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Phylogenetic System and Classification of the Family Scapaniaceae

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Phylogenetic System and Classification of the Family Scapaniaceae Ann. Bot. Fennici 39: 309–334 ISSN 0003-3847 Helsinki 11 December 2002 © Finnish Zoological and Botanical Publishing Board 2002 Phylogenetic system and classifi cation of the family Scapaniaceae Mig. emend. Potemkin (Hepaticae) Alexey D. Potemkin Department of Lichenology and Bryology, V.L. Komarov Botanical Institute, 2 Prof. Popov Street, St. Petersburg, 197376 Russia (e-mail: Vera@ iz6284.spb.edu) Received 15 February 2001, accepted 17 January 2002 Potemkin, A. D. 2002: Phylogenetic system and classifi cation of the family Scapaniaceae Mig. emend. Potemkin (Hepaticae). — Ann. Bot. Fennici 39: 309–334. The phylogenetic system and classifi cation of the family Scapaniaceae Mig. emend. Potemkin as a whole are presented for the fi rst time. The classifi cation of the family is signifi cantly revised. Scapaniaceae is classifi ed with 87 recent and one fossil species in 18 sections and three subgenera of one genus, Scapania (Dumort.) Dumort. emend. Potemkin. Subgenus Scapania has 82 species in 18 sections, subgenus Macrodiplo- phyllum (H. Buch) Potemkin has three species, and subgenus Macroscapania R.M. Schust. has two species. Earlier distinguished subgenera Plicaticalyx Müll. Frib. and Ascapania Grolle are merged in the sectio Plicaticalyx (Müll. Frib.) Potemkin, comb. & stat. nov., of the subgenus Scapania. The taxonomic composition of many sec- tions is considerably revised, and the species composition of all infrageneric groups is listed in a systematic arrangement. Alteration of the species composition and/or species differentiation (including keys) of the sections Ciliatae Grolle, Gracilidae H. Buch, Nemorosae (Müll. Frib.) H. Buch, Planifoliae (Müll. Frib.) Potemkin, Plicati- calyx, Scapania, and Verrucosae Potemkin of the subgenus Scapania as well as of the subgenera Macrodiplophyllum and Macroscapania are provided. New sections are described for Scapania karl-muelleri Grolle (Grolleoscapania Potemkin, sect. nova) and for S. schljakovii Potemkin and S. himalayica Müll. Frib. (Muelleria Potemkin, sect. nova). A number of new synonyms and index for identifi cation of taxonomic position of mentioned taxa of Scapania are provided. Key words: classifi cation, evolution, hepatics, phylogeny, Scapania, Scapaniaceae, taxonomy 310 Potemkin • ANN. BOT. FENNICI Vol. 39 Introduction Refi nements in the phylogenetic system of Scapania The paper presents a signifi cantly revised phy- logenetic concept of the monogeneric family After previous publication of the classifi ca- Scapaniaceae Mig. emend. Potemkin. It com- tion of the genus Scapania (Potemkin 1998a) prises all known evolutionary trends of Scapania I assembled all available data on the genus and (Dumort.) Dumort. and provides a phylogenetic analyzed them from the viewpoints of morphol- system and classifi cation of the entire family. ogy, ecology and distribution. Eight new species The paper is based on studies of all but a of Scapania were described (Potemkin 2000a, few unavailable recognized species of Scapania. 2000b, 2000c, 2001), among which S. kopone- Over 90 types mostly of critical taxa of Scapania nii Potemkin was important for re-evaluation were analyzed. Circumscription of all species is of relationships of the systematic groups of the based on a detailed analysis in about 60–100 genus. Scapania koponenii was segregated from morphological characters. The characters were plants distinguished earlier as Asiatic S. aspera evaluated from the viewpoints of the compara- M.&H. Bernet (Potemkin 1998a), and it was tive, evolutionary and functional morphology, demonstrated that its position is among the prim- taking into account their dependence on eco- itive species of the sectio Ciliatae Grolle rather logical conditions and variability throughout than in the sectio Aequilobae (Müll. Frib.) H. their range. Analysis of functional morphology Buch, to which S. aspera belongs. This altered is pertinent because of comprehensive morpho- the idea of a polymorphous “root species” of the functional integration of the gametophyte result- sectio Aequilobae and resulted in a delineation ing from its phenetically “developed” genotype. of the evolution of most sections of the subgenus All these approaches have provided the basis to Scapania from the common ancestor. analyze evolution in Scapania on the basis of Earlier the sectio Curtae (Müll. Frib.) H. functions of the gametophyte (e.g., assimila- Buch was distinguished as the simplest and prob- tion, water storage, light refl ection, sporophyte ably most primitive group of the genus (Potem- protection, etc.) and establishing that these are kin 1998a, 1999c). However, despite being becoming more effective in certain ecological richest in the genus with regard to an ensemble conditions by means of an integral interac- of unspecialized character expressions, the tion of diverse structures of the gametophyte. species of the sectio Curtae have a number of Analysis of functional morphology has lead advanced morphological characters (small size to understanding of evolutionary dynamics of of plants, reduced paraphyses, general suppres- characters within the genus, whereas approaches sion of branching with sporadic formation of of evolutionary morphology together with data lateral intercalary branches) and, advanced for on ecology and distribution have completed the the genus, a comparatively broad distribution evolutionary comprehension of the group. Space in territories with a continental climate. Moreo- constrictions do not permit full discussion of ver the distinctions of the species of the sectio details of all these methodological approaches Curtae are often uncertain. This lack of sharp to help me understand microevolution as well delimitation of the species speaks for a com- as taxonomic entities in hepatics. They were paratively young age of the group. On this basis already considered to some extent in my papers I distinguish the sectio Curtae as a young neo- (Potemkin 1998a, 1998b, 1999a, 2000d) and tenic sectio of Scapania with an origin probably will be further explored in forthcoming publica- induced by climatic changes of the Pleistocene. tions. General approaches of evolutionary mor- Close relationships of the sections Stephania phology of plants were considered by Takhtajan Amakawa & S. Hatt. ex Potemkin, Umbrosae H. (1954) and were applied to comprehension of Buch and Nemorosae (Müll. Frib.) H. Buch in macroevolution in hepatics by Schuster (1984a, their morphology and vicarious distribution pat- 1984b). tern led me to merge them in one, sectio Nemo- ANN. BOT. FENNICI Vol. 39 • Phylogenetic system and classifi cation of Scapaniaceae 311 Fig. 1. Phylogenetic scheme of Scapania (Dumort.) Dumort. Solid font = sections, hollow font = subgenera. Area of the contours roughly corresponds to number of species in the hypothesized evolutionary perspec- tive. Further explanation in the text. rosae. This very much expands the scope of the and its presumably Barbilophozia kunzeana- Nemorosae which, circumscribed in this way, like ancestor (Potemkin 1999c: p. 272). Among comprises the less specialized and probably most principal morphological characters forming this primitive recent species of the genus, Scapania hiatus, distinctions in branching pattern, leaf ligulata Steph. s. lato, as well as some advanced structure and ability to produce amphigastria are species, including S. nemorea (L.) Grolle, S. the most important. It should be noted that the umbrosa (Schrad.) Dumort., etc. This sectio species of the sectio Kunzeanae (C.E.O. Jensen) appears to be a “root group” from which most Grolle, to which Barbilophozia kunzeana of the other recent groups of Scapania might be (Huebe ner) Müll. Frib. belongs, develop chiefl y derived. terminal and exceptionally ventral intercalary Putting the sectio Nemorosae at the base branches, whereas primitive species of Scapania of the phylogenetic system of the genus sect. Nemorosae produce only ventral interca- (Fig. 1) shows a hiatus between the Nemorosae lary branches and no terminal ones. Species of 312 Potemkin • ANN. BOT. FENNICI Vol. 39 Scapania have a strongly fi xed bilobed leaf con- introduced synonyms, frequently used syno- dition and no underleaves while, on the contrary, nyms of recognized sections, names of species Barbilophozia kunzeana and related species are unavailable for study (given in brackets), and fl exible in leaf lobe number and freely develop taxonomic and systematic considerations. The underleaves. This hiatus should result from con- order of the taxonomic groups and species in siderable age of the genus Scapania, which may the classifi cation refl ects their relationships and extend to the age of Laurasia. their relative age. Distinction of sections of the The concept of sectio Nemorosae as the most subgenus Scapania often are not very sharply primitive group of the extant Scapania, as well defi ned because of considerable overlap of their as the idea of a considerable age of the genus, variability of separate characters (particularly is supported by discovery of S. hoffeinsiana the characters of the advanced species of large Grolle, the only known fossil Scapania, in Bit- and closely related groups). terfeld amber from Germany (Eocene, at least 50 m.y. BP) (Grolle & Schmidt 2001). This species appears to be close to S. umbrosa, an Subgenus Scapania s. lato advanced species of sect. Nemorosae. The phylogenetic system of the genus Scapa- = subgen. Ascapania Grolle, Buchiella R.M. Schust., Jense- nia S.W. Arnell, Kaalaasia H. Buch, Plicaticalyx Müll. Frib., nia is based on an understanding of
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