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Taxonomical and Distributional Notes on Polylepis (Rosaceae)

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Org. Divers. Evol. 6, Electr. Suppl. 1: 1 - 10 (2006) © Gesellschaft für Biologische Systematik URL: http://www.senckenberg.de/odes/06-01.htm URN: urn:nbn:de:0028-odes0601-8 Taxonomical and distributional notes on Polylepis (Rosaceae) Michael Kessler, Alexander N. Schmidt-Lebuhn* Department of Systematic Botany, Albrecht-von-Haller-Institute of Plant Sciences, Göttingen University, 37073 Göttingen, Germany * Corresponding author, e-mail: [email protected] Received 3 March 2005 • Accepted 4 April 2005 Abstract Polylepis pacensis M. Kessler & Schmidt-Leb. spec. nov. is described; P. flavipila (Bitter) M. Kessler & Schmidt-Leb., P. incarum (Bitter) M. Kessler & Schmidt-Leb., P. lanata (Kuntze) M. Kessler & Schmidt-Leb., and P. subtusalbida (Bitter) M. Kessler & Schmidt-Leb. are elevated from subspecies or varietal to species rank; P. triacontandra Bitter is reinstated as a species. Extensions to the known distribution ranges of three additional species of Polylepis are reported. An updated key to species in the genus is provided. Keywords: Polylepis; Rosaceae; Taxonomy; Andes Introduction The genus Polylepis Ruiz & Pav. (Rosaceae, Sangui- In the second revision of the genus, Simpson (1979), sorbeae) comprises species of trees inhabiting the high working with much more extensive herbarium materi- Andes from Venezuela to central Argentina. The num- al than available to Bitter, and based on personal field ber of species included by different authors varies bet- experience with several species of Polylepis, realized ween 15 and 33 (Bitter 1911; Simpson 1979; Kessler that many of the taxa recognized by Bitter represen- 1995a, b). The circumscription of taxa in the genus is ted variants of morphologically variable species. She difficult due to extensive morphological variability therefore reduced the number of species to 15, but later within populations, limited variability between many (Simpson 1986) treated P. tarapacana Phil. as distinct of the recognized species, and apparently extensive from P. tomentella Wedd. and mentioned that P. besseri hybridization (Simpson 1979, 1986; Kessler 1995a; Hieron. as defined by her in 1979 probably represented Romoleroux 1996; A.N. Schmidt-Lebuhn, M. Kess- a species complex. In a taxonomic revision of the Bo- ler & M. Kumar unpubl. data). Similar problems are livian species of the genus, Kessler (1995a) attempted found in a number of other genera of Rosaceae, e.g., to disentangle the taxa within this complex, recogni- the subfamily Maloideae (Campell & Dickinson zing two species, P. besseri and P. racemosa Ruiz & Pav., with three and two subspecies, respectively. The 1990) and the genus Acaena (Dawson 1960), which category of subspecies was used for morphologically is closely related to Polylepis. and geographically distinct taxa that were linked by The first taxonomic revision of Polylepis was pre- intermediate populations suggesting hybridogenic in- pared by Bitter (1911) who applied a typological spe- trogression. Within P. tomentella, Kessler (1995a) also cies concept and recognized 33 species, 9 subspecies, recognized three subspecies, in this case corresponding and 18 varieties, placing a different name on almost to geographically separate entities differing primarily every single herbarium specimen available to him. in size. A further member of the P. besseri complex Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Kessler & Schmidt-Lebuhn: Taxonomical and distributional notes on Polylepis (Rosaceae) 2 from southwestern Peru and northwestern Chile, P. ru- rience to adequately judge their status. These include gulosa Bitter, was also recognized as distinct (Kessler (A) a population in the Urubamba Valley of Cuzco that 1995b). was placed in P. besseri by Simpson (1979), and (B) In the ten years that have passed since Kessler‘s large stands on limestone hills north of Lake Titicaca. (1995a, b) study, much additional information on Po- The name P. pallidistigma Bitter may apply to this lat- lylepis has come to light, based especially on extensive ter taxon, but we have only seen a photograph of the collections mainly in Peru and Bolivia, and on molecu- type material and no recent collections from the type lar analyses of most taxa within the genus (Kerr 2003; locality. A.N. Schmidt-Lebuhn, M. Kessler & M. Kumar unpu- Because the taxonomic changes proposed by Kess- bl. data). These data suggest that hybridization takes ler (1995a, b) and by us imply that Simpson‘s (1979) place between probably all species of the genus as long key to Polylepis is outdated, we here provide a key to as they grow in geographical proximity to each other. the species recognized by us. Identification of speci- This applies even to morphologically clearly distinct mens of Polylepis is difficult due to the variability of species that were placed in different species groups the taxa and should best be attempted with several by Simpson (1979, 1986). Kerr (2003) also found evi- specimens from a given population. Ideally, young but dence for hybridization between Polylepis and the sis- mature branches should be studied, because older bran- ter genus Acaena. These findings have prompted us to ches often lose the hairs on stipular sheaths and leaves. reassess the taxonomic implications of hybridization in Hybrids are frequent and may be recognizeable only the genus. Based on a biological species concept (Mayr in extensive collection series also including the parent 1942), Kessler (1995a) placed morphologically distinct species. The two populations of uncertain taxonomic populations in a single species when they were linked status mentioned above (P. aff. besseri, P. pallidistig- by intermediate „hybrid“ populations, but our molecu- ma) were not included in the key because insufficient lar data shows that such „subspecies“ are genetically material was available to us. not closer to each other than species that have been unambiguously recognized as distinct species by all re- Key to the species of Polylepis searchers working on the genus (Bitter 1911; Simpson 1. Lower leaflet surfaces with sericeous indument, ra- 1979; Kessler 1995a; Romoleroux 1996). Rigorous ap- rely almost glabrous with a few silky hairs restric- plication of the biological species concept within Po- ted to veins ........................................................... 2 lylepis would result in the recognition of probably only – Lower leaflet surfaces with woolly, pannose, vill- a single species, an unsatisfactory course of action that ous, strigose, or small, matted, glandular indument, would not account for the conspicuous morphological or glabrous (some species with sericeous hairs on differences between many populations and clear bio- leaflet margins) .................................................... 6 geographic patterns. An alternative approach, and the 2. Hairs of lower leaflet surfaces dense, evenly co- one we favour, is to recognize morphologically, bio- vering the surface ................................................ 3 geographically, and ecologically distinct populations at – Hairs of lower leaflet surface confined to, or most species rank, even if there is evidence for hybridization prominent on, the veins ....................................... 4 with other species. Such a phylogenetic species con- 3. Lower leaflet surfaces often with a layer of cept (Nixon and Wheeler 1990) recognizes populations small, lanose hairs below the silky hairs; leaf- as species as long as genetic exchange with other po- let apices emarginate; 2–3(–4) pairs of leaflets pulations does not inhibit the independent evolutionary ............................................... P. lanuginosa Kunth development of these populations (Luckow 1995). – Lower leaflet surfaces lacking a lower layer of small As a result of this reappraisal of species limits in hairs; leaflet apices obtuse to acute; (2–)5–7 pairs of Polylepis, we here describe one new species, and raise leaflets ......................................... P. sericea Wedd. five taxa to species rank that were previously treated 4. Leaflet apices acute; 4–9 pairs of leaflets ................ as subspecies or varieties or placed in synonymy with ..................................................... P. pauta Hieron. other species. This increases the number of species in – Leaflet apices emarginate; 1–3 pairs of leaflets ... 5 Polylepis to 26. Taking into account three new country 5. Leaflets 0.6–1.3 cm long; fruits with irregular pro- records also reported here, the number of species recor- tuberances or blunt spines ....................................... ded per country now is 1 in Venezuela, 3 in Colombia .............................................P. pepei B.B. Simpson (1 endemic), 7 in Ecuador (2 endemics), 14 in Peru (3 – Leaflets 1.7–2.3 cm long; fruits with irregular flatte- endemics), 13 in Bolivia (4 endemics), 2 in Chile, and ned ridges ..................... P. subsericans J.F. Macbr. 4 in Argentina (1 endemic). Two further populations 6. Lower leaflet surfaces glabrous or with very short from Peru may also merit species rank, but we do not pannose, matted, glandular hairs that are not recog- yet have sufficient herbarium material and field expe- nizable as individual hairs with the unaided eye, Org. Divers. Evol. 6, Electr. Suppl. 1 (2006) Kessler & Schmidt-Lebuhn: Taxonomical and distributional notes on Polylepis (Rosaceae) 3 hairs shorter than the height of the leaf veins; leaf glandular hairs; leaflet apices emarginate; Colom- veins glabrous or puberulous, easily visible ........ 7 bia to NW Argentina .........................................
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    US007973216 B2 (12) United States Patent (10) Patent No.: US 7,973,216 B2 Espley et al. (45) Date of Patent: Jul. 5, 2011 (54) COMPOSITIONS AND METHODS FOR 6,037,522 A 3/2000 Dong et al. MODULATING PGMENT PRODUCTION IN 6,074,877 A 6/2000 DHalluin et al. 2004.0034.888 A1 2/2004 Liu et al. PLANTS FOREIGN PATENT DOCUMENTS (75) Inventors: Richard Espley, Auckland (NZ); Roger WO WOO1, 59 103 8, 2001 Hellens, Auckland (NZ); Andrew C. WO WO O2/OO894 1, 2002 WO WO O2/O55658 T 2002 Allan, Auckland (NZ) WO WOO3,0843.12 10, 2003 WO WO 2004/096994 11, 2004 (73) Assignee: The New Zealand Institute for Plant WO WO 2005/001050 1, 2005 and food Research Limited, Auckland (NZ) OTHER PUBLICATIONS Bovy et al. (Plant Cell, 14:2509-2526, Published 2002).* (*) Notice: Subject to any disclaimer, the term of this Wells (Biochemistry 29:8509-8517, 1990).* patent is extended or adjusted under 35 Guo et al. (PNAS, 101: 9205-9210, 2004).* U.S.C. 154(b) by 0 days. Keskinet al. (Protein Science, 13:1043-1055, 2004).* Thornton et al. (Nature structural Biology, structural genomics (21) Appl. No.: 12/065,251 supplement, Nov. 2000).* Ngo et al., (The Protein Folding Problem and Tertiary Structure (22) PCT Filed: Aug. 30, 2006 Prediction, K. Merz., and S. Le Grand (eds.) pp. 492-495, 1994).* Doerks et al., (TIG, 14:248-250, 1998).* (86). PCT No.: Smith et al. (Nature Biotechnology, 15:1222-1223, 1997).* Bork et al. (TIG, 12:425-427, 1996).* S371 (c)(1), Vom Endt et al.