FLORA OF NEW ZEALAND

BRUCHIACEAE

A.J. FIFE

Fascicle 27 MAY 2016 © Landcare Research New Zealand Limited 2016. Unless indicated otherwise for specific items, this copyright work is licensed under the Creative Commons Attribution 4.0 International license Attribution if redistributing to the public without adaptation: “Source: Landcare Research” Attribution if making an adaptation or derivative work: “Sourced from Landcare Research” See Image Information for copyright and licence details for images.

CATALOGUING IN PUBLICATION Fife, Allan J. (Allan James), 1951- Flora of New Zealand [electronic resource] : mosses. Fascicle 27, / Allan J. Fife. -- Lincoln, N.Z. : Manaaki Whenua Press, 2016. 1 online resource ISBN 978-0-478-34797-5 (pdf) ISBN 978-0-478-34747-0 (set) 1.Mosses -- New Zealand -- Identification. I. Title. II. Manaaki Whenua-Landcare Research New Zealand Ltd. UDC 582.344.2 (931) DC 588.20993

DOI: 10.7931/B15P40

This work should be cited as: Fife, A.J. 2016: Bruchiaceae. In: Heenan, P.B.; Breitwieser, I.; Wilton, A.D. Flora of New Zealand - Mosses. Fascicle 27. Manaaki Whenua Press, Lincoln. http://dx.doi.org/10.7931/B15P40

Cover image: suberectus, habit with capsule. Drawn by Rebecca Wagstaff from G. Brownlie 463, CHR 428081. Contents

Introduction...... 1 Typification...... 1 Taxa Bruchiaceae ...... 2 Trematodon Michx...... 2 Trematodon flexipes Mitt. ex Hook.f. & Wilson ...... 3 Trematodon mackayi (R.Br.bis) Broth...... 4 Trematodon suberectus Mitt...... 5 References ...... 7 Conventions ...... 8 Acknowledgements ...... 11 Plates ...... 12 Maps ...... 14 Index ...... 16 Image Information ...... 17

Introduction

The Bruchiaceae are a modest-sized family of about one hundred species and traditionally placed in general relationship with the Dicranaceae. Trematodon, the largest genus and the only one occurring in New Zealand, is most diverse in temperate regions of the southern hemisphere, and probably has about 80 species worldwide. Species of Trematodon are terrestrial in insolated and ephemeral sites and recognisable by their distinctive erect or curved capsules with long, narrow, often strumose necks that equal or exceed the urn length. The capsules are operculate and usually peristomate. Three species of Trematodon are accepted for the N.Z. flora. The other significant genus in the family, , also consists of terrestrial growing in disturbed terrestrial habitats. They also have long-necked capsules, but in Bruchia these are cleistocarpous. Bruchia is a less speciose genus, most diverse in North and South America; although it does not occur in N.Z. it has been recorded from Australia. The other three genera sometimes placed in this family are small genera of limited geographic distribution. The members of the Bruchiaceae are often interpreted to represent a sporophytic reduction series, with the operculate and peristomate members of Trematodon representing one end of the series and the cleistocarpous and smaller species of Bruchia the other evolutionary extreme. Throughout the family, species are defined primarily using sporophytic characters and they are rarely collected when sterile.

Typification

The following lectotypification is made in accordance with the International Code of Nomenclature for Plants, Algae and Fungi. Stirtonia mackayi R.Br.bis, Trans. & Proc. New Zealand Inst. 32: 149 (1900) Lectotype (designated here): N.Z., Stewart Island, wet turfy soil, June 1889 and Jan. 1892, R. Brown s.n., CHR 330901! Isolectotype (designated here): WELT M021258!

1 Bruchiaceae Plants small, probably annual, gregarious on soil. Stems unbranched or sparsely branched by innovation. Leaves often sheathing, ovate to lanceolate or subulate from a ± oblong base, entire or serrulate; laminal cells of upper leaf (including subula) subquadrate to rectangular to oblong-linear, firm-walled, smooth or prorate abaxially; laminal cells of leaf base similar or longer and thinner- walled; alar cells not differentiated. Costa well-developed, ending near apex or excurrent, often filling subula (if present). Brood bodies not known. Autoicous or dioicous. Setae very short to elongate, terminal; capsules erect or curved, immersed to long-exserted, obovoid to pyriform with a broad or narrow elongate neck, either cleistocarpous (in Bruchia) or opening by a functional operculum (in Trematodon). Peristome (if present) single, of 16 entire, perforate, or forked teeth, striate below. Operculum (if present) long-rostrate from a conic base. Calyptra mitrate or cucullate, not hairy, smooth or papillose. Spores rather large, mostly subreniform, variably ornamented. : The Bruchiaceae were treated as a subfamily (Trematodontoideae) of the Dicranaceae with three genera (Bruchia, Trematodon, and Wilsoniella) by Brotherus (1924). Crum (1994) took a narrow view of the family (including only Bruchia and Trematodon) and outlined his belief that these two genera represented a sporophyte reduction series “from operculate with peristome well- developed, rudimentary, or lacking in Trematodon to inoperculate and eperistomate in Bruchia and from seta and neck elongate to relatively short in Trematodon and very short in Bruchia.” Crum also presented arguments for placing genera sometimes placed here (Eobruchia and Pringleella) outside the family. Goffinet et al. (2009) presented a broader view of the family that included Bruchia and Trematodon as well as three additional and small genera of limited distributions (Pringleella, Eobruchia, and Cladophascum) in the family. The above description does not consider those lesser known genera. Only the widespread but predominantly southern hemisphere Trematodon occurs in N.Z. Trematodon is distinguished from the cleistocarpous Bruchia (which is recorded from mainland Australia by Scott & Stone (1976)) by having capsules with differentiated opercula, usually cucullate calyptra, exserted and long and slender capsules with necks that equal or exceed their urn in length, and variably developed peristomes.

Trematodon Michx., Fl. Bor.-Amer. (Michaux) 2, 289 (1803) Type taxon: Trematodon longicollis Michx. Plants mostly small, gregarious on soil. Stems unbranched or branching by innovation, beset below with smooth brown rhizoids, in cross-section with a central strand. Leaves erect-spreading, sheathing below, mostly abruptly shouldered and tapered from a ± oblong base to an elongate subula that is often equal or greater in length than the base. Laminal cells of leaf base ± oblong, thin-walled, non- porose, and smooth in N.Z. species; alar cells not differentiated. Costa often filling the subula, often poorly defined in leaf base, in cross-section (in lower subula) with a median band of guide cells. Autoicous or dioicous. Perichaetia terminal. Perigonia with inner bracts reduced and pigmented, surrounding several antheridia and filiform paraphyses. Setae elongate; capsules erect or curved, with a ± cylindric or obovoid urn and an elongate and narrow neck, operculate (dehiscent), not sulcate, often strumose; exothecial cells firm- or thick-walled, not in distinct bands; stomata numerous and superficial, restricted to neck; annulus vesicular in N.Z. species; operculum long-rostrate from a conic base. Peristome variably developed, sometimes absent or nearly so, the teeth (if present) often split into unequal branches that are fused apically, and sometimes perforate. Calyptra smooth and inflated, cucullate, or rarely ± mitrate. Spores subreniform or round. Taxonomy: A moderately large genus predominantly distributed in the southern hemisphere. Magill (1981, p. 111) considered the genus to contain c. 82 species. The genus is in need of large-scale taxonomic revision, which would likely result in a substantial reduction in the number of recognised species. The species are defined primarily using sporophytic characters and are rarely collected when sterile. Rod Seppelt (pers. comm., 14 Oct. 2015) has suggested that costal anatomy may be of taxonomic value regionally. Three species are accepted in the N.Z. flora. The genus was divided by Brotherus (1924, p. 174) into two subgenera based on the presence (subgen. Trematodon) or absence (or near absence) (subgen. Gymnotrematodon) of a peristome. Sainsbury (1955) emphasised calyptra form to define species regionally, but two of the three N.Z. species have calyptrae that vary in form. Etymology: According to Meagher (2011) tremato (perforated) + odon (tooth) alludes to the perforated peristome teeth.

2 1 Capsules apparently gymnostomous, with teeth reduced to a ± hyaline membrane scarcely exceeding the capsule mouth; spores >50 μm ...... T. mackayi 1' Capsules obviously peristomate; spores <40 μm ...... 2 2 Peristome teeth erect when dry, extending c. 300–510 μm beyond the capsule mouth; setae slender and straight or weakly flexuose; capsules strongly curved, c. 3.0–6.0 mm, the neck sometimes longer than the urn; calyptra cucullate; autoicous or possibly rhizautoicous ...... T. suberectus 2' Peristome teeth mostly reflexed when dry, extending c. 120–180 μm beyond the capsule mouth; setae stout and flexuose; capsules weakly curved or nearly straight, c. 1.5–1.9 mm, with the neck ± equal the urn; calyptra either mitrate or cucullate; dioicous ...... T. flexipes

Trematodon flexipes Mitt. ex Hook.f. & Wilson, Bot. Antarct. Voy. III. (Fl. Tasman.) Part II 173 (1859) Isotypes: Tasmania, Cumming’s Head, Western Mountains, s.d., Archer, HO 73757!, WELT M041289!

Plants small, yellow-green. Stems 1–4(–15) mm, unbranched or occasionally branched by innovation. Leaves erect-spreading, sheathing below, ± contorted when dry, distinctly shouldered and abruptly tapered from an oblong, concave and strongly sheathing base to a narrow subula that is completely filled by the costa and up to ⅔ the total leaf length, entire, to c. 2.5 mm (in perichaetium) but shorter (≤1 mm) on lower stem. Laminal cells of leaf base to c. 65(–75) × 18 µm. Costa (in leaves immediately below the perichaetium) occupying c. ⅙ the width of the base, completely filling the subula, in cross-section (lower subula) with median guide cells and abaxial and adaxial bands of smaller but non-stereid cells. Apparently dioicous (but male plants clustered near to female and possibly rhizautoicous). Setae mostly 2.5–5 mm, stout, flexuose, pale. Capsules c. 1.5–1.9 mm, weakly curved or nearly straight, erect or inclined, with the neck equal the obovoid urn, strongly strumose; exothecial cells oblong- polygonal; annulus persistent at mouth; operculum rostrate from a conic base, c. ½ capsule length. Peristome teeth variable in development, brown, inserted near mouth, broadly triangular to short- lanceolate, blunt or acute, mostly split or perforate c. halfway to base but sometimes undivided, extending 120–180 µm beyond mouth and c. 40–60 µm wide, coherent at base, strongly reflexed when dry, striate below. Calyptra mitrate or cucullate, covering the operculum and upper portion of urn. Spores 30–36 µm, coarsely baculate. Illustrations: Plate 1. Wilson 1859, tab. 172, fig. 6; Sainsbury 1955, pl. 14, fig. 2; Seppelt 2004, fig. 35. Distribution: NI: Taranaki (near Manganui Hut); SI: Nelson, Marlborough (Mt Stokes, Mt Fishtail), Canterbury (Arthur’s Pass), Otago (Pine Hill near Dunedin, Old Man Range, Rock and Pillar Range, Crown Range, Ben Lomond), Southland (Murchison Range, Borland Burn); M. Reported from A by Vitt (1979). Austral (but not known from South America). Tasmania*, Marion I.*. Reported from southern mainland Australia by Scott & Stone (1976, p. 164). Habitat: On mineral soil (often coarse gravel) or peat; often at stream or tarn margins, associated with cushion bogs, or occurring on granitic gravel in seepages. Sometimes in intermittent rain tracks. Observed at the Hope Range, Nelson L.D. forming emergent turves over granitic gravel in a very shallow rivulet with in subalpine tussock grassland. On South I. occurring from 450 (Pine Hill) to c. 1250 m (Mt Euclid, Nelson L.D.) but localities below c. 750 m are all near Dunedin. Often associated with Blindia robusta. Notes: The stems are commonly less than 5 mm, but occasional populations, perhaps growing in depositional environments, occur in which the stems ramify by innovation (e.g. A.J. Fife 7463 from Hope Range, Nelson L.D., CHR 406906). The extent of the division of the peristome teeth varies considerably within a population, and even within a single capsule. Although variable in these characters, T. flexipes is a well-defined species in a N.Z. context. Despite it having sometimes mitrate calyptrae and relatively short setae and capsules, the present species is a more robust than described for the genus Eobruchia W.R. Buck (Buck 1979) and detailed comparison to that exceedingly rare, mostly South American genus does not seem worthwhile in a flora context. Etymology: The species epithet refers to the flexuose setae.

3 Trematodon mackayi (R.Br.bis) Broth., Nat. Pflanzenfam. [Engler & Prantl] 1(3) 292 (1901) ≡ Stirtonia mackayi R.Br.bis, Trans. & Proc. New Zealand Inst. 32: 149 (1900) Lectotype: N.Z., Stewart Island, wet turfy soil, June 1889 and Jan. 1892, R. Brown s.n., CHR 330901! Isolectotype: WELT M021258! = Trematodon weymouthii E.B.Bartram & Dixon, Bot. Not. 1937: 65 (1937) Isotypes: Tasmania, on the ground, near Moore’s Lookout, West Coast, Tasmania, Mar. 1900, T.B. Moore, (herb. W.A. Weymouth 2868b), WELT M041287!, M041286!

Plants yellow-green. Stems 2–6 mm, unbranched. Leaves erect-spreading and ± twisted, sheathing below, contorted when dry, abruptly tapered from an oblong, concave and strongly sheathing base to a narrow subula that is completely filled by the costa and c. ⅔ the total leaf length, entire, mostly 4–7(–8.5) mm (in perichaetium) but much shorter below (≤2 mm). Laminal cells of leaf base mostly 75–105(–120) × c. 18–20 µm. Costa (in leaves immediately below the perichaetium) occupying c. ⅙ the width of the base, filling the subula, in cross-section (lower subula) with abaxial and adaxial bands of smaller, firm-walled (non-stereid) cells. Apparently dioicous. Setae 8–15(–23) mm, slender, nearly straight or weakly flexuose, yellow; capsules c. 3.8–5.0 mm, straight or curved, erect or inclined, with the neck equal or distinctly longer than the cylindric urn, strumose; exothecial cells elongate, not polygonal; annulus persistent at mouth; operculum rostrate from a conic base, c. ⅓ the capsule length. Peristome reduced to a hyaline membrane that scarcely exceeds the mouth, sometimes pigmented and with some striations at its upper edge. Calyptra cucullate or occasionally ± mitrate, covering the operculum and the urn. Spores (48–)54–66(–75) µm, coarsely baculate. Illustrations: Plate 1. Sainsbury 1955, pl. 14, fig. 1. Distribution: NI: N Auckland (Awanui), S Auckland (“Waikato bogs”); SI: Nelson (Collingwood, Mōkihinui, Granity, Whitehorse Bay, Four Mile River), Westland (Taylorville), Southland (Awarua Bog); St. Australasian. Tasmania*. Reported from mainland Australia by Scott & Stone (1976, p. 164). Habitat: Apparently restricted to lowland sites; usually on peat and often in pākihi; a large fraction of collections were made from recently burnt sites. From near sea level (Awarua Bog, Southland L.D.) to c. 150 m; often associated with Campylopus introflexus. Notes: The Berggren collection recorded by Sainsbury (1955, p. 94) from Ōhaeawai in N Auckland L.D. (a paratype of T. weymouthii) contains only three immature capsules and cannot be confidently named to species. In a N.Z. context this is not a notably robust species of Trematodon; Scott & Stone’s (1976, p. 164) emphasis upon the robustness of the present species is in comparison to the Australian gymnostomous species T. amoenum and is not applicable here. Its large spores and the highly reduced peristome make this species readily recognisable in a N.Z. context. It is perhaps the species with the largest spores in the entire genus. It is unclear whether the lectotype (CHR 330901) is from Brown’s June 1889 or January 1892 collection (see protologue of Stirtonia mackayi); the lectotype bears no date in Brown’s hand. The isolectotype in WELT (M021258) bears an annotation by Sainsbury: “Collection date was evidently Jany 1892 (See Trans. & Proc. N.Z. Inst. vol. 32 p. 149). This specimen (from near Traill’s house) should be considered as type, the earlier gathering being old & defective.” The CHR and WELT specimens appear to be derived from a single collection, and both have spores in the size range expected for this species. Two isotypes of T. weymouthii E.B.Bartram & Dixon (from Moore’s Lookout, Tasmania) are present in WELT. Although the protologue indicates that T.B. Moore was the collector of the type, the two duplicates in WELT are labelled as collected by L. Rodway (WELT M041287) or by W.A. Weymouth (WELT M041286). They contain three and two mature sporophytes respectively. Spores in WELT M041287 range from 54–63 µm in greater diam. and the peristome (observed without dissection of the capsule) consists of a membrane that scarcely exceeds the mouth. In other respects WELT M041287 appears to lie within the range of variability observed for T. mackayi in N.Z. On these grounds, Sainsbury’s (1955, p. 94) placement of T. weymouthii in the synonymy of T. mackayi is accepted here. A lectotype would most appropriately be selected from material in either the Bartram or the Dixon herbarium, but such material has not been seen. Etymology: The species T. mackayi was named in honour of Mr. A. Mckay [sic?], “geologist to the government of New Zealand” (Brown 1900).

4 Trematodon suberectus Mitt. in Hooker, Handb. New Zealand Fl., 415 (1867) Type: N.Z. Not seen = Trematodon arcuatus Mitt. in Hooker, Handb. New Zealand Fl., 415 (1867) Type: N.Z. Not seen = Trematodon cheesemanii Müll.Hal., Hedwigia 37: 110 (1898) Type: N.Z., Kermadec Group, Sunday (Raoul) Island, T.F. Cheeseman s.n., AK 121129!, AK 121130! = Trematodon integrifolius Müll.Hal., Hedwigia 37: 110 (1898) Type: N.Z., Marlborough, Kaikōura, 6 Dec. 1889, T.W.N. Beckett 277, CHR 585954!

Plants variable in size, bright or yellow-green. Stems c. 2–6 mm, branching by innovation. Leaves erect-spreading, sheathing below, contorted when dry, gradually tapered or abruptly shouldered from a ± oblong base to an elongate subula that is narrowly laminate below but filled by the costa above and ½–⅔ the total leaf length, entire except for a few blunt teeth at apex, mostly (3–)4.5–7 mm in perichaetium, much shorter below. Laminal cells of leaf base mostly 60–105 × 15–21 µm; laminal cells of lower subula rectangular, smooth. Costa (in leaves immediately below the perichaetium) occupying c. ⅙ the width of the base and completely filling the subula, in cross-section (lower subula) with both abaxial and adaxial stereid bands. Autoicous (or sometimes rhizautoicous). Setae c. (7–)13–15(–30) mm, slender and straight or weakly flexuose, yellow; capsules curved, mostly (2.5–)3–6(–7) mm, the neck equal or distinctly longer (to c. 1.5 × or rarely more) than the cylindric urn, weakly to strongly strumose; exothecial cells elongate; annulus apparently revoluble; operculum rostrate from a conic base, c. ⅓ the capsule length. Peristome teeth well-developed, red-brown, inserted near mouth, narrowly lanceolate, split into unequal branches nearly to the well-developed basal membrane and usually fused apically, often with other scattered perforations, 290–510 µm (excluding basal membrane), erect when dry, strongly striate. Calyptra cucullate, covering c. ½ the urn. Spores 21–33 µm, coarsely baculate-insulate. Illustrations: Plate 2. Sainsbury 1955, pl. 14, fig. 3. Distribution: K; NI: N Auckland (Te Aroha, Ōmahuta State Forest), S Auckland, Gisborne, Hawke's Bay, Wellington; SI: Nelson, Marlborough (Endeavour Inlet, Te Māhia, Kaikōura), Canterbury (Banks Peninsula) Westland, Otago, Southland (Milford Sound); St. Australasian. Mainland Australia*. Habitat: On bare soil including clay, pumice, sand, and gravel. Often on vertical banks and in roughly mown areas; in the latter forming turves to at least a square metre in extent. Also occurring on sides of drains, on spoil derived from excavations, and other disturbed habitats. From near sea level (Little Wanganui River, Nelson L.D.) to c. 1300 m (Copeland Valley, Westland L.D.). Trematodon suberectus is quite tolerant of thermally heated soil. On Kermadec Is T. suberectus is a conspicuous and abundant species on pumice associated with thermal fumaroles (W.R. Sykes, pers. comm., 10 July 2001) and its abundance there is reflected in the large number of Kermadec collections in N.Z. herbaria. A collection from “near an antimony mine” on the Coromandel Peninsula suggests that some populations may be tolerant of heavy metal concentrations. Often associated with Campylopus introflexus, Ceratodon purpureus, Dicranella jamesonii and Ditrichum difficile. Notes: The innovative branching of the stems in T. suberectus is often difficult to observe. The species is highly variable in overall stature and in capsule length; the degree of perforation of the peristome teeth also varies greatly. In most specimens the teeth are split from near their base to ½ to ⅔ their length, with the branches mostly fused at their apices. Even in single capsules, however, the degree of division varies greatly. In addition to the perforations between the two uneven branches, subsidiary perforations also occur. Such perforations are particularly numerous and conspicuous in material from Kermadec Is, including the Cheeseman material on which T. cheesemanii Müll.Hal. is based. Similar perforations can be seen in collections from scattered localities from the main islands, including T.W.N. Beckett 277 from Marlborough L.D. (the type of T. integrifolius). Clarification of the relationship between T. suberectus and T. longicollis Michx. is beyond the scope of this Flora. Trematodon longicollis is recorded to be widespread in the northern hemisphere and the name has been applied to material from P.N.G. (Norris & Koponen 1990), and elsewhere in the Pacific (Norris & Koponen 1990; Whittier 1976). Based primarily on capsule form, many North I. collections could justifiably be assigned to T. longicollis. Material from Waimangu, S Auckland L.D. (J. Child 2517, CHR 432707), has capsules with necks nearly twice the length of the urns, spores only 19–21 µm, and some plants clearly autoicous. Other collections that could justifiably be referred to T. longicollis have been seen from Hopuruahine River, Gisborne L.D. (WELT M021268), Wairoa County, Hawke’s Bay

5 L.D (WELT M021266), and Pōhāngina, Wellington L.D. (CHR 467074). Norris & Koponen (1990), without defining their concept of the “South Pacific”, state that “In the South Pacific, one should probably assign this name [T. longicollis] to any Trematodon with a fully developed peristome, and with at least some recurvature of the leaf margin.” In N.Z., material with well-developed necks and peristomes, and relatively small spores, represents one end of a morphological gradient, and it is doubtful that any clear distinction between representative T. suberectus and T. longicollis could be drawn using morphology alone. The names T. longescens Müll.Hal. and T. adaequans G.Roth have been applied to similar material from N.S.W. (e.g., CHR 588492, collected by Watts from Richmond River and named as T. adaequans by Brotherus). Monographic study, probably using non-traditional methods, would be needed to clarify the relationship of these species. At our present level of understanding, it is advisable to continue to apply the name T. suberectus to our most common and widespread species of Trematodon. The sexuality of T. suberectus is not always readily demonstrated; however, with care, some plants can usually be demonstrated to be autoicous. It is probable that in such plants the terminal meristem is consumed by the formation of the perigonium and that the perichaetial shoot arises by innovation. Dixon (1914, p. 36) observed that “the male inflorescence in T. suberectus appears to be sometimes on a short basal branch, sometimes on a separate but closely connected plant (i.e., rhizautoicous).” The rhizautoicous nature of some plants probably accounts for the varying reports of the sexuality of this species. There is no reason to question Dixon’s (1914, p. 35) conclusion that T. arcuatus Mitt. is conspecific with the present species; material so-named by Brotherus (CHR 588490) in herb. Beckett is assignable here. There is no type material of either T. suberectus Mitt. or T. arcuatus Mitt. in any N.Z. herbarium. Recognition: The distinctions from other N.Z. species of Trematodon are drawn in the key. In a N.Z. context, confusion is most likely with T. mackayi, particularly in older collections where the peristome is broken. Spore diameter is then the most reliable means of distinguishing these two species. As stated above, the relationship of the present species to T. longicollis Michx. is problematic. Etymology: The epithet suberectus refers to the less than completely erect stance of the capsules.

6 References

Brotherus, V.F. 1901–1909: Musci (Laubmoose) II Specieller Teil. In: Engler, A.; Prantl, K. (ed.) Die natürlichen Pflanzenfamilien. Teil 1. Abt. 3. Engelmann, Leipzig. 277–1246. Brotherus, V.F. 1924: Musci (Laubmoose). II. Spezieller Teil. In: Engler, A. (ed.) Die natürlichen Pflanzenfamilien. Edition 2. Bd 10. Engelmann, Leipzig. 143–478. Brown, R. 1900 ("1899"): Notes on the New Zealand Musci: on a proposed new genus. Transactions and Proceedings of the New Zealand Institute 32: 148–149. Buck, W.R. 1979: A re-evaluation of the Bruchiaceae, with the description of a new genus. Brittonia 31: 469–473. Crum, H.A. 1994: Bruchiaceae. In: Sharp, A.J.; Crum, H.A.; Eckel, P.M. (ed). The Flora of Mexico. Memoirs of the New York Botanical Garden 69: 84–89. Dixon, H.N. 1914: Studies in the bryology of New Zealand, with special reference to the herbarium of Robert Brown. Part II. Bulletin, New Zealand Institute 3(2): 31–74. Dixon, H.N.; Bartram, E.B. 1937: S. Berggren's New Zealand mosses. Botaniska Notiser 1937: 63–84. 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. Hooker, J.D. 1867: Handbook of the New Zealand Flora: a systematic description of the native plants of New Zealand and the Chatham, Kermadec's, Lord Auckland's, Campbell's, and Macquarrie's Islands. Part II. Reeve, London. Magill, R.E. 1981: Bryophyta, Part 1 Mosses, Fascicle 1, Sphagnaceae–Grimmiaceae. Leistner, O.A. (ed.) Flora of Southern Africa. Botanical Research Institute, Pretoria. Meagher, D. 2011: An etymology of Australian bryophyte genera. 2 – Mosses. Muelleria 29: 33–61. Michaux, A. 1803: Flora Boreali-Americana, sistens caracteres plantarum quas in America septentrionali collegit et detexit, Andreas Michaux. Vol. 2. Paris. Müller, C. 1898: Symbolae ad Bryologiam Australiae II. Hedwigia 37: 76–171. Norris, D.H.; Koponen, T. 1990: Bryophyte flora of the Huon Peninsula, Papua New Guinea. XXXV. Dicranaceae and Dicnemonaceae (Musci). Acta Botanica Fennica 139: 1–64. 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. 2004: The Moss Flora of Macquarie Island. Australian Antarctic Division, Kingston. Vitt, D.H. 1979: The moss flora of the Auckland Islands, New Zealand, with a consideration of habitats, origins, and adaptations. Canadian Journal of Botany 57: 2226–2263. Whittier, H.O. 1976: Mosses of the Society Islands. University Presses of Florida, Gainesville. Wilson, W. 1859 ("1860"): Musci. In: Hooker, J.D. 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. III. Flora Tasmaniae. Part II. Monocotyledones and acotyledones. Lovell Reeve, London. 160–221.

7 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 PK Poor Knights Islands

8 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

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

Supplementary Glossary for Bruchiaceae

9 Term Definition pākihi a type of wet heath, characterised by very infertile soils with an impervious horizon and little or no peat

10 Acknowledgements

Jessica Beever advised in many ways during the preparation of this treatment. Rod Seppelt read a draft and suggested several useful improvements. Rebecca Wagstaff made the line drawings with skill and patience. Peter Beveridge, Patrick Brownsey, David Glenny, Peter de Lange, and Barbara Polly provided many valuable collections for study. 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. Kate Boardman helped with electronic manipulation of images while Leah Kearns provided editing that improved the manuscript. The curators of the cited herbaria generously provided access and loans to their collections. I also thank the participants, over many years, of the John Child Bryological and Lichenological Workshops. 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: Trematodon. A–F: T. mackayi. A, capsule. B, calyptra. C, capsule with operculum, dry. D, leaves. E, perichaetial leaf. F, laminal cells of leaf base. G–L: T. flexipes. G, capsule. H, calyptra. I, capsule, dry. J, laminal cells of leaf base. K, leaves. L, perichaetial leaf. T. mackayi drawn from P.N. Johnson s.n., 21 Nov. 1985, CHR 242721. T. flexipes drawn from J. Child 6008, CHR 432677. Plate 2: Trematodon. A–I: T. suberectus. A, habit with capsule. B, calyptra. C, operculum. D, peristome detail. E, leaves. F, capsule. G, perichaetial leaf. H, laminal cells of leaf base. I, laminal cells at margin of shoulder. Drawn from G. Brownlie 463, CHR 428081. 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. Bruchiaceae 1, 2 Stirtonia mackayi R.Br.bis 4 Trematodon Michx. 1, 2, 2, 4, 6 Trematodon arcuatus Mitt. 5 Trematodon cheesemanii Müll.Hal. 5 Trematodon flexipes Mitt. ex Hook.f. & Wilson 3 Trematodon integrifolius Müll.Hal. 5 Trematodon mackayi (R.Br.bis) Broth. 4, 6 Trematodon suberectus Mitt. 5 Trematodon weymouthii E.B.Bartram & Dixon 4

16 Image Information

Image Creator Copyright Plate 1 R.C. Wagstaff © Landcare Research 2016 Plate 2 R.C. Wagstaff © Landcare Research 2016 Map 1 A.D. Wilton © Landcare Research 2014 Map 2 A.D. Wilton © Landcare Research 2014

17 18 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: Leah Kearns

19 ISBN 978-0-478-34797-5

9 780478 347975