Sea Slugs
Sea slugs is an informal group that comprise a remarkably diverse assemblage of gastropod molluscs, with more than 6,000 known species of marine heterobranch and related forms. They inhabit a wide range of environments, from shallow coastal waters and coral reefs to deep-sea habitats and the open ocean. The informal term sea slug is commonly applied to nudibranchs, sea hares, sacoglossans, side-gilled slugs and several groups of cephalaspideans, among others.

Despite their striking differences in body shape, colour, size and lifestyle, many of these animals share a characteristic soft-bodied organisation. The shell, when present, is often reduced, internal or concealed beneath the mantle, and it is completely absent in the adult stage of many groups. The head commonly bears sensory structures such as tentacles, rhinophores or other specialised appendages, while the muscular foot is generally adapted for crawling in benthic species or modified into swimming structures in some pelagic forms. Dorsal projections, including cerata and other papillae, may contribute to respiration, defence or the distribution of internal organs, although their presence and function vary among lineages.
The internal anatomy also reflects a common gastropod heritage. The circulatory system is generally open, with a heart that usually includes an atrium and a ventricle, while the nervous system consists of interconnected ganglia and nerve cords whose degree of fusion and organisation differs among groups. Sensory structures are often highly developed relative to the animals’ ecological needs. Chemoreception is particularly important in locating food, recognising potential mates and detecting environmental cues; it may involve rhinophores, cephalic tentacles, oral sensory structures or other specialised organs. Eyes are usually small, and statocysts provide information related to balance and orientation. The exact arrangement of these structures varies, and not every group possesses all of them in the same form.
Most sea slugs are simultaneous hermaphrodites, possessing both male and female reproductive functions, although reproductive anatomy and mating strategies differ considerably. Many deposit their eggs in distinctive gelatinous masses attached to a substrate, while others have different reproductive and developmental strategies. Development may include a planktonic veliger larva, which can contribute to dispersal, or direct development, in which juveniles hatch without a free-swimming larval stage. Planktotrophic larvae feed in the plankton, whereas lecithotrophic larvae rely mainly on yolk reserves. These developmental modes influence dispersal and recruitment, but their effects depend on larval duration, behaviour, currents and habitat availability.
The diversity of feeding strategies is equally remarkable. Marine heterobranchs may feed on algae, sponges, cnidarians, bryozoans, tunicates, other molluscs or a variety of other organisms. Many have highly specialised diets and possess feeding structures adapted to their prey. Some sacoglossans retain functional chloroplasts acquired from algae, a phenomenon known as kleptoplasty, while certain nudibranchs can sequester defensive compounds or stinging cells from their prey, something known as kleptocnidy. Other species rely on camouflage, warning colouration, mucus or behavioural responses to avoid predators.
The traditional taxonomic group Opisthobranchia is no longer recognised as a formal unit in modern gastropod classification. Its former members are now placed within a broader phylogenetic framework, particularly among the heterobranch gastropods. Euthyneura is a wider evolutionary grouping that includes many marine sea slugs, pteropods and numerous non-marine gastropods. Consequently, this guide uses sea slugs as a familiar common name while also considering other marine heterobranchs, including pelagic lineages, to reflect their evolutionary relationships and ecological diversity.
Main groups of marine heterobranchs
Marine heterobranchs include several distinct evolutionary lineages, some of which are commonly known as sea slugs.
- Nudibranchia: a highly diverse group of marine gastropods whose adults lack a shell. It includes dorid nudibranchs, often recognised by their dorsal gill plume, and cladobranch nudibranchs, many of which bear cerata or other dorsal appendages.
- Pleurobranchida: side-gilled sea slugs, characterised by a gill situated on the right side of the body beneath the mantle edge.
- Anaspidea: sea hares, generally with a broad body and well-developed parapodia in many species.
- Sacoglossa: a group that includes many algae-feeding species with a specialised radula; some species retain functional chloroplasts obtained from their food.
- Cephalaspidea and related lineages: a diverse assemblage including headshield sea slugs and bubble snails. Some retain an external shell, while others have reduced or lost it.
- Pteropoda: predominantly pelagic gastropods traditionally divided into thecosomes and gymnosomes. Thecosomes generally possess a shell or a reduced shell and use wing-like parapodia for swimming, whereas gymnosomes lack a shell as adults and are specialised predators, often feeding on other pteropods. Their pelagic lifestyle distinguishes them ecologically from most benthic sea slugs, but they are relevant to a broad account of marine heterobranch diversity.
- Other marine heterobranch lineages: additional groups include animals with specialised body forms and lifestyles. Their inclusion depends on the taxonomic and ecological scope adopted.
These groups do not all share the same anatomical features, and their classification continues to be refined as new morphological and molecular evidence becomes available. Euthyneura is a broader evolutionary grouping that encompasses several of these lineages, but it should not be treated as synonymous with sea slugs or with the traditional Opisthobranchia.
Acochlidiimorpha
Aplysiida
Acteonimorpha
Architectonicoidea
Cephalaspidea
Nudibranchia
Pleurobranchida
Pteropoda
Rhodopoidea
Ringiculimorpha
Runcinida
Sacoglossa
Umbraculida
