Pontohedyle

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]

Pontohedyle
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
Diversity[1]
at least 12 cryptic species (3 valid, at least 9 undescribed)
Synonyms[2]
Gastrohedyle Rankin, 1979 Mancohedyle Salvini-Plawen, 1973 Maraunibina Rankin, 1979

Distribution

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.

Description

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]

Species

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]

Cladogram

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	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

Ecology

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

References

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

1. 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. 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.

External links

• Jörger K. M. & Schrödl M. (2013). "How to describe a cryptic species? Practical challenges of molecular taxonomy". Frontiers in Zoology 10: 59. doi:10.1186/1742-9994-10-59.