FLORA OF NEW ZEALAND

CLIMACIACEAE

A.J. FIFE

Fascicle 13 – SEPTEMBER 2014 © Landcare Research New Zealand Limited 2014. This copyright work is licensed under the Creative Commons Attribution 3.0 New Zealand license Attribution if redistributing to the public without adaptation: “Source: Landcare Research" Attribution if making an adaptation or derivative work: “Sourced from Landcare Research"

CATALOGUING IN PUBLICATION Fife, Allan J. (Allan James), 1951- Flora of New Zealand [electronic resource] : mosses. Fascicle 13, / Allan J. Fife. -- Lincoln, N.Z. : Manaaki Whenua Press, 2014. 1 online resource ISBN 978-0-478-34769-2 (pdf) ISBN 978-0-478-34747-0 (set) 1.Mosses -- New Zealand -- Identification. I. Title. II. Manaaki Whenua-Landcare Research New Zealand Ltd.

DOI: 10.7931/J21Z4293

This work should be cited as: Fife, A.J. 2014: Climaciaceae. In: Heenan, P.B.; Breitwieser, I.; Wilton, A.D. Flora of New Zealand - Mosses. Fascicle 13. Manaaki Whenua Press, Lincoln. http://dx.doi.org/10.7931/J21Z4293

Cover image: dendroides, habit. Drawn by Rebecca Wagstaff from A.J. Fife 6797, CHR 405507. Contents

Introduction...... 1 Taxa Climaciaceae ...... 2 Climacium F.Weber & D.Mohr ...... 2 (Hedw.) F.Weber & D.Mohr ...... 3 References ...... 5 Conventions ...... 6 Acknowledgements ...... 8 Plates ...... 9 Maps ...... 10 Index ...... 12 Image Information ...... 13

Introduction

The Climaciaceae are a family of two genera, with the better known Climacium widespread in the northern hemisphere and including three species. The lesser known Pleuroziopsis is a monotypic genus confined to the northern Pacific Rim. Only one species of Climacium occurs in New Zealand. When well developed, Climacium dendroides looks, as befitting its epithet, like a small tree. While it is widespread in the northern hemisphere, until recently C. dendroides was known in N.Z. only from the South Island. It has, however, now been collected from a high elevation site on Mt Ruapehu in the central North Island. The collection history of this species is remarkably similar to other allegedly “bipolar” species in N.Z., with the earliest confirmed collection made in 1891 at Castle Hill, Canterbury, by T. Kirk. There appear to be two other South I. collections made in the 1890s, followed by an inexplicable collection gap of roughly 40 years’ duration. Subsequently, collections have been made since 1950 from nearly all parts of the South I. The absence of gatherings by numerous capable South I. collectors in the late 19th and early 20th centuries, and the post-1950 increase in the collection frequency of this species, strongly suggest that C. dendroides is an adventive that expanded its N.Z. range during the first half of the 20th century. While on the West Coast of the South I. C. dendroides can grow abundantly in damp roadside ditches and dry roadside gravels at low elevations, it is best developed and most often collected from seemingly unmodified wetland native vegetation at mid to upper elevations. The lack of congruity between its collection history and its occurrence in intact native vegetation, sometimes at remote localities, is perplexing and worthy of further investigation.

1 Climaciaceae : A family of two genera, with Climacium the more widespread and better known. Crum & Anderson (1981, p. 1227) defined the family by having dendroid branching of secondary stems arising from a subterranean “rhizome”, an abundance of filiform paraphyllia, ± ovate leaves with a single strong costa ending below the leaf apex, dioicous sexuality, and erect capsules with essentially hypnoid peristomes. Goffinet et al. (2009) place the Climaciaceae in the . Pleuroziopsis, a monotypic genus confined to the northern Pacific Rim, was segregated from the Climaciaceae by Ireland (1968); Goffinet et al. (2009) did not accept the segregate Pleuroziopsidaceae, and circumscribed the Climaciaceae in the same manner as did Brotherus (1925). No separate family description is provided here. Pleuroziopsis differs from Climacium mainly by having curved, more or less horizontal capsules and by peristome differences.

Climacium F.Weber & D.Mohr, Naturh. Reise Schwedens, 96 (1804) Type taxon: Climacium dendroides (Hedw.) F.Weber & D.Mohr Elements in the following description are translated from Brotherus (1925). robust, gregarious, dendroid, bright green to bronze, weakly glossy. Primary stems subterranean, rhizomatous, beset with scale-like and erect leaves and red-brown, smooth rhizoids. Secondary stems erect, with a well-developed stipe and frond, rarely unbranched or ± pinnately branched, with apex concealed or shortly protruding beyond apical branches, densely covered below with red-brown rhizoids, in cross-section ± angled, with several layers of thick-walled cortical cells and a central strand. Branches mostly simple, rarely ± pinnate, densely leaved, often unequal in length, tapered apically. Stipe leaves erect-appressed when moist, broadly ovate and cordate, sharply or bluntly mucronate, obscuring the stipe. Branch leaves erect-spreading when moist, little altered when dry, lanceolate-lingulate, acute, from an auriculate and ± decurrent base, weakly or markedly sulcate, serrate above; upper laminal cells smooth, narrowly rhomboid, moderately thick-walled, those below more linear and porose and orange near leaf base; alar cells lax, hyaline and thin-walled, forming a large and well-defined group. Costa single and strong, ending below the leaf apex, sometimes with few to several abaxial teeth near apex. Paraphyllia abundant, filiform or much-branched. Dioicous. Perichaetia numerous, on secondary stems and at branch bases, elongate and slender, with leaves erect and sheathing, the innermost abruptly narrowed to a narrow apex, entire, and shortly costate. Perigonia on secondary stems and branch bases, bud-like, with numerous antheridia and elongate brown paraphyses. Setae single or aggregated, elongate, purple, twisted to the right above; capsules erect and symmetric, ± cylindric, smooth, red-brown; stomata numerous, small, confined to the neck; annulus lacking; operculum rostrate from a conic base, falling with the columella attached. Peristome double; exostome teeth fused at base, linear-lanceolate, dark red-brown, bordered, closely articulate, coarsely papillose on the outer surface, the inner surface with numerous and conspicuous lamellae; endostome yellow or orange, arising from a very low membrane, segments linear, broadly perforate along the median line, and apically forked. Calyptra narrowly cucullate, split to the apex, smooth, enclosing the entire capsule. 15–28 µm, rust-coloured, finely granular. Notes: A genus of three species distributed in temperate northern hemisphere and adventive in Australasia. The species of Climacium grow primarily in swampy habitats. Etymology: According to Meagher (2011, quoting H.N. Dixon) the name Climacium derives from the Greek klimax (staircase, ladder), “alluding to the appearance of the processes of the inner peristome, the two halves of which are regularly united by projections between the perforations, giving somewhat the appearance of a ladder”.

2 Climacium dendroides (Hedw.) F.Weber & D.Mohr, Naturh. Reise Schwedens, 96 (1804) ≡ Leskea dendroides Hedw., Sp. Musc. Frond., 228 (1801) Type: Europe. Not seen. = Climacium novae-seelandiae Müll.Hal. ex Beckett, Trans. & Proc. New Zealand Inst. 25: 292 (1893) Holotype: N.Z., Canterbury, Castle Hill, Jan. 1891, T. Kirk, CHR 510623!

Plants ± glossy when dry, bright green, yellow-green, or rarely bronze. Secondary stems erect and dendroid with a well-differentiated stipe and frond, simple or rarely sparingly branched, mostly 30–70(–100) mm high, red-brown below, densely beset with smooth, much-branched, red-brown rhizoids. Stipe leaves erect-appressed when moist or dry, obscuring the stipe, broadly ovate and cordate, broadly acute or obtuse at apex, sharply mucronate, serrulate near apex, entire below, 2.3–2.8 × c. 2.0 mm. Frond to c. 45 mm diam. Branches variable in length, up to c. 22 mm, mostly unbranched. Branch leaves ovate-lanceolate, c. 1.6–2.2 × 0.8–1.0 mm, coarsely or sometimes weakly serrate in upper half, serrulate or entire below, weakly sulcate, auriculate and scarcely decurrent; upper laminal cells mostly 30–45 µm long, weakly porose; alar cells as per genus. Costa strong, failing c. 5–8 cells below apex, often ending in a single abaxial tooth. Perichaetial leaves squarrose. Perigonia and capsules not seen in N.Z. material. Illustrations: Plate 1. Brotherus 1925, fig. 479; Sainsbury 1955, pl. 49, fig. 1; Crum & Anderson 1981, fig. 611; Smith 2004, fig. 234, 1–4. Distribution: NI: Wellington (Upper Makatote River); SI: Nelson, Marlborough (Mt Tapuaenuku), Canterbury, Westland, Otago, Southland (S Arm of Lake Manapōuri, Lake Gunn). Adventive. Tasmania*, Australia (Victoria*), North America*, Europe*, E Asia*. A literature report from Tasmania was cited by Martin (1946), but this record was doubted by Dalton et al. (1991). However, L. Cave (pers. comm., Sep. 2004) has rediscovered the species in Tasmania. Habitat: On humic soil at margins of lakes, tarns, and streams, in seepage slopes, damp depressions in tussock grasslands (especially Chionochloa rubra or C. australis grasslands), and cushion bogs. In cushion bogs it is sometimes dominant atop hummocks. Occasionally in swampy areas or adjacent to watercourses in subalpine Nothofagus forest. This species can also occur abundantly in roadside ditches (as at the Taramakau Valley, Westland L.D.) and occasionally at low elevations on dry roadside gravels (as at Waimangaroa, Westland L.D.). On such dry gravels it can assume confusing irregularly branched and prostrate forms. Bryophyte associates include Aulacomnium palustre, Breutelia pendula, Campyliadelphus polygamum, Sanionia uncinatus, Sphagnum spp., and Syntrichia rubra. The single North I. collection (one stem only) was associated with Hylocomium splendens. On North I. known only from 1900 m elevation. On South I. occurring mostly at c. 470–1600 m, but extending to lower elevations (c. 30 m) in Westland and Southland and upwards to c. 2000 m in the ranges of western Otago (e.g., Remarkable Range). First record: The earliest confirmed collection was made in 1891 from Castle Hill, Canterbury L.D., by T. Kirk (CHR 510623, type of C. novae-seelandiae Beckett). An unknown collector gathered it in the Rock and Pillar Range in Otago L.D. in 1892 (CHR 623649) and Kirk made an undated but early collection at Lake Rotoiti in Nelson L.D. (CHR 623650). Subsequently, there is an inexplicable collection gap of roughly 40 years until 1931. Since 1950 the species has been collected in nearly all parts of South I. and there are c. 55 gatherings of this species in CHR. The absence of gatherings by numerous capable South I. collectors in the late 19th and early 20th centuries and the apparent rapid increase in the collection frequency of this species since 1950 strongly suggest that C. dendroides is adventive in N.Z. Given its alleged adventive status, its occurrence in seemingly unmodified native vegetation, sometimes in remote localities (such as at the cirque at the head of the Rahu River in the Victoria Range, Nelson L.D.), is difficult to explain. However, the paucity of pre-1950 collections of such a distinctive and conspicuous species leads to the conclusion that it is a 19th century introduction to the N.Z. flora. The habitat, general distribution, and collection history of C. dendroides exhibit similarities to those of Aulacomnium palustre. Notes: Although extremely variable in stature, the microscopic features of C. dendroides are very constant. Perichaetia are uncommon (seen only in lowland material from Westland L.D.). Shorter and more compact plants are associated with drier habitats. Horton & Vitt (1976) included N.Z. material in a detailed morphometric study of Climacium dendroides and C. americanum from North America, and concluded that N.Z. material falls within the range of variability of the former species. Dixon (1927, p. 241) reached an identical conclusion.

3 Recognition: Because of its frondose habit, C. dendroides could be confused with members of the genus Sciadocladus, particularly S. kerrii. The overall habit of these two plants is different, however, in part because the stipe leaves in C. dendroides are broadly acute to obtuse, mucronate, and erect- appressed, obscuring the stipe in both the dry and moist condition. The stipe leaves of all Sciadocladus spp. are acuminate, widely spreading when moist, and do not obscure the stipe. The branch leaves in C. dendroides are broader apically and have shorter upper laminal cells (c. 30–45 µm), while the branch leaves of S. kerrii are more acute and have longer (mostly 45–95 µm) upper laminal cells. The costae of C. dendroides lack multiple abaxial spines near their apex in contrast to species of Sciadocladus. Climacium dendroides is not known to fruit in N.Z., while species of Sciadocladus frequently fruit. Confusion between the terrestrial C. dendroides and the epiphytic Braithwaitea sulcata seems less likely; the latter differs in many ways including the pinnately branched and flattened nature of its fronds and the presence of filiform axillary gemmae. Confusion with other species in the N.Z. flora is not likely. Etymology: The epithet dendroides refers to the tree-like habit.

4 References

Beckett, T.W.N. 1893 ("1892"): Description of new species of Musci. Transactions and Proceedings of the New Zealand Institute 25: 289–297. Brotherus, V.F. 1925: Musci (Laubmoose). In: Engler, A. (ed.) Die natürlichen Pflanzenfamilien. Edition 2. Bd 11. Engelmann, Leipzig. 1–542. Crum, H.A.; Anderson, L.E. 1981: Mosses of Eastern North America. Columbia University Press, New York. Dalton, P.J.; Seppelt, R.D.; Buchanan, A.M. 1991: An annotated checklist of Tasmanian mosses. In: Banks, M.R.; Curtis, W.M. (ed.) Aspects of Tasmanian Botany - a Tribute to Winifred Curtis. Royal Society of Tasmania, Hobart. 15–32. Dixon, H.N. 1927: Studies in the bryology of New Zealand, with special reference to the herbarium of Robert Brown. Part V. Bulletin, New Zealand Institute 3(5): 239–298. Goffinet, B.; Buck, W.R.; Shaw, A.J. 2009: Morphology, anatomy, and classification of the Bryophyta. In: Goffinet, B.; Shaw, A.J. (ed.) Bryophyte Biology. Edition 2. Cambridge University Press, Cambridge. 55–138. Hedwig, J. 1801: Species Muscorum Frondosorum descriptae et tabulis aeneis lxxvii coloratis illustratae. Barth, Leipzig. Horton, D.G.; Vitt, D.H. 1976: Morphological characters, relative to distribution, and taxonomic considerations of the genus Climacium in North America. Canadian Journal of Botany 54: 1872–1883. Ireland, R.R. 1968: Pleuroziopsidaceae, a new family of mosses. Journal of the Hattori Botanical Laboratory 31: 59–64. Martin, W. 1946: Geographic range and internal distribution of the mosses indigenous to New Zealand. Transactions and Proceedings of the Royal Society of New Zealand 76: 162–184. Meagher, D. 2011: An etymology of Australian bryophyte genera. 2 – Mosses. Muelleria 29: 33–61. Sainsbury, G.O.K. 1955: A handbook of the New Zealand mosses. Bulletin of the Royal Society of New Zealand 5: 1–490. Smith, A.J.E. 2004: The Flora of Britain and Ireland. Edition 2. Cambridge University Press, Cambridge. Weber, F. 1804: Naturhistorisches Reise durch einen Theil Schwedens. Heinrich Dieterich, Göttingen.

5 Conventions

Abbreviations and Latin terms

Abbreviations Meaning A Auckland Islands A.C.T. Australian Capital Territory aff. allied to (affinis) agg. aggregate Ant Antipodes Islands a.s.l. above sea level auct. of authors (auctorum) B Bounty Islands C Campbell Island c. about (circa) cf. compare with, possibly the species named (confer) c.fr. with fruit (cum fructibus) Ch Chatham Islands comb. nov. new combination (combinatio nova) D’U D’Urville Island et al. and others (et alia) et seq. and following pages (et sequentia) ex from fasc. fascicle fide according to GB Great Barrier Island HC Hen and Chicken Islands Herb. Herbarium hom. illeg. illegitimate homonym I. Island ibid. in the same place (ibidem) incl. including in herb. in herbarium (in herbario) in litt. in a letter (in litteris) inter alia among other things (inter alia) Is Islands K Kermadec Islands KA Kapiti Island LB Little Barrier Island L.D. Land District or Districts leg. collected by (legit) loc. cit. in the same place (loco citato) l:w length:width ratio M Macquarie Island Mt Mount nec nor NI North Island no. number nom. cons. conserved name (nomen conservandum) nom. dub. name of doubtful application (nomen dubium) nom. illeg. name contrary to the rules of nomenclature (nomen illegitimum) nom. inval. invalid name (nomen invalidum) nom. nud. name published without a description (nomen nudum) non not N.P. National Park N.S.W. New South Wales N.T. Northern Territory (Australia) N.Z. New Zealand op. cit. in the work cited (opere citato) pers. comm. personal communication

6 PK Poor Knights Islands P.N.G. Papua New Guinea pro parte in part Qld Queensland q.v. which see (quod vide) RT Rangitoto Island S.A. South Australia s.coll. without collector (sine collectore) s.d. without date (sine die) sect. section SEM scanning electron microscope/microsopy sensu in the taxonomic sense of SI South Island sic as written s.l. in a broad taxonomic sense (sensu lato) s.loc. without location (sine locus) Sn Snares Islands s.n. without a collection number (sine numero) Sol Solander Island sp. species (singular) spp. species (plural) s.s. in a narrow taxonomic sense (sensu stricto) St Stewart Island stat. nov. new status (status novus) subg. subgenus subsect. subsection subsp. subspecies (singular) subspp. subspecies (plural) Tas. Tasmania TK Three Kings Islands U.S.A. United States of America var. variety vars varieties Vic. Victoria viz. that is to say (videlicet) vs versus W.A. Western Australia

Symbols

Symbol Meaning µm micrometre ♂ male ♀ female ± more or less, somewhat × times > greater than < less than ≥ greater than or equal to ≤ less than or equal to = heterotypic synonym of the preceding name ≡ homotypic synonym of the preceding name ! confirmed by the author * in distribution statements, indicates non-N.Z. localities from which material has been confirmed by the author

Technical terms conform to Malcolm, B.; Malcolm, N. 2006: Mosses and other Bryophytes: an Illustrated Glossary. Edition 2. Micro-Optics Press, Nelson. Abbreviations for Herbaria follow the standard abbreviations listed in Index Herbariorum.

7 Acknowledgements

Jessica Beever provided advice during the preparation of this treatment. Rod Seppelt read the manuscript and suggested many worthwhile improvements. Peter de Lange invited me to study his recent collections from Mt Ruapehu. Rebecca Wagstaff prepared the line drawings with patience and skill. Peter Heenan and Ilse Breitwieser encouraged me to submit this manuscript to the eFlora of New Zealand series. Sue Gibb, Aaron Wilton, and Katarina Tawiri converted the manuscript into a format suitable for electronic publication and Christine Bezar provided skilled editing. I thank the participants, over many years, of the John Child Bryological and Lichenological Workshops and the curators of AK and WELT for allowing me to study their specimen holdings. The preparation of this revision was supported by Core funding for Crown Research Institutes from the Ministry of Business, Innovation and Employment’s Science and Innovation Group. A.J. Fife Landcare Research, PO Box 69040, Lincoln 7640, New Zealand [email protected]

8 Plate 1: Climacium. A–G: C. dendroides. A, habit. B, paraphyllia. C, branch detail. D, branch leaves. E, branch leaf apex. F, stipe leaf. G, alar cells of branch leaf. Drawn from A.J. Fife 6797, CHR 405507. Map 1: Map of New Zealand and offshore islands showing Land District boundaries Map 2: Map of main islands of New Zealand showing Land District boundaries Index

Page numbers are in bold for the main entry, and italic for synonyms. Climaciaceae 1, 2 Climacium F.Weber & D.Mohr 1, 2 Climacium dendroides (Hedw.) F.Weber & D.Mohr 1, 3 Climacium novae-seelandiae Müll.Hal. ex Beckett 3 Leskea dendroides Hedw. 3

12 Image Information

Image Creator Copyright Plate 1 R.C. Wagstaff © Landcare Research 2014 Map 1 A.D. Wilton © Landcare Research 2014 Map 2 A.D. Wilton © Landcare Research 2014

13 14 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 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 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 hornworts, mosses, ferns and lycophytes, and seed 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.

Moss Set (ISBN 978-0-478-34747-0) The Moss Set covers indigenous and exotic mosses within the New Zealand Botanical Region. Authors Allan Fife and Jessica Beever intend to publish Flora of New Zealand Mosses as a book. However, they decided to make completed family treatments available through the eFloraNZ project in advance of being published in hardcopy, to enable immediate use. Editor-in-Chief: Ilse Breitwieser Series Editors: Peter Heenan (Principal), Ilse Breitwieser, Aaron Wilton Steering Committee: Ilse Breitwieser, Pat Brownsey, Peter Heenan, Wendy Nelson, Aaron Wilton Technical production: Aaron Wilton with Kate Boardman, Bavo de Pauw, Sue Gibb, Ines Schönberger, Katarina Tawiri, Margaret Watts Copy Editor: Christine Bezar

15 ISBN 978-0-478-34747-0

9 780478 347470