Belongs within: Littorinoidea.
Contains: Littoraria (Littorinopsis).
The Littorininae are a group of periwinkles found in the littoral fringe, characterised by the production of dimorphic spermatozoa including paraspermatic nurse cells, and egg capsules via a capsule gland (Reid 1989). Shells are turbinate, lacking an umbilicus and with a few-whorled operculum (Wilson 1993).
Arcane periwinkles
Published 15 June 2026
Around the world, the little periwinkles of Littorina and related genera are among the most abundant gastropods to be found in the upper part of the tidal zone of rocky shores. However, distinguishing species of Littorina can be no easy prospect, and is made even more of a challenge along shores of the eastern North Atlantic by the presence of the species L. saxatilis and L. arcana.
These two closely related species are often found intermingled in the same localities. Both exhibit significant variability in shell morphology and coloration. Some features may be more commonly associated with one than the other—for instance, ‘patula’-type shells without a flare at the base of the aperture are more common in Littorina arcana—but ultimately there seems to be no way to reliably distinguish the two on external features alone. The biggest difference between the two species is rather in their life history: L. saxatilis is ovoviviparous whereas L. arcana lays egg masses. As a result, they do exhibit differences in the structure of the pallial oviduct that can be identified through dissection—reliable, perhaps, but hardly convenient.
You may be wondering what caused these species to diverge if they are not found in different habitats. And you would not be alone in wondering this; the Littorina saxatilis complex has been and remains a much-studied subject for evolutionary studies. Some have wondered whether they might not truly be distinct, whether reproductive mode may switch in response to environmental factors. However, genetic studies have confirmed that the two taxa represent separate populations (e.g. Crossland et al. 1996), albeit populations that have only diverged recently, within the last three million years or so. And one interesting detail is that, though the species may happily intermingle as adults, breeding individuals of L. arcana migrate lower downshore in search of damper locales to lay their eggs (Hannaford Ellis 1985). Might the populations be less sympatric than they appear at first glance?
Systematics of Littorininae
<==Littorininae [Littorinae]
|--Cenchritis von Martens 1900R89
|--Nina Gray 1850BR17 [incl. Echininus Clench & Abbott 1942BR05; Echinininae]
| `--*N. cumingii (Philippi 1846) [=Trochus cumingii, *Echininus cumingii]BR17
|--Mainwaringia Nevill 1885R89
| `--*M. paludomoidea (Nevill 1884) [=Melania (*Mainwaringia) paludomoidea]P84
|--Macquariella Finlay 1927W93
| |--*M. hamiltoni (Smith 1898)P61 [=Paludestrina hamiltoniP61, Laevilitorina (*Macquariella) hamiltoniF26]
| |--M. aucklandica Powell 1933P61
| |--M. delli Powell 1955P61
| `--M. kingensis (May 1924) [=Natica kingensis]W93
|--Tectarius Valenciennes 1833BR05, BR17 (see below for synonymy)
| |--*T. coronatus Valenciennes 1832S98
| |--‘*Echinella’ grandinatus [=*Echinellopsis grandinatus]W93
| |--T. malaccanus (Philippi 1847) [=Littorina malaccana]H09
| |--T. muricatus (Linnaeus 1758)S98
| |--T. nodulosus (Gmelin 1791) [=Trochus nodulosus]H09
| |--T. pagodus Linnaeus 1758WG71 (see below for synonymy)
| |--T. pyramidalisH15
| `--T. rusticus (Philippi 1846)W93
|--Peasiella Nevill 1885R89
| |--*P. tantilla (Gould 1849) (see below for synonymy)R89
| |--P. conoidalis (Pease 1868) (see below for synonymy)R89
| |--P. infracostata (Issel 1869) (see below for synonymy)R89
| |--P. isseli (Semper in Issel 1869) (see below for synonymy)R89
| |--P. lutulenta Reid 1989R89
| |--P. minuta (Deshayes 1824) (see below for synonymy)R89
| |--P. pontileviensis (Morgan 1915) (see below for synonymy)R89
| |--P. reyti (Cossmann & Peyrot 1917) [=Pseudonina reyti]R89
| `--P. roepstorffiana (Nevill 1885) (see below for synonymy)R89
|--Nodilittorina von Martens 1897 [incl. Austrolittorina Rosewater 1970]W93
| | i. s.: N. antipoda (Philippi 1847)R96
| | ‘Littorina’ acutispira Smith 1892 [=L. (Austrolittorina) acutispira; incl. L. infans Smith 1892]W93
| | N. cincta (Quoy & Gaimard 1833)R96
| | N. porcata (Philippi 1846) [incl. Peasiella roosevelti Bartsch & Rehder 1939]R89
| | N. praetermissa (May 1908)R96 [=L. (Austrolittorina) praetermissaW93]
| | N. unifasciata (Gray 1826)R96 (see below for synonymy)
| |--N. (Nodilittorina)W93
| | |--*N. (N.) pyramidalis (Quoy & Gaimard 1833) (see below for synonymy)W93
| | |--N. (N.) australis (Quoy & Gaimard 1826) [incl. N. rugosa Menke 1843]W93
| | `--N. (N.) nodosa (Gray 1839)W93
| |--N. (Granulittorina Habe & Kosuge 1966)W93
| | |--N. (*G.) millegrana (Philippi 1848)W93 (see below for synonymy)
| | `--‘Granulittorina’ reticulata (Anton 1838)JG19
| `--N. (Tectinus) antoni (Philippi 1846) [incl. Echinus nodulosus var. major Usticke 1969]BC01
|--Melarhaphe Menke 1828R89 [=Melarapha Cristofori & Jan 1832R66, Melaraphe (l. c.)BR17; Melarhaphidae]
| |--M. neritoides (Linnaeus 1758)P61, R77 (see below for synonymy)
| |--M. acutispiraH15
| |--M. cincta (Quoy & Gaimard 1833)P61
| |--M. infansH15
| |--M. mauritianaH15
| |--M. mausseneti (Cossmann 1907)TTE93
| |--M. nebulosa (Lamarck 1822) (see below for synonymy)W62
| |--M. oliveri Finlay 1930P61
| |--M. punctata [=Littorina punctata]PP64
| |--M. rissoides (Deshayes 1861)TTE93
| |--M. scabraH15
| |--M. scutulata (Gould 1849)O27 (see below for synonymy)
| |--M. unifasciata Gray 1826WG71
| |--M. zelandiae Finlay 1926F26
| `--M. ziczac (Gmelin 1791) (see below for synonymy)A64
|--Littoraria Griffith & Pidgeon 1834R89
| | i. s.: L. aberrans (Philippi 1846)R01
| | L. flammea (Philippi 1847)R01
| | L. flava (King & Broderip 1832)AS06
| | L. irrorata (Say 1822)R01
| | L. rosewateri Reid 1999R01
| |--*L. (Littoraria) zebra [=Turbo zebra]W93
| |--L. (Littorinopsis)W93
| |--L. (Palustorina Reid 1986)W93
| | |--L. (*P.) melanostoma (Gray 1839)W93, R01 [=Littorina melanostomaW93]
| | |--L. (P.) articulata (Philippi 1846)W93 (see below for synonymy)
| | |--L. (P.) carinifera (Menke 1830)R01
| | |--L. (P.) sinensis (Philippi 1847) (see below for synonymy)R01
| | |--L. (P.) strigata (Philippi 1846) [=Littorina intermedia var. strigata]R01
| | `--L. (P.) sulculosa (Philippi 1846)W93
| `--L. (Prolittoraria)R01
| |--L. (P.) coccinea (Gmelin 1791)R01 [incl. L. limax Gray 1839W93, L. obesa Sowerby 1832W93]
| |--L. (P.) glabrata (Philippi 1846)R01
| |--L. (P.) mauritiana (Lamarck 1822)R01
| |--L. (P.) pintado (Wood 1828)R01
| `--L. (P.) undulata (Gray 1839)R01 (see below for synonymy)
`--Littorina Férussac 1822BR05
| i. s.: L. aberrans Philippi 1846PP78
| L. acicularis Stoliczka 1868S98
| L. albicarinata McLean 1970BC01
| L. anatina [=Paludina anatina]G40
| L. arcana Hannaford Ellis 1978CC96
| L. asperaC64
| L. coccinea (Martyn 1784) [=Limax coccineus]H09
| L. conspersaC64
| L. fabalisW04
| L. fasciataPB27
| L. filosaPRM96
| L. flavaG79
| L. irrorata Say 1822R77
| L. kurilaC64
| L. lineolata Orbigny 1842 [incl. L. carinata Orbigny 1842 non Sowerby 1819, L. jamaicensis Adams 1850]A64
| L. littoralisC60 [=Nerita litoralisJ64]
| L. mariae Sacchi & Rastelli 1966R77
| L. mauritiana (Lamarck 1822)F26 [=Phasianella mauritianaH09]
| L. melanacme Smith 1876H09
| L. muriatica [=Turbo muriaticus]G40
| L. muricataPB27
| L. neglectaCC96
| L. nigrolineata Gray 1839R77
| L. nodulosa [incl. L. dilatata, L. tuberculata]G79
| L. obesaF66
| L. pallescens Philippi 1846H09
| L. palliataH15
| L. philippii [incl. L. dubiosa, L. parvula, L. penicillata]C64
| L. picta Philippi 1846R77
| L. pintado Wood 1828R77
| L. pulchraPB27
| L. pullataC64
| L. saxatilisZLK11
| L. tenebrosaC64
| L. tessellata Philippi 1847A64 (see below for synonymy)
| L. ulvaeG40
| L. undulata Gray 1839H09
| L. variaPB27
|--L. keenaeW04
|--L. (Algamorda Dall 1918)O27
| |--L. (A.) castanea (Carpenter 1865) [=Paludinella castanea]O27
| |--L. (A.) newcombiana (Hemphill 1876) [=Paludinella newcombiana]O27
| `--L. (A.) subrotundata (Carpenter 1865) [=Assiminea subrotunda]O27
`--+--+--*L. (Littorina) littorea (Linnaeus 1758)O27, W04, P77 [=Turbo littoreusO27; incl. L. grandisC64]
| `--L. (Algaroda Dall 1918)O27
| `--L. (A.) squalida Broderip & Sowerby 1829O27, W04, O27
`--+--L. mandschuricaW04
`--+--L. breviculaW04
`--+--L. obtusataW04
`--L. (Littorivaga Dall 1918)O27
|--L. (L.) aleutica Dall 1872O27, W04, O27
|--L. (L.) atkana Dall 1886O27
|--L. (L.) groenlandica Menke 1830O27
|--L. (L.) planaxis Philippi 1847O27 [incl. L. patulaC64, L. tenebrata Phil. 1845C64]
|--L. (L.) rudis (Manton 1797)O27, R77 [=Turbo rudisO27]
`--L. (L.) sitchana Philippi 1845O27 (see below for synonymy)
Nomen nudum: Melarhaphe rupestrisR66
Littoraria (Palustorina) articulata (Philippi 1846)W93 [=Littorina intermedia var. articulataR01; incl. Littoraria blanfordi Dunker 1871W93]
Littoraria (Palustorina) sinensis (Philippi 1847) [=Litorina sinensis, Littorina intermedia var. sinensis; incl. Littorina adonis Yokoyama 1927, Littoraria adonis, Litorina strigata Lischke 1871 non Littorina intermedia var. strigata Philippi 1846, Littoraria scabra strigata, Littorina (Melaraphe) strigata, Littorinopsis strigata]R01
Littoraria (Prolittoraria) undulata (Gray 1839)R01 [incl. L. acuminata Gould 1849W93, L. columna Jonas in Philippi 1847W93, L. scabra concolor Weinkauff 1878W93, L. undulata contracta Nevill 1855W93, L. undulata subintermedia Nevill 1855W93, L. undulata sulcatula Nevill 1855W93, L. tenuis Philippi 1846W93]
Littorina tessellata Philippi 1847A64 [incl. Phasianella lineata Lamarck 1822 non Buccinum lineatum Gmelin 1791A64, L. lineataG79]
Littorina (Littorivaga) sitchana Philippi 1845O27 [incl. L. cincta Gld. 1847C64, L. sitchana var. modestaC64, L. subtenebrosaC64, L. sulcataC64]
Melarhaphe nebulosa (Lamarck 1822) [=Phasianella nebulosa, Littorina lineata var. nebulosa; incl. Litt. columellaris d’Orbigny 1840, Litorina exarata Philippi 1848, Litt. sayi Philippi 1846, Litt. tigrina d’Orbigny in La Sagra 1842]W62
Melarhaphe neritoides (Linnaeus 1758)P61, R77 [=Littorina neritoidesP61; incl. Turbo caerulescensR66, M. coerulescensR66, Paludina glabrata Pfeiffer 1828P61, BR17, *M. glabrataBR17]
Melarhaphe scutulata (Gould 1849)O27 [=Littorina (Melarhaphe) scutulataO27; incl. L. lepidaC64, L. pedroana Conrad 1855C64, L. plenaC64]
Melarhaphe ziczac (Gmelin 1791) [=Littorina ziczac; incl. L. debilis Philippi 1846, L. dispar Montagu 1815, L. dorbignyana Philippi 1847]A64
Nodilittorina unifasciata (Gray 1826)R96 [=Littorina unifasciataR96, L. (*Austrolittorina) unifasciataW93; incl. L. acuta Menke 1843W93, L. mauritiana crassior Philippi 1847W93, L. diemensis Quoy & Gaimard 1833W93, L. diemenensis pseudolaevis Nevill 1885W93]
Nodilittorina (*Granulittorina) millegrana (Philippi 1848)W93 [=Littorina millegranaW93; incl. N. chaoi Yen 1936W93, N. erronea Nevill 1885W93, N. eudeli Sowerby 1915W93, Littorina granicostata Smith 1887JG19, W93, N. granocostata Reeve 1857W93, N. insularis Smith 1889W93, N. melanacme Smith 1876W93, N. novaezelandiae Reeve 1857W93, N. radiata Eydoux & Souleyet 1852W93, N. ventricosa strubelli von Martens 1897W93, N. vidua Gould 1859W93]
*Nodilittorina (Nodilittorina) pyramidalis (Quoy & Gaimard 1833) [=Littorina pyramidalis; incl. N. cecillei Philippi 1851, N. malaccana Philippi 1847, N. monilifera Eydoux & Souleyet 1852, N. nodulosus Gmelin 1791 non Solander 1766, N. trochiformis Dillwyn 1817 non Brocchi 1814, N. trochoides Gray 1839, N. vilis Menke in Philippi 1846]W93
Peasiella conoidalis (Pease 1868) [=Trochus conoidalis, Risella (Peasiella) conoidalis; incl. Echinella gaidei Montrouzier in Souverbie & Montrouzier 1879, R. (P.) gaidei]R89
Peasiella infracostata (Issel 1869) [=Risella infracostata; incl. Cyclostrema fuscopiperata Turton 1932, R. (Peasiella) tantillus var. subinfracostata Nevill 1885]R89
Peasiella isseli (Semper in Issel 1869) [=Risella isseli; incl. R. isseli var. carinata Pallary 1926, R. isseli var. mauritiana Viader 1951, Trochus sismondae Issel 1869, R. isseli var. undata Pallary 1926]R89
Peasiella minuta (Deshayes 1824) [=Trochus minutus, Bembicium minutum, Risella minuta; incl. Xenophora bouryi Cossmann 1888]R89
Peasiella pontileviensis (Morgan 1915) [=Xenophora (Haliphaebus) pontileviensis, Circulus pontileviensis; incl. Tornus orthezensis Cossmann & Peyrot 1918, C. orthezensis]R89
Peasiella roepstorffiana (Nevill 1885) [=Risella (Peasiella) roepstorffiana; incl. R. balteata Preston 1908, R. (P.) templiana var. nigrofasciata Nevill 1885, R. (P.) templiana var. subimbricata Nevill 1885, R. (P.) templiana Nevill 1885]R89
*Peasiella tantilla (Gould 1849) [=Trochus tantillus, Bembicium tantillum, Risella (Peasiella) tantilla, R. (P.) tantillus; incl. Margarita angulata Adams 1853, Trochus diminutivus Reeve 1862, Risella parvula Dunker 1861, R. (Peasiella) tantillus var. parvula]R89
Tectarius Valenciennes 1833BR05, BR17 [incl. Echinella Swainson 1840R89, Echinellopsis Rovereto 1899W93, Fectaria Philippi 1846W93, Hamus Klein in Adams & Adams 1858W93, Pagodella Swainson 1840W93, Pagodus Gray 1839W93; Tectariinae]
Tectarius pagodus Linnaeus 1758WG71 [=*Fectaria pagodusW93, *Hamus pagodusW93, *Pagodella pagodusW93, incl. *Pagodus pagodusW93]
*Type species of generic name indicated
References
[A64] Abbott, R. T. 1964. Littorina ziczac (Gmelin) and L. lineolata Orbigny. Nautilus 78 (2): 65–66.
[AS06] Andrade, S. C. S., & V. N. Solferini. 2006. The influence of size on the radula of Littoraria angulifera (Gastropoda: Littorinidae). Malacologia 49 (1): 1–5.
[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.
[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.
[BC01] Boyko, C. B., & J. R. Cordeiro. 2001. Catalog of Recent type specimens in the Division of Invertebrate Zoology, American Museum of Natural History. V. Mollusca, part 2 (class Gastropoda [exclusive of Opisthobranchia and Pulmonata], with supplements to Gastropoda [Opisthobranchia], and Bivalvia). Bulletin of the American Museum of Natural History 262: 1–170.
[C64] Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 33: 517–686.
[C60] Cox, L. R. 1960. Gastropoda: general characteristics of Gastropoda. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I84–I169. Geological Society of America, and University of Kansas Press.
[CC96] Crossland, S., D. Coates, J. Grahame & P. J. Mill. 1996. The Littorina saxatilis species complex—interpretation using random amplified polymorphic DNAs. In: Taylor, J. D. (ed.) Origin and Evolutionary Radiation of the Mollusca pp. 205–209. Oxford University Press: Oxford.
[F26] Finlay, H. J. 1926. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.
[F66] Fraser, L. 1866. Communication of a list of mollusks collected by R. Swinhoe, Esq., in Formosa. Proceedings of the Zoological Society of London 1866: 146.
[G40] Gray, J. E. 1840. A Manual of the Land and Fresh-water Shells of the British Islands, with figures of each of the kinds. By William Turton, M.D. A new edition, thoroughly revised and much enlarged. Longman, Orme, Brown, Green, and Longmans: London.
[G79] Guppy, R. J. L. 1879. First sketch of a marine invertebrate fauna of the Gulf of Paria and its neighbourhood. Part I.—Mollusca. Journal of Conchology 2: 151–172.
Hannaford Ellis, C. J. 1985. The breeding migration of Littorina arcana Hannaford Ellis, 1978 (Prosobranchia: Littorinidae). Zoological Journal of the Linnean Society 84: 91–96.
[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.
[H15] Hedley, C. 1915. Presidential address. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 1–77, pls 1–7.
[JG19] James, D. J., P. T. Green, W. F. Humphreys & J. C. Z. Woinarski. 2019. Endemic species of Christmas Island, Indian Ocean. Records of the Western Australian Museum 34 (2): 55–114.
[J64] Jeffreys, J. G. 1864. The Upper Tertiary fossils at Uddevalla, in Sweden. Report of the British Association for the Advancement of Science 33 (Notices and Abstracts): 73–79.
[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).
[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d’Endoume, Bulletin 31 (27): 5–137.
[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.
[P84] Ponder, W. F. 1984. A review of the genera of the Iravadiidae (Gastropoda: Rissoacea) with an assessment of the relationships of the family. Malacologia 25 (1): 21–71.
[PP78] Poorman, F. L., & L. H. Poorman. 1978. Additional molluscan records from Bahía de Los Angeles, Baja California Norte. Veliger 20 (4): 369–374.
[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.
[P77] Pratt, D. M. 1977. Homing in Urosalpinx cinerea in response to prey effluent and tidal periodicity. Veliger 20 (1): 30–32.
[PRM96] Preston, S. J., D. Roberts & W. I. Montgomery. 1996. The Mollusca: coelomate turbellarians or mesenchymate annelids? In: Taylor, J. D. (ed.) Origin and Evolutionary Radiation of the Mollusca pp. 313–325. Oxford University Press: Oxford.
[R77] Raffaelli, D. G. 1977. Observations on the copulatory behavior of Littorina rudis (Maton) and Littorina nigrolineata Gray (Gastropoda: Prosobranchia). Veliger 20 (2): 75–77.
[R89] Reid, D. G. 1989. Systematic revision of the Recent species of Peasiella Nevill, 1885 (Gastropoda: Littorinidae), with notes on the fossil species. Nautilus 103 (2): 43–69.
[R01] Reid, D. G. 2001. New data on the taxonomy and distribution of the genus Littoraria Griffith and Pidgeon, 1834 (Gastropoda: Littorinidae) in Indo-West Pacific mangrove forests. Nautilus 115 (4): 115–139.
[R66] Rosewater, J. 1966. Reinstatement of Melarhaphe Menke, 1828. Nautilus 80 (2): 37–38.
[R96] Rudman, W. B. 1996. A note on the egg capsule of Nodilittorina unifasciata from eastern Australia. Molluscan Research 17: 111-114.
[S98] Stilwell, J. D. 1998. Late Cretaceous Mollusca from the Chatham Islands, New Zealand. Alcheringa 22 (1): 29–85.
[TTE93] Tracey, S., J. A. Todd & D. H. Erwin. 1993. Mollusca: Gastropoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 131–167. Chapman & Hall: London.
[W62] Weisbord, N. E. 1962. Late Cenozoic gastropods from northern Venezuela. Bulletins of American Paleontology 62: 7–672.
[W04] Wilke, T. 2004. How dependable is a non-local molecular clock? A reply to Hausdorf et al. (2003). Molecular Phylogenetics and Evolution 30: 835–840.
[W93] Wilson, B. 1993. Australian Marine Shells vol. 1. Prosobranch Gastropods. Part One. Odyssey Publishing: Kallaroo (Western Australia).
[WG71] Wilson, B. R., & K. Gillett. 1971. Australian Shells: illustrating and describing 600 species of marine gastropods found in Australian waters. A. H. & A. W. Reed: Sydney.
[ZLK11] Zou, S., Q. Li & L. Kong. 2011. Additional gene data and increased sampling give new insights into the phylogenetic relationships of Neogastropoda, within the caenogastropod phylogenetic framework. Molecular Phylogenetics and Evolution 61: 425–435.