Studies in the Liverwort Family Metzgeriaceae (Metzgeriales) from Southern Africa. 1. Metzgeria Nudifrons Stephani and a Key to the Species
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Polish Botanical Journal 58(2): 449–455, 2013 DOI: 10.2478/pbj-2013-0045 STUDIES IN THE LIVERWORT FAMILY METZGERIACEAE (METZGERIALES) FROM SOUTHERN AFRICA. 1. METZGERIA NUDIFRONS STEPHANI AND A KEY TO THE SPECIES NO N KULULO PHE P HU & JACQUES VA N ROOY Abstract. This is the first of several projected articles on the taxonomy of southern African Metzgeriaceae H. Klinggr. A brief history of the genus Metzgeria Raddi in southern Africa, the only local representative, is presented. Family and genus descriptions, as well as a key to the species, are provided. Of the seven species currently recognized in southern Africa, M. nudifrons is the only species with gemmae on the dorsal surface of the thallus. It is also characterized by the distinctly broad costa, hairy male branchlets as well as single and straight hairs on the ventral face and margins of the thallus. Metzgeria nudifrons is described, illustrated and its geographical distribution mapped. Key words: Metzgeriaceae, Metzgeria nudifrons, South Africa, gemmae, taxonomy Nonkululo Phephu & Jacques van Rooy, National Herbarium, South African National Biodiversity Institute (SANBI), Private Bag X101, Pretoria 0001, South Africa;. e-mail: [email protected] & [email protected] IN T R ODUCT I O N The family Metzgeriaceae H. Klinggr. consists Metzgeria is a large genus of 60–80 validly of 2–4 genera world-wide (Wigginton 2004). It published species world-wide, with South America is the only family in the order Metzgeriales that as its centre of diversity with about 50 recognized is primarily epiphytic with most species growing species (Wigginton 2004; So 2004). Out of eight on tree bark or living leaves. Some species may recognized species in Africa, four are endemic to also grow on damp rocks or rarely on soil. They the continent (So 2004) and seven occur in southern generally grow in tropical to temperate rain and Africa. The basic morphological uniformity of mist forests in mountainous areas. It differs from Metzgeria has become a reason to group rather the other southern African families of the order than split the species. Recent revisions (e.g., So Metzgeriales (Perold 1999) in producing hairs 2002, 2004) have therefore resulted in a decrease on the ventral side and along the margins of in species numbers. In southern Africa the genus the thallus as well as the midrib, gemmae often ranges from coastal areas of the southern Cape present, male branch without and female branch and Kwa-Zulu Natal, inland to Swaziland and ex- with hairs, elaters 1-spiral, capsule wall 2-striated tending to the Wolkberg area in Limpopo. and archegonial neck reduced. The aim of this series is to revise the family In southern Africa, the family is only repre- Metzgeriaceae in southern Africa (South Africa, sented by one widespread genus, Metzgeria Raddi. Lesotho, Swaziland, Botswana, Namibia) in con- Metzgeria species are often intermixed with other tinuation of a revision of all liverworts in the region corticolous mosses and most are epiphytic. They (Perold 1999). The first paper in the series deals grow in wetter parts of forests or open habitats, with the taxonomy and nomenclature of Metzgeria rarely on soil but on tree trunks, branches, twigs, nudifrons Stephani, the only species with gemmae living leaves, and rocks. on the dorsal surface of the thallus. 450 POLISH BOTANICAL JOURNAL 58(1). 2013 HI STO ri CAL N OTES not resolve the nomenclature of the taxa. She rec- ognized 13 species in southern Africa, and sub- According to Pócs (1993, and references therein), sequently described a new species, M. nicomariei the African Metzgeriaceae were reviewed first Veltman & Potgieter (Veltman et al. 2000), which by Vanden Berghen (1948), who described two was reduced to synonymy of M. furcata by So new species and supplied a key to the Central (2004). African taxa. Kuwahara (1973, 1983, 1986) de- In the latest checklist of South African liv- scribed new species, new records, and placed erworts, Perold (2006) accepted five species of several African taxa in synonymy. As a result, Metzgeria, namely M. consanguinea Schiffner, the known distribution of several African Metz- M. furcata, M. leptoneura Spruce, M. madagassa geria species widened considerably. At the time and M. nudifrons, based on the revision of So of Pócs’s (1993) study, 21 Metzgeria species (2004), who also recognized M. quadrifaria from were known from sub-Saharan Africa, of which Lesotho. Phephu and van Rooy (2011) confirmed 14 occurred in Central and East Africa. He in- the occurrence of M. saxbyi in Swaziland and the cluded a tentative key to species, declaring that Western Cape. the African species of Metzgeria are badly in need of an up to date revision. The recent revisions MAT E ri ALS A N D M ETHODS by So (2002, 2004) have brought a measure of stability to the taxonomy and nomenclature of Traditional morphological-anatomical methods were Metzgeria in the region. However, she did not employed to study the species. Microscope slides were examine Metzgeria specimens in the National prepared using standard bryophyte preparation methods Herbarium, Pretoria (PRE), which houses the employing Hoyer’s Mounting Medium (Magill 1981). largest collection of southern African bryophytes, Specimens studied were obtained from PRE and BM but only some southern African collections in (herbarium acronyms according to Holmgren et al. 1990). overseas herbaria. Sim (1926) published the first synopsis of the family Metzgeriaceae in South Africa. He recog- TAXO N O mi C T R E AT M E N T nized four species of Metzgeria, namely M. perro- Metzgeriaceae H. Klinggr. tana Stephani, M. furcata (L.) Corda, M. conjugata Lindb. and M. muscicola Stephani. The next revi- Höh. Crypt. Preuss.: 10. 1858. Sim (1926), Arnell (1963), sion of southern African Metzgeriaceae was by Ar- Schuster (1992), Wigginton (2004), Costa (2008). nell (1963) who recognized 13 species in the region (South Africa, Swaziland, Lesotho and Namibia): TY P E GE N US : Metzgeria Raddi M. vialacea (Ach.) Dumort, M. elliotii Stephani, Plants thalloid, yellowish green. Thallus narrow, M. capensis S. W. Arnell, M. tabularis Stephani, thin, prostrate, erect or pendent and flattened, ei- M. perrotana Stephani., M. hamata Lindb. (illegiti- ther irregularly or dichotomously branched; lamina mate), M. camerunensis Stephani, M. madagassa 1-stratose, apex truncate (or obtuse), margins Stephani, M. limbatosetosa Stephani, M. nudifrons bearing bristle-like hairs; costa well defined, with Stephani, M. quadrifaria Stephani, M. saxbyi large epidermal cells; hairs bristle-like, on thallus Pearson and M. muscicola Stephani. Most of these margins and ventral surface of costa and thallus. species are now synonyms of the seven currently Oil bodies absent or very small, homogeneous. recognized species in the region while M. violacea Ventral scales absent. Asexual reproduction by does not occur in the southern African region (So gemmae. Dioicous or monoicous. Specialized male 2004; Perold 2006). Like Sim, Arnell did not list and female branches (gametangia) on very short the specimens examined. branchlets scattered along the ventral side of the Veltman (1998) revised the southern African costa, antheridia inside a globose sac, archegonia species of Metzgeria for her M.Sc. degree but did hidden under a thallus flap, sporophyte surrounded N. PHEPHU & J. van ROOy: STUDIES IN THE METZgERIACEAE FROM SOUTHERN AFRICA. 1 451 by fleshy, hairy calyptras; capsule spherical to PR OV I S I O N AL KEY TO THE S P EC I ES OF METZGERIA ovoid, splitting into four valves when mature, in SOUTHE rn AF ri CA walls 1–2-stratose; elaters single-spiral, attached to elatophore at valve apices. Spores small, minutely 1. Plants monoicous, marginal hairs single . papillose. ................................. M. saxbyi 1.* Plants dioicous, marginal hairs single or geminate ........................................ 2 Metzgeria Raddi 2. gemmae only on dorsal surface of thallus (best Jungermanniogr. Etrusca: 34. 1818. Schuster (1992), So seen on fresh material, could be lacking on old (2004), Wigginton (2004), Costa (2008). herbarium material), discoid and oval; costa very LECTOTY P E : M. glabra Raddi, nom. illeg. [≡ Junger- broad . M. nudifrons * mannia furcata L., M. furcata (L.) Dumortier], fide So 2. gemmae often absent, if present on margins of (2004). thallus; costa fairly narrow . 3 3. Hairs abundant, throughout margin and ventral costa, Plants bright to pale yellowish green, dried paired, distinctly curved; thallus markedly convex herbarium specimens sometimes blue-green or ............................. M. leptoneura whitish green. Thallus thin, linear, 4–60 mm long, 3.* Hairs sparse to often absent, on ventral face of costa, up to 2.5 mm wide; branching irregular or di- curved or straight; thallus not markedly convex ... 4 chotomous, sometimes with intercalary branches 4. Thallus with tapered and acute apices (dimorphic); arising from ventral face of costa; apex obtuse or gemmae concave . M. consanguinea * acute, retuse or truncate, sometimes with slime- 4. Thallus with obtuse apices only, furcate; gemmae papillae; margins plane or recurved, border ab- fairly flat .............................. 5 sent; costa well defined, composed of rows of 5. Costa of 2–3(–5) dorsal and 3–(4–6) ventral cells, thallus margins sometimes inrolled ............. large, thin-walled dorsal and ventral epidermal ............................. M. quadrifaria cells and thick-walled internal medullary cells; 5.* Costa of 2(–3) dorsal and 2–3(–5) ventral cells, wing 8–32 cells from midrib to thallus margin, thallus margins plane ...................... 6 cells hexagonal, thin-walled, trigones absent or 6. Marginal hairs usually single, rarely geminate (less minute. Hairs present, one-celled, on thallus mar- than 5%); thallus often sparsely hairy; hairs often gins, ventral side of costa and often on ventral straight, marginal and costal hairs often about same surfaces of lamina, single or paired (geminate), length; costa of 2 dorsal and 2(3–4) ventral cells straight or curved. gemmae borne on margins or .............................