6===Taxonomic Revision Asplenium===

New Zealand Journal of Botany. 1977, Vol. 15: 39-86. 39

A taxonomic revision of the New Zealand species ofAsplenium

P. J. BROWNSEY

Department of Botany, Victoria University of Wellington, Private Bag,Wellington, New Zealand*

(Received 16 August 1976)

ABSTRACT

A taxonomic revision of the fern genus Asplenium in the New Zealand region has beencarried out. Fourteen species and four subspecies are recognised. One species, A. terrestre.and three subspecies, A. obtusatum subsp. northlandicum, A. flaccidum subsp. haurakiensc,and A. terrestre subsp. maritimum, are described for the first time. Typification, synonymy,range of morphological variation, chromosome number, habitat, and distribution are describedfor each taxon. A dichotomous key to the species is presented.

The origin and development of the New Zealand Asplenium flora is briefly discussed.It has its closest affinities with that of temperate Australia from where most of the specieshave probably been derived. However, some species such as A. trichomanes, A. polyodon,and A. obtusatum have affinities with Northern Hemisphere, tropical, or South Americanfloras and have therefore ultimately reached New Zealand from much further afield.

The genus displays a higher mean level of ploidy (5.9) in this country than in any otherarea for which records exist; of the 18 sexually reproducing cytotypes, 9 are tetraploid,1 is hexaploid, and 8 are octoploid. Most of the tetraploid species have distributions extendingoutside New Zealand and they are therefore considered ancestral. The higher polypoids arealmost all endemic and are thought to have originated within New Zealand from the tetraploidtaxa.

INTRODUCTION

Asplenium with over 700 species distributedthroughout the world is one of the largest generawithin the Filicopsida. The family Aspleniaceae isbelieved to be of relatively recent origin and is stillin an active state of evolution with most of thegenera and many of the species rather poorly defined(Lovis 1973, 1977). In recent years an intensivecytogenetic and biosystematic investigation of theEuropean representatives of the family has resultedin a much better understanding of the evolution andtaxonomy of this group of ferns. It is now knownthat hybridisation and chromosomal doubling havebeen of prime importance and that reticulate evolu-tion through auto- and allopolyploidy has producedseveral morphologically very similar but geneticallyquite distinct species (see reviews by Vida 1972,Lovis 1977).

Present address: National Museum, Private Bag,Wellington, New Zealand.

To ascertain whether Asplenium has developedin a comparable manner in other parts of the world,and perhaps to provide some evidence for the geo-graphic origin of this enormous genus, a similarbiosystematic investigation has been started on theNew Zealand representatives. It will be many yearsbefore the evolution of these species is as clearlydocumented as that of their European counterparts,but the preliminary results of this investigation haveprovided sufficient information for a much neededtaxonomic revision of the genus in New Zealand.Many authors (e.g., Hooker 1855, Allan 1961) haveremarked on the plasticity and difficulties of definingspecies of Asplenium in this country, but, apart froman investigation by Knowlton (1957) which wasterminated by his premature death, and somechromosome counts on individual species (Brownlic1954, 1958, 1961), no detailed study of the genushas been made in recent times. The present paperpresents new cytological and morphological evidencewhich more clearly delimits the taxa.

40 New Zealand Journal of Botany 15, 1977

MATERIALS AND METHODS

Plants were collected from throughout the NewZealand region and grown in cultivation in Welling-ton under approximately uniform environmentalconditions. Herbarium material was collected at thesame time and voucher specimens are deposited inCHR, WELT, and WELTU.

Meiotic chromosome preparations were obtainedby fixing young sporangia and staining accordingto the method described by Manton (1950, pp.295-6). Mitotic preparations were obtained by themethod outlined in Roy & Manton (1965). Materialwhich has been examined cytologically is listedimmediately after the chromosome number given foreach taxon or cytotype.

Measurements of the exospore were made fromspores mounted in gum chloral (a sample of 50spores was measured for each population). The rangeof mean values for a specified number of populationsis given for each species and subspecies, the figuresin brackets indicating the extreme sizes of individualspores. Rhizome scales have been drawn by cameralucida from permanent preparations in Canada balsamafter dehydrating in a series of ethanols and clearingin xylol.

Distribution maps covering the North, South, andStewart Islands have been prepared from herbariummaterial examined in AK, AKU, CHR, WELT,WELTU, the personal herbaria of Mr A. E. Esler(Botany Division, c/o Plant Diseases Division, DSIR,Auckland), Dr J. E. Braggins and Mr A. E. Wright(University of Auckland), and my own personalcollections. Lists of material examined have not beengiven, but the sheets in the above herbaria havebeen appropriately annotated.

Also, the occurrence of the different species onall the main islands in the New Zealand region hasbeen listed in the same abbreviated fashion as inAllan (1961); i.e., K, Kermadec Islands; N, NorthIsland; S, South Island; Ch., Chatham Islands; St.,Stewart Island; Sn., Snares Islands; A, AucklandIslands; C, Campbell Islands; Ant., Antipodes Island.

In determining the taxonomic rank to be assignedto particular taxa, precedents set in naming the Euro-pean Asplenium flora have been followed. It is hopedthat some equivalence in the treatment of this cos-mopolitan genus in different areas will thus beachieved. Hence, the level of subspecies has beenapplied to taxa which are either cytologically orecologically distinct but which can be clearly dis-tinguished morphologically only by means of micro-characters. The term variety is reserved for occas-

ional variants, or distinct forms confined to limitedgeographical areas. Species which show markedpolymorphy in many of their populations have notbeen taxonomically subdivided, but their ranges ofvariation are illustrated.

Wherever possible details of the type specimenare given. For those species first described byForster, lectotypes have been designated because itis not possible to identify the holotypes. Carolin(1963) indicates, that for species in the "Prodromus",specimens in BM and GOET have the greatest claimfor selection. These have been examined. Specimensfrom BM have been chosen as the lectotypes becausethey are of generally superior quality, those in GOETfrequently consisting only of fragments of a frond.

KEY TO THE NEW ZEALAND SPECIES

OF ASPLENIUM — see opposite page.

1. ASPLENIUM FLABELLIFOLIUM Cav. Descr.PL: 257 (1802)—as A. flavelifolium* (Fig. 1)

A. flabellifolium var. ramosum Col. Trans. N.Z.Inst. 20: 228 (1888).

•The spelling of the specific epithet in Cavanilles'original description is flavelifolium but this was latercorrected to flabellifolium by Swartz (1806; pp. 88,273). Eichler (1965) suggested that "The originalspelling . . . ought to be retained, as it is difficultto justify an alteration of this kind which is neithera correction of a typographic nor of an orthographicerror". However, the derivation of the original nameflavelifolium is obscure, unless it means yellow-leaved. This seems unlikely because (i) the leavesare not particularly yellow in this species, (ii) thereis no mention of yellow leaves in the originaldescription - indeed, they are stated as being brightgreen, (iii) if Cavanilles had wanted to use anepithet meaning yellow-leaved he would surely haveused flavifolium. Thus, the name flavelifolium iseither nonsensical or quite inappropriate, and it isextremely improbable that Cavanilles intended toproduce such a meaningless epithet for his newspecies. It seems much more likely that he meantthe name to refer to the very characteristic fan-shaped leaves of this fern which he specificallymentioned in the protologue. The name he intendedwas almost certainly flabellifolium but in publishingit as flavelifolium two kinds of errors may havebeen perpetrated - a typographic error in whichCavanilles' handwritten "b" was mistaken for a"v", and an orthographic error in which one " 1 "was used instead of two. Article 73 of the Inter-national Code of Botanical Nomenclature sanctionsthe correction of both types of error and the changefrom flavelifolium to flabelifolium to flabellifoliumis therefore entirely within reason. Hence, I haverejected Eichler's arguments for retaining the nameA. flavelifolium and followed the now widelyaccepted practice first established by Swartz.

Brownsey—N.Z. species of Asplenium 41

KEY TO THE NEW ZEALAND SPECIES OF ASPLENltjM

/ Fronds simply pinnate 2Fronds 2 — or more pinnate / /

2 Fronds < 3 cm wide 3Fronds > 3 cm wide 6

3 Stipe and rachis scaly; usually < 8 pairs pinnae 4Stipe and rachis without scales; usually > 8 pairs pinnae 5

4 Fronds thick and fleshy; pinnae with scales on underside only; plant of exposedsea cliffs 4. A. obtusatum

Fronds not thick and fleshy; pinnae often with scales on both surfaces; plant ofcalcareous soils 7. A. lyallii

5 Stipe and rachis green; fronds lax and rooting at tip 1. A. flabellifoliumStipe and rachis brown; fronds stiff and erect 2. A. trichomanes

6 Rachis dark chocolate brown throughout; pinnae irregularly and doublyserrate 3. A. polyodon

Rachis green, at least on the upper surface (sometimes turning pale brown whendried); pinnae entire or regularly serrate 7

7 Pinnae linear, > 10 times as long as broad 12. A. flaccidumPinnae < 10 times as long as broad 8

8 Pinnae prominently serrate; sori extending right to margin between serrations6. A. scleroprium

Pinnae entire or minutely serrate; sori not reaching margin 99 Lowest pinnae usually lobed or with at least one distinct pinnule; sometimes with

scales on both surfaces; plant of calcareous soils 7. A. lyalliiLowest pinnae not lobed or incised, lacking scales on upper surface 10

10 Rachis and foliar scales with very long hair-like apices (Fig. 9K-0); pinnae firmbut not thick and fleshy, glossy above, accuminate to ± acute 5. A. lucidum

Rachis and foliar scales ovate to narrowly ovate with short hair-like apices(Fig. 9A-J); pinnae thick and fleshy, dull above, obtuse to ± acute

4. A. obtusatum/ / Fronds bearing bulbils 9. A. bulbiferum

Fronds not bearing bulbils 1212 Rhizome creeping; fronds thin, glossy above 8. A. lamprophyllum

Rhizome not creeping; fronds dull above 1313 Sori remote from margin, 4-10 mm long 7. A. lyallii

Sori submarginal, or, if remote from margin, < 4 mm long 1414 Fronds 3-4 pinnate at base; ultimate segments long and narrow, but broadened

in region of sorus; plant of Kermadecs 14. A. shuttleworthianumFronds 2-pinnate at base, or, if 3-4 pinnate, ultimate segments not broadened in

region of sorus 1515 Fronds 3-4 pinnate at base; ultimate segments linear, mostly < 1 mm wide; pinnae

and pinnules crowded and overlapping; stipe stout 11. A. richardiiFronds 2-3 pinnate at base; not with all above combination of characters 16

16 Fronds thin; sori 2-4 mm long 17Fronds thick and leathery or fleshy; sori 2-10 mm long 18

(A. flaccidum agg.)17 Fronds with < 15 pairs pinnae (if c. 15 pairs pinnae, pinnules with distinct but very

slender stalks) 10. A. hookerianumFronds with > 15 pairs pinnae (if c. 15 pairs pinnae, pinnules sessile or on very

short broad stalks) 9. A. bulbiferum18 Spores* pale yellow, surface granular but with few or no prominent ridges

(Fig. 29 M-P) 19Spores* dark brownish yellow, surface with many very prominent ridges

(Fig. 30 O-R) 2019 Fronds limp and flaccid; plant usually epiphytic 12a. A. flaccidum subsp. flaccidum

Fronds firm and erect; plant of exposed sea cliffs, confined to east coast regionsnorth of lat. 38° 12b. A. flaccidum subsp. haurakiense

20 Fronds thick and fleshy; plant of exposed sea cliffs13b. A. terrestre subsp. maritimum

Fronds leathery, but not fleshy; plant of forest floor or rocky inland areas13a. A. terrestre subsp. terrestre

•Spores mounted in gum chloral or other suitable medium and viewed at c. X 200.


Fig. 1 Fronds of Asplenium flabellifolium ( x 1). Octoploid cytotype (A-C), hexaploid cytotype (D-F). A.NZ 191, Eastbourne, Wellington; B, NZ 834, Turakirae Head, Wellington; C, NZ 847, Seatoun, Wellington;D, NZ 177, Kapiti Island; E, NZ 887, Rangitoto Island; F, NZ 789, Wharariki Beach, NW Nelson.

TYPE: In MA (not seen).

DESCRIPTION: Rhizome short, erect, bearing darkbrown subulate scales with long filiform apices upto 6 X 0.5 mm (Fig. 4M-S). Stipes 1-10 cm long,green, slender, lacking scales. Laminae linear,7-25 X 1-2.5 cm, light green, flaccid and often pros-trate, pinnate. Raches green, weak, lacking scales,frequently extending beyond the uppermost pinnaeand rooting at their apices. Pinnae 8-25 pairs,gradually decreasing in size from base to apex, fan-shaped, crenate-dentate, cuneate at base, lowest 5-15X 5-20 mm. Sod radiating along veins, up to 6 mmlong. Spores (37)43-51(62) Mm long, (30)36-41(51)fim wide (8 populations), perispore with numerouscrests and ridges (Fig. 29C-D).

CHROMOSOME NUMBER: 2n = 210 or 211 (Fig. 31A).NZ 887, Rangitoto Island; NZ 216, Stronvar, Waira-rapa; NZ 177, 178, Kapiti Island; NZ 789, Whara-riki Beach, NW Nelson.

2n = 277-280 (exact number uncertain) (Fig. 31B).NZ 181, 186, Eastbourne, Wellington; NZ 314, Karori,Wellington; NZ 834, Turakirae Head, Wellington;NZ 847, Breaker Bay, Wellington; NZ 775, PortHills, Christchurch.

HABITAT AND DISTRIBUTION: N, S. A. flabellifoliumis found in scattered localities from the Bay of Islandsto Otago (Fig. 2), but it is absent from large areasof the west coast of both islands and from the farsouth. It grows from sea level to 1 250 m in dry rockyground, on lava and scoria fields, in grassy areasbelow scrub, and sometimes in open beech forest; itnever occurs in deep forest. Its greater abundance inopen scrub and the margins of agricultural land,particularly in Canterbury and the Wairarapa, sug-gests that it may be a species which has considerablyextended its range with the advent of Europeancolonisation. Wherever it occurs, it appears as a pros-trate plant and spreads rapidly by its rooting tips.


Fig. 2 Distributions ofAsplenium flabellifolium andA. polyodon.

A. flabellifolium

A. polyodon

COMMENT: A. flabellifolium is a very distinctivespecies confined to New Zealand and parts of southernAustralia and Tasmania. It was first investigatedcytologically by Brownlie (1958) who reportedn. = c.27O in plants from Lyttelton, indicating thatthis was a sexual c.l6-ploid species (the base number(x) in Asplenium is 36).

More recently, Lovis (1973) investigated a collec-tion from Australia and showed that there werec.212 univalents present at meiosis. Prothalli raisedfrom spores of this collection proved to be apogam-ous and Lovis therefore pointed out that the taxonwas an approximately hexaploid apomict of theAsplenium aethiopicum type*. Because plants of A.flabellifolium from Australia and New Zealand arevery similar morphologically, this evidence clearly

*i.e., reduction of the chromosome number is circum-vented by nuclear restitution after asynaptic meiosis(Braithwaite 1964) and not by a failure of mitosisin the formation of the spore mother cells, as inmost apomictic ferns (Manton 1950, Walker 1966).

suggested the possibility that the material investigatedby Brownlie was not a sexual c. 16-ploid species butan approximately octoploid apomict.

During the present investigation, plants of A.flabellifolium from several localities have beenexamined at meiosis and, in every case, a very dis-tinctive diakinesis has been observed in which onlyunivalents are present (Fig. 31 A, B). This confirmsthe fact that A. flabellifolium is an apomictic speciesin New Zealand as well as Australia. Two separatechromosome numbers have been recorded, 2n = 210~211 and 2n = 277-280. The lower number clearly cor-responds to that reported in Australian material, but,although there is some slight doubt as to whether theexact number is 210 or 211, it seems certain that itis not as high as the 212 suggested by Lovis (1973).The number of the higher cytotype has been deter-mined with less accuracy, but it is sufficiently clearto indicate that in reporting n = c.27O Brownlie (1958)mistook the univalents in this apomictic species forbivalents, thereby implying a level of plbidy twiceas high as is the case.

44 New Zealand Journal of Botany 15. 1977

The two cytotypes of A. flahellifolium representapproximately hexaploid and octoploid levels ofploidy, but it is now certain that their chromosomenumbers do not correspond to exact multiples of36. Thus, one would anticipate 216 and 288 chromo-somes respectively, but the actual numbers of 210-211 and 277-280 indicate that some aneuploidy hasoccurred. One rather large chromosome is regularlypresent in both cases and, as suggested by Lovis(1973), some of the missing chromosomes may havebeen absorbed by fusion.

No constant morphological distinction of any kindhas been observed between the two different cyto-types and neither are they obviously restricted toparticular localities. Hence, they have not beenaccorded any separate taxonomic status.

2. ASPLENIUM TRICHOMANES L Sp. PI.:1080 (1753).

DESCRIPTION: Rhizome stout, erect, bearing darkbrown subulate scales up to 5 mm long. Stipes 1-10 cmlong, dark brown, shining, stiff, lacking scales exceptat the very base. Laminae linear, 5-30 cm long, darkgreen, sub-coriaceous, pinnate. Raches dark brown,shining, stiff, lacking scales. Pinnae sub-sessile, 15-25(or more) pairs, oblong to ± orbicular, crenate-serrate to ± entire, 2-10 X 1-5 mm. Sori up to 3 mmlong.

CHROMOSOME NUMBER: TWO distinct chromosomenumbers have been reported for this species in NewZealand, n = 72 and n = 108 (Brownlie 1954; Lovis1956, 1958).

HABITAT AND DISTRIBUTION: N, S. A. trichomanesdisplays a considerable range of morphological andcytological variation over its virtually cosmopolitandistribution. A world-wide biosystematic investigationof this intricate group is in preparation (Lovis, unpub-lished data). However, in New Zealand A. tricho-manes is a very distinctive fern and is apparentlyquite unrelated to any other native species with thepossible exception of A. flabellifolium. In a detailedbut unpublished study of A. trichomanes in NewZealand, Lovis (1956) showed that two differentcytotypes were present which could only be dis-tinguished morphologically by the structure of therhizome scales and by the size and markings of thespores (mean values: 34-39 /nn long in the tetraploidand 39-47 pm long in the hexaploid). The tetraploid.which in Europe is the most common cytotype, israre here and apparently confined to local areas ofthe North Island. The second cytotype, a hexaploid.is the characteristic form in New Zealand. It iswidespread in the South Island from sea level to

900 m, occurring on both limestone and greywackerock, but in the North Island it is confined toscattered localities on calcareous soil. Outside NewZealand it is known from only a single locality inAustralia (Gudgenby Creek, Australian CapitalTerritory — Lovis, pers. comm.).

3. ASPLENIUM POLYODON Forst. f. Prodr.: 80(1786) (Fig. 3)

Trichomanes adiantoides L. Sp. PI. 2: 1098(1753)

Asplenium falcatum Lam. Encyc. 2: 306 (1786)(torn, abort.

Tarachia polyodon (Forst. f.) Presl. Epim. Bot.:77 (1851)

Asplenium adiantoides (L.) C. Chr. Ind. Fil.:99 (1906) non Lam. 1786, non Raddi 1819.non Raoul 1844

Asplenium adiantoides var. polyodon (Forst. f.)C. Chr. Ind. Fil.: 126 (1906)

Asplenium forsterianum Col. Tasm. J. nat. Sci.2: 171 (1846)

Asplenium falcatum var. caudatum (Forst. f.)Allan, Fl. N.Z.: 71 (1961)

TYPE: Not located (not in BM or GOET).

DESCRIPTION: Rhizome stout, short creeping, denselycovered in red-brown, narrowly triangular scales upto 10 X lmm. Stipes 10-30 cm long, dark brown,still", densely covered in scales similar to but smallerthan those of the rhizome (Fig. 4A-E). Laminaelanceolate, 25-50 (or more) X 10-20 cm, dark greenand glossy above, paler and dull below, frequentlypendulous, pinnate. Raches dark chocolate brown,very scaly. Pinnae 25 (or more) pairs, narrowlyangular-ovate to ovate, sometimes with a largerounded basal acroscopic lobe, acuminate, doublyserrate, 5-10 x 1—2 cm, scaly and with prominentveins on underside. Sori often slightly curved awayfrom the midrib, up to 2 mm long. Spores (26)30-37(40) fim long, (15)18-21(25) /an wide (6 popula-tions), perispore with a few very prominent ridges(Fig. 30A-B).

CHROMOSOME NUMBER: 2n = 144. NZ 679, SpraggsBush, Waitakere Ranges, Auckland; NZ 377, Bay ofMany Coves, Marlborough Sounds.

Tetraploid chromosome numbers have also beenrecorded for this species in Australia and Ceylon(Manton & Sledge 1954), Malaya [Malaysia] (Manton1954), and the Himalayas (Bir 1960).

HABITAT AND DISTRIBUTION: K. N, S, Ch., St. A.polyodon occurs widely in forested areas from NorthCape to Stewart Island, though it is now absentfrom most of the east coast of the South Island

Brovvnsey—N.Z. species of Asplenium 45

A

Fig. 3 Fronds of Asplenium polyodon ( x | ) . A. NZ 864. Carter's Bush. Wairarapa; B. NZ 928, KapitiIsland; C. NZ 1069. Lighthouse Track. Stewart Island.

Q RFig. 4 Camera lucida drawings of scales ( x 20), each taken from a different population. A-E, stipe scalesfrom Asplenium polyodon; F-L, stipe scales from A. lyallii (F & I from bipinnate plants, others from pinnatespecimens); M-S, rhizome scales from A. ftabellifolium (M, N, P from hexaploid cytotype; O, Q, R, S fromoctoploid cytotype).


where forest has been cleared (Fig. 2). It is foundprincipally as an epiphyte from sea level to c. 900 m,but it also grows on fallen logs and even on theground. In the far south, A. polyodon growsterrestrially in coastal forest and scrub, and oftendevelops into very large plants. A substantialdistance separates the plants in these localities fromthose in Westland which are more typically epiphytic,but whether this apparent disjunct distribution is realneeds further investigation. There is nothing to suggestthat the terrestrial plants are morphologically differentto the epiphytic ones apart from their larger size.

COMMENT: Until recently this fern has been widelyreferred to as either A. adiantoides (L) C. Chr. (e.g.Cheeseman 1925) or A. falcatum Lam. (e.g., Allan1961, Crookes 1963). However, Morton (1967) hasshown that the correct name is A. polyodon Forst. f.The confusion has arisen because the original descrip-tion of this species (as Trichomanes adiantoides L.by Linnaeus in "Species Plantarum'") was based ontwo separate elements - one from Ceylon illustratedby Burmann ("Thesaurus Zeylanicus", t.43, 1737),and one from Africa illustrated by Plukenet ("Phyto-geographia", t.123, 1692). These are now knownto be quite distinct species.

Lamarck was the first to realise this, but unfor-tunately compounded the existing confusion by retain-ing Linnaeus' specific epithet for the African plantwhich he called Asplenium adiantoides Lam. How-ever, it seems that the type of Trichomanes adian-toides L. was actually the plant from Ceylon (Morton1967) to which Lamarck erroneously gave the newname Asplenium falcatum Lam. and cited the Bur-mann illustration, but which he should have calledA. adiantoides (L.) Lam. The name A. falcatumLam. is therefore illegitimate because the type of theLinnean species Trichomanes adiantoides was quotedin synonymy.

Christensen tried to remedy this error by proposingthe new combination Asplenium adiantoides (L.) C.Chr. (Ind. Fil.: 99, 1906) for the Ceylon material(i.e., that which Lamarck had called A. falcatum).However, this name is also illegitimate, being a laterhomonym three times over (A. adiantoides Lam.Encyc. 2: 306, 1786; A. adiantoides Raddi, Opusc.Sci. Bol. 3: 291, 1819; A. adiantoides Raoul, Ann.Sci. nat. Ser. Ill 2: 115, 1844).

The oldest acceptable name for this species com-plex is therefore A. polyodon, first proposed byForster in his "Prodromus" but until recently quotedin synonymy with either A. falcatum Lam. or A.adiantoides (L.) C. Chr. The species extends through-out much of the Indo-Malaysian, Australian, andPacific regions and is very variable. However, Skotts-berg (1942) made a thorough investigation of the

taxon throughout its range of distribution and con-cluded that plants from Mauritius, Timor, thePhillipine Islands, New Zealand, Australia, and NewCaledonia were basically similar.

Within the New Zealand region, further nomen-clatural confusion has been caused by the existence ofa rather narrow-pinnuled form of A. polyodon onthe Kermadec Islands. This material has sometimesbeen recognised as A. caudatum Forst. f. (e.g.,Cheeseman 1925) or taxonomically distinguished as anew variety, A. falcatum var. caudatum (Forst. f.)Allan (e.g., Allan 1961, Crookes 1963). However,it is clear from examination of Forsters material inBM and GOET that A. caudatum is a distinct specieswhich has deeply dissected, more or less parallel-sided pinnae bearing sori only c. 4-5 mm long. Thereis no evidence to suggest that Forster's A. caudatumwas collected in New Zealand and plants comparableto it do not grow in this region today. Copeland(1932) indicates that the type locality was probablyin the Society Islands and that this species has adistribution extending from the New Hebrides toHawaii. The plants on the Kermadecs are thereforeprobably best regarded as one extreme of the rangeof variation in the polymorphic A. polyodon complex.

4. ASPLENIUM OBTUSATUM Forst. f. Prodr.:80 (1786) (Figs 5, 6)

A. obliquum Forst. f. Prodr.: 80 (1786)A. ohtusatum var. obliquum (Forst. f.) Hook. f.

Fl. Antarct. 1: 108 (1844)A. apice-dentatum Homb. et Jacq. Voy. Pole

Sud: pi. 1A (1853)A. lucidum var. obliquum (Forst. f.) Moore,

Ind. Fil.: 142 (1859)

TYPE: Lectotype in BM; New Zealand, coll. J. R.and G. Forster.

DESCRIPTION: Rhizome stout, often forming a hardwoody mass above ground, bearing brown, shiny,ovate, acuminate scales up to 15 X 4 mm. Stipes2-30 cm long, dark brown at base, green above, stout,covered in ovate or narrowly triangular scales withacute or acuminate apices. Laminae ovate to lanceo-late, 4-40 X 2-20 cm, dull green, very thick andfleshy, pinnate. Raches green, stout, prominentlygrooved and scaly. Pinnae 4-18 pairs, narrowly ovateto oblong, obtuse to acute, crenate-serrate to ± entire,cuneate at base, 10-100 X 7-25 mm, scaly on theunderside. Sori up to 10 mm long, not reaching tolamina edge. Perispore of spores slightly winged butwith few crests or ridges (Fig. 29E-H).

New Zealand Journal of Botany 15, 1977

Fig. 5 Fronds of Asptenium obtusatum subsp. obtusatum ( x J). A, NZ 1045, Curio Bay. SE Otago; B, NZ1059, Maori Bay, Stewart Island; NZ 471, Cape Foulwind, Westport; D, NZ 1170, Ship Cove, S Westland;E, NZ 1058, Maori Bay. Stewart Island; F, NZ 996, Tomahawk Beach, Dunedin.


4a. A. OBTUSATUM Forst. f. subsp.OBTUSATUM (Fig. 5)

DIAGNOSIS: Stipes 4-30cm long, covered in ovatescales (Fig. 9A-E). Laminae usually ovate, 6-40X 5-20 cm. Pinnae 5-15 pairs, 20-100 X 10-25 mm.Spores (35)40-43(49) Mm long, (21)24-27(33) ^mwide (5 populations).

CHROMOSOME NUMBER: n = 72 (Fig. 31C). NZ 977,

Cape Terawhiti, Wellington; NZ 472, Cape Foulwind,Westport; NZ 465, Punakaiki; NZ 998, TomahawkBeach, Dunedin.

The same chromosome number has also beenrecorded for this taxon from Stewart Island (Brownlie1958) and Tristan da Cunha (Manton & Vida 1968).

HABITAT AND DISTRIBUTION: N, S, Ch., St., Sn., Ant,

A, C. (Fig. 7). A. obtusatum subsp. obtusatum iscommonest in the South Island where it occurs moreor less continuously around the west and south coastsfrom Cape Farewell to Moeraki Point; there is alsoan outlying population on Banks Peninsula. InNelson, the Marlborough Sounds, and on the eastcoast as far south as Banks Peninsula it is apparentlyreplaced by A. lucidum (Fig. 7). In the Cook Straitregion there is a population on Brothers Island, butthe only recently confirmed record from the NorthIsland is at Cape Terawhiti*.

In the far south, A. obtusatum is abundant aroundthe coasts of the islands of Foveaux Strait and onthe sub-Antarctic islands.

In the northern part of its New Zealand range,subsp. obtusatum is a rather small plant found grow-ing in isolated populations only on the most exposedcoastal rocks. In the south, however, it growsluxuriantly not only on exposed cliffs but also furtherback in the shelter of the coastal vegetation whereit attains a height of 1 m or more and often formsan important association with Hebe elliptica (Forst.f.) Pennell and Blechnum durum (Moore) Chr.Nowhere, however, does it extend beyond theinfluence of salt spray.

COMMENT: The Forster material of A. obliquumin BM and GOET has been examined. In none ofits characters does it exceed the range of variabilityin A. obtusatum subsp. obtusatum and, hence, plantsreferred to it by Forster and later workers shouldbe regarded as merely large forms of A. obtusatum.Field work in localities from which such plants have

been collected indicates that there is often a com-plete range of forms within a few yards, from smallplants in the most exposed sites to much largerplants sheltered by scrub vegetation. The differencesin morphology are entirely due to changes in theenvironment, and, because the various forms areregularly found together in close proximity, thereseems little justification for maintaining any taxo-nomic distinction between them.

4b. A. OBTUSATUM Forst. f. subsp. NORTH-LANDICUM Brownsey subsp. nov. (Fig. 6)

Planta plerumque parvior quam subsp. obtusatum.Stipites 2-8 cm longi, paleis anguste triangularibusvestiti (Fig. 9F-J). Laminae lanceolatae vel ovatae,4-20 X 2-7 cm. Pinnae 4-18 jugatae, 10-40 X7-12 mm. Sporae (39)45-52(60) /on longae, (25)29-33(38) /ITO. latae (Fig. 29G-H). Numerus somaticuschromosomatum 288; in meiosi paria 144 observantur(Fig. 31E).

HOLOTYPUS: CHR; NZ 667, Karekare, Auckland(NZMS 1 N41:000432). Growing in crevices on thecliffs at the south end of the beach. Coll. P. J.Brownsey, 19.3.1974. (CHR 308924)

DIAGNOSIS: Generally a smaller plant than subsp.obtusatum. Stipes 2-8 cm long, covered in narrowlytriangular scales (Fig. 9F-J). Laminae lanceolate toovate, 4-20 X 2-7 cm. Pinnae in 4-18 pairs, 10-40X 7-12mm. Spores (39)45-52(60) Mm long, (25)29-33(38) /j.m wide (6 populations).

cm

*In WELT there are old herbarium specimens fromKapiti Island and from Somes Island in WellingtonHarbour.

10-

5-

0 J

Fig. 6 Fronds of Asplenium obtusatum subsp.northlandicum ( x 1). A, B, NZ 667, Karekare,Auckland; C, NZ 603, Little Barrier Island.

50 New Zealand Journal of Botany IS, 1977

CHROMOSOME NUMBER: n = 144 (Fig. 31E). N7. 667,

Karekare, Auckland.

HABITAT AND DISTRIBUTION: K, N. Apart from onelocality at Karekare, A. obtusatum subsp. north-landicum is found only on the east coast of theNorth Auckland peninsula extending from the ThreeKings to Slipper Island off Coromandel (Fig. 7). Itis separated from the tetraploid subspecies by virtuallyall the remaining coastline of the North Island. Itgrows only on coastal rocks or amongst scrubexposed to salt spray, but it is a rather small plantand is not as important a constituent of the coastalvegetation as the southern subspecies. It is so farnot known outside New Zealand.

COMMENT: The disjunct distribution of A. obtusatumin New Zealand is accompanied by a change inthe chromosome number, populations in thesouth being tetraploid and those in the north

octoploid, but the gross morphology of the twocytotypes is very similar. The tetraploid differs signi-ficantly only in having stipe scales which are morebroadly ovate (cf. Fig. 9A-E, F-J) and spores whichare smaller than those of the octoploid (cf. Fig. 29E.G). Despite their geographic separation, the twocytotypes can only be distinguished morphologicallyby these micro-characters and hence they have notbeen accorded full species status.

A. obtusatum occurs commonly around thesouthern coastlines of Australia and South Americaas well as on many islands of the southern Atlanticand Pacific Oceans. Material from Tristan da Cunhais known to be tetraploid (Manton & Vida 1968)and therefore presumably corresponds to subsp.obtusatum. Collections from elsewhere have yet tobe cytologically examined, but, because of the dis-covery of two distinct cytotypes in New Zealand, itis very desirable that this species be more thoroughlyinvestigated on a world-wide basis.

A. obtusatum

• subsp.northlandicum

• subsp.obtusatum

A. lucidum

Fig. 7 Distributions ofAsplenium obtusatum andA. lucidum.


5. ASPLENIUM LUCIDUM Forst. f. Prodr.: 80(1786) (Fig. 8)

A. oblongifolium Col. Tasm. J. nat. Sci. 2: 171(1846)

A. lucidum var. paucifolium Hook. f. Handbk.N.Z. Fl.: 371 (1864)

A. iTurvillei Mett. Kuhn Linn. 36: 95 (1869)A. obtusatum var. lucidum (Forst. f.) Hook, et

Baker Syn. Fil.: 207 (1873)A. obtusatum var. integrifolium Szyszyl. in

Wawra, Itin. princ. S. Coburgi: 122 (1888).

TYPE: Lectotype in BM; coll. G. Forster (typelocality given in "Prodromus" is New Zealand).

DESCRIPTION: Rhizome stout, often forming a hardwoody mass above ground, bearing pale brown,shiny, ovate, acuminate scales up to 30 X 7 mm.Stipes 8-20 cm long, dark brown, stout, denselycovered in narrow scales with very long filiformapices (Fig. 9K-O). Laminae oblong to elliptic,18-100 X 10-35 cm, dark green and glossy above,pinnate. Raches brown below, green above, stout,slightly ridged, scaly. Pinnae 4-15 pairs, lanceolateto narrowly oblong or ovate, acuminate, crenate-serrate to ± entire, cuneate at base, 4-15 X 1-3 cm,frequently covered in very small hair-like scales onthe underside. Sori up to 20 mm long, not reachinglamina edge. Spores (27)33-38(44) fim long, (18)20-24(26) /im wide (5 populations), perispore a com-plex reticulate network (Fig. 29A-B).

CHROMOSOME NUMBER: n=72 (Fig. 31D). NZ 885,

Rangitoto Island; NZ 462, Greymouth.

Brownlie (1958) reported n = 72 in material ofthis species from Queen Charlotte Sound.

HABITAT AND DISTRIBUTION: K, N, S, Ch. (Fig. 7).

A. lucidum is found commonly in the North Islandwhere it occupies a wide variety of habitats fromcoastal cliffs to deep forest and occurs both terres-trially and epiphytically. It is abundant in coastalregions but becomes less frequent towards the vol-canic plateau. In the South Island it is reasonablycommon in coastal areas of NW Nelson and in theMarlborough Sounds, but is rare further south appar-ently reaching no further than Greymouth on theWest Coast and Banks Peninsula in the east*. Itsabsence from the central North Island and from thefar south is probably due to an inability to toleratecold.

*I have found many herbarium specimens originat-ing from further south and labelled as A. lucidum,but these have all proved to be A. obtusatum subsp.obtusatum.

Cheeseman (1925) and Allan (1961) state thatA. lucidum has been recorded on Norfolk Island andLord Howe Island. However, collections from LordHowe Island have been variously identified as A.lucidum, A. milnei Carr., and A. obtusatum (fordetails, see Oliver 1917). Clearly, further investi-gation is required before the existence of A. lucidumoutside the New Zealand region can be confirmed.

COMMENT: Taxonomists have previously relied onthe texture, colour, and shape of the pinnae to dis-tinguish A. obtusatum from A. lucidum. However,these characters vary greatly in different environ-ments and considerable confusion has surroundedthe identity and distribution of the two taxa, evento the extent that they have sometimes been reducedto varieties of the one species. However, there is nodoubt that two quite separate species do exist andthat in most cases they can be satisfactorily distin-guished. The principal source of confusion appearsto have been the very large plants of A. obtusatumwhich occur in the far south and which superficiallyresemble A. lucidum in their general habit and pinnashape. However, A. lucidum is clearly distinct in thatit has thinner pinnae with acuminate apices and avery glossy upper surface; the stipe scales are narrowwith extremely elongated hair-like apices (Fig.9K-O); the spores have a very characteristic patternconsisting of a complex reticulate network (Fig.29A-B) and they are also slightly smaller (meanvalues 33-38 X 20-24 /tm). A. obtusatum is charac-terised by fleshy pinnae which have obtuse apicesand are usually dull green; the stipe scales arebroadly ovate without long hair-like projections (Fig.9A-E); the spores have a slightly winged and ridgedperispore (Fig. 29E-F), and they are somewhat larger(mean values 40-43 X 24—27 /im in subsp. obtusatum)than those of A. lucidum. On this basis it is nowbelieved that A. lucidum is entirely replaced in thefar south by A. obtusatum subsp. obtusatum, and thatA. lucidum extends only as far as Greymouth andBanks Peninsula in the South Island. On the southernedge of its distribution, particularly on the WestCoast, plants of A. lucidum are unusually small(Fig. 8A) and their stipe scales sometimes lack verylong hair-like apices. Nevertheless, their spore charac-ters indicate that they belong to A. lucidum and notto A. obtusatum.

The two species may also be confused in the farnorth where both A. lucidum and A. obtusatumsubsp. northlandicum occupy similar habitats on seacliffs of the Hauraki Gulf area. The stipe scales arenot always a reliable taxonomic guide in this regionbecause those of A. obtusatum subsp. northlandicumare narrower and have longer apices than those ofsubsp. obtusatum (cf. Fig. 9F-J, A-E). Never-theless, the important difference in chromosome

52 New Zealand Journal of Botany IS. 1977

c

DFig. 8 Fronds of Asplenium lucidum (x i ) . A, NZ 809, Takaka Hill, Nelson; B, NZ 913, Kapiti Island;C. NZ 1171, Buller Gorge, Nelson; D, NZ 532, Little Barrier Island.

Brownsey—N.Z. species of Aspleniutn

Fig. 9 Camera lucida drawings of stipe scales ( x 20), each from a different population. A-E, Aspteniumobtusatum subsp. obtusatum; F-J, A. obtusatum subsp. northlandicum; K-O, A. lucidum; P-R, A. scleroprium.


number between A. lucidum and A. obtusatum subsp.northtandicum is manifested in the much larger sporesize of the latter (mean values 33-38 X 20-24 /j.min A. lucidum, 45-52 X 29-33 ,um in A. obtusatumsubsp. northlandicum), and the two taxa may bereliably distinguished on this basis alone.

6. ASPLENIUM SCLEROPRIUM Homb. et Jacq.Voy. Pole Sud: pi. ID (1853) (Fig. 10)

A. flaccidum var. aucklandicum Hook. f. Fl.Antarct. 1: 109 (1844)A. lucidum var. scleroprium Moore, Ind. Fil.:

142 (1859)A. lucidum var. aucklandicum (Hook, f.) Allan,

Fl. N.Z.: 1012 (1961)A. aucklandicum (Hook, f.) Crookes, N.Z. Ferns

ed. 6: 334 (1963)

TYPE: Not located.

DESCRIPTION: Rhizome short, stout, erect, bearingred-brown, ovate, acuminate scales up to 25 x 4 mm.Stipes 15-50 cm long, green on upper surface, brownbelow, stout, covered in narrowly triangular scaleswith long filiform apices (Fig. 9P-R). Laminaelanceolate, 15-50 X 8-20 cm, dark green, thick andleathery, pinnate. Raches green, stout, prominentlygrooved and scaly. Pinnae 6-17 pairs, lanceolate,acuminate, often broadened and lobed towards cuneatebase, deeply and regularly crenate-serrate, 5-10 X1-2 cm, scaly on underside. Sori reaching the laminamargin in sinuses, not more than 10 mm long. Spores(38)48-53(61) Mm long, (24)28-33(42) pm wide (5populations), perispore slightly winged, but with fewridges (Fig. 29I-J).

CHROMOSOME NUMBER: n=144 (Fig. 32C). NZ1010, 1012, Bluff, Southland.

HABITAT AND DISTRIBUTION: S, St., Sn., A. A.scleroprium is an endemic species known only froma very few collections most of which have beenmade on the Auckland Islands; others originatefrom the Snares, from one locality on the mainlandat Bluff, and from Ernest, Big South Cape, andCodfish Islands around Stewart Island. (The site ofan early collection at Half Moon Bay was notrelocated during recent field work on Stewart Island.)The species is confined to coastal areas and oftengrows alongside A. obtusatum on cliffs and in scrubexposed to salt spray.

COMMENT: Accounts of the coastal vegetation onislands in the far south (e.g., Fineran 1966a, b; 1969;1973) frequently mention a very variable mixture ofplants which have usually been referred to the A.

obtusatum complex. It is now clear that most ofthese plants are forms of A. obtusatum subsp.obtusatum varying according to their degree ofexposure on the cliff. However, there is no doubtthat a second species, A. scleroprium, also occurs inthis habitat. Previously, this taxon has been con-sidered a variety of both A. lucidum and A. flaccidum,and it has also been confused with A. obtusatumsubsp. obtusatum. However, by virtue of its octoploidchromosome number, it is cytologically distinct fromall these taxa, and may be further distinguished byits larger spore size. In gross morphology it has somesimilarity to both A. flaccidum and A. terrestre (cf.Figs 10, 21, 25) but its pinnae are generally broaderand less deeply divided, and they are characteristic-ally glossy green on the upper surface. From A.lucidum (Fig. 8) and the two subspecies of A.obtusatum (Figs 5, 6) it may be distinguished byits narrower more lanceolate pinnae which havedeeper and more regular serrations, and because thesori extend to the edge of the pinna in the sinuses;in A. lucidum and A. obtusatum the sori stop shortof the lamina margin.

7. ASPLENIUM LYALLII (Hook, f.) Moore Ind.Fil.: 143 (1859) (Fig. 11)

A. lucidum var. lyallii Hook. f. Fl. N.Z. 2: 33(1855)

A. anomodum Col. Trans. N.Z. Inst. 15: 309(1883)

A. lucidum var. anomodum (Col.) Cheeseman,Man. N.Z. Fl.: 991 (1906)

TYPE: Holotype in K; Otago, Middle Island, N.Z.,coll. Lyall.

DESCRIPTION: Rhizome stout, ascending, bearingbrown subulate to narrowly triangular scales up to25 X 2 mm. Stipes 3-20 cm long, pale brown at baseand on underside, green elsewhere, covered withscales similar to but smaller than those of therhizome (Fig. 4F-L). Laminae lanceolate to elliptic,4—40 X 2—20 cm, dark shiny green to dull grey-green,pinnate to bipinnate. Raches normally green, occa-sionally brown on the underside, often grooved,scaly. Pinnae 2-16 pairs, ovate to narrowly oblong,obtuse to acuminate, serrate to ± entire, stalked,1-10 X 0.5-5 cm, often covered on both surfaceswith tiny scales. Lowermost pinnae normally at leastpartially lobed or divided at the base, sometimescompletely pinnate; pinnules stalked, lanceolate toelliptic, obtuse, serrate to entire, up to 3 X 1.5 cm.Sori up to 10 mm long, not reaching lamina edge.Spores (36)40-54(60) Mm long, (23)27-35(40) Mmwide (13 populations), perispore prominently wingedand ridged (Fig. 29K-L).


Fig. 10 Fronds of Asplen-ium scleroprium ( x | ) .A-C. NZ 1016. Bluff.

CHROMOSOME NUMBER: n=144 (Fig. 32D). NZ 268,

Manaia, Taranaki; NZ 744, Woodside Creek, Marl-borough; NZ 277, Kaikoura Peninsula; NZ 854, DanRogers Gully, Banks Peninsula; NZ 986, Castle Hill.Canterbury.

These results agree with those of Brownlie (1958)who reported n=144 in populations from Lytteltonand Weka Pass, Canterbury. No evidence has beenfound to confirm the finding by Knowlton (1957)of 2n = c.l44 in populations of A. anomodum fromPuketautahi. Hawke's Bay and 2n = 216 in A. lyalliifrom the Taieri River mouth, Otago.

HABITAT AND DISTRIBUTION: N, S, Ch., St. A. lyalliiis an endemic species largely confined to calcareoussoils and found at various sites along the drier eastcoast and in limestone upland areas of the interior(Fig. 12). In the North Island it occurs principallyin the mountain areas of Hawke's Bay and alongthe Wairarapa coast, but there are also scatteredlocalities on the Egmont coast and at Te Kuiti, itsnorthernmost limit. In the South Island there aremore extensive populations in the limestone areas ofNW Nelson and Canterbury, and coastal populationsin Marlborough, SE Otago, Stewart Island, and partsof Fiordland. It occurs from sea level to 1 400 m. On

GFig. 11 Fronds of Asplenium lyallii ( x i ) . A, NZ 801, Takaka Hill, Nelson; B, NZ 786, Castle Hill, Canter-bury; C, D, NZ 763, Mt Pleasant, Port Hills, Christchurch; E, NZ 730, Woodside Creek, Marlborough; F,NZ 722, Woodside Creek, Marlborough; G, NZ 504. Manaia, Taranaki.


A. lyallii

Fig. 12 Distribution of Asplenium lyallii.

the coast it grows in open forest or forest remnants,particularly amongst the roots of Corynocarpuslaevigatus J. R. et G. Forst. At higher altitudes it isfrequently found in crevices or beneath overhangsof limestone rock.

COMMENT: AS understood here, A. lyallii includesall the forms which have previously been assignedto the two taxa A. anomodum and A. lucidum var.lyallii. The latter was first described by Hooker(1855) who indicated that it differed from A.lucidum only by possessing bipinnate fronds. How-ever, Moore (1858-9) evidently attached greatersignificance to this character and raised the taxonto specific rank as A. lyallii.

A. anomodum is a later name coined by Colenso(1883) to describe some apparently new and dis-tinctive pinnate plants. Colenso presumably believedthat his A. anomodum was entirely different to A.lyallii because, although he pointed out characters by

which A. anomodum could be distinguished fromboth A. obtusatum and A. hookerianum, he foundit unnecessary to mention A. lyallii.

However, as more collections were made and theknown ranges of variability of A. anomodum andA. lyallii increased, it became evident that they wereclosely related and that the distinctions betweenthem were not so clear. Subsequent authors havetreated this situation in various ways. Cheeseman1906) reduced both taxa to varieties of A. lucidum.although in the second edition of his "Manual"(Cheeseman 1925) he reinstated A. anomodum as afull species distinct from A. lucidum var. lyallii inthe size, shape, and cutting of the frond. Allan(1961) retained A. anomodum but rejected A. lyalliialtogether, suggesting that such forms might behybrids between A. lucidum and A. bulbiferum. Onthe other hand, Crookes (1963) maintained all threetaxa, A. lucidum, A. lyallii, and A. anomodum, asfull species, though she admitted that the distinctionbetween the later two was somewhat doubtful.

During the present investigation, many wild popu-lations have been examined and many plants broughtinto cultivation. In the wild, both pinnate and bipin-nate plants have been found growing together inhabitats ranging from coastal cliffs to inland lime-stone mountains. In cultivation, a wide range ofthese morphological forms have reverted to a sur-prisingly uniform condition in which the fronds areessentially pinnate with only the lowest pinnae par-tially lobed or incised. Pinnate and bipinnate plantsshow the same ranges of variation with respect tospore pattern, spore size (mean values 40-54 X 28-35itim in pinnate forms, 42-50 x 27-33 nm in bipin-nate forms), and the shape of the stipe scales (Fig.4F-L). All the poulations investigated have beenuniformly octoploid. Hence, it seems clear that plantspreviously referred to the two species A. lyallii andA. anomodum rea'ly belong to one extremely poly-morphic taxon, and that the later name, A.anomodum, should be rejected as a synonym of A.lyallii.

Nevertheless, there is some suggestion that plantsof this complex growing in coastal areas, particularlyon non-calcareous soils, have dark shiny green frondswhich are larger, more divided, and with scales con-fined to the lower surface, whereas plants of lime-stone upland areas have du'l grey-green fronds whichare smaller, less divided, and have scales on bothsurfaces. It is possible that these forms should betreated respectively as var. lyallii and var. anomodum.However, the association between frond form andhabitat is not strong, and many exceptions to thistendency have been found making it difficult tojustify any taxonomic distinction between them.


cmr2O

-10

- O

ifrj£\WF^

AFig. 13 Fronds of Asplenium lamprophyllum ( x I ) . A, NZ 560, Little Barrier Island; B, NZ 529. SmithsBush, Auckland.

Brownsey—N.Z. species of Asplenium

Hence, I prefer to regard A. lyallii as a single poly-morphic species without any further formal sub-division in rank.

8. ASPLENIUM LAMPROPHYLLUM CarseTrans. N.Z. Inst. 56: 81 (1926) (Fig. 13)

TYPE: Lectotypc in CHR; lava flats, Mt. Wellington,Auckland, coll. H. Carse, 30.5.1924, 27.10.1927.

DESCRIPTION : Rhizome creeping, up to 15 cm long,pale green with a few scattered scales, stoloniferous.Stipes 6-20 cm long, pale green above, brown below,deeply grooved, covered in small, very dark, triangularto ovate scales which have thick cell walls (Fig.17F-J). Laminae lanceolate to elliptic, 15-60 X7-20 cm, light green, glossy above, thin, bipinnate.Raches green, scaly, prominently grooved. Pinnae12-20 pairs, lanceolate to narrowly ovate, acuminate,stalked, 3-10 X 0.5-2.5 cm, scaly on underside; lowerpinnae themselves pinnate, upper ones pinnatifld.

59

Pinnules sessile or shortly stalked, elliptic, oftendeeply serrate, 10-20 X 5-15 mm. Sori 3-10 mm long,nearer mid-vein than margin. Sporangia orange-brown. Spores (30)38-42(50) Mm long, (20)24-29(35)^m wide (4 populations), perispore very pale and notprominently patterned (Fig. 30E-F).

CHROMOSOME NUMBER: n=72 (Fig. 32F). NZ 652,

Hikurangi, Whangarei; NZ 681, Smith's Bush, Auck-land.

The same chromosome number was recorded byBrownlie (1961) for material from New Plymouth.

HABITAT AND DISTRIBUTION: N. A. lamprophyllum

is an endemic species almost entirely confined toregions north of latitude 38°N, with only veryscattered populations extending as far south asWanganui on the west coast (Fig. 14). It frequentlycarpets extensive areas on rocky ground or clay banksin shady forest, growing from its creeping rhizomeand spreading by stolons. It occurs from sea levelto 600 m.

r* 1 ?. .

A. lamprophyllum

u r

Fig. 14 Distributions ofA splenium lamprophyllumand A. bulbijerum.

A. bulbiferum


COMMENT: A. lamprophyllum is sometimes con-fused with A. bulbijerum but can easily be distin-guished from this species by its creeping rhizome,thick-walled stipe scales (cf. Fig. 17A-E. F-J),glossy frond, absence of bulbils, long orange sori.rather featureless spores (cf. Fig. 30E-F, K-L)and the fact that it is never more than bipinnatelydivided (cf. Figs 13, 15). It also frequently smellsof oil of wintergreen when crushed.

9. ASPLENIUM BULBIFERUM Forst. f. Prodr.:80 (1786) (Figs 15. 16)

TYPE: Lectotype in BM; New Zealand, Dusky Bay.coll. J. R. and G. Forster.

DESCRIPTION: Rhizome short, stout, erect, bearingovate scales up to 15 X 5 mm. Stipes 5-30 cm long,brown on underside, green above, stout, covered insmall brown ovate scales. Laminae lanceolate toelliptic, 15-120 X 7-30 cm, bi- to tripinnate, some-times bearing bulbils. Raches green, scaly, promin-ently grooved. Pinnae 15—30 (or more) pairs, ovateto narrowly ovate, acuminate, shortly stalked, 3-20X 1-5 cm, scaly on underside, basal pair pointingdownwards when fresh. Secondary pinnae sessile orshortly stalked, very narrowly elliptic to ovate orelliptic, obtuse, deeply serrate or sometimes almostpinnate, decreasing in size from base to apex, basalacroscopic pinnule often enlarged (up to 4 X 1 cm).Ultimate pinnules narrowly oblong, ± entire tocrenate-serrate, up to 1 cm long. Sori numerous,broad, submarginal, 2-4 mm long. Perispore of sporesirregularly ridged (Fig. 30K-N).

9a. A. BULBIFERUM Forst. f. subsp.BULBIFERUM (Fig. 15)

DIAGNOSIS: Stipe scales ovate, lacking long apicalprojections (Fig. 17A-E). Laminae pale green, usuallybulbiferous. Spores (33)41-48(55) /an long, (21)28-32(37) fim wide (3 populations).

CHROMOSOME NUMBER: n = 72 (Fig. 32A). NZ 31,

Wilton Bush, Wellington; 2n=144, NZ 707, BullStream, Akatarawa Range, Wellington.

Browniie (1954) also reported n = 72 in thisspecies.

9b. A. BULBIFERUM Forst. f. subsp.GRACILLIMUM (Col.) Brownsey, comb. nov.

(Fig. 16)

BASIONYM: A. gracillimum Col. Trans. N.Z. lnst.22: 453 (1890)

TYPE: Holotype in WELT; Dannevirke. coll. W.Colenso. (WELT P3348)

DIAGNOSIS: Very similar to subsp. bulbijerum butdiffering in the following characters. Stipe scales withapices drawn into long filiform projections (Fig.17K-O). Laminae dark green, rarely bulbiferous.Spores (33)41-48(55) i>m long, (21)28-32(37) ^mwide (3 populations).

CHROMOSOME NUMBER: n=144 (Fig. 32B). NZ 661.

Karekare, Auckland; NZ 55, Lake Rotoaira, Tonga-riro; NZ 1164, Carter's Bush. Gladstone, Wairarapa.

HABITAT AND DISTRIBUTION: N, S, St., Ch.. A. (Fig.

14). A. bulbijerum occurs commonly in woodedareas from North Cape to Stewart Island, and isabsent only from the drier parts of South Canter-bury and Central Otago where forest has beenremoved. It is a characteristic component of theground-layer in most forests and grows from sealevel to 1 100 m. It has not been possible to prepareseparate distribution maps for the two subspecies ofA. bulbijerum because herbarium specimens can onlybe distinguished by reference to micro-morphologicalcharacters. However, it seems likely from personalfield observations that both subspecies are equallywidespread and probably occur sympatrically inmany areas.

COMMENT: The A. bulbijerum complex is wide-spread and extremely polymorphic, and it has there-fore attracted considerable attention from taxonomistswho have proposed many different names for itsvarious forms. It is now clear that, at least in NewZealand, there are two different cytotypes in thiscomplex, and that some morphological distinctionis associated with the increase in chromosome num-ber. Thus, the tetraploid form is usually bulbiferous.has fronds which are often a pale green, stipe scaleswhich lack long filiform apices (cf. Fig. 17A-E,K-O) and spores which are smaller (mean values34-37 x 23-26 ^m). By contrast, the octoploid formis rarely bulbiferous, has darker green fronds, moreattenuated stipe scales and larger spores (mean values41-48 X 28-32 /xm). On this basis, the two cytotypeshave been referred to separate subspecies.

A. bulbijerum was first described by Forster, andexamination of his material in BM and GOET indi-cates that plants collected by him were all bulbi-ferous and had stipe sca'es and a spore size typicalof the tetraploid cytotype. The frond shape of hisspecimens also corresponds to that of known tetra-ploid plants. The name A. bulbijerum subsp. bulbi-jerum is therefore retained for this cytotype.

At the octoploid level there is a much greaterrange of frond dissection than amongst the tetraploid


c

AFig. 15 Fronds of Asplenium bulbiferum subsp. bulbiferum ( XGeorge Park, Wellington; C, NZ 35, Wilton Bush, Wellington.

I ) . A, NZ 906. Rapid; B. NZ 284, Keith

plants (cf. Figs 15. 16). At one extreme thereare forms with rather broad and only partially incisedsecondary pinnae (e.g.. Fig. 16A) and, at the other,forms which have their secondary pinnae completelydivided into rather narrow ultimate pinnules (e.g.,Fig. 16D). However, it is clear from examination ofmany herbarium specimens that these types havemany intermediates (e.g.. Fig. 16B, C) and thatthere is a complete morphological continuum betweenthe two extremes. It is proposed, therefore, that

the octoploid cytotype should be treated as a singlepolymorphic subspecies encompassing all thesevarieties. However, individual populations rarelyinclude all of these forms and the more extremefrond types are usually found in different localities.It is possible therefore that some morphologicaldifferentiation may be related to ecological factors,but until it can be more thoroughly investigated inrelation to environmental conditions this subspecieswill not be further subdivided.


Fig. 16 Fronds of Aspleniunt bulbiferum subsp. gracillimum ( x })• A, NZ 1071, Lighthouse Track, StewartIsland; B, NZ 231, Stronvar, Wairarapa; C. NZ 656. Karekare, Auckland; D, NZ 71, Lake Rotoaira, Tongariro.

Browrtsey—N.Z. species of Asplenium 63

K

Fig. 17 Camera lucida drawings of stipe scales ( x 20), each taken from a different population. A-E, Aspleniumbulbiferum subsp. bulbiferum; F-J, A. lamprophyllum; K-O, A. bulbiferum subsp. gracillimum; P-S. A.richardii; T-Z'. A. hookerianum (T-X, A. hookerianum, Y-Z', A. colensoi).


The most appropriate name for the octoploidsubspecies appears to be a new combination. A.bulhiferum subsp. gracillimum (Col.) Brownsey.Colenso's type specimen of A. gracillimum is verysimilar to the frond illustrated in Fig. 16A and therange of forms in his herbarium includes some similarto Fig. 16B. The spore size and stipe scales ofColenso"s material also fall within the range of varia-tion exhibited by plants now known to be octoploid.

Frond forms similar to that in Fig. 16D havebeen referred by some authors to A. bulbiferum var.tripinnatum Hook. f. (e.g., Field 1890, plate XXII1.fig. 6; Crookes 1963, p. 345). However, examinationof Hooker's type material indicates that his conceptof var. tripinnatum was quite different, the frondbeing considerably more divided than anything illus-trated in Fig. 16. It has very long narrow tertiarypinnae which are up to 15 mm long and 4 mm wide,and which are themselves divided into segments.Furthermore, the sori are up to 5 mm long which issignificantly greater than for any other specimen ofA. bulbiferum I have examined. The possibility there-fore exists that Hookers specimen may be a hybrid,but because it is unfortunately immature this hypo-thesis cannot be tested by reference to the spores,and the taxon must remain of equivocal status.

Whatever its status may eventually prove to be,there is no doubt that A. bulbiferum var. tripinnatumhas been widely misinterpreted by later workers, andthat forms such as Fig. 16D have been incorrectlyreferred to it. Because these varieties have apparentlyreceived no other taxonomic recognition, subsp.gracillimum is interpreted here on a broader basisthan by Colenso and includes the entire range ofvariation illustrated in Fig. 16. Hence, this taxonis more clearly defined by reference to chromosomenumber and spore size than it is by the degree offrond dissection.

One other name which has been proposedwithin the A, bulbiferum aggregate deserves furthermention. A. laxum R.Br. was described from Tas-mania as a species very similar to A. bulbiferumbut differing in that it lacked bulbils. Examinationof the type specimen in BM, however, indicates thatthe plant has aborted spores and is therefore veryprobably a hybrid. Other collections made by Brownin Australia and referred to A. laxum have goodspores, but clearly the name must now be appliedto the hybrid, whatever its parentage may be. Therepresentatives of the A. bulbiferum complex inAustralia need further investigation in the light ofthe present revision of New Zealand material.

10. ASPLENIUM HOOKERIANUM Col. Tasm. J.nat. Sci. 2: 169 (1846) (Fig. 18)

A. colensoi Col. Tasm. J. nat. Ssi. 2: 170 (1846)A. adiantoides Raoul Ann. Sci. nat. Ser. Ill 2:

115 (1844) non Lam. 1786A. adiantoides Raoul var. minus Hook f. in Hook.

Ic. PI. 10: t.983 (1854)A. adiantoides Raoul var. colensoi Hook. f. in

Hook. Ic. PI. 10: t.984 (1854)A. adiantoides Raoul var. hookeriana Hook. f.

Fl. N.Z. 2: 35 (1855)A. hookerianum var. colensoi Moore Ind. Fil.:

137 (1858)A. richardii var. colensoi Hook. Sp. Fil. 3: 197

(1860)A. ornatum Col. Trans. N.Z. Inst. 22: 452

(1890)A. svmmetricum Col. Trans. N.Z. Inst. 31: 264

(1899)

TYPE: Holotype in WELT; between Matamata andMaungatautari, coll. W. Colenso, January 1842.

DESCRIPTION: Rhizome short, erect, bearing numer-ous brown, ovate, acuminate scales up to 10 X 2 mm.Stipes 2-10 cm long, pale brown below, green above,densely covered in small, subulate to narrowly ovatescales with filiform apices (Fig. 17T-Z). Laminaelanceolate to rhombic, 4—25 X 1-15 cm, dark green,thin, normally bipinnate but often pinnate whenyoung and almost tripinnate in well-grown speci-mens. Raches green, slender, and very scaly. Pinnae5-15 pairs, very narrowly ovate to ovate, obtuse toacuminate, long-stalked, 1-8 X 0.5-2 cm, basal pairpointing upwards when fresh. Pinnules stalked, 3-10pairs, linear to suborbicular, crenate to deeplyincised (or almost pinnate), 3-12 X 3-10 mm, tendingto lie at 90° to plane of frond in well-grown speci-mens. Sori sub-marginal on narrow pinnules, remotefrom margins on broad segments, 1-3 mm long.Spores (25)31-37(42) /mi long, (19)23-27(33) ^mwide (9 populations), perispore very prominentlywinged and ridged (Fig. 30G-H).

CHROMOSOME NUMBER: n=72 (Fig. 33C). NZ 67.Lake Rotopounamu, Tongariro; NZ 118, OkahukaraForest, Tongariro; NZ 213, 277, Stronvar, Wairarapa;NZ 912, Kapiti Island; NZ 848, Khandallah, Welling-ton; NZ 318, Karori, Wellington; NZ 711, Linden.Wellington; NZ 796, Takaka Hill, Nelson.

Brownlie (1954, 1958) recorded n = 72 for bothA. hookerianum and A. colensoi.

HABITAT AND DISTRIBUTION: N, S, St. A. hookerianumis one of the most common and widespread speciesof Asplenium in New Zealand, having been recordedfrom Kaitaia to Stewart Island (Fig. 19). However,there are no recent collections of the species fromfurther north than Kaipara Harbour and its present


cm15-1

IO-

5-

OJ

Fig. 18 Fronds of Asplenium hookerianum ( x 1). A, NZ 1145, Carters Bush, Wairarapa; B, NZ 933, KapitiIsland; C, NZ 194, Stronvar, Wairarapa; D, NZ 306, Karori. Wellington; E. NZ 234. Stronvar, Wairarapa;F, NZ 769, Sugarloaf. Port Hills, Christchurch; G. NZ 942, Whangiamoana, Wairarapa.

66 New Zealand journal of Botany 15. 1977

A. hookerianum

Fig. 19 Distributions ofAsplenium hookerianum andA. richardii.

A. richardii

occurrence in the North Auckland peninsula needsinvestigation. Tt grows from sea level to 1 250 m andis found particularly on clay banks or rocky outcropsin scrub and open forest, or on the ground in dis-turbed forest remnants.

Outside New Zealand, this species has apparentlybeen recorded only once. Tn 1874, F. v. Muellercollected a specimen (in MEL) from rocks at 3 000—4 000 ft (900-1 200 m) on the Upper Hume River,Australia. Willis (1962) states that this locality ismost likely to have been on the slopes of MtKosciusko, on the New South Wales side of theMurray River. Clearly this record needs confirmation,but, even if the species is now extinct in Australia,it suggests that A. hookerianum once had a muchmore widespread distribution extending beyond NewZealand.

COMMENT: AS understood here, A. hookerianum isan extremely polymorphic species and one whichincludes several previously described taxa. Of these.

the one which has received the most widespreadrecognition is A. colensoi, a form with very narrowpinnules. This name was first mentioned by Colensoin a letter to W. J. Hooker (Hooker 1842), but notvalidly published until 2 years later when descrip-tions of both A. colensoi and A. hookerianumappeared together (Colenso 1846). Moore (1858-1859) later reduced A. colensoi to a variety of A.hookerianum and this combination has been acceptedby most subsequent authors, although Crookes(1963) retained both taxa as full species.

Colenso later described two more species, A.ornatum and A. symmetricum. However, examinationof the type specimens (in AK and WELT respec-tively) indicates that these are merely forms of hisearlier species A. hookerianum and A. colensoi, andshould therefore be rejected.

The real problem concerns the true status of A.hookerianum and A. colensoi. In the wild, these twoforms are very frequently found together in the same


populations and, indeed, it is rather unusual to findextreme forms of either type growing alone. Thepolymorphism in frond shape is particularly notice-able amongst young plants, different individualsproducing either rather broad pinnules, or muchnarrower pinnules. However, as they reach maturity,both in the wild, and especially in cultivation, thereis a strong tendency for plants of both forms toproduce fronds of an intermediate shape, possessingneither the very broad pinnules of A. hookerianumnor the very narrow ones of A. coletuoi. Further-more, the two taxa show the same rather narrowrange of variation with respect to spore pattern, sporesize (mean values 31-35 X 23-27 iim in A. hookeri-anum, and 31-37 X 23-26 /im in A. colensoi) andthe shape of the stipe scales (cf. Fig. 17T-X, Y-Z').Cytologically they are both uniformly tetraploid.Hence there can be no justification for maintainingtwo separate species. Because the two names werepublished simultaneously neither has priority; accord-ingly I have chosen to retain only A. hookerianumbecause this is the more widely used and thereforemore appropriate name.

It is possible that fronds bearing very narrowpinnules should continue to be recognised as a dis-tinct variety of A. hookerianum. However, in practiceit is frequently very difficult to assign a particularspecimen to one or other taxa on this basis. Thus,many plants have very narrow pinnules on theiryoungest fronds, but gradually acquire broader pin-nules as they get older (and vice versa). Further-more, although there is a considerable range ofvariation with respect to frond dissection, theextremes of variability are often found together andare not obviously correlated with particular habitatsor geographical locality. Recognition of var. colensoias a separate taxon is therefore difficult to justifyand it has been rejected.

A. hookerianum is usually a much smaller, moredelicate plant than A. bulbiferum, but confusionbetween large plants of the former and small plantsof the latter sometimes arises. Large specimens ofA. hookerianum can be distinguished by the presenceof very delicate but nevertheless quite distinct stalksto the pinnules. Small plants of A. bulbiferumgenerally have sessile pinnules, or very short broadstalks. A. bulbiferum also often has a narrow frond(more than twice as long as broad) which is flattenedin one plane, and a basal pair of pinnae which pointdownwards in fresh specimens. A. hookerianum hasfronds which are often less than twice as long asbroad (Fig. 18), pinnules tending to lie at 90° tothe plane of pinnae and raches, and a lower pair ofpinnae which point upwards.

11. ASPLENIUM RICHARDII (Hook, f.) Hook, f.Fl. N.Z. 2: 35 (1855) (Fig. 20)

A. adiantoides Raoul var. richardii Hook. f. inHook. Ic. PI. 10: t.977 (1854)

A. raoulii var. richardii Mett. U. e. Farngatt.:118 (1859)

TYPE: Holotype in P; New River, Southern Island,N.Z., herb. A. Richard.

DESCRIPTION: Rhizome short, stout, erect, bearingdark brown subulate scales up to 20 X 2 mm. Stipes5—15 cm long, brown on underside, green above,densely covered in subulate scales with filiform apices(Fig. 17P-S). Laminae ovate to narrowly ovate,10-25 X 4-12 cm, dark green, relatively thin, normallytripinnate. Raches green, very scaly, slightly grooved.Pinnae 10-15 crowded and overlapping pairs, ovateto narrowly ovate, sub-acute, stalked, 2-8 X 1-4 cm.Secondary pinnae stalked, ovate, 10-20 X 10-15 mm,again pinnate or pinnatifid. Ultimate segments linear,acute or sub-acute, up to 8 mm long. Pinnae andpinnules not flattened in one plane but spreading inthree dimensions. Sori 2-4 mm long, submarginal.Spores (38)44-49(55) pm long, (27)32-36(40) Mmwide (6 populations), perispore prominently wingedand ridged (Fig. 30I-J).

CHROMOSOME NUMBER: n=144 (Fig. 32E). NZ 422,Hooker Valley, Mt Cook.

This number was also recorded by Iirownlie(1954).

HABITAT AND DISTRIBUTION: N, S. A. richardii is an

endemic species almost entirely confined to mountaindistricts of the South Island, being found in theNorth Island only on Mt Honokawa, Mt Ruapehu,and parts of the Kaimanawa Range (Fig. 19). Itextends throughout the mountain regions of the SouthIsland, growing on rocky outcrops, in crevices,amongst boulders, and on stream banks particularlyunder beech forest. It is found on both limestoneand greywacke rock and occurs up to 1 400 m, butis more frequent east of the Main Divide.

COMMENT: A. richardii can be confused with well-grown specimens of A. hookerianum, a species towhich it is closely related but from which it is clearlydistinguished by its octoploid chromosome numberand its larger spores (cf. Fig. 30G-H, I-J).

A. richardii can also be confused with A. terrestre,a species with which it often grows in close associa-tion. The two taxa are both octoploid and havevery similar spores, but A. richardii is characterisedby its thinner frond with long narrow pinnules. (Fig.20), its shorter sori (2-4mm), the dense covering

New Zealand Journal of Botany 15. 1977

Fig. 20 Fronds of Asplcnium richardii (X i). A, D. NZ 419. Black Birch Stream. Mt Cook; B. NZ 401.Glencoe Creek, Mt Cook; C, NZ 271, Cobb Valley, Nelson.

of scales on the stipe and rachis, and by its habitof producing pinnae and pinnules in three dimensions.A. terrestre has a more leathery frond with broadpinnules flattened into one plane (Fig. 25), longersori (2-7 mm), and fewer scales on the stipe andrachis.

12. ASPLEN1UM FLACCIDUM Forst. f. Prodr.:80 (1786) (Figs 21, 23)

Coenopteris flaccida Thunb. Nov. Act. Pet. 9:158, t.D, f.1-2 (1795)

C. novae-zelandiae Spreng. Schrad. J. Bot. 2: 269(1799)

Fig. 21 Fronds of Asplenium fiaccidum subsp. fiaccidum ( x i ) . A. NZ 38. Wilton Bush, Wellington; B.NZ 441. Haast, South Westland; C. NZ 387, Terenikau Valley. Tararuas; D. NZ 63, Lake Rotopounamu.Tongariro: E. NZ 1004. Lake Fergus. Fiordland.


Darea fiaccida Willd. Sp. PI. 5: 245 (1810)D. odontites Willd. Sp. PI. 5: 245 (1810)A. heterophyllum A. Rich. Essai Fl. N.Z.: 74

(1832)

TYPE: Lectotype in BM; New Zealand, Dusky Bay,coll. J. R. and G. Forster.

DESCRIPTION: Rhizome short, stout, erect, bearingdark brown subulate or ovate scales up to 20 X 2 mm.Stipes 5-20 cm (or more) long, brown on underside,green above, flaccid or firm and erect, sparinglycovered in small subulate or ovate scales. Laminaelanceolate to elliptic, 10-90 (or more) X 4-25 cm,dull green, thick, leathery and either limp andpendulous or stiff and erect, pinnate to bipinnate.Raches green, sparingly scaly. Pinnae in 5-20 (ormore) pairs, linear to narrowly ovate, acuminate,long stalked, 2-15 X 0.5-2 cm; degree of dissectionvery variable, sometimes only divided into very shortobtuse segments, sometimes pinnate with linear acutepinnules up to 15 X 2 mm. Sori submarginal, linear,2-10 mm long.

12a. A. FLACCIDUM Forst. f. subsp.FLACCIDUM (Fig. 21)

DIAGNOSIS: Rhizome bearing subulate scales. Stipes5-20 cm (or more) long, flaccid, bearing small subu-late scales with long filiform apices (Fig. 24A-E).Laminae lanceolate to elliptic, 15-90 (or more) X5-25 cm, leathery, limp and pendulous. Pinnae 5-20(or more) pairs, linear, 5-15 X 0.5-2 cm. Pinnulesvery variable in length, from oblong and obtuse tolinear and acute, up to 15 X 2 mm. Basal acroscopicpinnule occasionally much longer than that next to it.Sori 2-10mm long. Spores (31)36^44(50) /im long,(19)23-27(33) pm wide (10 populations), perisporewith a prominent wing but lacking ridges (Fig.29M-N). Plant usually epiphytic, but sometimesterrestrial.

CHROMOSOME NUMBER: n = 72 (Fig. 33A). NZ 857,Renata Ridge, Akatarawa Range, Wellington; NZ 32,36, Wilton Bush, Wellington; NZ 840, Seatoun,Wellington.

Brownlie (1954) reported n = 72 for A. flaccidumsens. lat.

HABITAT AND DISTRIBUTION: K, N, S, Ch., St. (Fig.22). A. flaccidum subsp. flaccidum is a common fernthroughout the forested regions of New Zealandfrom North Cape to Stewart Island, being absent onlyin the drier parts of Otago and Canterbury fromwhich the forest has been cleared. It grows principally

as an epiphyte on various native trees, occurringfrom sea level to the tree line, but is sometimes alsofound on the ground. Outside New Zealand its distri-bution includes parts of Australia and some of thePacific Islands.

12b. A. FLACCIDUM Forst. f. subsp.HAURAKIENSE Brownsey, subsp. nov. (Fig. 23)

Rhizoma paleis ovatis cum parietibus cellularumcrassissimis vestitum. Stipes 5-12 cm longus, erectus,firmus, paleis anguste vel late ovatis cum parietibuscellularum crassissimis vestitus (Fig. 24F-I). Laminaoblonga vel elliptica, 10-30 X 4-15 cm rigida, erecta.Pinnae 5-12 jugatae, lineares vel anguste ovatae,20-80 X 8-16 mm. Pinnulae oblongae obtusaeque vellineares acutaeque, usque ad 7 X 2 mm. Pinnulabasalis acroscopica multo longior quam pinnulavicina, usque ad 4 cm longa, pinnatifida. Sori 2-7 mmlongi. Sporae (38)43-49(54) urn longae, (20)26-31(38) pm latae; perisporae interdum alatae,plerumque non cristatae. Numerus somaticuschromosomatum 144; in meiosi paria 72 observantur.Habitat in rupibus orae.

HOLOTYPUS: CHR; NZ 894, Oneroa Beach, WaihekeIsland, Auckland (NZMS 1 N42:528688). Growingamongst coastal rocks. Coll. PJ . Brownsey, 1.9.1974.(CHR 308926)

DIAGNOSIS: Rhizome bearing ovate scales with verythick cell walls. Stipes 5-12 cm long, erect, firm,bearing narrowly to broadly ovate scales with verythick cell walls (Fig. 24F-I). Laminae oblong toelliptic, 10-30 X 4-15 cm, stiff, erect. Pinnae 5-12pairs, linear to narrowly ovate, 20-80 X 8-16 mm.Pinnules oblong and obtuse to linear and acute, upto 7 X 2 mm. Basal acroscopic pinnule frequentlymuch longer than that next to it, up to 4 cm long,and itself pinnatifid. Sori 2-7 mm long. Spores(38)43-49(54) Mm long, (20)26-31(38) Mm wide (10populations); perispore sometimes winged, usuallylacking ridges (Fig. 29 O-P). Plant of coastal rocks.

CHROMOSOME NUMBER: n=72 (Fig. 33B). NZ 640,Lion Rock, Bay of Islands; NZ 582, Shag Point,Little Barrier Island; NZ 894, Oneroa, WaihekeIsland.

HABITAT AND DISTRIBUTION: N. A. flaccidum subsp.haurakiense is found along the eastern side of theNorth Auckland peninsula from the Three KingsIslands to the Hauraki Gulf and Coromandel, extend-ing locally into the Bay of Plenty (Fig. 22). It is sofar not known outside New Zealand. Plants of thissubspecies are almost always found growing terres-trially and are restricted to coastal areas within theimmediate vicinity of salt spray, either on exposedrock or amongst coastal scrub. The species is replacedon Auckland's west coast by A. terrestre subsp.maritimum (Fig. 26).

Brownsej—N.Z. species of Asplenium 71

Fig. 22 Distribution ofAsplenium ftaccidum.

A. flaccidumsubsp. flaccidum

A. flaccidumsubsp.haurakiense

COMMENT: The present investigation has shown thatthe A. flaccidum aggregate comprises two quiteseparate cytotypes, tetraploid and octoploid, and thatthe associated morphological differences are sufficientto warrant the recognition of two distinct species,A. flaccidum and A. terrestre. Each of these can befurther subdivided into two subspecies distinguishedlargely on the basis of ecological differentiation.

Material of A. flaccidum in BM and GOETcollected by Forster has been examined. The name isretained here for plants which are usually epiphytic,have long pendulous leathery fronds, and which arenever more than bipinnately divided (Fig. 21). Theyhave small subulate stipe scales (Fig. 24A-E), sporeswhich are prominently winged but rarely ridged(Fig. 29M-N), and are tetraploid. There remains awide range of frond forms within this species, butit is a more clearly circumscribed taxon than hasbeen interpreted by other authors and it correspondsclosely to material named by Forster.

A distinct ecological race of this species is des-cribed as A. flaccidum subsp. haurakiense. This is

similar to subsp. flaccidum in having many of thesame frond characters, an almost identical sporepattern, and being tetraploid. It is distinguishedprimarily by its coastal habitat and restricted geo-graphical range, its firm erect habit, its larger spores(cf. Fig. 29M-N. O-P), and by its very thick-walled, ovate, stipe scales (cf. Fig. 24A-E, F-I).

13. ASPLENIUM TERRESTRE Brownsey, sp. nov.(Figs 25, 27)

Rhizoma breve, erectum, paleis fuscis subulatisusque ad 15 X 2 mm obtectum. Stipes 5—15 cmlongus, infra fuscus, supra viridis, paleis subulatisvel anguste triangularibus cum apicibus longisfiliformibus vestitus (Fig. 24J-N, Q-V). Laminalanceolata vel elliptica, 10-30 X 5-20 cm, atroviridis,coriacea aut crassissima carnosaque, cernua aut rigidaerectaque, bi- vel tri-pinnata. Rhachis viridis,squamata, manifeste canaliculata. Pinnae 7-20jugatae, inferiores ovatae, superiores perangusteovatae vel ellipticae, acutae vel acuminatae, 3-10 X0.5-3 cm, paleis minutis infra vestitae. Pinnulaesuperiores lineares acutae integrae sessilesque;inferiores ovatae vel anguste ellipticae, pinnatifidae

cm20-i

10-

DFig. 23 Fronds of Asplenium flaccidum subsp. haurakiense ( x t ) . A, NZ 641, Lion Rock, Bay of Islands; B.NZ 1132, Poor Knights Islands; C, D, NZ 578, Awaroa Stream, Little Barrier Island; E, NZ 890, Mataitai,Waiheke Island.


Fig. 24 Camera lucida drawings of stipe scales ( x 20), each taken from a different population. A-E, Aspleniumflaccidum subsp. flaccidum; F-I, A. flaccidum subsp. haurakiense; J-N, A. terrestre subsp. terrestre; O-P,A. shuttleworthianum; Q-V, A. terrestre subsp. maritimum.


vel pinnatae, petiolatae, usque ad 30 X 10 mm. Sorisubmarginales, 2-7 mm longi. Sporae (38)43-51(56)/tm longae, (25)29-34(38) /ira latae, perisporae mani-feste alatae cristataeque (Fig. 30 O-R). Numerussomaticus chromosomatum 288; in meiosi paria 144observantur.

HOLOTYPUS: CHR; NZ 96, Blowhard Bush, Napier-Taihape Road (NZMS 1 N123:861491). Growing onthe ground beneath mixed forest. Coll. P. J. Brownsey,30.12.1973. (CHR 308925)

DESCRIPTION: Rhizome short, erect, bearing darkbrown subulate scales up to 15 X 2 mm. Stipes 5-15cm long, brown below, green above, covered insubulate to narrowly triangular scales with longfiliform apices (Fig. 24J-N, Q-V). Laminae lanceolateto elliptic, 10-30 X 5-20 cm, dark green, leathery orvery thick and fleshy, slightly drooping or stiff andupright, bi- to tripinnate. Raches green, scaly,prominently ridged. Pinnae 7-20 pairs, those at thebase ovate, those above very narrowly ovate orelliptic, acute to acuminate, stalked, 3-10 X 0.5-3 cm,covered on underside in tiny scales. Upper pinnuleslinear, acute, entire and sessile; lower pinnules ovateto narrowly elliptic and pinnatifid or pinnate, up to30 X 10 mm. Sori submarginal, 2-7 mm long. Spores(38)43-51(56) f«n long, (25)29-34(38) ^m wide (14populations), perispore prominently winged andridged (Fig. 30 O-R). 288 chromosomes in thesporophyte; 144 bivalents at meiosis.

HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.

terrestre is an endemic species with a distributionconfined to the North and South Islands and extend-ing from Auckland to Bluff. It is a terrestrial plantfound growing on the forest floor, on bluffs androcky outcrops, or on sea cliffs.

13a. A. TERRESTRE Brownsey subsp. TERRESTRE(Fig. 25)

DIAGNOSIS: Laminae lanceolate to elliptic, 10-30 X5-15 cm, thick and leathery, usually slightly droop-ing. Pinnae 8-20 pairs, those at the base ovate, thoseabove very narrowly ovate or elliptic, often withlong relatively undivided acuminate tips. Upperpinnules linear and entire, lower pinnules narrowlyelliptic and pinnatifid to pinnate, up to 30 X 8 mm.

CHROMOSOME NUMBER: n=144 (Fig. 33D). NZ 121,Okahukara Forest, Tongariro; NZ 96, Blowhard Bush,Napier-Taihape Road; NZ 312, Karori, Wellington;NZ 270, Woodside Creek, Marlborough.

HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.terrestre subsp. terrestre is found principally in theforests of North Canterbury and Marlborough, but

extends locally to many parts of the South Island.In the North Island it occurs mainly on the volcanicplateau and in the mountains of the Hawke"s Bayarea, but also reaches Rotorua, Mt Egmont, andthe Rimutakas. It grows terrestrially in forested areasbut may also be found on bluffs and rocky outcropsup to 1 100 m.

13b. A. TERRESTRE Brownsey subsp.MARITIMUM Brownsey, subsp. nov. (Fig. 27)

Lamina pro ratione brevis lataque, elliptica, 10-25X 7-20 cm, crassissima, carnosa, rigida, erecta.Pinnae 7-15 jugatae, inferiores ovatae, superioresanguste ovatae, acutae. Pinnulae superiores linearesintegraeque, inferiores ovatae vel ellipticae pinna-tifidaeque, usque ad 20 X 10 mm. Numerus somaticuschromosomatum 288; in meiosi paria 144 observantur.

HOLOTYPUS: CHR; NZ 12, Ohiro Bay, Wellington(NZMS 1 Nl64:300142). Growing on greywackecoastal cliffs. Coll. P. J. Brownsey. 15.12.1973.(CHR 308927)

DESCRIPTION: Laminae relatively short and broad,elliptic, 10-25 X 7-20 cm, very thick and fleshy, stiffand upright. Pinnae 7-15 pairs, those at the baseovate, those above narrowly ovate, acute. Upperpinnules linear and entire, lower pinnules ovate toelliptic and pinnatifid, up to 20 X 10 mm.

CHROMOSOME NUMBER: n=144 (Fig. 33E). NZ 669,

Karekare, Auckland; NZ 508, Manaia, Taranaki; NZ12, Ohiro Bay, Wellington.

HABITAT AND DISTRIBUTION: N, S. (Fig. 26). A.terrestre subsp. maritimum has a distribution centredon Cook Strait where it occurs more or less con-tinuously around the Wellington coast from CapePalliser to Pukeraa Bay and Kapiti. It is common inthe Marlborough Sounds, and extends down theMarlborough coast as far as Banks Peninsula. Else-where it is known only from Cape Farewell, theTaranaki coast, and from the west coast of Auck-land. It is found on coastal cliffs within the influenceof salt spray, either on exposed rocks where plantsare rather stunted, or amongst coastal scrub wherethe plants grow much bigger.

COMMENT: A. terreslye is a species which has beenincluded within A. flaccidum by most previous authorsbut which is clearly distinct from that taxon becauseof its octoploid chromosome number, its more highlydivided frond (cf. Figs 21, 23, and 25, 27), and byits more prominently ridged spore pattern (cf. Figs29M-P, 30 O-R). It differs from the epiphyticA. flaccidum subsp. flaccidum in growing terrestriallyand having a more or less erect frond and a largerspore size. It can also be distinguished from A.


cFig. 25 Fronds of Asplenium terrestre subsp. terrestre ( x 1). A, NZ 1042, Bluff Hill, Southland; B. NZ771, Sugarloaf, Port Hills, Christchurch; C. NZ 724, Woodside Creek, Marlborough; D. NZ 403. GlencoeCreek, Mt Cook; E, NZ 109, Okahukara Forest, Tongariro.


A. terrestresubsp.terrestre

Fig. 26 Distribution ofAsplenium terrestre.

A. terrestresubsp.maritimum

flaccidum subsp. haurakiense by the lack of a greatlyelongated basal acroscopic pinnule on each pinna(cf. Figs 23. 27).

A. flaccidum subsp. flaccidum frequently occurstogether with A. terrestre subsp. terrestre in forestedareas; in parts of the Cook Strait region where forestextends to the coast it also overlaps with subsp.maritimum. In both these areas hybrids between A.flaccidum and A. terrestre are reasonably common.They will be described in more detail in anotherpublication, but it is sufficient to say here that thehybrids produce only aborted spores and are com-pletely sterile. The recognition of two separate speciesis therefore fully justified not only on morphologicaland cytological evidence but also on geneticalgrounds.

The distinction between the two subspecies ofA. terrestre is primarily ecological, subsp. terrestrebeing largely confined to inland areas, and subsp.maritimum to coastal regions, but there is also somedegree of morphological differentiation. Thus, subsp.maritimum has a much thicker and more fleshy frond,

and pinnae which are generally broader than thosein subsp. terrestre (cf. Figs 25, 27).

It is possible that A. terrestre was first detectedas a distinct species by Armstrong (1882) when hedescribed A. canterburiense from various localitiesin Canterbury. His description is very similar to thatgiven here for A. terrestre, but the only specimenof his which I have been able to locate* consists ofa single frond which has aborted spores and isclearly a hybrid, probably between A. terrestre andA. richardii. This hybrid is extremely similar to A.terrestre (Brownsey, in preparation) and probablyoccurs reasonably frequently wherever the twoparents are found together. Armstrong's collectionsfrom other localities cited in his original descriptionof A. canterburiense may have included plants ofthe parent species as well as the hybrid, but untilsuch specimens are located the name A. canter-buriense must obviously be retained for A. richardiiX terrestre.

"One sheet in the Armstrong Herbarium, ChristchurchBotanic Gardens, labelled Mt Arrowsmith, 4 000 It.coll. J. F. Armstrong. March 1869.

Fig. 27 Fronds of Asplenium terrestre subsp. marilimum (X {). A, NZ 929. Kapiti Island; B, NZ 509.Manaia, Taranaki; C, NZ 673, Karekare, Auckland; D. NZ 353. Titahi Bay. Wellington; E. NZ 838. Seatoun,Wellington; F. NZ 837, Seatoun. Wellington.


Fronds of A. terrestre subsp. terrestre may some-times be confused with those of A. bulbiferumsubsp. gracillimum (cf. Figs 16B, 25B). However,A. terrestre subsp. terrestre can usually berecognised by its thicker, more leathery frond, byits longer sori (2-7 mm), and sometimes by thepresence of long, relatively undivided, acuminate tipsto the pinnae. A. bulbiferum subsp. gracillimum hasa thinner frond, shorter sori (2-4 mm), and usuallyhas pinnae which are divided throughout their lengthinto more or less distinct pinnules.

up to 12 X 5 cm. Secondary pinnae linear to lanceo-late, up to 15 X 2 mm, often pinnatifid. Ultimatesegments linear, subacute, slightly expanded in theregion of the sori. Sori near tips of ultimate seg-ments, solitary, broad, submarginal, c.2mm long.Spores (46)51-63(71) ^m long, (32)34-44(50) Mmwide (3 populations), perispore prominently winged,granular, but not conspicuously ridged (Fig. 30C-D).

CHROMOSOME NUMBER: n=c.l44. NZ 1112, RaoulIsland, Kermadecs.

14. ASPLENIUM SHUTTLEWORTHIANUMKunze Farrnkr. 1: 26, t.14 (1840) (Fig. 28)

A. flaccidum var. shuttleworthianum Hook. f.Handbk. N.Z. Fl.: 274 (1864)

TYPE: BM; Pitcairn Island, coll. Cuming 1374.

DESCRIPTION: Rhizome short, erect, bearing red-brown, linear attenuate scales. Stipes up to 15 cmlong, green above, brown below, sparingly covered insmall linear scales with long filiform apices (Fig. 24O-P). Laminae oblong to elliptic, 15-90 X 10-25 cm,yellow-green, coriaceous, 3- to 4-pinnate. Rachesgreen, almost lacking scales, prominently grooved.Pinnae ovate to narrowly ovate, acuminate, stalked,

HABITAT AND DISTRIBUTION: K. In the New Zea-

land region, A. shuttleworthianum is known onlyfrom the Kermadecs where it apparently occupiesvarious habitats from coastal scrub to inland forest,and where it grows both terrestrially and epiphytic-ally. It also occurs on Pitcairn and other Pacificislands.

COMMENT: This species is closely related to A.terrestre but distinguished from it by a greater degreeof frond dissection (cf. Figs 25, 28), its verylarge spores (cf. Fig. 30C-D, O-P), and by thecharacteristic broadening of the ultimate pinnules inthe region of the sori. Because of its remote geo-graphic distribution it has been very little studied.

A

cm

2 On

10-

BoJ

Fig. 28 Fronds ofAsplenium shuttleworth-ianum ( x I ) . A, NZ 690,Raoul Island; B, NZ 1112,Raoul Island.


• •

B

K <

N oFig. 29 Exospore and perispore of spores of Asplenium species (all X 400).A-B, A. lucidum, NZ 552, Little Barrier Island; C-D, A. flabellifolium, NZ 178.Kapiti Island; E-F, A. obtusatum subsp. obtusatum, NZ 790, Wharariki Beach.NW Nelson; G-H, A. obtusatum subsp. northlandicum, NZ 687, Slipper Island.Bay of Plenty; I-J, A. scleroprium, CHR 195053. Ernest Island, off StewartIsland; K-L, A. lyallii, NZ 810, Takaka Hill, Nelson; M-N, A. flaccidumsubsp. ftaccidum, NZ 378, Bay of Many Coves, Marlborough; O-P. A. flaccidumsubsp. haurakiense, NZ 890, Mataitai. Waiheke Island.

DISCUSSIONThe New Zealand representatives of Asplenium

undoubtedly have their closest affinities with theflora of temperate parts of Australia. Some species,such as A. bulbiferum and A. flaccidum, form acharacteristic component of the forest vegetation inboth regions. Other species, however, have affinitieswith more distant floras. A. obtusatum, for example,is a common coastal plant in higher latitudes of theentire Southern Hemisphere; A. polyodon is primarilya plant of tropical regions, whereas A. trichomanesis of cosmopolitan distribution and has its closest

relatives in the Northern Hemisphere. Nevertheless,it is clear that most of these species must haveultimately arrived in New Zealand from Australia,presumably as a result of spores being blown acrossthe Tasman Sea in the predominating westerly air-stream.

A striking feature of Asplenium in New Zealandis the degree of polymorphism shown by its differentspecies, a fact which has been remarked on by manyworkers, and which is probably due to divergentevolution in the relatively impoverished flora of thisoceanic archipelago. The number of species which


B

s \/ •

D

J '

N

Fig. 30 Exospore and perispore of spores of Asplenium species (all X 400).A-B, A. polyodon, NZ 295, Keith George Memorial Park, Wellington; C-D,A. shuttleworthianum, NZ 690, Raoul Island; E-F, A. lamprophyllum, NZ 560,Little Barrier Island; G-H, A. hookerianum, NZ 796, Takaka Hill, Nelson;I-J, A. richardii, NZ 426, Hooker Valley, Mt Cook; K-L, A. bulbiferum subsp.bulbiferum, NZ 911, Kapiti Island; M-N, A. bulbiferum subsp. gracillimum, NZ231, Stronvar, Wairarapa; O-P, A. terrestre subsp. terrestre, NZ 794, Takaka Hill.Nelson; Q-R, A. terrestre subsp. maritimum, NZ 505, Manaia, Taranaki.

have successfully established themselves is rathersmall, but many of these have almost certainly beenable to spread into a greater variety of habitats thanin the continental and tropical floras from whichthey have migrated, and evolution in response to avariety of new environmental conditions has there-fore taken place.

Nevertheless, morphological diversification is notthe only change to have resulted from the evolutionof these species. Hybridism has also played its part,and the occurrence of intermediate forms has almostcertainly contributed to the inability of differentauthors to consistently define the limits of the NewZealand Asplenium species. Taxa whose distributions


/* i . •

B

Fig. 31 Permanent acetocarmine preparations of diakinesis (all X 1 000). A, Aspleniumflabellifolium, NZ 789, Wharariki Beach, NW Nelson, showing 210-211 univalents; B, A.flabellifolium, NZ 834, Turakirae Head, Wellington, showing 277-280 univalents; C, A.obtusatum subsp. obtusatum, NZ 472, Cape Foulwind, Westport, showing 72 bivalents; D,A. lucidum, NZ 462, Greymouth, showing 72 bivalents; E, A. obtusatum subsp. northlandicum,NZ 667, Karekare, Auckland, showing 144 bivalents.


A B

V 7 **• ,

Fig. 32 Permanent acetocarmine preparations of diakinesis (all X 1 000). A, Aspleniumbulbiferum subsp. bulbiferum, NZ 707. Bull Stream, Akatarawas, showing 72 bivalents; B,A. bulbiferum subsp. gracillimum, NZ 55, Lake Rotoaira. Tongariro, showing 144 bivalents;C, A. scleroprium, NZ 1012, Bluff, showing 144 bivalents; D, A. lyallii, NZ 986, Castle Hill,Canterbury, showing 144 bivalents; E, A. richardii, NZ 422, Hooker Valley, Mt Cook,showing 144 bivalents; F. A. lamprophyllum, NZ 652. Hikurangi. Whangarei. showing 72bivalents.


D

% ^ -

E

0

Fig. 33 Permanent acetocarmine preparations of diakinesis (all X 1 000). A,Asplenium flaccidum subsp. ftaccidum, NZ 36, Wilton Bush, Wellington, showing72 bivalents; B, A. flaccidum subsp. haurakiense, NZ 640, Lion Rock, Bay ofIslands, showing 72 bivalents; C, A. hookerianum, NZ 796, Takaka Hill, Nelson,showing 72 bivalents; D, A. terrestre subsp. terrestre, NZ 96, Blowhard Bush,showing 144 bivalents; E, A. terrestre subsp. maritimum, NZ 669, Karekare,Auckland, showing 144 bivalents.

overlap do occasionally hybridise, and, in disturbedareas, particularly where forest has been opened up,some species spread more prolifically than usualand may give rise to several hybrid individuals.However, unlike many of the polymorphic generaof trees and shrubs in New Zealand, hybrids betweeneven closely related species of Asplenium are usuallysterile, and the idea that there are abundant hybrid

swarms exhibiting various degrees of introgressionis quite erroneous. Most hybrid individuals can beimmediately recognised by their aborted and shrivelledspores and therefore provide a useful reference pointfor denning the limits of their parent species.

A crucial role in the evolution of Aspleniumin New Zealand has been played by polyploidy and


the present investigation has revealed several newand unsuspected cytotypes amongst the more com-mon species. One such example is the A. flaccidum/terrestre aggregate. Here, increase in chromosomenumber has been accompanied by morphologicaldivergence and the establishment of a sterility barrierbetween the two cytotypes. Clearly, the extreme poly-morphism of A. flaccidum remarked on by earlierauthors was partly due to the inclusion within theone taxon of two quite separate species which arethemselves now known to be fairly polymorphic. Asimilar situation almost certainly exists amongstother species aggregates for which more than onechromosome number has now been determined.

It is very significant that all the newly discoveredcytotypes are at the octoploid level. Moreover, it isremarkable that within the New Zealand Aspleniumflora there is not a single diploid species. The genus inthis country includes approximately equal numbers oftetraploids (9) and octoploids (8) and displays ahigher mean level of ploidy (5.9) than in any otherarea for which sufficient records exist (cf. Lovis1973). The situation in New Zealand contrastsstrongly with that in Europe where over half theAsplenium species are diploid and the mean level ofploidy is only 2.9. In fact, there is a gradual increasein the percentage of diploid species and a decreasein the mean level of ploidy as one moves northwardsfrom New Zealand through the tropics and into theNorthern Hemisphere. This has led Lovis (1973) tospeculate that Asplenium may have originated inGondwanaland with "the main diversification occur-ring after the genus had reached tropical latitudes,only subsequently migrating into northern temperateregions". Thus, Asplenium may now display itshighest level of ploidy in the areas where it hasexisted longest. Lovis supports this hypothesis bypointing out that, in contrast to such northern generaas Dryopteris and Cystopteris which are deciduousand eminently well adapted to the seasonal climateof northern Europe, Asplenium is evergreen andmany species may be found undergoing meiosisthroughout the year.

Nevertheless, many of the higher polyploid sexualspecies of Asplenium in New Zealand are endemicand have almost certainly originated here, whereasthe tetraploid species are very largely of wider distri-bution. Thus, there are three natural groups of closelyrelated species in New Zealand: (1) the A. flaccidumaggregate with thick and rather leathery bipinnatefronds, (2) the A. hookerianum/bulbiferum complexwith thin highly dissected fronds, and (3) the A.lucidumjobtusatum complex with thick fleshy pinnatefronds.

Within the A. flaccidum aggregate there are justtwo species*; of these, one (A. flaccidum sens, str.)is tetraploid and widely distributed in Australia, NewZealand, and Polynesia; the other (A. terrestre) isoctoploid and thought to be endemic to New Zea-land. Botanists familiar with the floras of bothAustralia and New Zealand have often remarked onthe polymorphism and wide ecological range of A.flaccidum in this country compared to its moreuniform morphology and constant epiphytic habitin Australia, suggesting that the terrestrial species(A. terrestre) is absent from that region.

Within the A. hookerianum/bulbiferum groupthere are two cytotypes of A. bulbiferum of whichthe tetraploid occurs in both Australia and NewZealand, but of which the octoploid is so far onlyknown from this country. Similarly, the tetraploidspecies, A, hookerianum, is known from both coun-tries, but its octoploid relative, A. richardii, isendemic to New Zealand.

Within the A. lucidumjobtusatum aggregate, thetetraploid cytotype of A. obtusatum is widely dis-tributed in the Southern Hemisphere; A. lucidum isalso thought to occur on a few islands outside theNew Zealand region. However, the various octoploidtaxa, A. obtusatum subsp. northlandicum, A. sclero-prium, and A. lyallii are all strictly endemic.

The remaining taxa are neither closely related toeach other nor to the three groups mentioned above.A. trichomanes has two cytotypes of which thetetraploid is cosmopolitan (though rare in NewZealand) but the hexap'oid is almost confined tothis country. A. lamprophyllum is a tetraploid speciesapparently restricted to the northern part of theNorth Island but closely related to A. cuneatumLam. of widespread tropical distribution. A. polyodonis also tetraploid and in New Zealand is at thesouthernmost limit of its range which extends throughthe Indo-Malaysian tropical region. The remainingspecies, A. flabellifolium, is apomictic and thereforenot strictly comparable to the sexual taxa, but it hashexaploid cytotypes in both New Zealand and Aus-tralia whereas the octoploid is so far only knownfrom this country.

The evidence therefore strongly suggests that theNew Zealand Asplenium flora has two components:an ancestral element of widely distributed, sexually-reproducing, tetraploid species, and an endemicelement of higher polyploid taxa which haveoriginated within New Zealand from these tetraploid

'Excluding A. shuttleworthianum which occurs onlyon the fringes of the New Zealand Botanical Region.


parents. The derivation of the individual octoploidswill be discussed in a future publication whenevidence from naturally occurring hybrids can alsobe considered. However, it seems likely that processesof auto- and allopolyploidy may have been as impor-tant in the evolution of the New Zealand Aspleniumflora as they are now known to have been inEurope and North America.

ACKNOWLEDGMENTS

This work has been carried out during the tenureof a Post-doctoral Research Fellowship supported byVictoria University of Wellington. I am extremelygrateful to Professors H. D. Gordon and J. K. Heyesfor the opportunity to work in the Department ofBotany, and for financial assistance in carrying outfield work.

I thank the following who have collected materialfor me during this investigation: Mr R. G. Bagnall,Drs J. E. and R. E. Beever, Drs A. E. and B. D.Bell, Mr B. D. Bell, Dr J. E. Braggins, Mr R. J.Chinnock, Mr H. D. Christophers, Dr G. B. Cone,Dr D. R. Given, Mr G. S. Hardy, Professor J. K.Heyes, Miss M. McDonald, Mrs E. Miller, Dr F. B.Sampson, Dr J. E. Sheridan, Mr C. J. Smuts-Kennedy,Mr B. V. Sneddon, Mr G. Stephenson, Mr G. Walls,and Mr A. E. Wright. I also thank my wife andmany friends who have assisted me with field work.

I am indebted to the curators of the followingherbaria who have sent valuable material on loan toWellington: AK, AKU, BM, CHR, GOET, K, P, W,WELT. I particularly thank Miss B. H. Macmillan(Botany Division, DSIR) for packing and dispatch-ing the entire collection of Asplenium in CHR, DrJ. E. Braggins, Mr A. E. Esler, and Mr A. E.Wright for allowing me to examine their personalherbaria, and Mr L. J. Metcalf of ChristchurchBotanic Gardens for sending me type specimens fromthe Armstrong Herbarium.

Finally, special thanks go to Mr R. Lucas forhis care in maintaining a large collection of plantsin cultivation, to Mrs J. Benfield for help in thepreparation of the illustrations, to Dr W. A. Sledgeand Dr J. D. Lovis for their criticism of the manu-script and valuable advice on matters of nomen-clature, and to Dr R. Ross of the British Museumfor his opinion on the correct name for A. flabelli-folium.

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- 1958: Chromosome numbers in New Zealandferns. Ibid. 85: 213-16.

• 1961: Additional chromosome numbers - NewZealand ferns. Ibid. (Botany) 1: \-A.

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- 1883: A description of four new ferns fromour New Zealand forests. Transactions andProceedings of the New Zealand Institute 15:304-10.

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- 1966b: Contributions to the botany of Cod-fish Island, Stewart Island. Transactions of theRoyal Society of New Zealand (Botany) 3:111-22.

- 1969: The flora of the Snares Islands, NewZealand. Ibid. 3: 237-70.

- 1973: A botanical survey of seven mutton-bird islands, south-west Stewart Island. Journalof the Royal Society of New Zealand 3:475-525.

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• 1954: Cytological notes on one hundredspecies of Malayan ferns. In Holttum, R. E."A Revised Flora of Malaya". Vol. II.Government Printing Office, Singapore. 643 pp.

MANTON, IRENE; SLEDGE, W. A. 1954: Observationson the cytology and taxonomy of the pteri-dophyte flora of Ceylon. Philosophical Tran-sactions of the Royal Society of London,Series B 238: 127-85.

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6===Taxonomic Revision Asplenium===