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September 1999] KRON ET AL.—PHYLOGENETICS OF ANDROMEDEAE 1293 TABLE 2. Characters used in morphology-based cladistic analysis of Andromedeae. Plesiomorphic characters are listed first, followed by apomorphic ones. (Polarity decisions based on trees rooted with Enkianthus.) 1. Secondary phloem without bands of fibers (0); secondary phloem with bands of fibers (1). 2. Pith heterogeneous (0); pith homogeneous (1); pith Calluna-type (2). [unordered] 3. Bud scales numerous to 2, if the latter then minute (0); bud scales 2, enlarged (1). 4. Leaves convolute in bud (0); leaves revolute in bud (1). 5. Leaves alternate (0); leaves whorled (1). 6. Leaf venation pinnate (0); leaf venation parallel (1). 7. Vein reticulum open, the secondary and tertiary veins not equally prominent (0); vein reticulum dense, the secondary and tertiary veins equally prominent (1). 8. Leaf margin serrate or serrulate (0); leaf margin entire (1). 9. Leaf margin lacking raised glands (0); leaf margin with raised glands (1). 10. Leaves deciduous (0); leaves persistent (1). [polarity questionable; see Table 3] 11. Leaf midrib (as seen in cross section) unifacial (0); leaf midrib bifacial (1). 12. Fiber strands absent from leaf mesophyll (0); fiber strands present in leaf mesophyll (1). 13. Leaf epidermis not lignified (0); leaf epidermis at least slightly lignified (1). 14. Leaf epidermal cells not elongated (as seen in cross section) (0); epidermal cells elongated (1). 15. Hypodermis lacking (0); hypodermis present (1). 16. Multicellular hairs on leaves present (0); multicellular hairs lacking (1). 17. Multicellular hairs with small glandular head (or hairs not glandular) (0); multicellular hairs with expanded and elongated, glandular head (1). 18. Peltate scales lacking (0); peltate scales present (with margin irregular-erose) (1); peltate scales present (with margin entire) (2). [unordered] 19. Stalk of multicellular hair multiseriate (0); stalk of multicellular hair biseriate (1). 20. Stomata anomocytic (0); stomata paracytic (1); stomata tetracytic (2). [unordered] 21. Inflorescences monotelic, i.e., terminal flowers present (0); inflorescences polytelic, i.e., terminal flowers lacking (1). 22. Inflorescences developing (and producing flowers and fruits) on shoots of the current season (0); inflorescences developing (and producing flowers and fruits) on shoots of the previous season (1). 23. Inflorescences within bud (0); inflorescences exposed during winter or dormant season (1). 24. Pedicel lacking bracteoles (0); pedicel with two bracteoles (1); pedicel with several bracteoles (2). [unordered] 25. Bracteoles distinct, variably positioned (0); bracteoles connate, apical (1). 26. Pedicel not articulated with flower (0); pedicel articulated with flower (1). 27. Calyx of imbricate lobes (0); calyx of valvate lobes (1). 28. Calyx lobes brown, dry at maturity (0); calyx lobes usually colorful, fleshy at maturity (1). 29. Adaxial calyx stomata lacking (0); adaxial calyx stomata present (1). 30. Corolla lacking unicellular hairs on inside surface (0); corolla with unicellular hairs on inside surface (1). 31. Corolla lacking multicellular hairs on outer surface (0); corolla with multicellular hairs on outer surface (1). 32. Abaxial corolla stomata present (0); abaxial corolla stomata absent (1). 33. Filaments straight (0); filaments S-shaped (1). 34. Filaments distinct (0); filaments connate (1). 35. Filaments free from corolla (0); filaments adnate to corolla (1). 36. Filaments with unicellular hairs (0); filaments papillose or minutely roughened (1); filaments smooth (2). [unordered] 37. Anthers inverting late in development (0); anthers inverting early in development (1). 38. Anthers with projections (awns or spurs) (0); such projections absent (1). 39. Projections borne on anther, erect to deflexed (0); projections borne at junction of anther and filament (1); projections borne on filament (2). [unordered] 40. Projections single (or lacking) (0); projections divided (1). 41. Tubules on anthers absent (0); tubules on anthers present (1). 42. Disintegration tissue on back of anthers absent (0); disintegration tissue present (1). 43. Disintegration tissue not extending onto the spurs (or spurs lacking) (0); disintegration tissue extending onto the spurs (1). 44. Anthers open by longitudinal slits (0); anthers open by ‘‘terminal’’ pores (1). 45. Pollen shed as monads (0); pollen shed as tetrads (or modified tetrads) (1). 46. Ovary superior (0); ovary inferior (1). 47. Ovary lacking multicellular gland-headed hairs (0); ovary with multicellular gland-headed hairs (or homologous hair types, e.g., scales) (1). 48. Locules not divided (0); locules incompletely divided by outgrowth of ovary wall (1). 49. Fruit dehiscent (0); fruit indehiscent (1). 50. Fruit lacking thickened sutures (0); fruit with thickened sutures that often separate from the valves in dehiscence (1). 51. Fruit a capsule (0); fruit a berry (1). 52. Seed with vascular bundle in the raphe (0); seed lacking vascular bundle in the raphe (1). 53. Seed lacking fringe formed from differentiated cells (0); seed with fringe due to expanded and/or elongated, bulging cells (1). 54. Seed lacking tails (0); seed with tails, i.e., expanded portion of testa at hilum and chalazal ends (1). 55. Seed lacking unilateral wing (0); seed with unilateral wing (resulting from a flattening of chalazal end of seed, with cells of the wing not differentiated from those of the body) (1). 56. Testa one-layered (except sometimes in the raphe) (0); testa with several cell layers (1). 57. Testa cells isodiametric (in surface view) (0); testa cells much elongated (1). 58. Methyl salicylate lacking (0); methyl salicylate present (1). 59. Basic chromosome number 11 (0); 12 (1); 18 (2). [unordered] 60. Toxic diterpenes of andromedotoxin type absent (0); toxic diterpenes of andromedotoxin type present (1).
1294 AMERICAN JOURNAL OF BOTANY [Vol. 86 TABLE 3. Character values for species used in morphology-based cladistic analysis. V condition variable; ? condition unknown or not applicable. Species Character 1 2 3 4 5 6 123456789012345678901234567890123456789012345678901234567890 Enkianthus campanulatus 000000000000000000000000000000000000000000000000000000000000 Prionotes cerinthoides 0100000001011100000000020000000?000211?0000010000001000000?0 Sprengelia incarnata 0100010101001101?0?200020000000?101211?000001000000100001010 Agarista populifolia 1201001V010010000000V10101000000100011?00101100000010000101? Agarista salicifolia 10010011010010100000110101000000100011?0010110000001000010?? Pieris floribunda 1000001001000000001011110110100100001010010110000001000010?1 Pieris formosa 100000000100100000101111011010010000101001011000000100001011 Pieris phillyreifolia 1000000001001000001011110110100110021010010110000001000000?1 Pieris nana 11001001010010000010111101100001000110100V0110000001000010?1 Craibiodendron yunnanense 11000001011110000010010101000000100111?0010110000001001010?? Lyonia ferruginea 1100000101101000011011010100001010011V2001111010010101001011 Lyonia lucida 110000010110100000101101011000101001102001111010010100001011 Lyonia ligustrina 101000000010V00010101101011000101000102001111010010100001011 Lyonia ovalifolia 111000010V10V000101011010110001010001020011110V0010100001011 Vaccinium macrocarpon 01000001010000000001?00101000000000011?010011100102100000010 Vaccinium meridionale 01000000010000100001110101000000000101010011101102100000010 Satyria warszewiczii 01000001011000000001110101000000010011?01001110010210000V0?0 Leucothoë fontanesiana 020000000100000000011111010000000001100001011000000110000001 Leucothoë racemosa 010000000000000000011111010000000002100001011000000100000001 Andromeda polifolia 0001000101001001?0?011010100010100001000000011000000100010011 Zenobia pulverulenta 100000000000000000001101010000000001100101011000000100000001 Chamaedaphne calyculata 0100001001000000020V11010100000V0002100010011000000100000001 Diplycosia acuminata 01000000010100100001V10111010000V00211?01001100000010000112? Gaultheria miqueliana 000000000100000000011101010100000001100101011000000100000101 Gaultheria shallon 0000010000100000000011101010101100000100101011000000100000001 Pernettya tasmanica 00000000010000100001?102010101000000100100011000101100000001 Tepuia cardonae 01000001110000100001100111000V0?000011?0100110001011000001?? Oxydendrum arboreum 0200000V000000000001100100000000000011?001011000000100001011 Note: 25: Enkianthus scored as ‘‘0’’ because it has free bracts, which are homologous to bracteoles. 28: the coding of this character for members of the Vaccinieae, e.g., Vaccinium and Satyria, is problematic; they are coded as ‘‘0’’ based on the assumption that the colorful, fleshy sepals in Gaultheria and Diplycosia developed independently from the fleshy fruits of Vaccinieae; developmental studies would be useful. 60: See also Hegnauer (1966) and Anderberg (1993). hairs (no. 47), and capsule with thickened sutures (no. 50). Paired appendages on the filament (no. 39-2) are an additional synapomorphy under DELTRAN optimization. Agarista is supported by the apomorphies of leaves revolute in the bud (no. 4), leaves with a dense vein reticulum (no. 7), and stamens lacking appendages (no. 38). Gaultheria is monophyletic if Pernettya (as represented by P. tasmanica) is included, based on their fleshy calyx lobes (no. 28). The Lyonia group is monophyletic in some trees, but is paraphyletic in others because Andromeda polifolia is placed within the group (as sister to Agarista). Characters supportive of the monophyly of the Lyonia group include bands of fibers in the phloem (no. 1), S-shaped filaments (no. 33), disintegration tissue on back of anthers (no. 42), and more or less elongated testa cells (no. 57). Members of this group also have leaves with lignified epidermal cells (no. 13) and anomocytic stomata (no. 20-0). Both features also occur in Andromeda and in some trees function as synapomorphies. Generic relationships within the Lyonia group are poorly resolved, but Craibiodendron is always sister to Lyonia (Figs. 1, 2), a relationship consistently supported by their homogeneous pith (no. 2-1) and bifacial leaf midrib bundles (no. 11). The Gaultheria group is never monophyletic, although some of these genera, i.e., Gaultheria, Pernettya, Zenobia, Leucothoë, and also Chamaedaphne, are linked in a few cladograms by the base chromosome number of 11 (no. 59). Gaultheria, Pernettya, and Zenobia often form a clade based on the presence of forked anther appendages (no. 40). The monophyly of Leucothoë is equivocal, but L. racemosa and L. fontanesiana are sometimes linked by their inflorescences that are exposed during the winter (no. 23), a feature that also evolved in Pieris (Fig. 2). Finally, although not a major focus in this investigation, we note that Sprengelia incarnata and Prionotes cerinthoides form a clade in all trees that is supported by the lignified leaf epidermis (character no. 13), elongated epidermal cells (no. 14), pedicel with several bracteoles (no. 24-2), and smooth staminal filaments (no. 36-2). These two species represent the epacrid clade (see Crayn et al., 1996; Powell et al., 1996; Kron, 1997; Kron et al., 1998). Molecular data—Analysis of the rbcL data resulted in 42 trees (L 503, CI 0.58, RI 0.50). The strict consensus (Fig. 3) indicates that few clades are well supported including: Vaccinieae (parjack 94), Lyonia (parjack 99), and the Pieris floribunda P. formosa clade (parjack 90). Most of the structure in this tree collapses in trees one step longer. By contrast, the result of the matK analysis is much better resolved, especially within the Lyonia group (Fig. 4). In the matK analysis (four trees obtained, L 980, CI 0.65, RI 0.62) Vaccinieae and members of the Gaultheria group form a clade
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