Notes on some New Zealand Plants and Descriptions of New Species.

Transactions and Proceedings of the Royal Society of New Zealand, Volume 70, 1940-41, Page 27

Notes on some New Zealand Plants and Descriptions of New Species. By G. Simpson and J. Scott Thomson. [Read before the Otago Branch, October 10, 1939; received by the Editor, October 18, 1939; issued separately, June, 1940.] Carmichaelia pilosa Col. ex Hook. f., Fl. Nov. Zel., 1 (1852), p. 50. This was based on specimens collected by Colenso on the “east Coast” North Island. Hooker separates it from C. odorata by (1) the covering of silky hairs, especially on the inflorescence and ovary, (2) the longer curved style, (3) the much shorter and broader pod, hairy till nearly ripe. Kirk (1899, p. 113) treated the plant as a variety of C. Odorata. Neither Kirk nor Cheeseman (1925, p. 521) had recognised specimens, and both, considering hairy pods a variable character in the genus, were apparently doubtful of the status of Colenso's species. While examining specimens for another purpose, the authors found C. pilosa to be well represented in herbaria. The following brief description will serve to separate the species: C. odorata. Branchlets 2 mm. wide, silky-pubescent at the tips, notches small. Leaves small, about 5 mm. × 5 mm., 5 foliate, silky pubescent; leaflets very small, about 2 mm. × 1 mm., equal. Racemes pubescent, 8–12 flowered, erect. Ovary glabrous. Pods about 7 mm. × 4 mm., broadly ovate to almost orbicular, with a long subulate beak. C. pilosa. Branchlets 3 mm. wide, everywhere silky-pubescent notches conspicuous, oblique, pointed. Leaves 5 foliate, 8 mm. × 6 mm., terminal one the larger, silky pubescent, leaves on young branchlets simple and of varying sizes. Racemes silky pubescent, densely more or less 12 flowered, erect. Ovary with minute papillae at the base of the stigma. Pods about 6 mm. × 3 mm., broadly ovate to orbicular, sparingly pilose till nearly ripe. Beak short, stout, subulate. We find the pilose character in the pod to be rare in the genus, but a constant character when it occurs. Specimens of C. pilosa, labelled as C. odorata, appear in the collections of Kirk, Cheeseman, Petrie, Cockayne, Aston and others, and it appears to be a common enough plant in river gorges of the Wellington Province. It is amply distinct from C. odorata, and we restore it to specific rank. The life-form of Rubus squarrosus Fritsch. Cockayne (1933, p. 545) discussed the polymorphy of this species (using the name R. cissoides* For the Nomenclature see Allan (1935, p. 229)) and its causes. He distinguished two extreme states: The liane-form: stems high-climbing, leaves with well-developed laminae (“lamina form”), leaf-stalks short, with weak armature of prickles; the bolster-form: stems entangled into bolster-like masses, leaves with greatly reduced laminae (“midrib form”), leaf-stalks long, with strong armature of prickles. He reviewed the field evidence and considered these forms to be epharmones induced by the diverse habitat conditions—forest for the liane-form, open ground for the

bolster-form. He reported, but did not accept the opposing views we communicated to him: “Their observations on plants in modified forest (more sunny and with drier atmosphere than in primeval forest) lead them to the conclusion that midrib leaves on young plants in the forest are attuned to more shade than the lamina leaves of the adult,” and “they consider all spreading growth is at first juvenile in form and that sunlight is needed to induce the adult growth.” “They view the laminae form as the treetop form, but found also in sunlight at forest margins. They agree, however, that the juvenile midrib form produces bolsters in fully exposed situations, and that those develop neither lamina leaves nor flowers, though they carry much ripened wood.” Having further studied the problem, we maintain our position. The divergent views may best be brought out by taking Cockayne's summary seriatim. (a) “Rubus cissoides is a heteroblastic species with a very distinct juvenile stage, in which the leaves are of midrib form.” We agree as to the heteroblasty in the sense that the climbing midrib state differs in an astounding manner from the form in which lamina leaves and flowers are produced. (b) “The juvenile is apparently in tune with forest conditions.” This is our view, with the omission of “apparently.” The new sappy strict growth, with its elongated, fiercely armed midribs terminating in narrow lanceolate laminae, lacks nothing to function as a forest liane, but this fact, though generally accepted, has not been sufficiently explored. (c) “Plants growing in the open on dry ground and exposed to full sunshine, with frequent high winds, produce bolster-like masses, and the leaves are all of the midrib form. Such plants do not flower, though they attain a large size and evidently a considerable age.” We consider this to be an important point. The assumption is that bolster-like netted masses of stems and leaves are produced only in exposure to sun and wind, and Cockayne describes this form as “wind-resisting.” A similar form is, however, also common enough in shade and in shelter from wind, and the midrib leaves are then larger and more strongly armed than are the leaves of tangled windswept—not “wind-resisting”—masses of the open. In dense forest or thickets, midrib leaves only are in evidence, reaching up and out to the light as a climbing growth, but where the forest is open and support lacking—though the ground may be deeply shaded—bolsters are formed. The modification towards the lamina form, if the extreme be considered, is directly as the amount of sunlight, not shade, but first the climbing urge is lessened by a hardening of the stems. Leaves are produced on ripened stems, never on sappy growth. Light is the important factor in the production of lamina growth, and on the flowering branches the leaves are always considerably modified towards the unarmed, coriaceous, lamina stage. Lateral branches free to hang pendant, or to lie over the canopy, produce lamina leaves and flowers both in bright sunlight and in

comparative shade, and the greater the exposure to light the greater the development of the leaves. Flowering laterals and midrib growth may be found together, in the same exposure and from closely placed nodes, or the climbing growth may appear on an offshoot from the lateral branch, and the climbing stimulus is ever rampant. Midrib growth appears everywhere from the ground to the end of shoots pushing beyond the treetop, it is the climbing growth of a sun-demanding liane. Flowering branches may also be produced on the bolster form wind or protected in some manner from the grasp of the midrib leaves of the mass. There the necessary conditions, both light and freedom, are provided, but elsewhere the bolster is a mass of growth striving to climb, but arrested in its development as a liane. New shoots, produced from the base of older growth, for a while reach outwards or upward until, arching by their weight, or battered by wind, they are caught up in the entanglement. This exposed bolster is, as a mass, an epharmonic form, the product of the exposed conditions, but the plant in this stage of growth is out of its real habitat and a relic of forest and shrub associations long since destroyed. (d) “When growing as a liane of the forest, R. cissoides develops a lamina leaf, flowering and fruiting abundantly.” That statement is a perfectly correct one, but, while Cockayne concludes that all such development is due to shade, the authors contend that in forest the plant is but fulfilling its functions as a liane, reaching into light, not shade, and developing directly as the degree of light its forest habitat provides. (e) “The balance between the leaf forms is very delicate, and the plant epharmones readily in one direction or the other, according to the degree of sunlight and of exposure, or even a part of it, experiences.” The authors agree with Cockayne that the balance is delicate, but the midrib leaves which he considers to be an epharmonic form of bright light are the constant in all climbing growth, and lamina-leaved lateral branches will appear from the nodes of older persistent midrib leaves in the fullest exposure to light. In an earlier paper (1901, p. 265) he comments on the fact that poor soil hastens, and a moist atmosphere retards, flowering growth, and he writes, “Here, then, is a case to the contrary: the plant of the moist wood produces fruit regularly, the plant of the barren windswept slopes rarely or never does do.” If, however, the bolster mass of such situations is considered as arrested climbing growth, an extreme state, in one direction, of a heteroblastic species, and the lateral branches on the treetop, with short petioled coriaceous leaves, as the other extreme, it is no longer a question of aridity or moisture, no longer a case to the contrary, but a plant functioning exactly as other plants do in like circumstances. The species epharmones readily to all degrees of light and shade, but only in the degree of modification from the early climbing form towards the extreme lamina form, and in exactly the opposite direction to that anticipated by Cockayne. A truly shade-demanding

plant, in its early growth, could not survive under the conditions in which the bolster masses of the open persist. In essence, then, Cockayne views Rubus squarrosus as a plastic species, epharmoning towards its juvenile midrib form in sunlight, towards the lamina leaf, flowering form in shade. Our view, from repeated examination of all stages, is that this species follows the normal development from the seedling stage, but this development may be arrested. The seedling form is followed by a midrib form adapted to climbing. This is succeeded by the lamina leaf adult flowering form on attainment of full sunlight. The midrib bolster form of the open is prevented from attaining the adult form by the exposed conditions of the habitat. Most examples, at least, of this form are relics, persisting on formerly forested or shrub-covered areas. The midrib form is the constant in all climbing and extending growth, and may appear under all grades of light and shade. Fully developed lamina leaves are not shade leaves, but belong to adult flowering branches. This flowering stage of the sunlit tree-tops is quite common, but flowers may also be produced where the light is only sufficient to produce an intermediate growth. Epilobium wilsoni Petrie, var. pallidum nov. Typi valde affinis sed caulibus foliisque pallide viridibus nusquam pubescentibus differt. Leaves and floral characters exactly as the type, stems and leaves pale green in colour, nowhere dark red or reddish. Common on moist limestone in the gorge of the Ure River, Marlborough. The variety seeds freely in cultivation, and the seedlings are all as the parent plant. Hebe fruticeti sp. nov. Frutex c. 1 m. altus, ramosus, ramulis gracilibus confertis adscendentibus; ramuli hornotini virides, nodis rufo-brunneis; ramuli annotini rufo-brunnei. Folia late sessilia, 3 cm. longa, 5 mm. lata. lineari-lanceolata, subacuta, subcoriacea, supra pallide viridia concava que, subtus convexa, costa manifesta. Racemi apicem ramulorum versus dispositi, 5–6 cm. longi, 1–5 cm. lati; bracteae inferiorae lineari-lanceolatae, calycem attingentes; bracteae superiorae breviores; calycis lobi lanceolati, acuti; corollae tubus vix calycem excedens, lobi lati patentes; fructus calyce 2 plo longior, late ovatus, subacutus vel styli base mucronata. A closely-branched, spreading, glabrous shrub 1–1·5 m. high; branches spreading shortly and ascending to nearly erect, slender, leafy at the tips, elsewhere rough by old leaf sears, old wood reddish-brown; young tips green ringed with reddish-brown at the nodes; leaves sessile by a widened base, 3 cm. long, 5 mm. wide, linear-lanceolate, sub-acute, sub-coriaceous, pale green and concave above, lighter green and convex beneath, midrib evident, margins entire; racemes in one or two pairs near the tips of the branches, 5–6 cm. long, 1–5 cm. wide, rhachis slender, puberulous, pedicels of the lower flowers 3 mm. long, about opposite, and the bracts linear lanceolate, as long as the calyx, upper pedicels shorter, 2 mm.; bracts about

equalling the pedicel or shorter. Flowers closely placed, spreading; calyx small, 2 mm. long, 4 partite, segments lanceolate, acute, margins membraneous, corolla white, tube short, less than 2 mm. long, not exceeding the calyx; lobes 3 mm. long, wide, rounded at the tip, spreading; stamens as long as the lobes, style longer. Capsule 4 mm. long, 2 mm. broad, twice as long as the calyx, broadly ovate, sub-acute or with the hardened style forming a short point. Habitat: subalpine scrub on basins at the head of the Estuary Burn, Lake Wanaka. Altitude 1000–1500 m. This is the common Hebe component of the subalpine scrub association in all upper basins in this habitat. It is closely allied to H. subalpina, which has larger and wider leaves. Specimens turn brown in drying. Type in Herbarium, Plant Research Bureau, Wellington. Hebe lapidosa sp. nov. Frutex c. 1 m. altus, erectus, ramosus, ramuli hornotini virides, bifariam pubescentes; ramuli annotini glabri, rufo-brunnei. Folia satis dispersa, suberecta vel patentes, c. 2 cm. longa, 6–8 mm. lata, obovato-oblonga, subacuta, breviter petiolata, crasse coriacea, rigida, supra clare viridia nitentia, subtus glauco-viridia; costa subtus prominente. Inflorescentiae trichotome spicatae in axillis penultimis ramulorum dispositi; spicae dense multiflorae; bracteae obtusae tubo corollae longiores. Calyx 4-lobatus, lobi tubo corollae attingentes, lineari-lanceolati, obtusi; fructus late oblongus vel fere orbiculatus, obtusus, compressus, calycem breviter excedens. A much branched, erect, glabrous shrub, about 1 metre high, older branches stout, erect, reddish-brown, rough by old leaf sears, leafy towards the tips; young wood at the tips green, ringed with brown at the nodes, finely bifariously scaly pubescent above the leaves. Leaves openly spaced, sub-erect or spreading, 2 cm. or longer, 6–8 mm. wide, obovate-oblong, sub-acute, narrowed at the base to a short petiole, coriaceous, light shining green and concave above, convex, glaucous green, and keeled by a yellowish prominent midrib beneath; margins yellowish, entire; petiole attached by a spreading base. Flowers in 2 to 6 short opposite compound spikes near the tips of the branches; spikes slightly exceeding the leaves, dividing above leafy bracts at a brownish node into 3 densely flowered branchlets; bracts shaped as the leaves, 6 mm. long, upper ones greenish with membranous margins, obtuse, longer than the calyx and corolla tube; calyx 4-partite, greenish, segments about equalling the corolla tube, 4 mm. long, linear lanceolate, acute, margins membranous; corolla white; tube about 4 mm. long; lobes widely obovate-lanceolate, obtuse, spreading, about 4 mm. long; stamens shorter and the style longer than the lobes. Capsule 4 mm. long, 2 mm. broad, broadly oblong

or almost orbicular, obtuse, compressed, slightly exceeding the calyx. Probably nearest to H. rigidula, but differing in its erect habit, longer, wider spaced leaves, less branched spikes and larger flowers. Habitat: stony debris on rock benches and flood beds in the gorge of the Dee River, Clarence Basin, Marlborough, 500–1500 m. altitude, exceedingly common in this habitat and probably mistaken here and elsewhere in the North-Eastern District for H. willcoxii, a plant of subalpine scrub, which it somewhat resembles. Type in Herbarium, Plant Research Bureau, Wellington. Hebe recurva sp. nov. Frutex ramosus, c. 1 m. altus, ramuli hornotini virides, bifariam pubescentes; ramuli annotini rufo-brunnei. Folia haud conferta, arcuato-patentes, 4–5 cm. longa, 4–6 mm. lata, lineari-oblonga, subacuta, in basim semi amplexicaulem angustata, tenuia, supra vix concava, costa manifesta. Racemi apicibus ramulorum versus dispositi, c. 6 cm. longi, multiflori; bracteae 2 mm. longae, anguste lineares, obtusae; pedicelli breves puberuli; calyx 4-lobatus, c. 2 mm. longus, obscure puberulus, lobis lineari-lanceolatis, subacutis, marginibus ciliolatis; corollae tubus haud 2 mm. diam., lobis 2 plo longior; lobi lati, patentes, marginibus incurvis, lobus anterior multo brevior. Fructus oblongus, compressus, subacutus, calyce 2 plo longior. A much-branched, spreading, glabrous shrub about 1 m. high, older branches reddish-brown, rough by old leaf scars, young branches green and leafy, faintly red-brown under the leaf pairs, finely bifariously pubescent above the leaves. Leaves open spaced, spreading, deflexed by arching, 4–5 cm. long, 4–6 mm. wide, linear-oblong, sub-acute, narrowed to a semi-amplexicaul base, thin, almost glandular, flat or slightly concave above; midribs slender and lighter coloured; margins entire. Flowering racemes in two alternate pairs near the tips of the branches, about 6 cm. long; rhachis slender, many flowered, puberulous, bracts 2 mm. long, narrow linear, obtuse; pedicels short, about 4 mm., puberulous; calyx 4-partite, about 2 mm. long obscurely puberulous; segments linear-lanceolate, sub-acute, deeply cut, margins ciliolate and membranous, corolla white, 7 mm. long, tube less than 2 mm. diam. and twice the length of the lobes; lobes broad, spreading, rounded at the tip; margins incurved, the anterior lobe much smaller; stamens slightly exceeding the style and much exceeding the lobes. Capsule 4 mm. long, 2 mm. broad, oblong, compressed, sub-acute, twice as long as the calyx. Habitat: rock platforms on the banks of the Aorere River near Bainham, Nelson—plentiful in this habitat and perhaps a common ruprestral species in similar situations in North-West Nelson. Type in Herbarium, Plant Research Bureau, Wellington.

Fig. 1.—Young climbing growth from shaded forest. Fig. 2.—Climbing and adult growth from forest margin in sunlight.

References. Allan, H. H., 1928. New Zealand Trees and Shrubs, Wellington. Allan, H. H., 1936. Notes on New Zealand Floristic Botany, etc., No. 6, Trans. N.Z. Inst., vol. 65, p. 221. Bird, J. W., 1916. Observations of the Lianes of the Ancient Forest of Canterbury Plains of New Zealand, Trans. N.Z. Inst., vol. 48, p. 315. Buchanan, J., 1882. On the Alpine Flora of New Zealand, Trans. N.Z. Inst., vol. 14, p. 342. Cheeseman, T. F., 1925. Manual of New Zealand Flora, ed. 2, Wellington. Cockayne, L., 1901. An Inquiry into the Seedling Forms of New Zealand Phanerogams and the Development, Part 4, Trans. N.Z. Inst., vol. 33, p. 265. — 1910. On a Non-flowering New Zealand Species of Rubus, Trans. N.Z. Inst., vol. 41, p. 325. — 1917. Notes on New Zealand Floristic Botany, etc., No. 2, Trans. N.Z. Inst., vol. 49, p. 56. — 1933. A case of Epharmony in a New Zealand Rubus, American Journal of Botany, vol. 20, no. 8. — and Allan, H. H., 1927. Notes on Floristic Botany, including Descriptions of New Species, etc., No. 5, Trans. N.Z. Inst., vol. 57, p. 48. Laing and Blackwell, 1939. Plants of New Zealand, ed. 4, p. 207, Christchurch. Poppelwell, D. L., 1915. Plant Covering of Garvie Mountains, with List of Species, Trans. N.Z. Inst., vol. 47, p. 120.