new records of marine gastropods from the lower ...

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..

405

AMeGhINIANA - 2017 - Volume 54 (4): 405 – 440

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.

407


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


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

409


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


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.

411


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.


(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


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


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


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


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.

417


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


419


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


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


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.


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

423


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


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-

425


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


427


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


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).

429


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-


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


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


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


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.

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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)


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


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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