Whale Barnacles Research Articles

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

ABSTRACTA 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of California and the Middle Pleistocene (Chibanian) of Oregon (United States West Coast), described in a 1972 unpublished MA thesis, are formally described and illustrated herein.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.

The occurrence of the coronuloid barnacle Chelonibia Leach, 1817 as an encruster on mammalian bone in the central Mediterranean Sea

Among the turtle and whale barnacles (Coronuloidea: Chelonibiidae, Coronulidae, †Emersoni-idae and Platylepadidae), the members of the chelonibiid species Chelonibia testudinaria (Linnaeus, 1758) are known as epizoic barnacles that can attach to a rather wide spectrum of substrates (pri-marily sea turtles, crabs and sirenians). At present, three living morphs of C. testudinaria have been recognised; of these, the less host-specific is the patula morph, which also displays a remarkably simple, unspecialised shell architecture. Here we report on several chelonibiid shells, referred to the patula morph of C. testudinaria, encrusting a cetacean scapula collected from the floor of the Adriatic Sea facing Salento (Apulia Region, southeastern Italy) and tentatively referred to Tursiops truncatus (Montagu, 1821). This is one of the few records worldwide of a coronuloid barnacle from an inanimate substrate, as well as the second as an encruster on mammalian bone. Such an unusual occurrence is then briefly discussed in the broader framework of the coronuloid commensalism and substrate habits.

A new chelonibiid from the Miocene of Zanzibar (Eastern Africa) sheds light on the evolution of shell architecture in turtle and whale barnacles (Cirripedia: Coronuloidea).

The fossil history of turtle and whale barnacles (Coronuloidea: Chelonibiidae, Platylepadidae, Coronulidae and †Emersoniidae) 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 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.

Paleontological Studies of Whale Barnacles in Taiwan Reveal New Cetacean Migration Routes in the Western Pacific Since the Miocene.

This paper reports a rediscovery of the first museum specimens of fossil whale barnacles from Taiwan. They are part of the material studied and figured by Ichiro Hayasaka in 1934. After examination of the material, which includes two cut-sections and one slice, the taxonomic assignment is revised to Coronula bifida Bronn, 1831. A petrographic study of the surrounding matrix shows that the matrix lacks slate and lithic fragments, indicating that the specimen was deposited in the pre-collision settings during the Miocene to early Pliocene. Figured specimens in Hatai's work in 1939 were examined for comparison. The distribution record of Coronula fossils shows that whales passed through the Taiwan Strait to Okinawa and moved northwards via the Pacific coast of Honshu or entered into the Sea of Japan. The fossil record in this region extends back to the upper Miocene in Yamagata prefecture (facing the Sea of Japan) and Boso Peninsula (facing the Pacific Ocean) in Honshu (~11.2 Ma-5.3 Ma). This is one of the oldest cetacean migration routes documented to date.

The oldest platylepadid turtle barnacle (Cirripedia, Coronuloidea): a new species of Platylepas from the Lower Pleistocene of Italy

Coronuloid barnacles are epibionts of several marine vertebrates (including cetaceans and sea turtles) as well as invertebrates, and are assigned to two families of turtle barnacles (Chelonibiidae Pilsbry, 1916 and Platylepadidae Newman & Ross, 1976) and one family of whale barnacles (Coronulidae Leach, 1817). Chelonibiids and coronulids have a scanty, albeit significant fossil record extending back to the Eocene and Pliocene, respectively; in turn, the fossil record of platylepadids is limited to a single record from the Upper Pleistocene. Here we report on an isolated carinolateral compartment of Platylepas Gray, 1825, the type genus of the family, from Lower Pleistocene (Gelasian) epibathyal deposits exposed at Milazzo (Sicily, Italy). This specimen is here designated holotype of a new species, †Platylepas mediterranea sp. nov. We argue that, like most extant members of Platylepas, †P. mediterranea sp. nov. lived partially embedded in the skin of a sea turtle. This record of an extinct platylepadid – the first from the Mediterranean region and the second worldwide – pushes back the fossil record of Platylepadidae to the lowermost Quaternary, thus possibly supporting an even earlier (e.g., Neogene) timing for the origin of this family and adding a new chapter to the evolutionary history of one of the most diverse and successful lineages of epizoic crustaceans.

Isotopes from fossil coronulid barnacle shells record evidence of migration in multiple Pleistocene whale populations

Migration is an integral feature of modern mysticete whale ecology, and the demands of migration may have played a key role in shaping mysticete evolutionary history. Constraining when migration became established and assessing how it has changed through time may yield valuable insight into the evolution of mysticete whales and the oceans in which they lived. However, there are currently few data which directly assess prehistoric mysticete migrations. Here we show that calcite δ18O profiles of two species of modern whale barnacles (coronulids) accurately reflect the known migration routes of their host whales. We then analyze well-preserved fossil coronulids from three different locations along the eastern Pacific coast, finding that δ18O profiles from these fossils exhibit trends and ranges similar to modern specimens. Our results demonstrate that migration is an ancient behavior within the humpback and gray whale lineages and that multiple Pleistocene populations were undertaking migrations of an extent similar to those of the present day.

Accumulations of Fossils of the Whale Barnacle Coronula bifida Bronn, 1831 (Thoracica: Coronulidae) Provides Evidence of a Late Pliocene Cetacean Migration Route through the Straits of Taiwan.

John Stewart Buckeridge, Benny K.K. Chan, and Shih-Wei Lee (2018) This paper describes a remarkably prolific accumulation of the whale barnacle Coronula bifida Bronn, 1831 in sediments of late Pliocene to earliest Pleistocene age from central Taiwan. Extant Coronula is host-specific to baleen whales; as such, this accumulation of Coronula fossils represents a site where cetaceans congregated during the Plio-Pleistocene - perhaps for breeding. Although whale bones are found at the site, they are rare and fragmentary; the relatively robust shells of Coronula are thus a useful proxy for establishing ancient cetacean migration routes.

New insights on ancient cetacean movement patterns from oxygenisotope analyses of a Mediterranean Pleistocene whale barnacle

New insights on ancient cetacean movement patterns from oxygenisotope analyses of a Mediterranean Pleistocene whale barnacle

Fossil whale barnacles from the lower Pleistocene of Sicily shed light on the coeval Mediterranean cetacean fauna

We report on three shells of whale barnacle (Cirripedia: Coronulidae) collected from Pleistocene shallow-marine deposits exposed at Cinisi (northwestern Sicily, southern Italy). These specimens are identified as belonging to the extinct species Coronula bifida BRONN, 1831. Calcareous nannoplankton analysis of the sediment hosting the coronulid remains places the time of deposition between 1.93 and 1.71 Ma (i.e., at the Gelasian-Calabrian transition), an interval during which another deposit rich in whale barnacles exposed in southeastern Apulia (southern Italy) formed. Since Coronula LAMARCK, 1802, is currently found inhabiting the skin of humpback whales [Cetacea: Balaenopteridae: Megaptera novaeangliae (BOROWSKI, 1781)], and considering that the detachment of extant coronulids from their hosts' skin has been mainly observed in occurrence of cetacean breeding/calving areas, the material here studied supports the existence of a baleen whale migration route between the central Mediterranean Sea (the putative reproductive ground) and the North Atlantic (the putative feeding ground) around 1.8 Ma, when several portions of present-day southern Italy were still submerged. The early Pleistocene utilization of the epeiric seas of southern Italy as breeding/calving areas by migrating mysticetes appears to be linked to the severe climatic degradation that has been recognized at the Gelasian-Calabrian transition and that is marked in the fossil record of the Mediterranean Basin by the appearance of northern guests such as Arctica islandica (LINNAEUS, 1767) (Bivalvia: Veneroida). The subsequent abandonment of the Mediterranean Sea by most species of mysticetes is likely to have resulted from the progressive emergence of shallow-water coastal environments that occurred in Calabrian and Middle Pleistocene times.

Cetopirus complanatus (Cirripedia: Coronulidae) from the late Middle Pleistocene human settlement of Pinnacle Point 13B (Mossel Bay, South Africa).

The late Middle Pleistocene cave site of Pinnacle Point 13B (PP13B, South Africa) has provided the archaeologically oldest evidences yet known of human consumption of marine resources. Among the marine invertebrates recognised at PP13B, an isolated whale barnacle compartment was tentatively determined as Coronula diadema and regarded as indirect evidence of human consumption of a baleen whale (likely Megaptera novaeangliae). In this paper we redetermine this coronulid specimen as Cetopirus complanatus. This record significantly extends the fossil history of C. complanatus back by about 150 ky, thus partially bridging the occurrence of Cetopirus fragilis in the early Pleistocene to the latest Quaternary record of C. complanatus. Since C. complanatus is currently known as a highly specific phoront of right whales (Eubalaena spp.), we propose that the late Middle Pleistocene human groups that inhabited PP13B fed on a stranded southern right whale. Therefore, the whale barnacle from PP13B suggests the persistence of a southern right whale population off South Africa during the predominantly glacial MIS 6, thus evoking the continuity of cetacean migrations and antitropical distribution during that global cold phase. Interestingly, the most ancient evidence of humans feeding on a whale involves Eubalaena, historically the most exploited cetacean genus, and currently still seriously threatened with extinction due to human impact.

The whale barnacle Cryptolepas rhachianecti (Cirripedia: Coronulidae), a phoront of the grey whale Eschrichtius robustus (Cetacea: Eschrichtiidae), from a sandy beach in The Netherlands.

An isolated compartment of a whale barnacle is herein described from Recent beach deposits in Zoutelande (Walcheren, The Netherlands). This specimen is identified as belonging to the extant coronulid species Cryptolepas rhachianecti, currently known as an epizoic symbiont of the grey whale Eschrichtius robustus. This find represents the first occurrence of C. rhachianecti outside the North Pacific, and the first one as a (sub)fossil. In view of the fact that E. robustus, which is currently confined to the North Pacific, is known as a subfossil from the northeastern Atlantic between late Late Pleistocene (c. 45,000 years ago) and historical (c. 1700 AD) times, we propose a similar (late Quaternary) age for the isolated compartment. The find indicates that the extinct late Quaternary northeastern Atlantic population of E. robustus was infected by Cryptolepas rhachianecti. Our find is, therefore, compatible with the hypothesis of an ancient grey whale migration route running between the subtropical/temperate waters of the northeast Atlantic (or Mediterranean Basin), and the cold waters of the Baltic Sea (or southern Arctic Ocean), through the southern North Sea. Finally, we discuss the systematic placement of the fossil barnacle species Cryptolepas murata and propose the possibility of its removal from the genus Cryptolepas pending further investigations.

A new whale barnacle from the early Pleistocene of Italy suggests an ancient right whale breeding ground in the Mediterranean

The fragmentary fossil history of whale barnacles (Cirripedia: Coronulidae) is mainly constituted by remains of Coronula spp. in Plio-Pleistocene deposits found along extant humpback whale migration routes, and especially in occurrence of breeding areas. Here we report the presence of a single compartment of Cetopirus along with remains of Coronula bifida in Lower Pleistocene open shelf deposits of Salento (South Italy). This is the first occurrence of the genus Cetopirus prior to the Late Glacial period (ca. 15–10ky before Present), and the specimen here studied is designated as the holotype of the new fossil species Cetopirus fragilis. Since Cetopirus is currently found exclusively inhabiting the skin of the right whales (Eubalaena spp.), the fossil material here studied suggests the persistence of Balaenidae in the Mediterranean Basin during the Early Pleistocene and supports the existence of a baleen whale migratory route active between the central Mediterranean and the North Atlantic during the Plio-Pleistocene.

Alutiiq Subsistence Economy at Igvak, a Russian-American Artel in the Kodiak Archipelago

Igvak was a Russian-American Company fur-hunting outpost ( artel ) on the south end of Afognak Island that was occupied from the 1790s to about 1830. Midden samples were recovered from deposits adjacent to the Alutiiq workers’ barracks as part of the Dig Afognak program. Although small amounts of European domesticates were identified, the bulk of the diet focused on traditional local foods. The dominant species included cod, harbor seals, and puffins, with a mix of intertidal invertebrates such as mussels, butter clams, and periwinkles. Also common were salmon, cormorants, sea otters, and sea lions. A single species of whale barnacle was identified. The high number of puffin remains may be related to the production of puffin-skin parkas for the Russian-American Company. The site was likely occupied year-round, but the presence of the bones of harbor seal pups and puffin and cormorant chicks confirms a summer occupation.

Jumping from turtles to whales: a Pliocene fossil record depicts an ancient dispersal of Chelonibia on mysticetes

The barnacles included in the superfamily Coronuloidea are epizoic symbionts of various marine vertebrates (including cetaceans, sirenians, and sea turtles) and other crustaceans (crabs and horseshoe crabs). Among Coronuloidea, the so-called turtle barnacles (Chelonibiidae) are known from Paleogene times, whereas the whale barnacles (Coronulidae) likely appeared in the late Pliocene (Piacenzian). Although a derivation from the turtle barnacles (and especially from the genus Chelonibia ) has been proposed, the evolutionary origin of Coronulidae remains to date obscure. In this work we reappraise a fossil record from upper Pliocene (Piacenzian) marine deposits at Casenuove (Empoli municipality, Tuscany, Italy) comprising various shells of Chelonibia testudinaria associated to a partial skeleton of a balaenid mysticete. Based on taphonomic and morpho-functional considerations, we discuss the hypothesis that the barnacles were hosted on the baleen whale, possibly on its callosities, which could have represented an analogous of the horny carapace of marine turtles. This record strongly suggests that the baleen whales can be added to the list of the possible hosts of the barnacles of the genus Chelonibia , thus hinting that the whale barnacles may have evolved from an ancient phase of dispersal of Chelonibia (or a similar ancestor) on mysticete cetaceans.

Past biodiversity: Historical Japanese illustrations document the distribution of whales and their epibiotic barnacles

In recent centuries, excessive fishing has been a major driver in the decline of marine animals. To manage and restore ecosystems, it is necessary to determine the richness of past biodiversity. This report describes the whales and barnacles recorded in classic monographs (called Honzou Gaku) written by Japanese naturalists in the 18th–19th centuries. Past Japanese fauna and flora were documented in these monographs with detailed illustrations; however, many such monographs have been neglected by modern biologists. The whale barnacles illustrated in these monographs were identified as Coronula diadema (Linnaeus, 1767), Coronula reginae Darwin, 1854, and Conchoderma auritum (Linnaeus, 1767). A case study of historical biodiversity is presented here, and shows just one example of the many organisms recorded in the Honzou Gaku. Such information has important implications for the use of historical works to assess past or lost biodiversity all over the world.

Occurrence of whale barnacles in Nerja Cave (Málaga, southern Spain): Indirect evidence of whale consumption by humans in the Upper Magdalenian

A total of 167 plates of two whale barnacle species (Tubicinella majorLamarck, 1802 and Cetopirus complanatusMörch, 1853) have been found in the Upper Magdalenian layers of Nerja Cave, Mina Chamber (Maro, Málaga, southern Spain). This is the first occurrence of these species in a prehistoric site. Both species are specific to the southern right whale Eubalena australis, today endemic in the Southern Hemisphere. Because of Antarctic sea-ice expansion during the Last Glacial Period, these whales could have migrated to the Northern Hemisphere, and reached southern Spain. Whale barnacles indicate that maritime-oriented forager human groups found stranded whales on the coast and, because of the size and weight of the large bones, transported only certain pieces (skin, blubber and meat) to the caves where they were consumed.

The Middle and Later Stone Age faunal remains from Diepkloof Rock Shelter, Western Cape, South Africa

The faunal sample from the Middle Stone Age (MSA) and overlying Later Stone Age (LSA) deposits of Diepkloof Rock Shelter (Western Cape Province, South Africa) includes at least 40 taxa, mostly mammals, but also tortoises, snakes, birds (especially ostrich represented by eggshell), and intertidal mollusks. The LSA sample contains only species that occurred nearby historically, including domestic sheep, which LSA people introduced to the region by 1800 years ago. In contrast, like other Western Cape MSA faunas, the Diepkloof MSA sample has more species and it is especially notable for five large extralimital grazing species. These imply a greater-than-historic role for grasses in the local vegetation, particularly in the post-Howiesons Poort (latest MSA) interval where the grazers appear most abundant. Extreme fragmentation and dark-staining impedes analysis of the MSA bones, but cut-marks, abundant burning, and numerous associated artifacts suggest that people were the main accumulators. Rare coprolites imply that carnivores could have contributed some bones, and concentrations of small mammal bones, particularly near the bottom of the MSA sequence, suggest a role for raptors. Tortoise bones are common throughout the sequence, and the MSA specimens tend to be especially large, as in other MSA assemblages. The LSA specimens are smaller, probably because LSA human populations were denser and preyed on tortoises more intensively. The most surprising aspect of the Diepkloof assemblage is its marine component. The coast is currently 14 km away and it would have been even more distant during much of the MSA when sea levels were often lower. Intertidal mollusks, particularly black mussels and granite limpets, are concentrated in the LSA and in the Late and Post-Howiesons Poort layers. Only LSA shells are complete enough for measurement, and the limpets are small as at other LSA sites. The implication is again for more intense LSA collection by relatively dense human populations. Both the LSA and MSA deposits also contain bones of shorebirds and Cape fur seals. Whale barnacles and occasional dolphin bones indicate that MSA people scavenged beached cetaceans.

Phylogenetic position and evolutionary history of the turtle and whale barnacles (Cirripedia: Balanomorpha: Coronuloidea)

Barnacles of the superfamily Coronuloidea are obligate epibionts of various marine mammals, marine reptiles and large crustaceans. We used five molecular markers: 12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA and Histone 3 to infer phylogenetic relationships among sixteen coronuloids, representing most of the recent genera of barnacles of this superfamily. Our analyses confirm the monophyly of Coronuloidea and that this superfamily and Tetraclitoidea are sister groups. The six-plated Austrobalanus clusters with these two superfamilies. Based on BEAST and ML trees, Austrobalanus is basal and sister to the Coronuloidea, but the NJ tree places Austrobalanus within the Tetraclitoidae, and in the MP tree it is sister to both Coronuloidea and Tetraclitoidae. Hence the position of Austrobalanus remains unresolved. Within the Coronuloidea we identified four clades. Chelonibia occupies a basal position within the Coronuloidea which is in agreement with previous studies. The grouping of the other clades does not conform to previous studies. Divergence time analyses show that some of the time estimates are congruent with the fossil record while some others are older, suggesting the possibility of gaps in the fossil record.

A checklist of turtle and whale barnacles (Cirripedia: Thoracica: Coronuloidea)

A checklist of published records of coronuloid barnacles (Cirripedia: Thoracica: Coronuloidea) attached to marine vertebrates is presented, with 44 species (including 15 fossil species) belonging to 14 genera (including 3 fossil genera) and 3 families recorded. Also included is information on their geographical distribution and the hosts with which they occur.

A rejoinder and addendum to Hayashi (2011) regarding the systematics and biology of the turtle and whale barnacles (Cirripedia: Balanomorpha: Coronuloidea)

Barnacles of the superfamily Coronuloidea are obligate commensals of motile marine animals (Newman & Ross, 1976). Excepting the coronuloid genusChelonibiaLeach, the species included in this superfamily occur strictly upon marine vertebrates, particularly sea turtles and whales (Fricket al., 2011). Chelonophilic (turtle-associated) and cetophilic (whale-associated) coronuloids produce highly-ornamented shells with elaborations that enable these barnacles to grasp the skin of the host (Fricket al., 2010a). These same ornamentations are also useful characters for elucidating taxonomic affinities within the Coronuloidea (Darwin, 1854; Pilsbry, 1916; Ross & Frick, 2007).