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Download by: [Hacettepe University] Date: 31 March 2017, At: 13:16
Alcheringa: An Australasian Journal of Palaeontology
ISSN: 0311-5518 (Print) 1752-0754 (Online) Journal homepage: http://www.tandfonline.com/loi/talc20
Carboniferous postglacial faunas in the late
Serpukhovian–Bashkirian interval of central-
western Argentina
G.A. Cisterna, A.F. Sterren, O. López Gamundí & M.M. Vergel
To cite this article: G.A. Cisterna, A.F. Sterren, O. López Gamundí & M.M. Vergel
(2017): Carboniferous postglacial faunas in the late Serpukhovian–Bashkirian interval of
central-western Argentina, Alcheringa: An Australasian Journal of Palaeontology, DOI:
10.1080/03115518.2017.1299795
To link to this article: http://dx.doi.org/10.1080/03115518.2017.1299795
Published online: 30 Mar 2017.
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Carboniferous postglacial faunas in the late Serpukhovian–
Bashkirian interval of central-western Argentina
G.A. CISTERNA, A.F. STERREN, O. LÓPEZ GAMUNDÍ and M.M. VERGEL
CISTERNA, G.A., STERREN, A.F., LÓPEZ GAMUNDÍ,O.&VERGEL, M.M., March 2017. Carboniferous postglacial faunas in the late Serpukhovian–
Bashkirian interval of central-western Argentina. Alcheringa, ISSN 0311-5518.
Typical glacial–postglacial sequences associated with the Late Palaeozoic Ice Age (LPIA) are recognized in the Calingasta-Uspallata Basin, central-
western Argentina, particularly in the Hoyada Verde and El Paso formations (late Serpukhovian–Bashkirian) at Barreal Hill (San Juan province).
Brachiopods and bivalves accompanied by gastropods, conulariids, nautiloids, corals and ostracods constitute the marine assemblages of the El
Paso Formation. They are assigned to the Aseptella–Tuberculatella/Rhipidomella–Micraphelia (AT/RM) fauna, characterized by two fossil assem-
blages: Aseptella–Tuberculatella, identified in the lower fossiliferous interval, and Rhipidomella–Micraphelia in the upper. The development of the
different invertebrate assemblages within the El Paso Formation, and their relationship with coeval suite in the Hoyada Verde Formation, can be
explained by a complex array of abiotic factors (substrate stability, turbidity, nutrient availability, variation in oxygen levels, poor circulation and
salinity variations in the water column) that were directly related to glacial retreat dynamics and coastal configuration. A restricted palaeofjord set-
ting is proposed for the depositional environment of the El Paso Formation in contrast to an exposed open marine coast with a gently sloping shelf
for the Hoyada Verde Formation. The study of the postglacial fauna of the El Paso Formation and its relationship with the Levipustula fauna in the
Calingasta-Uspallata Basin, help determine the main controls on the distribution of the postglacial faunas in other late Palaeozoic South American
basins, such as the Tepuel Genoa Basin in Patagonia and the Tarija Basin in Bolivia.
Gabriela A. Cisterna [gabrielacisterna@conicet.gov.ar],CONICET-UNLAR, Av. Dr. Luis M. de la Fuente s/n, La Rioja, 5300, Argentina; Andrea
F. Sterren [asterren@unc.edu.ar], CICTERRA (CONICET –Universidad Nacional de Córdoba), Av. Vélez Sarsfield 1611, X5016GCA, Córdoba,
Argentina; Oscar López Gamundí [orlg2003@yahoo.com], P1C Consultants 1121 Banks Street, Houston, TX 77006, USA; María del Milagro
Vergel [vergelmar@tucbbs.com.ar], CONICET –INSUGEO –Facultad de Ciencias Naturales e Instituo Miguel Lillo (UNT), Miguel Lillo 205,
4000, San Miguel de Tucumán, Argentina. Received 16.12.2015; revised 20.2.2017; accepted 22.2.2017.
Key words: Carboniferous, postglacial faunas, brachiopods, bivalves, Argentina.
STRATIGRAPHIC records of the Late Palaeozoic Ice
Age (LPIA), can be recognized in several parts of
Gondwana. In South America, the Late Carboniferous
postglacial transgression related to this event is
extended well into western Argentina, from the Tarija
Basin in the north, through the Precordillera (Calin-
gasta-Uspallata, Río Blanco and Paganzo basins), to the
Patagonian Tepuel Genoa Basin in the south (López
Gamundí 1989). In west-central Argentina, glacial
deposits of mid-Carboniferous age [late Serpukhovian–
early Bashkirian, Gulbranson et al.2010; also part of
the glacial episode II of López Gamundí (1997) and the
glacial palaeoclimatic stage (late Visean–early
Bashkirian) of Limarino et al.2014] are abundant in
the Calingasta-Uspallata Basin and the adjacent Paganzo
Basin. In the Calingasta-Uspallata Basin, exposures of
Carboniferous glaciomarine deposits are distributed
irregularly along the western side of Tontal Hill
(Fig. 1A). The stratigraphic succession is characterized
by important lateral facies changes as a consequence of
the progressive sedimentary infill across an irregular
pre-Carboniferous topography (Veevers & Powell
1987). Some sections incorporate a distinctive glacial–
postglacial transition from basal deposits of proximal
glacial–marine environments with sediment gravity
flows to postglacial marine fossiliferous mudstones
(López Gamundí & Martínez 2000). One of these typi-
cal glacial–postglacial sequences is exposed at Barreal
Hill (San Juan province), particularly expressed in the
late Serpukhovian–Bashkirian Hoyada Verde Formation
(Mésigos 1953), in which the postglacial deposits con-
tain the very well known Levipustula fauna (Cisterna &
Sterren 2008,2010), which was originally referred to
the Levipustula levis zone (Amos & Rolleri 1965).
Diverse invertebrate assemblages have been identified
in the glaciomarine deposits of the El Paso Formation
(Mésigos 1953), located south of Barreal Hill (Fig. 1B).
In the context of the sequence stratigraphic framework
proposed by López Gamundí & Martínez (2003), the
fossiliferous beds of the Hoyada Verde and El Paso for-
mations are grouped in a transgressive systems tract and
considered to be part of ‘Sequence I’. However, the fos-
sil assemblages of the El Paso Formation have impor-
tant compositional, taphonomical and palaeoecological
differences from those of the Hoyada Verde Formation,
© 2017 Australasian Palaeontologists
http://dx.doi.org/10.1080/03115518.2017.1299795
and other coeval successions (Leoncito, La Capilla,
Majaditas and Yalguaraz formations), and the age of the
El Paso Formation has been a matter of controversy.
This paper reviews the invertebrate fossil assem-
blages of the El Paso Formation, mainly brachiopods
and bivalves, in order to define a new fauna for the ear-
liest postglacial interval (late Serpukhovian–Bashkirian)
of the Calingasta-Uspallata Basin, and to elucidate the
fauna’s palaeoecological, biostratigraphic and palaeobio-
geographic implications. Age interpretations discussed
herein are in concert with the recent palynological
studies of the El Paso Formation (Vergel et al.2015).
Terminology and methodology
All specimens mentioned and described come from a
stratigraphic section located on the south side of Barreal
Hill (Figs 1, 2) centred on GPS coordinates (31°40′
42.7″S, 69°25′26.9″W). Sampling in the field was
undertaken bed by bed, supported by observations of
the taphonomic/palaeoecological aspects of the assem-
blages (Stanley 1970, Ausich & Waters 1982, Ausich
1983, Fürsich & Heinberg 1983, Bassett 1984, Kidwell
et al. 1986, Kidwell & Bosence 1991).
Brachiopods, bivalves and gastropods are the main
constituents of the two fossil assemblages identified in
the El Paso Formation, but conulariids, nautiloids, ostra-
cods and corals have also been considered to evaluate
the relative generic richness in the section studied. The
fossils are distributed throughout two stratigraphical
intervals: the lower fossiliferous interval, where eight
beds were sampled (i.e., beds 1 to 8), and the upper
interval, in which only one fossil bed was recognized
(bed 9; Fig. 2). Brachiopods and bivalves are the
Fig. 1. A, Geographic location of the Carboniferous postglacial faunas in the Calingasta-Uspallata Basin, central-western of Argentina;
B, Generalized geological map of the Barreal Hill and the El Paso Formation outcrops (modified from Mésigos 1953), showing the studied area.
2 G.A. CISTERNA et al.ALCHERINGA
dominant groups in the fossil assemblages of the El
Paso Formation and are specifically studied because of
their palaeoecological significance. The relative abun-
dance of specimens of these groups has been calculated
only in beds 8 (the greatest diversity has been recorded
in this bed) and 9. The relative abundance data are
based on counts of whole and fragmented specimens.
The total number of articulated specimens and the
greater number of disarticulated valves in each sample
(ventral–dorsal in brachiopods and right–left in
bivalves) were considered in this calculation.
The taxonomic composition and relative generic
abundance were analysed in order to characterize fossil
‘assemblages’sensu Fargrestrom (1964), i.e., ‘any
group of fossils from a suitably restricted stratigraphic
interval and geographic locality’. In our case, we
defined two fossil ‘assemblages’restricted to the lower
and upper stratigraphical intervals of the El Paso
Formation.
The concept of a ‘fauna’, an informal term to refer a
group of marine invertebrates associated with a
palaeoenvironment and/or a particular interval of time,
has been used by several authors in relation to the late
Palaeozoic in Argentina (Césari et al. 2007 and refer-
ences therein) and Australia (Runnegar 1979), i.e., the
Tivertonia–Streptorhynchus fauna, Buxtonia–Heteralosia
fauna, Costatumulus fauna, Levipustula fauna and
Eurydesma fauna. Likewise, in this study, the concept
of ‘fauna’refers to a group of marine invertebrates
(embracing several ‘assemblages’) that lived in a cold
environment associated with the postglacial phase of the
Carboniferous glacial event (i.e., the Levipustula
fauna and the Aseptella–Tuberculatella/Rhipidomella–
Micraphelia fauna). Because the Levipustula fauna
Fig. 2. Stratigraphical section of the El Paso Formation (modified from Mésigos 1953) with details of the fossiliferous interval and the stratigraphic
distribution of the studied fossil taxa.
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 3
characterizes the late Serpukhovian–Bashkirian Levipus-
tula levis Biozone (Amos & Rolleri 1965), the occur-
rence of a chronologically equivalent fauna in the El
Paso Formation also has biostratigraphic implications.
The fossils were prepared mechanically, using
pneumatic vibrators and thin needles. Photographs of
specimens were taken using flash illumination after
whitening with ammonium chloride. Images were cap-
tured using a Canon Power Shot S50 digital camera
mounted on a Leica MZ75 binocular magnifier. The
material illustrated (Figs 3–6) is housed in the palaeon-
tological collection of the Centro de Investigaciones
Paleobiológicas (CIPAL) of the Universidad Nacional
de Córdoba, Córdoba (Argentina), with repository
number prefixed CEGH-UNC.
A systematic revision of the brachiopods from the El
Paso Formation, based on the new material collected, is
provided in the Systematic palaeontology. The classifica-
tion of Brachiopoda adopted herein follows Brunton
et al. (2000) for Productida, Williams & Harper (2000)
for Orthida and Lee et al. (2006) for Terebratulida. Only
the brachiopods are described because, from a biostrati-
graphic viewpoint, they are diagnostic for the postglacial
Aseptella–Tuberculatella/Rhipidomella–Micraphelia fauna
herein informally designated AT/RM fauna.
Previous palaeontological studies of the
El Paso Formation
The first palaeontological data from the El Paso Forma-
tion were reported by Mésigos (1953), who indicated
the presence of gastropods in a mudstone-rich interval
within the upper part of the glaciomarine section. New
palaeontological records from the same stratigraphic
interval were provided by Taboada (1989), who recog-
nized an invertebrate fauna composed of brachiopods,
bivalves, gastropods, bryozoans and corals. Taboada
(1989) carried out the first formal description of some
components of this fauna [the gastropods Glabrocingu-
lum (Glabrocingulum)advena (Reed in Du Toit, 1927),
Glabrocingulum (Stenozone) sp., Yunnania subpygmaea
(D’Orbigny, 1850), Neoplatyteichum barrealensis (Reed
in Du Toit, 1927), Sinuitina gonzalezi (Sabattini, 1977),
Murchisonia (Murchisonia) sp. aff. M. tatei? (Elias,
1957); the bivalves Phestia sp. and the brachiopods
Rugosochonetes gloucesterensis (Cvancara, 1958),
Rugosochonetes sp. A, Rugosochonetes sp. B, and
Bulahdelia sp. cf. B.myallensis (Roberts, 1976)], and
defined the Rugosochonetes-Bulahdelia Biozone (late
Visean–early Namurian), which was geographically
restricted to Barreal Hill. On the basis of new fossil col-
lections, Simanauskas & Cisterna (2001), studied the
brachiopod faunas from the El Paso Formation and
reassigned the species previously identified by Taboada
(1989). These authors described the species Aseptella
sp. aff. A. patriciae Simanauskas, 1996b,Tuberculatella
peregrina (Reed in Du Toit, 1927), Rhipidomella? sp.
and Micraphelia indianae Simanauskas & Cisterna,
2001, and suggested that they could be of Late
Carboniferous–earliest Permian age. However, the
brachiopod faunas of the El Paso Formation lack well-
resolved indices to determine the precise age of this
unit. Fortunately, in recent years, palynological data
from the invertebrate-bearing beds (Vergel et al.2008,
2015) suggest a late Serpukhovian–Bashkirian age
based on assemblages referred to Subzone A of the
Raistrickia densa-Convolutispora muriornata Biozone
(Césari et al.2011).
Geological setting of the El Paso
Formation
Outcrops of the El Paso Formation are located on the
southernmost part of Barreal Hill, a broad north–south
anticline located 3 km east of Barreal village in San
Juan province, Calingasta-Uspallata Basin (Fig. 1A). An
angular unconformity separates the base of the El
Paso Formation from the underlying rocks of the
pre-Carboniferous (Ordovician?) metasedimentary base-
ment of the Hilario Formation (Mésigos 1953; Fig. 1B).
The El Paso Formation is overlain by the basal Tres
Saltos Member of the Pituil Formation (Sequence II of
López Gamundí & Martínez 2003), with an erosional
unconformity (Fig. 2). López Gamundí & Martínez
(2003) distinguished Sequence I in the El Paso Forma-
tion (and the equivalent Hoyada Verde Formation),
characterized by a basal section that consists of glacially
influenced deposits of a lowstand wedge, overlain by
thin-bedded diamictites (early transgressive systems
tract or TST), followed by ice-rafted detritus-free
mudstones with fossils (late TST).
Two diamictite-rich intervals characterize the lower
and upper parts of the El Paso Formation in the type
section (Mésigos 1953), and between these intervals, a
vertical alternation of pebbly sandstones, boulder con-
glomerates, pebbly (dropstone) shales, sandstones, silt-
stones and shales has been described (Martínez et al.
1998). Immediately above and below the diamictite
package in the upper part of the section, two fossilifer-
ous intervals have been identified (Figs 1B, 2). The
lower fossiliferous interval consists of a 30 m-thick
section of alternating mudstones and shales with fine-
grained sandstones corresponding to a transgressive sys-
tems tract (López Gamundí & Martínez 2003); several
beds with invertebrates have been sampled there. The
upper fossiliferous interval, distinguished by 10 m of
mudstone and ripple cross-laminated medium- to very
fine-grained sandstone with dropstones, is interpreted as
a highstand systems tract (HST) (López Gamundí &
Martínez 2003) and contains marine invertebrates and
plant remains. The taxonomic, biostratigraphic and
palaeoecological characteristics of the fossil assem-
blages in the two fossiliferous intervals are discussed
below.
4 G.A. CISTERNA et al.ALCHERINGA
Postglacial fauna of the El Paso
Formation
Composition and stratigraphic distribution. The post-
glacial fauna of the El Paso Formation includes bra-
chiopods, bivalves, gastropods, conulariids, nautiloids,
corals and ostracods, occurring in two intervals: the
lower, corresponding to the sampling beds 1–8, and the
upper equivalent to bed 9 (Fig. 2). In the lower interval,
the fossil assemblage is represented by brachiopods, gas-
tropods, bivalves and disarticulated crinoids that appear
mainly scattered, and locally concentrated as nests in
thick mudstone packages with calcareous concretions
that usually also contain orthoconic nautiloids, conulari-
ids and ostracods. The fossils have a continuous vertical
distribution throughout this interval, but eight sampling
beds have been defined incorporating the most important
faunal changes (Fig. 2). From beds 1 to 4, the assem-
blage is dominated mainly by brachiopods [Rhipido-
mella discreta sp. nov., Tuberculatella peregrina,
Micraphelia indianae,Micraphelia? sp. and ‘inarticu-
lates’(Orbiculoidea sp.)], accompanied by sparse
bivalves and gastropods. A conspicuous level with con-
cretions (20 cm thick) containing orthoconic nautiloids,
conulariids, brachiopods (Tuberculatella peregrina,
Micraphelia? sp. and Orbiculoidea sp.), bivalves (Myo-
fossa calingastensis González, 2002) and gastropods
(Murchisonia? sp. and Gastropoda indet., according to
Sterren et al.2013) has been identified between the beds
2 and 4. From bed 4, an important compositional change
is evident: generic diversity begins to increase gradually
to bed 8, at which it is highest; brachiopods (Rhipido-
mella discreta sp. nov., Micraphelia indianae,
Micraphelia? sp., Tuberculatella peregrina, Overtoniinae
indet., Aseptella sp. aff. A. patriciae, Linoproductoidea
indet., Beecheria patagonica Amos, 1958, athyrids
indet. and ‘inarticulates’(Lingulids indet. and Orbicu-
loidea sp.), bivalves (Nuculanella camachoi González,
1972,Quadratonucula?sp.,Nuculopsis? sp., Phestia
sp.,Myofossa calingastensis, Bivalvia indet.) and gas-
tropods [Ananias sp., Glabrocingulum (Stenozone) sp.
and Murchisonia? sp., listed by Sterren et al. 2013] are
present with the maximum number of species in relation
with the entire fossiliferous interval.
In the upper sandy interval (bed 9 of the section;
Fig. 2), the fossil assemblage is composed of marine
invertebrates and abundant fragmentary plants; the fos-
sils are more dispersed than in the lower interval. Bra-
chiopods are dominant (Micraphelia indianae,
Rhipidomella discreta,Orbiculoidea sp. and Beecheria
patagonica), accompanied by bivalves (Aviculopecten
barrealensis Reed in Du Toit, 1927,Myofossa calingas-
tensis and Schizodus? sp.), gastropods [Glabrocingulum
(Stenozone) sp., Murchisonia? sp., Gastropoda indet.]
and solitary rugose corals.
Brachiopod occurrences. The names given to the fos-
sil assemblages identified in the El Paso Formation
(Aseptella–Tuberculatella and Rhipidomella–Micraphelia
assemblages) are based on the occurrence of diagnostic
brachiopod genera. The first assemblage, in the lower fos-
siliferous interval, is characterized mainly by Aseptella
sp. aff. A. patriciae and Tuberculatella peregrina.In
terms of biostratigraphy and palaeobiogeography, Asep-
tella can be considered a key genus for this assemblage.
It is a small Plicatiferinae, characterized mostly by two
radiating rows of stout ventral spines on each flank. The
genus was originally based on material from the Meré
Beds (early Bashkirian–Namurian B–C), in the
Cantabrian Mountains, Spain, with the type species
Aseptella asturica (Martínez Chacón & Winkler Prins,
1977). From the same region, the oldest related species,
Aseptella beetsi, has been described from the uppermost
Visean–lower Serpukhovian beds of the Alba Formation
(Winkler Prins & Martínez Chacón 1998). Aseptella
patriciae Simanauskas, 1996b, from the Tepuel Genoa
Basin, Patagonia, represents the first record of Aseptella
in Argentina. Specimens from the El Paso Formation con-
stitute a markedly more robust form compared with the
Patagonian species, but this feature has been explained as
an adaptation to a high-energy environment (Cisterna &
Simanauskas 1999). Argentinean species can be clearly
distinguished from those of Spain based on their larger
size, particularly if adult specimens are compared. Tuber-
culatella peregrina, the other diagnostic species of the
Aseptella–Tuberculatella assemblage, is characterized by
elongated spine bases, a broad shallow sinus and a single
row of spines developed between the auricles and the
disk. Tuberculatella peregrina has been interpreted as a
form morphologically intermediate between the type spe-
cies, Tuberculatella tubertella Waterhouse, 1982,
described from the Late Carboniferous of Thailand, and
Tuberculatella laevicaudata (Amos, 1961) from the
Tepuel Genoa Basin (Simanauskas & Cisterna 2001).
The second fossil assemblage, identified in the upper
fossiliferous interval, is characterized by the presence
of Rhipidomella discreta sp. nov. and Micraphelia
indianae.Rhipidomella discreta is a particularly small
representative of Rhipidomellinae with a subtriangular
outline and conspicuous growth lines. The small size
appears to be characteristic of only a few species, such
as Rhipidomella plana Yang from the Carboniferous of
China (Yang et al.1977) and the Permian Rhipidomella
cordialis Grant from Thailand (Grant 1976). This new
species has also been recognized immediately above the
El Paso Formation, in the lowermost layers of the Pituil
Formation (Tres Saltos Member). Rhipidomella discreta
occurs also in the Carboniferous postglacial succession
of the Agua del Jagüel Formation (Mendoza province).
Micraphelia Cooper & Grant has been identified mainly
in the Permian successions of Texas (Cooper & Grant
1969,1975). Micraphelia indianae, described from the
El Paso Formation, is a small to medium-sized member
of Rugosochonetidae (see Systematic palaeontology),
moderately to strongly concavo-convex, semicircular to
subquadrate, with the hinge line slightly shorter than
the maximum width, and cyrtomorph intraversed
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 5
cardinal spines (Simanauskas & Cisterna 2001).
Micraphelia indianae is similar to the type species,
Micraphelia scitula Cooper & Grant, 1969, particularly
in its external features and also shares some characteris-
tics with Micraphelia pumilis Cooper & Grant, 1975
and Micraphelia subalata Cooper & Grant, 1975, from
west Texas (Simanauskas & Cisterna 2001).
A systematic revision of the representative
brachiopods of the fossil assemblages identified in the
El Paso Formation, based on new materials, is provided
in the Systematic palaeontology.
Biostratigraphic implications. The Aseptella–
Tuberculatella and Rhipidomella–Micraphelia assem-
blages constitute the postglacial fauna associated with the
late Serpukhovian–Bashkirian transgression and are
chronologically equivalent to the Levipustula fauna. This
new postglacial fauna is distributed regionally, being doc-
umented in several glacial–postglacial successions of the
Calingasta-Uspallata Basin in the Precordillera (Ciénaga
Larga del Tontal Formation to the north and Agua del
Jagüel Formation to the south, Fig. 1A), and the Tepuel
Genoa Basin in Patagonia (Pampa de Tepuel Formation;
Cisterna & Sterren 2015,2016). In the lower part of the
Ciénaga Larga del Tontal Formation (Casa de Piedra
Formation, Lech et al.1998), fossiliferous layers located
in a glaciomarine interval [‘diamictitic member’of Lech
& Milana (2006)] contain brachiopods and molluscs of
the Aseptella–Tuberculatella assemblage. Aseptella? sp.
and Productella sp. described by Lech & Milana (2006)
from this interval can be reassigned to Aseptella sp. aff.
A. patriciae and Tuberculatella peregrina, which occur
in the El Paso Formation (Simanauskas & Cisterna
2001). Further, the bivalve Nuculopis? sp. described from
the Ciénaga Larga del Tontal Formation is similar to
the species of this genus recognized in the El Paso
Formation.
Typical elements of the AT/RM fauna of the El Paso
Formation have also been identified in the glaciogenic
succession (Sequence I of Henry et al.2010) of the
Agua del Jagüel Formation (Martínez et al.2001,
Simanauskas & Cisterna 2001, Cisterna et al.2013). A
very-low-diversity fossil assemblage composed of
Rhipidomella discreta sp. nov. and Micraphelia indi-
anae, accompanied by the bivalve Nuculanidae indet.,
gastropods indet. and corals characterizes the fossilifer-
ous beds related to the glacial succession of the Agua
del Jagüel Formation (Stage 2 according Henry et al.
2010). It is important to note that some components that
characterize the fossil assemblages of the El Paso For-
mation have also been identified in Patagonia. Aseptella
patriciae,Tuberculatella?laevicaudata and Beecheria
patagonica have been described from the glaciomarine
succession of the Pampa de Tepuel Formation (Amos
1958,1961, Simanauskas 1996a,1996b). However,
Pagani & Taboada (2010) suggested that the provenance
of this material is not certain; hence the biostratigraphi-
cal value of this Patagonian fauna can not be applied to
its occurrence in the Precordillera.
Palaeoecological considerations. There are impor-
tant compositional differences within the El Paso
Formation postglacial faunal succession particularly
between the lower and upper fossiliferous intervals.
Nevertheless, brachiopods and bivalves are the most
characteristic components of both intervals, and the dis-
tribution of each group is constrained by different phys-
ical factors (Ausich & Waters 1982). Analysis of the
functional morphology of these groups clarifies the
palaeoecological controls operating in the aftermath of
the Carboniferous glacial event.
Brachiopods are abundant and diverse in the El Paso
Formation (Fig. 7A). The conditions at the sediment–
water interface influenced by sediment mobility and
grain size were some of the most important factors
controlling the distribution of the rhynchonelliformean
brachiopods (Ausich 1983). Organism–substrate rela-
tionships have been ascribed to several categories that
reflect varied adaptations to life habits, interpreted on
the basis of shell morphology (Rudwick 1970, Fürsich
& Hurst 1974, Bassett 1984, Sánchez & Tóffolo 1996).
The energy of the environment also influences the gen-
eral shape and thickness of brachiopod shells: thick
shells appear to be an adaptation to turbulent water, and
the presence of a large median sulcus, typical of some
spiriferids, would produce a maximum separation for
the inhalant and exhalant currents produced by the
lophophore avoiding re-circulation in extremely quiet
water (Fürsich & Hurst 1974). However, although the
substrate type has usually been considered one of the
primary palaeoecological controls on the distribution of
brachiopods, some authors have suggested that nutrient
availability may also have controlled their distribution
(Fürsich & Hurst 1974, Pérez Huerta & Sheldon 2006).
Moreover, hypoxic to anoxic conditions have been pro-
posed by Shi et al.(2015), to restrict severely the diver-
sification of benthos and to favour the proliferation of
small brachiopod taxa. These authors have also sug-
gested that bathymetrically more tolerant species are
less sensitive to depth control with respect to their
body-size change dynamics, in contrast to stenobathic
species, which tend to grow larger in shallower water
depths.
In the lower fossiliferous interval (Fig. 2), the bra-
chiopods are dominated by the productids Aseptella sp.
aff. A. patriciae,Tuberculatella peregrina, Overtoniinae
indet. and indeterminate Linoproductoidea, accompanied
by chonetids (Micraphelia indianae, Micraphelia? sp.),
orthids (Rhipidomella discreta sp. nov.), and the less
abundant ‘inarticulates’(Lingulids indet. and Orbicu-
loidea sp.), terebratulids (Beecheria patagonica) and
indeterminate athyrids. The uppermost part of this inter-
val (bed 8; Fig. 2) has the highest overall diversity
(Fig. 7B). The dominant group of productids is charac-
terized by species with markedly convex ventral valves
and numerous robust spines to stabilize the shell within
the sediments; a quasi-infaunal life strategy is suggested
for these organisms [i.e., living partially buried in soft
6 G.A. CISTERNA et al.ALCHERINGA
substrates after a short initial hard-bottom attachment
during their early ontogenetic stages (Rudwick 1970,
Bassett 1984)]. The influence of substrate type and
water depth on brachiopods seems to explain some mor-
phological adaptations in specific taxa, and the structure
and dominance in some associations, but the availability
of nourishment might play a more important role in the
palaeoecology of brachiopods according to Pérez Huerta
& Sheldon (2006). These authors suggested that pro-
ductid brachiopods dominated and were larger in deeper
water with less food supply because of the efficiency of
their lophophore. Reconstruction of inhalant and exha-
lant currents in productid brachiopods indicates that
they could create multi-directional inhalant currents per-
miting them to obtain food from a more extensive area
around the shell (Brunton et al.2000); this would be an
advantage under conditions of low productivity, when
nourishment is depleted (Pérez Huerta & Sheldon
2006).
In the upper part of the section (bed 9, Fig. 2), cho-
netids (Micraphelia indianae) and orthids (Rhipidomella
discreta) dominate the brachiopod assemblage, and
there are notably fewer ‘inarticulates’and terebratulids
(Fig. 7C). The morphology of the small orthid Rhipido-
mella indicates the presence of a functional pedicle dur-
ing life. Although the pedicle has generally been
considered to be an organ for attachment, its function is
more closely comparable with a motile appendage than
with a stalk: the pedicle adjusts the position of the
organism relative to its external environment (Richard-
son 2000). Sánchez & Tóffolo (1996) have also sug-
gested that the evidence of a functional pedicle in adult
stages can not be considered indicative of a particular
type of substrate. Micraphelia can be characterized as
an epifaunal liberosessile brachiopod with a concavo-
convex shell adapted to the ‘ski or snow-shoe effect’
(sensu Bassett 1984), by developing posteriorly directed
spines along the ventral cardinal margin. Chonetids usu-
ally close their pedicle opening early in ontogeny to
become recumbent in a convex-downward dish posture
with the spines spread across the soft sediments as a
‘snow-shoe’device (Bassett 1984). The spines have
also been interpreted as attachment structures, fixed to
hard objects distally by mucal adhesion from the mantle
tips (Bassett 1984, Racheboeuf 1990). In relatively
high-energy environments, this life strategy would have
prevented chonetids being flipped over into the convex-
up position (Bassett 1984).
Levinton (1970)defined palaeo-opportunistic species
by a several features, such as: a random distribution
pattern in individual beds with poor size sorting, occur-
rence in thin but widespread isochronous layers (indi-
cating brief invasion events), great abundance in facies
with which they are not usually associated and numeri-
cal dominance in a fossil assemblage (85–100%). In
this sense, Rhipidomella discreta and Micraphelia
indianae seem to have developed opportunistic strate-
gies in the upper fossiliferous interval of the El Paso
Formation. Moreover, a small biovolume and conserva-
tive calcium carbonate secretion, reflected in the reduc-
tion in ornamentation in these species, have also been
suggested to characterize particular palaeo-opportunistic
brachiopod species (Alexander 1977).
Bivalves, the other important component of the El
Paso Formation fossil assemblages (Fig. 7A), have dif-
ferent life strategies according to their relationships with
the substrate and feeding style (Stanley 1970). In the
lower fossiliferous interval, they are dominated by
infaunal bivalves, among which 57% are detritus feed-
ers (palaeotaxodontids), which normally dwell on soft
substrates under low-oxygen and turbidity conditions
(Stanley 1970). Epifaunal, byssate suspension-feeding
types represent 29% of the bivalves and belong to
Pectinida. Finally, only one genus of Anomalodesmata
was recognized: Myofossa, with the species M.calin-
gastensis (14%), characterized as an infaunal suspension
feeder. All palaeotaxodontids found in this interval are
small, providing an advantage with regard to burrowing
efficiency, but the presence of concentric ridges in the
shell would hinder penetration of the substrate. On the
other hand, the prevalence of infaunal detritus-feeding
bivalves is consistent with a scarcity of food particles in
suspension.
The three bivalve species recognized in the upper
part of the fossiliferous interval are suspension feeders:
the epifaunal byssate Aviculopecten barrealensis and the
infaunal Myofossa calingastensis and Schizodus? sp.
Myofossa calingastensis is interpreted as a slightly
mobile burrower with a very thin lateral sulcus or
groove that interrupts ribs on both sides (Mángano
et al.1998, Morris et al.1991, González 2002). The
presence of a gape at the back of the valves would be
advantageous because it facilitates inhalent and exhalent
currents with the valves closed (Liljedahl 1984). Schizo-
dus De Verneuil & Murchison, 1844, is interpreted as a
genus of relatively rapid, shallow burrowers in soft sub-
strates. It is noteworthy that a significant proportion of
infaunal bivalves throughout the unit retain the valves
partially or entirely articulated. This feature could be
related to the infaunal habit of these organisms, together
with little physical reworking and minimal transporta-
tion of the shells (Kidwell & Bosence 1991).
Palaeoenvironments. The development of the
different fossil assemblages within of the El Paso
Formation would have been influenced by the nature
of the transgressive episode related to the glacial retreat
in a particular sector of the coast. The Aseptella–
Tuberculatella assemblage occurs in the initial trans-
gressive interval characterized by sediment starvation
and relatively weak current circulation and is followed
by a maximum flooding interval representing the maxi-
mum landward expansion of marine conditions (López
Gamundí & Martínez 2003). The calcareous concretions
containing diverse molluscs (gastropods, nautiloids and
bivalves) and less abundant brachiopods occur in these
offshore to offshore-transition facies (turnaround point
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 7
from TST to HST) during maximum flooding. Intervals
with fossil-rich calcareous concretions can reveal epi-
sodes of deposition followed by periods of sediment
starvation (Loutit et al.1988, Brett 2003, Dattilo et al.
2008, Bondioli et al.2015). Moreover, the abundance
of quasi-infaunal productid brachiopods and infaunal
detritus-feeding bivalves in this lower fossiliferous inter-
val suggests an environment characterized by little sus-
pended material. The prevalence of deposit feeders is
consistent with the accumulation of organic detritus in
the sediment under low-energy conditions. Organic-rich
bottom conditions, common near or below the storm
wave base (Bondioli et al.2015), could have offset the
scarcity of particles in suspension, and they might have
played a key role in supporting diverse populations of
brachiopods and taxodontid bivalves. On the other
hand, the significant proportion of bivalves that remain
partially or entirely articulated in the lower interval can
be explained by the delayed decomposition of the liga-
ment that joins the valves in environments with low
oxygen or low temperatures (according to Kidwell &
Baumiller 1990).
The presence of the brachiopod Orbiculoidea,a
conspicuous component of the lower interval, has been
considered to typify dark basinal shale and mudstone,
and might have tolerated poor nutrient supplies or low-
oxygen conditions that were uninhabitable for other
taxa (Butts 2005). These conditions of low oxygen
levels and low sea-floor temperatures are consistent
with proximity to a glacier.
The Rhipidomella–Micraphelia assemblage of the
upper fossiliferous interval of the El Paso Formation
appears to be associated with shallower depositional
conditions, at the onset of the HST, characterized by
very fine-grained sandstones interbedded with parallel-
laminated and ripple cross-laminated medium- to very
fine-grained sandstones (López Gamundí & Martínez
2003) and mudstones. This interval represents inner
shelf to nearshore sedimentation with tractional (oscilla-
tory) currents generated by wave action and deposition
near fair-weather wave base. The benthic fauna is less
abundant and is dominated by brachiopods and bivalves
that indicate more stressful energy conditions compared
with the lower interval. In particular, Rhipidomella
discreta and Micraphelia indianae would have
developed opportunistic life strategies in this setting
linked to higher water energy and turbidity. These
environmental parameters would have facilitated the
development of suspension-feeding trophic types.
Brachiopods and bivalves would have been the domi-
nant low-level suspension feeders, and the solitary cor-
als, also recognized in this fauna, represent the dominant
high-level suspension feeders and would have developed
within the photic zone near the fair-weather wave base
(Butts 2005). The high percentage of articulated infaunal
bivalves (Myofossa calingastensis and Schizodus? sp.)
may be related to rapid burial of individuals (Aigner
1985, Peterson 1985), which would have used the
ligament during the reworking and burial (Fürsich &
Heinberg 1983). Sudden burial is supported by the
increase in clastic sediment input (higher sedimentation
rate) that commonly occurs in this type of environment.
The differences observed between the two fossil
assemblages of the El Paso Formation include a
decrease in taxonomic abundance (decline in bra-
chiopod and bivalve generic richness) and changes in
the ecological guilds: disappearance of the dominant
guild (mainly in the bivalves), and replacement of
infaunal deposit feeders by epifaunal suspension feed-
ers. This faunal turnover is related to fluctuations in the
depositional environment that ranged from sediment
starvation in the lower fossiliferous interval to higher
hydrodynamic energies as a response to shallowing
depositional conditions and more active circulation of
nutrients in the upper interval.
Discussion
The deglaciation process that took place in an inland
sea in this basin produced sedimentary successions
hosting diverse marine invertebrates. In the Calingasta-
Uspallata Basin (Fig. 1), as previously mentioned,
two coeval invertebrate marine faunas appear associ-
ated with the late Serpukhovian–Bashkirian glacial–
postglacial succession. The significant differences
between these faunas must be due to a complex array
of abiotic factors directly related to glacial retreat
dynamics (substrate stability, turbidity, nutrient avail-
ability, oxygen levels and current circulation in the
water column) and the coastal configuration.
The El Paso Formation was probably deposited in a
rather confined palaeogeographic setting, such as a
palaeofjord, as interpreted by several authors for other
areas of the Protoprecordillera (Agua del Jagüel and
Quebrada Grande; Henry et al.2010, Kneller et al.
2004). Ancient fjords carved in the southwestern
Gondwanan margin (Kneller et al. 2004, Dykstra et al.
2006, Desjardins et al.2010, López Gamundí 2010)
promoted the development of sub-environments stress-
ful for benthic colonization, including low salinity, high
sedimentation rates, variable degrees of substrate con-
solidation, oxygen depletion and high turbidity (Syvitski
et al.1987, Buatois & Mángano 2011). Low salinity is
to be expected along a glacially influenced coast via
freshwater discharge from glaciated areas (Pazos 2002,
Buatois et al.2006). However, the marine fauna of the
El Paso Formation indicates that salinity conditions
were normal, at least during certain deglaciation phases.
The occurrences of marl and calcareous concretions,
such as those identified in the lower fossiliferous inter-
val of the El Paso Formation, have been linked to con-
ditions of clastic starvation in the postglacial sequences
of the Agua del Jagüel Formation (López Gamundí
1997, Pazos 2002). Henry et al.(2010) indicated that
clastic sedimentation persisted during deposition of the
lower part of the Agua del Jagüel Formation but at
8 G.A. CISTERNA et al.ALCHERINGA
reduced rates, and they identified anoxic conditions
throughout the sequence. A similar scenario might have
occurred during accumulation of the lower interval of
the El Paso Formation where the components of the
Aseptella–Tuberculatella assemblage indicate stressed
conditions related to low oxygen and nutrients suggest-
ing relatively stagnant conditions in the palaeofjord.
The amount of dissolved oxygen may also depend on
other factors, such as the configuration of the basin and
the water-depth profile (Fonseca 2000, Shi et al.2015).
López Gamundí (2010) suggested that the uneven
lateral extent of the deglaciation-phase transgressive
shales might be linked to their deposition within a gla-
cially sculpted palaeotopography. This author also
indicated that in the early stages of this transgression
triggered by a rapid ice sheet recession, the most deeply
scoured areas of the postglacial shelf were the first to
be flooded. Colonization by benthic organisms would
have been infuenced by very different gradients of the
seafloor in a palaeofjord or an open-shelf environment.
In this manner, the mudstones of the El Paso Formation
hosting the AT/RM fauna might represent deposition in
a relatively isolated or restricted (palaeofjord) part of
the basin, where some palaeoecological conditions were
less favourable; and the Levipustula fauna might have
developed in other areas (particularly Hoyada Verde,
Leoncito, La Capilla, Majaditas, Yalguaraz) having an
open connection to the ocean and more favourable habi-
tats (Fig. 8). Although some authors have proposed an
open-sea setting adjacent to a palaeotopographic high as
a possible depositional environment for the Hoyada
Verde Formation (Alonso Muruaga et al.2013), the
palaeoecological features of its fauna and the relation-
ship with those documented from the El Paso Forma-
tion suggest that the relief of the coast adjacent to areas
accumulating the Hoyada Verde Formation would have
been less steep than in the El Paso area.
The postglacial Levipustula fauna of the Hoyada
Verde Formation occurs in a stratigraphic interval repre-
senting the transition from a maximum flooding interval
at the top of a transgressive systems tract to an HST
(López Gamundí & Martínez 2000). In contrast to the
fossil assemblages of the El Paso Formation, the Levi-
pustula fauna is dominated by bryozoans, spiriferid bra-
chiopods and epifaunal bivalves, all suspension feeders
that suggest different palaeoenvironmental conditions.
Palaeoecolgical studies carried out by Cisterna (1999)
and Cisterna & Sterren (2010) in this fauna indicate a
stable marine environment, such as an open shelf with
moderate bottom currents and nutrient availability
throughout the water column. The variations recognized
in these communities through the fossiliferous interval
would have been controlled by substrate types and
food-supply variations during the postglacial transgres-
sion (Cisterna 1999, Cisterna & Sterren 2010).
Identification of the main controls on the distribution
of these postglacial faunas could also be extended to
other late Palaeozoic South American basins affected by
the Carboniferous glacial event, such as the Tepuel
Genoa Basin in Patagonia and the Tarija Basin in
Bolivia.
Systematic palaeontology
(by Gabriela A. Cisterna)
Order PRODUCTIDA Sarytcheva and Sokolskaya,
1959
Suborder CHONETIDINA Muir-Wood, 1955
Superfamily CHONETOIDEA Bronn, 1862
Family RUGOSOCHONETIDAE Muir-Wood, 1962
Subfamily UNDULELLINAE Cooper & Grant, 1975
Micraphelia Cooper & Grant, 1969
Type species.Micraphelia scitula Cooper & Grant,
1969; Lower Permian of Texas, USA.
Micraphelia indianae Simanauskas & Cisterna, 2001
(Fig. 3K–O)
Remarks. This species was described by Simanauskas &
Cisterna (2001, p. 216, Figs 3i–s; 4). The diagnostic
features include a shell that is medium to large for
the genus, moderately to strongly concave–convex,
semicircular to subquadrate; hinge line slightly shorter
than the maximum width, which is measured in the
posterior half of the shell; cardinal spines cyrtomorph
intraversed, forming an angle of about 50–80° with the
hinge line.
The new specimens from the lower and upper
fossiliferous intervals of the El Paso Formation herein
referred to Micraphelia indianae are predominantly small
(2–10 mm long, 3–13 mm wide), or in a few cases of
medium (10.5–11.5 mm long, 14–15 mm wide) size.
Micraphelia? sp. (Fig. 4A–D)
Remarks. A group of specimens (Fig. 4A–D), consisting
mainly of internal moulds, from the lower fossiliferous
interval of the El Paso Formation, is assigned to
Micraphelia? sp. This material has many of the features
that characterize adult specimens of Micraphelia indi-
anae documented by Simanaukas & Cisterna (2001).
However, the specimens of Micraphelia? sp. are larger
(12–16 mm long, 17–23 mm wide), with a shell that is
markedly more transverse, especially in comparison
with specimens of Micraphelia indianae from the upper
fossiliferous interval (bed 9). The cyrtomorph intraverse
spines that characterize Micraphelia, have not been
observed in this material. Vascular trunks that are paral-
lel, deeply impressed and extended anteriorly to a short
and high median septum in the ventral interior
(Fig. 4C) are also characteristic of Micraphelia? sp.
Suborder PRODUCTIDINA Waagen, 1883
Superfamily PRODUCTOIDEA Gray, 1840
Family PRODUCTELLIDAE Schuchert, 1929
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 9
Fig. 3. Brachiopods. Rhipidomella discreta sp. nov.A,B, Holotype, internal and external mould of ventral valve, CEGH-UNC 26706a-b; C, Exter-
nal mould of ventral valve, CEGH-UNC 26707b; D, Internal mould of dorsal valve, CEGH-UNC 2610; E,F, Paratype, internal and external mould
of dorsal valve, CEGH-UNC 26712a-b; G, Internal mould of ventral valve, CEGH-UNC 26711; H, Dorsal valve, CEGH-UNC 26708; I, Internal
mould of dorsal valve, CEGH-UNC 26709; J, Incomplete internal mould of articulate specimen, CEGH-UNC 26713. Micraphelia indianae Sima-
nauskas & Cisterna, 2001.K,L, External mould of ventral and dorsal valve, CEGH-UNC 26714a-b; M, Ventral valve partially decorticated,
CEGH-UNC 26715; N, Internal mould of ventral valve, CEGH-UNC 26716; O, Internal mould of articulated specimen, CEGH-UNC 26717. Scale
bars = 1.5 mm for A–D, G, I, J; 1.25 mm for E, F, H; 1.75 mm for K–M, O; 3.25 mm for N. A–D; G–I, K–L, O, Lower fosiliferous interval, bed
8; E, F, J, M, N, upper fossiliferous interval, bed 9.
10 G.A. CISTERNA et al.ALCHERINGA
Subfamily OVERTONIINAE Muir-Wood & Cooper,
1960
Tribe LETHAMIINI Waterhouse, 2001
Tuberculatella Waterhouse, 1982
Type species.Tuberculatella tubertella Waterhouse,
1982; Late Carboniferous of Thailand.
Tuberculatella peregrina (Reed in Du Toit, 1927)
(Fig. 4E–L)
Remarks. Tuberculatella peregrina (Reed in Du Toit,
1927) was described from the lower fossiliferous interval
of the El Paso Formation by Simanauskas & Cisterna
(2001), who also discussed the generic diagnosis of
Tuberculatella and compared this taxon with related
Fig. 4. Brachiopods. Micraphelia? sp. A, Internal mould of ventral valve, CEGH-UNC 26721a; B, External mould of dorsal valve, CEGH-UNC
26720a; C, Internal mould of ventral valve, CEGH-UNC 26718; D, Internal mould of dorsal valve, CEGH-UNC 26719. Tuberculatella peregrina
(Reed in Du Toit, 1927). E,F, Internal and external mould of ventral valve, CEGH-UNC 26723a–b; G, Internal mould of ventral valve, posterior
view, CEGH-UNC 26725; H, Internal mould dorsal valve, CEGH-UNC 26726; I,J, Dorsal external mould, frontal and posterior views, CEGH-
UNC 26727; K, Incomplete ventral valve partially decorticated, CEGH-UNC 26722; L, Internal mould of ventral valve, CEGH-UNC 26724. Over-
toniinae indet. M,N, External and internal mould of ventral valve, CEGH-UNC 26728; O, External mould of ventral valve, CEGH-UNC 26729;
P, External mould of ventral valve, CEGH-UNC 26730. Aseptella sp. aff. A.patriciae Simanauskas, 1996. Q,R, External and internal mould of
dorsal valve, CEGH-UNC 26731a–b; S, External mould of ventral valve, CEGH-UNC 26732b; T, External mould of ventral valve, CEGH-UNC
26733. Scale bars = 4.5 mm for A–G, I, J, L, M–P; 3.5 mm for H; 6.5 mm for K; 2.75 mm for Q, R, T; 1.25 mm for S. All material comes from
the lower fossiliferous interval, bed 8.
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 11
genera. Additional material figured herein (Fig. 4E–L)
confirms the presence of Tuberculatella peregrina in the
El Paso Formation. This species has been characterized
mainly by the elongate spine bases, wide and shallow
ventral median sinus and a row of spines over the inner
anterior ears, i.e., between the ears and the disc (Reed in
Du Toit, 1927, pl. 13, fig. 2; Simanauskas & Cisterna
2001,p.217,fig. 5a). The specimen assigned to
Bulahdelia cf. myallensis Roberts, 1976 (Roberts et al.
1976), also described from the lower fossiliferous
interval of the El Paso Formation (Taboada 1989), has
been synonymized with Tuberculatella peregrina
(Simanauskas & Cisterna, 2001). The general morphol-
ogy of the shell and the density of the spines on the
ventral disc and ears, described by Taboada (1989,
p. 127, pl. 2, fig. 9), are similar to those of Tuberculatella
peregrina. Material described as Productella sp. by Lech
et al.(1998, p. 406, figs 3c–e), from the lower part of the
Ciénaga Larga del Tontal Formation, also appears to
belong to Tuberculatella peregrina. Ornament of the ven-
tral valve, general morphology and size of the ears, and
the spines on the ears are similar to those recognized in
Tuberculatella peregrina from the El Paso Formation
(Simanauskas & Cisterna 2001).
Although Tuberculatella laevicaudata (Amos, 1961)
described by Simanauskas (1996a, p. 381, fig. 3) from
the Tuberculatella Biozone of Patagonia has been com-
pared with Tuberculatella peregrina from the Precordil-
lera (Simanauskas & Cisterna 2001), a review of
additional material from the El Paso Formation suggests
that these species are quite different. Tuberculatella lae-
vicaudata is a small delicate species with an inconspic-
uous ventral sinus; the ornament and interior features
are less well defined than the more robust forms that
characterize Tuberculatella peregrina from the
Precordillera.
In relation to the type species, Tuberculatella tuber-
tella Waterhouse (1982, p. 44, fig. 2), Tuberculatella
pregrina has a similar general shell morphology with a
distinctive ventral median sinus; the internal features are
also similar, particularly the well-developed adductor
scars. Simanauskas & Cisterna (2001) have suggested
that Tuberculatella peregrina could be a form
morphologically intermediate between the type species,
Tuberculatella tubertella, and the delicate Tuberculatella
laevicaudata from Patagonia.
Taboada & Shi (2011) reservedly synonymized
Tuberculatella peregrina with Absenticosta brun-
toneileenae Taboada & Shi (2011, p. 105, fig. 12), also
described from the lower fossiliferous interval of the El
Paso Formation. However, Tuberculatella peregrina is
larger, with a more convex ventral valve, having a profile
clearly geniculated, and bearing conspicuous growth
lines. The presence of at least three spines on the ears that
are markedly coarse compared with those on the corpus
separates this species from Absenticosta bruntoneileenae.
Moreover, the conspicuous dorsal median septum in
Tuberculatella peregrina, one of the diagnostic features
of Tuberculatella (Brunton, 2007), was not described in
the material studied by Taboada & Shi (2011).
Overtoniinae indet. (Fig. 4M–P)
Remarks. Incomplete internal and external moulds of
ventral valves are assigned to Overtoniinae indet. The
valves are small to medium sized (about 11 mm long
and 16 mm wide) and gently convex with large and
well-defined ears. At least three coarse spines occur on
the ears. Rounded spine bases are well spaced on the
Fig. 5. Brachiopods. Linoproductoidea indet. A,B, Internal and external mould of ventral valve, CEGH-UNC 26734a-b; C, Internal mould of
ventral valve, CEGH-UNC 26735. Beecheria patagonica Amos, 1958.D, Internal mould of articulated specimen, dorsal view, CEGH-UNC 26736;
E,F, Internal mould of articulated specimen, dorsal and ventral views, CEGH-UNC 26737. Scale bars = 3.5 mm for A–D; 4.5 mm for E, F. A–D,
lower fossiliferous interval, bed 8; E, F, upper fossiliferous interval, bed 9.
12 G.A. CISTERNA et al.ALCHERINGA
disk and smaller than the spines on the ears. The shell
bears rugae and conspicuous growth lines. In compar-
ison with specimens from the same bed (bed 8 of the
lower fossiliferous interval) assigned to Tuberculatella
peregrina, they are smaller and more transversely
expanded with rounded spine bases, particularly on the
ears. The fine, distinctive pustules over the floor of the
valve that characterize the interior of Tuberculatella
have not been observed in the material assigned to
Overtoniinae indet.
Subfamily PLICATIFERINAE Muir-Wood & Cooper,
1960
Tribe PLICATIFERINI Muir-Wood & Cooper, 1960
Aseptella Martínez Chacón & Winkler Prins, 1977
Type species. Aseptella asturica Martínez Chacón &
Winkler Prins, 1977; Carboniferous (Bashkirian) of
Spain.
Aseptella sp. aff. A. patriciae Simanauskas, 1996b
(Fig. 4Q–T)
Remarks. Aseptella sp. aff. A.patriciae from the El Paso
Formation was described and figured by Cisterna &
Fig. 6. Bivalves. A,Nuculanella camachoi González, 1972. Internal mould of left valve, CEGH-UNC 26797. B,C,Quadratonucula? sp. Lateral
view of internal mould of left valves; B, CEGH-UNC 26800; C, CEGH-UNC 26801. D,ENuculopsis? sp. D, External mould of left valve,
CEGH-UNC 26804; E, Composite internal mould of right valve and external mould, CEGH-UNC 26805. F,G,Phestia sp., F, Composite internal
mould of left valve, CEGH-UNC 26807; G, Internal mould of left valve, CEGH-UNC 26808. H,Schizodus sp., composite internal mould of left
valve, CEGH-UNC 26816. I–J,L,Myofossa calingastensis González, 2002.I, Composite mould of left valve, CEGH-UNC 26814; J, Composite
mould of left valve, CEGH-UNC 19796; L, Dorsal view of articulated valves CEGH-UNC 26815; K–M,Aviculopecten barrealensis Reed in Du
Toit, 1927.K, Composite mould of right valve, CEGH-UNC 26818; M, Internal mould of left valve, CEGH-UNC 26819. Scale bar for all figures
= 2.5 mm except J (=5 mm). A–E, G, lower fossiliferous interval, bed 8; F, lower fossiliferous interval, bed 7; J, lower fossiliferous interval, bed
3–4; H, I, K–M, upper fossiliferous interval, bed 9.
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 13
Simanauskas (1999, p. 118, figs 2a–l), in a review of
Aseptella in Argentina. Additional specimens figured
herein (Fig. 4Q–T) clearly conform to that species. The
size and ornament of the specimens described and figured
as Aseptella? sp. by Lech et al.(1998,p.405,fig. 3h),
from the Precordillera, suggest that those specimens are
also attributable to Aseptella sp. aff. A. patriciae.
Superfamily LINOPRODUCTOIDEA Stehli, 1954
Linoproductoidea indet. (Fig. 5A–C)
Remarks. Fragmentary material (Fig. 5A–C) comprising
two internal moulds of ventral valves, one with the
countermould, is herein assigned to Linoproductoidea
indet. These specimens are of medium size (about
14 mm long and 16 mm wide), with the ventral valves
gently convex and umbo strongly incurved, having
ornament of fine costellae (about five per 2 mm on the
median–anterior region) and small ears with at least two
spines on the anterior part.
Order ORTHIDA Schuchert & Cooper, 1932
Suborder DALMANELLIDINA Moore, 1952
Superfamily DALMANELLOIDEA Schuchert, 1913
Family RHIPIDOMELLIDAE Schuchert, 1913
Sufamily RHIPIDOMELLINAE Schuchert, 1913
Rhipidomella Oehlert, 1890
Type species.Terebratula michelini Léveillé, 1835;
Early Carboniferous of Europe.
Fig. 7. A, Generic diversity of the postglacial fauna of the El Paso Formation; B, Abundance of specimens of brachiopods and bivalves in the sampled
bed 8 of the lower fossiliferous interval; C, Abundance of specimens of brachiopods and bivalves in the upper fossiliferous interval (bed 9).
14 G.A. CISTERNA et al.ALCHERINGA
Rhipidomella discreta sp. nov. (Fig. 3A–J)
2001 Rhipidomella? sp.; Simanauskas & Cisterna,
p. 214, figs 3a–h.
2016 Rhipidomella sp.; Cisterna & Sterren, figs O–Q, T.
Etymology. The name refers to the delicate nature of
this species.
Holotype. An internal mould of a ventral valve and its
countermould, CEGH-UNC (Fig. 3A–B).
Paratype. An internal mould of a dorsal valve and its
countermould, CEGH-UNC (Fig. 3E–F).
Type locality, unit and age. Eloy Creek, Barreal Hill, San
Juan province, Argentina; upper part of the El Paso For-
mation, Carboniferous (late Serpukhovian–Bashkirian).
Diagnosis. A particularly small species of the genus
(3.5–7 mm wide, 4–7 mm long, 1.5–2 mm thick); sub-
triangular to subcircular in outline with very short hinge
line. Gently convex profile with dorsal valve almost flat
anteriorly with a shallow sinus initiating from the
umbo. Costellae fine, with about five costae and costel-
lae per millimetre near anterior margin; conspicuous
concentric growth lines fine and dense.
Remarks. Material assigned to Rhipidomella discreta sp.
nov. was described and figured previously as Rhipido-
mella? sp. by Simanaukas & Cisterna (2001, p. 214,
fig. 3a–h).These authors provided a detailed description
of the Argentine species, but additional material col-
lected during subsequent fieldwork (Fig. 3A–J) con-
firmed the generic assignment and provided the
characters for the erection of the new species.
Rhipidomella discreta can be differentiated from the
type species by its small size, gently convex profile and
more conspicuous concentric growth lines. Compared
with other Rhipidomella species described from the late
Palaeozoic of South America, Rhipidomella discreta is
similar to Rhipidomella cora (d’Orbigny, 1842) from
the Permian of Perú (Newell et al.1953, p. 91, pl. 16,
fig. 1a–8) in general outline of the shell and its micro-
ornament, but the Argentinean species is notably smal-
ler. Rhipidomella penniana (Derby, 1874), described
from the uppermost Mississippian of the Amazon Basin,
Brasil (Chen et al.2005, p. 915, figs 4.14–4.32, 5.1,
5.2), is a much larger species.
There are few species of Rhipidomella with shells as
small as the new species. Among these are Rhipido-
mella plana Yang from the Carboniferous of China
(Yang et al.1977, p. 314, pl. 6, fig. 6a–c), and the Per-
mian Rhipidomella cordialis Grant, 1976 from Thailand
(Grant 1976, p. 37, pl. 2, figs 31–41; pl. 3, figs 1–53).
The Chinese species is similar to Rhipidomella discreta
in its small size, subcircular outline, short hinge line,
gently convex valves and the presence of fine and dense
concentric growth lines. However, the shallow dorsal
sinus that characterizes Rhipidomella discreta is absent
in Rhipidomella plana. Also, Rhipidomella cordialis
differs from the Argentine species particularly by its
marked biconvexity.
Order TEREBRATULIDA Waagen, 1883
Suborder TEREBRATULIDINA Waagen, 1883
Superfamily DIELASMATOIDEA Schuchert, 1913
Family BEECHERIIDAE Smirnova, 2004
Beecheria Hall & Clarke, 1893
Type species.Beecheria davidsoni Hall & Clarke, 1893;
Upper Carboniferous, USA.
Beecheria patagonica Amos, 1958 (Fig. 5D–F)
Fig. 8. Generalized interpretation of the depositional environment along a glacially influenced fjordal coastine showing the development of different
faunas (modified from Kneller et al.2004).
ALCHERINGA CARBONIFEROUS POSTGLACIAL FAUNAS 15
Remarks. Two internal moulds of articulated specimens
(Fig. 5D–F), one with its dorsal external countermould,
are assigned to Beecheria patagonica Amos, 1958. This
species has been characterized mainly by an elongate
globose shell with the ventral valve being more convex
than the dorsal, the beak incurving strongly, the dental
plates diverging and extending about one-seventh of the
valve length and the dorsal cardinal plate sessile
(Amos 1958, p. 104, pl. II, figs 1–9; Amos 1979, p. 95,
figs a–c). Specimens from the El Paso Formation have
more conspicuous growth lines than the type material,
particularly on the anterior part of the shell, and the
dental plates are subparallel; however, other diagnostic
features (size, general shape of the shell and internal
features) allow confident assignment to Beecheria
patagonica.
Beecheria sp. figured by Cisterna & Sterren (2010,
fig. 5J–K) from the Hoyada Verde Formation is a small
species with subtriangular shell and a conspicuous
dorsal septum compared with Beecheria patagonica.
Acknowledgements
This study was supported by CONICET (Consejo
Nacional de Investigaciones Científicas y Técnicas): PIP
0091 (GAC) and PIP 112-201101-00571 (AFS), and
CIUNT (Universidad Nacional de Tucumán): 26 G-531
(MMV). We are especially grateful to Dr Sun Yuanlin
(Department of Geology, Peking University of China)
for the translated diagnosis of the Chinese species
Rhipidomella plana Yang. Many thanks also to Santiago
Druetta (CICTERRA) for drawing Fig. 8, to the
reviewers (Dr Guang Shi and an anonymous reviewer)
and to the editors Stephen McLoughlin and Benjamin P.
Kear, for their valuable comments and suggestions. We
also thank Dr Beatriz G. Waisfeld (CICTERRA) for the
critical reading of the manuscript.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by Consejo Nacional de Investigaciones
Científicas y Técnicas [grand no PIP 0091, 112-201101-00571];
CIUNT (Universidad Nacional de Tucumán) [grant no 26 G-531].
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