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Palaeontologia Electronica
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Araya, Juan Francisco and Reid, David G. 2016. A new species of Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) from the
Quaternary of Chile. Palaeontologia Electronica 19.1.8A: 1-8
palaeo-electronica.org/content/2016/1426-a-new-echinolittorina
Copyright: February 2015 Paleontological Society
A new species of Echinolittorina Habe, 1956 (Gastropoda:
Littorinidae) from the Quaternary of Chile
Juan Francisco Araya and David G. Reid
ABSTRACT
We describe a new fossil littorinid species, Echinolittorina nielseni sp. nov., from
the Quaternary Caldera Strata, Región de Atacama, northern Chile. Fossils of littorin-
ids are globally rare because of their high-intertidal habitat on rocky shores. The new
species has a large, broad shell with strong spiral ribs and an angled periphery, differ-
ing from the two living littorinids currently found along the coasts of mainland Chile and
from all the extant species distributed in the southeastern Pacific. In comparison with
the living Chilean Echinolittorina peruviana, the new species shows stronger ribs and
more inflated whorls, but they share an unusual detail in the irregular arrangement of
spiral sculpture. We hypothesize that the new species may be ancestral or sister to E.
peruviana and discuss the adaptive significance of shell sculpture.
Juan Francisco Araya. Departamento de Geología, Universidad de Atacama, Copayapu 485, Copiapó,
Región de Atacama, Chile and Programa de Doctorado en Sistemática y Biodiversidad, Universidad de
Concepción, Concepción, Chile. jfaraya@u.uchile.cl
author: zoobank.org/Authors/443B4F42-FB13-42A6-B92B-1B0F835698A9
orcid.org/0000-0002-4087-964
David G. Reid. Mollusca Research Group, Department of Life Sciences, The Natural History Museum,
London SW7 5BD, United Kingdom. d.reid@nhm.ac.uk
Keywords: Quaternary; Pleistocene; SE Pacific Ocean; Littoraria; new species
Submission: 19 September 2015 Acceptance: 29 January 2016
INTRODUCTION
The shallow-water marine molluscs living in
northern Chile are relatively well known, having
been the subject of regional faunistic studies (e.g.,
Marincovich, 1973; Guzman et al., 1998; Ashton,
2007; Araya and Araya, 2015), while some mem-
bers of widespread genera have been included in
monographic revisions (e.g., McLean, 1984; Reid,
2002; Geiger, 2012). Nevertheless, new living spe-
cies continue to be discovered in the area, particu-
larly in the Región de Atacama (Osorio, 2012;
Araya, 2013). In comparison, there have been
fewer studies of fossil molluscs from coastal north-
ern Chile. Many are old works dealing with the
marine species from the Miocene to Pliocene
Bahia Inglesa Formation, with a characteristic tem-
perate to tropical fauna (Philippi, 1887; Möricke,
1896; Herm, 1969). Quaternary species from
ARAYA & REID: A NEW ECHINOLITTORINA
2
around Caldera were first mentioned by Herm
(1969) and have recently been the subject of
renewed study (Guzmán et al., 2000; Guichar-
rousse et al., 2015).
In contrast to the older fossils from northern
Chile, those of the Quaternary Caldera Strata
include mainly warm-temperate species, of which a
high proportion is extant on the nearby coasts. For
example, the molluscan assemblage at the locality
described here consists of 40 species, all but two
of which are still living, including the bivalve Mulinia
edulis (King and Broderip, 1832), the gastropod
Aeneator aff. fontainei (d’Orbigny, 1839) and the
chiton Acanthopleura echinata (Barnes, 1824).
This assemblage indicates a coastal, shallow-
water environment with sandy pockets and sparse
rocky outcrops (cf. Araya, 2013; Araya and Araya,
2015). These species have been extensively
reported from Quaternary deposits, while a few of
them also have Miocene records (Nielsen, 2013);
however, the extant muricid gastropod Conchole-
pas concholepas (Bruguière, 1789) and the scallop
Argopecten purpuratus (Lamarck, 1819) indicate
that the assemblage is no older than the Pleisto-
cene (Guzmán et al., 2000; Marquardt et al., 2000).
The systematics and phylogeny of the family
Littorinidae have been as thoroughly investigated
as those of any marine gastropod group (Reid et
al., 2012, and references therein) and they have
become a model system for the study of diversifi-
cation in the marine realm. In a dated molecular
phylogeny, most of the living littorinids are esti-
mated to have originated in the Oligocene to Mio-
cene, although 46% of the 60 extant Echinolittorina
species diverged more recently (Reid et al., 2012).
Littorinidae are represented in the living fauna of
northern Chile by two common species, Echinolit-
torina peruviana (Lamarck, 1822) and Austrolitto-
rina araucana (d’Orbigny, 1840) (Reid, 2002; but
see Reid et al., 2012 for current generic assign-
ments). The majority of littorinids, including these
two, are typically found in abundance at high inter-
tidal levels on wave-exposed rocky shores, where
the potential for fossil preservation is generally low.
Therefore, fossil records of the family are globally
extremely scarce (review by Reid, 1989). As
expected, the fossil record of these two species in
Chile is very limited, with a single Pleistocene
record (not figured) for Echinolittorina peruviana at
Caleta Patillos (20°45’S; 70°12’W), in northern
Chile (Rivadeneira and Carmona, 2008). For all
these reasons, the discovery of a distinctive new
species of Littorinidae in the Quaternary Caldera
Strata is of interest and is reported here.
MATERIALS AND METHODS
The type locality of the new littorinid species is
a small, low cliff to the east of El Morro Hill, one of
a series of low terraces 10 km south of Caldera,
Región de Atacama, northern Chile (Figure 1). It
consists of a semi-consolidated and fossiliferous
sedimentary formation composed of littoral marine
coquinas, sandstone, cherts, phosphorites and
diatomites (Marquardt et al., 2000; Godoy et al.,
2003). The upper layer, of variable thickness and
which contains the new species, is part of the Cal-
dera Strata and the presence of Concholepas con-
cholepas and Argopecten purpuratus indicates no
more than a Pleistocene age (Guzmán et al., 2000;
Marquardt et al., 2000). As described above, the
molluscan assemblage is consistent with a shallow
coastal environment of sand and sparse rocky out-
crops (cf. Araya, 2013; Araya and Araya, 2015). No
other littorinid taxa were present in the assem-
blage. The specific location where the new species
was found was protected from the open ocean by
the El Morro Hill, a massive 2 x 4 km outcrop
aligned almost N-S, parallel to the coast, and up to
323 m high (Herm, 1969; Godoy et al., 2003).
Sampling was carried out in January 2012 at
an eroded cliff. Bulk samples were obtained from
the upper 2 cm of the sediment and further sieved,
with some specimens picked out by hand. The
specimens described here have been deposited in
the collections of the Museo Nacional de Historia
Natural, Santiago, Chile (SGO.PI), in the Natural
History Museum, London, U.K. (NHMUK), and in
the Museo Paleontológico de Caldera, Caldera,
Chile (MPCCL). For the morphological descrip-
tions, the shell height parallel to the axis of coiling
(H), breadth (B) perpendicular to H and maximum
length of aperture (LA) were measured with Vernier
callipers (± 0.1 mm).
SYSTEMATIC PALAEONTOLOGY
Phylum MOLLUSCA Linnaeus, 1758
Class GASTROPODA Cuvier, 1795
Subclass CAENOGASTROPODA Cox, 1960
Family LITTORINIDAE Anonymous, 1834
Genus ECHINOLITTORINA Habe, 1956
Type species. Litorina tuberculata Menke, 1828,
by original designation.
Distribution. Worldwide tropical and warm tem-
perate seas (Reid et al., 2012). Eocene to Recent.
Echinolittorina nielseni sp. nov.
Figure 2.1-8
zoobank.org/DD2906C5-7FE3-4DFF-9C50-5A2471AADDA3
PALAEO-ELECTRONICA.ORG
3
v. 2015 Echinolittorina n. sp.; Araya and Reid, p.
85, figs 1.A-H.
Diagnosis. Large, broad Echinolittorina with
rounded whorls, concave spire profile, 7–9 strong
spiral ribs at and above angled periphery.
Description. Shell large (H = 15.7–22.2 mm).
Shape high turbinate (H/B = 1.29–1.44; H/LA =
1.54–1.83); spire whorls well rounded, suture dis-
tinct; spire profile concave near apex; periphery of
last whorl slightly but distinctly angled; rarely a
slight angulation at shoulder (Figure 2.6-7). Colu-
mella straight, moderately broad, slightly hollowed
and pinched at base; no eroded parietal area.
Sculpture of 7–9 low, broad, rounded, spiral ribs
above periphery (including the strongest rib at
periphery), becoming less distinct near suture; ribs
of unequal width and prominence, separated by
narrow grooves or incised lines. Base with 6–8
finer ribs. No spiral microstriae visible. A trace of
colour is preserved on one specimen, showing a
pale spiral band on the base (Figure 2.4). None of
the specimens showed fluorescence under ultravi-
olet light, which might have revealed original colour
patterns.
Type material. Holotype: SGO.PI 23.100 (Figure
2.1-2); Paratypes NHMUKPAL PI TG 26769–
26775 (Figure 2.3-8, seven specimens), MPCCL
14012016 (fifteen specimens). All the material col-
lected at type locality by J. F. Araya, January 15,
2012.
Type locality. Eroded cliff east of El Morro Hill,
about 10 km south of Caldera, Región de Atacama,
northern Chile (27°09’13”S; 70°55’33”W, 123 m);
Pleistocene.
Etymology. The name of the species honours our
friend Sven Nielsen (Universidad Austral de Chile,
Valdivia, Chile) for his extensive research on Chil-
ean fossil molluscs.
Remarks. These relatively large, thick, spirally-
sculptured shells with an entire aperture containing
a basal white band, and lacking an umbilicus,
unquestionably belong to the subfamily Littorininae
(reviewed by Reid, 1989) and can be compared
with members of its living genera. Superficially, the
shells resemble members of the littorinid genus Lit-
toraria; this assignment is suggested by the strong
spiral sculpture, the enlarged peripheral rib that
gives an angled profile to the final whorl, and the
somewhat short and wide columella. In the recent
fauna six Littoraria species occur on the Pacific
mainland coast of Central and South America, but
all have a tropical distribution and five are almost
entirely restricted to mangrove and halophytic-
grass habitats. The one rocky-shore species (Litto-
raria pintado [Wood, 1828]; see Reid, 1999a; Reid
et al., 2010) can be discounted because it is
smooth-shelled and relatively narrow with a
straight spire. The most southerly records of living
FIGURE 1. Map of the study region showing the type locality of Echinolittorina nielseni sp. nov. in Caldera, northern
Chile.
ARAYA & REID: A NEW ECHINOLITTORINA
4
FIGURE 2. Shells of Echinolittorina nielseni sp. nov. and of two extant littorinids: 1-8, Echinolittorina nielseni sp. nov.,
SGO.PI 23.100 (holotype) (1 and 2), NHMUK PAL PI TG 26769 (paratype) (3 and 4), NHMUK PAL PI TG 26770–
26773 (paratypes) (5-8); 9-13, Echinolittorina peruviana (Lamarck, 1822), NHMUK acc. no. 2176 (9 and 12), NHMUK
20001268 (10), NHMUK 20001269 (11), NHMUK unnumbered (13); and 14 and 15, Littoraria varia (Sowerby, 1832),
NHMUK 1850.4.23.273 (14) and NHMUK 1850.4.23.274 (15). 13 is a scanning electron micrograph showing shell
sculpture. 2, 8, 12, 13 are abapertural views and 4 is a basal view; others are apertural views. Collection localities:
Chile (El Morro, Atacama) = 1-8, Chile (Valparaíso) = 9 and 12, Perú (Lagunilla Beach, Paracas) = 13, Panama = 14
and 15; others are unknown.
PALAEO-ELECTRONICA.ORG
5
Littoraria species are between 3° and 5°S in north-
ernmost Peru (Reid, 1999a). Since the present fos-
sil sample is from 27°S in northern Chile and
accompanied by a fauna of temperate, rocky-shore
gastropods, their identification as a Littoraria spe-
cies associated with mangrove trees would be
extremely surprising. Of the living Littoraria spe-
cies, the new species most closely resembles Litto-
raria varia (Sowerby, 1832) (see Reid, 1999a)
(Figure 2.14-15). However, close comparison
shows that the sculpture of L. varia is stronger, the
primary spiral ribs more equal in width, and that
they are separated by wide grooves with second-
ary ribs and fine spiral microstriae, quite unlike the
rounded ribs separated by grooves that are little
more than incised lines in the new species. The
peripheral angle is marked by a strong rib in L.
varia, and there is sometimes a slight angle at the
shoulder as well (Figure 2.14), which are points of
similarity with the fossils (compare with Figure 2.6-
7). One of the fossil shells bears a spiral white
band on the base (Figure 2.4); this band is weakly
present (and then only within the aperture) in only
three of the 39 living Littoraria species (Reid,
1999b), but is present in a variety of other littorinid
genera (Reid, 1989), including all members of the
genus Echinolittorina (Reid, 2002, 2007, 2009,
2011). The sculptural details and presence of the
white basal band exclude the present shells from
Littoraria.
Turning to the modern rocky-shore fauna of
the eastern Pacific Ocean, there are 18 littorinid
species in the genera Echinolittorina and Austrolit-
torina (see Reid, 2002 for detailed descriptions and
Reid et al., 2012 for current generic assignments
based on molecular phylogeny). All have a basal
white band visible within the aperture and often
externally as well. None of the members of Echino-
littorina in the tropical eastern Pacific is similar to
the new species, being smaller, narrower, usually
with regular spiral grooves, and sometimes nodu-
lose or umbilicate. Two of the three temperate lit-
torinid species can also be discounted:
Austrolittorina fernandezensis (Rosewater, 1970) is
large (to 21 mm), but macroscopically smooth, and
is endemic to the Juan Fernández Archipelago, off
central Chile; Austrolittorina araucana (d’Orbigny,
1840) is common in Chile, again macroscopically
smooth, but does not exceed 13.8 mm in height.
The only other living littorinid in the southeastern
Pacific is Echinolittorina peruviana (Lamarck,
1822). This is a well-known species, common on
temperate shores from northern Peru to southern
Chile (5°05’S to 41°49’S; Reid, 2002; Castillo and
Brown, 2010), where it is found in local assem-
blages that are much the same as that in which the
fossil shells occurred. It too can reach a large size,
up to 23.8 mm high. However, shells of E. peruvi-
ana are usually smooth, relatively elongate (H/B =
1.31–1.88; H/LA = 1.46–1.93), the whorls are not
swollen, the columella is long and the anterior
apertural margin protrudes slightly (Figure 2.9-12)
(Reid, 2002)—characters which at first appear to
exclude close relationship with the new species.
However, a very few shells of E. peruviana (13/884
= 1.4% of shells examined from throughout geo-
graphical range) show “1–4 (rarely 6–8) faint
incised lines above periphery” (Reid, 2002: 147)
(Figure 2.9,12-13). Sometimes, even in smooth
shells, irregularities in the axial colour stripes show
a periodicity similar to that of the incised spiral lines
(compare Figure 2.11 and 2.12). The incised lines
are shown more clearly in an SEM of a well-pre-
served juvenile shell (Figure 2.13). The spacing of
the incised spiral lines in these rare shells of E.
peruviana is not as regular as the ‘primary spiral
grooves’ of typical tropical Echinolittorina species
(e.g., Reid, 2002), instead recalling the somewhat
unequal ribs of the fossils. It is, therefore, proposed
that E. nielseni sp. nov. is a member of Echinolitto-
rina and is most closely related to E. peruviana.
DISCUSSION
Generic assignment of littorinid shells in the
subfamily Littorininae, even of living species, can
be difficult because shell colour pattern and shape
are quite uniform across the subfamily, whereas
sculpture is variable, but prone to homoplasy
(Reid, 1989). Shells are, therefore, often an unreli-
able guide to affinity (even at generic level), so that
the advent of systematic anatomical studies (Reid,
1989) and of molecular phylogenetics (e.g., Reid et
al., 2012) have had profound effects on littorinid
classification. Littorinids mainly inhabit wave-swept
intertidal shores, and their fossils are consequently
extremely rare (Reid, 1989). These rare fossils are
seldom well preserved and their classification pres-
ents considerable difficulty. The 23 shells in this
sample are not significantly damaged by erosion or
abrasion, so their surface sculpture is clear, and
there is even a trace of colour pattern on one spec-
imen. While their generic placement in Echinolitto-
rina is not immediately obvious (as discussed in
detail above), our observations of sculptural details
and the basal white band make a strong case.
To assess whether the fossil shells differ suffi-
ciently from the living form to justify diagnosis as a
separate species, it is necessary to consider their
ARAYA & REID: A NEW ECHINOLITTORINA
6
respective ranges of morphological variation. Only
about 1.4% of living specimens of E. peruviana
bear faint incised spiral lines and none show the
strong, rounded spiral ribs of the fossil shells. The
living shells have less rounded whorls and are usu-
ally more elongate than the fossils, although there
is overlap in the range of the ratio H/B. There are
also subtle differences in the shape of the aperture.
It could be argued that, since the fossil shells origi-
nate from a single locality, they represent an iso-
lated population of extreme variants of E.
peruviana. However, all Echinolittorina species
have planktotrophic development and their larvae
are likely able to disperse for more than 1400 km in
ocean currents (Williams and Reid, 2004). Isolation
of a small local population, followed by genetic drift
or selection, is not likely and there is no evidence
for such effects in this genus. The type locality of
the new species (27°S) lies near the midpoint of
the modern range of E. peruviana (5–42°S), so at
least under current conditions the fossil locality is
not near the edge of the range of the living species.
Striking variation in shell sculpture on a small local
scale does occur in some species of Echinolitto-
rina, but is the result of ecophenotypic plasticity
(Yeap et al., 2001; Reid, 2007). This usually takes
the form of nodules and spiral striae in small indi-
viduals and spiral striae alone in large ones; the
development of nodules is apparently connected
with slow growth in unfavourable hot, dry micro-
habitats on horizontal rock surfaces. Reid (2002)
examined 63 samples and over 800 shells of E.
peruviana from throughout its range and reported
no cases of localized variation consistent with
ecophenotypic effects on shell sculpture. We,
therefore, conclude that description of the new
species is justified.
The great majority of Echinolittorina species
bear strong spiral striae, spiral ribs or rows of nod-
ules (Reid, 2002, 2007, 2009, 2011), as do most
other members of the Littorininae (Reid, 1989).
Outgroup comparison suggests, therefore, that the
sculpture of E. nielseni is plesiomorphic. The only
fossils of E. peruviana recorded from Chile were
not figured (Rivadeneira and Carmona, 2008). With
additional sampling of fossil shells it might be pos-
sible to address hypotheses such as whether E.
nielseni is the sculptured ancestor of the smooth
species E. peruviana, or whether the two are sister
species. If the latter, then geographical separation
can be predicted, because sister species of Echin-
olittorina are almost always allopatric, implying that
geographical isolation is required for speciation
(Williams and Reid, 2004). Strongly sculptured
(both ribbed and nodulose) littorinid species tend to
predominate in the tropics and at higher tidal levels
on the shore (Vermeij, 1973). Furthermore, the one
well-studied case of ecophenotypic plasticity in
Echinolittorina (Yeap et al., 2001) suggested that
strong sculpture might be functional in relation to
convective heat loss. At present, smooth-shelled E.
peruviana extends to higher latitudes than all but
two of the 60 worldwide living Echinolittorina spe-
cies (Williams and Reid, 2004; Reid et al., 2012).
We, therefore, hypothesize that if E. peruviana and
E. nielseni were once allopatric sister species, the
latter may have had the more northerly (more tropi-
cal) distribution. Alternatively, anagenetic change
from a sculptured ancestor to a smooth descen-
dant species could suggest adaptation to a cooling
climate. In the molecular phylogeny of Echinolitto-
rina, no living species was identified as sister to E.
peruviana; it was a basal member of an Eocene
clade (Reid et al., 2012), indicating a long period of
independent evolution or extinction of related spe-
cies.
The assemblage of fossil molluscs at the
present site indicates a coastal shallow-water envi-
ronment. The abundant shell remains of the
bivalve Mulinia edulis, which lives exclusively on
open sandy beaches, and the presence of species
of the gastropod genus Chlorostoma (C. atrum
[Lesson, 1830], C. luctuosa [d´Orbigny, 1841] and
C. tridentata [Potiez and Michaud, 1838]), and of
some polyplacophorans and limpets—known to
live exclusively on rocky shores—points to a mixed
area with rocky and sandy habitats, which at the
time was protected from the open Pacific Ocean by
the El Morro outcrop. With the exception of the new
Echinolittorina described herein and one Cyclocar-
dia species, all the molluscan taxa found at the site
are currently represented in the living fauna of the
area (Guicharrousse et al., 2015).
ACKNOWLEDGEMENTS
The authors are grateful to S. Nielsen (Univer-
sidad Austral de Chile, Valdivia, Chile) for his help
with literature and information on related species;
to the staff of the library of the Facultad de Cien-
cias and of the Departamento de Geología of the
Universidad de Chile, in Santiago, Chile for their
help with geological literature for the site; to M.E.
Araya (Caldera, Chile) for her essential help in the
field; to H. Taylor (NHMUK) for taking the photo-
graphs; to M. Guicharrouse and G. Roa (MPCCL)
for their help in preparing the map of Figure 1; and
to A. Beu (GNS Science, Lower Hutt, New Zea-
PALAEO-ELECTRONICA.ORG
7
land) and an anonymous reviewer for corrections
and suggestions that improved this work.
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