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Home , Austrolycopodium, Diphasium, Huperzia australiana, Huperzia selago, Lycopodiaceae, Lycopodiella

FLORA OF AND

LYCOPODIACEAE

P.J. BROWNSEY & L.R. PERRIE

Fascicle 27 – JUNE 2020 © Landcare Research New Zealand Limited 2020. Unless indicated otherwise for specific items, this copyright work is licensed under the Creative Commons Attribution 4.0 International licence Attribution if redistributing to the public without adaptation: "Source: Manaaki Whenua – Landcare Research" Attribution if making an adaptation or derivative work: "Sourced from Manaaki Whenua – Landcare Research" See Image Information for copyright and licence details for images.

CATALOGUING IN PUBLICATION Brownsey, P. J. (Patrick John), 1948– Flora of New Zealand : ferns and lycophytes. Fascicle 27, / P.J. Brownsey and L.R. Perrie. -- Lincoln, N.Z.: Manaaki Whenua Press, 2020. 1 online resource ISBN 978-0-947525-66-8 (pdf) ISBN 978-0-478-34761-6 (set) 1.Ferns -- New Zealand – Identification. I. Perrie, L. R. (Leon Richard). II. Title. III. Manaaki Whenua – Landcare Research New Zealand Ltd. UDC 582.381(931) DC 587.90993

DOI: 10.7931/kyrt-4p89

This work should be cited as: Brownsey, P.J. & Perrie, L.R. 2020: Lycopodiaceae. In: Breitwieser, I.; Wilton, A.D. Flora of New Zealand - Ferns and Lycophytes. Fascicle 27. Manaaki Whenua Press, Lincoln. http://dx.doi.org/ 10.7931/kyrt-4p89

Date submitted: 23 Sep 2019 ; Date accepted: 11 Dec 2019

Cover image: billardierei. Pendulous epiphytic with stems dividing dichotomously many times. Contents

Introduction...... 1 Taxa Lycopodiaceae P.Beauv...... 2 Bernh...... 4 (Herter) Holub ...... 5 Holub ...... 7 Lycopodiella cernua (L.) Pic.Serm...... 8 Lycopodiella diffusa (R.Br.) B.Øllg...... 11 Lycopodiella lateralis (R.Br.) B.Øllg...... 13 Lycopodiella serpentina (Kunze) B.Øllg...... 16 L...... 17 Lycopodium deuterodensum Herter ...... 19 Lycopodium fastigiatum R.Br...... 21 Lycopodium scariosum G.Forst...... 24 Lycopodium volubile G.Forst...... 27 Phlegmariurus Holub ...... 29 Phlegmariurus billardierei (Spring) Brownsey & Perrie ...... 30 Phlegmariurus varius (R.Br.) A.R.Field & Bostock ...... 34 Kunze ...... 38 Phylloglossum drummondii Kunze ...... 38 References ...... 41 Acknowledgements ...... 45 Maps ...... 46 Index ...... 48 Image Information ...... 49

Introduction

Lycopodiaceae is a large, cosmopolitan family that has its greatest diversity in humid montane forests and alpine areas of tropical regions, but also extends into the temperate zones. It is represented in New Zealand by five indigenous genera (Huperzia, Lycopodiella, Lycopodium, Phlegmariurus and Phylloglossum) and 12 species, of which only Phlegmariurus billardierei is endemic. Huperzia, characterised by its lack of strobili and the presence of bulbils, has a single terrestrial species in New Zealand that occurs in montane to alpine areas of both main islands. Lycopodiella includes four species: L. cernua, with looping stems, is a widespread tropical species common in the northern half of the North Island and extending to the West Coast of the South Island; L. diffusa and L. lateralis are found on pākihi and other poor soils in both islands and are characterised by strobili borne laterally on the aerial stems; and L. serpentina is a rare and threatened species found only on boggy soils around Kaitāia and between Thames and Hamilton. Lycopodium includes four mostly widespread species: L. scariosum and L. volubile are scrambling species with dimorphic sterile flattened in one plane; L. fastigiatum is a montane to subalpine species with incurved leaves and stalked strobili; and L. deuterodensum is a lowland species of the northern North Island with dendroid aerial stems and sessile strobili. Two widespread species are recognised in Phlegmariurus, both of them either epiphytic pendent of forest, or terrestrial plants of more open areas. Phylloglossum is a very distinctive monotypic of gumland and poorly drained soil in Northland and Auckland. Members of Lycopodiaceae in New Zealand are representative of the range of variation in the family as a whole, with terrestrial, epiphytic, scrambling and climbing species. They are characterised by monomorphic or dimorphic non-ligulate leaves with single, unbranched veins, and isotomous or anisotomous branching. Sporangia are homosporus, solitary on the adaxial surface of the , and usually aggregated into strobili.

1 Lycopodiaceae P.Beauv. in Lamarck & Mirbel, Hist. Nat. Veg. 4, 293 (1802) Type taxon: Lycopodium L. Vernacular name: clubmosses Terrestrial, epiphytic, scrambling or climbing plants of varied habit; either with indeterminate, horizontal and subterranean, creeping, looping or climbing stems, giving rise to determinate, erect or inclined, simple or branched, aerial stems, usually of different thickness to the horizontal stems; or with erect or pendent, dichotomously branched stems of equal thickness, lacking horizontal indeterminate stems; very rarely with a simple stem and a cluster of leaves growing from a subterranean tuber. Scales absent. either arising at varying intervals along the length of horizontal stems, or forming a single basal tuft on erect or pendent species. Leaves appearing to be spirally arranged, or regularly or irregularly whorled, or in longitudinal ranks, lacking a ligule, undivided, with a single unbranched vein; all similar or sometimes gradually reducing towards the apex, or rarely of two kinds and arranged in ranks (anisophyllous); margins entire or minutely ciliate. Bulbils rarely present on aerial stems, or usually absent. Sporophylls similar to the leaves, or modified and aggregated into distinct strobili, persistent or ephemeral, monomorphic. Strobili usually terminal on ultimate branches or rarely lateral on overtopping branches, erect or pendent, sessile or stalked. Sporangia solitary, in axils or on adaxial surface of sporophylls, monomorphic, homosporous. trilete, cream or yellow, foveolate, rugulate, reticulate and baculate (NZ), or scabrate (not NZ). : A family with five genera (Burnard et al. 2016) and c. 388 species (PPG 1 2016). Lycopodiaceae have long been recognised as one of three isolated families within the , distinguished by their terrestrial, epiphytic or climbing/scrambling habit, non-ligulate leaves, and homosporous sporangia, usually borne in strobili. Subdivision of the family has been subject to considerable disagreement, with anything from two to 16 genera having been recognised. Traditionally the monotypic Phylloglossum was accepted alongside the large genus Lycopodium, which was the classification used by Allan (1961) for New Zealand species. However, a detailed investigation of the family led Øllgaard (1987, 1989, 1990) to recognise four genera: Huperzia, Lycopodiella, Lycopodium and Phylloglossum, with four sections in Lycopodiella and nine in Lycopodium. In doing so, he rejected the much more divisive classifications of Rothmaler (1944) and Holub (1964, 1975, 1983, 1985, 1991), who recognised two families (Huperziaceae and Lycopodiaceae) and subdivided Huperzia into two genera, Lycopodiella into four, and Lycopodium into seven. PPG 1 (2016) has accepted the more divisive approach and extended the subdivision of Lycopodium to nine genera, with a total of 16 in the whole family. Molecular analysis of Lycopodiaceae based on four chloroplast loci in 119 taxa (Field et al. 2016) demonstrated that both the broader classifications of Øllgaard and the divisive scheme of PPG 1 are monophyletic, leaving it a matter of interpretation as to which should be accepted. Field et al. (2016) showed that, within the huperzioid clade, there is a clear choice between accepting one genus (Huperzia) or three (Huperzia, Phlegmariurus and Phylloglossum). There are good morphological and geographical grounds for recognising three genera (discussed below under each genus), and hence they are accepted here. Within the lycopodioid clade there is strong morphological and molecular evidence for distinguishing Lycopodiella and Lycopodium, but the evidence for subdividing them is less compelling. A monophyletic arrangement requires recognition of either two genera with about 50 species in each, or 13 small genera, of which three are monotypic and four have five species or less (PPG 1 2016). Morphological support for these smaller genera is weak, or depends on obscure characters, and only the two broader genera are accepted here. Furthermore, our approach minimises differences from schemes used recently in New Zealand.

1 Tiny plants with a subterranean tuber, a rosette of linear leaves, and a short on a leafless stalk...... Phylloglossum Small to large plants, lacking a subterranean tuber, bearing leaves on elongated stems, and strobili that are sessile or borne on leafy stalks...... 2

2 2 All stems tufted, erect or pendent, branching dichotomously and of equal thickness; lacking indeterminate, horizontal stems; roots forming a basal tuft; leaves and sporophylls similar, or sporophylls smaller and persistent; spores foveolate...... 3 Stems branching dichotomously, of unequal thickness; main stems indeterminate and horizontal, giving rise to branched or unbranched, determinate, aerial stems; roots arising at intervals along the horizontal stems; sporophylls strongly modified and ephemeral; spores rugulate, reticulate or baculate...... 4 3 Terrestrial plants; bulbils present in axils of leaves; sporophylls not aggregated into distinct strobili...... Huperzia Epiphytic or sometimes terrestrial plants; bulbils absent; sporophylls aggregated into ± distinct terminal strobili...... Phlegmariurus 4 Strobili lateral on aerial stems, tan-brown, dark red-brown or chocolate- brown...... Lycopodiella Strobili terminal on aerial stems, yellow-brown or orange-brown...... 5 5 Strobili erect and terminating branching aerial stems, or strobili pendulous and stalked...... Lycopodium Strobili erect and terminating unbranched aerial stems, or strobili pendulous and sessile...... Lycopodiella Distribution: A large, cosmopolitan family, absent only from the arid zones, which reaches its greatest diversity in humid montane forests and alpine areas of tropical regions (Øllgaard 1990). Twelve indigenous species in New Zealand, one endemic. Biostatus: Indigenous (Non-endemic). Table 1: Number of species in New Zealand within Lycopodiaceae P.Beauv. Category Number Indigenous (Endemic) 1 Indigenous (Non-endemic) 11 Total 12 Recognition: Lycopodiaceae comprise terrestrial, epiphytic or climbing/scrambling lycophytes bearing non-ligulate leaves, which are either monomorphic or dimorphic with single, unbranched veins. Sporangia are homosporous, solitary in the axils or on the adaxial surface of the sporophylls, and usually aggregated into strobili. In huperzioid lycophytes the branching is isotomous, with all dichotomies resulting in equally thick branches, and the roots form a basal tuft. In lycopodioid plants the branching is anisotomous, with indeterminate horizontal main stems and determinate aerial branch systems, and the roots arise along the lower side of the main stems. Notes: The anatomy and morphological variability of New Zealand species of Lycopodiaceae were studied in detail by Holloway (1910, 1919), along with observations on methods of vegetative reproduction (Holloway 1917) and investigations of their life cycles and generations (Holloway 1916, 1920). The spores of all New Zealand species were described and illustrated by Large & Braggins (1991).

3 Huperzia Bernh., J. Bot. (Schrader) 1800(2): 126 (1801) = Urostachys Herter, Beih. Bot. Centralbl. Abt. 2, 39: 249 (1922)

Type taxon: (L.) Bernh. ex Schrank & Mart. Etymology: Named in honour of Johann Peter Huperz (1771–1816), a German physician and botanist. Terrestrial plants. Stems erect, dichotomously branched, with branches of equal thickness, lacking horizontal stems. Roots arising from the stem stele and descending through the cortex to the stem base, forming a single basal tuft. Leaves spirally arranged, all similar; leaf margins entire or minutely toothed at the apex. Bulbils present in axils of some leaves. Sporophylls similar to the leaves, not aggregated into distinct strobili, persistent. Spores foveolate or fossulate, triangular in polar view with concave lateral margins and truncate corners, and foveolate sculpture on both proximal and distal faces. Taxonomy: A genus of 25 species (PPG 1 2016). The huperzioid clade within Lycopodiaceae includes three distinct genera: Huperzia, Phlegmariurus and Phylloglossum (Field et al. 2016). However, under earlier classifications Huperzia was more broadly circumscribed to include two of those genera, Huperzia and Phlegmariurus (Øllgaard 1987), or all three (Christenhusz et al. 2011). New Zealand species of Huperzia and Phlegmariurus were treated in Lycopodium by Allan (1961). The huperzioid clade is distinguished by isotomous branching, with all dichotomies resulting in equally thick branches; roots arising from the stem stele and descending through the cortex to the stem base, forming a basal tuft; sporophylls that are either similar to the leaves, or modified, aggregated into strobili and persistent (Øllgaard 1987, 1990; Field et al. 2016); and with foveolate or fossulate spores (Tryon & Lugardon 1991, figs 227.1–227.14; Large & Braggins 1991). Huperzia is distinguished from Phlegmariurus by bearing reproductive bulbils in the leaf axils (Øllgaard 1987, fig. 22), having sporophylls that are not aggregated into strobili, and producing distinctive spores that are triangular in polar view with concave, lateral margins and truncate corners, and foveolate sculpture on both proximal and distal faces (Tryon & Lugardon 1991, figs 227.1–227.3). They are typically small, terrestrial plants (Field et al. 2016). Huperzia is also easily distinguished from Phylloglossum​​​​, which is a tiny terrestrial plant with a subterranean tuber, a whorl of linear leaves, and a small strobilus borne on a leafless stalk. Distribution: Huperzia is a genus of 25 species, mostly occurring in forests and alpine habitats of the north temperate region, but with some representatives in high-altitude tropical floras, and a very few in the south temperate zone (Field et al. 2016); one, or possibly two, species in (Chinnock 1998; Field & Renner 2019). One indigenous, non-endemic species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 2: Number of species in New Zealand within Huperzia Bernh. Category Number Indigenous (Non-endemic) 1 Total 1 Cytology: Chromosome numbers in Huperzia s.s. vary from 2n = 90 to 2n = over 400. A common base number is x = 11, but there are many exceptions (Field et al. 2016). Notes: Many of the species now placed in Huperzia and Phlegmariurus were previously included in Urostachys. However, the type of Urostachys is Lycopodium selago L., now in Huperzia, and hence Urostachys is a synonym of Huperzia, not Phlegmariurus.

4 Huperzia australiana (Herter) Holub, Folia Geobot. Phytotax. 20: 70 (1985) ≡ Lycopodium australianum Herter, Bot. Jahrb. Syst. 43: Beiblatt 98: 42 (1909) ≡ Urostachys australianus (Herter) Herter ex Nessel, Bärlappgewächse 49 (1939) Lectotype (selected by Brownsey et al. 2018): I.D.L. [Insula Diemen’s Land = ], Gunn 328, 1833, K 001193763 (!online). = Urostachys cockaynei Herter ex Nessel in Nessel, Revista Sudamer. Bot. 6: 158, t. 7, f. 12 (1940) Lectotype (selected by Øllgard 1989): Neu-Seeland, Süd Island [New Zealand, South Island], R. Helms s.n., BONN-Nessel 38 p.p. (image!)

Etymology: australianus (Latin) – pertaining to Australia. Vernacular name: fir clubmoss Horizontal stems absent. Aerial stems erect or decumbent, unbranched or branching dichotomously 1–4 times, 35–480 mm long, 4–18 mm wide (including leaves). Sterile leaves monomorphic, spirally arranged, decreasing in size slightly towards the apex, appressed or spreading, narrowly ovate, apices acute or acuminate, margins entire or minutely toothed at apex, bases adnate, 3–10 mm long, 0.8–1.4 mm wide, dull green or tinged orange. Bulbils present on the stems. Sporophylls similar to sterile leaves, confined to upper half of stem, not aggregated into strobili. Distribution: North Island: Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Chatham Islands, Stewart Island, Auckland Islands, Campbell Island. Altitudinal range: 100–1900 m. Huperzia australiana occurs in montane and subalpine regions of the North Island from the Raukūmara Range and Rotorua south to the Tararua Range. It is mostly found from 500 m to 1750 m, but has been recorded down to 250 m in the western Waikato. There is a record from Waipoua Forest (AK 99780), but the locality is considered doubtful and has not been mapped. In the South Island the species occurs throughout in montane to alpine areas, usually between 450 m and 1900 m, but occasionally down to 100 m in Westland. It also extends to the Chatham Islands, Stewart Island, the Auckland Islands and Campbell Island. Also Borneo, Sumatra, Java, Celebes, Papua New Guinea, Fig. 1: Huperzia australiana distribution map based on databased records at Australia (New South Wales, , Tasmania) and Macquarie AK, CHR & WELT. Island. Biostatus: Indigenous (Non-endemic). Habitat: Huperzia australiana grows terrestrially, often on stream banks, in podocarp and beech forest, under mānuka, Olearia and Dracophyllum scrub, in subalpine shrub and herbfield, in tussock grassland, on boulders, on rock outcrops and in crevices, on scree and cliffs, and in boggy ground. It occurs on greywacke, limestone, marble, ultramafic rock, and on a range of volcanic rock types. Recognition: Huperzia australiana is easily recognised by the presence of bulbils in the axils of the leaves, and is the only species of Lycopodiaceae in New Zealand that is bulbiferous. It is also distinguished by its upright, terrestrial habit, and by the absence of distinct strobili. Notes: The name Lycopodium selago L. used in many earlier New Zealand Flora treatments is a misidentification of Huperzia australiana. Allan (1961) made the superfluous new combination Lycopodium australianum (Herter) Allan, believing incorrectly that Herter’s original basionym was Urostachys australianus, rather than Lycopodium australianum.

5 Fig. 2: Huperzia australiana. Mature plant Fig. 3: Huperzia australiana. Mature plant showing showing erect, sparingly branched stems. developing bulbils in axils of leaves.

Fig. 4: Huperzia australiana. Mature plant Fig. 5: Huperzia australiana. Mature plant showing growing amongst rocks. sporophylls not aggregated into strobili, bearing solitary sporangia on the adaxial surface.

Fig. 6: Huperzia australiana. Mature plant Fig. 7: Huperzia australiana. Stems bearing a growing amongst rocks. mixture of sporophylls and bulbils.

6 Fig. 8: Huperzia australiana. Stems bearing Fig. 9: Huperzia australiana. Close-up of well- unusually well-developed bulbils in the axils of developed bulbils. the leaves.

Lycopodiella Holub, Preslia 36: 20, 22 (1964) = Lepidotis P.Beauv. ex Mirbel in Lamarck & Mirbel, Hist. Nat. Vég. 3, 477 (1802) nom. rej. = Franco & Vasc. ex Vasc. & Franco, Bol. Soc. Brot. Ser. 2 41: 24 (1967) nom. cons. = Lateristachys Holub, Folia Geobot. Phytotax. 18: 441 (1983) = Holub, Folia Geobot. Phytotax. 18: 441 (1983)

Type taxon: (L.) Holub Etymology: From the related genus Lycopodium and the Latin diminutive suffix -ellus. Terrestrial plants. Main stems indeterminate, horizontal and creeping or looping, giving rise to determinate, erect, simple or branched, aerial stems, often of different thickness to the horizontal stems. Roots arising at intervals along the length of horizontal stems. Leaves spirally arranged, usually all similar; leaf margins entire or minutely ciliate. Bulbils absent. Sporophylls modified and aggregated into distinct strobili, ephemeral. Strobili terminal on ultimate branches or lateral on overtopping branches, erect or pendent, sessile or stalked. Spores rugulate. Taxonomy: A genus of c. 54 species (PPG 1 2016, including all species in the subfamily Lycopodielloideae). Following Øllgaard (1987, 1989, 1990) the lycopodioid clade within Lycopodiaceae is interpreted here to include just two genera, Lycopodiella and Lycopodium. However, more recent classifications further subdivide these taxa into four and nine genera, respectively (Field et al. 2016). The lycopodioid clade is distinguished by anisotomous branching, with dichotomies resulting in branches of unequal thickness; horizontal, indeterminate main stems that are creeping, looping or climbing, and give rise to aerial determinate branches; roots that arise at intervals along the length of the main stem; strongly modified sporophylls aggregated into strobili (Øllgaard 1987, 1990); and reticulate, baculate or rugulate spores (Tryon & Lugardon 1991, figs 229.1–12, 230.1–12; Large & Braggins 1991). Lycopodiella is distinguished from Lycopodium by its horizontal and subterranean, prostrate or looping stems, strobili that are terminal and either pendent and sessile, or erect on undivided stalks. In some species the strobili are lateral on overtopping branches. Spores are rugulate (Tryon & Lugardon 1991, figs 230.1–12; Large & Braggins 1991). In New Zealand, all species have spirally arranged leaves of one size. Four species of Lycopodiella (as circumscribed here) are present in New Zealand. In the classification of Field et al. (2016), L. diffusa and L. lateralis would be placed in Lateristachys, L. cernua in Palhinhaea, and L. serpentina in Pseudolycopodiella. New Zealand species of Lycopodiella were included in Lycopodium by Allan (1961).

7 1 Strobili mostly borne laterally on aerial stems, tan-brown, red-brown or chocolate-brown at maturity...... 2 Strobili borne terminally on aerial stems and branches, yellow-brown at maturity...... 3 2 Aerial stems prostrate, 25–160 mm long; strobili red-brown or chocolate- brown at maturity...... diffusa Aerial stems erect or scrambling upwards, 50–700 mm long; strobili tan- brown at maturity...... lateralis 3 Horizontal stems looping across the ground; aerial stems branched many times, the ultimate branches curved downwards; strobili pendent or inclined downwards ...... cernua Horizontal stems prostrate, not looping; aerial stems unbranched, erect; strobili erect...... serpentina Distribution: Lycopodiella is a genus of over 50 species occurring in moist temperate and tropical regions. The greatest diversity is in ; four species in South (Roux 2009), five in Australia (Chinnock 1998). Four indigenous, non-endemic species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 3: Number of species in New Zealand within Lycopodiella Holub Category Number Indigenous (Non-endemic) 4 Total 4 Cytology: Chromosome numbers in Lycopodiella are very variable. In sect. Lycopodiella (= Lycopodiella s.str.) most counts are diploid with n = 78; in sect. Caroliniana (= Pseudolycopodiella) n = 34, 35 and c. 69; and in sect. Campylostachys (= Palhinhaea) counts of n = 104, 108, 110, 136, c. 165 have been reported (Øllgaard 1987, 1990). Notes: The type of Lepidotis was first designated by Rothmaler (1944), who chose L. cernua (L.) P.Beauv. That choice was challenged by Pichi Sermolli (1971), who instead designated L., thereby making Lepidotis a synonym of Lycopodium, a view shared, among others, by Øllgaard (1989, 2012). However, Greuter & Troia (2014) dismissed Pichi Sermolli’s arguments and asserted that Lepidotis is a legitimate name typified by L. cernua, stating that even if Rothmaler’s typification was disregarded, a subsequent typification by Fuchs (1955) could not be rejected. As a consequence, the name Palhinhaea (L.) Franco & Vasc. was made illegitimate because it was based on the previously designated type of Lepidotis. Greuter & Troia therefore proposed the conservation of Palhinhaea against Lepidotis to preserve the use of Palhinhaea and remove the threat to Lycopodiella, a proposal now accepted, leaving Lepidotis a rejected name under Lycopodiella.

Lycopodiella cernua (L.) Pic.Serm., Webbia 23: 166 (1968) ≡ Lycopodium cernuum L., Sp. Pl. 1103 (1753) ≡ Lepidotis cernua (L.) P.Beauv., Prodr. Fam. Aethéog. 108 (1805) ≡ (L.) Vasc. & Franco, Bol. Soc. Brot. Ser. 2 41: 25 (1967) Lectotype (selected by Proctor 1977): “Habitat in Indiis”, no locality, collector or date, LINN- HL 1257.13! = Lycopodium polycephalum Colenso, Trans. & Proc. New Zealand Inst. 27: 401 (1895) ≡ Palhinhaea polycephala (Colenso) Holub, Folia Geobot. Phytotax. 20: 79 (1985) Holotype: New Zealand, presented 1897, W. Colenso, K 00109430!

Etymology: From the Latin cernuus (drooping), a reference to the habit of the strobili. Horizontal stems dichotomously branching, looping across the ground, rooting at intervals 40–670 mm apart (in herbarium specimens), 2–6 mm diameter; leaves widely spaced, incurved, linear, adnate. Aerial stems arising between rooting points, usually one main stem per loop, erect, branching monopodial, glabrous or rarely hairy; longest fertile aerial branching system 50–620 mm tall, 30–270 mm wide; unbranched portion of stem 10–140 mm long, branched portion 40–600 mm long. Branches curling downwards at apices. Sterile leaves monomorphic, spirally arranged or in rough whorls, decreasing in size slightly towards the apex, incurved, linear to narrowly triangular, apices acuminate, margins entire, bases adnate, 3–8 mm long, 0.4–0.8 mm wide, grey-green. Sporophylls appressed when young, somewhat spreading at maturity, ovate, long acuminate, margins ciliate, bases adnate or subpeltate, 1.6–3.0 mm long, 0.7–1.2 mm wide, aggregated into strobili. Strobili solitary, terminal on

8 branches, pendent or inclined downwards, sessile, yellow to yellow-brown, 4–20 mm long, 3–6 mm wide. Note: measurements given above are from herbarium specimens. Longer aerial stems, documented photographically, are known to occur in the wild up to 1800 mm tall. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Westland. Kermadec Islands, Three Kings Islands. Altitudinal range: 0–1050 m. Lycopodiella cernua occurs on Raoul and Macauley Islands in the Kermadec Islands, and in coastal and lowland areas of the North Island from Kaitāia to Mt Tongariro, extending locally into montane regions. There are outlying populations further south around Palmerston North. The species grows from sea level up to 1050 m on Mt Tarawera. In the South Island it extends from north-west Nelson along the west coast to Bruce Bay in south Westland, growing from near sea level, up to 300 m near Denniston. There is an isolated population at Tennyson Inlet in the Marlborough Sounds. Also widespread in tropical and subtropical regions of the Americas, Africa, , , Australia (, Northern Territory, Queensland, New South Wales), Norfolk Island, Lord Howe Island, and most islands of the South Pacific. Fig. 10: Lycopodiella cernua distribution map based on databased records at Biostatus: Indigenous (Non-endemic). AK, CHR & WELT. Habitat: Lycopodiella cernua grows terrestrially on track or roadside banks, on road cuttings and cliffs, on clay soils, on scoria, in gumland, on stabilised dunes, under mānuka, kānuka or Dracophyllum scrub, occasionally under rātā forest, and most abundantly in thermal areas on the edges of steam vents, hot springs and streams or steaming lakes. Recognition: Lycopodiella cernua is easily recognised by its looping horizontal stems; its dendroid aerial stems that arise on the looping stems between rooting points; its ultimate branches, which tend to curl downwards; and its pendent strobili. Cytology: 2n = c. 200 (de Lange et al. 2004). Notes: The combinations Lycopodium cernuum var. curvatum (Sw.) Nessel, Bärlappgewächse 354 (1939) and L. cernuum var. vulcanicum (Blume) Nessel, Bärlappgewächse 354 (1939), based in part on New Zealand material, are almost certainly misidentifications. The types of L. curvatum Sw. and L. vulcanicum Blume are from Jamaica and Java, respectively. Nessel (1939) made almost 40 spurious varieties and forms in L. cernuum which, although unhelpful nomenclaturally, did indicate the range of morphological variation in this species. Burnard et al. (2016) detected three different haplotypes in populations of Lycopodiella cernua from New Zealand, New Caledonia, Fiji, Malaysia, Japan and USA. The New Zealand specimens were identical to that from Japan but different to those from elsewhere. Burnard et al. (2016) suggested that different lineages may be adapted to temperate and tropical habitats. The species also exhibits morphological variation in the hairiness of the branches (Zhang & Iwatsuki 2013) and needs further investigation.

9 Fig. 11: Lycopodiella cernua. Branching aerial Fig. 12: Lycopodiella cernua. Branching aerial stem with mature yellow strobili, sessile at the stem with immature green strobili. ends of branches and inclined downwards.

Fig. 13: Lycopodiella cernua. Mature aerial stem Fig. 14: Lycopodiella cernua. Over-mature aerial and sessile yellow strobili. stem with brown strobili.

Fig. 15: Lycopodiella cernua. Close-up of aerial Fig. 16: Lycopodiella cernua. Close-up of aerial branches showing incurved leaves and branches showing incurved leaves and mature, immature, sessile, green strobili. sessile, yellow strobili.

10 Fig. 17: Lycopodiella cernua. Horizontal stems Fig. 18: Lycopodiella cernua. Plants growing looping across the ground, giving rise to amongst scrub on thermally heated soil. branched aerial stems between rooting points.

Lycopodiella diffusa (R.Br.) B.Øllg., Opera Bot. 92: 176 (1987) ≡ Lycopodium diffusum R.Br., Prodr. Fl. Nov. Holland. 165 (1810) ≡ Lycopodium laterale var. diffusum (R.Br.) Hook.f., Bot. Antarct. Voy. II (Fl. Nov.-Zel.) Part II, 54 (1854) ≡ Lepidotis diffusa (R.Br.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 67 (1944) ≡ Lateristachys diffusa (R.Br.) Holub, Folia Geobot. Phytotax. 18: 441 (1983) Lectotype (selected by Field 2020): Port del Esperance [Port de l’Esperance, Tasmania], R. Brown Iter Austral. No.130, BM 001038074! = Lycopodium ramulosum Kirk, Trans. & Proc. New Zealand Inst. 11: 456, t. 19b (1879) ≡ Lateristachys ramulosa (Kirk) Holub, Folia Geobot. Phytotax. 18: 441 (1983) ≡ Lycopodiella ramulosa (Kirk) B.Øllg., Opera Bot. 92: 176 (1987) Lectotype (selected by Allan 1961): Hokitika, W. Tipler, WELT P004323/A-C!

Etymology: From the Latin diffusus (diffuse, loosely spreading), a reference to the prostrate habit of this species. Vernacular name: carpet clubmoss Horizontal stems dichotomously branching, subterranean, 0.5–2.0 mm diameter, bearing roots at intervals; leaves widely spaced, spreading, narrowly ovate, adnate. Aerial stems prostrate or curving upwards at branch apices, dichotomously branching 1–5 times, glabrous; longest fertile aerial branching system 25–170 mm long, 20–250 mm wide. Sterile leaves monomorphic, spirally arranged, similar throughout, spreading or incurved, linear to narrowly ovate, apices acuminate, margins entire or minutely ciliate at apex, bases adnate, 2.5–6.0 mm long, 0.5–1.1 mm wide, dull green or tinged red- brown or purple-brown. Sporophylls appressed when young, spreading at maturity, ovate to broadly ovate, apices acuminate, margins ciliate, bases adnate, 2.2–3.5 mm long, 1.4–2 mm wide, aggregated into strobili. Strobili solitary, lateral or terminal on aerial stems, erect, sessile or on short leafy lateral branches up to 7 mm long, yellow-brown when young, chocolate-brown at maturity, 4–24 mm long, 1.5–6 mm wide.

11 Distribution: North Island: Volcanic Plateau, Gisborne, Taranaki. South Island: Western Nelson, Sounds-Nelson, Westland, Southland, Fiordland. Stewart Island, Chatham Islands, Auckland Islands. Altitudinal range: 10–1350 m. Lycopodiella diffusa is confined to montane and subalpine areas of the Gisborne, central North Island, Taranaki and Ruahine Ranges, occurring between 550 and 1350 m. In the South Island the species extends down the west coast from the Marlborough Sounds and north-west Nelson to Southland and Stewart Island. It ranges from 10 m near Whanganui Inlet, up to 870 m on the Heaphy Track at Mt Perry. It extends also to the Chatham Islands (CHR 187455, 417491; not recorded by de Lange et al. 2011) and Auckland Islands. Also Australia (Tasmania). Biostatus: Indigenous (Non-endemic). Habitat: Lycopodiella diffusa forms prostrate mats on pākihi or sandy soils, in mossy turf, on boggy ground, at the edge of tarns Fig. 19: Lycopodiella diffusa distribution map based on databased records at and on granite pavement, often with Juncus and Empodisma AK, CHR & WELT. spp., or under Gleichenia, low mānuka, kānuka, epacrid or subalpine scrub. Recognition: Lycopodiella diffusa is distinguished from all other species of Lycopodiaceae in New Zealand, except L. lateralis, by its strobili usually being borne laterally on the aerial stems, and characteristically chocolate-brown at maturity. The aerial stems grow prostrate with only the ends of the branches upturned, branching dichotomously up to five times. In L. lateralis the aerial stems are held erect or scramble upwards, and are generally much longer than in L. diffusa (50–600 mm, cf. 25–170 mm tall). The leaves of L. diffusa are generally incurved, and slightly shorter and wider than in L. lateralis (2.5–6.0 mm long, 0.5–1.1 mm wide; cf. 3.5–9.0 mm long, 0.3–0.9 mm wide). Notes: Burnard et al. (2016) detected several haplotypes in Lycopodiella diffusa and L. lateralis, but they did not correspond to the two different morphologies. Three samples with L. lateralis morphology shared the haplotype found in L. diffusa. Whether this is evidence of hybridisation between the two taxa or a result of ancestral polymorphism is unclear. Their status needs further investigation.

Fig. 20: Lycopodiella diffusa. Prostrate plants Fig. 21: Lycopodiella diffusa. Aerial stems with growing on pākihi soil. strobili borne laterally or terminally on the branches.

12 Fig. 22: Lycopodiella diffusa. Chocolate- Fig. 23: Lycopodiella diffusa. Sward of prostrate coloured strobili on aerial stems. plants, with chocolate-coloured strobili, growing on pākihi soil.

Fig. 24: Lycopodiella diffusa. Prostrate plants at Fig. 25: Lycopodiella diffusa. Prostrate plants at right, growing alongside erect plants of left, growing alongside erect plants of L. lateralis at L. lateralis at left. right.

Lycopodiella lateralis (R.Br.) B.Øllg., Opera Bot. 92: 176 (1987) ≡ Lycopodium laterale R.Br., Prodr. Fl. Nov. Holland. 165 (1810) ≡ Lepidotis lateralis (R.Br.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 67 (1944) ≡ Lateristachys lateralis (R.Br.) Holub, Folia Geobot. Phytotax. 18: 441 (1983) Lectotype (selected by Field 2020): Port Jackson [Sydney, New South Wales], R. Brown Iter Austral. No. 131, BM 001038073! = Lycopodium consimile Colenso, Trans. & Proc. New Zealand Inst. 16: 348 (1884) – as L. consimilis Lectotype (selected by Brownsey et al. 2018): Great Barrier Island, C.P. W[inkelmann], WELT P003375/A-B!

Etymology: From the Latin lateralis (lateral), a reference to the strobili that are characteristically borne laterally in this species. Horizontal stems dichotomously branching, subterranean, 0.6–3.0 mm diameter, bearing roots at intervals; leaves absent or widely spaced, spreading, narrowly ovate, adnate. Aerial stems erect or scrambling upwards, dichotomously branching 1–4 times, glabrous; longest fertile aerial branching system 50–700 mm tall, 35–200 mm wide. Sterile leaves monomorphic, spirally arranged, similar throughout, spreading and often reflexed, linear to narrowly triangular, apices acuminate, margins entire or minutely ciliate at apex, bases adnate, 3.5–9 mm long, 0.3–0.9 mm wide, dull green or tinged red-brown. Sporophylls appressed when young, spreading at maturity, ovate to broadly ovate, apices acuminate, margins ciliate, bases adnate, 1.8–3.0 mm long, 1.2–2.0 mm wide, aggregated into strobili. Strobili solitary, lateral or rarely terminal on aerial stems, erect, sessile or on short leafy lateral

13 branches up to 7 mm long, yellow-brown when young, tan-brown or chocolate-brown at maturity, 5–30 mm long, 2.5–5.0 mm wide. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Westland, Canterbury. Chatham Islands. Altitudinal range: 0–850 m. Lycopodiella lateralis grows best in lowland areas of the North Island, from Te Paki south to the Herangi Range and Bay of Plenty, growing from near sea level to 600 m, but reaching 750 to 850 m in Coromandel and the Kaimai Range. There are also localised populations in the southern Tararua and Rimutaka Ranges, but the species is apparently absent from much of Taranaki, the central North Island and the east coast. In the South Island it extends from north-west Nelson and the Marlborough Sounds to the Cascade Valley in southern Westland. It is confined to lowland areas from near sea level to 250 m, but reaches 500 m on the Stockton Plateau. It also reaches the Chatham Islands. Also Australia (, Queensland, New South Wales, Fig. 26: Lycopodiella lateralis distribution map based on databased Victoria, Tasmania) and New Caledonia. records at AK, CHR & WELT. Biostatus: Indigenous (Non-endemic). Habitat: Lycopodiella lateralis grows on gumland, serpentine, pākihi, sand and other poor soils, in Sphagnum bog, on peat or swampy ground with Gleichenia, Juncus and Empodisma spp., and sometimes in ditches, on damp rock faces or clay banks. It occurs in tussock, under mānuka, kānuka, Dracophyllum and low broadleaved scrub, and sometimes extends into podocarp or broadleaved forest. Recognition: Lycopodiella lateralis is distinguished from all other species of Lycopodiaceae in New Zealand, except L. diffusa, by its strobili being borne laterally on the aerial stems, and in being characteristically chocolate-brown at maturity. The aerial stems grow erect, or scramble upwards through other vegetation, branching dichotomously up to four times, while the horizontal stems are subterranean. In L. diffusa​​​​​​, the aerial stems are prostrate, with only the ends of the branches upturned, and are generally much shorter than in L. lateralis (25–170 mm, cf. 50–700 mm tall). The leaves of L. lateralis are generally spreading or reflexed, and slightly longer and narrower than in L. diffusa (3.5–9.0 mm long, 0.3–0.9 mm wide; cf. 2.5–6.0 mm long, 0.5–1.1 mm wide). Cytology: 2n = c. 122 (de Lange et al. 2004). Notes: Burnard et al. (2016) detected several haplotypes in Lycopodiella diffusa and L. lateralis but they did not correspond to the two different morphologies. Three samples with L. lateralis morphology shared the haplotype found in L. diffusa. Whether this is evidence of hybridisation between the two taxa or a result of ancestral polymorphism is unclear. Their status needs further investigation.

14 Fig. 27: Lycopodiella lateralis. Branching aerial Fig. 28: Lycopodiella lateralis. Branching aerial stems growing erect on pākihi soil. stems growing erect on swampy soil.

Fig. 29: Lycopodiella lateralis. Immature green Fig. 30: Lycopodiella lateralis. Maturing yellow- strobili borne laterally on aerial stems. brown strobili borne laterally on aerial stems.

Fig. 31: Lycopodiella lateralis. Mature Fig. 32: Lycopodiella lateralis. Solitary strobilus chocolate-brown strobili borne laterally on aerial borne laterally on an aerial stem. stems.

15 Lycopodiella serpentina (Kunze) B.Øllg., Opera Bot. 92: 176 (1987) ≡ Lycopodium serpentinum Kunze in Lehmann, Pl. Preiss. 2, 108 (1846) ≡ Pseudolycopodiella serpentina (Kunze) Holub, Folia Geobot. Phytotax. 18: 442 (1983) Lectotype (selected by Brownsey et al. 2018): Albany, Plantagenet, [Western Australia], Oct. 1840, Herb. Preiss No. 1881, P 01219525! (see Øllgaard 1989) = Lycopodium drummondii Spring, Mém. Acad. Roy. Sci. Belgique 24: 35 (1849) ≡ Lepidotis drummondii (Spring) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 66 (1944) Lectotype (selected by Brownsey et al. 2018): New Zealand, Colenso, K 001094155!

Etymology: From the Latin serpentinus (snake-like), a reference to the flexuous and prostrate horizontal stem of this species. Vernacular name: bog clubmoss Horizontal stems dichotomously branching, prostrate and appressed to surface, rooting at intervals 8–30 mm apart, 1–2 mm diameter; leaves imbricate, ascending, linear to narrowly triangular, adnate. Aerial stems erect, unbranched, glabrous; longest fertile stems 15–140 mm tall (including strobili). Sterile leaves monomorphic, spirally arranged, appressed or spreading, linear to narrowly triangular, apices acuminate, margins entire but minutely ciliate at apex, bases adnate, 2.5–5.0 mm long, 0.4–1.0 mm wide, dull green or rarely tinged orange. Sporophylls appressed when young, spreading at maturity, narrowly ovate to ovate, apices acuminate, margins ciliate, bases adnate or subpeltate, 2.5–4.0 mm long, 1.0–1.8 mm wide, aggregated into strobili. Strobili solitary, terminal on branches, erect, sessile, yellow-green, 6–95 mm long, 5–8 mm wide. Distribution: North Island: Northland, Auckland. Altitudinal range: 0–300 m. Lycopodiella serpentina occurs only in a few lowland sites near Kaitāia and again in lowland peat bogs between Thames and Hamilton. There is a poorly documented record from the “Te Paki Ecological Region and District” in the Colenso Herbarium (AK 1016), which has not been mapped. There is suitable habitat for the species in the area, but it has not been recorded since Colenso’s collection, and its presence is unconfirmed and imprecisely located. Also Australia (Western Australia, South Australia, Queensland, New South Wales, Victoria, Tasmania) and New Caledonia. Biostatus: Indigenous (Non-endemic). Lycopodiella serpentina was given a conservation status of Nationally Vulnerable by de Lange et al. (2018). Habitat: Lycopodiella serpentina grows firmly appressed to the ground in boggy areas, on peat, in poorly drained paddock, in gumland, and on sandy soils, often with Sphagnum and Fig. 33: Lycopodiella serpentina distribution map based on databased Empodisma or under stunted mānuka or Gleichenia. records at AK, CHR & WELT. Recognition: Lycopodiella serpentina is distinguished by its appressed, prostrate, horizontal stems that at intervals, and its unbranched aerial stems that terminate in solitary, unbranched strobili. The sterile leaves are also distinctive in having an ascending habit on the horizontal stems, but appressed or spreading on the aerial stems. Notes: The name Lycopodium carolinianum L. used in 19th century New Zealand Flora treatments is a misidentification of Lycopodiella serpentina. Øllgaard (1987) included both Lycopodiella serpentina and L. caroliniana within section Caroliniana (= Pseudolycopodiella Holub). However, Burnard et al. (2016) did not recover these two species as a clade, suggesting that some of the distinguishing morphological characters must be homoplastic within Lycopodiella, and that the current classification needs revision.

16 Fig. 34: Lycopodiella serpentina. Mature plants Fig. 35: Lycopodiella serpentina. Horizontal stems with prostrate horizontal stems appressed to the growing appressed to the ground, with erect aerial ground, and unbranched aerial stems with stems of L. lateralis in the background. unbranched terminal strobili.

Fig. 36: Lycopodiella serpentina. Erect aerial Fig. 37: Lycopodiella serpentina. Prostrate stems with unbranched terminal strobili. horizontal stems, branching dichotomously, lacking fertile aerial stems.

Lycopodium L., Sp. Pl. 1100 (1753) = C.Presl ex Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 64 (1944) = Pseudodiphasium Holub, Folia Geobot. Phytotax. 18: 440 (1983) = Pseudolycopodium Holub, Folia Geobot. Phytotax. 18: 441 (1983) = Holub, Folia Geobot. Phytotax. 26: 90 (1991)

Type taxon: Lycopodium clavatum L. Etymology: From the Greek lykos (a wolf) and pod (a foot), a reference to the appearance of some species. Vernacular names: clubmoss; mātukutuku; whareatua Terrestrial, scrambling or climbing plants. Main stems indeterminate, horizontal and either subterranean, creeping, or climbing, giving rise to determinate, erect or inclined, branched aerial stems, often of different thickness to the horizontal stems. Roots arising at intervals along the length of horizontal stems. Leaves either similar and spirally arranged, or of two or three kinds and arranged in ranks (anisophyllous); leaf margins entire or minutely ciliate. Bulbils absent. Sporophylls modified and aggregated into distinct strobili, ephemeral. Strobili terminal on ultimate branches, erect or pendent, sessile or stalked. Spores reticulate or baculate (NZ) or scabrate (not NZ).

17 Taxonomy: A genus of c. 58 species (PPG 1 2016, including all species in the subfamily Lycopodioideae). Following Øllgaard (1987, 1989, 1990) the lycopodioid clade within Lycopodiaceae is interpreted here to include just two genera, Lycopodiella and Lycopodium. However, more recent classifications further subdivide these taxa into four and nine genera, respectively (Field et al. 2016). The lycopodioid clade is distinguished by anisotomous branching, with dichotomies resulting in branches of unequal thickness; horizontal, indeterminate main stems that are creeping, looping or climbing, and give rise to aerial determinate branches; roots that arise at intervals along the length of the main stem; strongly modified sporophylls aggregated into strobili (Øllgaard 1987, 1990); and reticulate, baculate or rugulate spores (Tryon & Lugardon 1991, figs 229.1–12, 230.1–12; Large & Braggins 1991). Lycopodium is distinguished from Lycopodiella by its horizontal and subterranean, scrambling or climbing stems. Strobili are either erect and sessile or stalked, or pendent and in clusters. Spores are reticulate or baculate (Tryon & Lugardon 1991, figs 229.1–12; Large & Braggins 1991). In New Zealand, some species have sterile leaves flattened in one plane, of two different sizes. Four species of Lycopodium (as circumscribed here) are present in New Zealand. In the classification of Field et al. (2016) they would be placed in four different genera: L. deuterodensum in Pseudolycopodium, L. fastigiatum in Austrolycopodium, L. scariosum in Diphasium, and L. volubile in Pseudodiphasium.

1 Sterile leaves of distal branches flattened in one plane, of two or three markedly different sizes...... 2 Sterile leaves of ultimate branches spirally arranged around stem, all about the same size...... 3 2 Smaller leaves in two rows on dorsal surface and one row on ventral surface; strobili borne on separate fertile branches in pendulous clusters...... volubile Two or three rows of smaller leaves on ventral surface only; strobili erect and solitary on short-leaved stalks, terminal on aerial stems...... scariosum 3 Aerial stems 150–1900 mm long; sterile leaves on mature aerial stems tightly appressed; strobili sessile...... deuterodensum Aerial stems 40–520 mm long; sterile leaves incurved but never appressed; strobili borne on sparsely leaved stalks, 5–60 mm long...... fastigiatum Distribution: A genus of over 50 species, mostly occurring in north and south temperate regions or in alpine areas of the tropics; eight species in southern Africa (Roux 2009), four in Australia (Chinnock 1998). Four indigenous, non-endemic species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 4: Number of species in New Zealand within Lycopodium L. Category Number Indigenous (Non-endemic) 4 Total 4 Cytology: Chromosome numbers in Lycopodium are very variable. In sect. Lycopodium (= Lycopodium s. str.), sect. Annotina (= ), sect. Obscura (= ) and sect. Lycopodiastrum (= Lycopodiastrum) the base number is n = 34; in sect. Complanata (= Diphasiastrum) n = 23; in sect. Diphasium (= Diphasium) n = 34–36, and n = c. 90; and in sect. Magellanica (= Austrolycopodium) n = 31; (Øllgaard 1987, 1990).

18 Lycopodium deuterodensum Herter, Index Lycopod. 15 (1949) nom. nov. pro Lycopodium densum Labill. 1807 ≡ Lycopodium densum Labill., Nov. Holl. Pl. 2, 104, t. 251, f.1 (1807) nom. illeg., non Lycopodium densum Lam. 1778 ≡ Lepidotis densa Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 67 (1944) nom. nov. pro Lycopodium densum Labill. 1807 ≡ Pseudolycopodium densum (Rothm.) Holub, Folia Geobot. Phytotax. 18: 442 (1983) Lectotype (designated by Øllgaard 1989): Terra Van Diemen [Tasmania], Herb. Webbianum ex Herb. Labillardière, FI 004231 (!online)

Etymology: From the Latin deuter- (second) and densum (compact, condensed), an indication that deuterodensum is a replacement name for Labillardière’s original epithet densum, which reflected the habit of the species. Vernacular name: puakarimu Horizontal stems dichotomously branching, subterranean, 1–4 mm diameter, bearing roots at intervals; leaves colourless, imbricate, appressed, narrowly ovate, adnate. Aerial stems, erect, branching monopodial, glabrous; longest fertile stems 150–1900 mm tall, 40–280 mm wide; unbranched portion of stem 25–750 mm long, branched portion 110–1650 mm long. Branches either horizontal, curling upwards or erect. Sterile leaves monomorphic, spirally arranged, dull green or orange-brown, often black-tipped especially on unbranched portions of stems; leaves on unbranched portion of aerial stems appressed, narrowly ovate to ovate, apices long acuminate, margins ciliate and membranous, bases adnate, 3–6 mm long, 0.8–1.3 mm wide; leaves on fertile aerial branches appressed, narrowly ovate, apices acuminate, margins entire or minutely ciliate, bases adnate, 1.7–3.0 mm long, 0.4–0.6 mm wide; leaves on juvenile aerial branches often spreading and larger than on mature branches. Sporophylls appressed when young, spreading at maturity, ovate to broadly ovate, apices acute or acuminate, margins ciliate and membranous, bases adnate to subpeltate, 2–3 mm long, 1–2 mm wide, aggregated into strobili. Strobili solitary or occasionally paired, terminal on branches, erect or ascending, sessile, yellow to yellow-brown, 3–40 mm long, 2.5–7 mm wide. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki. Chatham Islands. Altitudinal range: 0–1050 m. Lycopodium deuterodensum occurs in coastal and lowland sites from Te Paki to East Cape and inland Taranaki, extending locally into montane areas. It grows from near sea-level up to about 1050 m on Mt Tihia and Mt Pihanga, south of Lake Taupō. It has been collected on the Chatham Islands (WELT P003684, P004311; not recorded by de Lange et al. 2011), and may now be extinct there. Also Australia (South Australia, Queensland, New South Wales, Victoria, Tasmania) and New Caledonia. Biostatus: Indigenous (Non-endemic). Habitat: Lycopodium deuterodensum grows most commonly as a terrestrial species in mānuka and kānuka scrub, but also occurs under kauri and broadleaved forest, and on gumland, stabilised dunes, podsols or clay soils. It often grows in extensive Fig. 38: Lycopodium deuterodensum colonies, either on swampy ground or under dry scrub. It is distribution map based on databased occasionally found in Eucalyptus plantations. records at AK, CHR & WELT. Recognition: Lycopodium deuterodensum is recognised by its dendroid aerial stems, which arise from subterranean horizontal stems; its ultimate branches, which are either erect or curl upwards; and its strobili, which are erect or ascending. On mature stems the sterile leaves are appressed, but juvenile stems sometimes have spreading leaves. Cytology: 2n = 68 (Murray & de Lange 2013).

19 Fig. 39: Lycopodium deuterodensum. Mature, Fig. 40: Lycopodium deuterodensum. Mature, branching aerial stems with appressed leaves. branching aerial stems with appressed leaves.

Fig. 41: Lycopodium deuterodensum. Juvenile Fig. 42: Lycopodium deuterodensum. Aerial stem aerial stem with spreading leaves in foreground, with single, sessile, erect strobili terminating and mature aerial stem with appressed leaves branches. Immature strobili are yellow with behind. appressed sporophylls.

Fig. 43: Lycopodium deuterodensum. Mature Fig. 44: Lycopodium deuterodensum. Over-mature strobili with spreading sporophylls on aerial brown strobili with spreading sporophylls on aerial stem. stem.

20 Fig. 45: Lycopodium deuterodensum. Close-up Fig. 46: Lycopodium deuterodensum. Mature of sessile immature strobili with appressed aerial stems growing along a track through scrub. sporophylls, and aerial branches with appressed leaves.

Lycopodium fastigiatum R.Br., Prodr. Fl. Nov. Holland. 165 (1810) ≡ Lycopodium clavatum var. fastigiatum (R.Br.) Benth., Fl. Austral. 7, 675 (1878) ≡ Austrolycopodium fastigiatum (R.Br.) Holub, Folia Geobot. Phytotax. 26: 91 (1991) Lectotype (selected by Field 2020): summit of Table Mountain, V.D.L. [Van Diemen’s Land = Tasmania], R. Brown Iter Austral. No. 129, BM 001038482! = Lycopodium curvifolium Colenso, Trans. & Proc. New Zealand Inst. 20: 234 (1888) nom. illeg., non Lycopodium curvifolium Grev. & Hook. 1831 ≡ Lycopodium arcitenentis Herter, Index Lycopod. 5 (1949) nom. nov. pro Lycopodium curvifolium Colenso 1888 Lectotype (selected by Brownsey et al. 2018): East Cape District, Chambers, Herb. Colenso, WELT P003376/A-B! = Lycopodium scopulosum Colenso, Trans. & Proc. New Zealand Inst. 20: 235 (1888) Holotype: New Zealand, com. W. Colenso 5/1890, K 001094140! = Lycopodium decurrens Colenso, Trans. & Proc. New Zealand Inst. 28: 617 (1896) nom. illeg., non Lycopodium decurrens R.Br. 1810 Type: not located (see Brownsey et al. 2018) = Lycopodium magellanicum var. berggrenii Herter ex Nessel, Revista Sudamer. Bot. 6: 170, t. 16, f. 86 (1940) ≡ Lycopodium berggrenii (Herter ex Nessel) Herter, Index Lycopod. 6 (1949) Holotype: Omatangi, Taupo, S. Berggren, Jan. 1875, S-P-18 (!online); isotype BONN-Nessel 201 (image!) = Lycopodium cochinchense Herter ex Nessel, Revista Sudamer. Bot. 6: 172 (1940) Lectotype (selected by Øllgaard 1989): “Cochinchina”, P.E. Faber s.n., 1884, BONN-Nessel 309 p.p. (image!)

Etymology: From the Latin fastigiatus (fastigiate, with branches clustered, parallel and erect), a reference to the habit of this species. Vernacular names: alpine clubmoss; mountain clubmoss Horizontal stems dichotomously branching, subterranean, 0.9–2.5 mm diameter, bearing roots at intervals; leaves colourless, widely spaced, appressed or spreading, ovate, adnate. Aerial stems erect or prostrate with ends upturned, not flattened in one plane, branching monopodial, glabrous; longest fertile stems 20–520 mm long (including strobili), 10–210 mm wide; unbranched portion of stem 5–240 mm long, branched portion 25–380 mm long. Branches ascending. Sterile leaves monomorphic, spirally arranged or roughly whorled, dull green or orange-brown; leaves on aerial stems incurved, linear or narrowly ovate, apices acuminate, margins entire, bases adnate, 2.5–7.5 mm long, 0.3–1.0 mm wide. Sporophylls appressed when young, spreading at maturity, ovate to broadly ovate, apices acute or acuminate, 2.4–4.5 mm long, 0.7–1.2 mm wide, with brown centres and minutely erose, membranous margins, bases peltate, aggregated into strobili. Strobili solitary or in clusters of 2–3

21 (rarely, one or more may be divided dichotomously), erect, yellow to brown, 5–100 mm long, 3.5–9.5 mm wide, borne on sparsely leaved stalks or rarely sessile; unbranched stalks 5–80 mm long, terminal on aerial branches, erect, bearing leaves in widely spaced rough whorls. Distribution: North Island: Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Chatham Islands, Stewart Island, Antipodes Island, Auckland Islands, Campbell Island. Altitudinal range: 5–1750 m. Lycopodium fastigiatum occurs in montane to subalpine regions of the North Island from Coromandel and East Cape, south to the Tararua and Aorangi Ranges. It grows from 420 m near Wellington, up to 1580 m in the northern Ruahine Range. In the South Island it occurs in montane to subalpine areas throughout, mostly from 450 m, up to 1750 m near Mt Cook. However, it is found down to 300 m in Fiordland and Southland, and at lower altitudes, almost to sea level, on the Chatham Islands, Stewart Island and all subantarctic islands except the Snares. Also Australia (New South Wales, Victoria, Tasmania).

Biostatus: Indigenous (Non-endemic). Fig. 47: Lycopodium fastigiatum distribution map based on databased Habitat: Lycopodium fastigiatum grows terrestrially in open records at AK, CHR & WELT. podocarp, beech and broadleaved forest, under mānuka, in Dracophyllum and Olearia scrub, on forest margins, on road or river banks, in tussock grassland, in subalpine shrubland, herbfield and bog, and in rock crevices and open rocky ground. It occurs on greywacke, limestone and a variety of volcanic rock types. Recognition: Lycopodium fastigiatum is recognised by its much-branched and diverging aerial stems bearing linear, incurved leaves, and strobili that are solitary or borne in clusters of two or three at the end of sparsely leaved stalks. It is distinguished from L. deuterodensum by its stalked rather than sessile strobili, and from L. scariosum and L. volubile by its diverging aerial stems with monomorphic leaves rather than flattened aerial stems with dimorphic leaves. Cytology: 2n = 60 (Murray & de Lange 2013). Notes: Lycopodium decurrens was described by Colenso (1896), but is illegitimate because of an earlier homonym (Brown 1810). It has traditionally been included in the synonymy of L. fastigiatum (Cheeseman 1906, 1925; Allan 1961) but no specimens of this taxon are extant in AK, K or WELT, and its identity remains uncertain. Lycopodium cochinchinense was described by Nessel (1940) from material said to have been collected in Cochinchina and from Rotorua, New Zealand. Øllgaard (1987, 1989) treated it as a synonym of L. fastigiatum and designated the specimen from Cochinchina as the lectotype, although he said it was probably mislabelled. Lycopodium fastigiatum occurs only in New Zealand and Australia. The name Lycopodium magellanicum P.Beauv., and combinations based on it, as used in earlier New Zealand Floras, are misidentifications of L. fastigiatum, but Lycopodium magellanicum var. berggrenii Herter ex Nessel is based on a New Zealand type and treated as a synonym of L. fastigiatum by Øllgaard (1987). The holotype was collected by Berggren from Omatangi, Taupō, in 1875 (Nessel 1940). Lycopodium fastigiatum var. colensoi Nessel was originally described from New South Wales and from Jacksons, Arthur’s Pass, New Zealand (Nessel 1939), but the description was invalid because no Latin diagnosis was provided. Nessel (1940) later validly described the variety, citing only the holotype collected by von Mueller from New South Wales (with specimens in K, P, W and Herb. Boissier), with no mention of the New Zealand specimen. The name is therefore based on an Australian type and not included in the synonymy here.

22 Fig. 48: Lycopodium fastigiatum. Small plant Fig. 49: Lycopodium fastigiatum. Prostrate plant showing prostrate aerial stems and stalked with strongly incurved, orange-brown leaves and strobili terminal on the branches. immature strobili with appressed sporophylls.

Fig. 50: Lycopodium fastigiatum. Plants with Fig. 51: Lycopodium fastigiatum. Plant with horizontal aerial stems spreading over rocks, strongly incurved, orange-brown leaves and bearing orange-brown, maturing strobili. immature strobili with appressed sporophylls.

Fig. 52: Lycopodium fastigiatum. Plant with Fig. 53: Lycopodium fastigiatum. Plant with erect mature strobili and spreading sporophylls. aerial stems, and immature strobili borne singly or in pairs at ends of branches.

23 Fig. 54: Lycopodium fastigiatum. Plant with Fig. 55: Lycopodium fastigiatum. Plant with erect erect aerial stems, and strobili borne singly or in aerial stems, and strobili borne in clusters of two or pairs at ends of branches. three, occasionally dividing dichotomously.

Lycopodium scariosum G.Forst., Fl. Ins. Austr. 87 (1786) ≡ Stachygynandrum scariosum (G.Forst.) P.Beauv., Prodr. Fam. Aethéog. 114 (1805) ≡ Diphasium scariosum (G.Forst.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 65 (1944) Lectotype (selected by Nicolson & Fosberg 2003): New Zealand, Messrs Forster, BM 001048389! = Lycopodium decurrens R.Br., Prodr. Fl. Nov. Holland. 165 (1810) ≡ Lycopodium scariosum var. decurrens (R.Br.) Hook., Cent. Ferns t. 166 (1854) ≡ Diphasium decurrens (R.Br.) Holub, Folia Geobot. Phytotax. 20: 78 (1985) Syntypes (lectotypification in progress; see Mabberley & Moore, in press): Mont. Tabul. ad fluv. Derwent [Table Mountain towards Derwent River, Tasmania], R. Brown Iter Austral. No. 125, BM, K! = Lycopodium lessonianum A.Rich., Voy. Astrolabe, Essai, 59 (1832) Lectotype (selected by Brownsey et al. 2018): Nlle Zélande, Herb. Richard, P 00522977 (!online) = Lycopodium distans Colenso, Trans. & Proc. New Zealand Inst. 20: 236 (1888) Lectotype (selected by Brownsey et al. 2018): high land north of Gisborne, W.K. Chambers, ex W. Colenso, Herb. Cheeseman, AK 1031!

Etymology: From the Latin scariosus (scarious, of thin, dry, membranous texture), a reference to the nature of the ventral leaves and sporophylls in this species. Vernacular name: creeping clubmoss Horizontal stems dichotomously branching, prostrate or scrambling above ground, 1.1–3.0 mm diameter, bearing roots at intervals; leaves spirally arranged, appressed or spreading, linear or narrowly ovate, adnate. Aerial stems erect or ascending, more or less flattened in one plane, branching monopodial, glabrous; longest fertile stems 45–600 mm long (including strobili), 10–220 mm wide; unbranched portion of stem 5–100 mm long, branched portion 40–500 mm long. Sterile leaves dimorphic, flattened in one plane, with 2 rows of larger lateral leaves and 2–3 rows of smaller ventral leaves, yellow-green; lateral leaves sigma-shaped, apices acute, margins entire, bases decurrent, 2.3–5.0 mm long, 0.8–1.5 mm wide, tending to curl upwards; ventral leaves linear or narrowly elliptic, apices acuminate, those near branch apices with scarious tips, margins entire, bases adnate to subpeltate, 2.0–3.0 mm long, 0.4–0.6 mm wide. Sporophylls appressed when young, spreading at maturity, ovate to broadly ovate, apices acuminate with long, scarious tips, 2.8–5.0 mm long, 1.1–2.2 mm wide, with coloured centres and minutely erose, membranous margins, bases subpeltate, aggregated into strobili. Strobili solitary or in pairs, erect, yellow to orange-brown, 7–63 mm long, 5–9 mm wide, borne on short-leaved stalks; stalks 10–195 mm long, terminal on aerial branches, erect, bearing leaves in rough whorls that are shorter than either the sterile leaves elsewhere on the aerial stems or the sporophylls.

24 Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Chatham Islands, Stewart Island, Antipodes Island, Auckland Islands, Campbell Island. Altitudinal range: 0–1430 m. Lycopodium scariosum occurs in lowland to montane regions of the North Island from the Tutamoe Range south to Wellington, extending locally into subalpine areas. It mostly grows in a zone from 300 to 1000 m, but occurs up to 1300 m in the Kaimanawa Ranges, and down to near sea level in some northern localities. In the South Island it occurs in lowland and montane areas, extending locally into subalpine areas, reaching 1430 m in the St Arnaud Range. It is absent only from some parts of inland Otago and is rare in eastern Marlborough and Canterbury. It has been recorded from Banks Peninsula (Wilson 2013), but no herbarium specimens have been located. It extends to the Chatham Islands, Stewart Island and all the subantarctic islands Fig. 56: Lycopodium scariosum except the Snares. distribution map based on databased records at AK, CHR & WELT. Also Borneo, Philippines, New Guinea, Australia (Victoria, Tasmania). Biostatus: Indigenous (Non-endemic). Habitat: Lycopodium scariosum occurs in open beech and podocarp/broadleaved forest, on forest margins and in clearings, under mānuka and kānuka, in subalpine scrub and tussockland, and on pākihi soils. It is a scrambling species that is commonly found on roadside cuttings and banks, on track banks, river banks, rocky ground, scree margins, and around boggy areas and tarns. Recognition: Lycopodium scariosum is a scrambling plant. It is one of two species of New Zealand Lycopodiaceae that have branches bearing dimorphic sterile leaves flattened in one plane. There are two rows of larger, lateral leaves and additional rows of linear leaves on the ventral surface only. It is easily distinguished from L. volubile, which also has flattened branches and dimorphic leaves, but has smaller linear leaves on both the ventral and dorsal surfaces. Lycopodium scariosum is further distinguished from L. volubile by its solitary, erect strobili borne on stalks. Cytology: 2n = 60 (de Lange et al. 2004).

Fig. 57: Lycopodium scariosum. Plants Fig. 58: Lycopodium scariosum. Plants with scrambling over a clay bank. characteristic yellow-green leaves produced in open habitats.

25 Fig. 59: Lycopodium scariosum. Green juvenile Fig. 60: Lycopodium scariosum. Branching aerial strobili and brown mature strobili produced stem with leaves flattened in one plane, and terminally on short stalks on aerial branches stalked strobili borne individually at ends of branches.

Fig. 61: Lycopodium scariosum. Dorsal surface Fig. 62: Lycopodium scariosum. Ventral surface of of aerial branches showing two rows of flattened aerial branches showing two rows of larger lateral lateral leaves only. leaves and two rows of smaller ventral leaves.

Fig. 63: Lycopodium scariosum. Plants Fig. 64: Lycopodium scariosum. Plants scrambling spreading densely under scrub with abundant over a steep clay bank by means of their long juvenile and mature strobili. horizontal stems.

26 Lycopodium volubile G.Forst., Fl. Ins. Austr. 86 (1786) ≡ Lepidotis volubilis (G.Forst.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 67 (1944) ≡ Pseudodiphasium volubile (G.Forst.) Holub, Folia Geobot. Phytotax. 18: 442 (1983) Lectotype (selected by Nicolson & Fosberg 2003): no locality, J. Banks, Herb. G. Forst. No. 319, BM 001048401!

Etymology: From the Latin volubilis (entwining), a reference to the scrambling and climbing habit of this species. Vernacular names: climbing clubmoss; waekāhu; waewaekoukou Horizontal stems dichotomously branching many times, scrambling widely or climbing, 0.8–2.0 mm diameter, bearing roots at intervals; leaves widely spaced, appressed, linear, peltate. Branches from main stems spreading horizontally, monopodial, glabrous, dimorphic; sterile branches flattened in one plane with larger lateral leaves and smaller dorsal and ventral leaves; fertile branches with short linear leaves appressed to stem, bearing pendulous strobili at their apices. Sterile leaves dimorphic, flattened in one plane, with 2 rows of larger lateral leaves, 2 rows of smaller leaves on the dorsal surfaces, and 1 row on the ventral surface, dull green; lateral leaves sigma-shaped, apices acuminate, margins entire, bases decurrent, 2.5–5.0 mm long, 0.5–1.5 mm wide, tending to curl downwards; dorsal leaves linear, apices acuminate, margins entire, bases adnate, 1.5–4.0 mm long, 0.1–0.4 mm wide; ventral leaves similar to the dorsal leaves but shorter, 1.0–1.9 mm long. Sporophylls deeply concave, virtually unchanged at maturity, broadly ovate with long acuminate scarious apices, 1.5–2.2 mm long, 0.7–1.3 mm wide, with coloured centres and minutely toothed membranous margins, bases adnate, aggregated into strobili. Strobili solitary or paired, sometimes dichotomously divided 1–3 times, pendent, yellow becoming brown with age, 5–160 mm long, 1.3–3.5 mm wide, terminal on branchlets. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Kermadec Islands, Three Kings Islands, Chatham Islands, Stewart Island. Altitudinal range: 0–1050 m. Lycopodium volubile occurs on the Kermadec and Three Kings Islands, and in lowland to montane areas throughout much of the North Island, but is rare in the Gisborne Ecological Province. It grows from near sea-level, up to 1050 m on Mt Pihanga south of Lake Taupō. In the South Island it occurs in lowland and montane areas throughout, but is uncommon in drier parts of inland Marlborough, South Canterbury and Otago. It reaches 900 m at Jordan Stream, Marlborough. It extends to the Chatham Islands and Stewart Island. Also Malay Peninsula, Borneo, Papua New Guinea, Solomon Islands, Vanuatu, New Caledonia. It is doubtfully recorded from Australia (Queensland) (Field & Renner 2019). Fig. 65: Lycopodium volubile distribution map based on databased Biostatus: Indigenous (Non-endemic). records at AK, CHR & WELT. Habitat: Lycopodium volubile is a scrambling and climbing plant found in open kauri, podocarp, broadleaved and beech forest, on forest margins, under mānuka and kānuka, in regenerating scrub and forest, among bracken and in tussock. It grows on road and track banks, on river banks, in ditches, on slip faces, on swampy or peaty ground, on stabilised dunes and in open areas in forest. Recognition: Lycopodium volubile is a scrambling or climbing plant commonly found in open forest or scrub. It is one of two species of New Zealand Lycopodiaceae that have branches bearing dimorphic sterile leaves flattened in one plane. There are two rows of larger lateral leaves and additional rows of linear leaves on the ventral and dorsal surfaces. It is easily distinguished from L. scariosum, which also has flattened branches and dimorphic leaves, but has smaller linear leaves only on the ventral surfaces. Lycopodium volubile is distinguished from all other species by having dimorphic sterile and fertile branches, the latter bearing clusters of pendulous strobili, paired or solitary at the end of each axis.

27 Notes: The name Lycopodium durvillaei Bory used by Richard (1832), and combinations based on it, is now regarded as a species of . Citations of L. durvillaei sensu Richard by New Zealand Flora writers are mostly misidentifications of L. volubile.

Fig. 66: Lycopodium volubile. Apex of aerial Fig. 67: Lycopodium volubile. Branched aerial stem showing leaves flattened in one plane. stem with leaves flattened in one plane.

Fig. 68: Lycopodium volubile. Dorsal surface of Fig. 69: Lycopodium volubile. Ventral surface of aerial branches showing two rows of larger, aerial branches showing two rows of larger lateral flattened, lateral leaves and two rows of smaller leaves and one row of smaller ventral leaves. dorsal leaves.

28 Fig. 70: Lycopodium volubile. Plants showing Fig. 71: Lycopodium volubile. Mature brown strobili flattened sterile branches and clusters of borne in clusters on branching stems. pendulous strobili borne on branching stems.

Fig. 72: Lycopodium volubile. Close-up of Fig. 73: Lycopodium volubile. High-climbing plant dichotomously divided strobili borne in pairs at in forest vegetation. the ends of aerial branches.

Phlegmariurus Holub, Preslia 36: 17, 21 (1964) Type taxon: Phlegmariurus phlegmaria (L.) T.Sen & U.Sen Etymology: From Phlegmaria, the epithet of the type species, and the Greek oura (tail), a reference to the fertile part of these plants that are in the form of a long tail (Pichi Sermolli 1971). Vernacular name: tassel ferns Terrestrial or epiphytic plants. Stems erect or pendent, dichotomously branched with branches of equal thickness, lacking horizontal stems. Roots forming a single basal tuft. Leaves spirally arranged, similar or gradually reducing towards the apex; leaf margins entire. Bulbils absent. Sporophylls similar to the leaves or slightly modified, aggregated into more or less distinct strobili, persistent. Strobili terminal on ultimate branches, pendent on epiphytic plants or curved downwards in erect plants, sessile. Spores foveolate or fossulate, triangular in polar view with convex lateral margins, and foveolate sculpture only on the distal surfaces. Taxonomy: A genus of c. 250 species (PPG 1 2016). The huperzioid clade within Lycopodiaceae includes Huperzia, Phlegmariurus and Phylloglossum (Field et al. 2016). Previously Huperzia and Phlegmariurus were combined in a single genus (Øllgaard 1987), or sometimes included also with Phylloglossum (Christenhusz et al. 2011). However, more recent studies have concluded that Phlegmariurus is distinct from Huperzia, and that three genera

29 should be recognised (Øllgaard 2012; Field & Bostock 2013; Field et al. 2016). New Zealand species of Huperzia and Phlegmariurus were all treated in Lycopodium by Allan (1961). The huperzioid clade is distinguished by isotomous branching, with all dichotomies resulting in equally thick branches; tufted roots; sporophylls that are either similar to the leaves, or modified, aggregated into strobili and persistent (Øllgaard 1987, 1990; Field et al. 2016); and foveolate or fossulate spores (Tryon & Lugardon 1991, figs 227.1–227.14; Large & Braggins 1991). Phlegmariurus is distinguished from Huperzia by lacking reproductive bulbils in the axils (Øllgaard 1987, fig. 22), having sporophylls that are aggregated into strobili (at least in New Zealand), and spores with convex lateral margins, and foveolate sculpture only on the distal surfaces (Tryon & Lugardon 1991, figs 227.5–227.14). They are either epiphytic pendent plants, or erect terrestrial or rupestral plants (Field et al. 2016). Phlegmariurus is easily distinguished from Phylloglossum, which is a tiny terrestrial plant with a subterranean tuber, a whorl of linear leaves, and a small strobilus borne on a leafless stalk. Allan (1961) recognised three species in Lycopodium that are here included in Phlegmariurus: L. billardierei, L. novozealandicum and L. varium. However, Brownsey et al. (1985) reduced these to one polymorphic species requiring further investigation. Morphological and molecular analysis of this group by Perrie et al. (2020) has shown that Phlegmariurus billardierei is clearly distinct from P. varius and endemic to New Zealand, but that Colenso’s Lycopodium novozealandicum is just one form of Phlegmariurus varius. Terrestrial and epiphytic forms of P. varius, which were originally recognised as distinct varieties (Brown 1810), are very closely related, and treated here as one species. They occur in Australia as well as New Zealand.

1 Transition from leaf to sporophyll abrupt, strobili distinct; stems (including leaves) 1.5–8 mm wide just before junction with strobilus; longest sporophylls 1.5–3 mm long; strobili 1.5–3 mm wide at widest point...... billardierei Transition from leaf to sporophyll gradual, strobili indistinct at proximal end; stems (including leaves) 6–18 mm wide just before junction with strobilus; longest sporophylls 2.5–11 mm long; strobili 3–17 mm wide at widest point...... varius Distribution: A large genus found mainly in the tropics and subtropics but with some species extending into the southern temperate zones (Field et al. 2016); nine species in Australia (Chinnock 1998). Two indigenous species in New Zealand; one endemic. Biostatus: Indigenous (Non-endemic). Table 5: Number of species in New Zealand within Phlegmariurus Holub Category Number Indigenous (Endemic) 1 Indigenous (Non-endemic) 1 Total 2 Cytology: Chromosome numbers in Phlegmariurus vary from n = 136 to n = c. 278. A possible base number is x = 17, but there are many exceptions (Field et al. 2016).

Phlegmariurus billardierei (Spring) Brownsey & Perrie in Perrie et al., New Zealand J. Bot. 58: 126 (2020 [2019]) ≡ Lycopodium billardierei Spring, Nouv. Mém. Acad. Roy. Sci. Bruxelles 15: 56 (1841) – as billardieri ≡ Lycopodium varium var. billardierei (Spring) Kirk, Trans. & Proc. New Zealand Inst. 16: 377 (1884) ≡ Urostachys billardierei (Spring) Herter ex Nessel, Bärlappgewächse 194 (1939) ≡ Huperzia billardierei (Spring) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 61 (1944) Lectotype (selected by Brownsey et al. 2018): Nlle Zélande [New Zealand], Voy. de l’Astrolabe, 1826–1829, Lesson, P 01228213!

Etymology: Named in honour of Jacques Julien Houttou de Labillardière (1775–1834), French botanist on D’Entrecasteaux’s voyage (1791–1795) to find La Pérouse’s ill-fated expedition. He made extensive collections of Australian plants from south-west Australia and Tasmania, which formed the basis of his Novae Hollandiae plantarum specimen, published 1804–1807. Vernacular names: hanging clubmoss; tassel ; īwituna

30 Horizontal stems absent. Aerial stems pendent, branching dichotomously 3–9 times, 150– c. 1800 mm long (very rarely only 120 mm long in exposed conditions), 10–40 mm wide (including leaves) at widest point, 1.5–8.0 mm wide near the junction with the strobili. Sterile leaves monomorphic, spirally arranged, decreasing in size towards the apex, spreading proximally, appressed distally, linear to narrowly ovate, apices acute, margins entire and not thickened, bases adnate, dull green; longest leaves 12–30 mm long, 1.2–3.0 mm wide; shortest leaves 3.5–8 mm long, 0.6–1.2 mm wide. Transition from sterile leaves to sporophylls ± abrupt. Sporophylls appressed when young, slightly spreading at maturity, ovate to broadly ovate, apices acute, keeled on back, margins entire, bases adnate, 1.5–3.0 mm long, 1.0–1.8 mm wide, green, aggregated into well-defined strobili. Strobili solitary or paired at end of sterile branches, each strobilus undivided or divided dichotomously 1–4 times, pendent, green when young, yellow at maturity, 10–260 mm long, 1.5–3.0 mm wide, sessile, usually strongly 4-angled. Note : measurements given above are from herbarium specimens. Longer stems are known to occur in the wild. AK 1255577 comprises parts of a stem that was said to have measured 2400 mm long, and CHR 291483 a stem said to measure 9 ft (2700 mm) long. Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Westland, Southland, Fiordland. Kermadec Islands, Three Kings Islands, Chatham Islands, Stewart Island. Altitudinal range: 0–1000 m Phlegmariurus billardierei occurs on the Kermadec and Three Kings Islands, and in coastal and lowland areas throughout the North Island from Te Paki south, extending locally into montane regions, but is uncommon in the central North Island. It grows from near sea level to almost 1000 m at Lake Waikaremoana and on Mt Taranaki. In the South Island it is largely confined to coastal and lowland areas of Marlborough, Nelson and Westland, with very scattered populations south of Ōkārito in Fiordland, the Catlins District, and on Stewart Island. The species grows from sea level, up to 760 m near Tākaka, and extends also to the Chatham Islands. Fig. 74: Phlegmariurus billardierei Biostatus: Indigenous (Endemic). distribution map based on databased Habitat: Phlegmariurus billardierei occurs most commonly as an records at AK, CHR & WELT. epiphyte in kauri, podocarp, beech and broadleaved forest, or under mature mānuka and kānuka. It has been recorded growing epiphytically on a variety of forest trees and climbers including Beilschmiedia tarairi, Cordyline sp., Dacrydium cupressinum, Dysoxylum spectabile, Griselinia littoralis, Knightia excelsa, Kunzea spp., Melicytus ramiflorus, Metrosideros excelsa, M. fulgens, M. kermadecensis, M. robusta, Nestegis montana, Olearia traversiorum, Podocarpus totara, Rhopalostylis sapida, Vitex lucens, Weinmannia racemosa and species of Nothofagaceae, often hanging from clumps of Astelia. It has occasionally been recorded on Cyathea dealbata and Dicksonia squarrosa, but is generally not found on tree ferns (unlike P. varius). Sometimes it occurs on the ground, on banks, on rock overhangs, in rock crevices, or at the base of trees. Recognition: Phlegmariurus billardierei is usually a pendent epiphytic species, but occasionally grows terrestrially on banks, on rock overhangs or in rock crevices. The stem divides dichotomously into branches of equal thickness and produces a basal tuft of roots. The sporophylls are smaller than the sterile leaves and aggregated into distinct 4-angled strobili. Phlegmariurus billardierei can be distinguished from P. varius by its generally longer and more divided stems, but most obviously by its strobili. These are distinct from the sterile leaves and clearly defined, unlike those in P. varius​​​​​​, where the transition from leaf to sporophyll is gradual. The longest sporophylls are shorter (1.5–3 mm, cf. 2.5–11 mm long), and the strobili are narrower at their widest point (1.5–3 mm, cf. 3–17 mm wide). The stem (including the leaves) is generally much narrower just before the junction with the strobili than in P. varius​​​​​ (1.5–8 mm, cf. 6–18 mm wide). A fastigiate form is occasionally found. Cytology: Unknown, but see under Phlegmariurus varius.

31 Notes: The names Lycopodium flagellaria Bory and L. phlegmaria L., and combinations based on them, used in early New Zealand Flora treatments are misidentifications of Phlegmariurus billardierei. Lycopodium billardierei was described by Spring (1841) from a plant collected at the Bay of Islands. It was transferred to Phlegmariurus by Perrie et al. (2020) Perrie et al. (2020) segregated Phlegmariurus billardierei from P. varius​​​​​​​ using morphological and genetic evidence. The two species are sympatric in New Zealand.

Fig. 75: Phlegmariurus billardierei. Pendulous Fig. 76: Phlegmariurus billardierei. Plant growing epiphytic plant with stems dividing terrestrially on a steep bank. dichotomously many times.

Fig. 77: Phlegmariurus billardierei. Plant Fig. 78: Phlegmariurus billardierei. Base of an growing terrestrially from a clump of Astelia. epiphytic plant.

32 Fig. 79: Phlegmariurus billardierei. Fig. 80: Phlegmariurus billardierei. Clearly defined Dichotomously dividing aerial stems with strobili with sporophylls much shorter than sterile spreading sterile leaves. leaves. The branches are narrow with appressed sterile leaves just above junction with strobili.

Fig. 81: Phlegmariurus billardierei. Strobili with Fig. 82: Phlegmariurus billardierei. Strongly angled sporophylls much shorter than the sterile leaves. strobili

33 Phlegmariurus varius (R.Br.) A.R.Field & Bostock, PhytoKeys 20: 49 (2013) ≡ Lycopodium varium R.Br., Prodr. Fl. Nov. Holland. 165 (1810) – as varium α umbrosum ≡ Huperzia varia (R.Br.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 247 (1874) ≡ Urostachys varius (R.Br.) Herter ex Nessel, Bärlappgewächse 192 (1939) Lectotype (selected by Chinnock 1998; see Mabberley & Moore, in press: Table Mt [Mt Wellington, Tasmania], R. Brown Iter Austral. 123, BM 001079913 (!online) = Lycopodium varium var. alpinum R.Br., Prodr. Fl. Nov. Holland. 165 (1810) Lectotype (selected by Chinnock 1998; see Mabberley & Moore, in press): Mont Tabul, Van Diemen’s Land, [Mt Wellington, Tasmania], R. Brown Iter Austral. 124, BM 001038064 (!online). = Lycopodium pachystachyum Desv. ex Poiret, Encycl. Suppl. 3, 544 (1814) Holotype: Nova Hollandia [Australia], Herb. Desvaux, P 01238033 (image!) = Lycopodium varium var. gracile Kirk, Trans. & Proc. New Zealand Inst. 16: 377, t. 29 (1884) ≡ Lycopodium billardierei var. gracile (Kirk) Cheeseman, Man. New Zealand Fl. 1036 (1906) ≡ Urostachys saturnii Herter, Index Lycopod. 80 (1949) nom. nov. pro Lycopodium varium var. gracile Kirk 1884 Lectotype (selected by Brownsey et al. 2018): Maitai Valley, Nelson, T. Kirk, April 1873, WELT P004444! = Lycopodium varium var. polaris Kirk, Trans. & Proc. New Zealand Inst. 16: 376 (1884) Lectotype (selected by Brownsey et al. 2018): Ulva, Patersons Inlet, T. Kirk, Jan. 1882, WELT P004247/A-C! = Lycopodium novozealandicum Colenso, Trans. & Proc. New Zealand Inst. 19: 275 (1887) – as novae-zealandicum ≡ Huperzia novozealandica (Colenso) Holub, Folia Geobot. Phytotax. 20: 75 (1985) – as novae- zelandiae Lectotype (designated by Brownsey et al. 2018): no locality, com. W. Colenso 5/90, K 00853048 (!online) = Urostachys helmii Nessel, Repert. Spec. Nov. Regni Veg. 36: 186 (1934) ≡ Huperzia helmii (Nessel) Holub, Folia Geobot. Phytotax. 20: 73 (1985) Lectotype (selected by Øllgaard 1989): Insel Greymouth [Greymouth, South Island], R. Helm, 1885, BONN-Nessel 465 (image!) = Urostachys varius var. polaris Herter ex Nessel, Revista Sudamer. Bot. 6: 376 (1940) nom. illeg., non Lycopodium varium var. polaris Kirk 1884 ≡ Urostachys polaris (Herter ex Nessel) Herter, Index Lycopod. 76 (1949) ≡ Huperzia polaris (Herter ex Nessel) Holub, Folia Geobot. Phytotax. 20: 76 (1985) Holotype: Antipodes Island, L. Cockayne 5442, 1903, BONN-Nessel (n.v.; see Øllgaard 1989)

Etymology: From the Latin varius (variable), a reference to the polymorphic nature of this species. Horizontal stems absent. Aerial stems erect or decumbent or pendent, branching dichotomously 1–7 times or rarely unbranched, 70– c. 1000 mm long, 12–31 mm wide (including leaves) at widest point, 6–18 mm wide near the junction with the strobili. Sterile leaves monomorphic, spirally arranged, decreasing in size towards the apex, spreading proximally and distally, linear or narrowly elliptic, apices acute to obtuse, margins entire and sometimes thickened, bases adnate, dull green; longest leaves 10–19 mm long, 1–3 mm wide; shortest leaves 4.5–10 mm long, 0.8–2.6 mm wide. Transition from sterile leaves to sporophylls gradual. Sporophylls appressed or spreading when young, spreading at maturity, ovate to narrowly ovate, apices acute to obtuse, keeled or not on back, margins entire, bases adnate, 2.5–11 mm long, 0.8–2.2 mm wide, green, aggregated into distinct or indistinct strobili. Strobili solitary or paired at end of sterile branches, each strobilus undivided or divided dichotomously 1–3 times, erect or curled downwards or pendent, green when young, yellow or green at maturity, 10–350 mm long, 3–17 mm wide, not angled to strongly 4-angled or rarely slightly bilaterally flattened.

34 Distribution: North Island: Northland, Auckland, Volcanic Plateau, Gisborne, Taranaki, Southern North Island. South Island: Western Nelson, Sounds-Nelson, Marlborough, Westland, Canterbury, Otago, Southland, Fiordland. Chatham Islands, Stewart Island, Antipodes Island, Auckland Islands, Campbell Island. Altitudinal range: 0–1500 m. Phlegmariurus varius occurs in lowland to montane and subalpine areas of the North Island, from Great Barrier Island south, with outlying populations in the Waipoua and Warawara Forests. It mostly grows above 300 m, reaching nearly 1300 m on Mt Hikurangi and Maungapohatu, but is found in a few localities down to near sea level. In the South Island it occurs in lowland, montane and subalpine regions throughout, ranging from near sea level, up to 1500 m on Mt Starveall, Nelson. It extends to the Chatham Islands, Stewart Island and all the subantarctic islands except the Snares. Also Australia (Queensland, New South Wales, Victoria, Tasmania) and Lord Howe Island. Fig. 83: Phlegmariurus varius distribution map based on databased Biostatus: Indigenous (Non-endemic). records at AK, CHR & WELT. Phlegmariurus aff. varius (a) (WAIK 7743; “tree fern”) was given a conservation status of Data Deficient by de Lange et al. (2018), but this form is included here within the range of diversity of P. varius. ​​​​​​ Habitat: Terrestrial forms of Phlegmariurus varius occur on the ground, on banks, on rock outcrops and rock faces, in rock crevices, in scrub or tussock, on boggy ground, or on fallen logs. The species also occurs as an epiphyte in podocarp, broadleaved and beech forest, or under mature mānuka. It has been recorded growing epiphytically on Cyathea medullaris, C. smithii, Dicksonia fibrosa, D. squarrosa, Elaeocarpus sp., Fuchsia excorticata, Griselinia littoralis, Metrosideros umbellata, Olearia colensoi, Prumnopitys taxifolia, Weinmannia racemosa, Vitex lucens, species of Nothofagaceae, and hanging from clumps of Astelia. It is unlike P. billardierei in occurring quite frequently on tree fern trunks. Recognition: As its name suggests, Phlegmariurus varius is a variable species that either grows erect on the ground and on rock outcrops, or is a pendent epiphytic species. The stem divides dichotomously into stems of equal thickness and produces a basal tuft of roots. The leaves are similar to the sporophylls and grade imperceptibly into strobili that are either rather poorly distinguished and sometimes slightly bilaterally flattened, or clearly defined and 4-angled at their distal ends but poorly distinguished at their proximal ends. In terrestrial plants, the strobili are often curled downwards towards the ground. By contrast, the strobili of P. billardierei are well defined and characteristically 4- angled throughout. The longest sporophylls in P. varius are generally longer than in P. billardierei (2.5–11 mm, cf. 1.5–3 mm long), and the strobili are wider at their widest point (3–17 mm, cf. 1.5–3 mm wide). The stem (including the leaves) is generally much wider just before the junction with the strobili than in P. billardierei (6–18 mm, cf. 1.5–8 mm wide). Cytology: 2n = 256 (Murray & de Lange 2013 – as Huperzia varia). The voucher specimen (AK 303460) is sterile and could be either Phlegmariurus varius or P. billardierei. Notes: Lycopodium varium was described by Brown (1810) from specimens he collected in Tasmania. The species included two infraspecific taxa denoted as α umbrosum and β alpinum, now var. umbrosum and var. alpinum, respectively. Brown did not mention the autonym var. varium, but this was automatically created with the valid publication of the two infraspecific taxa, and is taken as the α epithet (see Mabberley & Moore, in press). The epithet α umbrosum is superfluous and therefore illegitimate. Brown did not identify a holotype for L. varium, but Chinnock (1998) designated R. Brown Iter Austral. 124 from Table Mountain (BM 001038064) as the lectotype. However this is labelled “β alpinum” and refers to L. varium var. alpinum (see Mabberley & Moore, in press). The lectotype for L. varium is Robert Brown Iter Austral. 123 (BM 001079913), cited in error by Chinnock as 124. The name Lycopodium mirabile, based on a collection by Labillardière from Australia, was published by Willdenow (1810) two months before Brown’s L. varium. Strict application of the Code requires replacement of the well-known name L. varium Brown by L. mirabile (Willd.). However, Field &

35 Brownsey (2019) proposed conservation of L. mirabile with a conserved type so that its use is maintained for an Asian species to which it has consistently been applied, and the basionym L. varium is retained for the Australasian species. The long-overlooked name Lycopodium pachystachyum Desv. is a synonym of P. varius based on an Australian type, whereas Kirk’s L. varium var. polaris and var. gracile, Nessel’s Urostachys helmii, and Colenso’s Lycopodium novae-zealandicum are all based on New Zealand types. The latter name, which was used by Allan (1961), is an orthographic error that must be corrected to L. novozealandicum (Webb & Edgar 1999). The number of synonyms for Phlegmariurus varius is a reflection of the morphological variation in the species. Perrie et al. (2020) segregated Phlegmariurus billardierei from P. varius using morphological and genetic evidence. The two species are sympatric in New Zealand. Perrie et al. (2020) found no morphological or genetic evidence to further sub-divide P. varius despite its morphological variability.

Fig. 84: Phlegmariurus varius. Pendulous Fig. 85: Phlegmariurus varius. Pendulous epiphytic epiphytic plant growing high on a forest tree with plant growing on a trunk near ground level. dichotomously branching aerial stems.

Fig. 86: Phlegmariurus varius. Lax, pendulous Fig. 87: Phlegmariurus varius. Short, erect plant epiphytic plant growing on trunk of growing terrestrially at base of trunk with strobili Dicksonia squarrosa. curled over distally.

36 Fig. 88: Phlegmariurus varius. Short, pendulous Fig. 89: Phlegmariurus varius. Short, erect plant plant growing terrestrially on a vertical bank. growing terrestrially with stem apices curled over.

Fig. 90: Phlegmariurus varius. Dichotomously Fig. 91: Phlegmariurus varius. Ill-defined strobili dividing strobili merging imperceptibly with the with sporophylls increasing in length towards sterile leaves. junction with leaves. Branches are broad just above junction with strobili.

Fig. 92: Phlegmariurus varius. Strobili with Fig. 93: Phlegmariurus varius. Strobili with lax, spreading sporophylls almost as long as the spreading sporophylls. leaves at the transition point.

37 Phylloglossum Kunze, Bot. Zeitung (Berlin) 1: 721 (1843) Type taxon: Phylloglossum drummondii Kunze Etymology: From the Greek phyllon (leaf) and glossa (tongue), a reference to the shape of the leaves. Terrestrial plants. Stems erect, unbranched, growing from a subterranean tuber. Roots forming a single basal tuft. Leaves whorled at ground level, terete, linear. Bulbils absent. Sporophylls markedly different to the leaves, aggregated into distinct strobili, ephemeral. Strobili solitary, erect, terminal on leafless stalk. Spores foveolate or fossulate, triangular in polar view with convex lateral margins, and foveolate sculpture only on the distal surfaces. Taxonomy: A monotypic genus (PPG 1 2016). Phylloglossum is a highly specialised genus which has been recognised for over 150 years. It belongs to the huperzioid clade of Lycopodiaceae (Field et al. 2016) but is morphologically distinct from both Huperzia and Phlegmariurus. It is a tiny terrestrial plant with a subterranean tuber, a basal rosette of linear leaves, and a small strobilus borne on a leafless stalk, quite unlike any other member of the family. Distribution: A monotypic genus confined to Australia and New Zealand, where it is adapted to seasonally wet conditions, surviving the dry season as an underground tuber. One indigenous, non- endemic species in New Zealand. Biostatus: Indigenous (Non-endemic). Table 6: Number of species in New Zealand within Phylloglossum Kunze Category Number Indigenous (Non-endemic) 1 Total 1 Cytology: Blackwood (1953) reported a single count of n = 246 II + 10 I from Australian material of Phylloglossum drummondii. Notes: The gametophyte of Phylloglossum in New Zealand has been examined in detail by Holloway (1935) and Whittier & Braggins (1992, 2000).

Phylloglossum drummondii Kunze, Bot. Zeitung (Berlin) 1: 721 (1843) ≡ Huperzia drummondii (Kunze) Christenh. & H.Schneid. in Christenhusz et al., Phytotaxa 19: 20 (2011) Lectotype (designated by Brownsey et al. 2018): ad fluvium cygnorum Novae Hollandiae [Swan River, Western Australia], Drummond, Herb. Shuttleworthianum 993, BM, top left- hand numbered specimen, labelled package and labelled specimens (!online) = Lycopodium sanguisorba Spring, Mém. Acad. Roy. Sci. Belgique 24: 36 (1849) Lectotype (designated by Brownsey et al. 2018): Swan River [Western Australia], Drummond 993, Herb. Hooker., K 001094294!

Etymology: Named in honour of James Drummond (1784–1863), British botanist and plant collector in Australia (1829–1863). Horizontal stems absent; plant growing from a subterranean tuber. Stems erect, unbranched, 11–50 mm long. Leaves whorled at ground level, spreading, linear, apices acuminate, margins entire, bases adnate, 7–20 mm long, 0.5–1.0 mm wide, dull green. Sporophylls appressed when young, spreading at maturity, ovate, apices acuminate, margins entire, bases subpeltate to peltate, 1.2–3.0 mm long, 0.7–1.5 mm wide, aggregated into distinct strobili. Strobili solitary, terminal on leafless stalk, erect, green to yellow-brown, 3–9 mm long, 2–5 mm wide.

38 Distribution: North Island: Northland, Auckland. Altitudinal range: 0–240 m. Phylloglossum drummondii has been recorded in coastal and lowland sites from Te Paki to the Waikato River, with sporadic occurrences reported as far south as Banks Peninsula (de Lange et al. 2010). However, it is now confined to gumland areas in the far north around Kaitāia, growing from near sea-level up to 240 m on the Ahipara Plateau. According to Cheeseman (1906), the species was “said to have been gathered near Picton by Helms, and on Banks Peninsula by Armstrong, but I have seen no specimens”. The plant was recorded by Armstrong (1880) from Banks Peninsula, based on collections made by him and his father, but no voucher material has been found in AK, CHR, MEL or WELT. The origin of the Picton record is unknown. Also Australia (Western Australia, South Australia, Victoria, Tasmania). Biostatus: Indigenous (Non-endemic).

Phylloglossum drummondii was given a conservation status of Fig. 94: Phylloglossum drummondii Nationally Endangered by de Lange et al. (2018). distribution map based on databased records at AK, CHR & WELT. Habitat: Phylloglossum drummondii favours open gumland or sandy soil under mānuka and other scrub that has been recently burned, or poorly drained soils on clay pans and podsols. Recognition: Phylloglossum drummondii is the smallest member of New Zealand’s Lycopodiaceae and easily overlooked in the wild. It is an ephemeral species, appearing above ground only in winter and early spring. It grows from an underground tuber and is distinguished by a basal rosette of linear leaves, and an unbranched, leafless stalk that bears a single short strobilus. Notes: Braggins (1974) provided a very full description of the species, and its habitat, distribution and dispersal in New Zealand.

Fig. 95: Phylloglossum drummondii. Sterile plant Fig. 96: Phylloglossum drummondii. Fertile plant with rosette of linear leaves. with a young emerging strobilus.

39 Fig. 97: Phylloglossum drummondii. Fertile plant Fig. 98: Phylloglossum drummondii. Fertile plant with a single strobilus borne terminally on a with a single strobilus borne terminally on a leafless stalk. leafless stalk.

40 References

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41 Field, A.R.; Renner, M.A.M. 2019: Rediscovered or reconsidered: the presumed extinct ferns and lycophytes of tropical Queensland, Australia. Australian Systematic Botany 32: 111–122. Field, A.R.; Testo, W.; Bostock, P.D.; Holtum, J.A.M.; Waycott, M. 2016: Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum. Molecular Phylogenetics and Evolution 94: 635–657. Forster, J.G.A. 1786: Florulae Insularum Australium Prodromus. Dietrich, Göttingen. Fuchs, H.P. 1955: Urostachys (Herter 1909, 5/29) Herter 1922, 249 nomen genericum conservandum? Verhandlungen der Naturforschenden Gesellschaft in Basel 66: 33–48. Greuter, W.; Troia, A. 2014: (2292) Proposal to conserve the name Palhinhaea against Lepidotis (Lycopodiaceae). Taxon 63: 680–682. Herter, W.G.F. 1909: Beiträge zur Kenntniss der Gattung Lycopodium. Studien über die Untergattung Urostachys. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 43: Beiblatt 98: 1–56. Herter, W.G.F. 1922: Itinera Herteriana III. Heteropteridophyta austroamericana. (Equisetales Lycopodiales Selaginellales Isoetales austroamericanae). Beihefte zum Botanischen Centralblatt, Abt. 2, 39: 248–256. Herter, W.G.F. 1949: Index Lycopodiorum. Herbarium Herter, Montevideo, Uruguay. Holloway, J.E. 1910: A comparative study of the anatomy of six New Zealand species of Lycopodium. Transactions and Proceedings of the New Zealand Institute 42: 356–370. Holloway, J.E. 1916: Studies in the New Zealand species of the genus Lycopodium: Part 1. Transactions and Proceedings of the New Zealand Institute 48: 253–303. Holloway, J.E. 1917: Studies in the New Zealand species of the genus Lycopodium: Part II – Methods of vegetative reproduction. Transactions and Proceedings of the New Zealand Institute 49: 80–93. Holloway, J.E. 1919: Studies in the New Zealand species of the genus Lycopodium: Part III – The plasticity of the species. Transactions and Proceedings of the New Zealand Institute 51: 161–216. Holloway, J.E. 1920: Studies in the New Zealand species of the genus Lycopodium: Part IV – The structure of the prothallus in five species. Transactions and Proceedings of the New Zealand Institute 52: 193–239. Holloway, J.E. 1935: The gametophyte of Phylloglossum drummondii. Annals of Botany 49: 513–520. Holub, J. 1964: Lycopodiella, novy rod radu Lycopodiales. Preslia 36: 16–22. Holub, J. 1975: Diphasiastrum, a new genus in Lycopodiaceae. Preslia 47: 97–110. Holub, J. 1983: Validation of generic names in Lycopodiaceae: with a description of a new genus Pseudolycopodiella. Folia Geobotanica et Phytotaxonomica 18: 439–442. Holub, J. 1985: Transfers of Lycopodium species to Huperzia: with a note on generic classification in Huperziaceae. Folia Geobotanica et Phytotaxonomica 20(1): 67–80. Holub, J. 1991: Some taxonomic changes within the Lycopodiales. Folia Geobotanica et Phytotaxonomica 26: 81–94. Hooker, J.D. 1854–1855: The Botany of the Antarctic Voyage of H.M. Discovery Ships Erebus and Terror, in the years 1839–1843, under the command of Captain Sir James Clark Ross. II. Flora Novae-Zelandiae. Part II. Flowerless plants. Lovell Reeve, London. Hooker, W.J. 1854: A century of ferns. Pamplin, London. Kirk, T. 1879: Description of a new species of Lycopodium. Transactions and Proceedings of the New Zealand Institute 11: 456–457. Kirk, T. 1884: On Lycopodium varium, R.Br., and L. billardieri, Spring, with description of a new form. Transactions and Proceedings of the New Zealand Institute 16: 376–377. Kunze, G. 1843: Phylloglossum, genus novum ex ordine novo. Botanische Zeitung (Berlin) 1: 721–723. Labillardière, J.J.H. de 1806–1807: Novae Hollandiae Plantarum Specimen. Vol. 2. Huzard, Paris. Lamarck, J.B.A.P.M de; Mirbel, C.F.B. de 1802: Histoire naturelle des végétaux, classés par familles. Vol. 3. Crapelet, Paris. Lamarck, J.B.A.P.M. de; Mirbel, C.F.B. de 1802: Histoire naturelle des végétaux, classés par familles. Vol. 4. Crapelet, Paris.

42 Large, M.F.; Braggins, J.E. 1991: atlas of New Zealand ferns and fern allies. SIR Publishing, Wellington. Lehmann, J.G.C. 1846–1848: Plantae Preissianae. Vol. 2. Meissner, Hamburg. Linnaeus, C. 1753: Species Plantarum. Impensis Laurentii Salvii, Stockholm. Mabberley, D.J.; Moore, D.T.: The Robert Brown Handbook: a guide to the life and work of Robert Brown (1773-1858), Scottish botanist. Regnum Vegetabile: (in press). Murray, B.G.; de Lange, P.J. 2013: Contributions to a chromosome atlas of the New Zealand flora – 40. Miscellaneous counts for 36 families. New Zealand Journal of Botany 51: 31–60. Nessel, H. 1934: Neue Lycopodien, die von allen schonbekannten Arten durch ihren Habitus ganz besonders abweichend und auffallenden sind. Repertorium Specierum Novarum Regni Vegetabilis 36: 177–193. Nessel, H. 1939: Die Bärlappgewächse. Fischer, Jena. Nessel, H. 1940: Beiträge zur Kenntniss der Lycopodiaceen. Revista Sudamericana de Botánica 6: 156–175. Nicolson, D.H.; Fosberg, F.R. 2003: The Forsters and the Botany of the Second Cook Expedition (1772–1775). Regnum Vegetabile 139: 1–760. Øllgaard, B. 1987: A revised classification of Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Øllgaard, B. 1989: Index of the Lycopodiaceae. Biologiske Skrifter 34: 1–135. Øllgaard, B. 1990: Lycopodiaceae. In: Kramer, K.U.; Green, P.S. (ed.) The families and genera of vascular plants. 1 Pteridophytes and . Springer-Verlag, Berlin. Øllgaard, B. 2012: New combinations in Neotropical Lycopodiaceae. Phytotaxa 57: 10–22. Palisot de Beauvois, A.M.F.J. 1805: Prodrome des cinquième et sixième familles de l'Aethéogamie. Les mousses. Les lycopodes. Fournier, Paris. Perrie, L.R.; Shepherd, L.D.; Field, A.R.; Brownsey, P.J. 2020: Morphological and genetic evidence for the separation of Phlegmariurus billardierei from P. varius (Lycopodiaceae). New Zealand Journal of Botany 58: 118–128. (Published online: 2 Oct 2019) Pichi Sermolli, R.E.G. 1968: Fragmenta pteridologiae I. Webbia 23: 159–722. Pichi Sermolli, R.E.G. 1971: Names and types of the genera of fern-allies: Lycopodiaceae, Selaginellaceae, Isoetaceae, Equisetaceae, Psilotaceae, Tmesipteridaceae. Webbia 26: 129–194. Poiret, J.L.M. 1814: Encyclopédie méthodique. Botanique. Suppl. 3. Panckoucke, Paris. PPG 1 2016: A community-derived classification for extant lycophytes. Journal of Systematics and Evolution 54(6): 563–603. Proctor, G.R. 1977: Pteridophyta. In: Howard, R.A. Flora of the Lesser Antilles. Arnold Arboretum, Harvard University, Jamaica Plain, Mass. Richard, A. 1832: Essai d'une Flore de la Nouvelle Zélande. In: Lesson, A.; Richard, A. Botanique. In: Dumont d'Urville, J. Voyage de Découvertes de l'Astrolabe. Tastu, Paris. [1]–376. Rothmaler, W. 1944: Pteridophyten-Studien, I. Feddes Repertorium Specierum Novarum Regni Vegetabilis 54: 55–82. Roux, J.P. 2009: Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1–296. Spring, A.F. 1841: Monographie de la famille des Lycopodiacées, première partie. Nouveaux Mémoires de l’Académie Royale des Sciences et Belles-Lettres de Bruxelles 15: 1–110. Spring, A.F. 1849: Monographie de la famille des Lycopodiacées. Mémoires de l’Académie Royale des Sciences, Lettres et Beaux-arts de Belgique 24: 1–358. Trevisan de Saint-Léon, V.B.A. 1874: Sylloge sporophytarum Italiae. I. Prothallogamae. Atti della Società Italiana di Scienze Naturali 17: 213–258. Tryon, A.F.; Lugardon, B. 1991: Spores of the Pteridophyta. Springer-Verlag, New York. Vasconcellos, J. de Carvalho e; Franco, J. do Amaral 1967: Breves notas sobre Licopodiáceas. Boletim da Sociedade Broteriana (Series 2) 41: 23–25. Webb, C.J.; Edgar, E. 1999: Spelling New Zealand in the specific and infraspecific epithets of vascular plants. New Zealand Journal of Botany 37(1): 71–77. Whittier, D.P.; Braggins, J.E. 1992: The young gametophyte of Phylloglossum (Lycopodiaceae). Annals of the Missouri Botanical Garden 79: 730–736.

43 Whittier, D.P.; Braggins, J.E. 2000: Observations on the mature gametophyte of Phylloglossum (Lycopodiaceae). American Journal of Botany 87: 920–924. Willdenow, C.L. von 1810: Species Plantarum. Vol. 5 (1). G.C. Nauk, Berlin. Wilson, H.D. 2013: Plant life on Banks Peninsula. Manuka Press, Cromwell. Zhang, L.; Iwatsuki, K. 2013: Lycopodiaceae. In: Lycopodiaceae through Polypodiaceae. Vol. 2–3. In: Wu, C.Y.; Raven, P.H.; Hong, D.Y. (ed.) Flora of . Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis.

44 Acknowledgements

We thank the staff at AK, CHR and WELT for loans of specimens, and for databasing and providing spreadsheets of collection data. We are grateful to staff at CHR for the preparation of maps and for assistance in editing and formatting the text, to Ashley Field and David Mabberley for collaboration in the typification of several species, to Germinal Rouhan at P for locating and imaging the type specimen of Lycopodium pachystachyum, and to Ashley Field for reviewing the manuscript.

45 Map 1: Map of New Zealand and offshore islands showing Ecological Provinces Map 2: Map of New Zealand showing Ecological Provinces Index Lycopodium laterale R.Br. 13 Lycopodium laterale var. diffusum (R.Br.) Hook.f. 11 Page numbers are in bold for the main entry, Lycopodium lessonianum A.Rich. 24 and italic for synonyms. Lycopodium magellanicum var. berggrenii Austrolycopodium Holub 17 Herter ex Nessel 21 Austrolycopodium fastigiatum (R.Br.) Holub 21 Lycopodium novozealandicum Colenso 34 Diphasium C.Presl ex Rothm. 17 Lycopodium pachystachyum Desv. ex Poiret 34 Diphasium decurrens (R.Br.) Holub 24 Lycopodium polycephalum Colenso 8 Diphasium scariosum (G.Forst.) Rothm. 24 Lycopodium ramulosum Kirk 11 Huperzia Bernh. 1, 2, 4, 29, 30, 38 Lycopodium sanguisorba Spring 38 Huperzia australiana (Herter) Holub 5 Lycopodium scariosum G.Forst. 1, 22, 24, 27 Huperzia billardierei (Spring) Rothm. 30 Lycopodium scariosum var. decurrens (R.Br.) Huperzia drummondii (Kunze) Christenh. & Hook. 24 H.Schneid. 38 Lycopodium scopulosum Colenso 21 Huperzia helmii (Nessel) Holub 34 Lycopodium serpentinum Kunze 16 Huperzia novozealandica (Colenso) Holub 34 Lycopodium varium R.Br. 34 Huperzia polaris (Herter ex Nessel) Holub 34 Lycopodium varium var. alpinum R.Br. 34 Huperzia varia (R.Br.) Trevis. 34 Lycopodium varium var. billardierei (Spring) Kirk Lateristachys Holub 7 30 Lateristachys diffusa (R.Br.) Holub 11 Lycopodium varium var. gracile Kirk 34 Lateristachys lateralis (R.Br.) Holub 13 Lycopodium varium var. polaris Kirk 34 Lateristachys ramulosa (Kirk) Holub 11 Lycopodium volubile G.Forst. 1, 22, 25, 27 Lepidotis P.Beauv. ex Mirbel 7 Palhinhaea Franco & Vasc. ex Vasc. & Franco 7 Lepidotis cernua (L.) P.Beauv. 8 Palhinhaea cernua (L.) Vasc. & Franco 8 Lepidotis densa Rothm. 19 Palhinhaea polycephala (Colenso) Holub 8 Lepidotis diffusa (R.Br.) Rothm. 11 Phlegmariurus Holub 1, 2, 4, 29, 32, 38 Lepidotis drummondii (Spring) Rothm. 16 Phlegmariurus billardierei (Spring) Brownsey & Lepidotis lateralis (R.Br.) Rothm. 13 Perrie 1, 30, 30, 35 Lepidotis volubilis (G.Forst.) Rothm. 27 Phlegmariurus varius (R.Br.) A.R.Field & Lycopodiaceae P.Beauv. 1, 2, 4, 5, 14, 18, 25, Bostock 30–32, 34 27, 29, 38, 39 Phylloglossum Kunze 1, 2, 4, 29, 30, 38 Lycopodiella Holub 1, 2, 7, 16, 18 Phylloglossum drummondii Kunze 38, 38 Lycopodiella cernua (L.) Pic.Serm. 1, 7, 8 Pseudodiphasium Holub 17 Lycopodiella diffusa (R.Br.) B.Øllg. 1, 7, 11, 14 Pseudodiphasium volubile (G.Forst.) Holub 27 Lycopodiella lateralis (R.Br.) B.Øllg. 1, 7, 12, 13 Pseudolycopodiella Holub 7 Lycopodiella ramulosa (Kirk) B.Øllg. 11 Pseudolycopodiella serpentina (Kunze) Holub Lycopodiella serpentina (Kunze) B.Øllg. 1, 7, 16 16 Lycopodium L. 1, 2, 4, 7, 8, 17, 30 Pseudolycopodium Holub 17 Lycopodium arcitenentis Herter 21 Pseudolycopodium densum (Rothm.) Holub 19 Lycopodium australianum Herter 5 Stachygynandrum scariosum (G.Forst.) Lycopodium berggrenii (Herter ex Nessel) P.Beauv. 24 Herter 21 Urostachys Herter 4 Lycopodium billardierei Spring 30 Urostachys australianus (Herter) Herter ex Lycopodium billardierei var. gracile (Kirk) Nessel 5 Cheeseman 34 Urostachys billardierei (Spring) Herter ex Lycopodium cernuum L. 8 Nessel 30 Lycopodium clavatum var. fastigiatum (R.Br.) Urostachys cockaynei Herter ex Nessel 5 Benth. 21 Urostachys helmii Nessel 34 Lycopodium cochinchense Herter ex Nessel 21 Urostachys polaris (Herter ex Nessel) Herter 34 Lycopodium consimile Colenso 13 Urostachys saturnii Herter 34 Lycopodium curvifolium Colenso 21 Urostachys varius (R.Br.) Herter ex Nessel 34 Lycopodium decurrens Colenso 21 Urostachys varius var. polaris Herter ex Nessel Lycopodium decurrens R.Br. 24 34 Lycopodium densum Labill. 19 Lycopodium deuterodensum Herter 1, 19, 22 Lycopodium diffusum R.Br. 11 Lycopodium distans Colenso 24 Lycopodium drummondii Spring 16 Lycopodium fastigiatum R.Br. 1, 21

48 Image Information

Image Creator Copyright Licence Front L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ cover Fig. 1 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 2 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 3 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 4 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 5 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 6 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 7 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 8 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 9 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 10 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 11 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 12 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 13 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 14 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 15 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 16 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 17 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 18 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 19 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 20 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 21 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 22 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 23 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 24 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 25 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 26 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 27 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 28 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 29 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 30 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 31 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 32 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 33 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 34 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 35 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 36 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 37 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 38 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 39 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 40 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 41 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 42 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 43 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 44 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 45 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 46 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 47 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 48 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 49 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 50 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 51 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 52 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 53 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 54 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 55 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 56 K.Boardman © Landcare Research 2019 CC-BY 4.0

49 Fig. 57 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 58 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 59 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 60 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 61 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 62 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 63 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 64 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 65 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 66 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 67 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 68 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 69 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 70 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 71 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 72 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 73 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 74 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 75 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 76 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 77 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 78 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 79 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 80 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 81 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 82 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 83 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 84 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 85 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 86 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 87 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 88 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 89 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 90 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 91 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 92 L.R. Perrie © Te Papa CC-BY-NC 4.0 NZ Fig. 93 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 94 K.Boardman © Landcare Research 2019 CC-BY 4.0 Fig. 95 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 96 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 97 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Fig. 98 L.R. Perrie © Leon Perrie CC-BY-NC 4.0 NZ Map 1 A.D. Wilton © Landcare Research 2014 CC-BY 3.0 NZ Map 2 A.D. Wilton © Landcare Research 2014 CC-BY 3.0 NZ

50 Flora of New Zealand: PDF publications

The electronic Flora of New Zealand (eFloraNZ) project provides dynamic, continually updated, online taxonomic information about the New Zealand flora. Collaborators in the project are Manaaki Whenua – Landcare Research, the Museum of New Zealand Te Papa Tongarewa, and the National Institute of Water and Atmospheric Research (NIWA). The eFloraNZ presents new systematic research and brings together information from the Manaaki Whenua – Landcare Research network of databases and online resources. New taxonomic treatments are published as fascicles in PDF format and provide the basis for other eFloraNZ products, including the web profiles. eFloraNZ will have separate sets of PDF publications for algae, lichens, liverworts and , , ferns and lycophytes, and plants. For each eFloraNZ set the PDF files are made available as dated and numbered fascicles. With the advent of new discoveries and research the fascicles may be revised, with the new fascicle being treated as a separate version under the same number. However, superseded accounts will remain available on the eFlora website.

Fern and Set (ISBN 978-0-478-34761-6) The Fern and Lycophyte Set includes ferns and lycophytes indigenous to New Zealand, together with exotic species that have established in the wild. Species that are found only in cultivation are excluded. Editor-in-Chief: Aaron Wilton Series Editors: Ilse Breitwieser, Aaron Wilton Steering committee: Ilse Breitwieser, Pat Brownsey, Wendy Nelson, Rob Smissen, Aaron Wilton Technical production: Aaron Wilton with Kate Boardman, Bavo de Pauw, Sue Gibb, Ines Schönberger, Katarina Tawiri, Margaret Watts Copy Editor: Ray Prebble

51 ISBN 978-0-947525-66-8

9 780947 525668