Go to National Library of New Zealand Te Puna Mātauranga o Aotearoa
Volume 80, 1952
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A Revision of the Genus Dracophyllum: Supplement

[Read before the Wellington Branch. April 26, 1951; received by the Editor, April 26, 1951]


This paper lists all the known species of Dracophyllum, giving descriptions and distribution details supplementary to those given in a revision of the genus published in 1929. New species and subspecies described are: D. densum, Mount Rochfort; D. trimorphum, West Wanganui Inlet. D. filifolium Hook f var. centrale, Mount Ruapehu; D. longifolium (Forst) var. septentrionale, Ruahine Range; var. pluviale, Caswell Sound Name changes are D. acerosum Berggr. for D. acicularifolium (Cheesem.) and D. peninsulare Oliver; D. cosmelioides new name for D. gracile Br. and Gris.; D. pyramidale new name for D. recumiatum Oliver (not Colenso). D. rosmarinifolium Forst is identified and described.


This paper is intended to be read with the paper published under the title, “A Revision of the Genus Dracophyllum”, in the Transactions of the New Zealand Institute, vol. 59, pp. 678–714, 1929. It contains additional matter gathered during the intervening period of twenty-two years, and corrections to the first account. Since 1929 a great deal of collecting has been done by various botanists and much material added to the herbaria of the Dominion Museum, Auckland Museum, and the Botany Division of the Department of Scientific and Industrial Research. All this material has been used by the author. Additional information regarding species found outside New Zealand has also been included.

Thanks are acknowledged to the Government Botanists at Brisbane and Melbourne for supplying information regarding the collections under their charge, and to Professor A. Guillaumm, of the Museum National D'Histoire Naturelle, Paris, for supplying information regarding the types of New Zealand species in that institution.

Species of Dracophyllum

The number of species of Dracophyllum herein admitted is 49 (not including D. compactum). They are arranged according to the following list, in which they are divided into group of related species.

Subgenus Oreothamnus

Group of D. muscoides: D. muscoides, D prostratum, D. pronum.

Group of D. minimum: D. minimum.

Group of D. pearsoni: D. densum, D. politum, D. pearsoni.

Group of D. rosmarinifolium: D. rosmarinifolium, D. uniflorum, D. acerosum.

Group of D. subulatum: D. subulatum, D. palustre.

Group of D. kirkii: D. kirkii, D. pubescens, D. trimorphum.

Group of D. scoparium: D. scoparium, D. paludosum.

Group of D. urvilleanum: D. urvilleanum, D. lessonianum, D. filifolium.

Group of D. longifolium: D. longifolium, D oliveri.

Group of D. adamsii: D. adamsii

Group of D. sinclairii: D. sinclairii, D patens, D arboreum, D. viride.

Group of D. recurvum: D. recurvum.

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Subgenus Eudracophyllum

Group of D. menziesii: D. menziesii, D. townsoni, D. fiordense.

Group of D. secundum: D. strictum, D. ramosum, D. secundum, D. vieillardii, D. alticolum, D. amabile, D. cosmelioides, D. thiebautii.

Group of D. milliganii: D. sayeri, D dracaenoides, D. milliganii, D. fitzgeraldi.

Group of D. latifolium: D. latifolium, D. matthewsii, D. traversii, D. pyramidale.

Group of D. verticillatum: D. verticillatum.

Subgenus Cordophyllum

Group of D. involucratum: D. involucratum.

Dracophyllum muscoides Hook. f.

Dracophyllum muscoides Hook. f. Oliver. Trans. N.Z. Inst., 59, 685, 1929.

This species resembles D. prostratum in its spreading habit, but at its central point it grows more compactly, forming dense cushions of upright close branches. Its leaves are of similar shape, but considerably smaller. In some localities the ends of a few or many of the branches bear the leaves in dense expanded bunches, thus increasing the density of the cushion. These are due to an insect feeding on the terminal buds. Specimens with these enlarged branches are from Mount St. Mary, with most branches enlarged (D.S.I.R. 65382). Old Man Range, few branches enlarged (A M. 517), and Hunters Hill.

Type: Alps of Otago. alt. 7–8,000 ft. Hector and Buchanan, in Kew Herbarium. The type locality is more precisely given by Buchanan (Trans. N.Z. Inst., 14, 346, 1882) as Mts. Alta and Hector's Col. 5–7,000 feet alt. Hector and Buchanan, 1862. There are specimens collected by Buchanan from Otago Alps in the Dominion Museum.

Distribution. Additional localities: Mt. Berger, Garvie Mtns., Humboldt Mtns., Maungatua. J. F. Armstrong recorded this species from Rangitata and Ashburton valleys, two localities in Canterbury.

Dracophyllum prostratum Kirk

Dracophyllum prostratum Kirk: Oliver, Trans. N.Z. Inst., 59, 686, 1929.

Hybrid. On Mt. Maungatua this species hybridises with D. uniflorum (i.e. D. rosmarinifolium of my former paper).

Type: Mountains above Lake Harris, Otago, 4,000 feet, among Sphagnum, T. Kirk, in Dominion Museum.

Distribution. Additional localities: Mt. St. Mary, Mt. Edwards, Garvie Mtns., Lake Wakatipu. Mt Cuthbert. Key Summit. Mt. Hamilton. Bold Peak, Takahe Valley.

Dracophyllum pronum Oliver

Dracophyllum pronum Oliver, Trans. N.Z. Inst., 59, 686, 1929. D. rosmarinifolium of Hook. f. and Cheesem.

The essential characters of this species are the prostrate trailing habit with short, scattered branchlets, and the small, tufted, blunt-pointed leaves, giving on open appearance to the plants which often closely hug the ground. It differs from D. densum in its open habit, due to the branchlets being covered only with clean scales below the terminal tuft of leaves.

Type: Blimit Mtn., above Arthur's Pass, 6,000 ft. W. R. B. Oliver, in Dominion Museum.

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Distribution. Mostly on the eastern side of the main divide of the South Island. Additional localities: Dun Mtn., Mt. Arthur, St. Arnaud Range, Mt. Isobel, Lead Hill, Mt. St. Mary, Humboldt Mtns., Maungatua. Homer Valley, Mount Peel (Canterbury), Takahe Valley.

The localities Mt. Rochfort and Mt. William recorded for this species (l.c., p. 687) should be transferred to D. densum.

Dracophyllum minimum F. v. Muell.

Dracophyllum minimum F. v. Muell: Oliver. Trans. N. Z. Inst., 59, 685, 1929.

There are two species mixed in my account of D. minimum. The specimens with upright stems (Fig. 1, top) are the true D. minimum. The lower specimen is a Sprengelia from Lake Dora, Tasmania. D. minimum forms cushions of upright branches 6–10 cm. tall. Leaves closely appressed, 6–7 mm. long, acute, with broad base 2 mm. or more across. Flowers terminal, corolla tube 4 mm. long, anthers included, lobes spreading. D. minimum is near D. prostratum, but is of much more compact growth and has broader leaves.

Type: From Mt. La Perouse, Tasmania, A. Oldfield, in the Melbourne Herbarium.

Distribution. Additional localities: Cradle Mountain, Elden Bluff, Mt. Victoria.

Dracophyllum densum Oliver sp.n.

Frutex humilis, saepe prostratus. multo ramosus. Rami foliis aridis vestiti Folia linearia, apice obtusa, basi supra concava, apice supra plana, subtus carinata, 12 mm. longa, viridia; vagina breve, margine ciliato Flores solitarii, terminales. Sepala ovata, ciliata. Corolla tubo 5 mm. longo, lobis imbricatis, triangularibus. Antherae sessiles Ovarium obconicum, truncatum, 5-loculare. Capsula truncata, 2 mm. diam.

A low, often prostrate, much branched shrub, with dense foliage. Branches rough, the lower ones ridged, dark grey to dark brown above, reddish-brown below, the branchlets usually covered with dead leaves. Leaves linear, obtuse, basal half of upper surface concave, apical half flat, under surface ridged, sheath short, one sixth the length of the leaf, margin sheared away from blade, minutely ciliate. Leaf blade green or olive green, sheath brown with white margin Length of leaf 12 mm. (including sheath 2 mm.). Flowers solitary, terminating short, leafy branchlets. Bracts and sepals narrowly ovate, scarcely overtopping the flowers, the sheath running gradually into the lamina, margin ciliate. Corolla tube 5 mm. long, lobes imbricate, triangular, with a central ridge on the inner face, 1 ½ mm. long. Anthers sessile, at the throat of the corolla tube. Ovary obconic, truncate, 5-lobed. Capsule about 2 mm. across, truncate.

Type from Mount Rochfort, collected by W. R. B. Oliver, March 2, 1949, in Dominion Museum.

In habit and size much like D. politum, but leaves smaller, with much smaller sheaths, and drying green or olive green.

From D. pronum it differs in its more contracted and its semi-erect habit, more dense foliage, and larger leaves with more acute tips. Whereas D. pronum trials over the ground for 30 to 60 cm. or more, D. densum forms a low shrub, the outer branches being prostrate, the central ones erect but short. On Mount Davy, in the Paparoa Range, in rock crevices, the stems grow to a height of five or six feet.

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It differs from both pronum and uniflorum by its branchlets being clothed with dead leaves or their bases.

Distribution. Mt. Rochfort and Denniston Plateau, Mt. William, Mt. Davy.

Dracophyllum politum (Cheesem.) Ckne.

Dracophyllum politum (Cheesem.) Ckne.: Oliver, Trans. N.Z. Inst., 59, 687, 1929.

A stout, prostrate plant with rigid leaves whose old bases clothe the branches below the living leaves. In this respect it agrees with D. pearsoni, D. densum, D. prostratum, and D. muscoides to differ from D. pronum and the D. uniflorum group. Leaves slightly curved inwards, sheath large, without cilia or only odd ones, leaf blade concave above for about two-thirds, then flattened to the tip, which is rounded and flattened. Under surface obscurely keeled. Flowers solitary, subtended by very broad, blunt-pointed bracts. The leaves dry reddish-brown as they do in its near relative D. pearsoni. D. politum differs from D. densum in its stouter leaves with large sheaths, and in the reddish-brown bark and leaves.

Type: Mt. Maungatua, 3,000 ft. D. Petrie, in Auckland Museum.

Distribution. Additional localities: Wilmot Pass, Rock and Pillar Range.

Dracophyllum pearsoni Kirk

Dracophyllum pearsoni Kirk: Oliver, Trans. N.Z. Inst., 59, 687, 1929.

This species is close to D. politum, both of which differ from all other species of Dracophyllum in their branchlets being covered with reddish-brown leaves.

Type: According to Kirk, collected by W. L. Pearson on Stewart Island, exact locality doubtful, in Dominion Museum.

Distribution. Additional localities: Near South Cape, Rakiahua, Table Hill, all in Stewart Island. Facile Harbour, Dusky Sound.

Dracophyllum rosmarinifolium (Forst. f.) R. Br.

Epacris rosmarinifolium Forst. f., Fl. Ins. Austr. Prodr., 13, 1786. Dracophyllum rosmarinifolium (Forst. f.) R. Br., Prodr. Fl. Nov. Holl., 556, 1810. Richard, Fl. Nouv. Zel., 220, 1832 (Forster's full description). A. P. DeCandolle, Prodr. Syst. Nat. Veg., 7,770, 1938. Oliver, Trans. N.Z. Inst., 59, 690, 1929 (Mount Barber specimen only). Allan, Trans. Roy. Soc. N.Z., 69, 272, 1939.

The synonymy given in my former paper (p. 689) under the heading D. rosmarinifolium should be allocated to the three species there named as here accepted.

In 1786 G. Forster described Epacris rosamarinifolia in a very brief diagnosis, but a fuller description by the same botanist was published in Richard's work on the flora of New Zealand, 1832. The species has been misunderstood by botanists dealing with the New Zealand flora mainly because it was not collected after Forster's visit until 1887, when Reischek brought back some scraps from Chalky Inlet. In 1912 Petrie collected it on the Takitimu Mountains. But in neither of these cases was it recognised as Forster's species. In 1927, that is 140 years after Forster's visit, I brought back a short series of specimens from Mt. Barber, above Wilmot Pass, and these enabled me to recognise it as the plant described by Forster, although I mistakenly united it (l.c., p. 689) with the widely distributed species described by Hooker as uniflorum. Allan (l.c.) after inspecting Forster's specimens in the British Museum, declared that my account included two distinct subspecies or species. With this I agree, and so in this paper have restricted Forster's rosmarinifolium to the plant now known to occur only in the Fiordland region and some of the mountain ranges to the eastward. In Cheesman's Flora

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(1906 and 1925 editions), following Hooker, the name rosmarinifolium is applied to the species herein called pronum. On a similar erroneous identification, Cockayne synonymized politum with rosmarinifolium in the Vegetation of New Zealand, 1928, and Simpson (Trans. Roy. Soc. N.Z., 75, 191, 1945), argued that he was correct. The true rosmarinifolium, however, is an upright form entirely different from the prostrate species politum and pronum.

All this confusion is really due to the fact that Forster's species was not again collected for more than a hundred years after he visited Dusky Sound and was not recognised for another forty years.

It is necessary to give a description of D. rosmarinifolium, as all accounts under that name are either inadequate or confused.

An erect shrub, 50 cm. or more tall, with rather stout, upright stems terminated by a cluster of 10 to 15 branches arising at acute angles. Branches 1 ½ to 2 ½ mm. in diameter, clear for 4 to 5 cm., with regular transverse leaf scars, and terminated by a narrow bunch of leaves. Leaves linear, gradually tapering to an acute, sometimes blunt, point. Sheath with the sides meeting only a short distance from the base, upper portion slightly angled or cut away, light reddish-brown with paler ribs, margins minutely ciliate. Leaf blade 30 to 35 mm. long, 2 to 2 ½ mm. wide at base, sheath 7 mm. long. On non-flowering specimens, presumably young, leaf blade 50 by 2 ½ mm., sheath 12 mm. Flowers single, terminating short, stout branchlets clustered near the tops of the branches. The leaves or bracts beneath the flowers are short and broad, the sheaths of the upper ones being longer than the blades. Sepals narrowly ovate, margin pale and minutely ciliate. Corolla tube 6 mm. long with five broad, triangular, imbricate lobes. Stamens within the top of the corolla tube. Capsule depressed, splitting widely into five valves.

Although this species is nearest to D. uniflorum, it differs appreciably in the stoutness of all its parts. This is especially noticeable in the broad, robust bracts surrounding the flowers. The leaves are much wider and are blunter at the apex than in uniflorum.

Type: Dusky Sound, G. Forster, 1773, in British Museum or Gottingen University.

Distribution: On mountains at altitudes of 3,000 to 4,000 ft. or more in the south-west of the South Island. Dusky Sound, G. Forster, 1773; Chalky Inlet, A. Reischek, 1887; Takitimu Mountains, D. Petrie, 1912; Copland River, A. Wall, 1923; Mtns. above Hay River, W. A. Thomson, 1926; Mt. Barber, W. R. B. Oliver, 1927; Mt. Pakiri, G. Simpson, 1944; Garvie Mtns., G. Simpson; Key Summit, W. R. B. Oliver, 1944; Mtns. above Caswell Sound, A. L. Poole, V. D. Zotov, W. R. B. Oliver, 1949.

The plant from Key Summit is more slender in all its parts than those from other localities and has shorter, narrower leaves.

Dracophyllum uniflorum Hook. f.

Dracophyllum uniflorum Hook. f., Handbk. N.Z. Flora, 182, 1864. Cheesem., Man. N.Z. Fl., 427, 1906, and Ed. 2, 709, 1925 (excluding varieties). D. rosmarinifolium Oliver, not Forst., Trans. N.Z. Inst., 59, 689, 1929. D. uniflorum Hook. f. var. frondosum Simpson, Trans. Roy. Soc. N.Z., 79, 424, 1952.

Three varieties of this species can be defined. (1) The typical form is an erect shrub with the leaf tips ending in short acicular points. (2) Upright shrub with short, blunt-pointed leaves. (3) Var. frondosum Simpson. A trailing shrub with narrow, needle-pointed leaves. In the blunt-leaved form there is a tendency

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to trail when growing in open. rocky situations such as the Mineral Belt, Dun Mountain.

Type of species: Wairau Mtns., Marlborough. W. T. L. Travers, in Kew Herbarium. Var. frondosum. Deep Stream, Otago. G. Simpson, in Dept. Sci. Ind. Res., Wellington.

Distribution. (1) Typical form. Marlborough and East Nelson southwards to Otago and Fiordland. Localities listed on p. 690, except Mt. Holdsworth, Mineral Belt, an Takitimu Mtns. Additional localities: St. Arnaud Ra., Mt. Isobel, Lead Hill. Clarence Valley, Upper Awatere, Papanui (Armstrong, 1865), Sealy Ra., Garvie Mtns., Mt. Ida. Rough Peaks, Kirkliston Range, Swampy Hill, Alex Knob, Mt. Moltke. Mt. Greenland, Takahe Valley. Wilmot Pass. Caswell Sound, Dusky Sound, Preservation Inlet.

(2) Form with blunt-pointed leaves. Kaimanawa Ra., Ruahine Ra., Tararua Ra., Mineral Belt.

(3) Var. frondosum. Deep Stream. Maungatua, Mt. St. Mary, Garvie Mtns.

Dracophyllum acerosum Berggr.

Dracophyllum acerosum Berggren, Minn. Fis. Sallsk. Lund, 28, 15, 1877. D. uniflorum var. acicularifolium Cheeseman. Man. N.Z. Fl., 427, 1906. D. acicularifolium Cockayne, Rept. Scenery Pres., 4. 915. D. peninsulare Oliver, Trans. N.Z. Inst., 59, 690, 1929.

An erect shrub 1–2 m. tall, stems upright, branches and leaves diverging little from the stems. Leaves narrow linear, acicular, concave above, rounded below at basal end, ridged at apical end where the margin is minutely serrate: sheath oblong, shouldered and ciliate on the margin above. Leaf blade 70 × 1 ½ mm. (Kowai Valley) to 190 × 1 ½ mm., sheaths 20 × 5 mm. (Banks Peninsula). Flowers solitary, mainly clustered below the topmost leaves, each on a short branchlet surrounded by bracts. When the flowers fall away, short stalks are left on the branches. Lower bracts leaflike, about 25 mm. long. Sepals narrow ovate, acuminate, reaching to the top of the corolla tube. margins ciliate. Corolla tube 7 mm., lobes 2 ½ mm.

This species, recognised by its strict habit; long leaves and solitary flowers, has received three specific names, the earliest being Berggren's. There is no doubt as to the identification of Berggren's species, as there is, in the Dominion Museum, a specimen from Mount Torlesse collected by Berggren himself. In any case, there is no other long-leaved species in this locality. Cheeseman included Berggren's name as a synonym of D. uniflorum and at the same time described the variety acicularifolium from the nearby locality of Broken River. In my former paper I considered Cheeseman's type to be a hybrid and introduced the name peninsulare with type from Banks Peninsula, but I am now satisfied that Berggren's, Cheeseman's and my own plant all belong to the same species.

Type specimens: D. acerosum Berggr., collected by S. Berggren on Mount Torlesse, February, 1874, now preserved in Oslo. D. uniflorum var. acicularifolium Cheesem., collected by T. F. Cheeseman at Castle Hill, Broken River Basin, in Auckland Museum. D. peninsulare Oliver, collected by R. M. Laing on Mount Berard, Banks Peninsula. in herbarium. Department of Scientific and Industrial Research.

Distribution: Mount Torlesse, Ben More, Castle Hill, Mount Peel (Canterbury), Tapuaenuku, Mount Cook, Selwyn Gorge, Banks Peninsula, Craigieburn Mountains, Awatere Valley, Rangitata.

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Dracophyllum subulatum Hook. f.

Dracophyllum subulatum J. D. Hooker: Oliver, Trans. N.Z. Inst., 59, 691, 1929.

Type collected by J. C. Bidwill near the Rangitaiki River (not far from the present township of Galatea), January, 1842, in Kew Herbarium.

Distribution: Additional localities: Matamata (northern limit of species), Kotukuroa Creek, summit of Tarawera Mountain, Kirk, 1873. Kakaramea, Wairakei, Kaweka Range, Puketitiri, Rotoaira, Ngauruhoe volcano, Maraeroa, Mount Tauhara.

Dracophyllum palustre Ckne.

Dracophyllum palustre Cockayne, ex Oliver, Trans. N. Z. Inst., 59, 690, 1929. (New name for D. virgatum, preoccupied.)

This species is recognized by its strict slender stems, which are purplish-brown; by the filiform, blunt-pointed leaves; and by the single flowers borne mostly in groups on the main stems or on side branches. The leaves are usually from 20 to 25 mm. in length, though in plants from the Denniston Plateau the lower leaves are up to 50 mm. in length. The flowers are surrounded by rather broad bracts with conspicuous broad, white margins, ciliate along the edge, and purplish bases.

The following specimens are included here, though they differ somewhat, especially in habit, from the prevalent form. Okarito, edge of swamp in forest. (Oliver, Jan., 1950, Dom. Mus) Stems long, up to 1 m. in length, spreading over the ground. In this respect it is like D. uniflorum var. frondosum, but is more slender and the leaves have blunt tips. The leaves agree with those of D. palustre, but have shorter sheaths and do not exceed 26 mm. in length. There are no flowers. The prostrate habit may in part be due to the shady position in which the plants were found growing.

The nearest relation of D. palustre is D. subulatum, which species, however, is smaller in all its parts.

Type: The type specimen of Cheeseman's Dracophyllum uniflorum var. virgatum was collected by T. Kirk near Kumara in February, 1877, and is preserved in the Auckland Museum (AM. 7028).

Distribution. Additional localities: Lewis Pass, Rahu Saddle, Jacks Pass, Mt. Drana (Wakatipu), Okarito. The Mount Earnslaw record is from a garden and therefore needs verifying. Denniston Plateau (robust form).

Dracophyllum kirkii Berggr.

Dracophyllum kirkii Berggren: Oliver, Trans. N.Z. Inst., 59, 690, 1929.

This small, prostrate, broad-leaved species resembles D. pubescens, but it has narrower leaves and is glabrous. The leaves average 4 mm. in width above the sheath. Leaves in specimens from Alex Knob, blade 39 × 4, sheath 8 mm. D. kirkii scrambles over rocks at high altitudes. On Mount Moltke it ascends to 6,500 feet.

Type from Mount Torlesse, collected by S. Berggren, 1877, in Oslo.

Distribution. Additional localities: Chaffeys Creek (Nelson), Mount Fishtail, Lead Hill, Crawford Range, mountains west of Hawdar River, Mount Moltke, Alex Knob, Mount Alexander (Westland), Brownings Pass, Godley River, Mount Cook.

The specimens from Copland River, recorded on page 692 of my previous paper, belong to D. rosmarinifolium.

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Dracophyllum pubescens Cheesem.

Dracophyllum pubescens Cheeseman: Oliver, Trans. N.Z. Inst., 59, 692, 1929.

In this species the leaves are wide and taper gradually to an acute point. They are covered on both sides with a greyish mealy looking pubescence. The leaves are larger than in D. kirkii, in which they are glabrous. Flowering specimens are fairly uniform in size of leaf. 36 by 4 to 52 by 6 mm. in south-east Nelson, 40 by 4·5 mm. on Denniston plateau. From Mount Frederick nearby there is a specimen with juvenile leaves 63 by 7 mm. The species scrambles over rocky places from 2,000 to 5,000 feet altitude. Simpson notes on the Lead Hill specimens “shrub to 6 feet on bush edges” Seedlings from Burnett's Face have the first leaves up to 10 by 1 mm. and almost glabrous. The next leaves may be 15 by 2 mm.

Type from mountains near Westport, collected by W. Townson, in the Auckland Museum.

Distribution. Additional localities: Mount Peel (Nelson), Mount Williams (south-west Nelson), Mount Perry (Gouland Downs), Lead Hill, Denniston Plateau, Anatoki Range, Mount Zetland. Mount Lema, range north side of Hurunui River, 4,200 feet.

Dracophyllum trimorphum Oliver n.sp.

Frutex 2–3 m. altus. Planta juvenilis erecta, caulibus levibus, refescentibus, cicatricibus circularibus. Folia acuminata, glabra. viridia, 125 × 10 mm. Planta submatura cortice fusco, cicatricibus obsoletis. Folia acuminata, pubescentia, glauca, 80 × 6 mm. Planta matura cortice fusco, ramulis cicatricibus. Folia angusta, acuminata, 45 × 4·5 mm., pubescentia minute. Vaginae breves, minute ciliatae. Flores solitarii vel 2–3 racemosi. Bractae foliosae. Sepala anguste ovata, acicularia. Corollae tubus 4 mm. Antherae subexsertae. Ovarium obovatum, truncatum, stigmate capitato.

A small tree, 2–3 m. tall, passing through several leaf changes from juvenile to maturity. (1) Juvenile plants ½ to 1 m. tall, erect, unbranched, stems smooth, with reddish-brown bark and circular leaf scars averaging 7 mm. apart. Leaves long, wide and grass-like, glabrous, bright green, 125 by 10 mm. including sheath. (2) Submature plants to 1 m. or more tall, bark changing to reddish-brown with the leaf scars obliterated. Leaves acuminate, with glaucous tinge due to a fine powdery looking pubescence, 80 by 6 mm. including sheath, and smaller. (3) Adult plants have dark brown bark with scars only on the fine twigs below the leaves. These small branches are very slender and numerous. They bear the flowers. Leaves narrow, 45 by 2 ½ mm. including sheath, minutely pubescent on both sides like the leaves in the submature stage, but more seanty. Sheaths in all stages short, overlapping at base, ciliate on the shoulder, which is weak. Flowers single or in racemes of two to three flowers, surrounded by leaf-like bracts, the upper ones not extending beyond the top of the corolla. Sepals narrowovate, with acicular apexes. Corolla tube 4 mm. long. Filaments attached to the inside of the corolla tube, anthers slightly exserted. Ovary obovate, with truncate top, stigma as long as the corolla tube, capitate.

This remarkable plant is best placed in the group containing D. pubescens and D. kirkii. The young plants resemble those of D. longifolium, but the leaves are much shorter. The submature leaves remind one of D. pubescens, but they are larger in every way and, unlike pubescens, have long drawn out tips. The adult leaves resemble those of K. kirkii, but are longer and narrower and are somewhat pubescent.

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Fig. 1—Dracophyllum densum Oliver, Mount Rochfort. Type. Fig. 2—Dracophyllum pronum Oliver, Fog Peak, Torlesse Range.

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Dracophyllum trimorphum (Forst f.) R. Br., Mount Alexander, Caswell Sound.

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Dracophyllum accrosum Berggren, Kowai River,

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Dracophyllum trimorphum Oliver, West Wanganui Inlet. Type.

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Dracophyllum allicolum Damker, Mount Humboldt. New Caledonia. Type.

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Dracophyllum longifolium (Forst. f.) R. Br. subsp. septcntrionale Oliver, Mt. Maharahara, Ruahine Range. Right hand figure is type of subspecies.

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Type from West Wanganm Inlet collected by W. R. B. Oliver, December 28, 1949, in Dominion Museum.

Distribution. North-west Nelson. West Wanganm Inlet, growing on low seacliffs. Cape Farewell, in manuka scrub.

Dracophyllum scoparium Hook. f.

Dracophyllum scoparium J. D. Hooker: Oliver. Trans. N.Z. Inst., 59, 693, 1929.

Type specimen collected by J. D. Hooker. December, 1840, on Campbell Island, in Kew Herbarium (No. 1611).

Distribution. Campbell Island: Hooker. 1840; Kirk, 1890; Cockayne, 1903; Laing, 1907; Aston, 1909; W. R. B. Oliver, 1927; R. L. Oliver, 1944; Sorensen, 1947; Brockie, 1947.

Dracophyllum paludosum Ckne.

Dracophyllum paludosum Cockayne Oliver Trans N. Z. Inst. 59, 693, 1929.

Type specimen collected by L. Cockayne, February, 1902, in Dracophyllum-Olearia bog, on Tableland, Chatham Island, in Dominion Museum (No. 3567).

Distribution Chatham Islands. This species was first collected by H. H. Travers in October, 1863, on Chatham Island and Pitt Island. Specimens were sent to Mueller and are now in the Melbourne Herbarium, but that botanist did not distinguish them from D. arboreum, which he describes under the name scoparium. Cockayne, in 1901, collected the species in boggy ground on the tableland, at Whangamarino, and between Korako and Wharekauri.

Dracophyllum urvilleanum A. Rich.

Dracophyllum urvilleanum A Richard: Oliver, Trans N.Z. Inst., 59, 694, 1929.

D. urvilleanum can easily be distinguished from D. filifolium when in flower, as the recemes bear only 2 to 3 flowers and sometimes the flowers are single. The flowers, too, are narrower and shorter, the sepals being about 4 mm. long, whereas in D. filifolium they are about 5 mm. long Forster's description, quoted by Richard, gives the flowers as “2–3–4, rarius solitarii”. The leaves of D. urvilleanum are distinctly narrower than are those of D. filifolium.

Type specimen collected by Dumont D'Urville at Tasman Bay. Professor Guillaumin informs me that he could not find the type of D. urvillcanum in Paris He says that it may be at Caen, where D'Urville's herbarium is preserved.

Distribution. Additional localities: Wangapeka, Kaiuma, Mount Riley, Okaramio, Rai Valley, Mount Stokes. Pelorus River. Whangamoa Saddle.

Dracophyllum lessonianum A. Rich.

Dracophyllum lessonianum A. Richard Oliver, Trans. N.Z. Inst., 59, 696, 1929.

Hooker in 1864 reduced Richard's species lessonianum to a variety of urvilleanum. Cheeseman in 1906 adopted this classification and in giving the range as from Rotorua to Stewart Island excluded the type locality. Cheeseman's conception of lessonianum was thus different from Richard's and must have included specimens here accepted as filifolium, although he accepts this as another variety of urvilleanum. Undoubtedly lessonianum is closely allied to filifolium, but it is separated by its shorter leaves, long acuminate sepals and bracts and by its longer flowers. Its area of distribution does not overlap that of filifolium.

Type from Bay of Islands, collected by Dumont D'Urville in 1827; in the Museum Nationale D'Histoire Naturelle, Paris.

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Distribution. Additional localities: Peria, Te Paki, Spirits Bay, Houhoura, Whangaroa, Mangonui, Kaiaka, Matakana, Kerikeri, Wade.

Dracophyllum filifolium Hook. f.

Dracophyllum filifolium J. D. Hooker: Oliver, Trans. N.Z. Inst., 59, 695, 1929.

D. collinum Oliver, l.c., 696.

D. filifolium is close to D. lessonianum, differing mainly in the longer leaves and longer sepals. It is related to D. urvilleanum, but the leaves are shorter and the racemes larger and more definite, urvilleanum having single flowers or a primitive raceme of 2 to 4 flowers. These differences were pointed out by Richard when describing urvilleanum. With a much larger series of specimens than I had in 1928 I am unable to maintain my collinum as a separate species.

Different habits of growth are shown by members of the species filifolium in different localities. They may be defined as varieties.

Var. centrale var. nov. Frutex, foliis parvis, 40–80 mm. longis, racemis parvis, paucis floribus parivis. These small, upright, rather dense shrubs are common in scrub and bog. Leaves filiform, 40–80 mm. long. Flowers small in few-flowered racemes. Type from Mount Ruapehu collected by W. R. B. Oliver, February, 1949, in Dominion Museum.

Var. filifolium. This is the typical form very common on Mount Egmont and on the Ruahine and Tararua Ranges. Leaves long, up to 200 mm. Racemes rather large with more and larger flowers than centrale.

Var. collinum. D. collinum Oliver, l.c. In the South Island the leaves are of moderate length and the flowers, which have broad bracts, are borne in conspicuous clusters of racemes.

Type of D. filifolium from Ruahine Range collected by W. Colenso, about 1850, in Kew Herbarium. Type of D. collinum from mountains above Tinline Valley, collected by J. H. McMahon, November, 1924, in Dominion Museum.

Distribution: Var. centrale.—Ruapehu, Tongariro and Hauhungatahi mountains. Waimarino Plain. Ohakune (in bog). Var. filifolium.—The northern limits of its distribution are indicated by the following localities: Mount Kakaramea, Maungapohatu, Kaimanawa Range. Generally distributed on Mount Egmont and the Ruahine, Tararua and Rimutaka Ranges to Cook Strait. Var. collinum.—D'Urville Island, Mount Stokes, Mount Arthur, mountains above Tinline Valley, Mount Fyffe, Dun Range, West Wanganui Inlet, Whangamoa Saddle, Pelorus River, Tophouse, St. Arnaud Range.

Dracophyllum longifolium (Forst.) R. Br.

Dracophyllum longifolium (Forster) R. Brown: Oliver, Trans. N.Z. Inst., 59, 701, 1929.

To the synonymy given in my first account of the genus add: Dracophyllum cockayneanum DuRietz, Svensk. Bot. Tidskr., Bd. 24, 374, 1930.

This species is separated from all others of the genus by the following combination of characters: (1) Juvenile leaves long and broad, 180 × 6 mm. (Ruahine), 160 × 6 (Arthurs Pass), 225 × 6 (Tuhua), 227 × 7 (Milford Sound), 185 × 7 (Caswell Sound), 240 × 10 (Stewart Island), 220 × 12 (Auckland Island). (2) Adult leaves long and tapering, much narrower than juvenile leaves: 130 × 2·5 (Ruahine), 140 × 2 (Arthurs Pass), 140 × 2·5 (Tuhua), 232 × 3 (Milford Sound), 96 × 1·6 (Caswell Sound), 230 × 4 (Stewart Island), 215 × 6 (Auckland Island). (3) Flowers in racemes, often curving a little downwards, 40–50 mm. long.

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The nearest allies of D. longfiolium are the species included in the urvilleanum group, from which longifolium is easily separated by the larger, wider leaves in both juvenile and adult stages.

On account of the varying sizes of the leaves it is difficult to define subspecies, but recognisable forms are found in different stations. These may be described as varieties as follows:

(1) Var. septentrionale var. nov. Frutex. Folia juvenilia 180 × 6 mm., matura 130 × 2·5 mm. This is a distinct form confined to a locality far removed from the other members of the species. The racemes are small like those of filifolium, which species is found both to the north and south, but the leaves of septentrionale are wider than those of filifolium at all stages.

Type from Mount Maharahara, collected by R. M. Greenwood in Olearia colensoi scrub, June, 1946, in herbarium of Botany Division, Department of Scientific and Industrial Research, Wellington.

Distribution. Ruahine Range: Mount Maharahara; first collected by D. Petrie on Ruahine Range, 4,000–4,500 feet, January, 1889 (Dom. Mus.).

(2) Var. longifolium (Forst.) R. Br. The typical variety is a rather slender tree with long and broad juvenile leaves and long adult leaves which, along the shores of the Fiordland Sounds and in Stewart Island, hang downwards on slender brachlets. The measurements of leaves given above for the localities Arthurs Pass, Tuhua, Milford Sound and Stewart Island refer to this variety.

Distribution. South Island: generally distributed. Mount Arthur, Dun Mountain, Mount Tuhua, near Greymouth, Arthurs Pass, Mount Peel (Canterbury), Swampy Hill, Colac Bay, Bluff Hill, Milford Sound, Caswell Sound, Doubtful Sound, Dusky Sound, Preservation Inlet, Stewart Island.

(3) Var. pluviale var. nov. Arbor vel frutex. Folia juvenilia 185 × 7 mm., matura 96 × 1·6 mm. The distinguishing characters of this variety are the erect form with small strict leaves and straight erect racemes. The trunk is sometimes 30–40 cm. in diameter, though the tree is not tall, 2–4 m. Compared with the typical variety, both the juvenile and adult leaves are short. In the high country, 2,000 to 3,000 feet above Caswell Sound, it grows in boggy low forest.

Type from Leslie Valley, Caswell Sound, collected by W. R. B. Oliver, March 29, 1949, in Dominion Museum.

Distribution. Wet regions of the west of the South Island, south of S. lat. 43°. Caswell Sound, Wilmot Pass, Waiho Valley, Alex Knob, Mount Moltke, Takahe Valley.

(4) Var. cockayneanum (DuRietz). This variety was first defined, but not named, in my first account (p. 702). The broad, stiff and spear-like leaves with broad sheaths mark it off from the other varieties. DuRietz gave it the name cockayneanum with full specific rank, but as the size of the leaves of the Auckland Island plants varies a good deal and those of the Campbell Island plants are smaller, I think that it is better treated as a variety of longifolium.

Type from Campbell Island collected by W. R. B. Oliver, April 1, 1927, in Dominion Museum. (280.)

Distribution: Auckland Island, Campbell Island.

Dracophyllum oliveri DuRietz

Dracophyllum longifolium form (3), Oliver, Trans. N.Z. Inst., 59, 762, pl. 84, fig. b, 1929. D. oliveri DuRietz, Svensk. Bot. Tidskr., Bd. 24, 374, 1930. Simpson, Trans. Roy. Soc. N.Z., 75, 191, 1945.

– 12 –

In my first paper on Dracophyllum, I included this form under D. longifolium, briefly describing and figuring a specimen from Mount Rochfort. I now separate it from longifolium on account of the narrow juvenile leaves. Simpson has described plants, which he refers to D. oliveri, from Lake Manapouri. These have rather shorter leaves than the typical form from Mount Rochfort. The typical form may briefly be characterized as follows Shrub, 1–2 m tall. Leaves subulate, 50–75 mm. long, 1 ½–2 mm. wide above sheath, rigid, concave above, surface and margins minutely pubescent, sheath with wide-angled, ciliate shoulder. Racemes in clusters, 8–10 flowered, on short lateral branches. Bracts early deciduous. Sepals 6 mm. long, ovate, acuminate, glabrous or ciliate on margin Corolla with acute lobes.

There are differences in the habit in different localities. On Mount Rochfort the branches are rather spreading West Wanganui Inlet and Charleston specimens are more strict. The southern form, found near Lakes Te Anau and Manapouri, is smaller and the leaves are shorter, 30–50 mm. long, 2 mm broad.

Type from Mount Rochfort, 1,800 feet, collected by D. Petrie, February 5, 1913, in Dominion Museum.

Distribution. Mostly collected in North-west Nelson, but probably widely distributed on both sides of the Southern Alps. West Wanganui Inlet, Denniston Plateau and Mount Rochfort, Charleston district, Mount Davy (southern end Paparoa Range), east side of Lakes Te Anau and Manapouri, Mossburn.

Dracophyllum adamsii Petrie

Dracophyllum adamsii Petrie. Oliver. Trans N.Z. Inst. 59, 702, 1929.

The habit of this species suggests an alliance with D. viride, but the juvenile leaves are much narrower and more flaccid than in that species, and the adult leaves are narrower and longer. The sepals differ from all the members of the group of D. sinclairii in their short, broad form. It is best kept in a species group by itself.

Type from Awatere River collected by D. Petrie, January, 1897, in Dominion Museum.

Distribution. Additional localities. Mount Makarako (kaimanawa Range), Waiotapu, Wharekia (East Cape district).

Group of Dracophyllum sinclairii

The distinguishing features of this group are (1) the large juvenile leaves, (2) single racemes terminating leafy branches. To the three species, sinclairii, patens and viride, included in my former account (p 697). I now add arboreum.

Dracophyllum sinclairii Cheesem.

Dracophyllum sinclairii Cheeseman Oliver, Trans. N.Z. Inst, 59, 698, 1929 (D. squarrosum, not of R. Brown).

In my former account, I adopted Hooker's name squarrosum because Brown's squarrosum was founded as Epacris squarrosa and at the time Hooker wrote, 1844, had been transferred from Dracophyllum and placed in a separate genus, Sphenotoma, which name was introduced by Brown as a section of Dracophyllum. Dr. Summerhayes, in a letter to Mr. H. Carse, in 1930, points out that D. squarrosum was validly published by Brown and therefore Cheeseman was justified in rejecting Hooker's D. squarrosum. With this I regretfully agree and so restore Cheeseman's name sinclairii, about which there can be no confusion.

– 13 –

Type of D. squarrosum Hooker, and consequently of Cheeseman's D. sinclairii, from “Manukau Bay” (i.e. Green Bay, Manukau Harbour), collected by W. Colenso, in Kew Herbarium.

Distribution. Additional localities: Great Barrier Island (Kirk. 1867). Mercury Bay.

Dracophyllum patens Oliver

Dracophyllum patens Oliver, Trans. N.Z. Inst, 59, 698, 1929.

The broad, robust, dagger-like adult leaves are very different from the small, narrow leaves of D. sinclairii. In habit it is nearer D. viride, but that species has thin, grass-like leaves, and is a fair-sized tree. Actually the leaves are more like those of D. strictum, but the inflorescence is entirely different, being racemose in patens and paniculate in strictum.

Type from summit of Mount Hobson, Great Barrier Island, collected by W. R. B. Oliver, December 8, 1916, in Dominion Museum.

Distribution: Great Barrier Island. Mount Hobson, Whangaparapara. Table Mountain, Coromandel Peninsula.

Dracophyllum arboreum Ckne.

Dracophyllum arborcum Cockayne: Oliver, Trans. N. Z. Inst., 59, 694, 1929.

In the paper quoted I placed this species in the group of D. scoparium, mainly because of the ciliate leaf margins. Its life history is, however, entirely different from that of D. scoparium or D. palydosum. It passes through a large-leaved juvenile stage to an adult stage in which the leaves are long like those of D. longifolium. Its pubescence is quite like that of D. paludosum. It is probably best included in the group of D. sinclairii, with which species it has a general resemblance and similar life history.

Type from Chatham Island, collected by L. Cockayne, 1901, in Dominion Museum.

Dracophyllum viride Oliver

Dracophyllum viride Oliver, Trans. N.Z. Inst., 59, 699, 1929.

The candelabra-like habit of the tree and its thin leaves reacall D. adamsii. It differs in the larger, broader leaves and especially in the short flowering branches and much longer flowers with long acuminate sepals. It is allied to D. sinclairii.

Type from Spirits Bay, collected by W.R.B. Oliver, November 28, 1916, in Dominion Museum.

Distribution. Additional localities: Tapotopoto Bay, Kaitaia, Cape Reinga.

Dracophyllum recurvum Hook. f.

Dracophyllum recurvum J. D. Hooker: Oliver, Trans. N.Z. Inst, 59, 700, 1929.

Type from Mount Tongariro, collected by J. C. Bidwill, in Kew Herbarium.

Distribution. Additional localities: Wannarino Plain, Kaimanawa Range, Thames Goldfield (Kirk), Mount Karioi (near Raglan). Mount Pihanga, Rangipo Plam, Mount Hauhungatahi, Mount Kakaramea.

Dracophyllum menziesii Hook. f.

Dracophyllum menziesn J. D. Hooker: Oliver, Trans N.Z. Inst, 59, 705, 1929.

Type from Dusky Sound, collected by A. Menzies, 1791, in Kew Herbarium.

– 14 –

Distribution. Additional localities: Upper Hollyford Valley, Matukituki Valley, Mount Alexander (Caswell Sound).

Dracophyllum townsoni Cheesem.

Dracophyllum townsoni Cheeseman: Oliver, Trans. N.Z. Inst., 59, 705, 1929.

Type from Mount Buckland, Paparoa Range, collected by W. Townson, in Auckland Museum.

Distribution. Additional localities: Lead Hill, Mount Burnett, West Wanganui Inlet, Mount Davy (Paparoa Range), north of Greymouth, Mount Mantell.

Dracophyllum fiordense Oliver

Dracophyllum fiordense Oliver, Trans. N.Z. Inst., 59, 705, 1929.

On Mount Alexander, above Caswell Sound, in upper beech forest, there are trees of this species 3 m. tall, with trunks 10 cm. in diameter, while in the tussock country at 4,000 feet they are reduced to shrubs ½ m. tall.

Type from Wilmot Pass, collected by W. R. B. Oliver, March, 1927, in Dominion Museum.

Distribution. Additional localities. Mount Balloon, McKinnon Pass, Sutherland Falls, Milford Sound, mountains above Caswell Sound, Alex Knob (Waiho Valley).

Dracophyllum strictum Hook. f.

Dracophyllum strictum J. D. Hooker: Oliver, Trans. N.Z. Inst., 59, 707, 1929.

Type from Mount Tongariro, collected by J. C. Bidwill, in Kew Herbarium.

Distribution. Additional localities: Tarawera Mountain (Kirk), Wairoa (Tauranga), Mayor Island, Hendley (Hawkes Bay), Lake Rotoiti, Whakarewarewa, Kaitaringa, Mamaku, Waimangu. Kaipikari (inland from Urenui), Waverley, Mount Rochfort (Townson).

Dracophyllum secundum (Poir) R. Br.

Dracophyllum secundum (Poiret) R. Brown: Oliver, Trans. N.Z. Inst, 59, 707, 1929.

Haptotype from Port Jackson, R. Brown, in Melbourne Herbarium.

Distribution. Additional localities: Cowan Creek, Wolgan, Lithgow, Port Jackson, Mount Wilson, Bradwood District.

Dracophyllum ramosum Br. and Gris

Dracophyllum ramosum Brongniart and Gris: Moore, Jour. Linn. Soc. Bot., 45, 349, 1921.

Oliver, Trans. N.Z. Inst., 59, 707, 1929. Daniker, Vier. Nat. Gess. Zur., 78, 341, 1933.

Type from M'bee, collected by Vieillard, in Paris Museum. (No. 830.)

Distribution. Additional localities: Mount Dore, River Dumbea, Port Bouquet, Taom (all recorded by Moore); Mount Humboldt and Mount Koghi (Daniker), Hermitage.

Plate 94 of my former paper was inadvertently labelled New South Wales. It should be Baie du Sud. New Caledonia.

Dracophyllum vieillardii Lenorm.

Dracophyllum vieillardii Lenorm: Oliver, Trans. N.Z. Inst., 59, 708, 1929.

Type from Dombea, collected by Vicillard, in Paris Museum. (283b.)

Dracophyllum amabile Br. and Gris

Dracophyllum amabile Brongniart and Gris: Oliver, Trans. N.Z. Inst., 59, 708, 1929.

Type from mountains near Kanala, collected by Vieillard, 1861–1867, in Paris Museum. (829.)

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Distribution Additional locality: Prony.

Concerning the last three species, Professor A. Guillaumin has written to me as follows “Je suis persuadé—mon examen des plantes sur place me le confirme—que D. amabilc et D. vieillardii ne constitutent qu'une seule espèce avec D. ramosum, les rameaux végétatifs portant d'énormes bouquets de grandes fenilles et les florifères de toutes petites feuilles.”

Dracophyllum alticolum Daniker

Dracophyllum alticolum Daniker, Viert. Nat Ges. Zur., 78, 339, 1933.

The following description is abridged from the original diagnosis. A small prostrate shrub, the branches marked with annular leaf scars. Leaves densely clustered at the ends of the branches, elongate-triangular, 5–6 by 1 ½ cm., the lower end contracted. slightly sheathing, narrowing towards the obtuse apex, glabrous, coriaceoups, striate. Inflorescence racemose, terminal, 11–19 cm. long. the base protected by 5–6 short, imbricating leaves; pedicels of the flowers pubescent. Flowers in threes, 8–10 mm. long, pedicels 10 mm. long. Sepals triangular, acute, 10 by 2 ½ mm. pubescent outside, glabrous within. Corolla and stamens not seen. Ovary 4 by 3 mm., pubescent, 5-locular, style 4 mm.

This prostrate mountain Species belongs to the group of D. secundum, and perhaps comes nearest to D. vicillardii, agreeing in the leaves being shorter than the panicle and the flowers being disposed in threes. In alticolum the three pedicels are separate, in vieillardii they have a short common peduncle. In both species they are densely pubescent. In alticolum the leaves are very much broader than they are in vieillardii.

Type from Mount Humboldt, collected by A. U. Daniker, November 5, 1924, in University of Zurich. (D509.)

Distribution: New Caledonia; Mount Humboldt, 1,300 m.

Dracophyllum cosmelioides Oliver sp. nov.

Dracophyllum gracile Brongniart and Gris, Ann. Sci. Nat. Bot., 2. 156, 1864 (not D. gracile R. Br.). Oliver, Trans. N.Z. Inst, 59, 708, 1929. Daniker, Vier. Nat. Ges. Zur., 78, 340, 1933.

Brongniart and Gris' name gracile is rejected because of Brown's action in transferring Poiret's Epacris gracilis to Dracophyllum (1810). though this species is now accepted as belonging to Sphenotoma. A new name is accordingly provided, namely, the manuscript name of Pancher quoted by Brongniart and Gris as a synonym of their gracile.

Type of D. gracile from Lac Arnaud, New Caledonia, collected by Vieillard, in Paris-Museum. (828.)

Distribution. Additional localities: River Lacs, River Yate, Prony.

Dracophyllum thiebautii Br. and Gris

Dracophyllum thiebautii Brongniart and Gris: Oliver, Trans. N.Z. Inst., 59, 709, 1929.

Type from mountains near Arama, New Caledonia, collected by Thiebaut, 1865, in Paris Museum.

Dracophyllum sayeri F. v. Muell.

Dracophyllum sayeri F. von Mueller: Oliver, Trans. N.Z. Inst, 59, 710, 1929.

Type from Mount Bartle Frere, S. Johnson, 1892, in Melbourne Herbarium.

Distribution. Additional locality: Junction of Roots Creek and Mossman Falls, Mount Bartle Frere.

– 16 –

Dracophyllum dracaenoides Schltr.

Dracophyllum dracaenoides Schlecter: Oliver, Trans. N.Z. Inst., 59, 710, 1929.

Type from mountains near Ou Hinna, New Caledonia. I am unable to state where the type specimen is preserved.

Dracophyllum milligani Hook. f.

Dracophyllum milligani J. D. Hooker: Oliver, Trans. N.Z. Inst., 59, 710, 1929.

Type from Mount Sorrel, Macquarie Harbour. Tasmania, J. Milligan, in National Herbarium, Melbourne. (No 747.)

Distribution. Additional localities. Bathurst Harbour, Port Davey.

Dracophyllum fitzgeraldi Moore and Muell.

Dracophyllum fitzgeraldi Moore and Mueller: Oliver, Trans. N.Z. Inst., 59, 711, 1929.

Type from Lord Howe Island, collected by R.D. Fitzgerald, in Melbourne Herbarium. (No. 58, 1869.)

Dracophyllum latifolium A. Cunn.

Dracophyllum latifolium A. Cunningham: Oliver, Trans. N.Z. Inst., 59, 711, 1929. Kirk, Forest Fl. N.Z. 251, 1889. D. recuriatum Colepso, Trans. N.Z. Inst., 21, 92, 1889.

The type specimen of Colenso's D. recurvatum, preserved in the Dominion Museum, is D. latifolium. The original description of the panicle agrees with that of D. latifolium.

Type of D. latifolium from Kawakawa River, Bay of Islands, collected by A. Cunningham, 1826, in British Museum. Type of D. recurvatum from Lake Waikaremoana, collected by H. Hill, 1888, in Dominion Museum.

Distribution. Additional localities: Great Barrier Island, Waipoua, Kaiaka, Mount Tamahunga, Hunua Range. Te Moehau, Port Charles, Mount Pirongia, Whetu Matarau (East Cape district).

Dracophyllum matthewsii Carse

Dracophyllum mattheusii Carse: Oliver, Trans. N.Z. Inst., 59, 712, 1929.

Type from Taumatamahoe, near Kaitaia, 1,900 ft., collected by H. B. Matthews. October 2, 1913, in Canterbury Museum.

Distribution. Additional localities: Great Barrier Island, between Ahipara and Herekino, Port Charles.

Dracophyllum traversii Hook. f.

Dracophyllum traversii J. D. Hooker: Oliver, Trans. N. Z. Inst., 59, 712, 1929.

Type from Nelson Province, collected by W. T. L. Travers, in Kew Herbarium.

Distribution. Additional localities: West Wanganui Inlet, Taipo River (Nelson), Kelly's Hill. Mount Rochfort.

Dracophyllum pyramidale Oliver sp. nov.

Dracophyllum recurvatum Oliver, Trans. N.Z. Inst, 59, fig. 23, 712, 1929 (not of Colenso).

Type of Colenso's D. recurvatum is now preserved in the Dominion Museum and is D. latifolium. Colenso's name recurvatum consequently falls into the synonymy of latifolium, and a new name is accordingly required for this very fine species. The name pyramidale refers to the stout, upright, pyramidal panicle, a very conspicuous feature of this species. Its nearest relative is D. traversii.

Type from Little Barrier Island, collected by W. R. B. Oliver, October 8, 1928, in Dominion Museum.

– 17 –

Distribution. Additional localities: Tc Moehau, Mount Pirongia, Mount Tamahunga. The Waikaremoana locality given in my former account (p. 713) should be omitted.

Dracophyllum verticillatum Lab.

Dracophyllum verticillatum La Billardiere: Oliver, Trans. N.Z. Inst., 59, 713, 1929. Brongn, and Gris, Ann. Sci. Nat. Bot., 2, 157, 1864. Damker, Vier, Nat. Ges. Zurich, 78, 341, 1933.

Type probably at Florence, where Labillardiere's herbarium is preserved.

Distribution. Additional localities: River Pirogue, River Nomatch, Aramagipfel, Mount Koniambo, Mount Pame, Mount Koghi, Prony.

Dracophyllum involucratum Brongn. and Gris

Dracophyllum involucratum Brongmait and Gris: Oliver, Trans. N.Z. Inst, 59, 714, 1929. Daniker, Vier. Nat. Ges. Zur., 78, 340. 1933. Probably D. compactum Moore is a synonym (see next species).

Type from mountains near Yate. collected by Vieillard, in Paris Museum. (832).

Distribution. New Caledonia: Additional localities Yate, Plame des Lacs, Mount Panie.

Dracophyllum compactum Moore

Dracophyllum compactum Moore, Jour. Linn. Soc. Bot., 45, 349, 1921.

The following description is abridged from the original diagnosis. A tall shrub, almost 4 m., trunk sparsely branched. Leaves 30 by 2.5 cm., imbricate, elongate-lanceolate, acummate, rigid, glabrous, with sheathing base. Flowers arranged in compact whorls, on a strong, upright, hirsute rachis, 30 cm. long; peduncles one-flowered, entirely covered with imbricated bracts; bracts 2.5–4 cm. long, ovate-lanceolate, acute, ciliate. Sepals 3 mm. long, lanceolate, acute, ciliate. Corolla white, tube scarcely exceeding the calyx, 4 mm. long, lobes oblong, obtuse. Filaments adnate to the corolla tube, anthers 1.5 mm. long. Ovary depressosubglobose, 1 mm. in diameter. Stigma minute, 5-lobed.

Moore compares this species with D. verticillatum thus: “readily known from it by reason of the strictly racemose inflorescence, peduncles covered with imbricating bracts, the narrow bracts and sepals, and rclatively shorter corolla.” These are precisely the characters by which involucratum differs from verticillatum. Furthermore. Moore's whole description fits D. involucratum, and the type locality. Plame des Lacs, is one of the localities given by Daniker for D. involucratum. From the description it would appear, therefore, that D. compactum is the same as D. involucratum, but I have not had an opportunity of comparing specimens, so list them separately for the present.

Type from Plaine des Lacs, collected by R. H. Compton, 1914, in British Museum. (371.)

Distribution. New Caledonia: Plaine des Lacs.

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A Skeleton of Notornis hochstetteri Meyer from Waitati, Otago, And Notes on Distribution of the Bird in the South Island, N.Z.

Department of Zoology, University of Otago*

[Read before the Nelson Philosophical Society, October 15, 1950; received by the Editor, May 28, 1951]


An incomplete skeleton of Notornis hochstetteri found at Waitati and now in the Otago Museum is described and figured. A list is given of all localities in the South Island from which Notornis remains have been recorded, also a list of localities of hitherto unrecorded bones in the Dominion, Canterbury, Otago and Invercargill museums.

A hitherto undescribed, incomplete skeleton of Notorms hochstetteri Meyer in the Otago Museum has recently come to my notice. The only available information about its origin is that it was found in sand at Waitati and was presented to the Museum by T. H. Murdoch. The skull, lower jaw, pelvis, femur and two tibiae were registered in 1913 and the tarso-metatarsus in 1914; presumably the tarso-metatarsus was overlooked in 1913 and was found in 1914 on revisiting the locality.

The skull is well preserved, but lacks the quadrate, quadrato-jugal, jugal, pterygoids and prevomers; these easily displaced bones apparently became separated from it in the sand and were overlooked by the collector. The following are the measurements of the various structures:

Total length from post. surface of occipital condyle to ant. tip of beak 94.0mm.
Greatest breadth 39.5
Least width between temporal fossae 16.5
Tip of beak to its junction with the frontals 63.5
Least width between external nostrils 9.5
Greatest depth of beak from junction of frontals to post. angle of maxilla 30.0
Lower Jaw:
Total length of right ramus 87.0
Greatest vertical height at coronoid process 22.0
Length of symphysis 36.0
Greatest width at post. articular expansion 45.0
Width of right post. articular expansion 15.5
Length 101.5
Breadth of proximal end along axis of neck 20.5
Breadth of distal end 21.5
Circumference middle of shaft 25.1
Length 156.5
Breadth of proximal end 28.0
Breadth of distal end 17.0
Circumference of shaft 22.0

[Footnote] * Now at Entomological Research Station, Cawthron Institute, Nelson.

– 20 –
Length 93.0
Breadth of proximal end (transverse) 17.5
Breadth of proximal end (antero-posterior) 18.0
Breadth of distal end 19.5
Breadth of shaft 8.5
Greatest length 115.0
Greatest width 52.5
Width of sacrum 23.5

The bones show no sign of fire or other indication that they may be midden material, and it appears that the bird died a natural death where it was found. This then is another record of its distribution in the South Island.

Since N. hochstetteri Meyer is considered to be confined to the South Island, and to be distinct from N. mantelli Owen, which appears to have been confined to the North Island, it is considered worthwhile to append an up-to-date list of localities from which Notornis remains have been recorded in the South Island. This is done on the assumption that all Notornis remains in the South Island are referable to N. hochstetteri.

Date found Reference to literature
1849 Duck Cove, on Resolution Island, Dusky Sound Mantell 1850
1851 Secretary Island, Thompson Sound Buller 1873
1879 Mararoa Flat, Te Anau Parker 1881
* 1881 Cascade Creek (now Cascade Burn) Park 1888
* 1881 Matukituki River, South branch Park 1888
1884 Patience Bay, Te Anau Parker 1885
*1888 Mt. Hodge, Dusky Sound Park 1888
*1888 Docherty's Creek, Dusky Sound Park 1888
*1888 Mt. Pender, Dusky Sound Park 1888
*1888 Cooper's Island, Dusky Sound Park 1888
*1888 South side of Dusky Sound opposite Cooper's Island Park 1888
1892 Castle Rocks, Southland Hamilton 1892
1892 Earnscleugh, Central Otago Hamilton 1892
1892 Long Beach, near Dunedin Hamilton 1892
1898 Middle Fiord, Te Anau Benham 1899
1948 Takahe Valley, Te Anau Falla 1948

Dr. R. A. Falla has furnished me with a list of Notornis bones in the collection of the Dominion Museum, from which the following new records were obtained:

Part skeleton collected by Major Paton 1872 in a limestone cave, Aniseed Valley, Nelson. This skeleton is on loan from the Nelson Museum. Various leg bones labelled by A. Hamilton as from Ngapara, North Otago, no date, and Warrington, Otago, no date.

From Dr. R. S. Duff a list of the Canterbury Museum collection, including new records from Kapua, Waimate, a tarso-metatarsus labelled by F. W. Hutton 1895. Various leg bones, a mandible and premaxilla from Pyramid Valley, North Canterbury, collected by Scarlett, Eyles, Duff and Falla between 1940 and 1949.

Dr. H. D. Skinner, Otago Museum, informs me that the recently acquired J. Murray collection of bird bones from limestone caves at Forest Hill, Southland, includes several crania, mandibles, pre-maxillae and a tarso-metatarsus of Notornis.

[Footnote] * These records of Park's were challenged by Melland (Trans N. Z. Inst, 22, pp. 295–300), but from my own experience of the habits of Notornis and its unmistakable “booming” note it seems reasonably certain that bis records are correct.

Picture icon

Skull of Notornis hochstctteri found at Waitati, Otago.
Top: Ventral view. Bottom: Dorsal view.

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Skeletal remains of Notornis hochstetteri found at Waitati, Otago.

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Distribution Map of Occurrences of Notornis hochsletteri in the South Island of N.Z.

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Mr. David Teviotdale supplied me with a list of the Notornis material held in the Invercargill Museum, a skull from Wakapatu, 1938, premaxilla and lower mandible from Pahia, 1933, lower mandible, Tokonui Mouth, 1939, all collected by J. H. Sorensen. A skull from Back Beach, Greenhills, 1942, collected by Robert Gibb, pelvis and sternum from caves, Limehills, 1928, collected by Messrs. George jaquiery, Fowler and Woods, and a skull and upper mandible from Wakapatu, 1946, collected by D. Teviotdale.

Falla (1948) states “Sub-fossil remains from Te Aute and Martinborough in the North Island, and Waikari, Enfield, Castle Rocks and other South Island localities …” Waikari (Pyramid Valley) and Castle Rocks are referable to records mentioned above, whilst Enfield, Dr. Falla (in litt.) informs me, refers to information gleaned from H. O. Forbes' records.

Consideration of the twenty-nine localities recorded above shows that Notornis was distributed from one end of the island to the other. The frequency of the sub-fossil remains found in an area is not necessarily a true indication of the population density of the bird, but depends rather on the suitability of the area for preservation of remains, and the work done there by collectors. The preponderance of remains and the bird's present existence in the Southland-Otago area indicates, however, that it occurred there in greater numbers than elsewhere.


My thanks are due to the Directors of the Dominion, Canterbury and Invercargill Museums for the information acknowledged in the text, and especially to the Director of the Otago Museum, Dr. H. D. Skinner, for information supplied and permission to describe the Waitati specimen, and to Mr. R. Blick, Cawthron Institute, for photographs.


Benham, W. B., 1899. Notes on the Fourth Skin of Notornis. Trans. N.Z. Inst., vol. 31, pp. 146–150.

Buller, W. L., 1873. History of the Birds of New Zealand. London, 1st Edition.

Falla, R. A., 1948. The Re-discovery of the Takahe. N.Z. Sci. Rev., vol. 6, pp. 123–124.

Hamilton, A., 1892. On the Fissures and Caves at the Castle Rocks, Southland; with a Description of the Remains of the Existing and Extinct Birds found in them. Trans. N.Z. Inst., vol. 25, pp. 88–106.

Mantell, G. A., 1850. Notice of the Discovery by Mr. Walter Mantell in the Middle Island of New Zealand, of a Living Specimen of the Notornis, a Bird of the Rail Family, Allied to Brachypteryx, and hitherto unknown to Naturalists except in a Fossil State. Proc. Zool. Soc. Lond., part XVIII, pp. 209–212.

Park, James. The Takahe (Notornis mantelli) in Western Otago. Trans. N.Z. Inst, vol. 21, pp. 226–230.

Parker, T. J. On the Skeleton of Notornis mantelli. Trans. N.Z. Inst, vol. 14, pp. 245–258.

Parker, T. J. Notes on a Skeleton of Notornis, recently acquired by the Otago University Museum. Trans. N.Z. Inst, vol. 18, pp. 78–82.

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Studies on the Earthworm Fauna of New Zealand. III

Department of Scientific and Industrial Research, Wellington

[Read before the Wellington Branch, May 30, 1951; received by the Editor, May 30, 1951]


This paper describes the New Zealand species of the Subfamily Megascolecinae, and includes a general discussion on their phylogeny and classification. The following new species are described and figured. Megascolides viridis, Motuhora: M. ruber, Mangamuku Range; M. parvus, Hicks Bay; M. raglani, Raglan Range; M. fuscus, Rotorua district; M. irregularis, Tangowahine; M. alba, Maungakaramea; Megascolex novaezealandiae, Mangonui; Pheretima campestris, Kaitaia; Diporochaeta obtusa, Rotorua; Perionyx egmonti, Mount Egmont.

The Sub-family Megascolecinae

The sub-family Megascolecinae was established by Michaelsen (1900) as a division of the family Megascolecidae. As defined by Stephenson (1930) it includes all those species of earthworms referable to the family Megascolecidae and having the following characters:

“Chaetae either eight per segment, or numerous, and then either in regular chains or approximated in couples. Clitellum beginning with or in front of xiv. Male pores on xviii. Spermathecal pores, if present, one to seven pairs, in front of the testis segments. Usually one gizzard in front of the testis segments, sometimes two or three, exceptionally none. Mega- or micronephridial. Two pairs of testes and funnels in x and xi, or only one pair. Prostates one pair, tubular or racemose (Pheretima—prostates), each prostatic duct uniting with the vas deferens of the same side and opening in common with it (except in Diplotrema).”

The occurrence of the sub-family Megascolecinae in New Zealand raises some interesting zoogeographical considerations. The “root genus” of the sub-family is taken as Diplotrema, which is found in Queensland and New Caledonia, and from Diplotrema all the other genera have been derived. The genera of the sub-family are now found for the most part in the Indian, Malayan, Oriental and Australasian regions and it is reasonable to assume that any general migration of Megascolecinae into New Zealand must have come from the north. A northern origin for the group is also indicated by its distribution within New Zealand, since nearly all the species are found only in the northern regions of the North Island. The entry of the Megascolecinae into New Zealand would require the postulation of a former land connection of the northern portion of the North Island either directly to Australia or to New Caledonia and the Indo-Malayan islands. Such a land connection need not have been continuous at any one time; it would be sufficient that at times portions of the land connection should have been emergent while other portions remained submerged, and the Megascolecinae were able to progress in stages until they eventually reached New Zealand.

Thirty species, belonging to ten genera of the sub-family, have been identified from the New Zealand region by previous workers and eleven new species, belonging to five of the ten genera, are described in this paper. The following is a list of the ten genera known to occur in the New Zealand region, with notes on their distribution within the region:

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Plutellus Perrier (1873) Auckland Islands.
Pontodrilus Perrier (1874) Lake Wakatipu, South Island; Chatham Island.
Megascolides McCoy (1878) northern North Island.
Notoscolex Fletcher (1886) northern North Island (possibly introduced); Poor Knights Island.
Diporochaeta Beddard (1890) North Island; western South Island; Chatham Island.
Spenceriella Michaelsen (1907) northern North Island; Little Barrier Island.
Megascolex Templeton (1844) northern North Island; Norfolk Island.
Pheretima Kinberg (1866) northern and eastern North Island; Raoul Island (probably introduced).
Perionyx Perrier (1872) western North Island; Auckland Islands; Snares Island.
Didymogaster Fletcher (1886) distribution unknown; a single species, probably introduced.

None of these genera is peculiar to the New Zealand region. The genera may be identified from the following key:

a1 Eight chaetae on each segment
b1 Purely meganephridial
  c1 Gizzard vestigial or absent Pontodrilus
  c2 Gizzard well developed Plutellus
b2 Micronephridial, at least in part
  d1 Prostate tubular, with single continuous central duct Megascolides
  d2 Prostate racemose, with branching system of ducts
    e1 Single gizzard in v or vi Notoscolex
    e2 Two gizzards, one in vi and one in vii Didymogaster
a2 More than eight chaetae on each segment
f1 Meganephridial
  g1 Prostate tubular, with single continuous central duct Diporochaeta
  g2 Prostate racemose, with branching system of ducts Perionyx
f2 Micronephridial
  h1 Gizzard in v, vi or vii
    i1 Prostate tubular, with single continuous central duct Spenceriella
    i2 Prostate racemose, with branching system of ducts Megascolex
  h2 Gizzard between septa vii/viii and x/xi Pheretima

The relationships of the New Zealand genera are shown in the diagram below, commencing from the “root genus” Diplotrema:

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Genus Plutellus Perrier

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

“Chaetae eight per segment. Male pores paired or single; female pores mostly paired; spermathecal pores end at furrow 8/9 or on segment ix, a single pair or a series of two to five pairs, or five single pores. A gizzard in the region of segments v–vii. Purely meganephridial. Prostates tubular, with simple unbranched duct.” (Stephenson, 1930.)

A single species, P. aucklandicus Benham (1909), is recorded from the New Zealand region. The species was confirmed by Michaelsen (1923), who noted that short side ducts open into the prostatic duct and consequently the species would perhaps go into the genus Woodwardiella. However, Michaelsen decided that the species should remain in Plutellus, since the side ducts are small and the central duct is of more importance (cf. New Zealand species of Megascolides, discussed below).

Genus Pontodrilus Perrier

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

“Chaetae eight per segment. Male pores paired; female pores paired. Spermathecal pores two to four pairs, the last in furrow 8/9. Gizzard vestigial or absent. Purely meganephridial, nephridia wanting in front of the clitellar region. Two pairs of free testes and funnels. Prostates tubular, with simple unbranched canal.” (Stephenson, 1930.)

The most remarkable feature of this genus is that of the five known species, three have a littoral habitat and live in the intertidal zone, while the fourth is limnic and the fifth terrestrial. This is the only known genus in which Megascolecid worms have a littoral habitat.

Two species are known from the New Zealand region. They are P. matsushimensis var. chathamensis Michaelsen, collected from the shore at Chatham Island, and P. laustris Benham, the only known freshwater species, collected at depths of 300–1,200 feet from the waters of Lake Wakatipu in the western South Island. Benham (1900) raised the Chatham Island form to a species, P. chathamensis, but Stephenson (1930) considers it as a variety of P. matsushimensis Michaelsen.

Genus Megascolides McCoy

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

“Chaetae eight per segment. Spermathecal pores one to five pairs, the last in furrow 7/8 or 8/9 or on segment ix. One gizzard in the region of segments v and vi. Micronephridial at least in the anterior part of the body, often throughout. Prostates tubular, with a simple unbranched canal.” (Stephenson, 1930.)

The species here included in Megascolides have been referred by various authors to Notoscolex, Tokea (a genus which has now been discarded), Megascolides, and in one case (M. morlensem) to Megascolex. The differences between Megascolides and Notoscolex are slight and for diagnostic purposes the form of the prostatic duct is relied upon. A typical Megascolides has simple tubular prostates, each with an unbranched central canal running right through the gland and opening directly to the exterior through the prostatic duct, while a typical Notoscolex has polylobate prostates, each with a diffuse system of ducts ramifying through it and frequently opening into a cavity which is connected with the external pore by the prostatic duct. If all the species of Megascolides and Notoscolex had one or other of these “typical” arrangements of the prostatic ducts, there would be no difficulty in placing a species in its correct genus, but there is a great deal of variation and the two types of prostates grade into one another. It would seem desirable to fuse the two genera, but this would produce a genus

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with a very large number of species, and for convenience it is better that the two genera should be retained.

Benham (1904) established a genus, Tokea, for several species from the North Island of New Zealand which had all the characters of Megascolides, but differed from that genus in one respect. The prostate had a simple central canal, but when sections of the gland were examined, it was found that the gland cells were grouped into small clusters which discharged their secretion into a slight evagination of the central duct. Benham called these small evaginations of the central duct “canalicules”. It was unfortunate that he chose this term, since it really means a small canal and was taken as such by other workers, who placed Benham's species in Notoscolex. However, reference to the figures in Miss Sweet's paper on the structure of prostatic glands (1900), or to Benham's paper (1941) on Megascolides napierensis, in which he figures a cross-section of the prostate of Megascolides (Tokea) esculenta and details of the “canalicules”, will show what Benham meant by the term. I have examined sections of the prostatic gland of other genera of the family Megascolecidae and have found that these “canalicules” are present in the prostates of other genera which have tubular prostates, e.g. in Neodrilus they are present in large numbers. The position of Tokea in relation to Megascolides and Notoscolex has been a subject of much discussion and confusion. Stephenson (1930) summed up the position as it appeared to him, but his statements are in many respects contradictory. He believes that Tokea should go into Notoscolex, a view also adopted by Michaelsen, yet in one part of his monograph (p. 658) he refers to the edibility of certain species of Tokea (Megascolides).” There is one character of Tokea which all the previous workers seem to have overlooked in their discussion. That character is the presence of meganephridia in association with the micronephridia in the posterior segments of the body (see Benham's original description of the genus, 1904). Now, according to the accepted relationships of the genera of the Megascolecinae (as set out above), Megascolides is derived from Plutellus by a change from the meganephridial to the micronephridial condition. In many cases the change is not complete and meganephridia are found in association with micronephridia in the posterior segments of Megascolides. Notoscolex is derived from Megascolides by the branching of the prostatic duct and there are few records of species of Notoscolex with meganephridia and micronephridia co-existent in any of the segments. In view of this fact and the questionable existence of branches in the central prostatic canal of Tokea there can be little doubt that Tokea should be included in Megascolides.

Megascolides mortenseni was referred by Michaelsen (1923) to the genus Megascolex, but reference to the original description shows that it is actually a species of Megascolides. M. reptans and M. unipapillatus were referred by Ude (1905) to Notoscolex, but reference to his descriptions shows that they belong to Megascolides, since they have unbranched tubular prostates.

Twelve species of Megascolides have been recorded from the New Zealand region by previous workers and the following is a list of them.

  • M. napierensis Benham (1941).

  • M. (Tokea) maorica (Benham, 1904).

  • M. (Tokea) decipiens (Benham, 1905).

  • M. (Tokea) kirki (Benham, 1904).

  • M. (Tokea) huttoni (Benham, 1904).

  • M. (Tokea) urewerae (Benham, 1904).

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  • M. (Tokea) suteri (Benham, 1904).

  • M. (Tokea) sapida (Benham, 1904).

  • M. (Tokea) esculenta (Benham, 1904).

  • M. (Notoscolex) reptans (Ude, 1905).

  • M. (Notoscolex) unipapillatus (Ude, 1905).

  • M. (Megascolex) mortenseni (Michaelsen, 1923).

Seven new species, M. viridis, M. ruber, M. parvus, M. raglani, M. fuscus, M. irregularis and M. alba are described in this paper.

The two species M. maorica and M. decipiens are distinguished by the number and position of the tubercula pubertatis. I have found these structures to be extremely variable in number and position and consequently the distinction cannot be upheld. and M. decipiens must be considered synonymous with M. maorica.

Three species, M. huttoni, M. urewerae, and M. suteri, cannot be separated according to the descriptions given by Benham. I have discovered two species which agree fairly closely with Benham's descriptions of M. urewarae and M. suteri, and further notes on the two species are included in this paper. I have not found any species which can be referred to M. huttoni, although M. parvus n.sp., described in this paper, is very similar to M. huttoni. M. napierensis was founded on a single specimen collected from a nurseryman's glasshouse at Napier. I have collected many hundreds of specimens of endemic earthworms from the district surrounding Napier and have not found M. napierensis or any species of any other Megascolecine genus among them. I conclude that M. napierensis is to be regarded as introduced into that one glasshouse in Napier with soil or plants and that it may have come from any part of the world where Megascolides is known. M. mortenseni (Michaelsen) was founded on a single incomplete specimen from Palmerston North. No details of its environment are given, so it cannot be definitely decided whether it is introduced or endemic. It has not been recorded again from Palmerston North or from the surrounding district and it is possible that it was, like M. napierensis, an isolated introduced specimen.

The species of Megascolides known from New Zealand may be identified from the following key:

a1 Two pairs of hearts, in xii and xiii M. napierensis
a2 Three pairs of hearts, in x, xi and xii
b1 Two pairs of spermathecae, in viii and ix
  c1 Calciferous glands absent
    d1 Three pairs of vesiculae seminales, in x, xi and xii M. maorica
    d2 Four pairs of vesiculae seminales, in ix, x, xi and xii M. viridis
  c2 One pair of calciferous glands, in xiv M. kirki
  c1 Two pairs of calciferous glands, in xii and xiii M. ruber
  c4 Five pairs of calciferous glands
    e1 Four pairs of vesiculae seminales, in ix, x, xi and xii M. reptans
    e2 One pair of vesiculae seminales, in xii M. unipapillatus
b2 Three pairs of spermathecae, in vii, viii and ix*
  f1 Two pairs of vesiculae seminales. in ix and xii
    g1 Gizzard in v M. parvus
    g2 Gizzard in vi M. suteri
  f2 Four pairs of vesiculae semmales, in ix, x, xi and xii M. urewerae
a2 Four pairs of hearts

[Footnote] * M. huttoni cannot be definitely separated from any species under this heading, due to inadequacy of description.

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h1 Hearts in ix, x, xi and xii M. raglani
h2 Hearts in x, xi, xii and xiii
  i1 One pair of spermathecae M. fuscus
  i2 Two pairs of spermathecae M. irregularis
  i3 Three pairs of spermathecae
    J1 Spermatheca with elongate tubular sac and slender, pyriform diverticulum M. mortenseni
    J2 Spermatheca with stout ovoidal sac and ovoidal or globular diverticulum
      k1 No oesophageal glands M. sapida
      k2 Oesophageal glands as lateral dilations of the oesophagus in xv M. esculenta
a4 Five pairs of hearts, in ix, x, xi, xii and xiii M. alba

The following are descriptions of the seven new species M. viridis, M. ruber, M. parvus, M. raglani, M. fuscus, M. irregularis and M. alba. Types in my collection.

Megascolides viridis n.sp. (Plate 10, figs. 1–3)

Several specimens of this small species were collected from native forest on a mountainside seven miles west of Motuhora, inland from Gisborne. The most striking character of the species is its colour. Dorsally it is reddish-brown and ventrally it is pale bluish-green. The chaetae arise each from the centre of a small raised white spot. The clitellum does not differ in colour from the remainder of the body, but completely surrounds xiv to xvii, obliterating the intersegmental furrows between those segments.

The specimen on which this description is based is 54 mm. in length and 4.5 mm. in diameter and has 66 segments.

The prostomium is prolobous. There are eight chaetae on each segment, arranged in pairs. On xxiv the arrangement is as follows: ab = 1.5 mm.; cd = 2 mm.; bc = 2 mm.; aa = 2.25 mm., dd = 3 mm.

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

There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, a pore in line with chaeta a of each side in each intersegmental furrow. The female pores are on xiv, a pore on each side anterior and slightly medial to chaeta a. The male pores open to the exterior on xviii, one on each side, slightly lateral to the line joining the chaetae a of the adjacent segments. Each pore is on the apex of a small, rounded, white papilla. There are two small, oval, unpaired, median ventral tubercula pubertatis, one at 13/14, and the other at 18/19. No nephridiopores nor dorsal pores are visible.

Internal Anatomy (Plate 10, fig. 2)

The septa vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are thickened and muscular.

Alimentary Canal. The pharynx is small and muscular and occupies the first four segments. There is no proventriculus. The gizzard is slender, but has thick muscular walls, and is situated in v. The oesophagus extends from vi to xiv and there are no oesophageal glands. The intestine commences in xv.

Vascular System. The dorsal blood vessel is unpaired throughout its length. A slender, unpaired, supra-intestinal vessel passes through x–xii and from it arise three pairs of dilated hearts, a pair in each of x, xi and xii.

Reproductive System. There are two pairs of small compact testes, a pair in x and a pair in xi, one on each side close to the ventral nerve cord in each segment. A pair of ovaries occurs in a similar position in xiii. Each ovary consists of a cluster of thread-like structures arising from the anterior septum of xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix.

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In the specimen examined both the spermathecae of ix appear to lie on the right hand side of the ocsophagus, although the ducts open one on each side of the segment, as is normal. The left spermatheca has probably been twisted under the oesophagus by the contortion of the earthworm's body when it was killed. Each spermatheca consists of an ovoidal sac, opening by a long, muscular duct to the exterior, and a pyriform diverticulum opening into the medial aspect of the spermathecal duct, close to distal end of the duct. (Plate 10, fig. 3.) The prostates are simple tubular organs with unbranched axial canals. They are of unequal extent, the left prostate commencing from the duct, which passes through xviii and xix, and extending back through xx and xxi into xxii, where it terminates in a dorsally directed portion, and the right arising from its duct in xviii and extending around the lateral surface of the intestine in xix to terminate on the left dorso-lateral surface of the intestine in xix. (See Plate 10, fig. 2.) There are four pairs of vesiculae seminales, a pair in each of ix x, xi and xii Those of ix and xii are much smaller than those of x and xi.

A broad band of small micronephridial tubules extends across the ventrolateral and lateral aspects of the body-wall in each segment.

Remarks. Although the two species are superficially distinct, M. viridis very closely resembles M. maorica in its internal anatomy. It can be distinguished from M. maorica by its possession of four pairs of vesiculae seminales (M. maorica has only three pairs).

Megascolides ruber n.sp. (Plate 11, figs. 1–3)

A single specimen of this dark red earthworm was collected from Awapuka clay near the summit of the Mangamuku Range, North Auckland, in a large area of native rain-forest. The posterior portion of the specimen is missing, so the length cannot be determined. The average diameter is 4·5 mm.

The prostomium is epilobous. The clitellum covers xiv to xviii and is lighter in colour than the rest of the body. The chaetae are lumbricine in arrangement, eight per segment. On segment xxii their arrangement is as follows:

ab = 1.25 mm.; cd = 2 mm.; bc = 1.5 mm.; aa = 1·5 mm.; dd = 3 mm.

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

There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with chaeta b. There is a single, unpaired, median, ventral female pore on xiv slightly anterior to the line of the chaetae on that segment. A single pair of prostatic pores occurs on xviii. Each of the prostatic pores is situated in the centre of a cuplike depression, with a raised tumid edge, in line with the chaetal interval ab on each side. Chaetae a and b are absent on xviii.

Internal Anatomy (Plate 11, fig. 2)

There are no specially thickened septa.

Alimentary Canal. The pharynx is slender and muscular and occupies i to iv. A small spheroidal gizzard lies in v. The oesophagus extends from vi to xv and bears two pairs of prominent calciferous glands, a pair in xii and a pair in xiii. The intestine commences in xvi and is very thin walled.

Vascular System. The dorsal blood vessel is unpaired. There are three pairs of dilated hearts, a pair each in x, xi and xii.

Reproductive System. There are two pairs of small lobate testes, a pair in x and a pair in xi. A single pair of ovaries is present in xiii and these organs are of a peculiar form which serves to distinguish the species from others of the same genus. Each ovary consists of a number of moniliform threads clustered

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together at their attachment to the anterior septum of xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each is in the form of a large, thin-walled, ovoidal sac with a small pyriform diverticulum opening into the anterior aspect of its duct. (Plate 11, fig. 3.) The prostates are simple, unbranched, tubular organs lying ventral to the intestine, as is typical of the New Zealand species of Megascolides (Tokea). Each prostate arises from a narrow muscular duct in xviii. The left prostate is coiled in the form of a reversed S, terminating in xx, and the right prostate extends posteriorly from xviii through xix and xx into xxi, where it curves sharply towards the dorsal aspect of the intestine and terminates. There are two pairs of small racemose vesiculae seminales, a pair in xi and a pair in xii. Segments ix and × contain a mass of loosely packed sperm cells.

Slender micronephridial tubules occur in each segment except the first.

The species is easily distinguishable from other species of Megascolides. No other species has cuplike depressions surrounding the prostatic pores and the form of the ovaries is a diagnostic character found in no other species of the genus.

Megascolides parvus n.sp. (Plate 12, figs. 1–3)

A single specimen of this small species was collected from a small remnant of native forest (kahikatea, tawa, nikau) on a hillside three miles north-west of Hick's Bay, beside the Hick's Bay-Opotiki road. It is 26·5 mm. in length and 3·5 mm. in diameter and has 72 segments It is reddish-brown dorsally, paler ventrally, and has a buff clitellum surrounding xiv–xvii (four segments).

The prostomium is epilobous, without a posterior groove. There are eight chaetae on each segment. On xxiv their arrangement is as follows: ab = 1·25 mm.; cd = 1·75 mm.; aa = 1 5 mm.; bc = 1·75 mm.; dd = 2·25 mm.

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There are three pairs of spermathecal pores, a pair in each of the intersegmental grooves 6/7, 7/8 and 8/9, one on each side slightly medial to the line of chaeta a in each of the grooves. The female pores are on xiv, one on each side, very close to the ventral midline and close to the anterior margin of the segment. The male pores are on xviii, one on each side, in line with chaeta a of the adjacent segments. Each pore opens on the apex of a low, rounded papilla. There is an unpaired tuberculum pubertatis, taking the form of a tumid ridge, extending across the ventral midline in the intersegmental groove 19/20 from the line of chaeta a on one side to the same line on the other side and a pair of small oval tubercula on xviii, one on each side, posterior to the male papilla of the same side. I could find no dorsal pores.

Internal Anatomy (Plate 12, fig. 2)

Septa vii/viii, viii/ix, ix/x, x/xi, xi/xii and xii/xiii are slightly thickened.

Alimentary Canal. The pharynx occupies the first four segments. The posterior dorsal and lateral aspects of the pharynx are covered by masses of salivary gland tissue. There is a thick-walled globular gizzard in v. The oesophagus extends from vi to xiv and there is a pair of prominent rounded calciferous glands in xii. The intestine commences in xv.

Vascular System. The dorsal blood vessel is unpaired. There are three pairs of dilated hearts, a pair in each of x, xi and xii.

Reproductive System. There are two pairs of minute lobate testes, a pair in x and a pair in xi, and a pair of ovaries in xiii. The testes and ovaries are

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situated one on each side, attached to the anterior septum in their respective segments. There are three pairs of spermathecae, a pair in each of vii, viii and ix. Each spermatheca consists of a conical sac, opening by a muscular duct to the exterior, and a small spherical diverticulum opening by a very slender duct into the spermathecal duct. (Plate 12, fig. 3.) The prostates lie one on each side of the ventral nerve cord beneath the intestine and are very long, slender, tubular organs. The left prostate originates from a slender duct passing through xviii and xix and extends back from xx to xxxiv, turns and passes forward again through xxxiii into xxxii, where it terminates. The right prostate passes back only as far as xxxiii, where it terminates. There are two pairs of small, racemose, vesiculae seminales, a pair in ix and a pair in xii.

There are micronephridial tubules in each segment except the first. They form a compact mat of tubules on the lateral aspects of the body-wall in each segment.

Remarks. M. parvus closely resembles and may actually be identical with M. huttoni (Benham, 1904) The inadequacy of Benham's description makes it impossible to positively distinguish the present species from M. huttoni, but the following points of difference are noted:

(i) The male pores of M. parvus are situated on low, rounded papillae on xviii; those of M. huttoni are in slight circular depressions on xviii.

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(ii) M. parvus has a median oval tuberculum pubertatis at 19/20; M. huttoni has a similar tuberculum on the hinder margin of xviii.

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(iii) The spermathecal pores of M. parvus are intersegmental, in the grooves 6/7, 7/8 and 8/9; those of M. huttoni are on the hinder margin of vi, vii and viii.

(iv) The gizzard of M. parvus is short and thick-walled; that of M. huttoni is long, narrow and thin-walled.

(v) The micronephridia of M. parvus commence in ii; those of M. huttoni commence in iii.

(vi) The prostates of M. parvus extend straight back from xviii to xxxiii or xxxiv; those of M. huttoni extend forward from xviii to xvi.

None of these characters is of specific value, since none of them is invariable, but M. parvus does not show sufficient similarity to the scanty description of M. huttoni to be positively identified as M. huttoni, and I prefer to establish a new species for it.

Megascolides raglani n.sp. (Plate 13, figs. 1–3)

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Two specimens were collected, one from a forest remnant near the crest of the Raglan Range, south of the Hamilton-Raglan road, and the other from native forest near Moerangi, beside the Raglan-Kawhia road. Superficially the species is easily mistaken for Spenceriella shakespeari (Benham). It is about the same size and colour and has prominent, transversely disposed tubercula pubertatis, one at 17/18 and another at 18/19, similar to those found in S. shakespeari. However, the possession of only eight chaeta on each segment as compared with 48 in S. shakespeari readily distinguishes the two species. The strong similarity between the two species may be interpreted as an indication that S. shakespeari has been derived from M. raglani as a local variant.

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The specimen on which this description is based is 81 mm. in length and 5 mm. in diameter and has 101 segments. It is brick red dorsally and pale ventrally and has a buff clitellum which completely surrounds xiii–xviii. The clitellum is not greatly developed over xiii and xviii and the intersegmental furrows 13/14 and 17/18 are quite well marked. The prostomium is epilobous.

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There are eight chaetae on each segment, arranged in pairs. On xxiv their arrangement is as follows:

ab = cd = 1.75 mm.; aa = 2·25 mm; bc = 2.25 mm.; dd = 2·5 mm.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with the chaetal interspace ab. There is an unpaired median ventral female pore on xiv, anterior to the level of the chaetae. The male pores are on xviii and their arrangement and the arrangement of the structures associated with them is strikingly similar to the arrangement seen in Spenceriella shakespeari. A pair of small oval depressions with slightly raised white borders lies one on each side of xviii, and in each of these small depressions, in line with chaeta a of the adjacent segments, is a small male pore. There are two very prominent, transversely disposed, oval, glandular tubercula pubertatis, one in each of the intersegmental furrows 17/18 and 18/19, extending across the ventral midline. There are no other tubercula pubertatis.

Internal Anatomy (Plate 13, fig. 2)

The septa ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are muscular and thickened.

Alimentary Canal. The pharynx occupies the first four segments and posteriorly it is covered dorsally and laterally by salivary glands. The gizzard has thick muscular walls and lies in v. The oesophagus extends from vi to xv and there are no oesophageal glands. The intestine commences in xvi.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of hearts, a pair in each of ix, x, xi and xii. Those of ix are much more slender than those of the subsequent segments.

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Reproductive System. There are two pairs of testes, a pair in x and a pair in xi, and a single pair of ovaries in xiii. The testes and ovaries have a similar, compact, lobate form and are situated one on each side of the ventral midline, attached to the anterior septa of their respective segments. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each consists of a large sac, almost spheroidal, but with a finger-like lobe lying over its upper surface and projecting towards the midline. The sac opens to the exterior by a long, thickwalled duct, and an elongate pyriform diverticulum opens medially into the duct (Plate 13, fig. 3). In one specimen the right anterior spermatheca was situated in vii instead of viii, but its duct passed back through vii and opened to the exterior at the intersegmental furrow 7/8, as is normal in the species. The prostates are short, curved, tubular organs, lying one on each side of the ventral nerve cord. They open to the exterior through a narrow duct, one on each side of xviii, and the organs extend back from xviii to xix. There are three pairs of racemose vesiculae seminales, a pair in each of ix, xi and xii. Those of ix are small rounded organs, attached to the posterior septum of the segment, while those of xi and xii are larger organs, extending around the lateral aspects of the oesophagus and attached to the anterior septa of their respective segments.

Micronephridia occur in every segment except the first, as a mat of small, slender tubules, covering the lateral aspects of the peritoneum. In the eleven most posterior segments there are very small paired meganephridia, in addition to the micronephridia. The meganephridia do not appear to be functional. Each consists merely of a tightly coiled tubule with no obvious funnel or elimination duct, lying close to the ventral nerve cord on its side of the segment.

Megascolides fuscus n.sp. (Plate 14, figs. 1–3)

Several specimens of this large brown earthworm were collected from topsoil

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Fig. 1—Megascolides viridis. Ventral aspect, segments i–xxi.
Fig. 2—M. viridis. Dissection from the dorsal aspect.
Fig. 3—M. viridis. Left anterior spermatheca anterior aspect.

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Fig. 1—Megascolides ruber. Ventral aspect segments i–xxi.
Fig. 2—M. ruber. Dissection from the dorsal aspect.
Fig. 3—M. ruber. Left posterior Spermatheca, lateral aspect.

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Fig. 1—Megascolides parvus. Ventral aspect, segment i–xx.
Fig. 2—M. parvus. Dissection from the dorsal aspect.
Fig. 3—M. parvus. Right spermatheca of viii, posterior aspect.

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Fig. 1—Megascolides raglani. Ventral aspect, segments i–xxiii.
Fig. 2—M. raglani. Dissection from the dorsal aspect.
Fig. 3—M. raglani. Right posteior spermatheca. posterior aspect.

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Fig. 1—Megascolides fuscus. Vential aspect, segments i–xx.
Fig. 2—M. fuscus. Dissection from the dorsal aspect.
Fig. 3—M. fuscus. Left spermatheca, medial aspect.

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Fig. 1—Megascolides irregularis. Ventral aspect, segments i–xx.
Fig. 2—M. irregularis. Dissection from the dorsal aspect.
Fig. 3—M. irregularis. Left posterior spermatheca, anterior aspect.

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Fig. 1—Megascolides alba. Ventral aspect, segments i–xxxi.
Fig. 2—M. alba. Dissection from the dorsal aspect.
Fig. 3—M. alba. Left posterior spermatheca, posterior aspect.

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Fig. 1—Megascolex novae-zealandiae. Ventral aspect, segments i–xx.
Fig. 2—M. novae-zealandiae. Dissection from the dorsal aspect.
Fig. 3—M. novae-zealandiae. Right anterior spermatheca, lateral aspect.

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Fig. 1—Pheretima campestris. Ventral aspect, segments i–xix.
Fig. 2—P. campestris. Dissection from the dorsal aspect.
Fig. 3—P. campestris, Left posterior spermatheca anterior aspect.

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Fig. 1—Diporochaeta obtusa. Ventral aspect, segments i–xxi.
Fig. 2—D. obtusa. Dissection from the dorsal aspect.
Fig. 3—D. obtusa. Left posterior spermatheca, medial aspect.

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Fig. 1—Perionyx egmonti. Ventral aspect, segments i–xx.
Fig. 2—P. egmonti. Dissection from the dorsal aspect.
Fig. 3—P. egmonti. Right posterior spermatheca, medial aspect.
Fig. 4—P. egmonti. Right prostate, ventro-lateral aspect.
Fig. 5—P. egmonti Vertical section through prostate. (Diagrammatic).

acc gl., accessory gland, e.g., cerebral ganglion, ch., chaeta, el, clitellum; c.p.e., central prostatic cavity; d.b.v., doisal blood vessel, d.div., duct of spermathecal diverticulum, div., spermathecal diverticulum, f.p., female pore; g., gizzard, h., heart, int., intestine; l., lobule of piostate; m., mouth; m.f., “male field”, m g., mucus gland, mn., meronephridial tubule, m p., male pore, n, meganephridial tubule; o., ovary, oe., oesophagus; oe.gl., oesophageal gland, o f., ovarian funnel; p., prostate; pap., prostatic papilla, p.d, prostatic duct, peri. peristomium, ph, pharynx, pr, prostomium, pv., proventriculus; s, septum; sal.gl., salivary gland, sh., sheath; s.i.v., supra-intestinal vessel; sp., spermatheca; sp.d., spermathecal duct, sp.p, spermathecal pore; t., testis, t.f., testicular funnel; t.pub., tuberculum pubertatis, v.b.v., ventral blood vessel, v.n.c., ventral nerve cord, v.s., vesicula seminalis.

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and subsoil in Taupo sandy silt in areas of native rain forest and exotic forest (Pinus spp.) in the Rotorua district.

The specimen on which this description is based is 251 mm. in length and 10 mm. in diameter and has 268 segments. The clitellum is greyish-brown and is developed over the entire surface of xv and xvi and on the dorsal and lateral surfaces of xiv, xvii and xviii. The prostomium is epilobous. The first six segments are biannulate, segments vii–xviii are quadriannulate and the post-clitellar segments are biannulate. There are eight chaetae on each segment, arranged in pairs. On xxiv: ab = 1·25 mm.; cd = 2·25 mm.; aa = 4·4 mm.; bc = 4·5 mm.; dd = 10 mm.

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There is only one pair of spermathecal pores, at 7/8 in line with the chaeta b. There is a single pair of female pores slightly anterior and lateral to the chaeta a on xiv. The male pores open on the apices of a single pair of prominent rounded papillae on xviii. The chaeta a is absent on xviii and in its place on each side is the papilla bearing the male pore. There are no tubercula pubertatis. Dorsal pores occur in every intersegmental groove posterior to the clitellum. There are no nephridiopores.

Internal Anatomy (Plate 14. fig. 2)

The first septum occurs at the intersegment iv/v. The septa vii/viii, viii/ix, ix/x, x/xi and xi/xii are muscular and thickened.

Alimentary Canal. A rounded muscular pharynx occupies the first four segments. The pharynx is covered dorsally and laterally by a downlike mass of salivary glands. There is a thin-walled proventriculus in v and a stout muscular gizzard in vi. The oesophagus extends from vii to xvi and there are no oesophageal glands. The intestine commences in xvii, is thin-walled and has a small typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are four pairs of dilated hearts in x, xi, xii and xiii, rising from a supraintestinal vessel which extends only through those four segments.

Reproductive System. There are two pairs of testes in x and xi. The testes have a lobate form and their funnels are on the posterior septa of the segments. The two slender vasa deferentia of each side are prominent, lying side by side on the ventro-lateral aspect of the peritoneum They remain distinct from each other for most of their length, fusing just before they enter the prostatic duct, in xviii. There is a single pair of ovaries in xiii. Each ovary appears as a thin lace-like lamina attached to the anterior septum of the segment. The ovarian funnels are very prominent white structures on the posterior septum of xiii, and the short oviducts, which open to the exterior in xiv, are very much thicker than the vasa deferentia. A single pair of spermathecae occurs in viii. Each is a large, roughly ovoidal, thin-walled sac with a small, thick-walled digitate diverticulum opening into the anterior aspect of its duct. (Plate 14, fig. 3). The whole organ is confined to the eighth segment. The prostates are large, coiled, tubular organs, extending through the segments xviii, xix, and xx and opening to the exterior in xviii by wide muscular ducts. The prostatic duct of each side fuses with the corresponding vas deferens close to the body-wall in xviii. There are two pairs of large, finely racemose vesiculae seminales surrounding the lateral aspects of the oesophagus in xi and xii.

Micronephridia commence in ii. There is a compact cluster of micronephridial tubules on the ventro-lateral aspect of the peritoneum on each side of the segments. The individual tubules are very long and are twisted like corkscrews.

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Remarks. This species most closely resembles M. (Tokea) sapida Benham (1904), collected from Ruatahuna, between Rotorua and Lake Waikaremoana. It agrees with Benham's species in size, colour, and in many points of its anatomy, but the number and form of the spermathecae in the two species are markedly different. M. sapida has three pairs of spermathecae, each with a small globular diverticulum, while M. fuscus has only one pair of spermathecae and their diverticula are digitate. The number and form of the spermathecae and their diverticula are very reliable systematic characters and serve to distinguish the two species one from the other.

Megascolides irregularis n.sp. (Plate 15, figs. 1–3)

One specimen of this large worm was collected near Tangowahine, North Auckland. Other specimens have been collected from localities in the same area. The specimen on which the description is based is 117 mm. in length with 120 segments, and has an almost uniform diameter of 7·5 to 8 mm. It is purplish-red in colour dorsally, and pale ventrally. The clitellum is buff in colour and completely surrounds xiv to xvii, except for a small area on the ventral surface of xiv where the female pores occur.

The prostomium is epilobous. There are eight chaetae on each segment, but their arrangement is most irregular. They are not arranged in pairs, but occur four on each side of each segment, scattered indiscriminately. The distance between corresponding chaetae a on each segment is more or less uniform anterior to the prostatic pores, but varies considerably posteriorly.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9. On each side the pores lie in line with chaeta a. A single pair of female pores is present on xiv, very close together in a small pale area covering the ventral mid-line. On xviii is a pair of prostatic pores surrounded by tumid lips. Each pore is in line with the chaetae a on the same side. Across the ventral mid-line, at the anterior margin and at the posterior margin of this segment is a small tuberculum pubertatis.

Nephridiopores are not visible externally.

Internal Anatomy (Plate 15, fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi and xi/xii are thickened and muscular.

Alimentary Canal. A large muscular pharynx, to which is attached dorsally a mass of small salivary glands, occupies the first four segments. A small muscular gizzard lies in v. The oesophagus extends from vi to xv and has a single pair of large calciferous glands in xiii. The intestine commences in xvi, is fairly thick-walled and has a narrow typhlosole on its median dorsal aspect.

Vascular System. The dorsal blood vessel is single throughout and bears four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. Two pairs of small lobate testes lie close to the ventral mid-line, a pair in x and a pair in xi, and a pair of very small lobate ovaries lies close to the ventral mid-line in xiii. There are two pairs of spermathecae, a pair in viii and a pair in ix. Each spermatheca takes the form of an ovoidal sac with a wide muscular duct leading to the exterior. A small rounded diverticulum opens into the medial aspect of the duct close to its distal extremity (Plate 15, fig. 3). A single pair of small tubular prostates is present. Each prostate opens by a narrow muscular duct in xviii, but the organs are of unequal extent. The left prostate extends through xviii and xix to terminate in xx in

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a coiled portion. The right prostate extends posteriorly, without coiling, through xviii, xix and xx to xxi, where it terminates. The prostates are noteworthy in that they lie ventral to the intestine and close to the ventral nerve cord, thus disposed similarly to those of M. (Tokea) esculenta Benham, 1904. There are two pairs of racemose vesiculae seminales, a pair in ix and a pair in xii.

A band of micronephridial tubules is present in each segment except the first, but on each side of the first segment there is a cluster of tubules larger than those of the micronephridia which probably function as a slime gland, as in some species of Octochaetus (e.g. O. multiporus Beddard, 1892).

Remarks. The asymmetrical arrangement of the chaetae in this species is an unusual character, and one of which the author can find no previous record in this genus. It would seem to be a secondary modification of the paired arrangement normally found in the genus. The disposition of the vesiculae seminales is also unusual, and assists in distinguishing the species from others of the same genus.

Megascolides alba n.sp. (Plate 16, figs. 1–3)

Several specimens of this large earthworm were collected from Motatau clay in a remnant of native bush about a mile south-east along the main Whangarei-Auckland highway from the Maungakaramea road junction. The specimen on which this description is based is 90 mm. long, has 141 segments, and is 7.5 mm. in diameter for the greater part of its length It is completely unpigmented except for a slightly yellowish clitellum surrounding xiv to xvii. The transparency of the body-wall makes the species readily recognizable, since the prostates are visible through the body-wall as a pair of yellowish streaks lying close to the ventral nerve cord and extending from xviii to xxviii or xxix.

The prostomium is epilobous. Chaetae are lumbricine in arrangement, there being eight on each segment. In the mid body:

ab = 1·5 mm.; cd = 1·25 mm.; aa = 2·5 mm.; dd = 9 mm.; bc = 2 mm.

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There are two pairs of spermathecal pores, a pair at 7/8 and a pair at 8/9, in line with chaeta a. A single pair of female pores is present on xiv, each pore situated slightly medial to chaeta a. There is a single pair of prostatic pores on xviii. Each pore is situated in the centre of an oval tumid area which lies in line with the chaetal interval ab on each side. Chaetae are absent on xviii. Tubercula pubertatis are present. Each is in the form of a large, oval, white pad which lies across the ventral mid-line, one at 16/17 and one at 19/20.

Nephridiopores are not visible externally.

Internal Anatomy (Plate 16, fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi and xi/xii are muscular and thickened.

Alimentary Canal. A small muscular pharynx occupies the first four segments. There is a strongly muscular gizzard in v. The oesophagus extends from vi to xvi and lacks oesophageal glands. The intestine commences in xvii.

Vascular System. The dorsal blood vessel is single throughout. There are five pairs of hearts, a pair each in ix to xiii. the pair in ix being much smaller than the others.

Reproductive System. There are two pairs of lobate testes, a pair in x and a pair in xi, and a single pair of ovaries in xiii. Two pairs of spermathecae are present, a pair in viii and a pair in ix. Each is an ovoidal sac with a small medial

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digitiform diverticulum opening into the duct distally. (Plate 16, fig. 3.) There is a single pair of tubular prostates arising from narrow muscular ducts in xviii. They lie close to the ventral nerve cord ventral to the intestine and are of unequal length, the right prostate terminating in xxvii and the left in xxix. Two pairs of vesiculae seminales are present, a pair in x and a pair in xi.

Micronephridia occur in each segment except the first, in the form of a band of slender tubules on the lateral aspects of the body-wall.

The following are supplementary notes on the two species M. suteri (Benham, 1904) and M. urewerae (Benham, 1904).

Megascolides suteri (Benham), 1904

Syn. Tokea suteri Benham, 1904

Megascolides (Tokea) suteri Benham, 1941

The type specimen of M. suteri was collected from Auckland and Benham did not record the species elsewhere It has now been collected from the Rotorua-Taupo district and the East Coast district of the North Island. All the specimens have been collected in or under rotten logs. This habitat preference was not noted by Benham in the case of the type specimen.

The description given by Benham is insufficient to distinguish the species from M. huttoni (Benham) and M. urewerae (Benham), so some additional notes on the characters of the species are set out below.

External Features. There is a pair of female pores on xiv, one on each side, anterior and slightly medial to chaeta a. The tubercula pubertatis may be absent.

Intenal Anatomy. The pharynx occupies the first four segments. There is a short proventriculus in v, opening into the small, feebly developed gizzard in vi. The oesophagus extends from vii to xv and has a pair of slight lateral dilatations in xii. The intestine commences in xvi.

The dorsal blood vessel is unpaired throughout its length. A very slender, unpaired, supra-intestinal vessel commences in x and extends back to xiii. From the supra-intestinal vessel arise three pairs of hearts, a pair in each of x, xi and xii. The supra-intestinal vessel terminates in a number of slender branches, running across the dorsal aspect of the oesophagus in xiii.

There are two pairs of testes, a pair in x and a pair in xi. Each testis is a small, folded, laminar organ. A pair of ovaries occurs in xiii. The ovaries are larger, thicker, laminar organs. There are two pairs of racemose vesiculae seminales, a pair in ix and a pair in xii.

Micronephridia are found in each segment except the first. The tubules form a desnely packed mass over the ventro-lateral aspects of the peritoneum, but are less numerous laterally and absent dorso-laterally.

Megascolides urewerae (Benham), 1904

Syn. Tokea urewerae Benham, 1904

Megascolides (Tokea) urewerae Benham, 1941

A specimen which agrees fairly closely with Benham's description of M. urewerae was collected from native forest on a hillside about seven miles west of Motuhora. Some additional notes on the species are necessary, since the description given by Benham is insufficeint to distinguish the species from M. suteri (Benham) and M. huttoni (Benham). The following additional notes are therefore given.

External Features. The female pores are on xiv, one on each side, close to the ventral mid-line, anterior and medial to chaeta a. The spermathecal pores are

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on vi, vii and vii, slightly anterior to the furrows 6/7, 7/8 and 8/9. A pair of small oval tubercula pubertatis occurs on xviii, one on each side posterior to the pit containing the male pore, extending slightly over the intersegmental furrow 18/19.

Internal Anatomy. The intestine commences in xvi.

The dorsal blood vessel is unpaired throughout its length. Three pairs of hearts arise from the dorsal vessel, a pair in each of x, xi and xii.

There are four pairs of racemose vesiculae seminales, a pair in each of ix, x, xi and xii. The prostates lie under the intestine, one on each side of the ventral nerve cord, and extend straight back through the segments.

The remainder of the anatomy of the specimen agrees closely with the description given by Benham.

Genus Notoscolex Fletcher

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“Chaetae eight per segment. Spermathecal pores one, two or three pairs, the last in furrow 8/9 (in certain abnormal species 7/8). One gizzard in segment v or vi. Micronephridia present, sometimes with meganephridia also. Prostates with branched system of ducts.” (Stephenson, 1930.)

Only two species from the New Zealand region, N. equestris Benham (1942) and N. hakeaphilus Benham (1949) are assigned to the genus Notoscolex. Of these two it is possible that N. hakeaphilus is introduced. The two species are readily distinguished by the folloiwng differences:

(i) N. hakeaphilus is a large worm (up to 650 mm. in length); N. equestris is about 200 mm. in length.

(ii) N. hakeaphilus is brick red in colour dorsally, pale ventrally; N. equestris is coloured alternately dark brown and pale cream in narrow bands.

(iii) N. hakeaphilus has two pairs of spermathecae, a pair in each of viii and ix; N. equestris has three pairs, a pair in each of vii, viii and ix.

(iv) N. hakeaphilus has three pairs of hearts, a pair in each of viii, ix and x; N. equestris has four pairs, in x, xi, xii and xiii.

There are many other points of difference which are obvious from Behnam's descriptions of the species (q.v.).

Genus Spenceriella Michaelsen

“Chaetae more than eight per segment. Spermathecal pores one to three pairs. One gizzard in segment v. Micronephridial. Prostates tubular, with simple unbranched duct.” (Stephenson. 1930.)

Two species of the genus were described by Benham (1905(a)) and referred by him to the genus Diporochaeta. Both species were collected from Little Barrier Island. Michaelsen (1907) established the genus Spencieriella for some Australian species, and included Benham's species, in it, since they differ from Diporochaeta in being micronephridial while Diporochaeta is meganephridial. The two New Zealand species are Spenceriella gigantea and S. shakespeari. They may be identified from the following key:

a1 One pair of vesiculae seminales in xii. Last heart in xii. S. gigantea

a2 Two pairs of vesiculae seminales, in ix and xii. Last heart in xiii S. shakespeari

S. gigantea is a very large worm, measuring about 4 ft. 6 in. (1,370 mm.) in length and half an inch (11 mm.) in diameter. It has been recorded only from Little Barrier Island.

S. shakespeari is a small, stout, brick-red worm (about 100–150 mm. in length and 4–5 mm. in diameter). It is readily distinguished in the field by the unusual

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form and disposition of its tubercula pubertatis and the large number of chaetae on each segment. It was collected from Little Barrier Island by Benham, and I have collected it from the western coastal ranges of the Waikato and Taranaki districts.

Genus Megascolex Templeton

“Chaetae, at least in the middle and hinder regions of the body, numerous (more than eight) in each segment. Spermathecal pores usually one to five pairs, between segments iv and ix (the exceptions are constiuted by the few cases where the pores are fused in the middle line, or where they are numerous on each side in each segment) One gizzard in v, vi or vii. Micronephridial. Prostates with brached system od ducts.” (Stephenson, 1930.)

Of about 120 known species only two have been recorded from the New Zealand region. They are M. laingii Benham (1902) and M. novae-zealandiae, a new species, described below. M. laingii is known only from Norfolk Island and M. novae-zealandiae has been found only in the extreme northern portion of the North Island of New Zealand. A species recorded by Baird (1871) as Megascolex antarctica may have been intended as a species of Megascolex, but Baird's description is insufficient for the recognition of the species or genus. There is no other record of the genus from the mainland of New Zealand.

The following is a description of the new species M. novae-zealandiae. Type in my collection.

Megascolex novae-zealandiae n.sp. (Plate 17, figs. 1–3)

Five specimens of this large red earthworm were collected, three from Awapuka clay loam six miles from Mangonui, and two from a steep bush-clad hillside two miles and a half south-east of Ahipara. The specimen on which this description is based is 179 mm. in length and 9·5 mm. in diameter, and has 120 segments.

The prostomium is epilobous. Chaetae average about 36 per segment, but there may be more or less. They are perichaetine in arrangement and commence on ii. Their spacing on the segments is not regular. The clitellum is darker in colour than the adjacent segments, and surrounds xiv to xviii, and the anterior portion of xix.

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

Spermathecal pores are not visible externally. A single pair of female pores lies adjacent to the ventral midline on the anterior aspect of xiv. They cannot be seen with the naked eye, and are very close together. A single pair of male pores is present on xviii, each pore situated 1·75 mm. from the ventral midline and occupying the centre of a small oval light brown area. A prominent oval tuberculum pubertatis lies in the intersegmental groove 17/18, and another similar tuberculum in the groove 18/19. There are no nephridiopores.

Internal Anatomy (Plate 17, Fig. 2)

Septa v/vi, vi/vii, vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii and xiii/xiv are thickened and muscular.

Alimentary Canal. A muscular pharynx occupies the first four segments. The gizzard, which has thick muscular walls, is elongate and is confined to v. The oesophagus extends from vi to xv and lacks oesophageal glands. The intestine commences in xvi, is thin walled and has a marked typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. There are two pairs of free testes, a pair in × and a pair in xi, and a pair of ovaries in xiii. Two pairs of spermathecae are present, a pair

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in viii and a pair in ix. Each spermatheca is in the form of a large pyriform sac opening ventrally to the exterior by a wide muscular duct. A small ovoidal diverticulum opens by a slender duct into the anterior aspect of the spermathecal duct. (Plate 17, Fig. 3.) The spermathecal pores are in 7/8 and 8/9. A single pair of prostates is present. The left prostate commences from a short duct in xviii and extends anteriorly through xvii/xviii to terminate in xvii. The right prostate commences from a similar duct in xviii and extends posteriorly through the septa to terminate in xxiii, lying close to the ventral nerve cord throughout its length. Examination of sections of the prostate shows that the axial duct receives small ductules from the surrounding prostatic tissue.

Micronephridial tubules occur in a wide band on the lateral walls of each segment except the first. Anterior to the gizzard the tubules are very numerous and completely cover the lateral walls of the segments.

This species is distinct from M. laingii, since it has four pairs of hearts, while M. laingii has only three pairs; it has large prostates while M. laingii has small compact prostates, and in a number of other points of internal and external anatomy it is also markedly distinct from M. laingii.

Genus Pheretima Kinberg

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“Chaetae numerous in each segment. Spermathecal pores one to six pairs, between segments iii and ix One gizzard between 7/8 and 10/11. Micronephridial. Testes and funnels enclosed in testis sacs; prostates with branched system of ducts; penial chaetae almost always wanting.” (Stephenson, 1930.)

There are about three hundred and fifty known species, of which only two are recorded from New Zealand. The two New Zealand species are P. clerica Benham (1947), collected from the Gisborne district, and P. campestris n.sp., collected from North Auckland and Raoul Island and described in this paper. P. campestris may be regarded as introduced into Raoul Island, probably with plants and seeds taken from New Zealand within the last few years. P. campestris may not actually be a new species, but since I cannot obtain literature concerning most of the great number of species belonging to the genus and it does not coincide with any of the descriptions that I am able to find. I have established a new species for it.

The following is a description of P. campestris. Type in my collection.

Pheretima campestris n.sp. (Plate 18, figs. 1–3)

Two specimens were collected from Mangakahia loam in a top-dressed pasture on the Victoria Valley road near Kaitaia, and two specimens from Otonga peaty loam in a top-dressed pasture near Awanui, North Auckland.

In both cases the species was found together with the introduced species Octolasium cyaneum and Allolobophora caliginosa, living in the upper inch of the topsoil among the roots of the pasture grasses. It is easily distinguished from the introduced species by its yellow-brown colour and its rapid movement when disturbed.

The specimen on which this description is based is 87 mm. in length, has 100 segments and is 3·75 mm. in diameter. The chaetae are perichaetine in arrangement, about 48 on each segment in the midbody, and evenly spaced around the perimeter of each segment. The clitellum is darker in colour than the adjacent segments and completely surrounds xiv to xvi. There are no chaetae on the clitellar segments.

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Spermathecal pores are not visible externally. There is a single median ventral female pore, in the centre of a median pigmented area on xiv. A pair of male pores is present on xviii. Each of the male pores occupies the centre of a round white area on a small papilla and is 1·5 mm. from the ventral midline. There are twelve chaetae between the two male pores.

Internal Anatomy (Plate 18. Fig. 2)

There are no thickened septa.

Alimentary Canal. An elongate muscular pharynx lies in i to iv. The gizzard, as in P. clerica, lies posterior to vii/viii and there is no septum between viii and ix, nor between ix and x. The oesophagus commences slightly anterior to x/xi and extends to xiv. There are no oesophageal glands. The intestine commences in xv and the junction between the oesophagus and the intestine is quite distinct. A pair of flattened cornuiform intestinal diverticula arise from the ventral surface of the intestine in xxvi and extend anteriorly, one on each side of the ventral nerve cord, through xxv into xxiv.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of dilated hearts, a pair each in x, xi, xii and xiii.

Reproductive System. There are two pairs of testes, a pair in × and a pair in xi, each testis lying in a testis sac. There is a pair of small lobate ovaries in xiii. Although the specimen is obviously sexually mature, there are no prostatic glands, although there is a pair of coiled muscular ducts in xviii resembling prostatic ducts. The vas deferens of each side opens to the exterior by the corresponding “prostatic” duct. The author dissected another mature specimen and found the same condition of the male genital organs. The vas deferens simply expands in xviii into what appears to be a prostatic duct, which coils before opening to the exterior, and there is no sign of a prostatic gland. This may be due to a seasonal regression of the prostates, but it is similar to the condition described by Stephenson (1930, p. 370) in P. heterochaeta. There are four pairs of spermathecae, a pair each in vi, vii, viii and in ix. Each spermatheca is in the form of a flattened cornuiform sac which opens to the exterior by a wide duct. A long, very narrow duct arises laterally from the spermathecal duct close to the body-wall and terminates in a small spheroidal diverticulum. (Plate 18, Fig. 3.) There are two pairs of lobate vesiculae seminales, a pair in xi and a pair in xii.

Micronephridia commence in ii and occur in every other segment except x. Septa viii/ix and ix/x are absent, but the nephridial tubules corresponding to viii and ix are referable to the spermathecae corresponding to those segments. There are, however, no nephridial tubules corresponding to x.

Remarks. This species shows a close similarity to P. clerica Benham (1947), but can be distinguished from P. clerica by the following points of difference:

(1) There are 48 chaetae on each segment in the midbody; P. clerica has 36.

(2) The colour of this species is yellowish-brown; P. clerica is purplish-brown in colour.

(3) There are no small tubercula pubertatis beside the male pores; tubercula pubertatis are present in P. clerica.

(4) The vesiculae seminales lie in xi and xii; in P. clerica they lie in x and xi.

(5) The transition from the posterior region of the oesophagus into the intestine is quite distinct, the oesophagus terminating in xiv and the intestine

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commencing in xv; in P. clerica it is difficult to determine the limits of the oesophagus and the intestine.

(6) The intestinal caeca arise in xxvi; in P. clerica they arise in xxvii or xxviii.

(7) Benham does not mention the absence of the micronephridia corresponding to × in P. clerica; these micronephridia are absent in P. campestris.

(8) There are no prostates in P. campestris; prostates are present in P. clerica.

It is the accumulation of such small differences which serves to distinguish one species from another, and the differences in this case are of an order which leaves little doubt that P. campestris is a species distinct from P. clerica.

Genus Diporochaeta Beddard

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“Chaetae, at least in the middle and hinder parts of the body, numerous (more than eight) per segment. Spermathecal pores two to five pairs, the last in furrow 8/9. One gizzard in the region of segments iv–vi, seldom vestigial. Purely meganephridial. Prostates tubular, with simple unbranched duct.” (Stephenson, 1930.)

Of the forty-eight known species, three have been recorded from the New Zealand region by previous workers and one more (D. obtusa) is described below. The three previously recorded species are D. intermedia Beddard (1890), an aquatic species, from the west coast of the South Island, D. aquatica Benham (1903), an aquatic species from Lake Manapouri in the south-western region of the South Island, and D. chathamensis Benham (1900), from Chatham Island. D. intermedia and D. aquatica are the only known aquatic species of Diporochaeta. D. aquatica was found at depths of 350–500 feet beneath the lake surface (see Benham, 1903a).

The New Zealand species may be identified from the following key:

a1 Two pairs of spermathecae D. aquatica
a2 Three pairs of spermathecae
b1 Spermathecae without diverticula D. chathamensis
b2 Spermathecae with diverticula D. obtusa
a3 Four pairs of spermathecae D. intemedia

The following is a description of the new species D. obtusa. Type in my collection.

Diporochaeta obtusa n.sp. (Plate 19, figs. 1–3)

Two specimens of this species were collected from Taupo sandy silt in an area of exotic forest two miles south of Rotorua. Specimens have also been collected from Stokes Valley (Wellington), the East Cape district, Norsewood, Waiouru, Lake Waikaremoana, Awakino, Kawhia, and Raglan. Such a wide distribution is remarkable for an endemic earthworm species, and it is difficult to imagine how such a distribution could have been achieved in one species while other species have generally a very limited distribution.

The specimen on which this description is based is 21 mm, in length, with 65 segments. The diameter is 1·5 mm. and the body terminates bluntly at the mouth and anus. The prostomium is tanylobous. The chaetae are perichaetine in arrangement, eighteen per segment, and are evenly spaced around the body except that the ventral gap is slightly greater than any other. The clitellum covers only segments xiv–xvi and is continuous right around these segments. Dorsal pores are obvious, commencing at iv/v and occurring in each intersegmental groove posterior to this.

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Three pairs of spermathecal pores occur at 6/7, 7/8 and 8/9 in line with chaeta b. A pair of small round white markings lies close together, one on each

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side of the mid-ventral line on the anterior margin of ix, in close association with the spermathecal pores, and a single similar marking is present on viii on the right, and on vii on the left. There is a single pair of female pores on xiv, slightly anterior to chaeta a. There is a very prominent pair of white papillae on xviii, between the chaetae a and b on each side, and on the apices of these papillae the common male and prostatic pores open to the exterior. Tubercula pubertatis occur on xvii, xviii, and xix. Those of xvii and xix take the form of a pair of small papillae anterior to the chaetae a and b on xix and posterior to those chaetae on xvii. Those of xviii are large transverse bands on the anterior and posterior margin of the segment, extending from the line of chaeta a of one side to the same line on the other side. There is also on this segment a pair of small papillae immediately anterior to the chaeta c of each side.

The nephridiopores are lateral and occur in a single series on each side in line with the chaeta e.

Internal Anatomy (Plate 19, Fig. 2)

Alimentary Canal. The pharynx occupies the segments i–v, is not very muscular, and is covered dorsally by salivary glands. There is a small gizzard in vi. The oesophagus extends from vii to xvi and has three pairs of calciferous glands in xiii, xiv, and xv. The intestine commences in xvii and lacks a typhlosole.

Vascular System. The dorsal blood vessel is unpaired throughout its length. There are three pairs of hearts in x, xi, and xii, those of xii being extremely dilated.

Reproductive System. There is one pair of free polylobate testes in xi, close to the mid-ventral line, and a single pair of ovaries close to the mid-ventral line in xiii. There are three pairs of spermathecae in segments vii, viii, and ix. Each consists of a lobulated sac opening by a narrow duct to the exterior, with a small, smooth-walled diverticulum lying close to the medial aspect of the sac and opening into the spermathecal duct. (Plate 19, Fig. 3.) One pair of very large prostates occurs. Each of the prostates is a wide, convoluted, tubular structure extending through the segments xviii to xxiii, closely applied to the lateral wall of the gut and opening to the exterior by a narrow duct in xviii. There are no penial chaetae. There is only one pair of vesiculae seminales, in xii. There is, however, a diffuse mass of sperms lying loose in the coelomic cavity of xi, the same as was recorded by Benham in D. chathamensis.

Remarks. D. obtusa shows very close affinities to D. chathamensis and also closely resembles the species D. maplestoni Spencer and D. walhallae Spencer from Australia. The differences between D. obtusa and these three species are set out in the accompanying table:

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D. obtusa D. chathamensis D. maplestoni D. walhallae
No. of chaetae on each segment 18 16 14 20–24
No. of spermathecae 3 pairs 3 pairs 2 pairs 5 pairs
Spermathecal diverticulum present absent absent absent
No. of testes 1 pair 1 pair 1 pair 2 pairs
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Genus Perionyx Perrier

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“Chaetae numerous (more than eight) per segment, in rings which are often almost closed. Male pores often approximated in greater or less degree, and may be very close to the middle line. Female pores unpaired (? always). Spermathecal pores, like the male pores, often very near the middle line, the last pair in 7/8 or 8/9. Gizzard very frequently more or less vestigial, in v or vi. Meganephridial. Two pairs of testes and funnels. Prostates with branched system of ducts.” (Stephenson, 1930.)

Of about fifty species, four have been recorded from the Auckland Islands and one (P. heterochaeta) from Snares Island, within the New Zealand region. These five species are P. heterochaeta Benham, 1909), P. brachysoma (Benham. 1909), P. perionychopsis (Benham, 1909), P. helophilus (Benham, 1909), and P. duodecimalis Michaelsen (1923). The first four of these species were placed in the genus Diporochaeta by Benham, but were transferred by Michaelsen (1923) to Perionyx. One new species, P. egmonti, is described in this paper. It is the first species of the genus to be recorded from the mainland of New Zealand. When I first discovered it, on the slopes of Mt. Egmont, I thought that it might have been introduced with some plants from Australia or India, but I have subsequently found it in areas of native forest inland from Mt. Egmont, as far east as Raurimu, in the central North Island near Mt. Ruapehu, so it is unlikely that it has been introduced. It does not coincide with any of the descriptions of species from the Auckland Islands, Snares Island, Australia or India.

The following is a description of P. egmonti. Type in my collection.

Perionyx egmonti n.sp. (Plate 20, figs. 1–5)

This a very large and distinctive species of earthworm, fairly common on the eastern slopes of Mt. Egmont. After heavy rain specimens are frequently found crawling on the surface of the ground. The specimen described below was found on a path near the Stratford Mountain House after heavy rain. It is brick-red dorsally, paler ventrally, and the pink clitellum surrounds xiii to xvii. It is 138 mm. in length and 10 mm. in diameter and has 156 segments. The prostomium is epilobous. There are about 80 chaetae on each segment, evenly spaced except that the ventral gap is about twice that between the other chaetae and the dorsal gap is about twice the ventral gap.

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

The spermathecal pores are not visible externally. A pair of female pores occurs on xiv, a pore slightly anterior to the ventral-most chaeta of each side of the segment. The male pores are on xviii, one on each side in line with the second from ventral-most chaeta of the adjacent segments. Each pore is situated on the apex of a small papilla. The papillae lie on the lateral boundaries of an oval “male field” bounded anteriorly and posteriorly by the margins of xviii. Within this “male field” are three pairs of small, rounded tubercula pubertatis, a pair adjacent to one another across the ventral midline on the anterior border of the “male field”, a pair similarly placed on the posterior border of the “male field”, and a third pair lying one on each side, immediately anterior to the prostatic papillae. Another pair of smaller tubercula pubertatis lie one on each side of the ventral midline at 19/20. (Plate 20, Fig. 1.)

The nephridiopores cannot be seen externally.

Internal Anatomy (Plate 20, Fig. 2)

The septa vii/viii, viii/ix, ix/x, x/xi, xi/xii, xii/xiii, and xiii/xiv are thickened and muscular.

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Alimentary Canal. The pharynx is large and muscular and occupies the first four segments. A thin, lace-like salivary gland is attached to each side of the pharynx near its posterior margin. The gizzard is very well developed and strongly muscular and lies in v and vi. The oesophagus extends from vii to xvi. There are five pairs of dilated calciferous glands, a pair in each of vii, viii, ix, xiii, and xiv, and in vii there is a pair of accessory glands, similar in form to the salivary glands, but smaller in size, pressed against the lateral walls of the oesophagus and opening by a narrow duet into the ventro-lateral aspect of the oesophagus, independently on each side. The function of these accessory glands is unknown, but they probably secret some enzyme important in the metabolism of the worm. The intestine commences in xvii. It has thin, almost transparent walls and is very wide.

Vascular System. The dorsal blood vessel is unpaired. There are four pairs of dilated hearts, a pair in each of x, xi, xii, and xiii, arising from a slender supra-intestinal vessel which extends only from x to xiii.

Reproductive System. There are two pairs of testes, a pair in x and a pair in xi. The testes are small lobate organs situated close to the ventral midline on the anterior septa of their respective segments. A pair of ovaries occupy a similar position in xiii. The ovaries take the form of a small flattened sheet, attached to the anterior septum of the segment, one on each side. The free edge of the ovary, which projects into the coelomic cavity of the segment, is fimbriate. There are four pairs of spermathecae, a pair in each of v, vi, vii, and viii. Each consists of a simple ovoidal sac without diverticula, opening by a narrow duct to the exterior 0·75 mm. from the ventral nerve cord and close to the anterior septa of its segment. (Plate 20, Fig. 3.) The prostates are dorso-ventrally flattened, mushroom-shaped organs lying beneath the intestine, one on each side of the ventral nerve cord in xviii. Each prostate is enclosed in a transparent sheath and is divided, within the sheath, into a number of small lobules. (Plate 20, Fig. 4.) The prostate opens to the exterior by a narrow duct, arising from the mid-ventral region of the organ. In sections it is seen that the prostatic duct communicates with a central prostatic cavity and that from each lobule of the glandular tissue a small ductule opens into the central cavity. (Plate 20, Fig. 5.) In the region of the prostates are three pairs of rounded glandular masses, attached to the ventral aspect of the peritoneum. They are situated one on each side, at the anterior and posterior borders of xviii and at xix/xx. These positions correspond with the positions of the external tubercula pubertatis and the glands probably secrete a mucous substance which holds the worms together when they are in copulo. There are four pairs of botryoidal vesiculae seminales, a pair in each of ix, x, xi, and xii.

There is a pair of slender meganephridial tubules in each segment except the first. The nephridia open to the exterior through a small vesicle about 1 mm. from the ventral midline on each side, close to the anterior septum of each segment.

Genus Didymogaster Fletcher

“Chaetae eight per segment. Spermathecal pores three pairs, on segments ix, x, and xi. Two gizzards, in vi and vii. Micronephridial. Prostates with branched system of ducts.” (Stephenson, 1930.)

There is only one species of the genus (D. sylvaticus) and it has been recorded from Australia (New South Wales) and from New Zealand. Stephenson (1930) regards it as introduced into New Zealand.

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Baird, W., 1871. Megascolex antarctica, an Earthworm from New Zealand. Proc. Linn. Soc., vol. 11, p. 96.

Beddard, F. E., 1890. Proc. Zool. Soc., London, 1890, p. 56.

— 1892. Further Notes on New Zealand Earthworms. Proc. Zool. Soc., London. 1892, p. 668.

Benham, W. B., 1900. On Some Earthworms from the Islands Around New Zealand. Trans. N.Z. Isnt., vol. 33, p. 129.

— 1902. On a New Species of Earthworm from Norfolk Island. Trans. N.Z. Inst., vol. 35, p. 273.

— 1903. On Some New Species of Aquatic Oligochaeta from New Zealand. Proc. Zool. Soc., London, 1903, pt. 2.

— 1903a. A note of the Oligochaeta of New Zealand Lakes. Trans. N.Z. Inst., vol. 36, p. 192.

— 1904. On Some Edible and Other Species of Earthworms from the North Island of New Zealand. Proc. Zool. Soc., London, 1904, pt. 2, p. 220.

— 1905. Additional Notes on the Earthworms of the North Island of New Zealand. Trans. N.Z. Inst., vol. 38, p. 239.

— 1905a. An Account of Some Earthworms from Little Barrier Island. Trans. N.Z. Inst., vol. 38, p. 248.

— 1909. Report on the Oligochaeta of the Subantarctic Islands of New Zealand. Subantarctic Islands of New Zealand. Government Printer, Wellington.

— 1941. Megascolides napierensis a New Species of Earthworm. Trans. Roy. Soc. N.Z., vol. 71, p. 27.

— 1942. Notoscolex equestris, an Earthworm from the Poor Knights Island. Trans. Roy. Soc. N.Z., vol. 72, p. 220.

— 1947. Studies in Earthworms. XLII. The Occurrence of the Genus Pheretima in New Zealand. Trans. Roy. Soc. N.Z., vol. 76, p. 423.

— 1949. A Yard-long Earthworm, Notoscolex hakeaphilus. Trans. Roy. Soc. N.Z., vol. 77, p. 346.

Fletcher, J. J., 1886. Notes on Australian Earthworms. Proc. Linn. Soc. N.S.W., vol. 1, p. 554

Kinberg, J. G. H., 1866. Annulata nova. Kongl. Vetenskap. Akad. Forhandlingar. Stockholm, vol. 23, p. 102.

McCoy, F., 1878. Prodr. Zool. Victoria, vol. 1, dec. 1, p. 21.

Michaelsen, W., 1900. Das Tierreich, 10—Oligochaeta. Berlin, 1900.

— 1907. Oligochaeta. Die Fauna Sudwest Australiens, vol. 1. Jena, 1907.

— 1923. Oligochaeten von Neuseeland und den Auckland-Campbell Inseln nebst einigen anderen Pacifischen Formen. Dr. Th. Mortensen's Pacific Expedition, 1914–16. No. 17.

Perrier, E., 1872. Nouv. Arch. Mus. Hist. nat. Paris, vol. 8, p. 126.

— 1873. Arch. Zool. Exp. gen. Paris, vol. 2, p. 250.

— 1874. C. R. Acad. Sci., Paris, vol. 78, p. 1582.

Stephenson, J., 1930. The Oligochaeta. Clarendon Press, Oxford, 1930.

Sweet, Georgina, 1900. On the Structure of the Spermiducal Glands and Associated Parts in Australian Earthworms. Linn. Soc. Journ. Zool., vol. 29, p. 109.

Templeton, R., 1844. Proc. Zool. Soc., London, vol. 12, p. 89.

Ude, H., 1905. Terricole Oligochaeten von den Inseln der Sudsee und von verschiedenen andern Gebieten der Erde. Zeitt. fur Wiss. Zool., vol. 83.

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New Entities in the Chlorophyceae of New Zealand

[Read before the Auckland Institute, June 20, 1951; received by the Editor, June 20, 1951]


The following new genera and species from New Zealand are described and figured: Letterstedtia stipita, Campbell Is.; Letterstedtia ulvoidca, Dusky Sound; Lobata nov. gen—type species, Lobata phyllosa n.sp., Tauranga Bay; Lobata foliosa, Catlins district; Gemina nov. gen.—type species, Gemina enteromorphoidea n.sp., Stewait Is; Gemina ulvoidea, Stewart Is; Gemina letterstedtioidea, Auckland Is. Gemina clavata, Campbell Is; Gemina linzoidea, Stewart Is.; Capsosiphon aurea, Russell; Capsosiphonaceae nov fam.; Rama nov. gen.—type species Rama novae zelandiae (J. Ag.) n. comb.; Rama antarctica, Campbell Is.


Letterstedtia stipita sp. nov. (Fig. 1)

Thallo stipitato, stipite inter brevem et longum variante, in lobum expansum cuneatum procedente, stipite raro simplici, saepius ramoso, plantis ad 5 cm. longis, 5 cm. latis, parte expansa saepe lacerata; cellulis irregulariter dispositis 13–15μ diametro, 33 5μ altis in T.S.; membrana partis expansae 86μ lata. 5μ crassa extrinsecus; plantis viridibus.

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Fig. 1 Letterstedtia stipita
(a) Plant.
(b) Cells of blade.
(c) Membrane of stipe region.
(d) Membrane of blade.
(e) T. S. stipe region.

Thallus stipitate, stipe short to long, developing into an expanded cuneate lobe, stipe rarely simple, more often branched, plants up to 5 cm. long and 5 cm. wide, expanded portion often lacerate; cells irregularly arranged. 13–15μ diameter, 33 5μ high in T. S., membrane of expanded part 86μ wide, 5μ thick on the outside; plants green, not adhering to paper. Type specimen in Herb. P.R. B., No. 69,398, from Campbell Is. (west end)

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Distribution. Local: St. Clair, Campbell Is.

In the typical form this species is quite distinct from L. petiolata*, but in some states the only means of distinguishing it is to measure the cells and the membrane.


Letterstedtia ulvoidea sp. nov. (Fig. 2)

Thallo breviter stipitato. formante laminam medialem planam e qua expansiones lobatae a basi attenuata oriuntur. lamina mediali perforata, lobis orbicularibus, planta tota ad 20 cm. diametro; cellulis 11–19μ, plerumque 13–15μ diametro, interdum per ordines breves dispositis, 15μ altis in lobis, 27–30μ altis in parte basali arctiore, duabus seriebus cellularum in parte basali per cellulas interstitiales parietibus crassis separatis; membrana 38μ lata in lobis, parietibus non crassatis, 76–80μ latis in basi parietibus crassatis; colore grammo-viridi.

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Fig. 2—Letterstedtia ulvoidea (a) T.S stipe. (b) Membrane of stipe region. (c) Cells of blade (d) Membrane of blade.

Thallus briefly stipitate, forming a flat central sheet from which lobed expansions arise from an attenuate base, central sheet perforate, lobes orbicular, entire plant up to 20 cm diameter; cells 11–19μ. average 13–15μ diameter, sometimes in short rows, 15μ high in the lobes, 27–30μ high in narrower basal portion, the two layers of cells in the basal portion separated by interstitial cells with thick walls; membrane 38μ wide in lobes. walls unthickened. 76–80μ wide in base with walls thickened; grass-green in colour, adheres well to paper. Type specimen in Herb. P.R. B. No. 55,163, from Dusky Sound.

Distribution. Local: Dusky Sound, Kohaihai.

This species combines the habit of an Ulva with the microscopie structure of a Letterstedtia. It is evident that the demarcation between the two genera is somewhat obscure. If the structure of the petiolar portion is regarded as a characteristic feature of Letterstedtia, then this plant, although more like an Ulva in appearance, must be classed as a Letterstedtia.

Lobata gen. nov.

Fronde elongata, axiale parte lobos latos ferente, membranaceosa, plana, cellulis distromaticis, appressis in lobis marginalis, in parte axis separantibus; organis generationis in partis cellulis axiali.

Frond elongate, with axis portion bearing lateral lobes, membranaceous, flat, consisting of two layers of cells, appressed in the marginal lobes. separated in axis; reproductive bodies produced in certain cells of axis portion. Type species Lobata phyllosa.

[Footnote] * See Laing, Trans. N. Z. Inst., 1926, 57, p. 133.

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Lobata ph [ unclear: ] llosa (× [ unclear: ] )

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Top: Gemina ulioidca (× ⅔). Bottom Gemina [ unclear: ] (× ⅓).

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Lobata phyllosa sp. nov. (Plate 21, Fig. 3)

Fronde lanceolata vel lineari-lanceolata, simplicr vel ramosa, ad 59 cm. longa, 4 cm. lata, ex stipite brevi oriente; parte superiore frondis constante e medial· parte axiali lobos latos frondosos distichosos ferente; cellulis partis axialis 13μ plerumque diametro, 41μ altis in T.S. cum prolongationibus in regionem medialem; cellulis loborum 14–15μ plerumque diametro, 30μ altis in T.S. sine prolongationibus, membrana loborum 75μ crassa, externe ad 5μ crassa; membrana axis 150–206μ, pariete exteriore ad 22μ crasso; colore gramino-viridi.

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Fig. 3—Lobata phyllosa. (a, c) Membrane central region. (b) Membrane lateral portion. (d, e) Cells.

Frond lanceolate or linear-lanceolate, simple or branched, up to 59 cm. long and 4 cm. wide, arising from a short stipe; upper part of frond consisting of a central axis-like portion, bearing broad leafy distichous lobes; cells of axis portion 13μ average diameter, 41μ high in T. S. with prolongations into central region; cells of lobes 14–15μ average diameter, 30μ high in T. S. without prolongations; membrane of lobes 75μ thick, outer wall up to 5μ thick; membrane of axis 150–206μ. outer wall up to 22μ thick; colour bright grass-green; adheres well to paper. Type specimen in Herb. P.R.B., No 38.129, from Tauranga Bay (Westport).

Distribution. Local. Lyall Bay, Tauranga Bay, Kowhaihai, Karamea, Chalky Inlet, Chatham Is.

This species is very characteristic and can only be confused with L. foliosa, from which it differs in the brighter green colour, the greater length and narrower width, and in the greater thickness of the membrane. In so far as the axis portion differs from the lobes, the species is intermediate between Ulva and Letterstedtia. It was originally believed to be a new species of Ulva (see label in Pl. 21 photograph), but further study and the discovery of another species indicated that it should be removed from Ulva.


Lobata foliosa sp. nov. (Fig. 4)

Fronde lanceolata vel lineari-lanceolata, ad 30 cm. longa, 8 cm. lata, ramosa ex parte basali expansiore, a stipite brevi oriente; ramis constantibus e mediali

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parte axiali appendices lobatas ferente; cellulis partis axialis 5–11μ, plerumque 8μ diametro, 26μ altis in T.S.; cellulis loborum 11–18μ, plerumque 15–16μ diametro, 26μ altis in T.S.; membrana loborum 54μ crassa, parietibus vix crassatis; membrana axis 80–90μ crassa, pariete exteriore ad 11μ crasso, interdum cum prolongationibus e cellulis in materiam interstitialem; cellulis generationis ad mediale parte restrictis; colore gramino-viridi.

Frond lanceolate or broadly linear-lanceolate, up to 30 cm. long and 8 cm. wide, sometimes branched from a more expanded basal portion, arising from a short stipe; branches consisting of a central axis portion bearing lobed appendages; cells of axis 5–11μ, average 8μ diameter, 26μ high in T.S.; cells of lobes 11–18μ, average 15–16μ diameter, 26μ high in T.S.; membrane of lobes 54–65μ thick, walls scarcely thickened; membrane of axis 80–90μ thick, outer wall up to 11μ thick, occasional prolongations from cells into interstitial material; reproductive cells apparently restricted to axial region, swelling to 30μ; colour pale grass-green; adheres moderately to paper. Type specimen in Herb. P.R.B., No. 69,149, from Catlins District.

Distribution. Local: Levin, Waipapa boatshed, Gore Bay, Wellington Heads, Karaka Bay, Lyall Bay, Tauranga Bay, Tautuku Bay (Catlins District), Timaru, St. Clair, Auckland Is., Chatham Is.

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Fig. 4—Lobata foliosa. (a) Cells of lateral portion. (b) Cells of central portion. (c) Membrane of central portion. (d) Membrane of lateral portion. (e) T.S. of central portion with reproductive cells?

One of the most interesting features of this species was the presence, in one of the herbarium specimens, of large swollen cells in the axis portion (Fig. 4). These cells contained rounded bodies and it is believed that they represent the reproductive cells. If this is correct, we have here a species in which certain cells become modified for reproduction and are restricted to a portion of the thallus. Exactly similar cells were seen in a Californian specimen collected in the Gulf of California by Dr. Yale Dawson.

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Gemina nov. gen.

Thallo filamentoso vel foliaceoso, distromatico, stipite vel praedito vel non praedito; cellulis intra membranam parentalem in pares dispositis, rarius in ternas vel quaternas, longis et arctis in T.S.; organis generationis in partis cellulis ad basem.

Thallus filamentous or foliaceous, distromatic, with or without a stipe; cells in pairs within the parent membrane, less frequently in threes or fours, long, narrow and conical in transverse section; reproductive bodies in special cells at base of thallus. Type species: Gemina enteromorphoidea.

This is a remarkable genus because it combines the characters of Enteromorpha, Ulva and Capsosiphon. The species are all characterised by the regular paired arrangement of the cells, the pairs commonly being themselves arranged in short rows. G. enteromorphoidea has the exact appearance of an Enteromorpha whilst G. ulvoidea looks like a species of Ulva. In the cell arrangement the genus agrees with Capsosiphon, but the thallus is never tubular, always distromatic, and there are no areolae.


Gemina enteromorphoidea sp. nov. (Fig. 5)

Planta simplici, compressa, ad 37 cm. longa, 5 mm. lata, tubulosa, ad stipitem attenuata, per discum minutum scutatum adfixa; cellulis parvis 3.75–9·3μ, plerumque 5.5μ diametro, in pares vel quaternas distincte dispositis, paribus ipsis in series breves dispositis membranis singulis 36μ latis, pariete exteriore 5μ crasso, pariete interiore 4μ, cellulis 27μ altis; colore fusco-gramino-viridi vel oleagineo.

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Fig. 5—Gemina enteromorphoidea. (a) Membrane. (b–d) Cells in surface view. (e) T.S. of thallus

Plant simple, compressed, up to 37 cm. long and 5 mm. wide, tubulose, attenuate to the stipe, attached by a minute scutate disc; cells small, 3.75–9·3μ, average 5.5μ diameter, arranged in distinct pairs or fours, the pairs themselves in short rows; single membrane 36μ wide, the outer wall 5μ thick, the inner 4μ, the cells 27μ high in T.S.; colour deep grass-green or olive-green, adhering moderately to paper. Type specimen in Herb. Auckland Institute from Stewart Island.

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Distribution. Local: Stewart Is., Campbell Is.

This is a very remarkable species, particularly in view of the relationships with allied genera. The arrangement of the cells in pairs or fours is characteristic of the genus Capsosiphon, but there is no evidence of any parent mother cell membranes, nor are there any areolae. Except for the absence of areolae, the species would be very similar to Capsosiphon aurea (see p. 54). In fact, this species, C. aurea and C. fulvescens form a definite series from the normal Enteromorpha type of construction to the normal Capsosiphon type of construction.

The small size of the cells suggests that it is allied to Enteromorpha bulbosa, though it differs in the cell arrangement and also in the very elongate cells as seen in T.S. Another interesting feature is seen in the fusion of the central part of the tube to produce an Ulva-like arrangement. In this respect, together with Ulva atroviridis of South Africa, it forms a transitional form between the Enteromorpha and Ulva types of construction.


Gemina ulvoidea sp. nov. (Plate 22)

Thallo membranaceoso, frondem complanatam expansam ad 13 cm. longam, 7 cm. latam formante, perforato; cellulis parvis, 3.75–9.3μ diametro, in pares vel quaternas dispositis, paribus interdum in series breves dispositis; cellulis 27μ altis in T.S., membrana 70–76μ crassa, pariete exteriore ad 5μ crasso.

Plant simple, membranaceous, forming a flattened expanded frond up to 13 cm. long and 7 cm. wide, perforated; cells small, 3.75–9.3μ diameter, in pairs or fours, the pairs sometimes arranged in short rows; cells 27μ high in T.S., membrane 70–76μ thick, the outer wall up to 5μ thick. Type specimen in Herb. Auckland Institute from Stewart Island.

Distribution. Local: Lyall Bay, Stewart Is.

This remarkable form collected by Mr. V. W. Lindauer has the same microscopical features as the preceding species, but differs from it in habit. Another specimen which appears referable to this species has been collected from Lyall Bay.


Gemina letterstedtioidea sp. nov. (Fig. 6)

Thallo membranaceoso, compresso, ad 8 cm. longo, 1 cm. lato, ad stipitem attenuato; plantis integris vel perforatis, simplicibus vel ramosis, thallo primario vel ramis linearibus vel in lobum expansis; cellulis 7.5–11μ diametro, plerumque 8μ, in pares dispositis, paribus in series breves dispositis, 49μ altis in T.S., membrana 116μ lata, pariete exteriore ad 7μ crasso.

Thallus membranaceous, compressed, up to 8 cm. long and 1 cm. wide, attenuate to a stipe; plants entire or perforate, simple or branched, main thallus or branches linear or expanding into a lobe; cells 7.5–11μ diameter, average 8μ, in pairs, the pairs in short rows, 49μ high in T.S.; membrane 116μ wide, the

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Fig. 6. Gemina letterstedtioidea. (a) Membrane. (b) Cells.

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outer wall up to 7μ thick. Type specimen in Herb. P.R.B., No. 44,096, from Auckland Is.

Distribution. Local: Campbell Is., Auckland Is.

This member of the genus has a remarkable superficial resemblance to Letterstedtia, but microscopically the paired arrangement of the cells places it in this genus.


Gemina clavata sp. nov. (Fig. 7)

Thallo simplici, minuto, ad 1.5 cm. alto, 0·5 cm. lato, clavato, contorto, ad arctum stipitem repente attenuato; cellulis parvis 3.5–9μ diametro, 27μ altis, intra parietem parentalem cellularum in pares vel quaternas dispositis; membrana 70–76μ lata; colore fusco-viridi.

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Fig. 7—(I) Gemina clavata, plants. (II) G. linzoidea. (a) Cells. (b) Membrane.

Thallus simple, minute, up to 1.5 cm. tall and 0.5 cm. wide, clavate, contorted, attenuate sharply to a narrow stipe; cells small, 3.5–9μ diameter, 27μ high, in pairs or fours within the parent cell wall; membrane 70–76μ wide; brownish-green in colour. Type specimen in Herb. P.R.B., No. 62,450, from Campbell Is. Co-type in Auckland Institute.

Distribution. Local: Campbell Is.

The data on the herbarium sheet are not sufficient to indicate whether this is a habitat form or not. It differs so markedly from the other species that it seems unlikely that it can be merely a habitat variation.


Gemina linzoidea sp. nov. (Plate 22, Figs. 7 and 8)

Planta ad 27 cm. longa, 6 cm. lata, in parte latissima, paulatim in regionem stipitis decrescente; parte superiore frondis laciniata, integra vel perforata; cellulis in pares, rarius in trinas vel quaternas dispositis, membrana parentali circumventis; cellulis 5–9μ, plerumque 7μ diametro, 41–49μ altis in T.S., conicis, membrana 116–131μ, pariete exteriore 4–7μ crasso; colore oleagineo-viridi.

Plant up to 27 cm. long and 6 cm. at the widest part, narrowing gradually to a stipe region; upper part of blade laciniate, entire or perforate; cells in pairs, more rarely in three or fours, surrounded by parent membrane; cells 5–9μ, average 7μ diameter, 41–49μ high in T.S., conical; membrane 116–131μ, outer wall 4–7μ thick; olive-green; adheres imperfectly to paper. Type specimen in Herb. V. W. Lindauer from Stewart Is.

Distribution. Local: Stewart Island.

This species differs distinctly from all the others and is named for its superficial resemblance to Ulva linza. Present evidence suggests that it probably has a wider distribution. By chance a specimen, agreeing in all characters, was encountered in Leiden Herbarium (Herb. Lug. Bat., 939/23/232 ex Herb. Hauck) and labelled Phycoseris lobata B africana, though no description appears to

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have been published. The form of the plant was very similar to the one found in New Zealand and the microscopical anatomy was identical. The most important feature of this plant was the discovery of the reproductive cells towards the base of the thallus. Only certain cells appear capable of producing swarmers, instead of all the cells as in Ulva. (Fig. 8) In this respect the plants are like Lobata. These cells are very conspicuous because of their size. They are not sunk in the thallus as they are in Lobata.

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Fig. 8—Gemina linzoidea.
Top: Reproductive and vegetative cells seen in surface view.
Below: The same in section. (Leiden specimen.)

Capsosiphon Gobi

Capsosiphon aurea sp. nov. (Fig. 9)

Plantis filamentosis, tubularibus, ad 4 cm. longis, 2·5 mm. latis, colore fulvo vel aureo, per discum basalem adfixis; cellulis vel binis vel quaternis per ordines longitudinales et horizontales dispositis, catervis ordinum longitudinalium laxe conexis; cellulis vegetativis 5.5–11.2μ diametro, cellulis generatibus 11·2–13μ

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Fig. 9.
Capsosiphon aurea.
(a) Plant. (b) Cells of old thallus. (c) Cells of young branch. (d) Membrane. (e) Apex of young branch. (f) Plantling.

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dia., celluls filamentorum novellorum plerumque in pares dispositis, interdum elongatis, 3·75–7·5μ longis; membrana 15–21μ lata, cellulis 7·5–11·2μ dia., plus minusve sphericis; generatione per ? aplanasporas, et quaque cellula octonas.

Plants filamentous, tubular, up to 4 cm. long, 2.5 mm. wide, brown or golden in colour, attached by a basal disc; cells in twos and fours, arranged in longitudinal and horizontal series, groups of the longitudinal series loosely connected; vegetative cells 5·5–11.2μ diameter, reproductive cells 11.2–3μ diameter, cells in young filaments usually in pairs, sometimes elongate, 3·75 × 7·5μ long; membrane 15–21μ wide, cells 7·5–11·2μ diameter, more or less spherical; reproduction by ? aplanospores, 8 per cell. Type specimen in Herb. Auckland Institute from ditch at Russell (V.W.L. No. 9065).

Distribution. Local: Russell (Bay of Islands).

This plant is extremely interesting, because it is clearly a Capsosiphon, yet it differs in certain important respects from C. fulvescens, the only other known species. In C. fulvescens the pairs of cells are retained within the sheath of the parent cell in a Gloeocapsa-like form. In C. aurea there is no distinct parent mother cell sheath Both species are tubular, lubricous and with the longitudinal series of cells loosely connected. In C. fulvescens each row of cells separates, in C. aurea several rows separate together. Except for its tubular nature the construction of C. aurea is very akin to that of a Prasiola. Members of the genus Prasiola are always flat plates, whereas Capsosiphon is tubular.

Ecologically, C. fulvescens and C. aurea are similar, because both are found in streams where the plants are subjected to tidal influence twice daily.

In view of the similarities between C. aurea and Prasiola, it is considered desirable to remove Capsosiphon from the Ulvaceae and create a separate family, the Capsosiphonaceae, which lies between the Ulvaceae and the Prasiolaceae. The family also differs from the Ulvaceae in the method of reproduction, as there is no evidence of sexual or asexual swarmers.

Capsosiphonaceae Nov. Fam.

Thallo membranaceoso, tubulo, membrana monostromatica, per discum adherente; cellulis uninucleis cum uno chloroplastio, vel binis vel quaternis in seriebus longitudinalibus dispositis; generationis ad aplanosporas.

Thallus membranaceous, tubular, with wall of a single layer of cells, attached by a disc; cells uninucleate with a single chloroplast, arranged in twos and fours in distinct longitudinal series; reproduction by aplanospores.


Rama Gen. Nov.

Filamentis prostratis, contortis, unae series cellularum compositis, sparsis ramosis a basalem, ramis rhizoidalibus unicellularibus; chloroplastio reticulato, parieto, pyrenoidis numerosis.

Filaments prostrate, tangled, formed of a single series of cells, branched very sparsely at the base, with few rhizoidal outgrowths of single cells; chloroplast reticulate, parietal, with numerous pyrenoids. Type species, Rama novae zelandiae.

This genus has been created to take three species with sparse branching like a Cladophora, but with rhizoidal outgrowths like a Rhizoclonium. The species are not well known and the method of reproduction has not been studied. It contains two species that J. Agardh (1877) described from New Zealand and placed in the genus Chaetomorpha De Toni (1881) transferred these species,

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because of the branching, to Cladophora. The degree of branching is so slight and so restricted that they are unlike any other species of Cladophora. The species with nearest affinity appears to be C. verticillata.


Rama longiarticulata J. Ag. (Fig. 10)

De Toni, Syll. Alg., p. 332, 1889; J. Ag; De Alg. Nov. Zel. Mar., p. 2, 1877; Laing, Trans. N.Z. Inst., 57, 134, 1926.

Plant decumbent, attached at first by rhizoids, then free-floating; rhizoids few at the base, filaments sparsely branched at base, branches simple, very long, capillary, cells 33–40μ wide, 4–8 times longer than broad; light green; does not adhere to paper.

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Fig. 10.
Rama longiarticulata.
(a) Basal region.
(b) Rhizoid.
(c) Terminal portion of filament.

Distribution. Local: Banks Peninsula, Chatham Is. Endemic.

This species appears to lie on the border between Chaetomorpha and Rhizoclonium. There is no branching in the specimen examined,* although such is described by Agardh. It is on account of this reported branching that it is removed to the new genus. The attachment rhizoids have some similarity to those of Rhizoclonium, but they are restricted in their position. The species is much less like a Rhizoclonium than R. novae zelandiae and R. antarcticum.


Rama novae zelandiae (J. Ag.) nov. comb. (Fig. 11)

De Toni, Syll. Alg., vol. I, p. 332, 1889. as Cladophora novae zelandiae; J. Ag., De Alg. Nov. Zel. Mar., p. 2, 1877. as Lychaete novae zelandiae.

Thallus caespitose, decumbent, attached at first, then floating; lower filaments prostrate, with rhizoidal branches, sparsely branched, branches very long, rarely rebranched and with occasional rhizoidal outgrowths; cells 75–95μ wide, walls thick, 1 ½–2 ½ times as long as broad.

Distribution. Local: Russell, Otago. Warrington, and probably elsewhere. Endemic.

The problem of this species was solved by the discovery of a specimen identified by J. Agardh in the herbarium of the Auckland Institute. Until this specimen was found the author was of the opinion that Rhizoclonium hookeri existed in a simple and branched form. It now appears that R. hookeri has rather narrower filaments and is unbranched, whilst this species has broader filaments (75–95μ as against 47–70μ) and is branched at the base. The presence of the unicellular rhizoidal branches would seem to remove it from the genus Cladophora, where it was placed by De Toni, though in possessing branches and rhizoids it is approaching members of the subgenus Spongomorpha. It differs from species of Spongomorpha in being very loose.

[Footnote] * Plant collected by Berggren and identified by J. Agardh and now in the herbarium of the Auckland Institute.

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Fig. 11—Rama novae zelandiae. Basal region with rhizoid and branching.


Rama antarctica sp. nov. (Fig. 12)

Filamentis rigidis, interdum per cellulam basalem ecolorem immutatam adhaerentibus, valde pullis, in fusca aresccntibus, laxe intertortis, ad basim ramosis; cellulis 111–126μ diametro, 1–1½-cies longioribus quam latioribus, parietibus 15–22μ crassis, paulum lamellatis; ramis rhizoidalibus unicellularibus, e cellula materna orientibus, perbrevibus, saepe ad aliud filamentum adhaerentibus.

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Fig. 12.
Rama antarctica.
(a) Rhizoid.
(b) Branched thallus.
(c, d) Filaments united by rhizoids.

Filaments rigid, sometimes attached by a modified colourless basal cell, very dark green, becoming brownish when dry, loosely intertwined, branched near the base; cells 111–126μ diameter, 1–1½ times as long as broad, walls 15–22μ thick, slightly lamellate; rhizoidal branches present, unicellular, arising from parent cell, very short, often attached to a second filament; does not adhere to paper. Type specimen in Herb. P. R.B., No. 67,807, from Campbell Is.

Distribution. Local: Snares. Auckland Is., Campbell Is.

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This species is close to Rhizoclonium hookeri, from which it differs by its greater size and coarseness. There is also the branching of the filaments and the habit of the rhizoids to attach themselves to other filaments. The distribution of the species shows that it is quite clearly a subantarctic form.

I am grateful to D.S.I.R. for permission to examine material in the herbarium of the Plant Research Bureau, to the Auckland Institute for permission to examine material in the herbarium, to Canterbury University College for the loan of the Laing Collection, to Mr. V. W. Lindauer, of Russell, for much material and use of his herbarium, to Mr. L. W. Crawley, of the Classics Department of Auckland University College, for all the Latin diagnoses except that of the family Capsosiphonaceae and that of the genus Rama, and to Miss S. Baker, of my department, for taking the photographs.


Agardh, J., 1877. De Algis Novae Zelandiae Marinis. Lunds. Univ. Arssk., 14.

De Toni, G., 1889. Sylloge Algarum, vol. 1. Patavii.

Laing, R. M., 1926. A Reference List of New Zealand Marine Algae. Trans. N.Z. Inst., 57, p. 126.

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Additions to the Rotatoria of New Zealand. Part 4

[Read before the Canterbury Branch, June 6, 1951; received by the Editor, June 15, 1951]


This paper contains a list of nineteen species of the Rotatoria not previously recorded in New Zealand. Nearly all the specimens were collected from small, weedy lakes and acid water swamps in the South Island. Keratella carinata, described as a new species in a previous paper, in this series is now declared to be a synonym of K. javana Hauer.


From the samples that have been collected from the larger lakes of the South Island it is thought that a large portion of the rotifer fauna from these habitats has now been listed, and out of nine collections made during 1950–1 only two unrecorded species were found. Representative samples from North Island lakes, so far, do not show the wide variety of species found in the Southern lakes, but this may be due to the method of collection. Intensive collections will have to be made from the Northern lakes covering the seasonal changes of temperature and pH before any reliable indication can be gained of their populations. Alkaline pools and lakes generally have the largest populations, but acid swamps contain a larger variety of species. Two collections from North Canterbury swamps contained only two rotifers that had previously been recorded in this country, the remainder being unlisted.

The rotifers so far found in New Zealand are contained in the following papers: Russell, “A New Rotifer from New Zealand,” Jour. Royal Micr. Soc., 1944, vol. 64, pp. 121–3. Russell, “A Reference List of the Rotatoria of New Zealand with Ecological Notes,” Trans. Royal Soc. N.Z., vol. 75. pp. 102–23. Russell, “Additions to the Rotatoria of New Zealand, Part I,” Trans. Royal Soc. N.Z., vol. 76, pp. 403–8. Russell, “Additions to the Rotatoria of New Zealand, Part 2.” Trans. Royal Soc. N.Z., vol. 77, pp. 351–4. Russell, “Additions to the Rotatoria of New Zealand, part 3,” Trans. Royal Soc. N.Z., vol. 78, pp. 161–6. Russell, “The Rotatoria of the Upper Stillwater Swamp.” Canterbury Museum Records, vol. 5, no. 5.

At the suggestion of some rotifer workers the species have been listed in this paper in alphabetical order instead of being arranged in their systematic position.

When Part 3 of “Additions to the Rotatoria of New Zealand” was written a number of unidentified slides remained from Professor E. Percival's collections from the Southern Lakes; these are included in this paper and marked (P) under the heading, “locality”.

Genus Colurella

Colurella hindenburgi Steinecke, 1916

1916. Schrift. Phys-okonom. Ges Konigsberg i P. Jahrg. 57: 90. 97.

Locality. Lake Ellesmere, temp. 14°C, brackish water. Common. Lenth body 100μ, width 40μ, toes 52μ. Specimens generally were larger, and more acute posteriorly than those described by Steinecke.

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Genus Cupelopagis

Cupelopagis vorax (Leidy)

1857. Dictyophora vorax Leidy. Prod. Acad. Nat. Sc. Phila., vol. 9, p. 205.

Locality. Victoria Lake, and lily ponds in Wilson's Nursery, Christchurch. Temp. 21–23°C. pH 8. Abundant. Both the free-swimming immature forms and the fully developed sessile females were found. Over-all length of fully grown animal 550μ, maximum width of body 270μ.

Genus Cephalodella

Cephalodella physalis Myers, 1924.

1924. Rotifer Fauna Wis., pt 2. Wis. Acad. Sc. Art. Lett., vol. 23, pp. 484–5.

Locality. Acid water swamp near L. Katrine, North Canterbury. Temp. and pH unknown. Common. Length over all 160μ, toes 40μ, trophi 30μ.

Cephalodella tantilla Myers, 1924

1924. Rotifer Fauna Wis., pt. 2, Wis. Acad. Sc. Art. Lett., vol. 23, p. 486.

Locality. L. McGregor, Canterbury. Not common. Normal specimens. (P.)

Genus Eosphora

Eosphora anthadis Harring and Myers, 1920.

1920. Rotifer Fauna Wis., pt. 1, Wis. Acad. Sc. Art. Lett., vol. 20, pp. 641–2.

Locality. Acid water swamp near L. Katrine, North Canterbury. Not common. Length 500μ over all.

Genus Euchlanis

Euchlanis meneta Myers, 1930

1930. Rotifer Fauna Wis., pt. 5, Wis. Acad. Sc. Art. Lett., vol. 25, p. 378.

Locality. L. McGregor, Canterbury. Common. Length dorsal plate 160μ, width 140μ. Length ventral plate 140μ, width 100μ. Toes 70μ. The lateral sulci are very deep in most specimens which are longer and wider than those described by Myers. (P.)

Genus Filinia

Filinia terminalis (Plate)

Triartha terminalis Plate. 1886. Jenische Natura., vol. 19, p. 19.

Locality. L. Pearson, Canterbury. Temp 9°C. Common. It has also been found sparingly in other high-altitude lakes. I agree with Edmondson (1935) that F. terminalis (Plate) is not a synonym for L. longiseta Ehrenberg. In L. Pearson, specimens were found having a body-length of 400μ, and appendages 500μ and 480μ respectively in length. Apart from the much greater size of F. terminalis it differs from F. longiseta in both the anterior and posterior details and shape of body. (P.)

Genus Lecane

Lecane (Monostyla) pygmaea Daday, 1897.

1897. Lecane (Monostyla) pygmaea Daday. Math. Termész Ertes, vol. 15, p. 139.

Locality. Acid water swamp near L. Katrine, North Canterbury. Not common. Length over all 146μ.

Lecane arcula Harring, 1914.

1914. Proc. U.S. Nat. Mus., vol. 47, p. 539.

Locality. Acid swamp near L. Katrine. North Canterbury. Not common. Length over all 146μ.

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RussellAdditions to the Rotatoria of New Zealand

Lecane luna var. presumpta Ahlstrom 1938.

1938. Jour. Elisha Mitchell Sci. Soc., vol. 54, no 1, p. 97.

Locality. L. McGregor, Canterbury. Fairly common. Length over all 163μ. (P.)

Genus Lepadella

Lepadella latusinus var. americana Myers, 1934.

1934. Amer. Mus. Nov. no. 760, p. 7.

Locality. L. Ellesmere. Canterbury, a brackish lake, and in L. McGregor, Canterbury, where it occurred in Professor Percival's collections. L. latusinus was described by Hilgendorf (1899) from Taieri Beach, and was also found at Mt. Cook by Murray; it has since been found in many habitats. L. latusinus is a very variable species. and it is considered that a search would show a complete range of forms from L. latusinus to var. americana

Lepadella whitfordi Ahlstrom, 1938.

1938. Jour. Elisha Mitchell Sc. Soc., vol. 54, no. 1, p. 100.

Locality. L. McGregor, Canterbury. Not common. Length over all 133μ. The collar was clear but not pronounced. (P.)

Genus Mytilina

Mytilina trigona (Gosse)

Diplax trigona Gosse. 1851. Ann. Mag. Nat. Hist, sei. 2, vol. 8, p. 201.

Locality. L. McGregor, Canterbury. Not common. Total length 185μ, toes 70μ. (P.)

Genus Pompholyx

Pompholyx complanata Gosse, 1851.

1851. Ann. Mag. Nat. Hist, sei. 2. vol. 8, p. 203.

Locality. L. Pearson, Canterbury. Common. Length. 114μ. (P.)

Pompholyx sulcata Hudson, 1885.

1885. Jour. Royal Micr. Soc. p. 613.

Locality. L. Alice Temp. 8°C. pH 6. Not common. Total length 115μ.

Genus Proales

Proales longidactyla Edmondson, 1948.

1948. Amer. Micr. Soc., vol. LXVII, pp. 150–2.

Locality. Acid water swamp near L. Katrine, North Canterbury. Not common. Total length of animal 146μ, head 41μ, toes 20μ. The specimens differ in several particulars from the animal described by Edmondson: the head is longer, there are a number of longitudinal folds round the body, the toes are shorter with their distal points recurved, and in dorsal view the animal is somewhat bdelloidal in appearance. In trophi and other details the animals are identical. It should be noted that Edmondson's specimens came from the psammon.

Genus Scepanotrocha

Scepanotrocha rubra Bryee, 1910.

1910. Quek. Micr. Club. ser. 2; vol. II p. 78.

Locality. Acid swamp near L. Katrine, North Canterbury. Not common Length over all 212μ; 7–8 teeth on each ramus. This is the first time any member of this genus has been recorded in New Zealand.

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Genus Squatinella

Squatinella mutica (Ehrenberg)

Stephanops muticus Ehrenberg. 1831 Abh. Akad. Wiss. Berlin. p. 138.

Locality. Pools near Auckland, where it was collected by Mr. I. W. T. Munro, of Christchurch. Total length 110μ. Fairly common.

Genus Trichotria

Trichotria truncata (Whitelegge)

Dinocharis truncatum Whitelegge. 1889. Proc. Royal Soc. N.S.W, vol. 23, p. 315.

Locality. L. McGregor, Canterbury. Not common Length of body 110μ, width 90μ. foot 45μ, toes 90μ (P.)


This paper would not have been possible without the assistance I have received from a number of people throughout the Dominion who have sent me samples. I would like to thank Professor E. Percival, of Canterbury University College, for the use of his extensive back-country lake collections; Mr. H. Talbot, of Springfield, for collections from the Lake Katrine and other districts; Mr. B. T. Cunningham, of the Fisheries Laboratory, Wellinington, for samples from Lake Alice; Mr. I. W. T. Murno, of Christchurch, for collections from Auckland; Miss Violet Jolly, of Otago University, for samples from both North Island and Southern lakes. I am also indebted to Mr. K. M. Wilson, of Christchurch, for the use of the ponds at his plant nursery, where the life history of some sessile rotifers is being studied.


Edmondson, W. T., 1935. Some Rotatoria from Amzona Trans. Amer. Micr. Soc., vol. 54. no 4. p. 303.

Edmondson, W. T., 1948. Two New Species of the Rotatoma from Sandy Beaches. Trans. Amer. Micr. Soc, vol. 67, no 2, pp. 150–2

Hilgendorf, F. W., 1899. A Contribution to the Rotitera of New Zealand Trans. Royal Soc. N.Z., vol. 31, p. 131.

Russell, C. R. See introduction.

Correction to “Additions to the Rotatoria of New Zealand.” Part 3

Trans. Royal Soc. N.Z., vol. 78, pp. 161–6.

Keratella carinata n.sp.

Dr. Bruno Berzins, of Lunds University, Sweden, has very kindly forwarded me a description and sketch of Keratella cochlearis var. Javana Hauer, 1937 Archiv. Hydrobiol., supl. xv, p. 382. This animal was described from a single specimen collected from Java. Ahlstrom, Bull. Amer. Mus. Nat. Hist, vol. LXXX, p. 434, gives a description of Hauer's animal without a drawing, and raises it to specific rank as K. javana Hauer. Although K. carinata differs in the length and shape of the anterior and posterior spines, the dorsal patterns which are of an unusual type are identical, and from a re-examination of the material I have no hesitation in saying that K. carinata must be considered a synonym for K. javana Hauer.

I am much indebted to Dr. Berzins for sending me the description with a sketch of the type specimen.

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Studies on the New Zealand Amphipodan Fauna
No. 1—The Family Cyamidae: The Whale-louse Paracyamus boopis*

[Read before the Wellington Branch, June 19, 1951: received by the Editor, June 19, 1951]


The Whale Louse, Paracyamus boopis. is described and figured from specimens collected from a humpback whale at Te Awhaiti, South Island A list of Cetacea occurring in New Zealand and their Cyamid parasites as found elsewhere is given.

Since species of the Family Cyamidae are host-specific, their dispersal is dependent on the territorial range of their host. For the purposes of this paper it has been taken that Cyamids from whales caught within New Zealand territorial waters or from whales cast on New Zealand beaches belong to the New Zealand fauna.

Commercial whaling in New Zealand is now almost entirely confined to Megaptera nodosa, the humpback whale. As far as records are available, and my material shows, the New Zealand Cyamid fauna is limited to one species, Pracyamus boopis (Lutk.), the common parasite of the humpback. Whales taken at Te Awhaiti Whaling Station, Picton, are all, to a lesser or greater degree, infected with the lice. The infestation is generally heaviest around the genital apertures, but is found all over the body, particularly where there is barnacle infestation, Coronula diadema. The lice will readily attach themselves to persons coming into contact with them. At least two of the whales caught off Te Awhaiti in recent years were considered too badly infested for handling and were not processed. In other cases. where there are bare exposed surfaces, the lice have congregated and appear to have been responsible for breaking throught the pigmented layer of the skin. It seems unlikely that the vitality of heavily parasitised whales could remain unimpaired.

Records and Synonymy of New Zealand Specimens

The first record of whale lice in the New Zealand literature is a description by Chilton (1883) of Cyamus ceti (Linn. 1780). from Virgia breviceps (= Kogia breviceps).

A later reference to Cyamus ceti (Thomson and Chilton, 1886), and referring in part to the same specimens, states:

“Parasitic on whales (Virgia breviceps), C.C. It appears to be common on various whales (and sharks?). I have it from several localities in the New Zealand seas, G.M.T. On small humpbacked whale, Napier, A. Hamilton.”

The remark, “and sharks?”, is one for which I have seen no verification.

Cyamids from different localities are included in the late Professor Chilton's collection, and I have identified them as follows:

[Footnote] * This study is part of an investigation being carried out with the aid of a New Zealand University Research Fund Fellowship.

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Paracyamus boopis (Lutk.)

There are two tubes of this species, one labelled “Cyamus erraticus, Roussel de Vauzème. On whale. Parengarenga. T. F. Cheeseman”, and the other, “Cyamus ceti, Linn Napier. A. Hamilton.” These latter specimens are probably the ones referred to above as from Virgia breviceps.

Cyamus ceti Linn.

One tube labelled “Cyamus ceti, Linn. Whaler ‘Othello.' G.M.T.” I have been unable to establish a more definite locality for these. Stephensen (1942) maintains the species Cyamus ceti as described by Linnaeus, although Barnard (1932) considered Linnaeus' species to be a composite one, and rejected it in favour of Cyamus mysticeti (Lutk.). Cyamus ceti Linn has precedence.

Cyamus ovalis R. de Vauz.

One tube labelled “Cyamus ovalis R. de Vauzème. New Zealand From Indian Museum. No. 1464.” Mr. K. K. Tiwari, of the Zoological Survey of India, informs me that these were exchange specimens obtained from the Canterbury Museum, 2/8/1876, through Dr. J. von Haast. Mr. R. R. Forster has examined records at Canterbury Museum, and I have been through Dr. Haast's papers in the Turnbull Library. Apart from corroboration that a shipment of Crustacea was received by Mr. Wood-Jones, of the Indian Museum, about that date, no further information has been revealed.

Since the localities for the last two species are so indefinite, I can only note the presence of the specimens as possibly from New Zealand.

My specimens, from Megaptera nodosa, are undoubtedly Paracyamus boopis.

Genus Paracyamus G. O. Sars

Sars, 1895, p. 669. Stephensen, 1942, p. 451.

Sars' definition of the genus has been paraphrased by Barnard thus: “Gnathopods 1 and 2 unequal. Antennae 2, 4-jointed. Maxilla 2 with outer lobes. Maxilliped, palp well-developed in young, but rudimentary in adult. Dactylos and unguis of gnathopod 1 evenly tapering, the latter not distinct Branchiae single on both segments 3 and 4.”

Paracyamus boopis (Lutk.). (Plates 23, 24; Text-figures 1–7)

Cyamus boopis, Lutken, 1873, p. 262, pl. iii, Fig. 6.

Cyamus pacificus, Lutken, 1873, p. 264, pl. iii. fig. 7.

Paracyamus boopis, Barnard, 1932, p. 312.

Male, length 11 ½ mm.; width 4 mm.; first antenna, 5 ¼ mm.

Female. length 9 ½ mm.; width 4 mm.; first antenna, 3 ½ mm.

Antennae. First: Four-segmented Segments successively narrower; first and second subequal in length, third segment ¾ length of second; last segment less than half length of third, fringed with setae terminally and along superior distal margin Second: Four-segmented. Second segment slightly shorter than the narrower third segment, both with scattered long setae. Last segment less than half length of third, with strong bristles terminally and superiorly.

Maxillae. First: Of two lobes, the inner with 7 strong curved spines, at base of innermost spine is an area of fine bristles. Outer lobe small, not reaching as far as extremity of spines on inner lobe, projecting as a small setose palp.

Second: One lobe only present, terminally bifurcate to two small lobes; inner lobe the smaller and bearing two strong setae, outer with a number of setae.

Maxilliped. Adult: Inner lobes subtriangular, tending to ovate; with terminal tuft of strong setae. Outer lobes present, emerging from oblique outer margin

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Paracyamlis boopis (Lutk.)
(1) Adult male, dorsal view. (2) Adult female, vential view. (3) Vential surface of segments 5, 6, 7, male. (4) First antenna. (5) Second antenna. (6) Upper lip. (7) Maxilliped. (8) Second maxilla. (9) First maxilla. (10) Left mandible. (11) Right mandible. (12) Lower lip. (Appendages figured 3–12 not all to same scale. Scale in millimetres.)

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Paracyamus boopis (Lutk.)
(1) Gnathopod 1, male. (2) Gnathopod 2, male. (3) Peraeopod 3, male. (4) Gnathopod 1, female. (5) Gnathopod 2, female. (6) Gill of adult male. (7) Gnathopod 2, Juvenile male. (8) Gill of Juvenile male. (9. 10. 11) Ursome appendages.

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Series of maxillipeds from Paracyamus boopis Juveniles of various sizes.
(1) Juvenile, 1 mm long. (2) Juvenile, 1 ½ mm. long. (3) Juvenile, 2 mm. long. (4) Juvenile, 2 ⅔ mm. long. (5) Juvenile, 3 mm. long. (6) Juvenile, 3 ⅔ mm. long. (7) Juvenile, 4 ⅓ mm. long.

of inner lobes, small and somewhat globular, with similar distal setae. Juvenile: Total length 2 ⅔ mm. Palp present, four-segmented, attached to outer margin of outer lobe. First segment more than twice as large as outer lobe, second and third segments successively longer; dactylos subequal with third segment, stout, curved, tapering, with row of minute setules distally along inner margin.

Upper Lip. Dorsal margin with wide, shallow cleft dividing lip into two lobes, each finely bristled terminally and along inner margins.

Lower Lip. Inner and outer lobes present. Inner slightly cleft medially, raised on mound, both mound and lobe with strong bristles. Outer lobes much larger, subovate, inner and distal margins strongly setose.

Body. Male: General outline of body ovate in dorsal view. Head parallelsided. Segment 1 fused with and widening sharply from head, then rounding laterally and distally. Segment 2 subrectangular, nearly twice as long as head, and half as wide again, anterior and posterior margins excavated sinuously to projecting anterolateral and posterolateral corners. Lateral margins between projections also sinuous. Segments 3 and 4 more or less spindle-shaped, widening a little laterally, posterior angles bluntly produced. Segments 5 and 6, anterior and posterior margins parallel, broadly arched and broadly concave posteriorly; last segment subtriangular, slightly broader than long. Body segments 6 and 7 each with a single pair of sharp tubercles ventrally. Female: Ventral surface of segment 5 with pair of inwardly directed blunt processes; a single pair of pointed tubercles on each of segments 5, 6, and 7.

Mandibles. Molar process absent. Right mandible with cutting edge of upper process with six teeth, and lower two-toothed spine. Surface and margin immedi-

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ately below lower spine finely setulose, below that again a single setulose spine. Left mandible with upper process of cutting edge bearing three teeth, lower spine with five. Below spine, inner margin finely bristled, inner margin medially with setulose spine.

Gnathopod 1. Male: Basos sharply constricted proximally, twice as long as distal width; ischium subequal in width with basos, half as long as wide. Merus subovate, narrower than ischium, length twice width, distal two-thirds of anterior margin contiguous with proximal margin of carpus. Carpus subtriangular, width ⅔ length, anterior margin twice length of posterior. Propod subtriangular, anterior margin rounded, more than twice length of straight posterior margin which rounds sharply to transverse palm Palm with strong tooth medially, excavated between tooth and dactylos, which is curved and as long as palm. Tooth and outer margin of palm minutely serrate. Female: Propod narrower at distal end than in male; palm oblique, poorly defined, minutely denticulatc, small setae at dactylos base and on single median tooth.

Gnathopod 2. Male: Merus subrectangular, as long as wide; anterior margin produced medially to strong blunt projection reaching to proximal margin. Carpus smaller, subrectangular, very narrow. Propod twice as long as wide, posterior margin (palm) half as long as anterior, with two teeth, the proximal one a small blunt projection; the anterior, near base of dactylos, produced, acute and strong; the palm between teeth excavate; dactylos curved and reaching as far as proximal tooth. Female: Distal tooth of palm poorly defined, with five or so setae; both teeth denticulate; palm between teeth very little excavated, almost straight, non-denticulate.

Peraeopods 1 and 2. Segments 3 and 4 which bear the gills have no peraeopods, and the peraeopods on segment 5 are regarded as Peraeopods 3.

Peraeopod 3. Ischium, anterior margin with strong flange produced backwardly; a small subrectangular or conical projection arising medially near distal margin and projecting slightly posterior to base of flange. Merus subrectangular, anterior margin similarly produced. Carpus width twice its length, posterodistal angle sharply produced, distal margin with posterior half excavated to acute anterodistal angle; proximal angle forming broadly rounded projection. Propod narrowing distally, anterior margin convex, twice length of straight posterior margin (palm). Dactylos long, stout, strongly curved.

Peraeopods 4 and 5. Slightly larger, otherwise similar.

Urosome. This bears ventrally, in the male, a pair of simple conical projections (Pl. 24, fig. 9) which Sars refers to as “external sexual appendages.” Posterior to these is a single appendage (fig. 10), cleft terminally for a third of its length to form two globular lobes. the outer lateral margins of lobes richly covered with bristles, the bristles thinning out along peduncle margin. Corresponding to the telson is a small subcircular plate, with a simple conical projection either side (fig. 11), these possibly corresponding to uropods.

Gills. Male: Unsegmented; single, present on segments 3 and 4, and as long as body less segment 1. Each with pair of accessory appendages. Female: A little shorter, comparatively, than in male. Juvenile: Shorter and stouter, more balloonlike.

These specimens were taken by Mr. W. H. Dawbin at Te Awhaiti, in July, 1949, and are in my personal collection, slides 71 to 75. The female described above contained between 70 and 80 juveniles in her brood-pouch, ranging in

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length from 1 to 3 millimetres. The colour of the animals in life is brownish-white, in spirit yellowish. The eyes are brown, small, round and apart.


Of the previous descriptions of the species published, only two (Lutken, 1873; Sars, 1895) are full descriptions, and Lutken's is in Danish. Barnard (1932) described P. erraticus in some detail, and stated that P. boopis differed only in the ventral tubercles. Since this is the first paper of a series towards a revision of the New Zealand Amphipodan fauna, a detailed description of the New Zealand specimens is given. The species itself is very constant in form.

There are five species in the genus, P. boopis, P. erraticus, P nodosus, P. Physeteris, and P. gracilis; and what Stephenson (1942) calls “a rather uncertain variety,” P. boopis var. physeteris. P. physeteris has the gills tufted, whereas in the other four species they are not tufted but elongate-cylindrical. P. nodosus may be distinguished in turn by the rugged dorsal surface. In P. erraticus, P. gracilis, and P. boopis the dorsal surface is smooth, and in P. gracilis the ventral surface lacks the characteristic pointed tubercles of P. erraticus and P. boopis. The female of P. erraticus has one pair of tubercles on each of segments 5 and 7, and two pairs on segment 6; and the male has one pair on each of segments 5, 6, and 7. On the other hand, P. boopis has one pair of tubercles on segments 5, 6 and 7 in the female. and one pair on only segments 6 and 7 in the male.

Otherwise P. erraticus and P. boopis are similar Although both species have been recorded elsewhere from Megaptera nodosa, I have not as yet seen P. erraticus collected from New Zealand.

Short references to the occurrence of P. boopis on whales are made in whaling papers by Lillie (1915) and Matthews (1937, pp. 47, 48).

The maxilliped palp in the young of the genus Paracyamus is easily distinguished under a high power microscope, and the constancy of the size-degeneration ratio makes it an eminently satisfactory characteristic for distinguishing the genus from Cyamus.

The growth stages of the palp (Text-figs.) in P. boopis closely parallel those figured by Chevreux (1913, fig. 62) for P. erraticus. Up to a size of about 3·5 mm. the palp is well developed; above this the maxilliped shows the degeneration characteristic of the genus. This agrees with Chevreux's findings for P. erraticus:

“… ce palpe était encore complètement développé chez les exemplaires de 3 millimètres de longueur. In commence à s'atrophier et ne possède plus que deux articles chez un spécimen long de 4 millimêtres, pour en arriver à être très court et uniarticulé chez les adultes.”

The following list of Cetaceans occurring in New Zealand waters (Oliver, 1922) and their Cyamid parasites as found elsewhere (cf. Barnard, 1932, and Stephensen, 1942) suggests that further examination of these animals will add a number of species of Cyamidae to the New Zealand fauna.

Balaena australis (Southern Right Whale) Cyamus ovalis
Paracyamus erraticus
Paracyamus gracilis
Balaenoptera musculus (Blue Whale) Cyamus balaenopterae
Balaenoptera physalus (Fin Whale) Cyamus balaenopterae
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Megaptera nodosa (Humpback) Paracyamus erraticus
Physeter macrocephalus (Sperm Whale) Paracyamus boopis
Paracyamus physeteris
P. boopis var. physeteris
Delphinus delphis (Dolphin) Isocyamus delphini
Grampus griseus (Risso's Dolphin) Isocyamus delphini
Globicephala maelena (Blackfish) Isocyamus delphini
Pseudorca crassidens (Tasmanian Blackfish) Isocyamus delphini

The genera Paracyamus and Isocyamus are alike in having the maxilliped palp in the adult rudimentary or absent. They may be distinguished in that the second antennae of Paracyamus sp. are 4-segmented, and in Isocyamus they are 3-segmented. In Cyamus the maxilliped palp in adult specimens is 5-segmented. Cyamus ovalis has two pairs of branchia to each segment and C. balaenopterae has one pair.


I wish to thank Mr. W. H. Dawbin for making available to me specimens and information obtained through the courtesy of Messrs. Perano Brothers, of the Te Awhaiti Whaling Station; the Canterbury University College Council and Professor Percival for making available the late Professor Chilton's specimens and literature; to Dr. Torben Wolff, of the Zoological Museum, Copenhagen, through whose courtesy I obtained papers by Dr. Stephensen; Mr. K. K. Tiwari, of the Zoological Survey of India, and Mr. R. R. Forster, of the Canterbury Museum, for information on Cyamus ovalis specimens; and to Professor L. R. Richardson for his helpful eriticism during the preparation of this paper.

Literature Cited

Barnard, K. H., 1932. Discovery Reports. Amphipoda. Vol. V, pp. 1–326, plate I and text-figs.

Cheverux, E., 1913. Amphipodes. Deuxième Exp. Ant. Franc., 1908–1910, pp. 79–186, text-figs.

Chilton, C., 1883. Additions to the Sessile-eyed Crustacea of New Zealand. Trans N.Z. Inst., vol. 16, pp. 249–265, pl. 17–21.

Lillie, D. G, 1915. Cetacea. Brit. Antarctic (‘Terra. Nova') Exped., 1910. Nat Hist Reports, Zool. 1, no. 3, pp. 85–124.

Lutken, Chr. F., 1873. Bidrag till Kundskab om Arterne af Slaegten Cyamus Latr. eller Hvallusene. Vid. Selsk. Skr. 5 R. Maturv. math. Afd., X, no. 3, pp. 231–284, and I–III, pls. 1–4.

Matthews, L. Harrison, 1937. The Humpback Whale; Megaptera nodosa. Discovery Reports, vol. 17, pp. 47–48

Oliver, W. R. B., 1922. A Review of the Cetacea of the New Zealand Seas. Proc. Zool. Soc. Lond., pp. 557–585, pl. 1–4

Sars, G. O., 1895. An Account of the Crustacca of Nomcay. 1. Amphipoda Text and plates.

Stephensen, K., 1942. The Amphipoda of North Norway and Spitsbergen with Adjacent Waters. Fasc. IV. Tromso Mus. Skr., vol. 3, part 4, pp. 365–526, 26 text-figs.

Thomson, G. M., and Chilton, C., 1886. Critical List of the Crustacea Malacostraca of New Zealand. Trans. N.Z. Inst., vol. 18, pp. 141–159.

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A Preliminary Study of the Land Operculate Murdochia pallidum (Cyclophoridae, Mesogastropoda)

[Read before the Wellington Branch, August 28, 1951; received by the Editor, August 28, 1951]


This paper gives a detailed description of the digestive and reproductive systems of the New Zealand land operculate Murdochia pallidum (Family Cyclophoridae), with a discussion on the morphological changes involved in the transition from the sea to a land habitat The Cyclophoridae have accomplished the transition from the sea to a land habitat with relatively few modifications of their primitive structure.

The Cyclophoridae represent one of several groups of terrestrial molluscs that have evolved from the marine gastropods belonging to the Sub-class Prosobranchiata. They are wholly distinct from the later-derived pulmonate stock, remaining much more generalised in the structure of the animal, and being conveniently referred to collectively as the “land operculates.” Three such families are represented in the New Zealand fauna, the Hydrocenidae, which originated among the Neritacea; the Cyclophoridae; and the Realiidae. The two last-named families are generally held to have arisen from littorinid-like marine ancestors; they are probably only distantly related, though by Thiele (1935) the two groups, together with a long list of other terrestrial mesogastropods are loosely associated in the Stirps Archaeotaenioglossa. Of the structure of the Cyclophoridae we have little detailed knowledge, although from Bouvier's picture (1882) of the nervous system of Cyclophorus tigrinus, with persistent ladder-like pedal ganglionic cords, it has long been inferred that they are highly primitive among mesogastropods The New Zealand members of the Cyclophoridae belong to the genus Murdochia, and are all of small size, seldom exceeding 5 mm. in length. The family has a wide distribution in S. E. Asia and the East Indies, Central America and the West Indies; some of the tropical species are of temptingly large size, and would be especially suitable for the detailed work on living gut and genitalia, which will be required before a confident assessment of the group is able to be made. But in view of the present lack of anatomical work, and the interest of the genus Murdochia among neozelanic land mollusca, it was felt that the following observations would prove of some value. It is hoped in a later paper to expand the present account, especially as regards reproductive activities, and also to provide a description of the New Zealand genera Hydrocena and Realia.

Material of Murdochia pallidum was dissected alive after removal of the shell, fixed in Bouin's and dissected preserved, also sectioned at 8μ and stained in Delafield's haematoxylin and Van Giesen's picrofuchsin. Mr. C. B. Trevarthen kindly collected several batches of living Murdochia, and his assistance is gratefully acknowledged. The writer is also indebted to Dr. W. D. Reid, of the Plant Diseases Division, D.S.I.R., Owairaka, Auckland, for carrying out a culture trial with the spirochaetes from the stomach.

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The described species of Murdochia are 13 in number (Powell, 1946) and seem to have a rather localised distribution in New Zealand rain-forests. The species most commonly encountered near Auckland is M. pallidum (see Suter, 1913); it is 5 mm. in length, 3·25 mm. in greatest diameter, turbinate, conical, horny brown in colour, with the operculum silvery-white (Text-fig. 1). The most typical habitat is in the deep, ensheathing leaf bases of Rhopalostylis, and also of Collospermum and Freycenetia. Pfeiffer's term “cryptozoic” admirably describes the mode of life of these animals, avoiding the light and seeking moisture. This fauna includes a wide range of small invertebrates, such as pulmonate snails (flammulinids), slugs (Athoracophorus), arachnids and myriapods, and insects—especially coleoptera and collembola. Most of these are vegetarians, taking into the gut fragments of food of varied detrital nature, including pieces of decaying leaves and wood pulp, and much unrecognisable plant material. In many cases, including Murdochia, a high proportion of the food is made up of fungal mycelia scraped off the moist substratum with the radula. M. pallidum seems to be more or less a specialist in its diet, and a similar preference for fungi is reported in the case of the tropical American cyclophorids (de la Torre, Bartsch, and Morrison, 1942).

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Text-Fig. 1—The operculum, uppen surface.

The animal when removed from the shell is found to be quite unspecialised in its structure. The foot is long, truncate, and squarish in front, rounded behind. The margin of the sole is surrounded by a wide strip of black-pigmented epithelium, thrown into small, regular, puckered folds; the central tract of the plantar surface is covered with short-celled smooth epithelium, deeply enfolded in the middle of the sole to give exit to the secretion of the pedal gland. Other sources of mucous secretion are present, in the mantle cavity—a broad hypo-

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branchial gland extending along the right side of the pallial roof, unaltered in histological structure from that of marine prosobranchs; and a narrow zone of sub-epithelial gland cells encircling the mantle just within its margin; these probably serve to seal the closed operculum, during prolonged withdrawal of the animal into the shell. The colour of the exposed parts of the head and foot is greyish-black, and the head, snout and short, paired tentacles are jet black. The pallial wall is divided into regions, the respiratory area occupying the left half, the hypobranchial gland to the right, and behind it the yellowish renal sac. The visceral mass is largely occupied by the digestive gland, composed of a mass of stout reddish-brown lobules, investing the stomach, which is seen from the surface as a translucent white sac. The gonad is quite large, forming a spacious undivided sac, applied to the convex surface of the digestive gland. It is white in the male, deep yellow in the female.

The Digestive System

The snout is deeply cleft at the tip, and the mouth is a vertical slit opening directly into the buccal bulb, which occupies most of the cephalic haemocoele. The anterior half of the bulb is equipped with a pair of chitinous jaws, triangular plates close together in the dorsal mid-line, diverging ventrally, and secreted by the buccal epithelium. Their anterior edges project sharply from the buccal wall, serving to grip firmly an object during abrasion by the protractor strokes of the radula. Each jaw is broken up by diagonal striations into a series of lozenge-like teeth, slightly overlapping and with their acute tips forwards. The odontophore is short and wide, supported by a single median cartilage formed of large squarish or spherical vacuolated cells. The cartilage is deeply excavated behind, forming a chair-like depression through which the radula passes. The radular caecum is long and slender, passing backwards along the ventral side of the oesophagus, and becoming involved in the torsion of the gut, so that its slightly expanded tip finally curves around to the right side. The median and two lateral teeth are of the taenioglossan rasping type, the former with three bluntly rounded cusps, the latter with a row of five. The two pairs of marginals are sharp, curved and prehensile. The salivary glands are minute, sending forward a pair of well-defined ducts embedded in the connective tissue of the folds bounding the dorsal oesophageal channel. The salivary cells are stained uniformly purple with haematoxylin, having no apparent enzyme contents and probably serving merely for secretion of mucus. The oesophagus preserves no trace of its lateral glandular pouches, proceeding straight to the stomach, as a uniformly narrow, ciliated tube. Mucus cells are everywhere abundant, and the ciliated cells are largest along the broad dorso-lateral folds, keeping up a strong aboral current.

The stomach in Murdochia is a rather long tubular sac curved around the first visceral whorl. The oesophagus opens into its rounded posterior end, and the intestine issues from the narrow extremity of the stomach in front. The structure of the sac, is of the simplest type, showing two histological regions. The longer posterior portion is lined with a thin transparent cuticle, save for a strip of short-celled ciliated epithelium which runs forward in a groove along the ventral wall. This groove originates at the posterior tip of the stomach from two or three folds of ciliated epithelium leading from the mouth of the oesophagus. The anterior third of the stomach is slightly constricted off from the cuticulate portion, and forms a small thimble-shaped sac, meeting the posterior chamber at an obtuse angle. Its lining epithelium is of the densely ciliated type, characteristic of the style sacs of mesogastropods and lamellibranchs, the bristle-

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like cilia reaching as much as two-thirds the height of the stout columnar cells. Along the ventral wall of this sac, the shorter ciliated longitudinal tract is continued from behind, interrupting the denser cilia, and passing imperceptibly into the intestinal epithelium in front. There is a single digestive diverticulum, opening just behind the ciliated sac, on the ventral wall of the stomach, close alongside the longitudinal ciliated tract. Its mouth is surrounded by ciliated cells whose currents appear to aid the entry of fine particles, and perhaps also the outward passage of egesta. At the angle of the stomach and along the margin of the ciliated groove, the cuticle becomes heavily thickened, forming a stouter ridge representing the gastric shield of style-bearing molluses.

In the living Murdochia, the stomach is filled with a viscid mucous rope, continuous behind with the less coherent food string entering from the oesophagus, and passing in front directly into the narrower string occupying the lumen of the intestine. The mucous contents of the stomach are thus divisible into two portions. The ciliated anterior chamber is filled with a short, opaque rod, whitish and quite free from food particles. This is inserted behind into a longer cord of mucus, permeated with dark-coloured ingested material, and extending through the rest of the stomach. It is constantly augmented by the inward passage of particles from the oesophagus. In addition to a good deal of unrecognisable detritus, material of plant origin such as leaf fragments, broken fern sporangia, and—in particular—numerous short segments of fungal mycelia, are almost always present. Murdochia must be to some extent non-selective, using the radula for scraping up the rich nutritive film overlying the damp surfaces within the leaf sheaths and similar locations. The general movement of stomach contents is apparent from figure 5.

The chief impetus to particles is conveyed by the colourless anterior portion of the mucous rod, which is rotated by the transverse beat of the “style sac” cilia. This has the effect of rotating the whole attached mass of food contents within the cuticulate chamber, while in addition, the shorter cilia of the ventral tract evidently impart sufficient forward motion to carry the contents slowly towards the intestine. This movement must be very gradual, permitting of sufficient time for such preliminary digestion as may occur while the mucous rod is rotated within the stomach. As suggested by Graham (1939) in the case of the land operculate Pomatias elegans, the long posterior chamber of the stomach evidently serves as a storage region, and—though feeding seems to be almost continuous in Murdochia—may help to even out irregularities in the flow of food through the stomach.

It is evident that the functioning of the stomach in Murdochia typifies what must have been the simplest and earliest condition of the style sac form of molluscan stomach. The mass of whitish mucus within the style sac—if we may call it such—is an example of what the writer has called elsewhere (1951b) a “protostyle.” Its chief role appears to be to transmit rotatory movement, imparted by the style sac cilia, backwards to the rest of the stomach contents—a function which is of obvious importance in the activity of the stomach. By this means, ingested food is drawn through the stomach at a slow, uniform speed by the action of the very robust cilia, concentrated in the style sac. Stomach contents are thoroughly admixed with such digestive enzymes as may be present. The smallest, finely divided particles are repeatedly rotated alongside the opening of the digestive diverticulum, into which they are received, in part by ciliated action, in part probably by compression movements of the sparsely muscular

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Fig. 1—
Fig. 2—
Fig. 3—
Fig. 4—

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Fig. 5—The stomach in longitudinal Section, somewhat diagrammatic showing the disposition of contents and the arrangement of cilliary currents.
Fig. 6—Transverse section of stomach, through the region of the gastric shield, just behind the digestive divertrculum.

[ unclear: ]

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stomach wall. In addition, the larger, coarse particles near the periphery of the food string are swept across the longitudinal groove (IGR) and caught by its cilia to be rapidly carried forward into the intestine. Though the clear mucus of the protostyle must also be slowly carried into the intestine, where it forms the matrix of the faecal string, yet the strongest forward flow is along the ventral ciliated groove. This forms a channel whereby particles too large to enter the digestive diverticulum are able to by-pass the protostyle, and find their way directly into the intestine. The longitudinal groove in Murdochia is the sole representative of the ciliary sorting area in this snail. Pomatias, similarly, shows an almost total reduction of the sorting plicae typical of the stomachs of marine prosobranchs. The mode of feeding of marine microherbivorous gastropods, generally on a continuous stream of poorly sorted bottom deposits, is a habit calling forth a high development of sorting and grading mechanisms within the stomach. In terrestrial prosobranchs, such as Murdochia or Pomatias, the problem is not so much one of sorting, as of breaking up larger fragments of plant material and rendering them finally suitable for absorption by the digestive gland epithelium.

According to Graham (1939), the opaque mucous contents of the style sac in Pomatias constitute a true crystalline style: this is gradually thrust into the stomach towards the gastric shield, by a backwardly directed ciliary current within the style sac. No such current could be detected within the much smaller stomach of Murdochia; from the whole appearance of the mucous rod, including its direct merging into the faecal string, we may conclude it is not properly speaking a crystalline style. Amylase tests are not mentioned in Graham's account of Pomatias. In Murdochia the spotting tests for amylase were difficult to carry out satisfactorily because of minute size, but gave negative results. No doubt the chief function of the protostyle in Murdochia is its mechanical role of rotating the stomach contents.

It is unlikely that any digestion takes place in Murdochia in the part of the gut anterior to the stomach. Particles of food, including fragments of fungal hyphae, are too large to enter the digestive diverticulum intact, and recgnisable hyphal or other food particles were never detected either within the lumina of the digestive tubules, or within the cytoplasm of the digestive cells. The structure of the digestive gland is in no way dissimilar from that of other phytophagous prosobranchs, or lamellibranchs already investigated; ingestion of fine particles evidently occurs, followed by intracellular digestion. The question arises, how—and to what extent—are large particles, such as fragments of hyphae, first broken up within the stomach? There is little evidence that mechanical trituration occurs. The muscle layer of the stomach wall is not conspicuous, and the cuticle is for the most part not robust. It is possible that particles are pressed, by the rotation of the prostyle, against the stout ridge of cuticle (GSH) and thus become slowly comminuted, perhaps assisted by a squeezing action of the stomach wall as suggested by Yonge in the protobranchiate bivalves (1939).

Further, in the cyclophorids, such as Murdochia, which feed to a large extent on fungal mycelia (see also de la Torre et al., 1942), a high proportion of cellulose-like substances must be included in the diet. If this carbohydrate is to be successfully utilised, a digestive problem is presented that has in only a few cases been successfully surmounted by molluses. A possible clue to the means of cellulose break-down is obtained by microscopic examination of the opaque mucoid mass of the style sac. The colloidal matrix is here invariably packed with long, highly

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motile spirochaetes, progressing through the medium in constant vibratile movement, and especially densely concentrated at the boundary between clear mucus and food material. It may be suspected that these organisms, occurring in pure state with such regularity, are not without some role in the digestive process of the molluse.

Large spirochaetes assigned to the genus Cristispira have been briefly recorded as living “commensally” in the stomachs and crystalline styles of a fairly long list of lamellibranchs. Yonge (1926) refers to the presence of spirochaetes in the style substance of Ostraea edulis, while Breed, Murray and Hitchens (1948) list occurrences in no less than 13 families of bivalves. There would appear to be no previous record of spirochaetes from a gastropod style sac; nor has any explanation been hazarded as to the role they may play. Results of great interest may be expected from further inquiry into the spirochaetes of style sacs, especially from a survey of their occurrence among molluscs, and their powers of breaking down complex carbohydrates. An analogy might be suggested with Spirochaeta cytophaga, a free-living cellulose-digesting organism, whose activity has been investigated by Walker and Warren (1938). As regards the snail, Murdochia, some amount of simpler carbohydrate is probably available witin the lumen of the gut for direct absorption, after external bacterial or autolytic breakdown of cellulose and other plant substances; but the problem of the fungal hyphae—so conspicuous in stomach contents but apparently quite absent in the faecal pellets—remains to be solved.

The digestive diverticula in Murdochia contain both digestive and excretory cells. The former are wedge-shaped or clavate, filled with granular cytoplasm, consisting of colourless refractile particles easily liberated by cell maceration. Ingested material appears to be taken up in an extremely fine state of division, and the digestive cells contain little or no coloured egesta for return to the stomach. Enzyme droplets are not detectable.

The excretory cells are broad-based and pyramidal, brown-staining with Van Giesen's. They apparently extract chlorophyllous or other absorbed pigments from the blood, passing their contents to the lumen in the form of large droplets, sometimes equipped with little tails, or drumstick-shaped, as they emerge from the cell. The intestine is of simple structure throughout, describing a short S-shaped course after leaving the stomach, and crossing to the right side around the edge of the renal sac. The rectum is slightly wider (70μ in diameter).

Each faecal pellet occupies the whole width of the intestine, and is stoutly ovoid in shape, being nipped off from the mucous string by repeated peristaltic constrictions of the intestinal wall, which is provided with a narrow coat of circular muscle, several fibres thick. Muscular action plays the most important part in faeces formation, both in Murdochia and evidently in mesogastropoda in general. While it is broadly true—as stated by Yonge (1936 and elsewhere)—that the gut in microphagous prosobranchs performs little muscular action, yet the small but definite peristaltic movements of the intestine have generally been overlooked. As well as in Murdochia, peristalsis has been observed by the present writer in Struthiolaria (1951), Suterilla (unpublished) and in Serpulorbis (1951a). Murdochia is reminiscent of marine mesogastropods in the care taken for the consolidation of faecal pellets, as a precaution against the fouling of the pallial cavity in gill-breathing animals. Mucus is liberally secreted by fusiform gland cells, interspersed between the narrow ciliated cells of the intestinal epithelium.

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Text-fig. 2—The head, snout and penis. INTR, terminal invaginable intromittent portion of penis. PEN, basal portion of penis; SNT, snout; TEN, cephalie tentacle.

Reproductive System

The structure of the genitalia in Murdochia pallidum may be outlined briefly. In the male, the vas deferens is a narrow tube closed throughout its whole length, as far as the tip of the prominent cephalic penis. It leaves the thin-walled testis sac as a lightly coiled, non-glandular tube, which emerges ventrally to the renal organ, to run the whole length of the pallial cavity, just below the rectum, along the attachment of the mantle to the trunk musculature. The closure of the pallial portion of the vas deferens—primitively an open ciliated groove—is to be regarded as correlated with the terrestrial habit. The wall of the pallial vas deferens is thickened and glandular, as far forward as the base of the penis, its lumen narrow, rounded or slightly flattened. This portion of the male duct evidently performs the role of a “prostate” gland: its lining cells are stoutly columnar, filled with minute, non-staining secretion granules, and having rounded, basal nuclei; Regularly alternating with the glandular cells are narrow, darkly compressed, ciliated cells, about half the length of the secreting cells, and with dark-staining elongate nuclei. Their distal surfaces are narrow, and cilia are borne also by the glandular cells, the whole of the lumen being uniformly lined. The penis is attached to the head immediately behind the right tentacle, forming a stout muscular column, flattened or cylindrical, with strong annular rugae, and yellowish-white in colour. It is generally reflected back within the pallial cavity. lying transversely across the trunk, or along the right side. It is perforated throughout by a narrow ejaculatory duct, continuous with the vas deferens, proceeding to the distal tip of the penis, which becomes narrow and cylindrical, and forms the intromittent part of the male organ. This structure is freely extensible, and can be withdrawn into the haemocoelic blood space enclosed by the stouter basal portion of the penis. (Fig. 3.) It can be strongly protruded by the pressure of the muscular wall of the basal part of the penis on the blood contained in the haemocoelic space. The terminal intromittent portion of the penis is supported by a flexible rod of elastic connective tissue, which runs alongside, and finally extends completely around the ejaculatory duct. It is made up of long, vacuolated cells, radially inserted at the base of the

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epithelium of the ejaculatory duct. The basal part of the penis has a stout layer of muscle fibres, circular and longitudinal elements intermingled, and its external surface is traversed along one side by a broad tract of glandular epithelium, mucus being produced by rounded, subepithelial secretory cells, sending up narrow ducts between the cells of the columnar epithelium.

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Text-fig. 3—Transverse section of the penis, with the terminal intromittent portion invaginated.
CART, cartilage-like supporting rod; EP, external epithelium of the penis; EJ.D, ejaculatory duct; G.EP, tract of glandular epithelium, GLD, subepithehal gland cells; HMC, haemocoele occupied by sparse connective tissue, INV, Invagination

The female genital duct is divisible into two portions—an extremely narrow, much convoluted posterior part, leading forward from the ovary to open into the stouter, glandular-walled pallial oviduct. The functions of the pallial oviduct have not yet been properly ascertained. It appears to combine the secretory role of both the albumen gland and the capsule gland as recognised by Fretter (1946) for marine prosobranchs. But the two regions are not structurally separable in Murdochia.

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On leaving the ovary, the posterior portion of the female duct is at first very narrow, thin-walled and translucent, the cells lightly staining, and with long cilia; the duct is here strongly distensible with the passage of eggs. A little further forward the proximal oviduct becomes thrown into a number of coils and closely convoluted, opaque white in appearance. It is here noticeably muscular, its coat of circular fibres being well developed and the epithelial lining being thrown into 4–5 longitudinal folds by cell height differences. The epithelium contains plump, ovoid gland cells, with basal nuclei and mucous contents staining lightly in haematoxylin. Ciliated cells are also present, especially at the crests of the folds, centrally constricted and with rod-like central nuclei. The posterior-most part of the coiled oviduct is of different structure from the rest, and functions as a receptaculum seminis. The epithelium here is not folded, and the cells are of one type only, regularly columnar, clear or lightly staining, with a single row of basal nuclei. A coating of close-pressed sperm heads is found attached to the distal surfaces of the cells, the tails occupying the middle of the lumen. It is presumably in this part of the duct that fertilisation takes place before the addition of glandular secretions to the egg.

The pallial oviduct runs forward immediately below the rectum. It is pale yellow in colour, thick-walled and laterally compressed, thrown into a succession of deep transverse folds which project into the lumen as ridges of the glandular wall. The epithelium is extremely regular, the gland cells tall and narrowly columnar, with basal nuclei, and with small triangular or wedge-shaped ciliated cells inserted between their tips. The character of the secretion changes about halfway along the duct. Posteriorly the cells are filled with rows of large, finely granular vesicles or spherules, each occupying the whole width of a cell, and staining light pinkish-brown with Van Gieson's. Further forward, the secretion is in little droplets, much smaller, and staining deeper purplish, with a stronger affinity for haematoxylm. The aperture of the pallial oviduct lies far forward, just below the anus, and near the opening the histology changes. The wall becomes somewhat folded by differences in the length of the cells. The ciliated cells are longer with their cilia more prominent, and the contents of the gland cells—in the basal two-thirds—are tiny refractile spherules, staining deeply black with haematoxylin.


The Cyclophoridae have accomplished the transition from the sea to a land habitat with relatively few modifications of their primitive structure. They have occupied a peculiarly favourable ecological niche, and like all terrestrial molluscs—save a few xerophilic pulmonates—they are confined to conditions of high humidity. The retention of the operculum closing the shell aperture prevents water loss. The air-filled pallial cavity remains widely open anteriorly and there is, properly speaking, no development of a lung, respiration taking place merely through the smooth vascularised epithelium of the pallial roof; gill filaments are lost and there are no folded respiratory lamellae. The lumen of the renal organ is small and restricted; its opening into the pallial cavity was not traced in sections, and there is no forward prolongation running alongside the rectum to form a “ureter” leading to the pallial margin, as has become the case in pulmonates. The lining cells of the renal sac are filled with yellowish granules—it would be of interest to know whether the animal has become uricotelic, and whether there is a periodic discharge of renal contents into the mantle cavity, or whether, as seems more probable, the renal organ functions as a kidney of

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accumulation. The mode of reproduction has not yet been fully worked out, but the animals are bisexual and the genital ducts relatively simple, as in the lower prosobranchs in general. The landward migration of marine gastropods has become possible only in those that have evolved about the rhipidoglossan level, and in which the gametes are no longer discharged by way of the right kidney. Fertilisation now becomes internal, and a closed pallial genital duct has been developed. In the males the sperms are conducted forwards by a closed channel to a well-developed cephalic penis; in the female, a closed glandular oviduct, derived from secretory tissue of the pallial wall, runs forward to open alongside the rectum.

In Murdochia, the alimentary canal combines a primitive structure with certain specialised features. The absence of oesophageal pouches or other sources of enzyme secretion within the oesophagus is evidently an advanced feature, paralleling the condition described in the specialised Turritella (Graham, 1938). The salivary glands are likewise devoid of enzyme cells, and Murdochia would appear to have been singularly improvident in abandoning all its sources of extra-cellular digestive enzymes, probably before the landward migration of the family. Whether or not the spirochaetes in the protostyle have taken on an accessory role in digestion remains to be determined. The small size of the animal—as well as its primitiveness—may be a contributing factor to the simplicity of structure of the stomach. The loss of the ciliary sorting area is probably correlated with the terrestrial habit and the changed nature of the food. The primary function of the stomach is no longer sorting, but the breaking down, whether by digestion or mechanically, of particles of plant material still too large for entry into the digestive diverticulum. The extreme posterior position of the opening of the oesophagus into the stomach is to be regarded—following Graham (1949)—as a highly primitive feature. In almost all other mesogastropods it has migrated some distance forwards towards the intestinal aperture. The long, backwardly directed sac in the stomach of Pomatias (Graham, 1939), Littorina (Graham, 1949), Melarhaphe and Risellopsis (present writer, unpublished), would appear to be directly homologous with the cuticulate chamber of the stomach in Murdochia, after the forward migration of the oesophagus, to open shortly behind the style sac—intestinal chamber. In Murdochia, the formation of firm, mucusbound faecal pellets in the intestine is a relic of ancestry among marine prosobranchs, where the compacting of the faeces serves as a device for prevention of fouling of the respiratory water of the pallial cavity. In a terrestrial prosobranch it is probably equally necessary in the absence of a cleansing current passing continuously through the pallial cavity.

References to Literature

1. Bouvier, E. L., 1887. Systeme nerveux, morphologie generale et classification des Gasteropodes prosobranches. Ann. des Sci. Nat. (Zool)., 7, iii.

2. Breed, R. S., Murray, E. G. D., and Hitchens, A. P., 1948. Manual of Determinative Bacteriology, Bergey, 6th Edn., London.

3. de la Torre, C., Bartsch, P, and Morrison, J. P. E., 1942. The Cyclophorid Operculate Land Molluscs of America. U.S. Nat. Mus. Bull, Smithsonian Inst., Washington, D.C.

4. Fretter, Vera, 1946. The Genital Ducts of Theodoxus, Lamellaria and Trivia, and a Discussion on their Evolution in the Prosobranchs. J. Mar. Biol. Ass. U.K., 26, 312–351.

5. Graham, A., 1938. On a Ciliary Process of Food-collecting in the Gastropod, Turritella communis Risso. Proc. Zool. Soc. London A, 108, 453.

6. — 1939. On the Structure of the Alimentary Canal in the Style-bearing Prosobranchs. Ibid, B, 109, 75.

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7. Graham, A., 1949. The Molluscan Stomach. Trans. Roy Soc. Edin., 61, 737.

8. Morton, J. E., 1951. The Ecology and Digestive System of the Struthiolariidae (Gastropoda). Quart. Journ. Micr. Sci., 92, (1), 1–25.

9. — 1951a. The Structure and Adaptations of the New Zealand Vermetidae. Pt. 1. Genus Serpulorbis. Trans. Roy. Soc. N. Z.,

10. — 1951b. The Role of the Crystalline Style. Proc. Malacol. Soc. Lond (in the press).

11. Powell, A. W. B., 1946. New Zealand Shellfish, Auckland, 2nd Edn.

12 Suter, H., 1913. Manual of the New Zealand Mollusca. Govt. Printer, Wellington. (With Atlas of Plates, 1915.)

13. Thiele, J., 1931. Handbuch der systematischen Weichtierkunde, I. Jena, Fischer.

14. Walker, E, and Warren, F. L., 1938 Decomposition of Cellulose by Cytophaga. Biochem. J., 32, 1, 31.

15. Yonge, C. M., 1927. Structure and Physiology of the Organs of Feeding and Digestion in Ostraea edulis. J. Mar. Biol. Ass. U.K., 14, 295–386.

16. — 1936. On Some Aspects of Digestion in Crliary Feeding Animals. Ibid., 20, 341–346.

17. — 1939. The Protobranchiate Mollusca: a Functional Interpretation of their Structure and Evolution. Phil. Trans. Roy. Soc., Lond., B, 230, 79.

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Cladode Anatomy and Leaf Trace Systems in New Zealand Brooms

Department of Botany, University of Otago, New Zealand.

[Read before the Otago Branch, June 12, 1951; received by the Editor, June 25, 1951]


The anatomy of the stem of the New Zealand Leguminous genera Carmichaelia, Corallospartium, Notospartium and Chordospartium is described and figured. Special structures dealt with are the cortical parenchyma, secondary growth, cork and leaf trace systems. A discussion on the relationships of the genera and species follows. It is shown that the primitive condition as seen in Carmichaelia kirkii is a round, non-flattened stem with leaf traces trilacunate. All other species possess flattened stems. This flattening takes place at two points in the stem: (a) in the region of the stem between the lateral traces of successive nodes (in all flattened species), (b) in the region of the stem between the median traces and the lateral traces of the same node (in Carmichaeliella). The primitive trilacunate leaf traces derive extra traces either interpolating or flanking, or both. Furrow development is associated with the fibre development of the leaf trace bundles. The interrelation of the four genera is shown on a chart (Fig. 10). From the subgenus Kirkiella arose an ancestral stock in which extra traces were added on the flanks. From this stock, within the genus Carmichaelia, came forms in which further flattening was accompanied by the interpolation of traces. The subgenus Huttonella and Corallospartium have in common flanking trace systems and indehiscent pods. In Notospartium the stem is rounded, although its pith is distinctly flattened. The anatomical evidence strongly favours the view that Chordospartium is derived from the hybridization of Notospartium and Corallospartium. In the adult stems of Corallospartium and Chordospartium a varying number of ridges is present.


The Leguminosae is poorly represented in New Zealand and of the eight genera, the brooms constitute four. Of these, Notospartium Hook. f., Corallospartium L. B. Armstrong, Chordospartium Cheeseman are endemic, and the fourth, Carmichaelia R. Brown, has but one out-lying species, Carmichaelia exsul, found in Lord Howe Island (Cheeseman, 1925; Cockayne, 1928).

This paper deals with the stem anatomy and morphology of these brooms, taking special notice of:


The development of flattening in the stems in all four genera.


The development of shallow grooving in many species of Carmichaelia and the deeper furrowing in Chordospartium and Corallospartium.


Changes in stem anatomy and particularly in the leaf trace systems that accompany the flattening.

Some investigators (Dormer, 1945, 1946; Sinnott, 1914) have shown that certain features of shoot structure have phylogenetic significance, and on this basis an attempt is made to interrelate the New Zealand genera. Since many of the Carmichaelia species are difficult to identify, and also have localised distribution, the writer is much indebted to Mr. G. Simpson, who, having recently revised the genus (1945), aided in supplying her with authentic material.

The writer wishes to express her thanks to Dr. G. T. S. Baylis for helpful advice and criticism.

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General Description

A. Carmichaelia R. Br.

This genus is the largest. Cheeseman (1925) divided it into the sections Eucarmichaelia (18 species) and Huttonella (4 species). Previously Kirk (1899) gave the Huttonella group generic rank. Simpson (1945) has now divided it into 8 subgenera, and while discarding some species listed by Cheeseman, he recognises 40 species* and 11 varieties. The subgenera are separated according to pod characters. Preliminary examination was made of all Carmichaelia species except five as indicated below. For further anatomical study at least one species from each subgenus was selected, and in Carmichaeliella four were chosen to show a sequence in the flattening of the stems within that subgenus. The selected species were:


Carmichaelia R. Br.


Thomsonella Simpson—4 species.

Ca. grandiflora Hook. f. (Pl. 28 E).


Carmichaeliella Simpson—17 species (16 investigated).

Ca. solandri Simpson.

Ca. cunninghamii Raoul.

Ca. aligera Simpson.

Ca. williamsii Kirk (Pl. 27 A).


Kirkiella Simpson—1 species.

Ca. kirkii Hook. f.


Enysiella Simpson—2 species.

Ca. enysii Kirk (Pl 27 D).


Petriea Simpson—3 species (2 investigated).

Ca. petriei Kirk (Pl. 28 F).


Monroella Simpson—3 species.

Ca. hollowayi Simpson.


Suterella Simpson—2 species.

Ca. uniflora Kirk (Pl. 27 C).


Huttonella Kirk—8 species (5 investigated)

Ca. curta Petrie.


Notospartium Hook. f.—3 species.

No. carmichaeliae Hook. f.


Chordospartium Cheeseman—1 species.

Ch. stevensoni Cheeseman.


Corallospartium J. B. Armstrong—1 species.

Co. crassicaule J. B. Armstrong (Pl. 27 B).

Carmichaelia shows wide variety in the external form of the stem. Even in the mat-forming subgenera the stems vary considerably in texture and size. Monroella is the stoutest, and the branches may have one side convex and the other concave or almost flat. Enysiella is distinguished by its “cushion” form on a stocky main stem (Pl. 27 D), while Suterella with its more slender creeping underground stems, forms mats (Pl. 27 C). Of the erect shrubby subgenera, Kirkiella alone can be singled out as easily distinguished, this being the only species showing no trace of flattening in either young or old stems (Pl. 30 A. B). The rest of the subgenera have flattened young stems, even if at one year they have become rounded (Pl 30 C, D), but in several such as C. williamsii (Pl. 27 A)

[Footnote] * Simpson lists 41 species, but Ca. corymbosa not specific rank.

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the flattening always persists, except in the main stems of plants several years old. These extremely flattened forms are all members of Carmichaeliella, and a series of forms exists linking this condition with the terete Kirkiella. No Carmichaelia species have deeply furrowed stems, but many show grooving which internal examination proves to be associated with fibrous bands that run longitudinally in the cortex.

The stems of Notospartium are smooth and not dissimilar from those of the slender-stemmed species of Carmichaelia, but its longer branches give the plant a semi-weeping habit. Of the two genera with furrowed stems Chordospartium has the weeping habit of Notospartium, while Corallospartium (Pl. 27 B) is strongly contrasted in habit, having thick, slightly flattened stems.

Phyllotaxy of all New Zealand brooms is one half. Seedlings of all genera bear foliate leaves with 1–7 leaflets. Adult plants usually produce scale leaves (Text-fig. 1 A), but they may produce small foliate ones (3–7 leaflets) at the height of each growing season. This recurrent foliate is a pronounced feature of some Carmichaelias (e.g., C. grandiflora—Pl. 28 E), but is insignificant in Chordospartium and Corallospartium, where such leaves are few and unifoliate, and absent altogether in Notospartium. Apart from this normal retention of a leafy habit in those adults usually producing only scale leaves, there may be a development of reversion shoots which are entirely leafy (Text-fig. 1 B), but these also have not been seen in Notospartium.

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Text-Fig. 1—Ca. williamsii. A. Normal spring shoot (× 1). B. Shoot with reversion foliage. (× 1). C. Leaves 1–6 from shoot B. Leaves 1–2. × 2 Leaves 3–6, × 4.

When a laminate leaf is shed, its stipules persist as a scale, and from plants which are changing from production of laminate to production of scale leaves, it is possible to obtain a series of intermediate forms which show that the scale leaf itself is composed of more or less coalesced stipules, the rest of the leaf having failed to develop (Text-fig. 1 C, 1–6).

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Text-fig. 2—A. Epidermal strip of Ca. grandiflora stem (× 30). B. Epidermal strip of Ca. hollowayi stem (× 30). C. Epidermal strip of Ca. williamsii stem (× 30). D. T.S. stems Ca. grandiflora (× 15). E. T.S. half stem Ca. hollowayi (× 30). F. I-IV, T.S.'s half stems Ca. williamsii from nodal to internodal regions (× 15) M, Mx, median traces; L, LX, lateral traces; F, Fx, flanking traces; P, Px, interpolated traces; A.B., axillary bud; ex.f., extra cortical fibre.

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General Stem Anatomy


In all genera a thick cuticle is present. In the furrowed ones this is accentuated on the ridges (Text-fig. 3, Pl. 29 C). In the furrows, are heavily cutinised unicellular hairs characteristic of the Leguminosae (Solereder, 1908). Stomata are arranged in longitudinal areas even in the smooth or slightly grooved stems, e.g. Ca. grandiflora, Notospartium. The epidermal strips devoid of stomata are underlaid by bands of cortical fibre (Text-fig. 2 A and D). In a few species, such as those of Monroella subgenus and Ca. williamsii, the distribution of this fibre is more diffuse (Text-fig. 2 E and F) and the stomatal areas less well defined (Text-fig. 2 B and C). Successive developmental stages of Corallospartium stems show that the stomatal areas (Text-fig. 2 A) are the forerunners of stomatal furrows. As the initially smooth seedling develops, fibre inceases to form ridges and the stomatal areas are sunken to form the furrows (Pl. 29 C). Chordospartium (Pl. 29 B) resembles Corallospartium. Stomata in all species are orientated transversely.

Cortical Parenchyma

All cortical parenchyma is photosynthetic except the layer adjacent to the fibres, which forms colourless sheaths (Text-fig. 3). In seedlings and in summer leafy species such as Ca. kirkii and Ca. grandiflora (Pl. 28 A), the cortex is composed of chains or small plates of cells with large air spaces between. This condition along with the leafiness is an indication of a more mesophytic nature. Progressive reduction of the air spaces is shown by Notospartium (Pl. 28 B) and Corallospartium (Pl. 28 C). The majority of Carmichaelias are similar to Notospartium, while Chordospartium again resembles Corallospartium. The very narrow flattened stems of Ca. williamsii depart from the usual Carmichaelia condition in having both pith and cortex greatly reduced (Pl. 29 D).


Normally this is a colourless layer that delimits the cortex from the stele. Adjacent to the fibre caps of the bundles it may, however, be obliterated as the fibre cap unites with the adjacent patch of cortical fibre. The interfasicular sections of the endodermis then link with the parenchymatous fibre sheath and this with the hypodermis, so that a continuous line of colourless cells is seen surrounding the photosynthetic cortex (Text-fig. 3).


In all genera the pith of the first internode of the seeling is cylindrical, but only in Ca. kirkii is this form retained throughout life (Pl. 30 A and B). In all other species the pith flattens in the course of development (Pl. 30 C) and in the extremely flattened Carmichaelias (Text-fig. 2F, iv) it is almost obliterated. The pith of internodes normally forms ridges where leaf trace bundles have withdrawn, but this does not occur in very flat stems nor in Notospartium, since in both these, bundles do not move out of alignment with the rest before reaching the node.

Vascular Bundles and Fibre Strands

In stems of adult plants vascular bundles are arranged in a ring in Ca. kirkii (Text-fig. 4 A), in roughly an oval form in Notospartium (Pl. 29 A), Chordospartium (Pl. 29 B), Corallospartium (Pl. 29 C), and the majority of Carmichaelias (Pl. 30 C), and in a zig-zag or double line in the extremely flattened Carmichaelias (Text-fig. 2 F, IV, and Text-fig. 4 C). Except in Notospartium

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Text-fig. 3—T.S. Corallospartium stem (× 308). Portion of a ridge and intervening furrow with one of the trace bundles and its associated fibre. The fibre is sheathed by a clear parenchymatous sheath continuous with the hypodermis on the outside and the endodermis inside. cut, cuticle; ctx, cortex; ep ht, epidermal hair, f, fibre, hy, hypoderma; p.sh, parenchymatous sheath, st, stoma.

the “ring” and “oval” types have from six to seventeen of the bundles situated a little out of alignment. These are the leaf trace bundles (Text-fig. 4 A and Text-fig. 2 D and E). In Corallospartium and Chordospartium they occupy the cortical ridges (Pl. 29 C).

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Fibrous tissue occurs mainly in association with the vascular bundles, and in all species [except Monroella and the extremely flattened Carmichaeliella (Text-fig. 2 E and F)] the leaf trace bundles are usually accompanied by more fibre than the stem bundles. In Ca. grandiflora every bundle is accompanied by two distinct fibre strands, the outer of cortical fibre and the inner the fibre cap of the bundle (Text-fig. 2 D). In other species and genera cortical and bundle fibre commonly unite, crushing a portion of the colourless endodermis and parenchymatous sheaths to form a continuous fibrous band extending from the phloem to the epidermis or to the hypodermis, depending upon whether or not the latter also is crushed as the fibre develops. In Notospartium actual union of the fibre masses is rare, but they lie close together and the intervening tissue is entirely colourless. In Ca. kirkii only the fibre groups accompanying leaf-trace bundles coalesce. When Ca. kirkii is compared with other Carmichaelias it is apparent that flattening involves an increase in the circumference of the stem and its stele. In flattened species the stem bundles divide and some of these (as well as the leaf-trace bundles) develop fibrous bands traversing the cortex. In the very flattened species (Text-fig. 2 F, iv), fibre of almost every bundle extends across the cortex. A converse development is seen in Chordospartium and Corallospartium. In the furrows of their stems the entire cortex is photosynthetic, the only fibre being in the bundle-caps. Each ridge is largely formed by the fibre-cap of a leaf-trace bundle united with a mass of cortical fibre.

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Text-fig. 4—A. Ca. kirkii. T.S. stem (× 30). B. Ca. enysii, T.S. stem (× 30). Ca. corrugata. T.S stem (× 30). All sections are to show the distribution of fibre in the cortex and the leaf-trace bundles. Mx, My, median traces; Lx, Ly, lateral traces, P, inter polated traces; ex.f, extra cortical fibre.

In Enysiella and Suterella (Text-fig. 4 B and C) there are a few extra cortical fibre strands not associated with vascular bundles. Comparable strands occur also in C. williamsii. They are not present at the nodes, appearing only as the bundles diverge in the broadening internodes. External examination shows them as fine colourless lines. The most prominent run the whole length of the internode.

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those less well marked run for successively smaller distances (Text-fig. 2 F, i-iv). In Monroella there is much scattered fibre arranged without definite relation to the bundles (Text-fig. 2 E). In old stems of all species (Pl. 29 D-F) stone cells may occur in a similar irregular manner.

Secondary Growth in the Older Stems and Formation of Cork

In Ca. kirkii the cambial activity results in the stem increasing more or less evenly in girth (Pl. 30 A, B), but in some species, e.g. Ca. petriei, the old stems are round, while young stems are distinctly flattened (Pl. 30 C, D). This rounding off is due to greater cambial activity on the two flattened sides of the stem. The flattened pith remains as an indication that flattening originally occurred in the stem (cf. C and D, Pl 30) Most of the woody shrub Carmichaelias resemble Ca. petriei, except the excessively flattened ones (Ca williamsii). In these the cambium never adds substantially to the bundles on the edges of the cladode so that these regions persist as flanges after the median portion of the stem has rounded into a cylinder (Text-fig. 5 A-D).

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Text-fig. 5—A-D: A series of stems of Ca. williamsii of varying ages to show successive stages in the cambial activity and cork formation. Persistent flanges are present in C and D (× 8). ck, cork; cm, cambium; ep, epidermis; fl, flange

Cork formation is initiated only where there is mechanical rupturing of the epidermis and the outer soft cortical tissues, by stelar growth (Pl. 29 D). Isolated patches of wound cork form (Text-fig. 5 C), eventually uniting to form a continuous layer of cork (Text-fig 5 D). There is never any cork formation in persisting flanges since there is no stelar thickening in these to cause an initiating rupture; nor is there ever any supercession of the original cork-forming layers by a deeper cork cambium unless the original layers are somehow damaged. In Corallospartium and Chordospartium the weakest areas of the cortex are in the furrows Splitting and cork formation first occurs here (Pl. 29 E), but the.

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

(A) Carmichaelia williamsii. × 2/9 approx.
(B) Corallospartium crassicaule. × 2/5 approx.
(C) Carmichaelia uniflora. × 3/7 approx.
(D) Carmichaelia enysii. × ¾ approx.

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(A) Ca. grandiflora. (B) No. carmichaeliae. (C) Corallospartium. (D) Ca. williamsii.
A—D (× 50). Longitudinal sections through cortex of these species to show the gradual reduction in the air spaces in the first three, and the decrease in width of cortex in the last.
(E) Ca. grandiflora. × ⅕. (F) C. petriei. × ⅕.

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(A) Notospartium. (B) Chordospartium. (C) Corallospartium. A—C (× 16). Photomicrographs of young stems of approximately the same age. Notospartium and Chordospartium exhibit similar stem dimensions, but furrow development occurs in Chordospartium and Corallospartium. (D) Notospartium. (E) Corallospartium. (F) Chordospartium D—F (× 50). T. S. of older stems of Notospartium and Chordospartium to show the masses of stone cells in the cortex. The initial cork formation is shown in the non-furrowed Notospartium, as well as two stages of cork cambial activity in the furrowed species.
f and cf, cortical fibre; hr, stomatal hairs, ck, cork; cut, cuticle, st.e, stone cells.

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(A) Carmichaelia kuku (× 48). Young stem with round open cortex traversed by fibre from six leat-trace bundles Stem round
(B) Carmichaelia kuku (× 18) T. S. Old stem Note persisting round pith
(C) Carmichaelia petriei (× 48) T.S. Young stem with oval pith, iminature cortex and flattened stem
(D) Carmichaelia petriei (× 18). T. S. Old stem to show the oval pith, an indication of the initially flattened nature of the stem although the outline of this section is round l ti, leaf trace

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heavily cutinised epidermis on their ridges may persist for a considerable time as vertical lines on old stems, although cork cambium has extended across the ridges (Pl. 29 F).

Leaf Trace Systems

1. Relationship of Shoot Structure to Taxonomy and Phylogeny

Sinnott (1914) by comparative anatomical study showed the node with three leaf gaps in the stele (trilacunate) to be primitive. He showed the unilacunate node with one leaf gap to have been derived from the trilacunate condition either by gradual movement of the two lateral traces into the same position as the central one, or by the lateral traces disappearing, only the central trace persisting. Multilacunate nodes with more than three leaf gaps have been derived from the trilacunate node by one of two methods. In some, supernumerary traces have been added on the flanks of the original trilacunate insertion (i.e. flanking traces), while in others there appeared to be a diversion into the leaf of some of the traces normally destined to serve the axillary branch (interpolating traces). Neither Sinnott (1914) nor Dormer (1945) mentioned multilacunate nodes in which both flanking and interpolating traces were present, but this paper will describe nodes in which the additional traces are of both types. Sinnott (1914) found that the seedling of the unilacunate and multilacunate forms often showed the primitive trilacunate condition. The Leguminosae he studied are tri- or penta-lacunate. Dormer, although principally investigating the taxonomic value of the shoot structure in the Leguminosae, also showed the same principles to extend to many other families. In the following table he listed criteria that could be applied to problems of phylogeny.

Primitive Characters Advanced Characters
1. Spiral phyllotaxy 1. Phyllotaxy verticillate, + or distichous with two vertical rows.
* 2. Woody habit. 2. Herbaceous habit.
*3. “Open” bundle system. 3. “Closed” bundle system.
4. Leaf trace insertions in contact. *4. Leaf trace insertions separated or interlocked.
5. Leaf highly compound. *5. Leaf less compound
*6. Stipules present and free. 6. Stipules absent or *adnate to the petiole.
*7. Leaf with three traces. *7. Leaf with more or less than 3 traces.
*8. Pulvinus present. 8 No general foliar pulvinus.

“Closed” bundle systems are those in which nodal anastomoses connect bundles of the vascular system tangentially so that there is no necessity for a continuous cylinder of secondary tissues for tangential continuity. They are always present in plants of extreme herbaceous habit. No such anastomoses are present in the “open” systems, which are always associated with secondary growth. Insertions of leaves in “contact” are those with extreme lateral traces of successive leaves arising vertically above one another or adjacent to one another. Those “interlocked” have points of origin overlapping. The trilacunate condition is the most common in the Leguminosae and departures from this are not found in the highly standardised “closed” bundle system (Sinnott, 1914). On the other hand, multilacunate nodes have been found to be frequent in “open” bundle systems and it will be seen that the New Zealand broom genera provide further examples of this.

[Footnote] * Features present in one or other New Zealand broom species.

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2. Carmichaelia Leaf-trace Systems

A. Trilacunate. Apart from two very reduced forms which are mentioned later, only the round-stemmed Kirkiella (Text-fig. 4 A) is consistently trilacunate, but usually the other subgenera are trilacunate at the first seedling internode and often at every branch base where flattening is at a minimum, e.g., Text-fig. 6 A. C. E..

B. Types with Additional Traces “Flanking”—Hultonella, Monroella, Petriea. The flattening in these groups arises by the division of the stem bundles between the original lateral traces of two successive nodes (between Lx and Ly, Text-fig. 4 A). If these bundles divide at an equal rate on each side of the stem, then the flanking traces are added in pairs to form pentalacunate nodes (Text-fig. 6 F) On the other hand, if the division is greater on one side than upon the other, so that the stem becomes flat or slightly concave on one surface while convex on the other, the flanking traces may only occur on this convex side of the stem. Such a node is tetralacunate and asymmetrical, the median trace being distinguishable by the fact that it is at the edge of the cladode (Text-fig. 2 E).

C. Types with Additional Traces “Interpolating.” These are found only in the extremely flattened species of Carmichaeliella, the rest of the subgenus having the nodal structure discussed under Section D. The initial flattening in these species is also by the division of stem bundles between the lateral traces of two successive nodes, but this is superseded by more intensive division of the bundles that normally supply the axillary bud. This division of axillary bud bundles does not take place at the lowest nodes of branches which are almost round, but at successively higher internodes division and spacing of the bundles increases until the stem attains maximum width. It is constricted, however, at each node, and it is when the bundles are thus congested that the interpolating traces withdraw into the cortex (Text-fig. 6 D). The trilacunate node of the branch base becomes pentalacunate as the stem flattens, and finally septalacunate. But there may be asymmetrical nodes where interpolating traces only depart on one side.

D. Types with Additional Traces both Interpolating and Flanking. The species of Carmichaeliella (other than those of division C above)), Thomsonella, and most members of the subgenera Enysiella and Suterella, have extra traces of both types. In a few species nodes may have only extra traces flanking but usually there are a varying number of interpolating traces as well (Text-fig. 6 G), and nodes with interpolating traces only are very common. They have been found in Ca. solandri, Ca. arborea, Ca. flagelliformis, Ca. hookeri (Carmichaeliella); Ca. grandiflora (Thomsonella); Ca. corrugata (Suterella) (Text-fig. 4 C and Text-fig. 6 B). However, Group D do not exhibit such well-developed flanges as stems of Group C. It seems fairly obvious that such stems show an intermediate stage between the two types of stem flattening described under B and C, and between their corresponding nodal structure.

Of the six nodes of Ca. enysii that were examined, all were trilacunate (Text-fig. 4 B). Ca. uniflora, the smaller stemmed species of Suterella, was also trilacunate. As both these species are small reduced forms (Pl. 27 C and D), it is unlikely that the trilacunate node is phylogenetically significant in them. Further investigation of nodal anatomy may reveal that extra traces may sometimes be present, though the small stem dimensions make this unlikely.

3. Corallospartium and Chordospartium Leaf-trace Systems

In Corallospartium and Chordospartium extra traces are exclusively flanking and added progressively as the stem flattens by the division of the stem bundles

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Text-fig. 6—A. T.S. branch Ca. solandri at the base (× 16). B, T.S. branch Ca. solandri several nodes above the base (× 16), C. T.S. branch Ca. williamsii at the base (× 16). D. T.S. branch Ca. williamsii several nodes above the base (× 16). E. T.S. branch Ca. hollowayi at the base (× 40). F. T.S. branch Ca. hollowayi several nodes above the base (× 40). G. T.S. branch Ca. grandiflora several nodes above the base (× 16). These are selected examples to show the various types of leaf trace that are found in Carmichaeliella, Thomsonella, Monroella Abbreviations as Text-fig. 2.

between the lateral traces of two successive nodes. Text-fig. 7 is a diagram illustrating the leaf-trace system of a seedling of Corallospartium. The first internode (Text-fig. 8 B) shows the stem isodiametric and with four corners associated

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Text-Fig. 7—Diagram of the leaf trace system of a Corallospartium seedling. Stem bundles are not shown. N1-N8, Nodes 1–8. In1-In8, Internodes 1–8. For explanation see text.

with four leaf traces. At higher internodes the number of ridges increases, along with an increase in the number of flanking traces (Text-fig. 8 A). The median trace always runs through two internodes before departing to a leaf, but lateral traces may arise only one internode before supplying the leaf (Text-fig. 8).

In the adult stems a varying number of ridges are present, depending upon the number of trace bundles per leaf, and whether or not the leaf insertions are overlapping. In young stems and tapering branch tips the most remote pair or pairs of flanking traces overlap (Text-fig. 8 A, F5 and F6 overlap). Sometimes a ridge is seen to be dividing, this being associated with the division of a trace bundle (Text-fig. 8 C and D at X), while on the other hand, the suppression or reduction of a trace bundle, particularly when the stem is narrowing towards

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Text-Fig. 8—A T.S. stem of Corallospartium seedling at node 5 of Text-Fig. 7 (× 48). B. T.S. stem of Corallospartium seedling at node 1 of Text-fig 7 (× 48) C and D, T. S.'s through the node of an adult Corallospartium stem to show the division of one of the ridges at “x” associated with the division of the leaf-trace bundle (× 10). M, M1, M2, median traces; L, L1, L2, lateral traces; F, F1, F2, flanking traces.

the distal nodes, is accompanied by a partially formed ridge. Such features show the close relationship between ridge and leaf-trace development.

Although Corallospartium has been investigated both from developmental anatomy as well as from the anatomy of the adult, and Chordospartium only from investigation of the adult, both genera seem remarkably similar, despite the fact that they are so different in habit and stem dimensions. Chordospartium sometimes has fewer ridges in its stems, but this is apparently connected with its smaller dimensions (Pl. 29, B).

4. Notospartium Leaf-trace Systems

The nodes of stems of seedlings of various ages show that the trilacunate node becomes pentalacunate and septalacunate by addition of flanking traces. Occasionally an asymmetrical tetralacunate node is developed. Insertions are initially in contact, but in branches of the older seedlings the stem bundles between the last pairs of flanking traces of two successive nodes (Text-fig. 9 A between Fx and Fy) increase in number so that insertions become separated. Older seedlings of Carmichaelia species were not available for similar investigation in that genus, but from comparison of leaf insertions in adults of both genera, a similar change appears to occur in Carmichaelia.

One difficulty arises in the adult Nolospartium which is not met with elsewhere, this being the variability of the adult nodes both in number of trace

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Text-Fig. 9—A. T.S. stem of Notospartium seedling of one year (× 40), B and C. T.S. nodes of adult Notospartium (× 18). Abbreviations as for Text-fig. 2.

bundles and in their arrangement and departure from the stele. Although the oldest seedlings available for this investigation showed a distinct edge to the cladode, the adult stems are more rounded and consequently median trace bundles are impossible to determine (Text-fig. 9 B, C), particularly in a trace consisting of an even number of bundles. Moreover, the trace bundles are usually very late in branching from the stem bundles, so that they may run as distinct leaf-trace bundles for only a very short distance in the internode. This feature was also often noticed in the seedling (Text-fig. 9 A). The other feature peculiar to this genus was that at some seedling nodes there was apparent movement of the original three traces to one edge of the stem, while the axillary bud bundles usually separating them tended to move further in (Text-fig. 9 A). Under such circumstances the median and original lateral traces would depart from one gap in the stele instead of from three, this being a similar change to one described by Sinnott (1914) for the origin of a unilacunate from a trlacunate node. A trace with three bundles departing from one gap was not observed in an adult node, but there were many gaps from which two departed. In view of what was observed in the seedling, these pairs may be two approximated traces rather than a divided trace. To sum up, it seems clear that in Notospartium initially flanking traces are added as the stem flattens, but subsequently various changes peculiar to the Notospartium genus take place, so that the interpretation of the structure of the adult is not possible until plants of intermediate age are available for stages between that of a one-year seedling and the adult.


Text-fig. 10 is an attempt to reconstruct the history of the New Zealand brooms using cladode structure as the principle guide. Kirkiella is chosen as the ancestral type because of its unflattened stem and trilacunate node. From Kirkiella arose an ancestral stock in which extra traces were added on the flanks. Its most direct derivatives to-day are the Petriea, Monroella and Huttonella subgenera. From

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this stock in turn came forms in which further flattening was accompanied by the interpolation of traces, a modification which as time went on tended to supersede the older supplementation by flanking traces. Thomsonella and some of the forms of Carmichaeliella, e.g. Ca solandri, occupy a primitive place in this series, the bulk of their nodes having added traces of both types, while the greatly

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Text-Fig. 10—Suggested evolution of the cladode types in the New Zealand brooms. Explanation in text.

flattened members of Carmichaeliella show complete replacement of the flanking by the interpolating traces. They are connected with the Ca. solandri type by a complete series of intermediate forms. Enysiella and Suterella are linked to the ancestral Carmichaeliella stock in the belief that their trilacunate node is not primitive but the result of elimination of both interpolating and flanking traces as the habit became reduced to a mat form. The fact that Ca. corrugata (Text-fig. 4 C), the wider-stemmed Suterella species, does frequently have extra interpolated traces, upholds this idea.

Corallospartium and Huttonella have in common flanking trace systems and indehiscent pods. The indehiscent pod distinguishes Huttonella from the other Carmichaelias. However, Corallospartium and Huttonella became separated when Corallospartium, becoming more flattened, added further traces so that its leaf insertions met or interlocked.

The structure of Notospartium seedlings indicates a close relationship between Notospartium and those Carmichaelia species in which extra traces are flanking. In Notospartium adult, however, there is much greater multiplication of stem bundles between the lateral traces of successive leaves, causing leaf insertions to be further apart. The adult stem is more rounded at its edges than in Carmichaelias, although its pith is distinctly flattened.

Chordospartium's connection with the other genera has for long been a problem. the anatomical evidence presented here indicates a close relationship with Corallospartium, the only other genus with furrowed stems. Trace systems are very similar (Pl. 29 B, C), and, as noted by Cheeseman (1925), pods are almost identical. However, the inflorescence, habit, and wiry stems of Chordospartium are reminiscent of Notospartium (Pl. 29, A, B) and quite unlike the rigid erect habit of Corallospartium (Pl. 27 B, Pl. 29 C) Cheeseman, in fact, says that Chordospartium combined characters of both genera. The pod structure is

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common also to the Huttonella section of Carmichaelia, a fact which has led Laing and Blackwell (1940) to suggest a possible hybrid ancestry of Chordospartium from Notospartium and Carmichaelia. The other possibility implied even in the similarities listed by Cheeseman is its derivation from the hybridisation of Notospartium and Corallospartium. The anatomical evidence strongly favours this alternative and in this case the few Carmichaelian features of Chordospartium could easily be derived from either parent, since both are probably descended from Carmichaelia.

The writer hopes to collect further evidence from wood histology and cytology to elucidate the relationship of these three genera.

Literature Cited

Cheeseman, T. F., 1925. Manual of New Zealand Flora, 2nd Edition. Wellington.

Cockayne, L., 1928, Vegetation of New Zealand Leipzig.

Dormer, K. J., 1945. An Investigation of the Taxonomic Value of Shoot Structure in Angiosperms with Especial Reference to the Legummosae. Ann. Bot. N.S., 9, 141–153.

— 1946. Vegetative Morphology as a Guide to the Classification of the Papilionatac. New Phytol., 40, 145–161.

Kirk, T., 1899. Students' Flora of New Zealand. Wellington.

Laing, R. M., and Blackwell, E. W., 1940. Plants of New Zealand, 4th Edition, Whitcombe & Tombs.

Simpson, G., 1945 A Revision of the Genus Carmichaelia Trans. Roy Soc. N.Z., 75, 231–287.

Sinnott, E. W., 1914. Investigations on the Phylogeny of the Angiosperms. Anatomy of the Node as an Aid in Classification of the Angiosperms. Amer. Jour. Bot., 1, 303–322.

Solereder, H., 1908 Systematic Anatomy of the Dicotyledons, English Trans., Boodle and Fritsch, Clarendon Press, Oxford.

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The Occurrence of Aquatic Oligochaetes in Soil

[Read before the Wellington Branch, August 28, 1951; received by the Editor, August 28, 1951]


Two species of aquatic oligochaetes identified as Aeolosoma kashyapi Steph. and A. niveum. Leyd. are described and figured from soil from the crater floor of Raoul Island, Kermadec Group.


Two soil samples collected by Mr. A. C. S. Wright from the floor of the main crater on Raoul Island, in the Kermadec Group, yielded rich cultures of two microscopic aquatic oligochaetes. The species are cosmopolitan in distribution, but this occurrence is of interest in that they have not previously been recorded from the South Pacific area, nor have they been recorded previously in soils.

Their presence in two of the soil types (Tui silt loam and Green Lake loamy fine sand) of the floor of the crater is probably correlated with the existence of an old crater lake. There are abundant signs that a lake once filled the south-west part of the main crater, resulting from a minor eruption on the present site of Green Lake which ejected material blocking the natural drainage channels from the western and southern slopes of the internal watershed of the main crater (Fig. 1) Although normally found in aquatic environments, these worms can withstand dry conditions by encystment (Beddard, 1892a).

The two species described belong to the genus Aeolosoma Ehrenberg (1831), principal genus of the family Aeolosomatidae. The group has recently been described by Marcus (1944), who gives a key to the species. One of the Raoul Island species resembles Aeolosoma kashyapi Stephenson. The second species appears to be identical with A. niveum Leydig, for which no adequate descriptions have been previously given (Beddard, 1895; Michaelson, 1900). Marcus (1944) refers to a description by Ude (1929), but I have not been able to consult this source.

Aeolosoma kashyapi Stephenson, 1923. (Figs. 2a. 2b, 4)

The fully grown worm is about 0.8 to 1.0 mm. in length, with a broad ciliated prostomium, the activity of which aids the animal in ingestion and movement. There are up to 12 segments with a division between the seventh and eighth which marks the budding zone. n, the number of segments of the adult worm, is therefore seven and there is never more than the one daughter worm.

The prostomium is flattened and almost circular, being about 80μ in diameter. Ventrally it is wholly ciliated, the beat of the cilia directing debris and food particles to the buccal funnel, which is also strongly ciliated, as is the whole of the alimentary canal. The edge of the prostomium is marked by a non-ciliated rim, about 5μ in width, which adheres to the surface of the substrate. The outer margin of this rim is also ciliated. The peristomium, the first true segment of the worm, appears circular when viewed from the ventral surface. Ventrally the opening of the buccal funnel occupies most of the segment. It is surrounded by a broad wall, about 10–12μ in width, which does not meet anteriorly but fuses with the prostomium, forming a channel through which the food particles are driven by the cilia of the prostomium to the buccal funnel.

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Fig. 2—(a) Acolosoma kashyapi. Ventral aspect, showing internal anatoms. (b) A. kashyapi. Nephridium.
Fig. 3—(a) Acolosoma niveum. Ventral aspect. (b) A. niveum Nephridium.
Fig. 4—Typical seta.

b.f., buccal funnel, br., brain, b. v., vential blood vessel; b. z, budding zone, e. g., colour gland. int., intestine; n., nephridium; nst., nephrostome; oe., oesophagus: pr. prostomium. s. seta: st., stomach.

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The alimentary canal is characterised by a sinuous oesophagus which extends to the fourth segment, a broad stomach from the fourth to the sixth segment, and a narrow intestine which runs from the seventh segment to the end of the body. There are no well-developed inter-segmental septa, but the gut is attached by a number of bridges to the body wall. There is typically only one such bridge per segment, starting wtih the fourth segment. Between the seventh and eighth segments, where budding takes place, the gut and body wall are much more closely apposed, there being a definite union of connective tissue for the greater part of the zone. The anus is quite simple and terminal. Ciliary action suggests that it may be an auxiliary respiratory organ. The gut shows typical peristalsis.

Nephridia are present on both sides in segments IV to VII inclusive and in segment IX. The nephridium is a simple tube with a ciliated nephrostome coiled in the form of a loop (Fig. 2b), the halves of which are closely apposed and intimately connected, so giving the appearance of an oval body, 30 × 15μ in size. There may be more than one loop, but the oval shape is retained and is very characteristic. There is a non-contractile ventral blood vessel formed by the union of two circum-buccal vessels and a dorsal contractile vessel anteriorly. The brain was the only part of the nervous system observed. This organ, somewhat kidney-shaped and formed by the partial fusion of the two cerebral ganglia, can be observed from the ventral surface as lying in the posterior half of the prostomium. The posterior indentation of the brain is conspicuous, but the organ varies slightly in shape with different individuals. The integument is about 5μ in thickness and is conspicuous by the presence of numerous colour glands. These are also present in the prostomium and are deep orange. These colour glands are the most striking feature of the worm and form a striking contrast to the drab colour of glandless worms. The setae are of importance in establishing the specific status of the material They are embedded in a sac which projects from the body wall into the coelomic cavity, and to the tip of the sac protractor and retractor muscles are attached. The contraction of these muscles enables the setae to serve in the locomotion of the animal, supplementing the contractions of the body musculature.

The setae are all capilliform, fine and hair-like, and about 80–85μ long in the adult worm (Fig. 4.) They are identical in both the dorsal and ventral bundles. The number of setae in each bundle varies, there being up to 5 in the dorsal and up to 3 in the ventral. The third segment, however, has 4 setae ventrally and not 3 in the mature worm. The segments of the daughter bud have only 2 setac in both dorsal and ventral bundles.

No clitellum was observed; nor were any reproductive organs. Fission was the only observed mode of reproduction. In the material observed only one bud appeared to be formed at a time. The limitation of size to only 12 segments would, of course, only allow one daughter bud to be lormed at a time. The budding is quite typical, the thickening between the segments VII and VIII giving rise first to a slight protuberance, the join narrowing and finally being nipped off and the daughter bud then developing the elaborate prostomium of the adult and its full complement of setae.

Aeolosoma niveum Leydig (1865). (Figs 3a, 3b, 4.)

This worm, which is about 1 mm. in length, has up to 10 segments; the broad, flattened prostomium typical of the genus; the wide buccal cavity; and capilliform setae. The budding zone is again between segments VII and VIII; n is 7. The prostomium is flattened and completely eiliated ventrally, as is the outer rim of the prostomium. The peristomium surrounds the wide buccal cavity, and

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like the prostomium is somewhat broader than the rest of the body. It has the same broad wall and surrounding rim as the previous species. The cerebral ganglion is strongly indentated, showing the partial fusion of the supraoesophageal ganglia.

The nephridia, in distinction to those of the previous species, are not in the form of a loop. They consist of a simple tube with a ciliated nephrostome as in the previous species (Fig. 3b), but this lies and winds along the side of the gut. The ciliated nephrostome is somewhat bulb-like and the tube swells again at its posterior extremity. The setae are of the same kind and size in both dorsal and ventral bundles. They are fine, hair-like and capillary with a slight bend as in the previous species. They are up to 65μ in length. (Fig. 4.) Unlike the previous species there are the same number in both dorsal and ventral bundles of each segment. The maximum number is 4 which are present in III; the other segments have a maximum of 3 excepting the posterior daughter segments, which have only 2, as in the previous species.

There are no colour-glands in the integument and consequently this worm is colourless in distinction to the preceding species.

No reproductive organs were observed, multiplication by fission being the invariable rule.


Marcus (1944) recognizes nineteen species of Aeolosoma and sixteen of these are coloured worms A. kashyapi Stephenson is distinguished by an orange colouration, slightly curved capilliform setae, and the absence of ciliary pits.

In Fauna of British India (1923), Stephenson gives a brief diagnostic description of this species. He states, amongst other details, that n is either 7 or 8, he implies that the setae are uniform, and that they are about 60μ in length. Aiyer (1926) describes a worm which he calls A. kashyapi. He affirms that n is always 8, and that the setae are of two kinds, one long, c. 66μ in length, the other about half that length, viz. 38μ. Another distinction is that Stephenson remarks that the nephridia do not occur behind the eighth segment. Aiyer states that nephridia occur ‘in all succeeding segments.' Aiyer does not record the number of segments in his worms. The worm described by Aiyer appears to differ in possessing two types of setae, but notwithstanding this, both Aiyer and Marcus consider it to be A. kashyapi.

Marcus both describes and figures this species. He is the first and only author to describe its reproductive system. The only points of difference between Marcus' worm and the worm here described are first the structure of the buccal cavity and secondly the number of segments in the adult worm. This latter would appear to be an illusory distinction, since Marcus' figure clearly shows the division zone between the segments VII and VIII, although in his description he states that n is 8 and not 7. He further records that the number of segments in his worms varies from 7 to 12 and it seems strange that a worm should have less than n segments. The structure of the buccal cavity, which is not described by Marcus, is figured slightly differently from that of the present worm, in which this structure is very well developed. The distinction is not, however, great. In all other details the two worms are identical, although Marcus records a wide range of size dimensions and also great variation in the number and size of the setae Doubtless it is because of this variation that he includes Aiyer's worm in the species, although Aiyer described setae of two lengths.

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Fig. 1—Raoul Island. Keimadee Group, showing location of main Crater and soil sample sites.

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In the present material, n is always 7. The setae are uniform, but c. 80μ in length. Nephridia do not occur in the eighth segment, which for this worm is the first segment of the daughter worm.

Marcus recognizes three species of colourless worms. Aeolosoma niveum is the only one of the three with simple capilliform setate; the other two species have as well sigmoid or serrated setae. The setae of the present material are, as observed, similar to those of A. kashyapi.

Beddard's (1895) description of A. niveum is brief: “Prostomium not wider than following segments. Setae sigmoid and capilliform. Integumental globules colourless; only one pair of nephridia at end of oesophageal region. Very minute.” Beddard, however, has confused A. niveum, which has only one type of seta, with A. beddardi, which has two (1892b).

Michaelsen (1900) gives the following description of A. niveum: “Kopflappen vorn zugespitzt, nicht breiter als die folgenden segm. Oldrusen farblos, ungleich gross, unregelmassig zerstrent. Borsten vorn zu 3 oder 4, hinten zu 2 im Bundel, schwach S-forming gebogen, mindestens so lang wie die segm. Gehirn hinten schwach aus geschnitten. Erstes Nephridienpaar vor dem 3. Paar Borstenbundel. L 1–2 5 mm. segms der Einzeltiere 12–13.”

While this description is not very detailed, it does not differ from the worm described in this paper except in the number of body segments. Important details such as the site of fission, the number and shape of the nephridia, and the ciliation of the prostomium are not given. Accepting this identification, this is the first record of the worm outside Europe, with the possible exception of Leidy's Chaetodemus panduratus, a synonymy given by Beddard.


Aiyer, K. S. P., 1926. Notes on the Aquatic Oligochaeta of Travancore. II. Ann. Mag. Nat. Hist., Series 9, vol. 18, pp. 131–42.

Beddard, F. E., 1892a. Note upon the Encystment of Aeolosoma. Ann Mag. Nat Hist., Series 6, vol. 9, pp. 12–19.

Beddard, F. E., 1892b. On some Aquatic Oligochaetous Worms. Proc. Zool. Soc. London, 1892, pp. 349–61.

Beddard, F. E., 1895. A Monograph of the Order of Oligochaeta. Oxford, 1895.

Marcus, E., 1944. Sôbre oligochaeta limnicos do Brasil. Bol. Fac. Fil Ciên. Letr. Univ. S. Paulo., vol. 43, Zool. No. 8, pp. 5–135.

Michaelsen, W., 1900. Oligochaeta. Das. Tierreich, vol. 10. Berlin, 1900.

Stephenson, J., 1923. Oligochaeta. Fauna of British India series. London, 1923.

Ude, H., 1929. Oligochaeta. Tierivelt Deutschlands, vol. 15, pp. 1–165. G. Fisher, Jena, 1929.

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Sir William Blaxland Benham, K.B.E., F.R.S., (1860–1950)

In the face of the indelible personal impressions left by a friendship of some forty-four years, it is very difficult to write objectively about Sir William Benham; but, fortunately, I have also other sources from which to compile this notice; and my thanks are due to Miss Marion Fyfe, his Senior Lecturer in Zoology, for the use of a personal record left her by him; to Professor J. S. Tennant, of Nelson, who held the fort until Sir William's arrival in New Zealand in 1898; to Mr. J. W. Hayward, Registrar of Otago University, who made the University records available; and to Mr. L. W. James, Registrar of Marlborough College, England, for information on Sir William's school days. Other records are expected from England, but, as they have not come, I feel that further delay in issuing this notice is unwarranted.

In any attempt to review the life of this great man, one becomes involved in the zoological history of the past one hundred years; because, not only did Sir William's career cover that century (except for the first decade of it), but also he became an eminent disciple of the Darwin-Huxley influence which he carried to New Zealand, and so maintained at Otago the tradition already founded by his famous predecessor, T. J. Parker, himself a pupil of Huxley. That these influences remained deep rooted was to be seen (if in nothing else) from his habit of periodically and alternately hanging over his mantelpiece at the Museum the portraits of Darwin and Huxley; when asked why he did not hang them side by side and be done with it, he replied that, as man was blessed with only one digestive tract, he could not possibly eat, much less digest, two solid meals simultaneously.

Sir William was born (the sixth in a family of seven) on March 29, 1860, at Isleworth, Middlesex, just at the moment when Darwin's Origin of Species was yet hot from the press, and when the controversy (largely led by Huxley) was raging around it; it is not suggested that this in any way disturbed Sir William at the time, but it serves to fix the period. He knew nothing of his forebears beyond his paternal grandfather who had been a publisher in partnership with Reeve of Henrietta Street, Covent Garden, and who published a variety of books on Natural History—a noteworthy point, perhaps; his father, “Edward Benham (baptised ‘Ebenezer', but changed his name) was a London Solicitor” and President of the Metropolitan and Provincial Law Association at the time of his death in 1871; his mother, Mary Anne Shoppee, of Uxbridge, died in 1900. Sir William claimed that none of his “relations were in any way noteworthy nor took part in public life—just ordinary, fairly prosperous folk of the professional class”, and that he “was brought up in a happy and healthy environment in a large house with extensive grounds of about three acres at Syon Lodge, Isleworth, adjacent to the Duke of Northumberland's house”; he did not recall any circumstances or influence affecting his later history as a zoologist, none of his family or ancestors having been interested in science. Nevertheless, his childhood background would tend to direct him into some intellectual channel.

His schooling commenced in 1867 at a grammar school at Coleshill, Warwickshire, under the Rev. W. Banks; in 1871, the year of his father's death, he transferred to a school at Ewell, near Epsom, and thence to a small school at Meriden, Warwickshire (W. Carles)—a preparatory institution for boys about to enter the Public Schools.

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In September 1874, he entered Marlborough College, his memories of which were to remain amongst his happiest, as revealed by a letter written to the College in his latter years His house was C1, part of the old building once the mansion of the Seymour family and later the famous Castle Inn. He remained there for seven terms, leaving in December, 1876 During six of these terms he received promotion and finished in the Lower V.

We should note that he was a member of the College Natural History Society, which was the first of its kind in any school and flourished under the Rev. T. A. Preston—it continues to flourish. The activities of the Society were wide. In spite of the fact that Sir William disclaimed having been influenced by any of his teachers in any way in his choice of what was ultimately to become his life work, and though there is no record of his having done anything to have his name mentioned in the proceedings of the College Society, it is quite reasonable to believe (very young though he was) that the beginning of his zoological bent had its roots somewhere in that Society, which was of high standard and still is; because later, when the opportunity came and after he had felt his way about, he succumbed to the zoological spell readily enough.

At Marlborough he took classics, as well as chemistry and physics, under Mr. Rodwell (the only science taught), “not for any interest”. he averred, “but because thereby I escaped having to do Latin verse! As at that time I intended to enter the Indian Civil Service as an engineer. I took various extra subjects, such as mechanical drawing. German, etc.”, and “was well grounded in Latin and Greek, ignorant of History, English Literature and Grammar, with a smattering of French Grammar and elementary Mathematics”: he won no scholarship at any time in his career.

His training in drawing is reflected by the clearly detailed (often artistic) illustrations in his publications; he always stressed the value of what he called “good pictures”; he would carefully ink in a line, and then, aided with a lens, go over it and fill in the irregularities with a fine pen. His knowledge of Latin, Greek and German became invaluable to him and his students in later years. His confessed ignorance of English literature and grammar is not to be taken seriously, or, if it is, then he obviously rectified it. On the other hand, there is possibly some truth in the smattering of elementary mathematics, when one recalls his efforts to balance his petty cash! Nevertheless, his school record. though seemingly commonplace, is what we find often enough in the histories of so many famous people.

On leaving school in 1876, he lived with his family in London whilst being coached for the Indian Civil Service; but this plan was eventually abandoned, and he commenced study to become an analytical chemist at University College, Gower Street; after two or more years, however, this also was abandoned, and he turned to Geology (under Bonney) in its place as a third subject for the B.Sc., but. as there were no laboratory or museum facilities at University College, he transferred to the Imperial College (then the School of Science and Art), where he studied under Judd and Cole. His other subjects were Botany (Scott and Vines) and Zoology under Ray Lankester. whose influence shaped his career. He graduated B.Sc. at London University in 1883.

It was of that period he claimed, “I had found my real interest in Zoology, in which I received much encouragement from Ray Lankester; and after acting as Junior Demonstrator, he appointed me as a senior assistant after A. G. Bourne had been appointed to a Chair of Zoology in a university in India”; that was

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Sir William Blaxland Benham

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in 1886, when he also became Lecturer in Zoology at Bedford College for Women in London, a position he held until 1898, when he left for New Zealand.

So it is that we find Sir William not only launched on his zoological career, but also with the fundamental training in Geology and Botany which enabled him, as Professor of Biology at Otago, to carry on the teaching of Palaeontology until Dr. P. Marshall (Professor of Geology) took over the subject some time about 1907, and of Botany until 1923, when Dr. J. E. Holloway became lecturer in that subject. Though naturally very interested in Palaeontology, Sir William had very little leaning toward Botany beyond what was required for the needs of the curriculum.

Whilst yet a student, Sir William made a study of the male organs (then unknown) of Limulus, living material having been secured from the Westminster Aquarium; the result was his first publication in 1883, inaugurating that long series of outstanding zoological publications which regularly appeared, and which fittingly closed simultaneously with his life by the issue of his last paper in the very month of his death. We would note that, though Sir William usually used his initials “W.B.” we find “W.B.S.” on this first publication, the “S.” representing “Shoppee” after his mother. He sent a copy of his Limulus paper to Sir Richard Owen—that man with the “surgical smile” (Emerson), the sweetness of which reminded Jane Caryle of sugar of lead—whose memoir was the only source of knowledge of the animal at that time.

In 1885 Sir William commenced his life work on the structure of earthworms. upon which he became such a world authority; on his original studies he graduated D. Sc. in 1887. The British Museum provided him with material from all parts of the world, and so quickly did he came to the fore that he was invited to contribute the account on “Polychaet Worms” in the Cambridge Natural History, which remains the only detailed monograph on the subject in the English language, while a little later he contributed, at Lankester's request, to the latter's Treatise on Zoology, part IV, “Platyhelmes, Mesozoa and Nemertinea”. Indeed, so authoritative did his work become, that many of his published drawings of worms and other animals are reproduced in several standard zoological books by English. German and French authors.

After graduating, Lankester advised him to study the field of karyokinesis then being developed by Prof. Wilhelm Fleming at Kiel; this he did for some weeks, “but rather to Lankester's disappointment I did not pursue this work. It did not appeal to me as did that of work on larger material”, a turn of mind which is obvious in the range of subjects dealt with in his publications; I think the smallest was an insect larva, but the largest were certainly whales, about which he became very enthusiastic for a time in his seventies, though he eventually reverted to his worms, apropos which he said, “If one's fickle enough in youth to forsake one's first love, one returns to it in old age—for it will claim you in the end!” Such characteristic humour frequently cropped out, and he appreciated it in others even though he was the object of it—except when derisive; for example, when someone placed an almost spherical quartz pebble in a cage of tuatara lizards which he had under observation in the hope of securing the egg, he was far from pleased; on the other hand, when one of his early students met him after many years and said, “Every time I turn up a worm in the garden I see your face”. he was highly delighted with the rather naif bon-motism.

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On April 24, 1899, at St. Paul's Church, Camden Town, he married Beatrice Eadie, daughter of John Eadie, a London merehant. She died in 1909—a great loss to Sir William and to all who had the pleasure of knowing her; she was very considerate and helpful to the younger fry, especially to those making their first appearance before the Otago Institute, when she would restore what remained of their courage.

In 1890 Ray Lankester was appointed to the Chair of Comparative Anatomy at Oxford, and Sir William accompanied him to take up an appointment as Aldrichian Demonstrator in that subject; it was during that period that Sir William took his M.A. in 1897, “by decree of convocation”. In the same year Professor T. J. Parker, of Otago University, died, and Sir William applied for the vacant Chair of Biology and the Curatorship of the Museum; he was supported by eulogistic testimonials from several renowned zoologists, the consensus of opinion being that no worthier successor could be had to follow the distinguished Parker. On March 10, 1898, Cabled advice from London stated that Sir William had been selected; and it is of interest that Prof. G. Vines, one of the appointing commission, drew the attention of the Otago University Council to the wisdom of creating two chairs—Botany and Zoology—not to eventuate, however, until 1951. A letter written by Sir William at that time was legible, presenting none of the difficulties so characteristic in later years when a typewriter was given him, it is said—hopefully (by the University Council, perhaps)—though I am sure all his correspondents were pleased when he discarded the machine in favour of the pen; however, toward the close of his life, his writing tended to revert to the early style.

Sir William sailed for New Zealand by the s. s. Kaikoura on March 31, 1898, being thirty-eight years of age virtually to the day. Prof. J. S. Tennant, who acted during the interregnum, gives the following excellent picture of Sir William on handing over to him in May, 1898: “I found Benham a cheery young man, dapper, frank, and intensely intersted in everything I had to show him. Nothing escaped him, and I felt he could look after himself and manage even the toughest medical student”; and later, “in 1908 I was with him for two weeks on the Auckland Islands, and there he was a general favourite Two of our colleagues were rather pompous know-alls, and Benham delighted to draw them out (often further than they could safely go). On the other hand, he was most kind and helpful to all the younger men.” So there was the man we came to know—indefatigable, genial, energetic, kindly, and yet masterful as the occasion arose; a man of definite views, yet with a tolerance; a compact man (physically and mentally), exhibiting nothing of absent-mindedness—real or false; short of stature and rapid in gait.

He took up his work with zest. In the Museum one of his first acts was to plan for its future development, and using as a foundation the material provided by his distinguished predecesors (Hutton and Parker, respectively), he built up over the years a museum of Comparative Anatomy unrivalled in New Zealand, and considered by competent observers to be comparable with the Oxford Museum of Comparative Anatomy; on the other hand, he be no means neglected the ethnological side. He was a stickler for public order in the Museum and even proposed that a policeman be on duty; in a letter to the University Council he considered that the Janitor (Mr. James Mackenzie) was becoming old and who “in a few years will be too feeble to exert any sort of influence on noisy

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youths”—yet that janitor (a feature of the museum) managed quite well until he retired twelve years later of real old age.

Sir William was a fluent and quite informal speaker, whether on the platform or in conversation; he drew upon an orderly mind stored with knowledge as wide as it was deep in understanding; his induction from observed phenomena followed each step to his conclusion: indeed, he always claimed that Logie should have its place in any scientific training—a report, shortly before his death in 1950, on the identity of an earthworm, was an example of precise syllogistic reasoning. He was a great techer, and his lectures were a model; he impressively led his students (in whom he took a personal interest) through the rise of the animal kingdom, the whole plan of creation unfolding His flair for holding the attention of the most irresponsible student (more or less) was doubtless an important influence in the well-known orderliness of his classes; but the greatest influence. I am sure, was the universal respect in which he was held—as a man of integrity and of high standing in the scientifie world, to say nothing of his forcefulness, which sometimes was rendered the more potent by a barbed subtlety not quite obseured under the cloak of courtesy; but if he had caused hurt (and not necessarily unwittingly) he could in many kindly ways make amend so long as he had not to deal with a fool—whom he could suffer neither gladly nor sadly.

Quite apart from his teaching and Museum duties, he led a very full life. He was unremitting in his original researches; and the attached list of his publications (none a pot-boiler) reveals the scope of his interests, while many of them give one an insight into his outlook. When dealing with subjects beyond the scope of his own speciality (earthworms), it was characteristic of him to reveal his modesty and think of others; for example, when examining the viscera of the rare Notornis, he did as little damage as possible, because “I have no special knowledge of bird anatomy” and the specimen might pass “into the possession of a competent ornithologist”; again, he would frankly confess his uncertainty in a field comparatively new to him, as with certain echinids when “a recent discussion as to the proper names of cidarids warns me, an outsider, to beware of rushing in where Baxter, Clark, Mortensen, and others have trodden”; though he would console himself with that, “after all, names are only labels, and need not have any meaning”, but could be of use “for the purpose, at any rate, of reference”, so long as they were accompanied by good descriptions and “pictures”; and when he made a mistake he did not fear to make a handsome admission of it, a trait seen to advantage in what he referred to as “a Confession of Errors”, where “one little error in life, one little sin if you like, leads one into further errors” Frequently, too, his conversational style would creep into his writings to bring to the reader an intimately personal touch, sometimes enhanced by a rather unorthodox title, such as “A couple of abnormalities”

In the administrative affairs of the University he played an influential part. his “wide experience, sound judgement and impartiality of outlook, giving added weight to the opinions he expressed.” He was Chairman of the Professorial Board from 1904–1907. Professorial Representative on the Otago University Council 1914–1930, Dean of the Arts and Science Faculty 1920–1926, and Member of the New Zealand University Senate 1912–1924.

His activities in wider fields were many and varied; from the time of his arrival in New Zealand he took a leading part in the Otago Institute, giving long serive on the Council, and was President and represented the Institute

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on the Board of Governors of the New Zealand Institute for several years; he was President of Section D of the Australasian Association for the Advancement of Science at Hobart in 1902; he delivered many public lectures and served on the Committee of the Workers' Educational Association; he was a member of the Subantarctic Expedition in 1907, and one of the sub-editors of the Reports; a member of the London Eugenics Society Council, and founder and President of the New Zealand Society, “which collapsed in 1915 owing to the war then raging” (he explained, not without humour); a member and sometimes leader of the Dunedin Naturalists' Field Club (though he denied being a naturalist); he was a member of the Cawthron Commission in 1917 to draw up the original scheme for the functioning of the Cawthron Institute, and he delivered the Annual Cawthron Memorial Leeture in 1918; President of the New Zealand Institute (now Royal Society of New Zealand) 1917–1918; for many years a member of the Portobello Marine Research Station and Chairman of the Board from 1926–1948; as Secretary he was very active in organising, planning, and raising funds for the Hocken Wing to the Museum, and was Chairman of the Hocken Library Committee, 1934–1936; he served several years on the Council of the Overseas League; he was an original member, and Preident for two years. of the Otago University Club, in which he was a well-known figure frequently seen at the billiard table; and if any other diversity of interest in this great man is to be recorded, he was a regular and enthusiastic patron of the motion picture theatres for many years, and Patron of the Film Society.

His public honours were:

  • (1902) Corresponding Member of the Royal Society of Tasmania.

  • (1907) Fellow of the Royal Scoeity, London.

  • (1911) Hutton Memorial Medal (on the first occasion of the award).

  • (1919) Fellow of the Royal Society of New Zealand (one of the original Fellows).

  • (1935) Hector Memorial Medal.

  • (1937) Emeritus Professor of Biology, Otago University (he retired in 1936).

  • (1937) Hon. D.Sc., New Zealand University.

  • (1937) Coronation Medal.

  • (1939) Knight Commander of the Order of the British Empire.

Such is something of the man, a great figure in a great generation of zoologists; and inscribed in the Archives of the University of Otago is found: “His vigour and originality of mind, his high ideals and his devotion to the advancement of knowledge, have ensured a powerful influence on the University and general community. His kindly and gracious personality has won the warm regard and affection of all who knew him, and his name will long be remembered in the University he adored”. He died on August 21, 1950.


1883. On the testis of Limulus. Trans. Linn. Soc. Lond., (2) 2: 362–366.

1886. Studies on earthworms.—i. Quart. J. Micr. Sci., 26: 213–302.

— Studies on earthworms.—ii Quart. J. Micr. Sci., 27: 77–108.

1887. Studies on earthworms.—iii. Quart. J. Micr. Sci., 27: 561–572.

— Recent researches on earthworms. Rept. Brit. Assn., 1887, pp. 749–750.

1888. British earthworms. Nature, 38: 319.

— Note on a new earthworm. Zool. Anz., 1888, p. 72.

1889. The anatomy of Phoronis australis. Quart. J. Micr. Sci., (2) 30: 125–158; (also J. Roy. Micr. Soc., 6: 740–741).

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1890. “Atrium” or “Prostate”. Zool. Anz., 13: 368–372.

— The genera Trigaster and Benhamia. Ann. Mag. Nat. Hist., 6: 414–417.

— An attempt to classify earthworms. Quart. J. Micr. Sci., 31: 201–315.

1891. Note on a couple of abnormalities. Ann. Mag. Nat. Hist., (7) 7: 256–258; (abstract in J. Roy. Mior. Soc., 1891, p. 328).

— The nephridium of Lumbricus and its blood supply; with remarks on the nephridia in other Chaetopoda. Quart. J. Micr. Sci., 32: 293–334.

— Report on an earthworm collected for the Natural History Department of the British Museum by Emm Pasha in Equatorial Africa J. Roy. Micr. Soc., 1891, pp. 161–168.

— Notes on some aquatic Oligochaeta. Quart. J. Micr. Sci., 33: 187–218.

1892. Descriptions of three new specres of earthworms. Proc. Zool. Soc. Lond., 1892, pp. 136–152.

— Notes on two acanthodiilord earthworms from New Zealand. Quart. J. Micr. Sci., 33: 289–312.

— A new English genus of aquatic Oligochaeta (Sparganophilus) belonging to the family Rhinodrilidae. Quart. J. Micr. Sci., 34. 155–179.

— An earthworm from Ecuador (Rhinodrilus ccuadoriensis n.sp). Ann. Mag. Nat. Hist., (6) 9: 237–246.

— Note on the occurrence of a freshwater nemertine in England. Nature, 46: 611.

— British earthworms. Nature, 47: 102.

1893. Description of a new species of Moniligaster from India Quart. J. Micr. Sci., 34: 361–382.

— Note on a new species of the genus Nais Quart. J. Micr. Sci., 34: 383–386.

— The post larval stage of Arenicola marina. J. Mar. Biol. Assn., 3: 48–53.

1894. A description of the cerebral convolutions of the chimpanzee known as “Sally”; with notes on the convolutions of other chimpanzees, and two orangs. Quart. J. Micr. Sci., (2) 37: 47–86.

— Notes on a particularly abnormal vertebral column of the bullfiog; and on certain other variations in the Anuran column. Proc. Zool. Soc. Lond., 1894, pp. 477–481.

— On Benhamia caecifera n.sp. Proc. Phys. Soc. Edinb., pp. 31–35.

— Notes on the clitellum of the earthworm. Zool. Anz., 17: 53.

— On the blood of Magelona. Rept. Brit. Assn., Oxford, p. 696.

— On the classification of the Polychaeta Rept. Brit. Assn., Oxford, pp. 696–697.

1895. Male of Apus. Nature, 53. 175.

— Some Javan Perichaetidae. Ann. Mag. Nat. Hist., (6) 16: 40–50.

1896. On Kynotus cingulatus, a new species of earthworm from Imerina in Madagascar. Quart. J. Micr. Sci., 38: 445–463.

— The blood of Magelona. Quart. J. Micr. Sci., 39 1–18.

— Fission in Nemertines. Quart. J. Micr. Sci., 39: 19–32.

— The male of Apus cancriformis. Ann. Mag. Nat. Hist., 17. 120–122.

— Some earthworms from Celebes. Ann. Mag. Nat. Hist., (6) 18: 429–448.

— Earthworms and streamworms; review of Beddaid's monograph on the Oligochaeta. Nature, 53: 74–75.

— Archiannelida, Polychaeta, Myzostomaria. In the Cambridge Natural History, 2: 241–344.

1897. New species of Perichaeta from New Britain and elsewhere; with some remarks on certain diagnostic characters of the genus. J. Linn. Soc. Lond., 26: 198–225.

1899. Notes on the internal anatomy of Notornis Proc. Zool. Soc., Lond., 1899. pp. 88–96.

— Notes on the fourth skin of Notornis. Trans. N.Z. Inst., 31: 146–150.

— Notes on certain of the viscera of Notornis. Trans. N.Z. Inst., 31: 151–156.

Balanoglossus otagoensis n.sp. Quart. J. Micr. Sci., 42: 497–504.

— A re-examination of Hutton's types of New Zealand earthworm. Ann. Mag. Nat. Hist., (7) 3: 136–141; (also in Trans. N. Z. Inst., 31 156–163).

— Phosphorescent earthworms. Nature, 60: 591.

1900. Zoological results of trawling trials off the coast of Otago. Trans. N.Z. Inst., 32: 1–3.

Note on Cordyceps Sinclairii Berkeley. Trans. N.Z. Inst., 32: 4–8.

— Note on the occurrence of the genus Balanoglossus in New Zealand waters. Trans. N.Z. Inst., 32: 9–10.

— The structure of the rostellum in two new species of tapeworm from Apteryx. Quart. J. Micr. Sci., 43: 83–96.

1901. The Platyhelmia, Mesozoa, and Nemertini. In E. Ray Lankester's A Treatise on Zoology.

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1901. On the larynx of certain whales (Cogia, Balaenoptera, and Ziphius). Proc. Zool. Soc., Lond., 1901, 1: 278–300.

— On the anatomy of Cogia breviceps. Proc Zool. Soc., Lond, 1901, 2: 107–134.

Heteropleuron hectori, the New Zealand Lancelet. Quart. J. Micr. Sci., (2) 44: 273–280.

— The coelomic fluid in acanthodrilids. Quart. J. Micr. Sci., (2) 44: 565–590.

— On the New Zealand Lancelet. Trans. N.Z. Inst., 33: 120–122.

— An account of Acanthodrilus uliginosus Hutton. Trans. N.Z. Inst., 33: 122–129.

— On some earthworms from the islands from around New Zealand. Trans. N.Z. Inst., 33: 129–144.

1902. Note on the osteology of the short-nosed sperm-whale Proc. Zool. Soc., London, 1: 54–62.

— Note on an entire egg of a Moa, now in the Museum of the University of Otago. Trans. N.Z. Inst., 34: 149–151.

— An account of the external anatomy of a baby Rorqual (Blaenoptera rostrata). Trans. N.Z. Inst., 34: 151–155.

— Notes on Cogia breviceps, the lesser Sperm-whale Trans. N.Z. Inst., 34: 155–168.

1903. The geographical distribution of enrthworms and the palaeogeography of the Antarctic Region. Presidential address. Rep. Australas. Assn., 1902, pp. 319–343.

— On the eggs of the Moa. Ibis, 1903, pp. 632–634.

— On the remains of a gigantic fossil Cirripede from the Tertiary Rocks of New Zealand Geol. Mag., (4) x, (3), pp. 110–119.

— On some new species of aquatic Oligochaeta from New Zealand. Proc. Zool. Soc., Lond., 1903, ii. 202–232.

— Note on a neglected Tasmanian earthworm. Rep. Australas. Assn., 10: 383.

— On an new spectes of earthworm from Norfolk Island Trans. N.Z. Inst., 35: 273–274.

— On an earthworm from the Auckland Islands—Notiodrilus aucklandicus. Trans. N.Z. Inst., 35: 275–277.

— On the old and some new species of earthworms belonging to the genus Plagiochacta. Trans. N.Z. Inst., 35: 277–290.

1904. The sipunculids of New Zealand. Trans. N.Z. Inst., 36: 172–184.

— On a new species of leech (Hirudo antipodum) recently discovered in New Zealand. Trans. N.Z. Inst., 36: 185–192.

— A note on the Oligochaeta of the New Zealand lakes. Trans. N.Z. Inst., 36: 192–198.

— On an apparently new species of Regalecus (R. parkri). Trans. N.Z. Inst., 36: 198–200.

— On some edible and other new species of earthworms from the North Island of New Zealand. Proc. Zool. Soc., Lond., 1904, 2: 220–263.

— On some new species of the genus Phreodrilus. Quart J. Micr. Sci., 48: 271–298.

— On a new species of the genus Haplotaxis, with some remarks on the genital ducts of Oligochaeta. Quart. J. Micr. Sci., 48: 299–322.

— Phylum Annulata, in Hutton's Index Faunae Novae Zealandiae, pp. 276–285.

1905. Some earthworms from the North Island of New Zealand. Trans. N.Z. Inst., 37: 281–285.

— On the Ohgochaeta of the Southern Islands of the New Zealand Region. Trans. N.Z. Inst., 37: 285–297.

— Earthworms from the Kermadecs. Trans. N.Z. Inst., 37: 298–299.

— Note on the occurrence of the foraminiferan genus Ramulina in the New Zealand waters. Trans. N.Z. Inst., 37: 300.

— Further notes on the sipunculids of New Zealand. Trans. N.Z. Inst., 37: 301–308.

— The aquatic larva of the fly Ephydra. Trans. N.Z. Inst., 37: 308–312.

— Notes on some nudibranch molluses from New Zealand Trans. N.Z. Inst., 37: 312–320.

— Presidential address of the Otago Institute. Proc. N.Z. Inst., 37: 613–614.

1906. On a new species of Sarcophyllum from New Zealand. Zool. Anz., 31: 66–67.

— Carnivorous habits of the New Zealand Kea parrot. Nature, 73, no. 1902, p. 559.

— The olfactory sense in Apteryx. Nature, 74, no. 1914, pp. 222–223.

— Additional notes on the earthworms of the North Island of New Zealand. Trans. N.Z. Inst., 38: 239–245.

— On a large pterotrachaeid from the Pacific Ocean. Trans N.Z. Inst., 38: 245–248.

— An account of some earthworms from Little Barrier Island. Trans. N.Z. Inst., 38: 248–256.

1907. Notes on the flesh-eating propensity of the Kea (Nestor nolabilis). Trans. N.Z. Inst., 39: 71–89.

— New Zealand ctenophores. Trans. N.Z. Inst., 39: 138–143.

— Two new species of leech in New Zealand Trans. N.Z. Inst., 39: 180–192.

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1907. On a new species of pennatulid (Sarcophyllum bollonsi). Trans. N.Z. Inst., 39: 193–195.

— On the Oligochaeta from the Blue Lake, Mount Kosciusko. Rec. Austr. Mus., 6: 251–264.

— The evolution of the elephant. Proc. N.Z. Inst., 39: 547–548.

1908. An erioneous echinodermal identification. Ann. Mag. Nat. Hist., (8) 1: 104–108.

1909. Report on the Polychacta of the Sub-antarctic Islands of New Zealand. Sub-antarct. Isls. N.Z., 1: 236–250.

— Report on the Oligochaeta of the Sub-antarctic Islands of New Zealand Sub-antarct. Isls. N.Z., 1: 251–294.

— The echinoderms, other than holothurians, of the Sub-antarctic Islands of New Zealand. Sub-antarct. Isls. N.Z., 1: 295–305.

— Hydromedusae and Scyphomedusae from the Auckland and Campbell Islands. Subantarct. Isls. N.Z., 1: 306–311.

— Preliminary report on two Hirudinea from the Sub-antarctic Islands of New Zealand Sub-antarct. Isls. N.Z., 1: 372–376.

— Annelida and Sipunculoidea, New Zealand Government Trawling Expedition, 1907. Rec. Cant. Mus., 1: 71–82.

— Echinoderma, Scientific results of the New Zealand Government trawling expedition 1907. Rec. Cant. Mus., 1: 83–116.

1910. The discovery of Moa-remains on Stewart Island. Trans. N.Z. Inst., 42: 354–356.

1911. Stellerids and echinids from the Kermadec Islands. Trans. N.Z. Inst., 43: 140–163.

1912. Report on sundry invertebrates from the Kermadec Islands. Trans. N.Z. Inst., 44: 135–138.

1914. The nomenclature of the birds of New Zealand: being an abstract of Matthews and Iredale's “Reference List.” Trans N.Z. Inst., 46: 188–204.

1915. Notes on New Zealand polychaetes. Trans. N.Z. Inst., 47: 161–170.

— Preliminary report on the polychaetous annelids from the Kermadec Islands. Trans. N.Z. Inst., 47: 170–174.

— Oligochaeta from the Kermadec Islands Trans. N.Z. Inst., 47, 174–185.

— A remarkable case of bifurcation in Lumbricus rubellus. Trans. N.Z. Inst., 47: 185–188.

— On Lumbricillus macquariensis Benham Trans. N.Z. Inst., 47: 189–191.

— Report on the Polychaeta obtained by the F.I.S. Endeavour on the coasts of New South Wales, Victoria, Tasmania. and South Australia. Pt. 1 Fisheries Commonw. Australia, 3: 173–237.

1916. Report on gephyrcan Periapulus obtained by the F.I.S. Endeavour in Australian waters. Pt 2. iii. Fisheries Commonw. Australia, 4: 127–162.

— Notes on New Zealand Polychaeta (2). Trans. N.Z. Inst., 48: 386–396.

1917. Presidential address to the New Zealand Institute. Proc. N.Z. Inst., 49: 543–547.

1918. Presidential address to the New Zealand Institute. Proc. N.Z. Inst., 50: 338–342.

— The organisation of research in agriculture. Nature, Feb. 28, 1918.

— Biology in relation to agriculture. Cawthron Lecture, 1918.

1919. On the occurrence of two unusual blood-vessels in Hyla aurca. Trans. N.Z. Inst., 51: 30–34.

— Some earthworms from Stephen Island and d'Urville Island. Trans. N.Z. Inst., 51: 34–41.

1921. Polychaeta. Austral. Antarct Exped., 1911–1914. Sci. Reps. Ser. C (Zool. and Bot.), 6: 1–128.

1922. Oligochaeta of Macquarie Island. Sci. Rep. Austral. Antarct. Exped., 6, 1922, 38 pp.

Gephyrea inermia. Sci. Rep. Austral. Antarct. Exped, 6, 1922, 22 pp.

— On the occurrence of the Opah, or moonfish, in New Zealand waters. N.Z. J. Sci. Tech., 4, 316–318.

1927. Polychaeta. Brit. Antarct. “Terra Nova” Exped., 1910, Nat. Hist. Rep., Zool, 7: ii. pp. 47–182.

1928. On some alcyonariaus from New Zealand waters. Trans. Proc. N.Z. Inst., 59: 67–84.

1929. The pelagic Polychaeta. Brit. Antract. “Terra Nova” Exped., 1910, Nat. Hist. Rep., Zool., 7, iii, pp. 183–201.

— The inheritance of mental qualities. Presidential Address, Biology Section. 4th Science Congress of the New Zealand Institute. Proc. N.Z. Inst., 60: 27.

1930. Another specimen of the oar-fish (Regaleus). N.Z. J. Sci. Tech. 11: 428.

1932. Notes on Polychaeta two new species of the genus Goniada. Ann. Mag. Nat Hist., (10) 9: 553–567.

1934. The skeleton of a small Moa, Emeus huttoni Owen. Trans. Proc. Roy. Soc. N.Z., 64: 87–104.

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1935. A reptilian jaw from Kakanui, South Island, New Zealand. Trans. Proc. Roy. Soc. N.Z., 65: 232–238.

— The teeth of an extinct whale, Microcetus hectori n.sp. Trans. Proc. Roy. Soc. N.Z., 65: 239–243.

— George Malcolm Thomson (1848–1933). Trans. Proc. Roy. Soc. N.Z., 64: 413–421.

1937. Fossil Cetacea of New Zealand. ii. On Lophocephalus, a new genus of Zeuglodont Cetacea. Trans. Proc. Roy. Soc. N.Z., 67: 1–7.

— Fossil Cetacea of New Zealand. iii. The skull and other parts of the skeleton of Prosqualodon hamiltoni n.sp. Trans. Proc. Roy. Soc. N.Z., 67: 8–14.

— Fossil Cetacea of New Zealand. iv. Notes on some of the bones of Kekenodon onamata Hector. Trans. Proc. Roy. Soc. N.Z., 67: 15–20.

1939. Mauicetus: a fossil whale. Nature, 143: 765.

1941. Megascolides napierensis, a new species of earthworm. Trans. Proc. Roy. Soc. N.Z., 71: 27–31.

1942. Fossil Cetacea of New Zealand. v. Mauicetus, a genetic name substituted for Lophocephalus Benham. Trans. Proc Roy. Soc. N.Z., 71: 260–270.

Notoscolex equestris, an earthworm from the Poor Knights Islands. Trans. Proc. Roy Soc. N.Z., 72: 220–225.

— Octopodus Mollusca of new Zealand. i. The midget octopus of the coastal waters. Trans. Proc. Roy. Soc. N.Z., 72: 226–236.

1943. The octopodous Mollusca of New Zealand. ii. Trans. Proc. Roy. Soc. N.Z., 73: 53–57.

— The octopodous Mollusca of New Zealand. iii. The giant octopus Macrotopus maorum (Hutton)—in youth, adolescence and maturity. Trans. Proc. Roy. Soc. N.Z., 73: 139–153.

1944. The octopodous Mollusca of New Zealand. iv. Trans. Proc. Roy. Soc. N.Z., 73: 255–261.

— The octopodous Mollusca of New Zealand. v. Trans. Proc. Roy. Soc. N.Z., 74: 294–296.

1945. An earthworm with four penes, Conicodrilus genus novum. Trans. Proc. Roy. Soc. N.Z., 75: 23–28.

1947. Studies on earthworms. xlii. The occurrence of the genus Pheretima in New Zealand. Trans. Proc. Roy. Soc. N.Z., 76: 423–428.

1949. A yard-long earthworm, Notoscolex hakeaphilus. Trans. Proc. Roy. Soc. N.Z., 77: 346–350.

1950. On Cryptochaeta, a new genus of earthworms; and a nomenclatural muddle solved. Trans. Proc. Roy. Soc. N.Z., 78: 321–328.

— Polychaeta and Oligochaeta of the Auckland and Campbell Islands. Scientific Results of the N.Z. Sub-antarctic Exped., 1941–1945, Bul. 10.

Joint Authorship

1885 (with Lankester and Beck). The muscular and endoskeletal systems of Limulus and Scorpio. Trans. Zool. Soc. Lond., 11: 311–384.

1900 (with Thomson and Malcolm). Introduction to “An account of a large branchiate polynoid from New Zealand, Lepidonotus giganteus Kirk.” Proc. Zool. Soc. Lond., 1900, pp. 974–986.

1906 (with Dunbar, W. J.). On the skull of a young specimen of the ribbon-fish Regalecus. Proc. Zool. Soc. Lond., 2: 544–556.

1913 (with Cameron, Gladys). The nephridia of Perieodrilus ricardi and P. montanus. Trans. N.Z. Inst., 45: 191–198.

— (with Wilson, K). Footprints of the Moa Trans. N.Z. Inst., 45: 211–212.

David Miller.

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Bernard Cracroft Aston, C.B.E., F.R.S.N.Z., F.R.I.C.

By the death of B. C. Aston on May 31, 1951, New Zealand has lost another of her rapidly thinning band of pioneer scientists. Agricultural Chemistry was virtually a one-man job when Aston was appointed Chemist to the Department of Agriculture in 1899. To-day dozens of chemists in various institutios throughout New Zealand are engaged in its many ramifications.

Aston was born at Beckenham, Kent, England, in 1871, son of M. Aston, of a family well known in the Home Counties. He was educated at Christchurch Boys' High School, Dunedin Technical College, and Otago University. At Otago University, Aston was considerably influenced by Professor Black, and was also a contemporary and friend of J. W. Mellor, the celebrated ceramic chemist. Aston's first chemical appointment was as chemist to a cement company, but he was soon afterwards appointed a consulting analyst to the Government, with accommodation at the University.

On his appointment as Chemist to the Department of Agriculture, in 1899, Aston moved to Wellington, which thereafter remained his home. His earliest investigations were largely influenced by the researches of Chief Veterinarian J. A. Gilruth on ailments of livestock in New Zealand, particularly in relation to poisonous plants. Together with Easterfield, he isolated and described “tutin”, the principal toxic glucoside of the New Zealand species of Coriaria. Other native poisonous plants investigated included the Karaka, Nagio, Pimelia, Rangiora, and Pukatea, from the last of which a series of alkaloids was isolated.

While pursuing these researches Aston was also actively building up a service laboratory for the Department and in his first year personally analysed and reported on some ninety-three samples of the most varied nature. He also introduced and tested the use of Dyer's method for available plant food—phosphate and potash—in soils. In this and the following few years Aston laid the foundations of most lines of agricultural analysis practised in New Zealand during the next thirty-five years.

In 1903 Aston visited Britain, Canada, U.S.A. and Australia. He made other trips abroad in 1914 and 1926. The years 1906 to 1909 are especially noteworthy for the expansion of the Chemical Laboratory under Aston, both in personnel and equipment, and in services performed and lines of research inaugurated.

During this period Aston published important papers concerned with the objectives and methods of field experimentation, the need for a soil survey, and the possibility of establishing a beet sugar industry in New Zealand. In his annual reports is recorded a wealth of data on the soils of New Zealand, accumulated from flying surveys and investigations of agricultural problems. Results of many field experiments with fertilizers and crops, undertaken jointly with district officers of the Department are presented, and doubtless influenced subsequent fertilizer practice in the areas concerned.

Following much exploratory work, Aston established the first district testing laboratory for export butter in New Plymouth in 1908. Such laboratories are now an essential feature of the dairy industry.

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The retrenchement of 1910 and the repercussions of the First World War permitted little more than routine servicing during the second decade of the agricultural chemical laboratory's establishment. Towards the end of this period a number of investigations were commenced, some of which showed rapid development.

Aston was greatly interested in native plant products and an acute shortage of imported dyes provided him with an opportunity to study the dyeing properties of the native coprosmas. His findings have since been of considerable interest to home dyers.

The testing of wheat for miling and baking qualities, and of dairy products and requisites were two lines of chemical service to agricultural industries whose early development in Aston's laboratory was followed by the setting up of separate organisationis—the Wheat Research and Dairy Research Institutes.

During the third decade, investigations in Aston's laboratory became more specialized along lines of plant and animal nutrition and their dependence on soil conditions.

In particular, the problem of “bush sickness” in ruminants depastured on the pumice lands was the object of research, both by field experimentation and by chemical analyses. It proved unexpectedly complex and baffling in the light of existing knowledge. Practical remedies were however found in certain iron compounds used either as medicaments or licks. Although Aston put forward the hypothesis that iron deficiency in the pasture and the animal was the cause of the disease, he realised that many anomalies existed. He investigated alternative possibilities such as the animal's ability to utilise iron in certain states of combination only, or that a contaminating element in the crude iron compounds used might be responsible for the curative effect.

In 1931 limonite was found by Aston to be a practical remedy when used as a lick, but was soon shown to vary greatly in curative value. Shortly afterwards Filmer and Underwood of Western Australia began the researches on the fractionation of limonite which led ultimately to the identification of cobalt as the curative element.

When in 1926 Aston visited Britain, he was invited to deliver a paper on “Bush Sickness” to the British Association for the Advancement of Science. Here and subsequently at the Rowett Institute, he and his work met with an enthusiastic reception from Sir John Orr. Largely as a result of this contact, the Empire Marketing Board in the following year decided to make an annual grant to the New Zealand Government for research on problems of animal nutrition, Aston being appointed to direct the use of the funds During the next four or five years, until the Empire Marketing Board scheme was discontinued as a result of the financial depression, an extensive programme of work on aspects of the mineral contents of pasture in relation to the nutrition of grazing livestock was carried out in Aston's laboratory. Phosphorus, calcium, nitrogen, iodine, iron and managanese were the elements on which greatest emphasis was placed, but many others were determined during investigations of various suspected deficiency diseases. Most use has perhaps subsequently been made of the data from the iodine and phosphorus surveys, but the Quarterly Reports (unpublished) to the Empire Marketing Board contain a wealth of material for future reference.

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Aston's greatest contribution to agricultural science is probably his recognition and demonstration that a widespread ailment of livestock, so severe as to make pastoral farming precarious if not impossible, was due to deficiency of a “minor” mineral element, not itself required, at least in comparable amount, for pasture production. The further demonstration that limonite as a lick could cure or prevent the disease was the primary stimulus to the great and successful development of huge tracts of pumice lands into profitable dairy and sheep farms which has since taken place. The subsequent identification by other workers of cobalt as the trace element concerned has had far-reaching effects in nutritional theory and has further simplified and cheapened the practical remedy.

Seldom has the life of a chemist been so closely identified with the institution he created as in Aston's case. The laboratory was in a large measure himself. His colleagues and those that worked under him seldom failed to be infected by his enthusiasm. Many of the chemists now in senior positions in research institutions or chemical laboratories in New Zealand received their early training under Aston. Others have taken up similar positions abroad. His influence on agricultural chemistry has thus been far reaching.

As a man, Aston was direct in approach and staunch in friendship. Foolishness was apt to arouse his ire, but intelligent interest and honest work received ready recognition.

None could have been a more agreeable companion for a tramp or an expedition, and few had command of so much bush lore and knowledge of general natural history. Aston had a great love for New Zealand—mountains, farmlands, plants and animals—and the few honours he received were far from adequate recognition of a life spent in her service.


Although Aston was by profession a chemist, he was also a botanist, and over a period of about forty years an exceedingly active collector of plants. With more than an average amount of physical strength, he was able to cross mountain ranges and ascend their highest peaks with little apparent fatigue. His botanical investigations in Otago began about 1890 and continued until his removal to Wellington nine years later. He explored many parts of the province, some as far away from his place of residence, Dunedin, as Clifden and Colac Bay, and he visited Stewart Island.

Soon after his arrival in Wellington in 1899 he became interested in the plants of the Tararua Range which, at that time, was unknown botanically. In four years he made ten ascents of peaks in the southern portion of the range. His crossing from the Hutt Valley to the Otaki River in December, 1907, with A. M. Jones and W. B. Aston was among the earliest made. His route led over the Quoin, Alpha and Hector summits.

Aston next turned his attention to the southern parts of New Zealand. In November, 1907, he was a member of the expedition to the Subantarctic Islands organized by the Philosophical Institute of Canterbury. The party left the Bluff in the Government steamer Hinemoa on November 14. A day was spent on the Snares, where soil samples and plants were collected. At the Auekland Islands most of the party, inclnding Aston. landed at Camp Cove in Carnley Harbour,

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The others went on to Campbell Island. At the Auckland Islands, subsidiary camps were established at the head of North Arm and on Adams Island. Altogether thirteen days were spent at the Auckland Islands, during which time Aston managed to examine a variety of stations within about twelve miles of the main camp, and soil samples and plants were collected on all these excursions. Port Ross and disappointment Island were visited before the Hinemoa returned to New Zealand. Aston paid a second visit to the Auckland Islands in January, 1909. In Fiordland, Aston visited Doubtful Sound in 1909 and Milford Sound in 1911.

During four years beginning January, 1911, Aston made six excursions to the Ruahine-Kaimanawa mountain system. Some of these trips were very strenuous. In January, 1911, he and J. S. Tennant explored the Kaimanawa Mountains which, up to that time, were unknown botanically. They ascended Waipahi, 5,200 feet, and Korokarenga, 5,300 feet. As if this were not enough mountaineering for one week, they crossed the plain on the west side of the Kaimanawas and climbed to the top of Ngauruhoe, 7,500 feet.

In January, 1914, with R. A. Wilson and F. K. Hutchinson, Aston attempted to follow the route across the Ruahine Range between Waipawa and Mokai Patea, in the Rangitikei basin, that Colenso travelled over six times between 1847 and 1852. Aston's party followed near Colenso's track on the eastern side of the range, but on the western side it came down a different spur from that used by Colenso. One of the longest journeys made by Aston, accompanied by Wilson and Hutchinson, began on the Rangitikei River near Taihape in December, 1914. From here they ascended Aorangi, 4,250 feet, thence they continued to the northern end of the Ruahines and climbed the Kaimanawas to the summit of Makarako, 5,700 feet, and then returned to the Rangitikei.

Three trips (1913, 1915, 1916) were made by Aston to Tarawera Mountain, which violently erupted in 1886 destroying all vegetation from the summit practically to the base. Here was an opportunity to study the regrowth of plants in completely sterilized soil. Summarizing the results of this investigation (Journal of Ecology, vol. 4), Aston says: “Practically the whole of the plants found on the mountain are those the seed of which is spread … by means of wind and water and birds.” Aston's record will be indispensible for future botanists wishing to follow the changes in the vegetation since Aston's visit.

Aston now moved his botanical operations to the South Island and made three long trips in the Marlborough district. He explored the Flaxbourne, Medway, Ure, Nidd and Swale valleys, all in limestone country. On one of these excursions, December, 1915, he climbed to about 6,000 feet up the slopes of Tapuaenuku. Two months later, he, with J. A. Thomson, H. Hamilton and A. F. O'Donoghue, succeeded in reaching the summit of Tapuaenuku, 9,465 feet.

After this achievement, Aston's botanical excursions became less frequent and less strenuous Aston always carried a notebook in which he entered the names of the species as he came across them, giving a rough idea of locality and height above sea-level. Sometimes these notes were transcribed on to sheets of paper, and in the case of the Tararua, Ruahine and Tarawera expeditions, the results were published. A letter which Aston wrote to a friend in England about his ascent of Tapuaenuku was published in the Kew Bulletin. But most of Aston's notes are unpublished and likely to remain so, because they are incomplete

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Bernard Cracroft Aston, C.B.E., F.R.S.N.Z., F.R.I.C.

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as descriptions of plant associations. Only Aston could fill in the details of what he saw. In the course of his travels Aston collected sufficient specimens to form a moderate-sized herbarium, and this he presented to the Dominion Museum in 1922. Aston is commemorated in the names of species belonging to the genera Coprosma, Gentiana, Muehlenbeckia, Myosotis, Poa and Hebe.

Aston was a member of the New Zealand Institute and Royal Society of New Zealand for fifty-five years, first of the Otago Institute, 1896–1899, then of the Wellington Philosophical Society (later the Wellington branch of the Royal Society), 1899–1951. He served on the councils of both these societies. He was one of the four Government Representatives on the Board of Governors of the New Zealand Institute and the Council of the Royal Society, 1915–1951. He was secretary of the New Zealand Institute 1909–1931. Aston was a valued and active member of the Council. He will be remembered as one of the founders of the Society's Endowment Fund, and as its chief guardian. He thought that eventually the fund might reach such a size as would enable the Society to have its own building. Aston represented the Royal Society on the Tongariro National Park Board, 1926–1931, and on the Dominion Museum Management Committee, 1937.

Honours in science came to Aston through the Royal Society, which he served so well and so long, for his research work in chemistry and botany. He was one of the twenty Original Fellows of the New Zealand Institute gazetted on November 20, 1919. In 1925 he was awarded the Hector medal and prize for “the investigation of New Zealand chemical problems.” Lastly, in 1926–1927, he was elected President of the New Zealand Institute, the highest honour the Institute could confer, and was Vice-President 1930–1935. In 1948 the honour Commander of the Most Excellent Order of the British Empire (C.B.E.) was conferred on him by His Majesty the King.

Aston not only collected plant specimens for his herbarium, but he was an enthusiastic grower of native plants. On his property in Espin Crescent, Wellington, he formed a garden that contained a large number of species, and he lived long enough to see the trees grow to a large size.

Aston had a genial and generous nature which brought him many friends and great respect. He readily gave to inquirers the benefit of his wide scientific experience. His interests, however, were much wider than his pursuit of botanical and chemical knowledge. He was specially interested in the preservation of native plants and plant associations, and of our native birds. He served on the Council of the Forest and Bird Protection Society of New Zealand and was its President for several years. When the Dunedin Naturalists' Field Club was re-instituted in 1894, Aston was its secretary, and it was largely through his energy and enthusiasm that the club became a vigorous institution. Principally owing to his exertions, a very complete and accurate catalogue of the plants of Dunedin and its neighbourhood was published in 1896.

Aston's published work, other than on chemistry, included ten botanical papers contributed to the New Zealand Institute. All these were the results of his explorations on the mountains and lowlands of the North Island, Marlborough and Otago. Perhaps his most important paper was his description of the vegetation of Mount Tarawera, but his papers on the Tararua and Ruahine

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mountain ranges were pioneers in these fields. Numerous articles from Aston's pen appeared in various popular journals. Amongst them was a short series on New Zealand trees and other plants which appeared in Forest and Bird during 1936–1938. Altogether Aston's contributions to the advancement of botanical science in New Zealand are considerable and important.