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 Lucas, S.G. and Zeigler, K.E., eds., 2005, The Nonmarine Permian, New Mexico Museum of Natural History and Science Bulletin No. 30.

208
TAXONOMY AND BIOSTRATIGRAPHY OF CONCHOSTRACA (BRANCHIOPODA,
CRUSTACEA) FROM TWO NONMARINE PENNSYLVANIAN AND LOWER PERMIAN
LOCALITIES IN NEW MEXICO

THOMAS MARTENS1 AND SPENCER G. LUCAS2

1
Museum of Nature, Stiftung Schloss Friedenstein Gotha, PSF 100319, 99853 Gotha, Germany;
2
New Mexico Museum of Natural History, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104

Abstract—The conchostracans are small, bivalved branchiopod crustaceans and very common in most Upper
Carboniferous and Lower Permian nonmarine sediments. The first fossil conchostracans described from New
Mexico were those from the Atrasado (Wild Cow) Formation (Pennsylvanian). Here, we describe conchostracans
from two new localities: NMMNH locality 3922 at Placitas is in the upper part of the Abo Formation. NMMNH
locality 4667 is in the Atrasado Formation in Socorro County, and yields Lioestheria carinacurvata sp. nov. At
locality 3922, well preserved conchostracan carapaces are similar to the known species Lioestheria monticula
MARTENS, 1983 from Germany. This supports biostratigraphic correlation of the upper part of the Abo Forma-
tion in northern New Mexico with the Lioestheria monticula-Zone of the Tambach Formation (Upper Rotliegend)
of the Thuringian Forest, the Wadern Formation (Upper Rotliegend) in the Saar-Nahe-Basin (both in Germany)
and the Archer City Formation of the Bowie Group (Wolfcampian) of Texas.

INTRODUCTION
Conchostracans are one of the most important fossil faunal groups
in nonmarine sediments representing lacustrine to semiaquatic condi-
tions. This fossil group is widely distributed over all nonmarine facies
from the Devonian to the present time (Jones, 1862, Martens, 1996).
Because of the limited occurrence of suitable facies, Devonian
conchostracans were published from only around the Old Red conti-
nent. Carboniferous, Permian and Triassic conchostracans were pub-
lished from all of the present continents, mostly from Europe and North
America, and Jurassic and Cretaceous conchostracans were published
mostly from Siberia and China. In the Upper Carboniferous and Lower
Permian sediments of North America, conchostracans were described
by Raymond (1946), Copeland (1957), Tasch (1958, 1960, 1961a, 1961b,
1961c, 1962, 1964, 1975) and Martens (1985b, 1986).
Conchostracans are very common in different types of nonma-
rine fine-grained sediments (siltstone, mudstone, limestone) indepen-
dent of the color of the sediment (from black shale to reddish-brown
mudstone). The first fossil conchostracan fauna from New Mexico was
collected and described as Pseudestheria sp. from the Pine Shadow
Member of the Atrasado (= Wild Cow) Formation (Late Pennsylva-
nian, early Virgilian) at the Kinney Brink Company Quarry, Manzanita
Mountains in Bernalillo County (Kozur, Lucas & Hunt 1992).
In the present study we begin with a systematic examination of
the Upper Carboniferous and Permian conchostracans from New
Mexico. This examination will be part of the taxonomic revision of all
described Carboniferous/Permian species from North America and
Europe, discovered in the last 150 years. After revision of these species
and after the analysis of phylogenetic effects, we can use the biostrati-
graphic significance of the conchostracans for the correlation of the
Carboniferous and Permian formations around the world and for the
correlation of the Carboniferous/Permian boundary from marine strati-
graphic levels to nonmarine formations in both New Mexico and in
north-central-Texas.
For the taxonomic analysis, the quality of fossil conchostracan
preservation is most important. In the past, in most of the papers about
the taxonomy of conchostracans the authors described more or less well
preserved carapaces, but without three dimensional reconstruction draw-
ings, or they described poorly preserved, deformed or distorted cara-
paces with incomplete drawings and without reconstructions of the cara-
pace. Only three dimensional reconstruction drawings of all typical
characteristics and features of the carapace provide the basis for an
FIGURE 1. Location map of conchostracan sites in New Mexico discussed here.
exact determination of new species. Important characteristics of the
carapace are: (1) morphology of the dorsal margin (DM) (Figs. 2E-F,
3A, C, D); (2) sculpture elements (S) of the larval shell (LS) (Fig. 3B);
(3) ornament (O) of the larval shell (LS) and of the growth bands (GB)
(Figs. 2D, 4C, 5B, 6C); and (4) size and convexity of the reconstructed
carapace (C) (Figs. 4A, 7A).
Describing types of carapace deformation was the main cause
for the inflation of “new species” in the last 60 years. Furthermore, the
inflation of “new species” suggests that conchostracans are not useful
for biostratigraphic correlations over the Pangea continent. Careful taxo-
nomic analysis is most important before we think about biostratigraphic
correlation. Based on our examination of a recent conchostracan popu-
lation we must know that the shape of the conchostracan shell (cara-
pace) can be very variable in specimens at one locality.
PROVENANCE
The following descriptions are based on conchostracans collected
by one of us (SGL) at two localities in central New Mexico. NMMNH
locality 4667 is in the drainage of the Arroyo Tinajas in the Cerros de
Amado of Socorro County, New Mexico (sec. 25, T2S, R1E) (Fig. 1).
This locality is in a 3.5-m thick interval of thinly laminated black shale
of the Upper Pennsylvanian Atrasado Formation (Lerner et al., 2001;
Lerner & Lucas, 2002; Kues et al., 2002). Bracketing marine beds sug-
gest an early Missourian age. The shale also yields fossil plants,
darwinulid ostracods and fragmentary remains of palaeoniscoid fishes,
and has been interpreted as a coastal lake deposit (Lerner et al., 2001).
NMMNH locality 3922 is north of Placitas in Sandoval County

FIGURE 2. Terminology of conchostracan carapace (after Martens, 1983a, figs. 1-
3, 7): A: DM – dorsal margin, AM – anterior margin, VM – ventral margin, PM –
posterior margin, U – umbo, LS – larval shell (valve), GB – growth band, GL –
growth line, C: Cl – length of the carapace, Dl – length of the DM, ll – length of the
LS, dpl – length of the DMp, dll – length of the DMl, dal – length of the DMa, Ch –
height of the carapace, alfa – angle between the direction of the PM and the direction
of the DM, GBw – width of the growth band B: DMp – posterior part of the dorsal
margin between PM of the LS and PM of the adult carapace, DMl – dorsal margin
of the larval shell, DMa – anterior part of the dorsal margin between the AM of the
LS and AM of the adult carapace D: structure of the shell: GB – growth band, S –
sculpture, O – ornament, GL – growth line, Sh1…Sh4 – shell 1 to shell 4 of the
carapace, Ma – interior shell with blood vessels, Me – membrane E: Lioestheriidae:
structure of the DMp, GB – growth band F: Pseudestheriidae: structure of the DMp,
GB – growth band.
(sec. 14, T3N, R5E) (Fig. 1). This locality is in a thin dark gray shale lens
of the upper part of the Abo Formation (Lucas et al., 1999). It also yields
insects (blattoids) and plant remains and has been interpreted as a flood-
plain pond deposit of Wolfcampian age (Lucas et al., 1999).
TERMINOLOGY
For this paper, we use the terminology of fossil conchostracans
after Martens (1983a) to describe all features and important measure-
ments of the carapace (Fig. 2 and 3). Soft parts of the body and eggs are
not preserved at the two localities in New Mexico.
SYSTEMATIC PALEONTOLOGY
The taxonomy of fossil conchostracans is mainly based on the
characteristics of the thin carapace, especially on the sculpture of the
larval shell and on the ornamentation of the LS and GB. The soft body
209
FIGURE 3. Terminology of conchostracan carapace: A: Pseudestheriidae: dorsal
view of the carapace, left and right shell, LS – larval shell, B: Lioetheriidae: dorsal
view of the carapace, left and right shell, LS – larval shell with different sculpture
elements, C: Pseudestheriidae: DM with structure of the growth bands, D
Lioestheriidae: dorsal view of DM with growth bands, E: growing of the carapace of
recent population of Cyzicus cycladoides (Joly), from Italy (Martens, 1985), GBw
– width of the growth bands, GBn – number of the growth bands F: growing of
carapaces 1 to 7 of fossil conchotraca (Pseudestheria graciliformis Martens 1983a,
fig. 29): GBw – width of the growth bands, GBn – number of the growth bands.
and appendages are seldom preserved. For the family Lioestheriidae
Raymond, 1946, the taxonomy is mostly dependent upon the features
of the larval shell (its size and sculpture elements). The sculpture con-
sist of a small, convex cone, a convolute concave furrow or a convolute
convex radial carina – mostly of different size, connection and relief
intensity.
In most fossil conchostracans, the carapace is more or less pre-
served in lateral apsect. The outline of the carapace varies only from
elliptical over oval, subquadrate to subcircular or semicircular. There-
fore, differences in carapace outline are mostly not regarded as taxo-
nomic characters. More then 90 % of the carapace outline is made after
different influences of carapace deformation in the sediments after di-
agenesis or tectonic events. Like recent conchostracans, after three di-
mensional reconstruction of the fossil carapace, we distinguish cara-
paces from globular to a flat convex disc. To reconstruct original shape
of the carapace is the main result of the taxonomic analysis. It guaran-
tees that the author detailed enough features to create a new species or
to assign a known species name to the discovered conchostracans. Ver-
210
bal descriptions of new conchostracan species without drawings of char-
acteristic features are not enough for exact determination. The number
of growth lines is mostly an indicator for the age of the animal, and
indicates the individual development of a conchostracan specimen. The
development of the width of the growth bands from LS to the outside
(broad near the LS and narrow near the ventral margin) is typical for
the normal ontogenetic development of a conchostracan.
Institutional abbreviations: MNG = Museum of Nature, Gotha,
Germany; NMMNH = New Mexico Museum of Natural History, Albu-
querque, New Mexico, USA.
Class Branchiopoda LATREILLE, 1817
Order Conchostraca SARS, 1867
Family Lioestheriidae RAYMOND, 1946
Type genus: Lioestheria DEPÉRET & MAZERAN, 1912
(emend. KOZUR, MARTENS, PACAUD 1981)
Genus: Lioestheria DEPÉRET & MAZERAN, 1912
(emend. KOZUR, MARTENS, PACAUD 1981)
Type species: Estheria (Lioestheria)lallyensis DEPÉRET &
MAZERAN, 1912
Lioestheria carinacurvata sp. nov.
Figs. 4-6
Derivatio nominis: carina (lat.) = keel, curvatura (lat.) = arc
Holotype: NMMNH P-45704, carapace, left shell (Fig. 5A).
Paratypes: NMMNH P-45701, P-45703, P-45706, P-45708, P-
45711, P-45718, P-45719, P-45730, P-45732 (Fig. 4, 5, 6)
Type locality: NMMNH locality 4667.
Referred specimens: NMMNH P-45701-P-45732, dark bluish
gray shale with intensiveLY deformed carapaces parallel to the layers
(Fig. 4D).
Diagnosis: Carapace with subquadrate outline; large larval shell
with a long curved carina (type A) or a short curved carina (type B).
Maybe type A and type B are characteristic of sexually dimorphism.
Because carapaces with eggs are missing yet, it is not certain, which
type represents the female or male carapace. The ornament is very ar-
ticulate on the larval shell and growth bands. The type of ornament is
net-shaped to honey-comb-pattern-shaped. The GBw are normal de- FIGURE 4. Lioestheria carinacurvata: A: reconstruction of the carapace (type A
veloped, but near the posterior margin and much narrower than in most with long radial carina at the LS), 1 - lateral view, 2 – anterior view, 3 – dorsal view,
species of Lioestheria. scale = 1 mm, B: type B with short radial carina at the LS, scale = 1 mm, C: NMMNH
Measurements: Length: 3-3.5 mm, height: 2.0-2.5 mm P-45719, paratype, normal development of the growth bands from the LS (right) to
Remarks: The genus Lioestheria was first established by Depéret the carapace outline (left), scale = 0,5 mm, NMMNH P-45719, paratype, complete
& Mazeran, 1912 to describe small and good preserved conchostracans carapace of Fig. 3C, scale = 1 mm, D: NMMNH P-45715, carapace of more than
with a large larval shell from the Autun Basin (Grès de Lally, Autunian) 20 individuals with different preservation, scale = 1 mm.
in France. Today we know, that this genus is very common from the
uppermost Carboniferous (Stefanian) to the Lower (? Middle) Permian Formation, Upper Rotliegend, Lower Permian.
worldwide. More than 8 species are know and described together with New Mexico locality: NMMNH locality 3922.
some synonyms. The type species Lioestheria lallyensis Depéret & Description: The reconstruction of the carapace in Figure 7A-B
Mazeran, 1912 (L. paupera after Kozur) represents a species actually shows typical features: Carapace convex with a relatively small LS. LS
known from Germany, France and North-Central-Texas, USA (Mar- with sculpture element as a small cone on the right side near the ante-
tens, in preparation). Lioestheria carinacurvata has a characteristic rior margin; small radial sculpture elements recorded in Lioestheria
feature of the larval shell – a long (type A) or a short (type B) curved monticula from the type locality are not preserved in the specimens
carina – an isolated feature, that is restricted to this species only. from NMMNH locality 3922. Sexual dimorphism is unknown. Remains
of soft parts and eggs are known only from the type locality “Bromacker.”
Lioestheria cf. M. monticula MARTENS, 1983a See measurements in Table 1.
Remarks: The type species L. lallyensis (L. paupera) may be
Figs. 7-8
typical for a stratigraphic level near the Carboniferous/Permian bound-
Derivatio nominis: lat.: monticulus = little mountain, describes ary. The species Lioestheria pseudotenella Martens, 1983a and
the little cone on the LS. Lioestheria monticula Martens, 1983a, both known from the Thuringian
Holotype: Martens, 1983a, pl. XXXV, Fig. 1, 2, 4, 6: MNG- Forest in Germany, are important species for the Lower Permian. The
3613-36-2. characteristic features of Lioestheria cf. M. monticula from New Mexico
Type locality of holotype: Bromacker quarry near Tambach- are the convex sculpture on the growth bands and the small convex
Dietharz, Thuringian Forest, Germany; Tambach Sandstone, Tambach cone near the anterior margin of the larval shell. Radial sculpture ele-

FIGURE 5. Lioestheria carinacurvata, scale = 1 mm: A: NMMNH P-45704
holotype, left shell, sculpture of LS concavo, B: NMMNH P-45708, paratype, C:
NMMNH P-45730, paratype, D: NMMNH P-45731, E: NMMNH P-45705 , F:
NMMNH P-45707
ments and parts of the soft body are still unknown.
BIOSTRATIGRAPHIC CONCLUSIONS
Biostratigraphic projects in the Upper Carboniferous/ Lower Per-
mian non-marine sediments of Pangea still face two main questions (Mar-
tens, 1994): (1) which fossil groups can used for biostratigraphic corre-
lations of terrestrial sediments?; and (2) in which areas around the world
we can find a biostratigraphic “correlation bridge” for the Carboniferous/
Permian boundary between marine and terrestrial facies ? Martens (1984)
created the first conchostracan zone succession with a definition of a
lower and upper boundary. For the next examination we have to find long
profiles without any important stratigraphic hiatus. We give an example
for the first intercontinental correlation with Lower Permian
conchostracans. Lioestheria monticula and Lioestheria cf. M. monticula
TABLE 1. Measurements of conchostracans.
Cl (mm) Ø Cl
Lioestheria carinacurvata sp. nov. (present paper) 3.0 (holotype)
Lioestheria cf. L. Monticule MARTENS 1983° 2.0 – 3.5
Lioestheria monticule MARTENS 1983a 1.86 – 4.08 2.78
Lioestheria pseudotenella MARTENS 1983a loc. L 1.40 – 3.35 2.58
Lioestheria pseudotenella MARTENS 1983a loc. W 1.44 – 2.56 2.12
Lioestheria parvula MARTENS 1983a 1.40 – 2.56 1.77
Lioestheria lallyensis DEPÉRET & MAZERAN 1912 1.9 – 2.8
Lioestheria extuberata JONES & WOODWARD 1899 3.0
211
FIGURE 6. Lioestheria carinacurvata: A: NMMNH P-45701, paratype, scale = 1
mm, B: NMMNH P-45711, paratype, scale = 1mm, C: NMMNH P-45711, paratype,
ornament and sculture element of the LS, scale = 0,25 mm, D: NMMNH P-45718,
paratype, sculture element of LS, scale = 1 mm , E: NMMNH P-45703, paratype,
ornament and sculpture element of LS, GBn 1 to 4, scale = 1 mm, F: NMMNH P-
45732, paratype, sculpture element of LS, scale = 1 mm, G: NMMNH P-45706,
paratype, ornament and sculpture element of LS, scale = 1 mm
are known from the Tambach Formation (Thuringian Forest Basin, Ger-
many), the Wadern Formation (Saar-Nahe-Basin, Germany), from the
Archer City Formation (north-central Texas, USA, Hentz, 1988) and
from the upper Abo Formation (NMMNH locality 3922, New Mexico).
But the problem now is, that we don’t know the exact stratigraphic
extension of L. monticula. We need better profiles and more collecting in
this stratigraphic level to establish its range, but the distribution of this
conchostracan as currently understood suggests a Tambach-Archer City-
upper Abo correlation.
ACKNOWLEDGMENTS
We thank the Deutsche Forschungsgemeinschaft (DFG) for sup-
port of the research about the taxonomy and biostratigraphy of the
Mn GBn
7 (15 – 22)
15 - 18
71 13
118 18
17 11
24 14
>6
212
FIGURE 7. Lioestheria cf. M. monticula Martens, 1983: A: reconstruction of the
carapace of Lioestheria monticula MARTENS 1983: 1 - lateral view, 2 – anterior
view, 3 – dorsal view, scale = 1 mm, B: Lioestheria cf. L. monticula Martens, 1983:
reconstruction of LS with sculpture element, C: NMMNH P-45740, scale = 1 mm,
D: NMMNH P-45746 , N ,scale = 1 mm
Bromacker locality and we thank the Rotary Club Gotha for sponsor-
ship and the Kulturstiftung Gotha for support of the visit of the meet-
ing “The Nonmarine Permian” in October 2005 in Albuquerque, New
Mexico, USA.
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