Structural cladistic study of Cocoseae, subtribe Attaleinae (Arecaceae): Evaluating taxonomic limits in Attaleinae and the neotropical genus Syagrus

Abstract

Syagrus (Areaceae: Cocoseae) is a New World palm genus occurring through most of South America, with one species in the eastern Caribbean. Relationships within Syagrus, Cocoseae and Attaleinae (Arecaceae) are investigated via cladistic analysis of 130 structural characters in 69 ingroup and three outgroup species. The data resolve relationships and test generic limits of Attaleinae and among the Syagrus complex. Maximum parsimony analysis derived the following relationships: 1) two major clades (butioid and syagroid) are resolved; 2) Syagrus is polyphyletic as presently defined, with Allagoptera (incl. Polyandrococos), Cocos, Voanioala, and Attalea nested within it; 3) all genera studied except Syagrus and Butia are monophyletic; and 4) relationships in Syagrus broadly align with phytogeography and leaflet anatomy. The data suggest that Syagrus is polyphyletic, and presently accepted taxonomy may not predict monophyletic groups. The relationships predicted by the structural data differ from those resolved via molecular means in part, and also show some congruence.

Appendix

Characters included in the phylogenetic analysis. Zero was not used to designate a character state because it was incompatible with DELTA (Descriptive Language for Taxonomy).

FRUIT CHARACTERS; 1. Fruit shape. 1 = fruit (excluding beak) globose, subglobose, (length no more than 1.2 times diam); 2 = turbinate; 3 = depressed (shorter than wide); 4 = ovoid to oblong or obovoid; 5 = pear-shaped (but obpyriformis); 6 = ellipsoid (1.25 to 1.4 times diam); 7 = arrowly ellipsoid (greater than 1.6 times diam). 2. Fruit color. 1 = fruit color (when mature) green to light green; 2 = yellowish green; 3 = yellow; 4 = orange; 5 = reddish orange; 6 = red; 7 = brown; 8 = greenish brown; 9 = yellowish brown; 10 = purplish brown; 11 = purplish. 3. Cupule color in fruit. 1 = cupule (persistent perianth) color in fruit, beige, tan or light orange; 2 = light brown; 3 = brown; 4 = greenish brown; 5 = dark brown; 6 = dark reddish or purplish brown; 7 = grayish brown or gray. 4. Cupule diameter. 1 = cupule width or diam 1-2 mm wide; 2 = 5-19 mm wide; 3 = 20-30 mm wide; 4 = 32 or more mm wide. 5. Petal length in mature fruit cupule. 1 = petals shorter than the sepals in the mature fruit cupule; 2 = petals subequal to the sepals; 3 = petals longer than the sepals. 6. Cupule enlargement amount in fruit. 1 = size of calyx and corolla remaining unchanged between flowering and fruiting; 2 = cupule (calyx and corolla) enlarges not more than 1.5-1.8 times more than the original perianth; 3 = cupule enlarges 2-4 or more times the original perianth. 7. Cupule enlargement rate. 1 = corolla and calyx enlarging at the same rate or not enlarging; 2 = corolla enlarging faster than the calyx in fruit; 3 = calyx enlarging faster than the corolla in fruit. 8. Staminodial ring. 1 = staminodial ring with 3-6 staminodes which are sometimes slightly fused at the base producing a 6-lobed ring; 2 = 6 dentate; 3 = undulate; 4 = truncate but often split by the expanding fruit. 9. Epicarp texture. 1 = epicarp smooth; 2 = rough fibrous; 3 = granulate; 4 = longitudinally striate. 10. Epicarp indument. 1 = epicarp glabrous or waxy; 2 = lepidote or scaly; 3 = tomentose; 4 = floccose. 11. Epicarp indument distribution. 1 = distribution of indument on the epicarp, apex only; 2 = upper half; 3 = patches; 4 = scattered or evenly distributed overall. 12. Epicarp thickness. 1 = epicarp thickness about 0.1 mm thick (pliable or fragil when fruit is mature); 2 = 0.3 or more mm thick (hard and durable at maturity). 13. Epicarp and mesocarp dehiscence. 1 = epicarp and mesocarp indehiscent or splitting irregularly; 2 = epicarp and mesocarp dehiscing along 3 vertical sutures. 14. Mesocarp texture. 1 = mesocarp fleshy with few fibers, collapsing into a thin layer over the endocarp when dry; 2 = fleshy fibrous or pulpy remaining as a fibrous mat over endocarp when dry; 3 = fibrous, dry, with very little flesh; 4 = outer zone fibrous and inner zone fleshy; 5 = starchy, dry to moist, and usually white in color with a few fibers; 6 = spongy and oily and usually yellow or orange in color with few to many fibers. 15. Mesocarp color. 1 = mesocarp color white; 2 = yellow; 3 = orange. 16. Endocarp texture. 1 = endocarp texture bony, plasticized, nonporous, nonfibrous, very hard and brittle; 2 = woody, softer, porous, and with many fibers; 3 = bony or woody but very thin, brittle and fragile. 17. Endocarp color. 1 = endocarp tan to light brown; 2 = caramel, yellow brown to brown; 3 = cinnamon, red brown, chocolate to dark brown; 4 = black; 5 = gray. 18. Endocarp wall thickness. 1 = endocarpal wall thickness at apex and base about same thickness as the sides; 2 = apex thickest part; 3 = base thickest part; 4 = apex and base about same thickness, both thicker than sides. 19. Endocarp shape. 1 = endocarp shape obloid or depressed, with a length to width 5:6 (0.8) ratio; 2 = spheroid or very broad, 1:1 (1.0) ratio; 3 = broad, 6:5 (1.2) ratio; 4 = normal, 3:2 (1.5) to 2:1 (2.0) ratio; 5 = narrow, 3:1 (3.0) ratio. 20. Endocarp widest portion. 1 = endocarp widest near the middle (obloid, spheroid, ellipsoid); 2 = widest near the apex or top (obovoid); 3 = widest near the base (ovoid). 21. Endocarp upper lateral exterior surface texture. 1 = endocarp upper lateral exterior surface smooth or nearly so; 2 = surface shallowly grooved; 3 = surface irregular and deeply grooved; 4 = surface irregular and pitted. 22. Endocarp fibers on lower third. 1 = endocarp fibers absent or only scattered fibers on the endocarp surface and those on the lower third easily removed and never forming a persistent "skirt"; 2 = fibers strands forming a persistent "skirt" on the lower third of the endocarp which can not be easily removed. 23. Endocarp apex (beak) shape. 1 = endocarp apex with beak present as a gradually tapered circular to triangular protuberance; 2 = beak present as 3-lobed structure (sutures always alternate with the lobes in this state); 3 = beak present as a broad slightly raised umbonate shield; 4 = beak present as a persistent part of the fruit beak fused to the endocarp, often (but not always) with the most difficult to remove fibers at the apex; 5 = beak flat.24. Suture behavior at endocarp apex. 1 = sutures meeting at apex; 2 = meeting at apex but interrupted by a beak on the apex; 3 = stopping short of the apex (often as ridges); 4 = indistinguishable at the apex or even short of the apex. 25. Suture behavior on lateral exterior endocarp surface. 1 = endocarp sutures on the lateral exterior surface visible as a shallow to very distinct ridge; 2 = visible as a shallow ridge with a shallow furrow or only a furrow; 3 = visible as a discoloration and usually also the presence of coarser fibers and sometimes a slight furrow near the apex; 4 = indistinguishable from the rest of the endocarp surface. 26. Endocarp pores location. 1 = endocarp pores basal; 2 = located halfway between the middle and the base; 3 = located at or near the middle; 4 = just above the middle; 5 = apical. 27. Endocarp pores even or sunken. 1 = endocarp pores almost even with surface of the endocarp; 2 = pores slightly sunken below the normal surface of the endocarp. 28. Endocarp interior shape and surface texture. 1 = endocarp interior cavity smooth or roughened and circular or nearly so; 2 = smooth and triangular; 3 = irregular and often invading the margins of the endosperm; 4 = irregular and invading the margins of the endosperm slightly but especially at the pores. 29. Endocarp interior markings. 1 = endocarp interior cavity markings absent or not clearly apparent; 2 = monovittate (with one dark vertical band); 3 = divittate (2 vertical bands); 4 = trivittate (3 vertical bands). 30. Endocarp embedded fibers. 1 = endocarp fibers within the endocarp matrix absent or indistinct; 2 = clearly visible as darker or lighter colored scattered fibers. 31. Seed number. 1 = normal seed number of 1; 2 = normal number of 2 or more. 32. Seed shape. 1 = seed globose to subglobose; 2 = ovoid to obovoid; 3 = turbinate; 4 = depressed; 5 = elliptical; 6 = narrowly elliptical; 7 = irregular from invaded endocarp. 33. Seed free or adhering to pericarp. 1 = seed free from pericarp; 2 = adhering to the pericarp. 34. Endosperm texture. 1 = endosperm homogeneous; 2 = homogeneous sometimes penetrated by the testa; 3 = ruminate. 35. Endosperm central cavity. 1 = endosperm without a central cavity; 2 = endosperm with a central cavity. 36. Germination type. 1 = germination adjacent-ligular; 2 = remote-tubular. 37. Eophyll (first leaf) shape. 1 = eophyll (first leaf) simple lanceolate or ovate and undivided; 2 = bifid; 3 = pinnate. INFLORESCENCE CHARACTERS 38. Prophyll indument. 1 = prophyll glabrous to waxy; 2 = lepidote or scaly, farinose; 3 = tomentose; 4 = spiny. 39. Peduncular bract outer surface. 1 = peduncular bract smooth on the outer surface; 2 = sulcate on the outer surface. 40. Peduncular bract indument. 1 = peduncular bract glabrous to glaucous waxy; 2 = lepidote or scaly, farinose; 3 = tomentose; 4 = spiny. 41. Inflorescence sexuality. 1 = palm usually produces bisexual inflorescences, and rarely staminate inflorescences (under adverse or stressful conditions?); 2 = palm usually produces unisexual or androdioecious inflorescences with only staminate flowers, or with pistillate flowers and aborted non-functioning staminate flowers, or rarely, androgynous with functional staminate flowers and a few (fertile?) pistillate flowers. 42. Inflorescence branching. 1 = inflorescence branched to one order but basal branches often branched to two orders; 2 = branched to one order only; 3 = spicate. 43. Peduncular bract indument. 1 = peduncle glabrous or glaucous waxy; 2 = lepidote or scaly; 3 = hairy or tomentose; 4 = spiny. 44. Rachis indument. 1 = rachis glabrous or glaucous waxy; 2 = lepidote, scaly or farinose; 3 = tomentose. 45. Rachis vs. peduncule length. 1 = rachis shorter than the peduncle; 2 = about equal to the peduncle; 3 = longer than the peduncle. 46. Rachillae indument. 1 = rachillae glabrous or waxy; 2 = lepidote, thin indumentose, mealy, or farinose; 3 = sparsely hairy to villous tomentose; 4 = dense tomentose to woolly tomentose. 47. Rachis bract shape. 1 = rachis bracts deltoid; 2 = laterally broadened. 48. Staminate vs. pistillate flower portion of rachillae. 1 = Staminate flower (SF) portion shorter than pistillate flower (PF) portion; 2 = SF portion equals PF portion; 3 = SF portion longer than PF portion. STAMINATE FLOWER CHARACTERS: 49. Staminate flower color. 1 = flowers white or cream; 2 = pale yellow; 3 = bright yellow to dark yellow; 4 = orange; 5 = maroon to purple. 50. Staminate flower attachment. 1 = flowers sessile; 2 = stalked (at least some). 51. Staminate sepal shape. 1 = sepals lanceolate or linear; 2 = triangular or deltoid. 52. Staminate sepal arrangement. 1 = sepals imbricate; 2 = briefly connate; 3 = connate in a stalk-like base. 53. Staminate petal shape. 1 = petals lanceolate; 2 = ovate-lanceolate; 3 = ovate; 4 = obovate; 5 = narrow, needle-like, terete in cross-section. 54. Staminate petal arrangement. 1 = petals imbricate; 2 = imbricate at the base but valvate at the apices; 3 = valvate; 4 = connate. 55. Staminate petal texture. 1 = petals membranous to coriaceous; 2 = sclerenchymous to fleshy 56. Staminate petal indument. 1 = petals glabrous; 2 = scaly, lepidote or dotted; 3 = tomentose; 4 = patches of ramenta on the veins; 5 = tiny tubercles. 57. Staminate petal venation. 1 = petals with venation conspicuous; 2 = venation inconspicuous or obscure. 58. Staminate tip shape. 1 = petal tip obtuse to emarginate; 2 = broadly acute to acute; 3 = attenuate; 4 = acuminate. 59. Staminate number. 1 = stamens 6 in number; 2 = 7 or more in number. 60. Anther filament connation. 1 = filaments distinct; 2 = very briefly connate at base. 61. Anther filament orientation. 1 = filaments inflexed; 2 = not inflexed, straight or reflexed (bending outward). 62. Anther filament attachment. 1 = anthers dorsifixed; 2 = medifixed and versatile; 3 = basifixed. 63. Pistillode shape. 1 = pistillode trifid; 2 = 4 or more -fid; 3 = simple, bifid, or reduced to a trilobed knob. PISTILLATE FLOWER CHARACTERS: 64. Pistillate vs. staminate flower size. 1 = pistillate flowers slightly smaller than staminate flowers; 2 = equal or subequal to the staminate flowers; 3 = larger than the staminate flowers. 65. Pistillate flower shape at late bud stage. 1 = flowers globose or rounded; 2 = ovate or ovoid; 3 = pyramidal or conical; 4 = oblong. 66. Pistillate flower color. 1 = flowers greenish white to white; 2 = pale or light yellow; 3 = bright yellow to dark yellow; 4 = orange; 5 = reddish brown; 6 = maroon; 7 = purple to violet; 8 = green to taffy 67. Basal pistillate flower attachment. 1 = flowers at the base of the basal rachillae sessile; 2 = flowers stalked (pedicellate). 68. Pistillate sepal venations. 1 = sepal nerves distinctly visible; 2 = nerves indistinct or faintly visible; 3 = no nerves visible at all, a perfectly smooth surface. 69. Pistillate sepal texture. 1 = sepals sclerophyllous to fleshy; 2 = membranaceous to coriaceous. 70. Pistillate sepal indument. 1 = sepals glabrous or nearly so or glaucous waxy; 2 = lepidote, scaly, mealy; 3 = tomentose. 71. Pistillate sepal arrangement. 1 = sepals imbricate; 2 = valvate. 72. Pistillate sepaltip shape. 1 = sepal tips attenuate; 2 = acuminate; 3 = acute to apiculate; 4 = broadly acute to obtuse; 5 = rounded. 73. Pistillate sepal tip keeled. 1 = sepal tips keeled; 2 = sepals not clearly keeled. 74. Pistillate petals vs. sepal size. 1 = petals slightly shorter than sepals; 2 = nearly equal to the sepals; 3 = slightly longer than sepals. 75. Pistillate petal arrangement. 1 = petals valvate; 2 = imbricate; 3 = imbricate at base, valvate at apex; 4 = connate at the base, valvate at apex. 76. Pistillate petal shape. 1 = petals triangular; 2 = ovate to oblong. 77. Pistillate petal venation. 1 = petals distinctly nerved; 2 = faintly, obscurely or indistinctly nerved; 3 = no nerves visible, completely smooth. 78. Pistillate petal indument. 1 = petals glabrous or glaucous waxy; 2 = lepidote or scaly; 3 = tomentose. 79. Pistillate petal tip shape. 1 = petal tip acute; 2 = rounded or broadly obtuse. 80. Gynoecium shape. 1 = gynoecium cylindrical; 2 = ovoid; 3 = globose; 4 = cone-like; 5 = obpyriform, pear-shape. 81. Gynoecium indument. 1 = gynoecium glabrous; 2 = lepidote or scaly; 3 = tomentose; 4 = woolly; 5 = granulate or pilose. 82. Gynoecium indument distribution. 1 = gynoecium with only a few hairs near the upper edge of the staminodial ring, and often hidden by it; 2 = indument visibly covering the lower half including that which is under the staminodial ring; 3 = visibly covering all or at least two-thirds or only covering the upper half. 83. Gynoecium locule number. 1 = syncarpous gynoecium with 3 equially developed locules; 2 = more than 3 equally developed locules (up to 10); 3 = pseudomonomerous with 1 fertile locule. 84. Stigma branches. 1 = stigmas 3 distinct branches; 2 = stigma reduced 3-lobed; 3 = stigma one or indistinct. 85. Stigma shape. 1 = stigmas erect; 2 = recurved; 3 = (rarely) indistinct. 86. Stigma style indument. 1 = style glabrous; 2 = lepidote, scaly, or very thin indument; 3 = tomentose; 4 = white woolly. 87. Stigma connation. 1 = styles distinct; 2 = connate. 88. Staminodial ring shape. 1 = staminodial ring irregular with 3, 4-6 projections or staminodes; 2 = 3-6-dentate; 3 = undulate; 4 = smooth and truncate; 5 = ragged fringed edge and undulate or truncate. VEGETATIVE CHARACTERS: 89. Stem clustering vs. solitary. 1 = stem clustering (axillary shoots borne at base); 2 = stem solitary. 90. Stem erect vs. prostrate or subterranean. 1 = stem tall and erect with a visible above ground stem; 2 = short, prostrate, or subterranean stem. 91. Leaf base persistence. 1 = leaf bases persistent along the entire length of the trunk; 2 = leaf bases persistent for over 2/3 the length of the trunk; 3 = leaf bases persistent for 1/3 to 1/10 the length of the trunk; 4 = neatly abscising. 92. Trunk ornamentation. 1 = trunk ornamentation after the leaf bases have abscised exposing closely packed rough corky leaf scars with no or little apparent internodal areas; 2 = rough corky leaf scars with clearly visible obliquely ringed leaf scars, and internodal areas distinct but rough; 3 = irregular rough trunk with no prominent rings; 4 = smooth trunk with obliquely ringed leaf scars and very distinct internodes; 5 = smooth trunk with inconspicuous scars and internodes; 6 = spiny trunk with emergent cortical bundles. 93. Leaf attachment arrangement. 1 = leaves arranged spirally; 2 = in five (often vertical) rows. 94. Leaf demeanor. 1 = leaves spreading; 2 = erect-arching or ascending; 3 = stiffly erect. 95. Leaf sheath type. 1 = leaf sheath with warp and weft strands that persist within a friable, splitting and discontinuous membrane composed of chartaceous (stiff papery pieces) of old epidermis and mesophyll; 2 = warp and weft fibers persist but epidermis and mesophyll disintegrate resulting in a flexible but strong "burlap sack" type of sheath; 3 = warp strands persisting the longest but too friable or brittle (not woody enough) to leave anymore than a simple ragged margin on the edge of a very long pseudopetiole (apparent petiole), i.e. S. romanzoffiana; 4 = warp strands thick, flattened and sometimes woody, weft strands barely noticeable, thin, brittle and disintegrate along with a thin epidermis and mesophyll, resulting in regularly spaced persistent flat, stiff warp strands along the margin of the pseudopetiole, i.e. S. coronata; 5 = some of the warp strands are modified into thick-based, hook-like spines. Weft strands are very thin and disintegrate along with a very thin epidermis and mesophyll, resulting in persistent, regularly spaced, thick-based and spine-like warp fibers along the margins of the pseudopetiole, i.e. S. schizophylla and many species of Butia. 96.Sheath and petiole indument on abaxial surface. 1 = sheath and petiole glabrous, glaucous or waxy on the abaxial surface; 2 = lepidote or with short indument or scales; 3 = thick tomentose, hairy or woolly hairy; 4 = spiny as well as tomentose. 97. Petiole width just below leaf blade. 1 = true petiole width or width of pseudopetiole just below the leaf blade 0.3–1.5 cm wide; 2 = 1.5–3 cm wide; 3 = 3–4.5 cm wide; 4 = 5 cm or more wide. 98. Petiole adaxial surface shape. 1 = petiole (upper cross-sectional surface) adaxially channeled only; 2 = adaxially channeled with a middle ridge; 3 = adaxially flattened; 4 = adaxially arched or rounded. 99. Leaf segment abaxial surface indument. 1 = leaf segment abaxial surface other than midvein glabrous and green; 2 = abaxial surface with obvious coating of wax; 3 = abaxial surface tomentose or hairy. 100. Abaxial nerve indument. 1 = blade and nerves glabrous; 2 = abaxial nerves with ramenta only at the base of the nerve or extending up the nerve for a short distance; 3 = with ramenta at the base of the nerve and as scattered patches (at least 3) extending at least part way up the length of the nerve; 4 = with continuous ramenta or hairs all along the length of the nerve for most of its length, except near the tip. 101. Transverse veinlets. 1 = transverse veinlets or cross veins on leaflet segments obscure or inconspicuous; 2 = conspicuous, usually somewhat raised adaxially. 102. Adaxial leaf color. 1 = upper, adaxial surface dark green or dark bluish green; 2 = medium or bright green; 3 = light green or yellow green or grayish to silvery green. 103. Leaflet arrangement along the rachis. 1 = leaflets regularly distributed along the rachis; 2 = in clusters along the lower 1/3 or lower 1/2 of the rachis and evenly distributed on the upper part; 3 = in loose clusters along the rachis; 4 = in fairly distinct or tight clusters along the rachis. 104. Leaflet arrangement one side vs. other side. 1 = two sides of leaf forming a strong V-shape; 2 = two sides of leaf forming a very open V-shape; 3 = two sides of leaf lie in almost the same plane. This character is only applied if leaf segments along one side of the midrib are arranged in one plane. 105. Leaflet segment tip shape. 1 = leaflet tips attenuate; 2 = acuminate; 3 = acute; 4 = obtuse. 106. Leaflet segment tip notching. 1 = condition of leaf segment tip symmetrical and not split; 2 = symmetrical and shallowly bifid or notched; 3 = asymmetrical and bifid; 4 = asymmetrical and not spit, oblique. VEGETATIVE ANATOMICAL CHARACTERS: 107. Vascular bundle levels. 1 = one level of smaller vascular bundles present; 2- two levels present. 108. Vascular bundle arrangement. 1 = each vascular bundle arrangement level with one type of similar-looking vascular bundles; 2 = each level with two distinctly different-looking bundles (either in size, shape or attachment). 109. Adaxial-abaxial girders present. 1 = Adaxial-abaxial girders almost always present; 2 = almost always absent. 110. Vascular bundles attached by abaxial girders. 1 = some vascular bundles attached by abaxial girders; 2 = none attached in this manner. 111. Adaxial strands present. 1 = adaxial strands almost always present; 2 = almost always absent 112. Adaxial strand present and location of attachment. 1 = adaxial strands when present mostly attached to the epidermis; 2 = adaxial strands when present mostly attached to the hypodermis; 3 = adaxial strands when present mostly unattached, separated by other layers of cells. 113. Adaxial strand vs. smallest vascular bundle size. 1 = adaxial strands when present mostly smaller than the majority of the secondary abaxial veins; 2 = adaxial strands when present mostly larger than the majority of the secondary abaxial veins. 114. Adaxial strand shape. 1 = adaxial strands when present usually forming a flat almost continuous to briefly interrupted layer, 1 to 2-cells thick; 2 = when present few-celled irregularly shaped stands mostly 1-3 cells thick; 3 = when present usually many-celled strands broadly triangular, broadly parabolic, or squarish, usually as wide or wider than long; 4 = when present usually many-celled strands narrowly triangular, parabolic, or elongate-shaped, several cells longer than wide. 115. Smaller vascular bundles with abaxial girders. 1 = some smaller veins attached by adaxial girders; 2 = none attached in this manner. 116. Attachment of girderless vascular bundles. 1 = most smaller girderless veins attached to epidermis or hypodermis; 2 = only every other vascular bundle attached in this manner; 3 = most unattached. 117. Abaxial strand presence and attachment. 1 = abaxial strands when present mostly attached to the epidermis or hypodermis; 2 = abaxial strands when present mostly unattached, separated by other layers of cells; 3 = abaxial strands usually absent. 118. Largest vascular bundle with girders. 1 = largest veins mostly with girders; 2 = largest veins more or less alternating veins with and without girders; 3 = mostly without girders. 119. Largest vascular bundle attachment. 1 = largest vein mostly attached to both sides of leaf; 2 = mostly attached adaxially; 3 = mostly attached abaxially; 4 = mostly unattached. 120. Cuticle thickness. 1 = cuticle almost as thick to thicker than the epidermis; 2 = not so, thinner than the epidermis. 121. Hypodermis thickness. 1 = adaxial hypodermis one layered; 2 = adaxial hypodermis two layered. 122. Palisade layer differentiated. 1 = palisade layer conspicuously differentiated; 2 = not so. 123. Palisade layer with fiber strands. 1 = free (unattached) fiber strands present in palisade; 2 = no or rare free strands present. 124. Submarginal vein. 1 = submarginal vein conspicuous, much larger than normal secondary veins; 2 = submarginal vein absent, inconspicuous; 3 = equal to or smaller than secondary veins. 125. Adjacent submarginal vein (second in from the margin). 1 = adjacent submarginal vein, large and conspicuous; 2 = not conspicuous, smaller. 126. Large submarginal adaxial strand. 1 = large submarginal adaxial strand present and larger than secondary veins; 2 = large strand absent. 127. Cross-section of margin. 1 = cross section of margin slanted towards the abaxial side; 2 = slanted towards the adaxial side; 3 = rounded; 4 = squared off. 128. Marginal reddish callus. 1 = marginal reddish callus present at or close to the edge of the margin; 2 = absent. 129. Margin notch. 1 = margin not notched or swollen; 2 = notched or swollend adaxially; 3 = notched or swollen abaxially. 130. Marginal hypodermal cell layers. 1 = cells of hypodermal layers enlarge, swell, and sometimes become 2-3 layered just before the margin and often encompass the marginal strand fibers; 2 = hypodermal cells unchanged near margin or not as described.