GEOLOGICA CARPATHICA
, DECEMBER 2018, 69, 6, 573–592
doi: 10.1515/geoca-2018-0034
www.geologicacarpathica.com
Palaeoenvironmental analysis of the Miocene barnacle
facies: case studies from Europe and South America
GIOVANNI COLETTI
1,
, GIULIA BOSIO
1
, ALBERTO COLLARETA
2, 3
, JOHN BUCKERIDGE
4
,
SIRIO CONSANI
5
and AKRAM EL KATEB
6
1
Dipartimento di Scienze dell’Ambiente e della Terra, Università di Milano-Bicocca, Piazza della Scienza 4, Milano 20126, Italy;
giovanni.p.m.coletti@gmail.com
2
Dipartimento di Scienze della Terra, Università di Pisa, Via Santa Maria 53, Pisa 56126, Italy
3
Museo di Storia Naturale, Università di Pisa, via Roma 79, Calci 56011, Italy
4
Earth & Oceanic Systems Group, RMIT, Melbourne 3001, Australia
5
Dipartimento di Scienze della Terra, dell’Ambiente e della Vita, Università degli Studi di Genova, Corso Europa 26, Genova 16132, Italy
6
Department of Geosciences, University of Fribourg, Chemin du Musée 6, Fribourg 1700, Switzerland
(Manuscript received August 1, 2018; accepted in revised form November 28, 2018)
Abstract: Acorn barnacles are sessile crustaceans common in shallow-water settings, both in modern oceans and in
the Miocene geological record. Barnacle-rich facies occur from polar to equatorial latitudes, generally associated with
shallow-water, high-energy, hard substrates. The aim of this work is to investigate this type of facies by analysing, from
the palaeontological, sedimentological and petrographical points of view, early Miocene examples from Northern Italy,
Southern France and South-western Peru. Our results are then compared with the existing information on both modern
and fossil barnacle-rich deposits. The studied facies can be divided into two groups. The first one consists of very shallow,
nearshore assemblages where barnacles are associated with an abundant hard-substrate biota (e.g., barnamol). The second
one includes a barnacle-coralline algae association, here named “barnalgal” (= barnacle / red algal dominated), related to
a deeper setting. The same pattern occurs in the distribution of both fossil and recent barnacle facies. The majority of them
are related to very shallow, high-energy, hard-substrate, a setting that represents the environmental optimum for
the development of barnacle facies, but exceptions do occur. These atypical facies can be identified through a complete
analysis of both the skeletal assemblage and the barnacle association, showing that barnacle palaeontology can be
a powerful tool for palaeoenvironmental reconstruction.
Keywords: Carbonate Factories, Heterozoan, Barnamol, Barnalgal, Tertiary Piedmont Basin, Sommières Basin,
Pisco Basin.
Introduction
Acorn barnacles (Cirripedia: Sessilia) are common carbonate
producers in modern and fossil shallow-water shelf environ-
ments (Foster 1987; Foster & Buckeridge 1987; Doyle et al.
1997). Albeit often overlooked, this group of sessile, suspen-
sion-feeding crustaceans occur on any available surface in
shallow seas, including “mobile surfaces” like turtles and
whales (Ross & Newman 1967; Newman & Abbot 1980;
Scarff 1986; Seilacher 2005; Bianucci et al. 2006; Dominici et
al. 2011; Harzhauser et al. 2011; Collareta et al. 2016 a, b).
They are common at middle and high latitudes (Raymond &
Stetson 1932; Hoskin & Nelson 1969; Milliman 1972; Müller
& Milliman 1973; Farrow et al. 1978; Hottinger 1983; Domack
1988; Nelson et al. 1988; Scoffin 1988; Wilson 1988; Taviani
et al. 1993; Henrich et al. 1995; Frank et al. 2014; Buckeridge
2015), but they can also thrive at low-latitudes, especially
in nutrient-rich environments (Glynn & Wellington 1983;
Carannante et al. 1988; Halfar et al. 2006; Westphal et al.
2010; Michel et al. 2011; Reijmer et al. 2012; Klicpera et al.
2013; Reymond et al. 2016). The fossil record of encrusting
cirripedes starts in the Cretaceous (if primitive forms like
Archaeochionelasmus Kočí, Newman & Buckeridge, 2017 in
Kočí et al. 2017 are included), but it is only during the Neo-
gene that barnacles became really frequent in the shallow-
water environments that they presently master (Darwin 1854;
Newman et al. 1969; Foster & Buckeridge 1987; Doyle et al.
1997; Buckeridge 2015).
Worldwide, barnacle facies are particularly well represented
in the sedimentary sequences of the Neogene and the Quater-
nary (Sakai 1987; Donovan 1988; Kamp et al 1988; Nebelsick
1989, 1992; Hayton et al. 1995; Doyle et al. 1997; Betzler et
al. 2000; Nielsen & Funder 2003; Civitelli & Brandano 2005;
Aguirre et al. 2008; Nomura & Maeda 2008; Radwańska &
Radwański 2008; Massari & D’Alessandro 2012; Stanton &
Alderson 2013; Brandano et al. 2015; Buckeridge 2015;
Buckeridge et al. 2018). Based on the ecology of modern taxa,
these facies are generally interpreted as shallow-water,
high-energy, deposits. Although this interpretation is usually
reasonable, considering the modern distribution of barnacle-
rich sediments, barnacle facies clearly display a variability
that reflects environmental differences. The aim of this work is
to investigate the environmental factors that govern the deve-
lopment of barnacle facies, thus gaining further insights useful
for palaeoenvironmental reconstructions. To achieve this goal,
four Burdigalian (early Miocene) barnacle facies, from both
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, 2018, 69, 6, 573–592
the Northern and Southern hemisphere, were analysed and
compared by means of palaeontology, sedimentology and
petrography, highlighting their differences and their simila-
rities. The studied barnacle-rich skeletal assemblages are
located in the well studied successions of the Pietra da Cantoni
Basin in Italy, of the Sommières Basin in France, and of
the East Pisco Basin in Peru (Fig. 1; Vannucci et al. 1996;
Bicchi et al. 2006; Reynaud & James 2012; Coletti et al. 2015;
Bianucci et al. 2018; DeVries & Jud 2018; Di Celma et al.
2018b). Previous research provides a firm basis for the inter-
pretation of the barnacle facies of these basins, which has
received limited attention until now. These facies, despite
having in common abundant remains of barnacles, are charac-
terized by different skeletal assemblages and different
petrographic composition, thus suggesting different palaeoen-
vironmental settings. The results of this analysis are integrated
with the existing information on both modern and fossil bar-
nacle facies, to provide a general framework for this kind of
sedimentary rock.
Geological setting
Pietra da Cantoni Basin, Northern Italy
The Pietra da Cantoni Group was deposited in the eastern
Monferrato, a part of the Tertiary Piedmont Basin that evolved
from the late Eocene to the late Miocene, over the inner part of
the Alpine wedge (Novaretti et al. 1995; Rossi et al. 2009).
During the Aquitanian, the deformation caused by the rotation
of the orogenic wedge uplifted the eastern Monferrato and
resulted in the deposition of the Burdigalian to early Langhian
limestones of the Pietra da Cantoni (Clari et al. 1995; Novaretti
et al. 1995; Maffione et al. 2008). The group is divided into
two depositional sequences (Bicchi et al. 2006). The oldest
(“Sequence 1” sensu Bicchi et al. 2006) is related to the first
and localized marine transgression. The youngest (“Sequence
2” sensu Bicchi et al. 2006) accumulated at the beginning of
a transgressive trend in the area that lasted for most of
the Miocene. The second sequence is divided into two units.
The Lower Unit of Sequence 2 is characterized by coral-
line-algal-nodule rudstones and floatstones interbedded with
grainstones and rudstones rich either in large benthic fora-
minifera or in barnacles. Deposition occurred during
the Burdigalian (Novaretti et al. 1995; Ruffini 1995; D’Atri et
al. 1999, 2001). The Upper Unit of Sequence 2 is characte-
rized by foraminiferal oozes, testifying hemipelagic sedimen-
tation. A bed of condensed sediments, rich in glauconite and
phosphates, separates the two units (Schüttenhelm 1976;
Bicchi et al. 2006). This interval is related to a period of major
sediment starvation, caused by the drowning of the carbonate
factory (Coletti et al. 2015). Both units of Sequence 2 occur in
the outcrop of Uviglie (Fig. 1; 45°04’42” N, 08°24’48” E),
where the barnacle-rich facies has been investigated.
Sommières Basin, Southern France
The Alpine Molasse Basin was a seaway during the early
Miocene; it was about 100 km wide and 1000 km long and
connected the Western Mediterranean with the Western
Fig. 1. Location of the studied facies. A — World map including the location of the studied areas highlighted in the panels. B — Western
Europe, magnification of panel B included in panel A. C — Location of the Sommières Basin (France) and of the Pietra da Cantoni Basin
(Italy). D — Location of the outcrop of Uviglie (Italy). E — Location of the Boisseron outcrop (France). F — North-western South America,
magnification of panel F included in panel A. G — Location of the Ullujaya outcrop (Peru).
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MIOCENE BARNACLE FACIES FROM EUROPE AND SOUTH AMERICA
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, 2018, 69, 6, 573–592
Paratethys (Allen et al. 1985; Rögl 1998; Dercourt et al. 2000;
Reynaud & James 2012). This narrow sea was dominated
by strong tidal currents created by the amplification of
the Atlantic tide entering the basin from the south-west (Allen
et al. 1985; Harzhauser & Piller 2007). The Sommières Basin
was a small embayment within the Alpine Molasse Basin,
located near the junction of the seaway with the Mediterranean
and connected with the former through a flooded valley
(Reynaud & James 2012). Within the Sommières Basin, three
main units are recognized: the Sandy Molasse, the Sandy
Marls and the Calcareous Molasse (Demarcq 1970; Reynaud
& James 2012). These units represent two different Burdigalian
depositional sequences (Berger 1974; Reynaud & James
2012). The first sequence records the marine transgression in
the Sommières Basin: in particular, the Sandy Molasse Unit
represents the transgressive system tract, while the Sandy
Marls Unit is the highstand system tract (Reynaud & James
2012). The end of the deposition of the Sandy Marls Unit is
followed by an abrupt fall in sea level. The subsequent trans-
gression is represented by the Calcareous Molasse Unit, which
is included in the second sequence (Reynaud & James 2012).
The Sandy Molasse Unit, which deposited during the early
Burdigalian, is characterized by limestones composed mainly
of bryozoans, molluscs, barnacles, echinoids and coralline
algae (Berger 1974; Reynaud & James 2012). The barna-
cle-rich assemblages occur in the lowermost bed of the unit
that onlaps the basement of the basin (“Sub-facies A1” sensu
Reynaud & James 2012). The architecture of these deposits
is especially clear in the outcrop of Boisseron (Fig. 1;
43°45’42” N, 04°04’54” E), investigated in the present paper.
East Pisco Basin, South-western Peru
The East Pisco Basin was a Cenozoic semi-enclosed
forearc-embayment, protected by an archipelago of islands,
located on the southern coast of Peru (DeVries & Jud 2018;
Di Celma et al. 2018b). It has been mainly investigated for its
diverse and exceptionally-preserved Neogene fossil verte-
brates (including pinnipeds, sharks, crocodiles, seabirds, tur-
tles and bony fish) that characterize several outcrops west of
the Ica River (e.g., Bianucci et al. 2015, 2016 a, b, and previous
references therein; Lambert et al. 2014, 2015, 2017 a, b;
Landini et al. 2017 a, b, 2018; Marx et al. 2017; Gioncada et
al. 2018). The sedimentary successions were first described
in the 1990s by Dunbar et al. (1990) and DeVries (1998).
Within the Palaeogene succession, these authors recognized
the Caballas (middle Eocene), Paracas (middle to late Eocene
age) and Otuma (late Eocene to early Oligocene age) forma-
tions. These are followed by the Neogene Chilcatay and Pisco
formations. The lower Miocene Chilcatay Formation (investi-
gated over the years by Wright et al. 1988; DeVries & Schrader
1997; León et al. 2008; DeVries & Jud 2018) is
a focus of this paper. Recent surveys in the Western side of
the Ica River Valley have recognized within the Chilcatay
Formation distinct depositional sequences, separated by basin-
wide unconformities (Di Celma et al. 2017, 2018a). Based on
that, the Chilcatay Formation has been divided into the Ct1
and Ct2 allomembers (Di Celma et al. 2018b). Ct1 is further
subdivided into two facies associations: Ct1a, composed of
sandstones and conglomerates (less common than sandstones)
alternating with siltstones, and Ct1b, including clinobedded
coarse-grained mixed siliciclastic/bioclastic arenites. Both
facies associations are characterized by abundant barnacle
remains. The overlaying Ct2 deposits include massive and
intensely bioturbated sandstones, changing upwards into mas-
sive siltstones with dolomitized mudstone layers. A tephra
layer near the top of this sequence locates the Chilcatay
Formation in the Burdigalian — which agrees with diatom
and silicoflagellate biostratigraphy (Di Celma et al. 2017,
2018b). These allomembers are well exposed at Ullujaya
(Fig. 1; 14°35’06” S, 75°38’30” W), and are investigated in
the present work.
Material and methods
For the purposes of the present work, skeletal assemblages
where barnacle remains dominate or codominate the bioclastic
fraction of the rock are regarded as barnacle facies (with
codominance meaning the situation in which barnacles, within
reasonable confidence limits, are equally abundant to another
group of skeletal grains). Barnacle facies were studied in
the outcrops of Uviglie, Boisseron and Ullujaya in order to
describe their macroscopic texture and sedimentary structures
(Fig. 1). Special attention was given to the observation of fos-
sil barnacle assemblage
s
, focusing on their first-order taxo-
nomic composition. Barnacle preservation and distribution
were investigated following the methods of Doyle et al.
(1997), Nomura & Maeda (2008) and Nielsen & Funder
(2003), which are based on the fragmentation of shells and
whether or not the specimens are in life position. Representative
rock samples from each facies were collected for petrographic,
palaeontological and mineralogical analyses. A large number
of barnacle specimens were also collected in the studied out-
crops (including isolated opercula, shell fragments, complete
shells and multi-individual aggregates; Fig. 2). In most cases,
cirripede palaeontology relies on an analysis of shell frag-
ments as the co-embedding of adjacent valves and, even more,
the occurrence of complete shells is uncommon (Foster &
Buckeridge 1987). For this reason, barnacle taxa were studied
by integrating information from both complete specimens
(where available) and plates retrieved from the embedding
rock. From each facies, about 500 g of rock sample was disag-
gregated through freezing-thawing cycles and then wet-sieved
to isolate barnacle opercula and wall plates (following Aguirre
et al. 2008). After studying them under a stereomicroscope,
some selected specimens (67, including compartment frag-
ments, isolated wall plates and complete shells) were prepared
as thin sections to observe the internal microstructure of
the shells (Fig. 2), which can be useful for taxonomic identifi-
cation (Cornwall 1956, 1958, 1959, 1960, 1962; Davadie 1963;
Newman et al. 1969; Newman & Ross 1971; Buckeridge 1983).
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In particular, attention was paid to interlaminate figures (i.e.,
“lames épithéliales” sensu Davadie 1963) that are observable
in thin sections of the wall plates of the encrusting barnacles
with a calcareous basis. These interlaminate figures (e.g.,
Fig. 2) reflect the patterns made by the organic matrix during
the formation of the calcareous denticles at the basal ends
of the longitudinal septa of the wall plates that interdigitate
with the corresponding radial septa of the calcareous basis
(Newman et al. 1967). As a result of the uncertainties inherent
in fossil barnacle taxonomy, it was decided to simply differen-
tiate the different taxa occurring in the investigated facies.
When enough information for a reliable diagnosis was avai-
lable an identification is proposed; otherwise only a provi-
sional identification is attempted. The systematic taxonomy
used for the classification mostly follows Buckeridge (1983),
Zullo (1992) and Newman (1996).
The petrographic characteristics of the rocks and their
ske letal assemblages were analysed on 35 polished thin
sections. The different components were identified and quan-
tified using point-counting method with a minimum of 400
points per section (Flügel 2010). X-ray powder diffraction
analysis (XRD) method was used to estimate the carbonate
and siliciclastic fractions of the whole rock. Samples were first
ground in an agate mortar, and then mounted on zero-back-
ground silicon plates. The measurements were made with
a Philips PW1140 diffractometer equipped with CoKα radia-
tion (Kα1 wavelength 1.789 Å) operating at 40 kV and 20 mA.
Each sample was scanned between 3° and 70° 2θ with a step
size of 0.02° 2θ and an acquisition time of 1 s per step. X-ray
pattern treatment was carried out with Panalytical X’pert
HighScore Plus to identify the mineralogical phases and
a semiquantitative analysis was carried out using the Reference
Intensity Ratio (RIR
)
method (Chung 1974).
Results
Barnalgal, Pietra da Cantoni Group
Skeletal assemblage and mineralogical composition
The barnacle facies of the Pietra da Cantoni Group consists
of a massive rudstone with a slight yellowish to pinkish colour
(Fig. 3A–E). This peculiar colour of the rock is partly due to
the large number of barnacle plates that still retains their pin-
kish pigmentation. The rock is very porous, poorly lithified
and, except for some rare burrows (Fig. 3C), does not present
macroscopically evident sedimentary structures; only locally
flat skeletal elements exhibit a preferential orientation (Fig. 3F).
Besides barnacle plates, coralline-algal nodules (rhodoliths)
are abundant, especially towards the top of the interval. Most
of them are encrusted by barnacles (Fig. 3D). Sectioned rhodo-
liths show that barnacles are also present within the nodules,
alternating with layers of coralline algae. Thin section analysis
shows that the skeletal assemblage is dominated by barnacles
and coralline algae (mainly Hapalidiales; Table 1; Fig. 3F–G).
For this assemblage, the new term “barnalgal” is here pro-
posed, complementing the assemblages introduced by Hayton
et al. (1995). Bryozoans are relatively common (Table 1); their
association includes branched colonies, globular colonies
and disarticulated elements of the articulated bryozoan
Bifissurinella lindenbergi Keij, 1969. Benthic foraminifera are
present but less abundant (Table 1; Fig. 3F–H), the most com-
mon genera being Amphistegina d’Orbigny, 1826, Elphidium
Montfort, 1808, and Cibicides Montfort, 1808. Sphaerogypsina
Galloway, 1933, Miogypsina Sacco, 1893, Operculina
d’Orbigny, 1826, Nephrolepidina Douvillé, 1911, Eponides
Montfort, 1808, and Neoconorbina Hofker, 1951 are less com-
mon; Stomatorbina Dorreen, 1948, textulariids and miliolids
are rare. Echinoids are uncommon in the skeletal assemblage
and mostly occur as loose spines (Table 1). Molluscs are rare,
those present being mainly pectinids; serpulids and ostracods
are very rare (Table 1). The siliciclastic fraction is almost
non-existent, except for some rare pebbles and some quartz
grains (Table 1). XRD results support this observation, indi-
cating >> 95 % of carbonate minerals (Table 1).
Barnacle preservation
Cirripede shells are evenly distributed in the horizon; they
are generally disarticulated and the plates are slightly abraded
(Fig. 3E; grade 0 of Nielsen & Funder 2003). Complete shells
occur on the surface of rhodoliths, together with stubs (i.e.,
broken shells of which almost only the base is preserved) and
partially broken specimens (Figs. 3D; 4A–B). These speci-
mens have randomly-oriented openings, suggesting that they
are no longer in life position (Type B preservation of Nomura
Fig. 2. Generalized structure of a balanomorph barnacle shell. Lower
left corner present a model of a complete shell, partially modified
after Buckeridge (1983); upper left includes details of the opercular
plates; lower right includes details of a carinal plate and its internal
structure revealed by a transverse section above the basal margin
showing the interlaminate figures, one being enlarged in the upper
right.
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& Maeda 2008; displaced clusters of Doyle et al. 1997).
The specimens are moderately well-preserved, retain their
colour and only lack the loose and tiny opercula (Fig. 4B).
The latter are quite abundant in the rock, commonly slightly
abraded, even though some of them still preserve a pinkish
colour; sometimes they present gastropod predation holes.
Scuta and terga do not occur in equal abundance, the scuta
being much more common than the terga (Fig. 4C–D).
Barnacle identification
The shells are generally small, with the basal diameter in
adult specimens ranging from slightly under 0.5 cm to a little
over 1 cm (Fig. 4B). They are comprised of six mural plates
plus a calcareous, tubiferous basis. Externally the parietes pos-
sess strong longitudinal ribs. Where the colour is preserved,
the plates are light pink with white spots organized in
Fig. 3. Barnacle coralline assemblage (barnalgal) of the Lower Unit of Sequence 2 of Pietra da Cantoni Group (Uviglie, Italy). A — Simplified
stratigraphic column of the barnalgal of the Pietra da Cantoni Group. B — Overview of the Uviglie outcrop including a simple stratigraphic
column highlighting the different units and the barnalgal facies, BRG = barnalgal assemblage, Sq = sequence. C — Detail of a burrowing trace,
black arrow = trace. D — Barnacle-encrusted rhodolith, black arrow = barnacles. E — Detail of the texture of the facies, black arrow = Operculina.
F — Thin section of a sample of lower part of the barnalgal facies. G — Thin section of a sample of the upper part of the barnalgal facies.
H — Axial section of a Miogypsina specimen.
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longitudinal rows. Interlaminate figures are present; they have
short and almost straight main axes with few transverse
branches (Fig. 4E–F). The terminations of the branches are
crescent-moon to arrow-shaped. The scuta are
thick and triangular in shape, with the outer sur-
face characterized by transverse growth lines
(Fig. 4C). The terga are much thinner, with
a spur as long as one third of the total length of
the plate, removed from the basiscutal angle by
less than half its width, and with a very shallow
furrow (Fig. 4D). The size and the morphology
of the recovered opercula indicates that they all
belong to a single species. This consistency is
also reflected in the wall plates and their inter-
laminate figures, leading us to conclude that
only one barnacle species is present in this
facies. The morphology of the opercula, wall
plates and interlaminate figures (Fig. 4E–F)
conforms to a taxon of Balanidae closely allied
to the amphibalanine Amphibalanus amphitrite
(Darwin 1854).
Barnamol, Sandy Molasse Unit
Skeletal assemblage and mineralogical
composition
This facies consists of rudstone organized in low-angle
cross-bed sets (Fig. 5A–C). The rock is porous, especially at
the base of the succession, and exhibits a clear fabric due to
the common orientation of flat skeletal elements (Fig. 5D–F).
Barnacle plates and fragments of molluscs are very common.
Granule-sized bryozoan colonies are also visible on the rock
surface. According to point-counting analysis the skeletal
assemblage is dominated by barnacles and molluscs (conse-
quently the assemblage was classified as barnamol sensu
Hayton et al. 1995; Table 1; Fig. 5 D–I). Barnacles are espe-
cially important at the base of the interval, where they account
for most of the skeletal grains. Molluscs are more important
upwards, where they are locally more abundant than barna-
cles; they are mainly represented by ostreids and subordinated
pectinids. Echinoids are abundant, occurring as spines and test
fragments (Table 1). Bryozoans are common (both branching
and globular colonies are present; Table 1). Coralline algae
and benthic foraminifera are very rare, the latter being mostly
represented by Cibicides, Amphistegina and Miogypsina
(Table 1). The siliciclastic fraction is important and accounts
for 16.5 % of the detected elements (Table 1; Fig 5D–L).
It includes quartz grains and fragments of sedimentary rocks
(mainly limestones and sandstones). XRD results highlight
the pre sence of about 30 % of silicate minerals and 70 % of
carbo nates (Table 1).
Barnacle preservation
Wall plates are invariantly disarticulated and evenly distri-
buted within the rock (Type D of Nomura & Maeda 2008;
comminuted shell bed of Doyle et al. 1997). They are frag-
mented, disarticulated and heavily abraded, often exposing
Pietra da Cantoni
Group
Sandy Molasse
Unit
Chilcatay Formation
Barnalgal
Barnamol
Ct1a
barnacle facies
Ct1b
barnacle facies
Petrographic composition [Point counting]
Bioclastic components
95.0 %
68.5 %
22.0 %
24.0 %
Terrigenous components
0.5 %
16.5 %
65.0 %
30.5 %
Sparite
4.0 %
8.0 %
10.0 %
41.0 %
Micrite
0.5 %
7.0 %
3.0 %
4.5 %
Detail of the bioclastic fraction [Point counting]
Barnacles
41.5 %
34.5 %
71.5 %
80.0 %
Molluscs
0.5 %
40.5 %
8.5 %
7.5 %
Echinoids
1.0 %
16.5 %
18.5 %
8.0 %
Bryozoans
6.5 %
8.0 %
< 0.5 %
///
Coralline algae
47.0 %
< 0.5 %
///
///
Benthic foraminifera
3.5 %
< 0.5 %
1.0 %
4.0 %
Serpulids
< 0.5 %
< 0.5 %
< 0.5 %
< 0.5 %
Ostracods
< 0.5 %
///
///
///
Mineralogical composition [XRD]
Carbonates
97.0 %
72.0 %
16.0 %
31.0 %
Silicates
3.0 %
28.0 %
71 %
64.0 %
Salts
0.0 %
0.0 %
13.0 %
5 %
Table 1: Petrographic composition, skeletal assemblage and mineralogical content of
the examined barnacle facies.
Fig. 4. Barnacles of the barnalgal of Uviglie (Italy). A — A rhodolith
completely encrusted by barnacles (cf. Amphibalanus sp.), upper part
of the Lower Unit of Pietra da Cantoni Sequence 2 (Colma, Fig. 1).
B — Barnacle shells (cf. Amphibalanus sp.) growing on a rhodolith.
C — Internal and external surfaces of the same scutal plate.
D — Internal and external surfaces of the same tergal plate.
E — Interlaminate figures of a wall plate of cf. Amphibalanus sp.
F — Detail of the interlaminate figures of the plate depicted in panel E.
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their internal structure (Grade 1 and 2 of Nielsen & Funder
2003). None of the observed plate fragments preserve their
original colour. Very rare scuta, but no terga, were recovered
from the disaggregated rock sample. They are fragmented,
abraded and exhibit pitted surfaces probably due to incipient
pressure solution.
Barnacle identification
The preservation state of the cirripede remains greatly
hinders identification based on their macroscopic features.
The inner structure of the plates and the interlaminate figures
indicates that three different taxa are probably present:
Concavinae? indet., Balaninae gen. et sp. indet. 1, and
Balaninae gen. et sp. indet. 2 (Fig. 6). The presence of more
than one barnacle taxon is also supported by the scutal plates,
which exhibit at least two different morphologies: one charac-
terized by transverse growth lines only and another exhibiting
both transverse growth lines and longitudinal (i.e., radial)
striae (Fig. 6F–G, respectively). The specimens identified as
Concavinae? indet. have small parietal-tubes, sometimes
divided by septa and characterized by interlaminate figures
with a straight axis and few transverse branches with crescent-
moon- to arrow-shaped terminations (Fig. 6A). The presence
Fig. 5. Barnacle-mollusc assemblage (barnamol) of the Sandy Molasse Unit of the Sommières Basin (Boisseron, France). A — Simplified
stratigraphic column of the barnamol facies of the Sandy Molasse Unit. B — Overview of the Boisseron outcrop. C — Macroscopic texture of
the facies. D — Thin section of a sample from the barnamol facies with common orientation of the bioclasts. E — Thin section of a barna-
cle-rich sample of the barnamol facies. F — Thin section of a mollusc-rich sample of the barnamol facies. G — Detail of a Miogypsina speci-
men. H — Detail of a cross-section of a tubiferous barnacle wall plate exhibiting interlaminate figures.
H
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of representatives of the subfamily Concavinae is as also sug-
gested by the occurrence of scuta with both longitudinal striae
and transverse growth lines; however, it must be stressed that
the latter feature is also observed in the extant amphibalanine
Amphibalanus eburneus (Gould, 1841). Concavine barnacles
are known to occur in the Neogene deposits of the Alpine
Molasse Basin. De Alessandri (1906) pointed out the presence
of “Balanus” concavus Bronn, 1831 (=Concavus concavus) in
lower Miocene strata of the Upper Marine Molasse of
Switzerland, but under this specific name several taxa of spe-
cific and generic rank are lumped (Newman 1982; Zullo 1992;
Carriol 2000). More recently, concavine barnacles (belonging
to the genus Chesaconcavus Zullo, 1992) have been reported
by Carriol & Schneider (2008) and Carriol & Menkveld-
Gfeller (2010) from Burdigalian beds of the Upper Marine
Molasse of Bavaria and Switzerland, respectively. The speci-
mens referred to Balaninae gen. et sp. indet. 1 have larger
parietal tubes and interlaminate figures with a long axis and
numerous transverse branches with club-shaped terminations
(Fig. 6B–C). Similar interlaminate figures have been figured
by Nebelsick (1989: pl. 4, figs. 2–3) for Burdigalian barnacles
of the Zogelsdorf Formation of Austria (Alpine Molasse
Basin) and by Davadie (1963; pl. XIV) for recent specimens
of the subfamily Balaninae. The specimens belonging to
Balaninae gen. et sp. indet. 2 have large tubes and interlami-
nate figures with a bent axis and numerous transverse branches
with club-shaped terminations; differing from the other
groups, these branches present complex ramifications
(Fig. 6D–E).
Barnacle facies, Chilcatay Formation
Skeletal assemblage and mineralogical composition
Two slightly different barnacle facies can be recognized in
Ct1a and Ct1b facies associations, respectively (Fig. 7A–B).
The barnacle facies of Ct1a consists of poorly-sorted
coarse-sandstones to conglomerates with siliciclastic and bio-
clastic elements (Fig. 7A, C–E); the barnacle-bearing horizons
alternate with medium- to fine-grained sandstones, almost
devoid of bioclasts (Fig. 7A).The rock is massive, porous and
poorly cemented. Barnacles are very common, occurring as
clusters of shells, isolated individuals and large shell frag-
ments (Fig. 7D–E).According to thin section analysis, barna-
cles dominate the skeletal assemblage (Table 1; Fig. 7F–G).
Echinoids are also very common; molluscs are less abundant
and mainly represented by ostreids and pectinids (Table 1).
Benthic foraminifera are very rare (mainly Cibicides,
Peneroplis Montfort, 1808 and Nonion Montfort, 1808;
Table 1). Serpulids are uncommon in the skeletal assemblage,
but occasionally they occur in large clusters. Bryozoans are
very rare (Table 1). The siliciclastic fraction is important and
accounts for more than half of the components (Table 1;
Fig. 7F–G). It consists of fragments of igneous rocks from
the basement and ash-flow tuffs. XRD results indicate > 70 %
of silicate minerals and only lesser amounts of carbonates and
salts (mainly gypsum; Table 1).
The barnacle facies of Ct1b is organized into clinobeds of
well-sorted, coarse-grained mixed siliciclastic-bioclastic
deposits, which are more cemented and less porous than those
of the lower facies (Fig. 7A). On a macro-scale at the outcrop,
barnacles dominate the bioclastic fraction (occurring as clus-
ters, isolated individuals and shell fragments), but bivalves
also occur (Fig. 7 H–J). In thin section, barnacle remains are
the most abundant group of skeletal grains; however, the com-
mon presence of sparite-filled molds suggests that molluscs
were also important (Table 1; Fig. 7 K–L). Among the few
preserved bivalve specimens, ostreids and pectinids are domi-
nant. Echinoids are also abundant (Table 1). Benthic fora-
minifera are less common (mainly Cibicides and Peneroplis;
Table 1). Rare calcareous tubes also occur (Table 1). Based on
point-counting analyses, siliciclastic components are slightly
more important than bioclasts and they mainly consist of
pebbles and granules of igneous basement rocks (Table 1;
Fig. 7F–G, K–L). XRD results suggest that silicate minerals
account for > 60% of the rock, whereas carbonates only repre-
sent 30 % (Table 1). Minor amounts of gypsum are also present
(Table. 1).
Fig. 6. Barnacles of the barnamol of Boisseron (France).
A — Concavinae? indet., interlaminate figures; note the straight axis,
few branches and crescent-moon to arrow-shaped terminations.
B — Balaninae gen. et sp. indet. 1, interlaminate figures; note the
long axis and numerous branches with club-shaped terminations.
C — Balaninae gen. et sp. indet. 1, detail of the interlaminate figures.
D — Balaninae gen. et sp. indet. 2, interlaminate figures; note
the bent axis and numerous, ramified branches with club-shaped
terminations. E — Balaninae gen. et sp. indet. 2, detail of the inter-
laminate figures. F — External surface of a scutal plate exhibiting
transverse growth lines only. G — External surface of a scutal plate
exhibiting both longitudinal striae and transverse growth lines.
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Fig. 7. Barnacle facies of the Chilcatay Formation (Ullujaya, Peru). A — Simplified stratigraphic column of the barnacle facies of
the Chilcatay Formation. B — Overview of the Ullujaya outcrop including a simple stratigraphic column highlighting the different facies.
C — Macroscopic texture of the barnacle facies of Ct1a. D — Detail of the texture and clast composition of the Ct1a barnacle facies.
E — Detail of the texture and composition of the Ct1a barnacle facies. F — Thin section of a sample of the Ct1a barnacle facies. G — Detail
of sample of the Ct1a barnacle facies with barnacles and foraminifera. H — Macroscopic texture of the Ct1b barnacle facies. I — Detail of
the texture and bioclastic composition of the Ct1b barnacle facies. J — Detail of the texture and composition of the Ct1b barnacle facies.
K — Thin section of a sample of the Ct1b barnacle facies. L — Detail of the abundant sparite-filled molds in the Ct1b barnacle facies.
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Barnacle preservation
In both facies, the cirripede remains occur as displaced
clusters, displaced complete shells and wall plate fragments
(Fig. 8). Unlike the barnacle-coralline facies, the clusters are
generally detached from their original substrate (Type C of
Nomura & Maeda 2008; displaced clusters of Doyle et al.
1997).
Complete specimens are generally moderately well-pre-
served, but display evidence of abrasion and lack the opercula
(Fig. 8). The best preserved specimens retain their pigmen-
tation (Fig. 8A, D). The shells are often filled by sand.
In the barnacle facies of Ct1a, this filling is remarkably rich in
bioclastic fragments and has less abundant mineral grains than
the rock embedding the specimens. Disarticulated plates are
often abraded (Grade 1 of Nielsen & Funder 2003). Opercula
are rare — only a couple of abraded and fragmented scuta
have been recovered.
Barnacle identification
Based on macroscopic features of the wall plates and on
their internal microscopic structure, three barnacle taxa have
been recognized: cf. Austromegabalanus sp., Balanidae indet.,
and Concavinae indet. (Fig. 8). The specimens identified as cf.
Austromegabalanus sp. have a shell diameter approaching
4 cm (Fig. 8A–C); they are comprised of six wall plates with
broad, well-developed radii plus a calcareous, tubiferous,
basis (Fig. 8A). The best preserved specimens are longitudi-
nally striped, with pink-purple bands alternating with white
(Fig. 8A). In thin section, the radii are tubiferous and their
sutural edges bear transversely oriented septa with secondary
Fig. 8. Barnacles of the barnacle facies of Ullujaya (Peru). A — cf. Austromegabalanus sp., complete shell (from Ct1a). B — cf. Austro
megabalanus sp., interlaminate figures (from Ct1a). C — cf. Austromegabalanus sp., detail of the interlaminate figures (from Ct1a).
D — Balanidae indet., cluster of shells (from Ct1a). E — Balanidae indet., interlaminate figures (from Ct1a). F — Balanidae indet., detail of
the interlaminate figures (from Ct1a). G — Concavinae? indet.,cluster of shells (from Ct1b). H — Concavinae? indet., interlaminate figures
(from Ct1a). I — Concavinae? indet., detail of the interlaminate figures (from Ct1a).
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denticles on the lower side only (a diagnostic character of
the megabalanine genera included in the tribe Austro-
megabalanini; Newman 1979; Buckeridge 2015). The interla-
minate figures present an almost straight main axis with many
transverse branches characterized by multiple arrow-shaped
terminations (Fig. 8B–C). Austromegabalanus Newman, 1979
is known to occur along the coasts of southern Peru and in
the Miocene sediments of the Pisco Basin (e.g., Newman
1991; Carriol et al. 1987). The specimens referred to Balanidae
indet. are medium- to large-sized shells, comprised of six wall
plates and characterized by a longitudinally striated outer wall
with spiny-bulging protuberances creating a rough and irre-
gular external surface (Fig. 8D–F). Some individuals display
a pink colour on the plates (Fig. 8D). In the lowermost portion
of the shell the basal edges of the wall plates become inflected,
and grow horizontally inwards, thus producing an inward-
tapering calcareous membrane. This feature is often incom-
plete (i.e., a hole is present at the centre of the basal calcareous
membrane). In juvenile individuals that were overgrown by
adults the basal calcareous membrane is not developed.
The parietes of the wall plates are porous and multiple rows
of tubes are present (Fig. 8E). Large pores occur also in
the sheath (Fig. 8E). The radii are solid. Interlaminate figures
are arborescent, with a long straight axis and short transverse
branches with crescent-moon to kidney-shaped terminations
(Fig. 8F). Based on the general architecture of the six-plated
shell and the presence of complex interlaminate figures, these
specimens are provisionally identified as indeterminate
balanids. Among Balanidae, a porous sheath characterizes
Titobustillobalanus tubutubulus Carriol & Álvarez-Fernández,
2015 from the latest Pleistocene of Spain and various species
of the amphibalanine genus Fistulobalanus Zullo, 1984.
The specimens identified as Concavinae indet. are medium- to
small-sized, six-plated shells with a basal diameter ranging
between 1 and 2 cm (Fig. 8G–I). They appear to have been
strictly cluster-forming organisms, since even displaced single
individuals show an elongated, tubiferous, basal plate tapering
downwards (a morphology that is typically associated with
a gregarious growth habit, e.g., Newman & Ross 1976).
Some specimens display a faint pink to purple pigmentation.
The wall plates are porous and display a single row of tubes
divided by frequent transverse septa (Fig. 8H). Interlaminate
figures have a long and straight axis with numerous, closely-
spaced transverse branches characterized by arrow-shaped
terminations (Fig. 8I). These branches are often organized in
pairs, stemming from the same point of the main axis (Fig. 8I).
The very rare opercula retrieved in the sieved material may be
attributed to this group of specimens due to their size. They
consist of triangular scuta characterized by transverse growth
lines.
A further, currently unidentified, barnacle taxon was
observed only once in the Ct1a facies association. It is repre-
sented by small-sized individuals (around 0.5 cm in basal
diameter) forming a cluster which partially encrusts a pectinid
shell. These small-sized barnacles could also represent juve-
niles of one of the afore-mentioned taxa. The common
occurrence of pectinids with small barnacle attachment scars
(Anellusichnus circularis Santos, Mayoral & Muñiz, 2005) on
the outer surface could indicate that this type was less uncom-
mon than suggested by body fossils alone.
Discussion
Facies interpretation
Previous works based on coralline-algal and foraminiferal
assemblages suggest that both the Pietra da Cantoni and
the Sommières Basin barnacle facies developed in a tropical
setting (Vannucci et al. 1996; Coletti et al. 2015, 2017, 2018).
Vertebrate and mollusc assemblages indicate warm-temperate
conditions for the East Pisco Basin during the deposition of
the Ct1 beds of the Chilcatay formation (DeVries & Frassinetti
2003; Bianucci et al. 2018). The observation of the forami-
niferal genus Peneroplis, which is typical of warm and
warm-temperate water (Murray 2006), supports this hypo-
thesis. The skeletal assemblages observed in all the sites,
characterized by the abundance of filter feeding organisms
and the scarcity of symbiont-bearing taxa, in warm water
points to a nutrient-rich setting (Brasier 1995 a, b). This is also
in agreement with other works on the studied successions
(Dunbar et al. 1990; Reynaud & James 2012; Coletti et al.
2015, 2017; Bianucci et al. 2018).
Both barnacle facies of the Chilcatay Formation have a “low
diversity” skeletal assemblage, with barnacle and molluscs
(mainly epifaunal bivalves like ostreids and pectinids)
accounting for > 75 % of the skeletal fragments (Table 1).
It should be noticed that mollusc abundance was probably
reduced by selective dissolution during diagenesis; this parti-
cularly applies to the barnacle facies of Ct1b, where mol-
lusc-shaped, sparite-filled molds are common. The abundance
of barnacles and epifaunal bivalves points toward an exposed
setting since both groups are favoured by high water energy
(Farrow et al. 1978; Nebelsick 1992). A setting favourable for
barnacles is also supported by the diverse barnacle assemblage
with at least three common taxa. The preservation of barnacles
in both facies is similar. Whole individuals are present (either
as displaced clusters or single shells) and always separated
from their substrate. Some specimens still preserve their pig-
mentation but the majority of them do not. Opercula are
extremely rare. These characteristics suggest significant
reworking, compatible with a setting above fair-weather
wave-base. This hypothesis is in agreement with Bianucci et
al. (2018) and Di Celma et al. (2018b), which interpreted Ct1b
as a shoreface deposit, probably formed at a water depth of less
than 15–20 m (assuming a storm-wave base around 15–20 m
in accordance with the models of Hernández-Molina et al.
2000; Massari & Chiocci 2006). The same authors interpreted
Ct1a as an offshore deposit resulting from the downslope
transport of shoreface material (below 30–40 m of water depth
since the height of the clinoforms is about 15–20 m). Since
both facies have the same signature, it is likely that most of
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reworking occurred in the shoreface environment, with
the down slope movement only mildly affecting the associa-
tion. This is also supported by the sediment preserved within
the shells of the Ct1a barnacles, which is similar in composi-
tion to that of Ct1b (more bioclasts and less siliciclastic
particles).
The skeletal assemblage of the barnamol facies of the Sandy
Molasse Unit is also overwhelmingly dominated by barnacles
and epifaunal bivalves, suggesting a high-energy setting
(Table 1; Farrow et al. 1978; Nebelsick 1992). Extensive evi-
dence of dissolution was not observed in this facies, sugges-
ting that the mollusc abundance was not significantly reduced.
Suitable conditions for barnacles are suggested by the pre-
sence of a diverse cirripede assemblage. There are no complete
shells and well-preserved specimens in general are missing;
opercula are also extremely rare. The poor state of preserva-
tion of the barnacle remains and the cross-bedded structure of
this facies are consistent with a very shallow and proximal
marine environment (nearshore zone, less than 15 m of water
depth). This interpretation is in agreement with previous sedi-
mentological studies on the Boisseron outcrop of the Sandy
Molasse Unit (Reynaud & James 2012). The environment was
probably characterized by higher hydrodynamic energy than
that of the Chilcatay Formation.
The barnalgal of the Pietra da Cantoni significantly diverge
from the other barnacle facies. Siliciclastic elements are much
rarer (Table 1), ruling out the presence of nearby rocky cliffs.
Its skeletal assemblage is more diverse, coralline algae are
codominant, benthic foraminifera are more diverse, and there
are fewer molluscs and echinoids than in the other facies
(Table 1). There is no extensive evidence of dissolution sug-
gesting that the low abundance of molluscs is not a result of
diagenetic processes (although leaching of aragonite might
have occurred since gastropods are absent even thought
gastropods predation holes are present on barnacles). Unlike
the other facies only a single taxon of barnacle was recog-
nized. Differing from the other facies, most of the shells are
well preserved, retain their original colour and are associated
with their substrate (i.e., the rhodoliths). Opercula are abun-
dant and only limited reworking can be inferred by the unequal
ratio of terga and scuta (the heavier scuta being more common
than the lighter terga). These features suggest a less exposed
environment, below the fair-weather wave base (probably
between 20 and 40 m of water depth). This is in agreement
with previous studies that interpreted this material as a short-
distance mass-transport of inner-middle ramp material depo-
sited in a slightly deeper middle-ramp setting (below 50–60 m
of water depth; Schüttenhelm 1976; Coletti et al. 2015). Short-
distance transport and rapid burial, without further reworking,
are strongly supported by the preservation of colour in barna-
cles (Hollingworth & Barker 1991; Aguirre et al. 2008).
The detailed comparison of skeletal assemblages, petro-
graphic characteristics and barnacle preservation and diversity
clearly separates the barnalgal facies from the remaining three
facies. The latter have a diverse barnacle assemblage mainly
associated with epifaunal bivalves and echinoids, and
comprise an important siliciclastic fraction. The barnalgal do
not. The French and Peruvian facies are related to a very shal-
low, nearshore carbonate factory, developed along a high-
energy rocky coast. This is the ideal setting for encrusting
barnacles; they are perfectly adapted to systems where hard
surfaces are abundant. The environment of the barnalgal facies
deviates substantially from this optimum. There are less avai-
lable surfaces and most of them are coralline algae, which
directly compete with barnacles for space (as suggested by
the presence of alternating layers of barnacles and coralline
algae in the rhodoliths). This might explain why various dif-
ferent species of barnacles are found in the first group of facies
and only a single species characterizes the barnalgal facies.
Environmental controls on barnacle facies
Modern barnacle facies occur from the Poles to the Equator
and, notwithstanding this large latitudinal variation, they show
clear similarities (see Table 2 for the complete references list
and Fig. 9 for the locations). Barnacle facies usually contain
abundant molluscs (Table 2; Hoskin & Nelson 1969; Milliman
1972; Müller & Milliman 1973; Farrow et al. 1978; Scoffin
1988; Wilson 1988; Halfar et al. 2006; Westphal et al. 2010;
Michel et al. 2011; Reijmer et al. 2012; Reymond et al. 2016).
Echinoids are important contributors in most of the occur-
rences (Table 2; Hoskin & Nelson 1969; Farrow et al. 1978;
Scoffin 1988; Wilson 1988; Michel et al. 2011). At low lati-
tudes, the association with hermatypic corals is also possible
(Table 2; Glynn & Wellington 1983; Halfar et al. 2006;
Reymond et al. 2016). Coralline algae and benthic foramini-
fera are practically absent (Table 2; Hoskin & Nelson 1969;
Farrow et al. 1978; Scoffin 1988; Wilson 1988; Halfar et al.
2006; Westphal et al. 2010; Michel et al. 2011; Reijmer et al.
2012; Reymond et al. 2016). Barnacle facies have an impor-
tant siliciclastic fraction, which accounts for at least 10 % of
the grains, although it is typically much higher (Table 2;
Hoskin & Nelson 1969; Milliman 1972; Müller & Milliman
1973; Farrow et al. 1978; Scoffin 1988; Scoffin and Bowes
1988; Wilson 1988; Halfar et al. 2006; Westphal et al. 2010;
Michel et al. 2011; Reijmer et al. 2012; Frank et al. 2014;
Reymond et al. 2016). As far as the depositional environment
is concerned, most modern barnacle facies are related to shal-
low-water (less than 50 m and generally less than 20 m), high-
energy environments with nearby rocky outcrops (Table 2;
Hoskin & Nelson 1969; Farrow et al. 1978; Scoffin 1988;
Wilson 1988; Henrich et al. 1995; Halfar et al. 2006; Westphal
et al. 2010; Michel et al. 2011; Reijmer et al. 2012; Reymond
et al. 2016). Most barnacle facies are also related to plank-
ton-rich water (Table 2; Müller& Milliman 1973; Taviani et al.
1993; Henrich et al. 1995; Westphal et al. 2010; Michael et al.
2011; Reijmer et al. 2012; Klicpera et al. 2013; Frank et al.
2014; Reymond et al. 2016).
Those occurrences that significantly deviate from this gene-
ral model are located at polar latitudes and in bathyal settings
(Table 2). The barnacle facies of the Barents Sea are characte-
rized by almost pure carbonates, composed of barnacles and
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benthic foraminifera (Henrich et al. 1995). Unlike the exam-
ples discussed in detail herein, carbonate production is loca-
lized far away from the coast, on shallow submerged rocky
shoals colonized by kelp forests (Henrich et al. 1995). The Ross
Sea barnacle facies develop on hard substrates under high
energy conditions; however, they are dominated by Bathy
lasma corolliforme (Hoek, 1883) that lives in deeper water
than most other encrusting barnacles (Taviani et al. 1993;
Frank et al. 2014). Bathylasmatidae-dominated facies occur
also in even deeper, bathyal, settings (Table 2; Newman &
Ross 1976; Buckeridge 1999). They are found close to sea-
mounts and, unlike their shallow-water counterparts, they are
associated with fine grained sediments and deep-sea fauna
(deep-water corals, bryozoans and planktonic foraminifera;
Location
Age
Key
references
Barnacle taxa
Associated bioclasts
Texture
Clastic
particles
Environmental information
Antartica, Ross
Sea Shelf
Pleistocene–
Recent
Frank et al.
2014; Taviani
et al. 1993
Bathylasma
corolliforme
(Hoek, 1883)
Locally only composed by
barnacles; minor local contributions
from stylasterine hydrocorals,
bryozoans and benthic foraminifera;
rare bivalves and serpulids.
According to Taviani et al. (1993)
barnacles are mainly associated with
foraminifera
Coarse to
very coarse 0 to 40 %
Polar; more than 350 m of
water depth; high energy;
plankton-rich water
South Pacific
Ocean, South
Tasman Rise
Holocene-
Recent
Buckeridge
1999
Tetrachaelasma
tasmanicum
Buckeridge,
1999
Mainly solitary deep-water
scleractinian corals, but also
planktonic foraminifera
Very fine to
coarse
Present, both
rocks and
mud
Bathyal environment, 2100-
3600 m of water depth
South
Madagascar;
Indian Ocean
Recent
Newman &
Ross 1976
Tetrachaelasma
Newman & Ross,
1971
///
Coarse
///
Bathyal environment, 2000 m
of water depth
Barents Sea,
Spitsbergenbank Recent
Henrich et al.
1995
Balanus
crenatus
Bruguière, 1789
Mainly benthic foraminifera
Coarse
Pure
carbonate
Polar; between 6 and 20 m of
water depth; high energy;
plankton-rich water
Alaska,
Alexander
Archipelago
Recent
Hoskin &
Nelson 1969
///
Major contributions from molluscs
and echinoids; minor contributions
from benthic foraminifera; rare
ahermatypic corals, coralline algae
and bryozoans
Fine to
coarse
15 % to 60 % Cold sub-polar; macrotidal
setting
Canada,
Newfoundland
Recent/Sub-
recent
Müller and
Milliman
1973
///
Major contribution from molluscs
Coarse
50 % to 80 %
Cold sub-polar; 60 to 80 m of
water depth; plankton-rich
water
USA, locally
along Carolina
and Florida
Atlantic shelves
Recent/Sub-
recent
MacIntyre &
Milliman
1970;
Milliman
1972
Balanus calidus
Pilsbry, 1916
(temptative)
Mainly coralline algae (locally
codominant)
///
3 % to 74 %,
39 % on
average
Barnacles growing on a relict
algal ridge that borders the
upper slope and the outer shelf
edge (50-150m); strong
currents.
USA, locally
along Carolina
and Florida
Atlantic shelves
Recent/Sub-
recent
Milliman
1972
Balanus calidus
(temptative)
Mainly molluscs (locally
codominant)
///
3 % to 94 %,
45 % on
average
Barnacles growing on
submerged rocky outcrops
U.K., western
Scotland
Recent
Farrow et al.
1978; Scoffin
1988; Wilson
1988
Semibalanus
balanoides
(Linnaeus, 1767)
Codominance with molluscs;
important contributions from
echinoids; minor contributions from
serpulids and coralline algae; rare
bryozoans and benthic foraminifera.
According to Wilson (1988) the
contributions from serpulids can be
very important
Coarse
0 % to 50 %,
with most of
the values
around 20 %
Cool-temperate; less than 50 m
of water depth; hard substrate;
high energy
Mexico, Bahía
de Los Angeles
Recent
Halfar et al.
2006
///
Major contributions from molluscs
and hermatypic corals; minor
contributions from bryozoans and
echinoids; rare serpulids; very rare
coralline algae and barnacles
Fine to
coarse
30 % to 70 %
Warm-temperate; upwelling
currents; high nutrient
concentration (eutrophic); less
than 20 m of water depth; hard
substrate; high energy
Pacific side of
Panama, Gulf of
Panama
Recent
Reijmer et al.
2012
///
Mainly bivalves; common
gastropods; minor contributions
from echinoids, serpulids, benthic
foraminifera and coralline algae
Coarse
At least 10 %
Tropical; upwelling currents;
high nutrient concentration
(eutrophic); less than 50m of
water depth
Mauritania,
Banc D'Arguin
Recent
Westphal et
al. 2010;
Michel et al
2011;
Klicpera et al.
2013
Balanus Costa,
1778
Codominance with bivalves;
significant contributions from
echinoids; rare gastropods and
benthic foraminifera; very rare
bryozoans
Medium
30 % to 50 %
Tropical; upwelling currents;
high nutrient concentration
(eutrophic); sub-tidal, less than
20 m of water depth; high
energy
Chile, Galápagos
Archipelago
Recent
Glynn &
Wellington
1983;
Westphal et
al. 2010;
Reymond et
al. 2016
Megabalanus
tintinnabulum
(Linnaeus, 1758)
[“barnamol”] Important gastropods
and serpulids; minor contributions
from bryozoans, bivalves, echinoids
and coralline algae
Fine to
coarse
10 % to 70 %
Equatorial; strong upwelling;
less than 15 m of water depth;
mesotrophic
[“barnamolcor”] Codominance with
gastropods; important contributions
from coralline algae and corals;
minor contributions from echinoids,
serpulids, bryozoans and bivalves
10 % to 50 %
Equatorial; moderate
upwelling; less than 15 m of
water depth; oligotrophic
Table 2: Sub-recent and recent barnacle facies, including information on the position, barnacle assemblage, skeletal assemblage, texture,
clastic fraction and available environmental information.
586
COLETTI, BOSIO, COLLARETA, BUCKERIDGE, CONSANI and EL KATEB
GEOLOGICA CARPATHICA
, 2018, 69, 6, 573–592
Newman & Ross 1976; Buckeridge 1999). The barnalgal asso-
ciation described by Milliman (1972) along the South-eastern
Coast of the United States is also peculiar and differs from
most of the other barnacle facies. It is related to the recent
barnacle colonization of a relict algal ridge growing during
the last glacial period that now borders the outer shelf edge.
Neogene and Quaternary barnacle facies follow the same
pattern as their modern counterparts (Table 3; Fig. 9). The ske-
letal assemblage is mainly characterized by barnacles and
molluscs, with common bryozoans and echinoids, rare benthic
foraminifera and coralline algae; the siliciclastic fraction is
also important (Table 3; Sakai 1987; Kamp et al. 1988;
Nebelsick 1989, 1992; Doyle et al. 1997; Betzler et al. 2000;
Nielsen & Funder 2003; Aguirre et al. 2008; Nomura & Maeda
2008; Stanton & Alderson 2013; Brandano et al. 2015;
Buckeridge et al. 2018). They are generally interpreted as
high-energy shallow-water deposits related to non-oligotro-
phic conditions (Table 3; Buckeridge 1983; Sakai 1987; Kamp
et al. 1988; Nebelsick 1989, 1992; Doyle et al. 1997; Nielsen
& Funder 2003; Civitelli & Brandano 2005; Aguirre et al.
2008; Nomura & Maeda 2008; Stanton & Alderson 2013;
Brandano et al. 2015).
When considering the distribution of both modern and
fossil barnacle concentrations, the major factors controlling
the deve lopment of a barnacle-dominated carbonate factory
seem to be the presence of suitable hard substrates, hydro-
dynamic energy and nutrient availability. Although barnacle
accumulations are more common in the cool-temperate and
cold realms, it seems that the lack of nutrients is the main fac-
tor that limits their distribution in the tropical zone. Therefore,
an inshore rocky substrate characterized by high-energy con-
ditions and abundant nutrient-supply is very favourable to
the development of a barnacle-dominated carbonate factory.
This setting represents an environmental optimum for barna-
cles (Sanford & Menge 2001) and other hard-substrate orga nisms
(e.g., ostreids and pectinids), and can lead to the formation of
classical barnamol facies carbonates (sensu Hayton et al.
1995). Coralline algae might occur in this environment, but
mostly as thin adherent crusts, with a poor preservation poten-
tial. Foraminifera can also occur, but they prone to be swept
away by currents and deposited elsewhere (Farrow et al. 1978;
Nebelsick 1992). The barnacle-rich facies investigated in this
study comply with this model, except for the barnalgal facies.
The latter represents a more atypical setting, located in slightly
deeper water and where the surface available for barnacle
colo nization is more limited. Facies similar to those of
the Pietra da Cantoni Group, with barnacle and coralline algae
growing together, occur in the lower Miocene Latium-Abruzzi
carbonate platform, where they have been interpreted as inner
ramp deposits (Civitelli & Brandano 2005). Barnalgal assem-
blages are also reported in modern oceans, well below the fair
weather wave base (Table 2; Macintyre & Milliman 1970;
Milliman 1972). However, they do not represent an example
of barnacles and coralline algae growing together but rather
a case of barnacles colonizing a hard substrate provided by
relict coralline algal bioconstructions (Milliman 1972).
Although atypical barnacle facies are uncommon, they occur
in both modern oceans and in the sedimentary record, thus
indicating that dismissal of all barnacle concentrations as
the result of nearshore carbonate factories might lead to erro-
neous palaeoenvironmental reconstructions. A perfect exam-
ple includes Bathylasmatidae-rich facies, which are dominated
by barnacles while being related to bathyal settings
(Buckeridge 1975, 1999; Newman & Ross 1976). The study of
the barnacle assemblage, as demonstrated herein, can help
in identifying these atypical situations. Furthermore, while
the hard substrate where barnacle carbonate factories normally
develop is a major site of skeletal production, due to the high
hydraulic energy, few bioclasts are preserved there (Scoffin
1988; Henrich et al. 1995). Most of the material is swept away
and deposited in the closest sheltered areas (Scoffin 1988;
Henrich et al. 1995). Remarkable transport processes are also
reported in fossil barnacle facies, and can lead to significant
displacement of the fossils (e.g., Buckeridge et al. 2018).
Therefore, before making palaeoenvironmental assumptions
based on barnacles, their preservation should be carefully
evaluated in order to determine how far the material has been
reworked and transported prior to burial.
Conclusions
Four barnacle facies from three different Burdigalian suc-
cessions have been analysed and divided into two groups on
the basis of skeletal assemblages and barnacle preservation
and diversity. The first group includes the barnamol facies of
the Sandy Molasse Unit of the Sommières Basin (France) and
the two barnacle facies of the Chilcatay Formation of the East
Pisco Basin (Peru). These facies are overwhelmingly domi-
nated by shallow-water hard-substrate biota, including diffe-
rent species of barnacles, molluscs (mainly ostreids and
pectinids) and echinoids; they are also characterized by
an important siliciclastic fraction. Skeletal assemblages,
Fig. 9. World map including the location of the fossil (Neogene and
Quaternary) and recent/sub-recent barnacle facies. Circles represent
recent/sub-recent facies; squares indicates fossil facies; stars identify
the early Miocene facies investigated in this study.
587
MIOCENE BARNACLE FACIES FROM EUROPE AND SOUTH AMERICA
GEOLOGICA CARPATHICA
, 2018, 69, 6, 573–592
barnacle preservation and sedimentary structures suggest that
the bioclasts originated in a high-energy rocky shoreface envi-
ronment (less than 15–20 m of water depth), where reworking
of bioclasts was significant. The second group includes
the barnacle and coralline algae facies of the Pietra da Cantoni
Group of the Tertiary Piedmont Basin (Italy). For this peculiar
skeletal assemblage, rarely reported from the fossil record,
the new name “barnalgal” is proposed. The barnalgal is
charac terized by a single barnacle species, abundant coralline
algae, benthic foraminifera and almost no siliciclastic ele-
ments. Compared to the other facies, barnacle shells are also
better preserved. The related carbonate factory probably
developed in slightly deeper conditions (20–40 m of water
depth) than those responsible for the first group of barnacle
facies. In this setting, the only hard substrate available for bar-
nacles were rhodoliths, leading to the formation of this atypical
association. Reworking was also less important, reducing
the abrasion of the specimens and leading to the conservation
of abundant barnacle opercula in the sediment.
The analysis of both modern and fossil barnacle facies sug-
gests that the major factors controlling the development of
a barnacle-dominated carbonate factory are the availability of
Location
Age
Key
references
Barnacle taxa
Associated bioclasts
Texture
Clastic
particles
Palaeoenvironmental
interpretation
Southeastern
Japan,
Shikoku
Oligocene
to Miocene Sakai 1987
///
Very minor contributions from
molluscs, bryozoans, echinoids,
sponges and foraminifera
Coarse
Highly
impure
limestone
Shallow-water; high-energy
Austria,
Bohemian
Massif
Early
Miocene
Nebelsick
1989, 1992
///
Codominance with bivalves
(pectinids and ostreids), important
contribution from bryozoans and
serpulids
Very coarse
50 %
Shallow-water; near-shore; high-
energy
Central Italy
(several
sites)
Early
Miocene
Civitelli &
Brandano
2005
///
Molluscs, coralline algae,
echinoids, bryozoans, serpulids,
ostracods and benthic foraminifera
(both large- and small-sized forms)
Very coarse
Almost pure
limestone
Tropical; shallow-water inner
ramp; hard substrate; high
energy; high nutrient
concentration
Motutapu
Island, New
Zealand
Early
Miocene
Buckeridge
1975
Hexelasma
aucklandicum
(Hector, 1888)
Bryozoans, molluscs, corals
Fine
Abundant
Deep-water; low-energy
Japan,
Sendai
Early to
middle
Miocene
Nomura &
Maeda 2008
Arossia sendaica
(Hatai et al., 1976)
(frequent); Balanus
sulcatus Bruguière,
1789 (frequent); B.
crenatus Bruguière,
1789; B. rostratus
(Hoek, 1883)
Minor contributions from
molluscs, serpulids and bryozoans;
rare brachiopods, foraminifera,
echinoids and corals
Very coarse
>>50 %
Very shallow-water; high-energy
California,
Santa
Monica
Mountains
Early–
middle
Miocene
Stanton &
Alderson
2013
///
Codominance with bivalves; minor
contributions from echinoids,
serpulids, bryozoans; rare
gastropods and brachiopods; very
rare coralline algae
Fine to
coarse
Impure
limestone
Warm-temperate; inner shelf;
moderate-energy; hard substrate;
high productivity
Italy, Liguria Middle
Miocene
Brandano et
al. 2015
///
Bryozoans, serpulids, echinoids,
benthic foraminifera (including
symbiont-bearing Amphistegina
d’Orbigny, 1826), bivalves; minor
contributions from Halimeda
Lamouroux, 1812
Very coarse
From more
than 50 % to
around 20 %
Humid tropical; high terrigenous
input; near-shore shallow-water,
above fair-weather wave base;
high energy; high nutrient supply
Spain,
Almeria
Late
Miocene
Doyle et al.
1997
Megabalanus cf.
tintinnabulum
(Linnaeus, 1758)
Minor contributions from bivalves,
echinoids and bryozoans
Very coarse
>50 %
Very shallow-water
Spain,
Almeria
Late
Miocene
Betzler et al.
2000
///
Minor contributions from bivalves,
bryozoans; coralline algae,
echinoderms, gastropods; rare
ostracods
Very coarse
///
More than 15m of water depth;
moderate energy
Spain,
Almeria
Pliocene
Aguirre et al.
2008
Concavus concavus
(Bronn, 1831)
(dominant);
Perforatus
perforatus
(Bruguière, 1789);
Megabalanus Hoek,
1913
Mainly bivalves but also bryozoans Coarse to
very coarse
Up to 50 %
High energy; near-shore shallow-
water; high nutrient supply
Greece,
Rafina
Pliocene
Radwańska &
Radwański
2008
Concavus concavus
///
///
///
///
New
Zealand,
North Island
(several
sites)
Pliocene–
Pleistocene
Hayton et al.
1995; Kamp
et al. 1988;
Buckeridge
1983;
Buckeridge
2015;
Buckeridge et
al. 2018
Notobalanus
Newman & Ross,
1976;
Austromegabalanus
Newman, 1979;
Balanus Costa,
1778; Fosterella
Buckeridge, 1983
Mainly bivalves, but also
bryozoans; minor contributions
from echinoids and small benthic
foraminifera; rare coralline algae;
brachiopods and solitary corals
Coarse to
very coarse
10 % –20 %
for most of
the samples
Cold- to warm-temperate; high
nutrient concentration; sub-tidal,
less than 50–30 m of water
depth; hard or coarse-grained
substrate; high-energy seaway;
locally important downslope
transport of sediment
Table 3: Neogene and Quaternary fossil barnacle facies, including information on the position, age, barnacle assemblage, skeletal assemblage,
texture, clastic fraction and palaeo-environmental interpretation.
588
COLETTI, BOSIO, COLLARETA, BUCKERIDGE, CONSANI and EL KATEB
GEOLOGICA CARPATHICA
, 2018, 69, 6, 573–592
hard substrate, hydrodynamic energy and nutrient availability.
Consequently, the most favourable setting for a barnacle-
dominated carbonate factory is probably an inshore rocky sub-
strate characterized by high-energy conditions and abundant
nutrient supply. This situation can lead to the formation of
the typical barnacle-rich deposit, where abundant barnacles are
associated with other hard substrate organisms (e.g., the bar-
namol assemblage). The first group of facies recognized in
this paper, embracing the French and Peruvian case studies,
can be easily included in this category. The Italian barnalgal
facies, on the other hand, represents an atypical barnacle facies
related to situations that deviate from the aforementioned
environmental optimum. Similar atypical situations can be
identified throughout careful analyses of both the skeletal
assemblage and the barnacle association, highlighting
the importance of barnacle palaeontology for palaeoenviron-
mental reconstructions in shallow-water settings.
Acknowledgements: The authors are grateful to Claudio
Di Celma (Università degli Studi di Camerino) for his assis-
tance during the preparation of the manuscript. Special thanks
go to Alfredo Frixa (Eni Spa), Jean Yves-Reynaud (Université
de Lille) and Mario Urbina (Museo de Historia Natural de la
Universidad Nacional Mayor de San Marcos) for their help
during the field work in the Pietra da Cantoni area (Italy) and
Sommières Basin (France), and East Pisco Basin (Peru),
respectively. Roberto Badano (Università di Genova) is grate-
fully acknowledged for his support with the XRD analyses.
The authors acknowledge Silvia Spezzaferri (Université de
Fribourg) for her help with the identification of foraminifera.
The authors are also grateful to Daniela Basso and Elisa
Malinverno (both at Università di Milano Bicocca), Giovanni
Bianucci (Università di Pisa) and Thomas J. DeVries (Univer-
sity of Washington) for their suggestions and fruitful discus-
sions. Comments and suggestions by Milan Kohút (Ústav vied
o Zemi Slovenskej akadémie vied), Matúš Hyžný (Univerzita
Komenského v Bratislave), Tomáš Kočí (Národní muzeum v
Praze), Mathias Harzhauser (Naturhistorisches Museum
Wien), and an anonymous reviewer greatly improved the qua-
lity of this work. Fieldwork by Giulia Bosio and Alberto
Collareta in the East Pisco Basin was supported by a grant
from the Italian Ministero dell’Istruzione, dell’Università
e della Ricerca [PRIN Project, 2012YJSBMK] to Giovanni
Bianucci, Claudio Di Celma and Elisa Malinverno, and by
a grant from the Università di Pisa to Giovanni Bianucci
[PRA_2017_0032].
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