PhytoKeys 20: 33–51, doi: 10.3897/phytokeys.20.4007
New and existing combinations in Palaeotropical Phlegmariurus (Lycopodiaceae) and lectotypification of the type species Phlegmariurus phlegmaria (L.) T.Sen & U.Sen.
Ashley R. Field 1,2, Peter D. Bostock 2
1 Australian Tropical Herbarium, James Cook University, Smithfield, Q 4878, Australia
2 Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens Mt Coot-tha, Toowong Q 4066, Australia

Corresponding author: Ashley R. Field (ashley.field@jcu.edu.au)

Academic editor: T. Ranker

received 19 September 2012 | accepted 29 January 2013 | Published 26 February 2013


(C) 2013 Ashley R. Field. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


For reference, use of the paginated PDF or printed version of this article is recommended.

Abstract

The genus Phlegmariurus Holubis recognised more widely than originally proposed and is circumscribed to include both Neotropic and Palaeotropic epiphytic and terrestrial species of Huperzioid Lycopodiaceae that have isotomous shoots, lack bulbils in their sporophyllous shoots and have spores with convex lateral margins and foveolate-fossulate sculpture restricted to their distal surfaces. New combinations with Phlegmariurus are proposed for 81 species and existing combinations identified for 33 species originating from the Palaeotropics. This installs a generic circumscription that is consistent between the Neotropics and Palaeotropics. A lectotype is designated for the type species of the genus, Phlegmariurus phlegmaria (L.) T.Sen & U.Sen

Keywords

Lycopodiaceae, Huperzia, Phlegmariurus, Lycopodium, Lycopodiophyta

Introduction

The Lycopodiaceae is a globally distributed family of homosporous Lycopodiophytes in which the species exhibit shoot-forms associated with their habitat. Early classifications of this family usually included all of the species, irrespective of their shoot-form, together in the large and polymorphic genus Lycopodium L. One exception was the small and morphologically divergent Australian and New Zealand pygmy-clubmoss Phylloglossum drummondii Kunzewhich was historically placed in a genus of its own. Multiple genera and subfamilies corresponding with the different shoot-forms, gametophyte and spore types were recognised following multi-character systematic studies on the family (Bruce 1976, Holub 1964, Øllgaard 1975, 1979, 1987, 1989a, Wagner and Beitel 1992, Wilce 1972). Until recently, the most widely adopted of these classifications was the four-genus classification of Øllgaard (1987) which recognised Lycopodium L., Lycopodiella Holub, Huperzia Bernh.and Phylloglossum Kunze and was supported by a comprehensive nomenclatural index (Øllgaard 1989a).

Recent molecular phylogenetic investigations of the family and new morphological observations present a hypothesis that the relationship between Phylloglossum drummondii and Huperzia is equivocal and that the genus Huperzia sensu Øllgaard (1987) is paraphyletic or at best weakly supported(Christenhuszet al. 2011, Jiet al. 2008, Whittier 2006, Whittier and Braggins 1992, 2000, Whittier and Storchova 2007, Wikström and Kenrick 1997, Wikström and Kenrick 2000, 2001, Wikström et al. 1999, Yatsentyuk et al. 2001). A monophyletic Huperzia Bernh. is obtained if Phylloglossum drummondii is included within it sensu Christenhusz et al. (2011). This classification produces a Huperzia sens. lat. that is morphologically heterogeneous and is not supported by synapomorphies that can be readily identified in the field. It also does not reflect the ancient divergence times between the Huperzioid clades of Lycopodiaceae (Wikström and Kenrick 2001).

An alternative classification is to circumscribe Huperzia Bernh. narrowly sensu Wagner and Beitel (1992) to include only the wholly terrestrial species with erect shoots bearing gemmae in sporophyll axils and spores that are triangular in polar view with concave lateral margins, truncate corners and foveolate-fossulate sculpture on both proximal and distal surfaces and to recognise a sister genus, Phlegmariurus Holub, that is broadened to include all Palaeotropical and Neotropical species with erect or pendulous sporophyllous shoots that lack gemmae and that have spores with convex lateral margins and foveolate-fossulate sculpture restricted to their distal surface. This circumscription of Phlegmariurus includes a broad spectrum of both epiphytic and non-epiphytic species primarily from the tropical and subtropical regions globally. The necessary generic combinations with Phlegmariurus Holub were recently proposed for the Neotropical species (Øllgaard 2012a, b) but many necessary combinations do not exist among the Palaeotropical species. We propose these transfers so that Phlegmariurus Holub may be considered monophyletic.

Species exhibiting the characteristic features of Phlegmariurus were identified from the examination of 121 type specimens (including images) for species previously combined with Lycopodium L., Huperzia Holub and Urostachys Herter and from characters described in these species protologues. Species level relationships within Palaeotropical Phlegmariurus are poorly understood and their circumscriptions require a thorough molecular and morphological review. It is apparent that there are more species names than there are species, and following a review of type materials we propose a number of new synonyms for species we have studied closely. Molecular evidence suggests that form homoplasy is common, especially among tropical rainforest epiphytes (Wikström et al. 1999). This suggests that morphologically similar species may not always be closely related. We have taken a more conservative approach to synonymisation for the Phlegmariurus phlegmaria (L.) T.Sen & U.Senspecies complex as we consider this to be a complex that is rich in species rather than being a remarkably heterogeneous species.

Lecotypification of Phlegmariurus phlegmaria (L.) T.Sen & U.Sen

Epiphytic Lycopodiaceae were first discovered byEuropean botanists in the East Indies in the late 1600s. Several species were described and illustrated in:


Breyne (1678) Exoticarum Plantarum Centuria Prima (tab. 92 n.v.)

Rheede et al.(1703) Hortus Indicus Malabaricus XII (p. 27, tab. 14!)

Rumph (1741) Herbarium Amboinense (VI p. 91, tab. 41, fig. 1!)

Dillen (1741) Historia Muscorum (p. 450 tab. 61 fig 5, A, B & C.!)

Dillen (1741) Historia Muscorum (p. 450 tab. 61 fig 5, D & E.!)

Linné (1743) Flora Zeylanica number 386 (P. Hermann herb. vol. 4. p. 5!)


Linné described Lycopodium phlegmaria in Species Plantarum in 1753 and cited Dillen (1741 p. 450 tab. 61 fig 5), Linné (1743 p. 183 no. 386 & P. Hermann specimen vol. 4 p. 5), Breyne (1678 tab. 92), and Rheede et al.(1703 p. 27, tab. 14) in his protologue. Based on the limited material available, Linné obviously regarded all of these illustrations, and the P. Hermann specimen, as belonging to the same species. His brief description could be applied to almost any heterophyllous epiphytic species in the genus Phlegmariurus. Subsequently, the figures and P. Hermann specimen cited by Linné have been recognised as representing multiple species. In accordance with ICBN (2006) Div. 2, Chapt. 2, Sect. 2 Art. 9.9 a lectotype may be selected to restrict the type of the taxon Lycopodium phlegmaria L.to one species (McNeill et al. 2006).

Øllgaard (1989) indicated that Dillen tab. 61 fig 5 A, B & C was eligible for selection as lectotype but did not designate it. The illustrations of Dillen tab. 61 fig 5 ABCDE are based on two separate illustrations reproduced from earlier publications. Dillen tab. 61 fig. 5. A, B & C is reproduced from Rheede et al. (1703 XII p. 27, tab. 14) but Dillen tab. 61 fig. 5. D & E is reproduced from Breyne (1678 tab. 92). In 1810, Willdenow described Lycopodium mirabilis (Willd.) Holub for Dillen tab. 61 fig 5 D & E (i.e. the Breyne illustration) but retained the referenceto Dillen tab. 61 fig 5 A, B & C (i.e. the Rheede et al. illustration) for Lycopodium phlegmaria L. even though the epithet phlegmaria was introduced by Breyne (1678). In 1832, Lycopodium phyllanthum Hook. & Arn. was describedfrom new material (Pacific origin) which is in our opinion conspecific with the P. Hermann specimen (Indo-Asian origin). In the interest of conserving nomenclatural stability we believe that Dillen fig. 5, A, B & C. (1741) is the most suitable lectotype because it exhibits characters present in the diagnosis (in accordance with ICBN 2006 Div. 2, Chap. 2, Sect. 2, Art. 9.12.) and most closely accords with Lycopodium phlegmaria as it is regarded today (in accordance with ICBN 2006 Div. 2, Chapt. 2, Sect. 2 Art. 9 Recommendation 9.A.5.y) (McNeill et al. 2006).

Taxonomy

Phlegmariurus afromontanus (Pic.Serm.) A.R.Field & Bostock comb. nov. Basionym: Huperzia afromontana Pic.Serm., Webbia27:394. 1972. Type: Burundi. J.Lewalle 3468 (holo: FI! [FI3002]). IPNI ID: 77125024-1


Phlegmariurus aloifolius (Hook. & Grev.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium aloifolium Hook. & Grev., Icon. Filic.2:233. 1831. Synonym: Huperzia aloifolia (Hook. & Grev.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Type: India. N.Wallich 129 (holo: K). IPNI ID: 77125026-1


Phlegmariurus balansae (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium balansae Herter, Bot. Jahrb. Syst. 43: Beibl. 98:51. 1909. Synonym: Huperzia balansae (Herter) Holub, Geobot. Phytotax. 20(1): 71. 1985. Type: New Caledonia M.Balansa 688 (syn: P! [P523007, P523008, P523009]); New Caledonia. Pompéry s.n. (syn: P! [P523010]). IPNI ID: 77125028-1


Phlegmariurus bampsianus (Pic.Serm.) A.R.Field & Bostock comb. nov. Basionym: Huperzia bampsiana Pic.Serm., Bull. Jard. Bot. Natl. Belg. 55(1-2): 193. 1985. Type: Rwanda. J.P.Bamps 2964 (holo: LG; iso: P! [P462093], FI! [FI3007]). IPNI ID: 77125029-1


Phlegmariurus banayanicus (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys banayanicus Herter, Bot. Arch. 3:17. 1923. Synonym: Huperzia banayanica (Herter) Holub, Geobot. Phytotax. 20(1): 71. 1985.Type: Philippines. H.M.Curran 8003 (syn: P! [P5239029]); Philippines. E.B.Copeland 1450 (syn: P! [P523926]); Philippines. A.D.Elmer 9498 (syn: P! [P523019]); Philippines. F.W.Foxworthy 2449 (syn: P! [P523018]). IPNI ID: 77125031-1


Phlegmariurus bolanicus (Rosenst.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium bolanicum Rosenst., Repert. Spec. Nov. Regni Veg. 12:181 1913. Synonym: Huperzia bolanica (Rosenst.) Holub, Geobot. Phytotax. 20(1): 71. 1985. Type: Papua New Guinea. C.Keysser B32. Heterotypic synonyms: Urostachys kaysseri Herter ex Nessel, Revista Sudamer. Bot. 6:156. 1940; Huperzia kaysseri (Herter ex Nessel) Holub Folia Geobot. Phytotax. 20(1): 74. 1985. Type: New Guinea. C.Keysser s.n. (holo: B [B112944]). IPNI ID: 77125033-1


Phlegmariurus brachystachys (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium dacrydioides Baker. var. brachystachys Baker, Handb. Fern-Allies 18. 1887. Synonym: Huperzia brachystachys (Baker) Pic.Serm., Webbia 23: 162. 1968. Type: Cameroon. G.Mann 2041(holo: K!). IPNI ID: 77125034-1


Phlegmariurus brassii (Copel.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium brassii Copel., J. Arnold Arbor. 10:174 1929. Synonym: Huperzia brassii (Copel.) Holub, Geobot. Phytotax. 20(1): 71. 1985. Type: Papua New Guinea. L.J.Brass 1521 (holo: UC! [UC377989], iso: BRI! [AQ521250]). IPNI ID: 77125036-1


Phlegmariurus cancellatus (Spring) Ching, Acta Bot. Yunn. 4:122.1982. Basionym: Lycopodium cancellatum Spring, Mém. Acad. Roy. Sci. Belg. 24: 27. 1849. Type: Bhutan. W.Griffith 162 (holo: K).


Phlegmariurus carinatus (Desv. ex Poir.) Ching, Acta Bot. Yunn.4:120. 1982.Basionym: Lycopodium carinatum Desv. ex Poir., Encycl. suppl. 3:555 1813. Synonyms: Huperzia carinata (Desv. ex Poir.) Trevis., Atti Soc. Ital. Sci. Nat. 17(2) 1874. Type: Sri Lanka. Jussieu 639 (holo: P!, iso: B!). Heterotypic synonyms: Lycopodium acrostachyum Hook. & Grev., Icon Filic. 2:t 181 (1831). Type: Singapore, N.Wallich 117 (holo: P!). Lycopodium albescens Bailey, Queensl. Dept. Agric. Brisbane Bot. Bull. 14:16. 1896; Huperzia albescens (F.M.Bailey) Holub, Geobot. Phytotax. 20(1): 70. 1985. Type: Australia ex New Guinea. M.Sutter ex Capt. Michael s.n. (holo: BRI! [AQ258306], iso: K). Lycopodium laxum C.Presl, Reliq. Haenk. 1:83. 1825; Huperzia laxa (C.Presl) Trevis., Atti Soc. Ital. Sci. Nat. 17: 247. 1874; Phlegmariurus laxus (C.Presl.) Satou, Hikobia 12(3):268. 1997. Type: Philippines. Haenke s.n. (iso: K). Lycopodium pendulum Roxb.; Calcutta J. Nat. Hist. 4. 472 (1844). Type: C. Smith s.n. from Amboina (lecto: BR fide Morton (1974) Contr. U.S. Natl. Herb. 38: 283-396.). Lycopodium struthioloides C.Presl, Reliq. Haenk. 1: 82. 1830; Huperzia struthioloides (C.Presl) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 61. 1944. Type: Philippines. Haenke s.n. (holo: PRC). Lycopodium vanikorense Copel., J. Arnold. Arbor. 12:49. 1931; Huperzia vanikorensis (Copel.) Holub, Folia Geobot. Phytotax. 20:77. 1985. Type: Vanikoro [Vanuatu]. S.F.Kajewski 521 (holo: UC! [UC422661], iso: GH, K, MICH, MO! [MO22133], US).


Phlegmariurus cavifolius (C.Chr.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium cavifolium C.Chr., Dansk Bot. Ark. 7:190. 1932. Synonym: Huperzia cavifolia (C.Chr.) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. H.Perrier 8273 (holo: BM, iso: P! [P46639], K! [K351220]). Urostachys smithianus Nessel, Repert. Spec. Nov. Regni Veg. 36:185. 1934; Huperzia smithiana (Nessel) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: Mauritius: s.c. s.n. (syn: BONN-Nessel). IPNI ID: 77125038-1


Phlegmariurus coralium (Spring) A.R.Field & Bostock comb. nov. Basionym: Lycopodium coralium Spring in Miquel, Pl. Jungh. 3: 273. 1854. Synonym: Huperzia coralium (Spring) Holub, Folia Geobot. Phytotax. 20(1): 71. 1985. Type: Java [Indonesia]. Junghuhn 291 (holo: LG, iso: P). IPNI ID: 77125040-1


Phlegmariurus creber (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium crebrum Alderw., Bull. Jard. Bot. Buitenzorg ser. 2, 24:4. 1917. Synonym: Huperzia crebra (Alderw.) Holub, Geobot. Phytotax. 20(1) 71. 1985. Type: New Guinea. A.A.Pulle 907 (syn: BM, BO, BONN, K, L, U). IPNI ID: 77125042-1


Phlegmariurus cryptomerinus (Maxim.) Satou Hikobia 12 (3): 268. 1997 Basionym: Lycopodium cryptomerinum Maxim., Bull. Acad. Sci. St.Petersb. 15: 231. 1870. Synonym: Huperzia cryptomerina (Maxim) R.D.Dixit, J. Bombay Nat. Hist. Soc. 77(3): 541. 1980 [1981]. Type: Japan. Maximowicz s.n.


Phlegmariurus cunninghamioides (Hayata) Ching, Acta Bot. Yunnan.4(2): 120. 1982. Basionym: Lycopodium cunninghamioides Hayata, Ic. pl. formos. 4: 131. 1914. Synonym: Huperzia cunninghamioides (Hayata) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: Taiwan. T.Soma 1 (holo: TI).


Phlegmariurus curiosus (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys curiosus Herter, Notul. Syst. 15:355. 1958. Synonym: Huperzia curiosa (Herter) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. H.Humbert 23478 (holo: P! [P46637]). IPNI ID: 77125044-1


Phlegmariurus dacrydioides (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium dacrydioides Baker, Handb. fern-allies 17-18. 1887. Synonyms: Huperzia dacrydioides (Baker) Pic.Serm., Webbia 23: 162. 1968. 1968. Type: Natal. J.Buchanan s.n. (lecto: K! [K351207] designated by Pichi Sermolli 1986: 7). IPNI ID: 77125045-1


Phlegmariurus dalhousieanus (Spring) A.R.Field & Bostock comb. nov. Basionym: Lycopodium dalhousieanum Spring, Mem. Acad. Roy. Belg. 24: 25. 1849 Synonym: Huperzia dalhousieana (Spring) Trevis., Atti Soc. Ital. Sci. Nat. 17(2) 1874. Type: Penang [Malaysia]. Lady Dalhousie 36 (holo: K! [3960/59]). Heterotypic synonyms: Based on a morphological comparison of living and herbarium specimens from Indonesia, Malaysia, Papua New Guinea, Australia and Fiji we consider the following taxa to belong to the same widespread but extremely scarce species: Lycopodium caudifolium Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 1:14. 1911; Urostachys caudifolius (Alderw.) Herter ex Nessel, Bärlappgewächse 185. 1939; Huperzia caudifolia (Alderw.) Holub, Folia Geobot. Phytotax. 20(1): 71. 1985. Type: Borneo [Malaysia]. A.W.Niewenhuis 317 (holo: BO! [BO185337], iso: L). Lycopodium clarae Bailey, Queensl. Dept. Agric. Brisbane Bot. Bull. 7. 1893; Urostachys clarae (F.M.Bailey) Herter ex Nessel, Bärlappgewächse 185. 1939; Huperzia clarae (F.M.Bailey) Holub, Geobot. Phytotax. 20(1): 71. 1985. Type: Australia. E.Cowley 29 (holo: BRI! [AQ24829]). Lycopodium glaucum Cesati, Ann. Acc. Sc. Fis. et. Mat. Napoli VII, 35 (1876); Urostachys glaucus (Cesati) Herter, Revista Sudamer. Bot. 8: 86 (1949); Type: Malaysia. O.Beccari (holo: FI! [FI11821]). Lycopodium magnificum Brownlie, Nova Hedwigia Beih., 55 (Pterid. Fl. Fiji) 22. 1977; Huperzia magnifica (Brownlie) Holub, Folia Geobot. Phytotax. 26(1): 92. 1991. Type: Fiji. Fiji Dept. Agr. 18070 (holo: CHR! [CHR341159]). Lycopodium ingens Gepp ex Nessel, Bärlappgewächse 185 (1939) nom. nud.; Type: Sogori, New Guinea. H.O.Forbes s.n. (holo?: BM! [BM1038045]). IPNI ID: 77125047-1


Phlegmariurus delbrueckii (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys delbrueckii Herter, Bot. Arch. 3:19. 1923. Synonyms: Huperzia delbrueckii (Herter) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: Philippines. E.A.Mearns & W.J.Hutchinson 4654 (holo: NY! [NY127344], iso: MICH! [MICH1191095], P! [P01220906, P01220907], US! [US134352]). IPNI ID: 77125049-1


Phlegmariurus dielsii (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium dielsii Herter, Bot. Jahrb. Syst. 54:229. 1916. Synonym: Huperzia dielsii (Herter) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: New Guinea. Ledermann 11318 (syn: B, BM); R.Schlechter 19085 (syn: B! [B2112382] BONN-Nessel). IPNI ID: 77125051-1


Phlegmariurus durus (Pic.Serm.) A.R.Field & Bostock comb. nov. Basionym: Huperzia dura Pic.Serm., Webbia 27: 392. 1973. Type: Kenya. R.B.Faden et al. 70/180 (holo: FI! [FI3008], iso: EA! [EA2516, EA2517], iso: K! [K351206]). IPNI ID: 77125052-1


Phlegmariurus ellenbeckii (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys ellenbeckii Nessel, Repert. Spec. Nov. Regni Veg. 36:188, t. 175. 1934. Synonym: Huperzia ellenbeckii (Nessel) Pic.Serm., Webbia 23: 162. 1968. Type: Ethiopia. Ellenbeck s.n. (lecto: BONN-Nessel 544 p.p. designated by Pichi Sermolli 1968: 162). IPNI ID: 77125054-1


Phlegmariurus elmeri (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys elmeri Herter, Bot. Arch. 3:16. 1923. Synonym: Huperzia elmeri (Herter) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: Philippines. A.D.Elmer 6411 (isosyn: P! [P01220901]); J.Bermejos 372 (isosyn: P! [P01220903]); H.M.Curran 3906 (isosyn: P! [P01220905]); E.D.Merrill 5923 (isosyn: P! [P01220902]). IPNI ID: 77125056-1


Phlegmariurus fargesii (Herter) Ching, Acta Bot. Yunn. 4:125. 1982. Basionym: Lycopodium fargesii Herter, Bot. Jahrb. Syst. 43: Beibl. 98: 48. 1909. Synonyms: Huperzia fargesii (Herter) Holub, Folia Geobot. Phytotax. 20(1): 72. 1985; Urostachys fargesii (Herter) Herter ex Nessel, Bärlappgewächse 137. 1939. Type: China. P.G.Farges s.n. (holo: P! [P01234387], iso: MO! [MO255690]).


Phlegmariurus filiformis (Sw.) W.H.Wagner, Contr. Univ. Michigan Herb.20:242. 1995 Basionym: Lycopodium filiforme Sw., J. Bot. (Schrader) 2: 114. 1800 [1801]; Synonyms: Urostachys filiformis (Sw.) Herter, Index. Lyc. 61. 1949; Huperzia filiformis (Sw.) Holub, Folia Geobot. Phytotax 20:72 (1985). Type: “Insulae Sandwich oceani pacific" [Tahiti] in Syn. Fil. 1806. Heterotypic synonyms: Lycopodium setaceum Lam., Encycl. (Lamarck) 3(2): 653. 1792. Huperzia setacea (Hamilt. ex D. Don) Trevis., Atti Soc. Ital. Sci. Nat. 17:248. 1874. Type: Réunion. Commerson s.n. (holo: P! [P46612], iso: P! [P583966]). Lycopodium polytrichoides Kaulf., Enum. Filic. 6. 1824; Huperzia polytrichoides (Kaulf.) Trevis., Atti Soc. Ital. Sci. Nat. 17(2) 1874. Type: Sandwich Islands [Hawaii], Chamisso s.n. (lecto: B designated by Nessel 1939: 30). Lycopodium verticillatum L.f., . Suppl. pl. 448. 1782; Huperzia verticillata (L.f.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54: 60. 1944. Type: Réunion. P.Sonnerat per A.Thouin (holo: SBT! [SBT12294]).


Phlegmariurus flagellaceus (Kuhn) A.R.Field & Bostock comb. nov. Basionym: Lycopodium flagellaceum Kuhn, Forschungsr. Gazelle 4:15. 1889. Synonym: Huperzia flagellacea (Kuhn) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: New Guinea. Nauman s.n. (holo: B! [B112446], iso: BM). Note: Some authors have considered this taxon to share affinities with Phlegmariurus varius but we consider that it is closer to Phlegmariurus brassii (Copel.) A.R.Field & Bostock and the relationship between the two needs investigating. IPNI ID: 77125058-1


Phlegmariurus foliosus (Copel.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium foliosum Copel., Bernice P. Bishop Mus. Bull. 59:7, t.1. 1929. Synonym: Huperzia foliosa (Copel.) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: H.E.Parks 20651(holo: UC! [UC351029], iso: BISH! [BISH1000366]). IPNI ID: 77125060-1


Phlegmariurus fordii (Baker) Ching, Acta Bot. Yunn.4:126. 1982. Basionym: Lycopodium fordii Baker, Handb. Fern-Allies 17. 1887. Synonym: Huperzia fordii (Baker) R.D.Dixit Census Indian Pteridophytes 7. 1984. Type: China. Ford 4 (holo: K).


Phlegmariurus gagnepainianus (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys gagnepainianus Herter, Revista Sudamer. Bot. 8:23. 1949. Synonym: Huperzia gagnepainiana (Herter) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. H.Perrier 15405 (lecto: P! [P226738] designated by Tardieu (1971: 20). IPNI ID: 77125062-1


Phlegmariurus giganteus (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys giganteus Herter ex Nessel, Bärlappgewächse 124. 1939. Synonym: Huperzia gigantea (Herter) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: Philippines. E.A.Mearns & Hutchinson 1277 (iso: ?); New Guinea. MacGregor 3945 (iso: ?). IPNI ID: 77125064-1


Phlegmariurus gnidioides (L.f.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium gnidioides L.f., Suppl. pl. 448. 1782. Synonym: Huperzia gnidioides (L.f) Trevis., Atti Soc. Ital. Sci. Nat. 17:247. 1874. Type: Mauritius. P.Sonnerat per A.Thouin s.n. (holo: P). Heterotypic synonyms: Urostachys helmii Nessel, Repert. Spec. Nov. Regni Veg. 36:186. 1934; Huperzia helmii (Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: “Insel Greymouth 1885" (lecto: BONN-Nessel designated Øllgaard 1989: 92). IPNI ID: 77125065-1


Phlegmariurus goebelii (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys goebelii Nessel, Repert. Spec. Nov. Regni Veg. 36:188, t.175. 1934. Synonym: Huperzia goebelii (Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: West-Sumatra [Indonesia] Goebel s.n. (isosyn: UC! [UC413367], BONN-Nessel!); Kwala Relai [Malaysia]. M.Haniff & M.Nur 10225 (isosyn: BM! [BM1038044]). Heterotypic synonyms: Urostachys heroldii Herter ex Nessel, Revista Sudamer. Bot. 6:167. 1940. Synonym: Huperzia heroldii (Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: Borneo [Malaysia]. Hallier (holo: P?). IPNI ID: 77125067-1


Phlegmariurus guandongensis Ching, Acta Bot. Yunnan.4(2):123. 1982. Type: China. S.P.Ko 51291 (holo: PE).


Phlegmariurus gunturensis (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium gunturense Alderw., Bull. Jard. Bot. Buitenzorg ser. 2, 1:14. 1911. Synonym: Huperzia gunturensis (Alderw.) Holub, Folia Geobot. Phytotax. 20(1): 73. 1985. Type: Indonesia. A.E.Kerkhoven 15 (holo: BO [BO1826257]). IPNI ID: 77125069-1


Phlegmariurus hamiltonii (Spreng.) Á.Löve & D.Löve, Taxon26(2-3):324. 1977. Basionym: Lycopodium hamiltonii Spreng., Syst. veg. 5: 429. 1828. Type: Nepal. J.Buchanan s.n. (holo: BM).


Phlegmariurus harmsii (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys harmsii Herter ex Nessel, Revista Sudamer. Bot. 6: 166. 1940. Synonym: Huperzia harmsii (Herter ex Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: Caroline Islands [Micronesia] R.Kanehira 1316 (iso: NY! [NY127353], US [US1917849]). IPNI ID: 77125071-1


Phlegmariurus hellwigii (Warb.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium hellwigii Warb., Monsunia, 1:97. 1898. Synonym: Huperzia hellwigii (O.Warb.) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: Papua New Guinea. F.Hellwig 349 (holo: B! [B112544], iso: BONN-Nessel). IPNI ID: 77125073-1


Phlegmariurus henryi (Baker) Ching, Acta Bot. Yunnan.4(2):125 1982. Basionym: Lycopodium henryi Baker, Kew Bull. 1906:15. 1906. Type: China. Henry 11551 (holo: K).


Phlegmariurus hillianus (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys hillianus Nessel, Repert. Spec. Nov. Regni Veg. 36:16. 1934. Synonym: Huperzia hilliana (Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: India. H.C.Levinge s.n. (holo: K, iso: BONN-Nessel). IPNI ID: 77125075-1


Phlegmariurus holstii (Hieron.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium holstii Hieron. in Engler, Pflanzenw. Ost-Afrikas C:90. 1985. Synonyms: Huperzia holstii (Hieron.) Pic.Serm., Webbia 23: 163. 1968. Type: Usambara [Tanzania]. C.Holst 8814 (holo: B! [B112586], iso: K! [K351205], P! [P466630], BM! [BM785226]). IPNI ID: 77125077-1


Phlegmariurus horizontalis (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys horizontalis Nessel, Bärlappgewächse 225. 1939. Synonym: Huperzia horizontalis (Herter ex Nessel) Holub, Geobot. Phytotax. 20(1): 73. 1985. Type: Borneo [Malaysia]. A.W.Niewenhuis 311 (iso: L! [L57351], BONN-Nessel). IPNI ID: 77125079-1


Phlegmariurus humbertii (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys humbertii Nessel, Repert. Sp. Nov. Regni Veg. 48:170. 1940. Type: Madagascar. J.Kiese s.n. (holo: BONN-Nessel). IPNI ID: 77125546-1


Phlegmariurus humbertii-henrici (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys humbertii-henrici Herter, Revista Sudamer. Bot. 8:23. 1949. Synonym: Huperzia humbertii-henrici (Herter) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. H.Humbert 12224 (isolecto: P! [P466628, P466627, P466629] designated by Tardieu 1971: 29). IPNI ID: 77125081-1


Phlegmariurus jaegeri (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys jaegeri Herter, Revista Sudamer. Bot. 8:24. 1949. Synonym: Huperzia jaegeri (Herter) Pic.Serm., Webbia 23: 163. 1968. Type: Sierra Leone. Jaeger 1522 (holo: P! [P466622]). IPNI ID: 77125083-1


Phlegmariurus juniperistachyus (Hayata) A.R.Field & Bostock comb. nov. Basionym: Lycopodium juniperistachyum Hayata, Ic. Pl. Formos. 4:132. 1914. Synonym: Huperzia juniperistachya (Hayata) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: Taiwan. Kawakami & Nakahara 787 (holo: TI). IPNI ID: 77125085-1


Phlegmariurus lauterbachii (E.Pritz. ex Schumann & Lauterb.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium lauterbachii E.Pritz. ex Schumann & Lauterb., Südsee 149. 1900. Synonym: Huperzia lauterbachii (E.Pritz.) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: New Guinea. C.A.G.Lauterbach 3174 (holo: B, iso: BONN-Nessel). IPNI ID: 77125087-1


Phlegmariurus lecomteanus (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys lecomteanus Nessel, Repert Spec. Nov. Regni Veg. 36(12-15):187. 1934. Synonyms: Huperzia lecomteana (Nessel) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: Madagascar. (holo: BONN-Nessel 492). IPNI ID: 77125089-1


Phlegmariurus ledermannii (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium ledermannii Herter, Bot. Jahrb. Syst. 54:232. 1916. Synonym: Huperzia ledermannii (Herter) Holub, Geobot. Phytotax. 20(1): 74. 198. Type: New Guinea. C.Keysser 199 (iso: SUNIV!, B! [B113004]). IPNI ID: 77125091-1


Phlegmariurus lockyeri (D.L.Jones & B.Gray ), A.R.Field & Bostock comb. nov. Basionym: Lycopodium lockyeri D.L.Jones & B. Gray, Austrobaileya 2:126. 1985. Synonym: Huperzia lockyeri (D.L.Jones & B.Gray) Holub, Geobot. Phytotax. 26(1): 92. 1991. Type: Australia B.Gray 3541 (holo: QRS! [QRS77021]) IPNI ID: 77125092-1


Phlegmariurus longus (Copel.), A.R.Field & Bostock comb. nov. Basionym: Lycopodium longum Copel., Philipp. J. Sci. 60:100, t.3. 1936. Type: Solomon Islands. S.F.Kajewski 1953 (holo: PHN destroyed, iso: BM! [BM16725], MICH! [MICH1287171]). IPNI ID: 77125094-1


Phlegmariurus lonyangensis C.Y.Ma, Bull. Bot. Res., Harbin 10(3):58. 1990. Type: China C.Y.Ma 3246 (holo: AMMS).


Phlegmariurus macgregorii (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium macgregorii Baker, J. Bot. 28: 109. 1890. Synonym: Huperzia macgregorii (Baker) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: New Guinea. W.MacGregor 68 (holo: K!, iso: BM! [BM1038027, BM1038028]). Heterotypic synonyms: Lycopodium versteegii Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 24:4. 1917; Huperzia versteegii (Alderw.) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: (holo: New Guinea. G.M.Vertseeg 2510 (holo: U! [U7449]). Urostachys whartoniensis Nessel, Repert. Spec. Nov. Regni Veg. 38:62. 1935. Huperzia whartoniensis (Nessel) Holub, Geobot. Phytotax. 20(1): 78. 1985. Type: Papua New Guinea. L.J.Brass 4729 (iso: NY! [NY127367], iso: BRI! [AQ24833]). IPNI ID: 77125096-1


Phlegmariurus macrostachys (Hook. ex Spring) N.C.Nair & S.R.Ghosh, J. Econ. Taxon.Bot.12(1):194. 1988. Synonyms: Huperzia macrostachys (Hook. ex Spring) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: India. F.Adams s.n. (syn: K). Sri Lanka. Walker s.n. (syn: K).


Phlegmariurus mannii (Hillebr.) W.H.Wagner, Contr. Univ. Michigan Herb. 20:241 1995. Basionym: Lycopodium phlegmaria var. mannii Hillebr., Fl. Hawaiian Isl. 645. 1888. Type: Hawaii. H.Mann & W.T.Brigham 506 (iso: BISH! [BISH1005485], MO! [MO22681]).


Phlegmariurus marsupiiformis (D.L.Jones & B.Gray) A.R.Field & Bostock comb. nov. Basionym: Lycopodium marsupiiforme D.L.Jones & B.Gray, Austrobaileya 2:128. 1985. Synonym: Huperzia marsupiiformis (D.L.Jones & B.Gray) Holub, Geobot. Phytotax. 26(1): 92. 1991. Type: Australia. B.Gray 3841 (holo: BRI! [AQ419317], CNS! [QRS78343]). IPNI ID: 77125098-1


Phlegmariurus megastachyus (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium megastachyum Baker, J. Linn. Soc. Bot. 21:454. 1885. Synonym: Huperzia megastachya (Baker) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. R.Baron 2840 (holo: K! [K351218, K351219]). IPNI ID: 77125100-1


Phlegmariurus merrillii (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys merrillii Herter, Bot. Arch. 3:15. 1923. Synonym: Huperzia merrillii (Herter) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: Philippines. E.D.Merrill 965 (iso: UC! [UC1096474], U! [U7444], MICH! [MICH1191096]). IPNI ID: 77125103-1


Phlegmariurus milbraedii (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium milbraedii Herter, Hedwigia 49:90. 1909. Synonym: Huperzia milbraedii (Herter) Pic.Serm., Webbia 23:163. 1968. Type: Cameroon. Milbraed 3449 (holo: B). IPNI ID: 77125104-1


Phlegmariurus mingcheensis Ching, Acta Bot. Yunnan.4(2): 125. 1982. Synonym: Huperzia mingcheensis (Ching) Holub, Folia Geobot. Phytotax. 20: 74. 1985. Type: China. P.S.Chiou 2069 (holo: PE). See recent note on the nomenclature of this species (Zhang 2012).


Phlegmariurus minutifolius (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium coralium var. minutifolium Alderw., Malayan fern allies 44. 1915. Synonyms: Huperzia minutifolia (Alderw.) Holub, Folia Geobot. Phytotax 20(1): 74. 1985. Type: New Guinea. Ruchmat s.n. (holo: L). IPNI ID: 77125106-1


Phlegmariurus multifarius (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium multifarium Alderw., Bull. Jard. Bot. Buitenzorg, ser. 3, 5:226. 1922. Synonym: Huperzia multifaria (Alderw.) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Ternate [Moluccas, Indonesia]. V.M.A.Beguin 1329 (holo: BO [BO186327]). IPNI ID: 77125108-1


Phlegmariurus myrtifolius (G.Forst.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium myrtifolium G.Forst., Florae Insulae Australiae Prodromus. 1786. Synonym: Huperzia myrtifolia (G.Forst.) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: New Caledonia. G.Forster s.n. (iso: P-Luerss., UPS-Thunb). IPNI ID: 77125109-1


Phlegmariurus nanus C.Y.Ma, Bull. Bot. Res., Harbin 10(3):58. 1990. Type: China. C.Y.Ma 3274 (holo: AMMS).


Phlegmariurus neocaledonicus (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys neocaledonicus Nessel, Repert. Spec. Nov. Regni Veg. 36:187. 1934. Synonym: Huperzia neocaledonica (Nessel) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: New Caledonia. E.Viellard (lecto: BONN-Nessel designated Øllgaard 1989: 96). IPNI ID: 77125110-1


Phlegmariurus nilagiricus (Spring) A.R.Field & Bostock comb. nov. Basionym: Lycopodium nilagiricum Spring, Bull. Acad. Roy. Sci. Belg. 8:517. 1841. Synonym: Huperzia nilagirica (Spring) Dixit, J. Bombay Nat. Hist. Soc. 77(3):541. 1981. Type: India. Perottet (holo: G-Deless.). IPNI ID: 77125112-1


Phlegmariurus nummulariifolius (Blume) Ching, Acta Bot. Yunnan.4(2):125. 1982. Basionym: Lycopodium nummulariifolium Blume, Enum. pl. Javae 2: 263. 1828. Synonym: Huperzia nummulariifolia (Blume) Chambers, Jermy & Crabbe, Brit. Fern Gaz. 10: 176. 1971. Type: Java. C.L.Blume s.n. (isolecto: L! [L57363]).


Phlegmariurus nutans (Brack.) W.H.Wagner Contr. Univ. Michigan Herb.20:241. 1995. Synonym: Huperzia nutans (Brack.) Rothm., Feddes Repert. 54: 62. 1944. Type: Hawaii. Wilkes 10 (holo: US, iso: BISH! [BISH1005484]).


Phlegmariurus nylamensis (Ching & S.K.Wu) H.S.Kung & Li Bing Zhang, Acta Phytotax. Sin.37(1):52. 1999.


Phlegmariurus obtusifolius (P.Beauv.) A.R.Field & Bostock comb. nov. Basionym: Lepidotis obtusifolia P.Beauv., Prodr. Aethéogam. 109. 1805. Synonym: Huperzia obtusifolia (P.Beauv.) Rothm., Feddes Repert. 54:61. 1944. Type: Réunion. J.B.G.M. Bory de St. Vincent (holo: P! [P466623], iso: B-Wild.! [B-W19344]). Heterotypic synonyms: Lycopodium pachyphyllum Kuhn ex Herter, Bot. Jahrb. Syst. 43: Beibl. 98:51. 1909; Huperzia pachyphylla (Kuhn) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Madagascar. J.M.Hildebrandt 4141(holo: B! [B85521], iso: B! [B8522], P! [P466624, P466625, P46626]). IPNI ID: 77125167-1


Phlegmariurus oceanianus (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium oceanianum Herter, Bot. Jahrb. Syst. 43: Beibl. 98:52. 1909. Synonym: Huperzia oceaniana (Herter) Holub, Folia Geobot. Phytotax. 20:75.1985. Type: New Hebrides [Vanuatu] s.c. s.n. (syn: P). IPNI ID: 77125114-1


Phlegmariurus oltmannsii (Herter ex Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys oltmannsii Herter ex Nessel [Bärlappgewächse 231. 1939, nom. inval.] Revista Sudamer. Bot. 6: 168. 1940. Synonym: Huperzia oltmannsii (Herter ex Nessel) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Tahiti. J.McGillivray 1331. IPNI ID: 77125115-1


Phlegmariurus ophioglossoides (Lam.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium ophioglossoides Lam., Encycl. (Lamarck) 3:646. 1789. Synonym: Huperzia ophioglossoides (Lam.) Rothm., Feddes Repert. 54:62. 1944. Type: Mauritius. Commerson s.n. (holo: P-Lam). Note: Molecular phylogenetic evidence indicates that this Palaeotropic species belongs to a Neotropical radiation of Phlegmariurus. IPNI ID: 77125118-1


Phlegmariurus ovatifolius (Ching) W.M.Chu ex H.S.Kung & Li Bing Zhang, Acta Phytotax. Sin.37(1):52 (1999). Basionym: Huperzia ovatifolia Ching, Acta Bot. Yunnanica 3: 298. 1981. Type: China. Yunnan Complex Exped. 1504 (holo: PE).


Phlegmariurus parksii (Copel.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium parksii Copel., Bernice P. Bishop. Mus. Bull. 59:8, t.2. 1929. Synonym: Huperzia parksii (Copel.) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Fiji. H.E.Parks 20584 (holo: UC! [UC351026], iso: P! [P711367], BISH! [BISH1000370], US! [US1759872]). Heterotypic synonym: Urostachys bonapartei Nessel, Repert. Spec. Nov. Regni Veg.. 36:182 t.172 1934; Huperzia bonapartei (Nessel) Holub, Geobot. Phytotax. 20(1): 71. 1985. Type: Flora Vitiensis [Fiji] C.Weber s.n. (syn: BONN). IPNI ID: 77125120-1


Phlegmariurus patentissimus (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium patentissimum Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 24:4. 1917. Synonym: Huperzia patentissima (Alderw.) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Irian Jaya [Indonesia]. A.A.Pulle 462 (holo: BO, iso: BM, BONN-Nessel, L! [L57365], U). IPNI ID: 77125121-1


Phlegmariurus pecten (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium pecten Baker, J. Linn. Soc. Bot. 15:421. 1876. Synonym: Huperzia pecten (Baker) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. W.Pool s.n. (holo: K1 [K351214]). IPNI ID: 77125123-1


Phlegmariurus perrerianus (Tardieu) A.R.Field & Bostock comb. nov. Basionym: Huperzia perreriana Tardieu, Adansonia 10:20. 1970. Type: Madagascar. H.Perrier 8294 (holo: P). IPNI ID: 77125127-1


Phlegmariurus petiolatus (C.B.Clarke) H.S.Kung & Li Bing Zhang, Acta Phytotax. Sin.37(1): 52. 1999. Basionym: Lycopodium hamiltonii var. petiolatus C.B.Clarke, Trans. Linn. Soc. London, ser. 2, Bot. 1: 590. 1880. Synonyms: Huperzia petiolata (C.B.Clarke) R.D.Dixit, J. Bombay Nat. Hist. Soc. 77(3): 541. 1980 [1981]. Type: Khasia [Bangladesh]. C.B.Clarke s.n. (holo: BM).


Phlegmariurus phlegmaria (L.) T.Sen & U.Sen, Fern. Gaz.11(6):421. 1978. Basionym: Lycopodium phlegmaria L., Species Plantarum 2. 1753. Synonym: Huperzia phlegmaria (L.) Rothm., Feddes Repert. 54: 62. 1944. Type: India. s.c. s.n. (lecto: Dillen (1741) Historia Muscorum p. 450 tab. 61 fig 5, A, B & C.! here designated).


Phlegmariurus phlegmarioides (Gaudich.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium phlegmarioides Gaudich., Voy. Uranie, Bot. 1830. Synonym: Huperzia phlegmarioides (Gaudich.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54(1). 1944. Type: Moluccas [Indonesia] C.-B.Gaudichaud s.n. (holo: P!, iso: K!). Heterotypic synonyms: Urostachys kandavuensis Nessel, Repert. Spec. Nov. Regni Veg. 39:67. 1935; Huperzia kandavuensis (Nessel) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: Fiji. A.C.Smith 142 (holo: NY! [NY127357]). Lycopodium pseudophlegmaria Kuhn, Forschungsr. Gazelle 4: 16. 1889; Urostachys pseudophlegmaria (Kuhn) Herter ex Nessel, Bärlappgewächse 237. 1939; Huperzia pseudophlegmaria (Kuhn) Holub, Folia Geobot. Phytotax. 20: 76. 1985; Type: Naumann, Fiji Islands (holo - unknown). Lycopodium tetrapterygium F.M.Bailey, Proc. Roy. Soc. Queensland 1:150 (1885); Urostachys tetrapterygius (F.M.Bailey) Herter, Index Lyc. 86. 1949; Type: Daintree River, Australia Fitzalan s.n. (syn: BRI!) and Johnstone River, Australia Kefford s.n. (syn: BRI!). IPNI ID: 77125129-1


Phlegmariurus phyllanthus R.D.Dixit J. Bombay Nat. Hist. Soc. 77(3): 541. 1981. Synonyms: Huperzia phyllantha (Hook. & Arn.) Holub, Geobot. Phytotax. 20(1): 75. 1985. Type: Hawaiian Islands G.T.Lay & A.Collie 3321 (holo: BM! [BM1038070]). Heterotypic synonyms: Lycopodium pachystachyon Spring, Bull. Acad. Roy. Sci. Belg. 8:518. 1841. Huperzia pachystachyon (Spring) Roth., Feddes Repert. 54:62. 1944. Type: Hawaii. C.-B.Gaudichaud (iso: BM, L! [L57364], P). IPNI ID: 77125132-1


Phlegmariurus pichianus (Tardieu) A.R.Field & Bostock comb. nov. Basionym: Huperzia pichiana Tardieu, Adansonia sér. 2, 10: 21, t. 2, f. 5-8. 1970. Type: Madagascar. H.Perrier 17773 (holo: P! [P466620], iso: BM! [BM785232], P! [P466618, P466619, P226993]). IPNI ID: 77125134-1


Phlegmariurus proliferus (Blume) A.R.Field & Bostock comb. nov. Basionym: Lycopodium proliferum Blume, Enum. Pl. Javae 2:265. 1828. Synonym: Huperzia prolifera (Blume) Trevis., Atti Soc. Ital. Sci. Nat. 17(2). 1874. Type: Java [Indonesia], C.L.Blume s.n. (lecto: L! [L57380], iso: P!). IPNI ID: 77125135-1


Phlegmariurus pulcherrimus Á.Löve & D.Löve Taxon 26(2-3): 324. Basionym: Lycopodium pulcherrimum Wall. ex Hook. & Grev. Icon. Filic. 1(2): 38. 1827. Synonyms: Huperzia pulcherrima (Wall. ex Hook. & Grev.) Pic.Serm. Type: Nepal N. Wallich s.n. (iso: BM! [BM1038025]). IPNI ID: 77125138-1


Phlegmariurus ribourtii (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium ribourtii Herter, Bot. Jahrb. Syst. 43: Beibl. 98:53. 1909. Synonym: Huperzia ribourtii (Herter) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: Tahiti. Ribourt s.n. (syn: P, BONN-Nessel), Nadeau s.n. (syn: P). IPNI ID: 77125140-1


Phlegmariurus rubricus (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys rubricus Herter, Revista Sudamer. Bot. 8:25. 1949. Synonym: Huperzia rubrica (Herter) Tardieu, Adansonia sér. 2, 10: 18. 1970. Type: Madagascar. H.Humbert 3717 (holo: P! [P466617]). IPNI ID: 77125141-1


Phlegmariurus rupicola (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium rupicola Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 16:39. 1914. Synonym: Huperzia rupicola (Alderw.) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: New Guinea. Gjellerup 1080 (holo: BO, iso: L! [L57385]). IPNI ID: 77125145-1


Phlegmariurus salvinioides (Herter) Ching, Acta Bot. Yunnan. 4(2):122. 1982. Basionym: Urostachys salvinioides Herter, Bot. Arch. 3: 18. 1923. Synonym: Huperzia salvinioides (Herter) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: not designated, numerous Philippines specimens cited. Heterotypic synonyms: Lycopodium edanoi Copel., Philipp. J. Sci. 46:209. 1931. Synonym: Huperzia edanoi (Copel.) Holub, Geobot. Phytotax. 20(1): 72. 1985. Type: Philippines G.E.Edaño (holo: MICH, iso: NY! [NY127347]).


Phlegmariurus samoanus (Herter ex Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys samoanus Herter ex Nessel, Revista Sudamer. Bot. 6:163. 1940. Synonyms: Huperzia samoana (Nessel) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: Samoa. Dr Friedrich s.n. (lecto: BONN-Nessel designated Øllgaard 1989: 100). IPNI ID: 77125146-1


Phlegmariurus shangsiensis C.Y.Yang, Acta Phytotax. Sin.22(1):87. 1984. Type: China. W.T.Tsan 22289 (holo: SCBI).


Phlegmariurus schlechteri (E.Pritz.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium schlecteri E.Pritz. in Schechter, Bot. Jahrb. Syst. 39:14. 1906. Synonym: Huperzia schlechteri (E.Pritz.) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: New Caledonia. R.Schlechter 15174 (holo: B, iso: BM, K). IPNI ID: 77125148-1


Phlegmariurus setifolius (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium setifolium Alderw., Bull. Bot. Jahrb. Buitenzorg, ser. 2, 16:40. 1914. Synonym: Huperzia setifolia (Alderw.) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: Borneo. Teysman s.n. (iso: L! [L57387]). IPNI ID: 77125150-1


Phlegmariurus sieboldii (Miq.) Ching, Fl. Xisang 1:12. 1983. Basionym: Lycopodium sieboldii Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 184: 1897. Synonyms: Huperzia sieboldii (Miq.) Holub, Geobot. Phytotax. 20(1): 76. 1985. Type: Japan. P.F.von Siebold s.n. (iso: L, US! [US134339]).


Phlegmariurus sooianus Lawalrée, Acta Bot. Acad. Sci. Hung. 19: 195. 1973. Synonyms: Huperzia sooiana (Lawalrée) Holub, Folia Geobot. Phytotax. 20: 77. 1985. Type: Zaire. Evrard 3580 (holo: BR).


Phlegmariurus squarrosus (G.Forst.) Á. Löve & D. Löve, Taxon 26:324. 1977. Basionym: Lycopodium squarrosum G.Forst., Fl. Ins. Austr. 479 (1786). Type: Tahiti. J & G. Forster s.n. (isosyn: BM! [BM1048443] and P!). Synonyms: Huperzia squarrosa (G.Forst.) Trevis.; Atti Soc. Ital. Sci. Nat. 17: 247. 1874; Urostachys squarrosus (G.Forst.) Herter; Bot. Arch. 3: 14. 1923.Heterotypic synonyms: Lycopodium acutifolium Desv. ex Poir., Encycl. suppl. 3: 599. 1813 [1814]. Type: Mauritius. s.c. s.n. (holo: P! [P4666615]). Lycopodium epiceifolium Desv. ex Poir., Encycl. suppl. 3:559. 1913; Huperzia epiceifolia (Desv. ex Poir.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Type: Mauritius. s.c. s.n. (holo: P! [P0466614]). Urostachys madagascariensis (Nessel) Herter, Index Lyc. 68. 1949; Type: Madagascar. H.Perrier 8276 (holo: P). Lycopodium pseudosquarrosum Pamp., Bull. Reale Soc. Tosc. Ortic. 111, 13: 99. 1908. Type: Indonesia. Pampanini s.n. (iso: P).


Phlegmariurus staudtii (Nessel) A.R.Field & Bostock comb. nov. Basionym: Urostachys staudtii Nessel, Repert. Spec. Nov. Regni Veg. 48:170. 1940. Type: Cameroon. Staudt 476 (lecto: P, isolecto: BONN-Nessel). IPNI ID: 77125151-1


Phlegmariurus strictus (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium strictum Baker, J. Bot. 11:271. 1882. Synonym: Huperzia stricta (Baker) Tardieu, Adansonia sér. 2, 10: 20. 1970. Type: Madagascar. Parker s.n. (holo: K). IPNI ID: 77125152-1


Phlegmariurus subfalciformis (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium subfalciforme Alderw., Bull. Jard. Bot. Buitenzorg. ser. 2, 28:44. 1918. Synonym: Huperzia subfalciformis (Alderw.) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Papua New Guinea. R.Schlechter 13775 (syn: B! [B0114953]). IPNI ID: 77125153-1


Phlegmariurus subulifolius (Wall. ex Hook. & Grev.) S.R.Ghosh, Pterid. Fl. E. India66. 2004. Basionym: Lycopodium subulifolium Wall. ex Hook. & Grev., Icon. Filic. 1. 1827. Synonym: Huperzia subulifolia (Wall. ex Hook & Grev.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Type: Nepal. N.Wallich s.n. (holo: K). IPNI ID: 77125155-1


Phlegmariurus subtrifoliatus (Brownlie) A.R.Field & Bostock comb. nov. Basionym: Lycopodium subtrifoliatum Brownlie, Nova Hedwigia Beih., 55 (Pterid. Fl. Fiji) 24. 1977. Synonyms: Huperzia subtrifoliata (Brownlie) Holub Folia Geobot. Phytotax. 26(1): 93. 1991. Type: Fiji. G.Brownlie 1287 (holo: CHR! [CHR341227]).


Phlegmariurus talamauanus (Alderw.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium talamauanum Alderw., Bull. Jard. Bot. Buitenzorg, ser. 2, 28:45. 1918. Synonym: Huperzia talamauana (Alderw.) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Sumatra [Indonesia]. Bünnemeijer 814 (holo: BO, iso: BONN-Nessel, L! [L57395]). IPNI ID: 77125156-1


Phlegmariurus terrae-guilelmii (Herter) A.R.Field & Bostock comb. nov. Basionym: Lycopodium terrae-guilelmii Herter, Bot. Jahrb. Syst. 54:229. 1916. Synonym: Huperzia terrae-guilelmii (Herter) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: New Guinea. Ledermann 9602 (holo: B). IPNI ID: 77125157-1


Phlegmariurus tetrastichus (Kunze) A.R.Field & Bostock comb. nov. Basionym: Lycopodium tetrastichum Kunze, Bot. Zeit. (Berlin) 6:99. 1848. Synonym: Huperzia tetrasticha (Kunze ex Alderw.) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Java [Indonesia]. Zollinger 2460 (iso: G, HBG, L! [L57313]). IPNI ID: 77125159-1


Phlegmariurus tetrastichoides (A.R.Field & Bostock) A.R.Field & Bostock comb. nov. Basionym: Huperzia tetrastichoides A.R.Field & Bostock, Austrobaileya 7(4): 711. 2008. Type: Australia. A.R.Field & O.Rawlins 1139 (holo: BRI, iso: CNS). IPNI ID: 77125158-1


Phlegmariurus toppingii (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys toppingii Herter, Bot Arch. 3:15. 1923. Synonym: Huperzia toppingii (Herter) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Philippines. D.L.Topping 1096 (iso: BONN-Nessel, NY! [NY127365], US! [US134342]). IPNI ID: 77125160-1


Phlegmariurus tournayanus Lawalrée, Acta Bot. Acad. Sci. Hung. 19:197. 1973. Synonym: Huperzia tournayana (Lawalrée) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Congo. J.Louis 4012 (iso: P! [P507738]).


Phlegmariurus trifoliatus (Copel.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium trifoliatum Copel., Bernice P. Bishop Mus. Bull. 59:7. 1929. Synonym: Huperzia trifoliata (Copel.) Holub, Geobot. Phytotax. 20(1): 77. 1985. Type: Fiji. H.E.Parks 20225 (holo: UC! [351031], iso: NY! [NY127366]). IPNI ID: 77125161-1


Phlegmariurus trigonus (C.Chr.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium trigonum C.Chr., Dansk Bot. Ark. 7:191. 1932. Synonym: Huperzia trigona (C.Chr.) Tardieu, Adansonia sér. 2, 10: 20. 1970. Type: Madagascar. H.Perrier 8270 (holo: P! [P466613]). IPNI ID: 77125162-1


Phlegmariurus ulicifolius (Sw.) S.R.Ghosh, Pterid. Fl. E. India 64. 2004. Basionym: Lycopodium ulcifolium Sw., Syn. Fil. 177. 1806. Synonym: Huperzia ulcifolia (Sw.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Type: India orientalis.


Phlegmariurus varius (R.Br.) A.R.Field & Bostock comb. nov. Basionym: Lycopodium varium R.Br., Prodr. 165. 1810. Synonym: Huperzia varia (R.Br.) Trevis., Atti Soc. Ital. Sci. Nat. 17:247. 1874. Type: Australia. R.Brown Iter Austral. 124 (isolecto: BM!, K!). Heterotypic synonyms: Lycopodium billardierei Spring, Bull. Acad. Roy. Sci. Belg. 8:516 1841; Huperzia billardierei (Spring) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54(1) 1944. Type: Nova Hollandia [Australia] La Billardière s.n. (syn: P-Jussieu!). Lycopodium novae-zelandicum Colenso, Trans. & Proc. New Zealand Inst. 19: 275. 1887; Huperzia novae-zelandica (Colenso) Holub, Folia Geobot. Phytotax. 20:75 (1985); Type: New Zealand (holo: WELT). Lycopodium flagellaria Bory in Duperry, Voy. monde, Bot. 1: 248 t. 26. 1829. Type: Novelle Irlande [Papua New Guinea] Durville & Lesson s.n. (syn: ?); d’Offack s.n. (syn: P). Lycopodium pachystachyum Desv. ex Poir., Encycl. suppl. 3: 544. 1813 [1814]. Type: Novelle Hollande [Australia]. s.c. s.n. (holo: P). IPNI ID: 77125163-1


Phlegmariurus whitfordii (Herter) A.R.Field & Bostock comb. nov. Basionym: Urostachys whitfordii Herter, Bot. Arch. 3:14. 1923. Synonym: Huperzia whitfordii (Herter) Holub, Geobot. Phytotax. 20(1): 78. 1985. Type: Philippines. H.N.Whitford 798 (holo: PNH lost 1945, iso: MICH! [MICH1191097], US! [US134345]). Heterotypic synonyms: Lycopodium magnusianum Herter, Hedwigia 49:91. 1909. Huperzia magnusiana (Herter) Holub, Geobot. Phytotax. 20(1): 74. 1985. Type: Philippines. M.S.Clemens (holo: B!). IPNI ID: 77125164-1


Phlegmariurus xiphophyllus (Baker) A.R.Field & Bostock comb. nov. Basionym: Lycopodium xiphophyllum Baker, Handb. fern-allies 12.1887. Synonym: Huperzia xiphophylla (Baker) Tardieu, Adansonia sér. 2, 10: 20. 1970. Type: Madagascar. R.Baron 4512 (holo: K! [K351213]). IPNI ID: 77125165-1


Phlegmariurus yandongensis Ching & C.F.Zhang, Bull. Bot. Res., Harbin3(3): 2. 1983. Type: China. K.Y.Shing 256.


Phlegmariurus yunnanensis Ching, Acta Bot. Yunnan. 4(2): 121. 1982. Type: China. K.M.Feng 7274 (holo: PE).

References
Bruce JG (1976) Development and distribution of mucilage canals in Lycopodium. American Fern Journal, 63, 481–491.
Christenhusz MJM, Zhang X-C, Schneider H (2011) A linear sequence of extant families anf genera of lycophytes and ferns. Phytotaxa, 19, 7–54.
Holub J (1964) Lycopodiella, novy rod radu Lycopodiales. Preslia, 36, 16–22.
Ji S-G, Hou K-K, Wang J, Pan S-L (2008) A molecular phylogenetic study of Huperziaceae based on chloroplast rbcL and psbA-trnH sequences. Journal of Systematics and Evolution, 46, 213–219.
McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nocholson DH, Prado J, Silva PC, Skog JE, Wiersema JH, Turland NJ (2006) International code of botanical nomenclature (Vienna Code): Adopted by the Seventeenth International Botanical Congress Vienna, Australia. July 2005. Regnum Vegetabile, 146.
Øllgaard B (1975) Studies in the Lycopodiaceae I. Observations of the structure of the sporangium wall. American Fern Journal, 65, 19–27.
Øllgaard B (1979) Studies in the Lycopodiaceae II. The branching patterns and infrageneric groups of Lycopodium sensu lato. American Fern Journal, 69, 49–61.
Øllgaard B (1987) A revised classification of the Lycopodiaceae s.l. Opera Botanica, 92, 253–178.
Øllgaard B (1989a) Index of the Lycopodiaceae. Biologiske skrifter. Royal Danish Academy of Sciences and Letters.
Øllgaard B (2012a) New combinations in Neotropical Lycopodiaceae. Phytotaxa, 57, 10–22.
Øllgaard B (2012b) Nomenclatural changes in Braziliand Lycopodiaceae. Rodriguésia, 63, 479–482.
Wagner WH, Beitel JM (1992) Generic classification of modern North American Lycopodiaceae. Annals of the Missouri Botanic Gardens, 79, 676–686.
Whittier DP (2006) Gametophytes of four tropical, terrestrial Huperzia species (Lycopodiaceae). American Fern Journal, 96, 54–61.
Whittier DP, Braggins JE (1992) The young gametophyte of Phylloglossum (Lycopodiaceae). Annals of the Missouri Botanic Gardens, 79, 730–736.
Whittier DP, Braggins JE (2000) Observations on the mature gametophyte of Phylloglossum (Lycopodiaceae). American Journal of Botany, 87, 920–924.
Whittier DP, Storchova H (2007) The gametophyte of Huperzia selago in culture. American Fern Journal, 97, 149–154.
Wikström N, Kenrick P (1997) Phylogeny of Lycopodiaceae and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. International Journal of Plant Sciences, 158, 862–871.
Wikström N, Kenrick P (2000) Phylogeny of epiphytic Huperzia (Lycopodiaceae): paleotropical and neotropical clades corroborated by rbcL sequences. Nordic Journal of Botany, 20, 165–171.
Wikström N, Kenrick P (2001) Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by use of nonparametic rate smoothing. Molecular Phylogenetics and Evolution, 19, 177–186.
Wikström N, Kenrick P, Chase M (1999) Epiphytism and terrestrialization in tropical Huperzia (Lycopoidaceae). Plant Systematics and Evolution, 218, 221–243.
Wilce JH (1972) Lycopod spores, 1. General spore patterns and the generic segregates of Lycopodium. American Fern Journal, 52, 65–79.
Yatsentyuk S, Valiejo-Roman KM, Samigullin TH, Wikstrom N, Troitsky AV (2001) Evolution of Lycopodiaceae inferred from spacer sequencing of chloroplast rRNA geners. Russian Journal of Genetics, 37, 1274–1280.
Zhang L-B (2012) The nomenclature of the lycophyte species Phlegmariurus mingcheensis Ching (Huperziaceae). Taxon, 61, 665–666.