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Volume 81, 1953
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[Read before Biology Section, Wellington Branch, July 9, 1952; received by the Editor, July 11, 1952.]


The fossil and living species of Monodilepas are leviewed and re–described. A new subspecies, Monodilepas monilifera cookiana is described and skinneri is classed as a subspecies of monilifera. Distribution and relationships are discussed.


The determination of mollusca from the Cook Strait region in preparation for a check list has shown that in a number of cases generic revisions are necessary before relationship can be decided. The present paper dealing with the keyhole limpets is the first of a series of such revisions.

The writer gratefully acknowledges the loan of material from Mr. A. W. B. Powell and Dr. C. A. Fleming.

Family: Fissurellidae
Genus:Monodilepas Finlay, 1927
Genotype:(o.d.) Lucapina monilifera Hutton.

Finlay (1927, p. 343) proposed Monodilepas for Hutton's Lucapina monilifera noting that the type of sculpture differentiated it from the Australian genera Amblychilepas Pilsbry, Sophismalepas Iredale and Cosmetalepas Iredale. Study of the radula of monilifera has now shown that Monodilepas is allied to Cosmetalepas although the type of sculpture differs. Finlay (1927, p. 343) mentioned an undescribed fossil form from Clifden. This record is based upon three broken specimens from band 6A which are still in the Finlay Collection. They are too damaged for description, but enough remains to show that they are close to monilifera Hutton. The genus has therefore been present in New Zealand seas since the Altonian (Lower Miocene).

Notes On The Radula Of Monodilepas monilifera Hutton (Text Fig. A)

The radula of a specimen from 50 fathoms off Oamaru has been examined.

Formula: ∞.(1.4).1.(1.4).∞×20.

There are twenty rows of teeth each consisting of a weak median tooth with four weak inner laterals and a relatively enormous outer lateral and a large number of very narrow marginals attached to a folded base, on each side. Central broad but very delicate, inner laterals narrow, the outermost almost vestigial. These teeth can play little part in mastication. Outer lateral massive, tricuspid, overlapping the inner laterals and the central. Marginals very fine, numerous attached at base to a folded plate which runs parallel to the axis of the radula. In all respects the radula of Monodilepas monilifera Hutton is in close agreement with that of the Australian Cosmetalepas concatenatus (Crosse and Fischer) as illustrated by Torr (1914, Pl.19. No.3).The two genera are separated by the nature of the sculpture, which consists of small sunken pits in Cosmetalepas. There can be little doubt, however, that the two genera are otherwise closely related.

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Text Figure A.—Radula of Monodilepas monilifera monilifera (Hutton).

Four species have previously been recognised from New Zealand waters. In the present paper an additional subspecies of monilifera is described from the Cook Strait area, and skinneri is classified as a subspecies. All the forms are closely related and from what can be seen of the Altonian representative there has been very little change since the Miocene. Slight differences are of more specific value in conservative genera, and for this reason all the New Zealand forms have not been classed as subspecies oi monilifera. Monodilepas diemenensis Finlay possesses a number of distinctive features and is therefore separated as a full species. Monodilepas otagoensis Finlay is a puzzling form in that no specimens apart from the type series appear to have been collected, and it would appear to occur within the geographical range of monilifera. Its distinguishing features are well marked, but only small shells are known. It is here accepted provisionally as a full species, at least until its adult characters, its true distribution, and its variations are better known.

One remarkable feature of the genus is that each form appears to occur in a restricted area. Monodilepas monilifera monilifera is relatively abundant in the Foveaux Strait area, with occasional sporadic occurrences to the north as far as Oamaru, M. diemenensis occurs in enormous numbers at times, near Cape Maria van Diemen, but very few specimens have been recorded to the south, the Cook Strait form appears to occur within a restricted area, while M. monilifera skinneri is, of course, confined to the Chatham Islands.

In order to make comparisons more objective, a number of measurements were made of each shell studied, and four indices were calculated from them. These measurements were total length, width, height, distance from anterior end to internal edge of foramen, and longitudinal diameter of the foramen. Where the sculpture was well enough preserved the number of radials was counted. The indices calculated were:

Width Index: The breadth of the shell expressed as a percentage of the length.

Height Index: The height expressed as a percentage of the length.

Foramen Position Index: The distance from the anterior end to the foramen expressed as a percentage of the length.

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Text Figure B.—1–3, Monodilepas monilifera monilifera (Hutton) (Holotype); 4–6, Monodilepas diemenensis Finlay; 7–9, Monodilepas skinneri Finlay (Holotype). 10–12, Monodilepas otagoensis Finlay (Holotype); 13–15, Monodilepas monilifera cookiana n.subsp. (Holotype); 16–18, Monodilepas monilifera monilifera (Hutton) Castlecliff.

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Foramen Index: The longitudinal diameter of the foramen expressed as a percentage of the length.

Monodilepas monilifera monilifera (Hutton). Text fig. B, figs. 1–3, 16–18; Pl. 16, figs. 5, 6.

Lucapina monilifera (Hutton), 1873. Cat. Mar. Moll., p. 42.

Megatebennus moniliferus (Hutton), 1893. Macleay Mem. Vol. p. 72, Pl. 8, fig. 76.

Fissuridea monilifera (Hutton); Suter, 1913. Man. N. Z. Moll., p. 105, Pl. 8, fig. 8.

Monodilepas monilifera (Hutton); Finlay, 1927. Trans. N. Z. Inst., vol. 57, p. 343.

As recognised here, this species ranges from the Snares to Cook Strait. The nominate subspecies appears to be restricted to the Forsterian. Its major characters are the attenuate anterior outline, the subtriangular muscle scar, netted sculpture with the concentric and radial ridges subequal, foramen of moderate size. The shell is fairly variable in its proportions. The number of radials varies considerably as interstitial radials are added as the shell grows. The measurements and ranges of indices give some indication of the degree of variability.

Localities. Stewart Island; Foveaux Strait; 50 fathoms off the Snares Islands; 100 fathoms off Puysegur Point, South–West Otago, 1 juvenile shell (Powell, 1927); 50 fathoms E.S.E. of Oamaru.

Fossil shells of this type have been seen from Castlecliff (Cu 3 of Fleming, 1947); muddy clay, Pukeora, Waipukurau; Okauawa Creek, Hawke's Bay, and Kai–Iwi. At first sight these fossil specimens appear separable from monilifera monilifera (Hutton) on the basis of larger size, prolongation of the anterior part of the shell, the finer sculpture and the relatively smaller foramen. Analysis of a series of living monilifera show that all these trends can be followed through the size series and that the extreme developments seen in the larger fossil shells are nothing but a consequence of larger size. Comparison of scatter diagrams comparing total length and development of the shell anterior to the foramen corroborate this point. As the shell of monilifera grows, the anterior part of the shell elongates in comparison with total length, extra radials are intercalated between the primary radials and the length of the foramen becomes shorter compared to the total length. No living shells have been seen as large as the largest fossil shells. Size differences of the order found here are not sufficient for subspecific separation. There seems, therefore, no constant feature by means of which these Pliocene shells may be separated from monilifera monilifera (Hutton).

Measurements And Indices Of Recent Shells
Length Width Height Anterior end to foramen Length of foramen Radials
Holotype 15.9 11.6 3.5 59 3.6 115
Stewart Island 15.3 10.7 3.6 6.1 3.3 worn Stewart Island 12.0 8.9 2.3 5.0 2.7 91
Stewart Island 13.0 8.1 3.1 5.0 2.8 64
Stewart Island 12.8 9.5 2.8 4.4 2.8 worn
Stewart Island 9.5 6.6 1.9 3.8 1.8 70
Stewart Island 16.2 11.7 3.0 6.5 3.3 109
Stewart Island 12.0 8.4 2.6 4.8 2.2
Stewart Island 9.5 7.3 distorted 2.8 81
Oyster beds 19.0 15.8 4.0 distorted
Foveaux Strait, 18 fathoms 22.2 16.2 5.1 8.5 4.6 worn
Foveaux Strait, 18 fathoms 10.4 7.3 2.8 4.0 2.1
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Fig. 1.—Monodilephas monilifera skinneri Finlay. Holotype. 21.7 × 14.0 mm.
Fig. 2.—Monodilephas otagoensis. Finlay. Holotype 10.0 × 8.1 mm.
Fig. 3.—Monodilephas monilifera cookiana n. subsp. Holotype 11.5 × 8.0 mm.
Fig. 4.—Monodilephas diemenensis Finlay. Holotype. 9.0 × 7.0 mm.
Fig. 5.—Monodilephas monilifera monilifera (Hutton).Castlecliff Specimen in collection of N. Z. Geological Survey. 23.0 × 16.0 mm.
Fig. 6.—Monodilephas monilifera monilifera. (Hutton). Holotype. 15.9 × 11.6 mm.

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The ranges for the indices (means in parenthesis):—

Width Index 64–74 (72)
Height Index 18.5–26 (21)
Foramen Position Index 34–42 (39)
Foramen Index 19–29 (22)

Measurements And Indices Of Fossil Shells

Length Width Height Anterior end to foramen Length of foramen Radials
Castlecliff Cu 3 23.0 16.0 5.3 9.9 3.7 91
Castlecliff Cu 3 20.4 15.0 5.9 9.6 4.3 86
Castlecliff Cu 3 12.8 9.5 3.5 5.5 2.4 69
Castlecliff 19.6 14.8 5.4 8.0 3.0 89
Castlecliff 21.9 15.5 5.4 9.9 4.0 114
Pukeora 15.0 10.1 3.7 6.1 2.8 95
Kai Iwi 28.5 20.0 8.2 12.3 6.4 98
Okauawa, H.B. 15.9 11.0 7.0 4.0 75

Ranges of Indices:

Castlecliff. Non–Castlecliff Total Fossil. Fossil and Recent Combined.
Width Index 69–75 (72) 67–70 (69) 67–75 (71) 64–75 (72)
Height Index 23–29 (26) 25–29 (27) 23–29 (28) 18.5–29 (24)
Foramen Position Index 41–47 (44) 40–44 (42) 40–47 (43) 34–47 (37)
Foramen Index 15–21 (18) 18–25 (22) 15–25 (19) 15–29 (21)

Monodilepas monilifera cookiana n. subsp. Text fig. B, figs. 13–15; Pl. 16, fig. 3.

Shell small (under 12 mm.), outline oval, with only very slight anterior attenuation. Sculpture consisting of strong radials crossed by weaker raised concentric ribs which result in a reticulation over the whole shell. The number of radials varies from 57 to 66 (66 on the holotype). The foramen is slightly longer in relation to the length of the shell than in monilifera, and the shell is distinctly higher. Obviously related to monilifera monilifera but distinguished by the smaller size and the oval outline with slight anterior attenuation.

Dimensions of Holotype. Length, 11.5 mm.; width, 8.0 mm.; height, 3.1 mm.

Locality. Dominion Museum Bottom Station 164, 40° 52' S., 174° 46' E. in 62 fathoms.

Holotype. (M. 5689) and three paratypes (M. 5690) in Dominion Museum.

Length Width Height Anterior end to foramen Length of foramen Radials
Holotype 11.5 8.0 3.1 4.6 2.9 66
Paratype 9.8 6.9 2.9 3.5 2.7 worn
Paratype 9.6 7.0 2.5 3.2 2.4 worn
Paratype 8.5 6.7 2.3 3.8 2.1 57
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Rages of Index:

Width Index 70–79 (71)
Height Index 25–30 (27)
Foramen Position Index 33–45 (36)
Foramen Index 25–28 (26)

Monodilepas monilifera skinneri Finlay. Text fig. B, figs. 7–9, Pl. 16, fig. 1.

Monodilepas skinneri Finlay, 1928. Trans. N.Z. Inst., vol. 59, p. 236, Pl. 43, fig. 59.

The distinguishing features of skinneri are the subquadrangular outline, the subparallel lateral margins, the comparatively narrow shell and the subordination of concentric sculpture to the radials. The foramen is comparatively long. In many ways it is the least variable of the species, as the comparatively narrow ranges of the indices show. There is some variation in outline, however, and several of the shells examined have an attenuated anterior margin. For this reason it is here classed as an allopatric subspecies of monilifera.

Locality. Chatham Islands.

Length Width Height Anterior end to formamen Length of foramen Radials
Holotype 21.7 14.0 5.0 9.4 5.0 78
Kaingaroa 28.5 18.6 7.2 10.7 81 worn
Kaingaroa 23.9 14.6 6.4 10.0 6.7 worn
Kaingaroa 18.7 11.5 3.2 7.6 4.1 worn
Kaingaroa 17.0 10.0 4.0 6.8 4.2 86
Chathams 21.3 13.7 5.1 8.3 6.2 worn
Chathams 17.8 11.4 4.1 6.9 4.7 83

Range of Indices:

Width Index 59–65 (63)
Height Index 17–27 (23)
Foramen Position Index 37–43 (40)
Foramen Index 22–29 (26)

Monodilepas diemenensis Finlay. Pl. 16, fig. 4. Text fig. B, figs. 4–6.

Monodilepas diemenensis Finlay, 1930. Trans. N.Z. Inst., vol. 61, p. 222.

The shells of this species possess two features that make them readily distinguishable from all the other populations. The internal callous of the foramen tends to be oval or irregularly hexagonal, in contrast with the subtriangular shape found in other species. When the shell is placed upon a flat surface the posterior margin is raised. The foramen is markedly larger than in other forms, and tends to be placed nearer the anterior margin. The characters of the posterior margin are constant in all shells seen (several hundred specimens). Such a shell modification should be due to quite marked differences in the animal (unfortunately unknown), and is of marked systematic import. The genus can no longer be maintained to be a cold–water type. In fact the genus as a whole has a wide temperature tolerance, although the individual species are restricted to comparatively small areas and have a markedly discontinuous distribution. There can be no useful discussion of this problem until more is known of larval stages and dispersal methods.

Locality. Cape Maria van Diemen, common; Discovery Stations 933 and 934 off Three Kings Islands in 260 and 92 metres respectively (Powell, 1937); Whangaroa Buoy moorings (Bollons Collection).

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Length Width Height Anterior end to foramen Length of foramen Radials
Cape Maria van Diemen 9.0 7.0 1.8 3.0 2.5 69
Cape Maria van Diemen 12.0 9.0 2.1 3.9 4.0 76
Cape Maria van Diemen 12.9 9.7 2.9 4.8 3.8 81
Cape Maria van Diemen 12.7 9.2 2.4 4.0 3.7 59
Cape Maria van Diemen 12.9 10.0 3.0 4.0 3.9 80
Cape Maria van Diemen 13.8 9.8 3.2 5.0 3.9 78
Cape Maria van Diemen 13.6 10.2 2.8 5.1 3.9 74
Cape Maria van Diemen 13.7 9.8 3.0 4.7 4.2 91
Cape Maria van Diemen 16.4 12.1 5.4 5.7 6.0 103
Cape Maria van Diemen 18.5 13.3 3.8 6.6 5.3 worn

Ranges of Indices:

Width Index 71–77 (74)
Height Index 17.5–33 (22)
Foramen Position Index 31–38 (34)
Foramen Index 28–37 (30)

Monodilepas otagoensis Finlay. Pl. 16, fig. 2; Text-fig. B, figs. 10–12.

Monodilepas otagoensis Finlay, 1930. Trans. N.Z. Inst., vol. 61, p. 222; Pl. 42, fig. 6.

Only the subadult holotype and three juvenile paratypes appear to be known. The measurements and indices of the holotype are as follows:—

Length, 10.0 mm.; width, 8.1 mm.; height, 2.2 mm.; distance from anterior end to foramen, 4.0 mm.; length of foramen, 1.5 mm.; number of radials, 80.

Width index, 81; height index, 22; foramen position index, 40; foramen index, 15.

The major distinguishing features are the very broad shell (the width index is higher than for any other specimen of the genus examined), the very small foramen, and the rounded anterior outline. These features are all present in the juvenile paratypes and present a sum of differences which make this form quite distinctive. Finlay mentioned the depressed form of the shell and the coarse sculpture in his original specific diagnosis, but these features are not marked when series of other forms are examined. Because of the marked differences in the shell and the fact that the only locality falls within the distribution area of monilifera, otagoensis is here accepted as a full species in spite of the small series of shells available for examination.

Known only from the type locality, 50 fathoms off Otago Heads.

Literature Cited

Finlay, H. J., 1927. A Further Commentary on New Zealand Molluscan Systematics. Trans. N.Z. Inst., vol. 57, pp. 320–485.

Fleming, C. A., 1947. Standard Sections and Subdivisions of the Castlecliffian and Nukumaruan Stages in the New Zealand Pliocene. Trans. Roy. Soc. N.Z., vol. 76, pp. 300–326.

Torr, C. M., 1914. Radulae of Some South Australian Gastropods. Trans. Roy. Soc. S. Austr., vol. 38, pp. 362–368.

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George Simpson, 1880–1952.

Born in Dunedin, Simpson was a son of a master-builder of that city. After school days he entered the business of his father in one of the oldest and largest building concerns of the country. Later he succeeded to the control and his profession led him to gain a considerable knowledge of the timbers of indigenous trees and of forestry. His ability and knowledge were soon recognized by a large circle with kindred interests. To his business acquaintances he was perhaps best known as an outstanding valuer. As a Crown Valuer he assessed values during the period of the operation of the Land Sales Act. He carried out his duties strenuously and resolutely, making his decisions without fear or favour.

With his life-long friend, the late John Scott Thomson, he shared a love of the open and of the garden. From their many trips into the field they brought back numerous treasures, and their gardens soon became well-known for the interest and variety of the indigenous plants they grew. The technical knowledge they gained of methods of cultivation was gladly shared with all interested. It was natural that they should seek a deeper knowledge of the plants themselves, apart from their aesthetic value; they became interested in ecological and taxonomic botany. It was equally natural that they should fall under the influence of the late Leonard Cockayne, one always eagerly on the look out for recruits to his beloved science. To them Cockayne always affectionately referred as “The Firm,” and indeed the intimate association of the two is reflected in all their horticultural and botanical work. Papers were always published jointly. Both were good photographers, and brought the camera to the aid of their work. Lectures and public talks would always be illustrated by remarkably good lantern slides. Both saw the need to make taxonomic work a matter of field and garden study, not merely a matter of herbarium specimens. Their expeditions covered a great part of the southern areas of the South Island, later to be extended to excursions throughout the island. Those privileged to join in any of these adventures soon learned to enjoy and profit by their open-heartedness, their fund of humour and bonhomie. Nights in the camp after a day's hard work live in our memories. A fitting award was that of the Loder Cup, in 1935, for they did very much towards the “protection and cultivation of the incomparable flora of the Dominion.”

Simpson gave full service to the administrative side of science as a life member of the Otago Branch of our Society. He was a member of its Council from 1927–1940, 1943–1946, 1951–1953; President in 1931; Vice-President in 1936 and 1932. He represented his branch on our Council in 1939–1940, and was a member of the Management Committee of the Otago Museum for a number of years.

Nor were Simpson's interests and activities confined to a narrow range. His unspoken motto was: “Where I can help, there I shall help.” He was a member of the Dunedin Amenities Society for many and President for several years. A popular Rotarian, his help to that organisation was given unobtrusively and effectively. Generous to a fault, his right hand did not know what his left was doing.

He, with his friend, was elected a member of the New Zealand Alpine Club in 1909. Not a mountaineer in the strict sense, he had a clear head and a good

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sense of practicable routes. He was elected on his botanical qualifications, and his talks on mountain plants were attractive and lucid. But he perhaps rendered his best services to the Dunedin Section of the Club as a highly qualified builder and master of the ways of dealing with controlling authorities. In 1940 he played a leading part in the erection of the Mount French bivouac; in 1941 of the Jumboland Hut; in 1946 of the Mount Aspiring Hut. I may be permitted to quote an anecdote related in the tribute paid by Mr. J. A. Sim, so revealing as it is of the nature of the man. “There are, I suppose, in every club, those rare individuals who do not seek the limelight: who seldom if ever raise their voices in public; but who are always there when there is hard work to be done, provided there is not too much fuss about it. Such a one was George Simpson, and as a master-builder, at a time when some doubts existed whether the primary object of the club was mountaineering or hut building, his services were invaluable. In 1940 he was a member of the Easter working party which built the Mount French bivouac under the direction of our present President, Harry Stevenson. Harry, not knowing the capabilities of his party, and seeing amongst his helpers an elderly looking fellow who might need a little coaching, began with an introductory talk on some of the more simple principles of building construction, including the right way to hold and use a hammer. It is not difficult to picture George Simpson with his quiet suppressed smile, the wrinkles creasing round his twinkling eyes, taking his instruction in all seriousness. It is typical, too, that it would not be from George that Harry would learn of the faux pas which he has not yet been allowed completely to forget.” Nor will his friends of the Otago Home Guard soon forget his help and companionship.

The range of his botanical work, or as he would have preferred to say, “of the firm,” is sufficiently attested by the bibliography attached. Special note should, however, be taken of his revision of the genus Carmichaelia, including the grouping into the subgenera Thomsoniella, Carmichaeliella, Kirkiella, Enysiella, Petriea, Suterella, Monroella, Huttonella.

This subdivision was based on the structure and mode of dehiscence of the pods. During his work he had grown in his garden specimens of almost all the species and searched the original localities. He recognized 39 species and 11 varieties, against the 21 and 9 of Cheeseman's Manual of 1925. Apart from Carmichaelia, some 46 species and 40 varieties were described in various papers in our Transactions.

Rewards came unsought, but were appreciated. He was elected a Fellow of the Linnean Society in 1930, and of our Society in 1949. He found his chief reward in the increased zest for life given by his deepening interest in botany. He will long be remembered for his ever ready help to any fellow-worker, whether young or old. He would go to no end of trouble to collect specimens, provide living plants or make his own knowledge available. With his friend he assiduously collected lichens, their total collections amounting to over 3000 specimens, fully annotated. These have been in large part worked up by Zahlbruckner in his Lichenes Novae-Zelandiae of 1941, and are still engaging the attention of later lichenologists.

Active to the last, Simpson was attending meetings, gathering specimens, and making notes to within a few days of his death. He left unfinished a revision of the genus Ourisia. New Zealand botany owes much to its devoted amateurs—

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a band in which George Simpson takes his quiet place. He had “warmed both hands before the fire of life.”

H. H. Allan.


(All papers till end of 1943 in conjunction with J. S. Thomson.)

1926. Results of a brief botanical excursion to Rough Peaks Range. N.Z. Journ. Sci. & Tech. 8, 373–378.

1926. A wild Hebe community in New Zealand. Genetica 8, 375–388.

1928. On the occurrence of the silver southern-beech. (Nothofagus Menziesii) in the neighbourhood of Dunedin. T.R.S.N.Z. 59, 326–342.

1930. The Red Hills Country. N.Z. Alpine Journ. 4, 84–90.

1932. The Vegetation of South Island, New Zealand. Vegetationsbilder. R.22, H.5/6. With L. Cockayne.

1932. Some New Zealand indigenous-induced weeds and indigenous-induced, modified and mixed plant–communities. Journ. Linn. Soc.—Bot. 49, 13–45. With L. Cockayne.

1936. Notes on hydrogen–ion concentration of forest soils in the vicinity of Dunedin. T.R.S.N.Z. 66, 192–200.

1936. Rate of growth of cultivated specimens of Raoulia buchanani T. Kirk. T.R.S.N.Z. 66, 329.

1938. An artificial scree in a New Zealand garden. Journ. Roy. N.Z. Inst. Hort. 8, 8–24.

1938. Some characteristic South Island mountain plants. N.Z. Alpine Journ. 7, 1–12. Reprinted in Quart. Bull. Alpine Garden Soc. of Great Britain, 7, 1939, 297–315.

1938. The Dunedin Sub-District of the South Otago Botanical District. T.R.S.N.Z. 67, 430–442.

1940. Notes on some New Zealand plants and descriptions of new species. T.R.S.N.Z. 70, 27–33.

1941. Records of plant stations. T.R.S.N.Z. 71, 82–98.

1942. The effect of severe winter conditions on plant-life in the Dunedin Botanical Sub–district. Journ. Roy. N.Z. Inst. Hort. 11, 73–77.

1942. Notes on some New Zealand plants with descriptions of new species. T.R.S.N.Z. 72, 21–40.

1943. Notes on some New Zealand plants and descriptions of new species. T.R.S.N.Z. 73, 155–171.

1945. Notes on some New Zealand plants and descriptions of new species. T.R.S.N.Z. 75 187–202.

1945. A revision of the genus Carmichaeha. T.R.S.N.Z. 75, 231–287. (Cockayne Memorial Paper No. 1.)

1952. Notes on some New Zealand plants and descriptions of new species. T.R.S.N.Z. 79, 419–435.