Penion

Penion is a genus of large marine snails, commonly known as siphon whelks, classified within the mollusc family Buccinidae, the true whelks.[1][5][6][7]

Penion
Temporal range: Early Paleocene to Recent, 66.0–0.0 Ma
A siphon whelk Penion ormesi,[1] collected from Golden Bay in New Zealand
Scientific classification
Kingdom:
Phylum:
Class:
(unranked):
Superfamily:
Family:
Genus:
Penion

Species

See text

Synonyms
  • Austrosipho Cossmann, 1906
  • Berylsma Iredale, 1924[3]
  • Largisipho Iredale, 1929
  • Verconella Iredale, 1914[4]

Description

An apetural view of a shell of Penion maximus

Siphon whelks are large, benthic marine snails, or whelks.[7][8][9][10][11]

Penion are commonly called siphon whelks because they have a very long siphon. Species typically have a large, pointed operculum.[12] Radulae have 3 or 4 cusps on lateral teeth and 3 cusps on central teeth.[12]

Males have a long, dorso-ventrally flattened penis, and correspondingly females have a large pallial oviduct and albumen gland.[12] However, geometric morphometric investigation of P. chathamensis indicates that secondary sexual dimorphism is not prominent for shell shape or size.[10]

Shells of Penion vary significantly in shape, size and colouration, making the distinction of species difficult.[1][9][10][11][12] Shells are fusiform with a tall spire of roughly equal height to the aperture and siphonal canal combined.[12] Protoconch morphology is also highly variable, from 1.5 - 4.0 whorls in height.[12] The siphonal canal of the shell is often long to protect the elongated siphon.[12] Small shells (or fossils) can be confused with those of Aeneator or Antarctoneptunea.[11][13]

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.[1] Shells originally recognised as P. fairfieldae can be distinguished from P. chathamensis using shell size, but not using shell shape.[1]

Ecology

Egg capsules of Penion sulcatus at Kawau Bay, Hauraki Gulf

Most species occur in deep water,[10][11] and inhabit soft sediments on the continental shelf.[14]

All species of Penion have a wide ranging diet, and are detritivores and carnivores that actively hunt prey.[9][10] P. sulcatus is known to feed on mussels and Dosina zelandica zelandica.[9][15] 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.[10] Two extant species are currently classified in waters surrounding Australia[12] and five extant species are documented from New Zealand.[1][6][8][11]

Numerous fossil species are recorded in New Zealand,[13] Australia,[12] Argentina and Chile,[16][17] and Antarctica.[18][19][20] In New Zealand, many fossils are found in Wanganui Basin sediments.[13]

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.[21]

Evolution

An embedded fossil of Penion crawfordi from Cape Palliser, New Zealand

Penion is currently classified within Buccinidae, the family of true whelks.[6][22] Alternatively they have been referred to the family of Buccinulidae.[6][22]


Antarctoneptunea

Kelletia

Penion

Aeneator

Buccinulum

Cladogram for Penion and related genera.[6][11][22]


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][11][22] Radulae and opercula morphology is similar between Penion and Kelletia.[12] 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.[11]

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.[1] 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.[1]

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.[23][24]

Species

Penion mandarinus with eggs at Kennon Cove in Flinders
Penion sulcatus near Goat Island, New Zealand with a damaged operculum

Species and subspecies in the genus Penion include:

Nomina dubia

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

Species brought into synonymy

Reclassified species

References

  1. 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.
  2. 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.
  3. 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.
  4. Iredale, T. 1914. On some invalid molluscan generic names. Proceedings of the Malacological Society of London 11: 170-178.
  5. Vaux, Felix (2017). Evolutionary lineages and the diversity of New Zealand true whelks (Doctor of Philosophy thesis). Massey University. hdl:10179/13113.
  6. 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.
  7. 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.
  8. Powell A. W. B., New Zealand Mollusca, William Collins Publishers Ltd, Auckland, New Zealand 1979 ISBN 0-00-216906-1
  9. 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
  10. 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.
  11. 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.
  12. 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.
  13. Beu, Alan G.; Maxwell, P.A. (1990). "Cenozoic Mollusca of New Zealand". New Zealand Geological Survey Paleontological Bulletin. New Zealand Geological Survey Bulletin. Lower Hutt, New Zealand: New Zealand Department of Scientific and Industrial Research. 58. ISSN 0114-2283.
  14. Dell, R.K.. 1962. New Zealand Marine Provinces - do they exist? Tuatara, 10: 43 - 52. Online Copy courtesy of New Zealand Electronic Text Collection
  15. Wilson, B. 1994. Australian Marine Shells. Prosobranch Gastropods. Kallaroo, WA : Odyssey Publishing Vol. 2 370 pp.
  16. 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.
  17. 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.
  18. 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
  19. Beu, Alan G. (2009). "Before the ice: Biogeography of Antarctic Paleogene molluscan faunas". Palaeogeography, Palaeoclimatology, Palaeoecology. 284 (3–4): 191–226. doi:10.1016/j.palaeo.2009.09.025.
  20. 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. doi:10.1371/journal.pone.0114743. PMC 4262473. PMID 25493546.
  21. 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.
  22. 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.
  23. Green, R.C.; Pullar, W.A. (1960). "Excavations at Orongo Bay, Gisborne". The Journal of the Polynesian Society. 69 (4): 332–353.
  24. 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.
  25. 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..

Further reading

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