Art. XXXVI.—A Review of the New Zealand Actiniaria known to Science, together with a Description of Twelve New Species.

By F. G. A. Stuckey, M.A., Wellington.

New Zealand waters, a circumstance which seems to point to the conclusion that there are in this region no abyssal or pelagic forms.

Methods.

Killing.—Several methods were used.

(1.) Electrocution with a strong current (14 amperes) was tried, but the results were fair only. I am of opinion that this method is not good for histological purposes, on account of the destruction of tissue brought about by electrolysis.

(2.) In another method the animals, after being expanded in a minimum of sea-water in small glass vessels, were dropped (vessels and all) into a boiling saturated solution of perchloride of mercury.

(3.) The animals were stupefied with magnesium-sulphate in sea-water (Tullberg's method), and then either chloroformed or dropped into boiling perchloride of mercury.

The last two methods yielded the best results.

Fixing, Imbedding, and Cutting.—All specimens were fixed by immersion for a longer or shorter period, according to size, in perchloride of mercury. The larger specimens were filled with the fixing fluid through the mouth by means of a glass syringe immediately after death. The mercury having been removed with iodine, the specimens were dehydrated in the ordinary way, and finally imbedded in paraffin. Sections were cut with a Cambridge rocking microtome, and, in the case of some of the large species, with a sliding microtome.

Staining.—Mass staining with borax-carmine yielded results which were satisfactory, but not so good as those obtained by staining on the cover-slip. Saffranin and methylin blue yielded good results, the nuclei staining well. Fuchsin was found a fine stain for nuclei and nematocysts. Kleinenberg's hæmatoxylin gave good definition of mesoglœa. The best results were got with Erlich's hæmatoxylin. This was used full strength (Squire's formula), or diluted with water in various proportions. All yielded excellent results. After staining on the cover-slip the sections were washed in “tap” water, in some cases being immersed in the water for several days. I am convinced that the best results with this stain are obtained by using it full strength, and then washing out for four or five days in water.

List Of Species Described.
Grade I. Paramera.
Order Edwardsiidea.
Genus Edwardsia (Quatrefages).

1. Edwardsia tricolor (Farquhar), nom. nov.

2. Edwardsia ignota, sp. nov.

Grade II. Cryptoparamera
Order Actiniidea.
Suborder Malacactiniæ.
Group Hexactiniæ.
Fam. Antheadæ.
Genus Actinia (Browne).

Genus Anemonia (Risso).

Fam. Sagartidæ.
Genus Sagartia (Gosse).

Genus Thoe (Wright).

Fam. Paractidæ.
Genus Paractis (Andres).

Genus Tealidium (Hertwig).

Fam. Corallimorphidæ.
Genus Corynactis (Allman).

Fam. Phyllactidæ.
Genus Cradactis (McMurrich).

Fam. Bunodidæ.
Genus Bunodes (Gosse).

Genus Leiotealia (Hertwig).

Fam. Alicidæ.
Genus Phlyctenactis (gen. nov.).

Genus (?).

General Results.

One new genus has been erected and one abandoned genus re-established. Twelve new species have been described, four species have been placed in other genera than those in which they were placed by their original describers, and one species has received a new specific name. For all these changes reasons are given in the text.

In general, the anatomy of all species is normal. Actinia tenebrosa shows great variation in the number and arrangement of its mesenteries.

The histology of the Actiniaria has been so fully worked out by the Hertwigs that little remains to be said on the subject. The structure of ectoderm and endoderm showed no departure from the normal type, with the exception of the ectodermal pits found in Leiotealia thompsoni. The mesoglœa, however, showed some considerable variation. In the majority of cases it presented the usual features—namely, “a homogeneous matrix in which fibres are imbedded.” In other cases lacunæ were found to a greater or less extent, especially in the tentacles of Phlyctenactis retifera, where the mesoglœa was found to be highly reticular. There are fine sphincter muscles in Leiotealia thompsoni and Cradactis plicatus, more especially in the former.

A peculiar arrangement of the “trefoil” of the mesenterial filament was noted in Actinia tenebrosa and Corynactis haddoni.

With regard to gonads, it is a common occurrence for observers to fail to find them in their specimens, either because the animals are immature, or because the reproductive cells are found at certain seasons only. In the papers enumerated hereunder, describing British anemones and those collected in various expeditions, out of 163 species described gonads were found in sixty-nine only. Of the twenty-three examined by me, and described in this paper, gonads were located in nine. In two cases the animals were found to be viviparous: one had ova in brood-pouches; another was observed to divide in a manner for which I suggest the name “lateral fission.”

Books And Papers Referred To In This Paper.

Dixon, A. F. and G. Y. “Notes on Bunodes thallia, Bunodes verrucosa, and Tealia crassicornis.” Proc. Roy. Dub. Soc., vol. vi, n.s., pt. 6. 1889.

Duerden, J. E. “The Actiniarian Family Aliciidæ.” Ann. and Mag. Nat. Hist., 6th series, No. 115. 1897.

Farquhar, H. “Preliminary Account of some New Zealand Actinians.” Linn. Soc. Journ. Zool., vol. xxvi. 1898.

Gosse, P. H. “Actinologia Britannica.” 1860.

Haddon, A. C. “Revision of the British Actiniæ,” Part I. Trans. Roy. Dub. Soc., vol. iv, ser. ii, No. v. 1889.

Haddon, A. C., and Shackleton, A. M. “Revision of the British Actiniæ,” Part II. Trans. Roy. Dub. Soc., vol. iv, ser. ii, No. xii. 1891.

Haddon, A. C., and Shackleton, A. M. “Report on Zoological Collections made in Torres Strait.” Trans. Roy. Dub. Soc., vol. iv, ser. ii, No. xiii, 1891.

Hertwig, R. “Report on the Actiniaria dredged by H.M.S. “Challenger” during the Years 1873–76.” “Challenger” Report, vol. vi. 1882.

Hertwig, R. Supplement to Report, &c. “Challenger” Report, vol. xxvi. 1888.

Hickson, S. J. “Cambridge Natural History,” vol. i.

Hutton, F. W. “The Sea Anemones of New Zealand.” Trans. N.Z. Inst., vol. xi. 1878.

McMurrich, J. P. “Report on the Actinian Collections made by the U.S. Fish Commission steamer ‘Albatross’ during the Winter of 1887–88.” Proc. U.S. Nat. Mus., vol. xvi. 1893.

Moseley, H. N. “On New Forms of Actiniaria dredged in the Deep Sea.” Trans. Linn. Soc. Lond., Zoology, ser. ii, vol. i, pt. 5. 1877.

Parker, G. H. “Longitudinal Fission in Metridium margenatum.” Bull. Mus. Har. Coll., vol. xxxv, No. 3. 1899.

Torrey, H. B. “Anemones, with Discussion of Variation in Metridium.” Proc. Wash. Acad. Sci., vol. iv. 1902.

Description Of Species.
Genus Edwardsia (Quatrefages).

1. Edwardsia tricolor (nom. nov.), Farquhar, 1898.

This species (fig. 1) was described by Farquhar under the name Edwardsia elegans. I have changed the specific name to tricolor, the name elegans having been in use for another species since 1849. Mr. Farquhar has fully described the external appearance of this species, so I shall only briefly state my own observations. The body is brown, wrinkled, and covered with a rough cuticle, which is thin at the upper end. It is marked longitudinally by 8 double brown lines. There is a strong constriction between body and capitulum. The lower part of the capitulum is bulbous, and is divided into 8 segments by double white lines, continuous with the brown lines of the scapus (fig. 2). In colour each segment is brown, with a large white spot. Above this is a narrower white portion. The tentacles

Fig. 1.

Fig. 2.

Fam. Antheadæ.

Genus Anemonia (Risso).

“Antheadæ without acrorhagi.”

4. Anemonia olivacea (Hutton).

This species was described by Hutton as Anthea olivacea. It is referred to as Anemonia olivacea by Farquhar, who is probably right in so naming it, the older generic name having precedence.

Pedal-disc.—Ectoderm and endoderm are thin. The mesoglœa is thick and fibrous, the fibres being arranged in wavy branching lines. Numerous nuclei appear in the mesoglœa.

Column.—Smooth. Colour, olive-green. There are light longitudinal lines near the base. The ectoderm and endoderm present the usual characters, and the mesoglœa resembles that of the pedal-disc. It varies in thickness, thick and thin places alternating in my sections.

Tentacles.—About 50 in number, arranged in four cycles. In colour they are pellucid-green, of the same shade as the body. There is a slight constriction near the end, and the tip is darker in colour. They are conical and slender, and their length is equal to the diameter of the oral disc. They

Fig 5.

are not completely retractile. The ectoderm and endoderm are thick and the mesoglœa thin. The musculature is very weak.

Oral Disc.—The colour is brownish-green. The mouth is set on a slight elevation, from which radiate yellowish lines. The structure of the disc closely resembles that of the tentacles.

Stomodæum.—This is pinkish-white in colour, and there are 2 siphonoglyphs. In cross-section the stomodæum is seen to be strongly folded.

Sphincter.—Weak, endodermal. Its shape in cross-section is semi-ovate, the flattened side closely applied to the body-wall.

Mesenteries.—There are 24 pairs, every second pair being perfect. Two pairs are directives. The musculature is strong, especially on the perfect mesenteries. The muscle-banners extend almost from wall to stomodæum.

Gonads.—None seen in my specimens.

Size.—Diameter, about 20 mm.; height, rather less.

Habits.—This species is found in rock-pools, often inhabiting quite small “pot-holes” on the upper part of the rocks. Its colour is strongly protective.

Distribution.—Of the genus—English Channel, Atlantic Mediterranean, Patagonia, Torres Strait, Pichelingue Bay (Delage and Herouard); also New Zealand. Of the species—Dunedin (Hutton), Cook Strait.

Fam. Sagartidæ.

“Hexactiniæ furnished with acontia.”

In thus defining the Sagartidæ I have followed what seems to be the general tendency of recent authors. Hadden has somewhat fully discussed the classification of sagartians, though he arrived at no very definite conclusion. In using the term “sagartians” he says, “By the latter term I include all those Actiniæ which possess acontiæ.” McMurrich (“Albatross” Report, 1893) says, “It seems at present convenient to associate all Actiniæ with acontia in a single family.” Most authors, however, in defining the family still include the provision “sphincter muscle mesoglœal.” I should like to point out, on the authority of G. C. Bourne, that Actinoloba, a sagartian genus, has an endodermal sphincter.

Subfam. Sagartinæ

“Sagartidæ with naked ectoderm. Cinclides usually present.”

Genus Sagartia (Gosse).

“Sagartidæ with smooth column; no verrucæ; no special arrangement of cinclides; margin tentaculate; tentacles concealed in contraction, the sphincter being fairly strong” (McMurrich).

5. Sagartia albocincta (nom. nov.), Hutton.

A description of this species is published at p. 372 of this volume.

Distribution.—Of the genus—Arctic Ocean, North Sea, English Channel, Atlantic, Mediterranean, Cape Horn, Pacific coast of South America, East Indies, Australia (Delage and Herouard); also New Zealand. Of the species—Dunedin (Hutton), Plimmerton (H. B. Kirk), Cook Strait.

6. Sagartia nutrix (sp. nov.)

This is the brown sagartian mentioned by Farquhar as probably forming the food of the butter-fish. It is found only on Lessonia, where it establishes itself in colonies.

7. Sagartia vagrans (sp. nov.).

Pedal-disc.—The ectoderm is thick, and presents no unusual feature. The mesoglœa is much “vacuolated,” and the endoderm is thin.

Column.—The colour is dirty-white and olive-brown in alternate longitudinal lines, or the whole may be dirty-white, or grey, or even pink. There is a circular muscle in the wall throughout its entire length carried on regular folds of the mesoglœa.

Tentacles.—These are very fine and threadlike. Their colour is salmon-pink, though in some specimens they are white. They are of different lengths in the different cycles. There are several cycles, but the tentacles are so numerous and crowded that it is impossible to make out how many cycles there are. Judging from cross-sections I should say there are four cycles. In structure the tentacles resemble those of the last species (S. nutrix), but there is a specially well-developed nervous layer.

Oral Disc.—Colour olive-brown generally, but there is considerable variation. The structure is identical with that of the tentacles, and closely resembling the same part of S. nutrix.

Stomodœum.—The colour is a rich pink, with darker-coloured red longitudinal lines. The inner edge of the mouth is also red. The stomodæum is freely everted, when the red lines, together with the edge of the mouth, form a rosette-like design on the disc. There are 2 siphonoglyphs.

Sphincter Muscle.—This is mesoglœal in character. There is only a slight thickening of the mesoglœa under the bases of the tentacles, but the whole body-wall has the character (more or less) of a diffuse mesoglœal sphincter. In contraction the tentacles and oral disc are much infolded, the wall completely overarching them.

Mesenteries.—There are 48 pairs, 12 pairs being perfect, the arrangement of the cycles being 6 + 6 + 12 + 24 = 48. Two pairs are directives. (Plate XXVII, fig. 2.)

Gonads.—None were present in my specimens.

Acontia.—These are extruded through lens-shaped cinclides, which are invisible except at the time of discharge. These cinclides appear to be on the white lines only, and to be limited to a zone encircling the middle third of the body.

Locality.—Professor H. B. Kirk has brought me specimens from Plimmerton. I myself have found the species in large numbers on the breastwork of the Thorndon Esplanade, Wellington Harbour, and on the piles of the baths. This anemone adheres so strongly that it is impossible to remove it without damage. It is attached also to shells of mussels, and this makes it possible to obtain good specimens.

Habits, &c.—This animal lives well in captivity. Some that I kept in an aquarium wandered about at will.

Size.—My largest specimens were 40 mm. high and 20 mm. in diameter, but I am told there are larger ones on the piles of the Queen's Wharf, Wellington.

Affinities.—Probably closely allied to S. albocincta.

Distribution.—Of the genus—As above. Of the species—Plimmerton (H. B. Kirk), Wellington Harbour.

Genus Thoe (Wright).

On page 122 of the “Actinologia Britannica,” Gosse, after referring to the genus Sagartia, says, “A group less typical I consider to be formed

visible cinclides except when the acontia are ejected. The colour is white.

Sphincter Muscle.—There is a spindle-shaped thickening of the mesogloea in the upper part of the column, causing a slight bulging of the wall. In the sphincter the muscle spaces are very close together, but otherwise this muscle is of the usual sagartian type.

Tentacles.—There are 96, in four cycles, 12 + 12 + 24 + 48 = 96. In colour they are opaque-white, like the column; in shape, conical and tapering. The length is rather more than half the diameter of the disc. At the base of each tentacle is an incomplete ring of yellow pigment, the open part directed inwards and the outer circumferences of the rings close together, so as to make an almost continuous ring of yellow round the top of the column under the tentacles (fig. 7). Nematocysts are numerous in the tentacles. The ectoderm and endoderm are relatively thick, the mesoglœa being reduced to a mere streak. The muscle-bearing processes on both sides are few in number. The nerve-layer is compact, and appears almost as a line in cross-sections. There are numerous large irregular spaces in the ectoderm.

Oral Disc.—Colour white, as in the tentacles and column. Mouth is set on a peristome. The stomodæum is white, and there are 2 siphonoglyphs.

Mesenteries. — Owing to the difficulty of killing and fixing this species I was unable to determine accurately the number and arrangement of the mesenteries.

Acontia: — These are emitted reluctantly, some through the mouth, others from cinclides which are placed on the column just under the yellow line at the bases of the tentacles. The acontia are long, and contain very large numbers of nematocysts, together with many nerve-cells. The processes of the nerve-cells run between the closely packed nematocysts (fig. 8).

Habits.—This species lives in the chinks among the “roots” of Lessonia. It is impatient of light.

Dimensions.—In full expansion it is 12 mm. high and 10 mm. in diameter. The pedal-disc is rather wider than the column.

Distribution.—Of the genus—British Isles, New Zealand. Of the species—Island Bay.

Genus Halcampactis (Farquhar).

This genus, which I consider to be closely allied to Chondractinia or to Paraphellia, will probably become the type for a new subfamily of sagartiads. By its author it is ascribed to the family Halcampidœ. At the same time he considers it as constituting a link between that family and the Sagartidœ. I have not been able to find Farquhar's Halcampactis mirabilis, though I made one search in company with Mr. Farquhar, and have been out several times myself and with Professor Kirk, the original discoverer of the animal. I have, however, seen the original sections of H. mirabilis, but until I have been able to rediscover the species described by Farquhar I prefer to simply place the genus provisionally in the Sagartidœ. While searching for Halcampactis mirabilis I found three specimens of the species to be now described, a species which probably belongs to the same genus.

Fig. 7.

Fig. 8.

9. Halcampactis dubia (sp. nov.).

The body is pink in colour and vermiform in shape. The epidermis is smooth, with very minute papillæ on the upper portion. The body is divisible into capitulum, scapus, and physa, the division between the last two being slight. A red line marks the division between scapus and capitulum. The capitulum, which is capable of being entirely invected, bears 24 pink tentacles, in two cycles. The scapus is marked by 6 double longitudinal white lines. The physa is adherent and flattened. It is thin, the filaments and acontia being plainly visible through it. The animal does not emit the acontia, but if a slit be made in the body they stream out readily. They are of the usual character, and the nematocysts are closely packed. I have not yet worked out the anatomy and histology of this species.

Dimensions.—The largest specimen was 25 mm. long and about 3 mm. in diameter.

Distribution.—Of the genus—The coast near Wellington. Of the species—Island Bay.

Fam. ParactidÆ.

I adopt Hertwig's definition as modified by McMurrich: “Actiniœ usually with numerous perfect mesenteries; circular muscle strong, imbedded in the mesoglœa; acontia wanting.”

Genus Paractis (Andres).

“Paractidœ with smooth body-wall, without papillæ or marginal spherules; tentacles slender, and not exceptionally numerous; nearly equal in length and strength; margin not lobed; sphincter widening somewhat abruptly in its upper part, and occupying, near the margin, nearly the entire thickness of the mesoglœa.”

10. Paractis ferax (sp. nov.).

Pedal-disc.—Adherent to the under-sides of stones, or to rocks, in dark places. The chief histological feature is the thick mesoglœa, which is indeed a noticeable feature of the whole animal. The fibres are wavy, and there are many lacunæ. The ectoderm is of the usual type, about half as thick as the mesoglœa. The endoderm is thin.

Column.—It is generally cylindrical in shape, but is occasionally constricted at various parts, giving the animal a vase-shaped contour. The column bears, especially in contraction, fine transverse wrinkles. Its height can be varied very considerably, but is generally equal to two or three times the diameter. The colour is milky white. In contraction the animal is drawn into a hard rounded ball of the size of a small marble. The ectoderm and endoderm are of the normal type. The mesoglœa is thick, about equal to the ectoderm, but at intervals it becomes thin, as shown in fig. 9. This feature seems to be characteristic. In other places, numerous enough for the structure to be considered characteristic, the mesoglœa forks, sending out a process which is lined on both sides by ectoderm, and which lies close to the body-wall (fig. 10).

Fig. 9.

Fig 10.

Distribution.—Of the genus—Mediterranean, Red Sea, Indian Ocean, Atlantic, Tierra del Fuego, New Zealand, Pacific, China Sea. Of the species—Island Bay, Ohiro Bay.

Genus Tealidium (Hertwig).

Paractidœ having the tentacles placed in several rows of uniform size in the same row, and having the body-wall covered with fine papillæ. All mesenteries perfect and gonophoric except the directives. (Hertwig's definition modified.)

This genus was erected by Hertwig for the reception of a form found in the “Challenger” material, a form which agreed with Paractis except in the warty character of the body-wall. In Hertwig's species (Tealidium cingulatum) the wall bulges outwards, forming a girdle below the tentacles. This he attributes to the great development of the sphincter. In the species described below the same thing is seen, though I have no direct evidence that it arises from the cause ascribed by Hertwig. Probably this feature may come to be regarded as of generic value.

11. Tealidium cinctum (sp. nov.).

Pedal-disc.—Adherent in cleft of rock, making the animal difficult to remove. The mesoglœa is fibrous, and contains numerous lacunæ.

Column.—The upper part is brownish in colour, the lower part dirty-white. The upper part is covered with verrucæ in vertical rows; small shells, &c., are attached to these verrucæ. The lower part, which is imbedded in a cleft of the rock, is without the verrucæ, and is channelled by fine furrows. The verrucæ act as suckers, by which the animal covers itself with bits of shell and other débris. In full expansion the column bulges outwards, forming a circular swelling just under the bases of the tentacles. The ectoderm is somewhat irregular, and there appear here and there spaces between the cells. The nerve-layer is feebly developed. The mesoglœa is well developed, and contains small lacunæ. It runs into the ectoderm in the form of conical papillæ.

Tentacles.—These are 48 in number, apparently in four cycles. Length, about 16 mm. They are pellucid, with a mauve-pink shade. There are white transverse markings on the inner sides. The ectoderm is of the same irregular character as that of the column, but the intercellular spaces are smaller. The nervous layer is well developed, being several cells deep. There is a fair development of the ectodermal muscles, but the endodermal musculature is weak. The lumen of each tentacle is filled with what appears to be hypertrophied endoderm.

Oral Disc.—The colour is pale brown, with a ring of green round the mouth, and a broken ring of yellow round the green. The histological features resemble those of the tentacles.

Sphincter Muscle.—This is mesoglœal and diffuse, extending through the whole wall of the column. There is, however, a decided thickening under the edge of the disc, and this probably constitutes the true sphincter. Hertwig describes a similar sphincter in Antholoba reticulata.

Mesenteries.—There are 24 pairs, 2 pairs being directives. All reach the stomodæum, and are about equal in development. All are gonophoric except the directives. I have made this a generic feature, since Tealidium cingulatum (Hertwig), the only other known species, has all its mesenteries perfect and gonophoric (? directives). The musculature is well developed. (Plate XXIV, fig. 2.)

Fam. Corallimorphidæ.

Fam. PhyllactidÆ.

Fam. BunodidÆ.

and taper to a fine point. In colour they are olive-brown, with from 2 to 6 white spots on the inner side. They are completely retractile, and very sensitive. The nerve-layer is distinct, and the ectodermal muscles are strong. The endomermal muscles are weak. The lumen is filled with zooxanthellæ.

Oral Disc.—Flat; peristome raised. Colour olive-brown. Stomodæum white, often everted. The histological features resemble those of the tentacles.

Sphincter Muscle.—Endodermal. Closely resembles that of B. aureoradiata.

Mesenteries.—Closely resemble those of B. aureoradiata.

Gonads.—I saw none in my preparations.

Dimensions.—My largest specimens were 15 mm. high, and about the same in diameter, but I am told they grow much larger.

Distribution.—Of the genus—As above. Of the species—Dunedin, Wellington.

Genus Leiotealia (Hertwig).

20. Leiotealia thompsoni (nom. nov.), Coughtrey.

This species is described in this volume in a separate paper.

Distribution.—Of the genus—Magellan Strait, Chili, Kerguelen Island, Arctic Ocean (Delage and Herouard); also New Zealand. Of the species—Dunedin (Coughtrey); Cook Strait, from Plimmerton to Seatoun.

Fam. AliciidÆ.

“Hexactiniœ with large, flat, contractile base. Tentacles simple, subulate, entacmæous. Column with simple or compound hollow outgrowths or vesicles over more or less of its surface, arranged mostly in vertical rows. No cinclides. Sphincter muscle endodermal and diffuse, variable in amount of development. Perfect mesenteries few or numerous. No acontia.” (Duerden.)

Up to the present time this family contains four genera — Alicia, Cystiactis, Bunodeopsis, and Thaumactis. I have several specimens of a form taken at Island Bay and one specimen taken at Stewart Island. Both these forms certainly belong to the Aliciidœ, but do not seem to be referable to any one of the hitherto recognised genera. They differ from Alicia in their larger number of perfect mesenteries; from Cystiactis in the large number of short tentacles; from Bunodeopsis in the disposal of the phlyctenia over the whole of the column; from Thaumactis in the lenticular form, and absence of a pedal-disc in the latter genus. For these reasons I propose for these two species a new genus, Phlyctenactis. The differences between this genus and the other four are shown by the following table:—

Table of the Genera of the Family Aliciidæ.

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Genus Phlyctenactis (gen. nov.).