The New Zealand Forms of Pterostylis R. Br.
[Read before the Auckland Institute, June 18, 1947; received by the Editor, July 1, 1947; issued separately, February, 1949.]
The pollination of the genus is described by Cheeseman (Trans. N.Z. Inst., 5, 1873, 352). Two Australian species (Pt. crypta Nich. and Pt. celans Rupp) are now known to be cleistogamous (Nicholls, Vict. Natr., 61, 1945, 207 t), and the endemic Pt. humilis Rog. is self-fertile without being cleistogamous (Hatch, Trans. R.S.N.Z., 75, 1945, 369). Vegetative reproduction, by the formation of additional tubers, occurs in most species. Pt. trullifolia Hook. f. is unique in developing a special non-flowering form devoted to vegetative increase.
Origin and evolution. The evidence (such as may be deduced from a study of comparative anatomy and of the present distribution of the various groups and species) suggests that Pterostylis originated in the south, differentiating along two arcs, through Tasmania and Australia to New Guinea, and through New Zealand to the Chathams and New Caledonia. In addition, Australian species have been carried from time to time by winds both to New Zealand and to New Caledonia. The genus appears to have arisen from a form with a basal rosette and a many-flowered terminal raceme, represented in recent times by the Australian rufa group. From simple plants of this type, by a process of suppression and adaptation all the known forms could have evolved. It is believed that the foliata group consists of the recent representatives of the form which gave rise to the australis group in New Zealand and some other groups in Australia. The remainder of the New Zealand species are Australian forms, either of fairly recent arrival such as Pt. nana, nutans, mutica and barbata; or old-established, locally differentiated forms such as Pt. humilis, furcata and trullifolia.
The groups and species are arranged in what the writer considers to be their probable phylogenetic sequence. The descriptions and illustrations are in most cases drawn up from living material examined by the writer. In a couple of instances photographs of living plants were used to make the illustrations and the dissections taken from dried material. The only species entirely drawn up from dried material is Pt. areolata Petr.
The writer is very much indebted to Dr. W. R. B. Oliver, Dr. H. H. Allan, Rev. H. M. R. Rupp, Mr. George Simpson, Miss B. E. G. Molesworth, Mr. Cedric Smith and Mr. F. W. Bartlett, who provided a vast amount of material and many invaluable notes.
Pterostylis R. Br.
Glabrous or puberulous terrestrial herbs, with small globular tubers and slender succulent roots, Leaves orbicular to narrow linear-
lanceolate, sessile or petiolate, basal or cauline or both, sheathing at the base. Flowers solitary or several in a terminal raceme, varying from pale green to deep red and more or less striate. Dorsal sepal and petals conniving to form an erect or incurved hood over the column. Dorsal sepal obtuse to filiform-caudate. Petals variously falcate, subacute to acuminate, as long as or shorter than the dorsal sepal. Lateral sepals erect or deflexed, conjoined at the base, the free lobes obtuse, linear, subulate or filiform-caudate. Labellum filiformterete or variously laminate, glabrous to densely plumose, mounted on an articulate sensitive claw and with a basal appendage which may be variously penicillate or entire. Column erect or inclined, free or partly adnate to the dorsal sepal. Stigma near the centre of the column, linear-lanceolate to globose, 2-celled, viscid. Column-wings variously lobed, the lower usually larger than the upper. Anther terminal, more or less horizontal, 2-celled. Pollinia 4, free, pollen granular or powdery, caudicle absent. Rostellum erect between the bases of the anther-cells.
A genus of some 67 species occurring throughout Tasmania, Australia, New Guinea, New Zealand, New Caledonia, and the Chathams.
Key to the New Zealand Species.
|1. Labellum filiform-terete, plumose||barbata|
|Labellum laminate, glabrous or pubescent||2|
|2. Labellar appendage entire, flowers several||mutica|
|Labellar appendage penicillate, flowers normally solitary||3|
|3. Labellum acuminate||4|
|4. Leaves in a rosette, flower conspicuously drooping||nutans|
|Rosette only present in the juvenile, mature leaves varying from orbicular-petiolate to bracteate-sessile||6|
|6. Leaves 2–3, subrosulate, ovate||venosa|
|Leaves numerous, polymorphous||7|
|7. Lateral sepals with an emarginate central lobe||8|
|Lateral sepals with an acute, unlobed sinus||trullifolia alobula|
|8. Petiolate leaves with the veining conspicuously embossed||trullifolia gracilis|
|Petiolate leaves with a smooth upper surface||trullifolia rubella|
|5. Upper third of the labellum symmetrically twisted, entire||9|
|Labellum-tip flat, entire or emarginate||10|
|Labellum-tip unevenly constricted||11|
|9. Flower large, leaves subrosulate, more or less ovate||furcata typioa|
|Flower small, leaves cauline, linear-lanceolate||furcata linearis|
|10. Leaves in a rosette, lateral sepals with an inflexed lobe||nana|
|Leaves strictly cauline, lateral sepals with an acute sinus||12|
|Leaves both subrosulate and cauline||13|
|12. Plant very slender, sepalar caudae very short||graminea|
|Plant robust, sepalar caudae very long||14|
|14. Dorsal cauda suberect, lateral caudae spreading, erect||banksii typica|
|Dorsal cauda incurved, lateral caudae strongly recurved||banksii patens|
|13. Flower large, dorsal sepal filiform-caudate, incurved||oliveri|
|Flower small, dorsal sepal acuminate, horizontal||areolata|
|11. Leaves rosulate, stigma globose||humilis|
|Leaves subrosulate, stigma linear||foliata|
|Leaves strictly cauline||15|
|15. Labellum reddish, lateral sepals caudate, erect||australis|
|Labellum greenish, lateral sepals acuminate||montana typioa|
|Labellum brownish, lateral sepals caudate, spreading||montana rubrioaulis|
1. Pterostylis barbata Ldl., Swam River App., 1839, 53.
Pt. squamata Hook. f. (not of R. Br.).
Up to 25 cm. high. Leaves numerous, subrosulate or rising up the stem, up to 3 cm. long by 5 mm. broad, sessile or shortly petiolate, oblong- to narrow-lanceolate, acute or acuminate. Stem bracts 2–3 below the similar floral bract, loosely sheathing, acute. Flower solitary, erect, up to 4 cm. high, pale green with dark green veins. Dorsal sepal acuminate, longer than the erect petals, the tip incurved. Lateral sepals narrow-linear, connate for half their length and deflexed against the ovary. Labellum exserted, pendulous filiformterete, plumose with long golden hairs and tipped with a large, variously lobed purple callus. Column slender, stigma large, obovate. Column-wings with recurved, narrow-subulate upper lobes much exceeding the anther and narrow, densely ciliate, forward spreading lower lobes.
Distribution. Australia—all States except Queensland. New Zealand—2, 3a, b, c, occasional throughout; 7, Kaitoke, B. C. Aston; Wallaceville, 10, 1946, H. Hodgson; Day's Bay, E. H. Atkinson.
Flowers October–November, sea-level–1,000 ft., solitary or in small groups in scrub or along forest margins, rare. Closely related to the Westralian Pt. turfosa Ldl., with which it forms an isolated section of the genus, believed to be an offshoot of the ancestral rufa group, very far back. Probably windborne originally across the Tasman.
2. Pterostylis mutica R. Br., Prodr., 1810, 328.
Pt. tristis Col., Trans. N.Z. Inst., 18, 1886, 271.
Up to 10 cm. high. Leaves 1–8, dsensely rosulate, acute, petiolate or the upper few sessile, up to 2 cm. long by 6 mm. broad, semi-succulent or membraneous, the lower surface silvery, glabrous, the upper surface glaucous-yellow, the veins conspicuously embossed. Stem bracts 1–6, closely sheathing, erect, acute, repand, brownish-grey. Flowers 1–5 in a terminal raceme, brownish-green, up to 5 mm. high. Dorsal sepal cumulate, obtuse. Petals finely ciliate. Lateral sepals broad, obtuse, deflexed, conjoined almost to the tips. Labellum broad, obtuse, emarginate or entire. Appendage entire, the tip turned inwards. Column erect, stigma high, short and broad. Column-wings with the upper lobes reduced to vestiges, the lower lobes bluntly triangular, turned out toward the petals. The stem elongates considerably with maturity. Australian specimens frequently reach a height of 30 cm. and may produce as many as 10 flowers.
Distribution. Australia—common throughout. New Zealand—5, National Park, 10, 1921, H. B. Matthews; 12, 1920, H. Carse; Waiouru, 12, 1944, E. D. Hatch; 6, Waipawa River, 1885, H. Hill; 12a, 13, 15, occasional throughout; 16, Stewart Island, D. Poppelwell.
Flowers October–January, sea-level–3-000 ft., solitary or small colonies in grass, rather rare. Closely related to the Australian Pt. cycnocephala R. Fitzg., which differs mainly in the labellar appendage being turned outwards. Probably windborne originally across the Tasman.
3. Pterostylis nutans R. Br., Prodr., 1810, 327.
Pt. matthewsii Cheesmn., Trans. N.Z. Inst., 47, 1915, 46.
A compound species with 2 jordanons in Australia. Only the typical form occurs in New Zealand.
Glabrous or occasionally puberulous, up to 15 cm. high. Leaves 1–8, rosulate, petiolate, up to 5 cm. long by 3 cm. broad, oblong to oblong-ovate, acute. Stem bracts 2–3 below the similar floral bract, closely sheathing, acute. Flower solitary, drooping conspicuously, green with faint striae. Dorsal sepal incurved, acute, slightly longer than the petals. Lateral sepals with a wide sinus, the caudae narrow-linear, spreading and shortly exceeding the galea. Labellum broad-acuminate, recurved along its whole length, greenish-red, more or less pubescent. Column incurved, the linear stigma very long. Column-wings with no upper lobes, the lower broadly triangular, ciliate, obtuse.
Distribution. Australia—abundant throughout. New Zealand—2, Kaitaia, 8, 1912, H. B. Matthews; 3a, Whangaparoa, 10, 1942, L. M. Cranwell.
Flowers August–October, sea-level–1,000 ft., small colonies in scrub, very rare. Related to the Australian-New Caledonian Pt. curta R. Br. The group, which has affinities and probably a common origin with the foliata group, is well developed in Australia and contains some half a dozen other species. Probably windborne originally across the Tasman.
4. Pterostylis nana R. Br., Prodr., 1810, 327.
Pt. puberula Hook. f., Fl. Nov. Zel., 1, 1853, 249.
Up to 15 cm. high. Leaves 3–8, broadly petiolate, ovate, acute, up to 15 mm. long by 7 mm. broad. Stem bracts 1–3 below the similar floral bract, acute, closely sheathing, sometimes foliaceous. Flower solitary, erect, golden-green, up to 3 cm. high. Dorsal sepal shorter than the petals, obtuse or shortly acute, the tip horizontal. Lateral sepals with a very wide shallow sinus, with a small inflexed lobe. Caudae erect, much exceeding the galea. Labellum oblong, pinkish, sometimes pubescent, the obtuse tip slightly deflexed. Appendage short, trifid. Column erect, stigma elliptical, rather long. Column-wings with acuminate upper lobes shorter than the anther. Lower lobes broadly oblong, with inturned ciliate margins.
Distribution. Australia—all States except Queensland, abundant. New Zealand—3a, Silverdale, 10, 1946, F. W. Bartlett (originally common on the scrub-covered gumlands of Auckland's western suburbs and north to the Kaipara Harbour, this species has been wiped out by the advance of civilisation); 3b, c, occasional; 7, Silverstream, 10, 1944, L. B. Moore; 10, Westport, W. Townson, Collingwood, T. F. Cheeseman.
Flowers September–October, sea-level–1,000 ft., large colonies in scrub. Related to the Australian Pt. pyramidalis Ldl., the group contains 6 or 7 other species, widespread throughout Australia. Probably windborne originally across the Tasman.
5. Pterostylis foliata Hook. f., Fl. Nov. Zel., 1, 1853, 249.
Pt. gracilis Nich., Vict. Natr., 43, 1927, 324 t.
Up to 40 cm. high. Leaves 1–6, subrosulate or spreading up the stem, sessile or shortly petiolate, up to 5 cm. long by 1 cm. broad, elliptic-to linear-oblong or almost orbicular, acute or obtuse, membraneous or semi-succulent. Floral bract foliaceous, lanceolate, acute. Flower solitary, up to 25 mm. high. Dorsal sepal acuminate, longer than the petals, the tip horizontal or slightly incurved. Lateral sepals filiform-caudate, exceeding the galea by as much as 8 mm. Labellum linear-oblong, recurved and channelled, the tip unevenly constricted. Column typical, stigma linear-lanceolate. Column-wings with narrow subulate upper lobes as high as the anther. Lower lobes broad, obtuse, hardly reaching the stigma. With maturity the upper leaves of the rosette tend to scatter up the stem, which elongates considerably above the floral bract.
Distribution. Australia—Tasmania. Victoria, not common. New Zealand—4, Lake Tutira, Guthrie-Smith; 5, Rotorua, 1, 1922, K. W. Allison, 11, 1946, F. A. Springhall; 7, not uncommon throughout the Tararua and Ruahine Ranges; 9, “Marlborough,” T. Kirk, J. H. MacMahon; 12a, Ashburton, H. H. Allan; 15, occasional in the Dunedin District.
Flowers November–January, sea-level–3,000 ft., small colonies near swamps, in scrub and along forest margins. Nowhere abundant. The foliata group consists of 4 rather closely related species (Pt. foliata—N.Z., Tasm., Vict.; Pt. hildae Nich.—N.S.W., Queensld., N. Caledonia; Pt. neocaledonica and bureauviana Schltr.—N. Caledonia) which show signs of having evolved along 2 arcs—through Tasmania-S.E. Australia-Queensland-New Caledonia on the west, developing from foliata through hildae to neocaledonica, and through New Zealand-New Caledonia on the east, developing from foliata to bureauviana.
The group is a local development with no parallel in the other areas where the genus occurs. It consists of 8 jordanons, all of which have been known to hybridise. In some isolated areas, particularly in Stewart Island for instance, the hybrid population far exceeds the numbers of true species. Another polymorphous area is the Waimakariri River basin. It has been suggested that the group arose from the ancestral foliata stock, spreading north and east, differentiating largely by epharmony until it is at present abundant in one form or another throughout New Zealand and the Chathams, extending from sea-level to the upper limits of the subalpine scrub. All the forms (with the occasional exception of Pt. graminea, a highly specialised form) have distinct rosulate juveniles, which recapitulation is but further proof that the group originated as an offshoot of a strictly rosulate form. All the members of this group occasionally occur in a very depauperated condition, the leaves being much reduced and the plant practically all flower.
6. Pterostylis australis Hook. f., Fl. Nov. Zel., 1, 1853, 248.
Pt. banksii var. B. Hook. f., Handbk. N.Z. Fl., 1864, 268.
Up to 20 cm. high. Leaves 1–6, narrow- to broad-linear, acuminate, up to 10cm. long by 2 cm. broad. Floral bract foliaceous. Flower up to 3 cm. high, rather narrow. Dorsal sepal shortly caudate, longer than the petals, the tip horizontal or suberect. Lateral sepals shortly caudate, hardly exceeding the galea. Labellum linear-oblong, pale red, the tip unevenly constricted. Column typical, stigma elliptical. Column-wings with acuminate upper lobes as high as the anther. Lower lobes rounded, narrow-oblong.
Distribution. Endemic—5, not uncommon in Nothofagus forests about the base of Mount Ruapehu, 12, 1944, E. D. Hatch; 7, occasional throughout the Tararua and Ruahine Ranges; 12, 15, abundant in montane and subalpine areas; 16, not uncommon throughout Stewart Island, 11, 1946, C. Smith; also abundant in the Chatham Islands.
Flowers December–January, sea-level–4,500 ft., scattered on the forest floor, in tussock or in scrub.
7. Pterostylis montana Hatch spec. nov.
Pt. australis affinis, subsimilis. Circiter 15 cm. alta. Folia 1–5, patula, linearo-lanceolata, acuminata, saepe repanda. Flos brevis. Sepalum dorsale acuminatum, apex horizontals. Sepala lateralia acuminata, lobae breves. Labellum viride, recurvum, apex impariter constringit. Columna Pt. australis similis, superioribus lobis acuminatis, inferioribus lobis angusto-oblongis incurvis.
A compound species of 2 jordanons. Cheeseman included them in Pt. graminea, but they differ from Hooker's species in having constricted labella and spreading leaves (characters which incidentally they share with Pt. australis), whereas graminea sens. strict. has a symmetrical labellum and erect leaves.
(a) Pt. montana var. typica Hatch.
Pt. graminea Cheesmn. in part (not of Hook. f.).
Up to 15 cm. high. Leaves 1–5, linear-lanceolate, acuminate, often repand, up to 4 cm. long by 4 mm. broad. Floral bract foliaceous.
Flower solitary, up to 3 cm. high. Dorsal sepal acuminate, longer than the petals, the tip horizontal. Lateral sepals acuminate, the lobes shortly exceeding the galea. Labellum green, recurved, the tip unevenly constricted. Column typical, stigma prominent, elliptical. Column-wings with acuminate upper lobes as high as the anther. Lower lobes incurved, narrow-oblong. With maturity the lateral sepals tend to fall away from the galea.
Distribution. Endemic—5, not uncommon about the Nothofagus forests on Mount Ruapehu, Matthews, Hatch; 15, Lake Manapouri, 1, 1946, Geo. Simpson; 16, abundant throughout Stewart Island, 12, 1946, C. Smith.
Flowers November–January, sea-level–4,500 ft., scattered on the forest floor, common. Probably derived from Pt. australis. Almost certainly confused with graminea sens. strict., and probably abundant in most subalpine areas in the North and South Islands, but has so far only been definitely recorded from the localities given above. Holotype in Herb. Hatch, No. 564, Halfmoon Bay, Stewart Island, 11, 1946, C. Smith. The accompanying illustration can be regarded as the hypotype of the species.
(b) Pt. montana var. rubricaulis (Matth.) Hatch comb. nov.
Pt. graminea var. rubricaulis Matth. ex Cheesmn., Man. N.Z. Fl., 1925, 351.
Up to 15 cm. high. Leaves 1–4, distant, spreading, linearlanceolate, acuminate, up to 6 cm. long by 8 mm. broad. Floral bract foliaceous. Flower solitary or rarely 2, up to 3 cm. high. Dorsal sepal acuminate, longer than the petals, the tip suberect. Lateral filiformcaudate, spreading, the lobes much exceeding the galea. Labellum recurved, brownish, the tip unevenly constricted. Appendage short, stout, much divided. Column typical, stigma linear-lanceolate, rather broader than the column and slightly swollen at its lower end. Columnwings with the upper lobes subulate, suberect, as high as or shorter than the anther. Lower lobes oblong, twisted, the inner margins finely pubescent. Anther rather short.
Distribution. Endemic—3a, abundant throughout the Waitakere Ranges and on the east coast as far north as the Whangaparoa Peninsula; 3b, occasional throughout the Hunua Ranges.
Flowers July–September, sea-level–1,000 ft., scattered on the forest floor. Probably derived from montana typica. The accompanying illustration can be regarded as the hypotype of the variety.
8. Pterostylis oliveri Petr., Trans. N.Z. Inst., 26, 1894, 270.
Up to 30 cm. high. Leaves 1–6, up to 10 cm. long by 4 cm. broad, glaucous, oblong-ovate to -lanceolate, acute, conspicuously reticulated. Floral bract foliaceous, acuminate. Flower solitary or rarely 2, up to 6 cm. high. Dorsal sepal strongly incurved, the tip of the very long cauda frequently reaching the ovary. Lateral sepals with a shallow sinus, the caudae very long, erect or recurved. Petals narrow-falcate, the acuminate tips incurved. Labellum rather long, membraneous, recurved abruptly at the tip. Column typical, stigma linear-lanceolate. Column-wings with the upper lobes acuminate, slightly higher than the anther. Lower lobes incurved, narrow and very long, often descending below the stigma. The leaves vary from rosulate to cauline in developing to maturity. The juvenile is a lax rosette of 2–4 elliptical leaves on rather long slender petioles.
Distribution. Endemic–11, common about the headwaters of the Otira and Waimakariri River, particularly so in the Arthur's Pass area; 12a, Bealey River, T. Kirk.
Flowers December–January, 1,000–4,000 ft., scattered in scrub and along forest margins. Something of an intermediate form between Pt. banksii and the foliata group.
Holotype in the Dominion Museum, Wellington, Kelly's Creek, Otira River, 1,100 ft., 1, 1892, D. Petrie.
9. Pterostylis areolata Petr., Trans. N.Z. Inst., 50, 1918, 210.
Up to 18 cm. high. Leaves 1–4, up to 5 cm. long by 1 cm. broad, oblong- to elliptic-lanceolate, acute, conspicuously reticulated. Floral bract foliaceous, elliptic, acute. Flower proportionately large, green with reddish striae, up to 4 cm. high. Dorsal sepal acuminate, a little longer than the petals. Lateral sepals with a narrow sinus, the lobes shorter than the galea. Labellum narrow-linear, flat or recurved. Column as in Pt. oliveri, but the lower lobes of the column-wings rather shorter. The leaves go through similar developmental stages as do those of Pt. oliveri.
Distribution. Endemic—9, Awatere Valley, L. Cockayne; 12a, Waimakariri Valley, T. Kirk.
Flowers December–January, 2,500–3,000 ft. Probably derived from Pt. oliveri, but fresh material is needed before its exact relationships can be ascertained. In the absence of living material, the accompanying illustration has been copied exactly from the Holotype.
Holotype in the Dominion Museum, Wellington. Base of Shingle Peak, Awatere Valley, L. Cockayne.
10. Pterostylis banksii R. Br., ex A. Cunn., Bot. Mag., 1832, t, 3172.
A compound species of at least 2 jordanons. Pt. auriculata Col., Trans. N.Z. Inst., 22, 1890, 489, may eventually prove to be a variety of Pt. banksii. Until more data are available from the Invercargill area it is best included in the typical form.
(a) Pt. banskii var. typica Hatch.
Pt. emarginata Col., Trans. N.Z. Inst., 15, 1883, 328.
Pt. speciosa Col., ibid., 22, 1890, 488.
Pt. subsimilis Col., ibid., 28, 1896, 611.
Up to 50 cm. high. Leaves 1–8, strictly cauline, linear-lanceolate, acuminate, erect, up to 20 cm. long by 8 mm. broad. Floral bract foliaceous. Flower solitary or rarely 2, up to 8 cm. high. Dorsal sepal longer than the petals, filiform-caudate, the tip horizontal or suberect. Lateral sepals filiform-caudate, spreading, much exceeding the galea. Labellum linear-oblong, recurved, subacute or emarginate. Column typical, stigma linear-lanceolate. Column-wings with subulate upper lobes higher than the anther. Lower lobes broadly oblong, twisted, finely ciliate.
Distribution. Endemic—abundant throughout lowland areas in the North and South Islands. The Chatham Islands form appears to be quite distinct and may eventually warrant varietal rank.
Flowers October–November, sea-level–2,000 ft., scattered in scrub or on the forest floor.
Francis Bauer's illustration in Curtis's Botanical Magazine, 1832, t, 3172, can be regarded as the hypotype of the species.
(b) Pt. banksii var. patens (Col.) Hatch, Trans. R.S.N.Z., 75, 1945, 370.
Pt. patens Col., Trans. N.Z. Inst., 18, 1886, 270.
Similar to the type in general characteristics. Leaves shorter, broader and more spreading. Dorsal cauda strongly incurved. Lateral caudae recurved abruptly, the tips often meeting behind and below the ovary. The caudae naturally pass through the horizontal and suberect stages in developing from the bud, and transitional forms may often be found bearing a superficial resemblance to the typical form.
Distribution. Endemic—5, abundant in tussock and along forest margins about the central volcanoes, 12, 1944, E. D. Hatch; 7, high country throughout the Kaimanawa and Ruahine Ranges, 12, 1945, E. D. Hatch; 11, abundant in the vicinity of Arthur's Pass, 1, 1930, J. Brownlee; 16, abundant throughout Stewart Island, 12, 1946, C. Smith. Will probably prove to be quite common throughout the South Island high country.
Flowers December–January, 2,000–5,000 ft., scattered in grass and scrub and along forest margins. Probably derived from Pt. banksii typica.
11. Pterostylis graminea Hook. f., Fl. Nov. Zel., 1, 1853, 248.
Up to 25 cm. high. Leaves 1–7, very long, slender, linearacuminate, up to 12 cm. long by 6 mm. broad. Floral bract foliaceous. Flower solitary or rarely 2, up to 2 cm. high. Dorsal sepal acuminate, longer than the petals, the tip incurved or horizontal. Lateral sepals shortly caudate, hardly exceeding the galea. Labellum linear-oblong, obtuse, entire, recurved at the tip. Column strongly inclined, stigma elliptical. Column-wings with acuminate upper lobes as high as the anther. Lower lobes oblong, rounded.
Distribution. Endemic—not uncommon in lowland areas throughout the North and South Islands.
Flowers September–November, sea-level–1,500 ft., scattered in scrub or on the forest floor. Probably derived from a very early form of Pt. banksii.
An. Australian group well adapted to life in subalpine bogs. Pt. alpina Rog. and Pt. furcata Ldl. both appear to have originated from Pt. falcata Rog. Pt. furcata has extended to New Zealand and differentiated again into two quite distinct forms.
12. Pterostylis furcata Ldl., Gen. et Spec. Orch., 1840, 390.
A compound species of 2 jordanons, one in both Australia and New Zealand and the other confined to New Zealand.
(a) Pt. furcata var. typica Hatch.
Pt. mioromega Hook. f., Fl. Nov. Zel., 1, 1853, 248.
Pt. polyphylla Col., Trans. N.Z. Inst., 22, 1890, 489.
Up to 20 cm. high. Leaves 1–8, repand or flat, lower few usually subrosulate, petiolate, the upper cauline, sessile, or all may be cauline, oblong-ovate to -lanceolate, acute, up to 4 cm. long by 15 mm. broad. Floral bract foliaceous. Flower very pale, solitary, up to 4 cm. high. Dorsal sepal acuminate, longer than the petals, the tip horizontal or recurved. Lateral sepals filiform-caudate, exceeding the galea by as much as 15 mm. Labellum linear-oblong, obtuse, recurved and slightly
Fig. 1—Pt. trullifolia var. rubella nat. size. a, column and labellum from side; b, labellum from above; c, stigma; d, lateral sepals; j, special form adapted to vegetative reproduction; k, juvenile.
Fig. 2—Pt. trullifolia var. gracilis nat. size. Dissections as for var. rubella.
Fig. 3—Pt. trullifolia var. alobula nat. size. e, labellum from above; f, stigma; g, lateral sepals; h, intermediate stage.
twisted, the midrib channelled conspicuously on either side. Column slightly inclined, stigma narrow-obovate. Column-wings with acuminate upper lobes as high as the anther. Lower lobes narrow-oblong, the inturned margins finely ciliate. As a bud, and when just opened, the flower is beautifully symmetrical, but with maturity it goes to pieces, the segments falling away in all directions, so that at a stage when most Pterostylis flowers are in their prime, this species presents an untidy appearance. Very susceptible to epharmony, changing from subrosulate to cauline with environment. The juvenile is a close rosette of 3–5, ovate, petiolate leaves.
Distribution. Australia—Tasmania, Victoria, South Australia, New South Wales, nowhere abundant. New Zealand—2, Kaitaia, R. H. Matthews; 3c, Coromandel Harbour, Jolliffe; 5, abundant in Hypolaena bogs from Rotorua to Murimotu, particularly so along the western slopes of Mount Ruapehu, Allison, Matthews, Hatch; 6, Ngaire, T. F. Cheeseman; 7, Wairarapa, W. Colenso; Chatham Islands, 12, 1886, W. Travers.
Flowers December–January, sea-level–3,000 ft. Probably windborne across the Tasman originally.
(b) Pt. furcata var. linearis Hatch var. nov.
Pt. mioromega Cheesmn. in part (not of Hook. f.).
Pt. furcata typica affinis. Differentis in parvitas; folia lineata acuminata; sepala brevissima; stigma late cordatum.
Up to 18 cm. high. Leaves 1–4, cauline, sessile, linear-lanceolate, acute, up to 6 cm. long by 1 cm. broad. Flower similar to but much smaller than the type, the lateral sepals acuminate and hardly exceeding the galea. Stigma broadly cordate. The juvenile is a close rosette of 2–4 broad-elliptic, shortly petiolate leaves.
Distribution. Endemic—5, National Park, 12, 1921, H. B. Matthews; Murimotu, D. Petrie; 12, 1944, E. D. Hatch.
Flowers December, 3,000–4,000 ft. Abundant in Hypolaena bogs, not so far found in company with the type jordanon. Probably derived from Pt. furcata typica.
Holotype in Herb. Hatch, No. 565, Murimotu, 12, 1944, E. D. Hatch. The accompanying illustration can be regarded as the hypotype of the variety.
An Australian group of 4 or 5 species centred round Pt. obtusa R. Br. New Zealand has 4 jordanons originating from the same source.
13. Pterostylis trullifolia Hook. f., Fl. Nov. Zel., 1, 1853, 249.
A compound species of 3 closely related jordanons. The original description gives no indication as to which of the forms was the specific type. The species in one form or another has been recorded from the Districts 2, 3, 4, 5, 7 and 10. Much field work is yet needed to ascertain which jordanon occurs in which area. The districts and localities given under the descriptions are only those who have been confirmed by the writer.
(a) Pt. trullifolia var. rubella (Col.) Hatch comb. nov.
Pt. rubella Col., Trans. N.Z. Inst., 18, 1886, 271.
Up to 20 cm. high. Leaves 1–8, basal-petiolate or cauline-sessile, heterophyllous, ranging from ovate-orbicular through linear-lanceolate to bracteate. Floral bract narrow-linear, acute. Flower solitary or rarely 2, up to 3 cm. high. Dorsal sepal acuminate, as long as or longer than the petals, the tip incurved. Petals strongly incurved, horizontal to the dorsal sepal, producing a conspicuous “cobra-hood” effect. Lateral sepals with an emarginate central lobe, the filiform caudae much exceeding the galea. Labellum gradually narrowed to a rounded point. Appendage trifid, slender and much incurved. Column erect, stigma elliptical. Column-wings with the upper lobes subulate, higher than the anther. Lower lobes oblong, obtuse, with finely ciliate, inrolled margins.
Distribution. Endemic—2, not uncommon; 3a, abundant throughout; 4, Lake Tutira, H. Guthrie Smith; 7, Hutt Valley, 6, 1946, J. A. Healy; Foxton, 6, 1945, Wgtn. Bot. Soc.
Probably derived from an early form of Pt. obtusa and windborne originally across the Tasman.
(b) Pt. trullifolia var. gracilis Cheesmn., Trans. N.Z.I., 47, 1914, 46.
Similar to var. rubella, more slender, often taller. Petiolate leaves with the reticulated veining conspicuously embossed. Basal rosette more inclined to persist with maturity. Flowers much smaller, the petal tips frequently crossed or divergent.
Distribution. Endemic—2, 3a, b, c, common throughout; 5, Rotorua, 7, 1934, K. W. Allison.
Probably derived from var. rubella.
(c) Pt. trullifolia var. alobula Hatch var. nov.
Pt. trullifolia rubella affinis, subsimilis. Differentis in sinus sepalium lateralium acutus, nulla loba.
Habit and size of var. rubella. Flower darker green, rather erect. Sinus of the lateral sepals acute, without a central lobe. Labellum narrower, with a slightly swollen, truncate or even crenulate tip. Probably derived from Pt. obtusa.
Distribution. Endemic—2, 3a, abundant throughout.
The species as a whole flowers from April-September, sea-level–1,000 ft. Large colonies in scrub, open mossy places or forest. Hybrids between gracilis and alobula are frequent in areas where they overlap.
Holotype in Herb. Hatch, Laingholm, 7, 1945, E. D. Hatch, No. 566. The accompanying illustration can be regarded as the hypotype of the variety.
Development. The juvenile rosette develops into the bracteate mature form over about 3 seasons. The degree of development depends entirely upon the size of the tuber. When a plant by the removal of leaves or other mutilation is not able to develop a normal-sized tuber, the following season finds it producing a form more juvenile than that which preceded it. The vegetative form is worth remarking. It consists of a pseudobulb at the surface, bearing 2 orbicular leaves on very long petioles, and 3–5 very long roots with terminal tubers.
14. Pterostylis venosa Col., Trans. N.Z. Inst., 28, 1896, 610.
Up to 7 cm. high. Leaves 1–3, sessile, elliptic-lanceolate to oblongovate or almost orbicular, obtuse or acute, up to 6 cm. long by 3 cm. broad. Floral bract foliaceous, acute. Flower solitary, up to 15mm. high. Dorsal sepal acuminate, slightly longer than the petals. Lateral sepals shortly caudate, the tips often recurved. Labellum recurved, gradually narrowed into a blunt point, or the upper half abruptly constricted into a long subulate tip. Column short, the stigma linearlanceolate. Column-wings with acuminate upper lobes as high as the anther. Lower lobes broadly oblong, obtuse.
The subrosulate habit is little changed with maturity, the lengthening of the stem taking place mainly above the floral bract.
Distribution. Endemic—7, not uncommon throughout the Tararua and Ruahine Ranges; 8, Endeavour Inlet, J. H. MacMahon; 10, montane areas in the vicinity of Westport, W. Townson; 11, not uncommon in the vicinity of Arthur's Pass, 1, 1931, L. M. Cranwell.
Flowers December–January, 2,000–4,000 ft., colonies in tussock or scrub, along forest margins or alpine herb field. The Arthur's Pass form differs somewhat from that found in the North Island and on Mount Frederick. It will probably be found that 2 jordanons exist within the present conception of the species. Probably derived from Pt. trullifolia alobula, although specimens collected recently from Mount Egmont suggest affinities with Pt. foliata Hook. f. also.
Holotype in the Dominion Museum, Wellington, east side of the Ruahine Ranges, 1, 1894, A. Olsen.
15. Pterostylis humilis Rog., Trans. Roy. Soc. S. Austr., 46, 1922, 151.
Hatch, Trans. R.S.N.Z., 75, 1945, 369.
Pt. confertifolia Allan, Trans. N.Z. Inst., 56, 1925, 32.
Up to 10 cm. high. Leaves 1–6, sessile or petiolate, rosulate, oblong-ovate, obtuse, up to 9 cm. long by 2 cm. broad, glaucous above, silvery beneath, turning a bright yellow in the autumn. Floral bract foliaceous, frequently larger than the leaves, oblong acute. Flower solitary, rather broad, up to 18 mm. high, translucent with thin green striae. Dorsal sepal erect, longer than the petals, the acuminate tip horizontal. Lateral sepals erect with a very wide sinus, the lobes inrolled and hardly exceeding the galea. Labellum linear-oblong, reddish-brown, recurved, the tip constricted. Column erect, free from the dorsal sepal. Stigma very large, globose, protruding on either side of the column and forward beyond the vertical line or the anther. Column wings with 2 upper lobes, the anterior small, acute, the posterior larger, rounded. Lower lobes rather long, narrow-acute, incurved.
The stem elongates considerably with maturity, scattering the leaves up the stem. Juveniles may have petiolate or sessile leaves.
Distribution. Endemic—5, occasional about the base of Mount Ruapehu, Matthews, Attwood, Hatch; Kaimanawa Ranges, 1, 1945, E. D. Hatch; 7, occasional on the upper levels of the Ruahine Ranges, H. H. Allan.
Flowers December, 4,000–5,000ft., large, closely massed colonies in deep mosses at the upper limit of the subalpine scrub. Probably derived from the Australian Pt. cucullata R. Br. and windborne originally across the Tasman.
Lectotype in the Auckland Museum, “Haunted Whare,” Mount Ruapehu, 1, 1921, H. B. Matthews (this is one of the fruiting plants, from the tubers of which Matthews cultivated the specimens later described by Rogers as Pt. humilis). The accompanying illustration can be regarded as the hypotype of the species.*
[Footnote] * Since writing the above the following new localities have been confirmed: 6, Mt. Egmont and foothills—Pt. montana typica, Pt. trullifolia alobula, Pt. banksii patens, Pt. australis, Pt. venosa, Pt. humilis—O. E. Gibson, B. Irwin, E. S. Richardson; 10, Wakapuaka (Nelson)—Pt. trullifolia alobula, A. W. Wastney; 11, Kelly River—Pt. banksii patens, Pt. montana typica—P. Haddon-Jones.