The aggregating anemone (''Anthopleura elegantissima''), or clonal anemone, is the most abundant
species of
sea anemone
Sea anemones are a group of predation, predatory marine invertebrates of the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemones are classifi ...
found on rocky, tide swept shores along the Pacific coast of North America. This
cnidarian
Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter.
Their distinguishing feature is cnidocytes, specialized cells that th ...
hosts endosymbiotic
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
called
zooxanthellae that contribute substantially to
primary productivity in the
intertidal zone
The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species o ...
. The aggregating anemone has become a model organism for the study of temperate cnidarian-algal symbioses.
Description

The polyps of the aggregating anemone reach up to eight cm across the oral disk with approximately 100 tentacles in three or four rows around the margins of the oral disk. Most are olive to bright green (depending on the species of algal symbionts present) with tentacles tipped in pink. Individuals that live in
microhabitats that are deficient in
photosynthetically active radiation (PAR), such as under docks or in caves, lack symbionts and are pale yellow to white in color.
Range

Pacific coast of North America from
Alaska, United States to
Baja California, Mexico. Most common in protected waters.
Life history
This species of anemone is capable of reproducing both sexually and asexually. As adults, aggregating anemones release
gametes into the water that join to form genetically unique individuals that settle on intertidal rock. This genetically distinct individual can then proliferate through
binary fission. Some argue that this is not true reproduction but actually a form of growth. Fission is often prompted in the autumn by a decrease in the abundance of food and follows sexual spawning in summer. Over time, a single individual can generate a large colony of genetically identical
polyps. Because of its ability to grow in this manner, the genetic entity of a colony is potentially immortal on an ecological time scale.
Natural history
Photosynthetic symbiosis
Aggregating anemones host
endosymbiotic, photosynthetic algae in their tentacles, oral disk and column of the polyps. In addition to a
chlorophyte
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a ...
, two species of
dinoflagellate
The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
''Symbiodinium'', ''S. muscatinei'' and ''S. californium'' are known to inhabit the anemone. These algae are generally referred to as
zooxanthellae (dinoflagellates) and zoochlorellae (chlorophytes) and many polyps concurrently host more than one type of alga within their tissues. The rate of occurrence of each alga is determined by temperature and light regimes of anemone habitats. Zoochlorellae are typically found at higher latitudes and in deeper intertidal habitats than zooxanthellae. Of the two species of zooxanthellae that inhabit the anemone, ''S. californium'' is restricted to southern California due to its intolerance of cold temperatures whereas ''S. muscatinei'' is tolerant of a broad range of temperature and irradiance levels.
The
mutualistic relationship of these organisms requires adaptations of each partner. The algal symbionts convert inorganic carbon into carbohydrates for use by each partner and release oxygen to the animal host in the process. To accommodate the algae, the anemone must provide concentrated
carbon dioxide to their intracellular photosynthetic guests as well as
photosynthetically active radiation (PAR) to fuel the photosynthetic process. This restricts the symbiotic organism to
euphotic habitats and requires consistent exposure to high levels of
ultraviolet radiation (UVR). To protect from potentially damaging UVR, the algae provide mycosporine-like amino acids (MAAs) that act as sunscreen for themselves and the host. The anemones, in turn, produce
antioxidant
Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubricant ...
s called superoxide dismutases to protect against reactive oxygen that causes oxidative stress.
Agonism between colonies

The aggregating anemone is agonistic toward other individuals with different genetic disposition. When one colony of genetically identical polyps encounters a different genetic colony, the two will wage territorial battles. Aggregating anemones have specialized tentacles called acrorhagi that are used solely to deter other colonies from encroaching on their space. When a polyp makes physical contact with a non-clonemate, it extends the acrorhagi to attack the competing anemone with stinging cells called
nematocytes
A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
. Acrorhagi of the attacking anemone leave behind a 'peel' of the
ectoderm
The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from t ...
and nematocysts that causes tissue necrosis in the receiving animals.
A study of two colonies on a boulder removed from the shore and brought into a laboratory revealed that hostilities between neighboring colonies follow the tides. As water rushed into the tank, warrior polyps inflated their acrorhagi, tripled their body length and began reaching into an empty swath of rock between the colonies. Occasionally, a polyp from one of the colonies would move into the spatial zone between the two colonies, acting as a scout, and would be attacked by the warrior polyps of the other clone. If the scout polyp received enough stings, it would be attacked by its clonemates upon return to its own colony. The return of an attacked scout to the clone with acrorhagial peel may serve to communicate the presence and identity of neighboring clones to the interior of the colony.
[Ayre DJ, Grosberg RK (2005) Behind anemone lines: factors affecting division of labour in the social cnidarian Anthopleura elegantissima. Animal Behaviour 70:97–110]
Predators
The few known predators of the aggregating anemone include a species of nudibranch (''
Aeolidia papillosa)'',
leather star, and
mosshead sculpin.
Footnotes
External links
{{DEFAULTSORT:Aggregating Anemone
Actiniidae
Animals described in 1835