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New Zealand Journal of Botany ISSN: 0028-825X (Print) 1175-8643 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzb20 Additions to the Lejeuneaceae flora of New Zealand: new species from the Kermadec Islands and range extensions of New Zealand species into the South Pacific MAM Renner & PJ de Lange To cite this article: MAM Renner & PJ de Lange (2011) Additions to the Lejeuneaceae flora of New Zealand: new species from the Kermadec Islands and range extensions of New Zealand species into the South Pacific, New Zealand Journal of Botany, 49:3, 421-433, DOI: 10.1080/0028825X.2011.580765 To link to this article: https://doi.org/10.1080/0028825x.2011.580765 Published online: 28 Sep 2011. Submit your article to this journal Article views: 203 View related articles Citing articles: 1 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalinformation?journalcode=tnzb20

New Zealand Journal of Botany Vol. 49, No. 3, September 2011, 421433 Additions to the Lejeuneaceae flora of New Zealand: new species from the Kermadec Islands and range extensions of New Zealand species into the South Pacific MAM Renner a,b * and PJ de Lange c a Herbarium, Auckland War Memorial Museum, Auckland, New Zealand; b National Herbarium of New South Wales, Sydney, Australia; c Ecosystems and Species Unit, Research and Development, Department of Conservation, Auckland, New Zealand (Received 7 October 2010; final version received 7 April 2011) Cheilolejeunea trifaria is recorded for New Zealand from two locations: Raoul Island in the Kermadec Islands group and Pitt Island (Rangiauria) within the Chatham Islands Group, the latter representing the world southern limit for this species. Thysananthus spathulistipus and Lejeunea exilis are also recorded for New Zealand; both having their only known occurrences within the New Zealand Botanical Region on the Kermadec Islands. Stenolejeunea acuminata is transferred to Lejeunea as L. hawaikiana nom. nov. This species (previously regarded as a New Zealand endemic) is, along with Lejeunea schusterii, newly recorded from the Kermadec Islands and Cook Islands groups. Lejeunea schusterii, a validly published name for Rectolejeunea denudata, is recognized here to reflect the increasingly supported view that most Rectolejeunea species are better accommodated within Lejeunea. Lejeunea schusterii (as Rectolejeunea denudata) had previously been regarded as a New Zealand endemic. Keywords: liverwort; Cheilolejeunea trifaria; Lejeunea exilis; Lejeunea schusterii; Lejeunea hawaikiana; Rectolejeunea denudata; Stenolejeunea acuminata; Thysananthus spathulistipus; new records; new combinations; New Zealand Flora Introduction The Kermadec Islands group (29815?31830?S, 177855?1798W) is the northernmost extension of the New Zealand Botanical Region (sensu Allan 1961; de Lange & Rolfe 2010). Of the 29 or so islands, islets, rock stacks and emergent reefs that make up the islands, Raoul Island at 2943 ha and 518 m.a.s.l is the largest and highest. Although the vascular flora of the island is well documented (see Sykes 1977; Sykes & West 1996), the bryophyte flora is poorly known. Campbell (in Sykes 1977) recorded 20 hornwort and liverwort taxa, and Beever et al. (1996) 52 mosses. Recently, So (2001) reinstated Plagiochila pacifica Mitt. as endemic to Raoul Island (the same species was also described again as the endemic P. kermadecensis Engel & Merrill by Engel & Smith-Merrill 1999), and Renner (2005) recognized a further new endemic, Radula cordiloba Taylor subsp. erigens M.A.M.Renner & Braggins, from Raoul and also noted the presence of R. javanica Gottsche on the island. Aside from these additions, nothing further seems to have been published about the bryophytes of the island. In 2009, as measures towards formalizing a proposal for World Heritage Status for the Kermadec Island group (see Anon 2006), the New Zealand Department of Conservation began a process of reviewing information about the biota of that island group. As part of that process it was recognized just how little was known about the bryophytes of Raoul Island. *Corresponding author. Email: matt.renner@rbgsyd.nsw.gov.au ISSN 0028-825X print/issn 1175-8643 online # 2011 The Royal Society of New Zealand DOI: 10.1080/0028825X.2011.580765 http://www.tandfonline.com Published online 28 Sep 2011

422 MAM Renner and PJ de Lange For a more meaningful assessment of the island s natural values it was decided that a better sampling the bryophytes was needed. To that end, one of us (Peter J. de Lange, hereafter PdL) briefly visited Raoul Island during May 2009. While specimens gathered from the visit are still being identified, preliminary listings suggest that there are now c. 136 hornworts and liverworts, and 95 mosses on the island (PdL unpubl. data). Only the two liverworts (P. kermadecensis and R. cordiloba subsp. erigens) are regarded as endemic, and no new species have been recognized so far. As part of this identification process, specimens have been distributed to a range of specialists, withthe result that the senior author recognized three lejeunaceous liverworts new to the New Zealand Botanical Region: Cheilolejeunea trifaria (Reinw., Blume & Nees) Mizut., Lejeunea exilis (Reinw., Blume & Nees) Grolle and Thysananthus spathulistipus (Reinw., Blume & Nees.) Lindenb. One of these (C. trifaria)had previously been recognized by the senior author from another 2008 gathering made by PdL from Pitt Island (Rangiauria) in the Chatham Islands. Here we formally document these taxa and provide descriptions based on the New Zealand gatherings. Information about the ecology and conservation status of these species (from a New Zealand perspective) is also provided. Recent collecting activities in Rarotonga resulted in the discovery there of two species of Lejeunea previously known as Rectolejeunea denudata R.M.Schust. and Stenolejeunea acuminata R.M.Schust., that were considered endemic to New Zealand (Schuster 2000a, b). We report these range extensions here and provide a new combination for the latter in Lejeunea. Both are also reported here for the first time from the Kermadec Islands. New records for New Zealand Cheilolejeunea trifaria (Reinw. Blume & Nees) Mizut., J. Hattori Bot. Lab. 30: 174 (1967). Jungermannia trifaria Reinw., Blume & Nees, Nova Acta Phys.-Med. Acad. Caes. Leop. - Carol. Nat. Cur. 12: 226 (1825). Lejeunea trifaria (Reinw., Blume & Nees) Nees, Syn. Hepat. 361 (1845). Euosmolejeunea trifaria (Reinw., Blume & Nees) Steph., Hedwigia 27: 292 (1888). TYPE: Indonesia. Java. Specific locality unknown, Blume s.n. & s.d. (L, STR). DESCRIPTION (Fig 1): Forming extensive pure mats. Shoots blackgreen when fresh, fading to brown in herbaria, large for genus, to 40 mm long and 8901200 mm wide, branching infrequent, shoot system monomorphic, lateral branches same stature as parent branch. Stems withthin, continuous, yellow-pigmented secondary thickening on external walls, cortical cells in 78 rows withbulging trigones at cell angles, medulla cells in 914 rows, smaller than cortical cells, with confluent nodular trigones at cell angles. Dorsal leaf-free strip absent. Branching Lejeunea-type, collar with small leaf-like bracts, persistent. Lobes rotund, 670850 mm long and 600820 mm wide, falcate, more or less plane, imbricate, margin continuously rounded, stem not visible between lobes in dorsal view. Lobe margins crenulate due to medial wall thickenings on outer walls of marginal cells. Surface of lobe cells bearing a single low papilla over eachcell. Vitta and ocelli absent. Lobules small relative to lobe size, c. 1/ 20 the lobe area, trapeziform, 130155 mm long and 90100 mm wide, carinal region weakly and broadly inflated, keel shallowly but continuously curved. Acroscopic margin not inrolled, straight. First lobule tooth indistinct, more or less incorporated into the lobule margin, but partially free on interior margin from adjacent lobule cells, unicellular, obscuring base of second lobule tooth. Second lobule tooth prominent, perpendicular to first lobule tooth, unicellular, cell apex acute. Lobule papilla attached to lobule margin in notch between first and second teeth. Underleaves rotund, 56 wider than the stem, shallowly bifid, sinus narrowly V-shaped, subauriculate, broadest at midpoint, margins entire, margins plane,

Additions to the Lejeuneaceae flora of New Zealand 423 Figure 1 Cheilolejeunea trifaria drawn from P.J. de Lange K55 & D.C. Havell, AK 305163. A, Underleaves. B, Detail of lobule. C, Stem transverse section. D, Branching diagrams. E, Lobe margin. F, Ventral view of shoot. G, Medial lobe cells. H, Antheridial bracts. I, Gynoecium on short lateral branch. J, Antheridial bracts on short lateral branch. Scale bar: D 5 mm; F,H,I,J 600 mm; B,C 60 mm; A 240 mm; E,G 36 mm. 340760 mm long and 470850 mm wide. Underleaf insertion strongly arched across two ventral cortical cell rows. Asexual reproduction absent. Autoicous. Gynoecia terminal on short lateral branches bearing zero to several pairs of vegetative leaves below the gynoecial bracts. Gynoecia usually without a subfloral innovation, rarely a single Radula-type innovation. Gynoecial bracts in one pair, asymmetrical, lobe of larger bract 550590 mm long and 270 300 mm wide, obovate, lobule of larger bract 410460 mm long and 130150 mm wide, apex truncate, lobe of smaller bract 430475 mm long and 220240 mm wide, obovate, lobule of smaller bract 340370 mm long and 90120 mm wide, elliptical. Bract underleaf oblong, apex truncate, 570620 mm long, 160180 mm wide, fused withbract lobules on bothsides. Antheridial bracts in 24 pairs produced intercalary on leafy shoots that continue vegetative growth. Antheridial bracteoles 280310 mm long and 230250 mm wide, obovate, entire. Antheridial bract lobes 220360 mm long and 390520 mm wide, lobules 220270 mm long and 260320 mu wide, keel deeply curved, apex triangular, hypostatic. Perianths and sporophytes not known in New Zealand. REPRESENTATIVE SPECIMENS: Kermadec Islands, Raoul Island, Ravine 8, P.J. de Lange K55 & D.C. Havell, 9 May 2009, AK 305163 (Duplicates: CHR, F, NSW); Ravine 8, up from Ladder, P.J. de Lange K333 & D.C. Havell, 9 May 2009, AK 313212. Chatham Islands, Pitt

424 MAM Renner and PJ de Lange Island (Rangiauria), Flower Pot Glory Road, Tupangi Stream (upper tributary), P.J. de Lange s.n. & P. B. Heenan, 30 May 2008, F (separated from AK 303450 a gathering of Radula multiflora). RECOGNITION: The recognition of Cheilolejeunea trifaria in the New Zealand Botanical Region brings the number of Cheilolejeunea (Spruce) Schiff. species recorded for New Zealand to 10, based on Glenny (1998) and Zhu (2006). Cheilolejeunea trifaria is easily recognized by its (1) huge imbricate underleaves that are six to eight times as wide as the stem, and rotund leaf lobes, which together make the plant appear isophyllous; (2) low, dome-shaped papillae on the dorsal surface of the leaf-lobe cells; and (3) small, wedge-shaped lobules, which are wider (as measured parallel with the stem) than long (as measured perpendicular to the stem), and which are also characteristic according to Mizutani (1961). The consistent presence of at least one subfloral innovation was highlighted as characteristic of Euosmolejeunea A.Evans by Evans (1906). The combination of these highly accessible characters should prevent this species being confused withany other in New Zealand. Distinctions between C. trifaria and other species of Cheilolejeunea subg. Euosmolejeunea were discussed by Mizutani (1961). DISTRIBUTION: Cheilolejeunea trifaria is known in New Zealand from two collections made from Raoul Island in the Kermadec group in May 2009, and a single May 2008 gathering from Pitt Island (Rangiauria). The Chatham Islands group is the world southern limit for C. trifaria. Outside New Zealand, C. trifaria is a widely distributed pantropical species found throughout the neotropics (Gradstein & da Costa 2003), and paleotropics (Mizutani 1961), including Java, from where the type specimen of C. trifaria was collected by Blume (Zhu et al. 2002), China (Zhu et al. 2002), and Australia (Thiers 1997). In Australia, C. trifaria occurs in tropical and subtropical rainforest of north-eastern New South Wales and Queensland. ECOLOGY: On Raoul Island, C. trifaria was collected from two stations along the same ravine system (Ravine 8). In bothcases, plants grew under a dense canopy of Metrosideros kermadecensis and Myrsine kermadecensis which are typical of the dry forest association of Raoul Island (see Oliver 1910; Sykes 1977). In the two places it was seen, it grew on coarse andesitic breccia at the apex of an ephemeral 5-m waterfall (P.J. de Lange K55 & D.C. Havell, AK 313212 reduced to a mere trickle at the time of collection, but obviously prone to sudden, extreme flashflooding during high rainfall); and on breccia boulders (AK 305163) within the ravine stream bed. In both sites it was recorded as common, and whether or not this is truly an accurate assessment of its abundance, it is likely that this species will be found in similar sites elsewhere on the island. Associated species were often sparse, probably because the habitats of C. trifaria tend to be prone to sudden flashflooding, and also to periods of extreme drought. Nevertheless, associates at the waterfall site included Anthoceros sp. (common), Ectropothecium sandwichense (Hook. & W.Arnott) Mitt. (common), Fissidens asplenioides Hedw. (common), F. leptocladus Rodw. (common), Heteroscyphus multispinus (E.A.Hodgs. & Allison) J.J. Engel & R.M.Schust., Lejeunea anisophylla Mont., L. exilis, L. gracilipes (Taylor) Steph., Lopholejeunea plicatiscypha (Hook.f. & Taylor) Steph. (common), Reboulia hemisphaerica subsp. australis R.M.Schust. (common), Schistochila kirkiana Steph., Triandrophyllum subtrifidum (Hook.f. & Taylor) Fulford & Hatcher, and Trichocolea rigida R.M.Schust. At the ravine stream bed site, associated bryophytes included Anthoceros sp., (common) Calymperes tahitense (Sull.) Mitt., Chiloscyphus helmsianus (Steph.) J.J.Engel & R.M.Schust. (common), Cyclodictyon blumeanum (C.Mu ll.) O.Kuntze, Echinodium umbrosum (Mitt.) Jaeg. (common), Ectropothecium sandwichense (common), F. asplenioides (common), F. leptocladus (common), Frullania pentapleura Taylor, Heteroscyphus allodontus (Hook.f. & Taylor)

Additions to the Lejeuneaceae flora of New Zealand 425 J.J.Engel & R.M.Schust., H. argutus (Reinw., Blume & Nees) Schiffn. (common), Lejeunea gracilipes, Lopholejeunea plicatiscypha (common), Plagiochila annotina Lindenb., P. banksiana Gottsche, P. pacifica Mitt., Phaeoceros carolinianus (Michx.) Prosk. (common), Radula cordiloba subsp. erigens, Riccardia aff. wattsiana (common) and Telaranea lindenbergii (Gottsche) J.J.Engel & G.L.S.Merrill var. lindenbergii. On Pitt Island (Rangiauria), Chatham Islands, C. trifaria occurred as scattered shoots admixed with Lejeunea anisophylla within a mat of Radula multiflora Gottsche ex Schiffn. (see Renner & de Lange 2009) that was growing on a rotting Olearia telmatica Heenan & de Lange log and surrounding clay shaded by overhanging matagouri (Discaria toumatou Raoul) bushes. Other associated bryophytes included Campylopus clavatus (R.Br.) Hook.f. & Wils., Lejeunea anisophylla and Solenostoma inundatum (Hook.f. & Taylor) Mitt. ex Steph. CONSERVATION STATUS: On Raoul Island, C. trifaria was recorded as common at in Ravine 8 and the gatherings from there are copious. However, the steep topography of the island, its size (2943 ha) and the length of time PdL spent there (2.5 days) was hardly sufficient to form any definitive impression of the species abundance. Nevertheless, ravine habitats are common on Raoul Island, and aside from Ravine 8, none of them have been thoroughly explored and documented from a bryological point of view. On the Chatham Islands, we know even less about the distribution of the species, which was collected by accident amongst a larger gathering of Radula multiflora. The occurrence of this species in both the Kermadec and Chatham Island groups is mirrored by that of other bryophytes such as Calymperes tenerum C.Mu ll., Pyrrhobryum paramattense (C.Mu ll.) Manuel and Syrrhopodon armatus Mitt. and Lejeunea anisophylla (see Fife & de Lange 2009; Renner & de Lange 2009). These distribution patterns were discussed briefly by Fife & de Lange (2009) who suggested that, with respect to Calymperes tenerum, it should be searched for in northern New Zealand. Calymperes tenerum has subsequently been discovered at Te Paki, NorthIsland, New Zealand (de Lange & Fife 2010). The highly disjunct distribution of C. trifaria follows so closely what is now known of the distribution of Calymperes tenerum, that we think it highly probable that it too will eventually be found in northern New Zealand. In summation then, as the species is known from three gatherings (one accidental) from two highly disjunct locations in two remote island groups within the New Zealand Botanical Region (sensu Allan 1961) and as no thorough survey for it has been undertaken in either island group, we suggest that a threat assessment of Data Deficient sensu Townsend et al. (2008) is appropriate. This assessment should be qualified SO (Secure Overseas), as the species is widely distributed and, at least in Australia, is well represented within legally protected areas. ADDITIONAL NOTES: The first reporting of Cheilolejeunea subg. Euosmolejeunea for New Zealand was by Campbell (1977) based on Sykes 671/K (CHR 161835) from Raoul Island. This material has since been examined by Zhu (2006) who concluded it was C. mimosa (Hook.f. & Taylor) R.M.Schust. We concur withzhu s determination. Cheilolejeunea mimosa does not belong to Cheilolejeunea subg. Euosmolejeunea, making the records of C. trifaria reported here the first genuine occurrence of this primarily tropical subgenus within the New Zealand Botanical Region. Cheilolejeunea trifaria is the type species of Cheilolejeunea subg. Euosmolejeunea (Schuster 1980; Zhu & Lai 2005). Euosmolejeunea is one of the most distinctive subgenera of Cheilolejeunea. It is characterized by having (1) large, imbricated, shallowly bifid, reniform underleaves with cordate bases and strongly arched, U-shaped insertion; (2) ovate leaf lobes; (3) small, inconspicuous, non-elongate unicellular first lobule tooth; and (4) perianths always with one or two

426 MAM Renner and PJ de Lange subfloral innovations. Of these characters, the large imbricate underleaves, in combination withthe rounded leaf lobes, give species of this subgenus the appearance that they bear three rows of equal leaves. Lejeunea exilis (Reinw., Blume & Nees) Grolle, J. Hattori Bot. Lab. 46: 353 (1979). Jungermannia exilis Reinw., Blume & Nees, Nava Acta Caes. Leop. Nat. Cur. 12: 227 (1825). Jungermannia cucullata Reinw., Blume & Nees var. b exilis (Reinw., Blume & Nees) Nees., Enumer. Pl. Crypt. Jav. Hepat. 57 (1830). Lejeunea cucullata (Reinw., Blume & Nees) Nees var. b exilis (Reinw., Blume & Nees) Gottsche, Lindenberg & Nees, Syn. Hepat. 390 (1845). Eulejeunea cucullata (Reinw., Blume & Nees) Schiffn. var. b exilis (Reinw., Blume & Nees) Schiffn., Consp. Hepat. Arch. Ind. 254 (1898). Microlejeunea exilis (Reinw., Blume & Nees) Bischler Bonner & H.A.Mill., Nova Hedwigia 3: 452. (1962). TYPE: Indonesia. Java (STR, PC, S, W). Microlejeunea lancistipula Steph. Sp. Hepat. 5: 830 (1915). Lejeunea lancistipula (Steph.) H.A.Mill., Bonner et Bischler. Nova Hedwigia 14: 66 (1967). TYPE: New Guinea. Stephanson, Aug. 1883, Kürnbach 1023 (G14864). Microlejeunea subacuta Horik. J. Sci. Hiroshima Univ. Ser. B. Div. 2, Bot. 2: 275 (1934). Drepanolejeunea subacuta (Horik.) H.A.Mill., Bonner & Bischler. Nova Hedwigia 4: 560 (1963). TYPE: Taiwan, Taito, Mt. Chipon, epiphyllous, 30 Dec. 1932, Y. Horikawa 10478. (holotoype: HIRO). Byssolejeunea abnormis Herzog Hedwigia 80: 84 (1941). Lejeunea abnormis (Herzog) R.M.Schust. J. Hattori Bot. Lab. 25: 4 (1962). non Lejeunea abnormis (Gottsche) Steph. in Renauld & Cardot Rev. Bryol. 18: 57 (1891). Lejeunea byssiformis Grolle et Mizut. in Mizutani J. Hattori Bot. Lab. 43: 131 (1978). Microlejeunea abnormis (Herzog) Inoue & H.A.Mill. Bull. Nat. Sci. Mus. Tokyo 11: 9 (1968). TYPE: Indonesia. Java. Pangerango prope Thibodas, 1330 m, Schiffner 2676 p.p. (holotype: JE). DESCRIPTION (Figs 2, 3): Forming diffuse patches of thread-like shoots, either pure or admixed within other bryophytes, mid-green when fresh, fading to pale yellowbrown in herbaria. Shoots to 10 mm long and 220 400 mm wide. Sparingly and irregularly branched. Shoot system monomorphic, lateral branches same stature as parent branch. Stems without secondary thickening on external cell walls, cortical cells in 7 rows withweak triangular trigones at cell angles, medulla cells in 35 rows, smaller than cortical cells, withweak triangular trigones at cell angles. Dorsal leaf-free strip present, one cell row wide, Branching Lejeunea-type, collar low basal ring, persistent. Leaves dimorphic. Normal leaf lobes parabolic 200250 mm long and 150200 mm wide, obliquely-patent, remote, margins straight to slightly curved, entire, apex acute. Surface of lobe cells withfine granular ornamentation. Reduced leaf lobes lanceolate. Vitta and ocelli absent. Stem visible between lobes in dorsal view. Lobules on normal leaves large relative to lobe size, c. 2/3 the lobe area, broadly ovate, inflated, keel curved in interior half then straight, antical margin straight, not obscured in ventral view, first lobule tooth unicellular, cell curved outward to point away from shoot apex, lobule arch U-shaped, of three cells, lobule second tooth obsolete, lobule papilla attached to lobule margin at base of first lobule tooth. Lobules on reduced leaves two to five cells only. Underleaves small, widely remote, 0.52.0 wider than the stem, broadest toward apex of lobes, variable, large underleaves deeply bifid, withdivergent lobes one to two cells wide at base and two to three cell tiers high, sinus gaping, smaller underleaves unlobed, lanceolate. Underleaf insertion transverse across two ventral cortical cell rows in

Additions to the Lejeuneaceae flora of New Zealand 427 Figure 2 Lejeunea exilis drawn from Renner 2105, NSW872056 (Australian material). A, Dorsal shoot view. B, Ventral shoot view. C, Gynoecium withbracts and subfloral innovation. D, Underleaves. Scale bar 300 mm. bifid underleaves, and a single ventral cortical cell in lanceolate underleaves. Asexual reproduction by ribbon-like gemmae produced from leaf lobe margins. Dioicous (?). Gynoecia terminal on shoots, subtended by a single Radula-type subfloral innovation bearing Lejeuneoid innovation sequence. Gynoecial bracts in one pair, subsymmetrical lobes ovate, acute, lobules oblong bearing two prominent teeth, bract underleaf oblong, bifid, margins of lobes denticulate or smooth, fused with bract lobules on bothsides. Males, perianths and sporophytes not seen in New Zealand material. REPRESENTATIVE SPECIMENS: Kermadec Islands, Raoul Island: R.C. Cooper 44333A, 1956, AK 297625; W.R. Sykes 671/K p.p., 23 Nov 1966, CHR 593620; Moumoukai Track, P.J. de Lange K208 & D.C. Havell, 8 May 2009, AK 313197; Moumoukai near summit, P.J. de Lange K258 & D.C. Havell, 8 May 2009, AK 313699; Moumoukai Summit, P.J. de Lange K249 & D.C. Havell, 23 May 2009, AK 313692; Moumoukai Eastern side of Summit, P.J. de Lange K288 & D.C. Havell, 8 May 2009, AK 313705; Moumoukai Summit (crater rim), P.J. de Lange K272 & D.C. Havell, 8 May 2009, AK 313724; Ravine 8, Hebe Site, P.J. de Lange K388 & D.C. Havell, 9 May 2009, AK 313216; Ravine 8, Hebe Site, P.J. de Lange K388 & D.C. Havell, 9 May 2009, AK 313878; Ammunition Store Bush, P.J. de Lange K28, 10 May 2009, AK 313179. RECOGNITION: Lejeunea exilis can be recognized by (1) the diminutive stature; (2) the widely remote leaves, that are acute and dimorphic; (3) the large inflated lobules twothirds the lobe size on normal leaves, and furnished with a unicellular first lobule tooth that curves outward away from the shoot apex; and (4) the frequent production of lanceolate, unlobed underleaves. Due to its small size and pointed leaf lobes, Lejeunea exilis is easily confused with Drepanolejeunea (Spruce) Schiffn. and Harpalejeunea

428 MAM Renner and PJ de Lange Figure 3 Lejeunea exilis on Rhopalostylis baueri frond (specimen from Moumoukai Track, Raoul Island). (Spruce) Schiffn. emend. Grolle & R.L.Zhu from which it is distinguished by (1) the frequent production of unlobed underleaves, (2) the absence of ocelli in leaf lobes, (3) the lack of teethon leaf lobe margins, and (4) the occasional presence of ribbon-like thalloid gemmae produced from leaf-lobe margins. Within Lejeunea, L. exilis is most likely to be confused with L. primordialis (Hook.f. & Taylor) Taylor; however, L. primordialis differs in that the leaves have rounded to obtuse lobes (not acute); the lobules are one-quarter the lobe area (not two-thirds); by the production of copious leafy cladia (not present in L. exilis); and by the underleaves that are always bifid (never withlanceolate underleaves), withthe underleaf sinus not gaping. These two taxa are also allopatric. Lejeunea primordialis is a southtemperate species common in the southern half of the North Island and throughout the South Island, Stewart Island (Rakiura), the Chatham Islands and New Zealand s subantarctic Islands. It is, otherwise, only sparsely distributed throughout the upper half of the North Island, usually in association withcloud forest on mountain peaks. Lejeunea primordialis has not been collected on the Kermadec Islands. DISTRIBUTION: In New Zealand, L. exilis is only known from Raoul Island in the Kermadec Islands group. Outside the New Zealand Botanical Region, this species is widely distributed throughout the paleotropics including China, Japan, Taiwan, Phillipines, Malaysia, Indonesia, Papua New Guinea, Australia, Norfolk Island and throughout Oceania, including the Caroline Islands (Zhu & Grolle 2003). ECOLOGY: Lejeunea exilis is a widespread species occupying a range of habitats from damp andesitic breccia seepages and ravine walls, lignum, bark as well as branchlets and foliage, most especially those of the indigenous palm Rhopalostylis baueri (Seem.) H.Wendl. & Drude and fern Hymenophyllum demissum (G.Forst.) Sw. It is also a frequent contaminant of larger bryophytes, including Echinodium umbrosum and Plagiochila pacifica (which are, arguably, the most common of the larger bryophytes on Raoul Island). In these habitats it is frequently associated with (and mostly threaded through) - Cheilolejeunea spp., Drepanolejeunea spp., Frullania spp., Metalejeunea spp., Lejeunea anisophylla and L. gracilipes and Lopholejeunea pilcatiscypha (Taylor) Steph. CONSERVATION STATUS: Until recently, L. exilis was known in the New Zealand Botanical Region (Allan 1961; de Lange & Rolfe 2010) only from a handful of chance gatherings from Raoul Island. Following the May 2009 visit upon which this paper is based, L. exilis was found to be a widespread and common species over those parts of Raoul Island visited by PdL. Plants were found in the two main vegetation types distinguished by Oliver (1910) and Sykes (1977), and were a common bycatch in other liverwort gatherings. There are no threats to this species on Raoul, beyond volcanic eruptions, which are natural phenomena that this species has had to contend withand whichare of no valid conservation concern here. Thus, from the accumulated herbarium specimens now available and the field observations, it is our view that L. exilis merits a threat listing of At Risk/ Naturally Uncommon sensu Townsend et al. (2008). To this suggested listing the qualifiers OL (One Location), SO (Secure Overseas) should be added. We see no reason to add the qualifier DP, as the spread of gatherings now available and the habitats occupied suggest that this minute species is widespread in any forested habitat on Raoul Island.

Additions to the Lejeuneaceae flora of New Zealand 429 ADDITIONAL NOTES: Lejeunea exilis was first recognized from the New Zealand Botanical Region by Rui-Liang Zhu in 2003 when it was separated from a Raoul Island, November 1966, Sykes gathering of Plagiochila pacifica. However, aside from Zhu s annotation on the specimen New to New Zealand, the record seems to have gone unremarked in the literature. Thysananthus spathulistipus (Reinw., Blume & Nees.) Lindenb., Syn. Hepat. 287 (1845). Jungermannia spathulistipus Reinw., Blume & Nees. Acta Phys. Med. Acad. Caes. Leop.- Carol. Nat. Cur. 12: 212 (1824). Lejeunea spathulistipa (Reinw., Blume & Nees.) Dum. Rec. d Observ. 12 (1835). Frullania spathulistipa (Reinw., Blume & Nees.) Nees Nat. eru. Leber. 3: 211 (1838). Phragmicoma spathulistipa (Reinw., Blume & Nees.) Mitt. in Seeman Fl. Vit. 412 (1873). TYPE: Java. Bantam. Leback. Blume s.n. & s.d. (STR, W, G). DESCRIPTION (Fig. 4): Forming diffuse pure patches. Shoots browngreen when fresh, fading to brown in herbaria, small for subfamily, to 20 mm long and 9801580 mm wide, branching infrequent and irregular, shoot system monomorphic, lateral branches same stature as parent branch. Stems with external and internal walls heavily and irregularly thickened by larger trigones of yellowbrown pigmented secondary wall, 1825 cortical cells and 3040 medulla cells the same size as the cortical cells. Dorsal leaf-free strip absent. Branching Lejeunea-type, collar persistent. Lobes ovate, 720920 mm long and 500660 mm wide, widely spreading, in situ strongly falcate due to reflexed basiscopic margin, contiguous to imbricate, when dry lobes adpressed to and surrounding stem. Lobe apex obtuse to acute, usually withapiculus and several small accessory teeth, shallowly arched, curvature increasing toward the stem insertion, stem not visible between lobes in dorsal view. Lobe margins irregularly crenulate. Surface of lobe cells smooth. Lobules small relative to lobe size, c. 1/20 the lobe area, oblong 190230 mm long and 95130 mm wide, carinal region weakly and broadly inflated, keel straight most of its length, slightly curved toward keellobe junction. Acroscopic margin not inrolled, curved at internal base otherwise straight until the notch within which the lobule papilla is situated. Lobule typically bearing no obvious teeth, occasionally one obsolete toothpresent, margin exterior to papilla straight to arched, also lacking obvious dentition. Underleaves unlobed, obovate to obtrapeziform, truncate, broadest at or above midpoint, margins entire or irregular, apex weakly denticulate, withone or bothlateral margins reflexed completely or in part on larger underleaves, 23 wider than the stem, 220340 mm long and 215355 mm wide. Underleaf insertion transverse across seven or eight ventral cortical cell rows. Asexual reproduction absent. Dioicous? (Female plants not seen.) Antheridial bracts in 24 pairs produced intercalary on leafy shoots that continue vegetative growth. Antheridial bract lobes 220360 mm long and 390520 mm wide, lobules 220270 mm long and 260320 mm wide, keel deeply curved, apex triangular, hypostatic. Antheridial bracteoles 280310 mm long and 230250 mm wide, obovate, entire. REPRESENTATIVE SPECIMEN: Kermadec Islands, Raoul Island: Fishing Rock Road, Ammunition Store Bush, P.J. de Lange K27, 10 May 2009, AK 313178. RECOGNITION: In the field, the combination of incubously inserted leaves, presence of lobules, the unlobed underleaves, and brown pigmentation are characteristic of Lejeuneaceae subf. Ptychantoideae. Within this subfamily, Thysananthus spathulistipus can be recognized by (1) the falcate leaf lobes with acute and denticulate lobe apices and deflexed basiscopic margins; (2) the obovate underleaves, with a weakly denticulate apex, and reflexed lateral margins on larger underleaves; (3) the lobules one twentieththe lobe area, withan indistinct or redundant first lobule tooth, and shallowly curved keel; and (4) the stems lacking a hyalodermis, with coarse confluent trigones in bothcortical and medullary layers.

430 MAM Renner and PJ de Lange Figure 4 Thysananthus spathulistipus drawn from P.J. de Lange K27, AK 313178. A, Leaves. B, Lobule detail. C, Medial lobe cells. D, Ventral shoot view. E, Dorsal shoot view. F, Lobe apex. G, Underleaves. H, Stem transverse section. I, Antheridial bracts. Scale bar: A,D,E,I 600 mm; B,F,H 60 mm; G 240 mm; C 36 mm. DISTRIBUTION: In the New Zealand Botanical Region known only from Raoul Island (Kermadec Islands group). Thysananthus spathulustipus is a widespread and common species found throughout the Paleotropics, including Australia (Thiers & Gradstein 1989). ECOLOGY: Known only from a permanently shaded, seasonally dry north-facing andesitic breccia cliff within the dry forest type of Oliver (1910) and Sykes (1977). This forest type is dominated by Kermadec pohutukawa (Metrosideros kermadecensis W.R.B.Oliv.) with an understorey of Myrsine kermadecensis Cheeseman, and, at the site where Thysananthus was collected, other common understorey associates included Melicytus aff. ramiflorus J.R.Forst. & G.Forst., and Pisonia brunoniana Endl. Within the breccia cliff habitat, T. spathulistipus formed large, mostly monospecific carpets in the damper recesses and overhangs toward the base of cliffs. Here it was occasionally found admixed with L. gracilipes and Lopholejeune a plicatiscypha which were partially shaded by Asplenium shuttleworthianum Kunze. CONSERVATION STATUS: Although known from only a single gathering from Raoul Island, T. spathulistipus was abundant at this location and as the type of habitat in which it was found is common on Raoul, it is likely that it will prove widespread on that island. Aside from periodic volcanic activity there are no active threats to this species. Raoul Island, as part of the Kermadec Islands Nature Reserve, is afforded the highest possible legal protection in New Zealand, withaccess strictly controlled by permits which have fixed limits on the numbers of people on the island, their length of stay and places they can visit. Nevertheless, until such time as a proper survey can be conducted it seems appropriate to regard this species as Data Deficient sensu Townsend et al. (2008). This assessment should

Additions to the Lejeuneaceae flora of New Zealand 431 be qualified SO (Secure Overseas) as the species is known to be widely distributed, throughout Australia and Oceania (Thiers & Gradstein 1989). ADDITIONAL NOTES: Ten species of subf. Ptychanthoideae in five genera are now known for New Zealand: Acrolejeunea (three species), Archilejeunea (one species), Lopholejeunea (three species), Ptychanthus (one species) and Thysananthus (two species). Five of these species are apparently endemic to New Zealand, if the sole Australian record of Lopholejeunea colensoi is based on mislabelled material (see Thiers & Gradstein 1989). Range extensions into the Pacific Lejeunea hawaikiana M.A.M.Renner & de Lange nom. nov. Stenolejeunea acuminata R.M.Schust. J. Hattori Bot. Lab. 89: 156 (2000) non Lejeunea acuminata (Lehm. & Lindenb.) Lehm. & Lindenb. Syn. Hepat. 3: 354 (1845). TYPE: New Zealand, NorthIsland, Northland, Radar Bush R. M. Schuster 95503, (holotype: herb. R. M. Schuster). NOTES: So derstro m et al. (2010) proposed Stenolejeunea be synonymized with Lejeunea, and proposed two new combinations for Javan taxa. Molecular evidence also supports the nesting of Stenolejeunea within the Lejeunea generic-complex (Renner 2008). We follow this synonymization and propose the above combination for the regional representative of Stenolejeunea Because the epithet acuminata is preoccupied in Lejeunea by L. acuminata (Lehm. et Lindenb.) Lehm. & Lindenb., a new epithet for Stenolejeuna acuminata Lejeunea schusterii is needed. We have chosen hawaikiana to acknowledge the cultural links between New Zealand and Rarotonga that are reflected in the known distribution of this liverwort. REPRESENTATIVE SPECIMENS: Cook Islands, Rarotonga: Vaimanga, Ara Tapu Road, P.J. de Lange CK40 & T. J. P. de Lange, 25 Jul 2010, AK 316667. Kermadec Islands, Raoul Island: Matatirohia, Water Tanks Track, Selaginella Site, P.J. de Lange K414, 9 May 2009, AK 305160. New Zealand, NorthIsland, North Auckland: Radar Bush, south of Mt Te Paki, J. E. Braggins 95065, 6 Feb 1995, AK 254990; Radar Bush, south of Mt Te Paki, J. E. Braggins 95066A, 6 Feb 1995, AK 254991; Karikari Peninsula, Matai [Maitai] Bay, Takanini Point, M.A.M. Renner 02/02b, 2 Jan 2002, AK 280234; Whangaroa Harbour, Totara North, near Sherman Stream, D. Glenny 10947 & M.A.M. Renner, 12 Jan 2009, CHR 606681; Maunganui Bluff, M.A.M. Renner 398a, 2 Apr 2003, AK 286387. Chatham Islands: Chatham Island (Rekohu), Te Hapupu, Te Hapupu Bush, P.J. de Lange CH1388 & P. B. Heenan, 1 Jun 2008, AK 302884. Lejeunea schusterii Grolle Haussknechtia 8: 60. (2001). Rectolejeunea denudata R.M.Schust. J. Hattori Bot. Lab. 89: 143 (2000). TYPE: New Zealand, NorthIsland, Northland, lowland coastal forest near Portland, R.M. Schuster 95-141, (holotype: herb. R.M. Schuster). NOTES: Rectolejeunea A.Evans sensu Schuster is a heterogeneous assemblage of unrelated entities, and is polyphyletic (Renner 2008). While the generic type, Rectolejeunea berteroana (Gottsche ex Steph.) A.Evans, groups with Lepidolejeunea R.M.Schust. in the phylogeny of Wilson et al. (2007), most remaining species are probably nested within the Lejeunea generic complex. Morphological characters indicating a close relationship between Rectolejeunea berteroana and Lepidolejeunea include the presence of leaf ocelli, the presence of secondary thickening in the stem medulla, the sideways-pointing first lobule tooth, and the production of rhizoids from leaf lobe margins. By contrast, most species assigned to Rectolejeunea have more in common with species of Lejeunea in their lack of ocelli, lack of secondary thickening in the stem medulla and normal first lobule tooth. Considering the type species of Rectolejeunea, the phylogeny of

432 MAM Renner and PJ de Lange Wilson et al. (2007) and the morphological characters described above, Lepidolejeunea would be a synonym of Rectolejeunea, while all species of Rectolejeunea lacking ocelli and sideways-pointing lobule teethwould need reinstating in or transferring to Lejeunea. Grolle (2001) has pre-empted this move for Rectolejeunea denudata, and as this reflects our view of relationships we follow him here. In contrast, we believe that maintenance of Rectolejeunea ocellata R.M.Schust. within Rectolejeunea is appropriate. Lejeunea schusterii (as Rectolejeunea) was considered a New Zealand endemic (Schuster 2000a). In May 2009 it was discovered on Raoul Island (Kermadec Islands), a find that strongly suggested it would be found elsewhere in the south Pacific. In July 2010, PdL found it in the Cook Islands group on Rarotonga. We suspect that with diligent searching it will be found elsewhere in Oceania. REPRESENTATIVE SPECIMENS: Cook Islands, Rarotonga: Vaimanga, Ara Tapu Road, P.J. de Lange CK40 & T.J.P. de Lange, 25 Jul 2010, AK 316668. Kermadec Islands, Raoul Island: Moumoukai Track, P.J. de Lange K51 & D.C. Havell, 8 May 2009, AK 313187. New Zealand, NorthIsland, NorthAuckland: NorthCape, NorthCape Scientific Reserve, Surville Cliffs, P.J. de Lange 8097, 19 Oct 2009, AK 307123; LeighMarine Laboratory Reserve, Goat Island, J.E. Braggins 86249C, 1 Nov 1986, AK 258966; Goat Island Marine Reserve, M.A.M. Renner 2847, 28 Jun 2007, AK 300070; Omaha, Omaha causeway, J.E. Braggins 95028, 22 Jan 1995, AK 254903; Million Bay, halfway between Baddleys and Campbell Beaches, P.J. de Lange 7877, 23 Jan 2009, AK 304944. Chatham Islands: Pitt Island (Rangiauria), Waipaua, Waipaua Stream, P.J. de Lange CH1706 & P.B. Heenan, 30 May 2008, AK 303422. Acknowledgements We thank Anders Hagborg (Early Land Plants Today, Field Museum, Chicago) for drawing our attention to Grolle s nomen novum. PdL wishes to acknowledge the considerable help of David Havell and Chauncey Ardell, both of Auckland Conservancy, Department of Conservation, while in the field on Raoul Island, and the crew of RNZN Resolution for transport to and from the Kermadec Islands. Thanks also to Amanda Baird, Kenny Dix and Ken Hunt of the Department of Conservation, Chatham Islands for assistance during visits to those islands, and to Peter Heenan and Gary Houliston for helping in field work while on Pitt Island (Rangiuria). Both authors thank John Braggins for providing the image of Lejeunea exilis taken from freshmaterial provided by PdL from Raoul Island in May 2009. We also thank three anonymous referees for their suggested improvements. References Allan HH 1961. Flora of New Zealand. Vol. 1. Wellington, Government Printer. 1085 p. Anon 2006. Our world heritage a tentative list of New Zealand cultural and natural heritage sites. Wellington, Department of Conservation. 79 p. Beever JJ, Fife AJ, West CJ 1996. Mosses of the Kermadec Islands, northern New Zealand. New Zealand Journal of Botany 34: 463471. Campbell EO 1977. Hepaticopsida (Hepaticae) and Anthocerotopsida In: Sykes WR ed. Kermadec Islands Flora an annotated checklist. Wellington, New Zealand Department of Scientific and Industrial ResearchBulletin 219. Pp. 200 205. de Lange PJ, Fife AJ 2010. Calymperes tenerum in northern New Zealand. Australian Bryological Newsletter 58: 89. de Lange PJ, Rolfe JR 2010. New Zealand indigenous vascular plant checklist. Wellington, New Zealand Plant Conservation Network. Engel JJ, Smith-Merrill GL 1999. Austral Hepaticae 31. Two new species of Plagiochila (Dum.) Dum. (Plagiochilaceae) from New Zealand. Journal of the Hattori Botanical Laboratory 87: 295300. Evans AW 1906. Hepaticae of Puerto Rico. VI. Cheilolejeunea, Rectolejeunea, Cystolejeunea, and Pycnolejeunea. Bulletin of the Torrey Botanical Club 33: 125. Fife AJ, de Lange PJ 2009. Calymperes tenerum Mu ll.hal. (Calymperaceae) on the Chatham Islands, New Zealand. Australian Bryological Newsletter 57: 1416. Glenny D 1998. A revised checklist of New Zealand liverworts and hornworts. Tuhinga 10: 119149.

Additions to the Lejeuneaceae flora of New Zealand 433 Gradstein SR, da Costa DP 2003. The Hepaticae and Anthocerotae of Brazil. Memiors of the New York Botanical Garden 87: 1318. Grolle R 2001. Miscellanea Hepaticologica 291300. Haussknechtia 8: 5969. Mizutani M 1961. A revision of Japanese Lejeuneaceae. Journal of the Hattori Botanical Laboratory 24: 115302. Oliver WRB 1910. The vegetation of the Kermadec Islands. Transactions of the New Zealand Institute 42: 118175. Renner MAM 2005. Additions to the Radula (Radulaceae: Hepaticae) floras of New Zealand and Tasmania. Journal of the Hattori Botanical Laboratory 97: 3979. Renner MAM 2008. Morphological variation informs evolutionary relationships within the Lejeunea tumida Mitt. aggregate (Lejeuneaceae: Jungermanniopsida). Unpublished PhD thesis, University of Sydney, Sydney. 492 p. Renner MAM, de Lange PJ 2009. Radula multiflora Gottsche ex Schiffn a new record for the Chatham Islands, New Zealand. Australian Bryological Newsletter 57: 1213. Schuster RM 1980. The Hepaticae and Anthocerotae of NorthAmerica. Volume 4. New York, Columbia University Press. 1334 p. Schuster RM 2000a. Studies on Lejeuneaceae, II. Rectolejeunea Evs. emend. Schust. (Lejeuneoideae). Journal of the Hattori Botanical Laboratory 89: 113150. Schuster RM 2000b. Studies on Lejeuneaceae, III. Revisionary studies on Stenolejeunea Schust. Journal of the Hattori Botanical Laboratory 89: 151171. So ML 2001. Studies on Plagiochila subgenus Metaplagiochila (Plagiochilaceae, Hepaticae). Australian Systematic Botany 14: 677688. So derstro m L, Gradstein SR, Hagborg A 2010. Checklist of the hornworts and liverworts of Java. Phytotaxa 9: 53149. Sykes WR 1977. Kermadec Islands Flora an annotated checklist. Wellington, New Zealand Department of Scientific and Industrial ResearchBulletin 219. 216 p. Sykes WR, West CJ 1996. New records and other information on the vascular flora of the Kermadec Islands. New Zealand Journal of Botany 34: 447462. Thiers BM 1997. Cheilolejeunea in Australia: description of new taxa and key. Journal of the Hattori Botanical Laboratory 82: 321328. Thiers BM, Gradstein SR 1989. Lejeuneaceae (Hepaticae) of Australia. I. Subfamily Ptychanthoideae. Memiors of the New York Botanical Garden 52: 179. Townsend AJ, de Lange PJ, Norton DA, Molloy J, Miskelly C, Duffy C 2008. The New Zealand Threat Classification System manual. Wellington, Department of Conservation. 35 p. Wilson R, Gradstein SR, Schneider H, Hienrichs J 2007. Unravelling the phylogeny of Lejeuneaceae (Jungermanniopsida): evidence for four main lineages. Molecular Phylogenetics and Evolution 43: 270282. Zhu R-L 2006. On Cheilolejeunea tenella (Jungermanniopsida: Lejeuneaceae), a poorly known species from Singapore. Annales Botanici Fennici 43: 484487. Zhu R-L, Grolle R 2003. Taxonomy and distribution of Lejeunea exilis (Lejeuneaceae, Hepaticae). Annales Botanici Fennici 40: 101106. Zhu R-L, Lai M-J 2005. Cheilolejeunea decursiva (Sande Lac) R.M.Schust. and C. ventricosa (Schiffn.) X.L.He (Lejeuneaceae, Hepaticae) new to Australia. Journal of the Hattori Botanical Laboratory 98: 193203. Zhu R-L, So M-L, Wang Y-F 2002. The genus Cheilolejeunea (Hepaticae, Lejeuneaceae) in China. Nova Hedwigia 75: 387408.