Below, we present a summary of 182 comparative anatomical, morphological, cytological, and related characters that have been proposed useful (e.g., Zimmerman 1985; Terrazas and Arias 2002) in the systematic analysis of the subfamily Cactoideae, Tribe Cacteae. Our goals are to 1) provide accurate comparative data for members of Tribe Cacteae, 2) examine the inferential ability of these comparative characters, relative to comparative DNA sequence data, in parsimony and maximum likelihood analyses. Character scores for morphological traits will be based on comparative, revisionary and monographic studies (Britton and Rose 1919-1923; Bravo 1978; Gibson and Horak 1978; Gibson 1989a, 1989b, 1991; Taylor and Zappi 1989; Bravo and Sanchez-Mejorada 1991; Taylor 1991; Zappi 1994; Barthlott and Taylor 1995; Arias et al. 1997; Nyffeler 1998; Arreola 2000; Barthlott and Hunt 2000; Leuenberger and Eggli 2000; Anderson 2001) and augmented by the observations of the collaborative group.

 

  1. Stem shape I (subjective form): 0 = cylindroid above ground, the subterranean portion relatively small; 1 = intermediate; 2 = short, the above ground part hemispherical or flat-topped, the stem largely subterranean.
  2. Stem shape, II (subjective ridgidness): 0 = stiff, firm, erect or potentially so; 1 = intermediate; 2 = soft, flaccid, decumbent or potentially so. [Species-specific, frequently-used trait taxonomically, potentially difficult to establish owing to seasonal differences in turgor]
  3. Diameter (at 1/2 length) of sexually mature and age-specific stems: Continuous variable, 1–1250 mm.
  4. Length of sexually mature and age-specific plants: Continuous variable, 5–5000 mm
  5. Number of mature stems per adult plant: Continuous variable, 1 to ± 100. Members of Cacteae generally reach sexual maturity with only one mature stem per plant, thus age-class distribution within a given population will greatly assessment of branching.
  6. Number and percentage of immature stems per plant: Continuous variables, 1 to ± 100, and 0–100 %.
  7. Stem shape, III: Sharply-defined narrow stipe at junction between stem and root system; 0 = no, 1 = yes (e.g., “Gymnocactus” spp. and some “Echinomastus” spp.)
  8. Maximum root diameter, and percentage of stem diameter: Continuous variables, 1–80 mm, ± 5% to nearly 100%. [Edaphics may influence the root morphology; plants grown from seed in cultivation may not be representative.]
  9. Roots distended with mucilage cells, and more or less parallel to the soil surface: 0 = no, 1 = yes (e.g., Ancistrocactus, Coryphantha).
  10. Root vascular system: 0= fibrous wood, 1 = wide band tracheids and fibrous wood, 3 = only wide band tracheids
  11. Pith diameter, as percentage of lesser stem diameter (excluding tubercles/ribs) and absolute measure: Continuous variables, ±5%–±50%. All seedlings begin with narrow pith, thus there is expected to be an ontogenetic effect.
  12. Medullary vascular system: 0 = absent, 1 = present. Seedlings initially lack a medullary vascular system.
  13. Woodiness: 0 = very woody; 1 = intermediate; 2 = scarcely woody, the stele remaining a loose reticulum of separate vascular bundles.
  14. Stem vascular system: 0= fibrous wood, 1 = wide band tracheids and fibrous wood, 3 = only wide band tracheids
  15. Mucilage cells in pith and cortex: 0 = absent, 1 = rarely present/very sparse, 2 = present/abundant.
  16. Laticifer development: 0 = none, 1 = subaxillary thin-walled regions in cortex, 2 = subaxillary ducts with semi-milky latex, 3 = ducts mostly subaxillary, but latex milky, 4 = ducts widespread in cortex and often in pith, slender, the latex milky. Laticifers in Cactaceae, restricted to Mammillaria.
  17. Mucilage ducts: 0 = none, 1 = present (e.g., Ariocarpus).
  18. Mucilaginous periderm: 0 = none, 1 = present (e.g., Neolloydia)
  19. Apparent color of epidermis: 0 = dark green, 1 = olive-green or grass-green, 2 = gray-green or blue-green, 3 = pale blue or gray. (The epidermis itself is transparent, thus surface color is the combined influence of the cuticle, hypodermis and underlying chlorenchyma.
  20. Stomata deeply recessed: 1 = unrecessed, 2 = moderately recessed (e.g., Ariocarpus fissuratus), 3 = strongly recessed (e.g., Ariocarpus agavoides).
  21. Epidermal cell wall thickness: 0 = thin, 1 = moderate, 2 = thick and with other modifications (e.g., Leuchtenbergia).
  22. Epidermis cell size (relative to hypodermis, in cross-section): 0 = smaller than hypodermal cells, 1 = equal to hypodermal cells, 2 = larger than hypodermal cells.
  23. Epidermal papillae: 0 = cells flat, 1 = cells bulging, 2 = cells each bearing a central papilla.
  24. Cuticle thickness: continuous character; 0 = normal (less than or equal to the epidermal cell wall thickness), 1 = immense (thicker than the entire epidermis; e.g., Ariocarpus spp.).
  25. Epidermis puffy-looking. watery, easily bruised: 0 = no, 1 = yes (e.g., Ortegocactus, Neolloydia matehualensis).
  26. Hypodermis, number of cell layers: 0 = none, or poorly differentiated, 1 = one, strongly differentiated, 2 = two to four, 3 = four to many (e.g., Ferocactus spp.). Seedlings and greenhouse-grown plants often have a poorly differentiated hypodermis, regardless of the normal condition for adult plants, thus cultivated plants are unreliable sources of this feature.
  27. Hypodermis primary cell wall thickness: 0 = cells thin-walled, 1 = cells moderately collenchymatous, 2 = cells strongly collenchymatous. Seedlings and greenhouse-grown plants often have a poorly differentiated hypodermis, regardless of the normal condition for adult plants, thus cultivated plants are unreliable sources of this feature.
  28. Druses large and lenticular in pith and cortex: 0 = no, 1 = yes.
  29. Abundance of druses in hypodermis: 0 = none, I = few or unpredictable, 2 = abundant in every cell layer, 3 = abundant in upper cell layer, absent from others.
  30. Abundance of druses in cortex: 0 = more numerous than in hypodermis, 1 = about as abundant as in the hypodermis, 2 = fewer than in the hypodermis.
  31. Types of crystals or crystal aggregates in hypodermis: 0 = spheroidal druses, 1 = single large prismatic crystals, 2 = numerous tiny crystals. (e.g., Anderson and Boke 1969).
  32. Tubercle length (= rib height): Continuous variable, 0.5–150 mm. (This is the distance that the podarium protrudes, not the length of its basal abaxial-adaxial axis. In most species, podarium length is ± perpendicular to the longitudinal axis of the stem.)
  33. Tubercle diameter (= rib width): Continuous variable, 2 to ± 40 mm (This distance should be taken mid-way between the base and apex of a tubercle (half-way to the crest of rib), not at the poorly-defined base of a podarium.
  34. Podarium decurrent or protuberant: 0 = strictly protuberant, 1 = slightly decurrent, appearing erect, 2 = strongly decurrent, 3 = extremely decurrent (e.g., Stenocactus). Seedlings of all of the ribbed taxa are tuberculate.
  35. Podarium ribbed versus tuberculate: 0 = tubercles, 1 = crenate or notched ribs, 2 = entire ribs.
  36. Inter-areolar distance: Continuous variable, used thus far only for ribbed taxa: 0 to ± 40 mm between areoles along the crest of a rib. (Measurement should be taken between the centers of the areoles.)
  37. Podarium shape acute or rounded: 0 = acute tubercles or carinate ribs, 1 = obtuse tubercles or obtuse ribs, 2 = broadly rounded low tubercles or low rounded ribs. (Referring to the shape of the podaria as seen in a stem-section.)
  38. Podarium angular cross-section: 0 = terete, 1 = weakly angular, 2 = sharply triangular, 3 = triangular, and with additional modifications (e.g., Ariocarpus spp.). (Applicable to ribbed taxa not evident.)
  39. Phyllotaxy: 0 = areoles in 3-and-5 or 5-and-8 spirals (as viewed from the apex or from the sides, respectively); 1 = areoles in 5-and-8 or 8-and-13 spirals; 2 = areoles in 8-and-13 or 13-and-2l spirals; 3 = areoles in 13-and-21 or 21-and 34 spirals, or even more numerous.
  40. Number of ribs: Continuous variable, from 4 to ± 120. (Not applicable to tuberculate taxa; may be correlated with phyllotaxy.
  41. Development of areolar trichomes: 0 = unusually short and/or caducous, 1 = average, 2 = longer than average and/or long-persistent, 3 = extremely long and conspicuous.
  42. Color of areolar trichomes: 0 = white, I = cream-colored, 2 = tan.
  43. Stellate trichomes present on epidermis: 0 = lacking, 1 = present (e.g., Astrophytum).
  44. Basal outline of spine-cluster: 0 = circular or broadly elliptical, 1 = narrowly elliptical, 2 = linear.
  45. Basal length of spine-cluster: Continuous variable, ±0.5–20 mm.
  46. Areole morphology: Three-dimensional array of qualitative character-states, presumably best coded as several linear characters: 0 = Acharagma-type, 1 = Ferocactus-type, 2 = Mammillaria-type, 3 = Ortegocactus-type, 4 = Protomammillaria-type, 5 = Escobaria-type, .6 = Coryphantha macromeris-type, 7 = Neolloydia-type, 8 = Pachycereus-type, 9 = Ariocarpus-type.
  47. Areolar gland presence/absence: 0 = always present and conspicuous, 1 = potentially present but always sparse and ephemeral, 2 = apparently never present.
  48. Areolar gland morphology: 0 = peg-shaped (similar to ordinary spines), 1 = capitate.
  49. Axillary spines presence/absence: 0 = absent, 1 = very few or seldom present, 2 = always present and conspicuous. (This character referrs to typical spines, not glands, and presence is restricted to Mammillaria.)
  50. Axillary spines morphology: 0 = acicular, I = filiform. (only in Mammillaria)
  51. Central spines appressed: 0 = none appressed, 1 = several not appressed, 2 = all but the principal inner central spine appressed, 3 = all including the principal inner central spine appressed.
  52. Radial spines appressed: 0 = not appressed, 1 = weakly appressed or only half of them appressed, 2 = average (e.g., Coryphantha), 3 = very tightly appressed, in contact with the surface of the underlying podarium (e.g., Pepecyphora).
  53. Spine-epidermis vestiture of adults: 0 = none (epidermal cells flat or concave), 1 = cells having short papillae, 2 = cells with long papillae, 3 = cells with modified papillae (branched, or extremely long, flattened, or restricted to one surface of the spine).
  54. Spine-epidermis vestiture of seedlings: 0 = none, 1 = slight, 2 = conspicuous, 3 = plumose or modified.
  55. Radial spine cross-section, 1: 0 = terete, 1 = angular or striate, 2 = dorsiventrally compressed but not papery, 3 = dorsiventrally compressed, papery, 4 = laterally compressed.
  56. Spines secondarily absent: 0 = no, 1 = almost (spines of adults rudimentary), 2 = yes. Surfaces of largest spines appearing sponge-like: 0 = no, 1 = yes. (Requiring scanning electron microscopy; e.g., Turbinicarpus schmiedickeanus.)
  57. Curvature of principal central spines: 0 = straight, 1 = slightly curved, 2 = strongly curved.
  58. Curvature of principal central spines tip: 0 = tip straight, I = tip slightly curved, 2 = tip strongly curved (spine hooked).
  59. Spines, or some of them, hairlike: 0 = no, 1 = yes.
  60. Spines blunt: 0 = no, 1 = yes.
  61. Spines of seedlings unusually wide and basally constricted: 0 = no, 1 = yes. (e.g., Coryphantha.)
  62. Number of inner central spines per areole: Meristic character, 0–12 (The mean per plant is less variable, and the population mean could be used as a continuous variable).
  63. Number of outer central spines per areole: Meristic character, 0–23 (The mean per plant is less variable, and the population mean could be used as a continuous variable).
  64. Number of radial spines per areole: Meristic character, 0–120 (The mean per plant is less variable, and the population mean could be used as a continuous variable).
  65. Total spines number per areole: Meristic character, 0–130 (Redundant but useful because it includes sub-central spines, a minor class not included above, and it does not require interpretation of spines as centrals or radials.)
  66. Number of central spines per areole on seedlings: Meristic character, 0–5+.
  67. Number of radial spines per areole on seedlings: Meristic character, 0–20+.
  68. Central spine arrangement: 0 = radiating arrangement, 1 = intermediate or variable, 2 = bird's-foot arrangement.
  69. Length of shortest inner central spine per areole: Continuous variable, 0–140 mm
  70. Length of longest inner central spine per areole: Continuous variable, 0–170 mm
  71. Length of shortest outer central spine per areole: Continuous variable, 2–80 mm
  72. Length of longest outer central spine per areole: Continuous variable, 5–160 mm.
  73. Location of longest radial spines per areole: 0 = upper, 1 = ambiguous, lateral, or variable, 2 = lower.
  74. Length of longest radial spine per areole: Continuous variable, 3–40 mm.
  75. Maximum radial spine thickness per areole: Continuous variable, 0.01–4 mm
  76. Minimum radial spine thickness per areole: Continuous variable, 0.01–1.5 mm
  77. Maximum central spine thickness per areole: Continuous variable, 0.1–8 mm
  78. Spine color, radials contrasting centrals: 0 = no, I = moderately, 2 = strongly, the radials much whiter than the centrals.
  79. Spine color, central spine pigmentation: 0 = bright white, 1 = dull white, 2 = pale but colored, 3 = brightly colored, 4 = dark, 5 = black. (RGB color values from standardized images could be used.)
  80. Spine color, red/yellow dimorphism in central spines: 0 = red, 1 = yellow, 2 = intermediate, 3 = absent.
  81. Spine color, extent of dark tipping: 0 = none; 1 = only on larger spines; 2 = on both large and medium spines; 3 = on large, medium, and some radial spines; 4 = on all spines.
  82. Flower position: 0 = apical (produced from new growth), I = intermediate (apical but produced from year-old tubercles, or subapical), 2 = lateral (produced from older growth). (This is in part a phenological character. Note that lateral fruits do not necessarily imply lateral flowering, since in some species the remains of the flowers are displaced outward by vegetative growth from the stem apex during the two to twelve months required for fruit maturation.
  83. Flower diameter: Continuous variable, 3–90(–120) mm. Note that flower diameter is measured from floral bract-tip to floral bract-tip, when the flower is at its maximum extent.
  84. Flower length: Continuous variable, 6–65 mm.
  85. Pericarpel shape at anthesis: 0 = cylindroid, 1 = funnelform, 2 = spheroidal.
  86. Pericarpel constriction at anthesis: 0 = demarcated from the receptacular tube by a slight constriction, 1 = no external demarcation from the receptacular tube.
  87. Shape of receptacular tube: 0 = broad (campanulate or funnelform), 1 = narrow (narrowly funnelform), 2 = very narrow (tubular).
  88. Walls of receptacular tube: Continuous variable, 0.5–12 mm thick; 0 = very thick and stem-like (=10 mm), 1 = average (1–2 mm thick), 2 = thin, colorful, corolla-like (=1 mm).
  89. Floral symmetry: 0 = actinomorphic, 1 = weakly zygomorphic
  90. Total length of receptacular tube: Continuous variable, 4.5–22 mm long
  91. Length of nectar-chamber: Continuous variable, 0–16 mm. “Length” is the vertical depth of the chamber, not the extent of the secretory region itself.
  92. Length of sterile distal portion of receptacular tube: Continuous variable or percentage of total tube length, 0–18 mm, or 0–79 %
  93. Rudimentary bracts (scales) on pericarpel and receptacular tube: 0 = uniformly dense, 1 = sparse but some present, 2 = absent from pericarpel, 3 = absent from both pericarpel and most of receptacular tube.
  94. Number of series of sepaloid bracts: Rather arbitrarily defined meristic character, from 1 (e.g., Epithelantha micromeris) to ±12.
  95. Number of series of petaloid bracts: Meristic character, 1–3.
  96. Shape of scales on pericarpel: 0 = broad and rounded, 1 = deltoid, 2 = lanceolate or ovate, 3 = subulate or linear (including spine-tipped bracts of Echinocactus), 4 = rudimentary.
  97. Number of sepaloid bracts per flower: Meristic character, ±3–60. (Traditionally, this count excludes proximal receptacular scales that are shorter than wide or that otherwise intergrade with the sepal-like distal bracts. This lends a highly subjective element to this count.
  98. Shapes of sepaloid bracts: 0 = lanceolate, 1 = oblong, 2 = oblanceolate, 3 = spatulate.
  99. Apex of the sepaloid bracts, general shape: 0 = attenuate, 1 = acute, 2 = obtuse, 3 = rounded or erose.
  100. Apex of sepaloid bracts, extreme apices: 0 = acute, I = obtuse, 2 = rounded or truncate, 3 = retuse or emarginate.
  101. Tips of sepaloid bracts, centers of extreme apices: 0 = entire, 1 = mucronate, 2 = caudate.
  102. Spinose tips on sepaloid bracts and scales of pericarpel: 0 = absent, 1 = present (e.g., Echinocactus, Homalocephala, and Astrothytum).
  103. Pigmentation of sepaloid bracts: 0 = similar to receptacular tube, 1 = like receptacular tube except having brightly colored midstripes, 2 = brightly colored throughout, contrasting with receptacle. (This can be quantified by using RGB values of standardize images.)
  104. General color of sepaloid bracts: 0 = pinkish or reddish; 1 = brown, or mixed red-and-yellow, or dimorphic; 2 = yellowish. (This can be quantified by using RGB values of standardize images.)
  105. Margins of bracts: 0 = all entire or sporadically erose, 1 = outermost bracts consistently erose or denticulate, 2 = all sepaloid bracts sparsely fimbriate, 3 = all sepaloid bracts densely fimbriate, 4 = all bracts fimbriate.
  106. Number of petaloid bracts per flower: (5–)8–45.
  107. Lengths of petaloid bracts: Continuous variable, (1–)4.5–40 mm.
  108. Widths of petaloid bracts: Continuous variable from 1.0–14 mm. Shapes of petaloid bracts (regardless of width): 0 = lanceolate, 1 = oblong, 2 = oblanceolate, 3 = spatulate.
  109. Apex of petaloid bracts, general shape: 0 = attenuate, 1 = acute, 2 = obtuse, 3 = rounded or erose.
  110. Apex of petaloid bracts, extreme apices: 0 = acute, 1 = obtuse, 2 = rounded or truncate, 3 =retuse or emarginate.
  111. Tips of petaloid bracts, centers of extreme apices: 0 = entire, 1 = mucronate, 2 = caudate.
  112. Color of petaloid bracts: 0 = intense; 1 = pale, or color confined to a midstripe, or mixed with greenish; 2 = white or nearly so. (This can be quantified by using RGB values of standardize images.)
  113. Yellow/purple dichotomy of pigmentation in petaloid bracts: descriptive (This can be quantified by using RGB values of standardize images.) 0 = yellowish, or yellow-and-red with no admixture of purplish; 1 = polymorphic, or brown, or mixed pink-and-yellow; 2 = pink, or purplish red.
  114. Zonation of color within the flower: 0 = none (filaments same color as the petaloid bracts, or else unpigmented), 1 = filaments much brighter/darker than the petaloid bracts, 2 = entire center of flower brighter/darker than the petaloid bracts, 3 = multiple concentric zones of color (e.g., Thelocactus bicolor)
  115. Color of throat and upper filaments: Divisible into two binary characters: A.) 0 = white, yellow, or green; 1 = pink, or purplish red; B.) 0 = white, yellow, or green; 1 = red or orange. [Red, as opposed to purple, centers are strongly correlated with yellow petaloid bract color. To some extent, this is a duplication zonation, above.]
  116. Petaloid bracts becoming more or less reflexed: 0 = yes, 1 = moderately so, 2 = no, remaining erect even under optimal conditions.
  117. Timing of anthesis: 0 = hottest and brightest part of the day, 1 = vespertine, 2 = matinal anthesis.
  118. Number of ranks of stamens: Meristic character, 2–20. (This is the approximate number of stamens likely to be present in a thin (roughly two stamens' widths) longisection of one wall of the receptacular tube.
  119. Lengths of upper filaments: Continuous variable, 3–15 mm.
  120. Lengths of lower filaments: Continuous variable, 3–20 mm.
  121. Anther color, yellow vs. pink: 0 = yellowish, 1* = pink.
  122. Anther color, pale vs. dark: 0 = orange, 1 = orange-yellow or dark yellow, 2 = average yellow, 3 = very pale yellow.
  123. Filaments inserted in two distinct bands: 0 = no, 1 = yes (e.g., Coryphantha pulleineana).
  124. Anthers of uppermost stamens sterile: 0 = no, 1 = yes (e.g., Mammillaria).
  125. Gynodioecy 0 = no, 1 = yes (e.g., Mammillaria).
  126. Outgrowths (staminodia?) in throat of flower: 0 = no, 1 = yes (e.g., Leuchtenbergia, Ferocactus spp.,
  127. Longitudinal grooves along style : 0 = style lacking grooves, 1 = style grooved.
  128. Style epidermis pappillose: 0 = style epidermis smooth, 1 = style epidermis pappillose.
  129. Style length: 0 = style/stigmas among anthers, 1 = style exserted, stigmas beyond the anthers.
  130. Number of stigma lobes: Continuous or meristic variable (mean per plant or population or counts), 2–20.
  131. Stigma lobe length: Continuous variable, from ± 0.5–12 mm.
  132. Stigma lobe pigmentation: 0 = white, 1 = pale pigmentation, 2 = dark pigmentation.
  133. Stigma lobe color: 0 = green or yellow-green; 1 = brown, or mixed pink-and-green; 2 = red, pink, or purple.
  134. Stigma lobes spreading: 0 = spreading, at least after first day; 1 = erect, but not fused; 2 = erect, fused into a capitate mass.
  135. Loculus width (cross-sectional diameter): Continuous variable, ± 1–10 mm.
  136. Loculus shape (longitudinal section): 0 = oblate, or shield-shaped; 1 = round; 2 = elongate.
  137. Number of ovules per carpel: Continuous variable, 2–50. (This character is estimated from the number of ovules per loculus divided by the number of stigma lobes.)
  138. "Column" between loculus and nectary: 0 = essentially none, 1 = well-developed, 2 = always well-developed (e.g., some Mammillaria spp.).
  139. Fruit shape, irrespective of length/width ratio: 0 = ovoid, I = spheroid or ellipsoid, 2 = obovoid, 3 = clavate.
  140. Fruit diameter: Continuous variable, 2–25(–50) mm (e.g., Ferocactus spp.).
  141. Fruit length: Continuous variable, 2–60(–100) mm.

     

     

     

     

  142. Fruit succulence: 0 = juicy, 1 = succulent pericarpel but dry funiculi, 2 = pericarpel and funiculi both dry at time of fruit abscission.
  143. Fruit color: 0 = remaining green when ripe, I = turning dull reddish or dull greenish yellow at ripening, 2 = turning bright yellow, 3 = turning bright red.
  144. Dried floral remnant: 0 = persistent, I = weakly persistent, 2 = never persistent, cleanly abscising a short time after anthesis.
  145. Fruits "dimorphic" (undergoing final size- and color-change at non-synchronous times, often following long fruit dormancy): 0 = no, 1 = yes (e.g., Mammillaria spp.).
  146. Fruit/ovary deeply sunken into primary vegetative shoot? 0 = no, 1 = yes (e.g., Mammillaria spp.).
  147. Fruit dehiscence via longitudinal splitting: 0 = never, 1 = yes, at least under optimal conditions.
  148. Fruit dehiscence via basal pore: 0 = not basally dehiscent in any way, 1 = small basal pore, 2 = large basal pore.
  149. Fruit circumscissilly dehiscence: 0 = not circumscissile above the base, 1 = circurnscissile above base of ovary.
  150. Seed length: Continuous variable, (0.25–)0.8–3.5(–4.5) mm. Note that seed length, as defined here, refers to the longest dimension of the seed, not necessarily the length of an axis perpendicular to the plane formed by the hilum.
  151. Strophiole size: 0 = apparently none, 1 = conspicuous when moist, 2 = conspicuous even when dry.
  152. Seed shape, I: 0 = symmetry superficially radial, longest axis of the seed perpendicular to the plane defined by the hilum (= "hilum basal," of the cactus literature); 1 = intermediate; 2 = symmetry obviously bilateral, the longest axis of the seed either oblique or parallel to the plane defined by the hilum (= "hilum lateral," of the cactus literature).
  153. Testa color: 0 = black, 1 = brown when moist and fresh but changing to black when old and dry, 2 = bright red or yellow when fresh and moist, changing to brown when old and dry.
  154. Testa color, red versus yellow: 0 = reddish, I = yellowish. (Not applicable for taxa having black or nearly black seeds)
  155. Testa luster: 0 = glossy, 1 = semi-matt, 2 = matt
  156. Testa periphery: 0 = undifferentiated periphery 1 = keeled, but cells undifferentiated 2 = crested, with cells of the periphery enlarged.
  157. Testa multicellular sculpture: 0 = seeds smooth, 1 = seeds rugose, 3 = seeds ruminate.
  158. Testa hilum boarder: 0 = not expanded around the hilum, 1 = cells expanded around the hilum, 2 = cells greatly expanded around the hilum, 3 = cells expanded around and constricted above the hilum
  159. Testa cell size: 0 = uniform, 1 = gradually smaller towards the hilum, 2 = abruptly smaller and with less pronounced surface sculpting than lateral cells at the hilum (e.g., Pediocactus, Turbinicactus).
  160. Testa cell shape: 0 = isodiametric, 1 = elongate
  161. Testa anitclinal cell-boundary relief: 0 = channeled, 1 = inconspicuous, 2 = raised.
  162. Testa anitclinal cell-boundary curvature: 0 = straight, 1 = irregularly curved, 2 = undulate.
  163. Testa anitclinal cell-boundary undulation: 0 = S-type, 1 = U-type, 2 = V-type, 2 = omega-type.
  164. Testa anitclinal cell-boundary interstices: 0 = undifferentiated, 1 = minutely pitted, 2 = cratered.
  165. Testa periclinal wall relief: 0 = none, 1 = convex, 2 = concave, 3 = par-convex, 4 = par-concave. (Requires scanning electron microscopy.)
  166. Testa periclinal wall convex structures: 0 = low-domed, 1 = high-domed, 2 = low-conical, 3 = high-conical. (Requires scanning electron microscopy.)
  167. Testa periclinal wall par-convex structures: 0 = par-domed, 1 = par-conical, 2 = lpar-cylindrical, 3 = par-clavate. (Requires scanning electron microscopy.)
  168. Testa periclinal wall microrelief: 0 = none, 1 = verrucose (cell wall and/or cuticle), 2 = striate (cuticle only). (Requires scanning electron microscopy.)
  169. Testa periclinal wall cuticle microrelief: 0 = weakly striate, 1 = fielded-striate, 2 = regular and finely striate, 3 = strongly folded (“loose cuticle”). (Requires scanning electron microscopy.)
  170. Hilum-micropylar region size: 0 = small, 1 = medium, 2 = large.
  171. Hilum-micropylar region orientation: 0 = basal, 1 = oblique.
  172. Hilum-micropylar region position relative to rim: 0 = superficial, 1 = impressed, 2 = deeply impressed (testa often flared).
  173. Hilum and micropylar configuration: 0 = disjunct, 1 = conjunct but separated by sclerified tissue, 2 = fused.
  174. Hilum-micropylar region shape: 0 = oval, 1 = “curved-band” type, 2 = “keyhole” type.
  175. Seed appendages: 0 = none, 1 = mucilage sheath, covering the entire seed, 2 = mucilage sheath, restricted to the hilum area, 3 = strophiolar pad present, 4 = strophiole present.
  176. Perisperm: 0 = large, 1 = small but still containing starch, 2 = vestigial, not containing starch.
  177. Cotyledon shape: 0 = pointed, 1 = rounded. (Echinocactus exhibits both character-states.)
  178. Embryo shape: 0 = strongly curved, 1 = slightly curved, 2 = straight.
  179. First podaria elongate and discrete, immediately forming a rosette: 0 = no, 1 = yes (e.g., Ariocarpus and Obregonia, perhaps intermediate in Leuchtenbergia)
  180. Ploidy level: 0 = diploid, I = tetraploid, 3 = higher polyploid.
  181. Fruit phenology: 0 = short (ca. 2 months), 1 = average (ca. 4 months), 2 = long (ca. 6 to 12 months).
  182. Flowering phenology: 0 = mid-winter (e.g., Sclerocactus [Ancistrocactus] scheeri), 1 = early spring, 2 = late spring, 3 = summer, or opportunistically following rains, 4 = autumn.

Anderson and Boke 1969.
Wittler 1984.

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