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Circular respiratory opening (1) closed at posterior by overlapping side lobes. Translucent mantle covers whole body and extends beyond it as a broad periphery showing the black substrate (2). Atlantic Spain. © V. Urgorri.
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Corambe testudinaria H. Fischer, 1889
Synonyms: Corambe evelinae Er. Marcus, 1958; Quasicorambe testudinaria (H. Fischer, 1889); Neocorambe testudinaria (H. Fischer, 1889).
Current taxonomy: World Register of Marine Species
www.marinespecies.org/aphia.php?p=taxdetails&id=139408
GLOSSARY BELOW
Preface
Corambe species are mainly defined by morphological features and there is not universal agreement on what species should be recognised. This account follows what is currently, in January 2025, accepted by WoRMS. Molecular sequencing in the future may require alterations to what is accepted.
Description
Corambe testudinaria is a small dorid nudibranch reaching a maximum length of 8 mm, but finds usually vary between 1.5 mm and 7 mm (García, Urgorri & López, 1990). From above, its outline is an oval with a posterior respiratory opening which can be varied between a slit 01Ct flic.kr/p/2qHpCcy and a circle 02Ct flic.kr/p/2qHvmvD . The translucent mantle covers the whole body and extends well beyond it as a broad periphery showing the the bryozoan substrate 03Ct flic.kr/p/2qHvh96 which, when it is on bryozoa, matches the mantle pattern 04Ct flic.kr/p/2qHvmtz , 14Ct flic.kr/p/2qHpCLV & 17Ct flic.kr/p/2rBRido . When in motion, part of the head and the posterior tip of the foot may protrude slightly. The body is slightly convex with a low, slightly undulated profile 05Ct flic.kr/p/2qHu81r . The mantle has a thick layer of gelatinous connective tissue 15 Ct flic.kr/p/2qHu6UZ covered by a protective, transparent cuticle, which is sloughed and renewed periodically (García, Urgorri & López, 1990). Below the cuticle, and clearly visible through it, there is an irregular lattice of yellow or whitish lines 02Ct flic.kr/p/2qHvmvD . Each cell enclosed by the lines has one to four brown spots in it. On light specimens the lines are thin and the spots small. On darker specimens, the cells and spots are larger 02Ct flic.kr/p/2qHvmvD . The same pattern of lines and dots is visible through the ventral surface of the mantle 06Ct flic.kr/p/2qHvoXw often more clearly than dorsally through the external cuticle 20Ct flic.kr/p/2rBRq4y .
The rhinophores each have a cylindrical axis with a keeled fold on its posterior face. The axis is partially enfolded by an erect proximal lamella and a taller distal one 02Ct flic.kr/p/2qHvmvD . The quadrangular tip of the distal lamella may be extended forward like a tongue or folded back over the inner lamella. Each rhinophore arises from the head below the mantle through a hole surrounded by a collar 05Ct flic.kr/p/2qHu81r & 16Ct flic.kr/p/2rtg5JP .
Unlike most dorids, C. testudinaria lacks dorsal gills as they are located below the mantle in the posterior third of the pallial groove 01Ct flic.kr/p/2qHpCcy & 06Ct flic.kr/p/2qHvoXw . There are about four gills on each side of a 1 mm long animal (Fischer, 1889) and up to ten on a 5 mm long specimen. The gill sizes increase towards the posterior and each has up to six triangular lamellae on each side of its axis (García, Urgorri & López, 1990). The largest, most posterior, white gills 19Ct flic.kr/p/2rBRkf9 sometimes project into view in the gap at the posterior of the mantle 07Ct flic.kr/p/2qHvmg5 & 18Ct flic.kr/p/2rBKJ2c .
Ventrally, the head consists of a mouth and large oral veil which terminates on each side in an extendible digitiform tentacle 01Ct flic.kr/p/2qHpCcy & 06Ct flic.kr/p/2qHvoXw . The eyes are sunk deeply and not usually visible. The foot has an anterior notch by the mouth and a translucent whitish sole sometimes with a brown, approximately trefoil mark created by the internal digestive gland (García, Urgorri & López, 1990) 06Ct flic.kr/p/2qHvoXw .
Key identification features
The genus Corambe differs from other dorid seaslugs in having ventral gills and anus below the posterior of the mantle.
Corambe testudinaria
Features selected to differentiate from C. obscura are derived from comparison of 73 characters in Martynov & Schrödl (2011).
1) Posterior of mantle is split into two lobes which can vary configuration to form an exhalant slit or circular gap over the ventral gills 01Ct flic.kr/p/2qHpCcy , which may be visible through it 07Ct flic.kr/p/2qHvmg5 .
2) Mantle lattice of yellow or whitish lines mimics encrusting bryozoa 07Ct flic.kr/p/2qHvmg5 .
3) Body has low profile 05Ct flic.kr/p/2qHu81r and is slightly convex in cross section15Ct flic.kr/p/2qHu6UZ . Maximum length usually 7 mm, rarely 8 mm.
4) Four to ten (when animal 5 mm long) elongate gills in connected row on either side below posterior section of mantle 06Ct flic.kr/p/2qHvoXw .
5) Dorsal cuticle is thick 15Ct flic.kr/p/2qHu6UZ and is sloughed and renewed periodically.
6) Usually on encrusting bryozoa; typically on algae14Ct flic.kr/p/2qHpCLV such as Laminaria and Saccorhiza polyschides which may be attached to hard substrate or drifting free.
7) Lives in fully marine salinity.
8) Occurs on Atlantic coasts from Bretagne, France to Gulf of Cadiz, Spain and recorded in Ghana (Edmunds, 2007) 08Ct flic.kr/p/2qHu7W8 . Not yet found in UK; a record in May 2022 in Cornwall was a misidentified Marsenia perspicua.
Similar species
Corambe obscura (A.E. Verrill, 1870)
1) Posterior of mantle is not cleft to form a gap over the ventral gills 09Ct flic.kr/p/2qHvpzP .
2) Mantle pattern does not mimic encrusting bryozoa according to Martynov & Schrödl (2011) 10Ct flic.kr/p/2qHpC7P , but some C. obscura in Atlantic North America do 11Ct flic.kr/p/2qHpCfE .
3) Body has more raised profile than C. testudinaria 10Ct flic.kr/p/2qHpC7P . Maximum length 8 mm, usually shorter.
4) Single, stout, bilamellar gill either side of anus below posterior of mantle 09Ct flic.kr/p/2qHvpzP .
5) Dorsal cuticle is relatively thin and does not slough periodically (Martynov & Schrödl, 2011).
6) Lives in varied habitats on bryozoans encrusting algae and hard substrates; most often in sheltered locations.
7) Lives in marine salinity and especially in brackish water.
8) Originates from Atlantic coast of North America, but now in the Netherlands and the Black Sea 08Ct flic.kr/p/2qHu7W8 .
Marsenia perspicua (Linnaeus, 1758)
1) Anterior of mantle raised into a tubular inhalant siphon. No posterior slit or gap 12Ct flic.kr/p/2qHvmTC .
2) Mantle pattern extremely varied, often mimics organisms other than its tunicate prey. Some forms have network of pale lines enclosing small dots closely resembling Bryozoa and the pattern on C. testudinaria.12Ct flic.kr/p/2qHvmTC &13Ct flic.kr/p/2qHvnbS
3) Body has more raised profile than C. testudinaria 12Ct flic.kr/p/2qHvmTC . Maximum length usually 20 mm, rarely 30 mm.
4) Single, unipectinate gill hidden in mantle cavity behind anterior siphon. Head has distinct eyes, long cephalic tentacles and no oral veil 12Ct flic.kr/p/2qHvmTC
5) Dorsal layer of mantle is sloughed periodically. Internal shell sometimes visible through mantle 13Ct flic.kr/p/2qHvnbS .
6) Found on hard substrate.
7) Lives in marine salinity.
8) Occurs all round Britain and from Northern Norway to the Mediterranean.
Habits and ecology
C. testudinaria is usually found sublittorally, or on the lower shore, living epibiotically on bryozoans encrusting the surface of algae 14Ct flic.kr/p/2qHpCLV such as Saccorhiza polyschides and Laminaria spp. which are floating unattached or attached to hard substrate, including floats of mussel rafts (Iglesias, 2012). The irregular lattice of yellow or whitish lines on its mantle resembles closely the zooids of its bryozoan prey which, combined with the small size of C. testudinaria, makes it difficult to detect. It has been recorded on Conopeum reticulum (Linnaeus, 1767), Chorizopora brongniartii (Audouin,1826), Membranipora membranacea (Linnaeus, 1767) and Electra pilosa (Linnaeus, 1761).
C. testudinaria and its congeners have evolved to live on large algae which swing to and fro in waves and currents and thrash about in storms which may tear the algae away to drift loose with the bryozoa and C. testudinaria continuing existence on them. The low profile of C. testudinaria streamlines it against dislodgement by water movements 05Ct flic.kr/p/2qHu81r and three pairs of dorsoventral muscle bundles 15Ct flic.kr/p/2qHu6UZ , not found in non-corambid dorids, depress further and stabilize the body and may enhance the sucking power of the foot (Martynov & Schrödl, 2011). The positioning of the gills and anus below the mantle shields them from damage 19Ct flic.kr/p/2rBRkf9 by moving algal fronds rubbing and rasping each other. Further protection from abrasion is provided by the mantle having a thick layer of connective tissue and a surface cuticle 15Ct flic.kr/p/2qHu6UZ .
The cuticle prevents admission of oxygenated water through its surface and the usual dorsal gills of dorids are absent on C. testudinaria. A strong respiratory current is created by cilia on the ventral gills. Water enters laterally under the mantle and passes through the gills in the pallial groove before exiting with any faeces from the posterior anus through the hole or slit in the posterior of the mantle. Lacunae in the lateral body walls and upper foot layer provide an additional respiratory surface (Martynov & Schrödl, 2011). The cuticle, pallial groove and dorsoventral muscles are analogous with the shell, pallial groove and pedal retractor muscles of Patellid limpets.
C. testudinaria, like other nudibranchs, is a simultaneous hermaphrodite. The genital openings are on the right of the body below the mantle, just behind the position of the rhinophore. The penis is more or less conical. Maturation is rapid and breeding can commence within four weeks of post veliger settlement (Martynov & Schrödl, 2011).
The spawn is deposited as a transparent, gelatinous, clockwise spiral of 2 or 2½ turns attached by its broad flat face onto or close to, encrusting bryozoa 04Ct flic.kr/p/2qHvmtz . The spiral is 2.5 mm to 4.3 mm in diameter and the ribbon is approximately 0.9 mm wide. The opaque white capsules are usually 100 μm to110 μm in diameter and are arranged in a single layer of oblique rows of 5 to 8 capsules (García, Urgorri & López, 1990). Spawn has been observed in September in Portugal 04Ct flic.kr/p/2qHvmtz . The small body size and short life cycle from metamorphosis of veliger larvae to breeding and hatching of the next generation of veligers fit well with life in an ephemeral and unstable algal environment (Martynov & Schrödl, 2011).
Distribution and status
C. testudinaria occurs on Atlantic coasts from Bretagne, France to the Gulf of Cadiz, Spain (Bouchet & Tardy, 1976) and it is recorded from Ghana (Edmunds, 2007) 08Ct flic.kr/p/2qHu7W8 .First find in Britain was on 13 September, 2025 16Ct flic.kr/p/2rtg5JP ; a record in May 2022 in Cornwall was a misidentified Marsenia perspicua. Long distance ‘rafting’ dispersal is possible while it lives on drifting algae. Corambid species are locally and seasonally abundant members of kelp-frond communities in some regions (Martynov & Schrödl, 2011). The small size and cryptic appearance of C. testudinaria when on encrusting bryozoans may have led to it being overlooked and under recorded. Targeted search with magnification of algal fronds bearing bryozoans, including drifting ones stranded on shores 14Ct flic.kr/p/2qHpCLV , might produce more records.
Acknowledgements
I am indebted to Victoriano Urgorri for images and permission to use figures from García, Urgorri and López (1990). I gratefully thank Jake Baldry, Charlotte Cumming, Guy Freeman, Jeff Goddard, David Kipling, Mick Otten, Brenton Prigge and João Pedro Silva for valued images, information and literature.
References and links
Bouchet P. & Tardy J. 1976. Faunistique et Biogéographie des Nudibranches des cotes francaises de I'Atlantique et de la Manche. Ann. Inst. Océanog. Paris. 52 (2): 205-213. www.researchgate.net/publication/277331332_Faunistique_et...
Edmunds, M. 2007. Opisthobranchiate Mollusca from Ghana: Dendrodorididae and Corambidae. Journal of Conchology. 39(3): 261-263, fig. 1C.
Fischer, H. 1889. Note préliminaire sur le Corambe testudinaria. Boll. Soco Zool. France, 14 (10): 379-381.
gallica.bnf.fr/ark:/12148/bpt6k61324090/f406.image.r=soci...
García, F.J., Urgorri, V. and López, P.J. 1990. Redescripcion de Corambe testudinaria Fischer, 1889. (gastropoda, nudibranchia). Boll. Malacologico 26 (5-9): 113-124. Milano. www.biodiversitylibrary.org/part/184256
Gomoiu, M-T, & Skotka, M. 1997. A new gastropod - Opistobranch at the Romanian Black Sea. Geo-Eco-Marina ,2/1997. www.geoecomar.ro/website/publicatii/Nr.2-1997/16.MTGomoiu...
Iglesias, M.F. 2012. Corambe testudinaria (2 de 3), Asturnatura.com (accessed 15 January, 2025). www.asturnatura.com/fotografia/invertebrados/corambe-test...
Marcus, Er. 1958. Notes on Opisthobranchia. Boletim do Instituto Oceanográfico. 7 (1-2): 31-78. (C. testudinaria as C. evelinae pp.53-56 & figs. 50-55) www.researchgate.net/publication/262717656_Notes_on_Opist...
Marcus, Ev. & Marcus Er. 1967. Some opisthobranchs from Sapelo Island, Georgia, U. S. A. Malacologia,6 (1-2): 199-222. (C. burchi as Doridella burchi p. 205 & figs. 7-12). ia801302.us.archive.org/25/items/biostor-100932/biostor-1... & www.biodiversitylibrary.org/item/47351#page/215/mode/1up
Martynov, A.V. & Schrödl, M. 2011. Phylogeny and evolution of corambid nudibranchs (Mollusca: Gastropoda) Zoological Journal of the Linnean Society 163(2): 585 – 604.
Menéndez , J.L., Oliveros, J. 2004. Accessed 6th January 2025. Corambe testudinaria H. Fischer 1889. AsturNatura.com at www.asturnatura.com/especie/corambe-testudinaria.html.
Picton, B. & Morrow, C. 2023. Nudibranchs of Britain, Ireland and Northwest Europe. Oxford, Princeton University Press. [ C. testudinaria entry includes images of misidentified Marsenia perspicua which will be removed in second edition.]
Otten, M. 2021 (Accessed 7th January 2025.) Mick’s Marine Life. micksmarinelife.blogspot.com/
Roginskaya, I.S. & Grintsov, V.A. 1997. Range expansion of an alien invader - The nudibranch mollusc Doridella obscura Verrill, 1870 (Opisthobranchia: Corambidae) in the Black Sea. Veliger 40 (2):160-164.
www.biodiversitylibrary.org/item/134483#page/174/mode/1up
Van der Loos, L., Otten, M. and Oonk, B. 2021. ‘Uitgestorven’ schijfslak na 80 jaar weer in Nederland. / 'Extinct' disc slug returns to the Netherlands after 80 years. Nature Today. Accessed 7th January 2025.
www.naturetoday.com/intl/nl/nature-reports/message/?msg=2...
Glossary
μm = 0.001 mm
bilamellar = (adj.) having two lamellae.
Bryozoa = phylum of small aquatic invertebrates with boxlike exoskeletons which often form a single-layered sheet which encrusts hard surfaces and/or algal fronds.
cilia (sing. cilium) = microscopic linear extensions of membrane which can act as sensory organs or move in rhythmic waves to create locomotion, or move particles and liquids e.g. inhalant water currents.
congeners = species in the same genus
connective tissue = tissue that connects, supports, binds, or separates other tissues or organs, often an amorphous matrix of collagen or other fibres.
cuticle = tough but flexible, non-mineral, protective outer covering of an organism.
distal = away from centre of body or from point of attachment.
dorid = a sea slug in the infraorder Doridoidei; most with gills and rhinophores on the dorsum which is usually raised in a low to high dome.
dorsoventral = extending along an axis joining the dorsal and ventral surfaces.
epibiont = organism living on surface of another organism.
epibiota = (pl.) organisms living on surface of another organism.
epibiotic = (adj.) of organisms living on surface of another organism.
epibiotically = (adv.) in an epibiotic manner.
exhalant = (adj.) of something used to breathe out.
inhalant = (adj.) of something used for breathing in.
lacuna = (pl. lacunae) a small cavity or pit an anatomical structure.
lamella = (pl. lamellae) a thin layer, membrane, or plate of tissue.
mantle = (of nudibranchs) sheet of tissue forming part or all of dorsal body-surface.
oral veil = anterior extension of head into a flat sheet and/or finger-like processes.
pallial = (adj.) of, relating to, or produced by the mantle (pallium).
proximal = towards the centre of the body or point of attachment.
rhinophores = chemo-receptor tentacles on top of head of nudibranch, below mantle in corambids.
siphon = extension of mantle to form a tube for respiratory water current.
unipectinate = with central axis and series of filaments or lamellae on one side.
veliger = shelled larva of marine gastropod or bivalve mollusc which swims by beating cilia of a velum (bilobed flap).
zooids = individuals which multiply by budding or division to make up a colonial organism such as bryozoa.
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