambique, Saugeth., p. 78, pi. 18, f. 2.
Tette, Mozambique (17°S).
Records.— I^OVLQ found.
Remarks.— Yieim de Balsac and Meester (1977:17) list C h. velutina
Thomas, 1904 to occur in . . presumably also Kenya and southern
Somalia.”
Crocidura Jacksoni Thomas, 1 904
Crocidura jacksoniThom^s, 1904. Ann. Mag. Nat. Hist., (7)14:238, Sept. Ravine Station,
Kenya.
Crocidura jacksoni amalae Dollman, 1915. Ann. Mag. Nat. Hist, (8)15:516, May; Ann.
Mag. Nat. Hist, (8)16:376, Oct. 1915. Amala River, NyanzaProv., Kenya (restricted
by Moreau et at, 1946:396 to Amala River, 30 mi N Kenya-Tanzania border, 5500
ft, Kenya).
Records.— Station (Thomas, 1904:239); Amala River (Dollman, 1914^:309,
1 9 1 5<3:5 1 6, NMK); Isiola River (Hollister, 1 9 1 8:60); Kaimosi (Hollister, 1 9 1 8:60, USNM);
Kapiti Plains (Hollister, 1 9 1 8:60); Mtito Andei (Hollister, 1 9 1 8:60, USNM); Neumann’s
Boma (Hollister, 1918:60); Southern Guaso Nyiro (Hollister, 1918:60); Ulukenia (Hol-
lister, 1918:60); Voi (Hollister, 1918:60, Allen and Lawrence, 1936:43, USNM); Na-
rosura River (Dollman, 1914/7:309, NMK); Loita Plains (NMK); Ngong (NMK); Fort
Hall (NMK); Tsavo River (Dollman, 1914^z:88, NMK); Yala River (NMK); Sultan
Hamud (NMK); Shimba Hills (NMK); Peccatoni (Allen and Lawrence, 1936:43); Gol-
banti (Allen and Lawrence, 1936:43); Zuwani (Dollman, 19 14^z:88); Lengototo (Dollman,
1914/7:309).
Crocidura littoralis Heller, 1910
Crocidura littoralis Heller, 1910. Smithsonian Misc. Colt, 56(1 5):5, 23 Dec. Butiaba,
east shore of Albert Nyanza, Uganda.
— Kaimosi (Hollister, 1918:68; Dippenaar, 1980:129).
Remarks.— Hollister (1918:68) referred this series to C maurisca
but Dippenaar (1980:130) regards this species as known only by the
holotype and that the specimens from Kaimosi are C. littoralis.
332
Annals of Carnegie Museum
VOL. 55
Crocidura luna Dollman, 1910
Crocidura luna Dollman, 1910. Ann. Mag. Nat. Hist., (8)5:175, Feb. Bunkeya River,
Katanga, Zaire.
Crocidura fumosa schistacea Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3):
20, 7 April. Lukenya Mountain, Kenya.
Crocidura raineyi Heller, 1912. Smithsonian Misc, Coll., 60(1 2):7, 4 Nov. Mt. Gargues,
Kenya.
Crocidura fumosa selina Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:510, May; Ann.
Mag. Nat. Hist., (8)16:371, Oct., 1915. Mabira Forest, Chagwe, Uganda.
Crocidura luna umbrosa Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:514, May; Ann.
Mag. Nat. Hist., (8)16:360, Oct., 1915. Machakos, 5400 ft, Kenya.
Records. —Lukenya Mountain (Osgood, \9\0b:2Q)\ Mt. Gargues (Heller, 1912:7; Doll-
man, 1915^:373; Hollister, 1918:60); Machakos, 5400 ft (Dollman, 1915^:514, \9\5e:
361); Ulukenia Hills (Hollister, 1918:59); Kapiti Plains (Hollister, 1918:59); Kaimosi
(Hollister, 1918:59); Mt. Elgon (Rode, 1935:167).
Remarks. de Balsac and Meester (1977:18) list only C. /.
schistacea occurring in Kenya and point out that C. raineyi could be
a valid large subspecies. Hollister (1 9 18:59) listed specimens from Kai-
mosi in western Kenya as C /. selina.
Crocidura macarthuri St. Leger, 1934
Crocidura macarthuri St. Leger, 1934. Ann. Mag. Nat. Hist., (10)13:559, May. Merifano,
20 mi from mouth of Tana River, Kenya.
Recor^/5.— Merifano (St. Leger, 1934:559; Hutterer, 1986:28); lOmiEMoyal, onMurri
Road, 1500 m (Heim de Balsac and Meester, 1977:19; Hutterer, 1986:28); Ijara (Percy;
et al., 1953Z?:11; Heim de Balsac and Meester, 1977:19; Hutterer, 1986:28); Rojewero
Plains (Hutterer, 1986:28).
Crocidura macowi Dollman, 1915
Crocidura macowi Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:515, May; Ann. Mag.
Nat. Hist., (8)16:378, Oct. 1915. Mt. Nyiro, south of Lake Rudolf, Kenya.
Records.-MX. Nyiro (Dollman, 1915^:515, 1915^:378).
Crocidura monax monax Thomas, 1910
Crocidura monax Thomas, 1910. Ann. Mag. Nat. Hist., (8)6:310, Sept. Rombo, Mt.
Kilimanjaro, 6000 ft, Tanzania.
Records.— River (Dollman, 1914Z?:309).
Crocidura nanilla Thomas, 1 909
Crocidura nanilla Thomas, 1909. Ann. Mag. Nat. Hist., (8)4:99, Aug. Probably Entebbe,
Uganda.
Crocidura denti St. Leger, 1932. Ann. Mag. Nat. Hist., (10)9:240, March, Koliokwell
River, North Turkana Dist., Kenya (not C. jacksoni denti Dollman, 1915).
Crocidura rudolfi St. Leger, 1932. Ann. Mag. Nat. Hist., (10)10:487, Nov. (renaming of
C. denti St. Leger, 1932).
— Koliokwell River (St. Leger, 1932:241).
1986 Aggundey AND ScHLiTTER— Kenyan Insectivora AND Macroscelidea 333
Remarks.— de Balsac and Meester (1977:20) place C rudolfl
as a synonym of C. nanilla.
Crocidum parvipes Osgood, 1910
Crocidura parvipes Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser., 10(3): 19, 7
Apr. Voi, Kenya.
Crocidura parvipes nisa Hollister, 1916. Smithsonian Misc. Coll., 66(8):2, May. Kibabe,
Kisumu, Kenya.
Eusso Nyiro Post (NMK); Voi (Osgood, 1910Z?:19; Hutterer, 1986:31); Ki-
babe (Hollister, 1 9 1 66:2; Hollister, 1 9 18:47, USNM); Embu (NMK); Fort Hall (Hollister,
1918:47); Mt. Sagalla (Heller, 1912:9; Hollister, 1918:60; Heim de Balsac and Meester,
1977:12, USNM).
Remarks. — Three specimens of shrews from Mt. Sagalla are reported
as C. parvipes by Heller (1912:9) but are referred to C jacksoni by
Hollister (1918:63). Heim de Balsac and Meester (1977:12) refer this
record again to C. cyanea parvipes. Both C. c. parvipes and C c. nisa
are recognized as valid in Kenya by Heim de Balsac and Meester (1 977:
12). Hutterer (1986:31) considered C. parvipes to be distinct from C.
cyanea.
Crocidura ultima Dollman, 1915
Crocidura ultima Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:517, May; Ann. Mag. Nat.
Hist., (8)17:204, Feb., 1916. Jombeni Range, Nyeri Dist., 5000 ft, Kenya.
Records. —iomhQni Range, 5000 ft (Dollman, 1915<2:517, 1916:205).
Remarks.— de Balsac and Meester (1977:20) plaee C ultima
as a synonym of C. monax, but Dippenaar (1980:130) considers C.
ultima to be a distinct species known only from the holotype.
Crocidura viaria (1. Geoffroy Saint-Hilaire, 1834)
Sorex viarus 1. Geoffroy Saint-Hilaire, 1834. Voyage aux Indes-Orientales par C. Belan-
ger, ZooL, p. 1 27. Senegal (restricted by Hutterer, 1984:209, to region between Dakar
and St. Louis).
Crocidura hindei Thomas, 1904. Ann. Mag. Nat. Hist., (7)14:237, Sept. Machakos,
Kenya.
Crocidura suahelae Heller, 1912. Smithsonian Misc. Coll., 60(1 2):6, 4 Nov. Mazeras,
Kenya.
Crocidura beta Dollman, 1915. Ann. Mag. Nat. Hist., (8)15:513, May; Ann. Mag. Nat.
Hist., (8)16:78, July, 1915. Chania River, Kenya (restricted to Chania River, near
Nyeri and Fort Hall, Kenya by Allen, 1939:31).
Machakos (Thomas, 1904:237; Hutterer, 1984:211); Mazeras (Heller, 1912;
6; Hollister, 1918:50, USNM); Chania River (Dollman, 1915^3!:513, 1915c:78); Chan-
gamwe (Hollister, 1918:50, UNSM); Juja Farm (Lonnberg, 19126:54; Hollister, 1918:
46); Ulukenia Hills (Hollister, 1918:46); Nairobi (Loveridge, 1923:698, NMK); Kajiado
(NMK); Potha (NMK); Ngatana (Allen and Lawrence, 1936:42); Mombasa (Hutterer,
1984:211); Karati (Thomas, 1904:237).
334
Annals of Carnegie Museum
VOL. 55
Remarks. (1984) placed C. sericea under C. fuivastra
(Sundevall, 1 843) but regarded C hindei and C suahelae as conspecific
with C viaria.
Crocidura voi Osgood, 1910
Crocidura voi Osgood, 1910. Publ. Field Mus. Nat. Hist., ZooL Sen, 10(3): 18; 7 April.
Voi, Kenya.
Crocidura percivali Dollman, 1915. Ann. Mag. Nat. Hist, (8)15:513, May; Ann. Mag.
Nat Hist, (8)16:126, Aug., 1915. Jombeni Range, Nyeri Dist, 3500 ft, Kenya.
Records.— Yoi (Osgood, 1910Z?:18; Hutterer, 1986:30) Jombeni Range (Dollman, 1915a:
5 1 3; Hutterer, 1 986:30); Lakiundu River (Hollister, 1 9 18:50); Mt. Suswa (Hutterer, 1 986:
30).
Remarks.— Hutterer (1986) recognized Crocidura voi as a distinct
species, and included C. butleri from Sudan, C percivali from Kenya,
and C aridula from Sudan as synonyms.
Crocidura xantippe Osgood, 1910
Crocidura xantippe Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10(3): 19, 7
April. Voi, Kenya,
Records. — Voi (Osgood, 19 10/?: 19; Dollman, 1915^:375; Heim de Balsac and Meester,
1977:25); Taveta (Dollman, 1915^:375); Nyiru (Heim de Balsac and Meester, 1977:25);
Tsavo (Heim de Balsac and Meester, 1977:25),
Crocidura yankariensis Hutterer and Jenkins, 1980
Crocidura yankariensis Hutterer and Jenkins, 1980. Bull. British Mus. (Nat. Hist.), Zool.,
39:305. Futuk, 16 km E Yankari Game Reserve boundary, Nigeria (9®50'N, 10®55'E).
Records. — West of Lake Rudolf, Kakuma, 50-60 mi NW Lodwar (Hayman, 1937:
531; Hutterer and Jenkins, 1983:195).
Crocidura zaodon Osgood, 1910
Crocidura turba zaodon Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10(3):21,
7 Apr. Nairobi, Kenya.
Crocidura turba provoeax Thomas, 1910. Ann. Mag. Nat, Hist., (8)6:1 12, July. Aberdare
Mountains, 1 1 ,000 ft, Kenya.
Crocidura turba iakiundae Heller, 1912. Smithsonian Misc. Colt, 60(1 2):6, 4 Nov. Lak-
iundu River, near junction with Northern Guaso Nyiro, Kenya,
Crocidura turba kempi Dollman, 1915. Ann, Mag. Nat, Hist,, (8)15:511, May; Ann.
Mag. Nat. Hist., (8)16:134, Aug., 1915. KimPs, southern foothills of Mount Elgon,
6000 ft, Kenya (restricted by Moreau et aL, 1946:397).
Records.— Nmmhi (Osgood, 1910Z?:21; Thomas, 1910^:113; Lonnberg, 1918:175,
NMK); Aberdare Mountains, 1 1,000 ft (Thomas, 1910Z):1 13; Dollman, 19 15a: 133; Hol-
lister, 1918:54, NMK); Lakiundu River (Heller, 1912:6; Lonnberg, 1912l?:54; Hollister,
1918:54); Kinangop (Kollmann, 1913:140); Kirafs, Mt. Elgon, 5000-6000 ft (Dollman,
191 5^7: 134); Sirgoit Lake (Hollister, 1918:54); Sirgoit (Hollister, 1918:54); Kakamega
(Hollister, 1918:54); Kibabe (Hollister, 1918:54); Naivasha Plains (Hollister, 1918:54);
Mt. Kenia, west slope (Hollister, 1918:54); Isiola River, head (Heller, 1912:7; Hollister,
1918:54); Archer's Post (Heller, 1912:7; Hollister, 1918:54); Mt. Mbololo (Hollister,
1 9 1 8:54); Mt. Umengo (Hollister, 1 9 1 8:54); Nzoia River, Guas Ngishu plateau (Hollister,
1986 Aggundey and Schlitter— Kenyan Insectivora and Macroscelidea 335
1918:54); Mt. Sagalla (Hollister, 1918:54); Yala River (NMK); Nyeri (NMK); Kaimosi
(Hollister, 1 9 1 8:54; Allen and Lawrence, 1 936:42, NMK); Kenna (NMK); Kericho (NMK);
Maua (NMK); Man Forest (Toschi, 1 947: 1 2, NMK); Kasigau (NMK); Kisumu (Hollister,
1918:54, NMK); Chania River (NMK); Amala River (Dollman, 1915d:132, NMK);
Kabete (NMK); Fort Hall (Hollister, 1918:54, NMK); Laikipia plateau, 15 mi N Nyeri
(Hollister, 1 9 1 8:54); Lukosa River (Hollister, 1 9 18:54); Northern Guaso Nyiro (Dollman,
1915€/:132); Jombeni Range (Dollman, 191 5£/: 132); Donya Sabuk (Lonnberg, 1916:6);
Cherengani Hills (Ruxton, 1926:29); Zuwani Swamp (Dollman, 1914^2:88); Nanyuki
(Southern and Hook, 1963:512); Thomson’s Falls (Southern and Hook, 1963:512); Mt.
Kenya, 1 2,500 ft (Harmsen and Jabbal, 1 968: 1 59); Mt. Kenia, 2450 m (Lonnberg, \9\2b:
54); Luazomela River (Lonnberg, 1912^:54); acacia steppe south of Guaso Nyeri (Lonn-
berg, 1912Z):54).
Crocidura zaphiri simiolus Hollister, 1916
Crocidura simiolus Hollister, 1916. Smithsonian Misc. Coll., 66(8):3, May. Kisumu,
Kenya.
Records. —Kisumu (Hollister, 19166:3; Hollister, 1918:51); Kaimosi (Hollister, 1918:
51); Kibabe (Hollister, 1918:51).
Genus Suncus Hemprich and Ehrenberg, 1832
Suncus infinitesimus infinitesimus (Heller, 1912)
Pachyura infinitesima Heller, 1912. Smithsonian Misc. Coll., 60(1 2):5, 4 Nov. Rumruti,
Laikipia Plateau, 7000 ft, Kenya.
Records.— KummXi (Heller, 1912:5).
Suncus lixus aequatorius (Heller, 1912)
Pachyura lixus aequatoria Heller, 1912. Smithsonian Misc. Coll., 60(1 2):4, 4 Nov. Sum-
mit of Mt. Sagalla, 4000 ft, Taita Hills, Kenya.
Recor<75.— Summit of Mt. Sagalla, 4000 ft (Heller, 1912:4; Hollister, 1918:41).
Suncus murinus (Linnaeus, 1766)
Sorex murina Linnaeus, 1766. Syst. Nat., 12th ed., 1:74. Java, Indonesia.
Records. (Heller, 1912:5); Witu (Heller, 1912:5).
Remarks. — This introduced species probably has a more widespread
distribution, especially along the coast, than these few records repre-
sent.
Genus Sylvisorex Thomas, 1 904
Sylvisorex grand mundus Osgood, 1910
Sylvisorex mundus Osgood, 1910. Publ. Field Mus. Nat. Hist., Zool. Sen, 10:18, 7 Apr.
Kijabe, Kenya.
Records.— YA^dibQ (Osgood, 19106:18); west side Mt. Kenya, 7000, 8500, 10,000 ft
(Hollister, 1918:39); Mt. Kenya (Duncan and Wrangham, 1971:161).
Remarks.— Tv^o additional locality records, perhaps Mt. Elgon and
Cherangani Hills, are plotted by Kingdon (1974:81). The former may
be Butandiga, Uganda, reported by Allen and Lawrence (1936:41).
336
Annals of Carnegie Museum
VOL. 55
Sylvhorex megalum gemmeus Heller, 1910
Sylvisorex gemmeus Heller, 1910. Smithsonian Misc. Coll., 56(1 5):7, 23 Dec. Rhino
Camp, Lado Enclave, Uganda.
Sylvisorex sorelloides Lonnberg, 1912. Ann. Mag. Nat. Hist., (8)9:67, Jan.; Kungl.
Svenska Vet.-Akad. Handl., Stockholm, (2)48(5):51, pi. 3, f. 1, 1912. Steppe near
Itiolu River, Northern Guaso Nyiro, Kenya.
Records. — Acacia steppe, near Itiolu River, south of Northern Guaso Nyiro (Lonnberg,
191 2^2:67, 1912Z?:51); Kaimosi (Hollister, 1918:39; Allen and Lawrence, 1936:41); Kirui
(Allen and Lawrence, 1936:41),
Genus Myosorex Gray, 1838
Myosorex (Surdisorex) norae (Thomas, 1906)
Surdisorex norae Thomas, 1906, Ann. Mag. Nat. Hist., (7)18:223, Sept. East side of
Aberdare Range, near Nyeri, Kenya.
Records.— EsisX side of Aberdare Range, near Nyeri (Thomas, 1906:224); Aberdare
Mountains, 10,000 to 11,000 ft (Hollister, 1918:37); Nr Kiandongoro Gate, Aberdare
Mountains, 8400 ft (Duncan and Wrangham, 1971:160).
Myosorex (Surdisorex) polulus (Hollister, 1916)
Surdisorex polulus Hollister, 1916. Smithsonian Misc. Coll., 66(1):1, 10 Feb. West side
of Mount Kenya, 10,700 ft, Kenya.
Records.— side of Mt. Kenya, 10,700 ft (Hollister, 1916<a!:l); west side of Mt.
Kenya, 9000 ft to 12,000 ft (Hollister, 1918:37); Mt. Kenya, 12,500 ft (Harmsen and
Jabbal, 1968:160); Mt. Kenya, 3960 m (Coe and Foster, 1972:8); Naro Mom track, Mt.
Kenya, 10,500 ft (Duncan and Wrangham, 1971:160).
Order Macroscelidea
Family Macroscelididae
Genus Petrodromus Peters, 1 846
Petrodromus tetradactylus sangi Heller, 1912
Petrodromus sultani sangi Heller, 1912. Smithsonian Misc. Coll., 60(12):12. Mt. Mbol-
olo, Taita Hills, 4000 ft, Kenya.
Records. -Mt. Mbololo, 4000 ft (Heller, 1912:12; Hollister, 1918:29; Allen and Law-
rence, 1936:39; Loveridge, 1937:526); Voi (Corbet and Neal, 1965:68).
Remarks. — Corbet and Neal (1965:68) reported a skull in the Berlin
Museum labelled “Kibwezi” but were uncertain if the record originated
from the Chyulu Hills in Kenya or on Mt. Meru in Tanzania.
Petrodromus tetradactylus sultani Thomas, 1897
Petrodromus sultani Thom^is, 1897. Proc. Zool. Soc. London, p. 435. Mombasa, Kenya.
Records.— R.i\QT Mombaca (Guenther, 1881:164); Mombasa (Thomas, 1897:435;
Neumann, 1900:541; Davis et al, 1968:844, CM, NMK); Mazeras (Heller, 1912:12;
Hollister, 1918:29); Gede (Corbet and Neal, 1965:67, Rathbun, 1979:16, NMK); Msa-
baha (NMK); Sokoke Forest (CM, NMK); Watamu (NMK); Jilari (Corbet and Neal,
1965:67); Taveta (Thomas, 1910c:309; Corbet and Neal, 1965:68; Corbet and Hanks,
1986 Aggundey and Schlitter — Kenyan Insectivora and M acroscelidea 337
1968:70); Shimba Hills (Heller, 1912:13); Rabai Hills (Thomas, 1897:435); Mrima Hill,
30 mi SW Mombasa (Harmsen and Jabbal, 1968:158^
Genus Rhynchocyon Peters, 1847
Rhynchocyon chrysopygus Guenther, 1881
Rhynchocyon chrysopygusGuQnthQr, 1881. Proc. Zool. Soc. London, p. 164, pi, 14. River
Mombaca, Kenya [=Mombasa, Kenya, according to Moreau et aL, 1946:392].
Records. — 'R.vvqt Mombaca (Guenther, 1881:164); Gede (Corbet and Hanks, 1968:65;
Rathbun, 1978:11, 1979:16, NMK); Mida (NMK); Msabaha (NMK); Sokoke Forest
(Corbet and Hanks, 1968:65; Rathbun, 1978:12, CM, NMK); Takaunga (Corbet and
Hanks, 1968:65); Arbagundi, Golana River (Corbet and Hanks, 1968:65); Malindi (Cor-
bet and Hanks, 1968:65).
Remarks.-— yioYQ2iU et al. (1946:392) restricted Guenther’s type lo-
cality of River Mombaca to Mombasa, Kenya. Corbet and Hanks
(1968:65) further qualify the type locality by suggesting a rather vague
interpretation of Mombasa as the type locality. R. chrysopygus is pres-
ently known only from north of Mombasa. Rathbun (1979:9) reports
observations at Kombeni River and Boni Forest.
Rhynchocyon petersi petersi Bocage, 1880
Rhynchocyon petersi Bocage, 1880. J. Sci. Math., Phys. Nat., Lisboa, (1)7:159, pi. 4, f.
2 (“Envoye de Zanzibar,” restricted to mainland of East Africa, region opposite of
Zanzibar, by Dollman, 1912:131).
Records. — MKZQTSiS (Hollister, 1918:28); Rabai Hills (Corbet and Hanks, 1968:64);
Shimba Hills (Corbet and Hanks, 1968:64).
Genus Elephantulus Thomas and Schwann, 1906
Elephantulus {Elephantulus) rufescens (Peters, 1878)
Macroscelides rufescens Peters, 1878. Monatsb. K. Preuss. Akad. Wiss,, Berlin, p. 198,
pi. 1, f. 3. Ndi, Kenya.
Macroscelides boranosThomdiS, 1900. Proc. Zool. Soc. London, 1900:802. Mega, western
Boran Galla, southeast of Lake Rudolf, Kenya.
Elephantulus dundasi Dollman, 1910. Ann, Mag. Nat. Hist., (8)5:95, Jan, Harich, near
Lake Baringo, 3000 ft, Kenya.
Elephantulus phaeus Heller, 1910. Smithsonian Misc. Coll., 56(1 5):8, 23 Dec. Njoro O
Solali, Sotik Dist., Kenya.
Elephantulus delicatus Dollman, 1911. Ann. Mag. Nat. Hist., (8)8:652. Orr Valley, Mt.
Nyiro, Kenya.
Elephantulus puicher rendilis Lonnberg, 1912. Kungl. Svenska Vet.-Akad. Hand!., (2)48(5):
49, 26 June. Thera, below Chanler Falls, Northern Guaso Nyiro, Kenya.
Elephantulus rufescens mariakanae Heller, 1912. Smithsonian Misc. Coll., 60(1 2): 10, 4
Nov. Mariakani, Kenya.
Records.— ^di (Peters, 1878:199); Mega (Thomas, 1900:803); Njoro O Solali, Sotik
(Heller, 1910Z?:8; Hollister, 1918:33); Harich (Dollman, 1910:96); Orr Valley, Mt. Nyiro
(Dollman, 191 1:653; Hollister, 1918:36); Thera (Lonnberg, 1912Zj:51); Mariakani (Hel-
ler, 1912:10; Hollister, 1918:33); Mtito Andei (Hollister, 1918:33); Voi (Hollister, 1918:
33; Allen and Lawrence, 1936:40; Corbet and Hanks, 1968:86, NMK); Kabalolot Hill,
Sotik (Hollister, 1918:33); Lime Springs, Sotik (Hollister, 1918:33); Loita Plains (Hoi-
338
Annals of Carnegie Museum
VOL. 55
lister, 1918:33, NMK); Southern Guaso Nyiro (Hollister, 1918:35); Telik River, Sotik
(Hollister, 1918:35); North Loroghi (Hollister, 191 8:35); NyamaNyango (Hollister, 1918:
35); Northern Guaso Nyiro River (Hollister, 1918:36); Longaya Water, Marsabit Road
(Hollister, 1918:36); Archer’s Post (Corbetand Hanks, 1968:86); Taveta (Thomas, 1910c:
309; Corbet and Hanks, 1968:86); ^thangaini (NMK); Kilungu (NMK); Limoni (NMK);
Ngari Nyiro (NMK); Southern Kidong (NMK); Lemek (NMK); 1 1 mi N Entesekera
(NMK); Emali (NMK); Sultan Hamud (NMK); Samburu (NMK); Lokori (NMK); Ka-
ruiru (NMK); Tam Desert (NMK); Golbanti (NMK); Kampi ya Samaki (NMK); Kan=
jangareng(NMK); Mt. Mbololo (Allen and Lawrence, 1936:40); Lodwar (St. Leger, 1937:
525); Wenje (Percy et ah, 1 953a: 1 16, 1 18); River Kerio Suk (Ruxton, 1926:29); Baringo
(Thomas, 1910c:310); Zuwani Swamp (Dollman, 1914a:88); Nanyuki (Southern and
Hook, 1963:51 1); 12 mi NW Kerio River (Dollman 1914Z?:309); Kerio River (Lonnberg,
1918:175); below Chanler’s Falls (Lonnberg, 1912^:51); Bushwackers (Rathbun, 1979:
16).
Elephantulus (Nasilio) bmchyrhynchus (A. Smith, 1836)
Macroscelides brachyrhynchus A. Smith, 1836. Report of the Expedition for Exploring
Central Africa, p. 42. Country between Lake Lakatoo and the Tropic.
Macroscelides delamerei Thomas, 1901. Ann. Mag. Nat. Hist., (7)8:155. Athi River,
6000 ft, Kenya.
Nasilio brachyrhynchus albiventer Os%oo(X, 1910. Publ. Field Mus. Nat. Hist., Zool. Ser.,
10(2): 13. Lake Elementeita, Kenya.
Records. ~ Axhi River (Thomas, 1901:155); Engare Narok River (Hollister, 1918:31,
NMK); Loita Plains (Hollister, 1918:31, NMK); Southern Guaso Nyiro (Hollister, 1918:
31); Ulukenia Hills (Hollister, 1918:31); Bargunett River (Hollister, 1918:31); Engare
Ndare River (Hollister, 1918:31); Lesiweru River, Mem Road (Hollister, 1918:31); Nai-
vasha Station (Hollister, 1918:31, NMK); Nyuki River (Hollister, 1918:31); Olorgesailie
(Toschi, 1949:27, NMK); Lemik Valley (NMK); Rumumti (NMK); Wame Hill, Konza
(NMK); Amala River (NMK); Voi (Allen and Lawrence, 1936:40); Narrosurra River
(Kollmann, 1914:319); Suswa (Kollmann, 1914:319); Guasso Nyero (Kollmann, 1914:
319); Lengototo (Dollman, 1914^:309); Lake Elementeita (Osgood, 19 10a: 13).
Gazetteer
Locality names are listed in alphabetical order with variant names cross-referenced to
the standard names. Standard names are taken from the second edition of the official
standard names gazetteer for Kenya published in 1978 and approved by the United
States Board on Geographic Names. Most of the entities can be identified and located
on the 1978 version of the Kenya and Northern Tanzania Route Map published in
English, French, and German by the Survey of Kenya.
Coordinates for locality names were taken mostly from the Kenyan gazetteer listed
above. In addition, Loveridge (1937), Moreau et al. (1946), Chapin (1954), and Davis
and Misonne (1964) were consulted together with place modifiers in the original refer-
ences for published records. In the case of rivers, when no place modifiers were available
for the published records or on the specimen labels, coordinates are given for the river
mouth or confluence.
In a number of instances, more than one entity exists in Kenya for a place name. This
generally does not cause a real problem but does in the case of the locality cited at Ewaso
Ngiro and its variants, especially the older specimens labeled Guasso Nyiro. In this latter
instance, we have given coordinates for both the southern and northern Ewaso Ngiro
rivers in the gazetteer.
Aberdare Mountains 0°25'S, 36°38'E
Aberdare Range 0°25'S, 36°38'E
1986 Aggundey and Schlitter — Kenyan Insecti vora and M acroscelidea 339
Amala River
r02'S, 35N4'E
Amboseli
2°40'S, 37N7T
Arbagundi
Archer’s Post
0°39'N, 37“41T
Athi River
r27'S, 36®59'E
Bargunett River [=Burguret River]
0°01'S, 36°56'E
Baringo [=Mukiitan]
0°38'N, 36°16'E
Blue Post
Burguret River
0°0rS, 36°56'E
Busia
0°28'N, 34®06'E
Chanler’s Falls
0°47'N, 38®05'E
Changamwe
4‘’0rS, 39°38'E
Chania River
r02'S, 37W'E
Cherangani Hills
riS'N, 35°27'E
Chyulu Hills
2°35'S, 37°50'E
Donya Sabuk [=ol Doinyo Sapuk]
r06'S, 37N5'E
Eldoret
0°3rN, 35°17'E
Elgeyo Forest
0°46'N, 35°3rE
Email
2®05'S, 37®28'E
Embu
0°32'S, 37“27'E
Engare Nanyuki
0°2FN, 36®55'E
Engare Narok
r09'N, 36®35'E
Engare Ndare River [=Engare Ondare]
0“35'N, 37®23'E
Engare Ondare
0°35'N, 3T23'E
Eusso Nyiro Post [=Archer’s Post]
0‘’39'N, 37®4rE
Entasekera
rsrs, 35®5rE
Entesekera [= Entasekera]
rSl'S, 35“5rE
Ewaso Ngiro (Northern)
0°37'N, 36"55'E~
Ewaso Ngiro (Southern)
0°28'N, 39°55'E
0°35'S, 35M7'E==
Fort Hall [=Muranga]
2W'S, 36®07'E
0°43'S, 37“09'E
Cede
3H8'S, 40°0rE
Golbanti
2®27'S, 40®12'E
Guasso Nyero [=Ewaso Ngiro]
Guaso Nyiro River, Sotik District
[=Ewaso Ngiro (Southern)]
Harich [=Marich]
1®32'N, 35®27'E
Horr Valley
2®10'N, 36“55'E
Ijara
r36'S, 40®3FE
Ilkaputiei
r38'S, 37®00'E
Isiola River [=Isiolo River]
0®34'N, 37°35'E
Isiolo River
0°34'N, 37°35'E
Itiolu River [=Isiolo River]
0‘’34'N, 37“35'E
Jilari [=Jilore]
3°1FS, 39“54'E
Jilore
3nrS, 39°54'E
Jombeni [=Nyambeni]
0°13'N, 57‘’52'E
Jombeni Range [=Nyambeni Range]
0°20'N, 37®57'E
Juja Farm
TIPS, 37°07'E
Kabalolot Hill
ca. rOO'S, 35°23'E
Kabete
ri6'S, 36°43'E
Kagio
0°40'S, 37N3'E
Kaimoni [=Kaumoni]
r44'S, 37“35'E
Kaimosi
OW'N, 34®5rE
340 Annals of Carnegie Museum
VOL. 55
Kajiado
1°5LS,
36®47'E
Kakamega
0°17'N,
34°45'E
Kakuma
3°43'N,
34®52'E
Kampi ya Samaki
0°36'N,
36®01'E
Kanyakeni [=Kanyekine]
0°08'S,
37040'E
Kanyangareng
1°47'N,
35'08'E
Kanyekine
0°08'S,
37°40'E
Kapiti Plains [=Ilkaputiei]
1°38'S,
37“00'E
Kapsabet
0°12'N,
35®06'E
Karati
0‘’26'S,
37°27'E
Karuiro
0°37'S,
37°07'E
Karuiru [= Karuiro]
0°37'S,
37°07'E
Kasigau
3°50'S,
38°40'E
Kathera
0°03'S,
37°35'E
Kaumoni
1°44'S,
37°35'E
Kazere [= Kathera]
0°03'S,
37°35'E
Kenna [=Kinna]
0°19'N,
38°12'E
Kericho
0°22'S,
35°17'E
Kerio River
2°59'N,
36°34'E
Kibabe [=Kibabet]
0°11'N,
35°15'E
Kibabet
O^ll'N,
35°15'E
Kijabe
0°56'S,
36°34'E
Kikuyu
1°15'S,
34°40'E
Kilungu
R48'S,
37°22'E
Kinangop
0°44'S,
36°40'E
Kinna
0°19'N,
38°12'E
Kirui, Mt. Elgon [=Kirui’s]
0°50'N,
34M0'E
Kirui’s
0°50'N,
34°40'E
Kisumu
0°06'S,
34°45'E
Kithangaini
1°29'S,
37°23'E
Kitui
1°22'S,
38°01'E
Koliokwell River
Kutu
m
o
O
37°19'E
Lagari [=Lugari]
0°39'N,
34°53'E
Laikipia
0°25'N,
36°45'E
Laikipia Plateau
0°25'N,
36°08'E
Lake Elementeita [=Lake Elmenteita]
0'’27'S,
36°15'E
Lake Elmenteita
0°27'S,
36“15'E
Lake Ilpolosat
0°09'S,
36°26'E
Lake Nakuru
0°22'S,
36°05'E
Lake Olbollosat [=Lake Ilpolosat]
0°09'S,
36®26'E
Lake Olbollossat [=Lake Ilpolosat]
0“09'S,
36®26'E
Lake Sergoi
0°42'N,
35°25'E
Lakiundu River [=Ngaramara River]
0°36'N,
37°37'E
Lamu
2°16'S,
40“54'E
Lemek
1°06'S,
35°23'E
Lemek Valley
ro9's,
35®19'E
Lemik Valley [=Lemek Valley]
1°09'S,
35°19'E
Lengototo [=Lenkutoto]
1°39'S,
35°58'E
Lenkutoto
r39'S,
35‘’58'E
Lesiweru River
Lime Springs [=Maji Moto]
1°20'S,
35°42'E
Lodwar
3°07'N,
35°36'E
1986 Aggundey and Schlitter — Kenyan Insecti vora
AND Macroscelidea 341
Loita
r30'S,
35®4rE
Loita Plains
r20'S,
35°32'E
Lokichokio
4®2rN,
34“2rE
Lokori
r57'N,
36®0rE
Londiani
OHO'S,
35°36'E
Longaya Water
ca. r07'N,
37®38T
Lorogi
roo'N,
36“5rE
Luazomela River
0°29'N,
37®40'E
Lugari
0°39'N,
34°53'E
Lukenya
r3rs,
36"58'E
Lukenya Hills
r28'S,
37°03'E
Lukenya Mountain [=Lukenya Hills]
r28'S,
37°03'E
Lukosa River
0°12'N,
34°56'E
Machakos
rsrs,
37H6'E
Maji Moto
r20'S,
35"42'E
Malindi
40“07'E
Mariakani
3°52'S,
39“28'E
Marich
r32'N,
35®27'E
Marsabit
2°20'N,
37°59'E
Mau Forest
0°20'SA)M0'S,
35°25'E-
G6°05'E
Maua
0°14'N,
37°56'E
Mayo River
OHO'S,
37^0 1'E
Mazeras
BOSS'S,
39033, E
Mega
Merifano
2H9'S,
40®08'E
Meru
0°03'N,
37"39'E
Mianzini
ca. 0“55'S,
36®25'E
Mida
3H9%
39®58'E
Molo
OHS'S,
35°44'E
Mombasa
4“03'S,
39M0'E
Mt. Elgon
r08'N,
34°33'E
Mt. Garguez [=Warges]
0°57'N,
37®24'E
Mount Lololokwi [=01 Doinyo Sabachi]
0«50'N,
37°32'E
Mt. Kenia [=Mt. Kenya]
OHO'S,
37°20'E
Mt» Kenya
OHO'S,
37“20'E
Mt. Mbololo
3®17'S,
38°28'E
Mt. Nyiro [=0 1 Doinyo Ngiro]
2°08'N,
36"5rE
Mt. Sagalla
3“27'S,
38°35'E
Mt. Umengo
ca. 3“18'S,
38°19'E
Moyale
3"32'N,
39°03'E
Mrima Hill
4‘»29'S,
39H6'E
Msabaha
3H6'S,
40°03'E
Mtito Andei
2°4rs,
38H0T
Muguga
ril'S,
36°39'E
Mukutan
0°38'N,
36°16'E
Muranga
0®43'S,
37“09'E
Mweru
0“40'S,
37°05'E
Nairobi
ri7'S,
36®49'E
Naivasha
0M3'S,
36°26'E
Naivasha Plains
OMl'S,
36°27'E
Naivasha Station
0°43'S,
36®26'E
Nakatishu River
0®33'S,
36°38'E
342 Annals of Carnegie Museum
VOL. 55
Nandi [=Kapsabet]
0°12'N,
35°06'E
Nanyuki
0°0LN,
37°04'E
Naro Moru
OHIO'S,
37°0rE
Narosura River
1°33'S,
35°53'E
Narrosurra River [=Narosura River]
1°33'S,
35°53'E
Ndi
3®14'S,
38°30'E
Neumann’s Boma [=Samburu Game Lodge]
0°34'N,
37035, E
Ngari Nyiro [=Ewaso Ngiro]
Ngaramara River
0°36'N,
37°37'E
Ngare Nocbor
ca. 2°45'N,
36°45'E
Ngatana
2°13'S,
40°11'E
Ngong
1°22'S,
36°39'E
Njoro O Solali
0°28'S,
35°04'E
North Laroghi [=Lorogi]
FOO'N,
36°51'E
Northern Guaso Nyiro [=Ewaso Ngiro]
0®37'N, 36°55'E~
0°28'N,
39°55'E
Nyahururu
0°02'N,
36°22'E
Nyama Nyango [=Samburu Game Lodge]
0°34'N,
37°35'E
Nyambeni
0°13'N,
37°52'E
Nyambeni Range
0°20'N,
37°57'E
Nyeri
0°25'S,
36°57'E
Nyiru [=01 Doinyo Ngiro]
2°08'N,
36°51'E
Nyuki River [=Engare Nanyuki]
0°21'N,
36°55'E
Nzoia River
0°03'N,
33°57'E
01 Arabel
0“18'N,
36°18'E
01 Doinyo Ngiro
2®08'N,
36°51'E
01 Doinyo Sabachi
0®50'N,
37°32'E
01 Doinyo Sapuk
ro8's,
37°15'E
Olgerei River
r43'S,
35°18'E
Olijoro O Nyon River
ca. 0°57'S,
35°55'E
Olorgasailie
r34'S,
36°27'E
Orr Valley [=Horr Valley]
2°10'N,
36°55'E
Peccatoni
2°25'S,
40°43'E
Potha
r34'S,
37°10'E
Rabai Hills
Ravine Station
0°01'N,
35°43'E
River Kerio Suk
2‘’59'N,
36°07'E
River Mombaca [=Mombasa]
4®03'S,
39°40'E
Rojewero Plains
0°il'N,
38°10'E
Rumruti [=Rumuruti]
onh'N,
36°32'E
Rumuruti
0“16'N,
36®32'E
Samburu
3M6'S,
39°17'E
Samburu Game Lodge
0®34'N,
37°35'E
Selengai
2®irN,
37°10'E
Sera
LOl'N,
37°53'E
Sergoi
0°39'N,
35°23'E
Sergoit Lake [=Lake Sergoi]
0M2'N,
35°25'E
Shimba Hills
4®13'S,
39°25'E
Sirgoit [= Sergoi]
0®39'N,
35°23'E
Sirgoit Lake [=Lake Sergoi]
0“42'N,
35°25'E
Sokoke Forest
3‘’29'S,
39°50'E
Solai
o°orN,
36®09'E
Southern Guaso Nyiro [=Ewaso Ngiro]
2®04'S,
36'’07'E
1986 Aggundey and Schlitter— Kenyan Insectivora and Macroscelidea 343
Southern Kedong Valley
ca. 1°24'S, 36°27'E
Southern Kidong [= Southern Kedong Valley]
ca. r24'S, 36®27L
Sultan Hamud
2°0rS, 37°22'E
Takaungu
3“4rS, 39®5LE
Talek River
r26'S, 35W'E
Tara Desert
3“45'S, 39“08'E
Taveta
3“24'S, 37°4rE
Teiek River [=Talek River]
r26'S, 35®04'E
Thera [=Sera]
rOl'N, 37®53'E
Thika
r03'S, 37°05'E
Thomson's Falls [=Nyahuraru]
0°02'N, 36“22'E
Tsavo
2®59'S, 38°28'E
Tsavo River
2°59'S, 38“3rE
Ulukenia [=Lukenya]
1°31'S, 36°58'E
Ulukenia Hills [=Lukenya Hills]
1°28'S, 37°03'E
Upper Nzoia River
ca. 0®53'N, 35®22'E
Upper Ura River
ca. 19U0'N, 37®59'E
Voi
3«23'S, 38®34'E
Wambugu
0°35'S, 37®02'E
Wame Hill [-Wami Hill]
r39'S, 37“08T
Wami Hill
r39'S, 37°08L
Warges
0°57'N, 37“24'E
Watamu
3®21'S, 40°0rE
Wema
2U3'S, 40UrE
Wenje
1M7'S, 40“06'E
West Slope Mt. Kenya
OUO'S, 37°10'E
Witu
2®23'S, 40®26'E
Yala River
0°04'N, 34®09'E
Ziwani
3'’23'S, 37'’47'E
Ziwani Swamp
3°16'S, 37°47'E
Zuwani [=Ziwani]
3°23'S, 37°47'E
Zuwani Swamp [=Ziwani Swamp]
3°16'S, 37M7'E
Literature Cited
Aggundey, I. R. 1977, First record of Potamogaie velox in Kenya. Mammalia, 41:
368.
Aggundey, I. R., and D. A, Schlitter. 1984. Annotated checklist of the Mammals
of Kenya. I. Chiroptera. Ann. Carnegie Mus., 53:1 19-161.
Allen, G, M. 1939, A checklist of African mammals. Bull. Mus. Comp, ZooL, 83:1-
763.
Allen, G. M., and B. Lawrence. 1936. Scientific results of an expedition to rain forest
regions in eastern Africa. III. Mammals. Bull. Mus. Comp. ZooL, 79:31-126.
Allen, G, M,, and A. Loveridge. 1942. Scientific results of a fourth expedition to
forested areas in East and Central Africa. 1. Mammals. Bull. Mus. Comp, ZooL, 89:
147-214.
Allen, J. A. 1909. Mammals from British East Africa, collected by the Tjader Expe-
dition of 1906. Bull. Amer. Mus. Nat. Hist, 26:147-175.
1914. Mammals from British East Africa, collected on the third African ex-
pedition of the American Museum by William S. Rainsford. Bull. Amer. Mus. Nat.
Hist, 33:337-344.
■ — •. 1 922. The American Museum Congo Expedition Collection of Insectivora.
Bull. Amer. Mus. Nat. Hist, 47:1-38.
344
Annals of Carnegie Museum
VOL. 55
Anderson, J. 1895. On a new species of the genus Erinaceus from Somaliland. Proc.
ZooL Soc. London, 1895:414-421.
Chapin, J. P. 1954. Gazetteer for “The Birds of the Belgian Congo.” Bull. Amer. Mus.
Nat. Hist., 75B:638-738.
Coe, M. j., and J. B. Foster. 1972. The mammals of the northern slopes of Mt. Kenya.
J. E. African Nat. Hist. Soc. Natl. Mus., 131:1-18.
Corbet, G. B., and J. Hanks. 1 968. A revision of the Elephant-shrews, family Macro-
scelididae. Bull. Br. Mus. Nat. Hist. (Zool.), 16(2):47-1 1 1.
Corbet, G. B., and B. R. Neal. 1965. The taxonomy of the Elephant shrews of the
genus Petrodromus, with particular reference to the East African coast. Rev. Zool.
Bot. Africaines, 71:49-78.
Davis, D. H. S., R. B. Heisch, D. McNeill, and K. F. Meyer. 1968. Serological survey
of plague in rodents and other small mammals in Kenya. Trans. R. Soc. Trop. Med.
Hyg., 62:838-861.
Davis, D. H. S., and X. Misonne. 1964. Gazetteer of collecting localities of African
rodents. Doc. Zool. Mus. Roy. African Cent., Turveren, 7:viii + 1-100.
Demeter, A., and R. Hutterer. 1986. Mammals from Mt. Meru, Tanzania. Cim-
bebasia, Ser. A, 8:199-207.
Dippenaar, N. j. 1 980. New species of Crocidura from Ethiopia and northern Tanzania
(Mammalia: Soricidae). Ann. Trans. Mus., 32:125-154.
Dollman, G. 1910. New African mammals. Ann. Mag. Nat. Hist., (8)5:92-97.
. 1911. On new mammals from East Africa, presented to the British Museum
by Mr. A. Blaney Percival. Ann. Mag. Nat. Hist., (8)8:652-659.
. 1912. A new Elephant Shrew from the Island of Zanzibar. Ann. Mag. Nat.
Hist., (8)10:130-131.
. 1914(3. Notes on mammals collected by Dr. Christy in the Congo and by Dr.
Bayer in Uganda and British East Africa. Rev. Zool. Africaine, 4:75-90.
. \9\Ab. Notes on a collection of East African mammals presented to the British
Museum by Mr. G. P. Cosens. Proc. Zool. Soc. London, pp. 307-318.
. 1915(3. On the African Shrews belonging to the genus Crocidura. Ann. Mag.
Nat. Hist., (8)15:507-527.
. \9\5b. On the African Shrews belonging to the genus Crocidura. 11. Ann. Mag.
Nat. Hist., (8)15:562-575.
. 1915c. On the African Shrews belonging to the genus Crocidura. III. Ann. Mag.
Nat. Hist., (8)16:66-80.
. \9\5d. On the African Shrews belonging to the genus Crocidura. IV. Ann. Mag.
Nat. Hist., (8)16:124-146.
. 1915c. On the African Shrews belonging to the genus Crocidura. V. Ann. Mag.
Nat. Hist., (8)16:357-380.
. 1915/ On the African Shrews belonging to the genus Crocidura. VI. Ann. Mag.
Nat. Hist., (8)16:506-514.
. 1916. On the African Shrews belonging to the genus Crocidura. VII. Ann. Mag.
Nat. Hist., (8)17:188-209.
Duncan, P., and R. W. Wrangham. 1971. On the ecology and distribution of sub-
terranean insectivores in Kenya. J. Zool., London, 164:149-163.
Fischer, G. A. 1884. Bericht uber die im Auftrag der Geographischen Gesellschaft in
Hamburg untemommene Reise in das Massai-Land. Mtt. Geogr. Ges. Hamburg,
1882/83:30-99.
. 1885. Uebersicht der in Ostafrika gesammelten Vogelarten, mit Angabe der
verschiedenen Fundorte. J. Om., 1885(170):! 13-142, 1 map.
Granvik, H. 1924. On mammals from the eastern slopes of Mount Elgon, Kenya
Colony. Mammals collected by the Swedish Mount Elgon Expedition 1920. Lunds
Univ. Arsskr. Ny Foljd, Avd. 2, 2 1(3): 1-32.
1986 Aggundey AND ScHLiTTER™ Kenyan Insectivora AND Macroscelidea 345
Guenther, A. 1881. Notes oe the species of Rhynchocyon and Petrodromus. Proc.
ZooL Soc. London, 1881:163-164.
Harmsen, R., and L Jabbal. 1968. Distribution and host-specificity of a number of
fleas collected in south and central Kenya. J. E. African Nat. Hist. Soc. Natl. Mus.,
27(2): 157-161.
Hayman, R. W. 1937. Mammals collected by the Lake Rudolf Rift Valley Expedition,
1934. Postscript Ann. Mag. Nat. Hist, (10) 19:530-531.
Heim de Balsac, H., and J. Meester, 1977. Order Insectivora. Part 1:1-29, in The
mammals of Africa: an identification manual (J. Meester and H. W. Setzer, eds.),
Smithsonian Inst. Press, Washington, D.C.
Heller, E. 191 Oa. Descriptions of seven new species of East African mammals. Smith-
sonian Misc. Coll, 56(9): 1-5.
— = — . 19106. New species of insectivores, bats, and lemurs from British East Africa,
Uganda, and the Sudan. Smithsonian Misc. Coll., 56(1 5): 1-8,
” — — . 1910c. [Species description in footnote to Appendix B, p. 480] in Theodore
Roosevelt^s African game trails. An account of the African wanderings of an Amer-
ican hunter-naturalist, Charles Scribner’s Sons, New York, 529 pp.
■ — — , 1912, New races of insectivores, bats, and lemurs from British East Africa.
Smithsonian Misc. ColL, 60(1 2): 1-1 3.
Hollister, N. 191 6a. Descriptions of a new genus and eight new species and subspecies
of African mammals. Smithsonian Misc. ColL, 66(1): 1-8.
. 19166. Three new African shrews of the genus Crocidura. Smithsonian Misc.
ColL, 66(8): 1-3.
“ — - — 1918. East African mammals in the United States National Museum. Part 1.
Insectivora, Chiroptera, and Carnivora. Bull. U.S. Natl. Mus., 99:1-194.
Hutterer, R. 1983. Taxonomy and distribution of Crocidura fuscomurina (Heuglin,
1865). Mammalia, 47:221-227.
• 1984. Status of some African Crocidura described by Isidore Geoffrey Saint-
Hilaire, Carl J. Sundevall, and Theodor von Heuglin. Pp, 207-217, in Proc. 3rd Int.
Coll. EcoL Taxon. African Small Mammals, Antwerp, Belgium, 20-24 July 1981
(E. Van Der Straeten, W. N. Verheyen, and F. DeVree, eds.), Ann. Mus. Roy. Afr,
Centr,, Sc. ZooL, 237:1-227 (for 1983).
. 1986. African shrews allied to Crocidura fischeri: taxonomy, distribution and
relationships. Cimbebasia, Ser. A, 8(4):23“35.
Hutterer, R., and P. D. Jenkins. 1 983. Species-limits of Crocidura somalica Thomas,
1895 and Crocidura yankariensis Hutterer and Jenkins, 1 980. Z. Saugetier,, 48: 1 93-
201.
Kingdon, j. 1974. East African mammals. An atlas of evolution in Africa. Volume
II, Part A (Insectivores and Bats). Academic Press, London, xii + 392 pp.
Kock, D. 1978. Vergleichende Untersuchung einiger Saugetiere im sudlichen Niger
(Mammalia: Insectivora, Chiroptera, Lagomorpha, Rodentia). Seeckenberg BioL,
58:113-136 (for 1977).
Kollmann, M. 1913. Note sur les Mammiferes rapportes de FAfrique orieetale par
MM. Alluaud et Jeaenel. Bull. Mus. Nat. Hist. Nat., Paris, 19:138-144.
— — 1914, Liste des Mammiferes rapportes par M. Guy Babault du British East
Africa. Bull. Mus. Nat. Hist Nat, Paris, 20:319-321.
Lonnberg, E. 191 2a. Some new mammals from British East Africa. Ann. Mag. Nat.
Hist, (8)9:63-67.
. 19126. Mammals collected by the Swedish Zoological Expedition to British
East Africa 1911, Kungl. Svenska Vet Hand!., (2)48(5): 1-1 88.
— 1916. Mammals collected by H. R. H. Prince Vilhelm’s expedition to British
East Africa 1914. Ark. ZooL, 10(12):l-32,
346
Annals of Carnegie Museum
VOL. 55
. 1918. Mammiferes recueillis dans le region du Mont Elgon par le Dr. Bayer,
en 1914. Rev. Zool. Africaine, 5:172-192.
Loveridge, a. 1923. Notes of East African mammals, collected 1920-1923. Proc.
Zool. Soc. London, 1923:685-739.
. 1937. Scientific results of an expedition to rain forest regions in eastern Africa,
IX. Zoogeography and itinerary. Bull. Mus. Comp. Zool., 79:481-541.
Meester, J. 1974. Family Chrysochloridae. Part 1.3: 1-7, in The mammals of Africa:
an identification manual (J. Meester and H. W. Setzer, eds.), Smithsonian Inst. Press,
Washington, D.C.
Moreau, R. E., G. H. E. Hopkins, and R. W. Hayman. 1946. The type-localities of
some African mammals. Proc. Zool. Soc. London, 115:387-447,
Neumann, O. 1900. Die von mir in den Jahren 1892-95 in Ost- und CentraLAfrica,
speciell in den Massai-Landem und den Landem am Victoria Nyanza gesammelten
und beobachteten Saugethiere. Zool. Jb. Syst., 13:529-562.
Osgood, W. 1910<2. Diagnoses of new East African mammals, including a new genus
of Muridae. Publ. Field Mus. Nat. Hist., Zool. Ser., 10:5-13.
. 1910A Further new mammals from British East Africa. Publ. Field Mus. Nat.
Hist., Zool. Ser., 10:15-21.
. 1910c. Two new African shrews. Ann. Mag. Nat. Hist., (8)5:369-370.
. 1936. New and imperfectly known small mammals from Africa. Publ. Field
Mus. Nat. Hist., Zool. Ser., 20:217-256.
Pagenstecher, H. a. 1885. Die von Dr. G, A. Fischer auf der in Auftrage de geogra-
phischen Gesellschaft in Hamburg, Untemommenen Reise in das Massai-Land
gesammelten Saugethiere. Jahrb. Wiss. Anst. Hamburg, 2:31-46.
Percy, H. E., R. C. Percy, and M. W. Ridley. 1953<2. An expedition to the Tana
River, Kenya colony. Oryx, 2:1 1 1-1 19.
Percy, R. C., H. E. Percy, and M. W. Ridley. 1953Z?. The water-holes at Ijara,
Northern Province, Kenya. J. E. Afr. Nat. Hist. Soc., 22:1-14.
Peters, W. 1878. Uber die von Hm. J. M. Hildebrandt wahrend seiner letzten ostaf-
rikanischen Reise gesammelten Saugethiere und Amphibian. Monatsb. K. Preuss.
Akad. Wiss., Berlin, pp. 194-209.
Rathbun, G. B. 1978. Evolution of the rump region in the Golden-rumped elephant
shrew. Bull. Carnegie Mus. Nat. Hist., 6:11-19.
. 1979. The social structure and ecology of elephant-shrews. Advances in Ethol-
ogy, 20:1-77.
Rhoads, S. N. 1896. Mammals collected by Dr. A. Donaldson Smith during his ex-
pedition to Lake Rudolf, Africa. Proc. Acad. Nat. Sci. Philadelphia, 1896:517-546.
Rode, P. 1935. Mammalia. In Mission Scientifique de fOmo, 2(9): 167-1 83.
Ruxton, a. E. 1926. On mammals collected by Captain C. R. S. Pitman, Game
Warden, Entebbe, Uganda. Ann. Mag. Nat. Hist., (9)18:28-33.
St. Leger, j. 1932. On two new mammals from Uganda. Ann. Mag. Nat. Hist., (10)9:
240-242.
. 1934. On a new species of Shrew from Merifano, East Africa. Ann. Mag. Nat.
Hist., (10)13:559-560.
. 1937. Mammals collected by the Lake Rudolf Rift Valley Expedition, 1934.
Ann. Mag. Nat. Hist., (10)19:524-530.
Southern, H. N., and O. Hook. 1 963. Notes on breeding of small mammals in Uganda
and Kenya. Proc. Zool. Soc. London, 140:503-515.
SwYNNERTON, G. H., AND R. W. Hayman. 1951. A checklist of the land mammals of
the Tanganyika Territory and the Zanzibar Protectorate. J. E. Afr, Nat. Hist. Soc.,
20(6/7):274-392 (for 1950).
Thomas, O. 1891. On a collection of small mammals made by Mr. F. J. Jackson in
eastern Africa. Proc. Zool. Soc. London, 1891:181-187.
1986 Aggundey and Schlitter—Kjenyan Insectivora and Macroscelidea 347
■ 1 897. [New African mammals. Exhibition of small mammals collected by Mr.
Alexander Whyte during his expedition to the Njika plateau and the Masuku moun-
tains, N. Nyasa]. Proc. ZooL Soc. London, 1897:430-436.
. 1 900. List of mammals obtained by Dr, Donaldson Smith during his recent
journey from Lake Rudolf to the Upper Nile. Proc. Zool. Soc. London, 1900:800-
807.
— — 1901. List of small mammals obtained from Mr. A. E. Pease, M. P., during
his recent expedition to Abyssinia, with descriptions of three new forms of Macro-
scelides. Ann. Mag. Nat. Hist, (7)8:154-156.
— — — . 1904. On Shrews of British East Africa. Ann. Mag. Nat Hist., (7)14:236-241.
— . 1906. Two new genera of small mammals discovered by Mrs. Holms-Tam in
British East Africa. Ann. Mag. Nat. Hist., (7)18:222-226.
■ . 1910fl. Further new African mammals. Ann. Mag. Nat Hist, (8)5:191-202.
— ■. 1910^. A new genus of Fruit-Bats and two new Shrews from Africa. Ann. Mag.
Nat Hist, (8)6:111-114.
— — — . 1910c. List of mammals from Mount Kilimanjaro, obtained by Mr. Robin
Kemp, and presented to the British Museum by Mr. C. D. Rudd. Ann. Mag. Nat.
Hist, (8)6:308-316.
Toschi, a. 1947. Su alcune comunita di vertebrati del massiccio del Mau (Kenya).
Ric. Zool. Appl. Caccia, SuppL, 2(2): 1-24.
— — — . 1949. Note ecologiche su alcuni mammiferi di Olorgasailie. Ric. Zool. Appl
Caccia, SuppL, 2(3):25~63.
True, F. W. 1892. An annotated catalogue of the mammals collected by Dr, W, L.
Abbott in the Kilima-njaro region, East Africa. Proc. U.S. Natl. Mus., 15:445-480.
Yates, T. L. 1984. Insectivores, elephant shrews, tree shrews, and dermopterans. Pp.
1 17-144, in Orders and families of Recent mammals of the world (Sydney Anderson
and J. Knox Jones, Jr., eds.), John Wiley and Sons, New York, viii +
453 pp.
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VOLUME 55 31 DECEMBER 1986 ARTICLE 15
DEVONIAN AND MISSISSIPPIAN CONULARIIDS OF
NORTH AMERICA. PART A. GENERAL
DESCRIPTION AND CONULARIA
Loren E. Babcock
Rodney M. Feldmann^
Research Associate, Section of Invertebrate Fossils
Abstract
The systematics, morphology and paleoecology of the new phylum Conulariida pro-
posed here, from the Devonian and Mississippian rocks of North America are reviewed
in this two-part work. Conulariids are identified by an elongate, pyramidal exoskeleton
comprising a framework of calcium phosphate rods, with or without spines and nodes,
covered by an integument made of thin layers of calcium phosphate and protein. Con-
ulariids were gregarious invertebrate animals that were attached to substrata by means
of calcium phosphate or chitinous stalks. These animals were exclusively marine and
limited to rocks of the Ordovician through the Triassic. Conulariids are found in all
types of marine facies and most probably had either a pseudoplanktonic or a benthonic
lifestyle.
Three valid genera of conulariids are recognized in the Devonian and Mississippian
of North America. These are Conularia Miller in Sowerby, 1821, Paraconularia Sinclair,
1940a and Reticulaconularia Babcock and Feldmann, n. gen. The genus Diconularia
Sinclair, 1952 is considered to be a junior synonym of Conularia. Prior to 1986, 69
species-level taxa of conulariids were described from the Devonian and Mississippian
of North America. Herein, and in Part B, 28 species are recognized as valid. Eleven
species are assigned to Conularia and are described in Part A.
‘ Address: Department of Geology, Kent State University, Kent, Ohio 44242.
2 Present address: Department of Geology, University of Kansas, Lawrence, Kansas
66045.
Submitted 2 June 1986.
349
350
Annals of Carnegie Museum
VOL. 55
Introduction
Conulariids, members of an extinct phylum of invertebrates, have
been described in the literature for a span of nearly 200 years. In that
time, well over 400 species, subspecies or varieties, and over 40 genera
have been named.
The purpose of this paper, and Part B of the same work, is to present
taxonomic and morphologic information obtained from an examina-
tion of conulariids from Devonian and Mississippian strata of North
America. Approximately 1 2% of all species-level conulariid taxa have
been described based upon specimens collected from these rocks. Spec-
imens described herein were collected only from the United States and
Canada; conulariids are not known to have been collected in Mexico.
This work involves: 1 , studies of intraspecific and interspecific variation
in morphology, ideally based upon large numbers of specimens; 2,
studies of the stratigraphic and geographic distributions of species; and
3, analyses of conulariid anatomy, functional morphology and paleo-
ecology, as the fossil record permits. Each taxon identified from De-
vonian or Mississippian rocks of North America is accompanied by a
new, or in some cases, the first, illustration, and a description high-
lighting points of morphology now considered of greatest taxonomic
value.
In the course of examining the Devonian and Mississippian forms,
it became clear that it was necessary to revise the terminology related
to morphology. In so doing, it was considered essential that conulariids
from a much wider range of geological and stratigraphical occurrences
had to be examined than those treated systematically herein. Thus, the
detailed morphology and terms are intended to be applicable to all
organisms referable to the Conulariida.
Part A of this work comprises a general description of the hard- and
soft-part morphology of conulariids, occurrences and paleoecology of
Devonian and Mississippian taxa of North America, a summary of the
taxa treated both in this paper and in Part B, a key to the Devonian
and Mississippian conulariid taxa of North America, and descriptions
of species of that group which are referred to the genus Conuiaria. Part
B of this work contains descriptions of species referable to the genera
Paraconularia and Reticulaconularia n. gen. and specimens described
in the literature as conulariids but which are here rejected from the
phylum. Locality descriptions and measurements of selected specimens
are included as appendices to Part B. Figures are numbered consecu-
tively in both Parts A and B in order to avoid cross-reference confusion.
Morphology
General. --When preserved in three dimensions, the exoskeleton of
a conulariid generally has a four sided, bilaterally symmetrical, elongate
1986
Babcock and Feldmann— North American Conulariida
351
pyramidal shape (Fig, 1.1). The profile may be modified by the de-
velopment of one or more exoskeletal constrictions (Fig. LI). The
exoskeleton, as preserved, generally ranges from 2 to 10 cm in length
in full grown individuals. In a few species, however, the exoskeleton
may attain a length in excess of 20 cm (Fletcher, 1938; Lamont, 1946;
Sinclair, 1948; herein, Fig. 1 1.2). In nearly all instances, the conulariid
exoskeleton diverges in width gradually and uniformly from a closed
apical end to an open apertural end. The apical end may be closed
either by a (morphological) apex (Fig. 1.1) or by an apical wall (Fig.
1.2). Presumed soft-parts consist of a single tubular structure that runs
internally along the length of the exoskeleton and at least one globular
body (Fig. 2.1).
Morphologic terms. --Tht literature on conulariid morphology in-
cludes important review papers by Slater (1907), Boucek and Ulrich
(1929), Kiderlen (1937), Richardson (1942), Sinclair (1948, 1952),
Moore and Flarrington (1956a, 1956/?) and Babcock and Feldmann
(1986). Much misunderstanding of conulariid morphology has arisen
from terms that have been improperly defined or undefined, from terms
that are ambiguous and from terms that imply systematic affinities.
Babcock and Feldmann (1986) proposed a set of morphological terms
for conulariids that described morphological features without intro-
ducing unnecessary connotations of genetic affinities. Below is a list of
morphological terms applied to conulariids, modified from Babcock
and Feldmann (1986), together with terms here considered synony-
mous and which relate to the morphology of members of the phylum
but are inappropriate in the light of advances made during this study.
AD APERTURAL SPINE —long spine projecting from near the adapertural side of a
rod, in the direction of the aperture.
AD APICAL SPINE— short spine projecting from near the internal adapical side of a
rod, in the direction of the apex.
ALIMENTARY TRACT— narrow, elongate, essentially tubular soft-part structure run-
ning the length of the central cavity. Synonyms: axial element, intestine.
ANGULATED CIRCULAR CURVE — style of rod articulation in which two abutting
rods on a face form a broad arcuate, adapically concave ridge, interrupted by a slight
adapertural point at the midline, and by gentle adapertural turns in the vicinity of
the facial margins.
APERTURAL CONSTRICTION— exoskeletal constriction located nearest the aper-
ture. Synonym: wrinkle.
APERTURE— opening at widest end of exoskeleton. Synonyms: base, mouth, opening.
APERTURAL TERMINATION— rounded or bluntly subtriangular extension of exo-
skeleton on each face at widest end of exoskeleton.
APEX (MORPHOLOGICAL APEX)— narrowest termination of exoskeleton, where the
four faces Join at a closed point. Synonyms: (biological) apex, summit. Compare
with hypothetical apex.
APICAL ANGLE— hypothetical angle formed by one face of the exoskeleton; measured
at the intersection of two lines each identified by tracing positions on the exoskeleton
exoskeleton
352
Annals of Carnegie Museum
VOL. 55
aperture
ridge
interridge area
midline
corner groove
corner angle
^^>«exoskeletal constriction
rod angle
face
apical wall
apical angle
hypothetical apex
Fig. 1.— Conulariid morphology, exhibited by a generalized Pamconularia. 1.1; exo-
skeleton with stalk attached. 1.2; apical region with stalk removed. Morphological terms
are explained in the text.
1986
Babcock and Feldmann— North American Conulariida
353
Fig. 2.~Conulariid morphology, exhibited by a generalized Paraconularia. 2.1; cutaway
view of exoskeleton showing internal soft-parts. Structure of the soft-parts in the apertural
region is problematic. 2.2; detailed view of two rods. Morphological terms are explained
in the text.
354
Annals of Carnegie Museum
VOL. 55
tangential to the facial margins and defining the maximum angle of separation.
Synonym: facial angle. See major apical angle and minor apical angle.
APICAL WALL —broadly rounded, adapically convex, portion of integument lacking
rods which completely covers the apical end of the exoskeleton when the apex itself
is missing. Synonyms: apical septum, apical diaphragm, basal limitation, diaphragm,
internal partition, Schott, septum.
CENTRAL CAVITY — region located internal to the four faces of the exoskeleton. Syn-
onym: body cavity.
CORNER ANGLE— longitudinal line in the marginal region of a face connecting points
of greatest inflection of the rods. Synonym: shoulder.
CORNER GROOVE — longitudinal invagination of exoskeleton connecting points where
pairs of rods from adjacent faces cross near the marginal terminations of those rods.
Synonyms: angular furrow, articulating suture, edge, lateral channel, longitudinal
channel, marginal furrow, marginal groove, side furrow.
EXOSKELETAL CONSTRICTION— depression, restricted in the longitudinal direc-
tion, traceable on all four faces of the exoskeleton in the same relative position.
Compare with apertural constriction.
EXOSKELETON (SKELETON)— four sided pyramidal structure, open at the widest
end and closed at the narrowest end, comprising rods joined by integument. Syn-
onyms: periderm, pyramid, shell, test.
FACE— one of four sides of the exoskeleton crossed by ridges; it is delimited by the
aperture, by the apex or the apertural wall and by two comer grooves. Synonyms:
side, surface, wall. See major face and minor face.
GLOBULAR BODY— large internal soft-part stmcture, subovoid in outline, located
near the aperture. Synonym: esophagus.
GOTHIC ARCH — style of rod articulation in which two adjacent rods on a face form
ridges that meet at an obtuse, adapically concave angle at the midline and proceed
away from the midline along lines subtly curved adapically.
HYPOTHETICAL APEX— point in space where two lines, traced along the mean di-
rection of the comer angles; meet; the hypothetical apex may or may not coincide
with the position of the (morphological) apex.
INFLECTED CIRCULAR CURVE — style of rod articulation in which two adjacent
rods on a face form a broadly arcuate, adapically concave ridge except in the vicinity
of the facial margins, where they turn gently adaperturally.
INFLECTED GOTHIC ARCH— style of rod articulation in which two adjacent rods
on a face form ridges that meet at an obtuse, adapically concave angle at the midline
and proceed away from the midline along lines subtly curved adapically except in
the vicinity of the facial margins, where they turn gently adaperturally.
INTEGUMENT— multilayered, presumably flexible, stmcture composed of calcium
phosphate and protein, within which rods and spines were embedded and held in
position. Synonyms: periderm, test.
INTERRIDGE AREA— roughly transverse band of integument located between two
facial ridges. Synonyms: intercostal space, interspace, transverse furrow, transverse
sulcus, space.
INTERRIDGE CREST —raised area, usually a linear ridge, located in an interridge area
and positioned at a right angle to a ridge; formed by integument covering an ad-
apertural or adapical spine. Synonyms: bar, intercostal longitudinal striation, vertical
striation, longitudinal bar, longitudinal striation.
INTERRIDGE FURROW— low area, usually linear, located in an interridge area, and
between two interridge crests.
INTERROD AREA— open region located between two rods; exposed only when integ-
ument is absent.
MAJOR APICAL ANGLE— apical angle subtended by a major face.
1986
Babcock and Feldmann— North American Conulariida
355
MAJOR FACE —wider of two adjacent faces.
MARGIN (FACIAL MARGIN)— longitudinal edge of a face, or a line connecting points
where two faces meet in a comer groove.
MIDLINE— longitudinal line connecting points where either two adjacent rods on a face
meet, or central to the facial terminations of each pair of adjacent rods if the rods
do not meet. The midline can be expressed as either a thin groove or a raised line
if the integument is preserved. The midline seems to be pigmented in some speci-
mens. Synonyms: central face furrow, central facial groove, facial groove, facial
midline, longitudinal carina, median groove, median line, mesial furrow, mid-line,
middle line, parietal line, septum, stmctural channel.
MINOR APICAL ANGLE— apical angle subtended by a minor face.
MINOR FACE— narrower of two adjacent faces.
NODE— minute, subcircular, raised surface on a rod or ridge. Synonyms: granule, papilla,
tubercle, pustule, wart.
RIDGE (FACIAL RIDGE)— raised line crossing a face from a comer groove to the
midline area, and formed by integument covering a rod. Synonyms: costa, crenu-
lation, ornamental rib, plica, plication, rib, riblet, transverse line, transverse rib,
transverse ridge, transverse striation.
ROD —narrow, elongate stmcture that is subcircular in cross section, composed of cal-
cium phosphate, and embedded within the integument; it is thickened near the
marginal termination, and tapers very gradually to a blunt point at the facial ter-
mination.
ROD ANGLE— angle subtended by a line connecting the two most distant points of a
rod along a longitudinal line and a line constmcted perpendicular to the facial margin
at the point where that ridge intersects the comer angle. Synonym: angle at the mid-
line.
ROD PAIR— two rods on a face whose distal ends meet, or approach closely, at the
midline.
SKELETON— See exoskeleton.
SPINE— solid, narrow, short or elongate stmcture, projecting from, and whose axis is
at a right angle to, a rod; tapers gradually to a sharp point distally. See adapical
spine and adapertural spine.
STALK— elongate stmcture, possibly chitinous, phosphatic, or chitinophosphatic, which
articulates proximally with a conulariid apex; distally, the stmcture seems to attach
to a substratum of uncertain nature.
Abandoned morphological terms. ~T\iQ following terms, previously
used in connection with the description of conulariids, are here con-
sidered inappropriate for various reasons including: 1, the structures
have been shown to be taphonomic in origin; 2, the structures were
described from organisms which should be excluded from the phylum
Conulariida; 3, the structures have been shown to be absent in conu-
lariids; or 4, the structures were described from dubious fossil material.
ANUS.
APPENDIX.
APERTURAL LOBE, Synonyms: apertural flap, apeitural lip, flap, laterales, lobe, lip,
mouth flap.
ATTACHMENT DISC.
BODY WALL.
EYE LENS.
HINGE.
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FEATHER EDGE.
KEEL.
LATERAL LINE. Synonym: longitudinal line, line near midline.
MOUTH PANEL.
NERVE CENTER.
NET-SHAPED EPITHELIUM.
ORNAMENTATION, Synonym: sculpture.
PLATE. Synonym: transverse plate,
SEPTUM: Synonyms: carina, internal longitudinal rib, internal rib, internal rod, internal
septum, longitudinal septum,
SKIN-MUSCLE LINING.
URULA.
WALL OPENING,
Aperture. --ThQ aperture (Fig. LI) appears to have been a simple
opening at the larger termination of the pyramidal exoskeleton. This
region has been the subject of much speculation. ‘"Flaps” or “lappets,”
partially or wholly closing the apertural region, were first described by
Miller {in Sowerby, 182 IX and subsequently by Etheridge (1901), Las-
eron (1912), Richter and Richter ( 1 930), Reed (1933), Kowalski (1935),
Kiderlen (1937), Sinclair (1948), Moore and Harrington (1956Z?), Bran-
isa (1965) and others. Kiderlen (1937), Termier and Termier (1949)
and Moore and Harrington (1956fi) proposed elaborate mechanisms
for closure of the apertural region involving the infolding of a flexible
exoskeleton. They assumed the line of flexure to be a straight line
normal to the midline. In one mechanism of closure, the exoskeleton
not only folded along a straight line perpendicular to a face, but also
collapsed like a bellows in the vicinity of the comer angles (Moore and
Harrington, 1956/?, Fig. 43.1), Moore and Harrington (1956/?, p. F57)
suggested that, in order for a conulariid to have been so flexible in the
apertural region, the line of flexure at the base of each “apertural flap”
was chitinophosphatic, while the remainder of the exoskeleton was
phosphatic. No chemical data were presented in support of this hy-
pothesis.
Specimens exhibiting closed or partially restricted apertures are com-
mon. Over 200 such specimens were observed in the course of this
study (for example, Figures 8.1, 10.1, 28.2). Among these, there is no
evidence of a consistent line of flexure (for example, Fig. 7. 1-7.2, 8.7=-
8.9, 10.2, 18.2, 22.2-22.3). Typically, the line along which a flap is
developed is not straight. Instead, the line of flexure often mimics the
style of rod articulation or seems to be arbitrary. No two adjacent faces
on the same specimen necessarily fold inward at the same position.
Within the same species, there is no consistency from individual to
individual either in the placement of a line of flexure or in the mode
of closure, as defined by Kiderlen (1937), Termier and Termier (1949)
or Moore and Harrington (1956/?). Furthermore, “apertural flaps” have
1986
Babcock and Feldmann— North American Conulariida
357
been observed at multiple sites on single conulariid exoskeletons (Bab-
cock and Feldmann, 1986, fig. IE). Their presence at various places
on the exoskeleton of different specimens indicates that they are taph-
onomic phenomena. Richter and Richter (1930), noted the extreme
flexibility of the holotype of Conularia tulipa from the Hunsruckshiefer
(Lower Devonian) of Germany. In their opinion, the conulariid exo-
skeleton was flexible enough to have collapsed under its own weight.
Infoldings of exoskeleton commonly found in the apertural region, and
less commonly elsewhere on a conulariid, are probably taphonomic
structures resulting from collapse of the exoskeleton after death. The
exoskeleton may not be quite as weak as suggested by Richter and
Richter, but it is certainly not as rigid as a mollusk shell.
Apex.— T\iQ apical end of a conulariid may be: 1, a narrow, blunt
point (Fig. 1.1); 2, truncated (Fig. 7.6); or 3, truncated, but having the
end covered by a smooth, convex, imperforate apical wall (Fig. 1.2).
The apex of a conulariid has been interpreted as a sharp point, as a
bluntly rounded structure, as a smooth, imperforate wall, or as a smooth
wall with a centrally located hole. Since the work of Kiderlen (1937),
conulariids have been thought of as metazoans having a sharp point
in the juvenile state. Presumably, the point was attached by an attach-
ment disc to a hard substratum. Support for Kiderlen’s argument was
provided by supposed conulariids which were previously described by
Ruedemann (1896i2, 1896/?). The specimens described by Ruedemann
seem to be tubes of Sphenothallus (Feldmann et al., 1986). Sphenoth-
alius has recently been shown to be unrelated to conulariids (Mason
and Yochelson, 1985).
Apices of conulariids are exceedingly rare. The apices are not pointed
as once interpreted, but are slightly rounded (Figs. 7.6, 16.6, 33. l-
33.2). During life, most or all conulariids were attached by an elongate
stalk (Figs. 1.1, 24.1-24.2, 32.5) to some substratum, during at least
part of the life cycle. The apex was sheathed by the proximal portion
of the stalk. Breakage at the proximal end of the stalk may possibly
explain why so few conulariids are observed that have their apices
intact.
Authentic conulariids with attachment discs have never been de-
scribed. Small, round, black, and presumably chitinous or chitino-
phosphatic, bodies attached to bryozoans or brachiopods have been
identified in various museum collections as conulariid attachment discs.
Often, the presumed base of a tube is preserved connected to such a
structure. Such tubes are circular or subcircular in cross section. These
problematic fossils probably represent attachment devices of some type
of organism, but a relationship to conulariids has not been demon-
strated.
Smooth, imperforate apical walls have been noted by many authors
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VOL. 55
(Miller in Sowerby, 1821; Hall, 1876, 1879; Slater, 1907; Richter and
Richter, 1930; Sinclair, 1948; Moore and Harrington, \956b\ Babcock
and Feldmann, 1984, 1986, herein. Figs. 4.1, 10.4, 11.2, 14.1, 14.4,
1 5.4, 19.1). These structures have been termed septa, apical diaphragms
or Schotten. None of these terms seems appropriate, either because of
its genetic implications or because of an incomplete description of the
morphology. Apical walls are single units of integument that cover the
convergent end of a conulariid close to the apical terminus. An apical
wall is not located at the apical terminus itself, but seems to be attached
to the interior of the faces on the exoskeleton, slightly adaperturally of
this region. The line of juncture of the apical wall with the remainder
of the integument always seems to occur near the narrowest portion of
an exoskeletal constriction. Apical walls appear to lack support from
rods or other structures, and may be bowed slightly in the adapical
direction. Multiple apical walls may be present in single individuals
(Eichwald, 1860; Steinmann and Doderlein, 1890; Slater, 1907; Sin-
clair, 1948).
References to conulariids with centrally perforated apical walls in-
clude Slater (1907), Richardson (1942), and Swartz and Richardson
(1945). A collapsed specimen of Conularia congregata exhibiting a
subcircular structure located centrally on the apical wall is illustrated
in Figure 4.1. This subcircular structure is interpreted as having resulted
from the compression of a thin, flexible apical wall against the apical
portion of some soft-part morphologic feature such as the alimentary
tract.
The function of the apical wall may have been to seal off the portion
of the central cavity in which the conulariid lived from the stalk and
older, unused portions of the body. In specimens preserving apical
walls, the most apicad portions of exoskeleton are not smooth (Figs.
1 1.2, 14.1). This indicates that conulariid exoskeletons like these may
have been tom from their stalks by current forces. It is also possible
that some few conulariids periodically may have shed unused portions
of the exoskeleton (Babcock and Feldmann, 1984).
Apical angle. — An apical angle is measured at the intersection of two
lines projected by tracing tangent lines along the trend of two adjacent
comer grooves (Fig. 1.2). Apical angles are typically in the range of 8°
to 26°.
The apical angle, as measured on a large segment of a specimen, may
differ, by several degrees, from the apical angle as measured on a small
section of the skeleton (see Appendix B in Part B). Small segments,
particularly at exoskeletal constrictions and near the apex, usually yield
somewhat larger apical angles than generalized apical angles, measured
over a large segment of an exoskeleton.
A difference in the acute apical angle between adjacent sides of a
1986
Babcock and Feldmann— North American Conulariida
359
conulariid, mostly attributed to compression, has been noted by nu-
merous authors, including Barrande (1867), Hall (1879), Slater (1907),
Boucek (1939), and Sinclair (1948). Studies of compressed and pre-
sumably uncompressed materials indicate that opposite sides of a con-
ulariid exoskeleton are paired (Babcock and Feldmann, 1984, 1986).
In cross section, a conulariid is typically rectangular, if only slightly so
(Figs. 30.4, 33.4). Each face subtends an apical angle equal to that of
the face opposite it, but different from either adjacent face. This suggests
that conulariids are bilaterally symmetrical, rather than tetramerally
symmetrical, metazoans (Babcock and Feldmann, 1984, 1986). Rhom-
boid-shaped conulariids may exist, but most forms that are thought to
have a rhomboidal cross section probably were described from subtly
compressed specimens.
Faces. —In most cases, the four faces of the conulariid skeleton are
essentially planar (Fig. 1.1). At least two forms, Mesoconularia ca~
huanotensis, from the Devonian of Bolivia (Branisa and Vanek, 1973)
and M. solitaria, from the Silurian of Czechoslovakia (Sinclair, 1 948),
have faces that are markedly curved. In Anaconularia anomala from
the Ordovician of Czechoslovakia, Kiderlen (1937) described a clock-
wise torsion of the exoskeleton, up to 40°. However, examination of
eleven specimens referable to this taxon (GSC 85063-85073) indicates
that these fossils, preserved in quartzite, are not twisted or compressed
in a uniform fashion. Therefore, the “torsion” which Kiderlen observed
may, in fact, be related to post-mortem diagenetic effects.
The two faces on a conulariid exoskeleton which subtend apical
angles equal to each other, but smaller than the remaining two faces,
are termed minor faces. Those two faces having larger apical angles
are termed major faces.
Integument.— T\iQ thin walls, or faces, of the exoskeleton are made
up of a multilayered calcium phosphate and protein integument (Ed
Landing, personal communication, 1984; based upon electron micro-
probe analyses of specimens of Paraconularia byblis and P. subulata
from locality 1 90). The precise number of layers and the extent to
which this number is consistent from species to species has yet to be
determined. In one example of Conularia desiderata, analyzed under
the scanning electron microscope, at least thirty very thin, but discrete,
layers of calcium phosphate were observed (Fig. 3.9). This stands in
marked contrast to previous interpretations of the histology of the
conulariid integument (for example, Sinclair, 1 940Z?, 1 948; Richardson,
1 942), in which only two or three layers were observed through use of
standard light microscopy.
— Rods (Fig. 2.2), embedded in the integument, are support
structures which cross each face transversely; they are composed of
calcium phosphate and are subcircular in cross section.
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Annals of Carnegie Museum
VOL. 55
1986
Babcock and Feldmann— North American Conulariida
361
Each rod crosses one half of each face transversely until its distal
end abuts with, or alternates with, the distal end of an adjacent rod.
Proximally, rods of adjacent faces articulate in a comer groove. When
covered by integument, a rod forms a thin ridge which traverses one
half of each face of a conulariid skeleton.
A rod may be equipped with numerous spines (Fig. 2.2) which project
adaperturally or adaperturally and adapically. When both types of spines
are present, adapertural spines seem to be longer than adapical spines.
The function of a spine was probably to provide a stronger framework
to support the integument (Figs. 2.2, 3.8). Most conulariid species
possess rods which have spines. Some, however, such as Paraconularia
planicostata (for example. Fig. 3.1) and P. subulata (for example. Figs.
3.2, 3.6), have integument supported by rods alone.
Rods may or may not possess small nodes on the external side of
the exoskeleton (Fig. 2.2). Nodes, if present, are arranged in a single
row along a rod. These structures occur slightly adapically of each
adapertural spine in all of the taxa reported herein (for example, Fig.
3.7). Thus, the number of adapertural spines and the number of nodes
are equal. In Paraconularia sorrocula, the nodes seem to be fused with
the adapertural spines, forming a single structure (Fig. 28.2).
The manner of rod articulation has long been used as a diagnostic
character at the species level (Hall, 1859; Barrande, 1867; Holm, 1893).
To delineate a ‘‘natural grouping” of the conulariids. Holm (1893)
identified four species groups, based partially upon ridge characteristics.
Richardson (1942) identified four modes of ridge arching among or-
ganisms which were then considered conulariids, presumably including
Fig. 3. — 3. 1-3.4; rod articulation styles. 3.1; USNM 33785, Paraconularia planicostata
(Dawson) showing gothic arch style; locality 165. 3.2; NYSM 3491, P. subulata (Hall)
showing inflected circular curve style; locality 203. 3.3; AMNH 33018, Conularia pyr-
amidalis Hall showing inflected gothic arch style; locality 1 17. 3.4; AMNH CU 282G,
C. elegantula Meek showing angulated circular curve style; locality 174. 3.5-3.10; scan-
ning electron micrographs; arrows point in apertural direction. 3.5; USNM 395833, P.
subulata (Hall); view along midline showing integument draped loosely over rods; locality
190;. 3.6; USNM 395834, P. subulata (Hall), rods at midline, integument lacking; locality
72. 3.7; USNM 395830, P. byblis (White); integument closely draped over rods; locality
190. 3.8; USNM 395832, C desiderata Hall, view of ridges, interridge crests and inter-
ridge furrows; rods have been broken away; locality 135. 3.9; USNM 395832, same
specimen as in Fig. 3.8, C. desiderata Hall, ridge with rod removed, showing multilayered
integument; locality 135. 3.10; USNM 395832, same specimen as in Fig. 3.8, C desi-
derata Hall, view showing ridges with rods broken away, interridge crests and interridge
furrows; locality 135. Bar scales equal 1 mm for Figs. 3. 1-3.4 and 0.1 mm for Figs. 3.5-
3.10.
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Annals of Carnegie Museum
VOL, 55
Fig. 4. — Conularia congregata Hall. 4.1; NYSM 3483, several paralectotypes from slab
exhibiting lectotype and twelve paralectotypes, preserved in black shale. Note that shells
1986
Babcock and Feldmann— North American Conulariida
363
Sphenothallus, now considered a worm (Mason and Yochelson, 1985;
Feldmann et al., 1986). Babcock and Feldmann (1986) recognized
four modes of rod articulation which produce patterns of ridges useful
as species-level taxonomic criteria. Their terminology for rod articu-
lation patterns is followed herein; examples of each style are given in
Figs. 3. 1-3.4.
One additional feature of rod articulation at the midline is of interest
for species-level determination of taxa. The two rods on a face which
meet, or approach closely, at the midline are termed a rod pair. If the
right rod of a rod pair is closest to the aperture when the specimen is
examined with the aperture upward, the pattern of articulation is right
superior (for example. Figs. 3.1, 3.2, 3.6). Conversely, if the left rod
of a rod pair is closest to the aperture, the pattern is termed left superior
(for example. Fig. 3.5). Some species can be distinguished from others,
in part, by the relative proportions of right superior to left superior
and to abutting rods on the major and minor faces.
An angle subtended by lines connecting the distal point of a rod with
the point at which it joins the comer angle and a line projected across
a face, perpendicular to the trend of the comer angle, from the point
where a rod joins the comer angle, is termed a rod angle (Fig. 1 .2). The
description of a rod angle, combined with information on the style of
rod articulation, is used herein as an effective way to help distinguish
conulariid species. Rod angles generally vary across an exoskeleton
within a few degrees. Notable changes in rod angles are often observed
at the narrowest points of exoskeletal constrictions.
A complex of structures is produced on the external surface of the
exoskeleton when the integument is draped over a framework of rods
and spines. When integument covers a spine, an interridge crest is
formed; when integument is draped between two adjacent spines, an
interridge furrow is produced. A specimen preserved with the integu-
ment draped loosely over the rod and spine framework may appear to
be of a different species than a specimen having integument diagenet-
ically compressed close to the framework. It may also appear to be
different from a specimen which lacks the integument altogether.
Midline.— A line, of variable distinctness, runs longitudinally down
the middle of each face (Fig. 1.1). This line, termed the midline, may
be a raised structure or a groove, and seems to be pigmented in some
of the inarticulate brachiopod Discina humilis Hall are attached to the conulariid exo-
skeletons; locality 154. 4.2; NYSM 3484, detail of ridge structure as preserved in a
counterpart specimen. 4.3; NYSM 3484, same specimen as in Fig. 4.2; locality 154. 4.4;
NYSM 3483; lectotype and three paralectotypes exposed on same slab as specimens in
Fig. 4.1; lectotype is indicated by an arrow. Bar scales represent 1 cm.
364
Annals of Carnegie Museum
VOL. 55
cases (Fig. 23.5). Wiman (1895), Knod (1908), and Liu (1981) have
published hand-drawn figures of three separate structures radiating
inward from the midlines of conulariids. Wiman (1895) described large
T- or Y-shaped ‘‘septa,” Liu (1981) figured elongate “septa” and Knod
(1908) figured localized thickenings of integument. Among specimens
examined from Devonian and Mississippian rocks of North America,
there seems to be no evidence of an elongate, inwardly-directed struc-
ture associated with the midline. However, a thickening of the midline,
on the internal side of the exoskeleton, has been observed on specimens
of Paraconularia ulrichana from the Devonian of Bolivia. In some
cases, there appears to be an invagination of the exoskeleton from the
interior at the midline (Fig. 14.2).
Corner groove. — A comer groove (Fig. 1 . 1) is the facial invagination
at a comer of the exoskeleton where two faces meet at right angles. An
individual rod articulates proximally with an adjacent rod in a comer
groove; rods on adjacent faces alternate in position along the comer
groove. No evidence of localized thickenings of integument interior to
the comer grooves, as described by Knod (1908), has been observed.
The points of articulation of the rods in the comer grooves seem to be
nodose swellings (Fig. 20.4).
Exoskeletal constriction. ■— An exoskeletal constriction (Fig. 1.1) is a
slight depression in the conulariid exoskeleton which is identifiable at
the same relative position on all faces. An exoskeletal constriction
appears to be continuous about the four faces of the exoskeleton (for
example. Figs. 5.3, 9. 3-9. 5, 10.1, 27.1-27.3). Trends of rod angles may
change slightly (Figs. 8.3, 8.5), rods may converge (Figs. 8.5, 1 1.3) or
an apical wall may be attached internally to the exoskeleton at the
narrow end of such a structure (Figs. 1 1.2, 14.1). Exoskeletal constric-
tions may be indications that conulariids grew by the incremental ad-
dition of new integument and rods at the aperture. Because the apertural
constriction, the last formed of the exoskeletal constrictions, is always
located slightly adapically of the aperture (for example. Fig. 10.1), it
is presumed that growth temporarily ceased near the widest portion of
the exoskeleton, located between two adjacent exoskeletal constric-
tions.
"" Septa.'' —ThQ term “septum” has been applied to three separate
morphologic features in conulariids: 1 , apical walls (for example, Miller
in Sowerby, 1821; Slater, 1907); 2, ridges (Slater, 1907); and 3, T- or
Y-shaped structures or unmodified elongate structures radiating inward
from the faces at their midlines (Wiman, 1895; Kiderlen, 1897; Liu,
1981). None of these seems to be an appropriate application of the
term. Most commonly, the word “septum” as applied to conulariids
means large T- or Y-shaped structures of the exoskeleton that project
inward from the midlines. These supposed hard-part structures were
described in Conularia loculata, from the Silurian of Sweden by Wiman
1986
Babcock and Feldmann— North American Conulariida
365
(1895) and were thought by Kiderlen (1 937) to be homologous to septa
composed of endodermal tissue in living scyphomedusans. These struc-
tures have not been observed, at least not to such a marked extent, in
any specimens other than those of Wiman. Wiman’s material was
illustrated only by drawings, and the specimens are now lost (W. A.
Oliver, Jr., personal communication). Thus, Wiman’s observations
cannot be replicated.
The cross sectional view of a specimen of Pamconularia subulata
illustrated in Fig. 33.4 exhibits a pattern of limonitic staining in the
central cavity which roughly approximates Wiman’s figures. The stain-
ing in this example is probably related to the preservation of incom-
pletely decomposed internal viscera. It is suspected that Wiman and
others may have been misled by some taphonomic feature such as this.
Soft-part morphology . — Much, speculation has surrounded the study
of the soft-parts of conulariids. Since the work of Kiderlen (1937),
Knight (1937), Moore and Harrington (1956^z, \956b) and Werner
(1966, 1967, 1969), conulariids have been interpreted as tentacled
creatures. Support for such interpretation is weak, being based on a
presumed homology of conulariids to medusoid cnidarians or upon a
grouping of true medusoid cnidarians with the conulariids (Kiderlen,
1937; Knight, 1937; Moore and Harrington, 1956<2, \956b).
Remains of presumed conulariid soft-parts were independently de-
scribed from European Devonian conulariids by Steul (1984) and from
North American Mississippian conulariids by Babcock (1985a) and
Babcock and Feldmann (1986). Babcock and Feldmann (1986), work-
ing only with exceptionally preserved three-dimensional specimens,
identified a single elongate tube that extends the length of the central
cavity and a large globular shaped structure near the aperture (Fig. 2. 1).
Steul’s (1984) work, based upon x-ray analyses of collapsed specimens
preserved in the Hunsruck Slate, revealed other structures which may
be preserved soft-parts, though the evidence is ambiguous.
The tubular and globular internal structures (Figs. 30.2=30.6), pre-
sumably representing remains of organ systems, appear to be reduced
in size compared to expected living organs and show no details of soft-
part anatomy. These structures may be somewhat contracted masses
of internal tissues. These structures may have been preserved, in outline
at least, as altered remains of partially digested food matter and/or
sediment left in the intestinal tract when the animals died. Other organs
that were originally present in the North American studied specimens
may have decayed.
Occurrences and Paleoecology
Conulariids have been reported exclusively from marine rocks rang-
ing in age from the Late Precambrian to the Recent (Caster, 1957).
VOL. 55
Fig. S. — Conularia desiderata Hall. 5.1; AMNH 2697, holotype; major face, preserved
in limestone; locality 121. 5.2; NYSM 3487, holotype of C continens var. rudis Hall,
1986
Babcock and Feldmann— North American Conulariida
367
This range, however, includes a variety of taxa now referable to other
groups. Occurrences here considered valid include specimens from
Lower Ordovician (Sinclair, 1948) through Upper Triassic (Gou and
Yang, 1985) rocks. While articulated conulariids are uncommon or
rare in many instances, conulariids are, nonetheless, pervasive faunal
elements in Middle Ordovician through Permian marine rocks. They
are rare in the Lower Ordovician and in the Triassic. Some Paleozoic
occurrences yield abundant conulariids.
Fossils identified as conulariids have been identified from all con-
tinents. However, the sole report of a conulariid from Antarctica (Cor-
dini, 1955) has been received with skepticism and may represent plant
material (Dalziel et aL, 1981).
The conulariid skeleton, or exoskeleton, is composed of a calcium
phosphate framework, made of rods, and usually having spines or
nodes. The framework is set in an interlayered integument made of
thin sheets of calcium phosphate and protein. Overall, the exoskeleton
was probably fairly delicate, and upon death of the animal, was readily
subject to collapse, occasional attachment of epibionts and decom-
position accompanied by disarticulation (Feldmann and Babcock, 1986).
Most occurrences of articulated conulariids involve rapid burial and
often, early diagenesis. For example, conulariids are abundant in the
Meadville and Wooster members of the Cuyahoga Formation in north-
eastern and central Ohio. Specimens in these units are usually found
in presumed tempestite beds or in siderite concretions. Specimens col-
lected from tempestite beds often seem to be current aligned (Fig. 30.7).
Specimens collected from siderite concretions, such as that illustrated
in Fig. 10.4, are weakly current aligned, if at all. In the Cuyahoga
Formation, concretions were probably produced through localized in-
creases in pH and lowerings of eH, causing precipitation of iron car-
bonate. The onset of siderite precipitation probably occurred soon after
burial of the animals.
Some occurrences of articulated conulariids involve “prefossilized”
specimens, or ones which have undergone early diagenesis, and which
have later been exhumed through bioturbation, winnowing of sedi-
ments by currents, or both. The specimens were later deposited in beds
major face, preserved in siltstone; locality 132. 5.3; NYSM 3487, same specimen as in
Fig. 5.2, comer view. 5.4; AMNH 2697, same specimen as in Fig. 5.1, minor face. 5.5;
NYSM 3487, detail of minor face. 5.6; NYSM unnumbered, detail of specimen with
integument draped loosely over rods; locality 124. 5.7; NYSM 3485, syntype of C.
continens Hall, preserved in black shale. 5.8; NYSM 3486, syntype of C. continens Hall;
detail of minor face; locality 125. 5.9; NYSM 3486, same specimen as in Fig. 5,8, nearly
complete, flattened specimen. Bar scales represent 1 cm.
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VOL. 55
Fig. 6. — 6. 1-6.3; Conularia delphiensis (Maroney and Orr) preserved in phosphatic con-
cretions at a stratigraphic discontinuity surface (Maroney and Orr, 1974). 6.1; lUPC
1986
Babcock and Feldmann— North American Conulariida
369
representing stratigraphic discontinuities (Maroney and Orr, 1974;
Baird, 1981; Baird and Brett, 1981; herein. Figs. 6. 1-6.3).
The general scarcity of articulated conulariid remains, at most lo-
calities, may be related to rapid disarticulation of the multielement
skeleton, predation and scavenging, and the constancy of water move-
ment near the sediment-water interface. The frequent occurrence of
articulated conulariids in low diversity faunas (Ruedemann, 1934; Jux,
1960; Babcock and Feldmann, 1986; Feldmann and Babcock, 1986)
may be largely a function of low biotic activity in these environments.
Many conulariids show indications that some breakdown of the exo-
skeleton has occurred, especially in the vicinity of the aperture (for
example. Figs. 5.7, 7.5, 7.7), where the integument may be greatly
reduced in thickness or lacking. In rare instances, poorly bioturbated
stratigraphic units yield specimens of conulariids in which the integ-
ument is lacking over much of the exoskeleton and disarticulated rods
have been displaced from their original positions (Fig. 21.4). In sedi-
ments that were probably well bioturbated, or sediments where water
movement took place rather constantly, articulated conulariids are rare.
It is possible that conulariid rods will be found in some Paleozoic and
Mesozoic rock units that are sampled for microfossils by insoluble
residue or other techniques. Disarticulated conulariid rods have been
occasionally misidentified as fossil fish bones (Feldmann and Babcock,
1986).
Many species of conulariids seem to have been geographically wide-
spread. For example, the Mississippian species, Paraconularia ches-
terensis, has a known geographic range from Alabama to British Co-
lumbia and, as with numerous other species, occurs in rocks of various
lithologies. These lithologies include mudstones and wackestones
(Chester Group of Illinois), gray shales (Borden Group of Indiana) and
siderite concretions (Cuyahoga Formation of Ohio). The occurrence of
this and other conulariid species in stratigraphic units of so vastly
dissimilar lithologies and environments of deposition, and their wide
geographic distribution, suggests that some species may have been
planktonic or pseudoplanktonic at some point in the life cycle. The
bilaterally symmetrical body plan (Babcock and Feldmann, 1984, 1986;
14470-1, holotype; view of flattened specimen and detail of minor face; locality 23. 6.2;
lUPC 14470-2, paratype; locality 23. 6.3; lUPC 14470-1, same specimen as in Fig. 6.1,
detail of ?minor face. 6.4-6. 5; C. desiderata Hall. 6.4; NYSM unnumbered, view of
?majorface; locality 132. 6.5; USNM 395832, detail of specimen with most of integument
removed, exposing broken rods and intact spines; locality 135. Bar scales in Figs. 6.1-
6.4 represent 1 cm; bar scale in Fig. 6.5 represents 5 mm.
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Annals of Carnegie Museum
VOL. 55
Steul, 1984), however, may be an indication that some species were
weakly nektonic.
Conulariids are usually referred to as solitary animals. Clusters are
rare, but a few have been figured by Slater (1907, Plate 2, fig. 1), Hall
(1876, Plate 28, fig. 1; 1879 Plate 24, fig. 1), Sinclair (1944, Plate 2,
fig. 5) and Babcock and Feldmann( 1984, p. 17; Babcock and Feldmann,
1986, figs. 4A, 4C). In all these clusters, individual specimens are
shown radiating about a central area. Apices are usually pointing inward
in these aggregations (for example. Fig. 24.3). Occasionally, apices of
such specimens are preserved intact (Fig. 24.1), yet in portions of a
somewhat disaggregated cluster illustrated by Hall (1876, 1879), here
illustrated in Figs. 4. 1 and 4.4, the apices are missing, perhaps because
of post-mortem decay. The apical walls are visible in some of these
specimens.
A slab exhibiting numerous P. chesterensis from the Borden Group
(Mississippian) of Indiana exhibits several small, black to reddish brown,
tubular structures, circular in cross section, attached to, or very near
the apices of, conulariids (Figs. 24. 1, 24.3). Combined with discoveries
of similar structures on conulariids from the Borden Formation (Mis-
sissippian) of Kentucky (Fig. 24.2) and from the Etherington Formation
(Mississippian) of British Columbia (Fig. 32.5), these structures are
interpreted as attachment stalks. The association of conulariids with
plant remains, perhaps algae (for example. Fig. 28.7), indicates that
some conulariids may have been pseudoplanktonic and were attached
to, or entwined with, planktonic algae. Alternatively, the conulariids
may have attached to plant remains which had previously settled to
the ocean floor.
Cluster associations involving numerous specimens, usually com-
prising single species, serve to indicate that some, if not all, conulariids
were gregarious, at least during some part of the life cycle. No evidence
of budding or any other asexual reproductive style exists.
Regardless of whether the conulariids were planktonic, pseudoplank-
tonic, benthonic, or even nektonic, attached or free swimming, the soft-
part organs in the region of the aperture of the exoskeleton probably
functioned for filter feeding. There is no evidence for aggressive food
gathering behavior.
The style of growth in conulariids was incremental, with addition of
rods and spines taking place at the aperture. Growth lines, such as those
seen on mollusks and brachiopods are not present. Exoskeletal con-
strictions, in exactly the same relative positions on all four faces, are
possible evidences of incremental growth. Additional evidence of in-
cremental growth at the aperture comes from some specimens showing
healed injuries (for example. Figs. 20.1, 29.5). Along the exoskeletal
1986
Babcock and Feldmann— North American Conulariida
371
constrictions of such examples, rods may be broken or the rod artic-
ulation patterns may be disrupted. This indicates two things: 1, that
the positions of apertural terminations changed and correspond to the
positions of apertural constrictions; and 2, that the apertural termi-
nations were easily fragmented but healing could occur during a suc-
ceeding growth phase.
Development of multiple apical walls may also have been a function
of an incremental growth pattern. When the internal cavity of an in-
dividual reached a certain volume, an apical wall was probably secreted
adaperturally of the apical terminus. Apical walls are commonly pre-
served among specimens referable to the genus Conularia but are rarely
found on specimens of Paraconularia. No specimens of Reticulacon-
ularia have been observed with apical walls in place.
Epibionts on conulariids include orbiculoid brachiopods (Hall, 1876,
1879; Moore and Harrington, 1956a; Babcock and Feldmann, 1984,
1986; herein, Figs. 4.1, 4.4, 5.9, 13.1, 13.5, 16.4, 28.1-28.2), en-
crusting bryozoans (Finks, 1955; herein. Figs. 22.4-22.5) and edrioas-
teroid echinoderms (Barrande, 1867; Moore and Harrington, 1956a).
Attachment of epizoans has been interpreted as either having occurred
during the life of the conulariid (Finks, 1955; Moore and Harrington,
1956a) or as having occurred after the death of the conulariid (Baird,
1981; Babcock and Feldmann, 1986). In all the specimens examined
in the course of this study, orbiculoids seem to have attached to the
conulariids after they died and came to rest on the sediment surface.
This is supported by the observation that orbiculoids are always found
attached to only two faces of a conulariid (Figs. 4.1, 4.4). Presumably,
these two faces were the only ones which projected above the sediment-
water interface. A specimen is illustrated herein (Fig. 25.5) of a bryo-
zoan encrusting a conulariid in the region of a comer groove. There is
no sign of damage to the bryozoan on this specimen, indicating that
encrustation by the bryozoan occurred after the death of the conulariid.
Systematic Paleontology
Summary of taxa.--A total of 69 trivial names have been applied
to Devonian or Mississippian conulariids of North America prior to
this paper. Of these, 54 taxa were published, and eight were described
in unpublished manuscripts. Herein and in Part B, 28 species are rec-
ognized as valid, of which five are new. However, primary type spec-
imens of 1 8 species were not available for study; therefore, the status
of these taxa was not assessed fully. In Part B, one species is removed
from the Conulariida. The conulariids described below and in Part B
are divided among three genera, Conularia, Paraconularia and Retie-
ulaconularia, n. gen. The genus Diconularia Sinclair, 1952 is regarded
372
Annals of Carnegie Museum
VOL. 55
as a junior synonym of Conularia Miller in Sowerby, 1821 and the
genus Adesmoconularia Driscoll, 1963 is considered to be a junior
synonym of Paraconularia Sinclair, 1 940a.
The list below summarizes all described species-level taxa of con-
ulariids from North American Devonian and Mississippian rocks and
indicates the status of each trivial name as currently recognized. Taxa
are arranged alphabetically according to their presently recognized sta-
tus. Junior synonyms and previously used combinations of each taxon
are listed below each valid name.
Conularia congregata Hall, 1876
Conularia delphiensis (Maroney and Orr, 1974)
Ctenoconularia delphiensis Maroney and Orr, 1974
Conularia desiderata Hall, 1861
Conularia continens Hall, 1876
Conularia continens var. rudis Hall, 1879
Conularia elegantula Meek, 1871
Conularia milwaukeensis Cleland, 1911
Conularia congregata var. milwaukeensis Cleland, 1911
Conularia multicostata Meek and Worthen, 1865
Conularia micronema Meek, 1871
Diconularia micronema (Meek, 1871)
Mesoconularia mcfarlani Sinclair, [1948] MS
Mesoconularia attica Sinclair, [1948] MS
Conularia pyramidalis Hall, 1859
Conularia huntiana Hall, 1859
Conularia lata Hall, 1859
Conularia subcarbonaria Meek and Worthen, 1865
Conularia intertexta Miller, 1894
Conularia spergenensis Miller and Gurley, 1893
Conularia tuzoi Clarke, 1907
Conularia desiderata var. tuzoi Clarke, 1 907
Conularia ulsterensis Howell, 1 942
Conularia undulata Conrad, 1 84 1
Conularia cayuga Hall, 1876
Conularia crebistria Hall, 1876
Paraconularia alternistriata (Shimer, 1926)
Conularia alternistriata Shimer, 1926
Paraconularia alpenensis Babcock and Feldmann, n. sp.
Paraconularia blairi (Miller and Gurley, 1893)
Conularia blairi Miller and Gurley, 1893
Conularia sedaliensis Miller and Gurley, 1896
Paraconularia indiana Sinclair, [1948] MS
Paraconularia byblis (White, 1862)
Adesmoconularia byblis (White), 1862
Conularia byblis V^hiXe, 1862
Paraconularia chagrinensis Babcock and Feldmann, n. sp.
Paraconularia chesterensis (Worthen, 1883)
Conularia chesterensis Worthen, 1883
Paraconularia missouriensis (Swallow, 1 860)
Conularia missouriensis Sw2l\\ov>/ , 1860
1986
Babcock and Feldmann= North American Conulariida
373
Conularia gratiosa Miller and Gurley, 1893
Conularia greenei Miller and Gurley, 1 896
Paraconularia sciotoviliemis Driscoll, 1963
Paraconularia okiahomaemis Babcock and Feldmann, n. sp,
Paraconularia planicostata (Dawson, 1868)
Conularia planicostata Dawson, 1868
Paraconularia recurvatus Babcock and Feldmann, n. sp.
Paraconularia salinensis (Whiteaves, 1891)
Conularia salinensis Whiteaves, 1891
Paraconularia sorrocula (Beede, 1911)
Conularia sorrocula Beede, 1911
Paraconularia subulata (Hall, 1858)
Conularia subulata 1858
Conularia victa White, 1862
Conularia Winchell, 1865
Conularia whitei Meek and Worthen, 1865
Conularia sampsoniMillQT , 1892
Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Paraconularia yochelsoni Babcock and Feldmann, n. sp.
Reticuiaconularia penouili (Clarke, 1 907)
Conularia penouili Clarke, 1 907
Conularia gaspesia Sinclair, 1 942
Reticuiaconularia sussexensis (Herpers, 1 949)
Conularia sussexensis Herpers, 1 949
Whereabouts of Type Material Unknown
Climacoconus viata Swartz and Richardson in Richardson, [1942] MS
Conularia crawfordsvillensis Owen, 1862
Conularia gracilis WTnck, 1888
(Name preoccupied by C. gracile Hall, 1847; changed to C. herricki by Miller, 1892.)
Conularia grandis Roemer, 1856
Conularia herricki Miller, 1892
Conularia jervisensis Shimer, 1 905
Conularia latoides Swartz and Richardson in Richardson, [1942] MS
Conularia marionensis Swallow, 1860
Conularia missouriensis var. hermansi Calvin, 1890
Conularia molaris White, 1876
Conularia novascotica Hartt, in Dawson, 1868
Conularia osagensis Swallow, 1863
Conularia pyramidalis var. parvinodis Swartz and Richardson in Richardson, [1942] MS
Conularia siphunculophora Swartz and Richardson in Richardson, [1942] MS
Conularia triplicata Swallow, 1860
Conularia verneuiiia Emmons, 1 846
Paraconularia welleri Sinclair, [1948] MS
Non-Conulariid
Conularia tenuicostata Branson, 1938
(Assigned, tentatively, to phylum Priapulida.)
Repositories. — Specimens are listed according to catalogue numbers
with the repositories abbreviated as follows:
374
Annals of Carnegie Museum
VOL. 55
AMNH
AMNH CU
BMS
CM
CMNH
FMNH P
FMNH PE
FMNH UC
GSC
ISGS
lUPC
KSU
NJSM
NYSM
oc
PU
RM(MU)
UCGM
UIPC
UK
UMC
UMMP
USNM
wvu
American Museum of Natural History, New York, New York
American Museum of Natural History, Columbia University Collection,
New York, New York
Buffalo Museum of Science, Buffalo, New York
Carnegie Museum of Natural History, Invertebrate Paleontology Collec-
tions, Pittsburgh, Pennsylvania
Cleveland Museum of Natural History, Invertebrate Paleontology Col-
lections, Cleveland, Ohio
Field Museum of Natural History, Paleontology Collection, Chicago, Il-
linois
Field Museum of Natural History, Invertebrate Paleontology Collection,
Chicago, Illinois
Field Museum of Natural History, University of Chicago Collection from
Walker Museum, Chicago, Illinois
Geological Survey of Canada, Ottawa, Ontario
Illinois State Geological Survey, Illinois State Museum, Champaign-Ur-
bana, Illinois
Indiana University Paleontological Collection, Bloomington, Indiana
Department of Geology, Kent State University, Kent, Ohio
New Jersey State Museum, Trenton, New Jersey
New York State Museum and Science Service, Albany, New York
Oberlin College Paleontological Collections, Oberlin, Ohio
Department of Geology, Princeton University, Princeton, New Jersey
Redpath Museum (McGill University), Montreal, Quebec
University of Cincinnati Geological Museum, Cincinnati, Ohio
University of Illinois, Department of Geology, Champaign-Urbana, Il-
linois
University of Kentucky, Department of Geology, Lexington, Kentucky
University of Missouri-Columbia, Columbia, Missouri
University of Michigan, Museum of Paleontology, Ann Arbor, Michigan
United States National Museum of Natural History, Washington, D.C.
West Virginia University, Department of Geology, Morgantown, West
Virginia
Treatment of manuscript names. — One of the most influential papers
regarding the systematics and morphology of conulariids is Sinclair
(1948), an unpublished Ph.D. thesis. In it were proposed many new
genus-level and species-level taxa. The undescribed genera identified
by Sinclair were subsequently published (Sinclair, 1952); however, the
majority of Sinclair’s new species have never been formally described.
Moreover, many of the species removed by Sinclair from Conularia,
as used in the sense of a form-genus, and placed in different genera,
have not been published in their revised state by Sinclair. Nevertheless,
various authors have used Sinclair’s combinations, often without ref-
erence to the authority for such usage.
It is our opinion that, because of the central importance of Sinclair’s
(1948) unpublished manuscript to the study of conulariid systematics,
Sinclair’s unpublished species-level names and his unpublished com-
binations should be included in the formal synonymies of the Devonian
and Mississippian taxa discussed below. For the sake of completeness,
1986
Babcock and Feldmann— North American Conulariida
375
we have also included in the present work the manuscript names of
Swartz and Richardson in Richardson (1942). By including these un-
published names in synonymy though, we do not intend to suggest that
these are available names. According to Article II(d)(ii) of the Inter-
national Code of Zoological Nomenclature, Third Edition (Interna-
tional Commission of Zoological Nomenclature, 1985), “a previously
unavailable name is not changed by its mere citation accompanied by
a reference to the work in which the name was published but was not
made available.” Also, in Article 1 1(e), the Code states that a, “name
first published as a junior synonym is not thereby made available unless
prior to 1961 it has been treated as an available name ...”
Phylum Conulariida Babcock and Feldmann, new phylum
—Animals generally possessing a four sided, steeply py-
ramidal exoskeleton; bilaterally symmetrical; integument composed of
calcium phosphate and protein, multilayered, moderately flexible; exo-
skeletai framework composed of calcium phosphatic rods arranged
transversely across each side face; adjacent rods abut or alternate at
midline of each face; rods of adjacent faces articulate in a groove at
junction of two faces; apical end closed either by a blunt point; one or
more smooth apical walls may be present internal to the exoskeleton
and aperturad the apex; apex sheathed by a ?chitinous, phosphatic, or
chitinophosphatic stalk; aperture simple and open. No internal hard-
part structures known; internal soft-parts comprise an elongate tube
extending most of the body length, in addition to one or more globular
shaped structures, all of uncertain function.
Remarks. — Approximately 40 genera of organisms have, at one time
or another, been grouped among the conulariids. Of these, six genera
have been excluded from the phylum to date. Based upon new infor-
mation on the architecture of the conulariid skeleton (Babcock and
Feldmann, 1986), it seems that even more genera have been de-
scribed than are warranted by fossil evidence. Upon further study,
several other genera are likely to be excluded from the phylum Con-
ulariida.
Organisms which are properly included within the phylum Conu-
lariida must possess a bilaterally symmetrical exoskeleton composed
of calcium phosphate rods and layered calcium phosphate and protein
integument. Herein, genera of conulariids are defined upon: 1, the
relative spacing of rods; 2, the relative proportion of rods that abut at
the midline to those that alternate; 3, the apical angles; 4, the presence
or absence of nodes and spines; and 5, the spacing of nodes and spines.
Species are distinguished using the characters upon which generic dis-
tinctions are made as well as the following: 1, the patterns of rod
articulation; and 2, the rod angles.
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Annals of Carnegie Museum
VOL. 55
Descriptions of conulariids represented in the Devonian and Mis-
sissippian rocks of North America follow. After examination of spec-
imens referred to the respective type species from Europe, Sinclair
(1940i2) proposed North American reference species. The following
diagnoses of Conularia and Paraconularia are based largely upon spec-
imens referable to the North American reference species. The new genus
Reticulaconularia is based upon specimens from the Devonian of east-
ern North America.
Specimens of Conularia are recognizable by their closely spaced rods,
by having more than 40% of the rods abutting at the midline, by having
small apical angles and by having both nodes and spines which are
closely spaced. Specimens referable to Paraconularia exhibit widely
spaced rods, fewer than 40% of rods abutting at the midline and small
apical angles. Nodes and spines may be present; if they are, they are
closely spaced. Conulariids referable to Reticulaconularia, n. gen. have
widely spaced rods which abut or alternate at the midline in proportions
which are not well established yet. They also have large apical angles.
Members of the genus Reticulaconularia are notable for the reticulate
appearances of the external surfaces of the faces (Figs. 33.1-33.5, 34. 1-
34.2, 34.4), a product of nodes and spines which are widely spaced. It
should be noted, however, that other conulariids, if preserved as ex-
ternal molds, may exhibit patterns similar to this '‘reticulate” appear-
ance (for example. Figs. 8. 5-8. 6). Such specimens are best studied with
latex casts.
Key to North American Devonian and Mississippian conulariids.—
The following is a key to all species of Devonian and Mississippian
conulariids from North America which are currently known. Some
categories are ambiguous due to the lack of complete or well preserved
specimens of some species, so it is not intended for use with specimens
of different ages or of other areas of the world.
1 . Faces have reticulate appearance 2
1 . Faces do not have reticulate appearance 3
2, Inflected gothic arch rod articulation present
Reticulaconularia sussexensis (Herpers)
2. Inflected gothic arch rod articulation not present
Reticulaconularia penouili (Clarke)
3. Number of rods that abut at midline less than 40% 4
3. Number of rods that abut at midline greater than or equal to 40% 19
4, Gothic arch rod articulation present ..................................... 5
4. Gothic arch rod articulation not present 9
5. Only gothic arch rod articulation present . . Paraconularia alternistriata (Shimer)
5. Gothic arch and another style of rod articulation present ................... 6
6. Gothic arch and inflected circular curve rod articulation present
Paraconularia oklahomaensis Babcock and Feldmann, n. sp.
6. Gothic arch and inflected gothic arch rod articulation present .............. 7
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Babcock and Feldmann— North American Conulariida
377
7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
18.
18.
19.
19.
20.
20.
21.
21.
22.
22.
23.
23.
24.
24.
25.
25.
26.
26.
27.
27.
Rods/cm fewer than or equal to 1 1 ....... Pamconularia planicostata (Dawson)
Rods/cm greater than 11
Paraconularia yochelsoni Babcock and Feldmann, n. sp.
Inflected gothic arch rod articulation present 9
Inflected gothic arch rod articulation not present 14
Only inflected gothic arch rod articulation present 10
Inflected gothic arch and another style of rod articulation present 11
Rods/cm fewer than 15
Paraconularia chagrinensis Babcock and Feldmann, n. sp.
Rods/cm greater than 15 Paraconularia sorrocula (Beede)
Rods/cm fewer than or equal to 1 2 12
Rods/cm greater than 12 13
Rods are slightly inflected at midline . . Paraconularia blairi (Miller and Gurley)
Rods are not inflected at midline Paraconularia subulata (Hall)
Number of rods that abut at midline fewer than 30%
Paraconularia salinensis (Whiteaves)
Number of rods that abut at midline greater than or equal to 30%
Paraconularia byblis (White)
Inflected circular curve rod articulation present; rods recurved near midline . . 15
Inflected circular curve rod articulation present; rods not recurved near midline
Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Rods/cm fewer than or equal to 1 2 17
Rods/cm greater than 12 16
Rods/cm fewer than 18 . . Paraconularia alpenensis Babcock and Feldmann, n. sp.
Rods/cm greater than or equal to 18
Paraconularia recurvatus Babcock and Feldmann, n. sp.
Rods/cm fewer than or equal to 7 Paraconularia chesterensis (Worthen)
Rods/cm greater than 7 Paraconularia missouriensis (Swallow)
Gothic arch rod articulation present Conularia pyramidalis (Hall)
Gothic arch rod articulation not present 19
Inflected gothic arch rod articulation present 20
Inflected gothic arch rod articulation not present 23
Rods/cm fewer than or equal to 30 21
Rods/cm greater than 30 22
Rods/cm fewer than or equal to 25 Conularia milwaukeensis Cleland
Rods/cm greater than 25 Conularia multicostata Meek and Worthen
Rods/cm fewer than or equal to 39 Conularia tuzoi Clarke
Rods/cm greater than 39 Conularia ulsterensis Howell
Angulated circular curve rod articulation present 24
Angulated circular curve rod articulation not present 27
Only angulated circular curve rod articulation present 25
Angulated circular curve and another style of rod articulation present ...... 26
Rods/cm fewer than 34 Conularia subcarbonaria Meek and Worthen
Rods/cm greater than 34 Conularia elegantula Meek
Rod angle less than or equal to 1 7° Conularia congregata Hall
Rod angle greater than 17° Conularia desiderata Hall
Rods undulose Conularia undulata Conrad
Rods not undulose Conularia delphiensis (Maroney and Orr)
Genus CONULARIA Miller, in Sowerby, 1821
Type species. — Conularia quadrisulcata Sowerby, by original desig-
nation; Silurian of England. Holotype is lost. North American reference
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Annals of Carnegie Museum
VOL. 55
spQciQS, fide Sinclair (1940a): Conularia niagarensis Hall, 1852 (Silu-
rian). Syntypes and plastosyntypes of C. niagarensis: AMNH 31625-
31628; plastosyntype of AMNH 31625: FMNH UC 60850.
Diagnosis. — Conulariids with rods that are generally closely spaced,
9-84 rods/cm. Fewer than 60% of rods alternate at midline; more than
40% abut; two adjacent rods on a face form a single arc across the face.
Apical angles small, 9-23°. Nodes, adapertural spines and adapical
spines usually present and closely spaced, 1-7/mm.
CONULARIA CONGREGATA Hall, 1876
Figs. 4. 1-4.4
Conularia congregata Hall, 1876, PI. 28, fig. 1; Miller, 1877, p. 141; Bigsby, 1878, p.
418; Hall, 1879, p. 214-215, PI. 34, fig. 1, PI. 34A, figs. 9-1 1; Miller, 1889, p. 390;
Harris, 1899, PI. 8, fig. 59; Clarke and Ruedemann, 1903, p. 565; Moore and
Harrington, 1956^, p. F61, fig. 46.2; Palmer and Brann, 1966, fig. 59; Babcock,
1985Z?, fig. 2; Babcock and Feldmann, 1986, fig. 4A.
Conularia s.l. congregata Hall. Sinclair, 1948, p. 286.
Conularia pyramidalis Hall. Sensu Babcock and Feldmann, 1984, p. 17.
Description. — ExosktlQion up to 11 cm in length. Major apical angle 17-18°; minor
apical angle 12-14°. Rod articulation exclusively of inflected circular curve style. Rods
almost always abut at midline; rod angle 9-13°. 16-21 rods/cm. 6-7 nodes/mm; 6-7
adapertural spines/mm; 6-7 adapical spines/mm; spines often not easily discernible.
Apical wall may be present.
Occurrences. Devonian of New York; localities 149-1 50 and
152-160. Maillieux (1933), Markovski and Nalivken (1934) and Xu
and Li (1979) reported C. congregata from Devonian rocks in Belgium,
the U.S.S.R. and the People’s Republic of China, respectively, but these
occurrences have not been confirmed.
Types. —LQcXoXypQ and twelve paralectotypes on one slab, NYSM
3483. Remains of at least thirteen specimens are preserved on James
Hall’s slab of syntypes (Figs. 4.1, 4.4). From this syntypic suite, the
best preserved of Hall’s figured specimens (Fig. 4.4) is here chosen as
the lectotype of C congregata. The remaining ten specimens are con-
sidered paralectotypes. Counterparts of the paralectotype specimens
shown in Fig. 4.1 are catalogued as FMNH unnumbered.
Remarks. — Conularia congregata Hall is most similar in morphology
to C. desiderata Hall. The similarities lie in overall size, apical angle
values, rod angle values and in the presence of inflected circular curve
rod articulation. The differences between the two taxa are subtle. In
specimens of C. congregata, few rods, generally fewer than 10%, al-
ternate at the midline; also, rods show very little or no inflection toward
the aperture near the midline. Among specimens referable to C. desi-
derata, as many as 15% of the rods may alternate at the midline;
specimens also show a strong adapertural inflection of the rods at the
1986
Babcock and Feldmann— North American Conulariida
379
midline. Some specimens, such as that illustrated in Fig. 5.6, appear
to be intermediate in morphology between C desiderata and C. con-
gregata.
Conularia congregata may also be confused with C pyramidalis Hall.
Conularia pyramidalis differs from both C congregata and C. desi-
derata is having inflected circular curve rod articulation in the apical
region and inflected gothic arch rod articulation elsewhere. There is no
evidence in C pyramidalis that the rods inflect near the midline.
Material examined.— AO specimens; housed in the AMNH, BMS,
FMNH, NYSM, USNM, and the private collection of Paul Zell.
CONULARIA DELPHIENSIS (Maroney and Orr, 1974)
Figs. 6. 1-6.3
Conularia sp. Kindle, 1901, p. 737, PL 123, fig. 8.
Ctenoconularia delphiensis Maroney and Orr, 1974, p. 3-6, fig. lA-F.
Description.— Exos\LQ\t\on up to 5.5 cm in length. Major apical angle and minor apical
angle about 1 5°. Rod articulation inflected gothic arch style in apical region and inflected
circular curve style elsewhere. Rods usually abut at midline; rod angle 10-11°. 26-42
rods/cm. 6-7 nodes/mm; adapertural spines and adapical spines not observed. Apical
wall not observed.
Occurrence. Devonian of Indiana; localities 22-23.
Fpppy. —Holotype, lUPC 14470-1; five paratypes, lUPC 14470-2
through 14470-6.
Remarks.— ThQ six specimens which comprise the type suite of C
delphiensis are badly preserved. Although no spines were actually ob-
served on any of the type specimens, all other morphologic features
are consistent with species of the genus Conularia as defined herein.
A specimen figured by Kindle (1901, pi. 123, fig. 8; USNM 62210),
an apparent external mold of C. delphiensis, clearly shows that nodes
are present, but spines are not evident. Spines may have been present
in this taxon, but have not been observed because of the poor pres-
ervation of the specimens studied. A cross sectional view of an un-
compressed specimen referable to this species has not been observed.
In their original description of C. delphiensis, Maroney and Orr
(1974) did not indicate why they chose to include the species in the
genus Ctenoconularia Sinclair, 1952. Moreover, they only compared
this species to Conularia congregata. Sinclair (1952, p. 141) noted that
the primary distinguishing characteristic of specimens referable to
Ctenoconularia was “strikingly slender shells.” This is certainly true
in the type species, Ctenoconularia obex Sinclair. Judging from Sin-
clair’s published figures (1952, figs. 56 A-C), the major and minor faces
subtend angles of 4° and 3®, respectively. In other respects, specimens
of Ctenoconularia are very similar to specimens of Conularia. ^^Cten-
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Annals of Carnegie Museum vol. 55
Fig. 7. — 7. 1-7.4; Conularia elegantula Meek. 7.1; AMNH CU 282G, holotype, comer
view; locality 174. 7.2; AMNH CU 282G, same specimen as in Fig. 7.1, comer major
1986
Babcock and Feldmann— North American Conulariida
381
oconularia"" delphiensis, which has apical angles of approximately 1 5°,
seems to expand from the apex too rapidly to warrant inclusion in the
genus Ctenoconularia.
Conularia delphiensis is similar in general morphology, including
similarity of rod articulation styles, to only one Devonian species from
North America, C. milwaukeensis. Conularia delphiensis differs, how-
ever, in having greater rods/cm values. Specimens here referred to C
delphiensis possess 26=42 rods/cm while specimens here referred to C.
milwaukeensis have fewer than 30 rods/cm.
Material examined. — ^ specimens; housed in the lUPC and the
USNM.
CONULARIA DESIDERATA Hall, 1861
Figs. 3.8=3.10, 5.1-5.9, 6.4=6.5, 16.6
Conularia =? Hall, 1859, p. 480.
Conularia desiderata Hall, 1861, PI. 72A, fig. 4; Richardson, 1942, p. 30-32, PI. 4, figs.
7, 8; Babcock and Feldmann, 1986, figs. IB, IH.
Conularia continens Hall, 1876, PI. 28, figs. 4-6; Miller, 1877, p. 141; Bigsby, 1878, p.
418; Hall, 1879, p. 212-214, PI. 33, fig. 6, PI. 34, figs. 3, 4, 6, PI. 34A, fig. 6; Lesley,
1889, p. 143, fig.; Miller, 1889, p. 390; Clarke and Ruedemann, 1903, p. 565; Moore
and Harrington, 1956a, fig. 25. 1; Moore and Harrington, 1956A p. F60, fig. 42. 10a-
b; Tasch, 1973, fig. 5.15H, Tasch, 1980, fig. 5.15H.
Conularia continens var. rudis Hall, 1879, p. 215-216, PL 34A, figs. 7-8; Miller, 1889,
p. 390; Clarke and Ruedemann, 1903, p. 565; Grabau, 1906, p. 331.
Conularia s.l. continens Hall. Sinclair, 1948, p. 286,
Par aconularia desiderata {Hail). Sinclair, 1948, p. 185.
Conularia s.l. rudis Hall. Sinclair, 1948, p. 284.
Description. —ETLOsk&lQton up to 10 cm in length. Major apical angle 14-27®; minor
apical angle 14-18°. Rod articulation inflected circular curve in early stages to angulated
circular curve in later stages. Rods almost always abut at midline; rod angle 7-17°. About
41 rods/cm in apical region; 14-27 rods/cm elsewhere. 3-4 nodes/mm; 3-4 adapertural
spines/mm; 3-4 adapical spines/mm. Apical wall not observed.
Occurrences. — and Middle Devonian rocks of New York and
Pennsylvania; localities 121, 124=126, 132-135, 142, 144, 232, 235-
236, and 239. Conularia continens, here referred to C. desiderata, has
been reported from Devonian rocks in Ohio (Claypole, 1903) and in
Indochina (Patte, 1 926), but the specimens upon which these references
were based were not studied. Woodward (1943) identified C continens
face. 7.3; AMNH CU 282G, same specimen as in Fig. 7.1, detail of major face. 7.4;
CMNH 44584, ?major face of flattened specimen; locality 175. 7. 5-7. 9; C. milwaukeensis
Cleland. 7.5; USNM 85988, holotype; locality 255. 7.6; USNM 78212; detail of apical
region. Note that no apical wall is present; locality 253. 7.7; MPM 20252, complete
specimen; locality 253. 7.8; MPM 20252, same specimen as in Fig. 7.7, detail of major
face. 7.9; MPM 22974; locality 253. Bar scales represent 1 cm.
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VOL. 55
Fig. %.-~Conularia pyramidalis Hall. 8.1; AMNH 33017, lectotype; comer view of flat-
tened specimen, preserved in calcareous shale; locality 117. 8.2; NYSM 3488, holotype
of C. huntiana Hall, comer view, preserved in calcareous shale; locality 1 1 8. 8.3; AMNH
1986
Babcock and Feldmann— North American Conulariida
383
var. rudis (=C delphiensis) in the Devonian of West Virginia, but
Woodward’s specimens were unavailable for study.
Types. — Holotype and plastoholotype of C delphiensis Hall, AMNH
2697; two syntypes of C continens Hall, NYSM 3485, NYSM 3486;
holotype of C continens var. rudis Hall, part and counterpart, NYSM
3487.
Remarks. — Conularia desiderata Hall is similar in appearance only
to C. pyramidalis Hall. Both species have rod angles in the range of
14-27°. Also, specimens of both taxa possess rods which appear to be
inflected at the midline. Features that are present in C. desiderata, but
not present in C pyramidalis include inflected circular curve to an-
gulated circular curve rod articulation style and rod angles of 14-27°.
The syntypes of C. continens Hall differ morphologically from the
holotype of C desiderata Hall only in being flattened. The holotype of
C continens var. rudis differs from the holotype of C desiderata in
being larger and better preserved. The type specimens of each taxon
have inflected gothic arch rod articulation in the region close to the
apex and inflected circular curve rod articulation elsewhere. The major
apical angle of the holotype of C desiderata, measures 23°; the minor
apical angle cannot be measured with certainty. The syntypes of C
continens have major apical angles of 16° and 19°. The major apical
angle of the holotype of C. continens var. rudis is 27°. All of the type
specimens have between 14 and 27 rods/cm and possess spines which
are difficult to observe in most specimens. Conularia continens Hall,
and C continens var. rudis Hall are, therefore, here considered junior
synonyms of C desiderata Hall.
Material examined. specimens; housed in the AMNH, BMS,
GSC, NJSM, NYSM, USNM and the private collections of Larry De-
cina and Paul Zell.
CONULARIA ELEGANTULA Meek, 1871
Figs. 3.4, 7. 1-7.4
Conularia elegantula Meek, 1871, p. 85-86; Meek, 1873, p. 228-229, PI. 23, fig. 4;
Miller, 1877, p. 141; Bigsby, 1878, p. 78; Whitfield, 1882, p. 242; Miller, 1889, p.
390; Babcock and Feldmann, 1986, fig. 2K.
Conularia s.l. elegantula Meek, Sinclair, 1948, p. 283.
33017, same specimen as in Fig. 8.1, detail of a minor face. 8.4; NYSM 3488, same
specimen as in Fig. 8.2, detail of a minor face. 8.5; NYSM 3490, holotype of C. lata
Hall, detail of major face; locality 122. 8.6; NYSM 3490, same specimen as in Fig. 8.5,
entire specimen, preserved as an external mold in fine-grained sandstone. 8.7; AMNH
33018; paralectotype, preserved in three dimensions, major face; locality 1 17. 8.8; AMNH
33018, same specimen as in Fig. 8,7, comer view. 8.9; AMNH 33018, same specimen
as in Fig. 8.7, minor face. Bar scales represent 1 cm.
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VOL. 55
Description.— Exoskoidon up to 7 cm in length. Major apical angle 20-30°; minor
apical angle about 1 7°. Rod articulation uniformly of angulated circular curve style. 6
nodes/mm on rods. 6 adapertural spines/mm; 6 adapical spines/mm. Rods usually abut
at midline; approximately 40% alternate left superior on major and minor faces; rod
angle 3-13°. 32-39 rods/cm. Apical wall not observed.
Occurrences. — Middle Devonian of Ohio; localities 174-176.
7>/?^.-Holotype, AMNH CU 282G.
Remarks. Conularia elegantula Meek is similar to C pyramidalis
Hall in the values for rod angles, the number of nodes/mm on the rods
and in the number of spines/mm. Conularia elegantula can be distin-
guished from C pyramidalis, and, indeed, all other species of Conularia
from the Devonian or Mississippian of North America by its rod ar-
ticulation architecture, which seems to be uniformly of angulated cir-
cular curve style. Partial specimens of C. desiderata exhibiting rods
from the apertural region can also be distinguished from specimens of
C elegantula by the curvature of the rods as they approach the comer
angles. The rods of specimens of C elegantula are noticeably more
inflected in the apertural direction than are rods belonging to specimens
of C. desiderata.
Material examined. —ThtCQ specimens; housed in the AMNH and
the CMNH.
CONULARIA MIL WA UKEENSIS Cleland, 1911
Figs. 7.5-7.9
Conularia congregata var. milwaukeensis Cleland, 191 1, p. 130, pi. 26, figs. 4-7; Teller,
1911, p. 251.
Mesoconularia milwaukeensis Sinclair, 1948, p. 119.
Conularia congregata milwaukeensis Cleland. Munthe, 1980, p. 6.
Description. — Exoskddon up to 5 cm in length. Major apical angle 12-15°; minor
apical angle 11-14°. Rod articulation uniformly of inflected gothic arch style. Rods abut
at midline; rod angle 5-18°. 18-24 rods/cm. 5-7 nodes/mm; 5-7 adapertural spines/
mm; 5-7 adapical spines/mm. Apical walls not observed.
Occurrence. —Middle Devonian of Wisconsin; localities 253-255.
7>/?c5. — Holotype, USNM 85988; five paratypes, USNM 78212.
Two paratypes, listed by Sinclair (1948) as MPM 244-245, could not
be found.
Remarks. — Conularia milwaukeensis may be distinguished from
similar appearing species such as C. elegantula Meek and C. desiderata
Hall by having only inflected gothic arch style rod articulation and by
attaining lengths of up to 5 cm, apparently without the addition of
apical walls. Conularia elegantula has narrow apical angles like C.
milwaukeensis, but its rod articulation style is exclusively angulated
circular curve. Likewise, C. desiderata has narrow apical angles, but
1986
Babcock and Feldmann— North American Conulariida
385
its rod articulation patterns include inflected circular curve style in the
apical region and angulated circular curve style elsewhere.
None of the examined specimens of C milwaukeensis preserves in-
tegument over the entire exoskeleton. Because of this, rods have ob-
viously been moved slightly from their original positions, making the
measurement of rod angles difficult. Also, because of the general lack
of preserved integument in most specimens, there is no evidence that
apical walls were developed in this species. Apical walls may have been
a feature of this species, but simply have not been preserved in the
samples studied.
Material examined, — 2>1 specimens; housed in the GSC, MPM, and
the USNM.
CONULARIA MULTICOSTATA Meek and Worthen, 1865
Figs. 9.1-9.5, 9.8, 10.1, 10.4, 12.1-12.5
Conularia multicostata Meek and Worthen, 1865, p. 252-253; Bigsby, 1878, p. 316;
Miller, 1889, p. 390; Weller, 1898, p. 190; Babcock and Feldmann, 1986, figs.
lA, IF.
Conularia micronema Meek, 1871, p. 84; Meek, 1875, p. 316, PI. 18, figs, la-d; Miller,
1877, p. 141; Bigsby, 1878, p. 316; Herrick, 1888<3. p. 94-95, PI. 2, figs. 18-19;
Herrick, 1888Z?, p. 49, PL 8, figs. 4-4a; Lesley, 1889, p. xv; Miller, 1889, p. 390,
fig. 643; Herrick, 1893, PI. 19, figs. 4-4a; Weller, 1898, p. 190; Grabau and Shimer,
1910, p. 13, figs. 1227c-f.
Mesoconularia multicostata (Meek and Worthen). Sinclair, 1948, p, 125.
Mesoconularia micronema (Meek), Sinclair, 1948, p. 124.
Mesoconularia mcfarlani Sinclair, [1948], p. 126-128, PI. 16, figs. 3-5.
Mesoconularia attica Sinclair, [1948], p. 125-126, PI. 9, fig. 2, PI. 17, figs. 9-11.
Diconularia micronema (Meek). Sinclair, 1952, p. 138-139; Moore and Harrington,
1956Z), p. F61, fig. 47.2.
Non Conularia trentonensis multicosta Ruedemann, 1912, p. 115-116; Ruedemann,
1930, p. 36; Goldring, 1935, p. 63.
Description.— Exosk&leion up to 25 cm in length. Major apical angle 20-24®; minor
apical angle 1 8-22®. Rod articulation of inflected gothic arch style in apical and most
other regions and of angulated circular curve style in apertural region of large specimens;
rods exhibiting inflected circular curve style articulation are mildly recurved near the
midline but they are angulated in the apertural direction at the midline. Rods abut at
midline; rod angle 9-17°. 26-60 rods/cm. Nodes prominent; 2-3 nodes/mm; 2-3 ad-
apertural spines/mm; 2-3 adapical spines/mm. Apical wall not observed.
Occurrences. — Mississippian of Indiana, Kentucky and Ohio;
localities 25, 71=73, 79-80, 193, 197-200, 205, 209, 214, 217-218,
223, 225, 227 and 228.
7>/7e'5'. — Holotype of Conularia multicostata Meek and Worthen is
lost; plastoholotype, with small fragments of fossil adhering, USNM
50157. Holotype of C. micronema Meek is apparently lost (Sinclair,
1948, p. 124); neotype, AMNH 6713. Specimen intended by Sinclair
(1948) to be holotype of Mesoconularia mcfarlani, UK 6089. Three
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VOL. 55
1986
Babcock and Feldmann— North American Conulariida
387
specimens which Sinclair (1948) intended to designate as types of M
attica, CM 34533-34534, GSC 87204.
Remarks. — Conularia multicostata Meek and Worthen is most sim-
ilar in morphology to C subcarbonaria Meek and Worthen. Like C
subcarbonaria, C. multicostata has prominent nodes and can have more
than 30 rods/cm. However, C. multicostata can be distinguished from
this and all other species of Conularia by the combination of its very
closely spaced rods, 26-60/cm, and its inflected gothic arch rod artic-
ulation in most places except the apertural region of large specimens.
More importantly, though, the rods of specimens referred to C sub-
carbonaria appear to be undulose. This is especially true for large
individuals (for example, Figs. 11.2-1 1.3).
In the apertural region of specimens of C. multicostata which are
large for the species, a unique form of angulated circular curve rod
articulation is present. This rod articulation pattern involves slight
recurvature beginning about 72 to V3 of the distance between the midline
and the comer angle; at the midline, the rods are slightly angulated.
Approximately 70% of specimens of C multicostata are found to have
well developed exoskeletal constrictions (for example. Figs. 9. 3-9. 5,
9.8, 10.1). These cannot be used as a species-level or genus-level taxo-
nomic criterion, however, since well preserved exoskeletal constrictions
have been observed in specimens belonging to nearly every taxon re-
ported herein and in Part B. This species is notable because it shows
good examples of this structure more frequently than any other known
species of conulariid from the Devonian or Mississippian of North
America.
According to Meek and Worthen (1865, p. 253), C. multicostata is
unique among conulariids in that it possesses rods so closely spaced
that “it is only under a good magnifier that the very minute crenulations
can be seen.” Later, Meek (1871, p. 84) described C micronema as a
Fig. 9. -—9. 1-9.5; Conularia multicostata Meek and Worthen. 9.1; USNM 50157, plas-
toholotype. Dark areas on photograph indicate areas where integument of the original
specimen still adheres; locality 203. 9.2; UK 6089, specimen intended by Sinclair (1948)
to be the holotype of Mesoconularia mcfarlani Sinclair, preserved in siderite concretion;
probably minor face; locality 73.9.3; AMNH 6713, specimen chosen by Sinclair (19421?)
as neotype of C micronema Meek, major face; locality 225. 9.4; AMNH 6713, same
specimen as in Fig. 9.3, comer view. 9.5; AMNH 6713, same specimen as in Fig. 9.3,
minor face. 9. 6-9. 7; C. subcarbonaria Meek and Worthen. 9.6; FMNH UC 6610, ho-
lotype of C intertexta Miller; detailed view of exoskeleton, locality 28. 9.7; FMNH UC
6610, same specimen as in Fig. 9.6, view of entire specimen. 9.8; C multicostata Meek
and Worthen, AMNH 6713, same specimen as in Fig. 9.3, detailed view of minor face.
Bar scales represent 1 cm.
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Fig. 10,— 10J--10.3; Conuiaria muiticostata Meek and Worthen, 10.1; USNM 50128,
comer view of specimen with pronounced exoskeletal constrictions; locality 225, 10,2-
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Babcock and Feldmann— North American Conulariida
389
conulariid distinct from all others because it possessed rods so closely
spaced that “it requires the aid of a magnifier to see them distinctly.”
In all morphological respects, the two species are remarkably similar,
assuming that AMNH 6713 is “typical” of the species C micronema
as suggested by Sinclair (1948, p. 142). Conularia micronema Meek is
here considered a junior synonym of C multicostata Meek and Wor-
then.
Two manuscript species, Mesoconularia mcfarlani Sinclair and M.
attica Sinclair, are here placed in synonymy with C multicostata for
the reason that they have very closely spaced rods and nodes. Values
for spacing of the rods and nodes are consistent with other specimens
referred to C. multicostata (see Appendix B in part B). The specimen
which Sinclair intended to designate the holotype of M. mcfarlani (Fig.
9.2) possesses an angulated circular curve style of rod articulation and
is very similar in morphology to specimens from the Mississippian of
Ohio which have been referred herein to C multicostata. The same
can be said for the three specimens (CM 34533-34534, GSC 87204)
referred to as M. attica in Sinclair’s (1948) unpublished thesis.
Conularia micronema was used by Sinclair (1 952, p. 138) as the type
species of the genus Diconularia. Sinclair (1952, p. 138-139) noted
that Diconularia was a probable form-genus which differs from Con-
ularia in routinely having closely spaced rods and an “accentuation of
the pustules.” Conularia, by contrast, was judged by Sinclair to exhibit
these features only in gerontic specimens. The genus Diconularia is
here included as a junior subjective synonym of Conularia because
there does not appear to be any consistent pattern of differences between
“Z). ” micronema and species included by Sinclair in the genus Conu-
laria. Examples supporting this argument are given below.
Conularia niagarensis Hall, the North American reference species
for the genus Conularia (Sinclair, 1940^z), has large, well-pronounced
nodes and rod spacing values in the range of about 12-24/cm. These
characters closely resemble those of the “typical” specimen of “D.”
micronema, AMNH 6713 (see Appendix B in Part B). The number of
rods/cm exhibited in AMNH 6713, 28-32, is well within the limits of
the genus Conularia as recognized herein. Species referable to Conu-
10.3; C. subcarbonaria Meek and Worthen. 10.2; UIPC 10680, holotype, minor face,
preserved in limestone; locality 13. 10.3; UIPC 10680, same specimen as in Fig. 10.2,
detail of minor face. 10.4; C. multicostata Miller and Gurley, USNM 50647, apical
region of specimen showing apical wall; locality 228. 10.5; C. subcarbonaria Meek and
Worthen, UIPC 10680, same specimen as in Fig. 10.2, major face. Bar scales represent
1 cm.
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Annals of Carnegie Museum
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Fig. ll. — Conularia subcarbonaria Meek and Worthen. 11.1; FMNH UC 6289, badly
weathered holotype of C. spergenensis Miller and Gurley, preserved in limestone; locality
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Babcock and Feldmann— North American Conulariida
391
laria from the Devonian and Mississippian of North America exhibit
9 to 84 rods/cm. Furthermore, no species of Conularia examined from
Devonian or Mississippian age rocks of North America are known to
consistently possess rods which are more closely spaced in large, pre-
sumably gerontic, individuals than in “average-sized” individuals.
Material examined. — 61 specimens; housed in the AMNH, CM,
CMNH, FMNH, GSC, and the USNM and the private collection of
Ron Fisher.
CONULARIA PYRAMIDALIS Hall, 1859
Figs. 3.3, 8.1-8.9, 16.7
Conularia pyramidalis Hall, 1859, p. 347-348; Hall, 1861, PI. 72A, figs, la-c; Miller,
1877, p. 141; Miller, 1889, p. 390; Whitfield and Hovey, 1899, p. 170-171; Rich-
ardson, 1942, p. 23-26, PI. 3, figs. 4-9; Sinclair, 1948, p. 106; Babcock and Feld-
mann, 1986, fig. 2E.
Conularia huntiana Hall, 1859, p. 348; Hall, 1861, PI. 72A, figs. 2a“b; Clarke and
Ruedemann, 1903, p. 566; Grabau, 1906, p. 151, fig. 65; Grabau and Shimer, 1910,
p. 13, fig. 1224; Sinclair, 1948, p. 106, PI. 9, fig. 1.
Conularia lata Hall, 1859, p. 479-480, PI. 70A, fig. 3, PI. 91, fig. 1; Bigsby, 1878, p. 78;
Miller, 1889, p. 390; Sinclair, 1948, p. 104.
Conularia huntana (sic) Hall. Miller, 1877, p, 141; Miller, 1889, p. 390.
Description.— Exoskelcton up to 20 cm in length. Major apical angle 17-23°; minor
apical angle 16-17°. Rod articulation gothic arch style in apical region and inflected
gothic arch elsewhere. Rods abut at midline; rod angle 5-22°. 7-17 rods/cm. 1-4 nodes/
mm; 1-4 adapertural spines/mm; 1-4 adapical spines/mm. Apical wall not observed.
Occurrences.— Lower Devonian of New Jersey and New York; lo-
calities 114, 116-120 and 122.
Types.— LeeXoXype, designated herein from James Hall’s syntypic
suite of four specimens, AMNH 33018, smaller of two specimens bear-
ing this number (Figs. 3.3, 8.1, 8.3); three paratypes, AMNH 33018,
larger specimen, and AMNH 33019, two specimens. Holotype of C
huntiana Hall, NYSM 3488; holotype of C. lata, NYSM 3490, plas-
toholotype, GSC unnumbered.
Remarks. — Conularia pyramidalis Hall is similar in size and apical
angles to C. desiderata Hall. Distinction between the two taxa is made
on the basis of differences in rod angles, 9-14® for C. pyramidalis versus
13-17® for C. desiderata. Additionally, when complete enough speci-
44. 11.2; FMNH UC 18494, larger of two specimens, flattened specimen with apical
wall preserved, preserved in calcareous shale; locality 38. 1 1.3; FMNH UC 18494, same
specimen as in Fig. 11.2, detail of major face. Note apparent convergence of rods at
exoskeletal constriction. 11.4; UIPC 10680, same specimen as in Fig. 10.2, detail in
region of comer groove; locality. 1 1.5; FMNH UC 6289, same specimen as in Figure
11.1, detail of exoskeleton. Bar scales represent 1 cm.
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5
1986
Babcock and Feldmann— -North American Conulariida
393
mens are studied, distinction between these two species can be made
on the basis of rod articulation style, gothic arch to inflected gothic
arch for C pyramidalis, compared to gothic arch to inflected circular
curve for C. desiderata. Differences in rod articulation styles imme-
diately serve to help distinguish C pyramidaiis Hall from C undulata
Conrad, though both may attain lengths greater than 1 5 cm. Conularia
pyramidaiis Hall is readily distinguished from C. elegantuia Meek by
having significantly fewer nodes and spines per unit space than are
present in C. pyramidaiis. The feature which immediately serves to
distinguish C. pyramidaiis from C multicostata Meek and Worthen is
the greater spacing between rods, 9- 14/cm in C pyramidaiis and up
to 32/cm in C. multicostata. Conularia multicostata and C subcar-
bonaria Meek and Worthen have more prominent nodes on the rods
than does C pyramidaiis.
Conularia huntiana Hall was distinguished from C pyramidaiis Hall
(Hall, 1859, p. 348) by its greater length, its smaller apical angles, the
greater convexity of its faces and differences in the appearance of ridges
and spines. The apical angles of the lectotype of C pyramidaiis are 1 8®
and 16°, and in the holotype of C. huntiana, the apical angles as mea-
sured are 13° and 11°. However, the holotype of C. huntiana is not
compressed to the extent that the lectotype of C pyramidaiis is. More-
over, the lectotype of C. pyramidaiis is a smaller specimen and pre-
sumably represents an earlier growth interval than does the holotype
of C huntiana. Therefore, a smaller set of apical angles is expected in
C. huntiana. Differences in convexity of the faces between the two taxa
is likely a result of differences in collapse of the exoskeletons after death
of the animals and/or differences in compression of the exoskeletons.
Differences in the appearance of the ridges and spines seem to be
functions of preservation: most of the lectotype of C pyramidaiis is
preserved as an internal mold, whereas the holotype of C huntiana
retains much integument. Where integument is present on the lectotype
of C pyramidaiis, it is identical to that on the holotype of C. huntiana
(compare Figs. 8.3 and 8.4). In both specimens, most of the rods are
broken out, leaving a “double ridge” arrangement of the integument,
Fig. 12. — Conularia multicostata Meek and Worthen. 12.1; GSC 87204, enlargement of
a specimen intended by Sinclair (1948) to be a paratype of Mesoconularia attka; locality
193. 12.2; CMNH 4684, external mold of a flattened specimen preserved in a siderite
concretion; locality 198. 12.3; CM 34533, specimen intended by Sinclair (1948) to be
the holotype of M attka; locality 193. Note that the apical wall is present. 12.4; FMNH
UC 540 14A, specimen figured by Herrick (1888a) as C. micronema; locality 228. 12,5;
GSC 87204, same specimen as in Fig. 12.1, view of a collapsed specimen preserved in
a siderite concretion; locality 193, Bar scales represent 1 cm.
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marking the former positions of the margins of the rods within the
integument.
Conularia lata Hall is included as a junior subjective synonym of
C pyramidalis largely because of qualitative similarities including near-
ly identical rod articulation styles. Also, values for apical angles and
rod spacing are similar in the lectotype of C pyramidalis and the sole
specimen referred to C. lata by Hall, NYSM 3490. Major and minor
apical angles for the lectotype of C. pyramidalis are 18° and 17°, re-
spectively. For the holotype of C. lata, they are 23° and 17°. Rods/cm
values for the lectotype of C. pyramidalis vary between 9 and 14 while
those of the holotype of C lata vary between 7 and 10. Values for rod
angles and node spacing on the rods for both specimens are also well
within the expected normal distribution for a species.
Material examined. ~A\ specimens; housed in the AMNH, CM,
CMNH, FMNH, GSC, NJSM, NYSM, USNM, and the private col-
lection of Paul Zell.
CONULARIA SUBCARBONARIA Meek and Worthen, 1865
Figs. 9.6-9.7, 10.2-10.3, 10.5, 11.1-11.5
Conularia subcarbonaria Meek and Worthen, 1865, p. 253; Meek and Worthen, 1873,
p. 520-522, PI. 19, figs. 4a-c; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Miller,
1889, p, 390; Keyes, 1894, p. 218; Weller, 1898, p. 191-192.
Conularia intertexta Miller {nomen nudum), \%91b, p. 692.
Conularia intertexta Miller, 1894, p. 317, PI. 10, fig. 4; Weller, 1898, p. 190.
Conularia spergenensis Miller and Gurley, 1893, p. 74-75, PL 8, fig. 2; Miller, 1897, p.
765; Weller, 1898, p. 191.
Mesoconularia subcarbonaria (Meek and Worthen). Sinclair, 1948, p. 123.
Mesoconularia intertexta (Miller). Sinclair, 1948, p. 123.
Diconularia micronema (Meek). Sensu Lane, 1973, p. 92-93, PI. 8, figs. 2-3.
Description.— ExoskeXQlon up to 35 cm in length. Major apical angle 10-21°; minor
apical angle 9-19°. Rods are undulose; rod articulation uniformly of angulated circular
curve style. Rods abut at midline; rod angle 4-6°. 1 7-3 1 rods/cm. Nodes prominent; 3-
5 nodes/mm on rods; no spines present. Apical wall may be present.
Occurrences. — Upper Devonian-Lower Mississippian of Illinois, In-
diana, Iowa and Missouri; localities 8-9, 13, 20, 24, 27-28, 37-38, 42,
44^46, 48, 51, 61, 64, 67, and 101.
Holotype, UIPC 10680, plastoholotypes USNM 50158,
FMNH UC unnumbered and GSC unnumbered. Holotype of C. in-
tertexta, FMNH UC 6610, plastoholotypes, USNM 68130 and GSC
unnumbered; holotype of C. spergenensis, FMNH UC 6289.
Remarks. — In the possession of prominent nodes along the rods, C.
subcarbonaria Meek and Worthen resembles the Mississippian species,
C. multicostata Meek and Worthen, and the Devonian species, C.
ulsterensis Howell. These taxa are readily distinguished, however, on
the basis of rod articulation style: C subcarbonaria possesses angulated
1986
Babcock and Feldmann-- North American Conulariida
395
circular curve rod articulation in which the rods are undulose, partic-
ularly in large individuals. On the other hand, C muiticostata has both
angulated circular curve and inflected gothic arch rod articulation and
C ulsterensis possesses only inflected gothic arch rod articulation. Ad-
ditionally, C ulsterensis is unique among the species of Conularia
studied in having a rod spacing of 45-84 rods/cm in adult specimens.
Like the Devonian species, C undulata Conrad, C subcarbonaria
possesses undulose rods. Conularia subcarbonaria can be distinguished
from C undulata in having more prominent nodes along the rods and
by having rods which nearly always abut at the midline. The rods of
C undulata alternate at the midline in approximately 1 0% of the cases
studied.
Conularia intertexta Miller is considered synonymous with C sub-
carbonaria because of general qualitative similarities between the types
of the two species and because of similarities in the value for number
of rods/cm (see Appendix B in Part B). However, the holotype specimen
(Figs. 9. 6-9. 7) of C. intertexta is very badly preserved, maHng any
species-level assignment somewhat doubtful. One characteristic of the
specimen is an apparently undulose set of rods. It is primarily because
of its undulose rods that C intertexta is here included in synonymy
with C subcarbonaria.
The holotype of C. spergenensis Miller and Gurley (Figs. 11.1, 1 1.5)
is, like the holotype of C intertexta, badly preserved. However, like
the holotype of the latter taxon, C spergenensis bears qualitative and
quantitative similarities to C subcarbonaria, most notably, undulose
rods. It too is therefore considered to be a junior subjective synonym
of C subcarbonaria.
Material examined. —46 specimens; housed in the AMNH, FMNH,
GSC, ISGS, lUPC, and the USNM.
CONULARIA rC/ZOJ Clarke, 1907
Fig. 15.3
Conularia desiderata var. tuzoi Clarke, 1907, p. 181, fig.; Clarke, 1908, p. 144, PL 11,
fig. 13; Dresser and Denis, 1944, p. 326.
Conularia tuzoi Clarke. Sinclair, 1948, p. 105,
Description. — based only upon holotype. Exoskeleton 1 1.3 cm in length.
Major apical angle approximately 10®; minor apical angle not observed. Rod articulation
style unknown in vicinity of apex and of inflected gothic arch style elsewhere; rods are
broadly inflected. Rods abut at midline; rod angle 9-10®. Nodes and spines not observed.
Apical wall not observed.
Type —Holotype, NYSM 9404.
Occurrence. — CowQT Mississippian of Quebec; locality 244.
Remarks. — The holotype of C tuzoi (Clarke) is badly preserved and
no additional specimens are known. A cross sectional view is not pre-
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Annals of Carnegie Museum
VOL. 55
Fig. \3. — Conularia undulata Conrad. 13,1; AMNH 41093, neotype, preserved as an
external mold in siltstone. Note rounded marks produced by orbiculoid brachiopods
1986
Babcock and Feldmann— North American Conulariida
397
served on the holotype. This taxon appears to differ from other species
of Conuiaria included in this paper in the possession of broadly rounded
inflections where the rods are of inflected gothic arch style.
Material examined, — 1 specimen, NYSM 9404.
CONULARIA ULSTERENSISllowGll, 1942
Figs. 15.4, 16.1-=16.3, 16.5
Conuiaria ulsterensis Howell, 1942, p. 91, figs. 10-11; Goldring, 1943, p. 208.
Mesoconularia ulsterensis (Howell). Sinclair, 1948, p. 117.
D^5m>£ioii.““Exoskdeton up to 3 cm in length. Major apical angle 14-19°; minor
apical angle 10-15°. Rod articulation uniformly of inflected gothic arch style. Rods usually
abut at midline; rod angle 1 1-15°. 45-84 rods/cm. Nodes prominent; 6-8 nodes /mm;
6-8 adapertural spines/mm; 6-8 adapical spines/mm. Apical wall not observed.
Typex— Holotype, PU 42071; two paratypes, PU 42072-42073.
Occurrences. — 'Lowqt Devonian of New York and Pennsylvania; lo-
calities 115, 231, 234 and 236. Specimens possibly referable to this
taxon have also been found in the Lower Devonian of Quebec; locality
242.
Remarks. — This species is distinct in its possession of up to 84 rods/
cm in adult specimens, the largest number recorded in the genus Con-
uiaria. It can also be distinguished by its prominent, closely spaced
nodes. In this form, nodes are spaced as closely as 7/mm. Some spec-
imens of C undulata Conrad may have as many as 7 nodes/mm, but
C undulata has neither inflected gothic arch rod articulation through-
out the exoskeleton nor prominent nodes.
The specimen of C. ulsterensis Howell illustrated in Fig. 16.2 is
preserved as an internal mold. It is unusual in that it clearly shows
that a longitudinal invagination existed along the integument internal
to the midline.
Material examined.--! specimens; housed in the CM, NJSM, and
the NYSM.
CONULARIA UNDULATA Conrad, 1841
Figs. 13.D13.5, 14.1-14.5, 15.1-15.2, 16.4
Conuiaria undulata Conrad, 1841, p. 57; Hall, 1861, p. 62-63; Bigsby, 1878, p. 62-63;
Hall, 1876, PL 29, figs. 1-7; Hall, 1879, p. 208-209, PI. 33, figs. 1-5, 7; PL 34A,
figs. 1-4; Miller, 1889, p. 390; Whitfield and Hovey, 1901, p. 326“”327; Grabau,
that were previously attached to the conulariid exoskeleton; locality 145. 13.2; NYSM
3493, preserved in siltstone, major face; locality 145. 13.3; NYSM 3494, preserved in
siltstone, major face; locality 145. 13.4; AMNH 41093, same specimen as in Fig. 13.1,
detail of major face. 13.5; AMNH 41093, same specimen as in Fig. 13.4, detail in region
of comer groove. Bar scales represent 1 cm.
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Annals of Carnegie Museum
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Fig. 14. — Conularia undulata. 14.1; AMNH 5439, specimen showing relation of apical
wall to the remainder of the exoskeleton. Note flange for connection at adapertural end
1986
Babcock and Feldmann— North American Conulariida
399
1899, p. 284-285, fig. 219; Grabau and Shimer, 1910, p. 13, fig. 1225-1226; Moore
and Harrington, \956b, p. F60, fig. 25.2; Babcock and Feldmann, 1986, fig. 21.
Conularia cayuga Hall, 1876, PL 28, figs. 2-3; Hall, 1879, p. 211-212, PI. 34, figs. 2, 5;
Miller, 1889, p. 390.
Conularia crebistria Hall, 1876, PI. 29, figs. 8, 9.
Conularia crebistriata {sic) Hall, 1879, p. 210-211, PI. 33, figs. 8-9, PL 34A, fig. 5; Miller,
1889, p. 390; Moore and Harrington, 1956A p. F60, fig. 42.9.
Mesoconularia undulata (Conrad). Sinclair, 1948, p. 118.
Conularia s.l. cayuga Hall. Sinclair, 1948, p. 285.
Ctenoconularia crebistriata {sic) (Hall). Sinclair, 1948, p. 241, PL 6, figs. 10-11.
Ctenoconularia crebistria (Hall). Sinclair, 1952, p. 142.
Conularia sp. cf. C. undulata Conrad. Kasznica, 1986, p. 14-15, fig. 2.
Description.— Exos\ie\Qion up to 15 cm in length. Major apical angle 10-18°; minor
apical angle 8-15°. Rod articulation uniformly of inflected circular curve style; rods are
undulose in the apertural Vi. Rods usually abut at midline; rod angle 10-18° in apical
region and 4-20° elsewhere. 17-32 rods/cm. 4 nodes/mm; 4 adapertural spines/mm; 4
adapical spines/mm. Apical wall may be present.
Neotype, AMNH 41093, plastoneotype, FMNH UC 694;
James Hall’s figured specimens, NYSM 3493, 3494, AMNH 5439.
Holotype of C. crebistria, AMNH 5440, plastoholotype, FMNH UC
679; holotype of C. cayuga, NYSM 3482, plastoholotype, FMNH UC
685.
Occurrences. Devonian of Maine, Maryland, New York,
Ontario, Pennsylvania, and Quebec; localities 90-91, 115, 128-129,
136-138, 140-141, 143, 145-148, 230, 236-238, and 241. Ulrich (1892)
has indicated that C undulata is present in the Devonian of Bolivia
and Reed (1904) has cited this taxon in the Devonian of South Africa.
These identifications are erroneous, and are described briefly below.
They will also be described in greater detail elsewhere. Cordini (1955,
p. 275, fig. 81) referred and figured some fossils found in Antarctica
as C. cf. C. undulata, but these have been subsequently identified as
plant remains (Dalziel et al., 1981).
Remarks.— Contdi&s, suite of syntypes is lost. However, judging from
his description of C. undulata (Conrad, 1841, p. 57) it is clear that the
species is based upon specimens now referable to either C. undulata
or C. pyramidalis Hall. Hall’s early figures and description of C. un-
dulata (Hall, 1876, Plate 29, figs. 1-7, explanation of Plate 29; 1879,
p. 208-209; Plate 33, figs. 1-5, 7, Plate 34A, figs. 1-4) have served as
bases for all subsequent studies on the species. Therefore, it is appro-
of apical wall; locality 145. 14,2; AMNH 5440, holotype of C. crebistria Hall, preserved
in calcareous shale, minor face; locality 136. 14.3; AMNH 5440, same specimen as in
Fig, 14.2, detail of minor face. 14.4; NYSM 3482; holotype of C. cayuga Hall, preserved
in calcareous shale; locality 136. 14.5; NYSM 3482, same specimen as in Fig. 14.4,
detailed view of minor face. Bar scales represent 1 cm.
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Annals of Carnegie Museum
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Fig, 15.-15.1-15.2; Conularia cf. C undulata Conrad. 15.1; NYSM 9410, detail of
exoskeleton; locality 241, 15.2; NYSM 9408, flattened and tectonically distorted spec-
imen; locality 90. 1 5.3; C tuzoi Clarke, NYSM 9404, holotype; locality 244. 1 5.4; NYSM
1986
Babcock and Feldmann— North American Conulariida
401
priate to select a neotype from among HalFs extant specimens. The
specimen chosen, AMNH 41093, is the subject of one of the best known
of Hairs figures of the taxon.
Two Devonian species are here considered junior subjective syn-
onyms of C undulata Conrad: C cayuga Hall and C crebistria Hall.
Conularia crebistria, according to Hall (1879, p. 210) differed from C
undulata in being “more slender in its mode of growth” and in having
more closely spaced rods. In terms of the arrangement of nodes and
spines, the two were judged to be “precisely similar.” The holotype of
C crebistria, AMNH 5440, is a badly preserved specimen exhibiting
one face (Figs. 13.2-13.3). The apical angle cannot be determined be-
cause the specimen has been distorted and the comer grooves are not
preserved on the surface of the slab. Like specimens of C. undulata,
this specimen has inflected circular curve rod articulation. The rods
are undulose in the vicinity of the aperture. The holotype of C. crebistria
has rod spacing values ranging from 24 to 30 rods/cm and node spacing
values of about 4 nodes/mm. No other values can be determined with
confidence from this specimen. The equivalent values for the lectotype
of C. undulata are 20-27 rods/cm and 6 nodes/mm. There seems to
be no significant difference in the features exhibited in AMNH 5440
from other specimens here referred to C. undulata.
The holotype of C cayuga, NYSM 3482, is a flattened specimen
preserved as an external mold (Figs. 13.4-13.5). An apical wall is
present. Hall (1879, p. 211-212) indicated that, in general, this spec-
imen is “not dissimilar to C undulata. ” However, subtle differences,
including “stronger” rods, wider spaced rods except in the apertural
region and the presence of “striae” between adjacent rods (=spines)
were used as key characters which served to distinguish this taxon from
C. undulata. Spines, of course, are present in C. undulata, just as they
are in the holotype of C. cayuga. The rods of NYSM 3482 have a
spacing of 17-22/cm, well within the expected range of values for
individuals of C undulata. Other quantitative determinants, given in
Appendix B in Part B, substantiate this conclusion. The rods are un-
dulose except near the apical wall and are articulated in angulated
circular curve style, similar to the neotype of C undulata. The rods
do not appear to be better pronounced than those shown in specimens
referred to C. undulata which are preserved as external molds, including
the neotype (Fig. 1 3.4). Thus, the holotype of C. cayuga is here referred
to Conrad’s species, C undulata.
9411, C. ulsterensis Howell, two specimens, one preserving an apical wall; locality 242.
Bar scales represent 1 cm.
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Annals of Carnegie Museum
VOL. 55
Fig. 16.— 16.1“16.3; Conularia ulsterensis Howell. 16.1; PU 420715 holotype; locality
115.1 6.2; PU 42072, paratype; internal mold showing longitudinal ridges at the midlines;
locality 115. 16.3; NJSM 12843, external mold; locality 236. 16.4; CM 34520, C un-
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Babcock and Feldmann— North American Conulariida
403
Sinclair (1948, p. 118) and Sinclair and Richardson (1954, p. 105)
stated that the species C grandis Roemer is synonymous with C. un-
dulata. However, this cannot be substantiated here because the spec-
imen upon which Roemer’s description is based (Roemer, 1856, p.
436, Plate 3, figs. 21a-b) was not available for study. It is likely that
the holotype of C. grandis is lost.
Conularia undulata is similar to C subcarbonaria Meek and Wor-
then in the possession of an undulose mode of angulated circular curve
style rod articulation. It differs from the latter form in having nodes
on the rods which are not prominent. Moreover, the rods of C sub-
carbonaria nearly always abut at the midline. In C undulata, as many
as 10% of the rods may alternate at the midline.
Two specimens from the Devonian of the Malvinokaffric Realm have
been misidentified as C. undulata. The first (Ulrich, 1892, p. 31-33,
PI. 3, figs. 6a-b), which was collected in Bolivia, is referable to C
albertensis Reed, judging from Ulrich’s well-executed figure. Conularia
albertensis differs from C. undulata in having gothic arch rod articu-
lation in the apical region and inflected gothic arch rod articulation
elsewhere, whereas C undulata has only inflected circular curve rod
articulation. The second specimen from the Malvinokaffric Realm which
was misidentified as C undulata was described and figured by Reed
(1904, p. 248-249, PL 31, figs. 1-la). It was collected from the Bok-
keveld beds of South Africa. A latex mold (UCGM 34720) of the
specimen has been examined. It is referable to C quichua Ulrich.
Conularia quichua and C. undulata both have undulose rods, but C
quichua has rods articulated in gothic arch fashion in the apical region
and in angulated circular curve style elsewhere.
Material examined.— 43 specimens housed in the AMNH, CM,
FMNH, GSC, NYSM, NJSM, USNM, and the private collections of
Gordon Baird, Robert Tinsley and Paul Zell.
CONULARIA sp.
Fig. 16.8
Conularia cf. huntiana Hall. Merriam, 1973, p. 35, PL 12, figs. 18-20.
Occurrence.— Upper Devonian of Nevada; Locality 108.
Remarks.— Remains of at least six conulariid specimens from the
Devonian of Nevada were figured and described by Merriam (1973,
dulata Conrad, portion of specimen with orbiculoid brachiopod attached; locality 91.
16.5; C. ulsterensisYiov<iQ\\, CM 34528. 16.6; C. desiderataUdlX, USNM 395827, juvenile,
with apex intact; locality 142. 16.7; C. cf. C. pyramidalis Hall, GSC 2598; locality 241.
16.8; C sp., USNM 159536; locality 108. Bar scales represent 1 cm.
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Annals of Carnegie Museum
VOL. 55
p. 35, Plate 12, figs. 18-20). Merriam identified these specimens as C.
cf. huntiana. The specimens (USNM 159536), one of which is figured
herein (Fig. 16.8), are poorly preserved and cannot be assigned to a
species at present.
The specimens in question possess an inflected circular curve rod
articulation. The rods are slightly deflected adaperturally at the midline.
This style of rod articulation is similar to that observed in specimens
of C. desiderata Hall.
Material examined. — 6 specimens; housed in the USNM.
Literature Cited
Babcock, L. E. 1985«. Hard and soft part morphology of selected Devonian and
Mississippian conulariids from the eastern United States. Geological Society of
America Abstracts with Programs, 17:278.
. 1985Z). Mahantango conulariid considered a hyolithid. Pennsylvania Geology,
16(3): 14-1 6.
Babcock, L. E., and R. M. Feldmann. 1984. Mysterious fossils. Earth Science, 37(3):
16-17.
—. 1986. The phylum Conulariida. In Problematic fossil taxa (A. Hoffman and
M. H. Nitecki, eds.), Oxford University Press.
Baird, G. C. 1981. Submarine erosion on a gentle paleoslope: a study of two discon-
tinuities in the New York Devonian. Lethaia, 14:105-122.
Baird, G. C., and C. E. Brett. 1981. Submarine discontinuities and sedimentary
condensation in the upper Hamilton Group (Middle Devonian): examination of
marine shelf and paleoslope deposits in the Cayuga Valley. New York State Geo-
logical Association, Guidebook, 53rd Annual Meeting, pp. 1 15-145.
Barrande, J. 1867. Systeme silurien du centre de la Boheme. lere Partie. Tome 3.
Classe des Mollusques, Ordre des Pteropodes. Prague and Paris, 179 pp.
Beede, J. W. 1911. The Carbonic fauna of the Magdalen Islands. New York State
Museum, Bulletin, 149:156-186.
Bigsby, j. j. 1878. Thesaurus Devonico-Carboniferous. The flora and fauna of the
Devonian and Carboniferous periods. . . London, 447 pp.
Boucek, B. 1939. Conularida. Pp. A1 13-A131, in Handbuch der Palaozoologie, Vol.
2A (O. H. Schindewolf, ed.), Berlin.
Boucek, B., and F. Ulrich. 1929. Etude sur la coquille du genre Conularia Miller.
Statniho geologickeho ustavu Ceskoslovenske Republicky, Vestnik, Roc., 5(2/3):
1-25.
Branisa, L. 1965. Los fosiles guias de Bolivia. Servicio Geologico de Bolivia, Boletin
6, 282 pp.
Branisa, L., and J. Vanek. 1973. Metaconularia cahuanotensis sp. n. from Bolivian
Lower Devonian. Vestnik Ustfedniho ustavu geologickeho, 48:95-96.
Branson, E. B. 1938. Stratigraphy and paleontology of the Lower Mississippian of
Missouri. Part 1. University of Missouri Studies, 13(3), 205 pp.
1944. The geology of Missouri. University of Missouri Studies, 19(3), 535 pp.
Calvin, S. 1890. Note on a specimen of Conularia missouriensis Swallow, with cren-
ulated costae. American Geologist, 5:207-208.
Caster, K. E. 1957. Problematica. In Treatise on marine ecology and paleoecology.
Volume 2, Paleoecology (H. S. Ladd, ed.). Geological Society of America, Memoir,
67:1025-1042.
Clarke, J. M. 1907. Some new Devonic fossils. New York State Museum, Bulletin,
107 (Geology 12) (New York State Education Department, Bulletin, 401:153-291.
1986
Babcock and Feldmann— North American Conulariida
405
1908. Early Devonic history of New York and eastern North America. New
York State Museum, Memoir 9, Part 1, 366 pp.
— — 1909, Early Devonic history of New York and eastern North America. New
York State Museum, Memoir 9, Part 2, 250 pp.
Clarke, J. M., and R. Ruedemann. 1903. Catalogue of type specimens of Paleozoic
fossils in the New York State Museum. New York State Museum, Bulletin, 65
(Paleontology 8) (University of the State of New York, Bulletin, 314), 76 pp,
Claypole, E. 1903. The Devonian era in the Ohio Basin. American Geologist, 32: 1 5-
41; 79-105; 240-250; 312-322; 335-353.
Cleland, H. F. 1911. The fossils and stratigraphy of the Middle Devonic of Wisconsin.
Wisconsin Geological and Natural History Survey Bulletin, 2 1 (Scientific Series 6),
222 pp.
Conrad, T. A. 1841. Fifth annual report on the palaeontology of the state of New-
York. State of New-York, Assembly Paper, 150:25-57.
CoRDiNi, I. R. 1955. Contribution al conocimiento del Sector Antarctico Argentino.
Instito Antartico Argentino, Publicacion, 1:273-277.
Dalziel, I. W. D., D. H. Elliot, D. I. Jones, J. W. Thompson, M. R. A. Thompson,
N. A. Wells, and W. J. Zinsmeister. 1981. The geological significance of some
Triassic microfossils from the South Orkney Islands, Scotia Ridge. Geological Mag-
azine, 118:15-25.
Dawson, J. W. 1868. Acadian geology. The geological structure, organic remains, and
mineral resources of Nova Scotia, New Brunswick, and Prince Edward Island. Sec-
ond edition. London, 694 pp.
Dresser, J. A., and T, C. Denis. 1944. Geology of Quebec. Volume 2: Descriptive
geology. Quebec Department of Mines, Geological Report 20, 544 pp.
Driscoll, E. G. 1963. Paraconularia newberryi (Winchell) and other Lower Missis-
sippian conulariids from Michigan, Ohio, Indiana, and Iowa. Contributions from
the Museum of Paleontology, University of Michigan, 18:33-46.
Eichwald, C. E. 1860. Lethaea Rossica, ou Paleontologie de la Russie. Tome 1, L’An-
cienne Periode. Part 2, pp. 681-1657. Stuttgart.
Emmons, E. 1846. Conularia verneulia. N.s. Emmons. American Quarterly Journal of
Science and Agriculture, 4:330,
Etheridge, R. 1901. Aperture of Conularia. Australian Museum, Records, 4(1): 5 2.
Feldmann, R. M,, and L. E. Babcock. 1986. Exceptionally preserved conulariids
from Ohio: reinterpretation of their anatomy. National Geographic Research, 2.
Feldmann, R. M., J. T. Hannibal, and L. E. Babcock. 1986. Fossil worms from the
Devonian of North America (Sphenothallus) and Burma (“Vermes”) previously
identified as phyllocarid arthropods. Journal of Paleontology, 60:341-346.
Finks, R. M. 1955. Conularia in a sponge from the west Texas Permian. Journal of
Paleontology, 29:831-836,
Fletcher, H. O. 1938. A revision of the Australian Conulariae. Australian Museum,
Records, 20:235-255.
Goldring, W. 1935. Geology of the Berne Quadrangle, with a chapter on glacial geology
by John H. Cook. New York State Museum Bulletin 303, 238 pp.
, 1943. Geology of the Coxsackie Quadrangle, New York, with a chapter on
glacial geology by John H. Cook. New York State Museum, Bulletin 332, 374 pp.
Gou, Zong-Hai, and Ji-Kai Yang. 1985. New material of Triassic Conularida from
Jiangyou of Sichuan, SW China. Acta Paleontologica Sinica, 24:358-360. [In Man-
darin.]
Grabau, a. W. 1899. Geology and palaeontology of Eighteen Mile Creek and the lake
Shore sections of Erie County, New York. Part 2, Palaeontology. Buffalo Society of
Natural Sciences, Bulletin, 6(2/4):93-403.
. 1 906. Guide to the geology and paleontology of the Schoharie Valley in eastern
New York. New York State Museum, Bulletin 92, pp. 76-386.
406
Annals of Carnegie Museum
VOL. 55
Grabau, a. W., and H. W. Shimer. 1910. North American index fossils, invertebrates.
Volume 2. New York, 909 pp.
Hall, J. 1847. Palaeontology of New-York. Volume 1. Containing descriptions of the
organic remains of the Lower Division of the New-York system (Equivalent of the
Lower Silurian rocks of Europe). Albany, 338 pp.
. 1852, Palaeontology of New-York. Volume 2. Containing descriptions of the
organic remains of the Lower Middle Division of the New-York System (Equivalent
in part to the Middle Silurian rocks of Europe). Albany, 362 pp.
-. 1858. Description of new species of fossils from the Carboniferous limestones
of Indiana and Illinois. Albany Institute, Transactions, 4:1-36.
. 1859. Geological survey of New York. Palaeontology. Volume 3. Containing
descriptions and figures of the organic remains of the Lower Helderberg Group and
the Oriskany Sandstone. Albany, 532 pp,
. 1861. Geological survey of New-York. Palaeontology. Volume 3. Part 2. Plates.
Albany, 141 plates.
. 1876. Illustrations of Devonian Fossils: Gasteropoda, Pteropoda, Cephalopoda,
Crustacea and Corals of the Upper Helderberg, Hamilton and Chemung Groups.
Albany, 7 pp.
. 1879. Geological survey of New York. Palaeontology. Volume 5. Part 2. Con-
taining descriptions of the gasteropoda, pteropoda and cephalopoda of the Upper
Helderberg, Hamilton, Portage and Chemung groups. Albany, 492 pp.
Harris, G. D. 1899. A key to the Upper Devonian of Southern New York designed
for teachers and students in secondary schools. Elementary Natural History Series,
No. 2. Ithaca, New York, 26 pp.
Herpers, H. 1949. A new conularid from the Esopus Formation, Sussex County, New
Jersey. New Jersey Department of Conservation and Economic Development, Mis-
cellaneous Geological Paper, 7 pp.
Herrick, C. L. 1888<2. Geology of Licking County, Ohio. Parts III and IV. The Sub-
carboniferous and Waverly groups. Denison University, Scientific Laboratories,
Bulletin, 3(1):13-110.
. 1888Z). Geology of Licking County. IV. List of Waverly fossils, continued.
Denison University, Bulletin, 4(1/2): 1 1-60, 97-123.
. 1 893. Observations on the so-called Waverly Group of Ohio. Geological Survey
of Ohio, Report, 7:495-515.
Holm, G. 1893. Sveriges Kambrisk-Siluriska Hyolithidae och Conulariidae. Sveriges
Geologiska Undersokning, Afhandlingar och uppsatser. Series C, No. 112, 172 pp.
Howell, B. F. 1942. New localities for fossils in the Devonian Esopus Grit of Ulster
County, New York. New York State Museum Bulletin, 327:87-93.
International Commission on Zoological N omenclature. 1985. International code
of zoological nomenclature. London, International Trust for Zoological Nomencla-
ture, 338 pp.
Jux, U. 1960. Montecaris lehmanni, a new crustacean from the Rhenish Devonian
and the problem of its systematic position. Journal of Paleontology, 34:1 129-1 152.
Kasznica, J. M. 1986. A conularid from the Mahantango Formation: first documented
report. Pennsylvania Geology 17(3): 13-1 6.
Keyes, C. R. 1894. Paleontology of Missouri (part II). Missouri Geological Survey,
Vol. 5, 266 pp.
Kiderlen, H. 1 937. Die Conularien. Uber Bau und Leben der ersten Scyphozoa. Neues
Jahrbuch fiir Mineralogie, Geologie und Palaontologie, 9e, usw., 77(B): 1 13-169.
Kindle, E, M. 1901. The Devonian fossils and stratigraphy of Indiana. Part 11. Pa-
leontology. Indiana, Department of Geology and Natural Resources, 25th Annual
Report, pp. 571-737.
1986
Babcock and Feldmann-- North American Conulariida
407
Knight, J. B. 1937. Conchopeltis Walcott, an Ordovician genus of the Conulariida.
Journal of Paleontology, 11:186-188.
Knod, R. 1908. Devonische Faunen Boliviens. Pp. 493-600, in Beitrage zur Geologic
und Palaontologie von Sudamerika, XIV (G. Steinmanii, ed.), Neues Jahrbuch fur
Mineralogie, Geologic und Palaontologie, usw., 25.
Kowalski, J, 1935. Les Conulaires. Quelques observations sur leur structure anato-
mique. Societe des sciences naturelles de I’Ouest de France, Bulletin, Serie 5, 5:28 1-
293.
Lamont, a, 1946. Largest British Conularia. Quarry Manager’s Journal, 29:569-570.
Lane, N, G. 1973. Paleontology and paleoecology of the Crawfordsville Fossil Site
(Upper Osagian; Indiana). University of California Press, Berkeley, 141 pp.
Laseron, C. F, 1912. Note on a new type of aperture in Conularia. Royal Society of
New South Wales, Journal and Proceedings, 45:247-249.
Lesley, J. P. 1889. A dictionary of the fossils of Pennsylvania and neighboring states
named in the reports and catalogues of the survey. Pennsylvania Geological Survey,
Report P 4, Volumes I~III, 437 pp.
Liu, Weizhou. 1981. Conulariid faunas from the Upper Paleozoic of central Jilin,
China. [In Mandarin.]
Maillieux, E. 1933. Terrains Roches et Fossiles de la Belgique. Second edition. Brus-
sels, 217 pp.
Markovski, B., and D. Nalivken. 1934. The Zadonsk and the Eletz beds. Union of
Soviet Socialist Republics, Geological, Hydrological and Geodetic Service, Trans-
actions, 313, 38 pp. [In Russian.]
Maroney, D, G., and R, W, Orr. 1974, Ctenoconuiaria delphiensis, a new species of
the Conuiata from the New Albany Shale (Upper Devonian) at Delphi, Indiana.
Indiana, Department of Natural Resources, Geological Survey Occasional Paper, 7,
6 pp.
Mason, C., and E. L. Yochelson. 1985. Some tubular fossils (Sphenothallm: “Vermes”)
from the middle and late Paleozoic of the United States. Journal of Paleontology,
59:85-95.
Meek, F. B. 1871. Descriptions of new species of invertebrate fossils from the Car-
boniferous and Devonian rocks of Ohio. Academy of Natural Sciences of Phila-
delphia, Proceedings for 1871, pp. 57-93.
- — — 1873. Descriptions of invertebrate fossils of the Silurian and Devonian systems.
Geological Survey of Ohio, Report, 1(2): 1-243.
. 1875, A report on some of the invertebrate fossils of the Waverly Group and
Coal Measures of Ohio. Geological Survey of Ohio, Report, 2(2):269-347.
Meek, F. B., and A. H. Worthen. 1865, Contributions to the paleontology of Illinois
and other western states. Academy of Natural Sciences of Philadelphia, Proceedings
for 1865, pp. 245-273.
— — . 1873. Descriptions of invertebrates from Carboniferous System. Geological
Survey of Illinois, 5:323-619.
Merriam, C. W. 1973. Paleontology and stratigraphy of the Rabbit Hill Limestone
and Lone Mountain Dolomite of central Nevada. United States Geological Survey,
Professional Paper, 808, 50 pp.
Miller, S. A. 1877. The American Paleozoic fossils: a catalogue of the genera and
species, with names of authors, dates, places of publication, groups of rocks in which
found, and the etymology and significance of the words, and an introduction devoted
to the stratigraphical geology of the Paleozoic rocks. Cincinnati, 253 pp.
— — — . 1889. North American geology and palaeontology for the use of amateurs,
students, and scientists. Cincinnati, 664 pp.
• 1 892f2. Palaeontology. Indiana Department of Geology and Natural Resources,
17th Annual Report, pp. 61 1-705.
408
Annals of Carnegie Museum
VOL. 55
. 1892Z). First appendix, 1892. Cincinnati, pp. 665-718.
. 1894. Palaeontology. Indiana Department of Geology and Natural Resources,
18th Annual Report, pp. 257-356.
. 1897. Second Appendix to North American Geology and Palaeontology, Oc-
tober, 1897. Cincinnati, pp. 719-793.
Miller, S. A., and W. F. E. Gurley. 1893. Descriptions of some new species of
invertebrates from the Palaeozoic rocks of Illinois and adjacent states. Illinois State
Museum of Natural History, Bulletin, 3, 81 pp.
. 1896. New species of Palaeozoic invertebrates from Illinois and other states.
Illinois State Museum of Natural History, Bulletin, 1 1, 50 pp.
Moore, R. C., and H. J. Harrington. 1956^1. Scyphozoa. Pp. F27-F38, in Treatise
on invertebrate paleontology. Part F, Coelenterata (R, C. Moore, ed.). Geological
Society of America and University of Kansas Press.
. 1956^. Conulata. Pp. F54-F66, in Treatise on invertebrate paleontology. Part
F. Coelenterata (R. C. Moore, ed.). Geological Society of America and University
of Kansas Press.
Munthe, J. 1 980. Catalog of fossil type and figured specimens in the Milwaukee Public
Museum. Milwaukee Public Museum, Contributions in Biology and Geology, No.
39, 27 pp.
Owen, R. 1862. Report of a geological reconaissance of Indiana, made during the years
1859 and 1860, under the direction of the late David Dale Owen, M.D., state
geologist. Indianapolis, 368 pp.
Palmer, K., and D. Brann. 1966. Illustrations of fossils of the Ithaca area. Ithaca,
Paleontological Research Institution, 28 pp.
Patte, E. 1926. Etudes paleontologiques relatives a la geologie de I’Est du Tonkin,
(Paleozoique et Trias). Service Geologique de ITndochine, Bulletin 15, 240 pp.
Reed, F. R. C. 1904. Mollusca from the Bokkeveld beds. South African Museum,
Annals, 4:239-174.
. 1933. Some new Ordovician species of Conularia from Girvan. Geological
Magazine, n.s. 70:354-358.
Richardson, E. S., Jr. 1942. A Middle Ordovician and some Lower Devonian con-
ularids, with two orthoceratids, from central Pennsylvania. Unpublished M.S. thesis,
Pennsylvania State College.
Richter, R., and E. Richter. 1930. Bemerkenswert erhaltene Conularin und ihre
Gattungsgenossen im Hunsriickschiefer (Unterdevon) des Rheinlandes. Sencken-
bergiana, 12:152-171.
Roemer, F. von. 1856. Pp. 433-788, in Lethaea Geognostica, usw. Third Revised
Edition, Lieferung II, Vol. 2 (H. G. Bronn and F. von Roemer, eds.).
Ruedemann, R. 1896a. Note on the discovery of a sessile Conularia. Article I. Amer-
ican Geologist, 17:158-165.
— . 1896Z). Note on the discovery of a sessile Conularia. Article II. American
Geologist, 18:65-71.
. 1912. The Lower Siluric shales of the Mohawk Valley. New York State Mu-
seum, Bulletin 162 (New York State Education Department, Bulletin 525), 151 pp.
. 1930. Geology of the Capital District (Albany, Cohoes, Troy and Schenectady
Quadrangles), with a chapter on glacial geology by John H. Cook. New York State
Museum, Bulletin 285, 218 pp.
. 1934. Paleozoic plankton of North America. Geological Society of America,
Memoir 2, 140 pp.
Shimer, H. W. 1905. Upper Siluric and Lower Devonic faunas of Trilobite Mountain,
Orange County, New York. New York State Museum, Bulletin 80 (Paleontology 10)
(New York State Education Department, Bulletin 330) pp. 173-269.
. 1926. Upper Paleozoic faunas of the Lake Minnewanka section, near Banff,
Alberta. Geological Survey of Canada, Bulletin, 42 (Geological Series, No. 45):
1-84.
1986
Babcock and Feldmann— North American Conulariida
409
Sinclair, G. W, 1940^2. The genotype of Conularia. Canadian Field-Naturalist, 54:
72-74.
— . 1940Z?. A discussion of the genus Metaconularia with descriptions of new
species. Royal Society of Canada, Section IV, Transactions, Series 3, 34: 101-121.
— . 1942^. A new species of Conularia"" from Gaspe. Le Naturaliste Canadien,
69:158-160.
. 1942^. The Chazy Conularida and their congeners. Annals of the Carnegie
Museum, 29:219-240.
1 944. Notes on the genera Archaeoconularia and Eoconularia. Royal Society
of Canada, Section IV, Transactions, Series 3, 38:87-95.
1948. The Biology of the Conularida. Unpublished Ph.D. thesis, McGill Uni-
versity, 442 pp.
1952. A classification of the Conularida. Fieldiana-Geology, 10:135-145.
Sinclair, G. W., and E. S. Richardson, Jr. 1954. A bibliography of the Conularida.
Bulletins of American Paleontology, 34, No. 145, 143 pp.
Slater, I. L. 1 907. A monograph of the British Conulariae. Palaeontographical Society,
41 pp.
Sowerby, J. 1821. The mineral conchology of Great Britain; or coloured figures and
descriptions of those remains of Testaceous animals or shells, which have been
preserved at various times, and depths in the earth. Volume 3, part 46. London.
Steinmann, G. and L. Doderlein. 1980. Elemente der Palaontologie. Verlag von
Wilhelm Engelmann, Leipzig.
Steul, H. 1984. Die systematische Stellung der Conularien. Giessener Geologische
Schriften, No. 37, 1 17 pp.
Swallow, G. C. 1860. Descriptions of new fossils from the Carboniferous and De-
vonian rocks of Missouri. Academy of Science of St. Louis, Transactions, 1:635-
660.
1863. Descriptions of some new fossils from the Carboniferous and Devonian
rocks of Missouri. Academy of Science of St. Louis, Transactions, 2:81-100.
Swartz, F. M., and E. S. Richardson, Jr. 1945. New structures in early Devonian
conularidae. Geological Society of America, Bulletin, 56:1206.
Tasch, P. 1973. Paleobiology of the Invertebrates. New York, 946 pp.
— . 1980. Paleobiology of the Invertebrates. Second Edition. New York, 975 pp.
Teller, E. E. 1911. A synopsis of the type specimens of fossils from the Palaeozoic
formations of Wisconsin. Wisconsin Natural History Society. Bulletin, 1 1:170-271,
Termier, H., and G. Termier. 1 949. Position systematique et biologic des Conulaires.
Revue Scientifique, 86(12), No. 3300, pp. 711-722.
Ulrich, A. 1892, Palaeozoische Versteinerungen aus Bolivien. Pp. 5-116, in Beitrage
zur Geologic und Palaeontologie von Sudamerika (G. Steinmann, ed.). Neues Jahr-
buch fiir Mineralogie, Geologic und Palaontologie, usw., 8.
Weller, S. 1898. A bibliographic index of North American Carboniferous inverte-
brates. United States Geological Survey, Bulletin, 153, 653 pp.
Werner, B. 1 966. Stephanoscyphus (Scyphozoa Coronatae) und siene direkte Abstam-
mung von den fossilen Conulata. Helgolander Wiss. Meeresunters., 13:317-347.
. 1967. Stephanoscyphus Allman (Scyphozoa Coronatae), ein rezenter Vertreter
der Conulata? Palaontologie Zietschrift, 41:137-153.
—. 1969. Neue Beitraege zur Evolution der Scyphozoa und Cnidaria. I Simposio
Intemacional de Zoofilogenia, Universidad de Salamanca, pp. 223-244.
White, C. A. 1862. Description of new species of fossils from the Devonian and
Carboniferous rocks of the Mississippi Valley. Boston Society of Natural History,
Proceedings, 9:8-33.
. 1876. Description of new species of fossils from Paleozoic rocks of Iowa.
Academy of Natural Sciences of Philadelphia, Proceedings for 1876, 28:27-34.
Whiteaves, J. F. 1891. The fossils of the Devonian rocks of the Mackenzie River
410
Annals of Carnegie Museum
VOL. 55
Basin. Geological and Natural History Survey of Canada, Contributions to Canadian
Palaeontology, 1:197-253.
Whitfield, R. P. 1882. On the fauna of the Lower Carboniferous limestones of Spergen
Hill, Ind., with a revision of its fossils hitherto published, and illustrations of the
species from the original type series. American Museum of Natural History, Bulletin,
i(3):39-97.
Whitfield, R. P., and E. O. Hovey. 1 899, Catalogue of the types and figures specimens
in the palaeontological collection of the Geological Department, American Museum
of Natural History. Part 11. Beginning with the Medina Sandstone. American Mu-
seum of Natural History, Bulletin, 11:73-188.
. 1901. Catalogue of the types and figured specimens in the paleontological
collection of the Geological Department, American Museum of Natural History.
Part IV. Carboniferous to Pleistocene, inclusive. American Museum of Natural
History, Bulletin, 11:357-500.
Wiman, C. 1895. Paleontologische Notizen 1-2. University of Upsala, Geological
Institution, Bulletin, 2(1 1), No. 3, pp. 109-1 17.
WiNCHELL, A. 1865. Descriptions of new species of fossils from the Marshall Group
of Michigan, and its supposed equivalent, in other states; with notes on some fossils
of the same age previously described. Academy of Natural Sciences of Philadelphia,
Proceedings for 1865, pp. 109-133.
Woodward, H. P. 1943. Devonian System of West Virginia. West Virginia Geological
Survey, 15, 655 pp.
WoRTHEN, A. H. 1 883. Description of some new species of fossil shells from the Lower
Carboniferous limestones and Coal Measures of Illinois. Geological Survey of Illi-
nois, 7:323-326.
Xu, Gueiyong, and Li Fenglin. 1979. New conulariid genera and species in China.
Acta Geologica Sinica, 2:91-98. [In Mandarin.]
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VOLUME 55 31 DECEMBER 1986 ARTICLE 16
DEVONIAN AND MISSISSIPPIAN
CONULARIIDS OF NORTH AMERICA.
PART B. PARACONULARIA, RETICULACONULARIA,
NEW GENUS, AND ORGANISMS REJECTED
FROM CONULARIIDA
Loren E. Babcock* ^
Rodney M. Feldmann*
Research Associate, Section of Invertebrate Fossils
Abstract
Descriptions of the species assigned to Pamconularia Sinclair, 1940 and Reticulacon-
ularia Babcock and Feldmann, new genus, as well as organisms rejected from the Con-
ulariida, are treated in Part B of this two-part work on the Devonian and Mississippian
conulariids of North America. Fifteen species of Pamconularia are considered valid, of
which five are new. The new taxa are P. alpenensis, P. chagrinensis, P. oklahomaensis,
P. wellsvillia, and P. yochelsoni. Adesmoconularia Driscoll, 1963 is considered a junior
synonym of Paraconularia. Two species are referable to Reticulaconularia Babcock and
Feldmann, new genus; Conularia penouili is selected as the type species.
Introduction
This paper is the second, and final, part of “Devonian and Missis-
sippian Conulariids of North America.” This work contains descrip-
‘ Address: Department of Geology, Kent State University, Kent, Ohio 44242.
2 Present address: Department of Geology, University of Kansas, Lawrence, Kansas
66045.
Submitted 2 June 1986.
411
412
Annals of Carnegie Museum
VOL, 55
tions of species referable to the genera Paraconularia and Reticula-
conularia n. gen., as well as specimens described in the literature as
conulariids but which are here rejected from the phylum. Locality
descriptions and measurements of selected specimens are included as
appendices A and B, respectively, herein. Figures are numbered con-
secutively in both Parts A and B in order to avoid cross-reference
confusion.
Genus PARACONULARIA Sinclair, 1940
Type species. — Conularia inequicostata Koninck, 1883, designated
by Sinclair (1940); Carboniferous of Belgium. Holotype: Musee Royal
d’Histoire Naturelle de Belgique, Brussels, Belgium. North American
reference species, Sinclair (1940): Conularia blairi Miller and Gur-
ley, 1893 (Mississippian). Lectotype of C blairi: UCGM 3985.
Diagnosis. —Corml^xiids with rods that are generally widely spaced,
4-35 rods/cm. More than 60% of rods alternate at midline; fewer than
40% abut. Apical angles small, 9-28°. Nodes, adapertural spines and
adapical spines may or may not be present; if present, they are usually
widely spaced, 2-6/mm.
PARACONULARIA ALPENENSIS
Babcock and Feldmann, new species
Figs. 17.1-17.3
Description. — T)Q^cnpX\on based only upon holotype. Exoskeleton 3.6 cm in length.
Major apical angle 21°; minor apical angle 16°. Rod articulation inflected circular curve
style; rods are slightly recurved near midline in apertural region. Rods abut or alternate
at midline; alternation pattern either right superior or left superior on major face, usually
left superior on minor face; rod angle 9-10°. 14 rods/cm. Nodes not observed; spines
absent. Apical wall not observed.
Occurrence.— Middle Devonian of Michigan; locality 92.
7>/7e -Holotype, GSC 85060.
Remarks.— taxon is similar to P. chesterensis (Worthen), P.
missouriensis (Swallow) and P. recurvatus Babcock and Feldmann, n.
sp. in the possession of rods that are recurved near the midline. Par-
aconularia alpenensis, however, exhibits rods that are not recurved in
the apical region, at least not in the holotype. None of the other three
taxa possess this characteristic.
It is not known whether P. alpenensis possessed nodes on the rods.
The holotype, and only known specimen, is weathered and lacks the
external surfaces of all the rods which are present. No spines are present.
The uniqueness of the rod articulation patterns are sufficient to distin-
guish this taxon from all other described taxa.
Material examined. — \ specimen, GSC 85060.
1986 Babcock and Feldmann— and Reticulaconularia
413
Etymology of trivial —Named for the Alpena Limestone, in
which the holotype was found.
PARACONULARIA ALTERNI STRIATA (Shimer, 1926)
Figs. 17.5-17.6
Conularia alternistriata Shimer, 1926, p. 84, PL 4, figs. 1 la-b.
Paraconularia alternistriata (Shimer). Sinclair, 1948, p. 190.
Description.— YOQsctiption based only upon holotype. Length 1.9 cm. Major apical
angle 1 1®; minor apical angle 10®. Rod articulation inflected gothic arch style; rods are
almost imperceptibly inflected near the comer angles. Rods always alternate at midline;
alternation pattern usually right superior on major face and usually left superior on minor
face; rod angle 9-10°. 28 rods/cm (extrapolated). Nodes and spines absent. Apical wall
not observed.
Occwrrpwc^. — Mississippian of Alberta; locality 4.
Type. — Holotype, GSC 5111.
Remarks. —Paraconularia alternistriata (Shimer) is similar in mor-
phology to specimens of P. yochelsoni Babcock and Feldmann, n. sp.
Both are of similar size, less than 3.5 cm in maximum length and both
exhibit similar forms of rod articulation style. The rod articulation
present on the holotype of P. alternistriata is here judged to be a form
of inflected gothic arch style. The rods in this specimen are inflected
so little, though, that the articulation could easily be confused for a
gothic arch style articulation pattern. This may simply be a function
of the small size of the holotype; a sample close to the aperture of a
larger specimen may yield a rod articulation pattern more distinctly of
an inflected gothic arch style. Paraconularia yochelsoni possesses rods
which are clearly articulated in an inflected gothic arch style close to
the apex and trending towards an inflected circular curve style near the
aperture. Paraconularia alternistriata is further distinguished from P.
yochelsoni in having a smaller apical angle, 10-11®, as compared to
1 5-20° in P. yochelsoni and, finally, in having greater rod spacing, 28
rods/cm as compared to 13-18 rods/cm.
The holotype, and only known specimen, of P. alternistriata exhibits
longitudinal folds in the integument between adjacent rods suggesting
that spines may have been present in this taxon. The folds are best
developed near the comer angles. Their occurrence seems to be erratic
and the spacing between adjacent folds is inconsistent. In all likelihood,
these folds do not represent integument folded over spines but simply
folds resulting from a contraction of the integument about the rods
and, perhaps, some shearing of the exoskeleton due to compression.
This phenomenon is relatively common among specimens of Para-
conularia, having also been observed in P. subulata (Fig. 3.2) and P.
missouriensis (Fig. 25.3).
Material examined. — 1 specimen, GSC 5111.
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1986 Babcock and Feldmann— PyiiMCOivt/i^i?//4 and Reticulaconularia
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PARACONULARIA BLAIRI (Miller and Gurley, 1893)
Figs. 18.1--18.5, 22.1=^22.3
Conularia biairi Miller and Gurley, 1893, p. 73-74, pi. 7, figs. 14-15; Miller, 1897, p.
765; Weller, 1898, p. 189; Chappars, 1936, p. 16; Branson, 1938, p. 110, PI. 14,
figs. 7-8; Branson, 1944, p. 216.
Conularia sedaliensis Miller and Gurley, 1896, p. 28, PL 3, figs. 4-5; Miller, 1897, p.
765; Weller, 1898, p. 191; Chappars, 1936, p. 16.
Conularia {Conularia) sedaliensis Miller and Gurley. Boucek, 1939, p. A 121.
Conularia (Paraconularia) biairi (Miller and Gurley). Sinclair, 1940, p. 74.
Paraconularia biairi (Miller and Gurley). Sinclair, 1948, p. 197; Moore and Harrington,
1956, p. F65, fig. 50.2.
Paraconularia sedaliensis Miller and Gurley. Sinclair, 1948, p. 201.
Paraconularia indiana Sinclair, [1948], p. 195, PL 18, figs. 1-2.
Paraconularia cf. newberryi (Winchell). Sensu Sinclair, 1948, PL 13, figs. 1-3.
Paraconularia missouriensis (Swallow). Sensu Babcock and Feldmann, 1984, p. 16-17.
Description. -~Exo%kQ\QXon up to 20 cm in length. Major apical angle 1 1-23°; minor
apical angle 10-20°. Rod articulation inflected gothic arch style in apical region and
inflected circular curve with a slight adapertural inflection at the midline elsewhere. Rods
almost always alternate at midline; alternation pattern usually right superior on major
and minor faces; rod angle 8-19°. 6-13 rods/cm. 2-3 nodes/mm; 2-3 adapertural spines/
mm; adapical spines absent. Apical wall not observed.
Types. —ThvQQ syntypes of C. biairi, UCGM 3984-3986, of which
UCGM 3986 (Fig. 1 8.5) is here designated the lectotype; UCGM 3984-
3985 are here designated the lectotype; UCGM 3984-3985 are here
considered paralectotypes. Four syntypes of C. sedaliensis, preserved
in five pieces, UCGM 1393, 1399; specimen intended by Sinclair (1948)
to be holotype of P. indiana, AMNH 25056.
Occurrences.— Towqt Mississippian of Illinois, Indiana, Iowa, and
Missouri; localities 11, 32, 35, 61, 94-98, 101, and 105. Laudon and
Bowsher (1941) reported this taxon in the Mississippian of New Mex-
ico, but their material was not available for study.
Remarks.— Paraconularia biairi (Miller and Gurley) is unique among
species of Paraconularia in having rods that exhibit inflected circular
Fig. 17.— 17.1; Paraconularia alpenensis Babcock and Feldmann, n. sp., GSC 85060,
holotype, minor face of specimen preserved in micrite; locality 92. 17.2; GSC 85060,
same specimen as in Fig. 17.1, comer view. 17.3; GSC 85060, same specimen as in Fig.
17.1, major face. 17.4; USNM 173926, Hyolithes sp., cmshed specimen of a hyolithid;
locality 240. 17.5-17.6; P. alternistriata Shimer. 17.5; GSC 5111, holotype, major face;
locality 4. 17.6; GSC 5111, same specimen as in Fig. 17.5, minor face. 17.7-17.8; P.
chesterensis (Swallow). 17.7; GSC 85061, a collapsed specimen preserved in siltstone;
locality 27. 17.8; GSC 85061, enlargement of same specimen as in Fig. 17,7. Note
inconspicuous spines on the rods. Bar scales in Figs. 17.1-17.4 and 17.7-17.8 represent
1 cm; bar scales in Figs. 17,5 and 17.6 represent 5 mm.
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Fig. IH.—Paraconularia blairi (Miller and Gurley). 18.1; UMC 4270, detailed view of
well preserved ?major face; locality 96, 18.2; UCGM 3985, syntype of Conularia se-
1986 Babcock and Feldmann— /l4i?^coAt/i:/4i?L4 and Reticulaconularia
417
curve style rod articulation and which are slightly inflected at the mid-
line. However, distinction between this species and P. subulata is often
difficult, especially in small specimens that preserve only inflected goth-
ic arch style rod articulation. Distinction between the two species can
be made on these criteria: 1 , P. blairi possesses distinct nodes on the
rods, P. subulata usually does not; 2, P. blairi has a slight adapertural
inflection of the rods near the midline except in the apical region, P.
subulata does not; and 3, P. blairi possesses 6-13 rods/cm whereas P.
subulata has 20-35 rods/cm.
Two species are here considered synonymous with P. blairi: P. se-
daliensis (Miller and Gurley) and P. indiana Sinclair, MS. The type
specimens of both are well enough preserved to compare all qualitative
and quantitative features of taxonomic interest. Values obtained by
measuring these specimens are given in Appendix B. In all respects,
P. sedaliensis and P. indiana are indistinguishable from the lectotype
and paralectotypes of P. blairi.
Material examined. — 34 specimens; housed in the AMNH, FMNH,
UCGM, UMC, and the USNM.
PARACONULARIA BYBLIS (White, 1862)
Figs. 3.7, 19.1-19.6, 23.2, 31.4
Conularia byblis White, 1862, p. 22; Miller, 1877, p. 141; Bigsby, 1878, p. 78; Herrick,
1888^2, p. 95; Miller, 1889, p. 390; Weller, 1898, p. 189; Weller, 1900^, p. 118-
1 19, PL 7, fig. 7; Weller, 1900Z?, p. 73; Grabau and Shimer, 1910, p. 14.
Conularia byblis White. Winchell, 1870, p. 257.
Conularia biblis (sic) White. Bigsby, 1878, p. 316.
Paraconularia byblis (V^hiXo). Sinclair, 1948, p. 200-201; Babcock and Feldmann, 1986,
figs. IE, 2B.
Adesmoconularia byblis (White). Driscoll, 1963, p. 40-41, PI. 3, figs. 1-7; Tasch, 1973,
fig. 5.14 Ga-b; Tasch, 1980, fig. 5.14 Ga-b.
Conularia! sp. Driscoll, 1963, p. 41, PL 3, fig. 8.
Description. —Exoskeleton up to 7 cm in length. Major apical angle 18-26®; minor
apical angle 10-19®. Rod articulation inflected gothic arch style in apical region and
inflected circular curve style elsewhere; rods are strongly inflected adaperturally at mid-
line; rod angle 12-18°. Rods generally abut at midline; 12-29 rods/cm. 1-2 nodes/mm;
spines seem to be absent. Apical wall may be present.
Occurrences. — Lower Mississippian of Indiana, Iowa, Kentucky and
Ohio; localities 29, 36, 39, 43, 50, 60, 62, 66, 71-72, 76, 77, 78, 81,
daliensis Miller and Gurley; locality 98. 18.3; UCGM 3985, counterpart of specimen in
Fig. 18.2. 18.4; UCGM 3984, paralectotype, ?minor face; locality 98. 18.5; UCGM 3986,
lectotype, a flattened specimen preserved in micrite; locality 98. Bar scales represent 1
cm.
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419
185, 195, 210, and 223-224. A specimen referred with question to this
species, UMMP 26735, is from Tennessee; locality 250.
—Holotype, UMMP 2167.
Remarks. —Paraconularia byblis (White) is distinguished from all
other species of Paraconularia by the combination of closely spaced
rods, 12-29/cm, the lack of spines and by rod articulation involving
an inflected gothic arch style in the apical region and an inflected
circular curve style elsewhere. There does not seem to be a species
present in the Devonian or Mississippian rocks of North America with
which this taxon could be easily confused if well preserved specimens
were available for study.
Driscoll (1963), designated C. byblis White as the type species of a
new genus, Adesmoconularia. Adesmoconularia, by Driscoll’s defini-
tion, is distinguished from Calloconularia Sinclair by a larger size and
a lack of “swelling” of the interridge areas near the comer angles in
Adesmoconularia. Adesmoconularia was deemed by Driscoll unlike
Paraconularia Sinclair in the lack of nodes and by the presence of an
apical wall in Adesmoconularia. To date, no other species have been
referred to the genus Adesmoconularia.
Examination of the holotype of Calloconularia strimplei Sinclair
(FMNH PE 1 42), the type species of the genus Calloconularia, reveals
that there is no expansion of the interridge areas near the comer angles.
Examination of the holotype of Conularia byblis White, type species
of the genus Adesmoconularia Driscoll, shows that nodes are present
on the rods, but they are very small and inconspicuous. The holotype
appears to have been considerably weathered, rendering the nodes
inconspicuous in most places on the specimen. Apical walls are present,
but rare, in specimens of Paraconularia. According to Driscoll’s di-
agnosis, then, size is the only criterion which distinguishes Adesmo-
conularia from Calloconularia', there is no distinction between Ades-
moconularia and Paraconularia. Therefore, Adesmoconularia Driscoll,
1963 is here considered a junior synonym of Paraconularia Sinclair,
1940.
Fig. \9.— Paraconularia byblis (White). 19.1; UMMP 2167, holotype, a weathered spec-
imen preserved in micritic limestone, comer view; locality 62, 19.2; CMNH 4492, small
specimen, preserved in shale and compressed along the faces and at the aperture; locality
219. 19.3; CMNH 2295, external mold preserving apical region; locality 195. 19.4;
UMMP 2167, same specimen as in Fig. 19.1, minor face. 19.5; UMMP 2167, same
specimen as in Fig. 19.1, detail of minor face. 19.6; CMNH 4691, ?major face of a
specimen preserved in shale; locality 185. Bar scales represent 1 cm.
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Fig. 20.— Paraconularia chagrinensis Babcock and Feldmann, n. sp. 20,1; CMNH 6717,
paratype, preserved in a dark gray phosphatic concretion. Note apparent healed wound;
1986 Babcock and Feldmann— and Reticulaconularia
421
Material examined. — S'i specimens; housed in the CM, CMNH, GSC,
FMNH, UMMP, and the USNM.
PARACONULARIA CHAGRINENSIS
Babcock and Feldmann, new species
Figs. 20.1-20.6, 21.1, 21.2
Description. ~~Exosk.Q\Qton up to 9 cm in length. Major apical angle about 28®; minor
apical angle 20-21°. Rod articulation exclusively of inflected gothic arch style. Rods
usually alternate at midline; rods, if they alternate, usually alternate left superior on both
major and minor faces; rod angle 9-12®. 16-20 rods/cm. 3-4 nodes/mm; nodes appear
to be subtle in apical region and prominent in apertural region; 3-4 adapertural spines/
mm; 3-4 adapical spines/mm. No apical wall observed.
Occurrences.— Devonian of Ohio; localities 178-184.
rj;/7^.y.-Holotype, CMNH 6633; 12 paratypes, CMNH 1247, 1272,
1427, 1622, 1674, 1788, 1818, 4030, 4292, 6717, 6807-6808.
Remarks.— Among species of Paraconularia, only P. chagrinensis
Babcock and Feldmann, n. sp. possesses the combination of wide apical
angles, 20-28°, adapertural and adapical spines, as well as rod articu-
lation which is exclusively of inflected gothic arch style. More striking,
however, is the pattern of nodes on the rods. Paraconularia chagri-
nensis is the only conulariid observed which appears to have nodes
which increase in size aperturally. Nodes are inconspicuous in the apical
region, but are prominent in the apertural region. The increase in size
of the nodes is not well shown in the holotype owing to the poor
preservation of the apertural region of this individual. The pattern is
well documented, however, in CMNH 6717 (Fig. 20.6).
When fragments of exoskeleton from the apertural region are found
alone, as is the case with the specimen illustrated in Figs. 20.3-20.4,
they are easily mistaken for species of Conularia such as C subcar-
bonaria or C. multicostata. This dilemma can be resolved only when
more complete material is found. Most conulariids from the small,
presumably phosphatic, nodules found in the Upper Devonian Chagrin
Shale of northeastern Ohio are preserved as fragmentary specimens,
which renders generic identification difficult. To date, only P. chagri-
nensis has been identified from this unit.
One paratype, CMNH 6717, is noteworthy not only for demonstrat-
ing the unique pattern of the nodes, but also for exhibiting an apparent
locality 179. 20.2; CMNH 1622, paratype; locality 184. 20.3; CMNH 1818, paratype;
locality 181. 20.4; CMNH 1818, same specimen as in Fig. 20.3, detail showing nodes
and spines. 20.5; CMNH 6633, holotype; locality 1 80. 20.6; CMNH 67 1 7, same specimen
as in Fig. 20.1, ?minor face. Note increase in size of nodes adaperturally. Bar scales
represent 1 cm.
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healed injury on one face (Fig. 20,1). On this specimen, rods in the
apertural region have been truncated and their broken ends rounded
slightly. Several rods have filled much of the region from where exo-
skeleton has been removed. These rods are oriented at a high angle to
the rods which comprise the remainder of the exoskeleton and are
complete with a midline distinct from the original midline on this face.
A small gap is left between the most adapical portion of the injury and
the most adapical rods which have filled the void. This region is filled
with integument that lacks embedded rods.
Material examined. --13 specimens; housed in the CMNH.
Etymology of trivial name. — Named for the Chagrin Shale, currently
the only known occurrence of this taxon.
PARACONULARIA CHESTEREN SI S {V^orthQn, 1883)
Figs 17.7-17.8, 22.4-22.7, 23.1-23.4,
23.7, 24.1, 24.3, 32.5
Conularia chesterensis Woithen, 1883, p. 325; Miller, 1889, p. 390, Worthen, 1890, p.
134, PI. 1 1, fig. 9a-b; Miller, 1897, p. 765; Weller, 1898, p. 189; Kent, 1982, p. 27.
Pamconularia chesterensis Worthen. Sinclair, 1948, p. 201-202; Babcock and Feldmann,
1986, fig. 4C.
Paraconularia newberryi (Winchell). Sensu Driscoll, 1963, PI. 2, figs. 6-9.
Paraconularia crawfordsvillensis (Owen). Sensu Lane, 1973, p. 93-95, PI. 8, fig. 1, PI. 9,
figs. 1-2.
Description. — Exoskeleton up to 20 cm in length. Major apical angle 14-21®; minor
apical angle 1 0-18°. Rod articulation exclusively of inflected circular curve style, recurved
near the midline. Rods alternate or abut at midline; rods, if they alternate, are not
preferentially right superior or left superior on either the major or minor face; rod angle
8-12°. 8-20 rods/cm. 4-5 nodes/mm; adapertural spines appear to be absent in apical
region, but small spines are sometimes present, 4-5/mm, in apertural region; adapical
spines absent. Apical wall not observed.
Occurrences. — Upper Mississippian of Alabama, Kentucky, Illinois,
Indiana, Iowa, Missouri, Nevada, Tennessee; localities 1, 10-12, 16-
21, 33, 35-36, 38-41, 47, 49-59, 70, 84-89, 104, 1 10, and 251. Spec-
imens referred questionably to this species have also been found in
British Columbia and Utah; localities 7 and 352.
Types. -nololypc, ISGS 2489.
Remarks. —Paraconularia chesterensis (Worthen) is similar to P. al~
penensis Babcock and Feldmann, n. sp., P. missouriensis (Swallow) and
P. recurvatus Babcock and Feldmann, n. sp. in having rods that are
recurved near the midline. Of these, P. alpenensis does not exhibit rods
that are recurved in the apical region, and both P. missouriensis and
P. recurvatus exhibit rods that are strongly recurved. The rods of P.
chesterensis tend to be slightly recurved. A rod pair in this taxon often
approximates the outline of a truncated pyramid (Figs. 22.4, 22.7).
In some cases, values for apical angles, rods/cm and rod angles may
1986 Babcock and FELDMANN"-F^iL4C<9ivt/i^^/^ and Reticulaconularia
423
be similar for specimens of P. chesterensis and P. missouriensis. If rod
articulation is also similar, distinction between the two may be made
on the basis of the spacing between nodes. Paraconularia chesterensis
possesses 4-5 nodes/mm and P. missouriensis possesses only 2-3 nodes/
mm.
Specimens referable to P. chesterensis which have been observed
with well-preserved spines are few in number. Moreover, it seems that
spines are only produced in the apertural regions of those individuals
that have them. When present, the spines are usually inconspicuous
and seem to be directed only in the apertural direction (for example,
Fig. 17.8). Many specimens referable to P. chesterensis (for example,
Fig. 23.2), however, seem to have small ridges developed in the integ-
ument between ridges. These ridges resemble interridge crests. Such
structures may indicate that adapertural, and perhaps even adapical,
spines are produced in areas other than the apertural region in this
taxon. No specimens exhibiting this have been observed to date. Thus,
the observations that only adapertural spines are present in P. ches-
terensis, and when present, that they occur only in the apertural region,
may be erroneous and owing to a lack of evidence to the contrary.
Material examined. — ?> 5 5 specimens; housed in the FMNH, GSC,
ISGS, lUPC, and the USNM.
PARACONULARIA MISSOURIENSIS 1860)
Figs. 21.3, 25.1, 25.2-25.5, 26.1-26.2, 32.1
Conularia missouriensis Swallow, 1860, p. 657; Miller, 1877, p. 141; Bigsby, 1878, p.
316; Miller, 1889, p. 390; Keyes, 1894, PI. 35, fig. la-b; Miller, 1897, p. 765; Weller,
1898, p. 190; Grabau and Shimer, 1910, p. 14; Branson, 1944, p. 246.
Conularia missouriensis Swallow?. Meek and Worthen, 1873, p. 541-542, PL 22, fig. 5;
White, 1880, p. 513, PL 6, fig. 4; Walcott, 1884, p. 264, PL 23, fig. 4.
Paraconularia (Swallow). Sinclair, 1948, p. 198-199.
Conularia gratiosa Miller and Gurley, 1893, p. 74, PL 8, fig. 1; Miller, 1897, p. 765;
Weller, 1898, p. 190.
Conularia greenei Miller and Gurley, 1896, p. 27-28, PL 3, fig. 3; Miller, 1897, p. 765;
Weller, 1898, p. 190, Cumings, 1906, p. 1367, PL 24, fig. 14.
Paraconularia greenei (Miller and Gurley). Sinclair, 1948, p. 194.
Paraconularia gratiosa (Miller and Gurley). Sinclair, 1948, p. 198.
Paraconularia sciotovillensis OtiscoW, 1963, p. 37-40, PL 1, figs. 9-12; Tasch, 1973, fig.
5.16, Table 5.2; Tasch, 1980, fig. 5.16, Table 5.2.
Ctenoconularial greenei (Miller and Gurley). Moore and Harrington, 1956, p. F65, fig.
51.4.
Conularia sp. Leary, 1985, PL 3, fig. 4.
Paraconularia cf. P. missouriensis (Swallow). Babcock, 1985a, p. 66-70, fig. lA-B.
Description.— up to 22 cm in length. Major apical angle 14-22°; minor
apical angle 10-18°. Rod articulation inflected circular curve style, strongly recurved
near the midline. Rods usually alternate at midline; if rods alternate, pattern is usually
left superior; rod angle 6-17°. 4-10 rods/cm. 2-3 nodes/mm; 2-3 adapertural spines/
mm; adapical spines absent. Apical wall not observed.
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Fig. 2 1 . — 2 1 . 1 ; Paraconularia chagrinensis Babcock and Feldmann, n. sp., CMNH 1788,
two small paratype specimens, presumably attached to the same object (obscured) and
1986 Babcock and Feldmann— il4i^4CDivc/i^^/^ and Reticulaconularia
425
Occurrences.— l^owQt Mississippian of Alberta, Illinois, Indiana, Iowa,
Kentucky, Missouri, and Ohio; localities 2, 5, 14-15, 31, 39, 69, 83,
94, 100, 103, 223 and 226-227. A specimen referred with question to
this species has been found in Alberta; locality 5.
Plaster cast of presumed holotype, FMNH UC 6639. Ho-
lotype of C. gratiosa, FMNH UC 6627, plastoholotype, USNM 67893;
holotype of C greenei, FMNH UC 6628, plastoholotype USNM 67880;
holotype of P. sciotovillensis, UMMP 26740.
—Considerable confusion has existed over the definition
of P. missouriensis (Swallow). This confusion of nomenclature is related
to at least two problems: 1, an ambiguous original definition of the
species, a definition which may have incorporated characters now iden-
tified as belonging to at least two species; and 2, a loss of Swallow’s
original specimens. Swallow’s type material was found in the “Car-
boniferous Limestone” of Cooper County, Missouri. This locality has
yielded at least two conulariid species; herein, they are identified as P.
missouriensis and P. blairi. It is possible, from Swallow’s description
(Swallow, 1860, p. 657), that specimens belonging to both forms were
used in the formulation of the original definition of P. missouriensis.
Some early authors, most notably Meek and Worthen (1873) and
Keyes (1894), used Swallow’s indication that the faces of P. missou-
riensis were “marked by flexuous, high, sharp plications” as the primary
determinative characteristic of the species. This concept of the species
is followed herein. A plaster cast of a specimen, marked “holotype?”
of P. missouriensis (FMNH UC 6639) is presumed to represent a cast
of the holotype of this species.
Paraconularia missouriensis is similar to P. chesterensis (Worthen)
in having an inflected circular curve style of rod articulation, with the
rods being reflexed near the midline. The degree of reflexure, however,
is greater in P. missouriensis. Paraconularia missouriensis can also be
distinguished from P. chesterensis by having a greater number of nodes/
mm on the rods, 4-5 nodes/mm as compared to 2-3 nodes/mm.
Other species of Paraconularia which have recurved rods include P.
alpenensis Babcock and Feldmann, new species and P. recurvatus Bab-
preserved in a phosphatic concretion. Arrow indicates a stalk; locality 1 84. 2 1 .2; CMNH
4294, partially disarticulated paratype; locality 183. 21.3; P. missouriensis (Swallow),
FMNHUC 1 125, view of major face; locality 14. 21.4; P. subulata(¥i?A\), USNM 395829,
preserved in a very dark gray, organic-rich shale. Note that no integument is present
and that rods are disarticulated. This specimen indicates that rods and integument are
separate components of the conulariid exoskeleton. Two specimens of ^^Linguld’" {=Bar-
roisellal) are visible in this photograph; locality 189. Bar scales in Figs. 21.2-21.4 rep-
resent 1 cm; bar scale in Fig. 21.1 represents 5 mm.
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7
1986 Babcock and Feldmann —Paraconularia and Reticulaconularia
427
cock and Feldmann, new species. Neither of these taxa have rod spacing
values less than 14 rods/mm or adapical spines on the rods and are
therefore easily distinguished from P. missouriensis. Moreover, P. ah
pensis possesses non-recurved rods in the apical region, unlike P. mis-
souriensis.
The species Conularia gratiosa Miller and Gurley, C. greenei Miller
and Gurley and P. sciotovillensis Driscoll are here included as junior
synonyms of C missouriensis because they all exhibit: 1 , similar values
for apical angles; 2, similar values for rod angles; 3, same rod articu-
lation style, including right superior rods if the rods alternate at the
midline; 4, similar values for rod spacing; 5, similar values for nodes/
mm; and 6, presence of adapical spines. Comparative values are given
in Appendix B.
Material examined. — 30 specimens; housed in the AMNH, FMNH,
GSC, ISGS, and the USNM.
PARACONULARIA OKLAHOMAENSIS
Babcock and Feldmann, new species
Fig. 27.5
Description. — Ty^scn^iion based only upon holotype. Exoskeleton 5.4 cm in length.
Major apical angle 19°; minor apical angle 17°. Rod articulation of gothic arch style in
apical end and of inflected circular curve style elsewhere. Rods usually abut at midline;
if they alternate, pattern is usually right superior on major face and left superior on
minor face; rod angle 12-13°. 24 rods/cm in apical region, 12 rods/cm elsewhere. Nodes
and spines absent. Apical wall not observed.
Occurrence.— UppQT Mississippian of Oklahoma; locality 229.
--Holotype, USNM 409811.
Remarks.— Par aconularia oklahomaensis Babcock and Feldmann,
n. sp. differs from all other described species of the genus in the com-
bination of rod articulation pattern, frequent rod abuttment, spacing
of rods, with 24 rods/cm in the apical region and 1 2 rods/cm elsewhere
and the lack of nodes and spines on the rods.
Fig. 22. — 22.1--22.3; Paraconularia blairi (Miller and Gurley). 22.1; AMNH 25056,
specimen intended by Sinclair (1948) to be holotype of P. indiana Sinclair, major face;
locality 32. 22.2; AMNH 25056, same specimen as in Fig. 22,1, comer view. 22.3;
AMNH 25056, same specimen as in Fig. 22.3, minor face. Note overturned apertural
termination. 22.4--22.7; P. chesterensis (Worthen). 22.4; lUPC 17414, bryozoan=en-
cmsted specimen; locality 1. 22.5; lUPC 17415, comer region of collapsed specimen
that has been encmsted by bryozoans subsequent to collapse; locality unknown, 22.6;
FMNH UC 23023; locality 38. 22.7; lUPC 11316; locality 57. Bar scales represent
1 cm.
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Fig. 23.-23.1-23.4; Paraconularia chesterensis (Worthen). 23.1; ISGS 2489, holotype,
preserved in micritic to sparry limestone, minor face; locality 10. 23.2; ISGS 2489, same
specimen as in Fig. 23.1, comer view. 23.3; ISGS 2489, same specimen as in Fig. 23.1,
1986 Babcock and Feldmann —Paraconularia and Reticulaconularia
429
This taxon is easily confused with P. subulata (Hall), which also lacks
nodes and spines on the rods and has inflected gothic arch rod artic-
ulation in the apical region and inflected circular curve rod articulation
elsewhere. The rods of P. oklahomaensis, however, are greatly inflected
in the vicinity of the comer angles in the apical region and almost
imperceptibly inflected elsewhere. In specimens of P. subulata, the
pattern of relative inflexure of the rods is reversed. Paraconularia okla-
homaensis is further distinguished from P. subulata in having 40-50%
of the rods abutting at the midline whereas specimens referred to P.
subulata seldom have more than 10% of the rods abutting.
Material examined. — 1 specimens, USNM 40981 1.
Etymology of trivial name. — Named for the State of Oklahoma.
PARACONULARIA PLANICOSTATA (Dawson, 1868)
Figs. 3.1, 27.1-27.4, 21 .6-21
Conularia planicostata Dawson, 1868, p. 307-308, fig. 117; Dawson, 1878, p. 307-308,
fig. 117; Bigsby, 1878, p. 316; Dawson, 1883, p. 416; Lesley, 1889, p. 145, fig.;
Beede, 1911, p. 174, 186; Bell, 1929, p. 98-100, PI. 32, figs. 1-2; Bamber and
Copeland, 1976, PI. 15, fig. 3.
Conularia planocostata (sic) Dawson. Miller, 1877, p. 141; Miller, 1889, p. 390; Weller,
1898, p. 191.
Conularia quadrisulcata Miller in Sowerby. Sensu Dawson, 1889, p. 87, fig.
Conularia sorrocula Beede. Sensu Bell, 1929, p. 100, PL 32, figs. 3-3a.
Conularia cf. tenuis Slater. Sensu Bell, 1929, p. 100, PL 32, figs. 4-5.
Paraconularia planicostata (Dawson). Sinclair, 1948, p. 199-200; Babcock and Feld-
mann, 1984, p. 16-1 7; Babcock and Feldmann, 1986, fig. 2G.
Connularia (sic) planicostata Dawson. Alison and Carroll, 1972, p. 17.
Description.— FxosV.Q\QXon up to 8 cm in length. Major apical angle 21-25°; minor
apical angle 18-22°. Rod articulation inflected gothic arch style in apical region and
gothic arch style elsewhere. Rods abut or alternate at midline; if they alternate, rod
pattern is usually right superior; rod angle 1 1-16°. 12-20 rods/cm. Nodes and spines
absent. Apical wall not observed.
Occurrences. —Lowqx to Upper Mississippian of Nova Scotia and
Quebec; localities 163-173 and 248.
— Holotype, RM(MU) 2749, plastoholotype, GSC unnum-
bered.
Remarks.— Paraconularia planicostata (Dawson) is distinguished
major face. 23.4; FMNH UC 25175; small specimen preserving apical region; locality
12. 23.5-23.6; P. missouriensis (Swallow). 23.5; FMNH UC 1125; specimen preserved
in micrite showing darkened areas in the integument along the midline and surrounding
the ridges. Darkened areas of integument in the vicinity of the midline have been in-
terpreted by numerous authors as remains of original color markings. Specimen not
coated with ammonium chloride; locality 14. 23.6; USNM 14425, original of Walcott
(1884, PL 23, fig. 4), locality 110. 23.7; P. chesterensis (Worthen), ISGS 2489, same
specimen as in Fig. 23.1, detail of minor face. Bar scales represent 1 cm.
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1
Fig. 24. — 24.1; Pamconularia chesterensis (Worthen), USNM 50150, portion of large
cluster of individuals preserved in siltstone and showing incomplete remains of stalks
1986 Babcock and pELDMANN—P^^coAt/i^i?/^ and Reticulaconularia
431
from other members of the genus by the combination of: 1 , its small
size, generally less than 7.5 cm in length; 2, its inflected gothic arch
and gothic arch rod articulation styles; 3, its very narrow apical angles,
8-14®; 4, its widely spaced rods, 12-20 rods/cm; and 5, its lack of nodes
and spines. Paraconularia planicostata is particularly notable, and eas-
ily distinguished from all other taxa described herein because it pos-
sesses gothic arch rod articulation up to 7.5 cm from the hypothetical
apex in the adapertural direction.
Dawson (1868, p. 308), in describing the species Conularia plani-
costata, compared the taxon to an apparent manuscript species, C
novascotica Hartt. Dawson considered this taxon, also from the Mis-
sissippian of Nova Scotia, to be a variety of C planicostata. The in-
tended holotype specimen of C novascotica is lost, but based upon
Hartt’s scant description {in Dawson, 1868), it is likely to be an example
of P. planicostata (Dawson).
Material examined. — 30 specimens; housed in the CM, GSC, NYSM,
RM(MU), and the USNM.
PARA CONULARIA RECUR VA TUS
Babcock and Feldmann, new species
Figs. 32.3, 32.6
Description. — up to 8 cm in length. Major apical angle about 16°; minor
apical angle about 1 5°. Rod articulation exclusively of inflected circular curve style, greatly
recurved near midline in apical region and slightly recurved near midline elsewhere.
Rods abut or alternate at midline; if they alternate, pattern is usually left superior; rod
angle 8-12°, 18-28 rods/cm. 2-3 nodes/mm; spines absent. Apical wall not observed.
Occurrence.— UpptY Devonian of Nevada; locality 109.
—Holotype, part and counterpart, USNM 409806. Three
paratypes, USNM 409807-409809, all present on the same slab as the
holotype. The paratype labelled as USNM 409808 is preserved as part
and counterpart.
Remarks. —Paraconularia recurvatus Babcock and Feldmann, n. sp.
is unique among members of this genus in having rods which are both
closely spaced and which are recurved near the midline. Three other
species of Paraconularia examined in this study have recurved rods,
namely, P. alpenensis Babcock and Feldmann, n. sp., P. chesterensis
(arrows), attached to possible plant matter. Specimen not coated with ammonium chlo-
ride; locality 27. 24.2; Paraconularia byblis (White), USNM 409800, specimen preserved
in siderite concretion and showing a stalk (arrow); locality 7 1 . Specimen not coated with
ammonium chloride. 24.3; P. chesterensis (Worthen), USNM 50150, same specimen as
in Fig. 24.1, view showing the complete aggregation of conulariids as exposed at the
surface of the slab. Bar scales represent 1 cm.
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(Worthen) and P. missouriensis (Swallow). Paraconularia recurvatus
differs from all of these forms in rod spacing. Specimens in the type
lot of P. recurvatus have 18“28 rods/cm while the holotype of P. al-
penensis has 14 rods/cm, specimens of P. chesterensis have 8™25 rods/
cm and specimens of P. missouriensis have 6-10 rods/cm.
In its overall appearance, it seems as though P. recurvatus could be
mistakenly included in the genus Conularia. However, the lack of
spines indicates that this species should be included in the genus Para-
conularia. All quantitative data (Appendix B) further support this con-
clusion.
Material examined.— A specimens; housed in the USNM.
PARACONULARIA (Whiteaves, 1891)
Figs. 28.3, 28.5»28.6
Conularia salinensis Whiteaves, 1891, p. 244, PL 32, figs. 9-9a.
Conularia s.l. salinensis Whiteaves. Sinclair, 1948, p. 287.
Description. — I>Qscnption based only upon holotype. Exoskeleton 3 cm in length.
Major apical angle approximately 24®; minor apical angle 21®. Rod articulation inflected
gothic arch style in apical region and inflected circular curve style elsewhere. Rods usually
alternate at midline; if they alternate, pattern is usually right superior on major and
minor faces; rod angle 1 3® in apical region, 8® elsewhere. 24 rods/cm. 3-4 nodes/mm;
3-4 prominent adapertural spines/mm; adapical spines absent. Apical wall not observed.
Occurrence.— Mhsisuppmn of Alberta; locality 3.
-Holotype, GSC 4292.
Remarks.— This taxon can be distinguished from other species of
Paraconularia by the combination of: 1 , inflected gothic arch and in-
flected circular curve styles of rod articulation; 2, apical angles of 8-
13°; 3, rod spacing of 24 rods/cm; 4, node spacing of 3-4 nodes/mm;
and 5, prominent adapertural spines. Paraconularia salinensis (Whit-
eaves) does not seem to be easily confused with any other conulariid
species described to date from the Devonian or Mississippian rocks of
North America.
Material examined. — 1 specimen, GSC 4292.
Fig. 25.— Paraconularia missouriensis (Swallow). 25.1; FMNH UC 6639, plaster cast of
presumed holotype specimen; locality 100. 25.2; ISGS 2619; oblique view of specimen
with three intumed apertural terminations. The fourth apertural termination is broken
off, but there is no indication of infolding. 25.3; UMMP 26740, holotype of P. scioto-
villensis Driscoll, same specimen as in Fig. 26.1, detail of major face; locality 226. 25.4;
FMNH UC 6628, holotype of Conularia greenei Miller and Gurley; minor face of a
specimen preserved in micrite. 25.5; FMNH UC 6628, same specimen as in Fig. 25.4,
comer view. Bar scales represent 1 cm.
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Fig. 26.— Pamconularia missouriensis (Swallow). 26.1; UMMP 26740, holotype of P.
sciotovillensis Driscoll, minor face of a somewhat distorted individual preserved in a
1986 Babcock and FELDMANN—P^i?^ and Reticulaconularia
435
PARACONULARIA SORROCULA (Beede, 1911)
Figs. 28.1-28.2
Conularia sorrocula Beede, 191 1, p. 184, 186, 2 figs.
Paraconularia sorrocula Beede. Sinclair, 1948, p. 199.
Description.— Exoskeleton up to 3 cm in length. Major apical angle 19-24°; minor
apical angle 1 7-22°. Rod articulation exclusively inflected gothic arch style with slight
adapertural inflection at midline. Rods usually alternate at midline, less frequently they
abut; rods, if they alternate, pattern is usually left superior on both major and minor
faces; rod angle 11-14°. 18-20 rods/cm. Nodes elongate and appear to be continuous
structures with adapertural spines; 5-6 nodes/mm; 5-6 adapertural spines/cm. Adapical
spines absent. Apical wall not observed.
Occurrence. —Mississippisin of Quebec; locality 247.
Typp. — Holotype, part and counterpart, NYSM 9414.
Remarks. —Paraconularia sorrocula (Beede) is unique among North
American Devonian or Mississippian examples of Paraconularia in
the possession of only inflected gothic arch rod articulation. It is also
the only conulariid species reported herein which has the nodes merged
with the adapertural spines without a significant change at the junction
of the two structures (Fig. 28.2). The nodes are not round or oblate in
outline as in other species of Paraconularia, but are elongate.
The holotype of P. sorrocula is curved in the apertural region. This
feature may have been present on the specimen in life, although this
cannot be confirmed owing to the crushed nature of the fossil.
Material examined. — specimens; housed in the NYSM.
PARACONULARIA SUBULATA (Hall, 1858)
Figs. 3.2, 3.5-3.6, 21.4, 29.1-29.10,
30.1-30.8, 31.1-31.5, 33.4
Conularia subulata Hall, 1858, p. 32; Miller, 1877, p, 141; Bigsby, 1878, p. 316; Whit-
field, 1882, p. 91, PI. 8, fig. 3; Hall, 1883, p. 372-373, PI. 31, fig. 3; Miller, 1889,
p. 390; Lesley, 1889, p. 146, fig.; Lesley, 1895, p. 1690, fig.; Weller, 1898, p. 192;
Whitfield and Hovey, 1901, p. 406-407; Cumings, 1906, p. 1366, PL 25, fig. 3.
Conularia victa White, 1862, p. 22-23; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Miller,
1889, p. 390; Herrick, 1893, PI. 19, fig. 3; Weller, 1898, p. 192.
Conularia newberryi Winchell, 1865, p. 130; Winchell, 1870, p. 258; Meek, 1875, p.
316-317, PL 18, fig. 2a-b; Miller, 1877, p. 141; Bigsby, 1878, p. 316; Hall, 1879,
PL 34A, fig. 12; Herrick, 1888^, p. 93-94, PL 6, figs. 13, 17, PL 8, fig. 9; Herrick,
siderite concretion; locality 226. 26.2; UMMP 26740, same specimen as in Fig. 26.1,
major face. 26.3; AMNH 28692, minor face of a specimen preserved in sparry limestone;
locality 30. 26.4; FMNH UC 6627, holotype of Conularia gratiosa Miller and Gurley,
preserved in micritic limestone, comer view; locality 30. 25.5; FMNH UC 6627, same
specimen as in Fig. 26.4, detailed view of a minor face. Bar scales represent 1 cm.
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1888Z?, PI. 8, fig. 5, PI. 10, figs. 27-28; Lesley, 1889, p. xv; Miller, 1889, p. 390;
Herrick, 1893, PL 19, fig. 5; Miller, 1897, p. 765; Weller, 1898, p. 191; Clarke and
Ruedemann, 1903, p. 566; Grabau and Shimer, 1910, p. 13, figs. 1227a-b; Tasch,
1973, fig. 5.16, Table 5.2; Tasch, 1980, fig. 5.16, Table 5.2.
Conularia whitei Meek and Worthen, 1865, p. 253-254; Bigsby, 1878, p. 316.
Conularia newberryi WinchelYI Herrick, 1887, p. 146-147, PI. 14, fig, 14.
Conularia victa White? Herrick, 1888Z), p. 47-48, PL 8, fig. 3.
Conularia whitii (sic) Miller, 1889, p. 390; Weller, 1898, p. 192.
Conularia sampsoni Miller, 1892<2, p. 690-691, PL 14, figs. 11-12; Miller, 1892^, p.
692; Holm, 1893, p. 125; Weller, 1898, p. 191; Branson, 1938, p. 110-111, PL 14,
fig. 9; Branson, 1944, p. 216.
Paraconularia subulata (Hall). Sinclair, 1948, p. 198; Babcock and Feldmann, 1986, figs.
IC-D, IG, 2F, 3A-C.
Paraconularia victa (White). Sinclair, 1948, p. 200.
Paraconularia newberryi (Winchell). Sinclair, 1948, p. 191-192; Driscoll, 1963, p. 34-
37, PL 1, figs. 1-5, PL 2, figs. 1-4.
Paraconularia whitei (Meek and Worthen). Sinclair, 1948, p. 192.
Paraconularia sampsoni Miller. Sinclair, 1948, p. 197.
Paraconularia sp. Feldmann, Coogan and Heimlich, 1977, fig. 2.50A.
""Conularia"" sp. Thompson, 1982, fig. 357.
Paraconularia cf. P. missouriensis {SwbWow). Sensu Babcock, 1985^z, figs. la-b.
Paraconularia missouriensis (Swallow). Sensu Babcock, 1985^2, fig, 2.
Paraconularia cf. P. subulata. Babcock and Feldmann, 1986, fig. 2H.
Description. --ExoskQlQXovi up to 17 cm in length. Major apical angle 17-22®; minor
apical angle 12-18°. Rod articulation inflected gothic arch style in apertural region and
inflected circular curve style elsewhere. Rods usually alternate at midline; if they alternate,
pattern is usually right superior on major face and usually left superior on minor face;
rod angle 15-18®. 20-35 rods/cm. Nodes absent or present; if present, they are incon-
spicuous, 2-3/mm; spines absent. Apical wall present.
Occurrences. --1.0SNQV Mississippian of Illinois, Indiana, Kentucky,
Montana, Ohio; localities 16, 26, 29, 38, 39, 72, 75, 82, 106-107, 185-
192, 194, 196-203, 205-208, 212-217, 220-222.
— Lectotype, designated herein from James Hall’s suite of three
syn types of C. subulata, AMNH 32403, smaller of two specimens
bearing this number; two paratypes, AMNH 32403, larger of two spec-
imens bearing this number, and AMNH 32404. Holotype of C victa,
Fig. 27.-27.1-27.4; Paraconularia planicostata (Dawson). 27,1; RM(MU) 2749, ho-
lotype; major face, locality 164. 27.2; RM(MU) 2749, same specimen as in Fig. 27.1,
comer view. Note exoskeletal constrictions. 27.3; RM(MU) 2749, same specimen as in
Fig. 27.1, minor face. 27.4; RM(MU) 2749, same specimen as in Fig. 27.1, detail of
major face. 27.5; P. oklahomaensis Babcock and Feldmann, n. sp., USNM 409801,
holotype, a flattened individual. 27.6-27.8; P. planicostata (Dawson). 27.6; CM 22667,
major face; locality 168. 27.7; CM 22667, same specimen as in Fig. 27.6, minor face.
27.8; CM 22667, same specimen as in Fig. 27.6, comer view. Bar scales in Figs. 27.1-
27.5 represent 1 cm; bar scales in Figs. 27-6-27.8 represent 5 mm.
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Fig. 28. —28. 1-28.2; Paraconularia sorrocula (Beede). 28.1; NYSM 9414, slab showing
two specimens, holotype to the right. Note orbiculoid brachiopods attached to, and
1986 Babcock and Feudmann—Paraconularia and Reticulaconularia
439
UMMP 2178, plastoholotype, GSC unnumbered; holotype of C new-
berryi, UMMP 245; holotype of C whitei, UIPC 10866; holotype of
C sampsoni, FMNH UC 6961, plastoholotype, USNM 68156.
Remarks.— Par aconularia subulata (Hall) is most similar in mor-
phology to P. oklahomaensis Babcock and Feldmann, n. sp. Both taxa
have inflected gothic arch rod articulation in the apical region and
inflected circular curve rod articulation elsewhere. Also, specimens
referable to both taxa may lack nodes and spines. Except for rod spacing
values, the quantitative measures are also very similar. Paraconularia
subulata has a rod spacing of about 20-35 rods/cm whereas the ho-
lotype of P. oklahomaensis has a value of 1 2-24 rods/cm. Paraconu-
laria subulata differs most substantially from P. oklahomaensis in hav-
ing very little infleetion of the rods in the vicinity of the apex and in
having a strong inflection of the rods elsewhere.
In addition to P. oklahomaensis, P. blairi (Miller and Gurley) bears
close similarity to P. subulata. Both taxa have inflected gothie arch rod
articulation adapically and inflected circular curve rod articulation ad-
aperturally. Apical angles and rod angles are nearly equal in the two
forms. Paraconularia subulata differs from P. blairi in its lack of nodes
on the rods or in having inconspicuous nodes, in the lack of any ad-
apertural inflection of the rods near the midline and in the possession
of a greater number of rods/cm, 20-35 as compared to 6-8.
The species P. victa (White), P. newberryi (Winchell), P. whitei and
P. sampsoni (Miller) are all included as junior synonyms of P. subulata
because they are indistinguishable from the lectotype and paralecto-
types of P. subulata. The type specimens of all these species bear subtle
nodes on the rods; all have apical angles in the range of 12-22®; all
have inflected gothic arch styles of rod articulation; all have 20-35
rods/cm; and all have rod angles of 1 5-18®. The holotype of P. sampsoni
possibly could be construed as a juvenile of P. blairi, but the lack of
rods which are slightly inflected at the midline makes assignment of
this specimen to P. subulata more reasonable.
Paraconularia subulata is one of the most abundant conulariids in
located near, the conulariids; locality 247. 28.2; NYSM 9414, same specimen as in Fig.
28.1, detail of holotype. 28.3; P. salinensis (Whiteaves), GSC 4292, holotype, minor
face; locality 3. 28.4; P. sp., CM 34531, a collapsed and poorly preserved specimen in
tan and dark red colored dolostone; locality 249. 28.5-2^6; P. salinensis (Whiteaves).
28.5; GSC 4292, same specimen as in Fig. 28.3, comer view. 28.6; GSC 4292, same
specimen as in Fig. 28.3, detail of minor face. 28.7; P. yochelsoni Babcock and Feldmann,
n. sp., external molds of two specimens attached to plant matter, holotype, UMMP
45499, to the right, paratype, UMMP 65509, to the left; locality 93. Specimen is not
coated with ammonium chloride. Bar scales represent 1 cm.
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Fig. 29.— Par aconularia subulata (Hall). 29. 1; AMNH 32404 (smaller of two specimens),
lectotype, minor face; locality 16. 29.2; UMMP 245, holotype of Conularia newberryi
1986 Babcock and Feldmann— and Reticulaconularia
441
the Lower Mississippian of the North American midcontinent. It is
often confused with other taxa, especially P. missouriensis, in museum
collections. The reason for this confusion is not clear. Some specimens
of this taxon, which superficially appear very similar to specimens of
P. biairi, may have been confused with P. missouriensis because of
ambiguity in Swallow’s original description of the latter. It is likely that
Swallow’s description was based upon specimens now referable to both
P. missouriensis and P. biairi.
Conulariids collected from the Bear Gulch Limestone of Montana
(CM 34507--34527 and 35000) and from the Cameron Creek Shale of
Montana (USNM 118731) are here assigned to P. subulata with little
reservation. The specimens differ from the type series of P. subulata
only in the uniform lack of nodes on the rods. However, samples of
many specimens referable to P. subulata from Illinois, Ohio, and else-
where indicate that nodes are frequently lacking in this taxon. Even
when nodes are present on such specimens, they are subtle.
Material examined. — I A9 specimens; housed in the AMNH, CM,
CMNH, FMNH, GSC, OC, UMMP, USNM, and the private collection
of Ron Fisher.
PARACONULARIA WELLSVILLIA
Babcock and Feldmann, new species
Figs. 33.3, 33.6-^33.8
Paraconularia sp. Babcock and Feldmann, 1986, fig. 2J.
Description. —Exosk&lQlon up to 13 cm in length. Major apical angle 14-18°; minor
apical angle 12-15°. Rod articulation gothic arch style in apical region and inflected
gothic arch style elsewhere; rods almost always alternate at the midline; if they alternate,
pattern is usually left superior on major face and usually right superior on minor face;
rod angle 26-31°. 4-5 rods/cm; 2-3 nodes/mm; 2-3 adapertural spines/mm; adapical
spines appear not to be present. Apical wall not observed.
Occurrence. Devonian of New York; localities 161“162,
Holotype, CM 35001; 12 paratypes, CM 34538-34550.
Remarks. —Paraconularia wellsvillia Babcock and Feldmann, n. sp.
Winchell, major face; locality 206. 29.3; UMMP 245, same specimen as in Fig. 29.2,
minor face. 29.4; AMNH 32404 (smaller of two specimens), same specimen as in Fig.
29.1, major face. The pitted material attached to the specimen is glue. 29.5; CMNH
5988, comer view of large, partially compressed specimen. Note healed injury near top
of minor face; locality 198. 29.6; UMMP 2178, holotype of C. victa White, ?minor face;
locality 63. 29.7; UMMP 2178, same specimen as in Fig. 29.6, comer view. 29.8; FMNH
UC 6961, holotype of C. sampsoni Miller, minor face; locality 94. 29.9; FMNH UC
6961, same specimen as in Fig. 29.8, comer view. 29.10; FMNH UC 6961, same spec-
imen as in Fig. 29.8, major face. Bar scale in Fig. 29.5 represents 1 cm; bar scales in
Figs. 29.1-29.4 and 29.6-29.10 represent 5 mm.
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is distinguished from all other species of Paraconularia known from
the Devonian and Mississippian rocks of North America in having
gothic arch rod articulation in the apical region and inflected gothic
arch rod articulation elsewhere. Superficially, this taxon resembles P.
yochelsoni in the general pattern of rod articulation near the aperture.
P. wellsvillia, however, possesses nodes and adapertural spines whereas
P. yochelsoni does not.
Of the 13 specimens examined and referred to P. wellsvillia, only
the holotype, CM 35001, shows well preserved nodes and spines (Figs.
33.7-33.8). Others, such as the specimen illustrated in Fig. 33.6, seem
to lack these structures. After examination of the holotype and 12
paratypes, it seems that two factors affect these profound preservational
differences: 1, degree to which the integument is fit around the rods,
nodes, and spines; and 2, type of lithology in which the specimen is
preserved. The Wellsville Formation, from which all specimens in the
type suite were collected, varies from a fine grained silty sandstone to
a micaceous siltstone to a micaceous shale. Preservation of a conulariid
tends to be better in a fine grained matrix.
The holotype of P. wellsvillia shows a wrinkling of the integument
only partially related to the pattern of nodes and spines (Fig. 33.8).
This wrinkling is also attributed, in part, to a tight fitting of the integ-
ument about the framework of the exoskeleton and slight displacement
of the framework.
Material examined. — 13 specimens; housed in the CM.
Fig. 30. —Paraconularia subulata (Hall). 30.1; USNM 409802, minor face of specimen
preserved in phosphatic concretion; locality 72. 30.2-30,3; USNM 395828, right and
left halves of a specimen preserved in a phosphatic concretion and showing internal soft-
parts; locality 72. 30.4; USNM 409802, same specimen as in Fig. 30.1, view from apical
end showing rounded cross section of soft-parts (arrow). 30.5; USNM 409802, same
specimen as in Fig. 30.1, x-ray photograph of specimen preserved in a phosphatic con-
cretion. The photograph was obtained using a Hewlett-Packard Faxitron Series x-ray
unit located in the Department of Anthropology, Kent State University. The specimen
is in the same orientation as in Fig. 30. 1 . Presumed internal soft-parts appear as a single,
elongate tube. 30.6; USNM 409803, specimen preserved in a phosphatic concretion with
some of the exoskeleton broken away, revealing remains of limonite-coated internal soft-
parts. 30.7; USNM 409804, two specimens preserved in same orientation in siltstone
block. Sole marks on reverse side of slab parallel the orientation of the conulariids and
indicate that these specimens have been current aligned; locality 220. 30.8; UIPC 10866,
holotype of Conularia whitei Meek and Worthen preserved in siderite; locality 20 1 . Bar
scales in Figs. 30.1-30.3 and 30.5-30.8 represent 1 cm; bar scale in Fig. 30.4 represents
5 mm. Specimens in Figs. 30.2-30.4 and 30.6 have not been coated with ammonium
chloride.
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Etymology of trivial we. —Named for the Wellsville Formation,
from which the holotype specimen was collected.
PARACONULARIA YOCHELSONI
Babcock and Feldmann, new species
Figs. 28.7, 33.1-33.2, 33.5
Paraconularia newberryi (Winchell). Sensu Driscoll, 1963, p. 34-40, PL 1, figs. 6-8.
Description. —Exoskeleton up to 3.5 cm in length. Major apical angle 17-20®; minor
apical angle approximately 18°. Rod articulation gothic arch style in earliest stages,
inflected gothic arch style in later stages. Rods usually alternate at midline; if they
alternate, pattern is usually right superior on both major and minor faces; rod angle 1 5-
20°. 13-18 rods/cm. Nodes appear to be absent; spines absent. Apical wall present.
Occurrence. —Lower Mississippian of Michigan; locality 93.
Types. — Holotype, UMMP 45499; two paratypes, UMMP 65509 on
the same slab as UMMP 45499, and UMMP 45500.
Remarks. —Paraconularia yochelsoni Babcock and Feldmann, n. sp.
is only similar in morphology to P. alternistriata (Shimer). Both species
seem to be less than 3.5 cm in maximum length, lack nodes on the
rods and have rod articulation patterns which appear to be similar.
Paraconularia yochelsoni, however, has larger apical angles, 1 5-20° as
compared to 10-1 1°, and fewer rods/cm, 13-15 as compared to 28.
The holotype, UMMP 45499, and paratype, UMMP 65509, speci-
mens of P. yochelsoni, are preserved as three dimensional specimens
lacking the integument except along the midline. The midline may
have been thickened in this taxon.
The holotype and paratype of P. yochelsoni are located on the same
slab as a large portion of black, carbonaceous matter composed largely
of densely packed, filamentous strands (Fig. 28.7). This material prob-
ably represents plant matter of some sort, perhaps a planktonic alga.
The two conulariids appear to be attached to the presumed plant matter
by stalks extending from their apices; only small traces of the stalks
remain in place. The two conulariids on this slab are radiating away
from the center of the dark mass.
Material examined. — 3 specimens; housed in the UMMP.
Etymology of trivial ^aw^.— Named for Ellis L. Yochelson, a dis-
tinguished student of problematic fossils.
Fig. 31.— Paraconularia subulata (Hall). 31.1; CM 34524, preserved in micrite; locality
106. 31.2; CM 35000, preserved in micrite; locality 106. 31.3; CM 34521, preserved in
micrite, locality 106. 31.4; NYSM 3491, preserved in siderite; locality 203. Original of
Hall’s ^Eonularia newberryE (1879, PI. 34A, fig. 12). 31.5; USNM 118731; locality 107.
Bar scales represent 1 cm.
VOL. 55
446
Annals of Carnegie Museum
Fig. 32. — 32.1; Paraconularia missouriensis (Swallow)?, GSC 85062; locality 5. 32.2; P.
sp., USNM 409805, fragment of specimen lacking integument preserved in soft blue-
1986 Babcock and FEi^uMAnN—PARACONULARiA and Reticulaconularia
447
PARACONULARIA sp.
Fig. 28.4
Occurrence.— Upper Devonian-Lower Mississippian transition of
South Dakota; locality 27.4.
Figured specimen. — CM 3453 1 .
Remarks.— A single specimen of conulariid was collected from the
Englewood Formation at Deadwood, South Dakota. It can be reliably
identified only to the genus level.
The specimen is badly crushed and incomplete, making it possible
to perform few qualitative or quantitative observations. The specimen
in question possesses 7 rods/cm and lacks nodes and spines; it is there-
fore referred to the genus Paraconularia. The specimen seems to have
a gothic arch style of rod articulation in the apical region and an in-
flected circular curve style elsewhere.
Species of Paraconularia examined in this work which may possess
7 rods/cm include P. blairi (Miller and Gurley) and P. missouriensis
(Swallow); P. chesterensis (Worthen) can have as few as 8 rods/cm and
P. wellsvillia Babcock and Feldmann, n. sp. has 4-5 rods/cm. Of these,
CM 34531 appears to be most similar to P. blairi or P. subulata (Hall)
in terms of rod articulation.
Material examined. — I specimen, CM 34531.
RETICULACONULARIA Babcock and Feldmann, new genus
— Conulariids with rods that are widely spaced, 12-39/
cm. 30-80% of rods alternate at midline; 20-70% abut. Apical angles
large, 22-59°. Nodes and adapertural spines present and widely spaced;
adapical spines not known.
Type species. — Conularia penouili Clarke, 1907; Lower Devonian of
Quebec. Holotype, NYSM 9412.
Remarks. — Species referable to Reticulaconularia differ from all oth-
er conulariids in having very large apical angles, 22-59° in the speci-
mens measured in this study. The wide spacing between adjacent rods
and between nodes, as well as between adapertural spines is also unique
to species of this genus. In specimens retaining the external surface of
the integument, this pattern of rods, nodes, and spines gives the exo-
gray calcareous shale; locality 177. 32.3; P. recurvatus Babcock and Feldmann, n. sp.,
USNM 409806. Holotype to right; a paratype (USNM 409807) is indicated by arrow;
locality 109. 32.4; P. byblis (White)?, UMMP 26735, a poorly preserved, collapsed
specimen; locality 250. Ori^nal of Winchell (1871, p. 257). 32.5; P. chesterensis (Wor-
then)?, GSC 49383, a juvenile specimen with stalk preserved. Specimen not coated with
ammonium chloride; locality 7. 32.6; P. recurvatus Bdhcock and Feldmann, n.sp., USNM
409806, same specimen as in Fig. 32.3, detail of holotype. Bar scales represent 1 cm.
448
Annals of Carnegie Museum
VOL. 55
1986 Babcock and Feldmann — Paraconularia and Reticulaconularia
449
skeleton a somewhat reticulate pattern. The genus is named for this
characteristic.
At present, three species of conulariids are referred with certainty to
Reticulaconularia: C. penouili from the Lower Devonian of Quebec,
C. sussexensis Herpers from the Lower Devonian of New Jersey, and
C baini Ulrich from the Devonian of Bolivia. This third species occurs
outside the geographic limits of this paper; we and others will redescribe
the taxon in a paper on the Devonian conulariids of Bolivia.
RETICULACONULARIA PiE^A^OU/L/ (Clarke, 1907)
Figs. 34.3-34.5; 35.3
Conularia penouili Clarke, 1907, p. 180--181, 2 figs.; Clarke, 1908, p. 144, PL 11, figs.
10-=11.
Conularia gaspesia Sinclair, 1942, p. 15 8-- 160, fig.
Conularia s.l. penouili Clarke. Sinclair, 1948, p. 283.
Description. —ExoskQlQton a curved pyramid, expanding slowly and non-uniformly
from the apex. Exoskeleton up to 6.7 cm in length. Major apical angle approximately
59°; minor apical angle approximately 22-30°. Rod articulation of inflected circular curve
style in apical ‘A, angulated circular curve style in middle Vs, and inflected circular curve
style in apertural '/a. Rods usually alternate at midline; if they alternate, pattern is usually
left superior on both major and minor faces; rod angle 0-8°. 12-21 rods/cm. 1-2 nodes/
mm. 1-2 adapertural spines/cm; adapical spines probably not present. Interridge furrows
broadly rounded into longitudinally oblong pits. Apical wall not observed.
Occurrence. — Lowqy Devonian of Quebec; localities 228 and 246.
Type'y. —Holotype, NYSM 9412; topotype of Conularia gaspesia,
GSC 87242.
Remarks. —Reticulaconularia penouili (Clarke) is distinguished from
R. sussexensis (Herpers) by the following features. First, R. penouili
has 12-21 rods/cm and R. sussexensis has 11-14 rods/cm. Greater rod
spacing in R. penouili, combined with wide spacing between the nodes
and between the adapical spines has produced large, oblong, hollowed
Fig. 33,-33. 1-33.2, Paraconularia yochelsoni Babcock and Feldmann, n. sp. 33, 1 ; UMMP
45499, holotype; comer view; locality 93. 33.2; UMMP 45499, same specimen as in
Fig. 33.1, major face. 33.3; P. wellsvillia Babcock and Feldmann, n. sp., CM 34502,
paratype, preserved in siltstone; ?minor face; locality 161. 33.4; P. subulata (Hall), KSU
1 172, cross section showing weakly bilateral, four-sided nature of the exoskeleton. Spec-
imen not coated with ammonium chloride; locality 216. 33.5; P. yochelsoni Babcock
and Feldmann, n, sp, UMMP 45500, paratype; locality 93. 33.6-33.8; P. wellsvillia
Babcock and Feldmann, n. sp. 33.6; CM 34503, detail of paratype preserved as an internal
mold in siltstone and not exhibiting interrod ridges or interrod furrows; locality 162.
33.7; CM 35001, holotype, a collapsed specimen preserved in silty shale; locality 161.
33.8; CM 3500 1 , same specimen as in Fig. 33.7, detail of a minor face. Bar scales represent
1 cm.
450
Annals of Carnegie Museum
VOL, 55
Fig, 34.-34. 1-34.2; Reticulaconulariasussexensisi^QrpQT^). 34. 1; NJSM 10806, smaller
of two specimens, major face of small specimen preserved in somewhat metamorphosed
1986 Babcock and Yeldmann—Paraconularia and Reticulaconularia
451
out interridge furrows on the exterior surface of the exoskeleton. This
feature is not exhibited, to this extent, on any other known species of
conulariid.
Secondly, the values obtained for rod angles are consistently smaller
for R. penouili than they are for R. sussexensis, 0-8° compared to 1 1-
14°. Reticulaconularia sussexensis exhibits a greater number of rods
which abut at the midline than does R. penouili. Up to 70% of rods
abut in specimens of R. sussexensis while 20-30% of rods abut in the
holotype of R. penouili. Unlike R. sussexensis, R. penouili shows an
alternation between a circular curve style of rod articulation and an
angulated circular curve style. Reticulaconularia sussexensis exhibits
only an inflected gothic arch style of rod articulation. Finally, judging
from the available sample, specimens of R. sussexensis seem to have
a smaller maximum length than R. penouili. The maximum recorded
hypothetical length of a specimen of R. sussexensis is about 2.5 cm.
The hypothetical length of the holotype of R. penouili is 6.7 cm.
The holotype of Conularia gaspesia Sinclair, which was said to have
been deposited in the RM(MU) (Sinclair, 1942, p. 160) apparently
never was deposited in that museum (Ingrid Birker, written commu-
nication, 1985), and is now presumed to be lost. However, we have
found, in the Sinclair collection housed in the GSC, a specimen labelled
as C. gaspesia (GSC 87242; Fig. 35.3). The handwriting on a label
accompanying the specimen is unmistakably that of G. Winston Sin-
clair. This specimen is of further value because a label glued to the
specimen indicates that it was collected from Lower Devonian Grande
Greve Limestone on the Gaspe Peninsula; thus, the specimen is a
topotype.
This topotype specimen of C. gaspesia, GSC 87242, exhibits one
well preserved face and has all of the salient morphological character-
istics that the holotype of R. penouili posseses. Among the character-
istics shown by the topotype of C. gaspesia are large, oblong, hollowed
out interridge furrows, just as are present in the holotype of R. penouili.
A good photograph of the holotype of C. gaspesia (Sinclair, 1 940, fig.)
shows these same features. Measurements taken on the topotype spec-
imen are given in Appendix B. The only way in which GSC 87242
differs from the holotype of R. penouili, NYSM 9412, is that it is not
siltstone; locality 1 12, 34,2; NJSM 10806, same specimen as in Fig, 34.1, comer view.
Note prominent exoskeletal constriction. 34. 3-34.5; R. penouili (Clarke). 34.3; NYSM
9412, holotype, detail of minor face; locality 246. 34.4; I^SM 9412, same specimen as
in Fig. 34,3, minor face of specimen preserved in micrite. 34.5; NYSM 9412, same
specimen as in Fig. 34,3, major face. Bar scales represent 1 cm.
452
Annals of Carnegie Museum
VOL. 55
Fig. 35.-35.1-35.2. Reticulaconularia smsexensis 35.1; NJSM 10750, exter-
nal mold of apical region of very small paratype specimen; locality 112. 35.2; NJSM
10751, largest of three specimens, paratype, preserved as an external mold; locality 1 12.
35.3; R. penouiii (Clarke), GSC 87242, ?minor face of topotype of Conularia gaspesia
1986 Babcock and Felumann —Paraconularia and Reticulaconularia
453
curved in the apical region. However, NYSM is a somewhat crushed
or collapsed individual. One final interesting point is that both NYSM
9412 and GSC unassigned were collected from the Grande Greve Lime-
stone on the Gaspe Peninsula. Based upon this topotype specimen of
C gaspesia, this taxon is here placed in synonymy with R. penouili.
Material examined.— 2 specimens; housed in the GSC and the NYSM.
RETICULACONULARIA SUSSEXENSIS {ULqvpqxs, 1949)
Figs. 34.1-34.2, 35.1-35.2, 35.4
Conularia sussexensis Herpers, 1949, p, 1-7, PL 1, 2.
Conularia gaspesia Sinclair. Sensu Herpers, 1950, p. 619.
Description. —FxoskeXeXon up to 2.5 cm in length. Major apical angle 24-28°; minor
apical angle 22-25°. Rod articulation uniformly of inflected gothic arch style. Rods usually
abut at midline; if they alternate, they usually occur left superior on major face and right
superior on minor face; rod angle 11-14°. Approximately 39 rods/cm (extrapolated) in
apical region; 14-18 rods/cm elsewhere. 2 nodes/mm; 2 adapertural rods/mm; adapical
spines absent. Apical wall not observed.
Holotype, NJSM 10749; four paratypes,NJSM 10750, NJSM
10751 (three specimens).
Occurrences. — Lowqt Devonian of New Jersey; localities 111, 112,
113.
Remarks.— ThQ present samples of R. sussexensis (Herpers) differ
from the holotype of R. penouili (Clarke) in that the former: 1, is of
smaller size; 2, has no curvature to the exoskeleton; 3, has smaller
apical angles, especially on the major face; 4, has a greater number of
rods/cm; 5, has larger rod angles; 6, shows greater than 30% of the
rods abutting at the midline; and 7, shows only an inflected gothic arch
style of rod articulation. These differences are discussed more fully in
the remarks accompanying the description of R. penouili, above.
Material examined. — \ 0 specimens; housed in the NJSM.
Organisms Previously Assigned to Conulariida,
Here Rejected from the Phylum
Phylum Mollusca
Class Hyolitha
HYOLITHES sp.
Fig. 17.4
Conularia sp. Ellison, 1965, p. 48-49, PL 4, fig. 1.
Hyolithes sp. Babcock, 1985^, p. 14-16, fig. 1.
Sinclair, preserved in micritic limestone; locality 242. 35.4; R. sussexensis (Herpers),
NJSM 10749, ?major face of holotype, preserved in somewhat metamorphosed siltstone;
locality 112. Bar scales represent 1 cm.
454
Annals of Carnegie Museum
VOL. 55
Fig. 36. — 36.1-36.3; ^'‘Conularia" tenuicostata Branson, here interpreted to be a ?pri-
apulid worm. 36.1; UMC 4271, holotype; locality 99. 36,2; UMC 4271, detail of same
1986 Babcock and Feldmann—P^/?^ and Reticulaconularia
455
Occurrence.— Middle Devonian of Pennsylvania; locality 240.
Figured sped men. — U SNM 173928.
Remarks. —Ellison’s figured specimen (1965, plate 4, fig. 1) is a small
conical shell expanding slowly and uniformly from a bluntly rounded
apex. The specimen possesses thin, closely spaced, raised lines, con-
centric about the apex. Crushing has produced a long, irregular line
down the middle of the shell. A ligula, or an apertural extension of the
shell on the dorsal side, is present, clearly indicating that the specimen
is a hyolith, not a conulariid.
Phylum Priapulida?
Figs. 36.1-36.3
Conularia tenuicostata Branson, 1938, p. 1 1 1, PL 14, figs. 5-6; Branson, 1944, p, 216.
Mesoconularia tenuicostata (Branson). Sinclair, 1948, p. 128.
Occurrence. — Fowtr Mississippian of Missouri; locality 99.
Type. -Holotype, UMC 4271.
Remarks.— The holotype, and only known specimen of Conularia
tenuicostata Branson, possesses tiny nodes or pustules arranged in closely
spaced rows which appear to run essentially perpendicular to the long
axis of the fossil. The rows of nodes or pustules are not supported by
calcium phosphate rods. Additionally, the specimen is flattened and
micrite replaced. This type of preservation is unlike that expected of
an animal composed of calcium phosphate, such as a conulariid. Rath-
er, the preservation is similar in appearance to the preservation of
objects having a tough cuticle, such as Plectodiscus discoideus (Rauff),
a chondrophorine cnidarian from the Hunsriick Slate (Devonian) of
West Germany (Yochelson et aL, 1983).
Branson’s specimen possesses indiscrete ringlike segments delimited
by thin, latitudinally arranged crests 0.5 to 0.7 mm apart. These seg-
ments are each covered with closely spaced, latitudinally arranged rows
of minute papillae which are strikingly similar to the cuticle of living
priapulid worms such as Priapulus and Tubiluchus. However, there is
not enough of the holotype preserved to determine whether the animal
possessed spines, a common feature of living priapulids. Thus, this
fossil is referred to the phylum Priapulida with reservation.
The specimen in question preserves only a small portion of cuticle,
specimen as in Fig. 36.1, showing ridges delimiting annular segments. 36.3; detail of
same specimen as in Fig. 36.1, showing surface structure. 36.4-36.5; Oracanthus sp.
36.4; USNM 409810; locality 6. 36.5; USNM 409810; detail of specimen in Fig. 36.5,
interpreted to be remains of fish spine. Bar scales in Figs. 36.1 and 36.4-36.5 represent
1 cm; bar scales in Figs. 36.2 and 36.3 represent 1 mm.
456
Annals of Carnegie Museum
VOL. 55
7.5 mm long and 8.1 mm wide. It is broken at its upper and lower
margins, presumably between adjacent segments. This mode of pres-
ervation supports the interpretation that the specimen possessed a
multielement covering, capable of readily fragmenting or tearing.
Phylum Chordata
Class Vertebrata
Order Pisces
ORACANTHUS sp.
Figs. 36.4-36.5
Conularia newberryi Winchell? Sensu McKee and Gutschick in McKee and Gutschick,
1969, p. 125-172.
Occurrence.— Mississippian of Arizona; locality 5.
Figured specimen. — External mold preserved in dolostone and latex
mold, USNM 409810.
Remarks.— This figured specimen mimics a conulariid in having
nodose structures arranged in rows, crossing the surface transversely.
The rows, however, are discontinuous, and form chevron-shaped pat-
terns in some places. The rows seem to be composed of semidiscrete
pits arranged in side-by-side fashion. As the specimen is an external
mold, the “pits” would correspond to nodes. These is no evidence on
the specimen of either a midline or a comer groove. The specimen is
undulated near the left margin, however.
This specimen is here regarded as the spine of a gyracanthid shark.
Michael E. Williams, of the Cleveland Museum of Natural History,
has viewed this specimen and has noted that small portions of dentine
adhere to it in places, confirming that it is a vertebrate fossil. It is his
suggestion that this specimen be referred to the genus Or acanthus.
Acknowledgments
Parts A and B of this work are based upon Babcock’s M.S. thesis, completed at Kent
State University. The study was suggested by Ellis L. Yochelson, U.S. National Museum
of Natural History, Washington, D.C. Alan H. Coogan and Barry B. Miller read various
drafts of this paper. Others who have assisted in this study through collecting specimens,
loaning specimens, providing reference materials or aiding with electron microprobe or
x-ray analyses include the following: Donald Baird, Princeton University; Gorden C.
Baird, SUNY College at Fredonia; Roger L. Battin, American Museum of Natural His-
tory; Gordon L. Bell, Jr., Red Mountain Museum; Ingrid Birker, Redpath Museum
(McGill University); Daniel B. Blake, University of Illinois at Urbana-Champaign; Thomas
H. Bolton, Geological Survey of Canada; Arthur J. Boucot, Oregon State University;
Ernest H. Carlson, Kent State University; Robert L. Carroll, Redpath Museum (McGill
University); John L, Carter, Carnegie Museum of Natural History; Mitchell J. Ciccarone,
Canton, Ohio; Frederick J. Collier, United States National Museum of Natural History;
G. Arthur Cooper, United States National Museum of Natural History; Murray J. Cope-
land, Geological Survey of Canada; Roger J. Cuffey, Pennsylvania State University; Larry
Decina, Drexel Hill, Pennsylvania; Ding Baoliang, Nanjing Institute of Geology and
1986 Babcock and Felumann —Paraconularia and Reticulaconularia
457
Mineral Resources, Nanjing, People’s Republic of China; Ruth L. Elder, Oberlin College;
Niles Eldredge, American Museum of Natural History; Frank R. Ettensohn, University
of Kentucky; David F. Factor, Hiram, Ohio; Howard R. Feldman, American Museum
of Natural History; Daniel C. Fisher, University of Michigan; Ron Fisher, Homer, Ohio;
Raymond C. Gutschick, University of Notre Dame; Kurt F. Hallin, Milwaukee Public
Museum; Joseph T. Hannibal, Cleveland Museum of Natural History; Alan Stanley
Horowitz, Indiana University; Thomas W. Kammer, West Virginia University; Gerald
J. Kloc, University of Rochester; Albert D. Kollar, Carnegie Museum of Natural History;
Ed Landing, New York State Geological Survey; Ralph L. Langenheim, University of
Illinois at Urbana-Champaign; Richard S. Laub, Buffalo Museum of Science; Richard
L. Leary, Illinois State Museum; Pierre J. Lesperance, Universite de Montreal; Richard
Lindemann, Skidmore College; Richard Lund, Adelphi University; Royal H. Mapes,
Ohio University; Dagmar Merino, Yacimientos Petroliferos Fiscales Bolivianos, La Paz,
Bolivia; Jonathon Mortin, University College of Swansea; Matthew H. Nitecki, Field
Museum of Natural History; James C. Ohman, Kent State University; William A. Oliver,
Jr., United States Geological Survey, Washington, D.C.; David C. Parris, New Jersey
State Museum; the late Eugene S. Richardson, Jr., Field Museum of Natural History;
Gabrielo Rodrigo, Museo Nacional de Historia Natural, La Paz, Bolivia; Edgar Roeser,
Cleveland Museum of Natural History; Colin T. Scrutton, University of Newcastle-
upon-Tyne; Robert Segedi, Cleveland Museum of Natural History; Peter F, Sheehan,
Milwaukee Public Museum; Thomas M. Stanley, St. Joe Mining, Deadwood, South
Dakota; James T. Stitt, University of Missouri-Columbia; Mario Suarez-Riglos, Yaci-
mientos Petroliferos Fiscales Bolivianos, Santa Cruz, Bolivia; Robert Walker, Kent State
University; Steven C. Ward, Kent State University; Lawrence A. Wiedman, Monmouth
College; Michael E. Williams, Cleveland Museum of Natural History; Margaret T. Wil-
son, Kent State University; and Paul Zell, State College, Pennsylvania.
We would especially like to thank Margaret Wilson for her unflagging support and
encouragement of the research and for her review of the final manuscripts; without her,
these papers would have been impossible. Ellis L. Yochelson and Richard H. Lindemann
reviewed the manuscripts for the Annals of the Carnegie Museum. This study supported
in part by American Association of Petroleum Geologists Grant-in-Aid no. 582-12-01
and by a Grant-in-Aid of Research from Sigma Xi, The Scientific Research Society, both
to Babcock. A portion of the cost of publication has been provided by the Office of
Research and Sponsored Programs, Kent State University. Part A is contribution 313
and Part B is contribution 314 of the Department of Geology, Kent State University,
Kent, Ohio 44242.
Literature Cited
Alison, D,, and R. Carroll. 1972. Catalogue of type and figured fossils in the Redpath
Museum, McGill University. Redpath Museum, McGill University, 173 pp.
Babcock, L. E. 1985^2. A new Ordovician conulariid from Oklahoma? Oklahoma
Geology Notes, 45:66“70.
. 1985Z?. Mahantango conulariid considered a hyolithid. Pennsylvania Geology,
16(3);14-16.
Babcock, L. E., and R. M, Feldmann. 1984. Mysterious fossils. Earth Science, 37(3):
16-17.
— 1986. The phylum Conulariida. In Problematic Fossil Taxa (A. Hoffman
and M, H. Nitecki, eds.), Oxford University Press.
Bamber, E. W., and M. j. Copeland. 1976. Carboniferous and Permian faunas. Pp.
623-632, in Geology and Economic Minerals of Canada. Part B (Chapters VIII-
XIII and Index) (R. J. W. Douglas, ed.). Geological Society of Canada Economic
Geology Report 1 .
458
Annals of Carnegie Museum
VOL. 55
Beede, J, W. 1911. The Carbonic fauna of the Magdalen Islands. New York State
Museum, Bulletin, 149:156-186.
Bell, W. A. 1 929. Horton-Windsor District, Nova Scotia. Geological Survey of Canada,
Memoir 155, 268 pp.
Bigsby, J. J. 1878. Thesaurus Devonico-Carboniferous. The flora and fauna of the
Devonian and Carboniferous periods. . . London, 447 pp.
Boucek, B. 1939. Conularida. Pp. A1 13-A131, in Handbuch der Palaozoologie, Vol.
2 A (O. H. Schindewolf, ed.), Berlin.
Branson, E. B. 1938. Stratigraphy and paleontology of the Lower Mississippian of
Missouri. Part 1. University of Missouri Studies, 13(3), 205 pp.
~. 1944. The geology of Missouri. University of Missouri Studies, 19(3), 535 pp.
Chappars, M. S. 1936. Catalogue of the type specimens of fossils in the University of
Cincinnati Museum. Ohio Journal of Science, 36:1-45.
Clarke, J. M. 1907. Some new Devonic fossils. New York State Museum, Bulletin,
107 (Geology 12) (New York State Education Department, Bulletin 401): 153-291.
. 1908. Early Devonic history of New York and eastern North America. New
York State Museum, Memoir 9, Part 1, 366 pp.
Clarke, J. M., and R. Ruedemann. 1903. Catalogue of type specimens of Paleozoic
fossils in the New York State Museum. New York State Museum, Bulletin, 65
(Paleontology 8) (University of the State of New York, Bulletin, 314), 76 pp.
CuMiNGS, E. R. 1906. Gasteropoda, Cephalopoda and T rilobita of the Salem Limestone.
Indiana Department of Geology and Natural Resources, 30th Annual Report, pp.
1335-1375.
Dawson, J. W. 1868. Acadian geology. The geological structure, organic remains, and
mineral resources of Nova Scotia, New Brunswick, and Prince Edward Island. Sec-
ond edition. London, 694 pp.
. 1878. Acadian geology. The geological structure, organic remains, and mineral
resources of Nova Scotia, New Brunswick, and Prince Edward Island. Third edition,
London, 694 pp.
1883. Preliminary notice of new fossils from the Lower Carboniferous lime-
stones of Nova Scotia and Newfoundland. Canadian Naturalist and Quarterly Jour-
nal of Science, n.s., 10(7):41 1-416.
. 1889. Handbook of geology. Montreal, 250 pp.
Driscoll, E. G. 1963, Paraconularia newberryi (Winchell) and other Lower Missis-
sippian conulariids from Michigan, Ohio, Indiana, and Iowa. Contributions from
the Museum of Paleontology, University of Michigan, 18:33-46.
Ellison, R. L. 1965. Stratigraphy and paleontology of the Mahantango Formation in
south-central Pennsylvania. Pennsylvania Geological Survey, Fourth Series, Bulletin
G 48, 298 pp.
Feldmann, R. M., A. H. Coogan, and R. A. Heimlich. 1977. Southern Great Lakes.
Dubuque, Kendall/Hunt Publishing Company, 24 1 pp.
Grabau, a. W., and H. W. Shimer. 1910, North American index fossils, invertebrates.
Volume 2. New York, 909 pp.
Hall, J. 1858. Description of new species of fossils from the Carboniferous limestones
of Indiana and Illinois. Albany Institute, Transactions, 4:1-36.
-. 1879. Geological survey of New York, palaeontology. Volume 5. Part 2. Con-
taining descriptions of the gasteropoda, pteropoda and cephalopoda of the Upper
Helderberg, Hamilton, Portage and Chemung groups. Albany, 492 pp.
-. 1883. Description of Spergen Hill fossils. Indiana, Department of Geology and
Natural History, 12th Annual Report, pp. 319-375.
Herpers, H. 1949. A new conularid from the Esopus Formation, Sussex County, New
Jersey. New Jersey Department of Conservation and Economic Development, Mis-
cellaneous Geological Paper, 7 pp.
1986 Babcock and Feldmann—Paraconularia and Reticulaconularia
459
1950. An Onondagan fannule in New Jersey, Journal of Paleontology, 24:6 1 1-
619.
Herrick, C. L, 1887. Sketch of the geological history of Licking County. No. 2, Ad-
ditional fossils from Coal Measures at Flint Ridge. Denison University, Scientific
Laboratories, Bulletin, 2:144-148.
— . 1888<3. Geology of Licking County, Ohio. Parts III and IV. The Subcarbonif-
erous and Waverly Groups. Denison University, Scientific Laboratories, Bulletin,
3(1):13-110.
—— — . 18886. Geology of Licking County. IV. List of Waverly fossils, continued.
Denison University, Bulletin, 4(*/2):l 1-60, 97-123.
. 1893. Observations on the so-called Waverly Group of Ohio. Geological Survey
of Ohio, Report, 7:495-515.
Holm, G. 1893. Sveriges Kambrisk-Siluriska Hyolithidae och Conulariidae. Sveriges
Geologiska Undersokning, Afhandlingar och uppsatser. Series C, No. 112, 172 pp.
Kent, L. S. 1982, Type and figured fossils in the Worthen Collection at the Illinois
State Geological Survey. Illinois State Geological Survey, Circular 524, 65 pp.
Keyes, C. R. 1894. Paleontology of Missouri (part II). Missouri Geological Survey,
Vol. 5, 266 pp.
Lane, N. G. 1973. Paleontology and Paleoecology of the Crawfordsville Fossil Site
(Upper Osagian; Indiana). University of California Press, Berkeley, 141 pp.
Laudon, L. R., and a. L. Bowsher. 1941. Mississippian formations of Sacramento
Mountains, New Mexico. American Association of Petroleum Geologists, Bulletin
25:2107-2160.
Leary, R. L. 1985. Fossil noncalcareous algae from Visean (Mid Mississippian) strata
of Illinois (U.S.A.). Dixieme Congres International de Stratigraphie et de Geologie
du Carbonifere (Madrid), Compte Rendu, Volume 2:307-317.
Lehmann, U., and G. Hillmer. 1983. Fossil Invertebrates. Cambridge University
Press, 350 pp.
Lesley, J. P. 1889. A dictionary of the fossils of Pennsylvania and neighboring states
named in the reports and catalogues of the survey. Pennsylvania Geological Survey,
Report P 4, Volumes I-III, 437 pp.
“. 1895. A summary description of the geology of Pennsylvania. Pennsylvania
Geological Survey, Final Report, 3:1629-2152.
McKee, E, D., and R, C. Gutschick. 1969. History of the Redwall limestone of
northern Arizona. Geological Society of America, Memoir 114, 726 pp.
Meek, F. B. 1875, A report on some of the invertebrate fossils of the Waverly Group
and Coal Measures of Ohio. Geological Survey of Ohio, Report, 2(2);269-347.
Meek, F. B., and A. H, Worthen. 1865. Contributions to the paleontology of Illinois
and other western states. Academy of Natural Sciences of Philadelphia, Proceedings
for 1865, pp. 245-273.
. 1873. Descriptions of invertebrates from Carboniferous System. Geological
Survey of Illinois, 5:323-619.
Miller, S. A. 1877. The American Paleozoic fossils: a catalogue of the genera and
species, with names of authors, dates, places of publication, groups of rocks in which
found, and the etymology and significance of the words, and an introduction devoted
to the stratigraphical geology of the Paleozoic rocks. Cincinnati, 253 pp.
- — - — — . 1889, North American geology and palaeontology for the use of amateurs,
students, and scientists. Cincinnati, 664 pp.
. 1 ^92a. Palaeontology. Indiana Department of Geology and Natural Resources,
17th Annual Report, pp. 611-705.
“. 18926. First appendix, 1892. Cincinnati, pp. 665-718.
— -. 1897. Second appendix to North American geology and paleontology, October,
1897. Cincinnati, pp. 719-793.
460
Annals of Carnegie Museum
VOL. 55
Miller, S. A., and W. F. E. Gurley. 1893. Descriptions of some new species of
invertebrates from the Palaeozoic rocks of Illinois and adjacent states. Illinois State
Museum of Natural History, Bulletin, 3, 81 pp.
. 1896. New species of Palaeozoic invertebrates from Illinois and other states.
Illinois State Museum of Natural History, Bulletin, 11,50 pp.
Moore, R. C., and H. J. Harrington. 1956. Conulata. Pp. F54=-F66, in Treatise on
invertebrate paleontology. Part F. Coelenterata (R. C. Moore, ed.). Geological Society
of America and University of Kansas Press.
Shimer, H. W. 1926. Upper Paleozoic faunas of the Lake Minnewanka section, near
Banff, Alberta. Geological Survey of Canada, Bulletin, 42 (Geological Series, No.
45): 1-84.
Sinclair, G. W. 1940. The genotype of Conularia. Canadian Field-Naturalist, 54:
72-74.
. 1942. A new species of '‘^Conularia’'’ from Gaspe. Le Naturaliste Canadien,
69:158-160.
— . 1948. The Biology of the Conularida. Unpublished Ph.D. thesis, McGill Uni-
versity, 442 pp.
Swallow, G. C. 1860. Descriptions of new fossils from the Carboniferous and De-
vonian rocks of Missouri. Academy of Science of St. Louis, Transactions, 1:635-
660.
Tasch, P. 1 973. Paleobiology of the invertebrates. Data retrieval from the fossil record.
New York, 946 pp.
. 1980. Paleobiology of the invertebrates. Data retrieval from the fossil record.
Second edition. New York, 975 pp.
Thompson, I. 1982. The Audubon Society field guide to North American fossils. New
York, Alfred A. Knopf, 846 pp.
Walcott, C. D. 1884. Paleontology of the Eureka District. United States Geological
Survey, Monographs, 8, 298 pp.
Weller, S. 1898. A bibliographic index of North American Carboniferous inverte-
brates. United States Geological Survey, Bulletin, 153, 653 pp.
. 1 900a. Kinderhook faunal studies. 11. The fauna of the Chonopectus Sandstone
at Burlington, Iowa. Academy of Science of St. Louis, Transactions, 10:57-129.
— . 1 900Z). The succession of fossil faunas in the Kinderhook beds at Burlington,
Iowa. Iowa Geological Survey, 10:59-79.
White, C. A. 1862. Description of new species of fossils from the Devonian and
Carboniferous rocks of the Mississippi Valley. Boston Society of Natural History,
Proceedings, 9:8-33.
. 1880. Fossils of the Indiana rocks. Indiana Department of Statistics and Ge-
ology, 2nd Annual Report, pp. 471-552.
Whiteaves, J. F. 1891. The fossils of the Devonian rocks of the Mackenzie River
Basin. Geological and Natural History Survey of Canada, Contributions to Canadian
Palaeontology, 1:197-253.
Whitfield, R. P. 1882. On the fauna of the Lower Carboniferous limestones of Spergen
Hill, Ind., with a revision of its fossils hitherto published, and illustrations of the
species from the original type series. American Museum of Natural History, Bulletin,
l(3):39-97.
Whitfield, R. P., and E. O. Hovey. 1901. Catalogue of the types and figured specimens
in the paleontological collection of the Geological Department, American Museum
of Natural History. Part IV. Carboniferous to Pleistocene, inclusive. American Mu-
seum of Natural History, Bulletin, 1 1:357-500.
WiNCHELL, A. 1865. Descriptions of new species of fossils from the Marshall Group
of Michigan, and its supposed equivalent, in other states; with notes on some fossils
of the same age previously described. Academy of Natural Sciences of Philadelphia,
Proceedings for 1865, pp. 109-133.
1986 Babcock and Felumann—Paraconularia and Reticulaconularia
461
1870. Notices and descriptions of fossils, from the Marshall Group of the
western states, with notes on fossils from other fossils. American Philosophical
Society, Proceedings, 11:245-260.
WoRTHEN, A. H. 1883. Description of some new species of fossil shells from the Lower
Carboniferous limestones and Coal Measures of Illinois. Geological Survey of Illi-
nois, 7:323-326.
1890. Description of fossil invertebrates. Geological Survey of Illinois, 8:69-
154.
Yochelson, E. L., G. D. Stanley, and W. Sturmer. 1983. Plectodiscus discoideus
(Rauff): a redescription of a chondrophorine from the Early Devonian Hunsriick
Slate, West Germany. Palaontologie Zeitschrift, 57:39-68.
Appendix A— Locality Index
Collector, year of collection if known, description of locality, stratigraphic assignment and
conulariid taxa present
Alabama
1 A. S. Horowitz. “Keyes site,” Skyline, Alabama. Formation unknown; probably
Chesterian Series. Paraconularia chesterensis.
Alberta
2 Anonymous. Near Banff, Alberta. Formation unknown and series unknown; Mis-
sissippian Subsystem. Paraconularia missouriensisl
3 R. G. McConnell, 1890. Athabasca River, La Saline, Alberta. Formation unknown;
Chautauquan Series. Paraconularia salinensis.
4 H. W. Shimer, pre-1926. Lake Minnewanka, Alberta. Formation and series un-
known; Mississippian Subsystem. Paraconularia alternistriata.
5 F. Beales. Upper part of Job Creek, western Alberta. Upper Rundle Formation, 345
m from the base of the formation; Chesterian Series. Paraconularia missouriensisl
Arizona
6 R. C. Gutschick and P.C.H., 1954. Top of mesa on point between Rock and Blye
Canyons on 7BarV Ranch, WF Cattle Company, south of Peach Springs and Cher-
okee Point, Arizona. Chert in Member 2 of Redwall Limestone, about 53 m above
base of Redwall Formation; ?Osagean Series. No conulariids collected; Oracanthus
spine.
British Columbia
7 D. Scott, 1962. Spur on northeast comer of Mt Hosmer, 14.5 km northeast of
Femee and 14.5 km southwest of Natal, British Columbia. Lower Etherington
Member of the Rocky Mountain Formation of the Rundell Group; Chesterian Series.
Paraconularia chesterensisl
Illinois
8 Anonymous. Kinderhook, Pike County, Illinois. Kinderhook Group; Kinderhook-
ian Series. Conularia subcarbonaria.
9 W. F. E. Gurley; Anonymous. Hamilton, Illinois. Keokuk Limestone; Osagean
Series. Conularia subcarbonaria.
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Annals of Carnegie Museum
VOL. 55
10 Anonymous. Chester, Randolph County, Illinois. Chester Limestone; Chesterian
Series. Paraconularia chesterensis.
1 1 Sloss. Pike County, Illinois. Burlington Limestone; Osagean Series. Paraconularia
chesterensis, P. blairi.
12 S. Weller, 1912. About 2.4 km south of Marigold, Illinois. “Lower Okaw, Marigold
Oolite” (=Burlington Limestone?); Osagean Series. Paraconularia chesterensis.
13 W. F. E. Gurley; Anonymous. Hamilton, Illinois. Keokuk Limestone; Osagean
Series. Conularia subcarbonaria.
14 W. F. E. Gurley. Madison County, Illinois. St. Louis Limestone; Meramecian series.
Paraconularia missouriensis.
1 5 Anonymous. Warsaw, Madison County, Illinois. St. Louis Limestone; Meramecian
Series. Paraconularia missouriensis.
16 Anonymous. Alton, Illinois. St. Louis Formation; Meramecian Series. Paraconu-
laria subulata, P. chesterensis.
17 W. F. Gurley. Madison County, Illinois. St. Louis Limestone; Meramecian Series.
Paraconularia chesterensis.
1 8 S. Weller, 1912. About 3.2 km east of Waterloo, Illinois, Illinois. Renault Limestone;
Chesterian Series. Paraconularia chesterensis.
19 S. Weller, 1918. About 4 km southeast of Vienna, Illinois. Glen Dean Limestone;
Chesterian Series. Paraconularia chesterensis.
20 A. S. Horowitz, 1966. Debris from slope below quarry in Mississippi River bluffs
above Illinois State Highway 3 Bypass, SW'A, SW‘/4 Sec.29, TVS, R6W, Chester,
Randolph County, Illinois, Chester 7.5' Quadrangle. Menard Limestone; Chesterian
Series. Paraconularia chesterensis, Conularia cf. C. subcarbonaria.
21 L. F. Rauchfrise; Anonymous. Pope County, Illinois. Chester Group; Chesterian
Series. Paraconularia chesterensis.
Indiana
22 E. M. Kindle. Delphi, Indiana. Sellersburg Formation; Chautauquan Series. Con-
ularia delphiensis.
23 D. G. Maroney and R. W. Orr, pre-1974. Delphi Limestone Company Quarry,
north side of U.S. Highway 421, northwest edge of Delphi, SWV4, SW‘/4 Sec. 19,
T25N R2W, Carroll County, Indiana, Delphi 7.5' Quadrangle. 0-10 cm thick phos-
phatic pebble bed at base of New Albany Shale; Chautauquan Series. Conularia
delphiensis.
24 D. E. Hattin? Probably Indiana. Probably Harrodsburg Formation; Osagean Series.
Conularia subcarbonaria.
25 D. E. Hattin. Hattin location S-776. Indiana. Borden Group; Meramecian Series.
Conularia multicostata.
26 Anonymous. Curiosity Hollow, near Martinsville, Indiana. New Providence For-
mation; Osagean Series. Paraconularia cf. P. subulata.
27 Anonymous. Crawfordsville, Indiana. “Keokuk Group” (=Borden Group); Osagean
Series. Conularia subcarbonaria, Paraconularia chesterensis.
28 W. F. E. Gurley. West Point, Indiana. “Keokuk Group” (=Borden Group); Osagean
Series. Conularia subcarbonaria.
29 G. K. Greene; Washburn. New Albany, Indiana. “Knobstone Group” (=Borden
Group); Osagean Series. Paraconularia byblis, P. subulata.
30 Anonymous. Spergen Hill, Indiana. St. Louis Limestone; Meramecian Series. Par-
aconularia missouriensis.
31 W. F, Gurley; Anonymous. Edwardsville, Indiana. “Keokuk Group” (=Borden
Formation); Osagean Series. Paraconularia missouriensis.
32 Klippart. New Providence, Indiana. Carwood Member, Borden Formation; Osagean
Series. Paraconularia blairi.
1986 Babcock and Feldmann—Paraconularia and Reticulaconularia
463
33 E. O. Ulrich. Crawfordsville, Indiana. Borden Formation (“Keokuk Group”); Osa-
gean Series. Paraconuiaria chesterensis.
34 Anonymous. Near Providence, Indiana. Borden Formation; Osagean Series, Par-
aconuiaria blairi.
35 C. Rominger? Crawfordsville, Montgomery County, Indiana. Near middle of Bor-
den Group; Osagean Series. Paraconuiaria chesterensis.
36 S. Makvrat. Bed of Gnaw Bone Creek just south of Indiana Highway 46, just east
of Gnaw Bone, Indiana, Nashville 7.5' Quadrangle. New Providence Shale according
to collector; more likely Carwood or Locust Point Formation of Borden Group
according to A, S. Horowitz (written communication, 1984); ?Osagean Series. Par-
aconuiaria cf. P. byblis, P. chesterensis.
37 R. Fields and J. Harris, 1981. Section on old Indiana Highway 37, SE'A, NW*/},
NW*/4 Sec.21, T9N, RIW, Monroe County, Indiana, Bloomington 7.5' Quadrangle.
Ramp Creek Member of Harrodsburg Limestone, 3-7 m above contact with Borden
Group; ?Osagean Series. Conularia subcarbonaria.
38 J. Hall?; Washburn; Anonymous. Crawfordsville, Indiana. “Keokuk Group” (=Bor-
den Group); Osagean Series. Conularia subcarbonaria, Paraconuiaria chesterensis,
P. subulata.
39 W. F. E. Gurley; G. Robb, pre-1923. New Albany, Indiana. “Knob or Knobstone
Shale” (=Borden Group?); Osagean Series. Conularia multicostata, Paraconuiaria
byblis, P. chesterensis, P. missouriensis, P. subulata.
40 W. F. E. Gurley. West Point, Indiana. “Keokuk Group (=Borden Formation)”;
Osagean Series. Paraconuiaria chesterensis.
41 R. L. Anstey et al., 1968. Bed of Indian Creek on O. C. Bennett or Ben Wilson
property, approximately .8 km north of Indiana Highway 234, NE'A, NW ‘A, SE'A
Sec, 8, T17N, R5W, Montgomery County, Indiana, ?Edwardsville Formation of
Borden Group; Osagean Series. Paraconuiaria cf. P. chesterensis.
42 C. Rominger? Crawfordsville, Montgomery County, Indiana. Near middle of Bor-
den Group; Osagean Series. Conularia subcarbonaria.
43 G. Campbell. Floyds Knob Hill, Highway 150, near center of NE'A 28-25-6E {sic),
3.2 km northwest of New Albany, Indiana. Kenwood Formation; ?Osagean Series.
Paraconuiaria byblis.
44 W. F. E. Gurley. Spergen Hill, Indiana. Probably St. Louis Limestone, though
possibly Salem Limestone; Meramecian Series. Conularia subcarbonaria.
45 W. F. E. Gurley. Spergen Hill, Indiana. St. Louis Limestone or Salem Limestone;
Meramecian Series. Conularia subcarbonaria.
46 Anonymous. Salem, Indiana. Salem Limestone; Meramecian Series. Conularia sub-
carbonaria.
47 A. S. Horowitz, N. G. Lane et al. Outcrop along west branch of Mosquito Creek,
0.48 km west and 0.24 km north of southeast comer of Sec. 25, T5S, R5E, ap-
proximately 4.5 km east of Laconia, Harrison County, Indiana; Laconia 7.5' Quad-
rangle, Somerset Shale; Chesterian Series. Paraconuiaria chesterensis.
48 J. J. Galloway, 1949. Galloway location L40C, ravine beginning at railroad, 1,2
km northwest of Harrodsburg and mnning northwest 0.8 km to old Indiana Highway
7, SEVa, SWV4 Sec. 20, T7N RIW, Monroe County, Indiana, Clear Creek 7.5'
Quadrangle. Lower part of Harrodsburg Formation; Meramecian Series. Conularia
cf. C. subcarbonaria.
49 J. J, Galloway, 1949. Galloway location 1.70A, old quarry, 4 km northwest of
Dolan, Monroe County, Indiana. Site is probably an abandoned Quarry 2,4 km
northwest of Dolan shown on Modesto 7.5' Quadrangle topographic map; WV2,
SW*/4, NWV4 Sec. 34, T ION RIW, approximately 0.4 km east of old Indiana High-
way 37 (A.S. Horowitz, written communication, 1984). Lower part of Harrodsburg
Formation; Meramecian Series. Paraconuiaria chesterensis.
50 D, E. Hattin, 1965. Roadcut on State Highway 46, east of Gnaw Bone, approxi-
464
Annals of Carnegie Museum
VOL. 55
mately NV2, NV2 Sec.25, T9N, R4E, Brown County, Indiana, Nashville 7.5' Quad-
rangle. Carwood Formation of Borden Group; Chesterian Series, Paraconularia cf.
P. byblis, P. chesterensis.
5 1 Anonymous. Spergen Hill, Indiana. Spergen Limestone; Meramecian Series. Con-
ularia subcarbonaria, Paraconularia chesterensis.
52 Anonymous. Spergen Hill, Indiana? Spergen Hill Limestone?; Meramecian Series?
Paraconularia chesterensis.
53 Anonymous. Hendricks County, Indiana. Formation unknown; probably Chesterian
Series. Paraconularia chesterensis.
54 W. F. E. Gurley. Evansville, Indiana. “Chester Group”; Chesterian Series. Para-
conularia chesterensis.
55 A. S. Brockley and T. G. Perry, 1954. Debris from Mulzer Brothers Quarry, SWy4,
NE‘/4 Sec. 3, T2S, R2W, Crawford County, Indiana, Taswell 7.5' Quadrangle. “Glen
Dean Limestone” (=Lower Tar Springs Formation); Chesterian Series. Paraconu-
laria chesterensis.
56 A. S. Horowitz. Spoil heaps from Mulzer Brother Quarry, north and south of county
road, approximately 1.5 km south of junction of Indiana Highways 145 and 164
and approximately 4 km north of Eckerty and .4 km east of Indiana Highway 145,
SW'/4, SE‘/4 Sec. 10, T2S, R2W, Crawford County, Indiana, Taswell 7.5' Quadrangle.
Glen Dean Limestone; Chesterian Series. Paraconularia chesterensis.
57 A. S. Horowitz et al., 1956-1982. Railroad cuts on west side of Baltimore and Ohio
Railroad spur leading to National Gypsum Company quarry, near Shoals, SW*/*,
NE‘/4 Sec. 28, T3N R3W, Martin County, Indiana, Huron 7.5' Quadrangle. “Gol-
conda Formation” (=Indian Springs Member of the Big Clifty Formation); Ches-
terian Series. Paraconularia chesterensis.
58 A. C. Brookley and T. G. Perry, 1954. Debris from abandoned Lutgring Quarry,
9.6 km east of Branch ville, NW'A, SW*/4 Sec. 18, TIS, RIW, Perry County, Indiana,
Branchville 7.5' Quadrangle. “Glen Dean Limestone” (=Lower Tar Springs For-
mation); Chesterian Series, Paraconularia chesterensis.
59 Haines. Washington County, Indiana. Formation unknown; probably Chesterian
Series. Paraconularia chesterensis.
60 J. Below, 1965. Dam site 4. 8-6.4 km north off Route 46, 4.8 km east of Gnaw
Bone, Indiana. Borden Group; Osagian Series? Paraconularia byblis.
Iowa
61 W. F. Gurley. Le Grand, Iowa. Formation unknown; Kinderhookian Series. Con-
ularia subcarbonaria, Paraconularia blairi.
62 C. A. White? Burlington, Iowa. English River Sandstone of the Kinderhook Group;
Kinderhookian Series. Paraconularia byblis.
63 C. A. White? Burlington, Iowa. “Upper Division of the Burlington Limestone”;
probably Osagian Series. Paraconularia subulata.
64 Anonymous. Iowa City, Iowa. Cedar Valley Limestone; Chautauquan Series. Con-
ularia subcarbonaria.
65 Anonymous. Probably Iowa City area, Iowa. Probably Cedar Valley Limestone;
Chautauquan Series. Conularia subcarbonaria.
66 Anonymous. Burlington, Iowa. Burlington Limestone; Osagean Series. Paraconu-
laria byblis.
67 Fenton. Southwest of Waverly, Iowa. Cedar Valley Limestone, lower part; Osagean
Series. Conularia subcarbonaria.
68 Anonymous. Burlington, Iowa. Burlington Limestone; Osagean Series. Conularia
subcarbonaria.
1986 Babcock and FELDMANN-”/l4i?^CDA^c/z^i?L4 and Reticulaconularia
465
69 Anonymous. Keokuk, Iowa. Keokuk Formation; Osagean Series. Paraconularia
missouriensis.
70 S. Weller? Keokuk, Iowa. Keokuk Limestone, “bed 1 1”; Osagean Series. Paracon-
ularia chesterensis.
Kentucky
71 C. E. Mason, 1984. Outcrops along Interstate 64, 8.2 km east of the junction with
Kentucky Route 32, near Morehead, Rowan County, Kentucky. Float from lower
few meters of Nancy Member of Borden Formation; Osagean Series. Conularia
multicostata, Paraconularia byblis.
72 C. E. Mason, T. M. Stanley, and L. E. Babcock, 1984. Nancy Member of Borden
Formation; phosphate pebble bed about 1 m above top of “dysaerobic fauna”;
Osagean Series. Spillway to Cave Run Lake, Daniel Boone National Forest, Bath
County, Kentucky, Salt Lake 7.5' Quadrangle, Conularia multicostata, Paraconu-
laria byblis, P. subulata.
IZ Anonymous. Natural Bridge, Kentucky. Borden Formation, probably Nancy Mem-
ber {fide F. R. Ettensohn, written communication, 1985); Osagean Series. Conularia
multicostata.
74 Anonymous. Marion County, Kentucky. Borden Formation?; Osagean Series? Par-
aconularia byblis.
75 Anonymous. Lebanon, Kentucky. “Waverly Formation” (=Borden Group?); prob-
ably Osagean Series. Paraconularia subulata.
76 Anonymous. About 2.5 km east of Lebanon, Kentucky. New Providence Formation;
Osagean Series. Conularia subcarbonaria, Paraconularia byblis.
11 T. W. Kammer. Kammer location 10885, St. Francis, Kentucky, Nancy Member
of the Borden Formation; Osagean Series. Paraconularia byblis.
78 T. W. Kammer. Kenwood Hill, Louisville, Kentucky. New Providence Shale Mem-
ber of the Borden Formation; Osagean Series. Paraconularia byblis.
79 Anonymous, Kentucky? Waverly Group equivalent?; Osagean Series? Conularia
multicostata.
80 U. P. James. Boyle or Marion County, Kentucky. “Waverly Group” (=Borden
Group?); probably Osagean Series. Conularia multicostata.
8 1 Anonymous. Knob just south of Louisville, Kentucky. New Providence Formation;
?Osagean Series. Paraconularia byblis.
82 G. Robb, pre-1923. Marion County, Kentucky. “Keokuk Formation, Knob Shale”
(=Borden Formation?); Osagean Series. Paraconularia subulata.
83 Anonymous. Elizabethtown, Kentucky. St. Louis Limestone; Meramecian Series.
Paraconularia missouriensis.
84 A. S. Horowitz. Pond north of Kentucky Highway 1576, about 2 miles east of
Morrilla, Jackson County, Kentucky. Pennington Formation; Chesterian Series.
Paraconularia chesterensis.
85 A. S. Horowitz. Near Colesburg, Hardin County, Kentucky. Somerset Shale Member
of Salem Limestone; Meramecian Series. Paraconularia chesterensis.
86 S. Weller, 1920. 5.2 km south of lola, Kentucky (GK 12). Glen Dean Limestone;
Chesterian Series. Paraconularia chesterensis.
87 Anonymous. About 1 .5 km west of Mongomery Switch, Caldwell County, Kentucky.
Claystone bed of upper Chester Formation; Chesterian Series. Paraconularia ches-
terensis.
88 A. S. Horowitz, 1966, etc. Pond above road leading to Pearson Farm glade (road
not on topographic map), near junction of road with Kentucky Highway 1576,
approximately 3.0 km east ofjunction of Kentucky Highway 1 576 and U.S. Highway
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Annals of Carnegie Museum
VOL. 55
421, approximately 3 km east of Morrill, Jackson County, Kentucky, Big Hill 7.5'
Quadrangle. Lower part of Pennington Formation, just above top of Bangor Lime-
stone; Chesterian Series. Paraconularia chesterensis.
89 A. S. Horowitz, 1969. Roy Norton Farm, glades on north, west and south slopes
of tributaries on north side of Broad Run, SO-lOO m south of Kentucky Highway
434, approximately 2.6 km west-southwest of Colesburg, Hardin County, Kentucky,
Colesburg 7.5' Quadrangle. Somerset Shale Member of the Salem Limestone; Ches-
terian Series. Paraconularia chesterensis.
Maine
90 Anonymous. Presque Isle stream. Chapman Plantation, Maine. Chapman Sand-
stone; series unknown, Devonian System. Conularia cf C undulata.
Maryland
91 F. M. Swartz. West Maryland Railroad tracks, Corrigansville, Maryland. Upper
part of the Shriver Chert; Ulsterian Series. Conularia undulata.
Michigan
92 Anonymous. Alpena, Michigan. Alpena Limestone; Brian Series. Paraconularia
alpenensis.
93 E. W. Hard. U.S. Gypsum Company Quarry, Sec. 27, T27N, R7E, near Alabaster,
Michigan. Michigan Formation, gray gypsiferous limestone bed 1.3 m below 5.3
m thick bed of mottled white gypsum; Osagean Series. Paraconularia yochelsoni.
Missouri
94 Sampson; Faber. Sedalia, Missouri, Chouteau Limestone; Kinderhookian Series.
Paraconularia blairi, P. missouriensis.
95 J. S. Williams, 1930? Easley, Missouri. Chouteau Limestone; Kinderhookian Series.
Paraconularia blairi.
96 E. B. Branson. Providence, Missouri. Chouteau Limestone; Kinderhookian Series.
Paraconularia blairi.
97 E. B. Branson, 1930. Providence, Missouri. Couteau Limestone; Kinderhookian
Series. Paraconularia blairi.
98 Anonymous. Pettis County, Missouri. Chouteau Limestone; Kinderhookian Series.
Paraconularia blairi.
99 E. B. Branson. Browns, Missouri. Chouteau Limestone; Kinderhookian Series. No
conulariids identified; ?priapulid worm, ^‘‘Conularia"'’ tenuicostata.
100 W. F. E. Gurley. Boonville, Missouri. Keokuk Limestone; Osagean Series. Para-
conularia missouriensis.
101 Anonymous. Carthage, Missouri. Formation unknown; Meramecian Series. Con-
ularia subcarbonaria.
102 Van Home. Foot of La Beaume Street, St. Louis, Missouri. St. Louis Limestone;
Meramecian Series. Paraconularia blairi, P. chesterensis.
103 Anonymous, Kansas City, Missouri. “Coal Measures”; probably Osagean or Mer-
amecian Series. Paraconularia missouriensis.
1 04 Anonymous. Carthage, Missouri. Keokuk Limestone; Osagean-Meramecian Series,
Paraconularia chesterensis.
105 Anonymous. Little Rock, St. Genevieve County, Missouri. St. Louis Limestone;
Meramecian Series. Paraconularia blairi.
1986 Babcock and Feldmann --Paraconularia and Reticulaconularia
467
Montana
106 R. Lund et al., 1978-1984. Potter’s Creek Dome, approximately 50 km southeast
of Lewistown, Fergus County, Montana. Bear Gulch Limestone of Heath Formation;
Chesterian Series. Paraconularia subulata.
107 W, H, Easton? Delpine, Meagher County, Montana. Cameron Creek Shale of the
Big Snowy Group; Chesterian Series. Paraconularia subulata.
Nevada
108 Merriam. Simpson Park Range, Nevada. Rabbit Hill Limestone; Ulsterian Series.
Conularia sp.
109 A. J. Boucot, 1984. West face of Red Hill, Eureka County, Nevada. “Fish bed” of
the Denay Limestone; Senecan Series. Conularia recurvatus.
110 C. D. Walcott. Eureka District, Nevada. Formation and series unknown; probably
Chesterian Series. Paraconularia chesterensis.
New Jersey
111 D. Parris and K. Cruikshank, 1980. On Weider Road, near County Road 521,
Montague Township, Sussex County, New Jersey. Esopus Formation; Ulsterian
Series. Reticulaconularia sussexensis.
1 12 H. Herpers. Montague, Sussex County, New Jersey. Esopus Formation; Ulsterian
Series. Reticulaconularia sussexensis.
113 H. Herpers, 1948. Millville, Montague Township, Sussex County, New Jersey.
Esopus Formation; Ulsterian Series. Reticulaconularia sussexensis.
1 14 D. Parris, K. Cruikshank et al., 1 984. 1.7 km southwest of Wallpack Centre, roadcut
across from Batteli’s Campground, Wallpack Township, Sussex County, New Jersey.
Port Ewen Formation; Ulsterian Series. Conularia pyramidalis.
New York
115 R. M. Fulle. First Esopus outcrop south-southeast of Hurley, on left fork of road,
about 1 .6 km from Hurley, Ulster County, New York. Esopus Formation; Ulsterian
Series. Conularia ulsterensis.
116 Anonymous. Schoharie, New York. New Scotland Limestone; Ulsterian Series.
Conularia pyramidalis.
1 1 7 Anonymous. Clarkesville, Schoharie County, New York. “Lower Helderberg Group”
(=New Scotland Limestone); Ulsterian Series. Conularia pyramidalis.
1 18 Anonymous. Probably Clarksville area, Schoharie County, New York. Helderberg
Group?, possibly New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
1 1 9 Anonymous. NYSM locality 2969, near Helderberg, New York. Manlius Limestone;
Ulsterian Series. Conularia pyramidalis.
120 Anonymous. Countryman’s Hill, New Salem, New York. Coeymans Limestone or
New Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
121 Anonymous. Knox, Albany County, New York. Oriskany Sandstone; Ulsterian
Series. Conularia desiderata.
122 Anonymous. Schoharie, New York. Oriskany Sandstone; Ulsterian Series. Conu-
laria pyramidalis.
123 F. M. Swartz. Clarksville, New York. “Lower Helderberg Group,” probably New
Scotland Limestone; Ulsterian Series. Conularia pyramidalis.
124 Anonymous. NYSM location 2, about 2 km south of Bridgewater, New York.
Marcellus Shale; Erian Series, Conularia desiderata.
468
Annals of Carnegie Museum
VOL. 55
125 Anonymous. Bridgewater, New York. Marcellus Shale; Erian Series. Conularia
desiderata.
1 26. Anonymous. Probably from near Bridgewater, New York. Probably from Marcellus
Shale; Erian Series. Conularia desiderata.
127 Anonymous. Vicinity of Hamilton, New York. Lower part of Hamilton Group;
Erian Series. Conularia desiderata.
128 Anonymous. Morrisville, New York. Hamilton Group, probably Solsville Member
of the Marcellus Formation; Erian Series. Conularia undulata.
129 Anonymous. About 1.2 km northwest of Solsville, Madison County, New York.
Hamilton Group; Erian Series. Conularia undulata.
130 P. Zell, 1982, Swamp Road quarry, near Morrisville, New York, Morrisville IS
Quadrangle. Marcellus Formation; Erian Series. Conularia pyramidalis.
131 L. E. Babcock, J. T. Hannibal, and R. M. Feldmann, 1984. Borrow pit on east side
of Swamp Road, 4,2 km north of Morrisville, New York, Morrisville 7.5' Quad-
rangle. Solsville Member of the Marcellus Formation; Erian Series. Conularia pyr-
amidalis.
132 Anonymous. Schoharie County, New York. Hamilton Group, possibly Schoharie
Formation; Erian Series. Conularia desiderata.
133 M. Kopf. 4 km east of Alexander, New York. Centerfield Limestone Member,
Ludlowville Formation; Erian Series. Conularia desiderata.
134 1. H. Reimann. Spring Creek, Alden, New York. Ledyard Shale Member of Lud-
lowville Formation; Erian Series. Conularia desiderata.
135 G. J. Kloc, 1983. Lake Erie shore, south of the Wanakah Water Plant, Wanakah,
Erie County, New York, N42°44'50" W78°54'13". Nautilus Bed of the Wanakah
Shale Member of the Ludlowville Formation; Erian Series. Conularia desiderata.
136 Anonymous. Genesee Valley, New York. Hamilton Group; Erian Series. Conularia
undulata.
1 37 Anonymous. Norton’s Landing, Cayuga Lake, New York. Hamilton Group, possibly
King Ferry Shale Member of the Ludlowville Formation; Erian Series. Conularia
undulata.
138 Anonymous. NYSM location 428, Shurger’s Glen, near Norton’s Landing, Cayuga
Lake, New York. Hamilton Group; Erian Series. Conularia undulata.
139 Anonymous. NYSM location 437, Shurger’s Glen, near Norton’s Landing, Cayuga
Lake, New York. Hamilton Group; Erian Series. Conularia sp.
140 G. C. Baird, ca. 1980; L. E. Babcock, 1983. Banks and bed of Bamum Creek, below
high falls 0.4 km west (upstream) from New York Route 89 overpass, Sheldrake
Quadrangle, New York. Bamum Creek Bed of the King Ferry Shale Member of the
Ludlowville Formation, approximately 1 0 m above top of the Pleurodictyum zone;
Erian Series. Conularia undulata.
141 G. C. Baird, ca. 1980. Bed of Sheldrake Creek, below high falls 0.48 km northeast
(downstream) from New York 89 overpass, Sheldrake Quadrangle, Seneca County,
New York. 2 m below Bamum Creek Bed, in the King Ferry Shale Member of
Ludlowville Formation; Erian Series. Conularia undulata.
142 G. A. Cooper, ca. 1930. Hamilton, New York, Cooper location 8Qa. Upper part
of Pompey Formation; Erian Series. Conularia desiderata.
143 Anonymous. NYSM location 558, Norwich, Chenango County, New York. Ham-
ilton Group; Erian Series. Conularia undulata.
144 Anonymous. NYSM location 611, Schoharie County, New York. Hamilton Group;
Erian Series. Conularia desiderata.
1 45 Anonymous. Near Cazenovia, New York. Hamilton Group; Erian Series. Conularia
undulata.
1 46 Anonymous. Cazenovia, New York. Hamilton Group, possibly Moscow Formation;
Erian Series. Conularia desiderata, C. undulata.
1986 Babcock and pELDMANN—P^ij^coAc/z^i?/^ and Reticulaconularia
469
147 P, Zell, 1983. Thompson Hill Road quarry, near Earlville, New York, Earlville 1.5'
Quadrangle. Moscow Formation; Erian Series. Conularia undulata.
148 R. M. Linsley; L. E. Babcock, J. T. Hannibal, and R. M. Feldmann, 1984. “Earlville
trilobite quarry,” off Morris Road, near Morrisville, New York, Earlville 7.5' Quad-
rangle. Upper part of Moscow Formation; Erian Series. Conularia undulata.
149 Anonymous. Folsomdale, New York. Rhinestreet Shale; Senecan Series. Conularia
congregatal
150 Anonymous. Ithaca, New York. Ithaca Formation; Senecan Series. Conularia con-
gregata.
151 Anonymous. NYSM location 347, Ithaca, New York. “Chemung Group”; Senecan
Series. Conularia sp.
1 52 Anonymous. NYSM location 390, west side of Cayuga Lake inlet, New York. Ithaca
Shale; Senecan Series. Conularia congregata.
153 Anonymous. NYSM location 392, 1.5 km southeast of Ithaca, New York. Ithaca
Group; Senecan Series. Conularia congregata.
1 54 J. W. Hall and G. B. Simpson, 1 870. NYSM location 425, Ithaca, New York. Ithaca
Shale, lower part of formation; Senecan Series. Conularia congregata.
155 Anonymous. NYSM location 514, Catskill Turnpike, 3-4.5 km east of Stamford,
Delaware County, New York. Ithaca Group; Senecan Series. Conularia congregata.
156 J. W. Hall and C. Van Deloo, 1866. “Mr, Cornell’s Quarry,” 1.5 km northeast of
Ithaca; also from Cemetary quarry and Cascadilla Creek, Ithaca, New York. Ithaca
Shale; Senecan Series. Conularia congregata.
157 D. D. Luther, 1900. NYSM locality 2439, West Hill, near Naples, New York.
“Naples Group”; Senecan Series. Conularia congregata.
1 58 Anonymous. South Hill, Ithaca, New York. Ithaca Shale; Senecan Series. Conularia
congregata.
1 59 P. Zell, 1 982-1 983. Collins Hill Road quarry, near Sherburne, New York, Sherburne
7.5' Quadrangle. Hamilton Group; formation unknown; probably Senecan Series.
Conularia congregata.
160 L. E. Babcock, E. L. Yochelson, and W. T. Kirchgasser, 1982. Big Sister Creek,
Angola, Erie County, New York. Float in Angola Shale; Chautauquan Series. Con-
ularia cf. C. congregata.
161 E. B. Hall. E. B. Hall locality I, Wellsville, New York. Wellsville Formation; Chau-
tauquan Series. Paraconularia wellsvillia.
162 E. B. Hall. E. B. Hall locality XVIII, Almond, New York. Wellsville Formation;
Chautauquan Series. Paraconularia wellsvillia.
Nova Scotia
1 63 Anonymous. Cape Breton, Nova Scotia. Lower Windsor Group; Osagian-Chesterian
Series. Paraconularia planicostata.
1 64 W. Dawson. Irish Cove, Cape Breton, Nova Scotia. “Lower Carboniferous” lime-
stone, probably Windsor Group; Osagian-Chesterian Series. Paraconularia plani-
costata.
165 Anonymous. Windsor, Nova Scotia. Lower Windsor Group; Osagian-Chesterian
Series. Paraconularia planicostata.
166 D. G. Kelley, 1954. GSC location 24841, about 100 m east of comer of Route 5
and Buckwheat Road, Nyonza, Cape Breton Island, Nova Scotia. Lower Windsor
Group; Osagian-Chesterian Series. Paraconularia planicostata.
167 D. G. Kelley, 1954. GSC location 24844, limestone at bridge on Lewis Mountain
Road, 0.8 km from Route 19, Cape Breton Island, Nova Scotia. Lower? part of
Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
470
Annals of Carnegie Museum
VOL. 55
168 Anonymous. Nova Scotia. Lower Windsor Group; Osagian-Chesterian Series. Par-
aconularia planicostata.
169 Anonymous. Cape Breton, Nova Scotia. Windsor Group; Osagian-Chesterian Se-
ries. Paraconularia planicostata.
170 M. J. Copeland, 1962. Shore of Bros d’Or Lake, Irish Cove, Cape Breton, Nova
Scotia. Windsor Group; Osagian-Chesterian Series. Paraconularia planicostata.
171 Anonymous. Brookfield, Colchester County, Nova Scotia. Lower Windsor Group;
Osagian-Chesterian Series. Paraconularia planicostata.
172 Anonymous. Harts County, Nova Scotia. Basal Windsor Group; Osagian-Cheste-
rian Series. Paraconularia planicostata.
173 Anonymous. Maxner Point, Nova Scotia. Probably lower part of Windsor For-
mation; Osagian-Chesterian Series. Paraconularia planicostata.
Ohio
1 74 Anonymous. Delaware, Ohio. “Comiferous Group,” probably Delaware Limestone;
Ulsterian Series. Conularia elegantula.
175 G. Meszaros, pre-1982. Rathbone, Ohio. Columbus Limestone; Ulsterian Series.
Conularia elegantula.
176 Hyatt Brothers, Dublin, Franklin County, Ohio. Columbus or Delaware Limestone;
Ulsterian Series. Conularia elegantula.
177 Anonymous. Quarry 4 km southeast of Sylvania, Ohio. Silica Shale; Erian Series.
Conularia sp.
178 G, Meszaros, pre- 1 982. Leroy, Ohio. Chagrin Shale; Chautauquan Series, Conularia
multicostata, Paraconularia chagrinensis.
179 E. Roeser, 1978. Float along Mill Creek, at and near Camp Koinonia, Lake and
Ashtabula counties, Ohio, north and south of Ross Road bridge. Chagrin Shale;
Chautauquan Series. Paraconularia chagrinensis.
180 D. Strock, 1985? Along Mill Creek, from Hidden Valley Park to the church camp.
Lake County, Ohio. Chagrin Shale; Chautauquan Series. Paraconularia chagrinen-
sis.
181 C. Talerico, 1984; L. E. Babcock et al., 1984. Float along Mill Creek, between Ross
Road bridge and small dam upstream of Ross Road, Ashtabula County, New York.
Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
182 T. Stanley, 1984. Mill Creek, within 165 m downstream (north) of Ross Road
bridge. Lake and Ashtabula counties, Ohio. Chagrin Shale; Chautauquan Series.
Paraconularia chagrinensis.
183 M. E. Williams, 1981; S. McKenzie, pre-1982. Float along Mill Creek, Ashtabula
County, Ohio. Chagrin Shale; Chautauquan Series. Paraconularia chagrinensis.
184 M. E, Williams, 1981; J. Hannibal etal., 1985. Stebbins Gulch, Holden Arboretum,
Geauga County, Ohio, Contact between the Chagrin Shale and the Cleveland Shale
Member of the Ohio Formation; Chautauquan Series. Paraconularia chagrinensis.
185 A. J. Weiss, 1984, Landfill on north side of Ohio Route 82, approximately 1.2 km
west of 1-77 interchange, Broadview Heights, Ohio. Cuyahoga Formation, Meadville
Shale Member; Kinderhookian Series. Paraconularia byblis, P. subulata.
186 J. Hall? Alexander, Licking County, Ohio. “Berea Shale” (=Sunbury Shale Sub-
member, Orangeville Member, Cuyahoga Formation); Kinderhookian Series, Par-
aconularia subulata.
187 Herrick? Alexander, Licking County, Ohio. “Berea Shale” (=Sunbury Submember
of the Orangeville Member of the Cuyahoga Formation?); Kinderhookian Series.
Paraconularia subulata.
188 L. E. Babcock, 1984. Sunbury Shale Submember of the Orangeville Member of the
Cuyahoga Formation; 1-3 cm thick silty zone with abundant pyrite at Sunbury-
1986 Babcock and Feldmann—P/I^ coiV[/iv4i?L4 and Reticulaconularia
471
Berea Sandstone contact; Kinderhookian Series. Quarry Rock picnic area, north of
Chagrin River, South Chagrin Reservation, east of Solon Road, Bentleyville, Cuy-
ahoga County, Ohio. Paraconularia subulata.
189 M. Ciccarone, 1984. Sunbury Shale Submember of the Orangeville Member of the
Cuyahoga Formation; Kinderhookian Series. Quarry Rock picnic area, north of
Chagrin River, South Chagrin Reservation, east of Solon Road, Bentleyville, Cuy-
ahoga County, Ohio. Paraconularia subulata.
190 L. E. Babcock, 1984. Sunbury Shale Submember of the Orangeville Member of the
Cuyahoga Formation; approximately 1 m above the top of the Berea Sandstone;
Kinderhookian Series. Quarry Rock picnic area, north of Chagrin River, South
Chagrin Reservation, east of Solon Road, Bentleyville, Cuyahoga County, Ohio.
Paraconularia subulata.
191 G. Meszaros, pre-1982. Weymouth, Ohio. Meadville Shale Member of Cuyahoga
Formation; Kinderhookian Series. Paraconularia subulata.
192 J. Burke, W. J. Hlavin et al., 1967. North Branch of Rocky River near bridge at
junction of Bagdad and Hood Roads, Bagdad, Ohio. Meadville Shale Member of
Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
193 Anonymous. Probably northeast Ohio. Probably Meadville Member of the Cuy-
ahoga Formation; Kinderhookian Series? Conularia multicostata.
1 94 Anonymous. Voorhes Cemetary outcrop, west of Lodi, Ohio. Meadville Shale Mem-
ber of Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
195 R. W. Scott. Lodi, Medina County, Ohio. Cuyahoga Formation, probably Meadville
Member; Kinderhookian Series. Paraconularia byblis.
196 G. Meszaros, pre-1982. Lodi, Ohio. Meadville Shale Member of the Cuyahoga
Formation; Kinderhookian Series. Conularia multicostata, Paraconularia subulata.
197 R. Fisher. Creeks in and near Lodi, Medina County, Ohio. Meadville Member of
the Cuyahoga Formation; Kinderhookian Series. Conularia multicostata, Paracon-
ularia subulata.
198 R. Segedi et al., 1974. Streambed olf Pawnee Road, about 200 m south of U.S.
Route 224, Lodi, Medina County, Ohio. Meadville Shale Member of Cuyahoga
Formation; Kinderhookian Series. Conularia multicostata, Paraconularia subulata.
199 R. Segedi. About 50 m east of bridge on Pawnee Road, just south of U.S. Route
224, Lodi, Medina County, Ohio. Float and in situ specimens from the Meadville
Member of the Cuyahoga Formation; Kinderhookian Series. Conularia multicos-
tata, Paraconularia subulata.
200 L. E. Babcock, 1985. West fork of East Branch of Black River, south of Route 224,
near intersection with Pawnee Road, Homer Township, about 3 km west of center
of Lodi, Medina County, Ohio, Lodi 7.5' Quadrangle. Meadville Shale Member of
Cuyahoga Formation; collected in situ in lowermost bed of siderite concretions
upstream of Pawnee Road bridge; Kinderhookian Series. Conularia multicostata,
Paraconularia subulata.
20 1 A. H. Worthen. Richfield, Ohio. “Kinderhook Formation” (=Cuyahoga Formation,
probably Meadville Member); Kinderhookian Series. Paraconularia subulata.
202 Anonymous. NYSM location 110, Richfield, Summit County, Ohio. “Waverly
Group” (probably Meadville Member of the Cuyahoga Formation); Kinderhookian
Series. Paraconularia subulata.
203 Anonymous. Richfield, Ohio. Waverly Group, probably Meadville Member of the
Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
204 L. E. Babcock, 1984. Float in Meadville Member of the Cuyahoga Formation;
Kinderhookian Series. Tributary to the Cuyahoga River, Furnace Run Metro Park,
west off Route 2 1 , about 1.5 km south of Summit County-Cuyahoga County bound-
ary, Summit County, Ohio. Paraconularia cf P. byblis.
205 Anonymous. Richfield, Ohio. “Waverly Group” (=Meadville Member of the Cuy-
472
Annals of Carnegie Museum
VOL. 55
ahoga Formation); Kinderhookian or Osagean Series. Conularia multicostata, Para-
conularia subulata.
206 A. Winchell. Cuyahoga River gorge, Cuyahoga Falls, Summit County, Ohio. “Near
top of the Waverly Group, water limestone below conglomerate” (=Cuyahoga For-
mation, possibly Meadville Member); Kinderhookian-Osagean Series. Pamconu-
laria subulata.
207 J. Weiss, 1961. Gravel in Akron, Ohio area? Possibly Cuyahoga Formation; Kin-
derhookian or Osagian Series. Paraconularia subulata.
208 Anonymous. “Rocky River bed,” probably Cuyahoga Formation; Kinderhookian
or Osagean Series. Medina, Ohio. Paraconularia subulata.
209 Anonymous. Bagdad, Ohio. Probably Cuyahoga Formation; Kinderhookian or Osa-
gean Series. Conularia multicostata.
210 C. L. Herrick. Near Lyon Falls, Richland County, Ohio. 18-24 m above “Con-
glomerate n”: Osagean Series. Paraconularia byblis.
211 W. P. Cooper, 1890; Anonymous. Portsmouth, Ohio. Waverly Group, probably
the Cuyahoga Formation; Osagean Series. Paraconularia missouriensis.
2 1 2 Anonymous. Wooster, Wayne County, Ohio. “Near the top of the Waverly Group”
(=Wooster Member of the Cuyahoga Formation); Osagean Series. Paraconularia
subulata.
213 Anonymous. Wooster, Wayne County, Ohio. Near top of Waverly Group, possibly
the Wooster Member of Cuyahoga Formation; Osagean Series. Paraconularia su-
bulata.
214 G. Meszaros, pre- 1 982. Wooster, Ohio. Cuyahoga Formation, Wooster Shale Mem-
ber; Osagean Series. Conularia multicostata, Paraconularia subulata.
215 H. E. Wilson. 7.2 km south of Loudonville, Ashland County, Ohio. Probably Woos-
ter Member, Cuyahoga Formation; Osagean Series. Paraconularia subulata.
216 F. Plutte, 1 964. About 4.5 km south of Loudonville, Ashland County, Ohio. Wooster
Member of the Cuyahoga Formation; Osagean Series. Paraconularia subulata.
217 L. E. Babcock, 1984. East facing borrow pit on west side of Route 3, 0.9 km north
of junction with Route 97, just south of Loudonville, Ashland County, Ohio, Greer
7.5' Quadrangle. Wooster Member of the Cuyahoga Formation; Osagean Series.
Conularia multicostata, Paraconularia subulata.
218 J. Hall? Water works in Newark, Ohio. Waverly Group, “base of Division IIP’;
probably Cuyahoga Formation; probably Osagean Series. Conularia multicostata.
219 G. Meszaros, pre- 1982. Rushville, Ohio. Allensville Member of Logan Formation;
Osagian Series. Paraconularia byblis.
220 Bowsher, Savage, and Allen, 1952, Approximately 300 m of elevation below Old
Maid’s Kitchen, in gully about 110 m west of Ohio Edison Dam on north side of
Cuyahoga Gorge, Akron, Summit County, Ohio. Float in Meadville Shale Member
of the Cuyahoga Formation; Kinderhookian Series. Paraconularia subulata.
22 1 Anonymous. Ohio. Possibly Cuyahoga Formation; Mississippian, possibly Osagean.
Conularia multicostata, Paraconularia subulata.
Ill Anonymous. Ohio. “Lower Waverly Group,” Cuyahoga Formation; Osagean Series.
Paraconularia subulata.
223 Stout and Girty, 1 897. Dixon’s Mill, on the Little Scioto River, about 5 km northeast
of Sciotoville, Scioto County, Ohio. “Waverly Group,” probably Wooster Member
of Cuyahoga Formation; Osagian Series. Conularia multicostata, Paraconularia
byblis, P. missouriensis.
224 Carman, Stout, and Carney. Sciotoville, Ohio. Upper part of Cuyahoga Formation,
16.5-23 m below base of the Logan Formation; Osagian Series. Paraconularia
byblisl, P. missouriensis.
225 F. B. Meek; Anonymous. Sciotoville, Ohio. “Waverly Group,” probably Cuyahoga
Formation; Osagean Series. Conularia multicostata.
226 E. B. Andrews, 1869. Sciotoville, Ohio. Uppermost Cuyahoga Formation, Black
1986 Babcock and Feldmann— /’A?L4C6>NCZL4i?/^ and Reticulaconularia
473
Hand Member or lowermost Logan Formation, Byer Member; Osagean Series.
Paraconularia missouriensis.
227 G. Meszaros, pre-1982. Sciotoville, Ohio. Portsmouth Shale; Osagean Series. Con-
ularia muliicostata, Paraconularia missouriensis.
228 Cooper. James Hall’s location 385, Moot’s Run, Licking County, Ohio. Cuyahoga
Formation, probably Wooster Member; Osagean Series. Conularia multicostata.
Oklahoma
229 G. A. Cooper et al., 1952. NE'A Sec. 7 T22N R20E, 5.3 km south of Adair, Mayes
County, Oklahoma. Fayetteville Formation; Chesterian Series. Paraconularia okla-
homaensis.
Ontario
230 C. S. 9.6 km west of Cayuga, Ontario. Upper part of Oriskany Sandstone; Ulsterian
Series. Conularia undulata.
Pennsylvania
23 1 F. M. Swartz. Near Curtin, Pennsylvania. Shriver Chert; Ulsterian Series. Conularia
ulsterensis.
232 F. M. Swartz. Intersection of Delaware and New York State Railroad, 0.8 km west
of mill of Mimsruk Paper Company Experimental Mills, near Curtin, Pennsylvania.
Shriver Chert; 1.05 m below Oriskany Shale; Ulsterian Series. Conularia cf C.
desiderata.
233 F. M. Swartz, 1937. Float on roadcut on road leading through gap east of War-
fordsburg, Pennsylvania. Shriver Formation?; 0.3”1 .7 m above conglomeratic sand-
stone at middle of Shriver-like beds; Ulsterian Series. Conularia ulsterensis.
234 F. M. Swartz, 1937. Roadcut along road leading through gap east of Waifordsburg,
Pennsylvania. 0.3- 1.5 m above conglomeratic sandstone at middle of Shriver-like
beds; Ulsterian Series. Conularia ulsterensis.
235 F. M. Swartz. Road leading north from Schellsburg, Bedford County, Pennsylvania.
Onondaga Shale; Erian Series. Conularia cf C desiderata.
236 S. Albright, 1 98 1 ; B. White, 1 983. Large roadcut on north side of Johnny Bee Road,
about 0.2 km north of intersection with road to Dingmans Falls, Delaware Town-
ship, Pike County, Pennsylvania. Mahantango Formation, approximately Center-
field biostrome level; Erian Series. Conularia desiderata, C. ulsterensis, C. undulata.
237 D. Parris, 1982. Large roadcut on north side of Johnny Bee Road, about 0.2 km
north of intersection with road to Dingmans Falls, Delaware Township, Pike Coun-
ty, Pennsylvania. Manhantango Formation; Erian Series. Conularia undulata.
238 L. Klensch and J. Valenti, 1981. Roadcut across from Bushkill Country Store, 2
km from U.S. Route 209, Lehman Township, Pike County, Pennsylvania. Mahan-
tango Formation; Erian Series. Conularia undulata.
239 L. Decina, 1983. Roadcut on north side of Pennsylvania Route 895, approximately
0.8 km west of Auburn, Schuylkill County, Pennsylvania. Mahantango Formation;
Erian Series. Conularia desiderata.
240 Anonymous. Huntingdon, Huntingdon County, Pennsylvania. Frame Shale Mem-
ber of the Mahantango Formation; Erian Series. No conulariids collected; Hy-
olithes sp.
Quebec
24 1 R. B., 1 862; Anonymous. Grande Greve, Gaspe, Quebec. Grand Greve Limestone;
Ulsterian Series. Conularia cf C desiderata, C. cf C. undulata.
474
Annals of Carnegie Museum
VOL. 55
242 Anonymous. High Falls, Dartmouth River, Gaspe Peninsula. Grande Greve Lime-
stone; Ulsterian Series. Reticulaconularia penouili.
243 Anonymous. Little Gaspe, Quebec. Grande Greve Limestone; Ulsterian Series.
Conularia cf. C. desiderata.
244 Anonymous. Perce Rock, Gaspe, Quebec. Grande Greve Limestone; Ulsterian Se-
ries. Conularia tuzoi.
245. Cape Barre, Quebec. “Cape Barre beds” (=Cape Bon Ami Formation); Ulsterian
Series. Conularia cf. C. desiderata.
246 Anonymous. Gaspe Peninsula, Quebec. Float block of limestone, probably Grande
Greve Limestone; Ulsterian Series. Reticulaconularia penouili.
247 J. W. Beede? Magdelen Islands, Quebec. Formation and series unknown; Missis-
sippian Subsystem. Paraconularia sorrocula.
248 J. W. Beede? Cape le Tron, Grindstone Island, Magdelen Islands, Quebec. For-
mation and series unknown; Mississippian Subsystem. Paraconularia planicostata.
South Dakota
249 L. E. Babcock, 1984. “Slagpile section,” overlooking bridge of Route 14A over
Whitewood Creek, SW ‘A Sec. 13, T5N R3E, Deadwood, Lawrence County, South
Dakota, Deadwood 7.5' Quadrangle. Englewood Formation, dolostone about 20
cm above top of shale-dolostone transitional zone. Paraconularia sp.
Tennessee
250 J. M. Safford. Hickman County, Tennessee. “Waverly Group”; Kinderhookian or
Osagean Series. Paraconularia byblisl
251 A. S. Horowitz, 1966, etc. Roadcuts on both sides of Interstate Highway 40, 8.3
km west of junction of Interstate Highway 40 and U.S. Highway 70 at Monterey,
Putnam County, Tennessee, Monterey 7.5' Quadrangle. Top of Pennington For-
mation; “Kinkaid level,” zone of Pterocrinus tridecibrachiatus Gutschick, just below
a quartz pebble conglomerate of the Pennsylvanian System; Chesterian Series. Par-
aconularia chesterensis.
Utah
252 C. D. Walcott? Divide Bet, American Fork and Snake Creek, Wasatch Mountains,
Utah, Kinderhookian or Osagean Series. Paraconularia chesterensis?
Wisconsin
253 E. E. Teller. Milwaukee, Wisconsin. Probably Milwaukee Formation; Erian Series.
Conularia milwaukeensis.
254 E. E. Teller? Estabrook Park, Milwaukee, Wisconsin. Lindwurm Member of the
Milwaukee Formation; Erian Series. Conularia milwaukeensis.
255 E. E. Teller. Milwaukee Cement Quarry, Berthelet, Wisconsin. Milwaukee For-
mation; Erian Series. Conularia milwaukeenensis.
Appendix B — Measurements
The following values are measurements of selected type and other conulariid specimens
considered representative of each taxon treated herein and in Part A. Species are listed
alphabetically according to species as they are recognized herein. Measurements are listed
in columns across each page. In cases where replicate measurements have been taken at
varying distances from the hypothetical apex, they are listed vertically under the appro-
priate columns. Terms are defined in the “Morphology” section. Abbreviations: L, length.
1986 Babcock and Feldmann --Paeaconularia and Reticulaconularia
475
cm; HL, hypothetical length, cm; AAMj, major apical angle, degrees; AAMn, minor
apical angle, degrees; N, nodes/mm; RLAMj, ratio of right superior : left superior : abut-
ting rods per ten on major face; RLAMn, ratio of right superior : left superior : abutting
rods per ten on minor face; D, distance from hypothetical apex at apicad limit of mea-
surement, applicable to values for R and RA; R, rods/cm; RA, rod angle, degrees; h,
holotype; plh, plastoholotype; 1, lectotype; n, neotype; p, paratype; pi, paralectotype.
Values given in parentheses are approximate; values followed by question marks are
accurate, but it is not certain that they have been properly classified as pertaining to
either the major or minor face.
Conularia congregata Hall
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
NYSM 3483 1
6.1
9.4
17
12
6
0:0:10
0:0:10
5.5
18
10
8.0
16
13
NYSM 3483 pi
6.7
10.7
17
14
7
0:0:10
0:0:10
5.5
17
10
8.5
21
9
NYSM 3483 pi
6.5
10.1
16
14
6
0:0:10
0:0:10
5.0
17
10
8.5
20
12
NYSM 3483 pi
6.3
9.9
17
13
6
0:0:10
0:0:10
8.0
16
10
Conularia delphiensis (Maroney and Orr)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
lUPC 14470-1 h
2.0
(4.8)
—
—
6
1:1:8
1:1:8
6.0
36
10
lUPC 14470-2 p
2.1
(5.1)
~
—
1:0:9
0:1:9
4.5
36
11
lUPC 14470-4 p
2.7
(3.4)
15?
-
-
0:0:10
0:0:10
1.6
42
lUPC 14470-6 p
-
-
-
7
-
-
“
-
Conularia desiderata Hall
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
AMNH 2697 h
1.6
2.3
23
20?
—
0:10:0
0:10:0
3.5
27
15
NYSM 3487
5.8
6.4
27
22
3
1:8:2
1:9:0
5.0
17
12
7.0
14
7
NYSM 3485
6.2
8.2
16
15
3
2:7:1
3:6:1
3.5
24
9
6.5
20
10
USNM 395827
0.9
0.9
22
21
0:0:10
0:0:10
0.6
41
16
Conularia elegantula Meek
SPECIMEN NO.
L
HL
AAMj
AAMn N
RLAMj
RLAMn
D
R
RA
AMNH CU 282G h
3.9
6.7
20
17?
6
0:4:6
—
1.0
39
8
4.0
37
13
CMNH 4584
2.7
3.2
(30?)
—
6
0:6:4?
—
3.5
32
3
CMNH 4648
3.1
(4.5)
-
5
-
-
(4.0) (24)
-
Conularia milwaukeensis Cleland
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
USNM 85988 h
3.1
5.0
15
14
6
0:0:10
0:0:10
6.0
18
5
MPM 20252
4.9
6.0
12
11
6
0:0:10
0:0:10
4.0
21
16
MPM 22974
4.5
4.6
15
13
5
0:0:10
0:0:10
4.0
24
18
6.0
18
8
8.0
20
9
476
Annals of Carnegie Museum
VOL. 55
Conularia multicostata Meek and Worthen
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
USNM 50157 plh
2.8
6.7
20
18
—
0:0:10
0:0:10
4.5
32
16
UK 6089
6.3
8.5
—
11?
3
—
0:0:10?
9.5
52
12
AMNH 6713
6.5
9.3
23
22
2
0:0:10
0:0:10
5.0
40
17
8.0
44
9
CM 34533
8.2
11.2
16
15
2
0:0:10
0:0:10
4.0
25
—
5.5
29
12
Conularia pyramidalis Hall
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
AMNH 33017 1
2.3
3.3
18
17
3
0:0:10
0:0:10
3.0
14
11
5.5
11
16
7.5
9
9
NYSM 3490
11.8
19.2
23
17
1
0:0:10
0:0:10
8.0
7
22
12.0
10
9
15.0
10
5
Conularia subcarbonaria Meek
and Worthen
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
UIPC 10680 h
8.6
11.9
21
19
4
0:0:10
0:0:10
8.0
18
4
FMNH UC 18494
24.5
33.5
10
9
4
0:0:10
0:0:10
18.0
27
5
25.0
31
6
FMNH UC 6289
7.7
-
-
-
3
0:0:10
0:0:10
-
-
-
FMNH UC 6610
3.1
-
-
-
2
-
-
-
60
-
Conularia tuzoi Clarke
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
NYSM 9404 h
6.7
11.3
(10)
—
—
(0:0:10)
—
6.0
26
9
9.0
36
10
Conularia ulsterensis Howell
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
PU 42071 h
1.8
2.5
18?
—
7
2:0:8?
—
4.5
45
13
CM 34528
1.9
3.2
17?
—
7
0:0:10?
—
2.2
60
12
2.8
84
11
CM 34529
1.8
(2.8)
(18)
(15)
6
0:0:10
0:0:10
(2.0)
62
12
Conularia undulata Conrad
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
AMNH 41093 1
10.0
15.5
13
10
6
1:0:9
1:0:9
6.0
21
18
10.0
20
11
13.0
27
4
AMNH 5440
8.3
—
(8)
4
—
—
-
30
-
NYSM 3482
10.2
11.8
18
15
5
0:0:10
0:0:10
4.5
17
9
9.5
24
10
CM 34532
7.5 (18.0)
21
14
5
0:0:10
0:0:10
18.0
32
9
Paraconularia alpenensis Babcock and Feldmann, new species
SPECIMEN NO.
L
HL
AAMj
AAMn N
RLAMj
RLAMn
D
R
RA
GSC 85060 h
1.5
3.6
21
16
_
2:2:6
4:2:4
3.0
14
9
4.0
14
10
1986 Babcock and Feldmann— and Reticulaconularia
All
Paraconularia alternistriata (Shimer)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
GSC 5111 h
0.8
1.9
11
10
0
10:0:0
3:7:0
1.0
28
11
Paraconularia blairi (Miller and Gurley)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
UCGM 3986 1
7.4
11.2
(22)
20
3
6:4:0
3:5:2
5.5
11
8
8.5
7
9
11.0
9
15
UMC 4270
3.4 (11.5)
23?
—
3
7:2:1?
-
(16.0)
6
19
UCGM 3985
6.8 (16.0)
16
2
-
7:3:1
12.5
6
-
UCGM 3984 pi
9.2
15.5
—
18
3
-
5:4:1
8.0
8
15
11.0
6
14
14.0
6
15
AMNH 25056
13.0
19.0
11
10
2
8:1:1
2:6:2
7.0
13
9
12.5
8
12
16.0
7
17
Paraconularia byblis (White)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
UMMP2167 h
6.7
10.0
22
(14)
1
1:4:6
—
6.5
15
12
CMNH 4691
2.8
3.8
26
—
1
1:7:2
—
2.5
28
13
USNM 409489
1.7
2.1
19
14
1
1:6:3
6:2:2
1.0
29
18
Paraconularia chagrinensis Babcock and Feldmann,
new species
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
CMNH 6633 h
3.9
4.9
28
21
4
0:9:1
0:9:1
2.0
20
12
3.0
18
9
4.2
16
10
CMNH 1622 p
3.4
8.3
--
20?
—
0:7:3
0:8:2
5.0
16
9
CMNH 1818 p
2.0
-
-
—
4
—
-
-
-
-
CMNH 1674 p
4.5 (10.0)
-
3
-
-
-
-
Paraconularia chesterensis (Worthen)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
ISGS 2489 h
11.7
17.2
14
10
4
4:5:1
4:2:4
7.5
11
10
10.0
10
9
14.0
8
11
USNM 50156
2.7
3.9
21
17
5
4:3:3
8:2:2
1.0
25
8
2.0
20
8
lUPC 17413
9.7
16.6
(14)
5
(4:5:1)
—
15.5
7
15
lUPC 11313
12.8
25.5
9
9
4
4:5:1
4:5:1
14.0
6
9
20.0
5
9
23.0
6
13
lUPC 6458
9.8
35.5
8
(7)
4
5:2:3
2:6:3
26.0
5
9
30.0
4
11
Paraconularia missouriensis (Swallow)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
FMNH UC 6639 plh
11.0
14.6
19
_
—
1:9:0
—
6.5
6
9
9.5
4
6
478
Annals of Carnegie Museum
VOL. 55
FMNH UC 6628
11.2
15.5
15
11 2
1:9:0
1:8:1
6.0
6
6
13.0
5
10
FMNH UC 6627
12.8
19.5
14
11 3
2:8:0
3:7:0
10.0
6
11
14.5
5
17
19.0
5
11
AMNH 28692
5.7
13.8
-
10 -
—
9:1:0
10.0
6
8
UMMP 26740
3.6
6.4
21
15 2
3:5:2
4:4:2
4.0
10
9
Paraconularia oklahomaensis Babcock and Feldmann, new species
SPECIMEN NO.
L
HE
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
USNM 409801 h
4.4
5.4
19
17 0
4:1:5
2:4:4
2.0
24
12
4.0
12
13
Paraconularia planicostata (Dawson)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
RM(MU) 2749 h
4.5
7.3
1 1?
9 0
6:3:1
5:4:1
3.5
19
12
6.0
13
15
GSC 7715
3.3
4.9
18
14 0
5:4:2
5:3:2
3.0
14
15
GSC 24644
1.6
5.3
13
12 0
3:6:1
5:3:2
3.5
20
12
Paraconularia recurvatus Babcock and Eeldmann,
new species
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
USNM 409806 h
4.8
7.8
16
15 3
2:5:3
2:4:4
4.0
18
12
6.0
(26)
8
USNM 409807 p
0.8
-
-
- 2
-
-
-
28
-
USNM 409808 p
1.1
2.6
16
- 3
3:6:1
3:5:2
1.5
26
8
USNM 409809 p
0.7
-
-
- 3
-
-
-
24
-
Paraconularia salinensis (Whiteaves)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
GSC 4292 h
2.6
3.0
(24)
21 4
7:2:1
5:3:2
1.0
24
13
1.5
23
8
Paraconularia sorrocula (Beede)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
NYSM 9414 h
2.3
2.9
23
- 6
1:7:2
—
0.8
20
11
1.2
19
14
Paraconularia subulata (Hall)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
AMNH 32404 1
0.6
0.8
21
18 4
5: 2:3
6:1:3
0.5
56
11
FMNH UC 6961
1.6
2.4
18
12 5
0:10:0
3:7:0
1.0
35
18
1.5
35
11
UIPC 10866
6.6
7.8
__
12 4
7:1:2
5.0
8
22
UMMP 2178
0.9
(5.5)
—
= 4
—
24
(19)
UMMP 245
0.5
1.4
22
(20) 4
0:7:3
6:3:1
1.0
30
11
OC 8309
5.4
12.8
12
10 2
1:8:1
4:5:1
11.0
8
15
Paraconularia
wellsvillia Babcock and Feldmann,
new species
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
CM 35001 h
5.7
12.5
(14)
(13) 3
1:8:1
5:4:1
8.5
4
26
10.0
5
31
1986 Babcock and Feldmann— il4iL4COivc/i^i?/y4 and Reticulaconularia 479
CM 34502 p
5.8 (12.5)
(18)?
3
3:6:1?
-
7.5
9
(24)
12.0
7
(18)
Paraconularia yochelsoni Babcock and Feldmann, :
new species
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
UMMP 45499 h
2.2
2.6
20
18
0
5:3:2
10:0:0
0.6
15
18
1.5
14
17
UUMP 45500 p
2.6
3.1
17
0
6:1:2
—
0.6
18
15
1.5
13
15
UMMP 65509 o
1.3
1.7
15
-
0
8:1:1
-
0.6
18
20
Reticulaconularia penouili (Clarke)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
NYSM 9412 h
4.6
6.7
59
(22)
2
2:5:3
3:5:2
3.0
14
4
4.5
21
3
6.0
14
0
GSC 87242
5.5
6.4
—
30?
2
—
5:4:1?
1.3
15
7
3.3
12
8
Reticulaconularia sussexensis
(Herpers)
SPECIMEN NO.
L
HL
AAMj AAMn N
RLAMj
RLAMn
D
R
RA
NJSM 10749 h
2.7
3.1
32
26
1
0:2:8
—
2.5
20
11
3.5
36
8
NJSM 10750 p
1.5
1.8
(27)
24
2
1:2:7
2:1:7
0.6
39
14
1.5 16 11
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VOLUME 55 31 DECEMBER 1986 ARTICLE 17
FLORAL MORPHOLOGY AND VASCULAR ANATOMY OF
AMIANTHIUM MUSCAETOXICUM (WALTER) A. GRAY
(LILIACEAEWERATREAE) WITH NOTES ON
DISTRIBUTION AND TAXONOMY
Frederick H. Utech
Abstract
Presentation of pedicel to stigma vasculature of the monotypic Amianthium muscae-
toxicum as a Veratrean example documents the perigyny, spiral insertion of floral parts
and an apocarpous gynoecium which are encountered. Total floral vascularization is
derived from three lower pedicel bundles. A spiral series of divisions and fusions in the
upper pedicel produces three compound outer tepal and three inner tepal bundles. From
the former, outer tepal medians, tepal laterals, stamen and dorsal bundles result, while
from the latter, inner tepal medians, tepal laterals and stamen bundles result. Due to
perigyny, their formation is above that at which the ventral and ovular supplies are
established. The ventral supply is spirally derived from continuing bundles following
the formation of the compound inner tepal bundles. Within each septal arm, a compound
lateral is associated with a compound ventral and both undergo radial divisions. Opposing
lateral and ventral products fuse as the perigynous condition ends and the three carpels
are freed. Each carpel has two simple ventrals, two laterals and a dorsal in the upper
perigynous zone and two fusion ventrals and a dorsal in the upper freed zone. Neither
tepal glands nor nectaries occur in this species. Epidermal cells characterized by ho-
mogenous tannins occur in the pedicel, receptacle wall, abaxial tepal surfaces, filaments
and freed carpels. Raphides commonly occur in cells along the lower margins of the
inner tepals.
Introduction
Amianthium muscaetoxicum (Walter) A. Gray is a monotypic species
of eastern North America with centers of distribution in both the Ozark-
Submitted 15 February 1986.
481
482
Annals of Carnegie Museum
VOL. 55
Fig. 1. — Distribution of Amianthium muscaetoxicum in eastern United States based on
published maps (Steyermark, 1963; Radford et al., 1968; Johnson, 1969; Smith, 1978;
Wherry et al., 1979) and collections at Carnegie Museum of Natural History (open circles).
Johnson (1969) surveyed 46 herbaria in his southeastern United States treatment of the
Liliaceae. Sampled populations are indicated by circled stars.
ian and Appalachian regions (Fig. 1). It occurs in mesic to dry wooded
slopes and coastal plain savannas and pinelands. Familial and subfa-
milial names associated with A. muscaetoxicum have changed during
the last century, but the tribal association with Melanthium, Schoen-
caulon, Stenanthium, Veratrum and Zigadenus “sensu lato” (including
Toxicoscordion, Anticela, Tracyanthus and Oceanoros) has remained.
Bentham and Hooker (1883) included these six genera in the tribe
1986
Utech —Amianthium
483
Veratreae, as did Engler (1889), Krause (1930), Melchior (1964), Thome
(1968), Takhtajan (1969), and Hutchinson (1934, 1959, 1973). Within
this tribal grouping, two lines —the Veratrum-Melanthium and the Zig-
adenus lines— are generally recognized (Anderson, 1940; Preece, 1956;
Zimmerman, 1958; Kupchan et al., 1961; Ambrose, 1975, 1980; Ster-
ling, 1982). Amianthium and Stenanthium are generally placed some-
where between these two evolutionary lines. Engler (1 889) initiated the
association of the Veratreae with the subfamily Melanthioideae. This
subfamily has been twice segregated as a separate family, the Melan-
thiaceae (Gates, 1918; Small, 1933; Dahlgren, 1980; Dahlgren and
Clifford, 1982; Dahlgren and Rasmussen, 1983; Dahlgren et al., 1985)
and the Colchicaceae (Baker, 1879).
Genera in the other tribes of the Englerian Melanthioideae share
separate styles and septicidal capsules with the genera in the Veratreae,
though the latter possesses unusual extrorse anthers with valvular de-
hiscence that open into peltate discs (Krause, 1930; Zimmerman, 1958;
Kupchan et al., 1961) and usually many bitegmic, basitropic and cam-
pylotropous ovules per carpel (Sterling, 1982), except for Amianthium
which has two, rarely four.
Amianthium of Asa Gray (1837) is a conserved generic name (Farr
et al., 1979; Voss, 1983) and its monotypic species, A. muscaetoxicum
(Walter) A. Gray (1837), has gone under various names since it was
first described by Walter in 1788 as Melanthium muscaetoxicum. Later
synonyms include: Melanthium laetum Solander in Aiton (1789), Me-
lanthium myoctonum J. F. Gmelin (1796), Helonias erythrosperma
Michaux (1803), Helonias laeta (Solander in Aiton) Kew-Gawler (Cur-
tis, 1805), Amiantanthus muscaetoxicum (Walter) Kunth (1843), Zig-
adenus muscaetoxicus (Walter) Regel (1883), and Chrosperma mus-
caetoxicum (Walter) Kuntze (1891). Excluded species of Amianthium
and their current status include: A. nuttallii A. Gray var. alpha (1837)
{Zigadenus nuttallii A. Gray ex S. Watson), A. nuttallii A. Gray var.
beta (1837) (Z. paniculatus (Nutt.) S. Watson), A. angustifolium A.
Gray (1837) (Z. densus (Desr.) Femald), A. leimanthoides A. Gray
(1837) (Z. leimanthoides A. Gray), and^. texanum (Bush) Gates (1918)
(Z. leimanthoides A. Gray).
Considerable biological information is implied in this species bi-
nomial. Amianthium is derived from the Greek amianthos for “un-
spotted” and anthos for “flower,” an allusion to a major generic char-
acter, that is the lack of glands or nectaries on the perianth. The specific
name, muscaetoxicum, translates literally as “fly poison.” In 1883, a
note in Gardeners’ Chronicle “stated that the root, when bmised and
mixed with honey, acts as a poison to flies.” While the species is
commonly known as “fly poison,” it has also been known as “crow
poison” and “swagger grass” (Muenscher, 1939, 1960). Several poi-
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sonous alkaloids are associated with the leaves and underground bulbs
of the species and are reported to cause death in cattle and sheep
(Pammel, 191 1; Marsh etal., 1918; Muenscher, 1939, 1960; Kingsbury,
1964). Amianthine, a steroidal or modified steroidal alkamine alkaloid
with a C27H41O2N formula, has been described from the roots and
leaves of A. muscaetoxicum as well as jervine and two unidentified
ester alkaloids (Neuss, 1953). The latter alkaloids were shown to be
responsible for the high toxicity of the species (Neuss, 1953). Although
numerous and highly poisonous alkaloids commonly occur throughout
the tribe Veratreae, amianthine has only been reported in Amianthium
(Kupchan et al., 1961; Willaman and Schubert, 1961; Hegnauer, 1 963).
Two meiotic chromosome counts of « = 16 have been reported for
A. muscaetoxicum: Ambrose (1975) from Bear Creek, Pennsylvania
and Preece (1956) from Big Laurel Gap, Yancey County, North Car-
olina. Excluding various polyploid multiples, Melanthium, Veratrum
and Zigadenus (^Zygadenus; Preece, 1956) all share n = 16 (Fedorov,
1969; Moore, 1973; Goldblatt, 1981). It is quite apparent that x ^ 8
is the basic number unifying the Veratreae.
In focusing on both the floral morphology and vascular anatomy of
A. muscaetoxicum a comparative model of a Veratrean gynoecium will
be established in this paper. Similar treatments for the other tribal
members of the Veratreae, such as Melanthium, Stenanthium, Vera-
trum and Zigadenus, are in preparation and will continue the studies
of Buxbaum (1925, 1927), Anderson (1940), El-Hamidi (1952), Am-
brose (1975, 1980) and Sterling (1982) on the vascularization of the
Veratrean carpel.
Materials and Methods
Flowering and fruiting inflorescences of Amianthium muscaetoxicum were collected
from two different populations— Pennsylvania: Clinton County, Mt. Tableland, ca. 4.5
mi E of Sinnemahoning, 20 July 1979, Utech 79-241 (CM), and North Carolina: Macon
County, ca. 3.5 mi N of Highlands, near Whiteside Mt., Nantahala National Forest, 28
July 1982, Utech and Ohara 82-270 (CM). The collected materials were fixed in acetic-
ethanol (1:3) for 10 h with subsequent storage in 70% ethanol. Standardized paraffin
sectioning (14-16 microns) and staining (saffarin-methylene blue) techniques (Johansen,
1940; Sass, 1958) were used on samples 00 flowers and 10 young fruits of varying ages)
from both populations. As an additional check on these serial sections, whole flowers
and fruits were cleared and stained in a NaOH-1% fuchsin mixture (Fuchs, 1963).
Composite photomicrographs (Figs. 3-5, 7-8, 10) present the vascular floral anatomy
and morphology of A. muscaetoxicum, whereas Figs. 6 and 9 are summary line diagrams
for the species. No teleological implications are intended in the descriptive ascent and
departure of the various floral bundles which are letter-coded for ease in comparison.
This coding parallels that used in our previous liliaceous studies (Utech, 1978^-1978^,
1979a, \919b, 1982, 1 984; Utech and Kawano, 1975, 1976, 1980, 1981).
Observations
Amianthium muscaetoxicum is a glabrous, subscapose, perennial
herb from a thick bulb (Fig. 2). At anthesis, the linear basal leaves are
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Fig. 2.— -Flowering individuals of A. muscaetoxicum in Macon County, North Carolina
(Utech and Ohara 82-270, CM) (scale indicated).
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Fig. 3. -Cross-sections from the pedicel and lower receptalar areas of A. muscaetoxicum
showing the spiral opening of the locules, the degree of perigyny and distribution of
epidermal tannin cells. A. Mid-pedicel section showing the formation of three compound
outer tepal (OT) bundles. B. Upper pedicel section showing formation of three compound
inner tepal (IT) bundles and departure of compound OT bundles. Due to spiral bundle
formation, a fusion bundle is opposite the upper OT bundle and a gap opposite the lower
left OT bundle (arrows). C. Transition between upper pedicel and lower receptacle areas
showing he opening of two locules, the departure of three IT bundles and the remaining
central bundles which form the ventral supply. D. Lower receptacle area with perigyny
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shorter than the stem. The few cauline leaves are progressively reduced
and bract-like. The bracteate raceme is at first ovoid to conic, but at
maturity it becomes cyclindric, measuring 4-14 cm long by 2-4 cm
wide. Initially the flowers are white, but following anthesis the persis-
tent tepals turn yellowish green or greenish purple and present a most
striking color pattern.
The flowering pedicels are normally 7-17 mm long and ascending.
In fruit, the pedicels are decidedly horizontal and much elongated.
Internally, the fruiting and flowering pedicels have the same bundle
number, though the vasculature in fruit is surrounded by a scleren-
chymatous sheath. Lower flowering pedicel cross-sections are broadly
triangular with three large, centrally arranged bundles. These three
bundles establish the complete floral vasculature and are located on
the radii which run from the middle of the “flat side” to the section’s
center. These three radii are designated the outer tepal (OT) radii. The
three radii from the “comers” to the center are designated the inner
tepal (IT) radii. Mid-pedicel cross-sections are characterized by broad
fluting and ridges (Figs. 3 and 9) which continue through the perigynous
zone.
The three lower pedicel bundles along OT radii undergo tri-parted,
radial divisions with three resulting bundles from each division. These
divisions occur at slightly different levels, that is they are not co-planar,
but rather in a spiral pattern. Within each division a gap is created by
the outward departure of a central bundle product. Three such central
bundles, designated compound outer tepal (OT) bundles (dorsal-com-
pound bundle; Sterling, 1982) depart along OT radii and remain free
of other vasculature. Eventually they establish the dorsals (D), the outer
tepal medians (OTM), the outer tepal laterals (OTL), and the outer
stamen (OS) bundles (Figs. 6, 9A-D).
The two resulting lateral bundles which are opposite a gap following
the tri-parted divisions fuse with similar adjacent laterals along the IT
radii. These fusion bundles are formed in a spiral pattern and undergo
a tri-parted division similar to that observed at a lower level among
the three original bundles. The central bundles of this second set of
divisions depart along the IT radii and establish the compound inner
tepal (IT) bundles (“zwischenbundel”; Sterling, 1982).
evident and three open locules with formation of the central ventral supply. E. Mid-
receptacle area showing the central hole and the ventral supply. F. Upper receptacle area
showing the spiral septal arm formation within the perigynous zone, inter-locular con-
nections and placental supply. Dorsal bundles are not established at this level (scale
indicated).
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Fig, 4.— Cross-sections from the lower receptacle to upper perigynous areas of A. mus-
caetoxicum. A. Lower receptacle section showing the departure of compound OT and
IT bundles, opening of two locules and formation of three bundles opposite IT bundles
which establish the ventral supply. B, Section above A showing three open locules and
the central hole. While three IT bundles depart, fusion and division occur among the
remaining septal arm bundles. C. Section above B showing three freed septal arms
between the inter-connected locules. Each septal arm has two laterals (L) and a compound
ventral (V) from which placental bundles (P) arise. D. Section above C showing ovule
supply via placental (P) bundles and the paired laterals and ventrals within the septal
arms. E. Section above D showing the division of the compound ventral bundle (V) and
the two laterals (L) within a septal arm. Papilloid nurse cells lining the inner septal arm
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From the second set of tri-parted divisions, the two remaining lateral
bundles fuse laterally with adjacent laterals, close the OT radii gaps
formed when the compound OT bundles departed and eventually es-
tablish the total ventral supply (Fig. 6). These three fusion bundles
undergo simple radial divisions in a spiral pattern. The two resulting
products of this radial division fuse laterally with a similar adjacent
lateral and form three fusion bundles along IT radii which close the
gaps formed during the departure of the compound inner tepal (IT)
bundles. Subsequent divisions among the compound OT and IT bun-
dles will be discussed later. A given pedicel cross-section will usually
show one bundle departing, another being formed and a gap in an area
where a third will be formed.
The gynoecium and its associated vasculature can best be described
in two parts: that in the lower perigynous zone and that in the upper
freed zone. Ventral supply formation, locule opening and ovule pla-
centation all occur within the lower perigynous zone (Figs. 3C“F; 4,
6). The dorsal bundles, on the other hand, which are derived from
compound outer tepal (OT) bundles are established in the upper limits
of the peripheral perigynous zone.
The ridged outline observed in the lower pedicel occurs up through
the upper perigynous region where the tepals and stamens are freed
(Figs. 3, 4F). Following the formation and departure of both the com-
pound OT and IT bundles, a triangular vascular zone (“stele”) remains
in the central area (Figs. 3C-D, 4A-B). The three large bundles along
IT radii establish the triangle’s comers. These three complex fusion
bundles which closed the gaps formed by the departure of compound
IT bundles (Fig. 6) generally have two phloem caps or poles. Once
these three comer bundles are established, the three locules which are
perpendicular to OT radii open spirally.
With locular opening, each of the three compound bundles along IT
radii undergoes a simple tangential division (Figs. 4B-C, 5A, C). Both
resulting bundles are along the IT radii. The outermost division product
has normally arranged xylem (adaxial) and phloem (abaxial), while the
inner product has reversed conducting elements. Phloem strands fre-
quently are observed to lag and may rarely and irregularly anastomose
with other bundles (Fig. 5A-B).
margins are weakly appressed. F. Upper periygnous section showing the spiral arrange-
ment of tepals, stamens and partially, freed carpels. Dorsals (D) are established at this
level. Paired lateral (L) and ventral (V) products within septal arms are indicated as are
outer tepal medians (otm), inner tepal medians (itm), outer stamen (os) and inner stamen
(is) bundles. Epitepaly between an inner tepal and inner stamen is shown in the upper
right (scale indicated).
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Fig. 5. -Cross-section of the middle to upper gynoecium of A. muscaetoxicum. A. Central
zone showing three appressed septal arm tips and radial arm separation with paired
lateral (L) and ventral (V) bundles. B. Section, same level as A, showing an outer stamen
(OS) bundle and dorsal (D). An inner carpellary wall indentation, not a notch, is associated
with each dorsal. C. Section above A showing the separated gynoecium and perianth.
Within a septal arm, a given ventral and opposing lateral fuse to form a new, compound
central in the same position as the lower, simple ventral. D. Section above C showing
two compound ventrals (V) and dorsal (D) per carpel. The inner septal arm tips are
further divided (arrow) compared with A. E. Section above D showing three free carpels
each with a dorsal (D) and two ventrals (V). F. Stylar zone with the dorsal (D) and two
ventrals (V) still present in each carpel (scale indicated).
The appearance at this level of a central opening or ‘"hole” along the
central floral axis (Figs. 3E, 4B) indicates inter-connection of the three
locules will follow. Openings develop from this hole along the OT radii
to each of the three locules (Fig. 3F). As openings inter-connect the
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491
three locules, three septal arms or wings are established along the IT
radii. Within these septal arms further divisions occur among the paired
ventral supply bundles. The outermost member of each septal bundle
pair, that is the one with normally arranged conducting elements, di-
vides radially and establishes two lateral bundles (L) (Figs. 4C-D, 5A,
6). (These bundles, here designated as laterals (L), could also be called
septal axials.) The innermost member of each septal bundle pair, that
is the one with reversed conducting elements, also undergoes a radial
division which establish two placental (P) or ovule supplying bundles.
The parental bundle which established the two placental (P) bundles
also undergoes a radial division and establishes two ventral (V) bundles.
Each septal arm at this level has a pair of laterals (L) with normally
arranged conducting elements and a pair of ventrals (V) with reversed
elements (Figs. 4C-F, 6).
Each locule has two, rarely four, bitegmic, basitropic, campylotro-
pous ovules. The placental (P) bundles supplying these ovules depart
quite horizontally. Usually one, rarely two, ellipsoidal, lustrous, dark
reddish brown to black seeds are found in each carpel. The wingless
seeds measure 1.5--2.0 mm wide by 4. 5-6.0 mm long. Dehiscence
begins in the upper stylar area and continues along the zones where
the septal arms or margins meet (Fig. 5D-F).
The three dorsals (D) are the last carpellary vascular elements to be
established and this occurs in the upper perigynous zone. The dorsals
are associated with a zone of parenchyma cells which protrude as a
locular indentation, not a notch (Figs. 4F, 5B, D-E, 7B). As the septal
arms are subdivided along IT radii (Figs. 4E-F, 5 A), the outer carpellary
wall is freed from the perigynous zone. Septal arm subdivision follows
the changing distribution of epidemal tannin cells which line the outer
carpellary wall margins (Fig. 5A, C, D). Within each subdivided septal
arm, two sets of vascular fusions occur. The two laterals (L) fuse with
two opposite ventral bundles (V) which are along the same radii. The
two resulting fusion bundles, here designated as ventrals (V), are in the
same location as the two lower, simple ventrals (V) (Figs. 5A, C-D,
6). It should be noted that while the ventrals prior to fusion and after
fusion are designated in the same way, there is a difference. The later
ventral is a terminal fusion product. With the formation of these fusion
ventrals (V), the three carpels are freed from one another (Fig. 5D-F)
in what can best be described as an apocarpous condition. Each freed
carpel has a dorsal (D) and two ventrals (V) which continue into the
stylar zone (Fig. 5F). There is no terminal carpellary fusion between
the ventrals or between the ventrals and the dorsal.
In most liliaceous species with a superior ovary, both the tepal and
stamen vasculatures are well established before the locules open. This
is not the case in A. muscaetoxicum (Figs. 3C-F, 9) and other members
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V
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493
of the Veratreae due to the basal perigynous condition. The six freed
perianth parts, that is, the three outer and three inner tepals which are
spirally inserted (Fig. 4F), are separate to their respective bases and
spread widely. The inner tepals which measure 5-7 mm are slightly
longer than the outer tepals. There are no glands or claws associated
with these perianth parts as is frequently reported in Melanthium and
Veratrum. The abaxial tepal surfaces of all six tepals are characterized
by a single epidermal layer containing homogenous tannins (Fig. 8D),
This epidermal tannin layer also occurs in the filaments and freed
gynoecium. In addition to these tannin cells, there are randomly scat-
tered tannin cells throughout the floral tissue. The adaxial tepal sur-
faces, on the other hand, lack these tannin cells and instead have
papilloid cells (Fig. 8D). These papilloid cells occur from the basal
regions of the freed tepals to the apices.
There are other differences between the inner and outer tepals (Figs.
7, 8). The inner tepals are differentiated from the outer in having a
short vertical zone of enlarged cells along the basal tepal margins (Figs.
4F, 8B, C). These enlarged cells frequently contain long raphides. Ep-
itepaly between the inner stamens and tepals is the rule (Figs. 4F, 8A,
B). Epitepaly between the outer stamens and tepals occurs, but it is not
as pronounced as the former.
Vascularization of the six tepals and six stamens occurs above the
level at which the ovules are supplied (Figs. 3C-F, 4F, 7-9). This is
due to the typical perigynous condition within the Veratreae. Six com-
pound bundles, derived via fusion and established in the pedicel and
lower receptacle levels, are ultimately responsible for the complete
vascularization of the tepals and stamens (Figs. 6, 9). These compound
bundles have been designated as the compound outer tepal (OT) and
compound inner tepal (IT) bundles, respectively, for they are located
along those respective radii. Insertion and departure of tepals and
stamens follow a spiral pattern (Figs. 4F, 6, 9). A vascular description
for one compound OT and IT bundle will illustrate the patterns for
both sets since they are free from each other.
In the upper perigynous zone, a compound OT bundle appears tri-
angular in cross-section (Figs. 7A, 9). Several complex subdivisions
occur within the compound OT bundles which results in the formation
Fig. 6. — Roll-out longitudinal summary diagram for the floral vasculature of A. mus-
caetoxicum. A spiral pattern is indicated, in part, by the shifted levels at which the
compound OT and IT bundles depart. Various text discussed bundles have been given
the following code: L = lateral, V = ventral (simple and compound) and P = placental.
Those bundles derived from the compound OT and IT bundles are not shown, see Figs.
7-9.
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495
of an outer tepal median (OTM), two outer tepal laterals (OTL), an
outer stamen bundle (OS) and a dorsal (D). The OTM is established
first and has normally arranged xylem (adaxial) and phloem (abaxial)
(Figs. 7A, 9B). The remaining product, after the departure of the OTM,
undergoes a rapid and complex subdivision in which two OTLs, an
OS and a D are established (Figs. 7, 9). The OTM, OS and D bundles
all lie along the same radius. It should be noted that the outer tepal
laterals are derived from the remaining product bundles, not the OTM.
Basally each outer tepal receives three bundles, an OTM and two OTLs.
The laterals undergo further radial divisions to establish additional
laterals (Fig. 7C). In freed outer tepals, a seven bundled condition is
typical, that is three OTLs + OTM + three OTLs.
The origin of the inner tepal median (ITM), the two inner tepal
laterals (ITL), and an inner stamen bundle (IS) from a compound inner
tepal bundle (IT) is similar to the outer series (Figs. 8, 9). The vascu-
larization of the inner tepals and stamens is in a spiral pattern. The
ITM, two ITLs and IS bundles have normally arranged xylem (adaxial)
and phloem (abaxial). Basally each inner tepal is supplied with three
bundles, an ITM and two ITLs. The laterals undergo further radial
divisions, as in the outer series, to establish additional laterals (Fig.
8C). In freed inner tepals, a five bundled condition, rarely seven, is
encountered, that is two ITLs + ITM + two ITLs. There is no fusion
between the laterals or between laterals and medians in either tepal set.
Tepal bundles follow a parallel course and end along tepal margins.
The six equal stamens are nearly as long as the inner tepals. The
filament’s epidermal layer contains tannins (Figs. 7D, 8C, lOA), while
the anther walls do not. The basifixed, extrorse anthers have a valvular
(lateral) dehiscence between the confluent thecae which open into a
peltate disc (Fig. 10). The endothecium has wall thickenings or bands
of the girdle type (Dahlgren and Clifford, 1982). This type of anther
and mode of dehiscence is characteristic of the Veratreae.
The difference between outer and inner stamen vascularization which
arose from compound OT and IT bundles, respectively, is that a dorsal
Fig. 7.— Vascularization of the outer tepals and stamens in A. muscaetoxicum. A. Mid-
perigynous zone showing the departure of an outer tepal median (OTM) from a compound
OT complex. B. Section above A showing the further division of the compound OT
bundle into two outer tepal laterals (OTL) and an outer stamen (OS) bundle. The locular
indentation associated with the dorsal (D) is evident. C. Upper perigynous section above
B showing further division among the outer tepal laterals (OTL). D. Section above C
showing the freed outer tepal and outer stamen (scale indicated).
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497
is associated with division of the OT parental bundle and there is no
counterpart with the IT compound bundle (Figs, 7B-D, SA-C, 9),
Discussion and Conclusions
In describing the vascular floral anatomy and carpel morphology
within selected members of the liliaceous tribe Veratreae, Anderson
(1940), El-Hamidi (1952), Ambrose (1975), Sterling (1982), and Utech
in this report have noted characteristic similarities and differences be-
tween the flowers and fruits of Amianthium muscaetoxicum and the
other members of the Veratreae. A common pattern of vascularization
appears to be consistently observed within the tribe, while major dif-
ferences which are frequently used to differentiate genera are reported.
These anatomical and morphological characters include the presence
or absence of sutural openings and a central carpellary hole at the
lowermost level of ovular insertion, the degree of carpellary separation
below the locular apex (an apocarpous tendency), the number of car-
pellary lateral or septal axial bundles and a hypogynous or perigynous
versus epigynous condition at the lowermost level of ovular insertion.
The floral vascular anatomy and carpel morphology of A. muscaetox-
icum as reported here will serve as a case study of the continuous
vascularization of a Veratrean gynoecium for further comparative work
within the tribe.
The pedicel to stigma vasculature of A. muscaetoxicum is most un-
usual in that perigyny, spiral insertion of floral parts and apocarpous
gynoecia are encountered. Through a complex series of divisions and
fusions the complete floral vasculature is derived from three lower
pedicel bundles. In the middle to upper pedicel a series of successive
divisions and fusions in a spiral sequence produces three compound
outer tepal (OT) bundles and three compound inner tepal (IT) bundles.
From each compound OT bundle (dorsal-composite bundle; Sterling
(1982)), an outer tepal median (OTM), several outer tepal laterals (OTL),
Fig. 8.— Vascularization of the inner tepals and stamens in A. muscaetoxicum with a
tepal surface comparison. A. Epitepaly between an inner tepal and stamen is shown as
an inner tepal median (ITM) is derived from a compound IT bundle. Subsequent division
of the IT bundle establishes two inner tepal laterals (ITL) and inner stamen (IS) bundle.
An epidermal tannin layer surrounds both the tepal and stamen. B. Section above A
showing epitepaly as well as specialized cells which frequently contain raphides along
the inner tepal margin (arrow). These cells are lacking in the outer tepals. C. Section
above B showing a freed inner stamen and tepal with specialized cells (arrow). D. Section
above C showing the difference in adaxial (tannins) and adaxial (pa = papilloid) tepal
surfaces. These papilloid cells occur throughout both adaxial surfaces and extend to the
tepafs tips (scale indicated).
so a
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& Dorsal
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an outer stamen (OS) and a dorsal (D) result. Similarly, from each
compound IT bundle (‘'zwischenbundel”; Sterling (1982)), an inner
tepal median (ITM), and an inner stamen (IS) bundle result. Due to
the perigynous condition, the formation of these tepal, stamen and
dorsal bundles is at a level above that at which the ventral and ovular
supplies are established.
The spiral origin of the ventral supply is via successive divisions and
fusions of continuing lateral branches following the formation of the
compound IT bundles. Within each of the three undivided septal arms,
that is, the perigynous zone where the three locules are not inter-
connected, a lateral bundle (L) (compound septal axial bundle; Sterling
(1982)) is associated with a ventral (V) (compound placental bundle;
Sterling (1982)). The lateral bundle has normally arranged xylem and
phloem while the ventral bundle has reversed conducting elements.
There is a radial subdivision within each septal arm of both the ventral
and lateral bundles. Each ventral bundle fuses with an opposite lateral
bundle as the perigynous condition ends and the three carpels are freed.
It is noteworthy that at a lower level each carpel has five bundles,
that is, a dorsal (D), two laterals (L) and two ventrals (V) while at a
higher and freed carpellary level there is only a dorsal (D) and two
fusion ventrals (V). Terminally, there is no fusion within a carpel be-
tween the dorsal and ventrals or between the ventrals. While the ter-
minology varies, the above observations are similar to those of Sterling
(1982). Anderson (1940) noted that the carpels in Amianthium, Me-
lanthium, Veratrum, and Zigadenus were supplied by a dorsal, two
laterals and two ventrals. The cross-section drawings of Amianthium
by Ambrose (1975; fig. 32G-K) were described as having one dorsal
and four ventrals. Simply counting carpellary bundles from selected
cross-sections is not adequate for detailed comparative purposes, rather
the continuity of the complete floral pattern including origins, fusions
and divisions must be followed.
The combination of spirally inserted floral parts as well as their
spirally derived vasculature coupled to both a perigynous and apocar-
pous gynoecium is most unusual among the “supposedly primitive
lilies.” Furthermore, there is a central carpellary hole associated with
the gynoecial base. This central hole is internally continuous with the
Fig. 9. “Line drawing showing cross-sections from the upper perigynous zone and the
resulting vascularization from both a compound OT bundle (A-D) and a compound IT
bundle (E-H). The outer tepal median (OTM), outer tepal laterals (OTL), outer stamen
(OS) and dorsal (D) bundles are all derived from the compound OT bundle, while the
inner tepal median (ITM), inner tepal laterals (ITL), and inner stamen (IS) bundles are
derived from the compound IT bundle.
500
Annals of Carnegie Museum
VOL. 55
1986
IJtecr—Amianthium
501
three locules and the open stylar canal. Such a central hole according
to Sterling (1982) could be taken to represent a partially closed suture
or a remnant of an open portion of the stylar canal. The three carpels
are essentially free above the perigynous zone. The inner septal wing
tips that form the stylar canal are weakly differentiated into papilloid
nurse cells. In this inner zone, dehiscence occurs exposing the single,
rarely two, wingless seeds of each carpel. Such inner septal margin
separation is a variation of the typical septicidal capsule dehiscence
where splitting occurs between two adjacent carpels along a common
septal radius.
Neither tepal glands nor nectaries occur in Amianthium though they
are commonly encountered throughout the Veratreae. The epidermal
layers of both abaxial tepal surfaces as well as that of the pedicel, the
complete perigynous zone, the filaments and the freed carpels are char-
acterized by cells with homogenous tannins. However, the adaxial sur-
faces of both the inner and outer tepals have a generalized epidermis
of small papilloid cells. Furthermore, as a possible defensive adapta-
tion, raphides are observed in cells along the lower, outer margins of
the inner tepals. Though there has been limited differentiation between
the adaxial and abaxial tepal surfaces, they are not specialized as nec-
taries or glands. Travis (1984) observed that foraging beetles are the
chief pollinators for A. muscaetoxicum and experimentally demon-
strated that this species is nearly self-incompatible and the fecundity
and fruit set levels are at least partly pollinator limited. Furthermore,
only a small percentage of seeds from self-pollinated plants are viable.
Acknowledgements
The author would like to recognize and thank the M. Graham Netting Research Fund
and the O’Neil Botany Field Fund of the Carnegie Museum of Natural Flistory for
supporting the field work and the lab related preparation of materials and photographs
in the new Biosystematics Laboratory. Mr. William W. Brown and Ms. Nancy J. Perkins
deserve special thanks for their artistic aid in figure production.
Literature Cited
Aiton, W. 1789. Hortus Kewensis. George Nicol, London, Ed. 1, 1:125.
Ambrose, J. D. 1975. Comparative anatomy and morphology of the Melanthioideae
(Liliaceae). Unpublished Ph.D. dissert., Cornell University, Ithaca, vii + 240 pp.
Fig. 10.— Cross-sections of an outer stamen in A. muscaetoxicum. A. Section through a
filament with a tannin epidermal layer and two lower anther thecae which lack tannins.
B. Section above A showing the central basifixed position of the filament. C. Section
above B showing the central position of the outer stamen (OS) bundle. D. Upper anther
zone section above C showing confluent thecae which open into a peltoid disc (scale
indicated).
502
Annals of Carnegie Museum
VOL. 55
— . 1980. A re-evaluation of the Melanthioideae (Liliaceae) using numerical anal-
yses. Pp. 65-81, in Petaloid monocotyledons (C. D, Brickell, D. F. Cutler, and M.
Gregory, eds.). Academic Press, London, ix + 222 pp.
Anderson, C. E. 1 940. Some studies on the floral anatomy of the Liliales. Unpublished
Ph.D. dissert., Cornell University, Ithaca, iv + 142 pp.
Baker, J. G. 1879. A synopsis of Colchicaceae and aberrant tribes of Liliaeeae. Journal
of the Linnean Society, Botany, 17:405-510.
Bentham, G., and J. D. Hooker. 1883. Genera Planatarum. L. Reeve, London, 3:
834-836.
Buxbaum, F. 1925. Vergleichende Anatomie der Melanthioideae. Repertorium Spe-
cierum Novarum Regni Vegetabilis, 29:1-80.
. 1927. Nachtrage zur vergleichenden Anatomie der Melanthioideae 1. Beihefte
zum Botanischen Centralblatt, 44:255-263.
Curtis, T. 1805. Helonias laeta. Channel-leaved Helonias. Curtis’s Botanical Maga-
zine, t. 803.
Dahlgren, R. 1980. A revised system of classification of the angiosperms. Journal of
the Linnean Society, Botany, 80:91-124.
Dahlgren, R., and H. T. Clifford. 1 982. The monocotyledons. A comparative study.
Academic Press, London, xiv + 378 pp.
Dahlgren, R., H. T. Clifford, and P. F. Yeo. 1985. The families of monocotyledons.
Structure, evolution, and taxonomy. Springer- Verlag, New York, xii + 520 pp.
Dahlgren, R., and F. N. Rasmussen. 1 983. Monocotyledon evolution: characters and
phylogenetic estimation. Evolutionary Biology, 16:255-395.
El-Hamidi, a. 1952. Vergleichend-morphologische Untersuchungen am Gynoeceum
der Unterfamilien Melanthioideae and Asphodelioideae der Liliaceae. Arbeiten aus
dem Institut fiir allgemeine Botanik, Universitat Zurich, ser. A, no. 4:1-50.
Engler, a. 1889. Liliaceae. Pp. 10-30, in Die naturlichen Pflanzenfamilien (A. Engler
and K. Prantl, eds.), Engelmann Verlag, Leipzig, 2:10-91.
Farr, E. R,, J. A. Leussink, and F. A. Stafleu. 1979. Index Nominum Genericomm
(Plantarum). Vol. l:Aa-Epochnium. Regnum Vegetabile, 100:67.
Fedorov, A. A. (ed.) 1969. Chromosome numbers of flowering plants. Nauk, Lenin-
grad, 926 pp. (in Russian)
Fuchs, C. 1963. Fuchsin staining with NaOH clearing for lignified elements of whole
plants or plant organs. Stain Technology, 38:141-144.
Gardeners’ Chronicle. 1883. Amianthium muscaetoxicum. Vol. 20:41, col. 2.
Gates, R. R. 1918. A systematic study of the North American Melanthaceae from a
genetic standpoint. Journal of the Linnean Society, Botany, 44:131-172.
Gmelin, j. F. 1796. Sy sterna vegetabilia, secundum classes differentiis. Bemuset, De-
lamolliere, Falque and Soc., Lyon, 1:587.
Goldblatt, P. (ed.) 1981. Index to plant chromosome numbers 1975-1978. Mono-
graphs in Systematic Botany, 5:300-321.
Gray, A. 1837. Melanthacearum Americae Septentrionalis Revisio. Annals of the
Lyceum of Natural History of New York, 4:105-140.
Hegnauer, R. 1963. Chemotaxonomie der Pflanzen. Birkhauser Verlag, Basel, 2:269-
359.
Hutchinson, J. 1934. The families of flowering plants. Vol. 2: Monocotyledons,
MacMillan and Co., London, Ed. 1, 234 pp.
, 1959. The families of flowering plants. Vol. 2: Monocotyledons. Clarendon
Press, Oxford, Ed. 2, 260 pp.
. 1973. The families of flowering plants. Vol. 2: Monocotyledons. Clarendon
Press, Oxford, Ed. 3, 268 pp.
Johansen, D. A. 1 940. Plant microtechnique. McGraw-Hill Book Co., New York, xi +
523 pp.
1986
Utech —Amianthium
503
Johnson, R. G. 1969. A taxonomic and floristic study of the Liliaceae and allied
families in the southeastern United States. Unpublished Ph.D. dissert., West Virginia
University, Morgantown, xii + 334 pp.
Kingsbury, J. M. 1964. Poisonous plants of the United States and Canada. Prentice-
Hall, Englewood Cliffs, pp. 448-466.
Krause, K. 1930. Liliaceae. Pp. 227-276, in Die naturlichen Pflanzenfamilien (A,
Engler and K, Prantl, eds.), Engelmann Verlag, Leipzig, 2(15a):227-390.
Kunth, C. S. 1843. Enumeratio Plantarum. Hinrichs, Tubingen, 4:180.
Kuntze, C. E. O. 1891. Revisio Generum Plantarum. Arthur Felix, Leipzig, 2:708.
Kupchan, S. M., J. H. Zimmerman, and A. Alfonso. 1961. The alkaloids and tax-
onomy of Veratrum and related genera. Lloydia, 24: 1-26.
Marsh, C. D., A. B. Clawson, and H. Marsh. 1918. Staggergrass {Chrosperma mus-
caetoxicum) as a poisonous plant. United States Department of Agriculture Bulletin,
no. 7101.
Melchior, H. 1964. Syllabus der Pflanzenfamilien. Vol. II: Angiospermen. Gebruder
Bomtraeger, Berlin, vi -T 666 pp.
Michaux, a. 1803. Flora Boreali-americana. Carol! Crapolet and Fratres Levrault,
Paris, 1:212.
Moore, R. J. (ed.) 1973. Index to plant chromosome numbers 1967-1971. Regnum
Vegetabile, 90:105-127.
Muenscher, W. C. 1939. Poisonous plants of the United States. MacMillan Co., New
York, pp. 50-51.
— . 1960. Poisonous plants of the United States. MacMillan Co., New York, Ed.
2, pp. 44-52.
Neuss, N, 1953. A new alkaloid from Amianthium muscaetoxicum Gray. Journal of
the American Chemical Society, 75:2772-2773.
Pammel, L. H. 1911. A manual of poisonous plants. Torch Press, Cedar Rapids, pp.
374-385.
Preece, S. j., Jr. 1956. A cytotaxonomic study of the genus Zigadenus. Unpublished
Ph.D, dissert., State College of Washington, Pullman, v -I- 167 pp.
Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the vascular flora of
the Carolinas. University of North Carolina Press, Chapel Hill, pp. 302-304.
Regel, E. 1883. Zigadenus muscaetoxicus. Gartenflora, 32:164, t. 1121, fig. 1.
Sass, j. E. 1958. Botanical microtechnique. Iowa State University Press, Ames, xi +
228 pp.
Small, J, K. 1933. Manual of the southeastern flora. Privately printed. New York, p.
278.
Smith, E. B. 1978. An atlas and annotated list of the vascular plants of Arkansas.
Privately printed, Fayetteville, iv + 592 pp.
Sterling, C. 1982. Comparative morphology of the carpel in the Liliaceae: Veratreae.
Journal of the Linnean Society, Botany, 84:57-77.
Steyermark, j, 1963. Flora of Missouri. Iowa State University Press, Ames, pp. 420-
421.
Takhtajan, a. 1969. Flowering plants. Origin and dispersal. Olive and Boyd, Edin-
burgh, X + 3 1 0 pp.
Thorne, R. F. 1968. Synopsis of a putative phylogenetic classification of the flowering
plants. Aliso, 6:57-66.
Travis, J. 1984. Breeding system, pollination and pollinator limitation in a perennial
herb, Amianthium muscaetoxicum (Liliaceae). American Journal of Botany, 7 1 :94 1-
947.
Utech, F. H. 1978^?. Floral vascular anatomy of Medeola virginiana L. (Liliaceae-
Parideae = Trilliaceae) and tribal note. Annals of Carnegie Museum, 47:169-191.
. 1978Z). Comparison of the vascular floral anatomy of Xerophyllum asphode-
504
Annals of Carnegie Museum
VOL. 55
hides (L.) Nutt, and X. tenax (Pursh) Nutt. (Liliaceae-Melanthioideae). Annals of
Carnegie Museum, 47:147-167.
. 1978c. Vascular floral anatomy of Helonias bullata L. (Liliaceae-Helonieae)
with a comparison to the Asian Heloniopsis orientalis. Annals of Carnegie Museum,
47:169-191.
— . 1978cf. Floral vascular anatomy of Pleea tenuifolia Michx. (Liliaceae-Tofiel-
dieae) and its reassignment to Tofieldia. Annals of Carnegie Museum, 47:423-454.
1978c. Floral vascular anatomy of monotypic Japanese Metanarthecium lu-
teoviride Maxim. (Liliaceae-Melanthioideae). Annals of Carnegie Museum, 47:455-
477.
. 1 919 a. Floral vascular anatomy of the Himalayan Theropogon pallidus Maxim.
(Liliaceae-Convallarieae). Annals of Carnegie Museum, 48:43-71.
— \919b. Floral vascular anatomy of Scoliopus bigelovii Torrey (Liliaceae-Pari-
deae = Trilliaceae) and tribal note. Annals of Carnegie Museum, 48:43-71.
. 1982. Floral vascular anatomy of the east Asian Rohdea japonica (Thunb.) A.
W. Roth (Liliaceae-Convallarieae). Journal of Phytogeography and Taxonomy, 30:
27-36.
. 1984. Floral vascular anatomy of Japonolirion osense Nakai (Liliaceae) and
its tribal relationship. Annals of Carnegie Museum, 53:447-461,
Utech, F. H., and S. Kawano. 1975. Biosystematic studies in Erythronium (Liliaceae-
Tulipeae) IL Roral anatomy of E. japonicum Decne. Botanical Magazine (Tokyo),
88:177-185.
. 1976. Biosystematic studies in Maianthemum (Liliaceae-Polygonatae) VIII.
Vascular floral anatomy of M. dilatatum, M. bifolium and M. canadense. Botanical
Magazine (Tokyo), 89:145-157.
-. 1980. Vascular floral anatomy of the Japanese Paris tetraphylla A. Gray (Lil-
iaceae-Parideae). Journal of Phytogeography and Taxonomy, 28:17-23.
. 1981. Vascular floral anatomy of the east Asian Heloniopsis orientalis (Thunb.)
C. Tanaka (Liliaceae-Helonieae). Botanical Magazine (Tokyo), 94:295-31 1.
Voss, E. G. (ed.) 1983. International Code of Botanical Nomenclature adopted by the
Thirteenth International Botanical Congress, Sydney, August 1981. Regnum Ve-
getabile, 111:324.
Walter, T. 1788. Flora Carolinana. J. Fraser, London, pp. 125.
Wherry, E. T., J. M. Fogg, Jr., and H. A. Wahl. 1979. Atlas of the flora of Penn-
sylvania. Morris Arboretum, Philadelphia, xxx + 390 pp.
WiLLAMAN, J. J., AND B. G. ScHUBERT. 1961. Alkaloid-bearing plants and their con-
tained alkaloids. United States Department of Agriculture Technical Bulletin, no,
1234.
Zimmerman, J. H. 1958. A monograph of Veratrum. Unpublished Ph.D, dissert..
University of Wisconsin, Madison, iii + 321 pp.
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