Polar Biol (2005) 28: 338–350
DOI 10.1007/s00300-004-0712-3
E RR A TU M
Verena Häussermann
The sea anemone genus Actinostola Verrill 1883: variability
and utility of traditional taxonomic features, and a re-description
of Actinostola chilensis McMurrich 1904
Published online: 16 February 2005
Springer-Verlag 2005
Abstract Species of the genus Actinostola are known for
high variability of features. Anatomy, histology and
cnidae of type specimens of five species from South
America and Antarctica originally described as members
of Actinostola and one species of Stomphia were compared to specimens of Actinostola chilensis collected
during this study. None of these traditionally used features clearly distinguish the examined Actinostola species. I therefore propose new distinctive taxonomic
features, including in vivo and in situ data. I provide an
emended diagnosis of the genus Actinostola and a
revised list of its species. I accept the synonymy of
A. excelsa, A. pergamentacea and A. intermedia with
A. crassicornis, and reject the synonymy of A. chilensis
with A. crassicornis and A. intermedia. I re-describe
A. chilensis in detail, including in situ information.
Specimens of A. chilensis inhabit exposed positions of
Electronic Supplementary Material Supplementary material is
available in the online version of this article at http://dx.doi.org/
10.1007/s00300-004-0712-3
The publisher regrets that during copy-editing of this article the
names of the authors of species were styled incorrectly. Therefore,
it was decided to publish this erratum which, due to the nature of
the paper, contains the whole original article, but now with the
names of species authors styled correctly.
The online version of the original article can be found at http://
dx.doi.org/10.1007/s00300-004-0637-x
V. Häussermann
Huinay Scientific Field Station, Chile
V. Häussermann (&)
Department Biologie II,
Ludwig-Maximilians-Universität München,
Karlstr. 23–25, 80333 München, Germany
E-mail: vreni_haeussermann@yahoo.de
URL: http://www.people.freenet.de/haeussermann
Present address: V. Häussermann
Departamento de Biologı́a Marina,
Universidad Austral de Chile, Avda. Inés de Haverbeck, casas 9,
11 y 13, Campus Isla Teja, Casilla 567, Valdivia, Chile
rocky substrate from 22 m depth down in south Chilean
fjords between Puerto Montt (4135¢35¢¢S, 7253¢W) and
Puyuhuapi (4431¢36¢¢S; 7232¢6¢¢W); the most conspicuous features are its relatively large size, bright-orange
colour, smooth, tough column and numerous and
clearly entacmaeic tentacles.
Introduction
The family Actinostolidae, with its approximately 20
genera, constitutes 1 of the 2 richest families of deepsea anemones (Fautin and Barber 1999). Due to the
depths at which most of its members are collected,
most species of Actinostola are collected by dredges and
bottom grabs and are known primarily from fixed
material (Fautin and Hessler 1989). Sampled specimens
are often damaged and poorly preserved. Publications
on Actinostolidae are generally scarce and widely
scattered in the older literature (e.g. Hertwig 1882;
Carlgren 1893, 1927, 1928; McMurrich 1893, 1904);
more recently, species have been described from deepsea hydrothermal and cold vents (Fautin and Hessler
1989; Fautin and Barber 1999). Although some species
of this family are known to extend to relatively shallow
waters, e.g. along the Patagonian coast of Chile and
Argentina, the northwest coast of North America and
in the Antarctic (McMurrich 1904; Carlgren 1959;
Riemann-Zürneck 1978; Fautin 1984), only two studies
describing these sea anemones alive or in their habitat
have been published (Ross and Sutton 1967; Ross and
Zamponi 1995).
The type genus Actinostola is especially rich in species, with most of its members known from polar and
subpolar regions. Species belonging to Actinostola are
extremely variable in many of the features that are traditionally used as specific characteristics (Carlgren 1893,
1921; Riemann-Zürneck 1971, 1978). Riemann-Zürneck
(1971) concluded that identifications based exclusively
on these features, and therefore the status of most
339
species, have to be treated as highly uncertain; she discerned an urgent need for new distinctive characteristics.
However, the features she suggested, such as shape of
preserved specimens and cnidae of unilobulate filaments,
have not been adopted in subsequent studies and nor
have alternative suggestions been presented (e.g. Doumenc 1984; Fautin 1984).
In the present paper, I compare type specimens of
South American and Antarctic species of Actinostola,
emend the diagnosis of the genus, and generate an up-todate list of its members (Appendix 1). I examine the
usefulness of traditionally used morphological features,
and propose new distinctive characteristics. I re-describe
Actinostola chilensis from fjords in southern Chile in
detail; this constitutes the first documentation of in vivo
and in situ information about an identified species of
Actinostola in the scientific literature.
Materials and methods
Between 1994 and 2004, Günter Försterra and I
observed and examined several tens of specimens of
A. chilensis along the Chilean coast from Lenca
(4135¢37.0¢¢S; 7242¢10.9¢¢W) to Puerto Chacabuco
(4527¢S; 7248¢W) (Fig. 1; Appendix 2) and preserved
15. Study sites are listed in Appendix 3. We studied
specimens in situ by means of scuba-diving. Some
specimens were kept in aquaria for several days for detailed examinations; photographs were taken both in
situ and in aquaria. For preservation, specimens were
relaxed with menthol crystals for 45–180 min and fixed
in 10–15% seawater formalin. Specimens were kept in
formalin for at least 4 months, before being transferred
to 70% alcohol. Parts of some specimens were preserved
in 96% ethanol for future molecular studies. For the
histological examinations, parts of seven specimens were
embedded in paraffin, sectioned at 8 and 9 lm, and
stained with Azocarmin triple staining (Humason 1967).
I examined cnidae from three living and three
preserved specimens with a light microscope (·1,000 oil
immersion); these were drawn or photographed and
measured. The discharge of fresh cnidae was provoked
with distilled water or 4% acetic acid solution. Semipermanent slides of discharged cnidae were prepared
using the technique of Yanagi (1999): a small amount
of tissue is put into a drop of 4% acetic acid or HCl
solution on a microscopic slide. After 2 or 3 min, the
liquid is drawn off carefully with a tissue. The tissue is
then suspended in a solution of 1:1 seawater:glycerin
that contains a few drops of phenol and formalin per
100 ml. The coverslip is applied and sealed several
times with nail coating. To test the value of cnidae as
distinctive characters (e.g. Fautin 1984), and especially
the value of cnidae of the unilobulate filaments (e.g.
Riemann-Zürneck 1978), I examined and compared the
cnidae of the type material of A. crassicornis, A. excelsa, A. pergamentacea, A. chilensis and A. intermedia
with the cnidae of a specimen of A. chilensis collected
Fig. 1 Type localities and distribution of Actinostola species
around southern South America [Actinostola chilensis A.ch.,
A. crassicornis A.cr., A. excelsa A.ex., A. intermedia A.in.,
A. pergamentacea A.pe.; triangles type localities, broken lines
distribution of A. chilensis (this paper) and A. crassicornis (sensu
Riemann-Zürneck 1978]
in the Chilean fjords (called A. chilensis Coll in Table 1), using type material of Stomphia selaginella as an
out-group. I compared cnidae types and mean size
values taking into account the standard deviation. Size
ranges of cnidae are values taken from single specimens
(Table 1). Nematocyst terminology follows that of
England (1991).
Specimens examined
Chile
Histological slides of transverse and longitudinal sections were deposited at the Zoologische Staatssammlung
340
Table 1 Size and distribution of cnidae from the type material of
A. crassicornis, A. excelsa, A. pergamentacea, A. intermedia,
A. chilensis and Stomphia selaginella compared with A. chilensis
collected in Chile (called A. chilensis Coll here; for cnidae see
Fig. 5) [r rare (less than 10 capsules found in 1 h search), f few (10–
30 capsules found in 1 h search), c common (30–50 capsules found
in 1 h search), v very common (>50 capsules found in 1 h search).
Tissue/cnidae type,
abundance
Tentacles
Spirocysts
A. crassicornisv
A. excelsav
A. pergamentaceaf
A. intermediav
A. chilensisc
A. chilensis Collv (B)
S. selaginellav
Basitrichs
A. crassicornisc
A. excelsac
A. pergamentaceac–v
A. intermediac
A. chilensisf–c
A. chilensisCollc(C)
S. selaginellav
Microbasic amastigophores
A. crassicornisf
A. excelsac
A. pergamentaceaf
A. intermediaf
A. chilensiss
A. chilensis Collr (D)
S. selaginella
Large b-mastigophores
A. crassicornisf
A. excelsaf–c
A. pergamentacea*
A. intermedias
A. chilensiss
A. chilensis Collr (A)
S. selaginellas
Column
Basitrichs
A. crassicornisv
A. excelsaf
A. pergamentaceaf
A. intermediac
A. chilensisf
A. chilensis Collc–v (E)
S. selaginellaf
Pharynx
Basitrichs
A. crassicornisf
A. excelsav
A. pergamentaceac
A. intermediac
A. chilensisf
A. chilensis Collc (F)
S. selaginella type 1f
S. selaginella type 2c
Microbasic amastigophores
A. crassicorniss
A. excelsac
A. pergamentaceaf
A. intermediac
A. chilensisf
A. chilensis Collc (G)
S. selaginellas
‘‘ml’’ and ‘‘mw’’ are the means, ‘‘dl’’ and ‘‘dw’’ are the standard
deviations (all in lm), ‘‘t’’ are the number of turns on the proximal
part of the tube, ‘‘p’’ is the proportion of animals examined with
respective type of cnidae present. No is the number of capsules
measured. Exceptional sizes in parentheses]. Note that three
b-mastigophores 49.5–50.4·5.4–6.3 lm were found in the filaments
of S. selaginella
Capsule length
(lm)
ml
dl
Capsule
width (lm)
mw
dw
18.9–52.2
27.0–54.0
18.0–46.8
(13.5) 22.3–67.5
18.9–52.2
23.4–62.1
24.3–51.3
35.6
38.8
33.2
42.8
37.2
42.5
40.0
9.13
6.46
7.29
12.8
9.19
9.44
6.90
1.8–5.4
2.7–5.4
2.7–5.4
1.8–5.4
2.7–5.4
2.7–5.4
2.7–4.5 (7.2)
3.7
3.9
3.9
3.9
3.8
4.1
3.9
0.77
0.78
0.84
0.86
0.82
0.86
0.84
23.4–32.4
23.4–33.3
18.9–32.4
(10.8) 24.3–34.2
24.3–32.4
19.8–33.3 (36.9)
25.2–31.5
A
18.0–24.3
20.7–27.0
18.9–24.3
16.2–24.3
20.7–21.6
19.8–22.5
–
28.8
28.0
23.3
29.1
28.3
27.6
28.4
1.84
1.92
3.09
3.91
2.07
3.01
1.35
2.7–4.5
1.8–3.7
1.8–3.6
(1.35) 1.8–2.7
1.8–3.6
1.8–3.6
1.8–2.7
2.8
2.5
2.7
2.3
2.8
2.7
2.3
0.38
0.44
0.24
0.43
0.33
0.38
0.37
21.8
23.6
21.9
21.0
1.42
1.72
1.28
2.22
4.2
3.8
4.5
4.2
0.56
0.42
0.39
0.55
21.0
0.86
3.6–5.4
3.2–4.5
3.6–5.4
3.6–5.4
4.5
3.6–4.5
–
4.4
0.34
47.0
45.3
3.46
3.25
4.5–6.8
5.4–8.1
–
6.3–7.2
6.3–7.2
4.0–7.0
4.5–5.9
5.9
6.5
0.55
0.60
40.5–56.7
36.0–52.2
–
49.5–51.3
46.8–49.5
43.0–47.0
49.5–54.9
18.9–23.4
18.0–22.5
17.1–21.6
19.8–27.9
18.0–22.5
14.4–21.6
18.0–22.5
(33.3)
(28.8)
(27.9)
(29.7)
(19.8) 22.5–30.6
18.9–27.9
(18.0) 22.5–28.8
(18.9) 24.3–31.5
18.0–27.0 (28.8)
(16.2) 22.5–28.8
17.1–24.3
28.8–36.0
A
18.9–23.4
19.8–25.2 (27.0)
20.7–24.3
19.8–25.2
18.0–24.3
17.1–24.3
22.5–25.2
t
p
6/6
7
?
6/6
5/6
4/6
21.4
20.0
19.6
23.3
20.4
18.3
20.9
2.86
1.66
1.17
1.81
1.90
1.54
2.26
1.8–2.7
1.8–2.7
2.3–2.7
1.8–2.7
2.7–3.6
2.3–3.6
1.8–3.6
2.5
2.5
2.6
2.4
2.7
2.9
2.6
0.40
0.28
0.21
0.39
0.15
0.42
0.29
26.1
23.8
24.5
28.7
25.3
26.9
20.3
31.7
2.68
2.29
2.28
2.36
2.60
2.49
1.91
1.86
1.8–3.6
1.8–2.7
1.8–2.7
1.8–2.7
2.3–3.6
2.3–3.6
1.8–2.7
1.8–2.7
2.7
2.6
2.5
2.5
2.9
2.8
2.3
2.6
0.32
0.26
0.29
0.37
0.38
0.26
0.34
0.21
21.7
22.4
22.4
22.7
21.2
21.7
24.5
1.83
1.59
0.95
1.57
1.98
1.46
0.93
4.5–5.4
3.6–5.4
3.6–5.4
4.5–5.4
4.5–6.3
4.1–5.4
3.6–4.5
4.8
4.6
4.2
5.0
5.5
4.8
4.2
0.33
0.54
0.51
0.46
0.59
0.39
0.40
5–6
5–7
7
6/6
6/6
6/6
No
40
41
45
39
42
43
41
40
41
53
72
69
66
40
25
40
29
14
3
7
0
25
41
0
3
3
6
5
23
43
45
40
35
45
44
43
52
46
43
48
42
41
42
8
40
25
33
22
38
8
341
Table 1 (Contd.)
Tissue/cnidae type,
abundance
Mesenterial filaments
Basitrichs
A. crassicornisc
A. excelsaf
A. pergamentaceac
A. intermediaf
A. chilensisf
A. chilensis Collf (J)
S. selaginellaf
Microbasic amastigophores
A. crassicornisf
A. excelsac
A. pergamentaceav
A. intermediac
A. chilensisv
A. chilensis Collc (K)
S. selaginellac
p-mastigophores
S. selaginella – type1c
S. selaginella – type2c
Pedal disc
Basitrichs
A. crassicornisf
A. intermediac
A. chilensisc
A. chilensis Collc (H)
Spirocysts
A. crassicornis
A. intermediar
A. chilensisf
A. chilensis Collf (I)
Capsule length
(lm)
ml
dl
Capsule
width (lm)
mw
dw
t
p
No
18.9–33.3
18.0–33.3
18.9–34.2
18.9–36.0
17.1–26.1
18.0–27.0 (35.0)
16.2–21.6
A
19.8–24.3
20.7–28.8
20.7–26.1
18.9–25.2
18.0–24.3
18.9–22.5
18.9–24.3
24.7
26.3
26.1
23.2
21.3
22.8
18.3
3.93
4.04
3.74
4.26
2.70
2.25
1.42
2.3–3.2
1.8–3.2
1.8–2.7
1.8–3.6
1.8–3.6
2.3–3.6
1.4–2.7
2.6
2.4
2.3
2.6
2.7
3.0
1.8
0.25
0.43
0.37
0.59
0.48
0.38
0.31
22.6
25.2
23.6
23.0
20.9
20.7
21.8
1.33
1.61
1.22
1.38
1.71
1.09
1.31
3.6–6.3
3.6–5.4
3.6–5.4
4.5–6.3
4.5–6.3
4.1–5.4
3.2–5.4
5.0
4.4
4.5
5.1
5.0
4.7
4.0
0.60
0.44
0.36
0.59
0.49
0.33
0.54
(36.9) 48.6–61.2
78.3–88.2
52.5
82.4
3.65
3.16
3.6–5.0
5.9–8.1
4.2
6.5
0.44
0.46
41
33
18.0–23.4 (27.0)
17.1–24.3 (30.6)
17.1–20.7
17.1–20.7
20.8
23.1
18.9
19.4
1.57
2.46
0.90
1.08
1.8–2.7
1.8–3.2
1.8–2.7
1.8–2.7
2.5
2.2
2.4
2.5
0.30
0.42
0.33
0.31
43
51
41
42
–
26.1–54.0
25.2–45.0
29.7–54.0
–
38.0
35.2
45.2
–
8.43
5.69
8.75
–
2.7–4.5
2.7–5.4
2.7–6.3
–
3.8
4.1
4.8
–
0.75
0.99
1.14
4–6
7
5–7
6/6
6/6
3/3
61
59
68
51
16
41
39
38
40
41
40
43
41
41
0
12
20
11
*A. pergamentacea: tentacles very badly preserved and decomposed
Munich (ZSM), at the Museum für Naturkunde of the
Humboldt-Universität zu Berlin (ZMB Cni 14227), as
well as at the Naturhistoriska Riksmuseet Stockholm
(SMNH-56586–56588 and SMNH-56595). A. chilensis
[all collected by G. Försterra (GF) and V. Häussermann
(VH)]; Punta Chaica, Seno Reloncavı́ (S53), 24 January
2000, 25 m (Ex. 284=ZSM 20030420); 22/24 January
2001, 22–30 m (Ex. 1, 2, 4, 5=ZSM 20030421); Punta
Llonco, fjord Comau (S60a), 11 April 2003, 28 m (ZSM
20030422); Caleta Gonzalo, fjord Reñihue (S61), 16
February 1998, 27 m (Ex. 253, 254=ZSM 20030423); 20
January 2000, 33 m (Ex. 267=ZSM 20030424); 24
March 2001, 25–33 m (Ex. 281=ZSM 20030425); 7
February 2001, 35 m (Ex. 230, 231=ZSM 20030426);
Caleta Gonzalo (S63), 19 January 2000, 25–30 m (Ex.
259=ZSM 20030427); S of Puyuhuapi (S90) 10 January
2000, 22–30 m (Ex. 233, 234=ZSM 20030428).
Examined type material (for localities see Appendix
4): A. chilensis McMurrich 1904, hermaphroditic (histological sections prepared); Calbuco, Chile, 4145¢S,
6513¢W (coordinates from Microsoft Encarta 2002) 29–
37 m, (holotype ZMB Cni 4204); A. intermedia Carlgren
1899, male (histological sections prepared), Cabo San
Vicente, Tierra del Fuego, Argentina, SW Atlantic,
274 m, 5437¢S; 6513¢W (coordinates from Microsoft
Encarta, 2002) (holotype SMNH-1184); A. crassicornis
(Hertwig 1882), fertile, SW Atlantic, 5220¢S, 680¢W,
101 m (station 313), and 5338¢S, 7056¢W, 18–27 m
(station 312) (paratypes SMNH-1183 and British Museum of Natural History 1889.11.25.3–4, 9 and 10);
A. excelsa McMurrich 1893, fertile, SW Atlantic,
4837¢S, 6546¢W and 5134¢S, 680¢W, 92–106 m (National Museum of Natural History, syntypes US
NMNH-17780); A. pergamentacea McMurrich 1893,
fertile, SW Atlantic, 4522¢S, 6420¢W, 94 m (station
2769) (syntypes US NMNH-17779); A. georgiana Carlgren 1927, fertile, Antarctic, 5429.3¢S, 343.9¢W, 567 m
(syntype SMNH-4015); A. clubbi Carlgren 1927, fertile,
Oates Land, Antarctic, 6721¢46¢¢S, 15521¢10¢¢W, 464 m
(holotype SMNH-4009); S. (Cymbactis) selaginella
(Stephenson 1918), one fertile, histological sections
available (1918.8.16.8), Ross Sea, Antarctic (BMNH
syntype 1918.5.12.15, and 1918.5.12.31–33). For original
drawings, see Fautin (2003).
Sampling sites where I found but did not collect
A. chilensis (see Fig. 1; Appendix 2; for a detailed
description of sites see Appendix 3): S53: 4138¢15,5¢¢S;
7240,8,3¢¢W; S57: 4140.353¢S, 7239.399¢W; S60c:
4209¢36¢¢S;
7226¢06¢¢W;
S60d:
4219¢40¢¢S;
7227¢04¢¢W; S60a: 4220¢28¢¢S; 7226¢54¢¢W; S60f:
4223¢15¢¢S;
7227¢38¢¢W;
S61:
4232¢46,6¢¢S;
7237¢0,2¢¢W; S 62: 4233¢S, 7236¢W; S63: 4233¢12,7¢¢S;
342
7235¢22,3¢¢W; S65: 4233,494¢S; 7236,271¢W; S83:
4347¢09,1¢¢S,
7255¢34,2¢¢W;
S85:
4358¢18,4¢¢S,
7307¢00,6¢¢W; S90: 4431,608¢S; 7232,107¢W; S96:
4526¢47,9¢¢S; 7249¢25,8¢¢W (identification uncertain).
Results
Family: Actinostolidae Carlgren 1932; Genus:
Actinostola Verrill 1883
Diagnosis after Carlgren (1949), with changes in bold:
Actinostolidae with body sometimes short, sometimes
cup-like, sometimes long, cylindrical. Column usually
thick, firm, slightly rugose to smooth, or with flat
tubercles produced by mesogloeal thickenings. Sphincter
mesogloeal; upper part of column can completely cover
tentacles. Tentacles short to medium-sized, inner considerably longer than outer; sometimes with mesogloeal
thickenings on aboral sides at the base; outside at tips
may be provided with microbasic b-mastigophores.
Longitudinal muscles of tentacles mesogloeal; radial
muscles of oral disc endodermal to mesogloeal. Two welldeveloped siphonoglyphs each connected to a pair of
directive mesenteries. Mesenteries hexamerously arranged. The two mesenteries in one and the same pair,
from third or fourth cycle, irregularly arranged, but as a
rule orientated so that the mesentery that turns its longitudinal muscle towards the nearest mesentery of the
preceding cycle is more developed than its partner.
Retractors of mesenteries diffuse, parietobasilar and
basilar muscles strong. Mesenteries of two first cycles
sterile.
Cnidae: spirocysts (in tentacles, may be found in pedal
disc), basitrichs (in all tissues), microbasic b-mastigophores (may be found in tentacles or rarely in filaments),
microbasic amastigophores A (in tentacles, pharynx and
filaments). Type species: Actinostola (Urticina) callosa
(Verrill 1882)
Re-description of Actinostola chilensis McMurrich 1904
Locality in parentheses if new material was collected.
A. chilensis
A. chilensis McMurrich
Non A. chilensis McMurrich
Non A. intermedia
A. intermedia Carlgren
? A. intermedia Carlgren
Non A. intermedia Carlgren
Non Catadiomene intermedia Carlgren
? A. callosa Verrill
Line missing
External anatomy
Differential diagnosis
Bright-orange, medium to large-sized with pedal-disc
diameter up to 80 mm, contracted animals shaped like
cylindrical stump of cone, with crater-like hole at apex.
Column smooth, without distinct fosse; up to more than
200 tentacles, outer considerably longer than inner,
mouth opening prominent (Fig. 2). Preserved specimens
white to slightly beige (Fig. 3). Hermaphroditic or with
separate sexes. Cnidae and internal anatomy similar
to that of other species of Actinostola. Specimens
of A. crassicornis from Argentina and specimens of
A. georgiana from Antarctica can be distinguished from
A. chilensis by the common presence of embryos in the
gastrocoel in A. crassicornis and A. georgiana.
Size
In life, pedal-disc diameter (to 80 mm); column diameter
(to 50 mm); column height (to 100 mm); oral-disc
diameter (to 85 mm); longest tentacles 40 mm; (preserved) pedal-disc diameter (to 65 mm); column diameter (to 60 mm); column height to 45 mm; oral-disc
diameter to 65 mm; longest tentacles 37 mm. Pedal-disc
diameter of most preserved specimens between 40 and
60 mm, much smaller individuals very rare.
Oral disc and tentacles
Oral disc round, insertions of mesenteries visible. Mouth
opening central, distinctly prominent, round to triangular, lips thick. Pharynx deeply furrowed, with two
distinct siphonoglyphs ending in distinct grooves. Up to
more than 200 tentacles on outer half of oral disc,
hexamerously arranged in up to seven cycles, last cycle
may be incomplete. Tentacles medium-sized to long,
longer than radius of oral disc, strongly entacmaeic,
outer tentacles very short, conical, each with slightly
rounded tip and with distal cinclis. Tentacles in preserved state short to medium-sized (Fig. 3), without
mesogloeal thickening at base.
McMurrich 1904, p 247 (Calbuco, Chile)
Stephenson 1920, p 557; Carlgren 1949, p 78
Clubb 1908, p 4 (Antarctica); Pax 1926
(Ross Sea, Antarctica); Fautin 1984, p 14 (Antarctica)
Carlgren 1899, p 31 (Tierra del Fuego, Argentina, Atlantic)
Carlgren 1959, p 29 (Seno Reloncavi and Golfo de Ancud, Chile);
Sebens and Paine 1979, p 230
Doumenc 1984, p 150 (Los Vilos to Coquimbo, northern Chile)
Carlgren 1927, p 58; Carlgren 1949, p 78; Riemann-Zürneck 1971,
p 161; Riemann-Zürneck 1978, p 66; Fautin 1984, p 14 (Antarctica)
Stephenson 1920, p 558
McMurrich 1893, p 167 (between Ecuador and Galapagos Islands)
343
Fig. 2a–d Specimens of Actinostola chilensis in situ. a Lateral view
column of two specimens; note posture of tentacles; fjord Reñihue,
32 m. b Oral view oral disc and tentacles; note insertions of
mesenteries; fjord Comau, 25 m. c Group of specimens in a
‘‘meadow’’ of Primnoella sp.; Seno Reloncavı́, 28 m. d Lateral view
completely contracted specimen; note spots with missing ectodermal tissue; Seno Reloncavı́, 28 m; real size
Column
As broad as high or higher than broad; proximally
slightly and distally strongly trumpet-like when expanded. Generally smooth, thick and firm; slightly tuberculate in large animals. Insertions of mesenteries
visible as fine longitudinal lines. Margin tentaculate, no
distinct fosse.
Pedal disc
Well developed, round, relatively thin. Limbus smooth.
Insertions of mesenteries visible.
Colouration
Fig. 3a,b Preserved specimens of Actinostola chilensis. a Lateral
view, b Oral view
Oral disc orange, slightly darker around the pharynx,
lips and pharynx light-orange, separated from oral disc
by a clear line; insertions of mesenteries visible as lighter
lines. Tentacles orange, column bright-orange, rarely
344
reddish-orange, in some large animals with white patches due to missing ectodermal tissue; most proximal
region in some animals lighter, in others limbus slightly
darker than rest of column.
Variability
Colour and appearance in situ of specimens in the
Chilean fjord region very uniform (Fig. 2), bright-orange; some specimens with white spots on column due to
scraped-off ectodermal tissue (Fig. 2d). Carlgren (1959)
described his specimens as pink, salmon-coloured to
bright-orange.
Internal anatomy
General
Mesenteries hexamerously arranged in up to seven cycles, first two (in one small individual) or three cycles
perfect. Mesenteries numerous and thin, more than 200.
Mesenteries from fourth cycle onward arranged
according to the Actinostola rule (Fig. 4g,h). Oral stoma
present in most perfect mesenteries; marginal stoma may
be present in larger perfect mesenteries. Actinopharynx
approximately half length of column, with deep longitudinal furrows proximally. Pharynx with two very
broad, proximally strongly prolonged siphonoglyphs
which nearly reach pedal disc and roll up at end. Two
pairs of short directives, connected to the siphonoglyphs
(Fig. 4f). Four of 15 specimens fertile, 2 male and 2 female. Because of the hermaphroditism of the holotype
(Fig. 4h,i), the species has to be defined as ‘‘hermaphroditic or with separate sexes’’. Diameter of eggs 225–
480 lm (in type of A. chilensis and ZSM 20030421)
(Fig. 4g–i) 50–110 lm respectively (ZSM 20030420).
Fourth to sixth cycle, rarely third cycle fertile, youngest
cycles sterile; on younger cycles often only stronger
mesentery of a pair fertile. Two oldest cycles of mesenteries and directives sterile. No evidence of asexual
reproduction.
Musculature
Sphincter (Fig. 4a–d) mesogloeal, long, reticulated or in
layers, in marginal region approximately 1/3–2/3 (in
some specimens up to 100%) breadth of mesogloea, either of constant breadth along the column (Fig. 4a,c) or
strongly tapering proximally (Fig. 4b,d). Circular musculature of body wall hardly visible, endodermal to
mesogloeal, arranged in layers. Longitudinal muscles of
tentacles strong, mesogloeal (Fig. 4e). Circular muscles
of oral disc endodermal to endo-mesogloeal. Mesenterial
retractors diffuse, thin, of equal breadth along mesentery
(Fig. 4f–h). Basilar muscles and parietobasilar muscles
strong; latter forms distinct fold in proximal-most region
of mesenteries (Fig. 4g)
c
Fig. 4a–i Actinostola chilensis. a,b Longitudinal sections through
marginal region. c,d Details of sections through mesogloeal
marginal sphincter. e Cross section through tentacle. f,g Cross
section through mesenteries at level of stomadaeum. h,i Type
material of A. chilensis: h cross section through mesenteries at
stomadaeum level, i ova and sperm (directives di, ectoderm ec, pair
of imperfect mesenteries im, mesogloea m, mesenterial filaments mf,
mesogloeal longitudinal muscle of tentacle mt, ova o, parietobasilar
muscle pb, lumen of actinopharynx ph, pair of perfect mesenteries
pm, retractor muscle r, reticulated pad rp, sperm s, siphonoglyph si,
mesogloeal sphincter sp)
Epithelia
Mesogloea very thick, up to 65 mm measured (column);
ectoderm very thin compared to mesogloea (Fig. 4f).
Batteries of spirocysts visible in tentacle ectoderm;
acidophil inclusions in ectoderm of column and filaments. Siphonoglyphs with strongly developed reticulated pads (Fig. 4f).
Cnidae
Spirocysts (in tentacles), basitrichs (in all tissues),
microbasic b-mastigophores (in filaments, and in some
specimens in tentacles), microbasic amastigophores A
(in pharynx, filaments, tentacles) (Fig. 5a–k). See
Table 1 for information on size and distribution of
cnidae.
Additionally, I found rare, exceptional cnidae in
single specimens: eight large basitrichs 49.5–63.9·3.6–
4.5 lm in the distal column, three b-mastigophores 24.3–
25.2·4.1–6.3 lm in the filaments of A. chilensis; four
b-mastigophores 40.5–51.3·5.4–6.3 lm in the column
and three b-mastigophores 80·11 lm in the filaments of
A. chilensis Coll.
The basitrichs of the unilobate filaments varied in
shape from an elongated capsule of equal breadth
(Fig. 5J1) to a capsule that narrows towards both ends
(Fig. 5J3); the length of the shaft varied from half
(Fig. 5J2) to the full length (Fig. 5J1) of the capsule. In
the filaments of a preserved specimen of A. chilensis
Coll, a fired cnida which looked like a p-mastigophore B
with a short ‘‘Faltstück’’ was found.
The tubule of fired microbasic amastigophores A of
pharynx (Fig. 5G2) and filaments (Fig. 5K2) has a
proximally thickened shaft equal in length to the
remainder of the tubule.
Distribution and zoogeography of Actinostola chilensis
A. chilensis can be found in more or less protected bays
and fjords of the northern part of the Chilean fjord
region from Seno de Reloncavı́ (4135¢35¢¢S, 7253¢W)
to the fjord Puyuhuapi (4431,608¢S; 7232,107¢W)
(Fig. 1, Appendices 2 and 3). I commonly found this
species in the Golf of Reloncavı́ (S53, S57), as well as in
345
the fjords Comau (S60) and Reñihue (S61–65), somewhat less commonly in the fjord Puyuhuapi (S90) and
around Raul Marin Balmaceda (S83, S85); I did not
find it along the exposed coasts around Bahia Tic Toc
(S77-S82). Dirk Schories (in litt. 2004) found it around
Melinka Island (S84a). While diving, I observed one
specimen in the fjord Chacabuco (S96), which probably
belonged to the same species. However, in contrast to
346
all the other specimens examined, it was salmon-coloured rather than orange. Most probably, A. chilensis
can be found further south than the fjord Puyuhuapi,
at least to Peninsula Taitao (4630¢–4657¢S). This large
peninsula divides the Patagonian Province and is considered a zoogeographical barrier (Lanzellotti and
Vasquez 1999; Häussermann 2004). I found this species
as shallow as 22 m, but more frequently between 32
and 45 m; Carlgren (1959) stated that the range extends
to 278 m.
Natural history of Actinostola chilensis
A. chilensis is a very eye-catching species that can be
found in exposed positions on rocky substrate attached
to bare rock (Fig. 2c; Appendix 5A), generally on the
edge of terraces or on top of elevations, but never under
overhangs (Fig. 2) and never with its oral disc
Fig. 5 Cnidae of Actinostola
chilensis. Letters A–K
correspond to cnidae listed in
Table 1 (Tentacles A large bmastigophore, B spirocyst, C
basitrich, D microbasic
amastigophore A; Column E
basitrich; Pharynx F basitrich,
G microbasic amastigophore A;
Pedal disc H basitrich, I
spirocyst; Filaments J basitrich,
K microbasic amastigophore A)
downward. One specimen was found attached to the axis
of a gorgonian of the genus Primnoella Gray 1858
(Appendix 5B). Specimens can be found either individually or, in favourable spots such as rocky elevations, in
aggregations of up to 15 individuals, in some cases
touching one another (Fig. 2a,c). During our survey,
water temperature in the habitat of A. chilensis ranged
from approximately 6C in winter to approximately
11C in summer; salinity was approximately 31&. In the
fjords, surface water to depths of 8 m is often brackish,
with minimum salinities lower than 10&; tidal amplitudes are up to 7.3 m (data measured in 2003 in Comau
fjord at Huinay Scientific Field Station).
Most specimens we observed were fully expanded in a
typical position with the inner tentacles directed upward
and the outer tentacles sideward (Fig. 2; Appendix 5a–
c); in current, the oral disc points downstream. Some
specimens were slightly or completely contracted, covering the tentacles with the column (Fig. 2d). In Chilean
347
fjords, I regularly found A. chilensis associated with the
gorgonian Primnoella sp. (Fig. 2c; Appendix 5B), with
the giant brachiopod Magellania venosa (Dixon 1789)
(Appendix 5C) and in the vicinity of the azooxanthellate
coral Desmophyllum dianthus (Esper, 1794) (Försterra
and Häussermann 2003).
Specimens of A. chilensis are easy to collect by scuba
diving because they come off the substratum readily and
without injury. When disturbed, animals completely
contract. Tentacles are sticky to the touch. In the
aquarium, specimens need 1 to several days to reattach,
requiring well-oxygenated, salty (non-superficial) water
with considerable current and, ideally, darkness before
they resume their in situ appearance. In unsuitable
conditions, they protrude the pharynx, eject food and
change position, e.g. by somersaults over the oral disc.
They can quickly and strongly change their shape, e.g.
from long trumpet-shaped over vase-or cup-shaped to
an almost perfect sphere. A. chilensis relaxes readily with
menthol, but extended exposure to this anaesthetic results in maceration of tissue and reduction in sensitivity
of cnidae to fire.
Discussion
Generic description of Actinostola
The proposed emendations to the diagnosis of Actinostola correct errors in its definition. One of the main
errors is the statement that the animals cannot fully
cover the tentacles with the column; this characteristic
was erroneously used to distinguish Actinostola from
Stomphia (Carlgren 1949). However, I found that living
specimens of A. chilensis are capable of fully covering
the tentacles with the column, and do so regularly
(Fig. 2d). I propose to delete the phrase ‘‘tentacles never
more numerous than the mesenteries at the base’’: of 13
specimens I examined, only 3 had fewer tentacles than
mesenteries at the base, and 9 had more; 1 had as many
tentacles as mesenteries at the base. Mesenteries are
numerous and thin and thus hard to count. I could not
confirm the presence of numerous perfect mesenteries: in
the examined specimens, generally 24 of approximately
100 pairs were perfect; in other species, up to 48 pairs
may be perfect. None of these emendations conflict with
the description of the type species A. callosa (Appendix
6).
Carlgren (1928) erected the genus Paractinostola
based on only minor differences between it and Actinostola: more tentacles than mesenteries at the pedal disc
and a more or less lobed oral disc. I agree with Riemann-Zürneck (1978) that both features may be present
in large members of some species of Actinostola and
concur with her synonymy of these genera. Sensu Carlgren’s (1949) generic identification key for the family
Actinostolidae, Ophiodiscus and Stomphia (Appendix
7A) are the only remaining genera within group IA,
which is characterized by possessing ‘‘mesenteries
distinctly arranged according to the Actinostola-rule’’.
The genus Ophiodiscus (Hertwig 1882) contains the two
species, Ophiodiscus annulatus and O. sulcatus, from
deep waters off North Chile; neither has been found
since the beginning of the twentieth century. In Ophiodiscus, mesenteries are divided into macro- and micronemes, and tentacles are arranged in a single corona.
Stomphia is distinguished from Actinostola by its cnidae
(see Table 1), by having a central rise on the pedal disc,
and 16 perfect, fertile mesenteries, and by always lacking
thickenings on the outer base of the tentacles (for in vivo
photographs see Appendices 5 and 7).
Taxonomic characteristics to distinguish Actinostola
species
Riemann-Zürneck (1971, 1978) showed that the following characteristics that had been used to denominate
species of Actinostola were very variable within A. callosa: marginal stomata, tentacular b-mastigophores,
thickenings at the outer base of the tentacles, thickenings
of the oral disc, and thickness of tentacular mesogloea.
The size and structure of the marginal sphincter vary
across the genus, but variation is more dependent on the
size of the specimens (Fig. 4a–d) than on taxonomy,
varying only slightly between species (Riemann-Zürneck
1971, 1978).
Riemann-Zürneck (1978) suggested the cnidae of the
unilobate filaments might distinguish species: she found
a second, smaller basitrich in those of A. crassicornis
that was lacking in A. intermedia. I have examined type
material of these species and did not find differences in
the cnidae of the unilobate filaments (Table 1). In
examining numerous filaments of the specimens I collected, I noticed that the basitrichs of these filaments
vary in shape and shaft length (see Fig. 5J) and that
some (Fig. 5J2, J5) resemble the microbasic p-mastigophores of the filaments of some actiniid anemones. I
observed intermediate stages of shape and shaft length
(Fig. 5J3), and thus could not distinguish distinct types
of cnidae. From my data, I conclude that neither types
nor sizes of cnidae are useful for distinguishing the
examined species. However, since fired cnidae reveal
more distinctive features than unfired ones, I have provided photographs of fired cnidae from A. chilensis
(Fig. 5), which could serve a comparative purpose in the
future if fired cnidae are examined for other species in
the genus.
Riemann-Zürneck (1978) proposed size and shape of
the preserved specimens as an additional feature, if a
large number of well-preserved specimens were available. The specimens I have examined showed extreme
variability in shape, and a single specimen may take the
shape of all described forms within minutes in the
aquarium. Due to the fact that the form of the preserved specimens strongly depends on the state of
relaxation, I reject this feature as a species-level characteristic.
348
Since preserved specimens of different Actinostola
species are extremely similar in anatomy and in the
shape, size and distribution of cnidae (Riemann-Zürneck 1978), the distinction of species has to be based on
other characteristics, such as reproductive mode (e.g.
brooded embryos in the gastrocoel, hermaphroditism),
texture of the column or range of variability in morphological features (e.g. maximum pedal-disc diameter,
maximum thickness of the columnar mesogloea, presence or absence of tentacular thickenings or marginal
stomata in all specimens from different depths). However, to detect these features, several specimens are
needed. This is problematic because many species of
Actinostola are only known from a few specimens. In
addition, many type specimens are more than 100 years
old and, at least in the case of some South American
species, are in rather poor condition. Thus additional
information is necessary to clarify species boundaries;
this additional information may include zoogeographic,
in situ and in vivo information. For a well-founded
distinction of the species of Actinostola found in subAntarctic and Antarctic waters, it will be necessary to
collect, examine and re-describe in detail specimens of
Argentinean and Antarctic species of Actinostola. In situ
and in vivo characteristics such as appearance (e.g.
length of tentacles or posture), variability of colour,
ecological niche, position in the habitat, mobility in the
aquarium and behaviour (e.g. reaction to disturbance)
that distinguish the species of Actinostola might be
useful to complete the few existing distinguishing characteristics so that species can be defined and an identification key can be made (for an example, see Appendix
6). This re-description of A. chilensis is a first step and
should be used as a comparative base for future work.
Synonymization of South American and Antarctic
species of Actinostola
Eight species of Actinostola have been described from
southern South America and Antarctica (Fig. 1;
Appendices 4 and 8). Several of these were later synonymized: Carlgren (1927) synonymized A. chilensis with
A. intermedia; Riemann-Zürneck (1978) synonymized
A. excelsa and A. pergamentacea with A. crassicornis, and
Fautin (1984) additionally included A. intermedia and
A. clubbi in the synonymy list of A. crassicornis, making
this species the most abundant and wide-spread species of
Actinostola in the southern hemisphere (Fig. 1; Appendix
8). Riemann-Zürneck (1978) and Fautin (1984) provided
discussions of the rationale for these synonymies, but
because they were based exclusively on anatomical and
cnidae data of questionable distinctive value for
preserved specimens, they should be reconsidered.
Riemann-Zürneck (1978) found that approximately
half of the specimens of A. crassicornis had embryos in
the gastrocoel. In contrast, none were found in any of
the type specimens of its putative synonyms A. excelsa
(three specimens), A. pergamentacea (five specimens),
A. intermedia (one specimen), A. chilensis (one specimen)
or A. clubbi (one specimen). Fautin (1984) regularly
found brooded young in the many USARP specimens
from Antarctic and sub-Antarctic waters that she
examined (Appendix 8). But since she assigned all
specimens from all locations to A. crassicornis based on
characteristics that are of minor distinctive value, her
description might refer to a species complex. The anatomy and cnidae of preserved specimens of A. intermedia,
A. excelsa and A. pergamentacea do not differ from
those of A. crassicornis. Due to the lack of distinctive
features and because of their overlapping distribution, I
accept the synonymy of A. excelsa, A. pergamentacea
and A. crassicornis. The fact that no embryos were
found in the few type specimens of A. excelsa and
A. pergamentacea is not proof that members of these
species do not brood young. However, a possible cooccurrence of brooding and non-brooding species of
Actinostola along the Argentinean shelf and the extension of A. crassicornis into the Antarctic Ocean require
further investigation.
I reject the synonymy between the Chilean species
A. chilensis and the Argentinean species A. crassicornis.
Neither Carlgren (1959), McMurrich (1904), nor I found
brooded young in any specimens of A. chilensis collected
in different regions, years and seasons in the fjords of
southern Chile. The holotype of A. chilensis is hermaphroditic, a unique characteristic among species of
Actinostola from South America. These differences in
the reproductive mode in combination with possible the
zoogeographical barriers such as the Peninsula Taitao
and the Strait of Magellan (Riemann-Zürneck 1986;
Lancellotti and Vásquez 1999) lead me to recognize
A. chilensis as a distinct species. A. chilensis is distinct
from several species of Actinostola found in the Antarctic Ocean and in Argentina; it differs significantly in
in vivo appearance, colour and texture of the column
(Rodriguez, in litt. 2003; Roux, in litt. 2003).
I accept the synonymy of A. intermedia and
A. crassicornis, but reject the synonymy of A. intermedia
and A. chilensis for zoogeographical reasons: the type
locality of A. intermedia was erroneously described as
‘‘Cabo San Vicente, Strait of Magellan’’ (Carlgren 1899)
and was therefore claimed to be in Chile. However, the
only ‘‘Cabo San Vicente’’ in this region is situated on the
Atlantic coast of Argentina, in Tierra del Fuego (Fig. 1),
a locality that lies within the known range of A. crassicornis. This separates the localities where specimens of
the Argentinean and Chilean species were found by
more than 1,000 kilometres and by zoogeographical
barriers.
Comparison of cnidae within South American species
of Actinostola
The cnidom of S. selaginella is clearly distinct from that of
the examined species of Actinostola: it lacks amastigophores A in the tentacles, and includes two types of
349
basitrichs in the pharynx and two types of microbasic
p-mastigophores in the filaments (Table 1). Taking into
account the standard deviation and the differences in sizes
of the examined specimens, the sizes of the following
cnidae do not differ significantly among the species of
Actinostola I compared: spirocysts and basitrichs of tentacles and all cnidae of column, pharynx and filaments.
The differences between mean values were approximately
equal to the standard deviations and thus not useful to
distinguish species. Spirocysts are distributed patchily and
thus were not encountered on every slide made from tissue
of the pedal disc of specimens of A. chilensis (both type
and Coll) or A. intermedia. The cnidae of the Chilean
specimens differ from those of the Argentinean ones in
minor details: the microbasic amastigophores A of the
tentacles and b-mastigophores of the filaments are scarcer, but have the same average sizes. The basitrichs of the
unilobate filaments of all of the Argentinean species show
a very broad size range, and there was no evidence for two
distinct size groups as proposed by Riemann-Zürneck
(1978); the mean value is very similar in all species of
Actinostola examined (Table 1).
Further records of Actinostola species along the west
coast of South America
Two other species identified as Actinostola have been
reported from the west coast of South America. Doumenc (1984) identified specimens collected from 350 to
450 m depth off central Chile, 30–3231¢S, as A. intermedia (which was at the time considered a synonym of
A. chilensis). McMurrich (1893) identified specimens
from 717 to 1,484 m depth between Ecuador and
Galapagos Islands, 024¢–037¢S as A. callosa. McMurrich (1893) considered the last species to be widely distributed and synonymous with A. crassicornis; Carlgren
(1927) rejected this synonymization, assigning the specimens from Ecuador to A. crassicornis rather than to
A. callosa, all other specimens of which are known from
the North Atlantic. Zoogeographical transition areas
such as the region between Coquimbo and Valparaı́so,
central Chile and around Paita, Peru (Brattström and
Johanssen 1983; Lancellotti and Vásquez 1999; Sullivan
Sealey and Bustamante 1999) are supposed to exist between the sites where I found A. chilensis and the two
other records further north along the west coast of
South America. However for some species, generally of
deep waters, these barriers may not apply. The abovementioned species descriptions do not offer enough
information for a well-founded identification, and an
examination of the specimens is necessary for a final
decision if these specimens belong to A. chilensis.
Therefore the records have to be treated as uncertain.
Acknowledgements I am particularly grateful to Günter Försterra
for his company and great help with diving and sampling during
the field trips, and to Meg Daly for very detailed and constructive
comments. Many thanks to Kensuke Yanagi for very helpful
remarks. Many thanks also to Björn Sohlenius, Carsten Lüter and
Stephen Cairns for the loan of type material, to Karin Riemann for
lending a field microscope and to Estefanı́a Rodriguez for helpful
discussions. Thanks go to Gerhard Haszprunar for providing
material, space and continuous support; to Carlos Gallardo,
Alejandro Bravo, Elena Clasing and Wolfgang Stotz for their
friendly support, and to Heide Felske for making available a
wonderful map. Many thanks go to the Huinay Foundation for a
scholarship for lodging, food and working space in the marine
biology station in Comau fjord, and to Rose and Fritz Häussermann for their manifold and continued help. This publication is
drawn from the doctoral thesis of the author, supported by 2 oneyear governmental scholarships ‘‘Förderung des wissenschaftlichen
und künstlerischen Nachwuchses’’ and ‘‘Förderung der Promotion
von Wissenschaftlerinnen’’ from the LMU Munich, and by a oneyear HSP III scholarship from the DAAD. This is publication
number 2 of the Huinay Scientific Field Station.
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