Anthopleura Elegantissima Class: Anthozoa, Hexacorallia

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Anthopleura elegantissima Class: Anthozoa, Hexacorallia

Aggregating or clonal anemone Order: Actiniaria, Enthemonae

Family: Actinioidea, Actiniidae

Taxonomy: Anthopleura elegantissima was 1980). The column has a groove below the originally described by Brandt in 1835 as Ac- tentacles (fosse) covered by a distinct fold tinia elegantissima. The subclass Zoantharia (parapet or collar). The anemone becomes a has been synonymized with Hexacorallia hemispheric glob when contracted (fig. 3), (Hoeksema 2015). and blends into its rocky intertidal habitat. It has a hydrostatic skeleton and will emit water Description Medusa: No medusa stage in Anthozoans when stepped on (Kozloff 1983; Ricketts et al. 1985). Polyp: Column: The column can be Size: The average diameter is about twice as high as the diameter when extended, 2.5-4 cm, though the maximum is 5 cm and is hemispherical when contracted. The (Fautin and Hand 2007). Specimens are of- entire column is covered with round verrucae ten larger in bays than on the open coast (tubercules) in longitudinal rows (Hand 1975). (Hand 1955). The illustrated specimen was Collar: The parapet is strong, 3.5 cm high, with a 4.5 cm disc diameter. with a well-developed fosse (groove) (fig. 2). Color: The tentacles are tipped with Oral Disc: The oral disc is a pink, purple or other colors; the illustrated large central area without tentacles on the top specimen had white, green, and maroon of the column. It is broad and flat, with radiat- tentacles, and a green disc with maroon ra- ing lines (mesenterial insertions). It is slightly dial lines. The column is usually green, and wider than the column, or of a similar width. sometimes shades to white at the base The mouth is in the center of the oral disc, (Fautin et al. 1987). The collar is green and and the lips may be swollen or flush with the acrorhagi are white to yellow (Fautin and surface of disc and are not ribbed. Hand 2007) (figs. 2, 3). Puget Sound forms Tentacles: There are more than are often pink and green (Ricketts et al. 24 tentacles (Fautin and Hand 2007). They 1985). Some of the green, especially in the are pointed, and about 1/4 as long as the di- tentacles, is caused by symbiotic algae cells ameter of the disc (fig. 3). There is no oral in- (Kozloff 1983); however, the majority of the ner ring of tentacles, and usually more than 5 coloring is from pigment cells produced by orders (rows) are present. There are no acon- the anemone to protect against UV rays. tia (thread-like defensive structures expelled Thus, in darker habitats the greens fade until through column wall). the anemone is white (Fautin and Hand Mesenteries: These are vertical 2007). body partitions. There are from 6 in young Body: The anemone has a strong specimens to more than 24 pairs in mature collar, broad flat disc, and slender pointed adults. They are visible at high magnification tentacles. The column has longitudinal rows as vertical lines on column, particularly near of tubercules, which are adhesive and cre- the base, and can be irregular, due to asexual ate a layer of attached shells and debris. fission (not shown). Body walls are soft and thin (Haderlie et al.

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: https://oimb.uoregon.edu/oregon-estuarine-invertebrates and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

Piazzola, C.D. 2015. Aequorea victoria. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR.

Pedal Disc: This species has cles (not pink-tipped or with radial lines on the a well-developed pedal disc that attaches to disc). The tentacles are in 5 or more rows the substrate. Its shape varies from circular (Haderlie et al. 1980). Its verrucae completely to very irregular (Hand 1955). The base is cover the column (they are not in rows). An- usually the same diameter as column. There thopleura artemisia has tubercules on the up- are no physa (bulbs) at the base. per 2/3 of its column only; the column is white Cnidae: There are several kinds of or pink below and usually gray or black cnidae in the tentacles, column, acrorhagi, above; tentacles are brightly colored and pat- actinopharynx and filaments (not shown); terned (red in Coos Bay). Anthopleura artemi- see Metridium (Hand 1955). sia is more likely to be found burrowing in a Acrorhagi: Also known as spherules, sandy or muddy substrate than A. elegantissi- these fighting tentacles are round, hollow ma, which can live close by. It also lacks the bodies covered with nematocysts. They are algae symbionts that are found in A. ele- inconspicuous at the top of the column just gantissima. outside the tentacles (fig. 2) (Hand 1955). A third species of Anthopleura, A. sola, Verrucae: These are simple, circular is very similar in appearance to A. elegantissi- tubercules (Fautin and Hand 2007). They ma. The primary difference between the two are adherent, and collect gravel, shell, and is that A. elegantissima is clonal while A. sola debris. This layer helps prevent desiccation is solitary. Though A. sola is not found locally, and protects the anemone from UV rays there are many examples in the literature of (MacGinitie and MacGinitie 1968; Ricketts et A. sola being misclassified as A. elegantissi- al. 1985). Verrucae on the collar are forked ma (Pearse and Francis 2000). and compound (see A. artemisia, fig. 3 in Other sand-dwelling anemones might this guide). Those on the column are ar- include Flosmaris grandis, a southern form, ranged in distinct longitudinal rows, are not which is vermiform and has a translucent or densely packed, and become fewer toward white column (Fautin and Hand 2007). Most the base ("limbus") (Hand 1955; Fautin and other elongated or tube-dwelling forms, i.e., Hand 2007). There are many cinclides Order Cerinatharia, are rarely intertidal in our (temporary or permanent pores at tips of area. verrucae) on the column (fig. 4). Ecological Information Possible Misidentifications Range: The type locality is Sitka (Sitchae Is- The genus Anthopleura can be distin- lands), Alaska (Brandt 1835). The range is guished from other estuarine anemones Alaska to southern California. (Metridium, Diadumene) by their acrorhagi Local Distribution: In Coos Bay, they can be inside the fosse under the tentacles, and by found in high abundance at Pigeon Point. the verrucae on their columns. Anthopleura Habitat: Specimens are found on rocky sub- always have a well-developed pedal disc strates in the mid to high intertidal, often in full and a flat, oral disc with a clear central area. sun, where it aggregates in beds of up to 20 Two other species of Anthopleura oc- m and 100,000 animals (Childress 1969; cur in this area: Anthopleura xanthogrammi- Fautin and Hand 2007). They are especially ca is a large open coast species occasional- prevalent in exposed-rocky habitats where ly found in the most marine parts of our estu- sand collects, and are more occasional on aries. It is very large, solitary (not aggregat- open coasts and exposed pilings (Ricketts et ing), with uniformly colored disc and tenta- al. 1985). When found in sand, A. elegantissi-

ma is attached to underlying rock, and can (Haderlie et al. 1980). Eggs are freely- be fully buried at times (Fautin and Hand spawned and brown, no larger than 250 µm 2007). Algae mats in the intertidal create (Sadro 2001), and covered with clusters of hospitable, moist habitats for the aggrega- spines (Fautin and Sebens 1987). Asexually, tions (Niesen 2007). Specimens can survive anemones divide via longitudinal binary fis- in polluted waters (Ricketts et al. 1985), and sion, producing aggregations of "clones" com- can hang from the roof of overhangs mon to this species (all are similar in colora- (Niesen 2007). tion and sex) (Hand 1955; Fautin and Hand Salinity: Collected at 30. 2007). Each division takes about two days to Temperature: Specimens are kept in lab at complete, and only the outer edge of the clon- 12°C. 20°C is considered high temperature al colony divides by fission (MacGinitie and and causes cnidarian bleaching (loss of MacGinitie 1968). symbiotic algae) (Richier et al. 2008). Larva: Sexual reproduction produces feeding Tidal Level: Found from 0 to +4.5 feet planula larvae. At three weeks larvae are ova- above mean lower low water level (Hand loid to cylindrical, covered in cilia, and no 1955). longer than 250 µm with a 70 µm apical tuft. Associates: Green algae (zoochlorellae) They actively swim using the cilia on their api- and dinoflagellates (zooxanthellae) live in cal tuft (Sadro 2001; Siebert 1974). They feed the anemone’s gut tissue and create some by releasing a mucus strand as they swim. of the green coloring; the algae provide The strand collects food particles, and the cil- some nutritional value to their host, though ia then pull the strand up to the mouth and the anemone still requires carnivorous meals ingest it (Siebert 1974). There are no symbi- (Ricketts et al. 1985; Kozloff 1983). The am- otic algae in the larvae; this relationship be- phipod Orchomenella recondita sometimes tween algae and A. elegantissima must be lives in the digestive cavity. Many organ- established at a later life stage (Siebert 1974). isms, including the amphipods Gibberosus Juvenile: Anemones are considered juveniles myersi and Macronassa macromera, the if they are less than 6.5 cm in diameter snail Epitonium tinctum, and the chiton (Sebens 1982b). They are common in intertid- Lepidochitona fernaldi, live in the aggrega- al mussel beds and less common but present tions (Chapman 2007; McLean 2007; Strath- in rock crevices. It is possible that they settle mann and Eernisse 1987). higher in the intertidal and migrate lower to Abundance: Anthopleura elegantissima is the tidepools as they grow (Sebens 1982b). the most abundant anemone on the coast Longevity: These anemones are reputed to (Ricketts et al. 1985), and is the most abun- be very long lived, and are especially suc- dant Anthopleura in Coos Bay. The peak of cessful as an aquarium animal (one particular the breeding season is Sept-Oct, and re- specimen died after about 80 years due to lab cruitment is late fall-winter (Fautin and Se- failure rather than old age) (Ricketts et al. bens 1987). 1985). Growth Rate: Specimens reach adult size Life-History Information two years after settlement (Sebens 1982b; Reproduction: There are both sexual and Fautin and Sebens 1987). The highest growth asexual reproductive cycles. Individual rate is concurrent with the lowest clonal divi- polyps and clonal aggregations are dioe- sion rate in the spring and summer, while the cious (Fautin and Sebens 1987). Sexual highest division rate and lowest growth rate spawning is in September (San Francisco) both occur in the fall and winter (Sebens

1982a). J. T. Carlton (ed.). University of California Food: Anthopleura elegantissima is a car- Press, Berkeley, CA. nivorous stationary hunter in the tidepools 4. CHILDRESS, L. F. 1969. Intra-specific ag- (Niesen 2007) that uses tentacles to capture gression and its relation to the distribution prey (Ricketts et al. 1985). It primarily eats pattern of the clonal sea anemone, An- crustaceans, such as copepods, amphipods, thopleura elegantissima. Ph.D. Stanford and isopods (Haderlie et al. 1980). Food University. preference seems to be genetically deter- 5. FAUTIN, D. G., and C. HAND. 2007. An- mined (Waters 1975). thozoa, p. 173-184. In: The Light and Predators: Specimens are eaten by varied Smith Manual: intertidal invertebrates from intertidal predators, including seastars. The central California to Oregon. J. T. Carlton nudibranchs Aeolidia papillosa and Hermis- (ed.). University of California Press, Berke- senda crassicornis attack the column ley. (McDonald 2007; Ricketts et al. 1985). The 6. FAUTIN, D. G., and K. P. SEBENS. 1987. snail Epitonium tinctum eats the tips of ten- Phylum Cnidaria, Class Anthozoa, p. 83- tacles (McLean 2007). 104. In: Reproduction and development of Behavior: Anemones at the edges of clonal marine invertebrates of the northern Pacif- groups will "attack" neighboring clonal ag- ic coast. M. F. Strathmann (ed.). University gregations with their acrorhagi, causing of Washington Press, Seattle. wounds; a corridor between clonal groups is 7. FAUTIN, D. G., A. E. SIEBERT, and E. N. thus maintained (Francis 1973; Ricketts et KOZLOFF. 1987. Class Anthozoa, p. 68- al. 1985). Symbiotic green algae may aid the 78. In: Marine invertebrates of the Pacific anemone in modifying phototaxis Northwest. E. N. Kozloff (ed.). University (Buchsbaum 1968) and in averting starva- of Washington Press, Seattle. tion (Kozloff 1983). Anemones contract, in- 8. FRANCIS, L. 1973. Intraspecific aggres- flate, and expel nematocysts or detach their sion and its effect on the distribution of An- pedal disc and move when their column is thopleura elegantissima and some related attacked by the nudibranch Aeolidia papil- sea anemones. Biological Bulletin. 144:73- losa (Waters 1975). 92. 9. HADERLIE, E. C., C. HAND, and W. B. Bibliography GLADFELTER. 1980. Cnidaria 1. BRANDT, J. F. 1835. Prodromus de- (Coelenterata): the sea anemones and al- scriptionis animalium ab H. Mertensio in lies, p. 40-75. In: Intertidal invertebrates of orbis terrarum circumnavigatione obser- California. R. H. Morris, D. P. Abbott, and vatorum. Google eBooks, Petropoli. E. C. Haderlie (eds.). Stanford University 2. BUCHSBAUM, V. M. 1968. Behavioral Press, Stanford. and physiological responses to light by 10. HAND, C. H. 1955. The sea anemones of the sea anemone Anthopleura elegantis- central California. Part II. The endomyar- sima as related to its algal symbiotes. ian and mesomyaroan anemones. Was- PhD. Stanford University. mann Journal of Biology. 13:37-99. 3. CHAPMAN, J. W., E. L. BOUSFIELD, 11. —. 1975. Class Anthozoa, p. 85-93. In: and D. E. BOWERS. 2007. Amphipoda: Light's manual: intertidal invertebrates of Gammaridea, p. 545-618. In: The Light the central California coast. S. F. Light, R. and Smith manual: intertidal inverte- I. Smith, and J. T. Carlton (eds.). Universi- brates from central California to Oregon. ty of California Press, Berkeley.

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