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NOTE on LUCERNARIANS, Port Erin, Isle of Man. W. I. Beaumont. 1893.

Notes on Work done at Port Erin (continued).

Mr. Beaumont also made some observations on Lucernarians, which will form the subject of a short paper by himself, to be laid before the Biological Society at an early meeting. Mr. Beaumont reports to me that he collected two species of Lucernarians under stones on the shore, between the boat jetty and the breakwater on the south side of Port Erin harbour. The one species was Lucemaria quadricornis, Muller; the other he identifies as Depastrum cyathiforme, Sars, and of this two varieties, a light reddish brown and a dark purple, occur — both being adult.

NOTE on LUCERNARIANS occurring in the neighbourhood of PORT ERIN, ISLE OF MAN. By W. I. Beaumont, EMMANUEL COLLEGE, CAMBRIDGE.


[Read May 12th, 1893.]

* The numbers in brackets refer to the List of Authorities at the end.

Three species of this interesting group have been found in the neighbourhood of Port Erin; of these, one - Depastrum cyathiforme - is fairly abundant, though apparently local in its distribution, and the examination of a number of specimens has enabled me to set at rest the divergent views which have been held with regard to it. The other two species have so far been found very sparingly: one is a well known and widely distributed form, the other will, I believe, prove to be a new species. The specimens were collected and examined while I was working at the L.M.B.C. Biological Station at Port Erin in the summer of 1892 and again in the spring of 1893.

Clark, in his Prodromus, (12)* divides the family Lucernaridae (most of the members of which were originally described under the generic name of Lucernaria) into two sub-families: -

1. Cleistocarpidae, characterised by the development of the gonads in "genital claustra" (mesogonial pouches and gastrogenital pouches of other authors) which are diverticula of the central enteric space or stomach. To this group belong Depastrum, Craterolophus, Halicyathus.

2. Eleutherocarpidae, in which mesogonial pouches or claustra are absent, the gonads being formed in the subumbral or axial wall of the 4 perradial gastral pouches (the 4 camerae of Clark). Lucernaria (as now restricted) and Haliclystus belong to this subfamily.

Between the publication of Clark's Prodromus (12) in 1863, and of Haeckel's "System der Medusen," 1879 (15) there had been found a number of species closely allied to the already known Lucernarians but having the stalk in a rudimentary condition, and thus forming a link with the ordinary free swimming medusae. For these Haeckel founded the family Tesseridae, which with the family Lucernaridae constitute his order Stauromedusae, the latter family coinciding with Clark's Lucernaridae, except that Depastrum is removed from it and placed among the Tesseridae on account of its supposed relationship to Depastrella, a species of the Tesseridae discovered by himself in the Canary Islands, which unlike the rest of the sub-family has a well developed stalk for attachment. Now although Depastrum has some points in common with Depastrella, yet in internal structure it differs very materially, and I prefer to follow Clark in retaining it among the Lucernaridae.

Sub-family - Cleistocarpidae.

Depastrum cyathiforme, (Sars.)

This species was discovered last year by myself and Mr. Gamble on the S. side of Port Erin Bay. I found it again during the recent vacation in the same locality, and also on the limestone rocks at Poyllvaaish. It is fairly abundant in both localities, attached to the under surface of stones, being apparently most plentiful above the Laminarian zone, though occurring in that zone also more sparingly, but according to my experience of larger size. It is very firmly attached, and I doubt whether it ever moves from the spot where it first attaches itself; I have never seen a detached specimen re-attach itself, indeed I do not think it is capable of doing so. In this respect it differs from most Lucernarians. This species has been the subject of no little confusion, into the discussion of which I must enter at some length.

It was first described and figured in 1846 as Lucemaria cyathiformis, by Michael Sars, in the "Fauna Littoralis Norvegias," (4). In 1858 Gosse in his " Synopsis of the British Actinias " (5) founded the genus Depastrum, based, I think, chiefly on specimens found by himself at Weymouth, which he regarded as identical with the Lucemaria cyathiformis of Sars. In 1859 Allman who had also discovered (in the Orkney Islands), what he considered the Lucemaria cyathiformis of Sars, in ignorance of Gosse's name, instituted a second one — Carduella (6) ? and in the following year gave a more detailed account of his species (7) with figures. Similarly a third generic name was founded by Milne Edwards (10) viz., Calicinaria. Gosse then pointed out (8) the claims of his name Depastrum to priority, he also gave further details with figures of the Weymouth specimens which he now elevated to specific rank as Depastrum stellifrons as it appeared to differ in certain points from Allman's species which retained the name Depastrum cyathiforme since he (Gosse) regarded it as identical with the Lucemaria cyathiformis of Sars. To this Allman replied (9) that the points of difference between Depastrum and Carduella were of generic not specific value merely, and that the name Carduella cyathiformis must stand for his own and the Norwegian form, which he also regarded as identical.

Thus the matter rested until the publication of Clark's Prodromus in 1863 (12) wherein the matter was practically cleared up, Clark having had the advantage of being able to compare specimens of the Orkney form from Allman with specimens of various ages sent to him by Sars from Norway. Now the points insisted on as distinguishing Depastrum from Carduella were these: - that while in both the tentacles are arranged in 8 groups round the margin of the disc oral surface of bell or umbrella) yet in Depastrum the tentacles are very numerous, and are arranged in each group in several rows or series, one within another, and further that they spring from the margin of the octagonal disc or from without it, but that in Carduella there are only about 5 tentacles in each group arranged in a single series and arising completely within the margin of the circular disc. Further in Carduella there is a single tentacle (primary tentacle of Haeckel, corresponding to the marginal anchors of Haliclystus) in each interval between the groups of other tentacles; this does not seem to have been observed by Gosse in Depastrum. Clark pointed out that, according to the figures and description of Sars, confirmed and extended by his own examination of the specimens sent to him by the latter, the Norwegian species when adult has the tentacles arranged in several rows (3 or 4), but that the younger individuals have at first only one row and later two. Further he pointed out that Allman was in error in asserting that the Orkney form always had its tentacles in one row only, for some of those sent to him, being presumably older than those described by Allman, had more than one row. He accordingly concluded that Carduella as defined by Allman was merely the young form of the Lucernaria cyathiformis of Sars. With regard to this point, the examination of a large series of specimens at Port Erin has amply confirmed the conclusions of Clark.

Now as to the Depastrum of Gosse, Clark inserts it provisionally as a distinct genus, but at the same time he gives some very good reasons for regarding it also as identical with Lucernaria cyathiformis, Sars. The difference in the number and arrangement of tentacles having been disposed of as far as the groups of secondary tentacles were concerned (as detailed above), there only remained the octagonal disc and absence of primary tentacles in Depastrum to separate it from Carduella.

With regard to the first point, as Clark points out, there is in Sars' figures (and in Airman's also) an octagonal area, the corners of which correspond to the intervals between the tentacular groups and whose outline is marked out by a distinct line of brown pigment; the tentacles arise from immediately without this line, and there can be little doubt that this is the octagonal margin of the disc spoken of by Gosse, but the real margin of the disc is formed by the circular muscle situated outside the origin of the tentacles, some of which are usually curved over it when the animal is fully expanded, the primary tentacles being invariably so apparently. This circular muscle is in life a delicate translucent structure and may have escaped Gosse's notice. Then as to the absence of primary tentacles in Depastrum. These in Carduella are similar in form and structure to the secondary tentacles of the groups, unlike those of Haliclystus auricula which have been modified into the so-called marginal anchors and function as adhesive organs. They are in Carduella not very conspicuous, being usually somewhat smaller than the secondary tentacles and constantly (as I have remarked above) reverted over the circular marginal muscle and closely applied to its surface. It is accordingly not unlikely these too escaped Gosse's observation, especially if his specimens had been kept long, as the tentacles seem apt to slough away when the animal is removed from its natural conditions. In addition I may mention that Depastrum, and apparently other members of this group also, is subject to much variation, especially in the number and arrangement of the tentacles; indeed of the many specimens examined at Port Erin very few had quite the typical arrangement. Sars also notices this and mentions one specimen with 7 pairs of gonads instead of the typical 4; the gonads seem more regular than the tentacles, but I have seen several individuals with 6 pairs, one of which had 13 groups of tentacles.

There can be, I think, little doubt that the Weymouth species is identical after all with Carduella and with the Lucernaria cyathiformis of Sars. And I may mention that a specimen recently found at Plymouth which has been kindly lent to me for comparison, differs in no way (externally) from the Port Erin form. That the latter is the Lucernaria cyathiformis there can hardly be a doubt, not only does it agree in external features with the figures of Sars and the description of Clark, but also as to its internal structure it is quite in accord with the account given by the latter (with one not very important exception).

This conclusion was at first accepted by Haeckel who inserts this species in his " System der Medusen" (15, p. 379) as Depastrum cyathiforme, Gosse, with Lucernaria cyathiformis and Carduella as synonyms; but in an appendix (15, p. 369) he departs from this view in consequence of having himself found on the Sutherland coast a form, agreeing with Allman's Carduella in having the tentacles in one row only, but which cannot be merely a young Depastrum since it is sexually mature, reproducing itself in that form with one row of tentacles. This, agreeing in its main structural features with the species Depastrella carduella discovered by himself in the Canary Islands (but at first called by him Carduella depastrella), Haeckel now names Depastrella allmani, giving Carduella cyathiformis Allman, as a synonym, and the Orkney Islands as one of its localities. Now whatever the Depastrella allmani from the Sutherland coast may be, there is very strong evidence that it is not the same species as the Orkney Carduella. First, since Clark had been able to compare undoubted specimens of the latter with specimens of Lucernaria cyathiformis from Sars, his opinion as to their identity must carry great weight; and secondly, Clark has given an account of the structure of this species, based on the above named specimens, the accuracy of which Haeckel acknowledges; and this account shows that these specimens differ materially in their internal organisation (more especially in the presence of mesogonial pouches) from the structure which is found in the genus Depastrella according to the type species from the Canary Islands - Depastrella carduella, of which he gives figures. He gives no figures of Depastrella allmani but describes it as having practically the same internal structure as Depastrella carduella, consequently if Haeckel's description of the Sutherland species be correct, it is obvious that its internal anatomy differs considerably from that of Carduella, however similar they may be in external features, and we may safely conclude that Carduella cyathiformis, Depastrum cyathiformis and Lucernaria cyathiformis are one and the same species.

Sub-family - Eleutherocarpidae.

Haliclystus auricula, (Rathke).

A small Haliclystus was found near Port St. Mary by M. Chopin, in 1891, on a lobster pot I believe, and is now in the Zoological Museum at University College, Liverpool. I have seen the specimen and, as far as I can judge, it is H. auricula, but it may possibly be H. octoradiata (Lamarck, 3). The shape and size of the marginal anchors seem to be those typical of H. auricula; the characters of the gonads I have not been able to make out satisfactorily.

With regard to these two species we again meet with confusion, but since I have seen specimens from Plymouth and Jersey which have all the distinguishing characters of the Haliclystus auricula described by Clark in his Prodromus (12) and again with great detail in a fully illustrated monograph in the Smithsonian Contributions, 1878, (14) I prefer to follow Clark in identifying his species with the European Lucemaria auricula, Rathke (2). Haeckel on the other hand considers it probable that Haliclystus auricula, Clark, is confined to the American side of the Atlantic, and identifies L. auricula, Rathke, with L. octoradiata, Lamk., which is described as a second European species by Clark. Rathke' s figures as far as they go support Clark's view.

It may be noted further that Lucemaria auricula, Rathke, was confused by a number of the earlier writers with L. auricula, Fabricius (1), one of the Cleistocarpidae [Manania auricula (Fab.) Clark, Halicyathus lagena (Muller) Haeckel].

Haliclystus, sp.

The third species of Lucernarian found at Port Erin differs materially from any species previously described as far as I know. The members of this family seem particularly liable to abnormalities affecting more especially the number and arrangement of the tentacles, as noticed under Depastrum, and I am loth to establish a new species on the scanty material at present available. After some consideration, I refer this species provisionally to the genus Haliclystus, for though it presents considerable divergence from the 3 species of that genus which have been already described, yet its structure, so far as I have been able to make it out, is in accordance with the generic definition both of Clark and Haeckel except as regards one point which is not I think of very great importance.

Three examples of this form were found last year: they were attached to the undersides of stones on the S. side of Port Erin Bay, where Depastrum also occurs. During the recent vacation I made a most careful search in the same locality but failed to find a single specimen. On the first occasion in my ignorance of the "points" of a Lucernarian, I did not observe in the living animal the presence of primary tentacles, but in one of the three specimens these are now plainly enough to be seen, but in the others which are smaller and were not preserved in an expanded condition I have been unable to ascertain whether these important organs are present or not. Now in Haliclystus auricula additional tufts of tentacles seem not uncommon, and accordingly, until I have seen more specimens I hesitate to conclude that what certainly appear to be genuine primary tentacles (retaining the original tentacular structure instead of being modified into marginal anchors as in H. auricula) are really normal structures.

If on the other hand these primary tentacles are merely individual abnormalities, and if I am right in my interpretation of its internal structure, then this species must be relegated to the genus Lucernaria (as at present restricted). In the approximation of its arms in pairs it approaches L. quadricornis, Muller, but its complicated gonads differ from those of that species. I at first took the Port Erin specimens to be small and somewhat abnormally shaped L. quadricornis, before I had discovered the primary tentacles ; they are referred to under this name in the L.M.B.C. Annual Report, 1892 (p. 33). I now append a description of the species: -

Haliclystus sp. (? n. sp.)

Umbrella somewhat conical, passing gradually into the stalk without any marked distinction. Subumbrella cavity very shallow. Stalk more or less round in transverse section with. 4 longitudinal grooves marking the position of the 4 interradial muscles, decreasing in diameter from its junction with the umbrella and then expanding again at its aboral end into the disc for Attachment. 1-chambered. (The other species of Haliclystus have the stalk 4 chambered.)

Arms 8, but so closely united in pairs, that there appear to be only 4, separated by 4 very wide perradial " bays," the interradial intervals being practically obliterated. Primary tentacles 8. The 4 perradial ones are smaller than the secondary tentacles but similar in form, they stand out horizontally from just within the margin of the umbrella. The 4 interradial ones spring from the point of junction of the paired arms, between the tufts of secondary tentacles and resemble the latter.

Secondary tentacles are grouped in tufts on the ends of the arms, about 7 on each, but in consequence of the fusion of the arms in pairs, there appear to be only 4 tufts of tentacles, each of about 15; really each consists of the tufts belonging to a pair of arms together with what I regard as the intervening interradial primary tentacle. The tentacles are somewhat club shaped, the head not being distinct from the stalk.

Gonads. These consist of numerous saccules forming 4 adradial bands in the endoderm lining the subumbral wall of the 4 perradial gastral pouches. These walls are very much folded, and in transverse sections the whole gastral cavity appears almost packed with the genital saccules.

Size. Height including stalk - about 7 mm. Width of umbrella - about 3 mm.

Colour. A rather dull pale yellow.


1. - 1780. 0. Fabricius - Fauna Groenlandica, p. 341.
2. - 1806. Rathke - Muller's Zool. Danica, vol. IV.
3. - 1816. Lamarck - Hist. Nat. anim. II.
4. - 1846. M. Sars - Fauna Littoral. Norweg. Fasc. I.
5. - 1858. Gosse - Ann. & Mag. Nat. Hist. vol. I, p. 419.
6. - 1859. Allman - Report Brit. Assoc, Aberdeen.
7. - 1860. " - Trans. Micros. Soc. VIII, p. 125 & PL
8. - " Gosse - Ann. & Mag. Nat. Hist. V, p. 481:
9. - " Allman - do. do. VI, p. 41.
10. - " Milne Edwards - Hist, des Corall. Ill, p. 459.
11. - 1862. Keferstein - Zeitschr. fur wiss. Zool. XII, p. 24.
12. - 1863. Clark. - Prod. Lucernar. Jour. Boston Soc. N.H.
13. - 1877. Taschenberg - Halle Zeits. Naturw, Bd.49, p.94.
14. - 1878. Clark - Lucernar. Monog. Smithsonian Contrib.
15. - 1879. Haeckel - System der Medusen.

Notes on this article by David Fenwick -

Haliclystus sp. (? n. sp.) here possibly = Lucernaria quadricornis.
Ref: Annual Report Millport Station. Dr. Russell. 1904.
Ref: XXV. Notes on Lucernaria quadricornis, Muller, and related species. By Richard Elmhirst, F.L.S., Superintendent of the Millport Marine Biological Station. p.224. The Annals and Magazine of Natural History. Vol. X -Ninth Series. 1922.

On researching Depastrum cyathiforme at Port Erin I have discovered images in the form of postcards that were taken a year or two after the discovery of the species at Port Erin. The images are important in that they display the habitat when the species was originally discovered at the location. Comparisons can now be made in relation to changes in habitat should the species not be found at this relatively small area of shore in the future. The image of Port Erin Breakwater would seem to be a particularly important image.

Port Erin breakwater 1894
Port Erin slipway / bay 1893
Port Erin slipway / Bradda Head 1895

Here I must thank The Francis Frith Collection for allowing the use of the above images of Port Erin. All the images here are copyright to The Francis Frith Collection -

Also see article by author -
Notes on stauromedusae at Port Erin and other
reported Isle of Man sites. David Fenwick. 2014.
A search for Depastrum cyathiforme.



Lucernarians Port Erin Isle of Man 1893 Liverpool Biological Society Image