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Volume 63, 1934
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The High-Tidal Mollusca of Rangitoto Island, Auckland; with Descriptions of a New Genus and Two New Species.

[Received by the Editor, 12th August, 1932; issued separately, 28th February, 1933.]

The writer is a member of an Auckland committee that in 1925 received a New Zealand Institute Research Grant for the purpose of making an ecological survey of the Waitemata Harbour. The present paper is based upon a few observations made and some specimens collected in the course of this investigation. Of special interest is the discovery of a new genus of the Ellobiidae, described in this paper, which, so far as is known, is restricted to the shores of Rangitoto Island.

A full report on the animal communities of the harbour is in preparation, and will be published at a later date.

The thanks of the writer are due to Miss L. M. Cranwell for the identifications of the seaweeds and maritime plants mentioned herein.

The foreshore along the southern coast of Rangitoto Island is formed of extensive lava-flows, many of which are broken up into loose blocks, and these are strewn upon a substratum of stiff mud with occasional patches of Zostera and small areas of comminuted shell.

The typical inter-tidal plant cover on this southern coast ranges from a low-tidal Carpophyllum brown-algal community, through a mid-tidal community of Corallina-Hormosira (which thins out above amongst a zone of rock-oysters, Ostrea glomerata), to a moderately broad high-tidal zone, which is dominated by the glass-wort, Salicornia australis Soland. Together with the Salicornia two other maritime plants are moderately common. They are Suadia maritima Dum. and Samolus repens Pers. It is the fauna of this Salicornia zone which extends from about high-water neap to a little above high-water spring tide levels, that is made the subject of this paper.

In the uppermost part of the high-tidal zone the Salicornia thins out, and is replaced by scattered clumps of tussock, Stipa teretifolia Stued.

This marks the extreme landward limit of marine mollusca at this locality, one species only, the common periwinkle, Melarhaphe oliveri, being known to occur. Of all the high-tidal molluscs this periwinkle has the greatest vertical range, being found throughout the Salicornia belt and for a little distance both above and below it.

Although the glass-wort does not directly support the animals associated with it, cover is provided for a small inconspicuous dark green alga, Rhizoclonium hookeri Keietz, which grows upon the shaded mud surface, and is eaten in a living state by two species of mollusca, the ever-present pulmonate, Ophicardelus costellaris, and

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a minute taenioglossid, Assiminea vulgaris. Associated with these molluscs are a few amphipods, isopods, and the crabs Hemigrapsus sexdentatus (M. Ed.) and Heterozius rotundifrons (M. Ed.).

In depressions among the rocks, masses of decaying wrack accumulate, made up for the most part of remnants of low-tidal brown algae Carpophyllum maschalocarpum (Turn.) Grev. and C. plumosum (A. Rich.) J. Ag., with intermingled Zostera nana.

This accumulation of decaying vegetative matter supports large numbers of half to full-grown individuals of the pulmonate mollusc Marinula filholi, while the decaying Zostera in particular is favoured by numerous Assiminea vulgaris.

Beneath the decaying algae are loose stones which rest upon or are partly imbedded in mud, and upon these occur large numbers of the young of Marinula filholi, together with Assiminea vulgaris and a second taenioglossid, Suterilla neozelanica, while underneath in contact with the mud a further three species are to be found. Two of these (one a new genus of the pulmonates) are organic-mud feeders which receive their sustenance by retrieving from the mud minute particles of decaying algae and animal detritus. The third species, a new Kellia, occurs in vast numbers, but the two pulmonates are by no means common.

At Rangitoto the food of Melarhaphe consists for the most part of an obscure dark greenish, almost black lichen, which is rendered quite inconspicuous owing to the dark-coloured lava upon which it grows.

In other areas Melarhaphe is often found in numbers feeding upon the sea-lettuce, Ulva, and also upon Enteromorpha, a small alga which forms a pale green coating upon high-tidal rocks wherever fresh-water seepage occurs. Of all the high-tidal molluscs only one, Melarhaphe, is able to withstand long periods of desiccation, and on sunny days the stones upon which it occurs often become quite hot.

Contact with the sea at the high-tidal levels is very brief, so the remaining seven species owe their existence primarily to the moisture retained in the mud shaded by the Salicornia and to the dampness caused by the more or less permanent accumulation of decaying wrack, which is retained owing to the roughness of the rocks. Very little of the debris that once touches the shore at Rangitoto is ever washed away again by succeeding tides, so the conditions are rendered more or less permanent.

This stability of the high-tidal conditions, which means an ever-present abundance of food, can be taken as the main reason why the Rangitoto high-tidal fauna is so much richer than that of any other high-tidal area so far studied in New Zealand.

The Salicornia zone at Hobson Bay, Auckland Harbour, in sharp contrast to that of Rangitoto Island, has only two species of mollusca, Ophicardelus costellaris and Melarhaphe oliveri, the species Marinula filholi, Suterilla neozelanica, Assiminea vulgaris, Kellia maoria, and Rangitotoa insularis all being absent. However, this paucity of species may be due to some extent to the fact that the salinity of the waters of Hobson Bay is known to be slightly lower than at Rangitoto.

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Summary of the Feeding Habits of the High-Tidal Mollusca.

1.

Ophicardelus costellaris feeds almost exclusively upon the living Rhizoclonium hookeri which grows on the mud under the shelter of Salicornia australis, but it has been observed also to eat decaying Hormosira.

2.

Melarhaphe oliveri prefers the open, and will endure much desiccation by exposure to strong sunlight. At Rangitoto it feeds upon a small dark-greenish lichen, but it is also known to eat Ulva and Enteromorpha.

3.

Assiminea vulgaris thrives equally well upon either living Rhizoclonium or upon decaying algae, especially Zostera.

4.

Marinula filholi and

5.

Suterilla neozelanica seem to live exclusively upon decaying algae, and at Rangitoto are found on and under stones beneath masses of decaying wrack.

6.

Kellia maoria occurs on the under sides of stones, and no doubt obtains its staple food from the organic-mud in which the stones are immersed.

7.

Rangitotoa insularis and

8.

Leuconopsis obsoleta are organic-mud feeders which occur sporadically on the under sides of stones that are in contact with mud.

In brackish situations Potamopyrgus antipodum is often found living upon the mud in association with mangroves. Caecum digitulum, abundant elsewhere, has not yet been observed at Rangitoto; it seems to require smooth, soft rock, of which there is none at this locality.

Assimineidae.

Subfamily Assimineinae.
Genus Assiminea (Leach) Fleming 1828.

(= Syncera Gray 1821, Med. Repos., p. 239. A nomen nudum, according to Thiele, 1927, p. 114.)

Type: (by subsequent designation, Gray 1847; for Assiminea). A. grayiana Leach = Nerita Syncera hepatica Gray 1821 (fide Gray 1857, p. 23 in Turton's British Shells).

Assiminea vulgaris (Webster).

  • 1905. Rissoia vulgaris Webster. Trans. N.Z. Inst., vol. 37, p. 277.

  • 1913. Rissoa (Setia) vulgaris (Webster). Suter, Man. N.Z. Moll., p. 216.

  • 1913. Barleeia chrysomela Melv. and Stand. Iredale, Pro. Mal. Soc., vol. 10, p. 370.

  • 1915. Notosetia vulgaris (Webster). Iredale, Trans. N.Z. Inst., vol. 47, p. 454.

  • 1915. Assiminea nitida (not of Pease 1865). Oliver, Trans. N.Z. Inst., vol. 47, p. 522.

  • 1924. Tatea hedleyi Brookes. Trans. N.Z. Inst., vol. 55, p. 153.

  • 1926. Assiminea nitida (not of Pease 1865). Finlay, Trans. N.Z. Inst., vol. 57. p. 379.

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The first reference of the New Zealand shells to their correct family was in Finlay (1926, p. 379), where on Iredale's authority our species was referred to the widely distributed Polynesian Assiminea nitida (Pease). Finlay also mentioned that this species has been recorded from the Kermadecs by Oliver.

The species nitida, however, is not an Assiminea, as has been demonstrated by Thiele (1927, p. 128), for the dentition and open umbilicus show that it belongs to the genus Paludinella.

The writer has examined the dentition of Rangitoto specimens, which are topotypes of Brookes's Tatea hedleyi, and it is almost inseparable from that of grayana, the English genotype (as figured by Thiele 1. c.), having the characteristic hinder denticles on the central tooth, and moreover the shell has the umbilicus closed, which is a characteristic feature of the genotype, but not of nitida, in which it is quite open. The open umbilicus in nitida was mentioned in Pease's original description, and this feature is clearly shown in specimens from Tonga in the Auckland Museum collection.

Furthermore, upon shell characters, Kermadec Island specimens are inseparable from New Zealand examples.

If Iredale's statement (l.c., 1913, p. 370) is correct, that the Loyalty Island Barleeia chrysomela, Melville and Standen (1896, J. Conch., vol. 8, p. 309), is identical with the Kermadec Island shells, then it may be necessary later to change the specific name of the New Zealand shell once more. However, in the absence of Loyalty Island specimens, and taking into account both the poor condition of the holotype (as mentioned by Iredale, l.c., p. 370) and the fact that Iredale misidentified the Kermadec specimens as nitida, the wisest course meanwhile is to select an available New Zealand name for the New Zealand shell. This name comes from an unexpected source, as was revealed by a comparison between Webster's holotype of Rissoia vulgaris and topotypes of Brooke's Tatea hedleyi, which proved to be identical. It is possible that the combination Assiminea vulgaris has been used before, but I have not succeeded in tracing any such combination, so Webster's name may be employed for the New Zealand and Kermadec Island Assiminea, the type locality for which becomes Waipipi, Manukau Harbour.

Dentition (Fig. 7): The dental formula of Assiminea vulgaris is 1 + 1 + 1 + 1 + 1 + 1 + 1. Central tooth with five cusps, and three denticles on each side, below. Inner lateral with six cusps and outer lateral with five. Marginal teeth with 15 pectinate cusps. The central tooth is a little broader, and the cusps more regular in size, otherwise the dentition of vulgaris is almost identical with that of the genotype.

Subfamily Omphalotropidinae.
Genus Suterilla Thiele 1927.

Type (original designation): Cirsonella neozelanica Murdoch.

Suterilla neozelanica (Murdoch).

  • 1899. Cirsonella? neozelanica Murdoch. Pro. Mal. Soc., vol. 3, pt. 6, p. 320.

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  • 1909. Acmella neozelanica (Murdoch). Thiele, Arkiv. fur Naturgeschichte 75, Jahrg., vol. 1, pt. 3, pp. 387–390.

  • 1913. Cirsonella neozelanica Murdoch. Suter, Man. N.Z. Moll., p. 155.

  • 1913. Acmella neozelanica (Murdoch). Suter, Man. N.Z. Moll. p. 1082.

  • 1927. Omphalotropis (Suterilla) neozclanica (Murdoch). Thiele, Zool. Jahrbucher, vol. 53, pp. 124 and 127.

Thiele (1927, pp. 124 and 127), in proposing Suterilla as a new section of the genus Omphalotropis, remarked upon the similarity of its dentition to that of Omphalotropis rather than to that of Acmella, under which genus he had placed it formerly (Thiele 1909, pp. 387–390). Thiele now shows that on dental characters the genus Acmella belongs to the subfamily Assimineinae, whereas Suterilla is typical of the Omphalotropidinae.

However, unlike Omphalotropis, which is exclusively terrestrial, Suterilla neozelanica is as much a marine species as its associates Marinula, Ophicardelus, and Kellia, and, moreover, is so distinct from Omphalotropis on shell characters, being small and globular without an umbilical keel, that it may well be given full generic status.

Ellobiidae.

Genus Rangitotoa n. gen.
Type: Rangitotoa insularis n. sp.

Odhner, in his systematic key of the Ellobiidae (1925, pp. 1–15), which is based largely upon dental characters, has divided the family into six subfamilies—the Carychiinae, Melampodinae, Pedipedinae, Pythiinae, Cassidulinae, and the Ellobiinae.

The first three of these form a major group, having the radula characterised by pectinate marginals with one or more ectocones, while the remaining three subfamilies are grouped together on the common character of blunt chisel-shaped marginals without ectocones.

The new genus Rangitotoa, here described, has a type of dentition characteristic of the first group of subfamilies, but the dentition is nevertheless distinctive, and not in accord with that of any genus belonging to the first group of Odhner's subfamilies.

The dentition suggests a position for these shells between the Carychiinae and the Melampodinae, nearer, no doubt, to the latter. The small dental formula, together with the obsolescent tricuspid nature of the central tooth, indicates a position not much in advance of the Carychiinae, and yet not as specialised as in the Melampodinae, in which all traces of the tricuspid central have vanished.

With regard to the laterals, the large mesocone* recalls the Melampodinae, but the presence of both a definite ectocone and an obsolescent entocone are features discordant for that subfamily.

The shell, however, is not unlike the West Indian Tralia, (Melampodinae), but it lacks the characteristic thickened process within the outer lip, which is a feature of that genus.

[Footnote] * The main or middle cusp of lateral and marginal teeth is termed the mesocone, while cusps situated on the inner side are known as entocones and those on the outside as ectocones.

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Rangitotoa insularis n. sp. (Fig. 1).

Shell minute, semi-transparent, imperforate, thin, and colourless. Outline ovate-cylindrical, spire conical, less than half height of aperture. Whorls seven, slowly increasing at first, including a minute smooth protoconch of one flattened whorl. Aperture long and narrow. Outer lip thin and sharp, without a ridge or any processes within. Inner lip with three plaits, uppermost weak and not plainly visible from without, situated at a little above half the height of aperture. Central plait large and prominently projecting; lower plait less prominent, merging below into the thickened narrowly rounded basal lip. Surface smooth except for faint irregular axial growth lines.

Dentition (Figs. 5 and 6): Radula very minute, the detail of the teeth being difficult to see even with a sixteenth oil immersion objective. Dental formula small (12–16) + 1 + (16–12). Central tooth with one large cusp and an obsolete denticle on each side of it; laterals 16, each with a mesocone and an ectocone; marginals 12, each with an entocone, a mesocone, and from two to four ectocones.

  • Height, 3.7 mm.; diameter, 1.9 mm. (holotype)

  • " 2.9 mm.; " 1.4 mm. (average size specimen).

Holotype: In Auckland Museum (collected by A. W. B. P., March 12, 1930.

Habitat: Rangitoto Island; type from near the “Beacon.” Found living only near high-water mark, on the under sides of stones which are partly in contact with mud. The writer has found this species to be distributed around the southern and western sides of Rangitoto Island, but so far has failed to discover it elsewhere.

Genus Marinula King 1831.
Type (by monotypy): Marinula pepita King.

Marinula filholi Hutton 1878.

This species shows considerable adaptability, and this accounts for its wide distribution around New Zealand shores. At Rangitoto filholi lives under stones and among decaying algae in muddy situations within the high-tidal zone, but it has been recorded by Oliver (1923, p. 499) from “between tide marks” at Island Bay, Cook Strait, and the writer has observed it on the open coast at Cape Foulwind, Manukau North Head, and Muriwai, living on cliff faces at above high-tide mark in situations where fresh-water seepage occurs.

Dentition: The dental formula (not previously recorded) is 121 + 1 + 121. The central tooth has one simple cusp, the laterals from two to three and the marginals from four to five cusps. The radula is very small, and the individual teeth are crowded and difficult to see.

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It is also of interest to record that Marinula striata Odhner occurs abundantly at Mason's Bay, Stewart Island. Formerly this species was known only from the Auckland and Antipodes Islands. The Mason's Bay specimens were found alive and in numbers by Mrs R. H. Harrison, of Stewart Island.

Genus Leuconopsis Hutton 1884.
Type (original designation): Leuconia obsoleta Hutt.

Leuconopsis obsoleta (Hutton 1878).

This species is found in the sheltered harbour bays towards high-tide, usually in company with Caecum digitulum and Onchidella patelloides. It occurs sometimes on the open coast, and at Muriwai it is associated with Marinula filholi, and lives on cliffs just above high-tide mark, in situations where fresh-water seepage occurs.

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Dentition (Figs. 3 and 4): Hutton, 1884, Trans. N.Z. Inst., vol. 16, p. 213, gave the formula 55 + 1 + 55/150 but he was unable to make out the form of the teeth. The radula is certainly very minute, the detail of the teeth being very difficult to see even with a sixteenth oil immersion objective. Dental formula small (16 + 39) + 1 + (39 + 16). Central tooth with one large elongated cusp, on each side of which is an obsolescent base of a former side cusp. The base of the central tooth is shaped like an inverted T. The laterals, which number 39, are each prominently bicuspid; both cusps, of which the mesocone is the larger, being long and slender and inclined outwards. Marginals 16, pectinate, each with an entocone, a mesocone, and an ectocone, all three of which are moderately long and of almost equal length.

Although the dental formula is smaller than is usual in the Pedipedinae, the character of both the teeth and the shell in Leuconopsis is most in accord with that subfamily.

Erycinidae.
Genus Kellia Turton 1822.

Type (subsequent designation, Herrmannsen 1846): Cardium rubrum

Mont.

Kellia, W. Turton. Conch. Insul. Brit., pp. 19, 56, 1822. Examples: Kellia suborbicularis (Montagu) and K. rubra (Montagu). Kellia, Turton, Herrmannsen, Indicis Gen. Malac., vol. 1, p. 568, 1846. “Typus: Cardium rubrum Mont.” Lasaea, Leach in Brown, Ill. Conch. Gt. Brit. Ed. 1, Pl. 20, Figs. 17–18, 1827. Sole example, L. rubra (Montagu).

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Grant and Gale (1931, p. 302) have shown that owing to Herr-mannsen's type designation in 1847 of Cardium rubrum Montagu, the well-known genus Lasaea becomes a synonym of Kellia, and also that the genus Chironia Deshayes 1839 is available for the well-known species “Kellia” suborbicularis.

Kellia maoria n. sp. (Fig. 2).

Shell small, ovate, thin, moderately inflated. Colour creamy-buff without any other colouration. Interior of the same uniform colour. Beaks low and broadly rounded. Anterior end slightly longer than posterior end; both broadly rounded. Margins sharp and smooth. Hinge well developed, but not so heavy as in most species of the genus. Teeth typical, a minute cardinal and two pairs of laminae in the right valve; a cardinal and an anterior and a posterior lamina in the left valve. Resilium large, extending from the cardinal along the lower edge of hinge-plate. Surface of valves crowded with somewhat irregular microscopic concentric threads, but without wrinkles or punctures.

The species is characterised by its uniform buff colour without darker tinting, lack of wrinkles, and punctures, and its almost straight dorsal margin which does not descend under the umbo as in australis (Lamk.) and rossiana vexata (Finlay).

Length 2.40 mm., height 1.95 mm., thickness (two valves) 1.50 mm. (holotype). Holotype in Auckland Museum (collected by A. W. B. P., December 7, 1930.

Habitat: Rangitoto Island, Auckland. On under sides of stones in the high-tidal zone (type). Awanui Heads (W. La Roche, 1922); Lyall and Island Bays, Wellington (A. W. B. P.); Chatham Islands; and Paterson Inlet (A. Hamilton collection).

Wherever it occurs this species is usually found in great abundance. It lives on the under-sides of stones which are in contact with mud, but only between high-water neap and high-water spring tide levels.

The genus Kellia is now represented in New Zealand by the following four species:—rossiana (Finlay 1928), rossiana vexata (Finlay 1928), hinemoa (Finlay 1928), and maoria, Powell 1933. Lasaea neozelanica Suter must be removed from this association of species, as it proves to be a synonym of Webster's Kellia bifurca, for which Finlay (1926, p. 463) has provided the genus Arthritica. The main characteristic of Arthritica according to its author is the formation in adolescence of two internal thickened limy patches. The writer has examined the holotype and over 100 of Webster's para-types of bifurca, and found that typical specimens were uncommon, the predominant form having the limy patches scarcely visible. Furthermore, the presence of limy patches was by no means restricted to the larger specimens. The type of L. neozelanica is a large thin-shelled specimen without the limy patches.

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At first it seemed possible to recognise two varieties—one, the typical bifurca, having limy patches combined with a faintly trapezoidal outline, and the other, with undeveloped limy patches and a more regularly ovate outline. However, the type of neozelanica and a few others from North and South Island localities combine the bifurca outline with the absence of limy patches. It is inevitable that the two be merged, for there is also much variation in the strength of the hinge-plate, convexity, and external sculpture.

References.

Finlay, H. J., 1926. A Further Commentary on New Zealand Molluscan Systematics. Trans. N.Z. Inst., vol. 57.

Finlay, H. J., 1928. The Recent Mollusca of the Chatham Islands. Trans. N.Z. Inst., vol. 59.

Grant, U. S., and Gale, H. R., 1931. Pliocene and Pleistocene Mollusca of California. Mem. San Diego Soc. Nat. Hist., vol. 1.

Odhner, N. H., 1925. Marinula juanensis, n. sp., nebst Bemerkungen über die Systematik der Ellobiiden. Arkiv for Zoologi, Stockholm. Band 17, h. 2, pp. 1–15.

Oliver, W. R. B., 1923. Marine Littoral Plant and Animal Communities in New Zealand. Trans. N.Z. Inst., vol. 54.

Thiele, J., 1927. Uber die Schneckenfamilie Assimineidae. Zool. Jahrb., Jena. Band 53, pp. 113–146.

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Description of Text Figures.

  • Fig. 1.—Rangitotoa insularis n. gen. and sp. Holotype × 18.

  • Fig. 2.—Kellia maoria n. sp. Holotype × 27.

  • Fig. 3.—Leuconopsis obsoleta (Hutton). Central and innermost lateral tooth of radula.

  • Fig. 4.—Leuconopsis obsoleta (Hutton). Marginal tooth of radula.

  • Fig. 5.—Rangitotoa insularis n. gen. and sp. Central and two lateral teeth of radula.

  • Fig. 6.—Rangitotoa insularis n. gen. and sp. Marginal teeth of radula, from left, ninth, second, and first respectively.

  • Fig. 7.—Assiminea vulgaris (Webster). Radula.