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Near Eastern Grass Seeds IDENTIFICATI NEAR EASTERN GRASS SEEDS Mark Nesbitt Illustrated by Jane Goddad Contents List of figures (excluding Genus catalogue) vii 2.5 Inflorescence Preface: user guide ix 2.6 Ecology and biogeography Acknowledgements xi 2.7 Near Eastern biogeography Abbreviations xiii 1 Introduction 1 3 Grass seed morphology 1 .l Previous work 3.1 The spikelet and floret 1.2 Identification criteria 3.1.1 Callus 1.2.1 Taxonomic system 3.1.2 Plantlets 1.2.2 Selection of species for study 3.1.3 Lemma and palea Native status Lemma and palea phytoliths Geography Charred lemma and palea remains Abundance 3.2 The caryopsis Reference material 3.2.1 Structure 1.2.3 A guide to taxonomic ranks 3.2.2 Shape Family Caryopsis length Subfamily Compression and thickness:breadth Tribe ratio Genus Narrowness and 1ength:breadth ratio Section Cross-section Species, subspecies and variety Ventral furrow 1.3 Archaeology and ethnobotany of grasses Dorsal ridging in the Near East: a brief overview Dorsal markings 1.3.1 Foraging Lateral groove Changing distribution of the grasses 3.2.3 Caryopsis hairs Wild grasses as human food 3.2.4 Sculpturing 1.3.2 Farming 3.2.5 Apex wrinkling Evolution of weeds 3.3 Embryo Weed ecology 3.3.1 Embryo size 3.3.2 Scutellum shape 3.3.3 Embryo type 2 lntroduction to grass biology 3.4 The hilurn 3.4.1 Hilum length 2.1 Habit 12 3.4.2 Hilum shape 2.2 Breeding systems 13 3.4.3 Hilum visibility 2.3 Polyploidy l3 3.5 Bran 2.4 Fungal symbiosis 14 3.5.1 Structure and terminology 3.5.2 Bran of uncharred modern caryopses and food remains 3.5.3 Bran of charred caryopses 3.5.4 Bran of waterlogged caryopses 3.5.5 Waterlogged bran fragments CONTENTS v 3.6 Endosperm 3.6.1 Starch 3.6.2 Endosperm texture 3.7 Presentation of data 3.8 Measurements 4 Using the key 4.1 The analytical key 4.2 The polyclave key Select bibliography General Near Easl Other Floras Additional references cited Notes Genus catalogue Format Appendix 1: Near Eastern grass genera Appendix 2: Caryopsis data Index togenera vi IDENTIFICATION GUIDE FOR NEAR EASTERN GRASS SEEDS Distribution and relationships Other genera Perennial, c. 450 species in temperate regions throughout the Festuca caryopses, particularly of E pratensis, are very similar to world; in highlands in the Near East. Hills, plains and meadows. those of L~liurn'~but are markedly smaller. Festuca grains are also Identification to species is highly problematic and often depends very similar to Micropyrum, another closely related genus. Micro- on study of leaf anatomy. The division into sections follows the pyruml%eeds have more parallel sides, not widening towards the Flom of Turkey. Section Bovinae is known to be closely related to apex. All three genera show the strong impression of the palea on Loliurn, particularly L. multiflorum. the ventral side. resulting in a very clearly defined (even if not deep) V-shaped ventral furrow. This feature makes these three Seed morphology genera highly distinctive. Two groups have been defined on the basis of differing hilum lengths. Too few species have been assessed to allow any sectional Bibliography pattern to be determined. Aiken, S. G. and S. E. Gardiner (1991) SDS-PAGE of seed proteins in Fes- tuco (Poaceae): taxonomic implications. Canadian lournal of Botany Glumes: light papery 69: 1425-32. Lemma andpalea: light papery, both adherent Aiken, S. G., S. E. Gardiner, H. C. M. Bassett. B. L. Wilson and L. L. Consaul (1998) Implications from SDS-PAGE analyses of seed pro- teins in the classification of taxa of Festuca and Lolium (Poaceae). Drymeja group Biochemical Systematics and Ecology 26: 51 1-33. Dorsal view: L:B 29-38; short embryo (17-32) Pahkinskiene, I., K. Amamthawat-T6nsson, M. W. Humphreys, V. Lateml view: Paplauskiene and R. N. Jones (1998) New molecular evidence on Ventral view: grooved; short linear hilum (40-45%) genome relationships and chromosome identification in fescue Cross-section: dorsally compressed (T:B 52-76) (Festuco)and ryegrass (Lolium). Heredity 81: 659-65. Length: short (2.02-2.79mm) Saint-Yves, A. (1926-1929) Contribution +3 l'btude des Festuca (subgen. Eu-Festuca) de l'orient, Asie et Region MBditerraneenne voisine. Pratensis group Candolleo 3: 321-466. Dorsal view: L:B 23-34; short embryo (27-37%) Lateral view: Ventral view: grooved; long linear hilum (69-93%) Cross-section: dorsally compressed (5046) Length: medium (2.91-3.18mm) 15. LOLZUM Group l Group 2 1. L. multiflorum Lam.'* 5. L. persicurn Boiss. & Hohen, ex Boiss.* 2. L. perenne L.' 6. L. remotum Schrank' 3. L. rigidum Gaudinl* 7. L. temulentum L. syn. L. loliaceum (Bory & Chaub.) Hand.-Mazz. i. var. temulentuml* 4. L. subulatum V~S.~ ii. var. arvense (With.) Liljebl. Distribution and systematics in the production of alkaloid toxins (still poorly characterised) that Perennial or annual, about eight species, native to Europe, North are poisonous to grazing animals and humans. The best documented Africa and temperate Asia; important as forage crops, lawn grasses instance is Lolium ternulentum, known since ancient times as a con- and weeds of crops. The genus is understudied and its phylogeny taminant of flour which can lead to poisoning of humans (p. 14). poorly understood, but a division into the two groups defined here is widely supported. ?tYo species, L. remotum and L. ternulentum, Seed morphology are only found in crop fields, of flax (and, recently. Ornithopus Glurnes: tough papery sativus) and cereals respectively. In the Near East, L. temulentum Lemma and palea: tough papery, both adherent, distinctly punctate is restricted to areas with a more Mediterranean-type climate Dorsal view: L:B 20-35; short embryo (21-40%) (Scheibe 1934).L. remotum is today mainly restricted to central Lateral view: grooved and northern Europe, but is archaeologically attested from the Near Ventral view: grooved: long linear hilum (72-92%) East and Aegean. The origin of these species remains unclear, but Cross-section: dorsally compressed (TB45-77) they probably share a common pre-agrarian ancestor. Isozymes and Length: medium (3.49-6.32mm) other genetic evidence suggest that L. persicum is the most closely related of the other species. It grows abundantly in central and eastern Turkey and the 'inner' Near East. It is also largely restricted to fields and may thus also represent an obligate segetal weed rather than the wild ancestor. This group of three species is well defined. Species relationships in group 1 are less certain, but the species probably form one group. L. perenne, L. rnultiflomm and L. subula- tum are closely related, as is the highly variable L. rigidum which has been proposed as the ancestral species for L. rnultiPomrn and L. perenne (which would be a surprising reversal of the more usual evolution from perennial to annual). L. multiflorum is believed to be native to Europe but is common on wasteground in the Near East. L, subulaturn is a relatively uncommon form closely related to L. rigidum but with larger spikelets. Endophytic fungi infect the fruits, stems and leaves of some spe- cies of Loliurn, Festuca and some other genera. The infection results 15.1 Lolium multifforum RMN 4549 (magnlficanon 10x1 54 IDENTIFICATION GUIDE FOR NEAR EASTERN GRASS SEEDS 15.5 LoliumpersicumGCH 3809 (magnification10x) 15.7 Loburn rrg~dumNM 943 (magnification 1Ox) 0 15.10 Lolium ternulenturn RMN 697 (magnification 1Ox) 0 L. rnultiflorum A persicurn Cl perenne A remotum X rigidurn 0 5 10 15 20 25 L (mm) Length and w~dthmeasurements of Lolium grains Notes dorsal surface, short (< 4.5mm in reference material). V-shaped scutellum on most but not all caryopses. The distinctive, 2. L. persicurn: not turgid, relatively flat dorsal surface, caryopses punctate, Iemma and palea often remain attached to charred variable in length but often over 5mm, overall distinctly larger material. Within the genus, well-preserved material can usually be than the first group. identified to one of four groups: 3. L. rernoturn: turgid (swollen, rounded), short caryopses 1. L. perenne/multiflorurn/rigidum:not turgid, relatively flat (< 4mm). GENUS CATALOGUE - POOlDEAE 55 4. L. ternulentum: turgid (swollen, rounded), caryopsis highly Charmet, G., C. Ravel and E Balfourier (1997) Phylogenetic analysis in variable depending on position of floret in spikelet, but usu- the Festuca-Lolium complex using molecular markers and ITS rDNA. ally over 4mm long. Theoretical and Applied Genetics 94: 1038-46. Variability in Lolium caryopsis size and shape and the effects of Hjelmquist. H. (1950) The flax weeds and the origin of cultivated flax. Bofaniska Notiser 2: 257-98. [pp. 285-9, Lolium remotum] charring make secure identification of small numbers of caryopses Kislev. M. E. (1980) Contenu d'un silo a bl6 de 1'6poque du fer ancient. difficult. The characteristics of the grains -and the presence of In Ell Keisan (1 971-1 976). Une cite phhicienne en Galilee, J. Briend more than one morphological group - will be far easier to detect in and J. B. Humbert (eds), pp. 361-79, pls 139-40. Paris: J. Gabalda. reasonably large samples of grains. [pp. 366-72. Lolium temulentum] Kroll, H. 1. (1983) Kastanas. Die Pflanzenfunde. Berlin: Volker Spiess, Other genera Prahistorische Archaologie in Siidosteuropa 2. [pp. 82-6, 102-3, L. See under Festuca13. Lolium grains are sometimes confused for temulentum, L. remotum] those of Hordeurn", but are easily distinguished by the V-shaped Loos, B. I! (1993)Morphological variation in Lolium (Poaceae) as a measure palea grooves on the ventral face. of species relationships. Plant Systematics and Evolution 188: 87-99. Morosov, V. A. (1929) On the grains of Loliurn remotum Schrnk.
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