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Tribes Anthospermeae and Paederieae

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Tribes Anthospermeae and Paederieae The Wood Anatomy of Rubiaceae tribes Anthospermeae and Paederieae By J. Koek-Noorman and Ch. Puff (Received April 27, 1982) Key Words: Rubiaceae, Anthospermeae, Paederieae. —Wood anatomy, secondary woodiness. wood anatomical the Abstract: Detailed descriptions are given for genera Anthospermum, Nenax, Phyllis, Carpacoce, Coprosma, Neogaillonia, Crocyllis, Plocama and Spermadictyon, and miscellaneous wood anatomical data on the and Aitchisonia. The genera Normandia, Pomax, Opercularia, Leptodermis wood anatomical variation within the is discussed. large genus Anthospermum Secondary woodiness is likely to occur in a number of Anthospermumspecies; other the have “normal” wood structure herbaceous. species of genus or are The Nenax and Carpacoce may also have secondary woodiness. Anthospermeae and the Paederieae differ from each other primarily in the fibre type: Anthospermeae have fibre tracheids, Paederieae libriform fibres. Except for the wood anatomical the delimitiation of the genus Phyllis , features largely support tribes and certain the within the two to a extent suggested groupings of genera two tribes. During the preparation of a paper on the delimitationof the tribes Anthospermeae and Paederieae(Puff 1982), it became obvious that very few wood anatomical data available the to are on genera belonging these tribes. Aims of this publication are, therefore, (1) to present wood anatomical of the and descriptions woody genera woody species belonging to genera with both woody and herbaceous representatives, out wood anatomical and to and (2) to point variation, (3) compare correlate the obtained wood anatomical data, as an independent data set, with the new classification of the tribes and the ideas on the relationships between genera and tribes as expressed by Puff (1982). Yet another target of the wood anatomical studies is to specifically of and allied of investigate woody species Anthospermum generain view possible secondary woodiness. While, on the basis of the commonly 2 PI. Syst. Evol., Vol. 143, No. I—2 18 J. Koek-Noorman & Ch. Puff theories of accepted “evolutionary trends”, one would expect woody species to be outfitted with characters of which at least some should be “primitive”, and herbaceous species with more “derived” features in their character set, the reverse seems to be the case in the large genus Anthospermum. Detailed of the under investigations genus (Puff, preparation) have shown that the (large) shrubby species are characterized by many morphological features which can clearly be considered while the herbaceous “derived”, species exhibit more traits. the “primitive” Similarly, genus Nenax, on the basis of ample evidence considered be allied and to closely to a “segregate” of Anthospermum, is derived in many of its characters but consists, without exception, of (dwarf) shrubs. The occurrence of secondary woodiness could explain this apparent contradiction. Materials and Methods Wood of studied. of samples 14 genera were Nine these were investigated in wood anatomical data on the detail; remaining genera are incomplete, mainly because of ofthe A list of the very small diameter samples. complete specimens studied, including information on growth form and habitat, is given in Table 1. 15 25 stained with Sections, pm thick, were saffranin. Macerations, obtained treatment with and by hydrogen peroxide glacial acetic acid were stained with astra-blue. Terms are used according to the Multilingual Glossary of Terms used in Wood Anatomy (Iawa 1964) with the exception of the terms “libriformfibre” and “fibre tracheid”. Fibre definitionsfollow Reinders their type (1935), as use for the of appears preferable descriptions rubiaceous woods (Koek-Noorman 1969). Mean vessel element and fibre lengths were calculated out of 25 measurements and lowest and minimum highest averages are given; and Table 1. List of Species Collector and institutioninstitution where voucher specimen is housed AnthospermumAnlhospermum herbaceum L.L. f. Puff 800106-1/2 WUVVU Puff 780208-2/3 WU A. randii S. Moore Puff 790129-2/2 WU Puff 780215-3/1 WU A. ternatum HiernHtern Puff 781222-1/1 WU Wood Anatomy of Anthospermeae etc 19 maximum values are given between brackets. Lowest and highest fibre length/vessel element length ratios are also included. The neither parenchyma strands were often very scarce. Therefore, of nor strand element averages lengths parenchyma length/vessel length—ratios were calculated. observed and The most frequently highest vessel diameters ray heights per the of and values within each species are taken, and range the lowest highest is In the difficult to trace in genus given. some species, rays were so tangential that cells could be this sections, no height (in or established; for reason, numbers of rays per mm were not counted. of Table 2 summarizes some wood anatomical characters the species nine in it also information belonging to the genera studied detail; includes on the diameter of the samples. Tribe Anthospemeae Anthospermum L. (Figs. 1 a-f) Growth rings absent to present, mostly formed by a zone of radially flattened fibres and less and narrower vessels; sometimes a tangential row of wider vessels in the first-formed wood, or some terminal parenchyma is present. radial Vessels nearly exclusively solitary or partly arranged in 2 to diameter 30-60 multiples, over 100per mm , angular round, up to urn to in A. vessel member (up 100 gm herbaceum), length 195-620 (148-775) Perforations but few reticulate observed in A. gin. simple, a plates galpinii. Intervascular pits alternate, vestured, round to oval, 3-5gm; in A. herbaceum irregular, oval to elongated. Vessel-ray and vessel- parenchyma pits similar to the intervascular pits. Spiral thickenings in Fibriform A. basuticum. vessel members present in many species. specimens investigated Locality Growth form Habitat wet -» moist —-*■ dry SOUTH AFRICA, Natal, Drakensberge, herb with a + woody xX Royal Natal National ParkRark base Zomba Plateau above MALAWI, Zomba Plateau as xX ZIMBABWE, c. 2020 km E of Headlands as above Xx MALAWI, Dedza Distr., Chongoni as above Xx Forest Reserve MALAWI, foot of Mts., s. Mafingi Mts., (short-lived) shrubby xX on Zambian border herb,herb, woody at basebase 2* 20 J. Koek-Noorman & Ch. Puff Table 1 (continued) Species Collector and institution where voucher specimen is housed A. streyi Puff (ined.) Puff 790426-3/1 WU A. dregei Bond.Sond. Puff 780810-2/1 WUwu A. comptonii Puff (ined.) Puff 790914-3/2 WU A. hirtum Cruse Puff 800918-4/1 WUwu A. welwitschii Hiern Puff 790210-2/1 WUwu Puff 790211-2/3 WUwu A. usambarense K. Schum. Puff 781220-1/13 WUwu Puff 781223-2/1 WUwu A. ammannioides S. Moore Puff 790124-1/1 wuWU A. vallicolum S. Moorf,Moore Puff 790128-1/1 WUwu A. emirnense Bak. Puff 800729-1/1 WUwu Puff 800730-2/1 WUwu A. ibityense Puff (ined.) Puff 800730-1/1 WUwu A. isaloense Hom. ex Puff (ined.) Puff 800814-1/1 WUwu A. Sprenoel A. tricostatum Puff wu A. spathulatum Bprenoel(=( = A. Bond.)Sond.) Puff 791226-4/1 WU Puff 800924-3/1 wuWU A. aethiopicum L. Puff 800918-1/4 WUwu A. littoreum L.L, Bolus Puff 790424-1/1 wuWU Puff 790116-1/1 WUwu Puff 790304-1/1 WUwu A. galpinii Schltr. Puff 790303-2/1 WUwu Puff 790415-4/1 wuWU Wood Anatomy of Anthospermeae etc 21 Locality Growth form Habitat -» moist -* wet -» -» dry SOUTH AFRICA, Natal, South Coast, cushion-forming xX Paddock Distr., Highlands Farm dwarf shrub NAMIBIA (SOUTHWEST AFRICA), as above xX southern Namib, Farm Kubub SOUTH AFRICA, Cape Prov., as above xX Witteberge, S of Matjiesfontein, Farm Fisantekraal SOUTH AFRICA, Cape Prov., straggling xX rd. dwarf shrub S of Elim, on Viljoenshof rd. shrub SOUTH AFRICA, Transvaal,Transvaal. (large)(large) shrub xX The Downs area SOUTH AFRICA, Transvaal, as above xX Westfalia Estate near Duivelskloof, Piesangkop MALAWI,MALAWI. Misuku Hills, as above xX Willindi Forest Reserve ZAMBIA, Nyika Plateau as aboveabove Xx W of above ZIMBABWE, Juliasdale, as / Vukutu Farm ZIMBABWE, Vumba Mts., Castle Beacon as above xX above MADAGASCAR, Lac Tritriva as Xx MADAGASCAR, Col des Mt. as above x MADAGASCAR, Col des Tapias, Mt. IbityUnity as above X MADAGASCAR, Mt. IbityIbity erect dwarf shrub xX Isalo MADAGASCAR,MADAGASCAR. Isalo Massif, shrub xX W of Ranohira SOUTH AFRICA, Cape Prov., (large) shrubshrub xX “Atlantis” development SOUTH AFRICA, Cape Prov., small, stunted shrub xX Witsenberg, Sneeugat Peak area SOUTH AFRICA, Prov., X Cape (large) shrubshrub x Stanford-Papiesvlei rd., Perdeberg SOUTH shrubshrub SOUTH AFRICA, Cape Prov., straggling xX Transkei side of Umtamvuna Bridge, Natal border above SOUTH AFRICA, Cape Prov., as xX Gonubie Mouth SOUTH Richards above SOUTH AFRICA, Natal, Bay as xX SOUTH AFRICA, Natal, Nqutu-Qudeni shrub Xx rd., Sigqokwana ridge SOUTH SOUTH AFRICA, Cape Prov,,Prov., Transkei, as above xX above Port St.Johns 22 J. Koek-Noorman & Ch. Puff Table I (continued) SpeciesSpecies Collector andand institution where voucher specimen is housed A. basuticum Puff (ined.) Puff 790113-5/3 WU Nenax dregei L. Bolus Puff 800831-5/1 WU N. microphylla (Sond.) Salter Puff 790112-4/1 WU Phyllis nobla L. Mendoza-Heuer 75—143 Z P. viscosa Webb. Berth. P. viscosa Webb. & Berth. Mendoza-Heuer s.n.s. n. Z Sond. Puff Carpacoce spermacocea (Retchb.) Bond. 791222-3/2 WU C. heteromorpha (Buek) L,L. Bolus Puff 800101-3/1 WU C. vaginellata Salter Puff 800913-4/3 WU Coprosma arborea i Kirk FPRL 3838 C. monlanamontana Hillebrand Stern & Herbst2981 US C. C. pubenspubens A. Gray Stern & Herbst 2924 US Stern & HHerbsterbst 2977 US Normandia neocalidonica Hook. f. SchlechterScHLECHTER 14846 Ww Pomax umbellalaumbellata Soland. Bauer s.n.s. n. Ww 417 Ww Opercularia spermacocea Labile.Labill. PretzelPritzbl 417 Neogaillonia aucheri (Guill.) Puff LamondLamond 139 E N. eriantha (Jaub. & Spach) Linch. Rechinger 54589 Ww E. Mey. Schumann Crocyllis anthospermoides E, Mey. ex Schumann Puff 780811-1/1 WU Plocama pendula Ait. Ridsdale & Baas 4 LL Spermadictyon suaveolens Roxb, vanSteenis 21125 LL Leptodermis virgata Edcew.Edoew.
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