New and existing combinations in Palaeotropical Phlegmariurus (Lycopodiaceae) and lectotypification of the type species Phlegmariurus phlegmaria (L.) T.Sen & U.Sen.

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


Introduction
Th e Lycopodiaceae is a globally distributed family of homosporous Lycopodiophytes in which the species exhibit shoot-forms associated with their habitat. Early classifi cations 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 Kunze which was historically placed in a genus of its own. Multiple genera and subfamilies corresponding with the diff erent 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 classifi cations was the four-genus classifi cation 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 (Christenhusz et al. 2011, Ji et 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). Th is classifi cation produces a Huperzia sens. lat. that is morphologically heterogeneous and is not supported by synapomorphies that can be readily identifi ed in the fi eld. It also does not refl ect the ancient divergence times between the Huperzioid clades of Lycopodiaceae (Wikström and Kenrick 2001).
An alternative classifi cation is to circumscribe Huperzia Bernh. narrowly sensu Wagner & 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. Th is circumscription of Phlegmariurus includes a broad spectrum of both epiphytic and non-epiphytic species primarily from the tropical and subtropical regions globally. Th e 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 identifi ed from the examination of 121 type specimens (including images) for species previously com-bined 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). Th is suggests that morphologically similar species may not always be closely related. We have taken a more conservative approach to synonymisation for the P. phlegmaria (L.) T.Sen & U.Sen species complex as we consider this to be 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 fi rst discovered by European botanists in the East Indies in the late 1600s. Several species were described and illustrated in: Breyne (1678) Rheede et al. (1703p. 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 fi gures 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 L. phlegmaria L. to one species (McNeill et al. 2006). Øllgaard (1989)