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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
1
Dipartimento di Scienze della Terra, Università di Pisa, via S. Maria 53, 56126 Pisa, Italy.
2
Museo di Storia Naturale, Università di Pisa, via Roma 79, 56011 Calci, Italy.
3
Koninklijk Belgisch Instituut voor Natuurwetenschappen, Operationele Directie Aarde en Geschiedenis van het
Leven, Vautierstraat 29, 1000 Brussel, Belgium.
4
Koninklijk Zeeuwsch Genootschap der Wetenschappen, Kousteensedijk 7, 4331 JE, PO Box 378, 4330 AJ
Middelburg, the Netherlands.
5
University of California Museum of Paleontology, 1101 Valley Life Sciences Building, Berkeley, CA 94720-4780,
USA.
*Corresponding author. Email: alberto.collareta@unipi.it
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Middle Pleistocene (Chibanian) of Oregon (United States West Coast), described in a 1972 unpublished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 flanges 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
significance is difficult to determine. Considering that Cetopirus is currently known as an obligate epi-
biont of right whales (including the North Pacific 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 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.
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. zulloi’have 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)to‘Polysyrinx zulloi’.
Below we describe Davis’new coronulid taxon on solid nomen-
clatural grounds and discuss its palaeontological significance.
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,1–8, 2022
https://doi.org/10.1017/S1755691022000214 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 file. The resultant STL file 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 file using their variant of quadric
edge-collapse decimation (Garland & Heckbert 1997). Both
the cropped CT-stack and decimated PLY files were uploaded
to MorphoSource.org.
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;fig. 2h) and Muhs et al.
(1992;fig. 3). Woodring et al.(1946) recognised the deposits
cropping out in this area as being entirely part of their ‘first ter-
race’of 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)
‘first terrace’represented 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 field 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 sandstone’approximately 100
feet along the cliff. Topographically, the site corresponds to the
area where Welton (2015) measured his Section 1 at Gold-
washer’s 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; Leffler 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 Welton’s(2015) sections 3
and 4, where the lower part of the Port Orford Formation is cov-
ered (see also Boessenecker, 2013a;fig. 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 field 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 refinement.
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; orifice
of the body chamber rounded–hexagonal, not larger than the
basal opening; opercular valves present, much smaller than the
orifice; 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.
https://doi.org/10.1017/S1755691022000214 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 flanges) that form a secondary outer lamina; primary
T-shaped flanges perforated by longitudinally elongated tubes
or tubules; secondary T-shaped flanges present in the form of
minute projectionsthat abut from the primary T-shaped flanges;
core of the ribs solidly calcified; ribs externally flattened, orna-
mented by weak transverse growth folds and fine 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 2–4)
?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); figs 35–37, 39, 40
Diagnosis: ACetopirus species in which the secondary
T-shaped flanges 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 flanges.
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 finds of Cetopirus polysyrinx sp. nov. along the Pacific coast of the USA. Geographical base map after wikimedia.org.
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.
3A NEW PLEISTOCENE SPECIES OF CETOPIRUS FROM THE WESTERN USA
https://doi.org/10.1017/S1755691022000214 Published online by Cambridge University Press
intersection of Westmont Drive and Gaffey Street and runs east
for 3/5 mile.’
Type horizon: Palos Verdes Sand Formation, Upper
Pleistocene.
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
find 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 zulloi’by 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 zulloi’by Davis 1972) consists
of a right carinolatus that is broken transversely to the ribs some
6–7 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 profile
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 https://doi.org/10.17602/M2/M432082 and
https://doi.org/10.17602/M2/M432154, respectively.
In both the holotype and the paratype, the primary T-shaped
flanges (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 flanges 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;fig. 4a). In turn, in the extinct species
Cetopirus fragilis Collareta et al.2016, the core of each flange is
calcified, and a few tubules occur apart from the main axis of the
flange, being aligned parallel to the outer wall and separated
from each other by relatively thick septa (Collareta et al.2016;
fig. 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 flange 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: flat 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 flanges, 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.
https://doi.org/10.1017/S1755691022000214 Published online by Cambridge University Press
rather depressed outline (Pastorino & Griffin1996); a narrow
radius (Pastorino & Griffin1996) that is strongly striated trans-
versely (Collareta et al.2016); and primary T-shaped flanges
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
flanges 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 identified 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 findas ‘100 ft. south
ofUCMPlocalityB7375.UCMPLoc.B7375:100–200 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;fig. 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 interspecific variability, the thin sutural
edges of UCMP 131832 are more difficult 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-
fin1996;figs 4–6). 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 finds 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, fig. 22) and Karasawa & Kobayashi (2022; pl. 2,
figs. 1–2) 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
frequently.
A cropped CT-stack and a 3D-model of UCMP 131832 are
available via MorphoSource at https://doi.org/10.17602/M2/
M432057 and https://doi.org/10.17602/M2/M432151, 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 flanges that are internally
multitubiferous and somewhat spongy (Fig. 4). By contrast,
the early Pleistocene species Cetopirus fragilis displays fewer
tubules within a more calcified secondary outer lamina, whereas
in the Recent form Cetopirus complanatus the primary T-shaped
flanges 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 flanges has an adaptive significance
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 flanges in penetrating
into a moulting substrate by cutting out thin strings of the
host’s 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 confirmed records exist to date involving
other hosts (Chemnitz 1785; Darwin 1854; Watson 1981; Scarff
1986; Holthuis et al.1998; Fertl & Newman 2018). A North
Pacific 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) identified this cetacean individual as belonging to
Eubalaena glacialis (Müller 1776), the North Pacific 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 Pacific 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.
5A NEW PLEISTOCENE SPECIES OF CETOPIRUS FROM THE WESTERN USA
https://doi.org/10.1017/S1755691022000214 Published online by Cambridge University Press
Tomokiyo’s 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;fide
Scarff 1986). (It mayalso be noted that one of the coronulid spe-
cimens figured in the ‘Isanatori Ekotoba’has the shell divided
into five compartments; curiously, the associated text also
describes whale barnacles as ‘pentagonal or hexagonal’in
shape.)
In light of the host specificity of extant whale barnacles, other
(sub)fossil finds 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 Pacific realm
is documented by a single specimen dredged from the sea floor
off Taiwan (Tsai & Chang 2019); furthermore, stratigraphically
older finds 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 Pacific 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
finds 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 confirm 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 Pacific 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 Zullo’s phyletic reconstruction, the composition
of the North-West Pacific 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,
2018b).
6. Acknowledgements
The present work is meant as a homage to the seminal cirripedo-
logical work by Clark William Davis (San Francisco State Uni-
versity).
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
Scientific), 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 fields 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 Office) 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 first author is supported
by a LinnéSys: Systematics Research Fund grant (funded by
the Linnean Society of London and the Systematics
Association).
8. Conflicts of interest
None.
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