Platydoris cinereobranchiata
Two Platydoris cinereobranchiata mating. Rhinophores on the right
Etymology of ‘rhinophore’ (Oxford English Dictionary)
Rhino-: Derived from the Ancient Greek word rhis (genitive rhinos), meaning ‘nose’.
-phore: Derived from the Ancient Greek -phoros (‘bearing’ or ‘carrying’), a derivative of pherō (‘to bear or carry’).
Literal meaning: A mixed Neo-Latin and Greek formation translating to ‘carrying noses’ or ‘nose-bearer’.
History: The term entered scientific usage in the 1860s (recorded around 1868) to describe the tentacle-like sensory structures on the heads of sea slugs.
And in other trivia: rhinophore occurs in modern written English 0.02 times for every one million words.
Which probably goes a long way to explain why it’s a word you may not recognise.
Elysia sp. Norfolk Island
Elysia with my hand for scale
Two Elysia showing their scroll-like rhinophores
In this blog post, I am going to take a look at some of the different rhinophores sported by sea slugs in Norfolk Island’s Emily and Slaughter Bays. It was a little green-coloured Elysia with fetching blue dots that sent me looking into all this. I’ve photographed this species plenty of times before – mollusc specialist Dr Richard Willan thinks it may even be an as-yet undescribed species – but while processing a close-up recently I suddenly noticed its rhinophores (left image, above). They looked like two tiny scrolls of tissue, rolled inwards along their length.
There were actually two Elysia in the photograph, mating, but for once that wasn’t what caught my attention. It was those scrolls. Inevitably, I started going back through my other sea-slug photographs.
I had been photographing rhinophores for years without paying terribly much attention to just how different they are. Once I did, there were scrolls, ridges, stacks of little plates, tubes and what look remarkably like rabbit ears. The variation is wonderful.
The ‘nose’ comparison is simplistic, of course. We smell volatile chemicals carried through the air. Sea slugs are detecting chemical information dissolved in seawater. Rhinophores are packed with sensory tissue and are particularly important for detecting cues at a distance. Depending on the animal, those cues can include food, predators and other members of the same species. Comparative work on sea-slug sensory organs has found considerable differences in their external shape, while the basic cellular organisation of the sensory system underneath is much more consistent.
There’s some great experiments showing the nudibranch Tritonia diomedea, with intact rhinophores, following the odour of prey upstream and turning away to evade predator odours. When researchers removed the rhinophores, those responses disappeared.
White-speckled sea hare - Aplysia argus
White-speckled sea hare - Aplysia argus
Sea hares have been particularly useful animals for investigating this. In Aplysia spp., the rhinophores appear to be important for distance chemoreception and also rheoreception – detecting water flow – whereas the oral tentacles at the front of the head seem to have a greater role in contact chemoreception and touch. Researchers have also identified that sea hares can detect water-borne pheromones – chemical signals associated with mating.
In the image of the white-speckled sea hare, you can clearly see a pair of rhinophores on top of its head and a second pair of structures, the oral tentacles, nearer the mouth. The rhinophores are often rolled or enrolled, and their resemblance to rabbit ears is where the common name ‘sea hare’ comes from. Once you know which bits you are looking at, it becomes much easier to see them in my blunt-end sea hare, Dolabrifera brazieri, dwarf sea hares - Aplysia concava, Stylocheilus striatus and white-speckled sea hare photographs.
Blunt-end seahare - Dolabella auricularia
Sea hare - Dolabrifera brazieri
Dwarf sea hare - Aplysia concava
Sea hare - Stylocheilus striatus
The Elysia version is different again. Rolled or grooved rhinophores are characteristic of the genus, although the details vary between species. In some taxonomic descriptions the groove runs almost the entire length; in others the tip itself is rolled.
Then there are the nudibranchs. In Goniobranchus, Halgerda, Dendrodoris, Roboastra and the pustulose wart slug, the rhinophores are much more obviously ridged or lamellate. Some look like a stack of very fine plates around a central stalk; others have broader, more widely spaced folds. The number, shape and arrangement of those lamellae can be sufficiently consistent to help taxonomists distinguish different species.
Those folds aren’t just for show. More folds mean more surface area exposed to the surrounding water – and therefore more area for sampling dissolved chemicals.
My pleurobranchs bring us back towards the scroll idea. The Forskål’s and Peron’s pleurobranchs have curled rhinophores, as do pleurobranchs more generally, although again the exact form differs. The umbrella sidegill has rolled tubular rhinophores tucked away at the front of what is otherwise a fairly improbable-looking animal.
Goniobranchus
Goniobranchus
Halgerda willeyi
Halgerda willeyi
Dendrodoris tuberculosa
Dendrodoris tuberculosa
Slender Roboastra - Roboastra gracilis
Slender Roboastra - Roboastra gracilis
Pustulose wart slug - Phylidiella pustulosa
Pustulose wart slug - Phylidiella pustulosa
Which made me wonder: why all these different shapes?
We know a good deal about what rhinophores detect, something about their sensory cells and nerves, and quite a lot about their anatomy. We also know that rhinophore form and structure varies strongly among different groups and is useful in identifying the animal. What I couldn’t find was good evidence that lets us look at the very fine lamellae on one species and say confidently: that shape evolved because it eats this food, lives in that habitat, or needs a more sensitive ‘nose’. Some of the variation may well have functional consequences. Some of it reflects evolutionary history. Almost certainly both are involved. But unfortunately it is not possible to assign a neat ecological explanation to every scroll, fold or frill.
PhD anyone? In the meantime, I’ll keep looking at rhinophores. I have a feeling I’ve only just started noticing them!
Pseudobiceros sp. with the lamellae on its rhinophores clearly visible
References
Cummins, S. F., Erpenbeck, D., Zou, Z., Claudianos, C., Moroz, L. L., Nagle, G. T., & Degnan, B. M. (2009). Candidate chemoreceptor subfamilies differentially expressed in the chemosensory organs of the mollusc Aplysia. BMC Biology, 7, 28. https://doi.org/10.1186/1741-7007-7-28 PubMed
Göbbeler, K., & Klussmann-Kolb, A. (2007). A comparative ultrastructural investigation of the cephalic sensory organs in Opisthobranchia (Mollusca, Gastropoda). Tissue and Cell, 39(6), 399–414. https://doi.org/10.1016/j.tice.2007.07.002 PubMed
Klussmann-Kolb, A., Croll, R. P., & Staubach, S. (2013). Use of axonal projection patterns for the homologisation of cerebral nerves in Opisthobranchia, Mollusca and Gastropoda. Frontiers in Zoology, 10, 20. https://doi.org/10.1186/1742-9994-10-20 PubMed
Lisova, E. D., & Vortsepneva, E. V. (2022). New data on nudibranchs rhinophore morphology and their spicule complex in Onchidoris muricata (Doridina, Gastropoda). Zoologischer Anzeiger, 296, 58–70. https://doi.org/10.1016/j.jcz.2021.11.003 ScienceDirect
Wertz, A., Rössler, W., Obermayer, M., & Bickmeyer, U. (2006). Functional neuroanatomy of the rhinophore of Aplysia punctata. Frontiers in Zoology, 3, 6. https://doi.org/10.1186/1742-9994-3-6 PubMed Central (PMC)
Wyeth, R. C., & Willows, A. O. D. (2006). Odours detected by rhinophores mediate orientation to flow in the nudibranch mollusc, Tritonia diomedea. Journal of Experimental Biology, 209(8), 1441–1453. https://doi.org/10.1242/jeb.02164
Side gill sea slugs
Forsskål's pleurobranch - Pleurobranchus forskalii
Forsskål's pleurobranch - Pleurobranchus forskalii
Peron's pleurobranch - Pleurobranchus peronii
Peron's pleurobranch - Pleurobranchus peronii
Umbrella slug - Umbraculum umbraculum
Umbrella slug - Umbraculum umbraculum