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Penion; Temporal range: Early Paleocene to Recent, 66.0–0.0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	A siphon whelk Penion ormesi, collected from Golden Bay in New Zealand
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Subclass:	Caenogastropoda
Order:	Neogastropoda
Family:	Austrosiphonidae
Genus:	Penion; P. Fischer, 1884
Species
	See text
Synonyms
	Austrosipho Cossmann, 1906; Austrosipho (Verconella) Iredale, 1914; Berylsma Iredale, 1924; Largisipho Iredale, 1929; Penion (Austrosipho) Cossmann, 1906; Verconella Iredale, 1914;

Penion is a genus of large marine snails, commonly known as siphon whelks, classified within the mollusc family Austrosiphonidae.^[2]^[5]^[6]^[7]^[8]^[9]

Description

Siphon whelks are large, benthic marine snails, or whelks.^[7]^[10]^[11]^[12]^[13]

Penion are commonly called siphon whelks because they have a very long siphon. Species typically have a large, pointed operculum.^[14] Radulae have 3 or 4 cusps on lateral teeth and 3 cusps on central teeth.^[14]

Males have a long, dorso-ventrally flattened penis, and correspondingly females have a large pallial oviduct and albumen gland.^[14] However, geometric morphometric investigation of P. chathamensis indicates that secondary sexual dimorphism is not prominent for shell shape or size.^[12]

Shells of Penion vary significantly in shape, size and colouration, making the distinction of species difficult.^[2]^[11]^[12]^[13]^[14] Shells are fusiform with a tall spire of roughly equal height to the aperture and siphonal canal combined.^[14] Protoconch morphology is also highly variable, from 1.5 - 4.0 whorls in height.^[14] The siphonal canal of the shell is often long to protect the elongated siphon.^[14] Small shells (or fossils) can be confused with those of Aeneator or Antarctoneptunea.^[13]^[15]

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.^[2] Shells originally recognised as P. fairfieldae can be distinguished from P. chathamensis using shell size, but not using shell shape.^[2]

Ecology

Most species occur in deep water,^[12]^[13] and inhabit soft sediments on the continental shelf.^[16]

All species of Penion have a wide ranging diet, and are detritivores and carnivores that actively hunt prey.^[11]^[12] P. sulcatus is known to feed on mussels and Dosina zelandica zelandica.^[11]^[17] 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.^[12] Two extant species are currently classified in waters surrounding Australia^[14] and five extant species are documented from New Zealand.^[2]^[6]^[10]^[13]

Numerous fossil species are recorded in New Zealand,^[15]^[18] Australia,^[14] Argentina and Chile,^[19]^[20] and Antarctica.^[21]^[22]^[23] In New Zealand, many fossils are found in Wanganui Basin sediments.^[15]

During the voyage of HMS Beagle, fossils of P. subrectus were among palaeontological samples collected by Charles Darwin from the mouth of the Santa Cruz River in Argentina.^[24]

Evolution

Penion is currently classified within Austrosiphonidae, a which is a family of large buccinoid whelks.^[9] Penion has previously been classified within the families Buccinidae and Buccinulidae.^[6]^[9]^[25]

Cladogram of Austrosiphonidae and Tudiclidae^[6]^[9]^[13]^[25]

	Antarctoneptunea

	Kelletia

	Penion

	Serratifusus

	Aeneator

	Buccinulum

	Euthria

	Tasmeuthria

Molecular phylogenetic trees based on mitochondrial genomic and nuclear ribosomal DNA 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.,^[6]^[9]^[13]^[25] as well as Serratifusus.^[9] Radulae and opercula morphology is similar between Penion and Kelletia.^[9]^[14] It is possible that the earliest known fossils of Penion belonging to P. proavitus from the New Zealand Paleocene (Teurian), actually represent a stem lineage that was the common ancestor of these three genera.^[6]

In Australian waters, the sister species P. mandarinus and P. maximus have overlapping geographic ranges (sympatry), and may have evolved from a common ancestor via niche differentiation based on prey size and water depth.^[13]

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.^[2] 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.^[2]

Human use

Penion shells are prized by shell collectors.

Shells found in middens of historic Māori settlements indicate that P. sulcatus may have been intentionally foraged as a food-source.^[26]^[27]

Species

Species and subspecies in the genus Penion include:

† Penion affixus (Finlay, 1930)^[15]
† Penion antarctocarinatus (Stilwell & Zinsmeister, 1992)
† Penion asper (Marwick, 1928)^[15]
† Penion australocapax Stilwell & Zinsmeister, 1992^[21]^[22]^[23]
† Penion bartrumi (Laws, 1941)^[15]
† Penion brazieri (Fleming, 1955)^[15]
Penion chathamensis (Powell, 1938)^[25]
† Penion clifdenensis (Finlay, 1930)^[15]
† Penion crassus Frassinetti, 2000^[19]
† Penion crawfordi (Hutton, 1873)^[15]
Penion cuvierianus (Powell, 1927)^[10]^[15]^[28]
† Penion darwinianus (Philippi, 1887)
† Penion diversus Frassinetti, 2000^[19]
† Penion domeykoanus (Philippi, 1887)^[14]
† Penion exoptatus (Powell & Bartrum, 1929)^[15]
† Penion finlayi (Laws, 1930)^[15]
† Penion gauli (Marwick, 1928)^[15]
† Penion haweraensis (Powell, 1931)^[15]
† Penion hiatulus (Powell, 1947)^[15]
† Penion huttoni (L.R. King, 1934)^[15]
† Penion imperfectus (Powell, 1947)^[15]
† Penion interjunctus (Finlay, 1930)^[15]
† Penion koruahinensis (Powell & Bartrum, 1928)^[15]
Penion lineatus Marshall, Hills & Vaux, 2018^[7]^[13]
† Penion longirostris (Tate, 1888)^[14]
† Penion macsporrani (Philippi, 1887)
Penion mandarinus (Duclos, P.L., 1831)^[14]
† Penion marwicki (Finlay, 1930)^[15]
Penion maximus (Tryon, G.W., 1881)^[14]
† Penion oncodes (Philippi, 1887)
Penion ormesi (Powell, 1927)
† Penion parans (Finlay, 1930)^[15]
† Penion patagonensis Reichler, 2010^[20]
† Penion petitianus (d'Orbigny, 1842)
† Penion proavitus (Finlay & Marwick, 1937)^[15]
† Penion roblini (Tenison Woods, 1876)^[14]
† Penion spatiosus (Tate, 1888)^[14]
† Penion subrectus (Ihering, 1899)^[14]^[24]
† Penion subreflexus (G.B. Sowerby I, 1846)^[14]
† Penion subregularis (d'Orbigny, 1852)
Penion sulcatus (Lamarck, 1816)^[10]^[15]^[25]
† Penion winthropi (Marwick, 1965)^[15]

Nomina dubia

Beu 2009 lists the following Antarctic fossil species as nomina dubia:

† Penion gazdzicki (Karczewski, 1987), previously Neptunea and Beringius, potentially Antarctoneptunea^[22]

Species brought into synonymy

Penion adustus (Philippi, 1845): synonym of Penion sulcatus (Lamarck, 1816)^[28]
Penion cuvierianus jeakingsi (Powell, 1947):^[10] synonym of Penion ormesi (Powell, 1927)^[2]
Penion dilatatus (Quoy & Gaimard, 1833): synonym of Penion sulcatus (Lamarck, 1816)^[28]
Penion fairfieldae (Powell, 1947):^[10]^[15] synonym of Penion chathamensis (Powell, 1938)^[2]

Reclassified species

Penion benthicolus Dell, 1956: accepted as Antarctoneptunea benthicola (Dell, 1956)^[6]

References

^ Fischer, P. 1884. Manuel de Conchyliologie et de Paléontologie Conchyliologique ou histoire naturelle des mollusques vivant et fossiles. Paris: F. Savy Vol. 7 pp. 609-688.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Vaux, Felix; Gemmell, Michael R.; Hills, Simon F.K.; Marshall, Bruce A.; Beu, Alan G.; Crampton, James S.; Trewick, Steven A.; Morgan-Richards, Mary (2020). "Lineage Identification Affects Estimates of Evolutionary Mode in Marine Snails". Systematic Biology. 69 (6): 1106–1121. doi:10.1093/sysbio/syaa018. PMID 32163159.
^ Iredale, T. 1924. Results from Roy Bell's molluscan collections. Proceedings of the Linnean Society of New South Wales 49(3): 179-279, pls 33-36.
^ Iredale, T. 1914. On some invalid molluscan generic names. Proceedings of the Malacological Society of London 11: 170-178.
^ Vaux, Felix (2017). Evolutionary lineages and the diversity of New Zealand true whelks (Doctor of Philosophy thesis). Massey University. hdl:10179/13113.
^ ^a ^b ^c ^d ^e ^f ^g Vaux, Felix; Hills, Simon F.K.; Marshall, Bruce A.; Trewick, Steven A.; Morgan-Richards, Mary (2017). "A phylogeny of Southern Hemisphere whelks (Gastropoda: Buccinulidae) and concordance with the fossil record". Molecular Phylogenetics and Evolution. 114 (2017): 367–381. doi:10.1016/j.ympev.2017.06.018. PMID 28669812.
^ ^a ^b ^c Marshall, Bruce A.; Hills, Simon F.K.; Vaux, Felix (2018). "A new species of Penion P. Fischer, 1884 from northern New Zealand (Mollusca: Neogastropoda: Buccinoidea)". Molluscan Research. 38 (4): 238–242. doi:10.1080/13235818.2017.1420398. S2CID 90709310.
^ 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
^ ^a ^b ^c ^d ^e ^f ^g Kantor, Yuri I.; Fedosov, Alexander E.; Kosyan, Alisa R.; Puillandre, Nicolas; Sorokin, Pavel A.; Kano, Yasunori; Clark, Roger; Bouchet, Philippe (2022). "Molecular phylogeny and revised classification of the Buccinoidea (Neogastropoda)". Zoological Journal of the Linnean Society. 194 (3): 789–857. doi:10.1093/zoolinnean/zlab031.
^ ^a ^b ^c ^d ^e ^f Powell A. W. B., New Zealand Mollusca, William Collins Publishers Ltd, Auckland, New Zealand 1979 ISBN 0-00-216906-1
^ ^a ^b ^c ^d 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 ISBN 978-1877257-60-5
^ ^a ^b ^c ^d ^e ^f Vaux, Felix; Crampton, James S.; Marshall, Bruce A.; Trewick, Steven A.; Morgan-Richards, Mary (2017). "Geometric morphometric analysis reveals that the shells of male and female siphon whelks Penion chathamensis are the same size and shape". Molluscan Research. 37 (3): 194–201. doi:10.1080/13235818.2017.1279474. S2CID 90288210.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Vaux, Felix; Crampton, James S.C.; Trewick, Steven A.; Marshall, Bruce A.; Beu, Alan G.; Hills, Simon F.K.; Morgan-Richards, Mary (2018). "Evolutionary lineages of marine snails identified using molecular phylogenetics and geometric morphometric analysis of shells". Molecular Phylogenetics and Evolution. 127 (October 2018): 626–637. doi:10.1016/j.ympev.2018.06.009. PMID 29913310. S2CID 49303166.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r 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.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y Beu, Alan G.; Maxwell, P.A. (1990). "Cenozoic Mollusca of New Zealand". NZGS Paleontological Bulletin. 58 (5): 799.
^ Dell, R.K. 1962. New Zealand Marine Provinces - do they exist? Tuatara, 10: 43 - 52. Online Copy courtesy of New Zealand Electronic Text Collection
^ Wilson, B. 1994. Australian Marine Shells. Prosobranch Gastropods. Kallaroo, WA : Odyssey Publishing Vol. 2 370 pp.
^ Hayward, Bruce W.; Stolberger, Thomas F.; Collins, Nathan; Beu, Alan G.; Blom, Wilma (2023). "A diverse Late Pliocene fossil fauna and its paleoenvironment at Māngere, Auckland, New Zealand". New Zealand Journal of Geology and Geophysics. doi:10.1080/00288306.2023.2243234.
^ ^a ^b ^c Frassinetti, Daniel (2000). "Upper Pliocene marine molluscks from Guafo Island, southern Chile. Part II. Gastropoda". Boletin del Museo Nacional de Historia Natural, Chile. 49: 131–161.
^ ^a ^b Reichler, Valeria A. (2017). "Estratigrafía y paleontología del Cenozoico marino del Gran Bajo y Salinas del Gualicho, Argentina y descripción de 17 especies nuevas". Andean Geology. 37 (1): 177–219. doi:10.4067/S0718-71062010000100008.
^ ^a ^b 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 ISBN 9781118667705
^ ^a ^b ^c Beu, Alan G. (2009). "Before the ice: Biogeography of Antarctic Paleogene molluscan faunas". Palaeogeography, Palaeoclimatology, Palaeoecology. 284 (3–4): 191–226. Bibcode:2009PPP...284..191B. doi:10.1016/j.palaeo.2009.09.025.
^ ^a ^b Crame, J. Alistair; Beu, Alan G.; Ineson, Jon R.; Francis, Jane E.; Whittle, Rowan J.; Bowman, Vanessa C. (2014). "The Early Origin of the Antarctic Marine Fauna and Its Evolutionary Implications". PLOS ONE. 9 (12): e114743. Bibcode:2014PLoSO...9k4743C. doi:10.1371/journal.pone.0114743. PMC 4262473. PMID 25493546.
^ ^a ^b Parras, Ana; Miguel, Griffin (2009). "Darwin's great Patagonian Tertiary formation at the mouth of the Río Santa Cruz: a reappraisal". Revista de la Asociación Geológica Argentina. 64 (1): 70–82.
^ ^a ^b ^c ^d ^e Hayashi, Seiji (2005). "The molecular phylogeny of the Buccinidae (Caenogastropoda: Neogastropoda) as inferred from the complete mitochondrial 16s rRNA gene sequences of selected representatives". Molluscan Research. 25: 85–98.
^ Green, R.C.; Pullar, W.A. (1960). "Excavations at Orongo Bay, Gisborne". The Journal of the Polynesian Society. 69 (4): 332–353.
^ Allen, Melinda S. (2012). "Molluscan foraging efficiency and patterns of mobility amongst foraging agriculturalists: a case study from northern New Zealand". Journal of Archaeological Science. 39 (2012): 295–307. doi:10.1016/j.jas.2011.09.013.
^ ^a ^b ^c 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.