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A forgotten cirripedological gem: a new species of whale barnacle of the genus Cetopirus from the Pleistocene of the United States West Coast

  • Royal Belgian Institute of Natural Sciences and Koninklijk Zeeuwsch Genootschap der Wetenschappen, Middelburg, The Netherlands

Abstract and Figures

A small lot of fossil whale barnacles from the Upper Pleistocene of California and the Middle Pleistocene (Chibanian) of Oregon (United States West Coast), described in a 1972 unpublished MA thesis, are formally described and illustrated herein. In that thesis, a new genus and species name were proposed; however, according to the International Code of Zoological Nomenclature, they have no taxonomic standing and are thus unavailable. Based on our reappraisal, two specimens in this lot belong to a new, extinct species that can be assigned to the purportedly extant genus Cetopirus. Cetopirus polysyrinx sp. nov. differs from congeners in that its secondary T-shaped flanges are multitubiferous internally, that is, they are perforated by a high number of irregularly-sized and irregularly-spaced tubules that result in a spongy aspect in transverse section. Whether or not this peculiar condition had any adaptive significance is difficult to determine. Considering that Cetopirus is currently known as an obligate epibiont of right whales (including the North Pacific form Eubalaena japonica (Lacépède 1818)), the host of C. polysyrinx sp. nov. was E. japonica or some other species of Eubalaena. The Plio-Pleistocene deposits of the Pacific coast of North America have yielded a rather idiosyncratic fossil whale barnacle fauna, inclusive of the genera Cetolepas, Cryptolepas and now Cetopirus, which seemingly contrasts with all other coeval assemblages worldwide, the latter being in turn dominated by Coronula spp.
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Spontaneous Article
A forgotten cirripedological gem: a new species of whale
barnacle of the genus Cetopirus from the Pleistocene of the
United States West Coast
Alberto COLLARETA1,2* , Mark BOSSELAERS3,4, Patricia A. HOLROYD5
and Ashley DINEEN5
Dipartimento di Scienze della Terra, Università di Pisa, via S. Maria 53, 56126 Pisa, Italy.
Museo di Storia Naturale, Università di Pisa, via Roma 79, 56011 Calci, Italy.
Koninklijk Belgisch Instituut voor Natuurwetenschappen, Operationele Directie Aarde en Geschiedenis van het
Leven, Vautierstraat 29, 1000 Brussel, Belgium.
Koninklijk Zeeuwsch Genootschap der Wetenschappen, Kousteensedijk 7, 4331 JE, PO Box 378, 4330 AJ
Middelburg, the Netherlands.
University of California Museum of Paleontology, 1101 Valley Life Sciences Building, Berkeley, CA 94720-4780,
*Corresponding author. Email:
ABSTRACT: A small lot of fossil whale barnacles from the Upper Pleistocene of California and the
Middle Pleistocene (Chibanian) of Oregon (United States West Coast), described in a 1972 unpublished
MA thesis, are formally described and illustrated herein. In that thesis, a new genus and species name
were proposed; however, according to the International Code of ZoologicalNomenclature, they have no
taxonomic standing and are thus unavailable. Based on our reappraisal, two specimens in this lot belong
to a new, extinct species that can be assigned to the purportedly extant genus Cetopirus.Cetopirus poly-
syrinx sp. nov. differs from congeners in that its secondary T-shaped anges are multitubiferous intern-
ally, that is, they are perforated by a high number of irregularly-sized and irregularly-spaced tubules that
result in a spongy aspect in transverse section. Whether or not this peculiar condition had any adaptive
signicance is difcult to determine. Considering that Cetopirus is currently known as an obligate epi-
biont of right whales (including the North Pacic form Eubalaena japonica (Lacépède 1818)), the host of
C. polysyrinx sp.nov.wasE. japonica or some other species of Eubalaena. The Plio-Pleistocene deposits
of the Pacic coast of North America have yielded a rather idiosyncratic fossil whale barnacle fauna,
inclusive of the genera Cetolepas,Cryptolepas and now Cetopirus, which seemingly contrasts with all
other coeval assemblages worldwide, the latter being in turn dominated by Coronula spp.
KEY WORDS: California, Cetopirus polysyrinx sp. nov., Chibanian, Cirripedia, Coronulidae,
epibiosis, Oregon, taxonomy, Upper Pleistocene.
1. Introduction
Fifty years ago, Clark William Davis, a graduate student at
San Francisco State University, USA, authored one of the
most relevant contributions to the modern understanding of
the skeletal anatomy and functional morphology of the extant
and extinct whale barnacles (Cirripedia: Coronulidae) in the
form of a MA thesis (Davis 1972) that is currently available
online (Collareta et al.2022a). In that thesis, entitled Studies
on the barnacles epizoic on marine vertebrates, Davis (1972)
informally described a new genus and species of whale barnacle,
Polysyrinx zulloi, on the basis of a handful of fossils from the
United States West Coast. However, both the names Polysyrinx
and P. zulloihave not been formally published and are thus
unavailable in light of the International Code of Zoological
Nomenclature (1999: articles 8 and 9; see also Dubois et al.
2013), whereas they are published herein as nomina nuda (Inter-
national Code of Zoological Nomenclature 1999: articles 11 and
13, and glossary). While we were unable to track down the author
of this particular MA thesis, we did succeed in tracing three out
of four specimens referred by Davis (1972)toPolysyrinx zulloi.
Below we describe Davisnew coronulid taxon on solid nomen-
clatural grounds and discuss its palaeontological signicance.
2. Material and methods
2.1. Institutional abbreviations
CAS: California Academy of Sciences (San Francisco, Califor-
nia, USA); LACM: Natural History Museum of Los Angeles
(Los Angeles, California, USA), in the Invertebrate Paleon-
tology (=LACMIP) department; UCMP: University of Califor-
nia Museum of Paleontology (Berkeley, California, USA).
2.2. Digital imaging
Two of the specimens studied herein were scanned by one of us
(PAH) on a Phoenix Nanotom-M nanofocus X-ray computed
tomography (CT) system in the Functional Anatomy and
© The Author(s), 2022. Published by Cambridge University Press on behalf of The Royal Society of Edinburgh. doi:10.1017/S1755691022000214
Earth and Environmental Science Transactions of the Royal Society of Edinburgh,18, 2022 Published online by Cambridge University Press
Vertebrate Evolution Laboratory at the University of California-
Berkeley at a resolution of 10 microns per pixel. Reconstructed
slices were cropped in Fiji 1.53q (Schindelin et al.2012), rendered
in three dimensions using Slicer 4.11 (Fedorov et al.2012) using
the SlicerMorph extension (Rolfe et al.2021), segmented and the
segmentation exported as an STL le. The resultant STL le was
imported into Meshlab (Cignoni et al.2008) and decimated to
approximately one-third the number of original faces to produce
a smaller, web-viewable PLY le using their variant of quadric
edge-collapse decimation (Garland & Heckbert 1997). Both
the cropped CT-stack and decimated PLY les were uploaded
3. Geological framework
The whale barnacle fossils dealt with in the present paper are
represented by three specimens, including the holotype and a sin-
gle paratype of a new species of Cetopirus, plus a third specimen
that is left herein in open nomenclature.
As reported by Davis (1972), one of these originated from
LACM locality 1210, San Pedro, Los Angeles County, Califor-
nia, USA (Fig. 1). Also referred to as LACMIP 1210, this local-
ity is described in the museum records as a bed of fossil shells in
unconsolidated sand thickening from 2 feet at west end to 6 feet
at east end; where it is underlain by single layer of cobble con-
glomerate and has some cross bedding in layers with fossils
above.This bed no longer appears to be accessible, but did
occur on the small bluff formed by the Gaffey Syncline, between
modern-day N. Gaffey Street and the West Basin in the north-
western part of the town of San Pedro, approximately where
Westmont Drive and industrial buildings sit today. The locality
was mapped in Wehmiller et al.(1977;g. 2h) and Muhs et al.
(1992;g. 3). Woodring et al.(1946) recognised the deposits
cropping out in this area as being entirely part of their rst ter-
raceof the Palos Verdes Sand Formation. Wehmiller et al.(1977)
recorded amino acid stereochemistry values from the bivalve
genus Saxidoma from LACM 1210 and assigned the locality to
the early part of the Marine Isotope Stage (MIS) 5. Muhs
et al.(1992) showed that faunas from Woodring et al.s(1946)
rst terracerepresented two temporally different highstands,
with LACM 1210 being assigned to the 125 ka highstand (i.e.,
MIS 5e). Based on the most recent synthesis of data bearing
on the age of MIS 5e sites along the west coast of North America
(Muhs et al.2021; Muhs 2022), the age of locality LACM 1210 is
estimated to be 130 to 115 ka.
According to Davis (1972), two additional specimens were
recovered from UCMP localities B7376 and B7380, near Cape
Blanco, Curry County, Oregon, USA (Fig. 1). With regard to
the former locality, it was described as in the same horizon as
B7375 and about 100 feet south. Both UCMP B7375 and
B7376 were collected by Jack A. Wolfe and Victor A. Zullo dur-
ing the same eld trip, but data on geology and location were
recorded only on the locality card for B7375. UCMP B7375
occurs in a well-sampled stretch of uplifted terraces exposed on
cliffs north of the mouth of the Elk River, Curry County,
Oregon, USA, that have produced diverse invertebrate assem-
blages and rarer marine mammals. The invertebrate assemblages
were the topic of an unpublished PhD thesis by Roth (1979) and
the vertebrate faunas were most recently reviewed by both
Boessenecker (2013a) and Welton (2015). Locality UCMP
B7375 was collected by Wolfe and Zullo in March 1960 from a
unit described in the UCMP records as 25 feet below the con-
glomerate in the lower buff sandstoneapproximately 100
feet along the cliff. Topographically, the site corresponds to the
area where Welton (2015) measured his Section 1 at Gold-
washers Gully, the northernmost of four sections, and contain-
ing the vertebrate localities UCMP V74042 and NMMNH
9241. The locality description suggests that the site is in the
Lower Brown Sandstone Member, aninterpretationalsoindicated
in Roth (1979). This unit falls within all modern interpretations of
the Port Orford Formation (e.g., Boessenecker 2013a;Wileyet al.
2014; Welton 2015), although older literature and records (e.g.,
museum locality registers; Lefer 1964;Davis1972; Roth 1979)
refer to this unit as part of the Elk River Formation.
UCMP B7380 occurs in the same stretch of uplifted terraces as
UCMP B7375 and B7376, between Weltons(2015) sections 3
and 4, where the lower part of the Port Orford Formation is cov-
ered (see also Boessenecker, 2013a;g. 1.4 for an outcrop photo-
graph). Fossils were collected in situ from the Upper Brown
Sandstone Member by Wolfe and Zullo in March 1960 and
again in 1961 by a class eld trip from the University of Califor-
nia, Department of Paleontology. UCMP B7380 is stratigraphic-
ally higher than B7375 and B7376, being separated from the
Lower Brown Sandstone Member by approximately 14 m of
the Blue Argillaceous Sandstone Member. Welton (2015) fol-
lowed Baldwin (1945,1959) in including the Upper Brown Sand-
stone Member in the Elk River Beds, whereas Boessenecker
(2013a) and Wiley et al.(2014) considered it to be part of the
Port Orford Formation. Whichever stratigraphic nomenclature
is used, molluscs from the Upper Brown Sandstone Member
(from LACM locality 3960) were dated to 500,000 ± 100,000
years via amino acid racemisation, and the rocks are reported
to have normal palaeomagnetic polarity (Wehmiller et al.
1978). Kennedy et al.(1982) also reported a potentially younger
age interpretation (early MIS 3) of a terrace at Cape Blanco on
the basis of amino acid racemisation, but also noted that the
fauna was otherwise indistinguishable from the cool-water
fauna associated with late stage 5 localities. Both studies indicate
that these rocks and their associated faunas are Chibanian (Mid-
dle Pleistocene) in age, although currently available data do not
permit greater renement.
In addition to the specimens described here, Roth (1979) noted
the presence of a single plate from this same area at UCMP local-
ity B7371, in the Upper Brown Sandstone Member. He referred
it to Cetopirus complanatus (Mörch 1853), but the specimen was
not formally catalogued and could not be located at the UCMP.
Roth (1979) also noted that the record of Coronula (Cetopirus)in
the Pleistocene of Oregon listed by Newman et al.(1969)was
likely based on his specimen, although we cannot exclude the
possibility that they were referring to the specimens described
here, especially as Zullo, being co-author of the paper by New-
man et al.(1969), was also one of the collectors. Interestingly,
the occurrence of Coronula complanata (=Cetopirus complana-
tus) in Late Pliocene [sic] deposits at Cape Blanco, Oregon was
also reported by Zullo (1969).
4. Systematics
Family Coronulidae Leach, 1817
Genus Cetopirus Ranzani, 1817
Type species: Lepas complanata Mörch, 1853, Late Pleisto-
cene to Recent (possibly extinct; see Bosselaers et al.2017).
Distribution: Early Pleistocene to Recent. Currently known as
an obligate epibiont of Eubalaena spp., living in temperate seas.
Diagnosis (emended after Collareta et al.2016): Body within a
depressed, often dome-shaped shell, consisting of six subequal
compartments; circumference subcircular in apical view; orice
of the body chamber roundedhexagonal, not larger than the
basal opening; opercular valves present, much smaller than the
orice; sheath short, smooth to somewhat grooved, whose
basal edge does not project freely; ala square and thin; com-
pound radius moderately to very thick, whose closely spaced,
copiously branching sutural septa originate from a main septum
2 ALBERTO COLLARETA ET AL. Published online by Cambridge University Press
running along the outer edge of the radius; external radius rather
narrow and transversely striated; paries thin, provided with broad
longitudinal ribs having T-shaped terminations (primary
T-shaped anges) that form a secondary outer lamina; primary
T-shaped anges perforated by longitudinally elongated tubes
or tubules; secondary T-shaped anges present in the form of
minute projectionsthat abut from the primary T-shaped anges;
core of the ribs solidly calcied; ribs externally attened, orna-
mented by weak transverse growth folds and ne longitudinal
striae, lacking transverse interlocking crenulations; apex of the
shell presenting four ribs forming three cavities in-between; sec-
ondary branching very symmetrical and frequent, occurring
near the apex of the shell and resulting in the basal edge of
each compartment presenting a tree-like aspect.
Cetopirus polysyrinx sp. nov.
(Figs 24)
?1961 Coronula complanata Zullo, p. 14
?1969 Coronula (Cetopirus)Newman et al., p. R289
1972 Polysyrinx zulloi[unavailable name, published herein
as a nomen nudum]Davis, p. 36 (partim); gs 3537, 39, 40
Diagnosis: ACetopirus species in which the secondary
T-shaped anges that comprise the primary outer lamina are
internally multitubiferous, being perforated by a high number
of irregularly-sized and irregularly-spaced tubules that provide
them with a spongy aspect in transverse section.
Holotype: LACMIP 1210.158 (=LACMIP Type 14915), iso-
lated right carinolatus.
Etymology: The new species name derivesfrom the Greek and
means many tubes, which appears very appropriate in light of
the multitubiferous condition of its primary T-shaped anges.
The same combination of Greek words was selected by Davis
(1972) as the (unavailable) genus name of his (unpublished)
new taxon. Thus, our choice also pays a due tribute to Davis
remarkable contribution to cirripedology.
Type locality: LACM locality 1210, San Pedro, Los Angeles
County, California, USA (Fig. 1). Davis (1972; 37) further
detailed the site as follows: North end of Los Angeles Harbor
District Yard, Torrance, Calif. Bed begins 2/5 miles due east of
Figure 1. Location of the nds of Cetopirus polysyrinx sp. nov. along the Pacic coast of the USA. Geographical base map after
Figure 2. Cetopirus polysyrinx sp. nov. (holotype), LACMIP 1210.158,
isolated right compartment (carinolatus) collected at San Pedro (Los
Angeles County, California, USA) from Upper Pleistocene deposits.
(a) external view; (b) basal view; (c) alar view; and (d) central view.
Photographs by Lindsay Walker.
intersection of Westmont Drive and Gaffey Street and runs east
for 3/5 mile.
Type horizon: Palos Verdes Sand Formation, Upper
Paratype: UCMP 131833, isolated right carinolatus.
Locality and horizon of the paratype: UCMP locality B7380,
near Cape Blanco, Curry County, USA (Fig. 1). Port Orford For-
mation, Chibanian. Davis (1972) further detailed the site of the
nd as follows: Fossils from bluff sandstone about 75 ft. above
beach in the northwest quarter of Sec. 18 (Cape Blanco quad.,
USGS 1954 ed. map).
Description, comparisons and remarks: Both LACMIP
1210.158 and UCMP 131833 conform to the typical outline of
Cetopirus compartments as detailed in the diagnosis above
(Figs 2,3).
The holotype (note that the same specimen was proposed as
the holotype of Polysyrinx zulloiby Davis 1972) consists of a
partly damaged right carinolatus lacking part of the radius as
well as the lowermost portion of the paries (although the periph-
ery is locally preserved) (Fig. 2). The sheath is distinctly grooved
in its radial portion (corresponding to the alar depression sensu
Collareta et al.2019), less prominently so in its alar portion
(Fig. 2d). The thin, small ala displays a serrated distal margin
(Fig. 2c). Both the sutural edges of the compartment are as
thick as the compartment itself (Figs 2c, d). Judging from its
overall morphology, LACMIP 1210.158 was part of a
dome-shaped shell.
The paratype (note that the same specimen was proposed as
one of the paratypes of Polysyrinx zulloiby Davis 1972) consists
of a right carinolatus that is broken transversely to the ribs some
67 mm below the basis of the sheath. The sheath is smooth
throughout (Fig. 3b)(Fig. 3). The distal margin of the ala is regu-
larly serrated (Fig. 3D). The sutural edges of the wall plate are
almost as thick as the wall plate itself (Figs 3c, d). In prole
view (Figs 3c, d), UCMP 131833 features a slightly sigmoidal
outer wall that hints at a truncated-conical, irregularly depressed
shell shape. This feature may be interpreted as an unusual growth
form due to substrate anomalies or clustering with other nearby
whale barnacle individuals. A cropped CT-stack and a three-
dimensional (3D)-model of UCMP 131833 are available via
MorphoSource at and, respectively.
In both the holotype and the paratype, the primary T-shaped
anges (sensu Collareta et al.2022b) are internally perforated by
a high number of irregularly-sized and irregularly-spaced tubules
that lend them a spongy aspect in transverse section (Fig. 4). In
the Recent species Cetopirus complanatus, each of these anges is
mostly hollow and its opposite [i.e., inner and outer] sides are
seen to be connected by shelly longitudinal plates(Darwin
1854), the resulting tubules being large and arranged in a single
row (Collareta et al.2016;g. 4a). In turn, in the extinct species
Cetopirus fragilis Collareta et al.2016, the core of each ange is
calcied, and a few tubules occur apart from the main axis of the
ange, being aligned parallel to the outer wall and separated
from each other by relatively thick septa (Collareta et al.2016;
g. 4b). The peculiar condition observed in C. polysyrinx sp.
nov. was described by Davis (1972; 40) as follows: [t]he outer-
most laminar complex of the ange possesses one to three longi-
tudinal tubes per septal pair, the majority being two. []In
paratype UCMP XXXXXb [ = 131833], a [] carinolateral
plate, [] the number of tubes varies from one to three, the
majority being two.
Davis (1972) regarded LACMIP 1210.158 and UCMP 131833
as representative of a newcoronuloid genus and species, which he
informally named as Polysyrinx zulloi. However, both these
specimens display characters that are regarded as diagnostic of
Cetopirus, namely: at and broad parietal ribs, which tend to
bifurcate symmetrically and lack transverse interlocking crenula-
tions (Darwin 1854); thin and square alae (Darwin 1854); a basal
edge of the sheath that does not project freely (Darwin 1854); a
Figure 3. Cetopirus polysyrinx sp. nov. (paratype), UCMP 131833, isolated right compartment (carinolatus) collected near Cape Blanco (Los Angeles
County, California, USA) in Middle Pleistocene (Chibanian) deposits. (a) external view; (b) internal view; (c) radial view; and (d) alar view. Photographs
by Bob Day.
Figure 4. Cetopirus polysyrinxsp. nov., close-up of the inner structure of
the primary T-shaped anges, indicated by arrowheads, as observable in
transverse cross-section of the shell (broken surfaces). (a) LACMIP
1210.158 (holotype), photograph by Lindsay Walker; and (b) UCMP
131833 (paratype), photograph by Bob Day. Divisions of the scale
bars equal 0.1 mm in panel (a) and 1mm in panel (b).
4 ALBERTO COLLARETA ET AL. Published online by Cambridge University Press
rather depressed outline (Pastorino & Grifn1996); a narrow
radius (Pastorino & Grifn1996) that is strongly striated trans-
versely (Collareta et al.2016); and primary T-shaped anges
that are internally perforated by longitudinal tubules (Darwin
1854). At the same time, these specimens differ from the other
congeners known to date by displaying secondary T-shaped
anges that are internally multitubiferous, and as such, some-
what spongy; furthermore, they differ from C. fragilis by virtue
of their thicker sutural edges. All things considered, LACMIP
1210.158 and UCMP 131833 are best interpreted as representing
a new, extinct species Cetopirus polysyrinx sp. nov. within the
Recent genus Cetopirus.
Davis (1972) included two additional specimens in the hypo-
digm of Polysyrinx zulloi, namely, an uncatalogued compart-
ment kept in the palaeontological collections of the CAS and
UCMP 131832. Whereas the former specimen originating
from UCMP locality B7375 seems to be lost (Christine Garcia,
pers. comm., December 2021), the latter is an isolated carinathat
seemingly differs from Cetopirus based on at least one character,
and as such, is provisionally identied herein as belonging to cf.
Cetopirus polysyrinx (see below).
cf. Cetopirus polysyrinx
(Fig. 5)
Referred material: UCMP 131832, isolated carina.
Occurrence: UCMP locality B7376, near Cape Blanco, Curry
County, USA. Port Orford Formation, Middle Pleistocene (Chiba-
nian). Davis(1972; 37) described the site of the ndas 100 ft. south
ofUCMPlocalityB7375.UCMPLoc.B7375:100200 ft. south of
large gully in Sec. 12 (Cape Blancoquad., USGS 1954 ed. map).
Remarks: While resembling Cetopirus polysyrinx sp. nov., and
especially the holotype, in overall outline and outer wall micro-
structure (Davis 1972;g. 38), UCMP 131832 also displays a
strongly grooved sheath and remarkably thin sutural edges
(Fig. 5). Whereas the occurrence of strong grooves on the sheath
may be considered to be interspecic variability, the thin sutural
edges of UCMP 131832 are more difcult to assess taxonomic-
ally. Indeed, in Recent Cetopirus shells, the compartmental
sutures are not that thin, not even in carinal plates (see e.g., the
carina of Cetopirus complanatus illustrated by Pastorino & Grif-
n1996;gs 46). As both LACMIP 1210.158 and UCMP
131833 resemble C. complanatus in terms of thickness of the
compartmental sutures, for the moment we do not include
UCMP 131832 within the hypodigm of C. polysyrinx sp. nov.
It is hoped that additional nds of this species will shed light
on this issue. The quest for new specimens of C. polysyrinx sp.
nov. may also include the Middle Pleistocene remains from the
Atsumi Group of Japan that were reported by Kobayashi et al.
(2008; pl. 1, g. 22) and Karasawa & Kobayashi (2022; pl. 2,
gs. 12) as belonging to Coronula sp., as they strongly recall
Cetopirus by displaying a short sheath, thick and narrow radii
that are externally striated, and low ribs that tend to branch
A cropped CT-stack and a 3D-model of UCMP 131832 are
available via MorphoSource at
M432057 and, respectively.
5. Discussion and conclusions
As already observed by Davis (1972), Cetopirus polysyrinx sp.
nov. is seemingly unique among extant and extinct coronulid spe-
cies in that its secondary outer lamina (sensu Collareta et al.
2022b) comprises primary T-shaped anges that are internally
multitubiferous and somewhat spongy (Fig. 4). By contrast,
the early Pleistocene species Cetopirus fragilis displays fewer
tubules within a more calcied secondary outer lamina, whereas
in the Recent form Cetopirus complanatus the primary T-shaped
anges are more hollowed out and plate-like septa alternate with
large tubules that run parallel to the outer wall. Whether the
development of tubules that alternate with longitudinal septa
within the primary T-shaped anges has an adaptive signicance
is not easily determined. It may have provided the basal edge of
the shell with additional grasping structures to supplement the
similarly shaped secondary T-shaped anges in penetrating
into a moulting substrate by cutting out thin strings of the
hosts skin (Collareta et al.2022b); alternatively, it could have
represented a solution for enhancing shell growthrates while saving
calciumcarbonateand preservingstructuralstrength,whichinturn
mayproveusefulif spacecompetitionand/orhydrodynamic energy
are high (Chan & Høeg 2015; Coletti et al.2019). In addition,
wheth ero r not C. fragilis,C. polysyrinx sp. nov.and C. complanatus
comprise a lineage characterised by a progressive hollowing ofthe
secondary outer lamina is also unclear to date.
The Recent species C. complanatus is currently known as an
obligate epibiont of right whales (Mysticeti: Balaenidae: Euba-
laena spp.), as no conrmed records exist to date involving
other hosts (Chemnitz 1785; Darwin 1854; Watson 1981; Scarff
1986; Holthuis et al.1998; Fertl & Newman 2018). A North
Pacic record of C. complanatus was provided by Scarff (1986),
who reported on shells of Coronula diadema (Linnaeus 1767),
Coronula reginae Darwin, 1854 and C. complanatus occurring
on a right whale off Half Moon Bay, California, USA. Although
Scarff (1986) identied this cetacean individual as belonging to
Eubalaena glacialis (Müller 1776), the North Pacic right whales
are currently known to comprise a separate species, Eubalaena
japonica (Lacépède 1818) (Rosenbaum et al.2000); the latter is
currently regarded as endangered (Cooke & Clapham 2018), as
its western and eastern populations could count no more than
300 and 30 individuals, respectively (Brownell et al.2001;
Wade et al.2010). Another record that seemingly hints at C. com-
planatus on modern North Pacic right whales is in Oyamada
Figure 5. cf. Cetopirus polysyrinx, UCMP 131832, isolated carina collected near Cape Blanco (Los Angeles County, California, USA) in Middle Pleis-
tocene (Chibanian) deposits. (a) external view; (b) internal view; and (c) right alar view. Photographs by Bob Day.
Tomokiyos book Isanatori Ekotoba, published in Japanese in
1829 and translated into English (albeit, allegedly, not without
errors) by Yamada (1983) (Omura 1986; Scarff 1986). Therein,
drawings of stalked and acorn barnacles occur beside illustra-
tions of the baleen and blubber of a right whale; the illustrated
cirripedes include a depressed form that the accompanying
description mentions as approximating a sake cup(which is
much reminiscent of the dome-like shell of C. complanatus,
although it may also apply to C. reginae) (Yamada 1983;de
Scarff 1986). (It mayalso be noted that one of the coronulid spe-
cimens gured in the Isanatori Ekotobahas the shell divided
into ve compartments; curiously, the associated text also
describes whale barnacles as pentagonal or hexagonalin
In light of the host specicity of extant whale barnacles, other
(sub)fossil nds of Cetopirus have been regarded as marking the
passage of ancient populations of Eubalaena (Álvarez-
Fernández et al.2014; Collareta 2016; Collareta et al.2016,
2017; Bosselaers et al.2017). Cetopirus polysyrinx sp. nov. also
was, in all likelihood, an obligate commensal of right whales,
either E. japonica or some other species of Eubalaena. Strength-
ening this interpretation, the occurrence of Eubalaena in the
Middle or Upper Pleistocene of the broad North Pacic realm
is documented by a single specimen dredged from the sea oor
off Taiwan (Tsai & Chang 2019); furthermore, stratigraphically
older nds referred to Eubalaena shinshuensis Kimura et al.
2007 and Balaenidae indet. are known from the Upper Miocene
to Lower Pliocene of Japan (Kimura 2009), whereas specimens
of cf. Eubalaena spp. are known from the Pliocene of central
California, USA (Boessenecker 2013b).
The coasts of North America represent a privileged location
for investigating the evolutionary history of the coronuloid bar-
nacles, not least because they are home to the earliest members
of this superfamily (Ross & Newman 1967; Zullo 1982; Collareta
& Newman 2020; Perreault et al.2022). Furthermore, the Plio-
cene and Pleistocene deposits of the Pacic coast of North Amer-
ica have yielded a rather idiosyncratic fossil coronulid fauna.
Indeed, two extinct whale barnacle species have been described
on the basis of and are currently limited to a few fossil
nds from the United States and Mexican West Coast, namely,
Cetolepas hertleini Zullo, 1969 and Cryptolepas murata Zullo,
1961. The former is a greatly enigmatic coronuline species of sub-
cylindrical shell shape that is known from the Upper Pliocene
(Vendrasco et al.2012) San Diego Formation of San Diego
County, California, USA (Zullo 1969). There is no hint, for
this extinct genus and species, of what sort of host it might
have been on. Cryptolepas murata was described from Upper
Pleistocene deposits exposed at San Quintín, Baja California,
Mexico, and is currently regarded as belonging to the same
genus as Cryptolepas rhachianecti Dall, 1872, an obligate epi-
biont of grey whales (Eschrichtius robustus (Liljeborg, 1861))
(Newman & Abbott 1980; Bradford et al.2011; Hayashi 2012;
Bosselaers & Collareta 2016; Taylor et al.2022). (Although Bos-
selaers & Collareta (2016) were hesitant about the genus-level
assignment of C. murata, the high-quality photographs provided
by Taylor et al.(2022) clearly conrm that this extinct species
belongs to Cryptolepas.) Remains of C. murata co-occur with
the holotype of C. polysyrinx sp. nov. in the Palos Verdes Sand
strata exposed at San Pedro, Los Angeles County, California,
USA (pers. obs. by AC, December 2021), thus evoking the
co-occurrence of grey and right whales off the Pacic North
American coast during the Late Pleistocene. Zullo (1961,
1969) envisaged C. murata as ancestral to C. rhachianecti, and
the former as originating through C. hertleini from a Cetopirus-
like ancestor (but see e.g., Davis 1972 for an alternative hypoth-
esis). If this inference is correct, the discovery of Pliocene repre-
sentatives of Cetopirus could be anticipated. Whatever the
likelihood of Zullos phyletic reconstruction, the composition
of the North-West Pacic coronulid fossil fauna including
Cetolepas,Cryptolepas and now Cetopirus seemingly contrasts
with all other coeval assemblages worldwide, which are domi-
nated by Coronula spp. (e.g., Beu 1971; Bianucci et al.2006a,
2006b; Buckeridge et al.2018,2019; Collareta et al.2018a,
6. Acknowledgements
The present work is meant as a homage to the seminal cirripedo-
logical work by Clark William Davis (San Francisco State Uni-
We warmly thank Lindsay Walker (formerly at LACM), Juliet
Hook (LACM), Austin Hendy (LACM), Bob Day (UCMP vol-
unteer), Christine Garcia (CAS) and Peter D. Roopnarine (CAS)
for their invaluable support throughout the present study. Not
only did they search for the specimens that have been studied
herein; they also provided precious curatorial information as
well as high-quality photographs.
John S. Buckeridge (Earth & Oceanic Systems Group, Royal
Melbourne Institute of Technology), Ray T. Perreault (Jarreau
Scientic), Giovanni Bianucci (Università di Pisa), Giovanni
Coletti (Università di Milano Bicocca) and the late William
A. Newman (Scripps Institution of Oceanography) deserve our
most sincere gratitude for many fruitful and illuminating discus-
sions in the elds of coronulid palaeontology and neontology. We
are sincerely indebted to John W.M. Jagt (Natuurhistorisch
Museum Maastricht), John S. Buckeridge and Andrew J. Ross
(National Museums Scotland), whose thorough and construct-
ive reviews greatly contributed to improve an early draft of this
paper. Not least, thorough support by Susie Cox (Earth and
Environmental Science Transactions of the Royal Society of Edin-
burgh Editorial Ofce) is also kindly acknowledged.
7. Financial support
Funding and training for computed tomography scanning was
provided by the Doris O. and Samuel P. Welles Fund of the
UCMP, Department of Integrative Biology, and the FAVE/
Tseng Laboratory. The research of the rst author is supported
by a LinnéSys: Systematics Research Fund grant (funded by
the Linnean Society of London and the Systematics
8. Conicts of interest
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8 ALBERTO COLLARETA ET AL. Published online by Cambridge University Press
... Remarks: Karasawa and Kobayashi (2022) reported the carinolateral/lateral and rostral compartments of an unnamed species of Coronula from the Middle Pleistocene Atsumi Group. After that, Collareta et al. (2022) suggested that these specimens had great affinities with the species of Cetopirus. However, compartments from the Atsumi Group have transverse interlocking crenulations of ribs, not seen in Cetopirus. ...
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Fossil whale barnacles (Balanomorpha: Coronulidae) deposited in the Mizunami Fossil Museum, are classified. Seven species in three genera of whale barnacles from the Pliocene–Pleistocene deposits of Japan are recorded and illustrated. Cetopirus complanatus (Mörch) from the Middle Pleistocene, Coronula reginae Darwin from the Lower Pleistocene (Calabrian), and Cryptolepas rachianecti Dall from the Lower Pleistocene (Gelasian) are first reported in the Japanese fossil records. Coronula Lamarck comprises five species, C. barbara Darwin from the Lower Pleistocene, C. bifida Bronn from the Lower Pliocene–Lower Pleistocene (Gelasian), C. diadema (Linnaeus) from the Lower Pleistocene (Calabrian)–Middle Pleistocene, C. reginae from the Lower Pleistocene (Calabrian), and C. sp. from the Middle Pleistocene. A compartment of an undetermined coronulid genus and species from the Lower Pleistocene (Gelasian) compared with that of Tubicinella major Lamarck and Cetolepas hertleini Zullo is also described.
After the success of the first two editions of the Palaeontological Virtual Congress in 2019 (first PVC) and 2021 (second PVC; Crespo & Manzanares 2019; Crespo & Citton 2021), we have decided to try to replicate the success with a third meeting of the PVC (Fig. 1). The appearance of new applications and technological advances has played a crucial role in paving the way for enhanced avenues of effective scientific communication. This became even more pronounced from more than two years of challenges stemming from the COVID-19 pandemic. Due to this crisis, online platforms gained more relevance and proved key to keeping up the drive for science communication and the dissemination of scientific results (Barral 2020).
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Barnacles (Cirripedia, Thoracica) from the middle Pleistocene Atsumi Group, deposited in the Mizunami Fossil Museum, are classified. Sixteen species in 13 genera of barnacles are recorded and illustrated. Platylepas hexastylos (Fabricius) (Platylepadidae) and three archaeobalanids, Acasta sulcata Lamarck, Euacasta dofleini (Krüger), and Pectinoacasta pectinipes (Pilsbry), are first reported from the Pleistocene of Japan. Adna Sowerby, a coral-inhabiting barnacle genus of Pyrgomatidae, is reevaluated. As a result, Pyrgomina Bałuk and Radwański is synonymized with Adna, and A. anglica (Sowerby), a sole included species of the genus becomes a senior synonym of Pyrgomina djanae Ross and Pitombo, Pyrgoma elargatum Seguenza, Pyrgoma oulastreae Utinomi, Pyrgomina seguenzai Bałuk and Radwański, and Pyrgoma sulcatum Philippi.
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The primary last interglacial, marine isotope substage (MIS) 5e records on the Pacific coast of North America, from Washington (USA) to Baja California Sur (Mexico), are found in the deposits of erosional marine terraces. Warmer coasts along the southern Golfo de California host both erosional marine terraces and constructional coral reef terraces. Because the northern part of the region is tectonically active, MIS 5e terrace elevations vary considerably, from a few meters above sea level to as much as 70 m above sea level. The primary paleo-sea-level indicator is the shoreline angle, the junction of the wave-cut platform with the former sea cliff, which forms very close to mean sea level. Most areas on the Pacific coast of North America have experienced uplift since MIS 5e time, but the rate of uplift varies substantially as a function of tectonic setting. Chronology in most places is based on uranium-series ages of the solitary coral Balanophyllia elegans (erosional terraces) or the colonial corals Porites and Pocillopora (constructional reefs). In areas lacking corals, correlation to MIS 5e often can be accomplished using amino acid ratios of fossil mollusks, compared to similar ratios in mollusks that also host dated corals. Uranium-series (U-series) analyses of corals that have experienced largely closed-system histories range from ∼124 to ∼118 ka, in good agreement with ages from MIS 5e reef terraces elsewhere in the world. There is no geomorphic, stratigraphic, or geochronological evidence for more than one high-sea stand during MIS 5e on the Pacific coast of North America. However, in areas of low uplift rate, the outer parts of MIS 5e terraces apparently were re-occupied by the high-sea stand at ∼100 ka (MIS 5c), evident from mixes of coral ages and mixes of molluscan faunas with differing thermal aspects. This sequence of events took place because glacial isostatic adjustment processes acting on North America resulted in regional high-sea stands at ∼100 and ∼80 ka that were higher than is the case in far-field regions, distant from large continental ice sheets. During MIS 5e time, sea surface temperatures (SSTs) off the Pacific coast of North America were higher than is the case at present, evident from extralimital southern species of mollusks found in dated deposits. Apparently, no wholesale shifts in faunal provinces took place, but in MIS 5e time, some species of bivalves and gastropods lived hundreds of kilometers north of their present northern limits, in good agreement with SST estimates derived from foraminiferal records and alkenone-based reconstructions in deep-sea cores. Because many areas of the Pacific coast of North America have been active tectonically for much or all of the Quaternary, many earlier interglacial periods are recorded as uplifted, higher-elevation terraces. In addition, from southern Oregon to northern Baja California, there are U-series-dated corals from marine terraces that formed at ∼80 ka, during MIS 5a. In contrast to MIS 5e, these terrace deposits host molluscan faunas that contain extralimital northern species, indicating cooler SST at the end of MIS 5. Here I present a review and standardized database of MIS 5e sea-level indicators along the Pacific coast of North America and the corresponding dated samples. The database is available in Muhs et al. (2021b;
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We report the finding of two partial specimens of Cryptolepas rhachianecti (Cirripedia, Coronulidae), a coronulid barnacle known only to inhabit the skin of gray whales ( Eschrichtius robustus ), in Pleistocene-aged sediments from the Canoa Basin, Ecuador. While the historical range of gray whales includes the North Pacific and North Atlantic, to our knowledge this is the first inferred evidence of a gray whale population having resided within the South Pacific. We describe the two Cryptolepas rhachianecti fossils, use isotopic analysis to investigate evidence of migration in their host whales, and discuss their implications for our understanding of gray whale evolutionary history.
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Large‐scale digitization projects such as #ScanAllFishes and oVert are generating high‐resolution microCT scans of vertebrates by the thousands. Data from these projects are shared with the community using aggregate 3D specimen repositories like MorphoSource through various open licenses. We anticipate an explosion of quantitative research in organismal biology with the convergence of available data and the methodologies to analyse them. Though the data are available, the road from a series of images to analysis is fraught with challenges for most biologists. It involves tedious tasks of data format conversions, preserving spatial scale of the data accurately, 3D visualization and segmentations, and acquiring measurements and annotations. When scientists use commercial software with proprietary formats, a roadblock for data exchange, collaboration and reproducibility is erected that hurts the efforts of the scientific community to broaden participation in research. We developed SlicerMorph as an extension of 3D Slicer, a biomedical visualization and analysis ecosystem with extensive visualization and segmentation capabilities built on proven python‐scriptable open‐source libraries such as Visualization Toolkit and Insight Toolkit. In addition to the core functionalities of Slicer, SlicerMorph provides users with modules to conveniently retrieve open‐access 3D models or import users own 3D volumes, to annotate 3D curve and patch‐based landmarks, generate landmark templates, conduct geometric morphometric analyses of 3D organismal form using both landmark‐driven and landmark‐free approaches, and create 3D animations from their results. We highlight how these individual modules can be tied together to establish complete workflow(s) from image sequence to morphospace. Our software development efforts were supplemented with short courses and workshops that cover the fundamentals of 3D imaging and morphometric analyses as it applies to study of organismal form and shape in evolutionary biology. Our goal is to establish a community of organismal biologists centred around Slicer and SlicerMorph to facilitate easy exchange of data and results and collaborations using 3D specimens. Our proposition to our colleagues is that using a common open platform supported by a large user and developer community ensures the longevity and sustainability of the tools beyond the initial development effort.
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The fossil history of turtle and whale barnacles (Coronuloidea: Chelonibiidae, Platylepadidae and Coronulidae) is fragmentary and has only been investigated in part. Morphological inferences and molecular phylogenetic analyses on extant specimens suggest that the roots of whale barnacles (Coronulidae) are to be found among the chelonibiid turtle barnacles, but the hard-part modifications that enabled early coronuloids to attach to the cetacean skin are still largely to be perceived. Here, we reappraise a fossil chelonibiid specimen from the Miocene of insular Tanzania that was previously referred to the living species Chelonibia caretta. This largely forgotten specimen is here described as the holotype of the new species †Chelonibia zanzibarensis. While similar to C. caretta, †C. zanzibarensis exhibits obvious external longitudinal parietal canals occurring in-between external longitudinal parietal septa that abut outwards to form T-shaped flanges, a character so far regarded as proper of the seemingly more derived Coronulidae and Platylepadidae. Along with these features, the presence of a distinct substrate imprint on the shell exterior indicates that †C. zanzibarensis grasped its host's integument in much the same way as coronulids and platylepadids, albeit without the development of macroscopic parietal buttresses and bolsters. Thin section analyses of the inner parietal architecture of some extant and extinct coronuloids conclusively demonstrate that vestiges of comparable external parietal microstructures are present in some living members of Chelonibiidae. This observation strengthens the unity of Coronuloidea while significantly contributing to our understanding of the evolution of the coronuloid shell structure in adapting to a diverse spectrum of hosts.
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(This is an 'informal' abstract. As the present publication is a short note, it has no abstract.) The fossil history of turtle and whale barnacles (Coronuloidea) is fragmentary and has only been investigated in part. In particular, fossils attributed to the currently monotypic family Chelonibiidae mostly consist of remains of the extant genus Chelonibia Leach, 1817 from Plio-Pleistocene strata worldwide. In geologically older deposits, the few published records of Chelonibiidae include some specimens of the Mio-Pliocene, Mediterranean/Paratethyan, archaic-looking genus Protochelonibia Harzhauser & Newman, 2011. A reappraisal of Chelonibia melleni Zullo, 1982 from the lower Oligocene of Mississippi (U.S. Gulf Coast) has revealed two characters that, among chelonibiids, are presently regarded as proper of Protochelonibia: 1-the rostral complex consists of obviously unfused compartments; 2-the parietes of both the rostrolaterals and rostrum exhibit acutely triangular outlines. Therefore, this species represents the oldest member of Chelonibiidae and a bona fide protochelonibiine, hence the new combination Protochelonibia melleni. This action allows us to extend the fossil record of Protochelonibia back for some 10-12 myr, to ~32-34 Ma, as well as to recognise that, in late Palaeogene times, the range of protochelonibiines extended over the broad Western Tethyan realm, on both sides of the North Atlantic. Interestingly, the U.S. Gulf Coast is also home to the only known specimen of Emersonius cybosyrinx Ross, 1967, an enigmatic form from the upper Eocene of Florida, which represents the geologically oldest (albeit surprisingly derived) coronuloid fossil worldwide. Therefore, we hypothesise that the early coronuloids had their centre of distribution somewhere in what is currently considered westernmost Tethys. More discoveries of Palaeogene fossils of Coronuloidea are needed to improve our understanding of the earliest turtle and whale barnacles.
Protochelonibia starnesi sp. nov. (Coronuloidea, Chelonibiidae) is described herein from the upper Rupelian Chickasawhay Formation of Mississippi (U.S.A.). This new "turtle barnacle" species shares similarities with both the extinct Protochelonibia melleni (Zullo, 1982) and the extant Chelonibia "manati" Gruvel, 1903. Similar to the latter, P. starnesi possibly dwelt on sirenians, or the soft skin of other vertebrates. Both P. melleni (whose first representative from the Marianna Formation of Mississippi is also described herein) and P. starnesi attained large shell sizes. Together, they represent the oldest unambiguous members of Coronuloidea, as well as the oldest chelonibiid taxa. As the oldest putative coronuloid is also known from the U.S. Gulf Coast, the Eocene marine deposits exposed over this broad region are a strong candidate to preserve remains of transitional forms.
We report on two clustering chelonibiid shells from Rupelian deposits of southwestern Germany. One of these specimens displays a tripartite rostral complex and disparietal radii that indicate the Oligocene species Protochelonibia melleni, which was known so far from isolated compartments only. A literature review reveals two additional, overlooked records of the rarely reported genus Protochelonibia, coming, respectively, from the Burdigalian of France and the Langhian of Austria. Both these historical finds likely represent the Miocene species Protochelonibia submersa. All together, these occurrences support the notion that the protochelonibiines had acquired a broad distribution as early as in Rupelian times, when P. melleni occurred along the proto-Gulf of Mexico and in the Western Paratethys. Both P. melleni and P. submersa grew in form of peltate shells that evoke a superficial adhesion to some kind of quickly moving hosts. The outer wall of the abraded German colony of P. melleni is comprised of pillar-like blocks of shelly material. In other coronuloids, similar yet more prominent septa abut outward to form T-shaped flanges and intervening longitudinal canals that facilitate the grasping of various kinds of penetrable substrates. The diminutive external longitudinal parietal septa of P. melleni are more likely to represent an early stage in the evolution of the coronuloid shell architecture than vestigial structures. New additions to the pre-Pliocene fossil record of Coronuloidea and a comprehensive phylogenetic analysis of the turtle and whale barnacles will hopefully clarify this and other crucial aspects of the origin and early evolution of these remarkable forms.