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Cellana
Cellana
Cellana
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Cellana
A shell of Cellana ornata
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Patellogastropoda
Superfamily: Lottioidea
Family: Nacellidae
Genus: Cellana
Adams, 1869
Type species
Nacella cernica
Adams, 1869
Synonyms
  • Bertinia Jousseaume, 1883
  • Granopatella Pallary, 1920
  • Helcion (Helcioniscus) Dall, 1871
  • Helcioniscus Dall, 1871
  • Nacella (Cellana) H. Adams, 1869 (original rank)

Cellana is a genus of sea snails or limpets, marine gastropod molluscs in the family Nacellidae, the true limpets.[1]

Distribution

[edit]

This genus occurs in the temperate and tropical Indo-Pacific oceans, Hawaii (where they are known as ‘opihi and considered a delicacy) and around Australia and New Zealand. Species are also found around the coasts of Japan, the Red Sea, Mauritius, Madagascar, South Africa and the sub-Antartarctic Islands. One species, Cellana radiata, is cosmopolitan.

These sea snails feed by grazing on green macroalgae growing on rocky substrate in the intertidal zone. Some of these limpets can live up to 7 years, however most do not get older than 2–3 years. They reproduce by broadcasting their spawn in large, yolky eggs in great numbers (between 20,000 for C. flava and C. denticulata to 230,000 for C. ornata).

Species

[edit]
Cellana conciliata
Cellana mazatlandica
Cellana rota

Species within the genus Cellana include:[1]

The sources given below, also mention the following species :

Species brought into synonymy

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Cellana

View on Grokipedia
from Grokipedia
Cellana is a of true limpets, marine gastropod mollusks in the family within the order Patellogastropoda, characterized by their conical shells, strong muscular foot for adhesion to rocks, and adapted for scraping . The genus was established by in 1869, with the Cellana livescens (formerly Nacella cernica), and currently encompasses over 35 accepted species, primarily identified through shell morphology, gill structure, and molecular data. These limpets are key components of intertidal ecosystems in tropical and warm-temperate waters, where they inhabit the mid- to low-intertidal zones, on , encrusting macroalgae, and spores while exhibiting limited larval dispersal (typically less than 20 days) but achieving broad distribution through occasional on floating debris. The is predominantly in range, spanning from to Hawaii in the east and from to in the north, with some species extending to and subantarctic islands; notable endemics include Hawaiian species like Cellana exarata (blackfoot ʻopihi), which are culturally significant as a delicacy and feature in traditional harvesting practices. Species exhibit ecological partitioning, such as zonation along intertidal gradients, and show phylogenetic divergence dating back approximately 24 million years to the Oligocene-Miocene boundary, reflecting historical biogeographic events in the .

Taxonomy and classification

Etymology and history

The genus Cellana was first described by British malacologist in 1869, in his paper "Descriptions of a new genus and fourteen new species of marine shells" published in the Proceedings of the . Adams introduced Cellana as a of Nacella, with the Nacella (Cellana) cernica H. Adams, 1869, designated by monotypy; this is now regarded as a junior of Cellana livescens (Reeve, 1855). Initially classified within the family , the broad group of true limpets, Cellana was recognized for its ribbed shells and distribution, distinguishing it from more temperate species. The family Nacellidae, encompassing Cellana and the genus Nacella, was formally established by Johannes Thiele in to accommodate these morphological differences, though early adoption was limited. Significant contributions to the of Cellana came from American conchologist Henry Augustus Pilsbry in the 1890s, who described numerous species and subspecies—such as Patella eucosmia (now Cellana eucosmia) and Patella orientalis (now Cellana orientalis)—in volumes 13 and 14 of his Manual of Conchology (1891–1893), refining classifications based on shell sculpture and radular features. The reclassification of Cellana into was solidified in the late through combined morphological and molecular analyses. Studies in the , including those establishing the subclass Patellogastropoda, highlighted distinctions in soft anatomy and separating from Patellidae. By the 2000s, phylogenetic research confirmed the of within the superfamily Lottioidea; for instance, a 2005 molecular study using mitochondrial 12S and 16S rRNA genes supported the Indo-Pacific Cellana as sister to Nacella, justifying the family's separation from Patelloidea. Modern taxonomic authority rests with the (WoRMS), which accepts Cellana in Nacellidae, with the genus comprising 39 valid species as of November 2025; the most recent major revisions to its classification occurred around 2010, incorporating data to resolve synonyms and distributions.

Phylogenetic relationships

Cellana is classified in the kingdom Animalia, phylum , class , subclass Patellogastropoda, order Nacellida, superfamily Nacelloidea, family Nacellidae. This placement reflects its position among true limpets, characterized by a cap-shaped shell and a distinct foot morphology adapted for intertidal . At the genus level, has several historical synonyms, including Bertinia Jousseaume, 1883, Granopatella Iredale, 1933, and Helcioniscus Dall, 1871, which were later subsumed based on morphological and molecular reassessments. There is also partial historical synonymy with Nacella, though current recognizes them as distinct sister genera within Nacellidae. Molecular evidence from 18S rRNA gene sequences and mitochondrial markers such as COI, 12S rRNA, and 16S rRNA has firmly established the phylogenetic position of Cellana. Studies in the 2000s, including analyses by Yoon et al. (2007) using 18S rDNA and by Nakano and Ozawa (2007) employing multiple mitochondrial genes, support the monophyly of Patellogastropoda and place Nacellidae as sister to Patellidae. These datasets reveal Cellana as closely related to Nacella in the Southern Hemisphere and to Patella species in the Mediterranean and Atlantic, with mitogenomic comparisons further confirming Nacellidae's basal position within Patelloidea. Divergence time estimates derived from molecular clock analyses of these markers indicate that the split between Cellana and Nacella occurred approximately 50 million years ago during the , with the crown age of Cellana dating back approximately 24 million years to the Oligocene-Miocene boundary, though estimates vary with fossil constraints. Earlier mitogenomic studies suggest the broader diversification of patellogastropod lineages began in the , with Cellana-Nacella splits occurring approximately 50 million years ago.

Description

Shell characteristics

The shells of Cellana species are characteristically conical and patelliform, featuring a low to medium apex positioned slightly anteriorly, typically at about one-third the distance from the anterior margin. The height-to-length ranges from 0.29 to 0.45, contributing to a low-profile that facilitates secure attachment to rocky substrates in wave-exposed environments. Basal diameters vary widely across the genus, generally spanning 2–10 cm, with species-specific maxima such as up to 9 cm for C. testudinaria. For instance, C. radiata commonly reaches 3–4 cm, though larger individuals up to 4.5 cm have been recorded. The external surface is ornamented with radiating ribs or costae, numbering 10–50 depending on the species, often accompanied by fine sculptured growth lines that enhance grip and structural integrity. Examples include 20–25 low radial ribs in C. radians and 30–40 strong ribs in C. tramoserica. The interior is smooth and iridescent, typically bluish-silver or white, with a prominent horseshoe-shaped muscle scar that reflects the attachment site of the foot. Exterior coloration shows considerable variation, ranging from olive-green to brown, frequently patterned with darker radial stripes or bands that align with the ribs. Species like C. tramoserica may exhibit orange-brown hues with reddish ribs, while C. radiata displays grayish-white to dark brown tones with contrasting darker costae.

Radula and soft anatomy

The of Cellana species is of the docoglossan type, characterized by a reduced central rachidian and 4–5 large lateral teeth per transverse row, enabling efficient rasping and scraping of algal films from rock surfaces. The rachidian is small and spear-shaped, while the lateral teeth feature robust cusps adapted for gripping and abrading substrata, with the often extending 4–5 times the length of the animal. This structure contrasts with more derived types in other gastropods by emphasizing fewer, stronger teeth for targeted rather than broad filtration. The soft body of Cellana limpets includes a large, muscular foot that facilitates strong to rocky substrates through a combination of generated by pedal musculature and properties of pedal , which forms a glue-like seal during periods of immobility. The mantle edge bears sensory tentacles and pallial lobes that extend around the shell margin, aiding in and contributing to the overall sealing mechanism against the substratum. The gills are bipectinate, arranged in a single row within the mantle cavity, which supports efficient during intermittent submersion in the . The visceral mass is compact and positioned dorsally within the shell, housing the digestive gland for processing algal material and the gonads for reproductive functions, with these organs showing seasonal variations in size tied to . The nervous system features a well-developed , a chemosensory organ in the mantle cavity that detects water-borne stimuli such as odors or pollutants, relaying signals via the osphradial to the central ganglia for behavioral responses. These anatomical features enable Cellana species to seal their shells tightly against the substrate using the foot and , thereby tolerating for up to several hours during low tide exposure by minimizing water loss through the mantle cavity. This , in conjunction with the protective conical shell, allows survival in harsh intertidal conditions.

Habitat and distribution

Geographic range

The genus Cellana is primarily distributed across temperate and tropical waters of the region, encompassing a vast area from the in the central Pacific to in the northwest, extending eastward to oceanic islands such as Juan Fernández off and westward to the . This range further includes southern extensions to , , , , and along the African coast, as well as sub-Antarctic islands like Campbell Island. Species such as C. exarata, C. talcosa, and C. sandwicensis are recorded in , while C. tramoserica and C. solida occur in , and C. flava, C. radians, C. stellifera, and members of the C. strigilis complex inhabit and its associated islands. In the western Indo-Pacific, C. eucosmia is found in the , and C. capensis along n shores. A notable exception to the predominantly regional patterns is C. radiata, which exhibits a more as a tropical outlier, spanning wider zones including the extensions from and the to the and . Subspecies such as C. radiata capensis extend this presence southward to , highlighting greater dispersal tolerance compared to other congeners. Endemism is particularly pronounced in isolated locales within this range, with hosting three to four endemic species (C. exarata, C. talcosa, C. sandwicensis, and occasionally C. melanostoma in the northwest islands), reflecting limited across oceanic barriers. Similarly, shows high regional endemism, including multiple species restricted to (e.g., C. solida in ) and (e.g., C. denticulata on the northeast coast). The genus's distribution has been shaped by historical expansions, primarily through post-Pleistocene long-distance dispersal facilitated by planktonic larval stages and potential rafting events, enabling colonization of remote islands despite generally limited dispersal capacity. No native populations occur in the Atlantic Ocean, underscoring the as the core biogeographic domain.

Environmental preferences

Species of the genus Cellana primarily occupy the mid- to upper intertidal zones on exposed rocky shores, typically ranging from 0.5 to 1.5 meters above mean low neaps, where they experience periodic submersion and emersion with the . This zonation allows them to exploit wave-surged environments that provide oxygenation while minimizing prolonged submersion risks. They are most abundant around mean on stable substrates, with smaller individuals often dominating higher elevations and larger ones lower down. Cellana limpets adhere to firm rock surfaces, boulders, or platforms, avoiding soft sediments or shifting substrates that could dislodge them. They prefer areas with moderate to high wave exposure, which enhances water flow and reduces stagnation, though they can tolerate semi-protected conditions. These microhabitats, often bare or lightly algaled rock, support their attachment and reduce competition from dense macroalgae. These limpets withstand typical marine salinities of 25–35 ppt, with broader tolerance to fluctuations up to 40 ppt in tide pools during emersion. Habitat temperatures range from 10–30°C across their distribution, reflecting adaptations to both subtropical and temperate conditions. resistance is achieved by clamping the shell tightly to the substrate, sealing a moist chamber beneath during . In these niches, Cellana species co-occur with and mussels, particularly in the upper intertidal, but they dominate open rock surfaces where space is not densely colonized by sessile competitors.

Ecology and life history

Feeding behavior

Cellana limpets employ a radula-based grazing mechanism to scrape , biofilms, and associated epibenthic organisms from intertidal rock surfaces. The radula's robust teeth, noted for their exceptional tensile strength among biological materials, enable effective rasping of tough substrates like crustose , facilitating the removal of thin algal layers without deeper excavation. This feeding strategy is primarily herbivorous, with no evidence of carnivory, and activity peaks during nocturnal or crepuscular periods to minimize risk and predation exposure during emersion at . The diet of Cellana species centers on , particularly diatoms such as Navicula spp., Amphora spp., Nitzschia spp., and Gryosigma spp., supplemented by green macroalgae including Ulva spp. (formerly Enteromorpha). Gut content analyses from field and laboratory studies reveal these components as dominant, with occasional ingestion of like sand grains, reflecting opportunistic scavenging during grazing bouts. Cyanobacteria such as Lyngbya spp. may also contribute minor nutritional value, but the overall reliance on algal biofilms underscores their role as key regulators of microalgal standing stocks in intertidal zones. Foraging patterns in Cellana are characterized by limited mobility within small ranges around preferred resting scars, where individuals return after tidal-driven excursions. These movements are density-dependent, with higher densities promoting that restricts extent and enhances homing to conserve energy. Grazing excursions typically occur when awash or immersed, covering distances up to 0.5–1 m from refuges in species like C. grata, but overall activity remains confined to optimize energy intake against environmental risks. Through persistent radular scraping, Cellana individuals achieve high grazing efficiency, thereby exerting top-down control on algal community structure and preventing dominance by foliose forms. This consumption rate, inferred from radular mark density and microalgal depletion studies, highlights their ecological impact in maintaining bare rock patches and influencing intertidal .

Reproduction and larval development

Species in the genus Cellana are dioecious broadcast spawners that undergo in . Sex ratios in populations are typically near 1:1, with males and females lacking obvious external dimorphism and requiring examination for determination. Spawning events are often synchronized, influenced by environmental cues such as lunar cycles and seasonal temperature increases, which promote release during periods favorable for larval survival. Females produce large numbers of yolky eggs, with output varying by and size; for example, C. flava females release approximately 20,000 eggs per spawning event, while C. ornata females can produce 200,000–360,000 eggs from larger individuals (40–49 mm shell length). Overall across the ranges from 20,000 to 230,000 yolky eggs per female, supporting high reproductive output despite partial spawning in some cases. Fertilized eggs develop into planktonic trochophore larvae within 1–2 days, which are non-feeding and lecithotrophic, relying on yolk reserves. These larvae transition to veliger stages and remain pelagic for 2–10 days (typically 5–10 days), during which dispersal occurs before settlement on intertidal rocky substrates. Settlement is followed by into juvenile limpets, which crawl to suitable microhabitats and begin benthic life. Individuals reach at 1–2 years of age or when shell length attains 1–2 cm, depending on growth rates influenced by environmental conditions. Lifespans vary across species but generally range from 2–7 years.

Species diversity

List of accepted species

The genus Cellana currently includes 39 accepted , all extant with no known extinct taxa. These species exhibit a primarily distribution, with regional concentrations such as 3 Hawaiian endemics (C. exarata, C. melanostoma, C. talcosa), 4 Australasian species (C. solida, C. strigilis, C. flava, C. howensis), approximately 5–6 endemics (e.g., C. enneagona, C. grata), and others like the South African C. capensis and the widespread C. radiata. identification is based on shell morphology, including rib count and form, apex position, and coloration.
Scientific NameAuthor(s)Year
Cellana analogiaIredale1940
Cellana ardosiaeaHombron & Jacquinot1841
Cellana capensisGmelin1791
Cellana conciliataIredale1940
Cellana craticulataSuter1905
Cellana cylindricaGmelin1791
Cellana denticulataMartyn1784
Cellana diraReeve1855
Cellana enneagonaReeve1854
Cellana eucosmiaPilsbry1892
Cellana exarataReeve1854
Cellana flavaF. W. Hutton1873
Cellana granostriataReeve1855
Cellana grataA. A. Gould1859
Cellana howensisIredale1940
Cellana karachiensisWinckworth1930
Cellana livescensReeve1855
Cellana mazatlandicaG. B. Sowerby I1839
Cellana melanostomaPilsbry1891
Cellana nigrolineataReeve1854
Cellana oliveriA. W. B. Powell1955
Cellana orientalisPilsbry1891
Cellana ornataDillwyn1817
Cellana priceiA. W. B. Powell1973
Cellana radiansGmelin1791
Cellana radiataBorn1778
Cellana rotaGmelin1791
Cellana sandwicensisPease1861
Cellana solidaBlainville1825
Cellana sonticaIredale1940
Cellana stelliferaGmelin1791
Cellana strigilisHombron & Jacquinot1841
Cellana taitensisRöding1798
Cellana talcosaA. A. Gould1846
Cellana testudinariaLinnaeus1758
Cellana toreumaReeve1855
Cellana tramosericaHolten1802
Cellana turbatorIredale1940
Cellana vitiensisA. W. B. Powell1973

Synonyms and taxonomic notes

The taxonomy of Cellana has undergone significant revisions due to historical over-description of species and subspecies based primarily on shell morphology during the , when numerous taxa were erected from variable specimens across the . For instance, the New Zealand C. strigilis complex was traditionally recognized as comprising six , reflecting perceived morphological differences in shell shape, sculpture, and coloration. However, 20th-century morphological studies began consolidating these, and molecular analyses in the further clarified relationships, reducing the complex to two distinct : C. strigilis (including synonyms such as C. strigilis flemingi and redimiculum) and C. oliveri (including synonyms like C. strigilis bollonsi). Similarly, and phylogenetic studies have resolved several synonymies in other . Shell color polymorphisms and have historically contributed to taxonomic confusion, leading to the description of variants or that are now regarded as intraspecific variation within Cellana species. For example, in C. nigrolineata, two distinct color morphs were once considered potentially separate taxa, but genetic and morphometric indicate they represent ecophenotypic responses to environmental factors rather than distinct lineages. Recent integrative approaches, including sequencing, have also revealed cryptic diversity in species like C. grata and C. toreuma, with allopatric clades in the prompting ongoing debates about species boundaries, though many previously split forms have been synonymized. At the genus level, Helcioniscus Dall, 1871, is recognized as a junior subjective synonym of Cellana H. Adams, 1869, based on priority and morphological overlap. No Cellana species are currently listed under IUCN criteria, but populations in Hawaii, particularly C. exarata and C. sandwicensis (known locally as ʻopihi), face significant pressure from overharvesting, leading to localized declines and regulatory monitoring to ensure sustainable use.

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