J. Mar. Biol. Ass. U.K.

(2000), 80, 719^724

Printed in the United Kingdom

An analysis of the nematocysts of the beadlet anemone

Actinia equina and the green sea anemone Actinia prasina
Phillip C. Watts*, A. Louise AllcockO, Sean M. Lynch and John P. Thorpe
Port Erin Marine Laboratory, School of Biological Sciences, University of Liverpool, Port Erin, Isle of Man, IM9 6JA.
*Present address: Laboratory 1.03, Donnan Laboratories, School of Biological Sciences, University of Liverpool, Crown Street,
Liverpool, L69 7ZD. E-mail: p.c. [email protected]
O
Present address: National Museums of Scotland, Chambers Street, Edinburgh, EH1 1JF

Numerous studies of their population genetics have reported incipient reproductive isolation among
sympatric populations of the common intertidal beadlet anemone Actinia equina. This has lead to certain
morphs being raised to speci¢c status. A study of the nematocysts of the green sea anemone Actinia prasina
and three genetically isolated morphs of A. equina was undertaken to establish that mean nematocyst
length could act as diagnostic phenotypic characters within a morphologically variable group. The results
support genetic and ecological evidence for the speci¢c status of the three red/brown coloured morphs of
A. equina. The data are discussed with respect to the ecology of Actinia and concepts of species, but more
work is required before the speci¢c status of A. prasina can be con¢rmed.

INTRODUCTION as the Up (upper-shore), Mid (mid-shore) and Low

(lower-shore) morphs. All three morphs have red or
The intertidal beadlet anemone Actinia equina (L.) is brown columns, but the Low morph can be readily
common on rocky intertidal and shallow subtidal habitats identi¢ed because it has a green/grey pedal disc bordered
throughout the British Isles. Its known geographic distri- by a blue line. The Up and Mid morphs both have red/
bution ranges from the Kola Peninsula, along the west pink coloured pedal discs and could only be identi¢ed
coast of Europe, throughout the Mediterranean and using a diagnostic allozyme locus until it was demon-
Black seas, to the west African coast (Stephensen, 1935; strated that there are nematocysts di¡erences between
Manuel, 1988; Shick, 1991). The colour of A. equina is them (Watts & Thorpe, 1998). The Low morph has been
highly variable and numerous di¡erent colour varieties or separated from A. prasina (also found mainly on the low
`morphs' have been described (Tugwell, 1856; Gosse, shore) on the grounds of column colour alone, but recent
1860; Stephensen, 1935), which has raised considerable genetic evidence (Perrin, 1993; Lynch, 1996) suggests that
debate as to whether certain morphs constitute di¡erent they may be conspeci¢c.
species. Some of the colour morphs were assigned speci¢c Cnidae (stinging cells) are characteristic of the
status by several early workers (e.g. Dalyell, 1848; Cnidaria and are essential to their mode of life. Within
Tugwell, 1856), but Gosse (1860) considered all the forms the Actiniaria two categories of cnidae can be readily
as varieties of a single species. discerned, the spirocysts and the nematocysts. Although
Carter & Thorpe (1981) used isozyme electrophoresis spirocysts are thought to be of limited systematic value
and reproductive evidence to separate Stephensen's (1935) (Cutress, 1955; Manuel, 1988), nematocysts may o¡er
Actinia equina mesembryanthemum and A. equina fragacea into useful characters for distinguishing between both higher
the di¡erent species Actinia equina (L.) and Actinia fragacea taxa and di¤cult taxonomic groups (Fautin, 1988;
(Tugwell) respectively. Due to the apparent lack of diag- Manuel, 1988). Several distinct size-classes of the same
nostic phenotypic characters, subsequent taxonomic work nematocyst type are often present within a tissue,
on A. equina has generally been based upon genetic although the functional signi¢cance of this is not known
analysis. Electrophoretic data and ecological di¡erent- (Mariscal, 1974). A preliminary comparison of nemato-
iation have led to a morph of A. equina being given speci¢c cyst distribution and abundance between a mixture of
status as Actinia prasina (Gosse) (Haylor et al., 1984). red/pink pedal disc morphs and A. prasina found no
Actinia prasina has a green column with a green pedal disc signi¢cant di¡erences (Solë-Cava & Thorpe, 1987, 1992).
bordered by a blue line around the limbus and occupies a A later study, however, revealed signi¢cant di¡erences in
low shore distribution (Haylor et al., 1984; Solë-Cava & nematocyst size between the red/pink and green/grey
Thorpe, 1987, 1992). pedal disc morphs of A. equina (Allcock et al., 1998).
Other research, however, focused upon pedal disc Subsequently, it was found that the size of the nematocysts
colour (e.g. Quicke et al., 1983; Quicke & Brace, 1984), from the acrorhagi allows phenotypic identi¢cation of the
which varies more discontinuously than column colour. Up and Mid morphs of A. equina (Watts & Thorpe, 1998).
Within A. equina, three genetically distinct morphs with No nematocyst studies, however, have described the size
di¡erent intertidal distributions were identi¢ed (Quicke distribution of nematocysts throughout the di¡erent
et al., 1983; Quicke & Brace, 1984) which are referred to tissues of the Up, Mid and Low morphs or have

Journal of the Marine Biological Association of the United Kingdom (2000)

720 P.C. Watts et al. An analysis of the nematocysts of Actinia spp.

Table 1. Mean nematocyst length (mm) for three genetic morphs of the beadlet anemone Actinia equina and the green sea anemone
A. prasina.

Actinia equina Actinia prasina

Tissue Upper-shore morph Mid-shore morph Low-shore morph

Nematocyst Type N Mean SD N Mean SD N Mean SD N Mean SD

Acrorhagi
Holotrich 240 41.85 3.37 240 52.49 5.26 240 51.48 4.11 240 50.38 3.72
Pharynx
Microbasic b-mastigophore I 240 13.67 0.61 240 15.54 0.90 240 13.38 0.58 240 12.86 0.51
Microbasic b-mastigophore II 240 23.95 1.08 240 25.84 1.21 240 24.46 0.62 240 20.74 0.62
Mesenteric ¢laments
Microbasic b-mastigophore I 240 13.00 0.83 240 14.65 0.97 240 12.74 0.40 240 12.88 0.72
Microbasic b-mastigophore II 68 19.77 1.82 138 19.46 0.59 162 17.56 0.62 193 17.80 0.46
Microbasic b-mastigophore ö ö ö 85 32.86 1.34 87 32.75 2.17 131 32.22 1.31
III
Microbasic p-mastigophore 240 19.36 1.31 240 23.55 2.08 240 21.26 0.69 240 21.25 0.83
Tentacles
Microbasic b-mastigophore I 116 12.37 0.64 152 14.57 0.97 219 13.41 0.45 213 12.86 0.44
Microbasic b-mastigophore II 240 20.13 0.65 240 22.59 0.96 240 19.97 0.75 240 20.74 1.06
Column
Microbasic b-mastigophore 240 14.67 0.51 240 17.08 1.22 240 14.17 1.24 240 14.99 0.76
Pedal disc
Microbasic b-mastigophore 240 14.12 0.35 240 15.80 0.17 240 12.55 0.42 240 13.04 1.05

N, number; SD, standard deviation.

speci¢cally investigated A. prasina. Given the extensive (Perrin, 1993), had been collected. Eight individuals of
taxonomic confusion within `A. equina' and the putative each of the Low morph and A. prasina were collected.
usefulness of nematocysts for taxonomy, further research Horizontal starch gel electrophoresis was employed to
seems warranted. The aims of this study, therefore, are to identify the Up and Mid morphs (for details of methods
provide a description of the nematocysts in A. prasina and see Solë-Cava & Thorpe, 1987 or Lynch, 1996). Gels were
the three morphs of A. equina and to relate these data to stained for malate dehydrogenase since this enzyme is
previous genetic work. diagnostic for these morphs (Quicke & Brace 1984;
Lynch, 1996). Eleven Up morph anemones were obtained
out of the mixed (Up and Mid morph) sample.
MATERIALS AND METHODS Because feeding and aggressive behaviour may reduce
Actinia prasina and the three morphs of Actinia equina nematocyst numbers all anemones were kept in individual
were sampled from Fleshwick Bay (OS grid reference SC pots, with the seawater replaced every 23 d, for at least a
202715), a sheltered bay on the west coast of the Isle of week before tissue samples were collected. Simple tissue
Man which contains both the sheltered boulder habitats squashes were made from small (51mm3) pieces of tissue
(typical of A. prasina and the Low morph) and more and stained using 0.1% methylene blue solution (see
exposed vertical rock faces (preferred by the Up and Mid Manuel, 1988). Nematocysts were viewed using a Zeiss
morphs). To standardize for size, the maximum and phase-contrast binocular microscope at 1000 and
minimum diameter of the pedal disc of each anemone measured using an eyepiece graticule calibrated against a
was measured using dial callipers. The length of certain stage graticule. As nematocyst length may vary with
nematocysts may vary with body size of the sea anemones tissue location (Cutress, 1955) all major types of nemato-
(Chintiroglou, 1996) or their weight (Chintiroglou & cysts were measured in tissue from speci¢c areas. Only
Simsiridou, 1997). Thus specimens were taken whose undischarged nematocysts were measured since large
average pedal disc diameter was 18^20 mm since this changes in volume may occur after discharge (Godknecht
appeared to be the most frequent size-class on the shore, & Tardent, 1988).
facilitating collection of rarer animals (A. prasina and the Six di¡erent tissue regions were studied: acrorhagi, tips
Up morph). Di¡erent morphs were collected as close of tentacles, central pharynx, mesenteric ¢laments, mid-
together as their ecological separation allowed whilst column and the centre of the pedal disc. Nematocyst types
anemones of the same morph were collected from a were identi¢ed following Cutress (1955). Within randomly
minimum of 2 m apart to minimize the chance of selected ¢elds of view 15 nematocysts of each type were
collecting clonemates. Since the Up and Mid morphs are measured. Two replicate samples were measured from each
indistinguishable on the shore 55 red pedal disc animals tissue for eight anemones of each morph. Where two or
were collected between mean tide level and mean high more discrete nematocysts sizes were present within a
water neap to ensure that enough of the Up morph, tissue they were counted separately and su¤xed I, II, etc.,
which is relatively uncommon around the Isle of Man according to size. Rarer types of nematocysts (i.e. where 15

Journal of the Marine Biological Association of the United Kingdom (2000)

 An analysis of the nematocysts of Actinia spp. P.C. Watts et al. 721

Table 2. Hierarchical analysis of variance of nematocyst length for three genetic morphs of Actinia equina and Actinia prasina.

Source of variation

Tissue Within tissue Within morphs Between morphs

Nematocyst Type F P F P F P

Acrorhagi
Holotrich 0.55 ns 69.16 50.001* 754.23 5
50.001*
Pharynx
Microbasic b-mastigophore I 1.07 ns 7.31 50.001* 144.30 5
50.001*
Microbasic b-mastigophore II 1.53 50.05 11.11 50.001* 63.86 5
50.001*
Mesenteric ¢laments
Microbasic b-mastigophore I 1.36 ns 9.00 50.001* 99.61 5
50.001*
Microbasic b-mastigophore II ö ö 3.94 50.001* 42.71 5
50.001*
Microbasic b-mastigophore III ö ö ö ö 0.13 ns
Microbasic p-mastigophore 1.00 ns 29.44 50.001* 383.91 5
50.001*
Tentacles
Microbasic b-mastigophore I ö ö 3.91 50.001* 55.87 5
50.001*
Microbasic b-mastigophore II 0.93 ns 8.67 50.001* 132.74 5
50.001*
Column
Microbasic b-mastigophore 0.98 ns 19.75 50.001* 267.80 5
50.001*
Pedal disc
Microbasic b-mastigophore 0.53 ns 11.93 50.001* 285.90 5
50.001*

F, F-value; P, probability; ns, not signi¢cant; *, indicates a signi¢cant (P50.05) result after a sequential Bonferroni correction for
multiple testing.

Table 3. Probability values for Tukey pairwise comparisons of mean nematocyst length between the upper- (U), mid- (M), and
lower-shore (L) morphs of Actinia equina and Actinia prasina (P).

Tissue Pairwise Comparison

Nematocyst Type U^M U^L U^P M^L M^P L^P

Acrorhagi
Holotrich 5
50.001 5
50.001 5
50.001 ns 50.05 ns
Pharynx
Microbasic b-mastigophore I 5
50.001 ns ns 5
50.001 5
50.001 ns
Microbasic b-mastigophore II 5
50.001 ns 50.05 5
50.001 5
50.001 ns
Mesenteric ¢laments
Microbasic b-mastigophore I 5
50.001 ns ns 5
50.001 5
50.001 ns
Microbasic b-mastigophore II ns 5
50.001 5
50.001 5
50.001 5
50.001 ns
Microbasic p-mastigophore 5
50.001 5
50.001 5
50.001 5
50.001 5
50.001 ns
Tentacles
Microbasic b-mastigophore I 5
50.001 50.01 ns 5
50.001 5
50.001 ns
Microbasic b-mastigophore II 5
50.001 ns ns 5
50.001 5
50.001 50.01
Column
Microbasic b-mastigophore 5
50.001 5
50.001 ns 5
50.001 5
50.001 5
50.001
Pedal disc
Microbasic b-mastigophore 5
50.001 5
50.001 5
50.001 5
50.001 5
50.001 50.01

ns, not signi¢cant.

could not be encountered readily) were speci¢cally data were analysed using a three level nested (hierarch-
searched for and all encountered were measured. ical) ANOVA (Zar, 1984), with the within-tissue variation
All data were analysed using MINITAB (1991) statis- nested within the between-anemone variation, and both
tical software. The size^ frequency of each nematocyst sources of variability being nested within that occurring
population was tested for normality using the NSCORES among the di¡erent morphs. Unbalanced data (i.e. data
procedure (Minitab, 1991); no signi¢cant (P40.01) depar- with unequal sample sizes) were analysed with a general
tures from normality were detected. Nematocyst length linear model of ANOVA; because these data included the

Journal of the Marine Biological Association of the United Kingdom (2000)

722 P.C. Watts et al. An analysis of the nematocysts of Actinia spp.

rarer nematocysts the within-tissue and within-morph it is probable that the distal parts were examined.
variance could not always be calculated. A table-wide Indeed, the di¤culty in exact dissection as well as their
type-I error rate of aˆ0.05 was maintained using a low abundance may explain why the microbasic b-masti-
sequential Bonferroni correction (Rice, 1989). Following a gophore III nematocyst population was not found by
rejection of the null hypothesis a Tukey multiple compar- Allcock et al. (1998). The lack of within-tissue variation
ison test (Zar, 1984) was calculated to locate which in nematocyst length, however, suggests that a tissue
morphs had di¡erent nematocyst parameters. sample from one area should be representative and
supports previous data (see Allcock et al., 1998).
Di¡erences in nematocyst type and distribution are
RESULTS only to be expected in distantly related taxa or among
Three categories of nematocyst were identi¢ed: the individuals at di¡erent stages of maturity (Cutress,
holotrich, microbasic b-mastigophore (ˆb-rhabdoid) and 1955). The absence of the microbasic b-mastigophore III
microbasic p-mastigophore (ˆp-rhabdoid). A few of the population from the mesenteric ¢laments of the Up
smaller microbasic b-mastigophores often showed a morph (Table 1) implies that it may be the most
gradation towards the structure of a basitrich. Since the distantly related taxon, but it is more probable that this
armature of the shaft could not be resolved and the nematocyst population was overlooked since it was rela-
majority of the nematocysts possessed a full central shaft tively rare. This perhaps demonstrates the di¤culty in
when undischarged they were all considered to be micro- only being able to study fresh tissue as it would
basic b-mastigophores; Manual (1988) regards basitrichs obviously be desirable to search further for this nemato-
and microbasic b-mastigophores to be synonymous. cyst population.
The qualitative distribution of nematocyst types Previous work has found no signi¢cant di¡erences
throughout the tissues revealed no di¡erences between between Actinia prasina and a mixture of red/pink pedal
the green Actinia prasina and the morphs of Actinia equina disc morphs of A. equina (Solë-Cava & Thorpe, 1987, 1992).
except that the microbasic b-mastigophore III population These analyses, however, may have been confounded by
is absent from the mesenteric ¢laments of the Up morph not distinguishing among the Up and Mid morphs and,
(Table 1). No signi¢cant (P40.05) variation in nematocyst furthermore, the sizes of the animals used were not stan-
size was found within tissues after correction for multiple dardized. Assuming that interbreeding will e¡ect a similar
testing (Table 2). All nematocyst populations tested size distribution of nematocyst lengths, the highly signi¢-
showed signi¢cant di¡erences in size between anemones cant di¡erences in nematocyst lengths among morphs
of the same morph after a Bonferroni correction provide support for previous genetic work suggesting that
(P50.05), although the within-morph variation for the there are a number of species within `Actinia equina' (see
microbasic b-mastigophore III nematocysts was not tested Quicke & Brace, 1983, 1984; Quicke et al., 1985; Solë-
because of small sample size (Table 2). Similarly, all Cava & Thorpe, 1987; Perrin, 1993; Lynch, 1996). It may
nematocyst populations, except the microbasic b-mastigo- be concluded that the Up, Mid and Low morphs of
phore III from the mesenteric ¢laments, demonstrated A. equina are all reproductively isolated and should there-
highly signi¢cant variation in size (P50.001) among fore be considered separate species.
morphs (Table 2) with all comparisons remaining signi¢- From genetic studies, Perrin (1993) suggested that the
cant (P50.05) after correction for multiple testing. Low morph was conspeci¢c with A. prasina. In addition,
The Mid morph has larger nematocysts than the Up these taxa have similarly coloured pedal discs bordered
and Low morphs and Actinia prasina for all but one nema- by a blue line, have a low strength of attachment to
tocyst population (Table 1); these di¡erences are signi¢- substrate (Quicke & Brace, 1983; Perrin, 1993), prefer low
cant for almost all of the pairwise comparisons (Table 3). shore cryptic habitats (Quicke & Brace, 1984; Haylor et
Actinia prasina and the Low morph showed consistent al., 1984; Perrin, 1993) and display reduced levels of
similarities in nematocyst lengths except for three nema- aggression (Brace & Reynolds, 1989). Furthermore, the
tocyst populations (Tables 1 & 2). The Up morph di¡ered analysis of nematocyst lengths between A. prasina and
from A. prasina and the Low morph in approximately half the Low morph resulted in the most similarities from
of the nematocyst populations (Table 3), but revealed no the pairwise comparisons of anemone taxa (Table 3).
pattern to the size di¡erentiation. Despite this and, contrary to the genetic data of Perrin
(1993), the signi¢cant di¡erences present in some type
of cnidae between the Low morph and A. prasina are
DISCUSSION indicative of them being distinct species. Although
The distribution of nematocyst types given here Perrin (1993) did not ¢nd any MDH heterozygotes
include those presented for Actinia equina by Solë-Cava & within A. prasina, more recent genetic data (S.M. Lynch,
Thorpe (1987, 1992), except that in this study microbasic unpublished data) have identi¢ed some `A. prasina' from
b-mastigophores were not considered to be present in the Fleshwick Bay as being heterozygous at the MDH-1
acrorhagi (Table 1). The presence of microbasic b-masti- locus. It is therefore possible that the `A. prasina' sample
gophores was noted to increase in smaller acrorhagi taken for this study also contained green-columned Mid
where it was di¤cult to make a clean dissection, and morphs. Atypical genotypes have also been noted in the
since the microbasic b-mastigophores found in this study red/brown column morphs (Quicke & Brace, 1983).
were of the same size as those from the neighbouring This result is important as it demonstrates that neither
column tissue they were considered to be contaminants. column nor pedal disc colour can be used reliably to
The exact part dissected from the mesenteric ¢laments discriminate among the morphs of British `A. equina'.
cannot be given due to their convoluted nature, although Since anemones cannot synthesize pigments, rather they

Journal of the Marine Biological Association of the United Kingdom (2000)

 An analysis of the nematocysts of Actinia spp. P.C. Watts et al. 723

assimilate them from their diet (Lubbock & Allbut, Dalyell, J.G., 1848. Rare and remarkable animals of Scotland repre-
1981; Shick, 1991) their colour is likely to be environ- sented from living subjects with practical observations on their nature.
mentally determined and cannot be used as a diagnostic London: Van Voorst.
character. Con¢rmation of the speci¢c status of Fautin, D.G., 1988. Importance of nematocysts to Actinian
taxonomy. In The biology of nematocysts (ed. D.A. Hessinger and
A. prasina, therefore, cannot be given and should await
H.M. Lenho¡ ), pp. 487^500. California: Academic Press.
a full genetic analysis with a concurrent description of Godknecht, A. & Tardent, P., 1988. Discharge and mode of
the nematocysts. It must be remembered that similari- action of the tentacular nematocysts of Anemonia sulcata
ties in nematocyst length do not necessarily imply that (Anthozoa: Cnidaria). Marine Biology, 100, 83^92.
the subjects are conspeci¢c (e.g. Schmidt, 1972 found no Gosse, P.H., 1860. Actinologia Britannica. A history of the British sea
signi¢cant di¡erences in nematocyst lengths between anemones and corals. London: Van Voorst.
A. equina and A. fragacea) but rather may simply re£ect a Haylor, G.S., Thorpe, J.P. & Carter, M.A., 1984. Genetic and
recent common origin. ecological di¡erentiation between sympatric colour morphs of
Since A. equina morphs have failed to breed under the common intertidal sea anemone Actinia equina. Marine
laboratory conditions, a biological species concept (Mayr, Ecology Progress Series, 16, 281^289.
1963) is hard to enforce and is also incompatible with Lambert, D.M. & Paterson, H.E.H., 1983. On `bridging the gap
between race and species': the isolation species concept and
hybrids or asexual species. The recognition species
an alternative. Proceedings of the Linnean Society, New South Wales,
concept (Lambert & Paterson, 1983) may seem more 107, 501^514.
appropriate for actiniid species; di¡erential aggressive Lubbock, R. & Allbutt, C., 1981. The sea anemone Actinia equina
behaviour among morphs (Brace & Reynolds, 1989) will tolerates allogenic juveniles but alters their phenotype. Nature,
have direct consequences for the breeding structure and London, 293, 474^475.
hence speciation within A. equina. Yet since aggressive Lynch, S.M., 1996. Empirical studies of allozyme based ecological
behaviour will almost certainly be coupled with the genetics of the Genus Actinia (Anthozoa: Actiniaria) and other
nematocysts, as will virtually all aspects of anemone marine invertebrates. PhD thesis, University of Liverpool.
ecology, the importance of nematocysts to coelenterate Manuel, R.L., 1988. British Anthozoa. London: Academic Press.
speciation and taxonomy is clearly paramount. Other Mariscal, R.N., 1974. Nematocysts. In Coelenterate biology: reviews
and new perspectives (ed. L. Muscatine and H.M. Lenho¡ ),
species concepts require a measure of phenotypic
pp.129^178. New York: Academic Press.
cohesion, which is clearly more problematic in Actinia Mayr, E., 1963. Animal species and evolution. Cambridge, MA:
since cnidarians are characterized by possession of a Harvard University Press.
tissue-grade construction (Shick, 1991). Although recently Minitab Inc., 1991. MINITAB reference manual, Release 8.
the usefulness of nematocysts has perhaps been ques- Rosemont, PA: Quickset Inc.
tioned (Fautin, 1988; Williams, 1996; Chintiroglou & Perrin, M.C., 1993. Aspects of the ecology and genetics of Actinia
Simsiridou, 1997), this study and others (Solë-Cava et. al., colour morphs. PhD thesis, University of Liverpool.
1985; Allcock et al., 1998) have demonstrated that the Quicke, D.L.J. & Brace, R.C., 1983. Phenotypic and genotypic
phenotypic resolution needed for recognizing cnidarian spacing within an aggregation of the anemone Actinia equina.
species can be acquired through use of the cnidom. Journal of the Marine Biological Association of the United Kingdom,
63, 493^515.
The authors would like to thank Chris Bridge for histological Quicke, D.L.J. & Brace, R.C., 1984. Evidence for the existence
and microscopical advice and Dr John Bishop for help with sta- of a third ecologically distinct morph of the anemone Actinia
tistics. Dr C. Ústman and an anonymous referee provided equina. Journal of the Marine Biological Association of the United
helpful comments on the manuscript. Kingdom, 64, 531^534.
Quicke, D.L.J., Donoghue, A.M. & Brace, R.C., 1983.
Biochemical genetic and ecological evidence that red/brown
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Journal of the Marine Biological Association of the United Kingdom (2000)