Contributions to a Knowledge of the Naturalized Flora of New Zealand. No. 4
[Read before the Wellington Branch, November 17, 1953, received by the Editor, November 19, 1953]
The alien species, Aphanes microcarpa (Boiss. et Reut.) Rothm. (Rosaceae) and Nasturtium microphyllum Boennh. ex Rchb. (Cruciferae) are recorded as new for New Zealand. Then previous inclusion under A. arvensis L. and N. officinale R.B.1 respectively is noted, the distribution is given, and reasons are advanced for the change from Alchemilla Scop to Aphanes L.
The rarity of the supposedly abundant alien. cronica agreslis L, is discussed.
1. Aphanes L. (Rosaceae)
Prior to consideration of the two species involved, an explanation for the reversion from the familiar Alchemilla Scop. to the Linnaean Aphanes appears warranted.
The species which constitute the genus Aphanes L. (sub-genus Aphanes (L.) Rothm. of Alchemilla Scop.) are small diffuse annuals, whereas the species belonging to Alchemilla [sensu stricto] are rhizomatous perennials. Apart from the difference in habit and duration, there are marked differences in the androecium—in Aphanes, the androecium consists of a single stamen (rarely two) with extrorse anther, mounted on the inner margin of the disc opposite a sepal, while in Alchemilla it consists of four stamens with introrse anthers, mounted on the outer margin of the disc and alternating with the sepals.
In view of these differences, which define Aphanes as a distinct generic entity, I have decided to revert to this genus for the species naturalized in New Zealand, a treatment in keeping with that of recent overseas workers.
Aphanes arvensis L. was originally recorded from New Zealand (locality not stated) by Hooker (1855: 321) as Alchemilla arvensis Sm., and later by the same author (1864A: 57) as collected by Travers from Tarndale Plains, South Island (as Alchemilla (Aphanes) arvensis L.) Bentham (1864: 433) refers to the occurrence of the plant in New Zealand, and commented on the nativity of the species thus “…… and is also found in New Zealand; but in the latter country, and in some, if not all, the Australian stations, very probably introduced from Europe.” The species was first recorded from the North Island by Kirk (1870: 137), it being noted as “Local” in the Waitemata district, Auckland Province.
The species which has till now been included under A. arvensis L., in New Zealand, and until recently in Europe, is* * A. microcarpa (Boiss. et Reut.) Rothm. described from Spain in 1842. This species went unrecognized in Northern Europe until Rothmaler in 1935 noted its occurrence in Sweden; it was then
[Footnote] * Species recorded for first time for New Zealand.
reported from Denmark by Gudjonsson in 1941, from Britain by Rothmaler in 1941, and more recently Walters (1949:163–9) has shown it to be widespread in Britain. This latter author gives an excellent account of the two species, and I have drawn on the information therein for data on the European distribution of A. microcarpa and details of Alchemilla [sensu stricto].
While the two species appear similar on cursory examination, I have found that A. arvensis is generally a stout plant, A. microcarpa more slender, and with longer internodes. Differentiation of the two species on habit character has definite limitations, and the best diagnostic characters are the nature of the lobes of the floral stipules and the fruiting urceoles surmounted by the persistent calyx.
In A. microcarpa, the floral stipules usually overtop the calyx lobes, the stipular lobes are long and “finger-like,” the urceole + calyx is 1.4–1.8 mm long, and the calyx teeth are small, convergent, with no distinct constriction (neck) at the point of origin. The fruit illustrated by Hyde (1945 617) for A arvensis is typical for A. microcarpa.
In A. arvensis, the calyx lobes usually project above the floral stipules, the stipular lobes are broadly triangular, the urceole + calyx is 2.2–2.5 mm. long, the calyx lobes are longer than in the preceding species and spread from a distinct constriction.
Previously published data on distribution of A. arvensis cover the two species, and to give a clear picture of distribution more material from many districts will require to be examined. Available material shows A. microcarpa to occur in both Islands, often locally abundant, while A. arvensis occurs through the South Island, and is apparently more restricted in the North Island. I have seen only two gatherings from the North Island, both of which, although undated, were probably collected during the nineteenth century.
The speces of Aphanes occur in dry waste places, low density pasture in dry situations, and in modified low tussock grassland, commonly in the nature of ephemeral volunteers occupying bare spaces, and of little economic significance As a bottom weed of cereal crops in some districts the species tend to be significant, in damp seasons, and where the binder is employed the mass of material in the butts of sheaves retards drying out. It is as a weed of lawns and playing-greens that A. microcarpa, often associated with Soliva spp., attains definite importance, under the vernacular name “Parsleypiert.”
A. Aphanes arvensis L.
Auckland Institute and Museum, Auckland: Raglan, T F. Cheeseman, date not stated (29072, 29073, 29074, 29075); Botany Division, D.S.I.R., Wellington: Tussock-grassland, Molesworth, Marlborough, H. H. Allan, 1943 (45338) (this material is atypical, but comes closer to this species than to A. microcarpa): tussock-grassland, confluence of Doubtful and Boyle Rivers, Canterbury, 1950 (74633); stony river-flat. Mason River, Waiau, 1944 (33596); dry soils near Christchurch, collector and date not stated, (5923) (ex Herb. T. Kirk); cornfields, Lincoln, collector and date not stated (58655) (ex Herb T. W. Kirk): dune area, Cromwell Flats, H. H. Allan, 1944 (48446). Dominion Museum, Wellington. Ruataniwha, Hawke's Bay. H. Orzym (No 64), date not stated.
B. Aphanes microcarpa (Boiss. et Reut.) Rothm.
Botany Division, D.S.I R., Wellington: near Taihape, V. D. Zotov, 1928 (872); Phormium nursery, Paiaka, near Shannon, 1943 (36267); dune pasture, Waikanae, collector and date not stated (45337); lawn, Upper Hutt, 1950 (74638); waste land, Upper Hutt, 1952 (74634); cultivated land, Wallaceville, 1952 (74634); grassy flats, Maruia River, about 8 miles west of Lewis Pass summit. 1950 (74636); on logging track, Lockington's sawmill, west of Lewis Pass summit, 1950 (74637); roadside, Omarama, H. H. Allan, 1936 (17350). Dominion Museum, Wellington: Birdling's Flat, Bank's Peninsula, T. Kirk, date not, stated (this collection, made in the nineteenth century, is the earliest I have examined from New Zealand. Examination of the material on which the records cited earlier in this paper are based may show that some belong to this species).
A further, as yet undetermined species of Aphanes was collected in New Zealand during the last century. There are specimens in the Kirk Herbarium at the Dominion Museum, Wellington, collected from Lyall Bay, Wellington, date not stated, and laid in by Kirk under A. vulgaris L. The plant is certainly not Alchemilla vulgaris, and is one of the annual species of Aphanes, but the material is not sufficiently good for an exact determination. I submitted fragmentary material from this collection to Dr. Walters, University of Cambridge, and he indicated that while it bears some resemblance to North American species, more material would be necessary for determination.
2. Nasturtium R.Br. (Cruciferae)
The common watercress (Nasturtium officinale R.Br. [sensu lato]) was first recorded from New Zealand by Hooker (1852: 14) when he noted “…. appears to be naturalized about Auckland,…,” and the distribution was widened by the same author (1864A 10) with the observation “…. abundantly naturalized in rivers, etc.” and further (1864A: 15) “…. a pest in the rivers about Canterbury “The through establishment of the species, at least locally, in these early years, is evidenced by Hooker's statement (1864B 124) that it cost more than £300 per annum to clear this weed in the Avon River, Christchurch. This local abundance may have prompted the colonists to introduce the Australian black swan in an attempt to control the weed, as is stated by Murphy (1951. 577), who reports that bird as “introduced at the behest of sportsmen or, as some New Zealanders allege, for the purpose of destroying the introduced watercress”
Field observations and later examination of herbarium material showed two distinct entities in our concept of N. officinale R. Br., one being N. officinale [sensu stricto], the other* * N. microphyllum Boenn. ex Rchb.
The two species are readily distinguished on macroscopic fruit characters—N. officinale the siliquae are relatively short and broad, pedicels stout, the seeds arranged in two distinct rows, and in N. microphyllum the siliquae are long and slender, often arcuate, the pedicels longer and more slender, and the seeds in one, or indistinctly two-rowed. The sculpturing of the seed coat is diagnostic—in N. officinale there are about 25 large depressions on each face, and in N. microphyllum about 100 small depressions on each face. Aside from the fruit
[Footnote] * Species recorded for first time for New Zealand.
and seed characters, the position of the mature anthers differs—in N. officinale that anthers are introrse, and in N. microphyllum the anthers are extrorse.
Overseas, the two species are distinct in autumn aspect—the stems and leaves of N. officinale remain green in autumn, while the stems and leaves of N. microphyllum turn purple-brown in autumn. I have been unable to examine the autumn aspect of N. officinale in New Zealand, but all stands of N. microphyllum examined show the characteristic purplish-brown autumn colour.
Further collections are required to provide more accurate data on distribution, but on the basis of field observations and herbarium material examined, N. officinale appears to be restricted to the North Island, while N. microphyllum occurs in both Islands, locally common in localities from the Manawatu and Wairarapa districts southwards.
A. Nasturtium microphyllum Boenn. ex Rchb.
Auckland Institute and Museum, Auckland: Auckland, T. F. Cheeseman, date not stated (28731, 28732, 28733). Botany Division, D.S.I.R., Wellington: Feilding, V. D. Zotov, 1928 (707, 708); Foxton Line, Manawatu, I. W. Davey, 1937 (18293); Tauherenikau, Wairarapa, V. D. Zotov, 1947 (59087); Wilton's Bush, Wellington, R. Mason, 1950 (70339); Waiwhetu Stream, Lower Hutt, R. Mason, 1950 (65623); Nelson, D. M. E. Merry, 1952 (81416); Wakefield, Nelson. W. O. Sly, 1952 (81415); north of Birchfield, Buller County, R. Mason and N. T. Moar, (81040, 81217); north of Charleston, Buller County, R. Mason and N. T. Moar, 1953 (81291); Cass, Canterbury, R. Mason, 1951 (71455); Lakeside, Leeston, P. R. Barrer, 1952 (81414); Washdyke, W. J. Kissock, 1952 (81418), Pleasant Point, W. J. Kissock, 1952 (81417); Invercargill, A. L. Poole, 1940 (23931, 29470). Dominion Museum, Wellington T. Kirk, 1893.
B. N. officinale R.Br.
Auckland Institute and Museum, Auckland: Taipa, North Auckland, R. Cooper, 1950 (36049). Botany Division, D.S.I.R., Wellington: Thompson's Lake, South Head, Manakau Harbour, N T. Moar, 1950 (70875); Whakataki River, Wairarapa, V. D. Zotov, 1945 (48680); Hutt River, near Upper Hutt, 1950 (81424, 81425); Cape Turakirae, R. Mason, 1952 (74081); near Barney's Hut, Palliser Bay, R. Mason, 1952 (74087). Represented in Seed Herbarium by two collections: Plant Research Station farm near Palmerston North, collector not stated, 1927; Orongorongo River, R. Mason, 1946.
A collection by Miss R. Mason from Ohariu Valley, Wellington Province (81586) appears to be one of the hybrid forms between the two species, but does not show the extreme sterility associated with overseas material. A series of specimens collected by my wife from Pakowhai, Hawke's Bay (81420, 81421, 81422, 81423) have siliquae longer and narrower than typical N. officinale, the seed testa depressions range between 40 and 50, and the seeds are arranged in two rows. Again in this material, there are less than 25 per cent of sterile ovules per fruit.
3. Veronica agrestis L (Serophulariaceae)
This European species was recorded originally from Auckland, on the authority of T. Kirk, by Hooker (1867: 761), and was subsequently recorded from the Waitemata district by Kirk (1870: 141). from the Wellington district
by Kirk (1878A: 375), from Otago by Kirk (1878B: 416), and as “A weed of frequent occurrence in most districts [Auckland Provincial District] by Cheeseman (1883:289).
On the distribution and abundance of the species in the present century, the main authorities comment as follows: Cheeseman (1906: 1083, 1925: 1077 “Kermadec Islands, North and South Islands: fields and waste places, abundant.”; Hilgendorf (1926. 158, 1948: 164) “…is abundant in fields and waste places throughout the islands.”; Allan (1940: 201) “Common in waste places, cultivated land and open pastures in both Islands.” The species has also been cited by various authors as a weed of waste places, arable land and pastures in many districts.
The above-quoted statements would indicate Veronica agrestis L. as a widespread and common species, yet on the basis of fifteen years of search in the field and examination of all available material in New Zealand herbaria, I have seen only two specimens of New Zealand origin, indicating that the species is by no means as common as has been suggested.
There is a specimen in the Botany Division Herbarium (ex Herb. T. Kirk) from Auckland, collector and date not stated (58027), although it was probably collected by Kirk himself, and may even be the basis for Hooker's original record. The second specimen I collected from cultivated land at Akaroa, 19th February, 1945 (33837), it being one which for several years I regarded as a whitish-flowered variant of V. persica Poir.
It is evident that the species has been confused both in the field and in the herbarium with V. persica Poir. and V. arvensis L., from both of which species it is readily distinguished V. arvensis with its sessile or subsessile fruits is easily separated from the other species which have distinctly stalked fruits. While closely related. V. agrestis is distinguished from V. persica by the fruiting pedicels being nearly as long as the leaves, the calyx lobes obtuse or sub-obtuse, the capsule slightly broader than long with narrow acute sinus, and the style about as long as the sinus, whereas in V. persica the fruiting pedicels are distinctly longer than the leaves, the calyx lobes acute, the capsule considerably broader than long with wide oblique sinus, and obliquely bent style much longer than the sinus.
The Akaroa specimens show mature fruits with turgid lobes and distinct keel, long glandular, and slightly shorter eglandular hairs on the keel, but virtually glabrous on the faces of the valves. Kirk's specimens are immature, but are hairy on the keel and valve faces, the more normal state in the species.
Considering the discrepancy between the published accounts of abundance, etc., and the virtual inability to find the species in vivo or in sicco, the position merits further consideration As far as absence of herbarium material is concerned, there is the possibility that, being a common species, collections were rarely made (this position exists with some common aliens), but this possibility is negatived by the reasonably large collections of the other common species of Veronica, and that what purported to be material of the species was either V. arvensis or V. persica. On this basis, I suggest that the species was, and still is not common, and that published accounts have been based on an erroneous concept of the species, as was instanced in Britain by Drabble and Little (1931: 180).
It could be, of course, that the behaviour of V. agrestis in New Zealand paralleled that in Britain, where there is evidence that it is becoming increasingly rarer—Edees (1945: 274–5) is quoted (Anon. 1947: 63) in respect of the species in Staffordshire as”.… formerly described as common but this ‘is not true to-day. In the last ten years… found only half-a-dozen times …. a decreasing species,….’”, and Good (1948: 196) with reference to Dorset, appears to imply a decrease in abundance of the species when he notes under V. persica Poir. “A common and widely distributed weed colonist chiefly on calcareous soils. Much more plentiful and frequent now than V. polita or V. agrestis.” Treating the present position of the species in the British Isles, Clapham et al. (1952: 886) not the species as “…… perhaps less common than formerly.”
For this premise of decreasing abundance to be valid in respect of the species in New Zealand, definite evidence of abundance in former years followed by a diminishing abundance during recent decades is necessary—such evidence is lacking by way of extant specimens, and the confusion in our concept of the species males it necessary to treat published records with doubt.
My experience with the species, based on the Akaroa occurrence, is that it appears fugitive in nature, since re-examination of the sites in February, 1949, revealed no trace of the plant, although conditions were unchanged. It is a casual, perhaps arising from periodic seed introductions, and at the present time is either rare or non-existent.
The remarks of Guthrie-Smith (1926: 275–6) concerning the habits of the plant he considered as V. agrestis, indicate a behaviour atypical of the other naturalized species of Veronica, and while the identification must be regarded with reserve, the behaviour seems more in keeping with that of V. agrestis. Of the occurrence at “Tutira,” Hawke's Bay, he notes the probable introduction in soil attached to roots of moss-roses about 1883, and (p. 276) that “the plant has never strayed far; it is a garden rather than a field species, and has for thirty years remained within a few dozen yards of the spot of its first appearance.”
My thanks are expressed to Mrs. P. Hynes, Auckland Museum and Institute, Auckland, and to Mrs. P. A. Williamson, Dominion Museum, Wellington, for the loan of herbarium material; to Messrs. P. R. Barrer, W. J. Kissock, D. M. E. Merry, and W. O. Sly, Field Officers of the Department of Agriculture, for collections of Nasturtium; to Dr. S. M. Walters, Botany School. University of Cambridge, for examination of, and comments on specimens of Aphanes submitted to him.
Allan, H. H., 1940. A Handbook of the Naturalized Flora of New Zealand N.Z.D.S.I.R Bull No 83 Wellington.
Anon. 1947. Plant Records B. E. C. Rept for 1945 vol. xiii. p. 63
Bentham, G, 1864. Flora Australiensis. Vol. II. London.
Cheeseman, T. F., 1883. The Naturalized Plants of the Auckland Provincial District. Trans, N. Z. Inst vol. XV, p. 289.
— 1906. Manual of the New Zealand Flora Edition I. Wellington.
— 1925 Manual of the New Zealand Flora Edition II. Wellington.
Clapham, A. R., Tutin, T. G., and Warburg. E. F., 1952. Flora of the British Isles. Cambridge, England.
Drabble, E., and Little, J. E., 1931. The British Veronicas of the Agrestis Group. Journ. of Botany, vol. 69, p. 180.
Edees, E. S., 1945. N. W. Nat., vol. 19. pp. 274–5 (not seen).
Good, R., D'O., 1948. A Geographical Handbook of the Dorset Flora. Dorchester. England
Guthrie-Smith, H., 1926. Tuta. The Story of a New Zealand Sheep-Station Edition II Edinburgh and London.
Hilgendorf, F. W., 1926. Weeds of New Zealand. Edition I.
— 1948. Weeds of New Zealand Edition IV. (revised J. W. Calder)
Hooker, J. D., 1852–55. Flora Novae-Zealandiae. London.
— 1864a. Handbook of the New Zealand Flora, Part I, London.
— 1864b. Note on the Replacement of Species in the Colonies and Elsewhere. Nat. Hist. Review, No. XIII, p. 124.
— 1867. Handbook of the New Zealand Flora. Part II. London.
Hyde, E. O. C., 1945. Weed Seeds in Agricultural Seed. Third Series. N.Z. J. Agric., vol. 70, p. 617.
Kirk, T., 1870. On the Naturalized Plants of New Zealand, Especially with Regard to those Occurring in the Province of Auckland. Trans. N.Z. Inst, vol. II, pp. 137. 141.
— 1878a. On the Naturalized Plants of Port Nicholson and the Adjacent District Trans. N.Z. Inst., vol. X, p. 375.
— 1878b. Contributions to the Botany of Otago. Trans. N.Z. Inst., vol. X, p. 416.
Murphy, R. C., 1951 The Impact of Man upon Nature in New Zealand. Proc. Amer. Phil Soc., vol. 5, p. 577.
Walters S. M., 1949. Aphanes microcarpa (Boiss. et Reut.) Rothm. in Britain. Watsonia, vol. I, pp. 163–9.