Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125 – 132

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Salterella in the Argentine Precordillera: an Early Cambrian

palaeobiogeographic indicator of Laurentian affinity
Ricardo A. Astinia,*, William A. Thomasb, Ellis L. Yochelsonc
a
CONICET, Cátedra de Estratigrafı́a y Geologı́a Histórica, Universidad Nacional de Córdoba, Av. Velez Sarsfield 1611,
Córdoba X5016GCA, Argentina
b
Department of Gelogical Sciences, University of Kentucky, Lexington, KY 40506-0053, United States
c
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0121, United States
Received 31 October 2003; received in revised form 3 June 2004; accepted 6 July 2004

Abstract

This paper documents Salterella maccullochi (Murchison) in the Precordillera of western Argentina in the present-day
southern hemisphere. The record of these Early Cambrian small conical-shaped fossils, assigned to the extinct phylum Agmata,
provides further palaeobiogeographic evidence to consider the Argentine Precordillera an exotic terrane to Gondwana that
originally rifted from Laurentia. For palaeogeographic reconstructions, Salterella can be considered as an organism equally
important and biogeographically as meaningful as the olenellid trilobites due to their limited dispersal and demonstrated link to
Laurentian shallow-water inner-shelf facies. It has not been found anywhere in Gondwana. Salterella existed for only a very
short interval of time, but spread almost instantaneously around Laurentia. On this basis, the Early Cambrian of the
Precordillera terrane remains the more conclusive palaeobiogeographical link to Laurentia. Salterella provides another line of
evidence supporting palaeontological, sedimentologic, stratigraphic, palaeomagnetic, and isotopic evidence for a Laurentia
origin of the Precordillera terrane.
D 2004 Elsevier B.V. All rights reserved.

Keywords: Salterella; Early Cambrian; Paleobiogeography; Argentine Precordillera; Laurentia

1. Introduction cone-shaped fossil of the extinct phylum Agmata.

The Lower Cambrian beds of the Argentine
Precordillera (Fig. 1A) contain Salterella, a small,
* Corresponding author.
E-mail addresses: raastini@efn.uncor.edu (R.A. Astini)8
geowat@pop.uky.edu (W.A. Thomas)8
yochelson.ellis@nmnh.si.edu (E.L. Yochelson).
0031-0182/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2004.07.008
During the past decade, a wide range of geological
studies (e.g., Astini et al., 1995; Kay et al., 1996;
Thomas and Astini, 1996; Astini, 1998; Rapalini and
Astini, 1998; Benedetto et al., 1999; Keller, 1999;
Thomas et al., 2001) has produced increasingly strong
evidence that the Argentine Precordillera was part of
the Laurentian continent into Early Cambrian time.
However, a contradictory view based on preliminary
126 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132

Fig. 1. (A) Location map of the Argentine Precordillera in the present-day Andean region (abbreviations: PR, Precordillera; F, Famatina; WP,
Western Pampean Ranges; FC, Frontal Cordillera; PC, Principal Cordillera. Locations: 1, Ancaucha block; 2, Cerro Totora area; 3, Villicum
Range in the eastern Precordillera). (B) Time-stratigraphic chart for Cambrian and Ordovician across the Argentine Precordillera (abbreviations:
CTF, Cerro Totora Formation; LLF, La Laja Formation; ZF, Zonda Formation; LFF, La Flecha Formation; LSiF, La Silla Formation; SJF, San
Juan Formation; LSF, Los Sombreros Formation; modified from Astini, 1998). Ancaucha block (1) with Salterella shown in the stratigraphy of
western Precordillera.

analyses of detrital zircons (Finney et al., 2003) has
recently questioned the Laurentian origin of the
Argentine Precordillera (for a thorough discussion,
see Aceñolaza et al., 2002; Astini and Rapalini, 2003).
Cambrian-age platform carbonates overlying Gren-
ville-age basement rocks in the Precordillera first
invited direct comparisons to the basement and cover
of Laurentia. Salterella is locally abundant in Lower
Cambrian beds of North America and other places
considered original parts of Laurentia. Except for the
new collections from the Precordillera, Salterella is
unknown in the present southern hemisphere and has
not been recorded in any Gondwanan location. The
purpose of this paper is to highlight the importance of
Salterella as a palaeobiogeographic indicator strongly
linking the Argentine Precordillera to Laurentia.
2. Location of Salterella collections
The specimens described and illustrated herein are
from the western Precordillera, where an Ordovician
slope-facies mudstone (Los Sombreros Formation)
contains large olistoliths of the early Paleozoic plat-
form-carbonate strata, as well as older synrift strata and
basement rocks (Fig. 1B). Salterella is identified in
limestone beds within the Ancaucha olistolith (Fig. 2)
of Lower Cambrian synrift strata (Astini and Thomas,
1999; Astini et al., 2000). The Ancaucha block
(olistolith) is more than 500 m in length and includes
a stratigraphic succession nearly 80-m thick, similar to
that exposed at the base of the autochthonous succes-
sion (Cerro Totora Formation) to the east (Fig. 1B). The
lower part of the block contains a clastic succession of
maroon and red coarse sandstones and siltstones with
subaerial features (e.g., wrinkle marks, mud cracks,
flat-top wave ripples). The succession grades upward
into a mixed carbonate-clastic rhythmic interval, where
Salterella and fragmented trilobites are common
constituents of graded and massive thin-bedded pack-
stones interbedded with muddy background sediments
(Fig. 2). Bed geometries are both tabular and lenticular.
One previous short report (Bordonaro and Martos,
1985) identified Salterella from the lower member of
the La Laja Formation in the eastern Precordillera
(Fig. 1B). The lower member of the La Laja
Formation includes shaly beds that grade up to a
thick carbonate sequence, indicating evolution of a
passive margin across the Precordillera terrane
(Astini et al., 1995; Keller, 1999). The Salterella
 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132 127

further, except to note that small hyoliths may be

confused with Salterella, especially if the specimens
have been sorted and transported.

3. Description of fossils

Salterella maccullochi (Murchison) from the Pre-

cordillera comprises a relatively thick calcium carbo-
nate integument with an exterior conical shape,
having an angle of expansion near 108, so that shape
resembles a pencil point (Fig. 3). Under low
magnification, the outer shell wall shows two prom-
inent divisions (sometimes better seen in cross
section, e.g., upper center in Fig. 3). The outer layer
has several more compact laminations, and from a
point about two thirds of the distance from apex to
aperture, the laminated section thins toward the rim at
the apertural opening (see longitudinal section left
center in Fig. 3). The thicker inner division of the
cone wall appears more amorphous. From a point
about two fifths of the distance between apex and
aperture, the inner division thins toward the aperature,
and it does not extend beyond the point where the
Fig. 2. Stratigraphic columnar section of the Salterella-yielding
Early Cambrian olistolith within the Los Sombreros Formation in the
western Precordillera (see location in Fig. 1). Note the abundance of
shallow and very shallow water features and the mixed nature of
sedimentation. Salterella is usually a more abundant component in
the thin carbonate layers (modified from Astini et al., 2000).

specimens are associated with olenellid trilobites,

indicating an Early Cambrian age (Bonnia–Olenellus
Zone). Bordonaro and Martos (1985) described
noncurved, sharp-edged, small, cone-like fossils,
which they identified as Salterella sensu lato. The
specimens range from 8 to 12 mm length and from 2
to 4 mm diameter, and have apical angles between
148 and 248. Bordonaro and Martos (1985) compared
the smaller specimens with Salterella mexicana
Fig. 3. Thin section from the Salterella packstones in the Argentine
Lochman (1952) and the larger ones with Salterella Precordillera showing Salterella maccullochi (Murchison) in
pulchella Billings (1861); both are now considered various orientations, including oblique and transverse cross sec-
subjective synonyms of Salterella maccullochi tions. The central tube in longitudinal V-shaped sections and in
(Murchison, 1859). Hyolithids from the same out- circular cross section may be seen. To the left, a longitudinal section
crops (Bordonaro and Martos, 1985) have maximum shows the thinness of the outer layer at the apertural margin and the
gradual thinning of the inner deposits toward the apertural cavity.
dimensions of 24-mm length and 7-mm diameter. Near the bottom left corner, a cross section of two imbricated
The only illustration provided is hand-drawn. We (telescoped) specimens is evident. Curved trilobite fragments are
have not restudied their material and cannot comment also present (CEGH-UNC 20220).
128 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132
outer layer begins to thin. Secondary deposits within
the cone are seen as concentric circles in cross section
and as a contrasting pattern within the lower part of
the longitudinal section. Laminae of secondary
deposits are at an angle to the outer cone wall.
Secondary deposits vary in length along the cone and
appear to increase in thickness slightly faster than the
cone grows. A central tube is present, which in a
perfectly oriented longitudinal section bisects the
secondary deposits (Fig. 3).
All specimens recovered from the western Pre-
cordillera belong to the species Salterella maccullochi
(Murchison) (Fig. 3). The outer wall slightly exceeds
10 mm in length and 2 mm in thickness. Growth lines
on the exterior surface are represented by transverse
rings preserved only in isolated external molds on
shale. Laminae in the inner more compact cone are
seen, in thin section, to curve concave upward
converging toward the outer wall. Neomorphism
obscures details among successive layers in some
specimens, but two size components seem to alternate.
A central tube filled with coarser calcite crystals,
extending down from the apertural cavity and
apparently not reaching the apex, is readily observed
in cross and longitudinal sections. The apertural rim is
sharp and well preserved, indicating little postmortem
reworking. No apertural cover can be observed. The
apertural cavity is small and occupies one third to one
fourth of the length of the shell. Secondary dolomite is
nearly absent in small specimens and increases with
increasing size, as has been noted in specimens from
elsewhere (e.g., Griffin and Yochelson, 1975; Osborne
and Yochelson, 1991; Yochelson and Kisselev, 2003).
4. Discussion
Although a number of specific names have appeared
in the literature, the genus Salterella Billings (1861)
(Phylum Agmata Yochelson, 1977; Family Salterelli-
dae Walcott, 1886) seems to contain only two species
(Yochelson and Kisselev, 2003). Salterella conulata
Clark (1925) (redescribed by Yochelson, 1970) is
known from only a few localities. Where S. conulata
is found in place, it is only a few meters below
Salterella maccullochi. This latter species is exceed-
ingly widespread and abundant. Only S. maccullochi is
represented in the specimens from the Precordillera.
All apparent variations seen in thin section in angle of
expansion, geometry of cross section, presence or
absence of a central tube, and width (Fig. 3) result from
random orientation of specimens relative to the plane of
the thin section. The configuration of the conch and the
secondary deposits, as well as other morphological
features, have been compared to topotype specimens of
S. maccullochi from northern Scotland to confirm
assignment of the Argentine specimens to this species.
Although individuals are abundant, no distinct
orientation is evident. Relative abundance has no
evident relation to preservation and fabric. Although
the specimens are wellpreserved, some are affected by
abrasion (Fig. 3), suggesting a shallow-water habitat.
Rare telescoping results in a cone-in-cone arrange-
ment. Fragments of trilobites, mostly oriented parallel
to bedding, are common but are a minor constituent of
the rock.
Fig. 3 shows longitudinal, oblique, and transverse
cross sections. The range of diameters in cross sections
is (the largest about 7 mm in length) mostly related to
the position at which the specimen is intercepted in
sectioning. Nevertheless, some cross sections show the
inner deposits in a bbull’s-eyeQ pattern, and others of
the same size do not, demonstrating that larger and
smaller specimens are intermixed.
Random arrangement of specimens within the
limestone, lack of size sorting, and intervals of
solution and nondeposition imply relatively gentle
water movement, suggesting that the fossils were not
transported a long distance.
5. Stratigraphic significance
In a review of Salterella as an Early Cambrian
index fossil (Yochelson, 1977), Fritz and Yochelson
(1988) considered geographic distribution and bio-
stratigraphic range within the Early Cambrian, con-
cluding that the genus might be restricted to the medial
part of the Bonnia–Olenellus Zone (525 Ma). The
association with olenellid trilobites in Argentina
(Vaccari, 1994) confirms the age and provides a more
restricted time line for correlation. Because Salterella
can easily be identified in the field with a hand lens, it
has the characteristics of an ideal bguide fossilQ.
In the northern Precordillera at Cerro Totora (Fig.
1A), the uppermost part of the Early Cambrian Cerro
 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132
Totora Formation has a lithological association similar
to the Salterella-bearing succession in the Ancaucha
block. The lower Cerro Totora Formation contains
abundant evaporites, interlayered with red clastic
rocks and overlain by Olenellus-bearing green shales
(Vaccari, 1990; Astini and Vaccari, 1996). The
physical stratigraphy of both the Ancaucha and Cerro
Totora successions is comparable to that of the Early
Cambrian Rome Formation in outcrop and subsurface
in the southern Appalachians. Thomas and Astini
(1999) have considered these two areas as counter-
parts from the rifted margin of Laurentia. Strontium
isotopic ratios of the evaporites in the Cerro Totora
and Rome Formations are indistinguishable and are
consistent with Early Cambrian sea water, further
confirming the Early Cambrian age of these succes-
sions (Thomas et al., 2001).
Salterella in the Rome Formation in Alabama
(Osborne and Yochelson, 1991) is restricted to several
thin beds within a range of not more than 4 m. Although
the stratigraphic position closely matches that in the
Ancaucha block in the Argentine Precordillera, and the
host sedimentary facies are nearly identical, some
cautions need to be raised in order not to suggest strict
facies specificity or taphonomic artifacts.
6. Environmental setting and life habitat
In the Argentine Precordillera, Salterella is mostly
concentrated in thin-bedded coarse skeletal packstones
with no obvious fabric and minor detrital content.
Salterella forms 50–60% of the total volume of some
beds. Conchs are generally not imbricated, and they
vary from random arrangement to poorly oriented
along the long axis. The limestone contains trilobite
spines, along with rare glauconite and phosphate
grains. The Salterella-rich limestones are found within
a silty bioturbated succession containing a few quartz-
rich sandstone beds with rippled tops. Symmetrical in-
plane geometry suggests oscillatory water flow.
Interference patterns on bed tops and scattered salt
casts indicate very shallow water. In addition to the
fossil shells in limestone, some of the silty–sandy beds
contain a few external molds of Salterella.
The packstones are interpreted as hydrodynamic
concentrations. However, in the absence of conclusive
indications of strong currents or vigorous storms, the
129
high concentration of individuals leads to the spec-
ulation that Salterella was gregarious. In North
America, Salterella has been found in limestones,
sandy limestones, and sandstones, collectively sug-
gesting a shallow-water habitat (Fritz and Yochelson,
1988). The shallow-water setting limits to a small
range the possibilities for the habitat of Salterella
during life. Inner-detrital shelf belts, carbonate shoals,
and lagoonal environments are so far the most likely
paleoenvironmental settings where these organisms
lived. Common transported and aligned individuals,
few of which are imbricated, support the interpretation
of a near-shore environment. No evidence supports an
infaunal or semi-infaunal habitat, and the organism
may have used tentacles to collect grains on which
bacteria and algae lived and placed them within the
cone to strip off the nutrients (Yochelson, 1977).
There is not necessarily any relationship between a
high degree of specialization and a short stratigraphic
range, but for Salterella, the two points may be linked
(Yochelson and Kisselev, 2003). In the Early Cam-
brian, apart from trilobites, there were few shelled
organisms in the near-shore environment and presum-
ably no competition for food or space with which this
animal had to contend.
7. Palaeogeography
In North America, Salterella maccullochi ranges
from Caborca, Mexico, and California, to the North-
west Territories of Canada, from Alabama to New-
foundland, and beyond North America through
Greenland and Scotland to Svalbard. Although pelagic
organisms tend to be widespread, benthic forms,
particularly those inhabiting inner-shelf areas, less
commonly have such a wide distribution, except in
times when supercontinents with long coastlines
existed.
To the best of our knowledge, Salterella maccul-
lochi has not been found anywhere in Gondwana
(Australia, New Zealand, India, China, Africa, or
South America apart from the Argentine Precordil-
lera) or in Siberia. Reports of Salterella from
Australia were rejected as spurious (Clark, 1925;
Spath, 1936; Kobayashi, 1937). Yochelson (1983)
placed Volborthella Schmidt (1888) in synonymy
with Salterella. New data (Yochelson and Kisselev,
130 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132

2003) now suggest that Volborthella is indeed both
slightly morphologically different and stratigraphi-
cally slightly older. The distributions of Salterella and
Volborthella apparently are mutually exclusive, in that
Salterella extends through North America, Greenland,
Scotland, and Svalbard, whereas Volborthella extends
across Europe from Norway through Estonia to
Poland (Yochelson, 1981).
Like the restricted Olenellus trilobite fauna (McKer-
row et al., 1992), Salterella evidently belongs to the
Laurentian faunal realm. Salterella existed for only a
very short interval of time, but spread almost
instantaneously around Laurentia. This genus may
have originated along present-day western Laurentia,
which was open to the sea before the opening of
Iapetus. Progressively, Salterella invaded the eastern
Laurentian margin after the Iapetus Ocean (Fig. 4) had
opened enough to represent a barrier to inner-shelf
forms. Evolution of the eastern Laurentia–Iapetus
passive margin began in the earliest Cambrian (~543
Ma) along the Blue Ridge rift (Thomas, 1991). A
widespread passive margin around Laurentia was
initiated at approximately 525 Ma (Thomas and
Astini, 1999; Cawood et al., 2001), but before that,
Salterella had expanded along the shallow Laurentian
shelf during late-stage rifting (Rome facies and
analogues). Salterella in Greenland, Svalbard, Scot-
land, the central and southern Appalachians, and the
Precordillera are all in similar shallow-water mixed
facies deposited before widespread passive-margin
carbonates spread along eastern Laurentia. Both
movement of Laurentia into low latitudes soon after
break-up (Torsvik et al., 1996; Cocks and Torsvik,
2002) and global eustatic rise led to a climax in
carbonate sedimentation, smoothing differences along
the Laurentian passive margin and helping to expand
the Iapetus shelf biotas (Erwin et al., 1997), possibly
leading to a faunal turnover and consequent demise of
Salterella.
For palaeogeographic reconstructions, Salterella
can be considered as an organism equally important
and biogeographically as meaningful as the olenellid
trilobites due to their limited dispersal and demon-
strated link to Laurentian shallow-water inner-shelf
facies. The olenellid trilobites of Laurentia show
minimal links with the Redlichiid trilobite Realm of

Fig. 4. Schematic paleogeographic reconstruction of Iapetus and bordering continents (modified from Cawood et al., 2001), showing the
Precordillera rifted block as suggested by Astini et al. (1995) and Thomas and Astini (1996). Paleoequator and paleopole according to Torsvik
and Rehnstrfm (2001) and Torsvik et al. (1996). Abbreviations: SP, South Pole; AM, Amazonia; ANT, Antartica; AUS, Australia, AV, Avalon;
BA, Baltica; C-SF, Congo-Sao Francisco; IND, India; K, Kalahari; LAUR, Laurentia; RP, Rı́o de La Plata; SIB, Siberia; WA, West Africa; Pc,
Precordillera; Ox, Oaxaca; Ar, Arequipa.
 R.A. Astini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 213 (2004) 125–132
Australia–Antarctica, both strongly marked for the
Early Cambrian (McKerrow et al., 1992). This means
that by then, separation of Laurentia and Gondwana
had to be sufficiently great for the Laurentian
ollenelids to develop in isolation (Dalziel, 1997).
From the paleontological viewpoint, the restricted
Olenellus trilobite fauna in Precordillera, as pointed
by Vaccari (1994) and Astini et al. (1995), is still an
irrefutable argument to hold that this region had a
connection with Laurentia during the Early Cambrian
(Borrello, 1971; McKerrow et al., 1992; Palmer et al.,
1996; Astini and Rapalini, 2003). As stated by
Benedetto (2003), the inner-shelf Olenellid trilobite
Realm in the Argentine Precordillera cannot be
explained by patterns of oceanic circulation, due to
the fact that dispersion of benthic organisms across
oceanic basins can only be achieved, at most, by a few
eurytopic genera but never by the fauna as a whole.
Independent studies on Mid and Late Cambrian
lingulate brachiopods (Holmer et al., 1999) and
trilobites (Vaccari, 1994; Bordonaro and Banchig,
1995) recovered from the Precordillera also suggested
strong affinity to those found in North America and
Greenland, reinforcing either the geographic continu-
ity or the close proximity of the Argentine Precordil-
lera and Laurentia during the rest of the Cambrian.
However, the Early Cambrian remains the more
conclusive on a palaeobiogeographical basis.
8. Conclusion
The distribution of Salterella further strengthens
the interpretation of a Laurentian provenance for the
Argentine Precordillera terrane. Salterella provides
another line of evidence supporting palaeontological,
sedimentologic, stratigraphic, palaeomagnetic, and
isotopic evidence for rifting of the Precordillera from
Laurentia during the Early Cambrian.
Acknowledgements
We are grateful to the Consejo Nacional de Inves-
tigaciones Cientı́ficas y Técnicas (PIP-2000-02971
CONICET), the Agencia Nacional de Promoción de
Ciencia y Tecnologı́a (PICT-2002/07-11741 FON-
CYT) and the Secretarı́a de Ciencia y Tecnologı́a of
131
the Universidad Nacional de Córdoba (62/03 SECyT-
UNC) that fund our Research Projects in west
Argentina. Part of the research in Argentina was supp-
orted by a grant from the National Science Foundation
(EAR-0229522). We thank Duncan McIlroy and
Gregory Edgecombe for their helpful reviews. Peter
Cawood and Juan Luis Benedetto provided encourage-
ment after reading an early version of the manuscript.
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