FLORA OF NEW ZEALAND

HEDWIGIACEAE

A.J. FIFE

Fascicle 14 – 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 14, / Allan J. Fife. -- Lincoln, N.Z. : Manaaki Whenua Press, 2014. 1 online resource ISBN 978-0-478-34771-5 (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/J2X63JTQ

This work should be cited as: Fife, A.J. 2014: Hedwigiaceae. In: Heenan, P.B.; Breitwieser, I.; Wilton, A.D. Flora of New Zealand - Mosses. Fascicle 14. Manaaki Whenua Press, Lincoln. http://dx.doi.org/10.7931/J2X63JTQ

Cover image: ciliata, portion of shoot with perichaetia and capsules, dry. Drawn by Rebecca Wagstaff from J. Child 5960, CHR 430718. Contents

Introduction...... 1 Taxa Hedwigiaceae ...... 2 Hedwigia P.Beauv...... 3 (Hedw.) P.Beauv...... 3 Hedwigidium Bruch & Schimp...... 4 Hedwigidium integrifolium (P.Beauv.) C.E.O.Jensen ...... 5 References ...... 7 Conventions ...... 9 Acknowledgements ...... 11 Plates ...... 12 Maps ...... 13 Index ...... 15 Image Information ...... 16

Introduction

The Hedwigiaceae are a family of 4 genera of rock-inhabiting pleurocarpous mosses. They are coarse, rigid, and monoicous with curved-ascendant stems and branches. The leaves lack costae and have short, thick-walled, and papillose cells. Their frequently produced capsules lack a peristome. The generic names Hedwigia and Hedwigidium both commemorate the 18th century German botanist Johannes Hedwig, often referred to as the “Father of Bryology”. Both Hedwigia and Hedwigidium are small genera of wide and predominantly temperate distribution; each is represented in N.Z. by a single species. They occur on rock in the drier and more easterly portions of the main islands. The largest genus in the family, Braunia, has a tropical distribution and does not occur regionally. While the relationships and circumscription of the family remain controversial, recent morphological studies have shown that all members have unusual globular, rather than filamentous, protonemata.

1 Hedwigiaceae Plants medium-sized to robust, creeping, yellow- or grey-green, usually on rock or tree bases. Protonema globular. Stems creeping or weakly ascendant, freely and irregularly branched, often producing slender microphyllous stolons, in cross-section lacking a central strand. Shoots julaceous or hoary when dry. Leaves crowded, appressed in lower portions (but often with spreading hair-points) when dry, spreading or secund when moist, mostly oblong-ovate, acute, acuminate, or piliferous, often hoary and plicate, ecostate, mostly recurved, entire or serrulate, often decurrent. Upper laminal cells quadrate, oblong, or rhombic, papillose on both surfaces, mostly incrassate and porose, becoming elongate towards leaf base; basal marginal cells differentiated, mostly quadrate in many rows. Paraphyllia absent. Pseudoparaphyllia absent or filamentous, reportedly papillose. Monoicous. Setae mostly short, occasionally elongate; capsules immersed (in N.Z. taxa) or exserted, erect, symmetric, mostly subglobose, gymnostomous; stomata superficial; annulus lacking; operculum plano-convex, umbonate, or long-rostrate. Calyptra mitrate or cucullate. Spores medium- sized. : The relationships and circumscription of the Hedwigiaceae have been, and remain, controversial. The Hedwigiaceae were placed in the order Isobryales by Brotherus (1925). He subdivided the family into 3 subfamilies with the Hedwigioideae the largest, with 4 genera. De Luna (1995) re-evaluated the limits of the family and its relationships; he considered the 4 genera (Hedwigia, Hedwigidium, Braunia, and Pseudobraunia) placed by Brotherus in the Hedwigioideae to comprise the monophyletic family Hedwigiaceae, which he considered supported by 2 unique synapomorphies: papillose pseudoparaphyllia and globular protonema. De Luna’s circumscription of the family agrees with that subsequently adopted by Goffinet et al. (2009), who placed the family in the order . Hedwigia and Hedwigidium are small genera with representatives in New Zealand. Braunia is a tropical genus of c. 23 species occurring on rock in the Americas, Africa, and Asia. Pseudobraunia is a monotypic genus confined to western North America. A fifth genus, Bryowijkia (=Cleistostoma), of eastern Asia, is also sometimes placed in this family. The distinctive globular protonemata, mainly composed of rounded cells, were illustrated by de Luna (1990), using cultured material. While the genus Rhacocarpus was placed in the Hedwigiaceae by Brotherus (1925), there is an increasing consensus that it deserves recognition at the family level, in large part because of its distinctive laminal cell ornamentation (see Barthlott & Schultze-Motel 1981). According to Crum (1994b, p. 667) Rhacocarpus resembles the Hedwigiaceae “only in lacking a costa and a peristome”. De Luna (1990) demonstrated that the protonema of Rhacocarpus was not globular. The removal of the Rhacocarpaceae from the Hedwigiaceae was advocated by de Luna (1995) and followed by Goffinet et al. (2009).

1 Shoots not julaceous when dry; leaves mostly ending in broad white tips, with margins plane throughout or weakly reflexed near base; upper laminal cells with 1–3 coarse, simple, forked, stellate, or peltate papillae on both surfaces; cells of leaf apices clearly differentiated from upper laminal cells, elongate-fusiform and with numerous unforked papillae; perichaetial leaves ciliate ...... Hedwigia 1' Shoots julaceous when dry; leaves lacking white or hyaline tips, with margins strongly reflexed nearly throughout; upper laminal cells with numerous simple and usually obscure papillae; cells of leaf apices not differentiated from upper laminal cells; perichaetial leaves entire ...... Hedwigidium

2 Hedwigia P.Beauv., Mag. Encycl. 9e Année, 5: 304 (1804) Type taxon: Hedwigia ciliata (Hedw.) P.Beauv. Plants medium-sized, hoary, grey-green above, often dark brown or nearly black in lower portions, lacking stolons; forming loose mats on dry rock. Protonema globular, irregular in shape, mainly composed of rounded cells (fide de Luna 1990). Stems curving away from the substrate, branching irregularly, in cross-section with incrassate outer cells grading into parenchymatous medullary cells, lacking a central-strand. Rhizoids sparse, scattered on stems, smooth, brown. Shoots appearing hoary due to widely spreading white hair-points when dry, not julaceous. Leaves appressed in their lower portions, erect, erect-spreading, or widely spreading when dry, spreading when moist, oblong- ovate or broadly ovate, acuminate or acute at apices, with short or elongate and white hair-points, weakly decurrent, concave and appearing smooth abaxially under a hand-lens, entire (except the hair- point), plane at margins or reflexed below; upper laminal cells irregularly subquadrate to rectangular, with very incrassate and sinuose walls, the adaxial surface with 1 to several conspicuous and simple, branched, stellate or peltate papillae, the abaxial surface less papillose, the laminal cells becoming longer and with more papillae in lower third of leaf; cells of the hair-point strongly differentiated or rarely not present, linear-fusiform, colourless, with thick and undulate walls and numerous non-stellate papillae, the terminal cell less papillose to ± smooth and either acute or truncate/papillose apically; central cells of leaf base more or less linear, non-papillose, porose, forming a large and conspicuous gold-brown group; marginal cells at base short and ± quadrate in several rows; costa absent. Autoicous. Perichaetia terminal on short branches, the inner leaves oblong-elliptic, ciliate or entire at margins. Perigonia gemmiform, scattered on stems, with bracts ovate, strongly pigmented, concave, and entire, surrounding numerous antheridia and filiform paraphyses. Setae very short and straight; capsules immersed, erect and symmetric, hemispheric when young, becoming cupulate after dehiscence, lacking a differentiated neck, smooth, gymnostomous, narrowed slightly at the transverse mouth; exothecial cells mostly oblong, rather thin-walled, not in alternating bands, with several rows isodiametric, thicker-walled, and more pigmented at mouth; annulus lacking; operculum umbonate. Calyptra mitrate. Spores reniform, coarsely lirate-murate, often with a faint proximal trilete scar. Taxonomy: Hedwigia has traditionally (Brotherus 1925) been treated to include a single species of nearly cosmopolitan distribution. Some authors (e.g., Sainsbury 1955; Smith 2004) have also included Hedwigidium integrifolium in Hedwigia, but this has received limited acceptance. Hedwigidium is maintained as a separate genus here. Recent northern hemisphere studies (Hedenäs 1994; Buck & Norris 1996) have recognised additional taxa in Hedwigia and advocated an emended and narrower concept of H. ciliata. Buck & Norris (1996) recognised three species, including the recently described H. stellata Hedenäs, from the west coast of North America. In their discussion they stated that an additional species, which they tentatively named H. nivalis (Müll.Hal.) Mitt., occurs from Arizona south to Argentina. They also speculated that this species might occur in Africa and Australasia. Etymology: The generic name honours Johannes Hedwig (1730–1799), a Leipzig physician and botanist whose posthumously published Species Muscorum Frondosorum (1801) provides the nomenclatural starting date for mosses.

Hedwigia ciliata (Hedw.) P.Beauv., Prodr. Aethéogam, 15 (1805) ≡ Anictangium ciliatum Hedw., Sp. Musc. Frond., 40 (1801) Type: Europe. Not seen. = Pilotrichum microcyatheum Müll.Hal., Bot. Zeitung (Berlin) 9: 564 (1851) ≡ Hedwigia microcyathea (Müll.Hal.) Paris, Index Bryol., 554 (1896) Syntypes: Tasmania. Not seen. = Hedwigia albicans Lindb., Öfvers. Kongl. Vetensk.-Akad. Förh. 21: 421 (1864) nom. illeg.

Plants, protonema, stems, and rhizoids as per genus. Leaves erect-spreading, widely spreading, or weakly secund when dry, spreading when moist, oblong-ovate, 2.0–2.2(–2.4) × 0.8–1.2 mm, otherwise as per genus; upper laminal cells irregularly subquadrate to short-rectangular, with very incrassate and sinuose walls, mostly (10–)12–15 × c. 6 µm, mostly with 1–2 conspicuous, peltate or stellate (or rarely simple) papillae on each surface (more prominent on adaxial surface), becoming longer (to at least 36 µm) and with more papillae in lower third of leaf; papillae of adaxial surface often appearing anastomosed under the light microscope, strongly branched and with the branches often unequal; cells of hair-point strongly differentiated, linear-fusiform, appearing white, with numerous non-stellate papillae and thick and undulate walls that obscure the lumina, the terminal cell less papillose than

3 adjacent cells or smooth, either acute or truncate/pluripapillose at tip; central cells of leaf base gold- brown, more or less linear, to 70 µm or greater, non-papillose, strongly porose, forming a large and conspicuous group; marginal cells at base much shorter than interior cells, subquadrate in c. 15–20 rows and gradually merging with the upper laminal cells; alar cells not differentiated. Autoicous. Perichaetia terminal on short branches, the inner leaves c. 3.0 mm, with numerous long, hyaline, and contorted marginal cilia. Perigonia 0.8–1.4 mm, scattered and usually numerous on stems, with entire leaves. Setae c. 0.4–1.5 mm; capsules immersed, c. 1 mm × 0.8 mm; exothecial cells as per genus; stomata not seen; annulus lacking; operculum as per genus. Calyptra mitrate, c. 1 mm, falling early. Spores reniform, mostly 27–30 µm, coarsely lirate-murate, often with a proximal trilete scar. Illustrations: Plate 1. Scott & Stone 1976, pl. 67; Crum & Anderson 1981, fig. 354; Hedenäs 1994, fig. 5 (as H. ciliata var. ciliata); Meagher & Fuhrer 2003, p. 61; Seppelt et al. 2013, pl. 9. Distribution: NI: N Auckland (Auckland City), S Auckland (Rotorua vicinity, Ātiamuri, Taupō vicinity), Wellington (Blue Rock Stream); SI: Nelson, Marlborough, Canterbury, Otago, Southland. Nearly cosmopolitan. Habitat: On either shaded or insolated dry rock (including greywacke, basalt, schist, and scoria); rarely on bark or “woody debris”. Forming loose mats to at least 30 cm diam. Restricted to dry sites and not known from wetter portions of either main island. This species almost certainly is under- collected from parts of the NI where it occurs from near sea level (Auckland City) to at least 670 m. On SI occurring from near sea level (Christchurch) to at least 1220 m (St Arnaud Range, Nelson L.D.). This species is commonly associated with Breutelia affinis, Campylopus clavatus, Grimmia pulvinata, Hedwigidium integrifolium, Racomitrium crispulum s.l., Schistidium apocarpum, and Triquetrella papillata. Various lichens, especially Cladia aggregata, are also common associates. Notes: A single variable taxon of Hedwigia is present in N.Z. Although some material, particularly collections from Canterbury, exhibit some of the morphological features that Hedenäs (1994) and Buck & Norris (1996) used to characterise H. stellata (especially widely spreading leaf apices and 1–2 papillae per upper laminal cell), the more familiar name H. ciliata is applied here to this material. When dry, the stance of the leaves in N.Z. material varies from widely spreading to erect, the number of upper laminal cell papillae (c. 100 µm below the base of the hair-point) varies from 1 to 3 or occasionally more, the morphology of individual papillae varies, and the nature of the terminal cell can be either truncate/papillose or acute, even within a single collection. This variability suggests that recognition of more than one taxon in N.Z. is inappropriate. The name H. microcyathea (Müll.Hal.) Paris has been applied to both Tasmanian and N.Z. collections. There is no reason to question Mitten’s (1859) placement of the Tasmanian basionym, Pilotrichum microcyatheum Müll.Hal., in the synonymy of H. ciliata. Dixon (1927, p. 240) concurred with Mitten, but saw only N.Z. material, whereas the type is Tasmanian. There is one Beckett collection (NZ 347, Birdlings Flat, Canterbury L.D., CHR 632080), named by Müller as H. microcyathea, which is unquestionably referable to H. ciliata. The Beckett collection is almost certainly a duplicate of the collection cited by Dixon. Recognition: In Hedwigia ciliata the leaf margins are plane or weakly reflexed near the leaf base when viewed under a hand-lens; this contrasts with the strongly recurved margins in Hedwigidium integrifolium. The presence of fusiform cells at the leaf tips (even when the hair-point is poorly developed) and the more strongly papillose laminal cells also clearly distinguish Hedwigia ciliata from Hedwigidium integrifolium. Hedwigia ciliata could be confused with Schistidium apocarpum. The latter species has well- developed and dark red peristome teeth, a well-developed costa, narrower vegetative leaves, upper laminal cells that are smooth or weakly papillose-projecting (but not stellate-papillose), and non-ciliate perichaetial bracts. Etymology: The species epithet ciliata refers to the characteristic ciliate margin of the perichaetial bracts.

Hedwigidium Bruch & Schimp., Bryol. Eur. 3, 155 (1846) The genus is considered here to be monotypic and the following species description applies. Taxonomy: The separation of Hedwigidium from Hedwigia is supported by the morphological studies of de Luna (1995); his concepts are followed here.

4 Brotherus (1925, p. 70) considered Hedwigidium to contain two species, but Magill & van Rooy (1998) placed the South African H. erosum (Müll.Hal.) Paris into synonymy, effectively making the genus monotypic. The status and typification of the genus Hedwigidium was discussed by Dalton et al. (2012). They considered Hedwigidium to be typified by the Irish H. imberbe (Sm.) Bruch & Schimp. and suggested that Hedwigidium be placed in synonymy with Braunia. If their synonymy is accepted, the nomenclatural implications for N.Z. material (which clearly does not belong to Hedwigia, as they suggest) are complicated and unclear. These problems are beyond the scope of this Flora, and are best resolved in a monographic context. Etymology: The generic name Hedwigidium alludes to a similarity to Hedwigia.

Hedwigidium integrifolium (P.Beauv.) C.E.O.Jensen, Skand. Bladmossfl., 369 (1939) ≡ Hedwigia integrifolia P.Beauv., Prodr. Aethéogam, 60 (1805) Type: presumably European. Not seen. = Braunia novae-seelandiae Beckett, Trans. & Proc. New Zealand Inst. 26: 275 (1894) Type material: N.Z., Canterbury, Selwyn Gorge, on dry rock, T.W.N. Beckett 417, Sep. 1892 (CHR 632074!).

Misapplications: Hedwigidium imberbe sensu Dixon (1927) Plants medium-sized, gold-green above, dark brown or nearly black in lower portions, stoloniferous, forming mats on dry rock. Protonema globular, irregular in shape, mainly composed of rounded cells (fide de Luna 1990). Stems creeping, branching irregularly, with numerous microphyllous stolons arising from lower portions, in cross-section with incrassate outer cells grading into parenchymatous medullary cells, lacking a central strand. Rhizoids numerous, scattered on stems and stolons, smooth, brown. Shoots julaceous when dry. Leaves crowded and appressed when dry, spreading when moist, broadly ovate or oblong-ovate, acute at apex, cordate-auriculate at base, ecostate, with two plicae running close to margins from the base nearly to the apex, entire, narrowly but strongly recurved on both sides nearly to apex, 1.4–1.8(–2.1) × 0.65–0.8 mm; upper laminal cells more or less rectangular, with strongly sinuose and incrassate walls, mostly (9–)12–15 × c. 6 µm, with a few to several low and inconspicuous papillae on each surface, becoming longer (to c. 30 µm) in lower third of leaf; cells at leaf apex scarcely differing from those of upper lamina; central cells at leaf base gold-brown, more or less linear, less incrassate and less sinuose, forming a large and conspicuous group; marginal cells at base subquadrate in c. 12–15 rows extending far up the margins and gradually merging with the upper laminal cells; alar cells gold-brown or orange, forming a large but ill- defined group, sometimes slightly inflated. Microphyllous leaves (of stolons) tapered to a slender and attenuate apex. Autoicous. Perichaetia terminal on stems or on short branches, the inner leaves oblong and acute, entire, c. 3.0 mm. Perigonia bud-like, c. 0.8 mm, lateral on branches and stems (sometimes <4 mm below perichaetia), the bracts ovate-acute, concave, pigmented, surrounding numerous antheridia and few filiform paraphyses. Setae c. 1 mm, straight; capsules immersed, erect and symmetric, broadly oblong-cylindric, c. 1.5 × 1.0 mm, lacking a differentiated neck, longitudinally sulcate, gymnostomous, narrowed slightly to the transverse mouth; exothecial cells mostly isodiametric to oblong, in distinct alternating bands of thin- and thick-walled cells; stomata few, superficial, restricted to extreme capsule base; annulus nil; operculum umbonate, broadly rounded at apex, c. 0.35 mm long × 0.7 mm diam. Calyptra mitrate and elongate, c. 1.5 mm, 3–4-lobed. Spores round, 27–33 µm, finely papillose-verrucate. Illustrations: Plate 1. Beckett 1894, pl. 26 (as Braunia novae-seelandiae); Scott & Stone 1976, pl. 68; de Luna 1995, figs 4–6, 21–26 (protonema only); Meagher & Fuhrer 2003, p. 61; Seppelt et al. 2013, pl. 10. Distribution: NI: N Auckland (Taupō Bay, Penrose), S Auckland (near Ātiamuri), Gisborne (Mt Whanakao, East Cape), Wellington (Whakapapa Dam); SI: Nelson (Pyramid Rock), Canterbury (Lowry Peak Range, Lees Valley, Banks Peninsula, Mt Somers, Little Mt Peel, Sebastopol), Otago (Mt Watkin and vicinity, Lee Stream, The Neck, Mt Misery, head of Lake Wakatipu). Anomalous. Tasmania*, Australia*, East Africa*, Europe*, northern South America*, West Indies*, Java*. Recorded from Mexico and Central America and western Europe (Crum 1994a), South Africa (Magill & van Rooy 1998), and also from Sri Lanka.

5 Habitat: On dry, non-calcareous rock (including schist, greywacke, and basalt); largely restricted to the drier parts of both main islands and best documented from Canterbury and Otago. The elevation range is poorly documented. On NI it occurs to at least 915 m (East Cape, Gisborne L.D.) and on SI from 200 m (Banks Peninsula) to at least 915 m (Little Mt Peel, Canterbury L.D.). Hedwigidium integrifolium is very often associated with Cladia aggregata and Hedwigia ciliata; other less frequent associates include Campylopus clavatus, C. introflexus, Racomitrium crispulum s.l., R. pruinosum, Rhacocarpus purpurascens, Frullania falciloba, and Usnea sp. Notes: Material with “short, glistening, occasionally reddish, cuspidate points”, described from type material of Braunia novae-seelandiae at Kew by Dixon (1927), has not been observed. Type material of this name in the Beckett herbarium (from which the Kew material was segregated) lacks red, cuspidate leaf points. I agree with Dixon (1927, p. 240) that B. novae-seelandiae is best considered a synonym of Hedwigidium integrifolium, provided the ample fruiting plants of Hedwigia ciliata in the type collection are excluded from the concept. Beckett recognised the heterogeneity of this collection and mentioned it in the protologue. Recognition: Hedwigidium integrifolium bears a superficial resemblance to Pseudoleskea imbricata; H. integrifolium is a of non-calcareous rocks, while P. imbricata is mostly found on limestone. Hedwigidium integrifolium is a more robust plant with ecostate leaves mostly >1.5 mm and rectangular, obscurely papillose upper laminal cells with incrassate and sinuose walls. The less robust P. imbricata, by contrast, has costate leaves <0.75 mm and short-rhomboid, rounded upper laminal cells that lack papillae (but are sometimes obscurely prorate). The two species differ in many other ways, including the nature of their basal leaf cells and their very different sporophytes. Confusion can occur between Hedwigidium integrifolium and forms of Hedwigia ciliata with poorly developed leaf hair-points. The terminal leaf cells of Hedwigidium integrifolium differ little from the adjacent upper laminal cells, while those of Hedwigia ciliata, even if the hair-points are poorly developed, are invariably elongate-fusiform and multi-papillose. The entire perichaetial leaves of Hedwigidium also contrast with the ciliate margins in N.Z. material of Hedwigia. Scott & Stone (1976, p. 355) noted that the leaves of Hedwigidium integrifolium do not “over-expand” upon wetting as do those of Hedwigia ciliata. They also correctly noted that the leaves of the stolons are prolonged into a multicellular hyaline hair-point.

Etymology: The species epithet integrifolium means “entire-leaved” and presumably refers to the perichaetial leaf margins.

6 References

Barthlott, W.; Schultze-Motel, W. 1981: Zur Feinstruktur der Blattoberflächen und systematischen Stellung der Laubmoosgattung Rhacocarpus and anderer Hedwigiaceae. Willdenowia 11: 3–11. Beckett, T.W.N. 1894 ("1893"): On four new species of New Zealand Musci. Transactions and Proceedings of the New Zealand Institute 26: 274–277. Brotherus, V.F. 1925: Musci (Laubmoose). In: Engler, A. (ed.) Die natürlichen Pflanzenfamilien. Edition 2. Bd 11. Engelmann, Leipzig. 1–542. Bruch, P.; Schimper, W.P.; Gümbel, W.T. 1837–1850: Bryologia Europaea seu genera muscorum Europaeorum monographice illustrata. Vol. 3. E. Schweizerbart, Stuttgart. Buck, W.R.; Norris, D.H. 1996: Hedwigia stellata and H. detonsa (Hedwigiaceae) in North America. Nova Hedwigia 62: 361–370. Crum, H.A. 1994a: Hedwigiaceae. In: Sharp, A.J.; Crum, H.A.; Eckel, P.M. (ed). The Flora of Mexico. Memoirs of the New York Botanical Garden 69: 662–666. Crum, H.A. 1994b: Rhacocarpaceae. In: Sharp, A.J.; Crum, H.A.; Eckel, P.M. (ed). The Moss Flora of Mexico. Memoirs of the New York Botanical Garden 69: 667. Crum, H.A.; Anderson, L.E. 1981: Mosses of Eastern North America. Columbia University Press, New York. Dalton, N.J.; Kungu, E.M.; Long, D.G. 2012: The misapplication of Hedwigia integrifolia P.Beauv. and identity of Gymnostomum imberbe Sm. (Hedwigiaceae, ). Journal of Bryology 34: 59–61. de Luna, E. 1990: Protonemal development in the Hedwigiaceae (Musci) and its systematic significance. Systematic Botany 15: 192–204. de Luna, E. 1995: The circumscription and phylogenetic relationships of the Hedwigiaceae (Musci). Systematic Botany 20: 347–373. 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. Hedenäs, L. 1994: The Hedwigia ciliata complex in Sweden, with notes on the occurrence of the taxa in Fennoscandia. Journal of Bryology 18: 139–157. Hedwig, J. 1801: Species Muscorum Frondosorum descriptae et tabulis aeneis lxxvii coloratis illustratae. Barth, Leipzig. Jensen, C.E.O. 1939: Skandinaviens Bladmossflora. Ejnar Munksgaard, Copenhagen. Lindberg, S.O. 1864: Dasymitrium, novum genus muscorum. Öfversigt af Kongl. Vetenskaps- Akademiens Förhandlingar 21: 421–423. Magill, R.E.; van Rooy, J. 1998: Bryophyta, Part 1 Mosses, Fascicle 3, Erpodiaceae-Hookeriaceae. Leistner, O.A. (ed.) Flora of Southern Africa. Botanical Research Institute, Pretoria. Meagher, D.; Fuhrer, B.A. 2003: A Field Guide to the Mosses & Allied Plants of Southern Australia. Flora of Australia Supplementary Series. Vol. 20. Australian Biological Resources Study, Canberra. Mitten, W. 1859 ("1860"): Description of some new species of Musci from New Zealand and other parts of the southern hemisphere, together with an enumeration of the species collected in Tasmania by William Archer Esq.; arranged upon the plan proposed in the "Musci Indiae Orientalis". Journal of the Proceedings of the Linnean Society. Botany. 4: 64–100. Müller, C. 1851: Die, von Samuel Mossman im Jahre 1850. in Van Deimen's Land, Neuseeland und Neuholland gemachte Laubmoossammlung. Botanische Zeitung (Berlin) 9: 545–552, 561- 567. Palisot de Beauvois, A.M.F.J. 1804: Prodrome de l'Aethéogamie (1) ou d'un Traité sur les familles de plantes dont la fructification est extraordinaire. Famille des mousses. Magazin encyclopédique, ou journal des sciences, des lettres et des arts 9e Année, 5(19): 289–330. Palisot de Beauvois, A.M.F.J. 1805: Prodrome des cinquième et sixième familles de l'Aethéogamie. Les mousses. Les lycopodes. Fournier, Paris.

7 Paris, E.G. 1894–1898: Index Bryologicus sive enumeratio muscorum hucusque cognitorum adjunctis synonymia distributioneque geographica locupletissimis. Paul Klincksieck, Paris. Sainsbury, G.O.K. 1955: A handbook of the New Zealand mosses. Bulletin of the Royal Society of New Zealand 5: 1–490. Scott, G.A.M.; Stone, I.G. 1976: The Mosses of Southern Australia. Academic Press, London. Seppelt, R.D.; Jarman, S.J.; Cave, L.H.; Dalton, P.J. 2013: An Illustrated Catalogue of Tasmanian Mosses. Part 1. Tasmanian Herbarium, Tasmanian Museum and Art Gallery, Hobart. Smith, A.J.E. 2004: The Moss Flora of Britain and Ireland. Edition 2. Cambridge University Press, Cambridge.

8 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

9 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.

10 Acknowledgements

Jessica Beever provided advice during the preparation of this treatment. Rod Seppelt read the manuscript and suggested many worthwhile improvements. 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 the herbaria cited 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]

11 Plate 1: Hedwigia and Hedwigidium. A–H: Hedwigia ciliata. A, habit with perichaetia and capsules. B, portion of shoot with perichaetia and capsules, dry. C, portion of shoot with perichaetium and capsule. D, capsule and perichaetial leaf. E, leaves. F, inner perichaetial leaf. G, upper laminal cells. H, mid laminal cells. I–M: Hedwigidium integrifolium. I, portion of shoot, dry. J, leaves. K, upper laminal cells. L, inner perichaetial leaf. M, leaf. Hedwigia ciliata: A–G drawn from J. Child 5960, CHR 430718; H drawn from A.J. Fife 11470, CHR 532712. Hedwigidium integrifolium: I–K drawn from J.T. Linzey 424, CHR 542750; L–M drawn from A.J. Fife 8801, CHR 475429. 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. Anictangium ciliatum Hedw. 3 Braunia novae-seelandiae Beckett 5 Hedwigia P.Beauv. 1, 2, 3, 4, 5 Hedwigia albicans Lindb. 3 Hedwigia ciliata (Hedw.) P.Beauv. 3, 3, 6 Hedwigia integrifolia P.Beauv. 5 Hedwigia microcyathea (Müll.Hal.) Paris 3 Hedwigiaceae 2 Hedwigidium Bruch & Schimp. 1-3, 4, 6 Hedwigidium integrifolium (P.Beauv.) C.E.O.Jensen 3, 4, 5 Pilotrichum microcyatheum Müll.Hal. 3

15 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

16 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

17 ISBN 978-0-478-34747-0

9 780478 347470