The Identifi cation of Medicinal A Handbook of the Morphology of Botanicals in Commerce

by Wendy Applequist, Ph.D. Foreword by Mark Blumenthal & Steven Foster Illustrations by Barbara Alongi Th e Identifi cation of A Handbook of the Morphology of Botanicals in Commerce

By Wendy Applequist, Ph.D.

Illustrations by Barbara Alongi

With a Foreword by

Mark Blumenthal Founder and Executive Director American Botanical Council and

Steven Foster President, Steven Foster Group Botanist and Author

A joint project of

Missouri Botanical Garden American Botanical Council St. Louis, Missouri Austin, Texas

2006 About the Author

Wendy Applequist earned her Ph.D. in systematics from Iowa State University and is an assistant curator in the William L. Brown Center for Plant Genetic Resources at the Missouri Botanical Garden. She conducts research on the botany of medicinal plants and plants native to Madagascar. She has previously published articles in several journals, including Systematic Botany, Taxon, Plant Systematics and Evolution, Evolution and Development, Pharmazie, Flora, and Adansonia. Th is is her fi rst book.

About the Artist

Barbara Alongi is a scientifi c illustrator based at the Missouri Botanical Garden. Her illustrations accompany many descriptions of new of plants as well as monographic treatments. She is an illustrator for the Flora of North America project. About the American Botanical Council

Th e American Botanical Council (ABC) is the leading independent, nonprofi t education and research organization using science-based and traditional information to promote the responsible use of . Th e member- supported organization serves all populations interested in herbal medicine: the general public, healthcare professionals, researchers, educators, industry, media and government. ABC’s vision is that the public makes educated, responsible choices about herbal medicine as an accepted part of healthcare. Founded in 1988, ABC supports this vision through its mission to provide science-based and traditional information to promote the responsible use of herbal medicine. ABC achieves its mission through: • Publication of its peer-reviewed journal • Continuing education materials for healthcare HerbalGram prrofessionals • HerbClip™ literature review service • Internships for healthcare professionals • HerbalEGram electronic newsletter • Safety guidelines for herbal products • Publication of books and literature • Seminars, presentations and workshops • Information on its website, www.herbalgram.org • Serving as a source of authoritative information to global print and electronic media For information: American Botanical Council P.O. Box 144345 Austin, TX 78714-4345 Phone: 512-926-4900 Fax: 512-926-2345 Toll free in the U.S.: 800-373-7105 Email: [email protected]. Website: www.HerbalGram.org

About the Missouri Botanical Garden

Th e oldest continuously operating botanical garden west of the Mississippi, the Missouri Botanical Garden of St. Louis is an award-winning horticultural and educational institution whose many public attractions run the gamut from an old- fashioned garden to a brand-new Children’s Garden. Behind the scenes, it is also one of the world’s most active centers for botanical systematic and fl oristic research, with about 150 full-time research staff who conduct fi eld studies in dozens of countries every year. Th e Garden’s mission is “To discover and share knowledge about plants and their environment, in order to preserve and enrich life”. Th e Garden’s William L. Brown Center for Plant Genetic Resources focuses its research specifi cally on the identifi cation, scientifi c study, and preservation of plants that are of direct use to humans.

vi Th e Identifi cation of Medicinal Plants: Table of Contents

Table of Figures ...... xi Foreword ...... xiii Acknowledgements ...... xvi Introduction ...... xvii Background ...... 1 Basics of Plant Morphology ...... 3 Practical Plant Identifi cation ...... 7 Botanical Nomenclature ...... 9 Description of Botanical Entries ...... 11 Botanical Entries ...... 13 L. (Yarrow) ...... 14 Actaea racemosa L. (Black Cohosh)...... 16 Adonis vernalis L. (Spring Adonis) ...... 18 Aesculus hippocastanum L. (Horse Chestnut) ...... 20 betulina (P. J. Bergius) Pillans, A. crenulata (L.) Pillans, A. serratifolia (Curtis) Spreeth (Buchu) ...... 21 Althaea offi cinalis L. (Marshmallow) ...... 22 Andrographis paniculata (Burm. f.) Nees (Andrographis) ...... 23 Angelica sinensis (Oliv.) Diels (Dong Quai) ...... 25 Apium graveolens L. () ...... 26 Arctostaphylos uva-ursi (L.) Spreng. (Uva-Ursi)...... 27 Arnica montana L. (Arnica) ...... 29 Artemisia absinthium L. (Wormwood) ...... 31 Artemisia annua L. (Sweet Wormwood) ...... 32 Astragalus mongholicus Bunge (Astragalus) ...... 34 Berberis aquifolium Pursh, B. nervosa Pursh, B. repens Lindl. (Oregon Grape) ...... 36 L. (Barberry) ...... 37 Betula pendula Roth, B. pubescens Ehrh. (Birch) ...... 38 Calendula offi cinalis L. (Calendula) ...... 39 Capsella bursa-pastoris (L.) Medik. (Shepherd’s Purse) ...... 41

A Handbook of the Morphology of Botanicals in Commerce vii Carum carvi L. () ...... 42 Caulophyllum thalictroides (L.) Michx. (Blue Cohosh) ...... 44 Centella asiatica (L.) Urb. (Gotu Kola) ...... 45 Chamaelirium luteum (L.) A. Gray (False Unicorn) ...... 46 Chamaemelum nobile (L.) All. (Roman ) ...... 48 Cichorium intybus L. (Chicory) ...... 49 Coriandrum sativum L. () ...... 50 Crataegus laevigata (Poir.) DC, Crataegus monogyna Jacq. (Hawthorn) ...... 51 Crocus sativus L. (Saff ron) ...... 54 Cucurbita pepo L. (Pumpkin) ...... 55 Cytisus scoparius (L.) Link (Scotch Broom) ...... 56 Dioscorea villosa L. (Wild Yam) ...... 57 Echinacea angustifolia DC. (Echinacea angustifolia); E. pallida (Nutt.) Nutt. (Echinacea pallida) ...... 58 (L.) Moench (Echinacea purpurea) ...... 60 Eleutherococcus senticosus (Rupr. & Maxim.) Maxim. (Eleuthero) ...... 62 Epimedium brevicornu Maxim., E. grandifl orum C. Morren, E. koreanum Nakai, E. pubescens Maxim., E. sagittatum (Sieb. & Zucc.) Maxim., E. wushanense T. S. Ying (Epimedium)...... 63 Equisetum arvense L. (Horsetail) ...... 66 Euphrasia offi cinalis L. (Eyebright) ...... 68 Fallopia japonica (Houtt.) Ronse Decraene ( Japanese Knotweed) ...... 70 Filipendula ulmaria (L.) Maxim. (Meadowsweet) ...... 71 Foeniculum vulgare Mill. () ...... 72 Frangula purshiana (DC.) J. G. Cooper (Cascara Sagrada) ...... 73 Galium aparine L. (Cleavers)...... 75 lutea L. (Gentian) ...... 77 biloba L. (Ginkgo) ...... 79 Glycyrrhiza glabra L. (Licorice) ...... 80 Hamamelis virginiana L. (Witch Hazel)...... 81 Hibiscus sabdariff a L. (Hibiscus) ...... 82 Hydrastis canadensis L. () ...... 84 Hypericum perforatum L. (St. John’s Wort) ...... 85 Hyssopus offi cinalis L. (Hyssop) ...... 86 Ilex paraguariensis A. St.-Hil. (Maté)...... 87 verum Hook. f. (Star ) ...... 89 Juniperus communis L. (Juniper) ...... 91 Mill. (English Lavender) ...... 92 viii Th e Identifi cation of Medicinal Plants: Ligusticum porteri J. M. Coult. & Rose (Osha) ...... 93 Linum usitatissimum L. (Flax) ...... 95 Lobelia infl ata L. (Lobelia) ...... 96 Lycopus europaeus L. (European Bugleweed) ...... 98 Lycopus virginicus L. (Bugleweed) ...... 99 Marrubium vulgare L. (Horehound) ...... 101 Matricaria chamomilla L. (Chamomile) ...... 104 Medicago sativa L. () ...... 105 Melissa offi cinalis L. subsp. offi cinalis () ...... 107 ×piperita L. () ...... 109 Mitchella repens L. (Partridge berry) ...... 110 europaea L. () ...... 112 Panax C. A. Mey. (Asian Ginseng); P. quinquefolius L. (American Ginseng) ...... 113 Passifl ora incarnata L. (Passionfl ower) ...... 115 Peumus boldus Molina () ...... 118 Phyllanthus amarus Schumach. & Th onn. (Phyllanthus amarus) ...... 119 Phyllanthus fraternus G. L. Webster, P. niruri L., P. urinaria L. (Phyllanthus) ...... 122 Pimpinella anisum L. (Anise) ...... 126 Plantago afra L., P. arenaria Waldst. & Kit., P. asiatica L., P. ovata (Psyllium) ...... 128 Plantago major L. (Plantain) ...... 130 Prunella vulgaris L. (Heal All) ...... 132 Rhamnus cathartica L. (Buckthorn) ...... 134 Rosmarinus offi cinalis L. () ...... 135 Rubus idaeus L. (Raspberry) ...... 136 Rumex crispus L. (Yellow Dock) ...... 138 L. (White Willow) ...... 139 Salvia offi cinalis L. (Sage)...... 140 Sambucus nigra L. (European Elder) ...... 142 Sanguinaria canadensis L. (Bloodroot) ...... 145 Schisandra chinensis (Turcz.) Baill. (Schisandra) ...... 146 Scutellaria laterifl ora L. (Skullcap) ...... 147 Senna alexandrina Mill. (Senna) ...... 150 Serenoa repens (W. Bartram) Small (Saw Palmetto) ...... 152 Sida cordifolia L. (Heart- Sida) ...... 153 Silybum marianum (L.) Gaertn. (Milk Th istle) ...... 155 Smilax aristolochiifolia Mill., S. febrifuga Kunth, S. regelii Killip & C. V. Morton (Sarsaparilla) ...... 156

A Handbook of the Morphology of Botanicals in Commerce ix Solidago virgaurea L. (European Goldenrod) ...... 158 Stellaria media (L.) Vill. (Chickweed) ...... 160 rebaudiana (Bertoni) Bertoni (Stevia) ...... 162 sylvatica Garden ex L. (Stillingia) ...... 163 (L.) Sch. Bip. (Feverfew) ...... 164 Taraxacum offi cinale F.H.Wigg. (Dandelion) ...... 166 Th ymus vulgaris L. (Th yme) ...... 168 Tilia cordata Mill., T. platyphyllos Scop., T. ×europaea L. (Linden) ...... 170 Trifolium pratense L. (Red Clover) ...... 172 Trigonella foenum-graecum L. () ...... 173 Turnera diff usa Willd. ex Schult. (Damiana) ...... 174 Tylophora indica (Burm. f.) Merr. (Tylophora asthmatica) ...... 175 Ulmus rubra Muhl. (Slippery Elm) ...... 177 Urtica dioica L. subsp. dioica (Stinging Nettle) ...... 178 Vaccinium macrocarpon Aiton (Cranberry) ...... 180 Vaccinium myrtillus L. (Bilberry) ...... 181 Valeriana offi cinalis L. () ...... 182 Viburnum prunifolium L. (Black Haw) ...... 183 Viscum album L. (European Mistletoe) ...... 184 Vitex agnus-castus L. (Chaste ) ...... 187 Vitis vinifera L. (Grape) ...... 188 Withania somnifera (L.) Dunal (Ashwagandha)...... 189 Zingiber offi cinale Roscoe () ...... 190

Appendix ...... 193 General References ...... 194 Glossary ...... 195

Index ...... 205

x Th e Identifi cation of Medicinal Plants: Table of Figures

Figure 1: Achillea millefolium ...... 15 Figure 25: Equisetum arvense, E. palustre ...... 67 Figure 2: Actaea racemosa, A. podocarpa, Figure 26: Euphrasia offi cinalis, E. stricta ...... 69 A. pachypoda, A. rubra ...... 17 Figure 27: Filipendula ulmaria...... 71 Figure 3: Adonis vernalis ...... 19 Figure 28: Foeniculum vulgare ...... 73 Figure 4: , A. crenulata ...... 21 Figure 29: Galium aparine, G. verum, G. odoratum ...... 75-77 Figure 5: Andrographis paniculata ...... 24 Figure 30: Ginkgo biloba ...... 79 Figure 6: Apium graveolens, Ammi majus, Ammi visnaga ...... 26 Figure 31: Hamamelis virginiana ...... 81 Figure 7: Arctostaphylos uva-ursi ...... 28 Figure 32: Hibiscus sabdariff a ...... 83 Figure 8: Arnica montana, A. chamissonis ...... 29-30 Figure 33: Hypericum perforatum ...... 85 Figure 9: Artemisia absinthium ...... 31 Figure 34: Hyssopus offi cinalis ...... 87 Figure 10: Artemisia annua ...... 33 Figure 35: Ilex paraguariensis ...... 88 Figure 11: Astragalus mongholicus ...... 35 Figure 36: Illicium verum, I. anisatum ...... 89 Figure 12: Berberis aquifolium, B. repens ...... 36 Figure 37: Lavandula angustifolia, L. latifolia ...... 92 Figure 13: Betula pendula, B. pubescens ...... 38-39 Figure 38: Ligusticum porteri ...... 94 Figure 14: Calendula offi cinalis ...... 40 Figure 39: Linum usitatissimum ...... 95 Figure 15: Capsella bursa-pastoris ...... 41 Figure 40: Lobelia infl ata ...... 96-97 Figure 16: Carum carvi, Cuminum cyminum ...... 43 Figure 41: Lycopus europaeus ...... 98-99 Figure 17: Centella asiatica ...... 45 Figure 42: Lycopus virginicus ...... 100 Figure 18: Chamaemelum nobile ...... 48 Figure 43: Marrubium vulgare, M. perigrinum, Ballota hirsuta, Figure 19: Cichorium intybus ...... 49 B. nigra ...... 102-103 Figure 20: Coriandrum sativum ...... 50 Figure 44: Matricaria chamomilla, Figure 21: Crataegus laevigata, C. monogyna....52-53 Anthemis cotula ...... 104-105 Figure 22: Echinacea pallida, E. angustifolia ...... 59 Figure 45: Medicago sativa ...... 106 Figure 23: Echinacea purpurea ...... 61 Figure 46: Melissa offi cinalis, Figure 24: Epimedium sagittatum, Nepeta cataria ...... 107-108 E. grandifl orum ...... 64 Figure 47: Mentha ×piperita, M. canadensis ...... 109

A Handbook of the Morphology of Botanicals in Commerce xi Figure 48: Mitchella repens ...... 111 Figure 67: Serenoa repens ...... 152 Figure 49: Olea europaea ...... 112-113 Figure 68: Sida cordifolia ...... 153 Figure 50: Panax quinquefolius ...... 114-115 Figure 69: Silybum marianum ...... 155 Figure 51: Passifl ora incarnata ...... 117 Figure 70: Solidago virgaurea, S. gigantea...... 159 Figure 52: Peumus boldus ...... 118 Figure 71: Stellaria media ...... 161 Figure 53: Phyllanthus amarus ...... 119-120 Figure 72: ...... 162 Figure 54: Phyllanthus fraternus, Figure 73: Tanacetum parthenium, T. vulgare ...... P. niruri, P. urinaria ...... 122-125 164-165 Taraxacum offi cinale ...... Figure 55: Pimpinella anisum, Figure 74: 166 Petroselinum crispum ...... 126 Figure 75: Th ymus vulgaris, T. zygis ...... 169 Figure 56: Plantago afra, P. major ...... 128 Figure 76: Tilia platyphyllos, T. cordata, T. europaea ...... Figure 57: Plantago major, P. lanceolata, × 170-171 Digitalis lanata ...... 131 Figure 77: Trifolium pratense ...... 172 Figure 58: Prunella vulgaris ...... 133 Figure 78: Trigonella foenum-graecum ...... 173 Figure 59: Rhamnus cathartica ...... 134 Figure 79: Turnera diff usa ...... 174 Figure 60: Rosmarinus offi cinalis ...... 135 Figure 80: Tylophora indica ...... 176 Figure 61: Rubus idaeus, R. fruticosus ...... 137 Figure 81: Urtica dioica, Urtica urens ...... 179 Figure 62: Salvia offi cinalis, S. fruticosa ...... 141 Figure 82: Viscum album subsp. album ...... 185 Figure 63: Sambucus nigra ...... 143 Figure 83: Vitex agnus-castus ...... 187 Figure 64: Sanguinaria canadensis ...... 145 Figure 84: Vitis vinifera ...... 188 Figure 65: Scutellaria laterifl ora, Figure 85: Withania somnifera ...... 189 Teucrium canadense ...... 148-149 Figure 86: Leaf characters ...... 198 Figure 66: Senna alexandrina, S. italica ...... 151 Figure 87: Common infl orescence types ...... 199

xii Th e Identifi cation of Medicinal Plants: Equisetum arvense L. Standardized Common Name: Horsetail

Other Common Names: Common Horsetail, Field Adulterants: E. arvense may be confused with other spe- Horsetail, Joint Grass cies of Equisetum. It is particularly important that E. ar- vense be distinguished from E. palustre L., as the latter Family: Equisetaceae species, which has been found as a contaminant of the former, is toxic when consumed by livestock. Diff erences : Equisetum is a pteridophyte (non-seed plant) ge- between the two include: nus of about 15 species, found nearly worldwide. Hybridization among similar species is not uncommon; E. ×litorale Kühlewein E. arvense E. palustre ex Ruprecht, a hybrid between E. arvense and E. fl uviatile, oc- Number of stem ridges (4–)8–10(–16) 4–10 curs throughout northern North America. Equisetum arvense is and stem sheath teeth extremely variable in gross morphology; Hauke (1966) estimates Position of branch Regular whorls along Only at midstem nodes; that over 200 infraspecifi c taxa have been described. However, whorls on stem whole length of stem other nodes lacking the features that distinguish these supposed varieties or forms branch whorls are often under environmental control, so that multiple forms Length of fi rst Longer than Shorter than subtending internode of each subtending stem stem sheath may appear in a single individual. branch sheath 3–4 4–6 Description: Perennial, rhizomatous herb with jointed stems Branch ridge number Central cavity of Absent; branches Present (observe near branching at the nodes; whorled, reduced to a sheath sur- branches solid base of branches) rounding the nodes. Reproductive stems and vegetative stems Stem sheath teeth Dark with Dark with conspicuous generally separate; reproductive stems brown, unbranched, inconspicuous white, membranous short-lived, with rounded cones at apex; cones borne on vegeta- light margins; often margins tive stems in occasional abnormal plants. Vegetative stems 2–100 cohering in pairs cm tall, 0.8–4.5 mm in diameter; internodes 1.4–4.5 cm long, Branch sheath teeth Lanceolate-attenuate Triangular with 4–16 ridges separated by valleys; in cross-section hollow, with central canal 1/3–2/3 diameter of stem (reduced in small References: stems), with large hollow spaces (vallecular canals) beneath val- leys and small carinal canals beneath ridges, closer to central Hauke RL. A systematic study of Equisetum arvense. Nova Hed- canal. Leaf sheaths on stems squarish in face view, 2–5(–10) wigia. 1966;13:81–109. mm high, 2–5(–9) mm broad; teeth 1–3.5 mm long, dark, nar- row, often cohering in pairs. Branches in regular whorls at most Hauke RL. A taxonomic monograph of Equisetum Subgenus nodes, ascending, solid, 3–4-ridged, with fi rst internodes longer Equisetum. Nova Hedwigia. 1978;30:385–455. than the subtending stem sheaths; sheath teeth attenuated. Hauke RL. Equisetaceae. In: Flora of North America Editorial Committee, eds. Flora of North America. Vol. 2. New York, NY: Parts in Commerce: Vegetative stems Oxford University Press; 1993:76–84. Identifi cation:

• First internode of each branch, except at the lowest nodes, longer than the subtending stem sheath • Branches solid, lacking central cavity • Branches occur in regular whorls on most or all of stem, not confi ned to midstem or lower part of stem • Branches 3–4-angled, normally not further branched • Stem sheath teeth (4–)8–10(–16), usually under 4 mm long, dark (not reddish), narrow, stiff (not papery), often cohering in pairs • Branch sheath teeth attenuate (not broadly triangu- lar)

66 Th e Identifi cation of Medicinal Plants: B 2 mm

A

3 mm C

E 2 cm 1 mm F

5 mm G D

Figure 25: a–d, Equisetum arvense; e–g, E. palustre.

A Handbook of the Morphology of Botanicals in Commerce 67 Pimpinella anisum L. Standardized Common Name: Anise

Family: Apiaceae (Umbelliferae) • Odor of crushed strong, aromatic • Taste characteristic, aromatic, pleasant Taxonomy: Pimpinella is a of about 150 Old World . Pimpinella anisum is one of 16 species that grows wild in , and has been widely cultivated for millennia. Syn- onyms include Anisum offi cinale DC., Anisum vulgare Gaertn., Apium anisum Crantz and Pimpinella aromatica Bieb.

Description: Annual herb. Stem (10–)30–50(–100) cm high, sometimes bearing small bristles. Leaves basal and cauline; bas- al leaves petiolate, simple, ovate or reniform, 2–5 cm long, the margins dentate to serrate; stem leaves with sheathing petioles, alternate, 1–2(–3)-pinnately or ternately compound, the leafl ets ca. 1.5–4 cm long, linear or ovate to rhomboid and toothed to deeply pinnatifi d. Infl orescence a compound umbel, long-pe- 2 mm duncled, with 7–15 rays, bractless or with 1 linear bract; umbel- lets 7–12(–15)-fl owered, without bracteoles or with few small linear bracteoles; fl owers small, white or yellowish, 5-petalled. Fruit a schizocarp of 2 mericarps, (2–)3–5(–7) mm long, ovoid, laterally compressed, constricted at commissure, with short scaly hairs; vallecular vittae usually 3, commissural vittae 2–4(–6); stylopodium conical.

Parts in Commerce: A Identifi cation: See glossary for explanation of the techni- cal terms pertaining to umbel fruits. • Schizocarp usually intact, not split into individual mericarps, and often still attached to the slender pedicel • (2–)3–5(–7) mm long • Ovoid or pear-shaped; apex narrowed and ending in conical stylopodium • Broad at commissure, but grooved on both sides between mericarps • Greenish or yellowish brown; ribs yellowish, paler than valleculae • Ribs delicate, threadlike, straight, at least as broad as high 0.5 mm • Pubescent with small yellowish scaly hairs; hairs may be worn off , but are easiest to observe in com- missural grooves or in valleculae near apex • Vittae usually at least 3 per vallecula, quite incon- spicuous; commissural vittae usually 2–4, observ- B able in cross-section or in separated mericarps as pale ridges on commissural face • Endosperm in cross-section fl at along commissural face Figure 55: a, Pimpinella anisum fruit; b, Petroselinum crispum fruit.

126 Th e Identifi cation of Medicinal Plants: Adulterants: Literature reports adulteration by Petroseli- References: num crispum (Mill.) A. W. Hill (), also important Arenas Posada JA, García Martín F. Atlas carpológico y in commerce, and Conium maculatum L. (poison hem- corológico de la subfamilia Apioideae Drude (Umbelliferae) en lock), a toxic plant that is no longer sold as medicinal. España peninsular y Baleares. Ruizia. 1993;12:1–245. Th ese share relatively small, ovoid fruits with threadlike ribs (although some material of C. maculatum has narrow, Cappellettii EM. Botanical identifi cation of Anise and Hem- lock fruits in powdered drug samples. Planta Med. 1979;39:88– elongated fruits), but they may be distinguished easily by 94. several morphological features, as well as great diff erences in aroma and taste: Matthews VA. Pimpinella. In: Davis PH. Flora of Turkey and the East Aegean Islands. Vol. 4. Edinburgh: Edinburgh University Press; 1972:352–364. Pimpinella Petroselinum Conium anisum crispum maculatum Tutin TG. Pimpinella. In: Tutin TG, Heywood VH, Burges NA, Pubescence Short scaly hairs, Hairless Hairless; et al., eds. Flora Europaea. Vol. 2. Cambridge: Cambridge Uni- often persistent minute teeth versity Press; 1968:331–333. only in grooves may be seen near apex and in valleculae of Tutin TG. Umbellifers of the British Isles. London: Botanical So- commissure immature fruits ciety of the British Isles; 1980. B.S.B.I. Handbook, No. 2. Primary ribs (in Straight; usually Straight; usually Tend to undulate dried fruits) at least as broad broader than high especially in Wichtl M, ed. Herbal Drugs and Phytopharmaceuticals, 3rd Eng- as high immature fruits; lish ed. Stuttgart: medpharm Scientifi c Publishers and Boca usually higher than broad; Raton, FL: CRC Press; 2004:42–44. often somewhat notched or toothed Vittae 2 or more on 2 on commissural Absent at fruit commissural face, face; 1 per maturity numerous and vallecula, very hard to observe in broad, giving valleculae valleculae brown color Commissure Fairly broad (but Constricted Constricted narrower than fruit) Endosperm at Flat Flat Deeply grooved commissural face in cross- section

A Handbook of the Morphology of Botanicals in Commerce 127