''Penion'' is a
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of large
marine snail
A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s, commonly known as siphon whelks, classified within the
mollusc
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Austrosiphonidae.
[MolluscaBase eds. (2021). MolluscaBase. Penion P. Fischer, 1884. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=488452 on 2021-12-24]
Description
Siphon whelks are large, benthic
marine snail
A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s, or
whelks.
[ Powell A. W. B., ''New Zealand Mollusca'', William Collins Publishers Ltd, Auckland, New Zealand 1979 ][Willan, R.C., de C. Cook, S., Spencer, H.G., Creese, R.G., O’Shea, S., Jackson, G.D. Phylum Mollusca. In: de C. Cook, S.C. (eds.), ''New Zealand Coastal Marine Invertebrates 1'', 406 – 407. Canterbury University Press, Christchurch, New Zealand ]
''Penion'' are commonly called siphon whelks because they have a very long
siphon
A siphon (; also spelled syphon) is any of a wide variety of devices that involve the flow of liquids through tubes. In a narrower sense, the word refers particularly to a tube in an inverted "U" shape, which causes a liquid to flow upward, abo ...
. Species typically have a large, pointed
operculum.
[ Ponder, W.F. 1973. A review of the Australian species of ''Penion'' Fischer (Neogastropoda: Buccinidae). Journal of the Malacological Society of Australia 2: 401–428.] Radula
The radula (; : radulae or radulas) is an anatomical structure used by mollusks for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food enters ...
e have 3 or 4 cusps on lateral teeth and 3 cusps on central teeth.
Males have a long, dorso-ventrally flattened penis, and correspondingly females have a large pallial oviduct and albumen gland.
However,
geometric morphometric investigation of ''
P. chathamensis'' indicates that secondary sexual dimorphism is not prominent for shell shape or size.
Shells of ''Penion'' vary significantly in shape, size and colouration, making the distinction of species difficult.
Shells are fusiform with a tall spire of roughly equal height to the aperture and siphonal canal combined.
Protoconch
A protoconch (meaning first or earliest or original shell) is an embryonic or larval shell which occurs in some classes of molluscs, e.g., the initial chamber of an ammonite or the larval shell of a gastropod. In older texts it is also called " ...
morphology is also highly variable, from 1.5 - 4.0 whorls in height.
The siphonal canal of the shell is often long to protect the elongated siphon.
Small shells (or fossils) can be confused with those of ''
Aeneator
An ''aeneator'' ( or ) was a specialized player of wind instruments who was attached to a Roman military unit. The word comes from Latin ''aēneus'' or ''ahēneus'', "brazen", from ''aes'', "copper alloy". While the size of individual Roman milit ...
'' or ''
Antarctoneptunea''.
Shell size can vary quite significantly among populations of ''Penion''. A species, ''Penion fairfieldae'' was formerly recognised, but recent genetic data has demonstrated that the species is indistinguishable from ''
Penion chathamensis''.
Shells originally recognised as ''P. fairfieldae'' can be distinguished from ''P. chathamensis'' using shell size, but not using shell shape.
Ecology

Most species occur in deep water,
and inhabit soft sediments on the continental shelf.
[ Dell, R.K. 1962. New Zealand Marine Provinces - do they exist? Tuatara, 10: 43 - 52]
Online Copy courtesy of New Zealand Electronic Text Collection
/ref>
All species of ''Penion'' have a wide ranging diet, and are detritivores
Detritivores (also known as detrivores, detritophages, detritus feeders or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
and carnivores that actively hunt prey. '' P. sulcatus'' is known to feed on mussels and '' Dosina zelandica zelandica''.[Wilson, B. 1994. ''Australian Marine Shells. Prosobranch Gastropods''. Kallaroo, WA : Odyssey Publishing Vol. 2 370 pp.] Members of the genus have similar niche placement to species of '' Buccinum'' in the Northern Hemisphere.
Distribution
''Penion'' species are restricted to the Southern Hemisphere. Two extant species are currently classified in waters surrounding Australia
Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
and five extant species are documented from New Zealand
New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
.
Numerous fossil species are recorded in New Zealand, Australia, Argentina
Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
and Chile
Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
, and Antarctica
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
.[Stilwell, J.D., Zinsmeister, W.J. 1992. Molluscan systematics and biostratigraphy, lower Tertiary La Meseta Formation, Seymour Island, Antarctic Peninsula. American Geophysical Union Antarctica Research Series 55: 126-128. DOI: 10.1029/AR055](_blank)
In New Zealand, many fossils are found in Wanganui Basin sediments.
During the voyage of HMS ''Beagle'', fossils of '' P. subrectus'' were among palaeontological samples collected by Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
from the mouth of the Santa Cruz River in Argentina.
Evolution
''Penion'' is currently classified within Austrosiphonidae, a which is a family of large buccinoid whelks. ''Penion'' has previously been classified within the families Buccinidae
The Buccinidae are a very large and diverse taxonomic family of large sea snails, often known as whelks or true whelks.Bouchet, P.; Gofas, S. (2010). Buccinidae. In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010) World Marine Mollusca database. Ac ...
and Buccinulidae
The Tudiclidae are Taxonomy (biology), taxonomic subfamily (biology), family in the superfamily Buccinoidea of large sea snails, often known as whelks and the like.MolluscaBase eds. (2023). MolluscaBase. Tudiclidae Cossmann, 1901. Accessed throu ...
.
Molecular phylogenetic trees based on mitochondrial genomic and nuclear ribosomal DNA
The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
sequence data indicate that ''Penion'' is closely related to two genera: '' Kelletia'' found in the north Pacific Ocean and '' Antarctoneptunea'', distributed in waters surrounding New Zealand and Antarctica., as well as '' Serratifusus''. Radulae and opercula morphology is similar between ''Penion'' and ''Kelletia''. It is possible that the earliest known fossils of ''Penion'' belonging to '' P. proavitus'' from the New Zealand Paleocene
The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
( Teurian), actually represent a stem lineage that was the common ancestor of these three genera.
In Australian waters, the sister species
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
'' P. mandarinus'' and '' P. maximus'' have overlapping geographic ranges (sympatry
In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
), and may have evolved from a common ancestor
Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...
via niche differentiation
In ecology, a niche is the match of a species to a specific environmental condition.
Three variants of ecological niche are described by
It describes how an organism or population responds to the distribution of resources and competitors (for e ...
based on prey size and water depth.
In New Zealand, the extinct species '' Penion exoptatus'', '' Penion clifdenensis'', and potentially also '' Penion marwicki'', may belong to the same evolutionary lineage as the extant species '' Penion sulcatus''. This hypothesis is based on geometric morphometric analysis of shell shape and size for all four taxa, as well as the analysis of morphometric variation exhibited all living species of ''Penion''.
Human use
''Penion'' shells are prized by shell collectors.
Shells found in middens of historic Māori
Māori or Maori can refer to:
Relating to the Māori people
* Māori people of New Zealand, or members of that group
* Māori language, the language of the Māori people of New Zealand
* Māori culture
* Cook Islanders, the Māori people of the Co ...
settlements indicate that '' P. sulcatus'' may have been intentionally foraged as a food-source.
Species
Species and subspecies in the genus ''Penion'' include:
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion affixus'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
, 1930)
* † '' Penion antarctocarinatus'' (Stilwell & Zinsmeister, 1992)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion asper'' ( Marwick, 1928)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion australocapax'' Stilwell & Zinsmeister, 1992
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion bartrumi'' (Laws, 1941)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion brazieri'' ( Fleming, 1955)
* '' Penion chathamensis'' ( Powell, 1938)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion clifdenensis'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
, 1930)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
''Penion crassus
''Penion'' is a genus of large marine (ocean), marine snails, commonly known as siphon whelks, classified within the mollusc family (biology), family Austrosiphonidae.MolluscaBase eds. (2021). MolluscaBase. Penion P. Fischer, 1884. Accessed throu ...
'' Frassinetti, 2000
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion crawfordi'' ( Hutton, 1873)
* '' Penion cuvierianus'' ( Powell, 1927)[ Ponder, W.F. 1975. Identity of ''Penion dilatatus'' (Quoy & Gaimard, 1833) (Mollusca: Buccinidae). New Zealand Journal of Marine and Freshwater Research 9 (4): 569-571.]
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion darwinianus'' (Philippi, 1887)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion diversus'' Frassinetti, 2000
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion domeykoanus'' (Philippi, 1887)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion exoptatus'' ( Powell & Bartrum, 1929)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion finlayi'' (Laws, 1930)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion gauli'' ( Marwick, 1928)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion haweraensis'' ( Powell, 1931)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion hiatulus'' ( Powell, 1947)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion huttoni'' (L.R. King, 1934)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion imperfectus'' ( Powell, 1947)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion interjunctus'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
, 1930)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion koruahinensis'' ( Powell & Bartrum, 1928)
* '' Penion lineatus'' Marshall
Marshall may refer to:
Places
Australia
*Marshall, Victoria, a suburb of Geelong, Victoria
** Marshall railway station
Canada
* Marshall, Saskatchewan
* The Marshall, a mountain in British Columbia
Liberia
* Marshall, Liberia
Marshall Is ...
, Hills & Vaux, 2018
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion longirostris'' (Tate, 1888)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion macsporrani'' (Philippi, 1887)
* '' Penion mandarinus'' (Duclos, P.L., 1831)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion marwicki'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
, 1930)
* '' Penion maximus'' ( Tryon, G.W., 1881)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion oncodes'' (Philippi, 1887)
* '' Penion ormesi'' ( Powell, 1927)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion parans'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
, 1930)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion patagonensis'' Reichler, 2010
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion petitianus'' (d'Orbigny, 1842)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion proavitus'' (Finlay
Finlay is a masculine given name, and also a surname. The given name is represented in Scottish Gaelic as Fionnlagh.
Given name Finlay
*Finlay Calder, Scottish rugby player
* Finlay Christie (comedian), British comedian
* Finlay Christie, Scottis ...
& Marwick, 1937)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion roblini'' (Tenison Woods, 1876)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion spatiosus'' (Tate, 1888)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion subrectus'' (Ihering, 1899)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion subreflexus'' (G.B. Sowerby I, 1846)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion subregularis'' (d'Orbigny, 1852)
* '' Penion sulcatus'' (Lamarck
Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1816)
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Penion winthropi'' ( Marwick, 1965)
''Nomina dubia''
Beu 2009 lists the following Antarctic fossil species as nomina dubia
In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application.
Zoology
In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
:
* †
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
''Penion gazdzicki'' (Karczewski, 1987), previously '' Neptunea'' and '' Beringius'', potentially '' Antarctoneptunea''
Species brought into synonymy
* ''Penion adustus'' (Philippi, 1845): synonym of '' Penion sulcatus'' (Lamarck, 1816)
* ''Penion cuvierianus jeakingsi'' ( Powell, 1947): synonym of '' Penion ormesi'' ( Powell, 1927)
* ''Penion dilatatus'' (Quoy & Gaimard, 1833): synonym of '' Penion sulcatus'' (Lamarck, 1816)
* ''Penion fairfieldae'' ( Powell, 1947): synonym of '' Penion chathamensis'' ( Powell, 1938)
Reclassified species
* ''Penion benthicolus'' Dell, 1956: accepted as '' Antarctoneptunea benthicola'' (Dell, 1956)
References
External links
Museum of New Zealand Te Papa Tongarewa, Taxon: ''Penion'' (Genus)
Auckland War Memorial Museum: ''Penion'' specimens
* ttp://www.molluscs.otago.ac.nz/ Checklist of the Recent Mollusca Recorded from the New Zealand Exclusive Economic Zone
Molluscs of Tasmania: Buccinidae - Siphonaliinae: Genus ''Penion'' Fischer, 1884 - “whelks”
Further reading
* Powell A. W. B., ''New Zealand Mollusca'', William Collins Publishers Ltd, Auckland, New Zealand 1979
{{Taxonbar, from=Q585580
Austrosiphonidae
Extant Miocene first appearances