Actiniaria Sea Anemones

Vreni Hussermann & Gnter Frsterra

215

Actiniaria Sea Anemones

Phylum Cnidaria

Vreni Hussermann & Gnter Frsterra

General Introduction

Class Anthozoa

With approximately 1,200 known species in 46 families

worldwide, sea anemones inhabit all oceans and habitats
from the deep sea to coral reefs, and also estuaries.
Especially in cold and temperate waters they can form an
important part of the macro-zoobenthos. Sea anemones
can range in size from a few mm to 1.25 m diameter. The
largest Chilean species can develop an oral disc diameter
between ten and twentycm but most species, especially
in the fjords, it varies between one and threecm.

External Morphology (Fig. 1)

Sea anemones are benthic sessile polyps without a
skeleton. They are solitary; with the exception of one
colonial species, Cereus herpetodes, which is known
from Chile. The proximal end of a sea anemone is
either rounded, in which case the species is buried in
soft substrate, or forms a more or less well-developed
flat pedal disc, which it uses to attach to hard substrate.
The column is smooth or provided with different

Order Actinaria

Subclass Hexacorallia

Fig. 1

Fig. 1. Schematic drawing of a sea anemone that represents a mesomyarian anemone on the left and an endomyarian anemone on the right (drawing Roman Babuscak, from Hussermann, 2003).

216

structures such as verrucae (see e.g. Parantheopsis

ocellata), tenaculi (see e.g. Phellia exlex), tubercles
(see e.g. Hormathia pectinata), vesicles (see e.g.
Phymactis papillosa), marginal spherules (=acrorhagi,
see e.g. Phymactis papillosa), marginal pseudospherules
(=pseudoacrorhagi, see e.g. Bunodactis spp.), or marginal
projections (see e.g. Isotealia antarctica). It can be subdivided into different regions (scapus and scapulus or
capitulum, e.g. Hormathia pectinata and Metridium
senile lobatum, respectively); and may possess a cuticle
(e.g. Phellia exlex, Hormathia pectinata). The margin
can be separated from the tentacles by a more or less
developed fosse, or be tentaculate (without fosse).
The oral disc (at the distal end) is usually
circular; in some species it can be drawn
out into lobes (e.g. Antholoba achates).
The tentacles are generally simple, hollow,
and usually arranged in hexamerously
alternating cycles. They arise from the
margin and/or the oral disc, and nearly
never possess spherical tips (acrospheres).
Some species possess special fighting
tentacles that can be everted or retracted,
they bear large amounts of special cnidae
used for defence. Colour and markings of
the animals are highly variable.

Fig. 2-7. Histological slides of longitudinal and transverse

sections of Chilean sea anemones.
Fig. 2. Longitudinal section of Anemonia alicemartinae (Family Actiniidae); note endodermal diffuse sphincter (sp) and
basilar muscles (bm); taken from Hussermann & Frsterra
(2001). Legend: ac acrorhagus; co column; me mesenteries;
od oral disc; pd pedal disc; t tentacles.
Fig. 3. Longitudinal section of upper part of column of Phymactis
papillosa (Family Actiniidae); from Hussermann (2004). Legend:
ac acrorhagus; ch channel; lm longitudinal muscle of column; m
mesogloea; v vesicle; sp sphincter (endodermal, diffuse)
Fig. 4. Endodermal, circumscribed sphincter (sp) of Isotealia
antarctica (Family Actiniidae).

Fig. 2
Fig. 3

Fig. 4

217

Key to the higher taxa included in this chapter (adapted after Carlgren, 1949)
Taxa with species from the Chilean Fjord Region in bold

Key to the suborders

Phylum Cnidaria

1) a) Ciliated tract of filaments absent.................................................. Protantheae (1 family; only 1 species in Chile)

b) Ciliated tracts of filaments present or absent. Medially fused mesenteries at proximal end of body.....................
................................................................................................................................... Ptychodacteae (2 families)
c) Ciliated tracts of filaments present. With or without basilar muscles...................................................................2
2) a) Mesenteries, after development of first 12 appearing in lateral endocoels, with their longitudinal muscles oriented
as in directives. No basilar muscles........................................................................ Endocoelantheae (2 families)
b) Mesenteries, after development of first 12 appearing in exocoels, as pairs whose longitudinal muscles face one
another.......................................................................................................................Nynantheae (3 infraorders)

Suborder Ptychodacteae

Class Anthozoa

1) a) Column short, smooth without outgrowth. Numerous tentacles simple and not retractile. 1 weak retractor
muscles per mesentery................................................................................................................Ptychodactiidae
b) Column elongate with vesicular outgrowth histologically identical to tentacles. Tentacles absent or only at
margin of disc. 2 weak retractor muscles per mesentery............................. Preactiidae (only 1 species in Chile)

Suborder Endocoelantheae

Subclass Hexacorallia

1) a) Mesenteries divisible into macro- and micronemes; 6 or 19 pairs of macrocnemes. Retractors of macrocnemes
restricted. 1 siphonoglyph..........................................................................Halcuriidae (only 1 species in Chile)
b) Mesenteries not divisible into macro- and microcnemes. Retractors of mesenteries weak; diffuse. 2

siphonoglyphs...................................................................................................................................Actinernidae

Suborder Nynantheae

Order Actinaria

1) a) Basilar muscles absent.......................................................................................................................................2

b) Basilar muscles present............................................................................................... Thenaria (3 superfamilies)
2) a) Aboral end of body rounded; rarely flattened or disc-like. Column rarely with ectodermal muscles and then
only in uppermost part. Muscles of mesenteries strong. Retractors usually strongly restricted to circumscribed;
parietobasilar muscles usually form well differentiated parietal muscle together with columnar part of
longitudinal mesenterial muscles...................................................................................... Athenaria (10 families)
b) Aboral end of body always disc-like; never rounded. Column with longitudinal muscles. Muscles of mesenteries
weak.......................................................................................................................... Boloceroidaria (2 families)

Infraorder Thenaria
1) a) Families without acontia. Sphincter endodermal or absent; exceptionally meso-endodermal to endo-mesogloeal. Endomyaria
b) Families with mesogloeal sphincter, but without acontia................................................................. Mesomyaria
c) Families with acontia or acontoid organs. Sphincter commonly mesogloeal; sometimes absent........ Acontiaria

Superfamily Acontiaria (13 families)

1) a) Sphincter mesogloeal; strong. Acontia with basitrichs only ........................................................ Hormathiidae

b) Sphincter mesogloeal; usually strong; Acontia with basitrichs and p-amastigophores........................................

...........................................................................................Sagartiidae and Metridiidae (only 1 species in Chile)6

Family Sagartiidae (14 genera)

1) a) Column with somewhat modified but adhesive verrucae ........................................................................Cereus
b) Column smooth. 1st cycle of mesenteries sterile . .............................................................................Anthothoe
c) Column with tenaculi with strong cuticle; forming stout papillae. Mesenteries (and directives) irregularly
arranged. Retractors of older mesenteries very strong; restricted ............................................................ Phellia
226

Superfamily Mesomyaria (4 families)

1) a) Mesenteries divisible into macro- and microcnemes....................................Isanthidae (only 1 species in Chile)7
b) Mesenteries not divisible into macro- and microcnemes. Arrangement of tentacles regular; in cycles. Younger
mesenteries not bilaterally arranged .............................................................................................Actinostolidae

Family Actinostolidae (23 genera)

1) a) Mesenteries distinctly arranged according to the Actinostola rule. Longitudinal muscles of tentacles and radial
muscles of oral disc mesogloeal. Oral disc not lobed.........................................................................Actinostola
b) Mesenteries not arranged according to Actinostola rule. Longitudinal muscles of tentacles and radial muscles of
oral disc ectodermal or mesogloeal . .................................................................................................................2
2) a) All or all stronger mesenteries fertile. Tentacles without nematocyst batteries or b-mastigophores. 12 pairs of
mesenteries perfect. Retractors strong; diffuse. Tentacles rather numerous. Body elongate..................Paranthus
b) The 3 oldest cycles of mesenteries sterile. Oral disc lobed. Sphincter very long ................................ Antholoba

Superfamily Endomyaria (12 families)

Family Actiniidae (55 genera)
1) a) Acrorhagi present in older specimens; in young specimens sometimes absent or very scarce. Sometimes
holotrichs also in other parts of column.............................................................................................................2
b) Acrorhagi absent. Column smooth (apart from marginal projections)................................................................5
c) Acrorhagi absent. Column with verrucae, vesicles or other protuberances........................................................7
2) a) Column smooth; without vesicles or verrucae. Acrorhagi at margin. Tentacles long; imperfectly retractile.
Sphincter not very strong; diffuse to restricted........................................... Anemonia ( North and Central Chile)
b) Column with non-adhesive vesicles, which are simple, compound or branched (more or less weak nematocyst
batteries).............................................................................................................................................................3
c) Column without vesicles, but with adhesive verrucae. With well developed marginal ruff (verrucae at margin
set on small lobes). Sphincter decidedly diffuse.............................Oulactis (North and Central Chile; 2 species)
3) Vesicles distributed over whole or almost whole column; simple or compound. Sphincter well developed;
distinctly diffuse.................................................................................................................................................4
4) a) Acrorhagi with holotrichs and very numerous and long basitrichs.......................................................Phymactis
b) Acrorhagi with holotrichs only.................................................................Phymanthea (North and Central Chile)
5) a) Tentacles with endodermal sphincter at their bases; long; numerous; occupying only outer half of oral disc.
Sphincter diffuse.................................................................................................................................. Bolocera8
b) Tentacles without endodermal sphincters at their base......................................................................................6
6) a) With ring of perforated marginal projections. Body not elongate. Mesenteries of 2 first cycles sterile. Sphincter
circumscribed........................................................................................................................................ Isotealia
b) Without ring of perforated marginal projections. All or all stronger mesenteries fertile. Tentacles very long;
longitudinally ridged with very long basitrichs. Sphincter endodermal; diffuse ............................. Boloceropsis8
7)
Column with verrucae.......................................................................................................................................8
8) a) Body elongate, cylindrical. No distinct sphincter. Most mesenteries perfect. Tentacles short. Verrucae well
developed..................................................................................................................................... Parantheopsis
b) Body as a rule not elongated. All stronger mesenteries, with or without directives, fertile. Outer tentacles shorter
than inner ones. Sphincter more or less curcumscribed to restricted; rarely weak. Verrucae well developed;
sometimes compound in distal part. Marginal pseudoacrorhagi........................................................Bunodactis
6 Metridiidae are described with b-mastigophores and p-amastigophores in the acontia; but examination of the cnidae showed the
presence of basitrichs and p-amastigophores.
7 Another species collected from the fjord region.
8 The cnidome of the examined specimens attributed to the genera Boloceropsis and Bolocera was identical and we found all
transitions from an endodermal sphincters at the tentacle base to no such sphincter. Therefore we cannot separate the species
Boloceropsis platei and Bolocera occidua at the moment.

227

Key to the described anemone species

White threads (acontia) are expelled

Tentacles can be thrown off

0(1)

Anthopleura

Bunodactis hermaphroditica

Bunodactis octoradiata

Parantheopsis ocellata

0/1

Phellia exlex

Hormathia pectinata

Anthothoe chilensis

Acontiaria sp. 2 (brown)

Acontiaria sp. 1 (yellow)

Metridium senile lobatum

Bolocera/Boloceropsis sp.

Actinostola chilensis

Antholoba achates

Paranthus niveus

Paraisanthus fabiani

Halcurias pilatus

0/1

0/1

Isotealia antarctica

Gonactinia prolifera

Dactylanthus antarcticus

Phymanthea pluvia

0(1)

Oulactis concinnata2

1 (0)

1(0)

Oulactis coliumensis3

Anemonia alicemartinae4

0 (1)

Adult with lobed oral disc

Blistered structures on margin, sometimes

hidden in small depression

Entire column with irregularly distributed

lobes

Column with rough, not slimy surface layer,

often covered with detritus, uppermost
column distinct, perfectly clean

Cereus herpetodes

Species

Column +-smooth and +- clean

Phymactis papillosa

Column with clear longitudinal rows of

distinctly coloured, often adhesive spots

Tentacles can be completely covered

Reaction to
annoyance

White or yellow longitudinal stripes

Margin

Uppermost part of column distinct, delicate

Column

Entire column completely covered with

blister-like smooth structures

Class Anthozoa

Phylum Cnidaria

Character

Order Actinaria

Subclass Hexacorallia

hermaphroditica

228

1-4

Common shallow-water species of North and Central Chile, see Hussermann & Frsterra (2001) and Hussermann (2003; 2004b).1: 18

Legend:

0 absent, 1 present, 0(1) generally absent, 1(0) generally present, 0/1 and 1/0 absent or present, num numerous. In columns size and tent

In situ habitus

Brooding species

Fighting tentacles may be present

Present in Peruvian Province (PP)

0/1

23

num.

0/1

23

num.

0/1

0/1

96

<48

1 (2)

3248

12

96

0/1

~100

0/1

(2)3

96

0(1?)

0/1

<100

0/1

2(3)

<100

0/1

<100

0/1

12

num.

1?

<~100

0 (1)

num.

0(1?)

num.

0/1

96

0/1

0 (1)

<~110

0/1

23

<96

23

96

<20

<20

num.

num.

Tentacle number

Pedal laceration

Length of longest tentacles

Transverse fission

Size (OD without tentacles)

In habitat elongate

0
(0)1

Longitudinal fission

Lower colum hidden in sand

Other
features

Entire body visible in situ

General morphology

6 or 12 tentacles closer to mouth

Mode of asexual
reproduction

Several mouth openings on oral disc

Oral disc

0/1

96

<100

8S33S; 2:18S40S; 3: 37S; 4: 18S37S.

tacle number: 1 small/short, 2 medium, 3 large/long.

229

Halcurias pilatus

233

Halcurias pilatus McMurrich, 1893

Subclass Hexacorallia

Class Anthozoa

Phylum Cnidaria

Common name: Orange decameric anemone;

Anmona decamera naranja
Description: External morphology: PD to ~1.5 cm ;
OD to ~1.3cm. Colour generally orange; more rarely
white; base of tentacles white; preserved specimens
whitish. Elongated anemone. Column smooth to slightly
corrugated; covered with white spots (cnida batteries, see
Fig.) in longitudinal lines; in some individuals uppermost
part seems more delicate. Up to 96 long tentacles; outer
considerably shorter than inner; can be fully covered
by column. Contracted shape round to oblong. Internal
morphology: Actinopharynx short; with 1 siphonoglyph.
2 pairs of directives. 10 pairs of mesenteries; all perfect.
Oral stoma. Histology: Sphincter mesogloeal; weak.
Retractors diffuse; restricted to inner half of mesenteries.
All mesenteries fertile. Cnidae: Spirocysts; basitrichs;
p-amastigophores; holotrichs. Possibility for confusion:

In vivo: Other orange, pink, or white anemones.

However, H. pilatus has a long column with white spots
and no acontia. Preserved: Other elongated anemones
(e.g. Paranthus niveus), but the decamerous mesenterial
arrangement and the holotrich batteries (3045 m) on
the column (see Fig.) distinguish H. pilatus from all other
anemones from this region.
Habitat: Rocky substratum. Column regularly hidden
in crevices or holes, dead barnacle and coral skeletons.
Depth: 5821m. Abundance: In channels locally very
abundant; more rarely in fjords. Generally aggregated.
Distribution: SE Pacific (NPZCPZ). Chile: 42S50S.
Biology: Globular reproductive products observed
in the tentacles. Sexes separate, no signs of asexual
reproduction.
Main reference: McMurrich (1893).

Hormathia pectinata (Hertwig, 1882)

Order Actinaria

Common name: Tubercular anemone;

Anmona de tubrculos
Synonymy: Phellia spinifera Hertwig, 1888 (at least pro
parte).
Description: External morphology: To 5cm high; PD
to ~4cm. Colour white to light peach-coloured or light
orange; actinopharynx brown to brown-red; preserved
specimens whitish to grey. Column tough; divided into
scapus and scapulus. Scapus with brown cuticle that can
be at least partially lost. Scapulus with tubercles with
small brown, chitinuous cuticle hats; with longitudinal
ridges ending in 12 larger coronal tubercles. 96 tentacles;
long; thin; can be fully covered by column. Mesenterial
insertions visible on OD. Mouth opening prominent; lips
thick. No cinclides on column. Internal morphology:
Actinopharynx deeply furrowed. 2 siphonoglyphs. 2
pairs of directives. Oral stoma. Mesogloea thick. Acontia
present. Histology: 48 pairs of mesenteries: 1st cycle perfect
and sterile; 2nd and 3rd cycle fertile. Parietobasilar muscles
strong. Sphincter mesogloeal; strong. Cnidae: Spirocysts;
basitrichs; p-mastigophores (Fig. 8I Introduction);
p-amastigophores. Acontia with basitrichs only (25
234

29 m; 5464 m; see Fig.). Possibility for confusion:

On first sight other larger Chilean anemones, but due to
the presence of acontia and a cuticle this species is easily
recognizable. Phellia exlex also has acontia and a cuticle
but is smaller, much darker, appears in groups and has
additional p-mastigophores in the acontia.
Habitat: Steep to overhanging rocky walls. Depth: 12
1,220m. Abundance: Infrequent in shallow water; most
individuals below 30m; never in groups. Distribution:
SW Atlantic (Argentina); SE Pacific (NPZCPZ); Strait
of Magellan. Chile: 32S33S?; 42S53S. Biology:
Specimens retract very fast due to light or vibration
stimuli. Comments: Typical deep-water emerging species.
Chilean shallow water specimens are considered dwarf
form. This form includes 2 morphs; some individuals
have a very firm PD, while in others it is delicate.
Morphological characters of hormathiids vary strongly,
even within species, and thus species are often difficult
to tell apart. Argentinean specimens often epibiotic.
Possibly also off Central Chile (32S33S, 350400m).
Main references: Hertwig (1882); McMurrich (1893);
Riemann-Zrneck (1973).

Hormathia pectinata

235

Phylum Cnidaria

Glossary
Acontium (pl.

Thin threads attached at one end to mesenteries, as a rule below the filaments, while the other

acontia)

end is free. They are laden with extraordinarily numerous nematocysts of variable categories.

Acrorhagus (pl.

Endocoelic structure outside the tentacle crown and adoral to the marginal sphincter

acrorhagi)

(further away from the mouth) in which the epidermis contains holotrichous nematocysts
and epitheliomuscular cells that are vacuolated at their bases (see Fig. 2&3). Called marginal
spherules by Carlgren (1949).

Acrospheres

The globular ends of certain tentacles, laden with numerous nematocysts. They are of
differentcategories in the Corallimorpharia, in the Actiniaria and seemingly in the Madreporaria

Class Anthozoa

also.
Actinopharynx (syn.

Throat, stomodaeum; the tube which leads from the mouth into the coelenteron; formed by

pharynx)

the epidermis.

Basilar muscles

Radial muscles running along both sides of the base of the mesentery, close to the pedal disc.
Present in most of the Actiniaria (see Fig. 2).

Basitrichs

Type of nematocyst. No V-shaped notch at base of unfired shaft; tubule with long proximal and
short distal spines. Rod-like basitrichs are very long and slender.

b-mastigophores

Type of nematocyst. No V-shaped notch at base of unfired shaft; discharged shaft or proximal
tube approximately same diameter as remaining tubule; proximal tubule with prominent
armature.

Capitulum

See Scapus.

Cinclides

Small apertures (or organized soft spots which will rupture readily) in the column through

Subclass Hexacorallia

which acontia can be expelled. Also present in some species without acontia.
Circumscribed

Shape of a muscle. When the muscles are very strongly concentrated and there is only one
mesogloeal lamella (or a few main lamellae close to each other) originating from the main
lamella of the mesentery (see Fig. 4).

Cnidae (see Fig. 8)

Spirocysts (only Anthozoa), nematocysts (all Cnidaria), and ptychocysts (only Ceriantharia).

Cnidom

List of types of cnidae of a species.

Column

Body-wall. Its musculature is usually entirely endodermal, at least in origin. The fibres of the
muscle-sheets on the endodermal face of the mesogloea are placed horizontally, and constitute
the endodermal circular muscle of the column. The column can possess a variety of structures
such as verrucae, vesicles, tubercles and tenaculi.

Diffuse

Shape of a muscle. Muscle with relatively short processes which are based upon a considerable

Order Actinaria

area of the mesoglea (see Fig. 3).

Directives

Pair of mesenteries situated in the directive axis; with their longitudinal muscles (retractors) on

(syn. directive

their outer sides turned towards the exocoels (see Fig. 6B).

mesenteries)
Distal

The upper end of a sea anemone where the oral disc is situated.

Ectoderm (syn.

External body layer; composed of cells (see Fig. 6C).

epidermis)
Endocoel

The space between 2 mesenteries belonging to one and the same pair.

Endoderm (syn.

Internal body layer; lining the body cavity and tentacles; composed of cells (see Fig. 6C).

gastrodermis)
Exocoel

The space between mesenteries belonging to different pairs.

Fighting tentacle

Specialized tentacle with different cnidome compared to regular tentacles; often longer and

(syn. catch tentacle)

in some cases with different colouration; it is inducible and involved in agonistic behaviour
(competition and defense).

262