Journal of South American Earth Sciences 108 (2021) 103209

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Journal of South American Earth Sciences

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Paleoclimate and changing composition of the Paleogene-Neogene shallow

Molluscan Assemblages of Patagonia
Claudia Julia del Río
Museo Argentino de Ciencias Naturales B. Rivadavia. A. Gallardo 470 (C1405DJR), Buenos Aires, Argentina

A R T I C L E I N F O A B S T R A C T

Keywords: A detailed compilation and updated taxonomy of the shallow molluscan assemblages of the Paleogene-Neogene
Paleoclimates of Patagonia was performed. Together with the Sr-chronostratigraphic framework developed during the last
Patagonia decade, five assemblages of Danian, late middle Eocene, latest Oligocene-early Miocene, middle Miocene and late
Marine molluscs
Miocene ages were recognized. Temporal variation in the composition of these assemblages reflects the suc­
Paleogene-Neogene
South Western Atlantic Ocean
cessive transient hyperthermal events superimposed to the global cooling trend in the sea surface temperatures
during Cenozoic times. Tropical or warmest subtropical temperatures in the region during the Danian and late
middle Eocene times coincide respectively with the Dan-C2 and the Latest Danian Event, and the Middle Eocene
Climatic Optimum events. Tropical taxa, represented by Tethyan and warm-Cosmopolitan genera constitute
27.6% and 31.3% of the fauna respectively. The declining participation of Paleogene warm-genera, comprising
just 15.3% of the fauna, coupled with the massive appearance of the subtropical Neogene Southern elements,
reveals a decrease in sea surface temperatures during the latest Oligocene-early Miocene interval until the
establishment of warm-temperate conditions in the area. Although lower than during the Paleogene, tempera­
tures should have been higher than today, in coincidence with the late Oligocene warming and the multiple
warming events in the early Miocene. The middle Miocene records the first large occurrence of Caribbean taxa
that will be continue being important components of the fauna during the late Miocene. The contribution of
warm-taxa to the faunas rose from 15.3% in the early Miocene to 23% and 22.8% in the middle and late Miocene
respectively. Both faunas point to the development of subtropical conditions in the area and match with the
record of three new warming peaks. One hyperthermal event is recognized at the Middle Miocene Climatic
Optimum, and the other two increments coincide with the transient peaks occurring at ~11 Ma and ~9 Ma.
Paleogene-Neogene heritage in modern faunas of the region is rather poor and comprises 36% of fossil survivor
genera. Taxa that managed to survive are mainly Cosmopolitan, Neogene Southern and Endemic genera that
mostly had appeared in the early Miocene and adapted to Recent cool and temperate-cool conditions of the
Southwesten Atlantic Ocean.

1. Introduction
Cenozoic molluscan faunas of Patagonia drew the attention of nat­
uralists since the middle 19th century, when d’Orbigny (1842) and
Sowerby (in Darwin, 1846) described the first fossil species. Later, these
assemblages were focused of detailed studies performed by Ortmann
(1902), Ihering (1897, 1904), and were finally summarized by Ihering
(1907) in his revision of the Tertiary molluscan faunas of Patagonia.
After these pioneering works, followed the analyses of the Cretaceous
and Cenozoic faunas carried out by Feruglio (1936) and Furque and
Camacho (1949), and the short contributions of Frenguelli (1932),
Feruglio (1954), Camacho (1953; 1957), Camacho and Fernández
(1956), Rossi de García (1959) and Rossi de García and Levy (1977),
among others.
Modern taxonomic studies refer to the faunas contained in the Car­
men Silva Formation (early Miocene) (Malumián et al.,1978), to that
yielded in the early Miocene San Julián Formation exposed at the
southern San Jorge littoral (Zinsmeister, 1981), to the Eocene bivalves
of the Río Turbio Formation described by Griffin (1991), to the late
Miocene molluscs of the Puerto Madryn Formation studied by del Río
(1992; 1994) and Brunet (1995; 1997), to the Danian gastropods of the
Cerro Dorotea Formation (Griffin and Hünicken, 1994) and the molluscs
contained in the early Miocene El Chacay Formation (Chiesa et al.,
1995). These works were accompanied by the revisions of specific

E-mail address: claudiajdelrio@gmail.com.

https://doi.org/10.1016/j.jsames.2021.103209
Received 20 October 2020; Received in revised form 4 February 2021; Accepted 5 February 2021
Available online 15 February 2021
0895-9811/© 2021 Elsevier Ltd. All rights reserved.
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 1. -Geographic location of main Paleogene-Neogene fossiliferous areas.1-Leticia Formation (late middle Eocene) exposed in the surrounding of Tolhuin; 2- Cerro
Dorotea Formation (Danian) and Río Turbio Formation (late middle Eocene) exposed at Río Turbio; 3- Man Aike Formation (late middle Eocene); 4– Salamanca
Formation (Danian) exposed between Puerto Visser and Bahía Malaspina; 5- exposures of the Salamanca Formation between Puesto Alvarez and Canadón Iglesias; 6-
Salamanca Formation at Sierra de los Guanacos; 7-Lefipan Formation (Danian) exposed between Piedra Parada and Paso del Sapo; 8- exposures of the Roca Formation
at the Colitoro Plateau (Danian); 9- Roca Formation at the north of the city of General Roca; 10- Roca Formation in the surroundings of Cerros Bayos; 11- Carmen
Silva Formation (early Miocene) exposed in northern Tierra del Fuego between Río Grande and San Sebastián; 12- Estancia 25 de Mayo Formation (early Miocene)
exposed in the surroundings of Lago Argentino; 13-15- fossiliferous areas situated between Lago Cardiel and Lago Buenos Aires where the El Chacay Formation (early
Miocene) is exposed; 16- area of exposures of the Monte Léon Formation (early Miocene) 50 km to the south of the mouth of the Río Santa Cruz; 17- Monte Léon
Formation at the mouth of the Santa Cruz River; 18- San Julián (latest Oligocene) and Monte Léon (early Miocene) Formations at the Bajo de San Julián; 19- ex­
posures of the San Julián Formation between Puerto San Julián and Cabo Curioso (latest Oligocene); 20–21 area of exposures of the San Julián Formation between
Cabo Tres Puntas (20) and Punta Casamayor (21) (early Miocene); 22- early and middle Miocene Chenque Formation exposed between Punta Maqueda and sur­
roundings of Comodoro Rivadavia; 23- upper section of the Chenque Formation at Valle Hermoso (middle Miocene); 24–26 area of exposures of the Camarones
Formation (middle Miocene) from vicinities of Camarones (24); Estancia Maciega - cabo Raso (25) and Estancia la Antonieta- Isla Escondida (26); 27- Gaiman
Formation exposed at Trelew - Gaiman; 28-30- area of exposures of the Puerto Madryn Formation (late Miocene) at península Valdés (28) and from Puerto Madryn
(29) northwards to Cueva Los Leones (30); 31- exposures of Facies Balneario La Lobería (Río Negro Formation) (early Miocene) between Viedma and Barranca Final;
32- Gran Bajo del Gualicho Formation (middle and late Miocene) at Salinas and Bajo del Gualicho.

taxonomic groups such as pectinids (Morra, 1985; Morra and Erdmann,
1986; del Río, 1995), trochids (del Río, 1985; del Río and Morra, 1985;
Morra and del Río, 1987; Reichler and Camacho 1994), anomiids
(Camacho, 1985), struthiolarids (Camacho and Zinsmeister, 1989),
limids (Parma et al., 1990), mitilids (Griffin, 1990); volutomitrids
(Olivera and Camacho, 1990), muricids (Olivera et al., 1994), veneroids
(del Río, 1997), oysters (Casadío, 1998), and nuculoids and arcoids
(Erdmann and Morra, 1985; del Río and Camacho, 1996; 1998).
After the initial efforts to explain their origin and biostratigraphic
meaning (Ihering, 1907; Feruglio, 1949), an attempt to elucidate the
composition and biostratigraphy of Neogene assemblages was accom­
plished by del Río (1988; 1990; 2004a), but the wealth of new taxo­
nomic and stratigraphic results obtained along the last 15 years,
modified in part the conclusions previously obtained. These data
involved the analyses of new assemblages of Danian (del Río et al., 2007;
del Río, 2012), late middle Eocene (Camacho et al., 2000a,b; Casadío

2
C. Julia del Río
et al., 2009) and middle Miocene (Reichler, 2010) ages. Moreover, a
renewed interest on the taxonomy of specific groups concerned pecti­
nids (del Río, 2004b, 2006, del Río et al., 2008; Pastorino and Griffin,
2018; Santelli and del Río, 2019a, b), oysters (Griffin et al., 2005),
muricids (Griffin and Pastorino, 2005; Herbert and del Río, 2005),
volutids (del Río and Martínez, 2006), mitilids (Griffin et al., 2008),
naticids (Griffin and Pastorino, 2013), veneroids (Pérez and del Río,
2013; Alvarez et al., 2020), crassatellids (Santelli and del Río, 2014),
carditids (Pérez and del Río, 2017a,b), micromolluscs (Griffin et al.,
1998; Griffin and Pastorino, 2012), struthiolarids (Lopez Cabrera and
Olivero, 2018), and other less abundant groups (Griffin and Pastorino,
2006a, b).
Besides, the ongoing numerical datings of the marine units involved
allowed addressing an accurate stratigraphic distribution of the fauna
and its correlation among Patagonian basins (Parras et al., 2008; 2012;
Cuitiño et al. 2012; 2015a,b; del Río et al., 2013; 2018).
Finally, prospection of new fossiliferous sites during the last 35 years
carried on by the author between Lago Cardiel and Lago Buenos Aires
(Western Santa Cruz Province), General Roca (Neuquen Province),
Colitoro Plateau (Río Negro Province) and Tierra del Fuego, has
particularly enhanced our knowledge of the Paleogene -Neogene faunas
of Patagonia.
Thus, the noticeably increased bulk of data compiled in reference to
the age and composition of Paleogene-Neogene molluscs, enables us to
outline the evolution of these faunas, their paleoclimatic meaning, and
their participation in the Recent shallow assemblages of the
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Journal of South American Earth Sciences 108 (2021) 103209
Fig. 2. Chronostratigraphic chart of Paleogene-
Neogene deposits in Patagonia. Marine units
addressed in the text (in blue). Information sources
dealing with ages and stratigraphic relationships
indicated by numbers: (1) after Barrio (1990) and
Malumián and Nañez (2011); (2) after Getino (1995);
(3) after Scasso et al. (2012); (4) after Anselmi et al.
(2004); (5) after Bellosi (2010) and Cuitiño et al.
(2015a); (6) after Cuitiño et al. (2012; 2015b), and
Torres Carbonell and Olivero (2019); (7) after Parras
et al. (2008); (8) after Torres Carbonell and Olivero
(2019)
Southwestern Atlantic Ocean.
2. Geological setting
Tha analyzed molluscan faunas are contained in marine rocks of
Danian, late middle Eocene, latest Oligocene-early Miocene, middle
Miocene and late Miocene ages exposed from the southernmost tip of
South America (Tierra del Fuego) to northern Patagonia, in the Austral,
Golfo San Jorge, Asfalto-Somuncura, Colorado-Neuquina and Valdés
basins, spanning about 17◦ of latitude (between 54◦ 30′ S and 37◦ 40’ S).
A brief summary of the fossiliferous units will be provided below.
Figs. 1 and 2 respectively display the geographic distribution of main
fossiliferous areas and stratigraphic relationships among units.
2.1. Danian
Fossiliferous exposures of Danian age are widely spread in Patagonia
and belong to the Cerro Dorotea, Roca, Lefipán, and Salamanca forma­
tions. Main fossiliferous sites of the Cerro Dorotea Formation are
exposed in the area of Río Turbio, between Cerro Dorotea and Estancia
San Jorge (Austral Basin).
A well-diversified, albeit poorly preserved fauna is contained in the
Roca Formation exposed at General Roca and Cerros Bayos (Colorado-
Neuquina Basin) and in the area between Los Menucos and Ingeniero
Jacobacci (Colitoro Plateau; Somuncurá-Asfalto Basin).
The Lefipán Formation is exposed in the middle valley of the Chubut
C. Julia del Río
River (Somuncurá-Asfalto Basin), yielding abundant molluscs in the
uppermost strata. Its main fossiliferous sites are located between Piedra
Parada and Paso del Sapo.
Poorly fossiliferous exposures of the Salamanca Formation are found
in the surroundings of Sierra de los Guanacos (Somuncurá-Asfalto
Basin), but the richest and best preserved molluscan faunas of this unit
correspond to those exposed from Puerto Visser northwards to Bahía
Malaspina (Atlantic coast), and to the confluence of the Chico and
Chubut rivers (between Puesto Alvarez and Cañadón Iglesias) (Golfo San
Jorge Basin).
For specific fossiliferous sites, descriptions of available lithological
sections and stratigraphic provenance of the Danian assemblage see
Celeste (1949), Masiuk (1967), Medina et al. (1990), Griffin and
Hünicken (1994), del Río et al. (2007, 2011), del Río (2012), and Scasso
et al. (2012).
Based upon calcareous nannofossils, foraminifera, and palynomorph
content, the Roca Formation is considered of earliest Danian age (Con­
cheyro and Nañez, 1994; Papu et al., 1999; del Río et al., 2011; del Río
and Martínez, 2015, and bibliography therein). Recently the Cerro
Dorotea Formation has been dated between 61.9 ± 0.3 Ma and 60.2 ±
1.3 Ma (Danian-Selandian) (Fosdick et al., 2019).
According to molluscan correlation, Paleocene faunas of the Austral,
Somuncurá-Asfalto and Golfo San Jorge basins were placed in the early
Danian by del Río and Martínez (op.cit.).
2.2. Late middle Eocene
Exposures of this age are much more geographically restricted than
the Danian ones and are confined to the southern sector of the Austral
Basin, where a low diverse assemblage is contained in the Leticia, Río
Turbio and Man Aike formations.
Fossiliferous strata of the Leticia Formation are exposed north of
Lago Fagnano, in the surroundings of Tolhuin (Tierra del Fuego), and
the scarse material included herein comes from Estancia Arroyo (for
placement of specific localities see Buatois and Camacho, 1993).
Most of the fossil material of the Río Turbio Formation considered
herein comes from exposures between Yacimiento Río Turbio and
Cancha Carreras, and was collected by M. Hünicken and described by
Griffin (1991), as well as molluscs collected by H. Camacho in the sur­
roundings of Estancia La Primavera. Fossiliferous sites of the Man Aike
Formation are exposed in the area placed between Lago Argentino in the
west, the Santa Cruz River to the south, and the Shehuen River to the
north. Camacho et al. (2000a, b) and Casadío et al. (2009) display
lithological sections.
The Arroyo Verde Formation exposed at Arroyo Verde (boundary
between Río Negro and Chubut Provinces) is also considered of Eocene
age but its fauna has been poorly known since the original description
made by Rossi de García (1959), Rossi de García and Levy (1977), and
Ravazzoli et al. (1982) Because of this, it was not included in the present
analysis.
According to stratigraphic relationships and molluscan content,
Camacho et al. (1998; 2000a) placed the Man Aike Formation in the
middle Eocene, and Malumián (1999), based on its foraminifera content,
in the latest middle Eocene. Casadío et al. (2009) obtained discrepant
87
Sr/86Sr ages from a single osyter specimen, ranging between 35.2 Ma
and 54.82 Ma, but in coincidence with Malumián (1999), placed it in the
late middle Eocene age. Recently, Fosdick et al. (2019) obtained U–Pb
ages that constrain the unit between 45.7 Ma and 40.5 Ma.
The Río Turbio Formation has been dated by Fosdick et al. (op.cit.)
who proposed a U–Pb age ranging between 47 Ma and 41 Ma (middle
Eocene) for the lower portion of the unit and 36 Ma – 26 Ma (late
Eocene-Oligocene) for the upper part which could correspond to the Río
Guillermo Formation. This does not agree with the age inferred from the
molluscan fauna which do not show any difference between those
yielded in the lower and upper portions of the unit, being completely
different from the Oligocene molluscs (San Julián Formation). For this
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Journal of South American Earth Sciences 108 (2021) 103209
reason, this assemblage is considered herein as late middle Eocene in
age.
The Leticia Formation was considered of late middle Eocene age
upon its foraminifera content and stratrigraphic relationships, and
correlated with the Man Aike and Río Turbio formations (Olivero and
Malumián, 1999) and later placed between 41.9 ± 0.71 Ma and 39.6 ±
0.82 Ma by Olivero et al. (2020).
2.3. Latest Oligocene-early Miocene
Main fossiliferous exposures of latest Oligocene or early Miocene age
are distributed along the coast of the Atlantic Ocean from San Sebastián
(Carmen Silva Formation, Tierra del Fuego) northwards to Puerto Santa
Cruz and Puerto San Julián (San Julián and Monte León Formations)
(Austral basin), reaching the surroundings of Comodoro Rivadavia
(lower Chenque Formation, Golfo San Jorge Basin; San Julián Forma­
tion, Mazarredo Sub-basin). In the western region of the Santa Cruz
Province (Austral Basin) they are exposed along a wide strip that
stretches along the Andean foothills between Lago Argentino and Lago
Buenos Aires (Estancia 25 de Mayo and El Chacay formations).
Latest Oligocene deposits are restricted to the San Julián Formation
exposed in the Bajo de San Julián and surroundings of Puerto San Julián
(Austral Basin), where they have been dated between 25.32 Ma and 23
Ma (Parras et al., 2012).
In reference to early Miocene marine rocks, numerical age obtained
by 87Sr/86Sr for the Monte León Formation, exposed in the type area
(mouth of the Santa Cruz River), ranges from 22.58 Ma to 21.68 Ma at
the base, and from 18.9 Ma to 17.51 Ma at the top (Parras et al., 2012).
In the Estancia 25 de Mayo Formation ages range between 20.05 Ma and
19.1 Ma (Cuitiño et al., 2012) while, the deposits of the El Chacay
Formation span the 20.31 Ma-17.87 Ma interval (Cuitiño et al., 2015a).
The lower section of the Chenque Formation embraces the 19.15 Ma –
15.37 Ma interval (Cuitiño et al., 2015b).
There is no numerical age for the Carmen Silva Formation and it has
been initially placed in the early Miocene-middle Miocene (Codignotto
and Malumián, 1981) and later considered of middle Miocene (Olivero
and Malumián, 2008; Malumián and Nañez, 2011), mainly based on its
foraminiferological content. del Río and Martínez (2006) placed it in the
early Miocene based on molluscan fauna and recently Torres Carbonell
and Olivero (2019) also considered the unit of early Miocene age
(Burdigalian). An early Mioccene age is supported in the present paper
in the light of the molluscan assemblage yielded by that unit.
del Río (2004a) proposed the presence of four informal Molluscan
Zones based on species content, three of them are recognized in the
Eastern sector of the Austral Basin: the PP Assemblage (late Oligocene),
and the early Miocene PA and RSP Assemblages; the fourth one, the JR
Assemblage, is identified in the Golfo San Jorge Basin. All mentioned
papers, and those of del Río and Camacho (1998), Chiesa and Camacho
(1995), del Río (2004a), and del Río and Martínez (2006), include de­
scriptions of lithostratigraphic sections and detailed placement of
fossiliferous localities.
2.4. Middle Miocene
Highly fossiliferous deposits of this age are recognized in the Golfo
San Jorge Basin in the environs of Comodoro Rivadavia - Monte Her­
moso Valley (upper part of the Chenque Formation), along a narrow
littoral strip from Camarones 120 km northwards to Cañadón Isla
Escondida (Camarones Formation), in the Gaiman Formation exposed
from Gaiman-Trelew area northwards to Cueva de los Leones (65 km to
the north of Peninsula Valdés), and in the Bajo and Salinas del Gualicho,
where exposures of the lower and middle sections of the Gran Bajo del
Gualicho Formation (middle Miocene) contain a rich molluscan
assemblage .
del Río (2004a) proposed the NVG Molluscan assemblage to include
the fauna yielded by the units mentioned above. Based on stratigraphic
C. Julia del Río
relationships and molluscan content it was firstly restricted to the latest
early Miocene - earliest middle Miocene (del Río, op. cit), partially in
coincidence with the relative ages provided by the associated palyno­
logical assemblage in the Chenque Formation (Barreda and Pala­
marczuk, 2000). New field research carried out by the author, allowed a
more accurate placement of the NVG Assemblage in the Parasequence 4
and 5 of Bellosi (1990), and probably in the uppermost horizons of
Parasequence 3 of the Chenque Formation. Although there are no nu­
merical datings of Parasequences 4 and 5, they are younger than the
87
Sr/86Sr age of 15.37 Ma estimated for Parasequence 3 (Cuitiño et al.,
2015b), and older than the beginning of the late Miocene transgression
dated in the 11.4 Ma at the base of the Puerto Madryn Formation that
overlays the Gaiman Formation at Cueva de los Leones (del Río et al.,
2018). Thus, the NVG assemblage covers the middle Miocene interval
and that is the age accepted herein for all the correlated units yielding
this association.
2.5. Late Miocene
The late Miocene transgression extended from northern Patagonia
northwards to the Uruguay and Brazil border. In the Patagonian region,
marine rocks of this age are continuously exposed along a coastal area
streching from Trelew and Puerto Madryn cities and Peninsula Valdés
(Puerto Madryn Formation) northwards to the San Matías Gulf (Bar­
ranca Final Formation and Facies La Lobería of the Río Negro Forma­
tion) and Bajo del Gualicho (uppermost strata of the Gran Bajo del
Gualicho Formation). These beds are highly fossiliferous and the
Aequipecten paranensis Zone was recognized (del Río, 1988). Recent
numerical datings in Patagonia demonstrated that the late Miocene
marine transgression spanned the 11.4 Ma (in the south) - 6,68 Ma (in
the north) interval (del Río et al., 2018).
For lithological sections and stratigraphic provenance of faunas see
del Río (1992, 1994), Reichler (2010), and del Río et al. (2018).
3. Paleontological analysis
The origin and relationships of Patagonian molluscan faunas were
matter of discussion since the middle of the 19th century. Danian as­
semblages were firstly related with New Zealand ones (Zittel, 1864), and
later considered to be highly endemic, partially related to Chilean ones,
and having given rise to younger Patagonian faunas (Feruglio, 1949). In
reference to younger assemblages, Ihering (1907) compared the latest
Oligocene-early Miocene ones with New Zeland and European faunas,
stating that the association now considered of middle Miocene age, had
an important “Antillean” component. In turn, Feruglio (op.cit.), consid­
ered that since Cretaceous times our faunas were completely differen­
tiated from Boreal assemblages, and that during the Miocene, they
maintained strong links with Chilean and Antarctic faunas. By that time,
they hardly resembled those of New Zeland and Australia, had even less
similitude with Indo-European faunas, and were not alike the North
American ones.
Subsequent studies showed that relationships with worldwide coe­
taneous assemblages are not so linear and simple as claimed by previous
authors, and the concept of elements of different origin as constitutents
of the Patagonian faunas was introduced. In this way, terms such as
Paleoaustral (Zinsmeister, 1982; del Río, 1990; Griffin and Hünicken,
1994), Tethyan (Camacho, 1992; Casadío, 1998), Caribbean (del Río,
1990) and Cenozoic Dispersers (Beu et al., 1997) taxa were employed in
the Argentinean paleontological literature, showing the existence of a
complex compositional pattern for our faunas.
3.1. Material and methods
The database used in the analysis performed herein includes 494
species and whenever possible, updated taxonomic assignments were
employed (Supplementary Data S1). Stratigraphic distribution for the
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Journal of South American Earth Sciences 108 (2021) 103209
225 recognized genera is displayed in Table 1. This table does not
include those taxa (with quotation marks in Supplementary Data S1)
only known from a single poorly preserved specimens with no visible
diagnostic characters, that precluded a proper taxonomic assignment, or
those species represented by specimens lost from collections and only
known from illustrations.
The author is aware that there are at least two large biostratigraphic
indicators (turritellids and buccinids) that are poorly studied, but
because they are abundant, they were included anyway in the analysis.
Material analized in the present paper includes the type collections of
J. Hatcher (studied by Ortmann in 1902), C. Ameghino (described by
Ihering in 1907), O. Nordenksjöld (analyzed by Steinmann and Wilckens
in 1908), and E. Feruglio (published by himself in 1936), and collections
assembled during last decades by the author. Repositories of studied
material are as follows (only acronyms of repository that houses mate­
rial illustrated is included):
- MGGC - Museo Geológico Giovanni Capellini (Bologna, Italy, Fer­
uglio collection);
- PRI - Paleontological Research Institution (Ithaca, USA; Hatcher
collection);
- MACN-Pi- Museo Argentino de Ciencias Naturales (Buenos Aires,
Argentina; Ameghino, Wichmann, Hünicken, Feruglio, Camacho,
Olivero, del Río collections);
- CPBA - Cátedra de Paleontología de la Universidad de Buenos Aires
(Buenos Aires, Argentina; Camacho, del Río collections);
- Naturhistoriska riksmuseet (Stockholm, Sweden) (Nordenskjold
collection); - Museo La Plata (La Plata, Argentina, Nordenskjold
collection)
- Museo Paleontológico de Bariloche (Río Negro, Argentina, Pose,
Smekal collections)
- Museo Paleontológico E. Feruglio (Trelew, Argentina, Brunet
collection)
- Natural History Museum (London, United Kingdom, Darwin
collection)
- Museo de Historia Natural de Santiago (Santiago, Chile, Philippi
collection) Santiago, Chile.
- Muséum national d’histoire naturelle (Paris, France, d’Orbigny
collection)
For our analysis, and according to present knowledge on temporal
and spatial distribution of the Patagonian taxa mentioned herein, the
following molluscan biogeographic categories are defined (Ca, SUS, T,
Pa, NS, A, E, as identified in Table 1):
- Caribbean elements (Ca): the term “Caribbean” refers to warm-taxa
inhabiting the Miocene Gatunian Bioprovince of Petuch (1982) and
Landau et al. (2008), developed in the Caribbean Basin, a unit
including areas of both Pacific and Atlantic Oceans. Some of these
elements could have had their origin in the late Oligocene and as
stated by Petuch (1988), some of them are West Tethyan faunal el­
ements that managed to survive in America.
- Southern United States elements (SUS): this category includes
members of theMaastrichtian-Eocene molluscan assemblages
restricted to the Gulf Coastal Plain and to the Pacific Slope of North
America (the Gulf Coastal Plain elements of del Río and Martínez,
2015, are now included in this category). As these areas were con­
nected with the Atlantic-Tethys Ocean sice the Caribbean Seaway
opened in the Oxfordian, (except for a short interval in the Late
Cretaceous when this passage was closed; Iturralde-Vinent, 2003),
faunal exchange was possible between both regions and some ele­
ments show strong tethyan affinities (Givens, 1989).
- Tethyan elements (T): these are the warm-water genera originated
and distributed in the Tethyan Realm during the Cretaceous-
Paleocene, also including those that appeared in the Oligocene-
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Table 1
Stratigraphic distribution of the molluscan genera and their distribution in the Recent Molluscan Bioprovince of the South Atlantic
Ocean. (R.BP. = Recent Bioprovinces; † = extinct genus; ** cosmopolitan genus restricted to tropical or warmest subtropical regions;/ =
genus present in the northern sector of the biogeographic unit). E = endemic; A = American; C= Cosmopolitan; Ca= Caribbean; SUS=
Southern United States; Pa= Paleoaustral; T = Tethyan; P= Pacific; NS= Neogene Southern; I–P= Indo-Pacific.

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7
C. Julia del Río
Neogene Paratethyan Sea (Steininger and Wesley, 2000; some of
them having survived into Recent times in the Indo-Pacific region).
- Paleoaustral elements (Pa): this term refers to those taxa originated
in the Maastrichtian Weddellian Province (see discussion in del Río
and Martínez, 2015) and that persisted in the Southern Hemisphere
continents during part of the Cenozoic, being especially abundant in
the Patagonian Dorotean, Rocaguelian and Salamancan Danian
bioprovinces defined by del Río and Martínez (op.cit.) where they
inhabited warm-temperate waters
- Neogene Southern elements (NS): genera that during the Neogene
were distributed across high and mid latitudes of the southern
Atlantic, Indian and Pacific Oceans. Most of them survived into
Recent times in circumpolar cool-temperate waters but were origi­
nally associated to subtropical faunas (see Section 5- Results and
Discussion). Although highly represented during the Neogene, it
seems that a few taxa would have appeared in the middle Eocene of
Patagonia. The NS is a new name for the Cenozoic Dispersers taxa of
Beu et al. (1997). Those authors coined the term to referto
post-Eocene molluscs with a southern circumpolar distribution that
would have been dispersed through the Circumpolar Antarctic Cur­
rent from New Zealand to South America as well as in the opposite
direction. This dispersal route was discussed by Casadío et al. (2010),
who claimed a migration through the West Antarctic Rift system was
also possible. I propose the use of the neutral term NS since it is
devoid of any reference to their origin, or a dispersal mechanism, ie.
the cold CCA, a rafting subtropical taxa, or through the West Ant­
arctic Rift system, or a vicariance process.
8
Journal of South American Earth Sciences 108 (2021) 103209
- American elements (A): genera that since Neogene times remained
restricted to both margins of the American continent.
- Endemic elements (E): taxa originated in southern South America
(Chile, Argentina)where they remained restricted until their extinc­
tion or survival into Recent Magellanic and Argentinean Provinces
and in the Subantarctic region.
- Pacific and Indopacific elements (P/I–P): these categories refer to
extremely uncommonly represented groups in Patagonia that during
the Paleogene-Neogene were only known from the North Pacific or
Indo-Pacific regions.
When dealing with paleoclimatic inferences, tropical waters are
those with temperatures above 25 ◦ C; subtropical between 20 ◦ C − 25 ◦ C;
warm-temperate between 20 ◦ C − 15 ◦ C; cool-temperate between 15 ◦ C
− 10 ◦ C, and subantarctic waters between 6 ◦ C − 10 ◦ C.
Regarding the composition of the fauna the term “tropical” refers to
taxa distributed in Circum-tropical and warmest Circum-subtropical
areas, i.e. Tethyan taxa and the related Paleogene SUS elements,
Caribbean genera, to those taxa restricted to the tropical region of
America since late Oligocene times, and to the Cosmopolitan taxa
restricted to tropical and warmest subtropical latitudes. Paleoaustral
elements were considered by del Río and Martínez (2015 and discussion
herein)) as warm-temperate taxa during Danian times, This assestion is
supported by the presence of Neogene Southern elements in the sub­
tropical late Oligocene – early Miocene of New Zealand (Beu and
Maxwell, 1990) associated to Paleoaustral taxa, which should make us
to consider them as subtropical elements. By the way, Woelders et al.
(2017) obtained isotopic paleotemperatures for northern Patagonia
C. Julia del Río
Fig. 3. A- Paleogeography during Danian times showing the distribution of main components of the fauna (paleogeographic map after Scotese, 2014a); B- His­
trograms of percentage of molluscan biogeographic categories.
during the Danian (see Discussion) pointing to the existence of sub­
tropical conditions in the area and thus sustaining the climatic infor­
mation provided by the molluscs. In this way, Paleoaustral taxa are
considered to represent warm-temperate to subtropical conditions in the
Paleogene of Patagonia.
3.2. Composition of the Molluscan Assemblages
Analysis of the composition of the Paleogene-Neogene molluscan
fauna of Patagonia allowed the identification of five assemblages of
Danian, late middle Eocene, latest Oligocene-early Miocene, middle
Miocene, and late Miocene ages.
Another assemblage was recognized since the beginning of 20th
century as probably belonging to the Pliocene (Feruglio 1933, 1954),
and recent numerical 87Sr/86Sr datings on pectinid valves, confirmed a
Zanclean age (4,89 Ma- 5,33 Ma) (del Río et al., 2013). Its fauna, clearly
different from the older ones, was studied by Feruglio (1954), and
except for a few taxa recently reviewed by Santelli et al. (2019a), Pérez
and del Río (2017a), and Alvarez et al. (2020) (i.e. Moirechlamys, Sca­
laricardita, Kolmeris, Carditella, and Eucallista), most of the fauna still
remains unknown, and for this reason it will not be included in the
present analysis.
Composition of each assemblage is depicted in Figs. 3, 5, 7, 10 and
12, and their commonest representants are illustrated in Figs. 4, 6, 8-9,
11, 13-14.
3.2.1. Danian Assemblage
Pioneer studies of Danian molluscs were carried out by Burckhardt
(1901), Ihering (1903), Feruglio (1936), and Petersen (1946). Modern
9
Journal of South American Earth Sciences 108 (2021) 103209
analyses include those of Griffin and Hünicken (1994), Casadío (1998),
Griffin et al. (2005), del Río et al. (2007), and del Río (2012). The
assemblage considered incorporates 35 genera including 50 species of
gastropods and 31 genera including 47 species of bivalves. The number
of Maastrichtian survivors is considerably high, comprising 33% of the
assemblage; they are represented by the bivalves Plicatula, Glycymeris,
Lahillia, Cuccullaea, Gregariella, Neilo, Australoneilo, Phygraea, Cubitostrea
and Turkostrea, and by the gastropods Struthioptera, Austroaporrhais,
Pseudotylostoma, Taioma, Fyfea, Priscaphander, Darwinices, Spirogalerus,
Austrophaera and Heteroterma.
Main components are Cosmopolitan (41.4%) and Paleoaustral
(29.3%) taxa, which along with Tethyan (10.5%) and SUS (8.5%)
genera, characterize the Danian fauna (Figs. 3–4).
Endemic genera are 10.4% of the fauna and are represented by
Austrophaera, Rocalaria, Pseudotylostoma and oysters, among others.
Oysters constitute the dominant molluscan group throughout almost all
basins. The genera Turkostrea and Cubistostrea have their oldest occur­
rence in the Maastrichtian of Patagonia where they survived into the
Danian (Griffin et al., 2005), and therefore should be considered as
Endemic elements during this interval. Both taxa are absent from the
middle Eocene assemblage, when they became Cosmopolitan genera
widely distributed in the Northern Hemisphere. As noted by Griffin et al.
(op.cit.), Cubitostrea re-appeared in Patagonia in the middle Miocene and
became extinct in the region by the end of the late Miocene.
Many Paleoaustral genera survived the K/Pg boundary when the
Cretaceous Weddellian Province disintegrated and became a charac­
teristic components of the Patagonian Danian fauna, being represented
by Cuccullaea, Lahillia, Neilo, Spirogalerus, Darwinices, Austroaporrhais,
Struthioptera, Pseudofax, Fyfea, Globisinum, Microfulgur, Zemacies and
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 4. Danian Assemblage-A- Pseudotylostoma romeroi Ihering (1903), General Roca MACN-Pi 693; B- "Nucula" pervicax Feruglio (1936), Holotype MACN- PI 5463,
Puesto Ramírez, Salamanca Formation; C- Neilo sp. MACN-Pi 5223, General Roca, Roca Formation; D- "Nucula" suboblonga (Wilckens, 1907), MACN-Pi 5215, General
Roca, Roca Formation; E-F- Pycnodonte (Phygraea) burckhardti (Böhm, 1903) MACN-Pi 5219, General Roca; G-H- Bathytormus chubutensis (Feruglio, 1936), Lectotype
MACN-Pi 5369, Puesto Ramírez, Salamanca Formation; I-J- Cubistostrea ameghinoi (Ihering, 1902) MACN-Pi 167, General Roca, Roca Formation; K-L - Sulcobuccinum
prominentis del Río (2012), MACN-Pi 5257 Puesto Carmelo Ibañez, Roca Formation; M- Claibornicardia paleopatagonica Feruglio, 1936, MACN-Pi 5197, Puesto
Ramírez, Salamanca Formation; N-Ñ- Heteroterma carmeloi del Río (2012), MACN-Pi 5250, Puesto Intermedio, Roca Formation; O–P- Gryphaeostrea callophylla
(Ihering, 1903) MACN-Pi 138, Roca Formation; Q-R- Rocalaria alani del Río (2012), Puesto Carmelo Ibañez, Roca Formation, Q- MACN-Pi 5307, R- MACN-Pi 5306; S-
Austrophaera patagonica Feruglio, Holotipo MACN-Pi 5281, Puesto Ramírez, Salamanca Formation. (All specimens x 1).

10
C. Julia del Río
Fig. 5. – A- Paleogeography during the middle Eocene showing the distribution of components of the fauna (paleogeographic map after Scotese, 2014a); B- His­
trograms of percentage of molluscan biogeographic categories.
Priscaphander (del Río and Martínez, 2015). Among Paleoaustral taxa it
should be included a particular group of large nuculid species distrib­
uted since the Late Cretaceous in the Southern Hemisphere. These are
“Nucula” hünickeni Zinsmeister and Macellari (1988) (Danian,
Antarctica), “Nucula” guillermensis Griffin, 1991 (Eocene, Patagonia),
“Nucula” frenguellii Feruglio (1936) and “Nucula” pervicax Feruglio
(1936) (Danian, Patagonia), “Nucula” grandis Malumián, Camacho and
Gorroño, 1978 (early Miocene, Patagonia) and “Nucula” suboblonga
Wilckens, 1907 (Cretaceous of Patagonia, New Caledonia and New
Zealand; Cretaceous - Danian of Antarctica). This group would have
managed to survive into modern times being possibly represented now
in Antarctic and Subantarctic regions by “Nucula” grayi, “N’. georgiana,
and “N.” livingstonensis.
Tethyan elements are represented by Bathytormus, Acesta (Plica­
cesta), and by the common oysters Gryphaostrea and Phygraea. Gry­
phaostrea has its oldest occurrence in the lower Cretaceous of the
Tethyan region and a wide distribution in the Northern Hemisphere
during the Paleogene. Phygraea, a Cosmopolitan genus in the Northern
Hemisphere since the Cretaceous, shows strong Tethyan affinities and is
recorded in the Maastrichtian-Danian of Patagonia and in the Maas­
trichtian of Antarctica (Zinsmeister and Macellari, 1988; Casadío,
1998).
The carditids Kaleia, Cardites and Calibornicardia could be also
considered as Tethyan elements, but this point deserves further discus­
sion. The three genera have been recently described by Pérez and del Río
(2017a) for the Danian of Patagonia, where they likely have their oldest
occurrence. Kaleia and Cardites were later recorded in the late Paleocene
and in the Miocene of Europe respectively, and Claibornicardia in the
middle Eocene of Europe and of the Gulf Coastal Plain of USA. At this
moment, and until European records are revised to assess a more
11
Journal of South American Earth Sciences 108 (2021) 103209
accurate temporal distribution, those genera may be considered as ele­
ments with Tethyan affinities.
Finally, the SUS elements are represented by Yoldia (Calorhadia),
Tejonia, Cidarina and Heteroterma, that along with Sulcobuccinum, Phy­
graea, Priscoficus, Claibornicardia and Rotunidicardia constitute common
components of the Paleogene assemblages of the southern United States
of America.
The Paleoaustral, Tethyan and SUS taxa provide the fauna with a
distinct signature differentiating it from the remaining Southern Hemi­
sphere coetaneous faunas, leading to its separation from the Weddellian
Bioprovince, a unit that did not reach into Patagonian regions during the
Danian. Latitudinal variation in the proportion of these elements
allowed the identification from south to north of the Dorotean, Sala­
mancan, and Rocaguelian Bioprovinces (del Río and Martínez, 2015).
3.2.2. Late middle Eocene Assemblage
It is one of the less diverse Patagonian faunas and is represented by
55 bivalve species and 34 bivalve genera and 14 gastropod species
belonging to 12 genera. Paleontologic studies are scarce, and include
descriptions by Feruglio (1936), Furque and Camacho (1949), Camacho
(1957), Griffin (1991), Buatois and Camacho (1993), Camacho et al.
(2000a,b), and Casadío et al. (2009). Almost 50% of Danian genera
became extinct, among these there were many Paleoaustral, Cosmo­
politan and Tethyan elements, and although those taxa continued
dominating the Eocene assemblage, there was a substantial decrease in
their participation compared with the previous interval. The most
typical and widely spread Danian taxa that became extinct are the
oysters, the carditids Claibornicardia, Cardites and Kaleia, the Paleo­
austral aporrhids Struthioptera and Austroaporrhais, the tudiclids Fyfea
and Microfulgur, and the Endemic genera Rocalaria and Pseudotylostoma.
C. Julia del Río
Fig. 6. Late middle Eocene Molluscan Assemblage- A- Neovenericor carrerensis Griffin, 1991,MACN-Pi (ex-CIRGEO) 128, Minas Río Turbio; B- Iheringinucula sp.
MACN-Pi 2336; C-D- Austrocallista sp., C- MACN –Pi 2305, MACN-Pi 6451; E- Ameghinomya sp. MACN-Pi 2305; F- Cucullaea MACN-Pi 2309.(All material from
Yacimiento Río Turbio, Río Turbio Formation, x1).
The late middle Eocene records the first occurrence of 23 taxa that
include mainly a new suite of endemic taxa that constitutes 22.5% of the
fauna, and that incorporates the genera Neovenericor, Ameghinomya,
Adelfia, Austrocallista, Valdesia, Fagnanoa, and probably the pectinid
Jorgechlamys. (Figs.5 and 6)
The proportion of the Paleoaustral taxa strongly decreased from
29.3% in the Danian to 17.5%, but were still important constitutens of
the fauna being represented by Australoneilo, Darwinices, Spirogalerus,
Lahillia, Cuccullaea, the “large nuculas”, the long-ranging genus Pter­
omyrtea, that will be recorded until the late Miocene, and the wide­
spread gastropod Perissodonta, a common taxa with a modern Southern
circumpolar distribution.
To the Danian SUS survivors (Heteroterma, Yoldia (Calorhadia), and
Tejonia), is now added the genus Clavipholas, making up 10% of the
fauna. The Tethyan taxa (10%) are marked by the appearance of Solena
(Eosolen) and Electrotoma. When dealing with genera of Tethyan affin­
ities one should mention the presence of Lopha hermini, a species from
the Man Aike Formation described by Feruglio (1936) and Casadío et al.
(2009), the generic assignment of which should be matter of a future
revision. The presence in Patagonia of Lopha-like genera can not be
12
Journal of South American Earth Sciences 108 (2021) 103209
explained at this time because the taxonomic validity of some of these
genera and their stratigraphic and geographic distribution are as yet
controversial. On one hand, Lopha s.str. is considered a monotypic genus
represented by its type species, the Recent Indo-Pacific L. cristagalli
(Harry, 1985; Huber, 2010) but at the same time it was recognized in the
Upper Cretaceous of the Western Interior in the United States (Ste­
phenson, 1956; Hook and Cobban, 2016). On the other hand, most
Cenozoic species formerly placed in this genus by Vokes (1977),
Woodring (1982), and Moore (1987), have been removed from Lopha s.
str. to other “Lopha-like” genera such Dendostrea and Myrakeenaor even
to Ostrea, based on phylogenetic results (Raith et al., 2015). Anyway, the
presence of a “Lopha –like” genus in Patagonia is relevant because it
points to an increment in the participation of tropical taxa with Tethyan
or Caribbean affinities in the region.
This interval records a minor participation of Indo-Pacific and Pacific
taxa (4.4%) and the first occurrence of Neogene Southern genera rep­
resented by Purpurocardia and Sigapatella, which constitutes 5% of the
fauna
C. Julia del Río
Fig. 7. Paleogeography during the late Oligocene-early Miocene showing the distribution of components of the fauna (paleogeographic map after Scotese, 2014b); B-
Histrograms of percentage of molluscan biogeographic categories.
3.2.3. Latest Oligocene-early Miocene Assemblage
Molluscs are abundant, well preserved, and along with the late
Miocene ones were amongst the first Argentine Cenozoic faunas to be
described (Sowerby, 1846; Ihering, 1897, 1899, 1907, 1914; Cossmann,
1899; Ortmann, 1902; Ihering, 1907; Steinmann and Wilckens, 1908).
Many modern taxonomic studies including early Miocene molluscs have
been already listed in the Introduction.
The latest Oligocene and early Miocene faunas are considered as one
assemblage because the latest Oligocene one does not include any genus
restricted to that interval and the first occurrences of 19 genera recorded
by this time extend their range into the early Miocene. It is the most
diversified assemblage of the Neogene of Patagonia and is represented
by 122 bivalve species (76 genera) and 116 gastropod species (75
genera). This interval attests one of the most dramatic Cenozoic faunal
turnovers in the Southwestern Atlantic Ocean, recording the first
occurrence of around 124 genera (20 in the latest Oligocene and 104 in
the early Miocene), among which 76 remained restricted to this
assemblage. (Figs 7, 8 and 9)
The presence of Paleoaustral, SUS and Tethyan genera abruptly
decreased from Paleogene times to 6.1%, 1.4% and 4.2% respectively.
Up to 21 late middle Eocene genera became extinct in the area (e.g. the
SUS elements Tejonia, Heteroterma, Yoldia (Calorhadia), Clavipholas, and
the Paleoaustral Spirogalerus, Darwinices and Australoneilo).
A high taxonomic diversification took place comprising the families
Pectinidae, Volutidae, Carditidae, Cardiidae, Muricidae and Turridae
and, to a lower extent Columbellidae, Skenidea, and Cancellaridae.
Pectinidae are now represented by new endemic genera: Pixiechlamys,
Reticulochlamys, Chokekenia and Zygochlamys, by the genus Swiftopecten,
a chlamydini with Pacific affinities, and by the NS Cyclochlamys, that
survived in the region into Recent times. Moreover, all Endemic volutids
(Miomelon, Adelomelon and Pachycymbiola) that now inhabit the
13
Journal of South American Earth Sciences 108 (2021) 103209
Southwestern Atlantic Ocean, appeared during this interval. Cardiids
are represented by short-live endemic genera now extinct in the area,
such as Iheringicardium, Patagonicardium and Hedecardium, and the
Recent American genus Trachycardium. Muricids are represented by
Urosalpinx, Xymene, Trophon, and a group of unstudied taxa related to
Crassilabrum and Ocenebra.
Only two Endemic taxa became extinct (Austrophaera and Adelphia),
and 21 genera arose in the region, increasing their proportion in the
assemblage to 21.2%. Apart from the pectinids and cardiids mentioned
above, the area witnessed the appearance of Iheringinucula, Osterheldia,
Neoinoceramus, Austroimbricaria, Neoimbricaria, Peonza and Struthioche­
nopus, all now extinct in the region, and the Endemic genera Proteopitar,
Retrotapes, Buccinanops and Antistreptus, all taxa still living in the
Southwestern Atlantic Ocean.
Tethyan elements are represented by Carolia (Parinomya), Limatula,
Cuccullaria and Darwinicardia, which together with the Eocene survivors
Solena (Eosolen) and Bathytormus make up 4.2% of the fauna.
Finally, a suite of Cosmopolitan genera from tropical and warmest
subtropical environments appeared in Patagonia (Barbatia, Crassatella,
Trachycardium, Ficus, Conus, Borsonia, Liotia, Retizafra, Sveltia, Scalptia,
Dalium), increasing the list of these elements from 27.5% in the late
middle Eocene, to 40.2% in this interval.
Undoubtedly, the most striking change of this fauna was the sudden
and massive occurrence of the Neogene Southern genera (NS), that
constituted the 24.6% of the fauna and provided the early Miocene
assemblage with certain similarities shared with the coeval New Zealand
fauna. Thirty five NS are recognized, some of which survived in the
Recent Southwestern Pacific Ocean (Austromitra, Zeacumina, Antizafra,
the Sassia zealta group [see Beu et al., 1997; Beu, 2010], the turrids
Antimelatoma and Fusiguraleus, the crassatellids Spisatella and Talabrica,
the mactrid Scalpomactra, and the genera Scaeoleda and Neopanis).
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 8. Latest Oligocene-early Miocene Assemblage- A- Patagonicardium philippi (Ihering, 1897) MACN-Pi 399, Yegua Quemada, Monte Léon Formation; B–C-
Miomelon gracilior (Ihering, 1896), MACN-Pi 889, Puerto Santa Cruz, Monte Léon Formation; D- Jorgechlamys centralis (Sowerby, 1846), MACN 269, Monte Espejo,
Monte León Formation; E- Trophon camachoi Griffin and Pastorino (2005), MACN-Pi 2182, El Unco, El Chacay Formation; F-G- Panopea nucleus Ihering (1899),
MACN-Pi 2727, Estancia Quien Sabe, Estancia 25 de Mayo Formation; H- Cucullaea alta (Sowerby, 1846), MACN 6484, Punta Guanacos, Monte León Formation; I-
Reticulochlamys borjasiensis (del Río, 2004b), Holotipo MACN-Pi 4641, Punta Borjas, Chenque Formation; J- Penion subrecta (Ihering, 1899) MACN-Pi 842, Puerto
Santa Cruz, Monte León Formation; K- Sassia bicegoi (Ihering, 1897), MACN-Pi 790, Puerto Santa Cruz, Monte León Formation; L- Glycymerita cuevensis (Ihering,
1897) MACN-Pi 5998, Las Cuevas, Monte León Formation; M- Pachycymbiola ameghinoi Ihering, 1896, MACN-Pi 6069, Las Cuevas, Monte León Formation; N-
Reticulochlamys zinsmeisteri (del Río, 2004b), MACN-Pi 4648, Monte Entrada, Monte León Formation; Ñ- Zygochlamys geminata (Sowerby, 1846), PRI 72678, Oven
Point, San Julián Formation (x1) (All specimens x 0.5).

14
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 9. Latest Oligocene-early Miocene Assemblage- A- Austrocallista iheringii (Cossmann, 1898), MACN-Pi 6391, Lago Cardiel, El Chacay Formation; B–C Neilo ornata
(Sowerby, 1846), MACN-Pi 6311, Punta Quilla, Monte León Formation; D- Valdesia collaris (Sowerby, 1846), MACN-Pi 1938, Estancia Primera Argentina, El Chacay
Formation; E- Cirsotrema rugulosa (Sowerby, 1846), MACN-Pi 622, Cabo Tres Puntas, San Julián Formation; F- Jorgechlamys juliana (Ihering, 1907), PRI 85900, Monte
Entrada, Monte León Formation; G- Swiftopecten iheringii del Río (1995), MACN-Pi 252, Yegua Quemada, Monte León Formation; H–I- Valdesia cuevensis (Ihering,
1897), MACN-Pi 2763, Sección 25 de Mayo. Estancia 25 de Mayo Formation; J-K- Zygochlamys jorgensis Ihering (1907), CPBA 14456, Bahía Solano, Chenque
Formation; L-M- Darwinicardia patagonica (Sowerby, 1846), L- MACN-pi 5769, Playa La Mina, San Julián Formation, M- MACN-Pi 362, Las Cuevas, Monte León
Formation (all specimens x1).

15
C. Julia del Río
Fig. 10. A- Paleogeography during the middle Miocene showing the distribution of components of the fauna (paleogeographic map after Scotese, 2014b); B-
Histrograms of percentage of molluscan biogeographic categories.
Others, such as Cyclochlamys, Lissarca, Puyseguria, Cyammomactra,
Eatoniella, Bulbus, Zeadmete, Cerithiopsilla, Kaitoa, and probably Austro­
toma, apparently interrupted their presence in the shallow waters of
Patagonia by the end of the early Miocene and resumed their occurrence
in the Recent of the Southern Circumpolar region. Others NS genera
became extinct in the region in the middle Miocene (Penion, Anti­
solarium) or in the late Miocene (Offadesma).
Pseuportlandia and Talabrica are considered herein as NS taxa, with
their oldest record in the Eocene of the Southern Hemisphere, appearing
in the Caribbean regions by the middle Miocene.
3.2.4. Middle Miocene Assemblage
This assemblage represents a second turnover of the Paleogene-
Neogene faunas, when almost 60% of the early Miocene genera
became extinct. It is much less diverse than the latest Oligocene-early
Miocene one, and is represented by 51 bivalve species (41 genera) and
25 gastropods species belonging to 28 genera. Thirtyfour genera are
shared with the latest Oligocene-early Miocene faunas, and 10 consist of
taxa restricted to the middle Miocene, among which there are the new
endemic short-live genera Argenthina and Ckaraosippur (Figs. 10 and 11).
The SUS elements fade away from the middle Miocene fauna, and the
Paleoaustral and Tethyan taxa kept on decreasing to 6.2% and 1.5% of
the fauna, respectively.
There were two events that provided the fauna with peculiar attri­
butes. On one hand, there was a drastic drop of the Neogene Southern
elements, from 24.6% in the latest Oligocene-early Miocene to 10.8% in
the middle Miocene. On the other hand, the first important occurrence of
Caribbean taxa (9.2%) is recorded represented by Amusium, Nodipecten,
Clementia, Torcula, Muracypraea, and Antinioche. This participation of
tropical genera in the region was reinforced by the presence of Pro­
fundimitra, Sconsia and the warm and warm-temperate waters genera
16
Journal of South American Earth Sciences 108 (2021) 103209
from the early Miocene (Sveltia, Ficus, Conus, Dalium).
3.2.5. Late Miocene Assemblage
After the pioneering taxonomic mentions of Sowerby (1846) and
Ihering (1907), followed the overall analyses of del Río (1992; 1994)
and Brunet (1995; 1997), and the revision of specific genera such as
Trophon, Offadesma, Madrynomya (Griffin and Pastorino, 2005, 2006a,
b), and the family Naticidae (Griffin and Pastorino, 2013). This fauna is
amongst the best studied, and its origin, composition and paleoclimatic
meaning was analyzed by del Río (1990), who recognized the presence
of Caribbean, Paleoaustral, Endemic, Cosmopolitan and Neoaustral el­
ements, stating that the late Miocene fauna (although considered of
middle Miocene age by then) had a tropical aspect like those of the
Recent Caribbean and Panamic Provinces. Martínez and del Río (2002)
proposed the Valdesian Bioprovinces restricted to northern Patagonia
-the focus of our analysis-, and the Paranaian Bioprovince, that extended
from Buenos Aires reaching northwards to the Uruguay and Brazil
border.
At the beginning of this interval, 64% of the middle Miocene genera
became extinct, including Tethyan, some Endemic, and almost all
Paleoaustral and Neogene Southern taxa. The fauna is represented by 47
bivalve species (41 genera) and 16 gastropod species (12 genera), and by
16 genera restricted to this interval. Cosmopolitan taxa constitute
46.1%, and Endemic elements 25% of the fauna. Among extinct genera
in the region are the Endemic pectinids Ckaraosippur, Zygochlamys and
Pixiechlamys, Austrocallista, Iheringinucula, Hedecardium, Austro­
imbricaria, Argenthina and the genus Neovenericor, that withdrew its
distribution northwards, where surived in the Paranaian Bioprovince.
Except for the genus Amusium, the Family Pectinidae went through a
striking extinction and no genus survived into the late Miocene. In turn,
the new Endemic pectinid Moirechlamys appeared, together with the
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 11. Middle Miocene Molluscan Assemblage- A-B- Pachycymbiola arriolensis del Río and Martínez (2006) MACN-Pi 4736, Puesto Arriola, Gran Bajo del Gualicho
Formation (x 0.5); C-D - Muracypraea posei (Figueiras, 1985), MACN-Pi 4811, Puesto Arriola, Gran Bajo del Gualicho Formation (x1); E-F- Nodipecten salis (del Río,
2006), MACN-Pi 1362, Gran Bajo del Gualicho Formation (x1); G- Clementia sp. CPBA-Pi (exCIRGEO) 1366, Gran Bajo del Gualicho Formation (x1); H- Glycymerita
camaronesia (Ihering, 1897), MACN-Pi 5170, Camarones, Camarones Formation (x0.5); I- Torcula hautali (Ihering, 1907) MACCN-Pi 731, Camarones, Camarones
Formation (x1); J- Neovenericor austroplata (Gardner and Bowles, 1939), MACN-Pi 1302 Puesto Astorga, Gran Bajo del Gualicho Formation (x0.5); K -Ckaraosippur
camachoi Santelli & del Río, 2019a,b, Holotype CPBA 8604, Puesto Salado, Formación Chenque, Puesto Salado (x1).

17
C. Julia del Río
Fig. 12. A- Paleogeography during the late Miocene showing the distribution of the fauna (paleogeographic map after Scotese, 2014b); B- Histrograms of percentage
of molluscan biogeographic categories.
species group related to “Aequipecten” paranensis, and also the Carib­
bean Leopecten (Figs 12, 13 and 14).
The Paleoaustral taxa were reduced to 2% of the assemblage, the
long-living genera that had characterized the Patagonian faunas (i.e.
Lahillia, Neilo, Cucullaea) became extinct, and the only Paleoaustral
survivor was Pteromyrtea. The Neogene Southern genera continued
decreasing in number (5.7%) and are represented by Offadesma, Pur­
purocardia and by “Cyclocardia” nortensis, a species related to the New
Zealand “Cyclocardia” awamoensis Group (del Río, 1986; 1994).
Typical middle Miocene taxa such as the tropical Ficus, Torcula,
Muracypraea, Sconsia, Profundimitra, Clementia, and Nodipecten were
replaced by a new Caribbean suite (7.7%) integrated by Chionopsis,
Lucinisca, Hexacorbula, Polymesoda, and Leopecten. Amusium and Anti­
nioche (although the latter taxon is not recorded in the Valdesian but in
the Paranaian Bioprovince), are the only middle Miocene Caribbean
survivors. Accompaying these genera are Cyrtopleura, Anomalocardia,
Crassinella and Dinocardium, taxa restricted since Neogene times to the
American continent. These elements increased their percentage from
3.1% in the middle Miocene to 9.6% in the late Miocene fauna.
3.3. Molluscs and paleoclimates
The study area is today placed between 54◦ 30′ S and 37◦ 40′ S,
mostly embracing the cool-temperate Magellanic Molluscan Province
(43o-57oS), with mean SSTs oscillating between 5 ◦ C in the south to 10
◦
C in the north, a transition zone (43o-41oS), and the warm-temperate
southern sector of the Argentinean Province (41o- 32oS), with SSTs be­
tween 10 ◦ C and 15 ◦ C (Huber, 2010).
From Danian times onwards there has been an increment in the
participation of Endemic and Cosmopolitan elements coupled with a
18
Journal of South American Earth Sciences 108 (2021) 103209
noticeable decline of Paleoaustral, SUS and Tethyan genera. During the
Danian the SUS, Tethyan and Paleoaustral taxa were significant com­
ponents and continued being relevant ingredients of the late middle
Eocene assemblage. Although the appearance the Neogene Southern
taxa had probably occurred in the middle Eocene, these elements typi­
fied the latest Oligocene-early Miocene interval, while the massive
occurrence of Caribbean taxa distinguished the middle and late Miocene
faunas.
The meaning of those faunal variations is linked to the changing sea
surface temperatures (SSTs) during the Palegene-Neogene interval. The
composition of the assemblages suggests the existence of warmer SSTs
than today in these latitudes, even more when taking into account that
during the Danian the Patagonian region was displaced 4◦ southwards
from its present position (Scotese, 2014a). Because of its northwards
Cenozoic drift towards its present position, cooler temperatures than
today should be expected.
In an overall cooling trend from the “greenhouse” conditions in the
Danian to the modern “icehouse” environment, the development of each
fauna is the response to transient global warming events superimposed
to the global cooling curve that defines the Cenozoic (Zachos et al.,
2001; Westerhold et al., 2020) (Fig. 15). The occurrence of Tethyan and
SUS elements of Tethyan affinities during Danian times alludes to the
development of high SSTs in the area, which is reinforced by the pres­
ence of tropical Cosmopolitan taxa. All those elements represent 27.6%
of the fauna, and their appearance in the Danian sea matches with the
hyperthermal Danian Dan-C2 and the Latest Danian (LDE) events. The
Dan-C2 event spanned the 65.24 Ma – 65.12 Ma interval, and implies a
warming event similar to that occurring in the Paleocene-Eocene
Thermal Maximum (PETM) (Quilliévéré et al., 2008), when SSTs oscil­
lated between 19 ◦ C and 27 ◦ C in the Southern Hemisphere (Hollis et al.,
C. Julia del Río
Fig. 13. Late Miocene Assemblage- A- Moirechlamys actinodes (Sowerby, 1846), CPBA 15158, Eje Tentativo; B- Glycymerita magna del Río (1992), MACN-Pi 5795,
Puerto Pirámides; C- Leopecten piramidesensis (Ihering, 1907), MACN –Pi 5735, Puerto Pirámides; D- Amusium paris del Río (1992), CPBA 15284, Eje Tentativo; E-F-
Dinocardium novus del Río (1992), CPBA 14597, Puerto Piramide.(All specimens from the Puerto Madryn Formation, x 0.5).
2012). These values agree with those documented by Woelders et al.
(2017), who proposed 24 ◦ C and 27 ◦ C for the K/P boundary and for the
earliest Danian, respectively, in northern Patagonia, when temperatures
were still increasing to reach the Dan-C2 event. The LDE was dated at
62.2 Ma and was cooler than the Dan-C2 event. While during the Dan-C2
event there was an increase of SSTs in high latitudesbetween 8 ◦ C and
10 ◦ C (Zachos et al., 2003), Jehle et al. (2019) estimated an increase of
2.6 ◦ C for the LDE in the Atlantic Ocean. Thus, the Danian fauna is now
considered warmer than previously thought (del Río and Martínez,
2015), with the development of tropical waters in the studied area. The
northwards increase in the SSTs proposed by those authors is reflected
by the higher proportion of the SUS and Tethyan elements in the
northern Rocaguelian Bioprovince than in the southern Dorotean
Bioprovinces.
The late middle Eocene fauna is geographically restricted to the
southernmost sector of the Austral Basin. It has a similar percentage of
tropical elements (31.3%) as the Danian one and its presence in the area
roughly responds to the Middle Eocene Climatic Optimum (MECO)
episode described by Bohaty and Zachos (2003) at 41.5 Ma in the
Southern Atlantic Ocean, when SSTs rose 4 ◦ C. This short-term warming
peak was more accurately precised in the 40.51 Ma – 40.21 Ma interval
(Rivero-Cuesta et al., 2019). The Danian SUS taxa survived into late
middle Eocene times and their presence in Patagonia reinforce the
persistence of warm or subtropical waters in the area. By the way, those
genera were components of the late middle Eocene warm-water fauna of
the Gosport Sand Formation (Gulf Coastal Plain), with temperatures
around 20.7 ◦ C ± 2 ◦ C (Haveles and Ivany, 2010). These values agree
with the global SSTs curve proposed by Hollis et al. (2012) and River­
o-Cuesta et al. (2019), with temperatures between 19 ◦ C - 22.5 ◦ C in the
19
Journal of South American Earth Sciences 108 (2021) 103209
Southern Hemisphere. In summary, Danian and late middle Eocene
faunas represent tropical and warmest subtropical waters, respectively
in Patagonia.
An almost uniterrupted sedimentary deposition is recorded during
Miocene times and the three Miocene assemblages identified offer an
opportunity to investigate the potential effects of climatic variations on
shallow marine communities.
The latest Oligocene-early Miocene assemblage stands for the first
faunal turnover in the Cenozoic, when an important loss of tropical taxa
and a massive appearance of subtropical genera would suggest cooling
seawaters in the region. The assemblage covers the 25.32 Ma – 17.51 Ma
interval in the southern Austral Basin, and the 19.15 Ma − 15.37 Ma
interval in the Golfo San Jorge Basin. After the climatic deterioration
during the Oligocene, there was an amelioration of climatic conditions
around 27 Ma and 24 Ma, leading to the late Oligocene warming peak,
the intensity of which, however, is discussed among researchers. Ac­
cording to Pekar and Christie-Blick (2008), pCO2 levels in the latest
Oligocene were near to those of present-day, Villa and Persico (2006)
proposed the existence of warm-temperate waters in the Kerguelen
Plateau, and Hartman et al. (2018) detected an increment of SSTs up to
25.1 ◦ C ± 5.2 ◦ C in Eastern Antarctica at 25.5 Ma, decreasing abruptly
towards the Mi glaciation.
During the early Miocene, abrupt and numerous alternating short
cooling (including minor glaciations) and warming events have been
ascertained (see Billups et al. 2002), but numerical values of tempera­
tures are almost unknown for this interval. In any case, the transient
warming episodes were slightly cooler than the late Oligocene warming
(Heydt and Dijkstra, 2006).
On one hand, our fauna shows a decline in the participation of
C. Julia del Río Journal of South American Earth Sciences 108 (2021) 103209

Fig. 14. Late Miocene Assemblage- A-B -Purpurocardia leonensis del Río (1986), MACN-Pi 6478, Cerro Prismático; C,H- Trophon leanzai Brunet (1997), MACN-Pi
4059, Puerto Piramides; D -Proteopitar mutabile (del Río and Martínez, 1998), MACN-Pi 6395, Puerto San José; E- Retrotapes ninfasiensis del Río (1997), MACN-Pi
6394, Cerro Prismático; F- Chionopsis australis del Río (1994), CPBA 14556, Puerto Piramide; G- Aequipecten paranensis (d’Orbigny, 1842), CPBA 11692, Puerto
Piramide. (All specimens from the Puerto Madryn Formation, x 1).

tropical and warmest subtropical taxa, that now are represented by
Barbatia, Crassatella, Trachycardium, Ficus, Conus, Borsonia, Liotia, Reti­
zafra, Sveltia, Scalptia, and Dalium, making up 15% of the assemblage.
On the other hand, the sharp boost of subtropical taxa (Neogene
Southern elements) happened during this interval and constitutes 24.6%
of the fauna. The decrease of tropical elements and their concurrence
with subtropical taxa could reveal the existence of lower SSTs than those
recorded in the late middle Eocene or middle Miocene intervals, but still
warmer than today in the Patagonian region.
To elucidate the degree of thermal resilience of the fauna or its cli­
matic response to brief cooling and warming events, an extremely ac­
curate stratigraphic provenance must be obtained, along with isotopic
and taphonomic analyses.
During the middle Miocene, the Neogene Southern elements
abruptly decreased to 10.8% of the fauna. The Tethyan elements
represent only 1.5% and the warm-taxa were replaced by the first
extensive manifestation of Caribbean genera, that increased from 2.5%
during the MECO to 9.2%. When taking into consideration the concur­
rence of Tethyan, Caribbean and tropical Cosmopolitan taxa, the per­
centage of warm-water genera increased to 23% of the fauna. The sea
surface temperatures suffered a new warming episode during the middle
Miocene Climatic Optimum (MMCO) at 17 Ma − 14.8 Ma (Sangiorgi
et al., 2018), oscillating between 19 ◦ C and 20 ◦ C in the Southern Pacific
Ocean, when the global mean temperatures were around 18.4 ◦ C (You
et al., 2009). This enhancing of the SSTs was probably the cause of the
massive occurrence of Caribbean taxa. One characteristic of the middle
Miocene molluscs is the development of unusually large and thick shells
when compared with the same species in the early Miocene. This is the
case of Retrotapes striatolamellata, Ameghinomya argentina, Austrocallista
iheringii, Cucullaea alta, Lahillia patagonica and Perissodonta ameghinoi. A
similar situation is seen in the early Miocene vs middle Miocene
congeneric species of the genera Glycymerita and Pachycymbiola which is
likely due to a faster growth in response to a higher primary production
in warmer waters than in early Miocene times (Jones et al., 1989;
Haveles and Ivany, 2010).
The late Miocene fauna studied herein comes from the Puerto
Madryn Formation exposed from Peninsula Valdés northwards to Cueva
de los Leones, where it has been dated between 11.4 Ma and 9.25 Ma
(del Río et al., 2018). By this interval, at least two new transient
hyperthermal peaks were recorded, one at ~11 Ma (Tortonian Thermal
Maximum of Westerhold et al., 2020), and the other one at ~ 9 Ma,
when temperatures of 23 ◦ C and 22 ◦ C were respectively estimated for
the southern seas placed between 30 oSL and 50 oSL, with a cooling peak
at 10 Ma of 19 ◦ C–20 ◦ C (Herbert et al., 2016). These temperatures

20
C. Julia del Río
Fig. 15. Paleogene-Neogene Molluscan Assemblages of Patagonia referred to
the average global marine temperature curve proposed by Westerhold et al.
(2020). Stratigraphic placement of the transient warming peaks in the global
cooling trend curve are according to the following authors: Dan-C2 (Quilliévéré
et al., 2008), LDE (Jehle et al. 2019); MECO (Rivero-Cuesta et al., 2019); latest
Oligocene warming peak (Westerhold et al., 2020); MMCO event (Sangiorgi
et al., 2018); Tortonian Thermal Maximum (Westerhold et al., 2020).
21
Journal of South American Earth Sciences 108 (2021) 103209
indicate that subtropical conditions subsisted through late Miocene
times, when 22.8% of Caribbean and Cosmopolitan tropical taxa were
recoded.
Martínez and del Río (2002) recognized an incipient thermal
gradient from the Valdesian to the Paranaian Bioprovinces. As temper­
atures were much warmer than today, it was concluded that the Cold
Falkland Current was not still fully operating in the area during the late
Miocene. In contrast to the proposal of Aguirre and Farinati (1999),
these paleobiogeographic units were not prefigurations of the present
Recent Argentinean and Magellanic provinces (Martínez and del Río, op.
cit.).
In summary, except for one numerical value of temperature for the
Danian of northern Patagonia (Wolders et al., 2017), no specific absolute
SSTs are known for Patagonian latitudes in the Southwestern Atlantic
Ocean. Anyway, as mentioned above, temperature values obtained for
southern latitudes indicate warmer temperatures than today, with
decreasing values during Paleogene-Neogene times in the region. On the
basis of the recorded warming peaks and on the porcentual participation
of warm or tropical-taxa in the five assemblages, we can estimate
tropical to warmest subtropical conditions in the Danian and late middle
Eocene Patagonian seas, warm-temperate waters for the latest
Oligocene-early Miocene fauna, and subtropical temperatures for the
middle and late Miocene assemblages, yet lower than those during
Paleogene times.
Of the 225 Paleogene-Neogene genera, 82 managed to survive in the
region (36.5% of total fauna) (Table 1). Thirteen taxa are Danian sur­
vivors, five come from the late middle Eocene, 51 from latest Oligocene-
early Miocene times, two from the middle Miocene, and 11 from the late
Miocene. Taxa that persisted into Recent times are mostly eurythermal
Cosmopolitan (52 genera), and to a fewer extent, survivors comprise 13
Neogene Southern taxa, 13 Endemic genera, three taxa restricted to the
American continent, and only one Paleoaustral genus.
The low Paleogene-Neogene heritage in the area is likely explained
because a high proportion of the fossil fauna included tropical and
subtropical taxa that became extinct in the region with the progressive
cooling of the waters from late Miocene onwards, withdrawing their
distribution to lower latitudes, where more suitable temperatures exis­
ted. In turn, subtropical Neogene Southern taxa had a different behavior,
as they almost dissapeared from the region once the SSTs increased at
the MMCO and re-appeared some time after the late Miocene. One
probable reason may be that the Neogene Southern taxa were unable to
adapt to increasing temperatures, being displaced to deeper environ­
ments. However, once cooler conditions were re-established after the
late Miocene, at least 13 genera shifted back to shallow waters, where
they managed to survive in the cool-temperate Magellanic Bioprovince
and Circumpolar region, becoming adapted to much cooler conditions
than those prevailing during early Miocene times.
The Paleogene-Neogene interval was characterized by high values of
endemism suggesting a tectonic stability in the region to allow the
development of endemic genera that survived into Recent times. In this
context, closely related co-generic species are represented, among
others, by the following species groups: Ameghinomya argentina (late
Oligocene – late Miocene) – Ameghinomya antiqua (Recent); Retrotapes
striatolamellata (late Oligocene) – Retrotapes ninfasiensis (late Miocene) -
Retrotapes exalbida (Recent); “Aequipecten” paranensis (late Miocene) –
“Aequipecten” tehuelchus (Recent); Anomalocardia entrerriana (late
Miocene) – Anomalocardia brasiliana (Recent); Trophon sowerby (late
Oligocene) – Trophon leanzai (late Miocene) – Trophon geversianus
(Recent) – Adelomelon cannada (early Miocene) – Adelomelon valdesiense
(late Miocene)- Adelomelon beckii (Recent); Pachycymbiola ameghinoi
(early Miocene) – Pachycymbiola brasiliana (Recent).
4. Conclusions
Five benthic molluscan assemblages are recognized as inhabiting the
shallow sea environments of Patagonia during Paleogene-Neogene
C. Julia del Río
times.
In an overall cooling trend during the Cenozoic, those faunas reflect
warmer sea surface temperatures than today in the region, agreeing with
superimposed transient warming events along the global curve of sea
temperatures. Cosmopolitan and Endemic taxa are dominant con­
stitutens of each assemblage.
The Danian and late middle Eocene faunas indicate the development
of tropical or warmest subtropical conditions in the region matching
with the Dan-C2 and Latest Danian events, and with the Middle Eocene
Cimatic Optimum. Apart from Cosmopolitan and Endemic genera, main
components were Paleoaustral, Southern United States and Tethyan
elements, and the total of warm or tropical-taxa reached 27.6% in the
Danianand and 31.3% in the late middle Eocene.
The latest Oligocene –early Miocene assemblage is characterized by a
drop of warm-taxa to 15.3% of the fauna, which was characterized by a
massive occurrrence of the subtropical Neogene Southern elements. The
SSTs would have corresponded to a warm-temperate ones, being higher
than in the Recent Magellanic Province, but lower than during previous
and subsequent Cenozoic intervals.
Middle and late Miocene assemblages were imprinted by the
appearance of Caribbean taxa, warm-water taxa increasing again since
the early Miocene to reach 23% of the fauna, suggesting the presence of
subtropical waters in the area in coincidence with the Middle Miocene
Climatic Optimum, the Tortonian Thermal Maximum hyperthermal
peak and with the warming event at 9 Ma.
Paleogene-Neogene heritage of the Magellanic and southern sector of
the Argentinean Province is rather low, only 36.5% of the fossil molluscs
managed to survive, and all tropical and most subtropical elements
withdrew their geographic distribution to lower latitudes with the
progressive cooling of the Patagonian sea since the late Miocene
onwards.
Most survivors are Endemic, Cosmopolitan and some Neogene
Southern genera that adapted to the Recent cool and temperate-cool
conditions in the region.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgement
I wish to express my special acknowledgement to my colleague and
friend S. Martínez by his critical reading of the MS.E. Olivero provided
help with the stratigraphy of Tierra del Fuego and one of the reviewers,
M. Griffin, has made very interesting comments and highly improved the
English language.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jsames.2021.103209.
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