NEW RECORDS OF MARINE GASTROPODS FROM THE LOWER CRETACEOUS OF WEST-CENTRAL ARGENTINA
CECILIA S. CATALDO
Instituto de Estudios Andinos Don Pablo Groeber, Consejo Nacional de Investigaciones Científicas y Técnicas-Universidad de Buenos Aires, Departamento de Ciencias Geológicas, Facultad de Ciencias Exactas y Naturales, Intendente Güiraldes 2160, C1428EGA, Ciudad Autónoma de Buenos Aires, Argentina.
Submitted: September 22nd, 2016 - Accepted: December 14th, 2016
To cite this article: Cecilia S. Cataldo (2017). New records of marine gastropods from the Lower Cretaceous of west-central Argentina. Ameghiniana 54: 405–440.
To link to this article: http://dx.doi.org/10.5710/AMGH.14.12.2016.3053
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Also appearing in this issue: New records of aquatic sloths from the Pliocene of Chile and the southern distribution of subtropical coastal environments.
New species from the Berriasian–Barremian of Patagonia reveal a gastropod fauna with strong Tethyan influence.
Palynology of La Veteada Formation with a abundant lycopsid tetrads as in extra-Gondwanan records of Lopingian–Triassic age.
AMEGHINIANA - 2017 - Volume 54 (4): 405 – 440
ARTICLES
ISSN 0002-7014
NeW ReCORDS OF MARINe GASTROPODS FROM The LOWeR CReTACeOUS OF WeST-CeNTRAL ARGeNTINA CECILIA S. CATALDO Instituto de Estudios Andinos Don Pablo Groeber, Consejo Nacional de Investigaciones Científicas y Técnicas-Universidad de Buenos Aires, Departamento de Ciencias
Geológicas, Facultad de Ciencias Exactas y Naturales, Intendente Güiraldes 2160, C1428EGA, Ciudad Autónoma de Buenos Aires, Argentina.
[email protected]
Abstract. Eleven new records of marine gastropods from several localities in west-central Argentina are hereby presented, including the following new species: Proconulus kotrus, Calliotropis (Riselloidea) ligosta, Lyosoma truquicoensis, Exelissa crassicostata, Cirsocerithium agriorivensis, Cataldia? binodosa, Vanikoropsis? leviplicata and Tornatellaea neuquina. These records were recovered from two Early Cretaceous units of the Neuquén Basin, the Mulichinco (Valanginian) and the Agrio (upper Valanginian–upper Hauterivian/lowermost Barremian) formations. The families Proconulidae, Calliotropidae, Neritidae, Cryptaulacidae, ?Pseudomelaniidae, Vanikoridae, Epitoniidae and Acteonidae are represented whilst one record is yet to be assigned at the family level. Key words. Gastropoda. Neuquén Basin. Argentina. Valanginian. Hauterivian.
Resumen. NUEVOS REGISTROS DE GASTRÓPODOS MARINOS DEL CRETÁCICO INFERIOR DEL CENTRO-OESTE DE ARGENTINA. Se dan a
conocer 11 nuevos registros de gastrópodos marinos procedentes de numerosas localidades en el centro-oeste de Argentina. Entre ellos se encuentran las siguientes especies nuevas: Proconulus kotrus, Calliotropis (Riselloidea) ligosta, Lyosoma truquicoensis, Exelissa crassicostata, Cirsocerithium agriorivensis, Cataldia? binodosa, Vanikoropsis? leviplicata y Tornatellaea neuquina. Estos registros proceden de dos unidades de edad cretácica temprana de la Cuenca Neuquina: las formaciones Mulichinco (Valanginiano) y Agrio (Valanginiano superior–Hauteriviano superior/Barremiano basal). En esta fauna se encuentran representadas las familias Proconulidae, Calliotropidae, Neritidae, Cryptaulacidae, ?Pseudomelaniidae, Vanikoridae, Epitoniidae y Acteonidae, con un registro no asignado aún a nivel familia. Palabras clave. Gastropoda. Cuenca Neuquina. Argentina. Valanginiano. Hauteriviano.
The latest Jurassic–early Cretaceous marine units of the
recovered only from the Agrio Formation, plus several still
one of the most diverse of the coeval basins of southern
of the same basin (pers. obs.).
so far. The upper Valanginian–lowermost Barremian Agrio
tropod fauna of the Mulichinco and Agrio formations of the
see Aguirre-Urreta et al., 2011).
gastropod records are reported, including eight new species.
Neuquén Basin have yielded a rich macroinvertebrate fauna,
South America and certainly the one that is the best known Formation is particularly highly fossiliferous (for a summary
Gastropods from the Tithonian–Berriasian interval of
undescribed gastropods from other early Cretaceous units herein, new systematic information regarding the gas-
Neuquén Basin is provided and eleven hitherto unknown Three records are however left in open nomenclature at
the Neuquén Basin were first reported by Behrendsen
the species level due to scarcity of material or poor preser-
Damborenea et al. (1979) and Manceñido and Damborenea
tropods Proconulus kotrus sp. nov. and Calliotropis (Rise-
(1891, 1892), haupt (1907), Weaver (1931), Camacho (1953),
vation. The new association is composed of the vetigas-
(1984) and yet studied with different degrees of detail.
lloidea) ligosta sp. nov., the neritiform Lyosoma truquicoensis
in total, more recent studies resulted in the synonymy of
Cirsocerithium agriorivensis sp. nov, Paosia? sp., Cataldia? bin-
Although these authors reported about 45 gastropod taxa
several of these records and the arising of new findings
(e.g., Cataldo, 2013; Cataldo and Lazo, 2016) whilst several
records are still under revision (see Supplementary Online Information). So far, more than 30 gastropod species were AMGhB2-0002-7014/12$00.00+.50
sp. nov., the caenogastropods Exelissa crassicostata sp. nov.,
odosa sp. nov., Vanikoropsis? leviplicata sp. nov. and Confu-
siscala sp., and the heterobranchs Tornatellaea neuquina sp. nov. and Tornatellaea? sp..
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MATERIALS AND PRESERVATION
follow Cox (1960). D, maximum diameter of teleoconch; H,
The new records herein presented comprise more than
height of teleoconch; H/D, height to diameter rate; Hap,
Neuquén Basin. Precise stratigraphic ranges are provided
of last whorl to height of teleoconch rate; PA, pleural angle.
500 specimens recovered from 17 localities across the
for each species based on the solid biostratigraphic frame-
height of aperture; Hlw, height of last whorl; Hlw/H, height
work produced by Aguirre-Urreta et al. (2005) and Aguirre-
LOCALITIES AND GEOLOGICAL SETTING
loose three-dimension shells with fine-grained fill as well
15 localities in the Neuquén province and two localities in
Urreta and Rawson (2012). The specimens constitute
The new records presented herein were collected from
as shells embedded in coquina and thin pavements marked
the Mendoza province (Fig. 1). Nine out of ten species were
recrystallized and, except for a few exceptions, exhibit a
species were retrieved from the Mulichinco Formation (Fig.
by different degrees of consolidation. The teleoconchs are
comparable level of taphonomic modification. The apices are, as a rule, broken off and, therefore, the protoconchs could
gathered from the Agrio Formation whilst the remaining 2–3). The species hereby discussed are rather uncommon
and restricted to one or a handful of localities but for a few
not be observed except those of the holotype of Calliotropis
exceptions and thus contrast with other known species
neuquina sp. nov. The fact that the delicate peristomes are
widespread within the basin (e.g., Cataldo, 2014).
(Riselloidea) ligosta sp. nov. and one paratype of Tornatellaea
also almost always chipped hinders taxonomic determination. Still, because abrasion is generally low, the details of the teleoconch sculpture prove well preserved in almost every specimen.
METHODS
All scanning electron microscope (SEM) images were
taken with a FeI Quanta 200 device in low vacuum mode in
the Servicio de Microscopía electrónica de Barrido y Micro-
análisis (SeMFi), Laboratorio de Investigaciones de Meta-
from these units that are very abundant and geographically Within the sedimentary fill of the Neuquén Basin, the
bearing units are part of the Mendoza Group and particu-
larly of its middle and upper sections (Fig. 2). In the Valanginian Mulichinco Formation, continental and volcaniclastic deposits coexist with marine shales and thick carbonates (Legarreta and Gulisano, 1989). The outcrops of this unit are developed mainly in central and northern Neuquén and
the facies grade vertically and laterally from continental to
marginal marine, being mostly marine in the northernmost
area (Schwarz et al., 2011), with sandstones and shales in-
lurgia Física Ingeniero Gregorio Cusminsky (LIMF), Facultad
terbedded with coquinas (Aguirre-Urreta et al., 2008). The
Argentina. Conventional photographs were taken with a
marine beds for which deposition begun with a basin-scale
de Ingeniería, Universidad Nacional de La Plata, La Plata,
Agrio Formation is a thick unit almost totally composed of
number of compact and DSLR plus macro lens cameras; all
transgression (Spalletti et al., 2011). The Pilmatué (lower)
Matthews’ (1973) and Bengtson’s (1988) criteria for the
posed of thick shales interbedded with sandstones, lime-
specimens were coated with sublimated ammonium chloride.
and Agua de la Mula (upper) members are typically com-
usage of open nomenclature were followed.
stones and coquinas within a shallow-platform marine
ted herein is housed in the Collección de Paleontología, Uni-
Avilé Member was the result of a temporary drop in the
Institutional acronyms. The material described and illustraversidad de Buenos Aires, Buenos Aires, Argentina (CPBA)
and the Colección de Paleoinvertebrados, Museo de Ciencias Naturales y Antropológicas J.C. Moyano, Mendoza, Argentina
setting (Spalletti et al., 2001). The middle, fluvial-aeolian relative sea level during the mid-hauterivian (Legarreta and Gulisano, 1989).
(MCNAM-Pi). Additionally, references to specimens housed
SYSTEMATIC PALEONTOLOGY
La Plata, La Plata, Argentina; PRI, Paleontological Research
Class GASTROPODA Cuvier, 1795
in other institutions are provided as follows: MLP, Museo de
Institution, Ithaca, USA; SIPB, Steinmann-Institut für Geolo-
Subclass VeTIGASTROPODA Salvini-Plawen, 1980
Measurements. Morphologic terminology and measurements
Family PROCONULIDAe Cox in Knight et al., 1960
gie, Mineralogie und Paläontologie, Bonn, Germany.
406
Superfamily TURBINOIDeA Rafinesque, 1815
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
Remarks. Gründel (2000) revised this family and its type genus taking into account diagnostic features previously
unnoticed, e.g., an intermediate sculpture between the pro-
toconch and the teleoconch, and stressing the diagnostic
importance of the morphology of the inner lip area.
FOSSIL LOCALITIES 1) Lagunitas Este (S34º36’, W69º30’) Paosia? sp. (1) 2) Lomas Bayas (S34º39’, W69º31’) Paosia? sp. (6) 3) Puerta Curaco (S37º22’, W69º56’) Tornatellaea? sp. (5) 4) Loma de la Torre (S37º20’, W69º48’) Vanikoropsis? leviplicata sp. nov. (2) 5) Chos Malal Norte (S37º21’, W70º23’) Calliotropis (Riselloidea) ligosta sp. nov. (1) 6) El Gasoducto (S37º25’, W69º56’) Cataldia? binodosa sp. nov. (1) 7) Arroyo Truquico (S37º28’, W70º17’) Lyosoma truquicoensis sp. nov. (27) 8) Puesto Canale (S37º36’, W70º2’) Vanikoropsis? leviplicata sp. nov. (1) 9) Pichaihue (S37º47’, W70º13’) Tornatellaea neuquina sp. nov. (15) 10) Agua de la Mula (S34º36’, W69º30’) Exelissa crassicostata sp. nov. (166) Cirsocerithium agriorivensis sp. nov. (23) Paosia? sp. (2) Cataldia? binodosa sp. nov. (2) Vanikoropsis? leviplicata sp. nov. (20) 11) Salado Norte (S38º10’, W70°5’) Cataldia? binodosa sp. nov. (59) 12) Salado Sur (S38º12’, W70º3’) Confusiscala sp. (1) 13) Agrio del Medio (S38º20’, W69º57’) Paosia? sp. (5) 14) Bajada Vieja (S38º25’, W70º0’) Exelissa crassicostata sp. nov. (135) Cirsocerithium agriorivensis sp. nov. (19) Paosia? sp. (1) Cataldia? binodosa sp. nov. (7) 15) Cerro Bayo (S38º41’, W70º4’) Confusiscala sp. (2) 16) Cerro Marucho (S39º26’, W70º10’) Cataldia? binodosa sp. nov. (1) 17) Aguada Florencio (S39º29’, W70º16’) Cataldia? binodosa sp. nov. (6) Figure 1. Map with fossil localities, indicating species and number of specimens (in parentheses) occurring at each site.
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Genus Proconulus Cossmann, 1918 (= Cochleochilus Cossmann, 1918)
Type species. Trochus guillieri Cossmann, 1885 (see also Cossmann, 1918, pl. 9, fig. 30), Bathonian, France; by original designation.
Remarks. According to the records reported by various authors (Cossmann, 1918; Cox in Knight et al., 1960; Calzada,
the basis of a subtle difference regarding inner lip construc-
tion: while there is a knob on the adapical end of the col-
umellar groove in Cochleochilus, in Proconulus there is no
such knob but a thickening of the adapical end of the right
wall that borders the columellar groove. however, as Gründel (2008) stated, the implications of such subtle difference
are yet to be clarified. With only a handful of species based
1989a; Szabó, 2008), this genus ranges from the Lower
on badly preserved material failing to conclusively exhibit
almost worldwide, though more widely distributed during
mann, 1918; Fischer and Weber, 1997), Cochleochilus is a
Proconulus is strongly similar in overall shape and sculp-
fig. 34) and Fischer and Weber’s (1997, pl. 20, fig. 7) figures
for trochiform, anomphalous shells with a prosocline and
like that of Proconulus, i.e., with a thickening instead of a
Jurassic (Sinemurian) to the Upper Cretaceous and occurs the Jurassic than during the Cretaceous.
ture to Cochleochilus, a genus erected by Cossmann (1918) round aperture as well as a columellar callus bearing a
the supposedly diagnostic inner lip morphology (see Coss-
rather enigmatic genus. Moreover, Cossmann’s (1918, pl. 9, of the type species illustrate an inner lip morphology just
knob. It is thus herein concluded that Cochleochilus is not
groove and a knob on its adapical end. The type species is
thoroughly supported by morphological features and should
d’Orbigny, 1850, sensu Fischer and Weber, 1997, p. 119, pl.
Another similar genus is Calliostoma Swainson, 1840,
Trochus diomedes d’Orbigny, 1850 (= Trochus cottaldinus
thus be regarded as a junior synonym of Proconulus.
20, figs. 6–7; see also Cox in Knight et al., 1960, p. 253, fig.
but, according to Cossmann (1918), it can be distinguished
diagnosis of Proconulus. however, this author considered
its abapical end and a notch that separates it from the
163.16), which, according to Gründel (2008), fits the revised
that Proconulus and Cochleochilus could be distinguished on
from Proconulus by its smooth columellar lip with a knob on basal lip.
Figure 2. Comparative stratigraphic chart of the units outcropping in southern Mendoza, north-central Neuquén and southern Neuquén. Based on Leanza and hugo (1997), Leanza et al. (2001) and Sruoga et al. (2005). Fm., formation; Gr., group; L., Lower; Mbr., member; pars., partially; U., Upper.
408
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
Proconulus kotrus sp. nov.
The construction of the columellar lip is comparable to
Figure 4.1–6
that of Proconulus coelotropis (Schmidt, 1905) and Proconu-
Diagnosis. Proconulus with very fine spiral threads, mature
Jurassic (upper Oxfordian) of Poland (see Gründel and Kaim,
lus viadrinus (Schmidt, 1905), both species from the Upper
whorls with concave-convex profile and suture dropping
2006, p. 128–130, figs. 6–7).
Description. Shell small, trochiform, slightly taller than wide,
an abapical displacement of the adapical suture, making the
ever lower, and broadly rounded periphery.
with moderately low apex and regularly expanding whorls. Protoconch not preserved. Teleoconch with three and a
half to four preserved whorls. Spire conical, representing
In the mature whorls of the holotype of P. kotrus there is
shell outline change from conical to slightly cyrtoconoid.
This is consistent to what whas pointed out by Dambore-
nea and Ferrari (2009) when comparing Lithotrochus Conrad,
approximately 25% of the total shell height. First and
1855 with Proconulus. Both paratypes present fewer whorls
concave, abapically bordered by poorly defined suprasutural
outline. The lateral profile of whorls is also marked by onto-
second spire whorls flat-sided; third whorl slightly more
spiral carina. Suture impressed, slightly channeled from
third whorl onwards where it drops ever lower. Last whorl
large, with sinuous profile: concave profile in adapical half
than the holotype and their shells therefore retain a conical
genetic variability in this species; that is, the younger the
whorl, the more noticeable its concave-convex lateral pro-
file will be. The sculpture in P. kotrus is composed of nu-
and convex towards a markedly rounded periphery. Base
merous, very thin spiral threads that are almost invisible to
callus. Aperture only partially known, apparently holosto-
The oldest records of Proconulus correspond to the Sine-
flat to slightly convex. Umbilicus closed, covered by inner lip
the naked eye.
mate, round and strongly prosocline. Outer lip not pre-
murian of hungary (e.g., Szabó, 2008). Most known species
narrow and shallow groove, slightly more excavated on its
other peri-Tethyan localities. Only a handful of early Cre-
served. Columellar lip wide and crescent-shaped, with adapical end. Inner margin of columellar lip raised as ridge
that thickens on its adapical end forming a transversely expanded protuberance. Sculpture of fine, equidistant spiral
were recovered from the Middle Jurassic of europe and
taceous species are so far known: Proconulus paskensis
Beisel, 1983 (p. 58, pl. 2, fig. 4), from the lower Valanginian of Siberia; Proconulus sp. (Mongin, 1979: p. 112, fig. 1), from
threads. Growth lines strongly prosocline, forming shallow
the Valanginian of Tunisia; Proconulus eixarchi Calzada and
forming growth rugae near aperture.
lower Aptian of eastern Spain; Proconulus hiraigensis Kase,
prosocyrt sinus just below adapical suture and then straight,
Forner, 2006 (p. 49, figs. 1–2), from the upper Barremian–
Occurrence. The species occurs at the Salado Sur locality,
1984 (p. 64, pl. 4, figs. 2–5, 11–12), from the upper Aptian–
C. schlagintweiti Subzone in the Agua de la Mula Member of
sis Collignon, 1972 (p. 12, pl. 1, figs. 10–13), from the Albian
central Neuquén, in late hauterivian beds belonging to the the Agrio Formation.
Material. Three specimens; holotype CPBA 21292.1 and two paratypes, CPBA 21292.2 and 21292.3.
Derivation of name. Latinized form of the Mapuche adjective
kotrü, salty, bitter, literal translation of the name of the type and so far only locality.
Dimensions. holotype: h= 11.1 mm; hlw= 8.3 mm; hap= 5.2
mm; D= 9.3 mm; D/h= 0.84; hlw/h= 0.75; PA= 49.7º.
Paratype (CPBA 21292.2): h= 10.6 mm; hlw= 8.3 mm; hap= 5.7 mm; D= 10.4 mm; D/h= 0.98; hlw/h= 0.78; PA= 61º.
Discussion. Proconulus kotrus sp. nov. matches the diagnosis
of Proconulidae and Proconulus in the sense of Gründel’s (2000) revision.
lower Albian of Miyako, Japan; and Proconulus chebeikiaen-
of Morocco; among others. These species differ from P. kotrus in having a different shell and/or whorl profile (e.g., P.
hiraigensis, P. chebeikiaensis, P. eixarchi, Proconulus sp.) and/or
in lacking a protuberance in the columellar lip (e.g., P. hirai-
gensis, P. paskensis, P. eixarchi).
The representatives of this genus were seemingly
present in the region since the early Jurassic, as indicated by
the record of Proconulus? argentinus Ferrari, 2015a, from the
Pliensbachian of Neuquén. This species differs from P. kotrus in presenting a conical instead of a cyrtoconoid adult shell
and flat-sided to slightly convex whorls instead of a concave-
convex whorl profile, as well as a more angular periphery.
Proconulus kotrus is very similar to the type species of
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
originally proposed Calliotropidae as a tribe of the eucyclinae and Bandel (2010) subsequently elevated it to family.
The main shell feature distinguishing eucyclids consists
of the fact that the first teleoconch whorls exhibit collabral ribs (e.g., Gründel, 1997, pl. 3, figs. 5–7; Kaim, 2004, figs. 6A4, 6-B2, 9-e5; Bandel, 2010, pl. 9, figs. D–G).
Genus Calliotropis Seguenza, 1903 Type species. Trochus ottoi Philippi, 1844, Pliocene–Pleistocene, Italy; by original designation. Currently living in the Mediterranean Sea and the Ne Atlantic Ocean (Kaim, 2004; Stilwell, 2005; Bandel, 2010).
Subgenus Riselloidea Cossmann, 1909a Type species. Risellopsis subdisjuncta Cossmann, 1908, Pliensbachian, France; by original designation.
Remarks. The great morphological variability and similarity
among the species of Calliotropis and Riselloidea was repeatedly pointed out by numerous authors over the last
decades. In some works they were treated as independent
genera whilst, in others, a synonymy was put forth (e.g., Gründel, 1997, 2000; Nützel and Senowbari-Daryan, 1999;
Nützel et al., 2003; Kaim, 2004; Stilwell, 2005; Szabó, 2008; Bandel, 2010). Some distortion was introduced by Wenz’s
(1938, fig. 552) illustration of the type species of Riselloidea,
Figure 3. Stratigraphic range of the species herein described. Ammonoid biozonation from Aguirre-Urreta et al. (2005) and AguirreUrreta and Rawson (2012). Discontinuous lines indicate inferred occurrences. LOW., lower; VAL., Valanginian; B., Barremian; other abbreviations as in Figure 2.
Cochleochilus as illustrated by Cossmann (1918, pl. 9, figs. 31–
34). Such resemblance, plus the fact that even the construc-
which is quite inaccurate and significantly differs from
Cossmann’s original (1908, pl. 2, figs. 10–12). Also, the remarkable longevity of Calliotropis, recorded from the lower Mesozoic upwards, constituted an underlying problem
which led to a great systematic instability both at the generic and the suprageneric levels. Recently, Ferrari et al.
(2014) proposed that Riselloidea is in fact a temporal subgenus of Calliotropis and the problem of the longevity of
tion of the inner lip seems equivalent in both species, supplies
Calliotropis and the similarity between both genera was
Superfamily SeGUeNzIOIDeA Verrill, 1884
Triassic (e.g., Cossmann, 1909a; Nützel and Senowbari-
further reasons to question the independence of Cochleochilus.
Family eUCYCLIDAe Koken, 1897
thus tackled.
Riselloidea is represented by records as early as Late
Daryan, 1999) although it was most diverse and widely distributed during the early and Middle Jurassic. Thus far, only
Remarks. Recently, Ferrari et al. (2014) revised the type
two species were reported from the Lower Cretaceous (see
posed a new classification scheme in which Calliotropidae
The diagnosis of Trochonodus Nützel, hamedani and
genus of eucyclidae and other related genera and prois synonymous of eucyclidae. hickman and McLean (1990)
410
discussion below).
Senowbari-Daryan, 2003 falls within the morphological
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
variability of Riselloidea. The differences between these
without prominent nodes, are based on Wenz’s (1938)
presents a sculpture dominated by collabral elements
subdisjuncta and other species of the genus, including the
genera mentioned by Nützel et al. (2003), i.e., that Riselloidea
aforementioned illustration of R. subdisjuncta. Actually, R.
Figure 4. 1–6, Proconulus kotrus sp. nov.; 1–3, 6, holotype, CPBA 21292.1; 1–3, apertural, lateral, and basal view; 6, SeM close-up of sculpture, the arrow points towards apex of shell; 4–5, paratype, CPBA 21292.2, abapertural and apertural view. 7–12, Calliotropis (Riselloidea) ligosta sp. nov., holotype CPBA 21291; 7, apertural view; 8–9, SeM images, abapertural and lateral view; 10, basal view; 11–12, SeM images, details of the protoconch and first teleoconch whorls. 13–16, Lysosoma truquicoensis sp. nov.; 13–14, holotype, CPBA 21364.1, abapertural and apical view; 15–16, paratype, CPBA 21364.6, abapertural and apertural view. Scale bars: 1–5= 2 mm, 6= 750 µm, 7–10= 1 mm, 11= 500 µm, 12= 250 µm, 13–16= 5 mm.
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
one described below, exhibit nodular sculpture (see Coss-
fainter and more irregular towards last whorl. Growth lines
odus is therefore herein considered a synonym of Riselloidea.
Occurrence. The only known specimen of this species was
mann, 1909a; Ferrari et al., 2014; among others). TrochonThe species described below is very similar to R. sub-
disjuncta, including the varieties described by Cossmann (1916), and, at the same time, resembles some extant and
straight, strongly prosocline.
retrieved from a late hauterivian bed close to the top of the
Agrio Formation, north of Chos Malal city. The bed is provi-
sionally attributed to the C. diamantensis/P. groeberi zones
fossil species of Calliotropis, including its type species.
of the Agua de la Mula Member.
ing the close relationship between Calliotropis and Riselloi-
Derivation of name. Latinized form with a change of endin-
Therefore, this species supplies further evidence support-
dea.
Calliotropis (Riselloidea) ligosta sp. nov. Figure 4.7–12
Diagnosis. Calliotropis without primary spiral threads or
node-rows on the base; third spiral node-row at periphery of
Material. holotype, CPBA 21291.
gof the modern Greek adjective λιγοστός (ligostós), scant,
scarce, referring to the paucity of material of this species.
Dimensions. holotype: h= 7.4 mm; hlw= 5.5 mm; hap= 3.8 mm; D= 6.7 mm; D/h= 0.9; hlw/h= 0.74; PA= 50.6º.
Discussion. The smooth, planispiral protoconch whorls and convex early teleoconch whorls with thin collabral ribs enable a safe attribution to the eucyclidae. Moreover, the
last whorl; periphery broadly angular; nodes large and round.
gradate to trochiform shell outline together with the spire
form, slightly taller than wide, with regularly expanding,
conspicuous while the adapical one is adjacent to the su-
Description. Shell small, broad and low, gradate to trochiimbricate whorls. Protoconch with approximately one to
one and a half planispiral and smooth whorls. Teleoconch
with four and a half angulated whorls. Spire low, repre-
whorls with two node-rows, the abapical one being more
ture, and the rudimentary collabral ribs, the canaliculate sutures and the prosocline aperture based on growth lines, as well as the straight columellar lip, all match the diagnoses
senting approximately 25% of the total shell height. First
provided for Calliotropis and Riselloidea.
thocline to slightly prosocline collabral ribs and one very faint
other species of the subgenus in presenting a broadly an-
strengthening gradually towards subsequent whorls, de-
rows below it, being inconspicuous, prosocline growth lines
whorl and a half of teleoconch convex, with ca. 30 thin orspiral thread delimiting the adapical third. Spiral thread
Calliotropis (Riselloidea) ligosta sp. nov. differs from most
gular and nodose periphery and lacking spiral cords or node-
veloping into conspicuous carina near mid-whorl, flanking
and very thin secondary spiral threads the only sculpture on
platform widening towards later whorls. Rounded nodes
C. (R.) ligosta is so characteristic that it calls for a formal
broad, flattish to slightly concave sutural platform. Sutural
appearing by third whorl, ca. 15 per whorl. Subsutural row of
nodes appearing by third whorl and a half; nodes smaller,
connecting with those of central row through faint, wide,
prosocline collabral ribs that nearly disappear by last
the base. Despite being represented by a single specimen, description. If more specimens were to be collected in the
future, its generic attribution and specific distinctiveness are unlikely to change.
The species described herein is considerably similar to
whorl. Lateral whorl face flat and vertical. Suture canalicu-
several recent species of Calliotropis, including its type
much less prominent nodes. Base convex, without primary
node-row at its periphery and a similar shell profile as well,
late. Last whorl expanded. Periphery broadly angular, with
spiral threads or nodes. Umbilical area covered by narrow inner lip callosity. Peristome not fully preserved. Aperture
apparently holostomate, round, prosocline. Outer lip unknown; parietal lip forming narrow, clearly delimited callus;
columellar lip straight to slightly concave and slightly thickened. Secondary sculpture of fine spiral threads covering all whorls, even superimposed to nodes. Nodes becoming
412
species. In fact, C. ottoi presents a third peripheral spiral
even though the profusion of concentric node-rows at its
base and the overall smaller size of the nodes distinguish it from the Argentine early Cretaceous species. Riselloidea subdisjuncta is also very similar to C. (R.) ligosta, especially
as regards its strongly angular whorl profile, its canaliculate
suture and the fact that it presents a third peripheral carina.
however, this peripheral carina is smooth while the one in C.
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
(R.) ligosta bears wide and somewhat spirally-elongate
sculpture is lost as well. however, its conical shape, its
cords at its base.
two node-rows adjacent to the sutures somewhat resem-
nodes. Riselloidea subdisjuncta also exhibits concentric spiral Only a few Riselloidea species recorded from the Lower
Cretaceous are referred to in the available literature. In ad-
slightly convex base, its moderately wide umbilicus and the ble those of Calliotropis.
Representatives of the genus and subgenus were
dition to establishing the genus, Cossmann (1908) pointed
present in the region from as early as the early Jurassic
record from the “Neocomian” of Germany. This species,
(Riselloidea) keideli Ferrari, Kaim and Damborenea, 2014 (p.
out that Wollemann (1908) had reported an almost certain
(Pliensbachian–Toarcian), with the record of Calliotropis
“Trochus” stillei Wollemann, 1908 (p. 174, pl. 12, fig. 3, pl.
1181, figs. 4.1–13, 5.1–8) from Neuquén and Chubut
considering its trochiform, anomphalous shell, the moder-
(R.) keideli in the profile of its spire whorls and the canalicu-
riphery of the last whorl and the unconspicuous prosocline
other aspects. For instance, the overall shell profile of C. (R.)
13, fig. 2), is certainly similar to R. subdisjuncta and, when
ately deep sutures it presents, the two node-rows in the pe-
provinces. Calliotropis (R.) ligosta is somewhat similar to C.
late suture. Yet, these two species prove very different in
collabral ribs interconnecting the nodes, can be assigned to
keideli is taller and pagodiform and its last whorl is more
adapical, subsutural row of nodes, and such presence can-
distinct spiral cords, including a node-row bordering the
Riselloidea. however, Wollemann (1908) did not mention an not be ascertained based on the illustrations either. For this reason and until the meaning of such difference is better
understood, it might be more cautious to regard this attri-
elongate and has a strongly convex base bearing five
pseudoumbilicus, thus resembling the genus Ambercyclus
Ferrari, Kaim and Damborenea, 2014. Also, the nodes are
smaller, sharper and more numerous than in C. (R.) ligosta.
bution as tentative. Riselloidea? stillei differs from C. (R.)
Similar records, such as C. (R.) cf. keideli and Calliotropis
between the mid-whorl and the peripheral node-rows.
in basically the same traits that characterize C. (R.) keideli.
ligosta in bearing less deep sutures and a concave surface In the same work, Wollemann (1908, p. 173, pl. 12, fig.
2) additionally illustrated “Trochus” quadricoronatus harbort,
1905, which is very similar to R.? stillei and could also be
attributed to Riselloidea. This species differs from C. (R.) ligosta in presenting a more conical, less trochiform and almost cyrtoconoid shell profile.
(Riselloidea) sp. (Ferrari et al., 2014), differ from C. (R.) ligosta
Subclass NeRITIMORPhA Koken, 1897
Superfamily NeRITOIDeA Rafinesque, 1815 Family NeRITIDAe Rafinesque, 1815
Remarks. It is because of a calcitic outer layer (see Kaim and
Beisel (1983, p. 56, pl. 2, fig. 3) described two poorly-
Sztajner, 2004, and references therein) that neritids are
and attributed them to Riselloidea sp. This record differs
tropods with entirely aragonitic shells present in the same
preserved specimens from the lower Valanginian of Siberia
from C. (R.) ligosta in exhibiting a row of paired nodes along
the periphery of the whorls and several smooth spiral ribs in
generally better preserved in comparison with other gaslithologies. however, Saul and Squires (1997) indicated that, in spite of the broad stratigraphic range of the family
the base.
(Triassic–Recent; Cossmann, 1925; Keen and Cox in Knight
compared with from nearby early Cretaceous basins of
derrepresented in the fossil record due to their preference
genus and subgenus in the Cretaceous of the southern
have been several reports of neritids in shoreface environ-
der I Island, Antarctic Peninsula, is somewhat similar. Am-
the early Cretaceous gastropod fauna of the basin (see
Calliotropis (Riselloidea) ligosta lacks similar forms to be
South America, actually representing the first record of the
world. Only one form from the Lower Cretaceous of Alexanphitrochus sp., as reported by Thomson (1971, p. 47, fig. 2j), is too badly preserved to attempt a generic revision, for the
apical whorls and aperture are missing and most of the
et al., 1960; Tracey et al., 1993), neritids are normally unfor rocky-shore environments. In the Neuquén Basin, there
ments, thus making this family one of the most diverse of Supplementary Online Information). The Neritidae attained a cosmopolitan distribution during the Jurassic.
413
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
Genus Lyosoma White, 1883
genus to confirm that the diagnostic smooth columellar
Type species. Neritina powelli White, 1876, Middle Jurassic, USA; subsequent designation by Fischer (1885).
so far attributed to it, a task beyond the scope of this paper.
Remarks. The identification of Jurassic and Cretaceous neritid genera is complex because the shell feature that bears the
apparently highest diagnostic value, i.e., the inner lip sep-
tum, is not always preserved (Kase, 1984). Shell sculpture in
Mesozoic neritids is generally not conclusive enough by its
edge of the inner septum is indeed present in all the species
Both Lyosoma and Lissochilus present an inner septum
without denticles on the free margin and prove alike in terms of the shell outline as well. Sohl (1965) distinguished Lissochilus from Lyosoma on the basis of a taller spire, a
more developed spiral sculpture and a bicarinate periphery.
own given that it can be highly convergent. examples of
So far, a few Lissochilus species have been recorded from
resemblance between Otostoma d’Archiac, 1859 and Lyo-
Supplementary Online Information) and, although their sys-
such inconvenience can be perceived when observing the
soma, as well as between the latter and Lissochilus Pëtho in zittel, 1882 (see below). The sculpture combined with the
nature of the margin of the apertural inner septum would
the Lower Cretaceous of west-central Argentina (see
tematics require further study, they all exhibit conspicuous nodular spiral sculpture.
Lyosoma powelli and L. enoda, both from the USA, are the
constitute a better diagnostic base.
oldest records of the genus (Middle Jurassic). The list of
blematic genus based on poorly preserved material, and
cludes two taxa from the Lower Cretaceous of europe
established. The authors additionally pointed out that the
“Senonian” of Brazil (White, 1888). White’s (1888) species
smooth columellar edge on the inner lip septum, could prove
thus far attributed to the genus.
Bandel and Kiel (2003) argued that Lyosoma is a pro-
whose systematic position has not yet been adequately
main difference between Lyosoma and Otostoma, i.e., a
an artifact of preservation. This may be certainly so, given that the inner septum is only known, according to Sohl
(1965), in two out of the more than 80 known specimens of
L. powelli. Nevertheless, Sohl (1965) added the lack of a ca-
Lyosoma species provided by Cossmann (1925, p. 206) inwhilst the youngest record consists of one species from the and the one described below are the only southern records
Lyosoma truquicoensis sp. nov. Figure 4.13–16
rina in the last whorl of the species of Otostoma to the list of
Diagnosis. Lyosoma with small- to mid-sized, smooth shell
its character as an independent genus. Afterwards, Calzada
the only sculpture, adapical third of last whorl raised above
differences with regard to Lyosoma and therefore affirmed
(2000) pointed out another diagnostic feature of Lyosoma:
a subsutural welt. Another species recovered from an approximately equivalent stratigraphic interval as that from
which L. powelli was collected is Lyosoma enoda Sohl, 1965
(p. 18, pl. 2, figs. 11–13, 15–24), from the Middle Jurassic of
with fine, slightly prosocline and prosocyrt growth lines as the peripheral carina and well-developed subsutural welt.
Description. Teleoconch medium-sized to small, nearly as
tall as wide, anomphalous, sub-globose in outline, with short and wide spire. Protoconch not preserved. Suture slightly impressed. Base wide and gently convex. Last whorl
Utah and Wyoming, USA. The inner septum is not yet known
very expanded, representing more than 90% of the total
of L. enoda with several specimens of the type species and,
platform and wide and convex subsutural welt. Shallow to
in this species. Sohl (1965) directly compared the specimens
on the grounds of a similar overall shape, concluded that
shell height; with narrow and nearly horizontal sutural well-marked spiral depression between subsutural welt
they were congeneric.
and peripheral carina; peripheral carina strong, wide and
similar to L. enoda and is closer to the revised diagnosis of
Aperture wide. Outer lip thin, angular at peripheral carina,
The new species described below proves remarkably
Lyosoma provided by Sohl (1965) than to any other neritid
genus. It is for these reasons that the species is herein in-
cluded within Lyosoma pending a thorough revision of the
414
rounded. Whorl side more or less evenly convex up to base.
convex adapically to carina and straight to slightly concave
abapically, nearly vertical up to union with wide basal lip. Inner lip callous, extended inwards as a wide septum with a
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
smooth columellar margin. Teleoconch smooth but for very
veloped subsutural welt and a spiral depression above the
Occurrence. This species has been recorded from a single
sinuous, prosocline collabral ribs that extend from the
Agrio Formation, at a single locality of central Neuquén:
White (1888) attributed one species from the “Senonian”
fine growth lines, slightly prosocline and prosocyrt.
level within the O. (O.) atherstoni Subzone at the base of the
peripheral carina, as well as a sculpture composed of thin, adapical suture up to the base of the last whorl.
Arroyo Truquico.
of Brazil to Lyosoma: L. squamosa White, 1888 (p. 179, pl. 10,
holotype CPBA 21364.1 and two paratypes: CPBA 21364.5
senting conspicuous, lamellous collabral ribs that become
Material. Twenty-seven specimens, CPBA 21364.1–27; and 21364.6.
figs. 25–27). This species differs from L. truquicoensis in pre-
more accentuated a quarter of a whorl before the aperture.
Derivation of name. Named after Arroyo Truquico, the type
Additionally, the lamellous ribs form spiny tubercles at the
Dimensions. holotype: h= 47.2 mm; hlw= 46.8 mm; hap=
1934 (p. 58, pl. 4, fig. 8), from the Upper Cretaceous of Peru,
locality in the Neuquén province.
intersection with the shoulder. Desmieria peruviana Olsson,
43.8 mm; D= 52 mm; h/D= 0.9; hlw/h= 0.99; PA= 104º.
is somewhat similar to the type species of Lyosoma in
25 mm; D= 30.7 mm; D/h= 0.98; hlw/h= 0.96; PA= 115º.
guished in terms of its coarse node rows along the subsu-
Paratype (CPBA 21364.5): h= 30 mm; hlw= 28.7 mm; hap= Discussion. Lyosoma truquicoensis sp. nov., which was re-
trieved form a single bed, is represented by several speci-
outline and sculpture. however, D. peruviana can be distintural welt and peripheral carina. It is worth noticing that the inner septum in D. peruviana is still unknown. Squires and
mens exhibiting characteristics corresponding with the
Saul (1993) attributed this species to another controversial
including the non-dentate columellar edge. This species
the basis of its external morphology.
features indicated in Sohl’s (1965) diagnosis of Lyosoma,
can be readily distinguished from other neritids present in
genus, i.e., Corsania Vidal, 1917 (see also Calzada, 2000), on The neritid record reported by Alarcón and Vergara
the Agrio Formation due to the absence of both spiral and
(1964, p. 115, pl. 4, fig. 10) as Lyosoma sp. aff. squamosa and
Lyosoma truquicoensis is most similar, both in shell out-
Antofagasta, Chile, bears a well-developed spiral sculpture
collabral sculpture.
line and lack of sculpture, to L. enoda from the Bathonian–
Bajocian of the USA. These species can be distinguished because L. truquicoensis is somewhat taller and the adapical
third of its last whorl is raised higher above the peripheral carina, and presents a more developed subsutural welt.
Cossmann (1925) attributed two early Cretaceous
species to Lyosoma. According to Sohl (1965), however, these species cannot be placed with certainty within the genus since their inner septum is in fact not known.
recovered from the Lower Cretaceous of the el Way area in
formed by several thin rows of nodes below the peripheral
suture and is therefore not consistent with the diagnosis of Lyosoma.
Subclass CAeNOGASTROPODA Cox in Knight et al., 1960 Superfamily CeRIThIOIDeA Fleming, 1822 Family CRYPTAULACIDAe Gründel, 1976
Remarks. Cryptaulacids are characterized by a protoconch
Lyosoma? capduri (Cossmann, 1907, p. 22, pl. 4, figs. 23–26)
consisting of one to one and a half smooth whorls followed
the peripheral carina and poorly-developed subsutural welt
The onset of the teleoconch is marked by the appearance of
differs from L. truquicoensis in presenting coarse nodes along
by bicarinate whorls in which the carinae gradually develop.
and spiral depression, as well as an apparently taller spire.
the collabral sculpture. Besides, the aperture exhibits some
differs from the Argentinian species in exhibiting a rather
gued that the concept of the family, originally proposed as
veloped spiral sculpture at and below the peripheral carina.
the close resemblance between Cryptaulax Tate, 1869 and
description for the species Lyosoma gonii Bataller, 1959,
which, according to him, may even be synonymous.
Lyosoma? ornatissima (Coquand, 1865, p. 74, pl. 5, figs. 4–5)
discoidal, much more depressed shell outline and a well-deRecently, Calzada and Corbacho (2015a) provided a new
which differs from L. truquicoensis in bearing a poorly de-
kind of basal modification (Guzhov, 2004). Bandel (2006) ar-
a subfamily of Procerithiidae, is difficult to sustain given Procerithium Cossmann in Chartron and Cossmann, 1902, In the Southern hemisphere, cryptaulacids have been
415
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
so far recorded from the Upper Triassic of Peru (haas, 1953;
2006). Guzhov (2004) defined the subfamily exelissinae
Gründel, 2001; Gründel and Parent, 2001, 2006; Ferrari,
which includes several other genera in addition to Exelissa,
Ferrari, 2015b), the Jurassic of Chile and Argentina (e.g., 2012), and the Lower Cretaceous of Argentina (this work).
This family was much less diverse in Cretaceous than in Jurassic faunas (Guzhov, 2004, tab. 1).
Genus Exelissa Piette, 1860 (= Kilvertia Lycett, 1863) Type species. Cerithium strangulatum d’Archiac, 1843 (p. 382, pl. 31, fig. 1a, b), Bathonian, France; by original designation.
Remarks. The earliest discussions regarding the type species
of Exelissa dealt with whether it presented an anterior canal. While d’Archiac (1843) described one, Piette (1860) studied
several topotypes and confirmed that the aperture was
holostomatous. Moreover, Lycett (1863), who defined Kil-
vertia based on the same type species thus establishing a
of the Cryptaulacidae based on Exelissa. This subfamily, differs from the Cryptaulacinae in presenting a last whorl of conspicuous morphology.
Considering that most protoconchs described for the
genus are morphologically more similar to Cryptaulax and
following the criteria of several recent authors (e.g., Guzhov,
2004; Gründel and Kaim, 2006; Gründel and Parent, 2006),
Exelissa is herein considered a representative of the Cryptaulacidae. Additionally, the Procerithium-like protoconchs
illustrated by Gründel (1999) for a few Exelissa species were described by Kaim (2004) as similar to the protoconch of a
Cryptaulax species from the Lower Cretaceous of Poland. Guzhov (2004) simultaneously reassigned those same species to a different genus.
According to Cossmann in Chartron and Cossmann
junior objective synonym of Exelissa, confirmed Piette’s
(1902), the earliest record of Exelissa corresponds to the
sometimes constricted at the base, and the peristome is
described hereunder, the stratigraphic range of the genus
(1860) notion and added that the last whorl is cylindrical and
hettangian (early Jurassic) and, considering the species
slightly thickened and expanded. According to Cossmann
can be extended up to the hauterivian (Lower Cretaceous).
straight to slightly slanted groove at the end of the columella
genus was widespread especially during the Jurassic, with
(1906), the basal lip only implies the presence of a short,
but does not actually confirm the presence of a real notch
Although uncertainty about certain records prevails, the
records across europe, North America, eastern Africa and
or canal.
southern South America.
phasizing the teleoconch morphology is followed. The
cerithioid genera in which the aperture is detached from the
present in several species of the genus could be added to
Francocerithium Nützel and Gründel, 2007 and Tomaszoviella
In this work, Kaim’s (2004) diagnosis of the genus em-
constricted nature and abapical shift of the last whorl
such diagnosis. This feature was stressed by Guzhov (2004)
Nützel and Gründel (2007) indicated that there are other
shell base, e.g., Cryptaulax, Rhabdocolpus Cossmann, 1906,
Kaim, 2001. Overall, all these genera differ from Exelissa
to establish the subfamily exelissinae.
in their much stronger nodose sculpture, their more nu-
has been to, although uncertainly, assign Exelissa to the
and the conspicuous spiral cords forming nodes at the in-
sculpture. Given that the protoconch of the type species
and round, orthocline to slightly opisthocline ribs that are
According to Gründel (1997), the traditional position
Procerithiidae on the grounds of teleoconch ontogeny and
remains unknown, the familial affinity of this genus within
merous opisthocyrt and othocline to prosocline collabral ribs
tersections with the ribs. Instead, Exelissa presents wide aligned along the coiling axis providing the spire with a
Cerithoidea has not yet been settled. The protoconchs of a
pyramidal aspect. Also, Cryptoptyxis Cossmann (1906) a
figs. 18, 22; Guzhov, 2004, pl. 4, figs. 4b, 10; Gründel and
elissa in having two columellar folds, a notch in its basal lip
few other Exelissa species (e.g., Gründel, 1999, p. 22, pl. 5,
Parent, 2006, fig. 3D) were described as similar to the protoconchs of either Procerithium or Cryptaulax. Consequently,
opinions on whether belongs within the families Cerithiidae,
Procerithiidae or Cryptaulacidae are divided and, more recently, the family Bitiidae was also suggested (Bandel,
416
pupoid shell with a detached aperture but differs from Exand a conspicuous subsutural ramp (see Cossmann, 1906; Gründel, 1999; Guzhov, 2004). Guzhov (2004) proposed that Cryptoptyxis is a junior synonym of Exelissa on the
grounds of a gradual evolution between the cyrtoconoid shell with seven or eight collabral ribs that characterizes
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
Exelissa and the conical shells with five or six collabral ribs observed in Cryptoptyxis.
Exelissa crassicostata sp. nov. Figure 5.1–6, 5.8
21293.8, 21792.1 and 21792.2.
Derivation of name. From the Latin adjectives crassa, thick, and costāta, ribbed, owing to the sculpture of thick ribs.
Dimensions. holotype (incomplete): h= 8.6 mm; hlw= 5.9 mm; hap= 2.4 mm; D= 4.25 mm; PA= 18º. Paratype (CPBA
21293.8, incomplete): h= 8.05 mm; hlw= 4.9 mm; hap= 2.6
Diagnosis. Exelissa with prominent, coarse collabral ribs and
mm; D= 3.55 mm; D/h= 0.44; hlw/h= 0. 61.
Description. Shell small, cyrtoconoid to pupiform. Apex
cyrtoconoid to pupoid shell outline, the nature of the sculp-
non-nodose spiral sculpture.
acute. Protoconch and first teleoconch whorls not pre-
Discussion. Despite lacking well-preserved apertures, the ture and the known portion of the aperture in the studied
served. Teleoconch with up to eight whorls preserved. Spire
specimens are most reminiscent of the type species of the
height. Suture canaliculated, within shallow depression
of the collabral ribs and the non-nodose overlapping spiral
pyramidal, high, representing nearly 60% of the total shell
abapical to conspicuous spire cord. Spire whorls convex, imbricate, scarcely overlapping. Sculpture of wide collabral
ribs, straight to opisthocyrt and orthocline to slightly prosocline, oblong in shape, with convex top, tapering off towards
both sutures. Ribs equidistant from each other, lined up
genus as well as of other Exelissa species. The prominence
sculpture are the key features that distinguish E. crassicostata from other congeneric species.
In Exelissa crassicostata sp. nov., the narrowing of the
last whorl and the abapical shift of the aperture are weak
and similar to those of Exelissa distans Cossmann, 1913 (see
forming longitudinal crests along coiling axis. Seven to eight
also Gründel and Kaim, 2006, p. 139, fig. 16). The reduction
timate and last whorls by intercalated, out-of-alignment
subtle. even though the small size and thickening and fla-
ribs per whorl on spire, increasing up to 10 to 12 by penuladditional ribs. Surface between ribs flat to concave. Con-
vexity of whorls decreasing with ontogeny. Ribs diminish in strength and become nearly indistinct towards periphery
of last whorl. Numerous fine, band-like spiral cords, 11 to 12 on penultimate whorl, not forming nodes at intersections
in strength of the sculpture towards the periphery is also
ring of the peristome are manifest, the aperture in E. crassicostata is not preserved sufficiently well so as to evaluate its degree of detachment from the last whorl.
The Exelissa species that are most similar to E. crassi-
costata are from the Jurassic and one example is the type
with collabral ribs, with finer spiral threads in between. Pe-
species, E. strangulata. According to Gründel’s (1997) revi-
in between. Suture slightly dropping below periphery by
(1913, p. 116, pl. 5, figs. 42–45), E. strangulata bears seven
riphery marked by two thicker spiral cords, surface concave penultimate whorl. Base concave, with fine spiral threads.
sion of the specimens originally published by Cossmann to eight collabral ribs initially orthocline and straight and
Last whorl shorter and narrower than spire whorls. Aper-
becoming slightly prosocline and opisthocyrt towards the
concave, somewhat thickened and flared; basal and outer
p. 93, pl. 4, figs. 12–13). Towards the last whorl, ribs lose
orthocline to slightly prosocline.
become more conspicuous. Such nodose sculpture distin-
ture partially known, small and seemingly round; parietal lip
lip unknown. Columella indistinct. Growth lines opisthocyrt,
last whorl, just as in the Argentinian species (Gründel, 1997,
much of their relief and the nodose, intersecting spiral cords
Occurrence. The species occurs in the lower hauterivian of
guishes E. strangulata from E. crassicostata.
mation.
and Germany, is also very similar to E. crassicostata. The
Bajada Vieja, O. (O.) laticosta Subzone, CPBA 21293.1–135;
secondary, and the absence of nodes at the intersections
central Neuquén, in the Pilmatué Member of the Agrio ForMaterial. A total of 301 specimens; 135 specimens from
125 specimens from Agua de la Mula, O. (O.) laticosta Sub-
Exelissa distans, from the Oxfordian of France, Poland
number of collabral ribs and spiral cords, both primary and are common to both these species. however, E. crassicostata
zone, CPBA 21792.1–125; 41 specimens from Agua de la
differs from E. distans in bearing coarser ribs and a concave
21293.6 and five paratypes: CPBA 21293.3, 21293.7,
Exelissa arcuatoconcava Gründel and Parent, 2001 (p. 15,
Mula, H. gentilii zone, CPBA 21793.1–41. holotype CPBA
base with two stronger spiral cords at the periphery.
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
fig. 3A–D) (see also Gründel and Parent, 2006, p. 506, fig.
3A–D) was recorded from the lower Tithonian of the
Neuquén Basin, in localities both in the Neuquén and Mendoza provinces. This species presents a slender shell
and its sculpture consists of eight to 10 opisthocline and
opisthocyrt collabral ribs that are stronger at their adapical
records presented above, E. crassicostata could represent
the first early Cretaceous record of the genus.
Genus Cirsocerithium Cossmann, 1906
end forming a round node. There are five to six spiral cords
Type species. Cerithium subspinosum Deshayes in Leymerie, 1842 (p. 14, pl. 17, fig. 12a, b), Albian, France; by original designation.
dose sections, and also four to five cords on the base. The
Cossmann, 1906, especially as regards the gradate spire,
on the whorl lateral face that break the ribs into several nodistinct sculpture distinguishes this species from E. crassi-
costata.
There are very few known Cretaceous species of Exelissa.
Remarks. This genus is very similar to Rhynchocerithium
the outer lip varix, the presence of the anterior canal and the
sculpture. These genera differ mainly in that the nodes in Rhynchocerithium are spine-like, especially right below the
Guzhov (2004) reported three: Pyrazus valanginensis Pche-
suture, as can be observed in the type species, R. fusiforme
Crimea; Cryptoptyxis barremicus Sayn, 1932 (p. 37, pl. 3, figs.
mann, 1906, pl. 6, figs. 18–21). Besides, the anterior canal
lintsev, 1965 (p. 142, pl. 27, fig. 10), from the Valanginian of 2–6, 34, 35, 36), from the Barremian of Spain; and Melania
dollfusi Choffat, 1901 (p. 94, pl. 1, figs. 22–23), from the “Senonian” of Portugal. Pyrazus valanginensis is characterized
by a conical to slightly cyrtoconical pyramidal shell with five
rows of collabral ribs along the spire and three spiral cords
(hébert and eudes-Deslongchamps, 1860) (see also Cossin Rhynchocerithium is more developed than in Cirsocerithium,
where it is more like a notch than a canal and does not project beyond the basal lip margin. Moreover, the outer lip in Cirsocerithium is not sinuous as it is in Rhynchocerithium.
Cossmann (1906) originally placed both Cirsocerithium
on each whorl. Its aperture is not preserved but would seem
and Rhynchocerithium within the subfamily Paracerithiinae
author did not mention a narrowing in the last whorl. Thus,
ture with a basal notch that is not projected beyond the
that it somewhat resembles the teleoconch morphology
this grouping with evidence from protoconch morphology.
cus is similar to E. crassicostata, especially regarding the
sculpture as well, ascribed the similar Rhynchocerithium to
to have a round peristome (Pchelintsev, 1965). The latter
its attribution to Exelissa remains doubtful, considering also
of Alamirifica Saul and Squires, 2003. Cryptoptyxis barremicoarse collabral ribs and the pair of stronger spiral cords at the periphery. however, the Spanish species bears a
(Family Procerithiidae), which is characterized by an aper-
basal lip margin (Guzhov, 2004). Bandel (2006) supported
however, Guzhov (2004), based on protoconch shape and
a different family, the Maturifusidae, and argued that
Paracerithium Cossmann in Chartron and Cossmann, 1902
broader shell and fewer collabral ribs. Given its seven rows
should be revised (see type species in Gründel, 1997;
dollfusi as illustrated by Choffat (1901) resembles that of
sented arguments opposing Guzhov’s (2004) grouping,
of collabral ribs and thin spiral cords, the teleoconch of M.
Exelissa. Nevertheless, its aperture is badly preserved and
seems to have an oval peristome. In view of the doubtful
hikuroa and Kaim, 2007). Nützel and Gründel (2007) prewhilst Tracey (2010) ranked Cirsocerithium in the Cerithii-
dae. Kiel (2006, fig. 3.12, 3.14) illustrated a Cryptaulax-
Figure 5. 1–6, 8, Exelissa crassicostata sp. nov.; 1, 6, 8, paratype, CPBA 21293.3; 1, SeM close-up of sculpture; 6, 8, apertural and abapertural view; 2, holotype, CPBA 21293.6, apertural view; 3–4, paratype, CPBA 21293.7, abapertural and apertural view; 5, paratype, CPBA 21293.8, lateral view; 7, 9–12, Cirsocerithium agriorivensis sp. nov.; 7, 9–10, 12, holotype, CPBA 21801.1; 7, 12, SeM close-ups of sculpture and aperture; 9–10, apertural and abapertural view; 11, paratype, CPBA 21801.2, apertural view. 13–17, 19, Cataldia? binodosa sp. nov.; 13–14, holotype, CPBA 21796, apertural and abapertural view; 15, paratype, CPBA 21797.1, abapertural view; 16–17, CPBA 21797.3, abapertural view and close-up of spiral sculpture; 19, CPBA 21797.4, abapertural view. 18, 20–22, Paosia? sp.; 18, 21–22, CPBA 21294, detail of growth lines on base, apertural and abapertural view; 20, CPBA 21794.1, apertural view. Scale bars: 1, 7, 12= 500 µm, 2–6, 8–11, 17= 1 mm, 13–16, 19= 4 mm, 18= 2 mm, 20–22= 5 mm. Arrows in 1, 7, 12, 17, 18 point towards apex of shells.
418
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
like protoconch for C. collignoni Kiel, 2006 and transferred
jada Vieja, O. (O.) laticosta Subzone, CPBA 21801.1–19; 20
morphology that Kaim (2004) had previously regarded as
CPBA 21802.1–20; two specimens from Agua de la Mula,
to Cirsocerithium two other species with similar protoconch
specimens from Agua de la Mula, O. (O.) laticosta Subzone,
representatives of Cryptaulax. In consideration of the afore-
H. gentilii zone, CPBA 21803.1–2; one specimen from Agua
within Cryptaulacidae. Yet, given its apertural morphology,
CPBA 21801.1, two paratypes: CPBA 21801.2 and 21801.3.
mentioned appreciations, the genus is herein included
it is however difficult to assess which of Guzhov’s (2004) subfamilies of the Cryptaulacidae would Cirsocerithium actually correspond to.
Cirsocerithium agriorivensis sp. nov. Figure 5.7, 5.9–12
Diagnosis. Cirsocerithium with convex whorl profile, sharp or-
de la Mula, C. diamantensis zone, CPBA 21804. holotype Derivation of name. From Agrio and the Latin noun rīvus,
river, because the type locality is by the Agrio River in cen-
tral Neuquén.
Dimensions. holotype: h= 10 mm; hlw= 5 mm; hap= 3.35
mm; D= 4.5 mm; D/h= 0.45; hlw/h= 0.5; PA= 22.1º.
Paratype (CPBA 21801.2, incomplete): h= 8.2 mm; hlw= 5.55 mm; hap= 4 mm; D= 5.1 mm.
Discussion. This Cirsocerithium species clearly displays a
thocline collabral ribs, four spiral cords, no subsutural cord
well-preserved apertural varix in several specimens. Addi-
instead of pointy nodes.
with the end of the columella also justifies the attribution
and secondary threads, and with faint swellings on the ribs
tionally, the shallow pout at the junction of the basal lip
Description. Shell small, cyrtoconoid to subfusiform. Apex
to this genus.
served. Teleoconch with up to seven whorls preserved. Spire
species apart from the type, C. subspinosum. Some species
Spire whorls strongly convex, slightly overlapping, without
agriorivensis sp. nov. are, for instance, C. aptiense (d’Orbigny,
whorl. Ribs sharp, straight, orthocline to slightly prosocline,
collected from the Aptian of Se France (Kollmann, 2005, p.
Surface between ribs concave. Ribs crossed by four equi-
upper Aptian–lower Albian of Japan (Kase, 1984, p. 130, pl.
acute. Protoconch and first teleoconch whorls not prehigh, representing 50% of total shell height. Suture grooved.
subsutural ramp. Sculpture of 10 strong collabral ribs per equidistant, out of alingment between succeeding whorls.
distant spiral threads forming faint swellings at intersections. Last whorl slightly taller than wide. Five visible spiral
The lack of a mid-whorl angulation sets the Argentinian
with a convex whorl profile similar to that of Cirsocerithium
1843) and C. reticulatum (Nagao, 1934). The former was
159, pl. 17, fig. 28) whilst the latter was retrieved from the
20, figs. 12, 13). Both C. collignoni Kiel, 2006 (p. 459, fig.
3.11–14) and Cirsocerithium lallieranum (d’Orbigny, 1843)
ribs. Periphery rounded, marked by one slightly thicker
were recovered from the Albian of Madagascar (Kollmann,
Base convex, slightly concave near columella. Columella
whorl profile. Cirsocerithium aptiense also bears four spiral
spiral thread followed abapically by another spiral thread.
straight. Aperture partially known. Peristome obliquely oval.
2005, p. 160, pl. 17, figs. 26–27) and present a subangular cords on the whorl side and a fifth one just below the adapi-
Outer lip evenly convex; apertural varix well-preserved,
cal suture, a feature that is present in several other species
suture to junction with basal lip, sculptured by prolongation
is not yet clear since all the other features characterizing C.
thick, straight, slightly prosocline, extended from adapical
but absent in C. agriorivensis. The meaning of such attribute
of spiral threads. Inner lip simple and concave. Shallow pout
agriorivensis agree well with the diagnosis of the genus.
true anterior canal.
nosis and stated that the subsutural cord is frequently
at junction of basal lip with end of columella, not forming Occurrence. This species occurs in the hauterivian of central
Neuquén, both in the Pilmatué and the Agua de la Mula
Cossmann (1906) included this feature in the original diagbeaded, as occurs in C. subspinosum.
Regarding the collabral sculpture, C. agriorivensis is
members of the Agrio Formation. In the Pilmatué Member,
characterized by low swellings at the intersections of spiral
costata sp. nov..
nent or even pointy nodes and yet this feature exposes
the species occurs in the same levels as Exelissa crassiMaterial. A total of 43 specimens; 19 specimens from Ba-
420
cords and collabral ribs. Other species present more promisome intraspecific variability. For instance, the syntypes and
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
several other specimens of C. subspinosum from europe bear
Superfamily and family indeterminate
pointy nodes along the mid-whorl carina of the last and
Genus Cataldia Calzada and Corbacho, 2015b
specimens from around the world present blunter nodes (see Abbass, 1973, p. 119, pl. 2, figs. 1–2; Kase, 1984, p.
Type species. Confusiscala caneroti Calzada, 1973 (fig. 1), upper Barremian–lower Aptian, Spain; by original designation.
24–25). The collabral ribs in C. aptiense are thicker and
species from the Lower Cretaceous of Spain. The available
three to four precedent whorls; however, several other
130, pl. 20, figs. 10–11; Kollmann, 2005, p. 159, pl. 17, figs.
wider, and there are deeper-set interspaces between spi-
ral cords. In C. reticulatum, as the specific epithet states, the
Remarks. This is a rather ill-defined genus based on a
material of the type and other species lacks complete adult
apertures. This is a recurrent problem with Mesozoic cerithi-
sculpture forms a reticulate pattern. The ribs are thinner
form gastropods that precludes establishing affinities at a
closely spaced towards the aperture while there are four
2009).
than in C. agriorivensis and become even thinner and more
main spiral cords with finer threads intercalated. Cirsoceri-
suprageneric level (Saul and Squires, 2003; Ozawa et al., The diagnosis provided by Calzada and Corbacho
thium collignoni exhibits two to three-noded spiral cords and
(2015b) mentioned a homeostrophic, paucispiral and
Two species from the Cretaceous of england provi-
teleoconch, slightly convex whorls, tuberculate collabral
more numerous secondary spiral threads and collabral ribs.
sionally included in the genus by Abbass (1973, p. 120, 121,
pl. 2, figs. 3–4, 8) differ from C. agriorivensis mainly as re-
gards sculpture. More specifically, C.? kirkaldyi Abbass, 1973,
from the Aptian, exhibits stronger collabral ribs and a su-
tural platform with secondary spiral threads while C.? nooryi Abbass, 1973, from the Cenomanian, lacks well-defined
smooth protoconch as well as a mid-sized cerithiform
ribs, spiral cords, a bicarinate periphery, a feebly convex base, a straight columella, a thickened parietal lip, an ovate
and slightly dilated aperture and a small angular callus
where the parietal and the outer lip meet. The protoconch
is too poorly illustrated so as to confirm its alleged morphology and the form of the aperture is largely presumed
collabral ribs and presents beaded spiral cords increasing in
since it is only partially known. Furthermore, Calzada and
Cossmann, 1906 (p. 242, pl. 13, figs. 6–7, 11), from the
is pyramidal or not. Although the holotype of C. caneroti
weak collabral and spiral sculpture. Cirsocerithium harborti
dicated in the original description of the species (Calzada,
number towards the last whorl instead. Cirsocerithium peroni Cenomanian of Algeria, differs from C. agriorivensis in its very
Corbacho (2015b) did not state whether the upper spire
presents a clearly pyramidal upper spire, the opposite is in-
(Wollemann, 1908, p. 179, pl. 13, figs. 5–7), from the Lower
1973). Other specimens attributed to C. caneroti do not ex-
orivensis in terms of its angular whorl profile and coarse
nor do the specimens of Cataldia mirambelensis (Vilanova,
Cretaceous of Germany, can be distinguished from C. agri-
hibit a pyramidal spire (Calzada, 1989b, pl. 1, figs. 8–9) and
pointy nodes, as observed in C. subspinosum. The sculpture
1863, pl. 8, fig. 17), another species from the Lower Creta-
Cenomanian of Germany, is similar to that of C. lallieranum,
Corbacho, 2015b). The importance of such feature relies
of Cirsocerithium quadricinctum Kollmann, 1979, from the
although the whorl profile in the former is more convex.
These two species are marked by thinner and more nu-
merous collabral ribs that are broken into conspicuous round nodes by the spiral cords.
Cirsocerithium agriorivensis is seemingly the earliest
record of the genus, as Cirsocerithium antiquum Glasunova,
1968, from the upper hauterivian of Russia, was recently
reinterpreted by Blagovetshenskiy (2015) as a representative of the epitoniidae. All the other records found in the literature are Barremian or younger in age.
ceous of Spain (Calzada, 1985; Viera, 1991; Calzada and
upon the fact that the material from the Neuquén Basin
lacks a pyramidal spire while other Cretaceous genera with
a similar teleoconch shape and sculpture, such as as Echi-
nobathra Cossmann, 1906, Pyrazopsis Akopyan, 1972 or Alamirifica, do bear one.
Cataldia is very similar to most batillariids, especially
Batillaria Benson in Cantor, 1842 and Zeacumantus Finlay,
1927. Cataldia is most similar to Batillaria in its conical to
cyrtoconoid shell outline. however, in this genus, spiral
elements of the sculpture are in general more developed than collabral elements, which are weaker and sometimes
421
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
inconspicuous. The periphery is also different, being round
thermore, the holotype illustrated by Calzada and Corbacho
The shell sculpture in Cataldia is more similar to that of
by Bandel (2006, pl. 10, fig. 11). Therefore, the attribution
and inconspicuous in Batillaria, and bicarinate in Cataldia.
(2015b) is similar to a juvenile of Campanile sp. illustrated
Zeacumantus since the collabral ribs are more conspicuous,
of this genus to any family known so far will not be pursued
to slightly opisthocline and straight to opisthocyrt, and
ture morphology.
wide and rounded as well as being subnodulous, orthocline
sometimes cut by spiral furrows. They are out of alignment between successive whorls. The numerous convex to flat
whorls, the straight columella, the bicarinate periphery and
the slightly convex base with or without fine spiral threads
herein in view of the lack of details on protonch and aperCataldia? binodosa sp. nov. Figure 5.13–17, 5.19
are also similar in both genera. Cataldia somewhat resem-
Diagnosis. Cataldia with flattish to convex whorls bearing
sculpture, especially in the non-pyramidal spire. The latter
cline to slightly opisthocline and straight to opisthocyrt,
or spine-like ribs whilst, in Cataldia, the last whorl is convex
two node-rows, the abapical one slightly more prominent
tural morphology in the type species of Pyrazus, P. ebeninus
Description. Shell mid-sized, conical to slightly cyrtoconical.
bles Pyrazus Montfort, 1810 as regards its spire outline and
can be distinguished by an angular last whorl with pointed
as the spire whorls and the nodes are rounded. The aper(Bruguière, 1792), is apparently much more complex than
broad, rounded and out-of-alignment collabral ribs, orthocrossed by shallow spiral depression that splits them into towards last whorl.
Protoconch and first teleoconch whorls missing. Teleoconch
that of Cataldia as it presents thick and flared outer and
with up to seven whorls preserved. Spire high, repre-
cerithial canal as well as a ventrolateral varix (Ozawa et al.,
canaliculated, bordered by suprasutural spiral cord. Whorls
inner lips with a wide siphonal notch and a conspicuous
2009). The apertural features so far known for Cataldia
seem much simpler, with a relatively thick and bent parietal
lip and, apparently, a simple and thin outer lip that would
explain its easy fragmentation. Most Cretaceous cerithiform
senting nearly 55% of the total shell height. Suture slightly
flat to convex. Sculpture of thick, wide collabral ribs on all
teleoconch whorls, 10–13 on penultimate whorl. Ribs or-
thocline to slightly opisthocline and straight to opisthocyrt
on upper spire, wider on middle part and tapering towards
records with a pyramidal spire had been assigned to Pyrazus
sutures. Ribs crossed by shallow spiral depression that
1984; Buitrón, 1986; Buitrón-Sánchez and López-Tinajero,
nodes slightly more prominent towards last whorl. Ribs
(e.g., Cossmann, 1906; Sayn, 1932; Olsson, 1944; Kase, 1995; Ayoub-hannaa and Fürsich, 2011; Kase et al., 2015) but Saul and Squires (2003) and Ozawa et al. (2009) ascribed
them all to different genera.
Cataldia resembles the spire of some species attributed
to Eustoma Piette, 1855 such as E. forneri Calzada, 1996. The main difference between Cataldia and Eustoma is that the
latter has very prominent pointed nodes in the last whorl
formed by fusion of the nodes in the spiral rows of the spire.
Calzada and Corbacho (2015b) ranked Cataldia within
the Batillariidae and, in that respect, Cataldia is consistent
splits them into two connected nodes. Abapical row of equidistant and out of alignment between succeeding
whorls. Interspaces concave and narrower than ribs. Number of ribs increasing slightly with ontogeny. Fine, subequal
spiral threads covering all whorls, not observed on base, 18–20 on penultimate whorl. Periphery bicarinate; carinae
smooth to slightly nodose. Base flat. Growth lines ortho-
cline and opisthocyrt. Columella straight. Aperture incomplete; parietal lip thick, not forming a callus, continuous with
columellar lip, thick and raised. Parietal lip meeting outer lip
in acute angle, slightly projecting adapically from peristome
with most of the diagnostic features listed for Batillariidae
but not forming true posterior channel and without parietal
resemblance as regards shell shape and sculpture, there
Occurrence. The material studied herein comes from six lo-
s.l. by Ozawa et al. (2009). however, other than the general
are no further grounds for this attribution. For instance, the
teleoconchs of representatives of other cerithiod families,
e.g. Potamididae, are also similar to that of Cataldia. Fur-
422
ridge. Basal and outer lips unknown.
calities in central Neuquén province, in late hauterivian– early Barremian strata near the top of the Agua de la Mula
Member of the Agrio Formation.
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
Material. A total of 76 specimens; one specimen from el
broad, rounded and out-of-alignment collabral ribs crossed
from Salado Norte, P. groeberi/S. riverorum zones, CPBA
adapical part of its outer lip being thickened and dilated as
Gasoducto, ?P. groeberi zone, CPBA 21296; 59 specimens 21798.1–6 and 21797.1–53; two specimens from Agua de
by spiral cords. The species is also characterized by the
well as extended towards the spire. even though Calzada
la Mula, P. groeberi zone, CPBA 21795 and CPBA 21796;
and Corbacho (2015b) tentatively included this species
21799.1–4 and CPBA 21297.1–3; one specimen from Cerro
served in other species of the genus. Ozawa et al. (2009) had
seven specimens from Bajada Vieja, S. riverorum zone, CPBA
within Cataldia, its outer lip morphology has not been ob-
Marucho, ?C. diamantensis/P. groeberi zones, CPBA 21298;
previously assigned this species to Pyrazopsis? and yet,
CPBA 21300. holotype CPBA 21796 and two paratypes:
would not fit into this genus either.
Derivation of name. From the Latin adjective nōdōsa ,
7e; see also Akopyan, 1974, p. 238, pl. 126, fig. 11), from
six specimens from Aguada Florencio, CPBA 21299.1–5 and CPBA 21797.1 and 21797.2.
meaning ‘having or covered with small knobs’, owing to the pair of nodes formed by the splitting of each collabral rib.
Dimensions. holotype (incomplete): h= 40.3 mm; hlw=
given its lack of a piramidal spire and beaded spiral cords, it Pyrazus quinquenodosus (Mertin, 1939, p. 210, pl. 6, fig.
the Lower Cretaceous of Germany and Armenia, also bears
a resemblance with C.? binodosa as regards teleoconch
shape and sculpture. Yet, its collabral ribs seem fainter than
21.45 mm; D= 18.8 mm; D/h= 0.46; hlw/h= 0.53; PA=
those observed in the Argentinian species.
D= 16.15 mm; D/h= 0.47.
to Potamides Brongniart, 1810, in its broadest sense, were
preserved in the available specimens, the teleoconch mor-
and Varela, 2012). Although the specimens are poorly pre-
17.6º. Paratype (CPBA 21797.1, incomplete): h= 31.4 mm;
Discussion. Although the aperture and early whorls are not
phology, especially the sculpture, suggests that the closest genus is Cataldia. The species resembles many batillariids
Two relatively abundant taxa provisionally attributed
described from the Upper Cretaceous of Patagonia (Griffin
served and present variability in the sculpture, slight similarities with the species herein described can be observed
and particularly Batillaria and Zeacumantus, as well as some
in terms of their convex whorls, their thick, opisthocyrt,
record restricted to the Cenozoic. Nevertheless, the assign-
bear. Additionally, these specimens differ from C.? binodosa
aperture in this genus is not fully known either and the
divide each rib into five rounded nodes as opposed to the
potamidids, but these are all recent genera with a fossil
ment of the species to Cataldia is only tentative since the
meaning of the apparently pyramidal spire of the type
species is unclear. The studied material is considerably
subnodulose collabral ribs and the bicarinate periphery they
mainly in that they display five conspicuous spiral cords that ribs split into two nodes that characterize C.? binodosa.
Uchauxia fraasi Alencáster, 1956 (non Blanckenhorn,
similar to the specimens of C. caneroti illustrated by Calzada
1890), from the Aptian of Mexico, greatly resembles C.?
specimen of figure 9, the observed collabral ribs are
species are similar in size, teleoconch outline, spiral and
(1989b, pl. 1, figs. 8–9). however, in the well-preserved
straighter than those of C.? binodosa and the adapical node row is slightly more prominent instead of the abapical one
binodosa and could also be congeneric with it as these
collabral sculpture, periphery and base. The Mexican species
can be however distinguished by its slightly larger number
as in the Argentinian species. Besides, the peripheral spiral
of collabral ribs per whorl and the more numerous fine spi-
in C.? binodosa, it can only be seen in the periphery of the
(Alencáster, 1956, pl. 6, fig. 6). Unfortunately, the aperture
differs from C.? binodosa in presenting seven to nine fine
it is clear that this species does not belong in Uchauxia
thread is visible thorughout most of the spire whorls whilst,
last whorl. The early hauterivian species C. mirambelensis spiral threads with finer intercalating lines.
ral threads that cover both its whorl face and its flat base
in U. fraasi sensu Alencáster (1956) is unknown. Anyways, Cossmann, 1906, which is more similar to Procerithium and
Pyrazus? scalariformis Nagao, 1934 (p. 257, pl. 35, figs.
Rhabdocolpus but for its conspicuous, short and twisted
figs. 18–19), from the upper Aptian–lower Albian of Japan,
Alencáster, 1956 (pl. 6, figs. 1, 2), although similar in spire
10, 10a,b, pl. 36, fig. 24; see also Kase, 1984, p. 137, pl. 20,
resembles the Argentinian species in its convex whorls with
anterior canal (Abbass, 1973). Conversely, Uchauxia? poblana
shape and sculpture, and judging by the prominent and
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
pointed nodes that form in its last whorl, seems to belong in Eustoma.
Superfamily PSeUDOMeLANOIDeA Fischer, 1885
height. Suture impressed. Spire whorls slightly convex. Last
whorl sub-cylindrical, with convex sides, diameter slightly
below mid-whorl. Whorls smooth but for very fine growth lines. Periphery gently rounded; base convex. Growth lines
Family PSeUDOMeLANIIDAe Fischer, 1885
orthocline and straight to slightly opisthocyrt near mid-
Remarks. Pseudomelaniidae is a problematic group based
obliquely oval in outline; outer lip and parietal lip joining in
turn, has a poorly known type. Often, species with mid-sized
lip simple, straight and vertical. Basal lip not fully preserved,
on a genus which is often a wastebasket taxon and that, in
to large, slender conical and anomphalous teleoconchs,
whorl, markedly prosocyrt at base. Aperture holostomate,
acute angle slightly dropping relative to suture line. Outer
with slightly thickened margin. Parietal lip flat, oblique and
smooth or with inconspicuous sculpture and with parasig-
slightly thickened. Columellar lip flat to concave, oblique to
and Campiche, 1862 (Gründel, 2001). Pseudomelaniids are
Occurrence. This species occurs in the upper part of the Agua
moidal growth lines are included in Pseudomelania Pictet
vertical, slightly thickened.
also distinguished by their inner lip callus without folds
de la Mula Member of the Agrio Formation; that is, in the C.
pseudomelaniids are, among the Mesozoic marine caeno-
both central Neuquén and southern Mendoza.
(Squires and Saul, 2004). That being said and thus far,
gastropods, the closest to the specimens from the Neuquén
Basin as regards teleoconch morphology.
Genus Paosia Böhm, 1895 Type species. Natica fadaltensis Böhm, 1895, Cenomanian, Italy; by original designation.
Remarks. Squires and Saul (2004) revised the genus, its
stratigraphic range and the geographic extension of its occurrence. According to these authors, Paosia may have
originated towards the end of the Jurassic in eastern europe and reached North America by the Aptian. So far, there were
no records of the genus in South America and most of the Austral world (Squires and Saul, 2004, fig. 2).
The diagnosis of Paosia indicates that it presents an
elongate-conical adult shell, a spire that can be high or low,
whorls that may bear no sculpture or rarely display fine spi-
ral grooves, a columella that usually has a callus, sinuous
growth lines and a basal lip curved anteriorly and forming a triangular projection (Squires and Saul, 2004). Paosia? sp.
diamantensis and P. groeberi zones. It was recorded from Material. Fifteen specimens; five specimens from Agrio del
Medio, CPBA 21794.1–5, P. groeberi zone; one specimen from Bajada Vieja, CPBA 21294, C. diamantensis zone; two
specimens from Agua de la Mula, CPBA 21295.1–2, P. groeberi zone; six specimens from Lomas Bayas, MCNAM-PI 24492, P. groeberi zone; one specimen from Lagunitas este,
MCNAM-PI 24493, P. groeberi zone.
Dimensions. CPBA 21294 (incomplete): h= 62.3 mm; hlw= 43.9 mm; hap= 30.2 mm; D= 29.3 mm; D/h= 0.47; hlw/h=
0.7; PA= 23.2°. CPBA 21794.1 (incomplete): h= 56.5 mm;
hlw= 36.25 mm; hap= 24.9 mm; D= 23.2 mm; D/h= 0.41; hlw/h= 0.64; PA= 29.5º.
Discussion. The Neuquén specimens resemble Paosia in their
teleoconch profile and the subcylindrical last whorl of the
most elongate species of this genus. They however differ in
that the aperture in most Paosia species is lower relative to
the height of the last whorl and their growth lines are more
sinuous than in the Argentinian species. Moreover, given that the basal lip of the available specimens of Paosia? sp. is
not fully preserved, it cannot be compared with the trian-
gular projection that is diagnostic of the genus.
Among all the known species of Paosia, Paosia? sp. re-
Figure 5.18, 5.20–22
sembles P. acuminata (Anderson, 1958) in its larger adult
Description. Shell medium-sized, slightly cyrtoconoidal, oval
its higher last whorl and its lack of sculpture. however, this
in outline. Protoconch and first teleoconch whorls not pre-
served. Teleoconch with up to five preserved whorls. Spire
moderately high, representing nearly 30% of the total shell
424
shell size, its taller spire outline, its narrower pleural angle,
species is based on a weathered holotype and the basal lip
is not preserved either.
Paosia? sp. also resembles some Recent or Cenozoic
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
fresh- or brackish-water genera of the cerithioid families
prosocline (Tunnell et al., 2010). Cossmann (1925) described
such families are not represented in the Cretaceous, the
however, Bandel (2006, pl. 2, figs. 8, 10) illustrated the pro-
Thiaridae, Pleuroceridae and Pachychilidae. Although most similarities among them and Paosia? sp. include the teleoconch shape, the apertural morphology and the sculpture.
the protoconch of vanikorids as smooth, slender and pointed.
toconch of a species of Vanikoro Quoy and Gaimard, 1832
by depicting three round whorls of which one was a smooth
This resemblance may be due to convergence since the
embryonic whorl while the remaining two were larval
sculpture and shell shape both inter- and intraspecifically.
The stratigraphic range of Vanikoridae depends on
shells in these families are actually often very variable in
whorls bearing beaded spiral cords.
The species herein discussed is very similar to the Recent
whether or not Vanikoropsis Meek, 1876 is included within
1877 (Cerithioidea, Thiaridae) in its elongate-oval cyrto-
as far back as the early Cretaceous.
freshwater genus Bayania hébert and Munier-Chalmas,
conoidal teleoconch outline, its tall and subcylindrical last
whorl, its broadly round periphery, its slightly convex spire whorls and its slightly opisthocyrt growth lines with a
prosocyrt sinus towards the base. Additionally, their aper-
(see discussion below). If so, its earliest records could date
Genus Vanikoropsis Meek, 1876
ture is similar; that is, oval with an acute adapical margin
Type species. Natica tuomeyana Meek and hayden, 1856 (p. 270), Maastrichtian, USA; by original designation.
and vertical outer lip and a straight to slightly concave
specimen lacking the apertural area and only presenting a
and a broadly rounded abapical one, as well as a simple
and slightly thickened columellar lip. The sutures in the type species of Bayania, B. lactea (Lamarck, 1806), and the
Remarks. The holotype of V. tuomeyana is an incomplete
part of the last whorl preserved and a considerably worn spire (Sohl, 1967; erickson, 1974). This specimen has a
Neuquén specimens are, when considering that they both
relatively thick shell, a large and globose last whorl, a wide
near the aperture, similar as well. The early spire whorls in
sculpture consists of spiral striae and collabral growth
present a slightly irregular trace in some parts and drop
B. lactea show collabral ribs and spiral cords and, although
conical and depressed spire and a narrow umbilicus. Its
rugae, i.e., thick, broad and irregular ribs. Whitfield (1876)
such whorls are not well preserved in the available speci-
described topotypes with a slightly taller spire and more
somewhat flared and forms a shallow sinus yet cannot be
within Vanikoropsis and therefore expanded the morpho-
mens, they seem to be smooth. The basal lip in B. lactea is
compared with that of the Neuquén specimens for their
conspicuous rugae. Sohl (1967) included thin-shelled species
logical range of the genus. According to erickson (1974),
basal lip is not fully preserved. Moreover, B. lactea is much
this genus resembles Spironema Meek, 1864 (Family Littori-
in shell height while other species are frequently below 10
of these genera based on more abundant and better material.
smaller than the specimens studied herein, reaching 30 mm
mm (Gougerot and Le Renard, 1983). Only a few Cretaceous
records of Bayania were reported from the Upper Cretaceous (Campanian–Maastrichtian) of southern India and
europe by Cossmann (1909b) and, furthermore, their attribution is dubious.
Superfamily VANIKOROIDeA Gray, 1840 Family ?VANIKORIDAe Gray, 1840
Remarks. According to Cossmann (1925), vanikorids are
characterized by an invariably open and deep umbilicus
whilst their teleoconch, normally small and globose, may be
highly variable in shape, and their peristome is round and
nidae), thus exposing the need for a so far unachieved revision Meek (1876) tentatively assigned Vanikoropsis to
Vanikoridae on the basis of its similarity with Vanikoro, type
genus of the family. These species are similar in their general
shell outline and their sculpture of wide spiral threads and
deep striae. Although several authors followed this classifi-
cation (e.g., Sohl, 1967; Thomson, 1971; erickson, 1974; Kase, 1984; Stilwell and henderson, 2002; Kollmann, 2005),
there were some exceptions (e.g., White, 1889; Pchelintsev,
1927; Beisel, 1983; Caze et al., 2011). Sohl (1967), in particular, emphasized the strong resemblance between Vanikoropsis and Vanikoro cancellata (Lamarck, 1822), the
former differing only in having a thicker shell, a lower spire
and a rounder aperture. Cossmann (1907, 1925) and Coss-
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
mann and Peyrot (1919) argued that the thick shell, broad
wide, moderately thick, with regularly expanding whorls.
the diagnosis of Vanikoridae. Accordingly, Cossmann (1925)
Teleoconch with four to five preserved whorls. Spire low,
aperture, inner-lip callus and closed umbilicus do not match
included the genus within the Ampullinidae. Sohl, 1967 (pl. 5, figs. 15, 16) re-illustrated the holotype of V. tuomeyana
displaying its actually narrow umbilicus.
Apical whorls apparently dome-shaped, paucispiral, smooth. representing approximately 25% or less of the total shell
height. Suture impressed to slightly canaliculated. Whorls
strongly convex. Last whorl large and swollen, slightly taller
The shell shape and sculpture in Vanikoropsis are evi-
than wide, maximum diameter slightly above mid-whorl.
should also consider protoconch morphology for a safer
callus. Aperture incomplete, holostomate, slightly proso-
dently convergent with other gastropod taxa. A revision
suprageneric placement. Sohl (1967) described the proto-
conch of V. nebrascensis (Meek and hayden, 1860), from the
Umbilicus closed, umbilical area partially covered by inner-lip
cline, peristome round to slightly ovate, subangular adapi-
cally, round abapically. Outer lip not preserved. Inner lip
Maastrichtian of Wyoming, USA (Sohl, 1967, p. B22, pl. 5,
flat to convex on parietal area, callus thick and narrow.
p. 170, pl. 15, figs. 1–3), as dome-shaped, paucispiral and
separated by spiral striae. Cords twice as broad as striae,
figs. 1, 5–10, 12, 14, 17; pl. 6, figs. 1–4, 11; erickson, 1974, smooth. Although poorly preserved, the protoconch of V.
demipleura Stilwell and henderson, 2002, from the Cenomanian of Australia, is also paucispiral.
Sohl’s (1967) criterion is herein adopted and the genus
Columella concave. Sculpture of wide, band-like spiral cords
nearly 20 on last whorl. Growth lines prosocline and straight.
Growth rugae on last and penultimate whorl; irregular,
parallel to growth lines, accentuated on adapical third of whorls. Periphery inconspicuous. Base convex, with spiral
is accordingly tentatively included within Vanikoridae.
cords and growth lines.
Jurassic/Cretaceous and Cretaceous/Paleogene boundaries
from the upper hauterivian, Agua de la Mula Member of
The stratigraphic record of Vanikoropsis crosses the
Occurrence. Vanikoropsis? leviplicata sp. nov. was recorded
(Rosenkrantz, 1970; Kollmann and Peel, 1983; Gerasimov,
Agrio Formation, in few localities in central and northern
early Cretaceous.
Material. A total of 23 specimens; 19 specimens from Agua
1992), and the genus was rather widespread during the Vanikoropsis? leviplicata sp. nov. Figure 6.1–7
Neuquén.
de la Mula, C. schlagintweiti zone, CPBA 21805.1–19; one
specimen from Agua de la Mula, C. diamantensis zone, CPBA
21302; two specimens from Loma la Torre, P. groeberi zone,
CPBA 21303.1–2; one specimen from Loma Rayoso (Puesto
Diagnosis. Vanikoropsis with moderately thick shell, low
Canale), H. neuquensis zone, CPBA 21304. holotype CPBA
and partially covered with callus, ca. 20 band-like spiral
Derivation of name. From the Latin adjectives levis, weak,
spire, moderately marked growth rugae, umbilicus closed cords and lacking secondary spiral striae.
Description. Shell small, turbiniform, slightly taller than
21304, two paratypes: CPBA 21302 and 21805.1.
subtle, and plicāta, folded, owing to the sculpture of weak growth rugae.
Figure 6. 1–7, Vanikoropsis? leviplicata sp. nov.; 1–2, holotype, CPBA 21304, abapertural and apical view; 3, paratype, CPBA 21805.1, apertural view; 4, paratype, CPBA 21302, abapertural view; 5, CPBA 21303.1, basal view; 6–7, CPBA 21303.2, SeM close-ups of sculpture and apical morphology. 8–9, Vanikoro? sp. in Stanton (1901), PRI 66880, abapertural and apertural view. 10–15, Confusiscala sp.; 10–11, 15, CPBA 21310.1, apertural, abapertural and basal view; 12–14, CPBA 21310, abapertural view and SeM close-ups of sculpture and aperture. 16–18, “Aporrhais? sp.” in Stanton (1901), PRI 66931, basal, apertural and abapertural view. 19–24, Tornatellaea neuquina sp. nov.; 19, holotype, CPBA 21311.2, abapertural view; 20, paratype, CPBA 21311.1, apertural view; 21–22, paratype, CPBA 21311.5, abapertural and abapertural-lateral view; 23–24, CPBA 21311.6, SeM close-ups of heterostrophic protoconch and sculpture. 25–26, Tornatellaea? sp.; 25, CPBA 21312.1, abapertural view; 26, CPBA 21312.4 (left) and CPBA 21312.5 (right), abapertural view. 27, Tornatellaea? andina, SIPB 9, apertural view. 28–29, Tornatellaea patagonica, lectotype, PRI 66932, abapertural and apertural view. Scale bars: 1–5, 17–18= 4 mm, 6–7, 13–14, 24= 500 µm, 8–9= 1 mm, 10–12, 15–16, 19–22, 25–29= 2 mm, 23= 250 µm. Arrows in 6, 13, 14, 24 point towards apex of shells.
426
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
Dimensions. holotype: h= 17.5 mm; hlw= 12.8 mm; hap= 7.7 mm; D= 16.7 mm; D/h= 0.95; hlw/h= 0.73; PA= 73.9º.
Pchelintsev (1927) described three species from the
lower Albian of Crimea. Vanikoropsis borissjaki Pchelintsev,
Paratype (CPBA 21302, incomplete): h= 16.5 mm; hlw=
1927 (p. 157, pl. 5, fig. 5) and V. multistriata Pchelintsev,
0.83; PA= 66.6º.
naticiform shell outline and their sculpture of thin spiral
13.8 mm; hap= 10.1 mm; D= 13.8 mm; D/h= 0.83; hlw/h=
Discussion. Although the aperture is not well-preserved in
the available specimens of V.? leviplicata, the remaining
teleoconch characters, especially the growth rugae, the pre-
dominant spiral sculpture, the thick shell and the narrow
1927 (p. 158, pl. 4, fig. 31) differ from V.? leviplicata in their cords, whilst V. communis Pchelintsev, 1927 (p. 158, pl. 4, fig. 21) presents a lower spire and a reticulate sculpture.
Vanikoropsis exerta Cossmann, 1925 (p. 41, pl. 5, figs.
30–31) and V. houdardi Cossmann, 1925 (p. 41, pl. 6, figs.
inner-lip callus, enable a tentative attribution to the genus.
15–17), from the Barremian and the Albian of France,
cata is most similar to V. nebrascensis, as described and
shell outline with a much lower spire. Sohl (1967) discussed
Amongst all the known species of Vanikoropsis, V.? levipli-
respectively, differ from V.? leviplicata in their naticiform
illustrated by Sohl (1967). The apical whorls in V.? leviplicata
the generic attribution of these and other species listed
demipleura as well.
from the Cretaceous of the San Juan Islands, USA.
are of the same aspect than those of the former and of V. The species herein described differs from the type
species in presenting a thinner shell, a higher spire and less
by Cossmann (1925), as well as of V. suciensis White, 1889, Three species were reported from the Lower Cretaceous
of Australia: V. jackii etheridge in Jack and etheridge, 1892
conspicuous growth rugae. Its differences with V. nebras-
(p. 486; see also etheridge, 1920, p. 11, pl. 2, figs. 33, 38),
umbilicus partially covered with callus and the fact that it
etheridge, 1907, p. 327, pl. 62, figs. 9–13) and V. decussata
ones towards the base of the last whorl. This latter feature
Leymerie, 1842). All three aforementioned species differ
censis mainly consist of its slightly lower spire, its closed lacks secondary spiral striae intercalated amongst the primary is, according to Sohl (1967), a highly variable attribute of V.
V.? stuarti etheridge, 1902 (p. 42, pl. 6, figs. 18–20; see also
etheridge, 1920 (p. 11, pl. 2, figs. 35; non Deshayes in from V. leviplicata as regards their naticiform shell outline,
nebrascensis.
their lower spire and the reticulate sculpture they present.
hibiting spiral band-like threads and striae only in the abapi-
from the Lower Cretaceous of Alexander I Island, Antarc-
platform (Stilwell and henderson, 2002, figs. 5.1–2, 5.4, 5.7).
casts with some shell remains. According to the author, the
Vanikoropsis demipleura differs from V.? leviplicata in ex-
cal half of the last whorl, and a narrow but marked sutural
Vanikoropsis decussata (Deshayes in Leymerie, 1842, p.
13, pl. 17, fig. 16), from the Albian of France and the Barremian of Japan (Kase and Maeda,1980, p. 315, pl. 35, figs.
1–2), is phaneromphalous and overall smaller than V.?
leviplicata. This species also differs from the Argentinian
one in its numerous conspicuous prosocline collabral ribs in
Thomson (1971, p. 51, fig. 2i) reported Vanikoropsis? sp.
tica. The material therein collected consists of two internal
recorded specimens are similar to V. nebrascensis, with a
turbinate shell, a moderately tall spire and a large and
swollen last whorl, all of which are features that are similar to those that characterize V.? leviplicata. however, the generic attribution of Vanikoropsis? sp. remains uncertain as
neither the growth rugae nor the umbilicus and aperture
the spire whorls and the adapical third of its last whorl.
are preserved in the shell remains.
fig. 5), from the Upper Jurassic/Lower Cretaceous of Russia
mian of Patagonia is represented by a single, small-sized,
Vanikoropsis neritoides (Trautschold, 1866, p. 12, pl. 2,
Stanton’s (1901) record of Vanikoro? sp. from the Barre-
(Gerasimov, 1992, p. 101, pl. 29, figs. 22–25), presents a
poorly preserved specimen (PRI 66880). The specimen’s
spire that is lower than that of V.? leviplicata. Additionally,
is similar to that of Vanikoro and Vanikoropsis. Unfortunately,
scultpture consisting of a reticulate pattern as well as a
Vanikoropsis valanginensis Beisel, 1983 (p. 71, fig. 26, pl. 4,
fig. 7), from the Valanginian of northern Siberia, differs from
the Argentinian species in having a lower spire and an open umbilicus.
428
overall shell shape with a low spire and a swollen last whorl
the shell material is mostly lost, with only a few remaining patches. Moreover, the visible sculpture does not fully match the typical sculpture of either genus since it presents
coarse and elongated nodes or ribs over the shoulder that
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
continue abapically into numerous, thin, prosocline collabral
each other and the differences amongst them are often in-
ribs (Fig. 6.8–9). Fine spiral threads are visible on the upper
correctly assessed due to the poor quality of the available
typical growth rugae of Vanikoropsis while the abapical
surface sculpture may aid in the distinction of species.
along the shoulder prove incompatible. Besides, the sutural
the lack of knowledge on the protoconch of the type species
a neritoid.
(2003, p. 456, fig. 4.1–2) illustrated the only protoconch
whorl side. The adapical nodes or ribs are different from the
sculpture is similar to that of Vanikoro although the nodes
platform is narrow and slightly concave, resembling that of
Superfamily ePITONIOIDeA Berry, 1910 Family ePITONIIDAe Berry, 1910
material. The spire apex, the aperture and a well-preserved The diagnoses of the genera are further complicated by
of many of them, including Confusiscala. Kiel and Bandel
thus far known for the genus, which presents a single inflated and smooth whorl.
Cossmann (1912) included a rather large shell size in the
extended diagnosis of Confusiscala. In fact, the type species
Remarks. The epitoniid adult turriculate teleoconch is easily
is about 60 mm high as reconstructed by Kollmann (2005)
lamelliform or variciform, straight or opisthocyrt collabral
including the one hereafter presented, are characterized
recognized due to the conspicuous presence of numerous
based on the lectotype. however, several other species,
ribs (Gardner, 1876; Cossmann, 1912; Guzhov, 2004). how-
by a considerable smaller shell height.
ticularly on early teleoconch whorls. The earliest record of
the observable differences are few and sometimes subtle.
cosmopolitan at least since the Aptian (Gardner, 1876;
bearing a reticulate sculpture and collabral ribs that may ex-
ever, such typical morphology is not always present, parthe family is from the Bathonian of France. The family is
Several genera are markedly similar to Confusiscala as
For instance, Amaea Adams and Adams, 1853 differs in
Cossmann, 1912; Cleeveley, 1980; Guzhov, 2002, 2004;
tend onto the basal disc (Squires and Saul, 2003). Turriscala
The general shell outline and the number, shape and
Clathroscala de Boury, 1889 differs in lacking a suprasutural
Squires and Saul, 2003).
alignment of collabral ribs as well as the spiral sculpture
are, according to Kilburn (1985), diagnostic features at the
genus level. Squires and Saul (2003) recommended that
the better known genera be more widely interpreted until
a better understanding of the epitoniid systematics is achieved.
Genus Confusiscala de Boury, 1909 Type species. Scalaria dupiniana d’Orbigny, 1843 (p. 54, pl. 154, figs. 10–13), Albian, France; by original designation.
Remarks. The most relevant diagnostic features of Confusis-
de Boury, 1889 presents a double peripheral cord, whilst spiral thread on the spire whorls (Cossmann, 1912). Claviscala de Boury, 1909 is characterized by a slenderer shell
outline than that of Confusiscala, as well as taller and flatter
whorls (Cossmann, 1912; Squires and Saul, 2003). Plicaceri-
thium Gerasimov, 1992 differs from Confusiscala in its
poorly-developed or even absent peripheral carina (Guzhov,
2002). Finally, the Cretaceous species of Opalia Adams and
Adams, 1853 strongly resembles Confusiscala (Squires and Saul, 2003). Opalia australis (Lamarck, 1822) exhibits whorls
that are less convex than those of C. dupiniana while its ribs are most prominent right below the adapical suture
instead of at or near mid-whorl, its peripheral cord is cre-
cala are its turriculate shell, its convex whorls, its deep su-
nated instead of smooth and its spiral sculpture is much
peripheral spiral cord, its base with thin spiral threads and
while C. dupiniana is early Cretaceous in age, it is possible that
tures, its variciform collabral ribs, its thin spiral threads, its
its round aperture (Cossmann, 1912). Still, these features may at least partially coincide with those of other epitoniid
genera. Cleeveley (1980) indicated that most Cretaceous epitoniid species bear strong collabral ribs, thin spiral
threads and a peripheral carina. In fact, a great part of the
species of Confusiscala described so far are very similar to
finer. Moreover, because of the fact that O. australis is Recent some Cretaceous species of Opalia belong in Confusiscala.
The stratigraphic range of Confusiscala is Valanginian–
Maastrichtian and possibly reaches the Oligocene (Cossmann, 1912; Corroy, 1925).
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AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
Confusiscala sp. Figure 6.10–15
Description. Shell small, turriculate. Apex of shell broken.
Teleoconch with up to four and a half whorls preserved. Spire high. Whorls well connected with each other, slightly
Besides, given their closer resemblance with C. dupiniana
than with O. australis, these specimens are therefore herein attributed to Confusiscala.
The species-level identity of the presented record is still
equivocal primarily due to the scarcity of the material and
lack of preservation of apertural and apical morphology. The
overlapped, almost imbricate, convex. Sutures canaliculated.
highly conservative morphology of epitoniids and the abun-
spire whorls. Periphery marked by spiral carina partially
material add to such uncertainties.
ribs per whorl. Ribs straight and slightly prosocline, narrow
cala species are known from the Lower Cretaceous. Some
ture up to peripheral carina, with maximum relief slightly
1862, p. 329, pl. 72, figs. 8, 9), C. menzeli (Wollemann, 1908,
Last whorl taller than wide, poorly differentiated from
concealed on spire whorls. Sculpture of 11 strong collabral
and raised above whorl surface, extended from adapical su-
dance of species described on the basis of poorly preserved
In addition to C. dupiniana, a small number of Confusis-
of them are Confusiscala cruciana (Pictet and Campiche,
above mid-whorl giving appearance of sutural platform.
p. 177, pl. 13, fig. 3), C. cf. novemvaricosa (Whitfield, 1891)
cline ridges along teleoconch; alignment more irregular on
(see Kase, 1984, p. 165, pl. 28, figs. 18a, b).
concave, wider than ribs. Nearly 30 very thin, subequal,
niana consist of the thicker and more numerous collabral
visible only with magnification. Base flat to concave, with
the adapical part of the whorls of C. dupiniana.
orthocline to slightly sinuous: opisthocyrt adapically and
land, presents less convex whorls than Confusiscala sp. and
opishocyrt at base. Growth lines occasionally grouped to-
collabral ribs of C. menzeli, from the Barremian of Germany,
Ribs irregularly aligned along spire axis and forming prosoearlier whorls and more regular on later whorls. Interspaces
closely spaced spiral threads on all whorls, crossing ribs, 20–25 fine concentric threads. Growth lines inconspicuous,
prosocyrt abapically; bent at periphery, prosocline and
gether forming low-relief rugae on base. Aperture not preserved. Parietal lip without callus. Columella straight to
(see Kase, 1984, p. 166, pl. 28, figs. 11–12) and C. cf. dupiniana
Some differences between Confusiscala sp. and C. dupi-
ribs and the alternation of finer and thicker spiral threads in Confusiscala cruciana, from the hauterivian of Switzer-
orthocline instead of prosocline ribs. On the other hand, the
are much more prosocline than those of Confusiscala sp.
Confusiscala cf. novemvaricosa, from the Aptian–Albian of
concave towards basal lip. Umbilicus closed.
Japan, has similar shell size and number of ribs as the Ar-
ties of central Neuquén, in upper hauterivian beds of the
opisthocline. The hauterivian–Barremian specimens from
Occurrence. Confusiscala sp. was recorded from two locali-
basal part of the Agua de la Mula Member of the Agrio For-
gentinian specimens yet its ribs are thicker and slightly
Japan that were attributed to C. cf. dupiniana exhibit more
mation.
collabral ribs per whorl than Confusiscala sp.
Bayo, S. riccardii zone, CPBA 21310.1–2; one specimen from
Lower Cretaceous of Alexander I Island, Antarctic Peninsula.
Dimensions. CPBA 21310.1 (incomplete): h= 17.9 mm; hlw=
single fragmentary and external cast specimen (Thomson,
Material. Three specimens total; two specimens from Cerro Salado Sur, C. schlagintweiti zone, CPBA 21800.
10.1 mm; D= 7.9 mm; D/h= 0.44; hlw/h= 0.56; PA= 16.3º.
Thomson (1971) described two epitoniids from the
The first one, Turriscala (Claviscala) sp., is represented by a
1971, p. 50, fig. 2g) and thus provides highly speculative
CPBA 21310.2 (incomplete): h= 12.2 mm; hlw= 7.1 mm;
diagnostic features. The author attributed such record to
16.9º. CPBA 21800 (incomplete): h= 7 mm; D= 3.5 mm;
sutures. Yet this feature is uncertain as it is only visible on
hap= 4.1 mm; D= 5.7 mm; D/h= 0.47; hlw/h= 0.58; PA= D/h= 0.5.
Claviscala on the basis of the spiral cords at each side of the
the largest preserved whorl and could represent a preser-
Discussion. The specimens described above match the
vation artifact. Moreover, the flattish aspect of the whorls
pleural angle, similar whorl profile, collabral ribs and spiral
Alexander I Island is also represented by a single, poorly pre-
general shell outline of C. dupiniana, have a comparable
threads, and share the presence of a conspicuous basal disc.
430
could be the result of crushing. The other epitoniid from
served specimen tentatively attributed to Proscala Coss-
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
mann, 1912 (Thomson, 1971, p. 51, fig. 2h). The preserved
suboval teleoconch, a relatively tall spire, spiral striae pitted
(Claviscala) sp. and the collabral ribs seem finer than those
adapically and widely rounded abapically oval aperture with
whorls seem taller and more convex than those of Turriscala
of Confusiscala sp. In both cases, a more detailed comparison with Confusiscala sp. is not possible.
Although Stanton (1901) reported Aporrhais? sp. from
or punctate by the intersection of growth lines, a narrow
a sinus or notch, a thickened outer lip crenulate or denticulate on the inner margin and callus on the columellar lip with
two oblique and acute folds (Cossmann, 1895; Kaim, 2004;
the Barremian of Patagonia, a direct examination of the
Gründel and Nützel, 2012). The genus is most similar to the
but rather an epitoniid. Furthermore, that which is observed
fact that the latter only presents one columellar fold (Kiel
specimen (PRI 66931) revealed that it is not an aporrhaid
in the specimen proves consistent with the diagnosis of
Confusiscala as it presents convex whorls, deep sutures,
type genus of the family, Acteon Montfort, 1810, but for the
and Bandel, 2001). Besides, in Acteon, the inner margin is
sharp and the basal lip is whole (Cossmann, 1895; Kollmann,
variciform collabral ribs, fine spiral threads and a basal disc
2005).
Confusiscala sp. in being larger and having more evenly
Deshayes and Milne edwards, 1838 (Family Ringiculidae),
collabral ribs per whorl. A different spiral pattern of sculp-
extends adapically and abapically upon the last whorl and
spiral threads on the whorl face intercalated with finer spi-
subdivided. Moreover, the outer lip is strongly thickened
with spiral threads (Fig. 6.16–18). It conversely differs from
convex whorls–maximum diameter at mid-whorl–and more
ture with two orders of threads, nearly 20 slightly thicker
ral lines and extending over the base, is also observed. Subclass heTeROBRANChIA Gray, 1840
Superfamily ACTeONOIDeA d’Orbigny, 1843 Family ACTeONIDAe d’Orbigny, 1843
Remarks. Recently, Gründel and Nützel (2012) revised the
systematics of the order Architectibranchia, including the
family Acteonidae, and provided revised diagnoses and a
key for the identification of genera. The most relevant teleoconch features of acteonids are, according to the emended
diagnosis provided by these authors, a moderately high
Although Tornatellaea is similar to Ringicula Deshayes in
the latter bears a prominent callus on the columellar lip that spire, and forms one to several folds frequently distally
an also extends axially (Kiel and Bandel, 2001; Kiel et al., 2002).
Other ringiculids, e.g., Avellana d’Orbigny, 1843, Oligoptycha
Meek, 1876 and Biplica Popenoe, 1957, differ from Tornatellaea in having a more globular shell profile with a lower spire (Cossmann, 1895; Squires and Saul, 2001).
The stratigraphic record of Tornatellaea is Aalenian–
Miocene and the genus was cosmopolitan (Cossmann, 1895; Kase, 1984; Gründel and Nützel, 2012). Tornatellaea neuquina sp. nov. Figure 6.19–24
spire, convex whorls, an absent or rarely present sutural
Diagnosis. Tornatellaea with moderately tall spire, swollen
aperture, callus on columella and one to three columellar
closely spaced spiral striae (ca. 30 on last whorl) with rhom-
ramp, a sculpture of punctate spiral striae, a long and narrow
last whorl, oblique columellar folds and numerous thin,
folds. According to Kiel and Bandel (2001), the outer lip is
bic pits formed by intersection with growth lines.
the number of parietal and columellar folds. Acteonids are
conical, moderately tall, representing 16–21% of total shell
not expanded and genera are distinguished on the basis of
cosmopolitan and their stratigraphic record is Bathonian– Recent (Gründel, 1975; Gründel and Nützel, 2012).
Description. Shell small, elongate oviform; spire broadly height. Apex very low, protoconch heterostrophic, mediaxial,
apparently smooth, 0.46 mm in diameter. Teleoconch with four to five rapidly-expanding whorls preserved, strongly
Genus Tornatellaea Conrad, 1860
convex, with maximum diameter slightly above mid-whorl.
Type species. Tornatellaea bella Conrad, 1860 (p. 294, pl. 47, fig. 23; see also harris, 1896, p. 188, pl. 7, fig. 3; harris, 1899, p. 6, pl. 1, fig. 6); eocene, USA; by original designation.
whorl swollen, taller than wide. Periphery convex, incon-
Remarks. The species of this genus are characterized by a
Sutures canaliculated. Very narrow sutural platform. Last
spicuous. Base regularly convex. Growth lines well-marked,
with slightly opisthocyrt sinus near adapical suture, ortho-
431
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
cline and prosocyrt on whorls lateral face and markedly
17), which can be readily distinguished from T. neuquina by
near aperture. Aperture nearly radial, wide, with oval-sub-
Other Tornatellaea species from the Lower Cretaceous
retrocurrent towards base. Growth lines more deeply set
its sub-spherical shell and its very low and broad spire.
oval section, adapical end acute and abapical end well
described in the literature are, for example, two species
area. Columella straight, with two subequal oblique folds.
and T. matura (Schröder, 1995) (see Kaim, 2004, p. 154–
Sculpture of numerous thin spiral striae finer than inter-
regards its taller spire, its less inflated last whorl and the
rounded. Inner lip with thin but conspicuous callus on parietal
Basal lip with shallow sinus. Outer lip badly preserved.
spaces, ca. 30 on last whorl. Striae with rhombic pits formed
by intersection with growth lines.
Occurrence. All the available specimens of Tornatellaea
neuquina sp. nov. were collected from a single bed at the
base of the Pilmatué Member of the Agrio Formation (upper
Valanginian), in the locality of Pichaihue, central Neuquén.
Material. A total of 15 specimens from Pichaihue, CPBA
from the Valanginian of Poland: T. gazdzickii (Schröder, 1995)
155, figs. 132–133). The former differs from T. neuquina as fact that its spiral sculpture is formed by two groups, one
adapical and one abapical, of a few pitted spiral striae.
Similarly, T. matura differs from T. neuquina in having a much
taller spire.
Kase (1984, p. 168, pl. 28, figs. 1–2) recorded Tornate-
llaea cf. ebrayi (de Loriol, 1882) from the Aptian–Albian of
Japan. The shells studied by Kase (1984) are slightly smaller
21311.1–15, P. angulatiformis zone. holotype CPBA 21311.2
than the shells of T. neuquina and present a proportionally
Derivation of name. Named after the Neuquén province.
instead of oblique columellar folds and thicker and less
and three paratypes: CPBA 21311.1, 21311.5 and 21311.6.
Dimensions. holotype: h= 7.5 mm; hlw= 5.9 mm; D= 5.2 mm; D/h= 0.69; hlw/h= 0.79; PA= 60.7º. Paratype (CPBA
taller and more conical spire as well as nearly horizontal closely spaced spire striae (ca. 13 on the last whorl).
Tornatellaea casanovai Calzada, 1989b (p. 16, pl. 2, fig.
21311.5, incomplete): h= 7.6 mm; hlw= 6.5 mm; D= 5 mm;
8a–d), from the lower Aptian of Spain, is represented by
Discussion. This species is herein attributed to Tornatellaea
neuquina in having more spiral striae (ca. 35 in the last
hlw/h= 0.85; PA= 69.7º.
on the basis of its oviform shell outline, its moderately tall
spire, its pitted spiral striae and the two simple columellar
very well-preserved specimens. This species differs from T. whorl) and a taller spire.
Tornatellaea albensis (d’Orbigny, 1843), from the hau-
folds it bears. Its spire, however, is lower than that of other
terivian of France, was revised by Kollmann (2005, p. 44,
species from the Neuquén Basin (see below).
more resemblance to the specimen of figures 6a,b, in any
species of the genus such as Tornatellaea? sp., another The genus has already been reported from the Lower
Cretaceous of southern South America. Tornatellaea patagonica Stanton, 1901 (p. 34, pl. 6, figs. 18–19) (Fig. 6.28– 29) was described on the basis of a few specimens from the
pl. 6, figs. 5–6). even though Tornatellaea neuquina bears
case, T. albensis displays a taller spire. According to Kollmann (2005), the outer lip in T. albensis is thin and lacks inner crenulations or denticles.
Tornatellaea mendozana Gründel, Parent, Cocca and
Barremian of Patagonia. Direct examination of the original
Cocca, 2007 was reported from the middle Tithonian of
specimens) confirmed that this species is very similar to T.
the basis of two oblique columellar folds. Later, Gründel and
material (PRI 66932, two specimens, and PRI 66933, three neuquina but for its larger shell (h= 12.7 mm) and relatively
lower spire (hlw/h= 0.92). Besides, T. patagonica presents
fewer and wider spiral striae (ca. 20 on the last whorl), its
whorls taper into each other at the adapical sutures instead
of buttressing, and its penultimate whorl is wider in relation
to the maximum diameter of the shell. Unfortunately, the
southern Neuquén province and attributed to the genus on
Nützel (2012) expressed uncertainty regarding the generic
attribution of this species because the spiral striae are not pitted. Then again, it is worth mentioning that T. mendozana
is a junior synonym of Acteon andinus haupt, 1907 (p. 206, pl. 10, figs. 4a,b), originally recorded from “Neocomian”–
Aptian beds (likely Tithonian, Vaca Muerta or Picún Leufú
outer lip is not preserved in the available specimens of T.
formations) at a southern Neuquén locality very close to the
gonia is Cinulia australis Stanton, 1901 (p. 34, pl. 6, figs. 16–
Information). The type series of A. andinus comprises four
patagonica. Another acteonoid from the Barremian of Pata-
432
type locality of T. mendozana (see Supplementary Online
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA
moderately well-preserved specimens under the collection
nuses, one below adapical suture and another one towards
number SIPB 9. The specimen illustrated by haupt (1907) is
the base. Growth lines more deeply set near the aperture.
dina differs from T. neuquina in its larger and more inflated
lower Valanginian outcrops of the Mulichinco Formation at
herein designated as lectotype (Fig. 6. 27). Tornatellaea? an-
Occurrence. This species has been so far recorded only from
last whorl with a narrow but conspicuous sutural platform
the Puerta Curaco locality, northern Neuquén.
half of the last whorl consisting of non-pitted spiral striae.
L. riveroi zone.
and its spiral sculpture that is only visible in the abapical even though other acteonoids have been reported from
the Upper Jurassic–Lower Cretaceous of the Neuquén Basin
Material. Five specimens, Puerta Curaco, CPBA 21312.1–5, Dimensions. CPBA 21312.1: h= 6.9 mm; hlw= 5 mm; D= 4.4 mm; D/h= 0.64; hlw/h= 0.72. PA= 60.1º. CPBA 21312.2:
(see Supplementary Online Information), their generic attri-
h= 6.8 mm; hlw= 5.1 mm; D= 4.3 mm; D/h= 0.63; hlw/h=
of discussion. For instance, Tornatellaea sp. was reported
Discussion. Unfortunately, it was not possible to observe
bution and relationship with other records remain a matter by Behrendsen (1891) from the Tithonian of Mendoza
province, but the autor failed to provide an illustration of
such record and the corresponding original material is not
0.75.
the adapertural side of the shells of Tornatellaea? sp., and
further details regarding the outer and inner lips, including
the number, placement and shape of the columellar folds,
found in the collection of the Geowissenschaftliches
remain thus unknown. Therefore, although the remaining
Behrendsen’s (1891, 1892) material. The record remains
material cannot be attributed to Tornatellaea with certainty.
zentrum der Universität Göttingen that houses the rest of therefore unrevised. The same occurs with Cinulia sp.,
teleoconch features are consistent with the genus, the In fact, the available specimens of Tornatellaea? sp. bear a
recorded by Behrendsen (1892) from the “Neocomian” of
close resemblance to T. neuquina in spite of displaying a
age of this record is similar to that of T. neuquina and their
differences and the stratigraphic gap between Tornatellaea?
central Neuquén (likely Valanginian, Agrio Formation). The
respective fossil localities are nearby (ca. 35 km apart). Damborenea et al. (1979) recorded Cinulia cf. andina from the
Berriasian of southern Mendoza but the specimen (MLP
taller spire and less spiral striae on the last whorl. These
sp. and T. neuquina constitute the reasons why these records are being for the moment kept as separate taxa.
14890) is so poorly preserved that no identification or com-
FINAL REMARKS
in order to clarify the identity of the abovementioned
marine benthic and nektonic invertebrates (e.g., bivalves,
parison is possible. In every case, new materials are required records.
Tornatellaea? sp. Figure 6.25–26
Over the last decades, research on different groups of
nautiloids) has revealed that exchange of taxa occurred between the Neuquén Basin and the Tethys and even remoter regions such as northern europe, North America, Australia
and eastern Africa (Lazo, 2007; Aguirre-Urreta et al., 2008; Fernández and Pazos, 2013). Such expansion is possibly the
Description. Shell small, elongate oviform. Spire conical,
result of the existence of open water connections between
Protoconch not preserved. Teleoconch with four to five
Tethyan taxa towards the west by means of the equatorial
moderately tall, representing 23–25% of total shell height.
strongly convex whorls preserved, maximum diameter slightly above mid-whorl; sutures canaliculated. Last whorl
the Neuquén Basin and the Pacific Ocean, and the spread of current (Aguirre-Urreta et al., 2008).
Regarding gastropods, recent progress on the knowledge
taller than wide, inflated, with a very narrow sutural plat-
of the composition of the early Cretaceous fauna of the
observed. Basal lip forming a wide, shallow sinus. Spiral
tic affinities and paleobiogeographic implications of the
form. Periphery and base regularly convex. Aperture not
sculpture of thin pitted striae, 20–25 on the last whorl, with
band-like interspaces. Growth lines orthocline and prosocyrt on the whorl side and with two shallow opisthocyrt si-
basin allows us to interpret, albeit preliminarily, the faunisrecorded gastropod association. So far, 20 species belonging to 19 genera and 14 families were described and/or
revised from the Mulichinco and Agrio formations (Cataldo
433
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440
and Lazo, 2012, 2016; Cataldo, 2013, 2014; this work) and
with the more provincial Late Cretaceous–Paleocene Aus-
to other units from the Lower Cretaceous of the Neuquén
Additionally, the findings hereby reported expand the
as many are still under study. The analysis was expanded
tral gastropod fauna.
Basin such as the Vaca Muerta, Chachao and huitrín forma-
geographic distribution of most of the recorded genera to
information can be treated at different taxonomic levels, i.e.,
Proconulus, Calliotropis (Riselloidea) and Exelissa, evidently
tions (see Supplementary Online Information). The available species, genus and family, thus offering different degrees
of resolution of faunistic influences.
When establishing comparisons with Berriasian–Barre-
southern South America. Some of these genera, namely
persisted in the Andean region since Jurassic times.
These preliminary observations call for more detailed
analyses to test paleobiogeographic hypotheses for south-
mian faunas from other regions, it can be seen that most of
ern South America. however, further advances in the revi-
ceous had a widespread distribution during the early Creta-
should be achieved prior to such attempt.
the genera represented in the Neuquén Basin Lower Creta-
ceous (e.g., Pleurotomaria Defrance, 1826, Harpagodes Gill, 1870, Paraglauconia Steinmann, 1929) with so far only one
genus with a distribution restricted to the western margin
of South America (Protohemichenopus Camacho, 1953). In
turn, other less widespread genera display a predominantly
Tethyan influence, especially from europe and Asia. Follow-
ing Sohl (1987), among the taxa recorded in the Neuquén
Basin, those that are more representative of the early Cretaceous Tethyan influence are the nerineoids, cassiopids and proconulids, as well as the genus Harpagodes.
At the species level, most taxa are endemic to west-
central Argentina but for a few species (Pleurotomaria gerthi Weaver, 1931, Harpagodes jaworskii Weaver, 1931, “Natica”
cf. bulimoides Deshayes in Leymerie, 1842) with possible
records in coeval sediments of neighboring Andean basins
such as, for instance, the Central Chile Basin (e.g., Corvalán, 1956; Corvalán and Pérez, 1958). A few other records bear similarities with coeval species from Peru, Venezuela and
the Argentinian Austral Basin (see Fritzsche, 1924; Jaworski, 1938; von der Osten, 1957). Only one species presents
records in a very distant region (Metacerithium turriculatum
Forbes, 1845; Cataldo and Lazo, 2016).
Finally, the gastropod fauna studied herein shares at
least family-level gastropod taxa with other basins along
the western margin of South America, including the Argen-
tinian Austral Basin, and the Antarctic Peninsula.
hence, these preliminary results suggest a strong Tethyan
influence during the Berriasian–Barremian with additional faunistic exchange through open seaways between the
Neuquén Basin and both neighboring and remote regions. The mixed nature of this gastropod association contrasts
434
sion of South American early Cretaceous gastropod records
ACKNOWLEDGEMENTS
The author wishes to acknowledge the contribution of the following people at different stages of this research: D.G. Lazo, M.B. AguirreUrreta, L. Luci, D.e. Fernández, R.M. Garberoglio, and L.C. Gaetano (IDeAN); M.A. Tunik (Universidad de Río Negro); A. Riccardi (MLP); C. Gee (SIPB); M. Reich (formerly at Geozentrum Universität Göttingen); G. Dietl and L. Skibinski (PRI); S. Calzada (Museo Geológico del Seminario de Barcelona). The SeM operators at LIMF-UNLP are thanked for their assistance. This manuscript benefited from the reviews of S. Nielsen (Universidad Austral de Chile) and A. Kaim (Instytut Paleobiologii PAN). Funding for this work was provided by Agencia Nacional de Promoción Científica y Tecnológica through project PICT 0464/2010 (M.B. Aguirre-Urreta), and by Universidad de Buenos Aires through project UBACyT GC 20020100100974 (M.B. Aguirre-Urreta). This is the contribution R-197 of the IDeAN.
REFERENCES
Abbass, h.L. 1973. Some British Cretaceous gastropods belonging to the families Procerithiidae, Cerithiidae and Cerithiopsidae (Cerithiacea). Bulletin of the British Museum (Natural History) Geology Series 23: 107–175. Adams, h., and Adams, A. 1853. The genera of Recent Mollusca, arranged according to their organization, Volume 1. J. Van Voorst, London, 256 p. Aguirre-Urreta, M.B., and Rawson, P.F. 2012. Lower Cretaceous ammonites from the Neuquén Basin, Argentina: a new heteromorph fauna from the uppermost Agrio Formation. Cretaceous Research 35: 208–216. Aguirre-Urreta, M.B., Rawson, P.F., Concheyro, G.A., Bown, P.R., and Ottone, e.g. 2005. Lower Cretaceous (Berriasian–Aptian) Biostratigraphy of the Neuquén Basin. In: G.D. Veiga, L.A. Spalletti, J.A. howell, and e. Schwarz (eds.), The Neuquén Basin, Argentina: A Case Study in Sequence Stratigraphy and Basin Dynamics. Geological Society of London Special Publications 252: 57–81. Aguirre-Urreta, M.B., Casadío, S., Cichowolski, M., Lazo, D.G., and Rodríguez, D.L. 2008. Afinidades paleobiogeográficas de los invertebrados cretácicos de la Cuenca Neuquina, Argentina. Ameghiniana 45: 591–612. Aguirre-Urreta, M.B., Lazo, D.G., Griffin, M., Vennari, V., Parras, A.M., Cataldo, C.S., Garberoglio, R., and Luci, L. 2011. Megainvertebrados del Cretácico y su importancia bioestratigráfica. In: h.A. Leanza, C. Arregui, O. Carbone, J.C. Danieli, and J.M. Vallés (eds.), Geología y recursos naturales de la provincia del Neuquén. Asociación Geológica Argentina, Buenos Aires, p. 465–488.
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA Akopyan, V.T. 1972. [On a new cerithioidean family]. Trudy Akademii Nauk Armyanskoy SSR, Nauky o Zemie 25: 3–14. [in Russian and Armenian]. Akopyan, V.T. 1974. [Atlas of fossil fauna of the Armenian SSR]. Izdatel’stvo Armyanskoy SSR, Yerevan, 424 p. [in Russian and Armenian]. Alarcón, B., and Vergara, M. 1964. Nuevos antecedentes sobre la geología de la quebrada el Way. Universidad de Chile, Instituto de Geología, Publicaciones 26: 101–128. Alencáster, G. 1956. Pelecípodos y gasterópodos del Cretácico Inferior de la región de San Juan Raya-zapotitlán, estado de Puebla. Paleontología Mexicana 2: 1–47. Anderson, F.M. 1958. Upper Cretaceous of the Pacific coast. Geological Society of America Memoir 71: 1–378. Archiac, A.d’. 1843. Description géologique du département de l’Aisne. Mémoires de la Société Géologique Française 5: 129–418. Archiac, A.d’. 1859. Note sur le genre Otostoma. Bulletin de la Société Géologique de France 16: 871–879. Ayoub-hannaa, W.S., and Fürsich, F.T. 2011. Revision of Cenomanian-Turonian (Upper Cretaceous) gastropods from egypt. Zitteliana 51: 115–152. Bandel, K. 2006. Families of the Cerithioidea and related superfamilies (Palaeo-Caenogastropoda; Mollusca) from the Triassic to the Recent characterized by protoconch morphology, including the description of new taxa. Paläontologie, Stratigraphie, Fazies. Freiberger Forschungshefte C 511: 59–138. Bandel, K. 2010. Relationships of the Triassic eucycloidea Koken, 1897 (Mollusca, Gastropoda) to modern genera such as Pagodatrochus, Calliotropis and Euchelus, based on morphology of the early shell. Bulletin of Geosciences 85: 435–486. Bandel, K., and Kiel, S. 2003. Relationships of Cretaceous Neritimorpha (Gastropoda, Mollusca) with the description of seven new species. Bulletin of the Czech Geological Survey 78: 53–65. Bataller, J.R. 1959. Primer suplemento a la “Sinopsis de las especies nuevas del Cretáceo de españa”. Boletín del Instituto Geológico y Minero de España 70: 1–77. Behrendsen, h.O. 1891. zur Geologie des Ostabhanges der argentinischen Cordillere. I Theil. Zeitschrift der Deutschen geologischen Gesellschaft 43: 369–420. Behrendsen, O. 1892. zur Geologie des Ostabhanges der argentinischen Cordillere. II Theil. Zeitschrift der Deutschen geologischen Gesellschaft 44: 1–42. Beisel, A.L. 1983. [Late Jurassic and early Cretaceous Gastropods of the North of Middle Siberia (systematic composition, paleoecology, stratigraphic and paleogeographic significance)]. Trudy Institut Geologii i Geofiziki Akademiya Nauk SSSR 484: 1–94. [in Russian]. Bengtson, P. 1988. Open nomenclature. Palaeontology 31: 223– 227. Berry, S.S. 1910. Review of: Report on a collection of shells from Peru, with a summary of the littoral marine Mollusca of the Peruvian zoological province, by W.h. Dall. The Nautilus 23: 130–132. Blagovetshenskiy, I.V. 2015. Gastropods of the family epitoniidae from the Lower Cretaceous of the Volga Region near Ulyanovsk. Paleontological Journal 49: 361–368. Blanckenhorn, M. 1890. Beiträge zur Geologie Syriens: Die Entwickelung des Kreidesystems in Mittel- und Nord-Syrien mit besonderer Berücksichtigung der paläontologischen Verhältnisse nebst einem Anhang über den jurassischen Glandarienkalk. L. Döll, Kassel, 130 p. Böhm, G. 1895. Die Gastropoden des Marmolatakalkes. Palaeontographica 42: 211–308.
Boury, e. de. 1889. Révision des Scalidae Miocènes et Pliocènes de l’Italie. Bulletino della Società Malacologica Italiana 14: 161–326. Boury, e. de. 1909. Catalogue des sous-genres de Scalidae. Journal de Conchyliologie 57: 255–258. Brongniart, A. 1810. Sur des terrains qui paraissent avoir été formés sous l’eau douce. Annales du Museum d’Histoire Naturelle 15: 357–405. Bruguière, J.G. 1792. Encyclopédie Méthodique, Histoire Naturelle des Vers, Tome Premier. Panckoucke, Paris, 757 p. Buitrón, B.e. 1986. Gasterópodos del Cretácico (Aptiano TardíoAlbiano Temprano) del Cerro de Tuxpan, Jalisco. Boletín de la Sociedad Geológica Mexicana 47: 17–31. Buitrón-Sánchez, B.e., and López-Tinajero, Y. 1995. Mollusk gastropods in a Lower Cretaceous rudist-bearing formation of Jalisco, west-central Mexico. Revista Mexicana de Ciencias Geológicas 12: 157–168. Calzada, S. 1973. Confusiscala caneroti n. sp. del Aptiense inferior de Chert (Castellón). Acta Geologica Hispanica 8: 131–133. Calzada, S. 1985. Sobre Confusiscala mirambelensis (Vilanova, 1868). Mediterránea, Serie de Estudios Geológicos 4: 79–84. Calzada, S. 1989a. Algunos tróquidos neocretácicos del Prepirineo catalán. Batalleria 3: 23–36. Calzada, S. 1989b. Gasterópodos del Aptiense inferior de Forcall (Castellón, españa). Batalleria 2: 322. Calzada, S. 1996. Gastrópodos hauterivienses de Caltellfort (Castellón). Batalleria 6: 67–76. Calzada, S. 2000. Sobre los géneros Otostoma, Lyosoma, Corsania y las especies Otostoma rugosum y Otostoma corsanum (Gastropoda mesozoica). Batalleria 9: 7–14; 43–44. Calzada, S., and Corbacho, J. 2015a. Sobre Lissochilus goñii, gasterópodo cretácico. Scripta Musei Geologici Seminarii Barcinonensis Series Palaeontologica 17: 3–6. Calzada, S., and Corbacho, J. 2015b. Sobre la atribución genérica de la especie Confusiscala caneroti. Scripta Musei Geologici Seminarii Barcinonensis Series Palaeontologica 18: 3–9. Calzada, S., and Forner, e. 2006. Un nuevo tróquido del Cretácico español. In: V.M. Cardona eixarch (ed.), Miscellània en homenatge a José Eixarch Frasno. Colección Aldebarán 62. Ajuntament de Forcall, Forcall, and editorial Antinea, Vinaròs, p. 49–51. Camacho, h.h. 1953. Algunas consideraciones sobre los Aporrhaidae fósiles Argentinos. Revista de la Asociación Geológica Argentina 8: 183–194. Cantor, T. 1842. General features of Chusan, with remarks on the flora and fauna of that Island. Annals and Magazine of Natural History 9: 265–278; 360–370; 481–493. Cataldo, C.S. 2013. A new early Cretaceous nerineoid gastropod from Argentina and its palaeobiogeographic and palaeoecological implications. Cretaceous Research 40: 51–60. Cataldo, C.S. 2014. The gastropod family Aporrhaidae in the Lower Cretaceous of the Neuquén Basin, west-central Argentina. Journal of Paleontology 88: 1222–1239. Cataldo, C.S., and Lazo, D.G. 2012. Redescription of Pleurotomaria gerthi Weaver, 1931 (Gastropoda, Vetigastropoda), from the early Cretaceous of Argentina: new data on its age, associated palaeoenvironments and palaeobiogeographic affinities. Ameghiniana 49: 75–95. Cataldo, C.S., and Lazo, D.G. 2016. Taxonomy and paleoecology of a new gastropod fauna from dysoxic outer ramp facies of the Lower Cretaceous Agrio Formation, Neuquén Basin, west-central Argentina. Cretaceous Research 57: 165–189. Caze, B., Merle, D., Le Meur, M., Pacaud, J.-M., Ledon, D., and Saint Martin, J.-P. 2011. Taxonomic implications of the residual
435
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440 colour patterns of ampullinid gastropods and their contribution to the discrimination from naticids. Acta Palaeontologica Polonica 56: 329–347. Chartron, C., and Cossmann, M. 1902. Note Sur l’Infralias de la Vendée et spécialement sur un gisement situé dans la Commune du Simon-la-Vineuse. Bulletin de la Société Géologique de France Quatrième Série 2: 163–203. Choffat, P. 1901. Recueil d’études paléontologiques sur la faune crétacique du Portugal. Espèces nouvelles ou peu connues. Quatrième série. Imprimerie de l’Académie Royale des Sciences, Lisbon, p. 107–171. Cleeveley, R.J. 1980. Two new British Cretaceous epitoniidae (Gastropoda): evidence for evolution of shell morphology. Bulletin of the British Museum (Natural History) 34: 235–250. Collignon, M. 1972. Les gastéropodes et les serpules crétacés du bassin côtier de Tarfaya (Maroc meridional). Notes du Service Géologique du Maroc 228: 9–29. Conrad, T.A. 1860. Descriptions of new species of Cretaceous and eocene fossils of Mississippi and Alabama. Journal of the Philadelphia Academy of Natural Sciences Series 2 4: 275–298. Coquand, h. 1865. Monographie de l’étage Aptien de l’Espagne. Arnaud et Cie., Marseille, 221 p. Corroy, G. 1925. Le Néocomien de la Bordure Orientale du Bassin de Paris. Bulletin de la Société des Sciences de Nancy 4: 171–506. Corvalán, J. 1956. Über marine Sedimente des Tithon und Neokom der Gegend von Santiago. Geologische Rundschau 45: 919–926. Corvalán, J., and Pérez, D.e. 1958. Fósiles Guías chilenos: Titoniano– Neocomiano. Manual Nº1. Instituto de Investigaciones Geológicas, Santiago de Chile, 48 p. Cossmann, M. 1885. Contribution à l’étude de la faune de l’étage Bathonien en France (Gastropoda). Mémoires de la Société Géologique de France 3: 1–374. Cossmann, M. 1895. Essais de Paléonconchologie Comparée, Première livraison. Printed by the author and Comptoir Géologique, Paris, 159 p. Cossmann, M. 1906. Essais de Paléonconchologie Comparée, Septième livraison. Printed by the author and F.R. de Rudeval, Paris, 261 p. Cossmann, M. 1907. Description des gastropodes et pélécypodes. In: e. Pellat, and M. Cossmann (eds.), Le Barrémien supérieur à facies urgonien de Brouzet-les-Alais (Gard). Mémoires de la Société Géologique de France 37: 6–42. Cossmann, M. 1908. Note sur un gisement d’âge Charmouthien à Saint-Cyr-en-Talmondois (Vendée). Bulletin de la Société Géologique de Normandie 27: 45–65. Cossmann, M. 1909a. Paléoconchologie: Note sur un gisement d’âge Charmouthien à Saint-Cyr-en-Talmondois (Vendée). Revue Critique de Paléozoologie 13: 27–28. Cossmann, M. 1909b. Essais de Paléonconchologie Comparée, Huitième livraison. Printed by the author and F.R. de Rudeval, Paris, 248 p. Cossmann, M. 1912. Essais de Paléonconchologie Comparée, Neuvième livraison. Printed by the author and J. Lamarre et Cie., Paris, 215 p. Cossmann, M. 1913. Contributions à la paléontologie Française des terrains jurassiques, III: Cerithiacea et Loxonematacea. Mémoires de la Société Géologique de France, Série Paléontologie 46: 1–263. Cossmann, M. 1916. Étude complementaire sur le Charmoutien de la Vendée. Mémoires de la Société Linnéenne de Normandie, Section Géologique 33: 113–159. Cossmann, M. 1918. Essais de Paléonconchologie Comparée, Onzième livraison. Printed by the author, Paris, 388 p. Cossmann, M. 1925. Essais de Paléonconchologie Comparée, Treizième livraison. Les Presses Universitaires de France, Paris, 345 p.
436
Cossmann, M., and Peyrot, A. 1919. Conchologie néogénique de l’Aquitaine. Actes de la Société Linnéenne de Bordeaux 70: 181–356. Cox, L.R. 1960. Gastropoda. General characteristics of Gastropoda. In: R.C. Moore (ed.), Treatise on Invertebrate Paleontology, Part I, Mollusca 1. Geological Society of America, Boulder, and University of Kansas Press, Lawrence, p. 84–169. Cuvier, G. 1795. Second Mémoire sur l’organisation et les rapports des animaux à sang blanc, dans lequel on traite de la structure des Mollusques et de leur division en ordre. Magazin Encyclopédique, ou Journal des Sciences, des Lettres et des Arts 8: 433–449. Damborenea, S.e., and Ferrari, S.M. 2009. el género Lithotrochus Conrad (Gastropoda, Vetigastropoda) en el Jurásico Temprano de Argentina. Ameghiniana 45: 197–209. Damborenea, S.e., Manceñido, M.O., and Riccardi, A.C. 1979. [Estudio paleontológico de la Formación Chachao, Informe Final. Yacimientos Petrolíferos Fiscales, Buenos Aires, 152 p. Unpublished.]. Defrance, J.L.M. 1826. Dictionnaire des Sciences Naturelles. e.g. Levrault, Strasbourg, 534 p. Deshayes, G.P., and Milne edwards, e. 1838. Histoire des Animaux Sans Vertèbres. Deuxième edition. Tome Huitième. J.B. Baillière, Paris, 660 p. erickson, J.M. 1974. Revision of the Gastropoda of the Fox hills Formation, Upper Cretaceous (Maestrichtian) of North Dakota. Bulletins of American Paleontology 66: 132–253. etheridge, R. 1902. A monograph of the Cretaceous invertebrate fauna of New South Wales. Memoirs of the Geological Survey of New South Wales 11: 1–98. etheridge, R. 1907. Lower Cretaceous fossils from the sources of the Barcoo, Ward and Nive Rivers, south central Queensland. Part I. Annelida, Pelecypoda and Gasteropoda. Records of the Australian Museum 6: 317–329. etheridge, R. 1920. Small Gasteropoda from the Lower Cretaceous of Queensland. Publications of the Geological Survey of Queensland 269: 8–21. Fernández, D.e., and Pazos, P.J. 2013. Xiphosurid trackways in a Lower Cretaceous tidal flat in Patagonia: Palaeoecological implications and the involvement of microbial mats in trace-fossil preservation. Palaeogeography, Palaeoclimatology, Palaeoecology 375: 16–29. Ferrari, S.M. 2012. The genera Cryptaulax and Procerithium (Procerithiidae, Caenogastropoda) in the early Jurassic of Patagonia, Argentina. Alcheringa 36: 325–339. Ferrari, S.M. 2015a. early Jurassic marine gastropods from Argentina: a palaeobiogeographical analysis based on Vetigastropoda. Journal of Systematic Paleontology 13: 919–941. Ferrari, S.M. 2015b. Systematic revision of Late Triassic marine gastropods from Central Perú: considerations on the Late Triassic/early Jurassic faunal turnover. Andean Geology 42: 71–96. Ferrari, S.M., Kaim, A., and Damborenea, S.e. 2014. The genera Calliotropis Seguenza and Ambercyclus n. gen. (Vetigastropoda, eucyclidae) from the early Jurassic of Argentina. Journal of Paleontology 88: 1174–1188. Finlay, h.J. 1927. A further commentary on New zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485. Fischer, P. 1885. Manuel de Conchyliologie et de Paléontologie Conchyliologique, ou Histoire Naturelle des Mollusques vivants et fossils. Fascicule 9. Librairie F. Savy. Paris, p. 785–896. Fischer, J.C., and Weber, C. 1997. Révision critique de la Paléontologie Française d’Alcide d’Orbigny Volume 2: Gastropodes Jurassiques. Masson and Muséum National d’histoire Naturelle, Paris, 300 p.
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA Fleming, J. 1822. The philosophy of zoology, a general view of the structure, functions and classification of animals, volume 2. Constable & Co., edinburgh, 618 p. Forbes, e. 1845. Catalogue of Lower Greensand fossils in the Museum of the Geological Society with notices of species new to Britain. Quarterly Journal of the Geological Society of London 1: 237–250, 345–355. Fritzsche, C.h. 1924. Neue kreidefaunen aus Südamerika (Chile, Bolivia, Peru, Columbia). In: G. Steinmann (ed.), Beiträge zur Geologie und Paläontologie von Südamerika, vol. 27. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilage-Band 50: 1– 56, 313–334. Gardner, J.S. 1876. On Cretaceous Gasteropoda. Family Scalidae. The Geological Magazine, New Series 3: 105–114. Gerasimov, P.A. 1992. [Gastropods from the Jurassic and Lower Cretaceous of European Russia]. Rossiyskaja Akademija Nauk, Moscow, 190 p. [in Russian]. Gill, T. 1870. On the Pterocerae of Lamarck, and their mutual relations. American Journal of Conchology 5: 120–139. Glasunova, A.e. 1968. [A new Neocomian member of Procerithiidae from the Volga Region]. In: B.L. Markovsky (ed.), [New Species of Ancient Plants and Invertebrates of the USSR, Volume 2, Part 1]. Nedra, Moscow, p. 295–296. [in Russian]. Gougerot, L., and Le Renard, J. 1983. Clefs de détermination des petites espèces de gastéropodes de l’eocène du Bassin Parisien. Cahiers des Naturalistes 39: 41–50. Gray, J.e. 1840. Shells of molluscous animals. In: British Museum (Corporate author), Synopsis of the contents of the British Museum, Forty-second edition, Second impression. Woodfall and Son, London, p. 106–156. Griffin, M., and Varela, A.N. 2012. Systematic palaeontology and taphonomic significance of the mollusc fauna from the Mata Amarilla Formation (lower Upper Cretaceous), southern Patagonia, Argentina. Cretaceous Research 37: 164–176. Gründel, J. 1975. Gastropoden aus dem Dogger. IV. euomphalidae, Pseudomelaniidae, Neritidae, Pyramidellidae und Acteonidae. Zeitschrift für Geologische Wissenschaften 3: 777–787. Gründel, J. 1976. zur Taxonomie und Phylogonie der BittiumGruppe (Gastropoda, Cerithiacea). Malakologische Abhandlungen Staatliches Museum für Tierkunde in Dresden 5: 33–59. Gründel, J. 1997. zur Kenntnis einiger Gastropoden-Gattungen aus dem französischen Jura und allgemeine Bemerkungen zur Gastropodenfauna aus dem Dogger Mittel- und Westeuropas. Berliner Geowissenschaftliche Abhandlungen, Reihe E, Paläobiologie 25: 69–129. Gründel, J. 1999. Procerithiidae (Gastropoda) aus dem Lias und Dogger Deutschlands und Polens. Paläontologie, Stratigraphie, Fazies. Freiberger Forschungshefte C 481: 1–37. Gründel, J. 2000. Archaeogastropoda aus dem Dogger Norddeutschlands und des nordwestlichen Polens. Berliner Geowissenschaftliche Abhandlungen, Reihe E, Paläobiologie 34: 205–253. Gründel, J. 2001. Gastropoden aus dem Jura der südamerikanischen Anden. Pälaontologie, Stratigraphie, Fazies. Freiberger Forschunshefte C 492: 43–84. Gründel, J. 2008. Remarks to the classification and phylogeny of the Ataphridae Cossmann, 1915 (Gastropoda, Archaeogastropoda) in the Jurassic. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 250: 177–197. Gründel, J., and Kaim, A. 2006. Shallow-water gastropods from Late Oxfordian sands in Kłęby (Pomerania, Poland). Acta Geologica Polonica 56: 121–157. Gründel, J., and Nützel, A. 2012. On the early evolution (Late Triassic to Late Jurassic) of the Architectibranchia (Gastropoda: hetero-
branchia) with a provisional classification. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 264: 31–59. Gründel, J., and Parent, h. 2001. Lower and Middle Tithonian marine gastropods from the Neuquén-Mendoza Basin, Argentina. Boletín del Instituto de Fisiografía y Geología 71: 13–18. Gründel, J., and Parent, h. 2006. Marine Jurassic gastropods of Argentina. III. Lower and Middle Tithonian of Picún Leufú and Cerro Lotena. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 8: 503–512. Gründel, J., Parent, h., Cocca, S.e., and Cocca, R., 2007. Marine Jurassic gastropods of Argentina. IV. A new species from the Middle Tithonian (Internispinosum Biozone) of Barda Negra, southern Neuquén-Mendoza Basin. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 245: 143–146. Guzhov, A.V. 2002. To the Revision of Some Jurassic Gastropods from Central Russia: 1. Genus Plicacerithium. Paleontological Journal 36: 338–342. Guzhov, A.V. 2004. Jurassic gastropods of the european Russia (Orders Cerithiiformes, Bucciniformes, and epitoniiformes). Paleontological Journal 38: 457–562. haas, O. 1953. Mesozoic invertebrate faunas of Peru. Bulletin of the American Museum of Natural History 101: 1–328. harbort, e. 1905. Die Fauna der Schaumburg-Lippeschen Kreidemulde. Abhandlungen der Preuβischen Geologischen Landesanstalt, Neue Folge 45: 1–111. harris, G.D. 1896. The Midway Stage. Bulletins of American Paleontology 1: 1–158. harris, G.D. 1899. The Lignitic Stage, Part II; Scaphopoda, Gastropoda, Pteropoda and Cephalopoda. Bulletins of American Paleontology 3: 1–128. haupt, O. 1907. Beiträge zur Fauna des oberen Malm und der unteren Kreide in der argentinischen Cordillere. In: G. Steinmann (ed.), Beiträge zur Geologie und Paläontologie von Südamerika, vol. 12. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie 23: 187–236. hébert, e., and eudes-Deslongchamps, e. 1860. Mémoire sur les fossiles de Montreuil-Bellay (Maine-et-Loire): I. Céphalopodes et Gastéropodes. Bulletin de la Société Linnéenne de Normandie 5: 153–240. hébert, e., and Munier-Chalmas, e. 1877. Recherches sur les terrains tertiaires de l’europe méridionale. Comptes rendus hebdomadaires des séances de l’Académie des Sciences 85: 125–128; 181–186. hickman, S., and McLean, J.h. 1990. Systematic revision and suprageneric classification of trochacean gastropods. Natural History Museum of Los Angeles County, Science Series 35: 1–169. hikuroa, D.C.h., and Kaim, A. 2007. New gastropods from the Jurassic of Orville Coast, eastern ellsworth Land, Antarctica. Antarctic Science 19: 115–124. Jack, R.L., and etheridge, R. 1892. The Geology and Paleontology of Queensland and New Guinea. Government Printer, Brisbane and Dulau & Co., London, 808 p. Jaworski, e. 1938. Gasterópodos del Cretácico Inferior de Colombia. In: e.A. Scheibe (ed.), Estudios geológicos y paleontológicos sobre la Cordillera Oriental de Colombia, parte 3. República de Colombia, Ministerio de Industrias y Trabajo, Departamento de Minas y Petróleos. P. and G. Gärtner, Bogota, p. 109–121. Kaim, A. 2001. Faunal dynamics of juvenile gastropods and associated organisms across the Valanginian transgression-regression cycle in central Poland. Cretaceous Research 22: 333–351. Kaim, A. 2004. The evolution of conch ontogeny in Mesozoic open sea gastropods. Palaeontologia Polonica 62: 1–183. Kaim, A., and Sztajner, P. 2004. The opercula of neritopsid gastropods and their phylogenetic importance. Journal of Molluscan
437
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440 Studies 71: 211–219. Kase, T. 1984. early Cretaceous marine and brackish-water Gastropoda from Japan. National Science Museum Monographs 1: 1– 263. Kase, T., and Maeda, h. 1980. early Cretaceous Gastropoda from the Choshi District, Chiba Prefecture, Central Japan. Transactions and Proceedings of the Palaeontological Society of Japan, New Series 118: 291–324. Kase, T., Kurihara, Y., Aguilar, Y.M., Pandita, h., Fernando, A.G.S., and hayashi, h. 2015. A new cerithioidean genus Megistocerithium (Gastropoda; Mollusca) from the Miocene of Southeast Asia: a possible relict of Mesozoic “eustomatidae”. Paleontological Research 19: 299–311. Kiel, S. 2006. New and little-known gastropods from the Albian of the Mahajanga Basin, northwestern Madagascar. Journal of Paleontology 80: 455–476. Kiel, S., and Bandel, K. 2001. About heterostropha of the Campanian of Torallola, Spain. Journal of the Czech Geological Survey 46: 319–334. Kiel, S., and Bandel, K. 2003. New taxonomic data for the gastropod fauna of the Umzamba Formation (Santonian-Campanian, South Africa) based on newly collected material. Cretaceous Research 24: 449–475. Kiel, S., Bandel, K., and Perilliat, M.C. 2002. New gastropods from the Maastrichtian of the Mexcala Formation in Guerrero, southern Mexico, part II: Archaeogastropoda, Neritimorpha and heterostropha. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 226: 319–342. Kilburn, R.N. 1985. The family epitoniidae (Mollusca: Gastropoda) in southern Africa and Mozambique. Annals of the Natal Museum 27: 239–337. Knight, J.B., Cox, L.R., Keen, A.M., Batten, R.L., Yochelson, e.L., and Robertson, R. 1960. Systematic descriptions. In: R.C. Moore (ed.), Treatise on Invertebrate Paleontology, Part I, Mollusca 1. Geological Society of America, Boulder, and University of Kansas Press, Lawrence, p. 169–310. Koken, e. 1897. Die Gastropoden der Trias um hallstadt. Jahrbuch der Kaiserlich-Königlichen Geologischen Reichsanstalt 46: 37–126. Kollmann, h.A. 1979. Gastropoden aus den Losensteiner Schichten der Umgebung von Losenstein (Oberösterreich) 3 Teil - Cerithiacea (Mesogastropoda). Annalen des Naturhistorischen Museums in Wien 82: 11–51. Kollmann, h.A. 2005. Révision critique de la Paléontologie Française d’Alcide d’Orbigny Volume 3: Gastropodes Crétacés. Blackhuys Publishers, Leiden, 239 p. Kollmann, h.A., and Peel, J.S. 1983. Paleocene gastropods from Nûgssuaq, West Greenland. Bulletin Grønlands Geologiske Undersøgelse 146: 1–115. Lamarck, J.-B. 1806. Mémoires sur les fossiles des environs de Paris, Suite 6. Annales du Muséum d’Histoire Naturelle 7: 53–430. Lamarck, J.-B. 1822. Histoire Naturelle des Animaux sans Vertèbres, Tome Sixième, 2me. partie. Printed by the author, Paris, 232 p. Lazo, D.G. 2007. early Cretaceous bivalves of the Neuquén Basin, west-central Argentina: Notes on taxonomy, palaeobiogeography and palaeoecology. Geological Journal 42: 127–142. Leanza, h.A., and hugo, C.A. 1997. Hoja Geológica 3969-III, Picún Leufú, provincia del Neuquén y Río Negro, República Argentina, Boletín 218. Servicio Geológico y Minero Argentino, Buenos Aires, 135 p. Leanza, h.A., hugo, C.A., Repol, D., Gonzalez, R., and Danieli, J.C. 2001. Hoja geológica 3969-I, Zapala, provincia del Neuquén, Boletín 275. Servicio Geológico y Minero Argentino, Buenos Aires, 128 p.
438
Legarreta, L., and Gulisano, C.A. 1989. Análisis estratigráfico secuencial de la Cuenca Neuquina (Triásico Superior–Terciario Inferior, Argentina). In: G.A. Chebli, and L.A. Spalletti (eds.), Cuencas Sedimentarias Argentinas. Serie Correlación Geológica 6. Instituto Superior de Correlación Geológica, San Miguel de Tucumán, p. 221–243. Leymerie, M.A. 1842. Suite du Mémoire sur le Terrain Crétacé du Departement de l’Aube, Seconde Partie. Mémoires de la Société Géologique de France 5: 1–34. Loriol, P. de. 1882. Études sur la faune des couches du Gault de Cosne (Niévre). Mémoires de la Société Paléontologique Suisse 9: 1–118. Lycett, J. 1863. Supplementary Monograph on the Mollusca from the Stonefield State, Great Oolite, Forest Marble, and Cornbrash. Palaeontographical Society Monographs 15: 1–129. Manceñido, M.O., and Damborenea, S.e. 1984. Megafauna de invertebrados paleozoicos y mesozoicos. 9º Congreso Geológico Argentino (San Carlos de Bariloche), Relatorio 2: 413–465. Matthews, S.C. 1973. Notes on open nomenclature and on synonymy lists. Palaeontology 16: 713–719. Meek, F.B. 1864. Check list of invertebrate fossils of North America; Cretaceous and Jurassic. Smithsonian Miscellaneous Collection 7: 1–40. Meek, F.B. 1876. A report on the invertebrate Cretaceous and Tertiary fossils of the upper Missouri country. U.S. Geological Survey of the Territories 9: 1– 629. Meek, F.B., and hayden, F.V. 1856. Descriptions of new fossil species of Mollusca collected by Dr. F.V. hayden in Nebraska Territory together with a complete catalogue of all the remains of invertebrate hitherto described and identified from the Cretaceous and Tertiary formations of that region. Proceedings of the National Academy of Sciences of Philadelphia 8: 265–286. Meek, F.B., and hayden, F.V. 1860. Descriptions of new organic remains from the Tertiary, Cretaceous and Jurassic rocks of Nebraska. Proceedings of the National Academy of Sciences of Philadelphia 12: 175–185. Mertin, h. 1939. Über Brackwasserbildungen in der Oberen Kreide des nördlichen harzvorlandes. Nova Acta Leopoldina Neue Folge 7: 139–263. Mongin, D. 1979. Mollusques du Crétacé Inférieur de Tunisie (Gastéropodes et Lamellibranches). Notes du Service Géologique de Tunisie, Travaux de Géologie Tunisienne 45: 107–153. Montfort, D. 1810. Conchyliologie Systématique et Classification Méthodique de Coquilles, Coquilles Univalves Non Cloisonnées, Tome second. F. Schoell, Paris, 676 p. Nagao, T. 1934. Cretaceous Mollusca from the Miyako District, honshu, Japan. Faculty of Science, Hokkaido Imperial University, Series 4 2: 17–277. Nützel, A., and Gründel, J. 2007. Two new gastropod genera from the early Jurassic (Pliensbachian) of Franconia (South Germany). Zitteliana Reihe A 47: 59–67. Nützel, A., and Senowbari-Daryan, B. 1999. Gastropods from the Upper Triassic (Norian-Rhaetian) Nayband Formation of central Iran. Beringeria 23: 93–132. Nützel, A., hamedani, A., and Senowbari-Daryan, B. 2003. Some Late Triassic Gastropods from the Nayband Formation in Central Iran. Facies 48: 127–134. Olsson, A.A. 1934. Contributions to the Paleontology of Northern Peru: the Cretaceous of the Amotape Region. Bulletins of American Paleontology 20: 1–140. Olsson, A.A. 1944. Contributions to the Paleontology of Northern Peru. Part VII. The Cretaceous of the Paita region. Bulletins of
CATALDO: eARLY CReTACeOUS GASTROPODS FROM ARGeNTINA American Paleontology 28: 163–304. Orbigny, A.d’. 1843. Paléontologie Française. Description zoologique et géologique de tous les animaux et rayonnés fossiles de France. Terrains Crétacés. Tome Deuxième, Gastéropodes. V. Masson, Paris, 456 p. Orbigny, A.d’. 1850. Prodrome de paléontologie stratigraphique universelle des animaux mollusques et rayonnés, Tome 1. V. Masson, Paris, 394 p. Ozawa, T., Köhler, F., Reid, D.G., and Glaubrecht, M. 2009. Tethyan relicts on continental coastlines of the northwestern Pacific Ocean and Australasia: molecular phylogeny and fossil record of batillariid gastropods (Caenogastropoda, Cerithioidea). Zoologica Scripta 38: 503–525. Pchelintsev, V.F. 1927. [The Jurassic and Lower Cretaceous fauna of the Crimea and the Caucasus]. Trudy Geologicheskogo Komiteta Novaya seriya 172: 1–367. [in Russian]. Pchelintsev, V.F. 1965. [Mesozoic Murchisoniata from the Crimean Mountains]. Nauka, Moscow-Leningrad, 215 p. [in Russian]. Philippi, R.A. 1844. Enumeratio molluscorum Siciliae, Volumen Secundum. e. Anton, halle, 303 p. Pictet, F.-J., and Campiche, G. 1861–1864. Description des fossiles du terrain Crétacé des environs de Sainte-Croix. Deuxième partie. 7me, 8me, 9me et 10me livraisons. h. Georg Librairie, Geneva, p 145–348. Piette, É. 1855. Observations sur les étages inférieurs du terrain Jurassique dans les départements des Ardennes et de l’Aisne. Bulletin de la Société Géologique de France Série 2 12: 1083– 1122. Piette, É. 1860. Sur un nouveau genre de Gastéropodes. Bulletin de la Société Géologique de France, Série 2 18: 14–16. Popenoe, W.P. 1957. The Cretaceous gastropod genus Biplica, its evolution and biostratigraphic significance. University of California Publications in Geological Sciences 30: 425–454. Quoy, J.R.C., and Gaimard, J.P. 1832. Voyage de découvertes de l’Astrolabe exécuté par ordre du Roi, pendant les années 1826–1827– 1828–1829, sur le commandement de M.J. Dumont d’Urville. Zoologie Tome Second, partie 1. J. Tastu, Paris, 320 p. Rafinesque, C.S. 1815. Analyse de la Nature, ou Tableau de l’Universe et des Corps Organisés. Printed by the author, Palermo, 224 p. Rosenkrantz, A. 1970. Marine Upper Cretaceous and lowermost Tertiary deposits in West Greenland. Meddelelser fra Dansk Geologisk Forening København 19: 406–453. Salvini-Plawen, L. 1980. A reconsideration of systematics in the Mollusca (Phylogeny and higher classification). Malacologia 19: 249–278. Saul, L.R., and Squires, R.L. 1997. New species of neritid gastropods from Cretaceous and Lower Cenozoic strata of the Pacific Slope of North America. The Veliger 40: 131–147. Saul, L.R., and Squires, R.L. 2003. New Cretaceous cerithiiform gastropods from the Pacific Slope of North America. Journal of Paleontology 77: 442–453. Sayn, G. 1932. Description de la faune de l’Urgonien de Barcelonne (Drôme). Travaux du Laboratoire de Géologie de la Faculté des Sciences de Lyon 18: 1–70. Schmidt, M. 1905. Über Oberen Jura in Pommern. Beiträge zur Stratigraphie und Paläontologie. Abhandlungen der Preußischen Geologischen Landesanstalt, Neue Folge 41: 1–222. Schröder, M. 1995. Frühontogenetische Schalen jurassischer und unterkretazischer Gastropoden aus Norddeutschland und Polen. Palaeontographica, Abteilung A 238: 1–95. Schwarz, e., Spalletti, L.A., Veiga, G.D., 2011. La Formación Mulichinco (Cretácico Temprano) en la Cuenca Neuquina. In: h.A. Le-
anza, C. Arregui, O. Carbone, J.C. Danieli, and J.M. Vallés (eds.), Geología y Recursos Naturales de la provincia del Neuquén. Asociación Geológica Argentina, Buenos Aires, p. 131–144. Seguenza, L. 1903. Molluschi poco noti dei Terreni Terziarii di Messina. Trochidae e Solariidae. Bolletino della Società Geologica Italiana 21: 455–465. Sohl, N.F. 1965. Marine Jurassic gastropods, central and southern Utah. U.S. Geological Survey Professional Paper 503D: 1–29. Sohl, N.F. 1967. Upper Cretaceous gastropods from the Pierre Shale at Red Bird, Wyoming. U.S. Geological Survey Professional Paper 393B: 1–46. Sohl, N.F. 1987. Cretaceous gastropods: Contrasts between Tethys and temperate provinces. Journal of Paleontology 61: 1085–1111. Spalletti, L.A., Poiré, D.G., Pirrie, D., Matheos, S.D., and Doyle, P. 2001. Respuesta sedimentológica a cambios en el nivel de base en una secuencia mixta clástica-carbonática del Cretácico de la Cuenca Neuquina, Argentina. Revista de la Sociedad Geológica de España 14: 57–74. Spalletti, L.A., Veiga, G.D., and Schwarz, e. 2011. La Formación Agrio (Cretácico Temprano) en la Cuenca Neuquina. In: h. Leanza, C. Arregui, O. Carbone, J.C. Danieli, and J.M. Vallés (eds.), Geología y Recursos Naturales de la Provincia del Neuquén. Asociación Geológica Argentina, Buenos Aires, p. 145–160. Squires, R.L., and Saul, L.R. 1993. A new species of Otostoma (Gastropoda: Neritidae) from near the Cretaceous/Tertiary boundary at Dip Creek, Lake Nacimiento, California. The Veliger 36: 259– 264. Squires, R.L., and Saul, L.R. 2001. New Late Cretaceous gastropods from the Pacific Slope of North America. Journal of Paleontology 75: 46–65. Squires, R.L., and Saul, L.R. 2003. New Late Cretaceous epitoniid and zygopleurid gastropods from the Pacific Slope of North America. The Veliger 46: 20–49. Squires, R.L., and Saul, L.R. 2004. The pseudomelaniid gastropod Paosia from the marine Cretaceous of the Pacific Slope of North America, and a review of the age and paleobiogeography of the genus. Journal of Paleontology 78: 484–500. Sruoga, P., etcheverría, M., Folguera, A., Repol, D., and zanettini, J.C. 2005. Hoja Geológica 3569-I, Volcán Maipo, Provincia de Mendoza, Boletín 290. Servicio Geológico y Minero Argentino, Buenos Aires, 114 p. Stanton, T.W. 1901. Volume IV, Palaeontology I, part I, The marine Cretaceous Invertebrates. In: W.B. Scott (ed.), Reports of the Princeton University Expeditions to Patagonia, 1896–1899, J.B. Hatcher in charge. The University of Princeton, Princeton and Schweizerbart’sche, Sttutgart p. 1–43. Steinmann, G. 1929. Geologie von Peru. Carls Winters Universitätsbuchhandlung, heidelberg, 448 p. Stilwell, J.D. 2005. A new species of Calliotropis (Mollusca: Gastropoda: Vetigastropoda: Trochidae: eucyclinae) from the eocene of Antarctica. Molluscan Research 25: 9–13. Stilwell, J.D., and henderson, R.A. 2002. Description and paleobiogeographic significance of a rare Cenomanian molluscan faunule from Bathurst Island, Northern Australia. Journal of Paleontology 76: 447–471. Swainson, W. 1840. A Treatise on Malacology, or The Natural Classification of Shells and Shell-fish. Longman, Orme, Brown, Green & Longmans, London, 416 p. Szabó, J. 2008. Gastropods of the early Jurassic hierlatz Limestone Formation; part 1: a revision of type collections from Austrian and hungarian localities. Fragmenta Palaeontologica Hungarica 26: 1–108.
439
AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440 Tate, R. 1869. Contributions to Jurassic Palaeontology 1: Cryptaulax, a new genus of Cerithiadae. The Annals and Magazine of Natural History 4: 417–419. Thomson, M.R.A. 1971. Gastropoda from the Lower Cretaceous sediments of south-eastern Alexander Island. British Antarctic Survey Bulletin 25: 45–58. Tracey, S. 2010. Chapter 10: Gastropods. In: J.R. Young, A.S. Gale, R.I. Knight, and A.B. Smith (eds.), Fossils of the Gault Clay. Palaeontological Association Field Guide to Fossils 12. The Palaeontological Association, London, p. 106–155 Tracey, S., Todd, J.A., and erwin, D.h. 1993. Chapter 8: Mollusca, Gastropoda. In: M.J. Benton (ed.), The fossil record 2. Chapman & hall, London, p. 131–167. Trautschold, h. 1866. zur Fauna des russischen Jura. Bulletin de la Societé Impériale des Naturalistes de Moscou 39: 1–24. Tunnell, J.Jr., Andrews, J., Barrera, N., and Moretzsohn, F. 2010. Encyclopedia of Texas Seashells, identification, ecology, distribution, and history. harte Research Institute for Gulf of Mexico Studies Series. Texas A&M University Press, College Station, 343 p. Verrill, A.e. 1884. Second catalogue of Mollusca recently added to the fauna of the New england coast and the adjacent part of the Atlantic, consisting mostly of deep-sea species, with notes on others previously recorded. Transactions of the Connecticut Academy of Arts and Sciences 6: 139–194. Vidal, L.M. 1917. Nota paleontológica sobre el Cretáceo de Cataluña. Asociación Española para el Progreso de las Ciencias, Congreso de Sevilla 5: 3–19. Viera, L.I. 1991. Confusiscala mirambelensis (Gastropoda) en el Neocomiense de Igea (La Rioja). Consecuencias. Munibe 43: 3–7. Vilanova, J. 1863. Ensayo de descripción geognóstica de la provincia de Teruel. Imprenta Nacional, Madrid, 312 p. von der Osten, e. 1957. A fauna from the Lower Cretaceous Barranquin Formation of Venezuela. Journal of Paleontology 31: 571–590. Weaver, C.e. 1931. Paleontology of the Jurassic and Cretaceous of West Central Argentina. Memoirs of the University of Washington 1: 1–595. Wenz, W. 1938. Gastropoda, Teil I: Allgemeiner Teil und Prosobranchia. In: O.h. Schindewolf (ed.), Handbuch der Paläozoologie, Band 6. G. Bornträger, Berlin, 1639 p. White, C.A. 1876. Invertebrate paleontology of the Plateau province, together with notice of a new species from localities beyond its limits in Colorado. In: J.W. Powell (ed.), Report on the geology of the eastern portion of the Uinta Mountains and a region of country adjacent thereto. Government Printing Office, Washington, p. 74–135.
440
White, C.A. 1883. Contributions to invertebrate paleontology, no. 7; Jurassic fossils from the Western Territories. Annual Report of the U.S. Geological and Geographical Survey of the Territories 12: 143–155. White, C.A. 1888. Contributions to the palaeontology of Brazil, comprising descriptions of Cretaceous Invertebrate Fossils mainly from the provinces of Sergipe-Pernambuco, Para, and Bahia. Archivos do Muséu Nacional do Rio de Janeiro 7: 1–273. White, C.A. 1889. On Invertebrate Fossils from the Pacific Coast. Bulletin of the United States Geological Survey 51: 9–63. Whitfield, R.P. 1876. Descriptions of new species of fossils. In: W. Ludlow, Report of a reconnaissance from Carroll, Montana Territory, on the Upper Missouri to the Yellowstone National Park and return, made in the summer of 1875. Government Printing Office, Washington D.C., p. 141–145. Whitfield, R.P. 1891. Observation of some Cretaceous fossils from the Beyrut district of Syria, in collections of the American Museum of Natural history, with descriptions of some new species. Bulletin of the American Museum Natural History 3: 381–441. Wollemann, A. 1908. Nachtrag zu meinen Abhandlungen über die Bivalven und Gastropoden der Unteren Kreide Norddeutschlands. Jahrbuch der Königlich Preuβischen Geologischen Landesanstalt zu Berlin 29: 151–193. zittel, K.A. 1882. Handbuch der Paläontologie. Abteilung 1, Paläozoologie, Band 2, Mollusca und Arthropoda. R. Oldenbourg, MunichLeipzig, 893 p.
doi: 10.5710/AMGh.14.12.2016.3053 Submitted: September 22nd, 2016 Accepted: December 14th, 2016
TABLE – Comprehensive list of gastropod records from the Tithonian–Barremian of the Neuquén Basin, with original data as well as revised taxonomy and occurrence. See Figure 2 for geographic and stratigraphic reference. Taxon Reference Original record Revised taxonomy Revised occurrence/Comments Acteon andinus Haupt (1907, p. “Neocomian”–Aptian Tornatellaea? andina Tithonian (Vaca Muerta Fm.). Haupt, 1907 * 206, pl. 10, fig. S Neuquén (Acteonoidea, Acteonidae) See Gründel et al. (2007, p. 144, fig. 2, under 4a,b) T. mendozana) and this paper. Alaria acuta Behrendsen (1892, “Neocomian” ?Mathildoidea, ?Mathildidae Early Valanginian (Mulichinco Fm.). Behrendsen, 1892 * p. 19, pl. 4, fig. N and central Neuquén 3a,b) Alaria acutecarinata Behrendsen (1891, Tithonian Nerineopsis acutecarinata Upper Hauterivian (Agrio Fm., Agua de la Behrendsen, 1891 * p. 413) S Mendoza (Campaniloidea, fam. unknown) Mula Mbr.). See Cataldo and Lazo (2016, p. 175, fig. 8). Alaria cf. glacus Haupt (1907, p. Tithonian Protohemichenopus Likely Vaca Muerta or Picún Leufú fms. d'Orbigny, 1843 ‡ 205) S Neuquén neuquensis Camacho See Cataldo (2014a, p. 1227, fig. 3). (Stromboidea, Aporrhaidae) Amauropsis sp. ‡ Weaver (1931, p. Upper Agrio Fm. Ampullina pichinka Cataldo Upper Hauterivian (Agrio Fm., Agua de la 380) Hauterivian and Lazo Mula Mbr.). Central Neuquén (Campaniloidea, Ampullinidae) See Cataldo and Lazo (2016, p. 173, fig. 7). Aporrhaidae sp. Damborenea et al. Vaca Muerta and Protohemichenopus See Cataldo (2014a, p. 1227, fig. 3.21). (1979, p. 71, pl. 11, Chachao fms. neuquensis (pars.) indet. α * Upper Berriasian– fig. 3) lower Valanginian S Mendoza Aporrhaidae sp. Damborenea et al. Agrio Fm. ?Protohemichenopus See Cataldo (2014a, p. 1227). indet. β * (1979, p. 72) Upper Hauterivian neuquensis (pars.) S Mendoza Capulus argentinus Haupt (1907, p. Tithonian Unrevised Likely Vaca Muerta or Picún Leufú fms. Haupt * 204, pl. 8, fig. 5a,b) S Neuquén Needs further study. Cerithium cf. heeri Weaver (1931, p. Agrio Fm. Paraglauconia (Diglauconia) Upper Hauterivian–lower Barremian (Agrio Pictet and Renevier, 385, pl. 42, fig. 284) Central Neuquén aff. peruana Fritszche Fm.). 1854 * (Cerithioidea, Cassiopidae) Cerithium sp. * Haupt (1907, p. “Neocomian”–Aptian Paleorissoina sp. Upper Hauterivian–lower Barremian (Agrio 206, pl. 10, fig. S Neuquén (Rissoidea, Paleorissoinidae) Fm.). 5a,b) Cinulia sp. ‡ Behrendsen (1892, “Neocomian” Unrevised (lost material) Likely Valanginian (Agrio Fm., Pilmatué Mbr.). p. 18) N Neuquén See text.
1
Cinulia? sp. *
Cinulia cf. andina Haupt, 1907 *
Manceñido and Damborenea (1984, p. 428, pl. 2, fig. 12, 13) Damborenea et al. (1979, p. 76)
Ortíz Fm. Tithonian–?Berriasian Río Negro
Acteonoidea gen. et sp. indet.
Also in the lower Hauterivian of Neuquén (Agrio Fm.).
Vaca Muerta Fm. Berriasian S Mendoza Chachao and Agrio fms. S Mendoza Agrio Fm. Central Neuquén
Acteonoidea? gen. et sp. indet.
See text.
Eunerinea mendozana Cataldo (Nerineoidea, Eunerineidae)
Uppermost Hauterivian–lowermost Barremian (Agrio Fm., Agua de la Mula Mbr.). See Cataldo (2013). Upper Hauterivian (Agrio Fm.). See Cataldo (2014a, p. 1229, fig. 4).
Cossmannea (Eunerinea) sp. indet. * Dicroloma cf. obtusa (Pictet and Campiche, 1864) * Dicroloma glaucus (d'Orbigny, 1843) *
Damborenea et al. (1979, p. 73, pl. 11, fig. 1) Weaver (1931, p. 386) Weaver (1931, p. 386)
“Neocomian” S Neuquén and S Mendoza
Emarginula sp. ‡
Tithoniano S Mendoza Ortíz Fm., Tithonian–?Berriasian Río Negro
Unrevised (lost material)
?Vaca Muerta Fm.
Campaniloidea, Ampullinidae
Needs further study.
Gastropoda gen. et sp. indet. I *
Behrendsen (1891, p. 414) Manceñido and Damborenea (1984, p. 428, pl. 2, fig. 8– 11) Damborenea et al. (1979, p. 77)
?Dimorphosoma weaveri
?Upper Hauterivian See Cataldo (2014a, p. 1229).
Gastropoda gen. et sp. indet. II *
Damborenea et al. (1979, p. 77)
?Mathildoidea,?Mathildidae (pars.)
Likely same species as “Alaria acuta” from Mulichinco Fm.
Harpagodes cf. americanus Imlay, 1940 * Harpagodes oceani (Brongniart, 1821) ‡ Homalopoma sp. nov. *
Damborenea et al. (1979, p. 70, pl. 11, fig. 4) Haupt (1907, p. 204) Manceñido and Damborenea (1984,
Agrio and ?Chachao fms. S Mendoza Vaca Muerta Fm. Upper Valanginian S Mendoza Chachao and Agrio fms. S Mendoza Tithonian S Neuquén Ortíz Fm. Tithonian–?Berriasian
Harpagodes jaworskii (Weaver) (Stromboidea, Aporrhaidae)
Upper Hauterivian–lower Barremian (Agrio Fm., Agua de la Mula Mbr.). See Cataldo (2014a, p. 1232, figs. 5, 6). Likely Picún Leufú Fm. See Cataldo (2014a, p. 1232). Needs further study. Possibly also in the Mulichinco Fm. of
Euspira sp. *
Dimorphosoma weaveri Cataldo (Stromboidea, Aporrhaidae) Protohemichenopus neuquensis
?Harpagodes jaworskii Turbinoidea, ?Colloniidae
See Cataldo (2014a, p. 1227, fig. 3).
2
Lissochilus sp. indet. * Lissochilus sp. nov. *
Natica cf. bulimoides (Deshayes in Leymerie, 1842) * Natica praelonga Deshayes in Leymerie, 1842 * Naticidae gen. et sp. indet. *
p. 427, pl. 2, figs. 6– 7) Damborenea et al. (1979, p. 69) Manceñido and Damborenea (1984, p. 427, pl. 2, figs. 2– 5) Weaver (1931, p. 372, pl. 43, figs. 289–290) Weaver (1931, p. 372) Damborenea et al. (1979, p. 75)
Nerinea cf. nerinaeformis (Coquand, 1862) *
Weaver (1931, p. 383)
Odostomopsis? sp. indet. *
Damborenea et al. (1979, p. 76)
Patella sp. *
Behrendsen (1891, p. 414) Damborenea et al. (1979, p. 68)
Pleurotomaria (Conotomaria) cf. pailleteana d’Orbigny, 1843 Pleurotomaria gerthi Weaver, 1931 * Pleurotomaria cf. provincialis
Weaver (1931, p. 367, pl. 42, fig. 281–283) Weaver (1931, p. 366)
Río Negro
Neuquén
Chachao Fm. Upper Valanginian S Mendoza Ortíz Fm. Tithonian–?Berriasian Río Negro
Lissochilus sp. a (Neritoidea, Neritidae)
Needs further study.
Lissochilus sp. b
Possibly also in the Agrio Fm. of Neuquén.
Upper Agrio Fm. Central Neuquén
Ampullina sp. (Campaniloidea, Ampullinidae)
Upper Hauterivian (Agua de la Mula Member, Agrio Fm.).
Lower Agrio Fm. S Neuquén
Harpagodes jaworskii
See Cataldo (2014a, p. 1234, fig. 6.3–4).
Chachao Fm. Valanginian S Mendoza Upper Agrio Fm. Upper Barremian– lower Aptian N Neuquén Chachao Fm. Upper Valanginian S Mendoza Tithonian S Mendoza Chachao and ?Agrio fms. S Mendoza
Ampullina sp.
Same species as in Agrio Fm. from Neuquén and Mendoza.
?Campaniloidea, ?Campanilidae
Uppermost Hauterivian–lowermost Barremian (Agrio Fm., Agua de la Mula Mbr.).
?Harpagodes jaworskii
See Cataldo (2014a, p. 1234, fig. 6.10).
Unrevised (very poorly preserved) Pleurotomariidae gen. et sp. indet.
?Vaca Muerta Fm.
Upper Agrio Fm. Barremian S Neuquén Quintuco Fm. Lower Valanginian
Pleurotomaria gerthi (Pleurotomarioidea, Pleurotomariidae) Bathrotomaria cf. provincialis (Pleurotomarioidea,
Upper Valanginian–lower Hauterivian (Pilmatué Mbr., Agrio Fm.). See Cataldo and Lazo (2012, p. 81, figs. 4, 5). Tithonian (Picún Leufú Fm.). See Cataldo and Lazo (2012, p. 87, figs. 5.5–8,
Upper Hauterivian (Agrio Fm., Agua de la Mula Mbr.)
3
d'Orbigny, 1843 * Pleurotomaria cf. jaccardi Pictet and Campiche, 1863 * Polinices? bodenbenderi (Behrendsen, 1891) * Protohemichenopus neuquensis Camacho, 1953* Pterocera sp. Tornatella sp. ‡ Trochus sp. indet. A *
Weaver (1931, p. 368) Manceñido and Damborenea (1984, p. 428, pl. 2, fig. 14–16) Camacho (1953, p. 191, figs. 1, 2) Burckhardt (1900) Behrendsen (1891, p. 413) Weaver (1931, p. 371)
Trochus sp. indet. B *
Weaver (1931, p. 371)
Turbo bodenbenderi Behrendsen, 1891 * “Turritella” sp. *
Behrendsen (1891, p. 413, pl. 25, fig. 5) Manceñido and Damborenea (1984, p. 427, pl. 2, fig. 1) Weaver (1931, p. 381, pl. 42, fig. 285)
Turritella cf. lineolata Roemer, 1841 * Turritellidae gen. et sp. indet α * Turritellidae gen. et sp. indet β *
Damborenea et al. (1979, p. 74, pl. 11, fig. 2) Damborenea et al. (1979, p. 74)
S Neuquén Lower Agrio Fm. S Neuquén
Pleurotomariidae) Pleurotomaria gerthi
5.12–13, tab. 1). See Cataldo and Lazo (2012, p. 81, fig. 5.9– 11).
Ortíz Fm. Tithonian–?Berriasian Río Negro
?Neritoidea, ?Neritidae
Needs further study.
Agrio Fm. Hauterivian N Neuquén “Neocomian” Central Neuquén
Protohemichenopus neuquensis
Tithonian S Mendoza Upper Agrio Fm. ?Aptian Central Neuquén “Lower Neocomian” S Mendoza
Unrevised (lost material)
Lower Tithonian–upper Hauterivian (Vaca Muerta, Mulichinco, Chachao and Agrio fms.). See Cataldo (2014a, p. 1227, fig. 3). Likely Mulichinco Fm. Non-described record, no material associated to it. Likely Vaca Muerta Fm. See text. Berriasian (Huitrín Fm., La Tosca Mbr.).
Tithonian S Mendoza Ortíz Fm. Tithonian–?Berriasian Río Negro Upper Agrio Fm. Barremian–Aptian Central Neuquén Chachao Fm. Upper Valanginian S Mendoza Agrio Fm. Upper Hauterivian S Mendoza
Unrevised (too poorly preserved) Cerithioidea, ?Turritellidae
?Harpagodes jaworskii
?Valvatoidea, ?Provalvatidae ?Pleurotomaria gerthi
?Berriasian–Valanginian Likely top of Vaca Muerta Fm. or base of Chachao Fm. Likely Vaca Muerta Fm. Also in Picún Leufú Fm. (upper Tithonian) of Neuquén.
Mesalia? kushea Cataldo and Lazo (Cerithioidea, Turritellidae) Cerithioidea, ?Turritellidae
Upper Hauterivian (Agrio Fm, Agua de la Mula Mbr.). See Cataldo and Lazo (2016, p. 178, fig. 9). Needs further study.
?Campaniloidea, ?Campanilidae
Uppermost Hauterivian–lowermost Barremian. Likely same species as “Nerinea cf.
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Upper Agrio Fm. Upper Hauterivian– Barremian Central Neuquén Agrio Fm. Upper Hauterivian S Mendoza Symbols: *= original material examined; ‡= original material lost Tylostoma jaworskii Weaver, 1931 *
Weaver (1931, p. 379, pl. 41, fig. 273, pl. 43, figs. 287– 288) Damborenea et al. (1979, p. 75)
Harpagodes jaworskii
Harpagodes jaworskii and Ampullina sp.
nerinaeformis” by Weaver (1931). Upper Valanginian–uppermost Hauterivian/lower Barremian (both mbrs. of Agrio Fm.). See Cataldo (2014a, p. 1232, fig. 6). Likely upper Hauterivian (Agrio Fm, Agua de la Mula Mbr.). See Cataldo (2014a, p. 1232, figs. 5, 6).
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