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a green slug
live Pontohedyle milaschewitchii
ot = oral tentacles,
ey = eyes,
dg = digestive gland.
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
(unranked): clade Heterobranchia
clade Euthyneura
clade Panpulmonata
clade Acochlidiacea
clade Microhedylacea
Family: Microhedylidae
Genus: Pontohedyle
Golikov & Starobogatov, 1972
at least 12 cryptic species (3 valid, at least 9 undescribed)

Gastrohedyle Rankin, 1979
Mancohedyle Salvini-Plawen, 1973
Maraunibina Rankin, 1979

Pontohedyle is a genus of sea slugs, acochlidians, shell-less marine gastropod mollusks in the family Microhedylidae. Sea slugs in this genus are highly simplified and uniform.[1]


The genus Pontohedyle shows a circumtropical distribution with a single derived species (Mediterranean/ Black Sea Pontohedyle milaschewitchii) inhabiting temperate waters.[1] In the absence of a fossil record for meiofaunal slugs, the only available estimate for divergence times derives from a molecular clock approach, calibrated with shelled heterobranch fossils.[1] Jörger et al. (2010)[3] estimated the origin of the genus Pontohedyle to the late Cretaceous, 84 mya (95% confidence interval ranging from 160–60 mya), providing a rough estimation of how much time was available for diversification and circum-global dispersal of Pontohedyle slugs.[1]

Pontohedyle has never been found in colder waters despite a well-studied meiofauna and hydrographic conditions similar to the Mediterranean.[1] The distribution of Pontohedyle might be constrained by ancestry from warm-water adapted animals.[1] Considering the estimated mid to late Mesozoic origin and the recent primarily tropical distribution pattern in Pontohedyle, it is most likely that this meiofaunal slug clade originated in Tethyan waters.[1]

Map of Pontohedyle distribution.


With its vermiform body, a putatively multi-functional radula, ‘simplified’ organ systems and a special fast and imprecise mode of sperm transfer, Pontohedyle reflects a meiofaunal slug lineage highly adapted to its interstitial habitat.[1]

Pontohedyle is morphologically well-defined genus of meiofaunal slugs.[1] Specimens of Pontohedyle are externally uniform and easily distinguishable from other acochlids by the lack of rhinophores and the bow-shaped oral tentacles.[1] Pontohedyle typically bear monaxone, rodlet-like spicules distributed randomly and frequently accumulated between the oral tentacles.[1] No diagnostic differences in external morphology or spicules could be detected between the collected populations apart from eyes externally visible or not.[1] Comparative SEM-examination of the available radulae revealed two types of the typically hook-shaped radula: a lateral tooth without a denticle (Pontohedyle verrucosa) or with a denticle (Pontohedyle milaschewitchii).[1]

Pontohedyle slugs have a well-adapted body plan that can be conserved for millions of years in a worldwide evolutionary success story.[1] Pontohedyle presents a stunning example of extreme morphological stasis and uniformity over long evolutionary timeframes, probably constrained by their simplified bodyplan and by the requirements of the meiofaunal habitat.[1]


The first discovered Pontohedyle species is well described and abundant Pontohedyle milaschewitchii from the Black Sea and Mediterranean.[1] The second discovered species is poorly known Western Pacific Pontohedyle verrucosa from the Solomon Islands.[1] In absence of distinguishing morphological characters Jörger et al. (2007)[4] synonymized the tropical Western Atlantic Pontohedyle brasilensis with its temperate congener Pontohedyle milaschewitchii.[1] Subsequently (2012) authors tested molecular markers to detect possible cryptic species and they reestablished Pontohedyle brasilensis as a valid taxon and they uncovered at least nine (undescribed) candidate species.[1]

There are at least nine candidate species plus three confirmed valid species:[1]

  • Pontohedyle brasilensis (Rankin, 1979)[1] - tropical, East Atlantic, subtidal, lateral radula tooth with denticle, eyes are sometimes externally visible.[1]
  • Pontohedyle milaschewitchii (Kowalevsky, 1901)[1][5] - temperate, Mediterranean and Black Sea, subtidal, lateral radula tooth with denticle, eyes externally visible.[1]
  • Pontohedyle verrucosa (Challis, 1970)[1][5] - tropical, Central Indo-Pacific and West-Pacific, intertidal, lateral radula tooth without denticle, eyes are sometimes externally visible.[1]
  • Pontohedyle sp. 1[1] - tropical, Central Indo-Pacific, subtidal, lateral radula tooth without denticle, eyes externally visible.[1]
  • Pontohedyle sp. 2[1] - tropical, East Atlantic, subtidal, lateral radula tooth with denticle, eyes externally visible.[1]
  • Pontohedyle sp. 3[1] - tropical, Central-Pacific, subtidal, unknown radula, unkknown eyes.[1]
  • Pontohedyle sp. 4[1] - tropical, Red Sea, subtidal, lateral radula tooth with denticle, eyes externally visible.[1]
  • Pontohedyle sp. 5[1] - tropical, Indian Ocean, subtidal, lateral radula tooth with denticle, eyes externally visible.[1]
  • Pontohedyle sp. 6[1] - tropical, Central Indo-Pacific and Central-Pacific and West-Pacific and Indian Ocean, subtidal, lateral radula tooth with denticle, eyes externally visible.[1]
  • Pontohedyle sp. 7[1] - tropical, West Atlantic, subtidal, lateral radula tooth with denticle, eyes not visible externally.[1]
  • Pontohedyle sp. 8[1] - tropical, Central-Pacific, subtidal, unknown radula, unkknown eyes.[1]
  • Pontohedyle sp. 9[1] - tropical, East-Pacific, subtidal, unknown radula, eyes externally visible.[1]


A cladogram based on sequences of mitochondrial 28S ribosomal RNA, 16S ribosomal RNA and cytochrome-c oxidase I (COI) genes showing phylogenic relations of the genus Pontohedyle:[1]


Pontohedyle sp. 1

Pontohedyle sp. 2

Pontohedyle sp. 3

Pontohedyle sp. 4

Pontohedyle sp. 5

Pontohedyle verrucosa

Pontohedyle sp. 7

Pontohedyle sp. 6

Pontohedyle sp. 8

Pontohedyle sp. 9

Pontohedyle milaschewitchii

Pontohedyle brasilensis


Pontohedyle (and many other meiofauna taxa) are rare and can be expected to have small effective population sizes.[1]


This article incorporates CC-BY-2.0 text from the reference[1]

  1. ^ 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 z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av Jörger K. M., Norenburg J. L., Wilson N. G. & Schrödl M. (2012). "Barcoding against a paradox? Combined molecular species delineations reveal multiple cryptic lineages in elusive meiofaunal sea slugs". BMC Evolutionary Biology 12: 245. doi:10.1186/1471-2148-12-245.
  2. ^ Bouchet, P.; Gofas, S. (2012). Pontohedyle Golikov & Starobogatov, 1972. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=138178 on 2013-02-16
  3. ^ Jörger K. M., Stöger I., Kano Y., Fukuda H., Knebelsberger T. & Schrödl M. (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology 10: 323. doi:10.1186/1471-2148-10-323.
  4. ^ Jörger K. M., Neusser T. P. & Schrödl M. (2007). "Re-description of a female Pontohedyle brasilensis (Rankin, 1979), a junior synonym of the Mediterranean P. milaschewitchii (Kowalevsky, 1901) (Acochlidia, Gastropoda)". Bonner Zoologische Beiträge 55(3/4): 283-290. PDF.
  5. ^ a b Schrödl M. & Neusser T. P. (2010). "Towards a phylogeny and evolution of Acochlidia (Mollusca: Gastropoda: Opisthobranchia)". Zoological Journal of the Linnean Society 158: 124-154. doi:10.1111/j.1096-3642.2009.00544.x.

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