Russian Devonian brachiopod genera Cyrtinoides and Komiella in North America

Abstract

The Middle Devonian brachiopod genus Cyrtinoides, described from the southern Urals, is a senior subjective synonym of Mucroclipeus, previously known only from eastern and western North America. The Middle and Late Devonian cyrtinid brachiopod genus Komiella, previously known only from the Timan Range of eastern Europe, is identified from west-central Alaska and Nevada. A new family Komiellidae is proposed. New species are Komiella gilberti, K. magnasulca, and K. stenoparva. Known species of both genera occupy carbonate platform foreslope facies or shelf basins, allowing open marine migration via peripheral biofacies.

Full text

J. Paleont., 67(6), 1993, pp. 952-958
Copyright ? 1993, The Paleontological Society
0022-3360/93/0067-0952$03.00
RUSSIAN DEVONIAN BRACHIOPOD GENERA CYRTINOIDES AND
KOMIELLA IN NORTH AMERICA
J. G. JOHNSON AND R. B. BLODGETT
Department
of Geosciences,
Oregon
State
University,
Corvallis 97331 and
U.S. Geological
Survey,
National Center/MS
970, Reston, Virginia
22092
ABSTRACT-The Middle Devonian
brachiopod
genus Cyrtinoides,
described from the southern
Urals,
is a senior
subjective
synonym
of Mucroclipeus,
previously
known
only from
eastern
and western North
America. The Middle
and Late Devonian
cyrtinid brachiopod
genus
Komiella,
previously
known
only from the Timan Range
of eastern
Europe,
is identified from west-central
Alaska
and
Nevada.
A new family Komiellidae is proposed.
New species are Komiella
gilberti,
K. magnasulca,
and K. stenoparva.
Known species of
both genera occupy
carbonate
platform foreslope
facies or shelf basins,
allowing
open marine
migration
via peripheral
biofacies.
INTRODUCTION
M
Ck 'IDDLE CAMBRIAN to Late Devonian biogeographic pat-
I..
? terns oftrilobites, brachiopods, corals, and conodonts
were basically similar in that they were composed of platform
and peripheral biofacies. Platform faunas tended to be isolated,
whereas peripheral faunas of the northern hemisphere were rel-
atively cosmopolitan and surrounded one or several platforms"
(Johnson, 1983, p. 1). For these reasons, widely distributed
Paleozoic brachiopod genera are no longer labeled cosmopolitan
until analysis shows that they cross significant biogeographic
boundaries. The two genera analyzed here gained wide distri-
bution within a single peripheral biofacies.
Locality descriptions are given in the Appendix.
SYSTEMATIC PALEONTOLOGY
Superfamily CYRTIOIDEA
Frederiks, 1924
[nom. transl. Pitrat, 1965
(ex Cyrtiinae Frederiks, 1924, p. 312)]
Discussion. -Although the ambocoeliid brachiopods have been
assigned to various superfamilies since they were first recognized
at subfamily rank by George (1931), we follow Pitrat (1965) in
recognition that the Ambocoeliidae were derived from a cyrtiid
stock, a conclusion recently affirmed by Johnson and Lenz (1992).
Family AMBOCOELIIDAE George, 1931
[nom. transl. Ivanova, 1959, p. 56
(ex Ambocoeliinae George, 1931, p. 42)]
Subfamily AMBOCOELIINAE George, 1931
[Ambocoeliinae George, 1931, p. 42]
Genus CYRTINOIDES
Iudina and Rzhonsnitskaia, 1985
Figure 1
Type species. - Cyrtinoides ajica Iudina and Rzhonsnitskaia,
1985, p. 82, P1. 2, figs. 10-12, P1. 3, figs. 1-3.
Subjective synonym. -Mucroclipeus Goldman and Mitchell,
1990, p. 85; type species, M. eliei Goldman and Mitchell, 1990,
p. 88.
Diagnosis. -The original diagnosis and comments by Iudina
and Rzhonsnitskaia (1985) can be paraphrased as follows (from
a translation by Mrs. Josephine Cooper): small, smooth, sharply
inequivalved shells with a semi-pyramidal ventral valve and
very slightly convex, almost flat dorsal. Beak of ventral valve
sharp, upright, area high, triangular, with a wide open delthyrial
aperture. In ventral valve dental plates and a tichorhinum are
united; in dorsal valve a cardinal process and crural plates di-
rected obliquely toward center of valve and almost reaching its
floor. Shell substance impunctate.
With the aid of material from New York and Nevada, de-
scribed previously (Goldman and Mitchell, 1990, p. 85-89, figs.
6, 15; Johnson et al., 1980, p. 99, P1. 1, figs. 19-26), we rein-
terpret the type species as lacking crural plates. The structures
interpreted as crural plates by Iudina and Rzhonsnitskaia ( 1985,
P1. 2, figs. 1-3, reprinted here as our Figure 1) are recumbent
crural bases. The tichorhinum of Cvrtinoides is open, without
being bisected by the median septum as is the case in Cvrtina.
Comparison. -No other ambocoeliid has a tichorhinum.
Species assigned. -Cnrtinoides ajica Iudina and Rzhonsnit-
skaia, 1985, Vucroclipeus eliei Goldman and Mitchell, 1990,
and Echinocoelia septata Johnson (in Johnson et al., 1980, p.
99, P1. 1, figs. 19-26).
Geographic distribution. -Western North America (Nevada),
eastern North America (western New York), and eastern Europe
(western slope of Southern Urals).
Stratigraphic distribution. -The type species is reported from
the Givetian Bol'shearshinsk series. In Nevada, Cyrtinoides sep-
tata ranges from the upper part of costatus Zone (Eifelian) to
Lower disparilis Subzone (upper Givetian). In New York, C'.
eliei is reported by Goldman and Mitchell (1990) from the Le-
vanna Shale Member of the Skaneateles Formation (lower Give-
tian).
Biofacies. -The Levanna Shale Member of the Skaneateles
Formation is a gray shale of probable dysaerobic, basinal bio-
tope (Brett et al., 1991). In Nevada, occurrences in the Denay
Limestone represent carbonate platform middle shelf and fore-
slope facies (facies belts 7 to 3 of Wilson, 1975; Johnson, 1990).
Biofacies of the Russian occurrence is uncertain.
Superfamily CYRTINOIDEA
Frederiks, 1912
[nom. transl. Johnson, 1966, p. 177
(ex Cyrtininae Frederiks, 1912)]
Family KOMIELLIDAE n. fam.
Diagnosis. -Nonplicate Cyrtinoidea with sessile jugum; lack-
ing a tichorhinum.
Genus KOMIELLA
Liashenko, 1985
Type species. -Komiella devonica Liashenko, 1985, p. 14, P1.
2, figs. 5, 6.
Diagnosis. -The original diagnosis and comments by Lia-
shenko (1985) can be paraphrased as follows (from a translation
by Mrs. Josephine Cooper): small, triangular or trapezoidal,
half-pyramidal, smooth shells with a high, half-pyramidal ven-
tral valve and flat dorsal valve, with narrow sulcate grooves on
both valves. Interarea high, triangular, flat in lower part, slightly
concave in upper, with a high narrow delthyrium. Teeth and
dental plates form a spondylium in ventral valve. Punctate shell
structure. Liashenko assigned Komiella to the Cyrtinidae.
Based on material from Alaska and Nevada, described below,
there are only a few changes and additions that need to be made.
Dental flanges, lacking adminicula (not dental plates), may join
952

JOHNSON AND BLODGETT-DEVONIAN BRACHIOPODS
the ventral median septum to form a spondylium or the three
plates may be free. The dorsal interior, not described by Liashen-
ko, has widely set, deeply incised sockets and a simple, knoblike
cardinal process, which may be supported on a short median
septum. Rudimentary crural plates have been observed. A ses-
sile jugum is located anteriorly.
Comparison. -No other cyrtinid genus is smooth-bisulcate;
no other genera of the superfamily lack a tichorhinum or have
a sessile jugum.
Species assigned.-Komiella devonica Liashenko, 1985; K.
gilberti n. sp.; K. magnasulca n. sp.; and K. stenoparva n. sp. In
addition, Spirifer trochiloideus Gortani, 1911, may belong to
Komiella, but it is unknown internally.
Geographic distribution. -Western North America (Alaska,
Nevada) and eastern Europe (Timan Range).
Stratigraphic distribution.--Upper part of Eifelian costatus
Zone to upper part of Frasnian (within M. N. zones 10-12 of
Klapper, 1989).
Biofacies.--the Lyaiol'sk series, in which the type species oc-
curs, is a basinal facies, according to G. Klapper (personal com-
mun.), who has studied conodonts from cores in that unit. The
new species from Alaska had a probable lagoonal biotope, as-
sociated with abundant snails and dasyclad algae. Amphipora
were evidently proximal because Amphipora beds occur strat-
igraphically close to the beds with Komiella. Both new species
from Nevada, named and described below, are from the Denay
Limestone, a carbonate platform foreslope facies (facies belts 5
to 3 ofWilson, 1975). With the possible exception of the Alaskan
specimens, the known occurrences of Komiella are in peripheral
biofacies (Johnson, 1990, p. 928).
KOMIELLA GILBERTI n. sp.
Figure 2
Diagnosis. -Komiella of trapezoidal outline, with faint bisul-
cation and fully formed spondylium.
Etymology. -For Wyatt G. Gilbert, formerly of the Alaska
Division of Geological and Geophysical Surveys, in recognition
of his contribution to the knowledge of the geology of west-
central Alaska.
Material. -Six articulated specimens, five ventral valves, and
one dorsal valve from 79RB12 (USGS 10063-SD), all silicified.
Exterior. -Very small, hemipyramidal (cyrtinaform), trape-
zoidal, equidimensional; well-defined, triangular, catacline to
procline ventral interarea; delthyrium narrow, triangular, and
open; dorsal interarea very small, ill defined; faint bisulcation;
surface smooth except for growth lines; commissure deflected
ventrally.
Ventral interior.-Dental flanges, lacking adminicula, join
ventral median septum to form spondylium, bisected medially
by dorsal edge of median septum.
Dorsal interior. -With widely set, deeply incised sockets and
simple, knoblike cardinal process supported by short median
septum; T-shaped sessile jugum located anteriorly.
Two illustrated specimens show the sessile jugum (Figure 2.14,
FIGURE
1-1-3, Cyrtinoides
ajica Iudina and Rzhonsnitskaia,
1985;
sections
copied from their Plate 3, figures
1-3. The top right
section,
at 2.4 mm, we interpret
to lack crural
plates. There were no figure
legends
with the original
illustrations.
2.18); each of these has fragmentary, arcuate segments of the
spiral brachidium still attached, with broken tips pointing ven-
trally.
Shell structure.
-Although the specimens are coarsely silici-
fled, several exhibit a porous texture that appears to be evidence
of endopunctae (Figure 2.11, 2.12, 2.19).
Comparison. -Komiella gilberti differs from the type species,
K. devonica Liashenko, in having a trapezoidal outline com-
pared to the nearly triangular outline of K. devonica. Otherwise
the two species are closely similar. Other comparisons are given
below, following the descriptions of two other new species.
Occurrence and age. -Cheeneetnuk Limestone (Blodgett and
Gilbert, 1983), McGrath A-5 quadrangle, west-central Alaska,
loc. 79RB 12 (USGS 10063-SD); early Eifelian. See Blodgett and
Rohr (1989, fig. 2) and Poncet and Blodgett (1987, fig. 2) for
location.
KOMIELLA MAGNASULCA n. Sp.
Figures 3, 4
indet. n. gen., n. sp. JOHNSON, KLAPPER,
AND TROJAN, 1980, table 3, p.
86, table 12, p. 91.
indet. spiriferid
n. gen. JOHNSON,
1990, fig. 24, p. 916.
Diagnosis. -Komiella of transverse outline, with strong bisul-
cation and dental flanges free of median septum.
Etymology. -Referring to the prominent median sulci on both
valves.
Material.--Thirteen specimens from VW-49 and one speci-
men from VW-56, all coarsely silicified.
Exterior. -Very small, hemipyramidal (cyrtinaform), trape-
zoidal, transverse; well-defined, triangular, catacline to procline
ventral interarea; delthyrium triangular and open; dorsal inter-
area very small, ill defined; strong bisulcation; surface smooth
FIGURE
2-1-19, Komiella
gilberti
n. sp., Cheeneetnuk
Limestone,
McGrath
A-5 quadrangle,
west-central
Alaska,
loc. 79RB12 (USGS 10063-
SD). All figures
x 10
except 19, x 19. 1-3, posterior, lateral,
and
anterior views of articulated
specimen,
USNM 466416, holotype;
4-7, posterior,
lateral,
anterior,
and dorsal views of articulated
specimen, USNM 466417; 8, 9, posterior
and dorsal
views of articulated
specimen,
USNM
466418; 10, anterior view of ventral
valve, USNM 466419; 11, 12, exterior
and interior
views of dorsal
valve, USNM 466420; 13, interior
view of dorsal
valve, USNM 466421; 14, oblique posterior
view of articulated
specimen,
USNM 466422; 15-17, posterior,
anterior,
and
interior
views of ventral
valve, USNM 466423; 18, 19, oblique posterior
view of interior
and dorsal view of articulated
specimen,
USNM 466424.
FIGURE
3-1-14, Komiella
magnasulca
n. sp., lower member
of the Denay Limestone,
northern
Antelope
Range,
Eureka
County,
Nevada, loc.
VW-49. All figures
x 10. 1-6, posterior,
anterior,
interior,
dorsal,
lateral,
and oblique interior
views of free ventral
valve, USNM 466425; 7-
10, posterior, anterior,
ventral, and lateral views, USNM 466426; 11-14, anterior,
oblique interior,
interior,
and posterior
views, USNM
466427.
953

JOURNAL OF PALEONTOLOGY, V. 67, NO. 6, 1993
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JOHNSON AND BLODGETT-DEVONIAN BRACHIOPODS
except for faint growth lines; commissure slightly deflected ven-
trally.
Ventral interior.--Dental flanges strong, curved, lacking ad-
minicula, free of median septum.
Dorsal interior. -With widely set, deeply incised sockets and
simple, knoblike cardinal process joined to crural bases by shell
thickening and supported by short median septum; short crural
plates converge medially.
A sessile jugum was not observed, but a small protuberance
at the anterior midline may be the site of its former attachment.
Shell structure. -Not observed.
Comparison. -Komiella magnasulca differs from the type
species, K. devonica Liashenko, in having a trapezoidal outline
compared to the nearly triangular outline of K. devonica. It
differs from Komiella gilberti in having a transverse outline and
stronger bisulcation and in the absence of a spondylium. An-
other comparison is given below, following the description of
Komiella stenoparva.
Occurrence and age. -Sample VW-49 is in the upper part of
the Eifelian costatus Zone; VW-56 is in the basal bed of the
Eifelian australis Zone.
KOMIELLA
STENOPARVA
n. sp.
Figure 5
n. gen. bilobate
cyrtinid
sp. JOHNSON, KLAPPE, MURPHY,
AND
TROJAN,
1986, table 5, p. 185.
indet. spiriferid
n. gen. JOHNSON, 1990, fig. 34, p. 921.
Diagnosis. -Komiella of minute size and narrow relative width.
Etymology. -Referring to the unusually small size and rela-
tively narrow outline of the valves.
Material.-One specimen from H64-3 (not previously re-
ported), two specimens from DCM III 39, one specimen from
DCM III 40, and one specimen from TA X-34 (Johnson, 1990,
fig. 34, p. 921), all silicified.
Exterior. -Minute, hemipyramidal (cyrtinaform), trapezoi-
dal, higher than wide; well-defined, triangular, procline ventral
interarea; delthyrium triangular and open; dorsal interarea very
small, ill defined; strong bisulcation; surface smooth except for
faint growth lines; commissure slightly deflected ventrally.
Ventral interior.-Not observed.
Dorsal interior. -With widely set, deeply incised sockets and
simple, knoblike cardinal process lacking support of median
septum; vestigial crural plates; sessile jugum not observed.
Shell structure. -Not observed.
Comparison. -Komiella stenoparva is smaller, relatively more
narrow, and has a higher ventral interarea than the other named
species.
Occurrence and age. - H64-3 and TA X 34 are both in the
upper Givetian Lower disparilis Subzone. DCM III 39 and 40
are not directly dated by associated conodonts, but are from the
same part of the section as H64-3, which is nearby, and belong
to the same fauna, which was assigned to the Choperella Com-
munity (Johnson, 1990, fig. 34, p. 921).
DISCUSSION
AND CONCLUSIONS
The newly recognized distributions of Cyrtinoides and Komi-
ella cross province and realm boundaries and might be regarded
simply as cosmopolites, but a label is not an explanation. Bio-
4
7 10
FIGURE
5-1-10, Komiella stenoparva n. sp., upper member of the
Denay Limestone, Eureka County, Nevada. All figures x 10. 1-4,
posterior, anterior, lateral, and dorsal views of articulated specimen,
USNM 466432, loc. DCM III 40, holotype, northern Roberts Moun-
tains; 5, 6, exterior and interior views of dorsal valve, USNM 466433,
loc. DCM III 39, northern Roberts Mountains; 7, 8, exterior and
interior views of dorsal valve, USNM 466434, USNM loc. 36732,
northern Antelope Range; 9, 10, exterior and interior views of dorsal
valve, USNM 466435. USNM loc. 17795, northern Roberts Moun-
tains.
facies analysis, described above for both genera, reveals that
they occupied carbonate platform foreslope facies (facies belts
5 to 3 of Wilson, 1975), predominately but not exclusively. They
were therefore in a peripheral biofacies (Johnson, 1983, 1990,
p. 926), free to migrate around the margins of continental shelves
or around the slopes of carbonate platforms that were built upon
them. In the example of the gray shale occurrences of western
New York, the shale basin must have had direct access to the
southern continental margin of North America as depicted in
an Eifelian lithofacies map (Klapper and Johnson, 1980, p. 430,
text-fig. 4).
ACKNOWLEDGMENTS
We thank Mrs. Josephine Cooper for translations from the
Russian and Rex Doescher for providing relevant literature. A
suggestion by John Carter about the identity of the sessile jugum
of Komiella was especially helpful. John Carter and Norman
Savage reviewed the manuscript. Blodgett acknowledges Wyatt
G. Gilbert, who graciously provided him with helicopter support
during field work in 1979 in the McGrath quadrangle, west-
central Alaska. Johnson's research was supported by NSF Grant
EAR-9204772.
REFERENCES
BLODGETr,
R. B., AND
W. G. GILBERT.
1983. The Cheeneetnuk Lime-
stone, a new Early(?) to Middle Devonian formation in the McGrath
FIGURE
4-1-19, Komiella
magnasulca
n. sp., lower member of the Denay Limestone,
northern
Antelope
Range,
Eureka
County,
Nevada, 1-14,
loc. VW-49, 15-19, loc. VW-56. All figures
x 10. 1-5, posterior, anterior,
oblique interior,
interior and dorsal
views of ventral
valve, USNM
466428, holotype;
6-11, posterior,
anterior, ventral,
interior,
oblique
interior,
and lateral views of ventral
valve, USNM 466429; 12-14, exterior,
interior, and anterior views of dorsal valve, USNM 466430; 15-19, posterior, anterior, oblique interior, interior, and ventral views, USNM
466431.
957
Iri
,?~~~

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ACCEPTED
6 AUGUST
1992
APPENDIX
LOCALITY DESCRIPTIONS
Alaskan locality
79RB12 (USGS 10063-SD). Lat. 62005'15"N, long. 155?07'27"W,
SE'i4, SW'4, NE/4, NWI/4 sec. 15, T34N, R32W, Seward Meridian,
McGrath A-5 quadrangle. Rubble crop of silicified fossils just above a
band of prominent white dolomite on the south side of an east-dipping
slope; collected by R. B. Blodgett, 1979.
Nevada localities, northern Antelope Range
TA V section: ascends east from a point 4,800 ft N, 2,300 ft W of
SE corner of section 21 to middle hill between hill 7602 and hill 7829.
Section plotted in Johnson et al. (1980, fig. 3) and Johnson et al. (1986,
fig. 3). VW-49 (110 ft in section, upper part of costatus Zone) and VW-
56 (139 ft in section, base of australis Zone), both from lower part of
the Denay Limestone; collected by J. L. Waldman, 1987.
TA X section: from top of lower member of the Denay Limestone,
ascends ENE from a point, elev. 7,720 ft, 2,550 ft N, 600 ft W of SE
corner of section 21, toward S slope of VABM 7829. Section plotted in
Johnson et al. (1986, fig. 4). TA X-34, USNM loc. 36732 (34 ft in
section, Lower disparilis Subzone), from lower part of upper member,
Denay Limestone; collected by J. P. Graham, 1982.
Nevada localities, northern Roberts Mountains
DCM III section: ascends west from a point, elev. 6,720 ft, 4,050 ft
S, 5,350 ft E, to a point, elev. 6,840 ft, 4,200 ft S, 4,450 ft E of NE
corner of section 22, T24N, R50E, Eureka County. Sample 39 at 448
ft in section; sample 40 at 533 ft in section, from upper part of upper
member, Denay Limestone; collected by M. A. Murphy, 1979-1981.
H64-3, USNM loc. 17795 (Lower disparilis Subzone), north side of
small saddle at elev. 7,200 ft, approx. 550 ft E of hill 7625 in SEI/4
of
T24N, R50E, between Willow and Dry Creeks, upper member of the
Denay Limestone, northern Roberts Mountains. Collected by M. A.
Murphy, 1975, 1976.
958