GM Llp ie ities seta ab Ee ine 0 A a 2 Sem DIG Ee PPCM EE DEE aN sy x ~ RN NY NN N X % a AS \ N x a % nN oN Lilian 4 an aR Cy mene wen ei halls al fae temo EVN RNASE EVENS SUSE . AQAA AN ERR KA AAG SY \ CRA SY SS \ QQ \\ Ww SY : ae AX SOs SNS RRO A : LAN \ \< \ QSOs \ RORY SS WAAKK NY SS SN AY AS N A AX \ NS ; ‘i NS SN ‘ . \ SSS ANS SSS \ SAN BERRY SS SS S AY AN \ SN SENN SY WAN SS \ SQW 2h AMARA ay feat Brit ‘ wea IOWA GEOLOGICAL SURVEY Bulletin No. 4 THE WEED FLORA OF IOWA Eby L eee ales GEORGE F. KAY, STaTE GEOLOGIST JAMES H. LEES, ASSISTANT STATE GEOLOGIST DES MOINES: PUBLISHED FOR IOWA GEOLOGICAL SURVEY 1913 DES MOINES ROBERT HENDERSON, STATE PRINTER JOHN M. JAMIESON, STATE BINDER 1. OF D. WAY 7 1914 GEOLOGICAL BOARD. Eismihxcellency, George W. Clarke: .)o2.. -.. 0.2.2. 0.055 Governor of Iowa EEKCHAMmme OLATIbere Mls] CAV CLV As atiaha io sire rat opoustonitcle anets: sysbadeceon je speielaraieitele Auditor of State PCN eMES OWI ELMe ss ley arrete ste ciee) vie susie President State University of Iowa ey monly Aes SATS OM rer c.0 seo) «scsi, si eueseieleve miso President Iowa State College (CPPCC Vic cis cc cies exe a ere eia. eee President Iowa Academy of Science ADMINISTRATIVE OFFICERS. GeOGe mE) RAY oso. oasis s Ce, RARE IAT, MMM OU POT at Rte a State Geologist 20083 IST Ieee oping dsich oon veo umes poeta Assistant State Geologist EIDE RUMEN EG wimaiiee ec AN ee es hoy uns oa SOCRetAry LETTER OF TRANSMITTAL Iowa Geological Survey. To Governor George W. Clarke and Members of the Geological Board: GENTLEMEN: I submit, herewith, a bulletin on The Weed Flora of Iowa and recommend that it be published for distribution among the people of the state. ‘The thanks of the whole state are due Dr. L. H. Pammel of the Iowa State College of Agriculture and Mechanic Arts for the preparation of a comprehensive and thorough report, represent- ing many years of careful and painstaking scientific work, on a subject that is most intimately related to agriculture, the industry that far exceeds in importance all other industries of our great state. The Survey wishes to express its thanks to Dean Curtis, Director of the Iowa Agricultural Experiment Station, for his co-operation, and for his kindness in permitting the Survey to include in the bulletin on weeds results of investigations which were carried for- ward by Doctor Pammel while connected with the Experiment Station. The Iowa Geological Survey had the honor, about ten years ago, to publish a complete monograph by Doctor Pammel on the Grasses of Iowa. This publication proved to be of great value, and it is with the fullest confidence that the bulletin on The Weed Flora of Iowa will be of equal if not of greater service to the agricultural and related interests of the state that it is now pre- sented for publication as Bulletin 4 of the lowa Geological Survey. I have the honor to be, Yours very sincerely, Grorce F. Kay, State Geologist. THE WEED FLORA OF IOWA By L. H. PAMMEL WITH THE COLLABORATION OF. CHARLOTTE M. KING J. N. MARTIN J. Cc. CUNNINGHAM ADA HAYDEN and HARRIETTE S. KELLOGG TABLE OF CONTENTS CHAPTER I. Page MS Cregeen LAIN alleneyeyeic et ecu wenenneietes cre Row ccna lreallel alone, seer ervunoe le clahelbie era wee 1 CHAPTER II. The General Characters of Seeds. BORSA ch i a eI Fey aN A/a Ml Dele ie 405 CHAPTER III. The Microscopic Structure of Some Weed Seeds....................-.. 503 CHAPTER IV. Morphology of Flowers and Leaves.......... Per aan ey a Asta cepa n aerate oak ne NUN oe 589 CHAPTER V. Cale rg mG lem OUS starccet ste ous tetera orca Gia reevencl seatm a: St Setar atireene slave: maecsie balers 627 CHAPTER VI. ECOULSH AMC OOt-StOCKS! Ob NVECUSm anne daistccde aelsule oss a ee cae one ome 641 CHAPTER VII. Number and Kinds of Weeds in Different Soils........................ 655 CHAPTER VIII. MMA UNO MISTS SHOeNVCCUS Hare pace cake se alors lara ecdtava seus oe aereractte, oe MNecciteensiale Ceeeiece 669 CHAPTER IX. NCCC am VIN eae OMmec erento auc aun ska Rhos Bas er eels Wendie) am ht e 685 CHAPTER X. ISA SG LAST WHY SRG) ai ssn ste see aCe Petco eae RS Te RS a A TTA CHAPTER XI. ENEMMIOO Sy Ole WICC US a eters seers seis ie o's p.oleisiors wate alee bea do weahe edhe ivan 783 CHAPTER XII. Weed! Bing! Seeel SIRNAS Se Ramee rec oe oleae et Re ier ry rer oc 791 CHAPTER XIII. ISUISIONPA BANG! TEMTOIbIOVSR FEN olalira yoke Hike Amiens arte Ele Gee ane a 817 GIOSSEIAT ~ iodo 5 5) 5 S!SfD ie as Ho Goa Eee EEO Eee ed es 862 PREFACE Weeds do an enormous damage to the crops of lowa. A con- servative estimate places the injury at $25,000,000 annually. This loss could be largely avoided if we had more concise information - on the subject, and if we could conserve: the matchless resources of our soil by keeping the weeds down, the farmers would be greatly benefited in a financial way. It would seem appropriate, therefore, to publish a volume of the Weed Flora of the state at this time. _ The need of a volume dealing with Iowa weeds as a feature of the flora of the state has long been felt by the public schools. Many papers have been published by the Iowa Agricultural Ex- periment Station, but these papers are mostly out of print. This work is much more comprehensive than anything heretofore pub- lished in this state. The title indicates that it is not a weed book, but rather ia weed flora. Much stress has been laid on the geograph- ical distribution of weeds. It is a contribution to the local flora of the United States. The chapter on the geographical distribution of weeds will be found of special interest to the phytogeographer. I desire to ex- press my thanks to Dr. M. L. Fernald, Messrs. R. I. Cratty, F. W. Paige, O. M. Olson, J. P. Anderson and Prof. B. Shimek for assist- ance in giving the geographical distribution of Iowa weeds. The chapter on the microscopic structure of weed seeds brings together in English much on the subject which has hitherto been inacces- sible to the student not familiar with German or French ltera- ture. The chapter on morphology will greatly help the student to_ understand the changes occurring in the development of the flower and the formation of the seed. The chapter on the use of weeds in medicine will be of interest to those who occasionally make use of wild plants for medicinal porposes. The chapter on seeds de- scribes a large number of weed seeds; this will be found of value to those engaged in a study of seeds. The chapter on various weed laws gives a summary of laws in various states in the Mississippi Valley. In compiling these xii PREFACE laws I have been aided by Mr. Small of the Iowa State Library, Mr. Moore of Wisconsin, Mr. Michel of South Dakota, Dr. Howard of Missouri and H. L. Bolley of North Dakota. In the preparation of this Weed Flora, I have been greatly assisted by Professor J. N. Martin, who has written the chapter on morphology of the plant; Miss Ada Hayden, who has written the chapter on dissemination; Professor J. C. Cunningham, who has written the chapter on roots and underground organs; Miss Charlotte M. King, who is the joint author of the chapters on gross characters and microscopic structure of seeds. Miss King is also responsible for many of the excellent drawings. The chapter on medicinal weeds was prepared by Miss Kellogg. I am greatly in- debted to her for painstaking editorial work and for preparing the bibliography and the index. In the matter of bibliography, it has seemed best to divide the subject into various sections so that the student may easily find the desired literature. The bibliography is not complete but enough papers are given to enable the student to find the important literature. JI am indebted to Dr. Clark, of the Canadian Seed Laboratory, for the privilege of using some of the admirable illustrations of the Canadian work on weeds, also to Dr. Ernest Bessey for illustrations from Beal’s Weeds of Michigan, and the classical Hillman seed figures in the Michi- gan bulletin, and to the Nevada Station for the use of the Hill- man cuts. Some of the Hillman and a few other figures have been taken from the government publications. I am also indebted to various publishers for figures which have been taken from sev- eral textbooks of botany, as the Bergen & Davis book published by Ginn & Co.; several botanical works like Thomé’s published in German, and to the Connecticut Experiment Station. A few fig- ures have been taken from the Botanical Gazette. Credit is given under each figure. The photographs were made by F. E. Colburn, photographer at the Iowa Agricultural Experiment Station, G. T. Hart and C. R. Quade. The clerical work of the volume was per- formed by Miss Bertha Herr and my daughter, Miss Harriet. Mr. Burlingmair assisted in a study of weeds in different fields. To Mr. James H. Lees for his assistance in proof reading and edi- torial work I am also indebted. To all I wish to express my sincere thanks. The reader will find it advisable to have several of the recent treatises on weeds. Mention may be made of my work ‘‘ Weeds of the Farm and Garden,’’ a general treatise of three hundred pages PREFACE xiii with numerous illustrations; the work of Fletcher and Clark of Canada; Bolley’s *‘North Dakota Weeds;’’ Blatchley’s ‘‘Indiana Weed Book;’’ W. J. Beal’s ‘‘The Weeds of Michigan.’’ A list of these publications and where they may be obtained will be found in the bibliography. Ames, Iowa, December 18, 1912. . : L. H. PAMMEL. Page 754 Page 837 Page 838 Page 842 Page 858 Page 861 Rosenberg, 1882 should be 1782. Patrick, P. E., should be Patrick, G. E. Blankenship should be Blankinship. L. T. Henderson anol be L. F. Henderson. Panton, J. H. Weeds of Ontario should be listed on page 845. T. L. Williams should be T. A. Williams. CHAPTER I. DESCRIPTIVE MANUAL L. H. PAMMEL CHAPTER I. KEY FOR FAMILIES. I. Plants without true flowers; not producing seeds...Pteridophyta. Stemsmyoimted, TUShliKe. jac ccs cero ders ec ne le ss ec Hquisetaceae. I. Plants with true flowers, stamens, and pistils and producing seeds. Spermatophyta. II. Ovules not borne in a closed ovary (Pine, Spruce).......... Gymnospermae. II. Ovules borne in a closed ovary (Rose, Willow, Corn, etc.).... III. ITI. Angiospermae. Stems endogenous without central pith; no annual rings; parts of the flower usually in threes; single cotyledon... : Monocotyledoneae. 1. Grasslike plants 2. 2. Flowers enclosed by chaff-like scales. Stems hollow; sheaths of leaves split...Gramineae. Stems solid; sheaths of leaves not split. Cyperaceae. © 2. Flowers not inclosed by chaff-like scales. .Juncaceae. 1. Plants not grasslike; flowers with a perianth of 6 DICCESGMSCAIMENISS Gah eienevatel svar cnoveusrate eben cis atcher cere Liliaceae. Stem formed of bark, wood, and pith, exogenous; ieaves netted-veined; embryo with a pair of cotyledons......... Dicotyledoneae. 1. Corolla absent 7, IPMGNOES WSN; Op SCUNPA TS Gia bonoodbodE Chenopodiaceae. 2. Plants not fleshy or scurfy 3. 3. Ovary free 4. 4, Flowers unisexual. Ovaryral=celle@ dl We erciesletere ate we eranestcleisns Urticaceae. Oviatyars-¢ clledeea anes eiciseo Huphorbiaceae. 4. Flowers perfect. Calyx and bracts greenish and scarious...... Amaranthaceae. Calyx generally corolla-like. 1. Fruit a i1-seeded achene..... Polygonaceae. 1. Fruit a 5-12 seeded berry. ..Phytolaccaceae. Se OVA MERTON te alsa eleve eles siedele re: wicvers Nyctaginaceae. 1. Calyx and corolla present. 2. Corolla of separate petals 3. 3. Plants fleshy, flowers yellow........ Portulacaceae. WEED FLORA OF IOWA 3. Plants not fleshy 4. 4. Pistil single 5. 5. Flowers regular. Stamens numerous, free...Ranunculaceae. Stamens numerous; sepals and petals pres- ent, inserted on calyx........... Rosaceae. Stamens 10; fruit a legume...Leguminosae. 4. Pistil compound 6. 6. Ovary free. Ovary l-celled ........... Caryophyllaceae. Ovary more than 1-celled 7. 7. Ovaries united into a ring...Malvaceae. 7. Ovaries not united into a ring 8. 8. Leaves simple. With punctate dots; stamens numer- OUSHARie aseeneener A rors ., Hypericaceae. Leaves not with punctate dots; sta- NNT Hosgaqasosnabdousads Cruciferae. Leaves compound, pinnately 3-foliate. Anacardiaceae. Leaflets 3, obcordate..... Oxalidaceae. Leaflets 5-7 pairs..... Zygophyllaceae. Leaflets 3, viscid or fetid herbs..... Capparidaceae. 6. Ovary adherent; flowers in umbels..... Umbelliferae. 6. Flowers not in umbels....... Onagraceae. 1. Calyx and corolla present, petals more or less united. 2. Flowers regular 3. 38. Plants with milky juice. Stamens united .............-.0e8- Asclepiadaceae. Stamens distinct ................... Apocynaceae. 3. Plants without milky juice. 4 Plants Gwinn eee ce eesierete eters Convolvulaceae. 4, Plants not twining 5. 5. Stamens 5 or more 6. 6. Style 2-cleft; flowers not in heads; fruit 2-4 seedlike nutlets.......... Boraginaceae. Fruit many seeded pod...Hydrophyllaceae. Flowers in heads; anthers in ring or tube about the style......... Compositae. 6. Style 1; fruit many seeded..... Solanaceae. 5. Stamens fewer than corolla lobes Caulescent, flowers blue ........ Verbenaceae Acaulescent, flowers greenish. Plantaginaceae 2 Flowers irregular. Stems 4-angled; ovary deeply 4-lobed...... Labiatae. Stems not 4-angled; ovary 2-celled. .Scrophulariaceae. DESCRIPTIVE MANUAL 4) PTERIDOPHYTA, FERNS AND THEIR ALLIES. This group of plants, sometimes called vascular cryptogams, Is represented in our flora by the maidenhair fern, brake, spleenwort, shield fern, ete. The Boston fern is frequently cultivated. Fic. 1. Common Horsetail (Hquisetum arvense). Roadsides, fields, common everywhere in Iowa. (Photographed by Colburn.) Fic. 1-A. Distribution of Common Horsetail. 6 WEED FLORA OF IOWA EQUISETACEAE, HORSETAIL FAMILY. This small family of rushlike plants contains a few species only and but one that is weedy. Common Horsetail (Equisetum arvense L.). Description—A rushlike perennial with running rootstocks and annual stems; branches in whorls; fertile and sterile plants, the fertile appearing early in spring with a terminal cone, yellowish in color, bearing the spore cases (sporangia) underneath a scale; spores provided with hygroscopic bands; sterile frond with whorled branches. Distribution — Widely distributed in North America, common in sandy moist fields, on railroad embankments; common in Story, Boone, Carroll, Crawford, Harrison, Woodbury, Clinton, Dubuque, Lee, Page, Polk, Cerro Gordo, Emmet, Webster, Marshall, Johnson, Winneshiek and Allamakee counties Extermination.—This is a most persistent perennial; only by giv- | ing frequent shallow cultivation after small grain is removed during the summer can the weed be kept in check. SPERMATOPHYTA, FLOWERING PLANTS. The seed plants have stamens and pistils and reproduce by seeds. Represented by the pine, spruce, hemlock, wheat, rye, corn, rose, maple, ash, aster, goldenrod, squash, ete. GYMNOSPERMAE, GYMNOSPERMS. The seeds are not inclosed in an ovary. Trees or shrubs gener- ally with needlelike or scalelike leaves; represented in Lowa by the red cedar, white pine, etc. Noné are weedy. ANGIOSPERMAEH, ANGIOCSPERMS. Ovules borne in a closed ovary. Represented by a large number of our native and cultivated plants, like wheat, corn, lily, rose, clover, tomato, ete. MONOCOTYLEDONEAE, MONOCOTS. Plants with endogenous stem, the woody fibers in bundles dis- tributed through the pith. Annual ring absent. Flowers gen- erally on the plan of 3; embryo with a single cotyledon. Corn, lily, onion, asparagus, blue grass, switch grass are representatives. DESCRIPTIVE MANUAL ts GRAMINEAE, GRASS FAMILY. This large family is of great economic importance, since it con- tains many of our food plants, including the well known cereals, \ tn Fic. 2. Johnson Grass (Sorghum halepense) ; a, sessile spikelets. A most trou- blesoume weed. (Lamson-Scribner, U. S. Dept. Agr.) Fic. 2-A. Distribution of Johnson Grass. Reported recently from southwestern owa. 8 WEED FLORA OF IOWA rye, wheat, oats, barley, corn, kaffir corn and millet, besides such forage grasses as blue grass, timothy, brome grass, foxtail, and a few ornamental plants, ike pampas grass, ribbon grass, ete. Johnson Grass (Sorghum halepense (.) Pers.). Description.—A_ stout perennial, with smooth, erect, simple culms, 3-5 feet high, and strong, creeping rovt-stocks; leaves elongated, 14-34in. wide, acute; ligule ciliate, and on the back where leaf-blade joins the sheath there is more or less pubescence ; panicle open, 6-12 in. long, the whorled branches naked below, the 3-5-flowered racemes clustered towards their extremities; pedicels of the staminate (rarely neutral) spikelets pilose with stout hairs; sessile spikelet broadly lanceolate, acute, 2-3 lines long, pale green or violet, becoming dark or nearly black at maturity; callus small, obtuse, shortly and sparsely barbate; first glume coriaceous, spar- ingly pubescent on the flattened back, 5-7-nerved; second glume similar and equaling the first, convex below, subcarinate above, acute, the hyaline inflexed margins ciliate; third glume a little shorter than the outer ones, membranous, faintly 2-nerved, the in- folded margins ciliate; fourth glume broadly oval, obtuse, nearly 1% shorter than the second, 2-lobed or bidentate at the apex, ciliate awned; awn 5-8 lines long; palea a little shorter than the glumes, nerveless, ciliate. Introduced and cultivated in many southern states for hay; in many places it has become a dangerous weed, difficult to exterminate. Distribution.—The weed is common in the south, often a most troublesome weed. It has been reported as persisting in the vicinity of Hamburg, Fremont county. Extermination.—Use the same methods as for quack grass. This may become a most troublesome weed. Finger Grass (Digitaria sangwinalis (L.) Secop.). Description—A much branched, leafy annual, 1-3 ft. high, spreading on the ground, with erect, smooth, spreading culms, fre- quently rooting at the lower joints, joints sometimes smooth, though more frequently bearded with deflexed hairs; sheaths loose, gener- ally pilose, hairy, ciliate on the margins, with a membranaceous ligule; leaves 2-4 in. long with rough margins, occasionally pilose at the base; flowers produced in digitate spikes, hence the common name finger grass; spikelets less than 14 in. long in pairs, 1 nearly sessile, the other with a stalk, each flower consisting of 2 sterile DESCRIPTIVE MANUAL Fic. 3. Common Crab Grass (Digitaria sanguinalis). Common in fields, gar- dens, meadows. Rooting at the joints. (Photographed by Colburn.) — Si a ator Mitchel ties ROMA SauPene * fe le Kossuth = ote a Ay . Bai AO z *. ‘|: CL |RAko + ° . is . mf . 2 : 2. 2 Sure . a Tel . + dubug Z ce . re ° 47 Sle ot a is ee : 5 rene eo bt t,]. Ce ole = +. es Geet ate = Se a hee eet = 2 Clinton C a - tr . eo bohvsbw eis e : es| Ma Ra eed lie + - = = Hieiscarine ONG fs z Keokuk = awaited mie + 5 ote ° a5 +> . =o ’ # Henn / | E ae + a A faye Decatur a c + | +. Fe + the he ee Se ee ee % Fic. 3-A. Distribution of Common Crab Grass. glumes and the flower proper; the first glume very small, the sec- ong about 14-24 as long as the spikelet, usually hairy on the mar: gin, the third glume somewhat longer than the fourth, which is 5- 10 WEED FLORA OF IOWA nerved and usually silky-villous along the marginal nerves, fourth glume smooth and acute; fruit minute, pitted and cross-striated, light straw color except where the sterile glumes remain attached, which are gray in color and minutely hairy. Distribution.—This European grass is cosmopolitan; abundant in the eastern and southern states and in California; common in all parts of Iowa, more particularly in gardens, corn fields, and streets. Extermination—tThis grass is much more difficult to remove than the foxtails because it roots so readily at the joints. Thorough cultivation will remove the weed. Do not allow it to go to seed. Chemical Composition—Common crab grass (Digitaria sanguin- alis) has been used as a forage plant in many parts of the United States and many chemical analyses have been made. Analyses are reported from Mississippi, Tennessee and lowa. The Iowa analysis reported by Weems is as follows: NATURAL CONDITION Albu~ Crude Nitrogen Sample Water Fat Protein minoids Fiber Ash free extract ‘he a ee 66.95 Ul 2e02 (1.98) 8.62 4.10 16.70 WATER FREE SUBSTANCE see ase 2 | eee eee | 3.34 | 7.61 | (5.98) | 26.11 | 12.41 | 50.53 Smooth Crab Grass (Digitaria humifusa Pers.). Description.—An annual 6 in.-2 ft. high, closely resembling D. sanguinalis in habit, but smooth throughout, excepting for a few hairs at the throat of the sheaths; spikelets 2-7, smaller than in D. sanguinalis, about 1 line in length; first glume very minute or obsolete; second and third glumes nearly equal in length, or the second a little shorter than the fourth, pubescent at the back. Distribution—Smooth erab grass is native to Europe but is now cosmopolitan; in eastern North America from New England to Texas and Mexico, Rocky mountains and Pacific coast; less com- mon in Iowa than common crab grass; rapidly spreading in the state, more particularly in gardens; common in lawns and pas- tures. SES: [Z = ex aS Sc === SS a xe Fic. 4. Fic. 4-A. Tt DEE SO _ S$ S Figure 4 Crab Grass (Digitaria sanguinalis). Figure 4A —— SEOQEE Wate: Fe . © | Mitchell ellie : 5 . S 45 . ° is MI so |e | elie \ . \ IaiHers| * « we Wig. 108-A. Distribution of White Sweet Clover. 189 190 WEED FLORA OF IOWA B bs 5 a 29 z # Ge o 4 = Water) 2.25225 ee ee 79.35 6.02 waren Ash) assess et oe ae ee ee eS ne 2.10 9.57 10.18 Hither. extract): =2222 28 ee ee en ee ae 0.53 2.42 2.52 Crude fiber (222s 22 ee ae eee eee ea eee 4.78 21.77 23.16 Crude: protein, | 23-222 2s ee ee el ee eee 3.96 18.00 19.15 Nitrogen, free. extractq. 2s seo ss ee Se eee 9.28 42.22 44.99 Black Medic (Medicago lupulina L.). Description—A procumbent, pubescent, annual; compound leaves trifoliate ; leaflets wedge-shaped, obovate, toothed at the apex; flowers yellow, in short spikes; pods kidney-form, 1-seeded. — Distribution.—Native from Europe, common in alfalfa fields in the Rocky mountains and on the Pacific coast; common in waste places in eastern North America; reported from a number of coun- ties In Jowa, as Wright, Kossuth and Story. | Extermination—This annual is not difficult to exterminate by giving thorough cultivation; care should be used in planting clover and alfalfa seed, because this weed seed is a common impurity of the latter. The seed also retains its vitality for some time. Chemical Composition.* FRESH OR AIR DRY MATERIAL. x : Nitrogen free Water | Ash | Protein Fiber Bench Fat 78.52 | 1.37 | 3.40 | 6.31 9.20 | iar bl WATER FREE SUBSTANOE. | 6.4 | 15.8 | 29.3 43.4 | 5.1 *Compiled by Jenkins and Winton, Bull. Off. Exp. Sta., Bull. 11. DESCRIPTIVH MANUAL iii Fig. 109. Black Medic (Medicago Wwpulina). Clover and alfalfa meadows. (Photographed by Hart.) Fig. 109-A. Distribution of Black Medic. 192 WEED FLORA OF IOWA Pink Parosela (Dalea alopecuroides Willd.). Description—An erect annual from 2-3 ft. high, with pinnately- compound leaflets; smooth flowers, whitish or light rose-color, in cylindrical spikes; calyx villous; seeds kidney-shaped. Distribution—F rom Minnesota to Alabama and the Rocky moun- tains, common in western Iowa and introduced eastward in Wright, Boone, and Story counties. Extermination.—It is sometimes found in clover seed grown in western Iowa, also in alfalfa seed from the west, therefore care should be used in the selecting of seed; succumbs readily to culti- vation. Fig. 110. Parosela (Dalea alopecuroides). Common in western Iowa and along railroads. (Photographed by Quade.) DESCRIPTIVE MANUAL 193 Fig. 110-A. Distribution of Parosela. Rattle-box or Milk Vetch (Astragalus canadensis L.). Description.—A tall, erect, smooth or pubescent perennial, 1-4 ft. high; leaves compound, leaflets 21-27, oblong; flowers greenish cream-colored in spike of variable length; pods crowded, smooth, terete, occasionally somewhat suleate; the seeds separate from the pod, rattling, hence the common name. Distribution—Common in the northern Mississippi valley, east to New York and south to Georgia. Common on borders of thick- ets, woods and native meadows. Extermination.—This weed is easily exterminated by cultivation. 13 194 WEED FLORA OF IOWA Fig. 111. Rattle-box (Astragalus canadensis). Woodland pastures. (Photographed by Charlotte M. King.) Purple or Stemless Loco Weed (Oxytropis lamberti Pursh.). Description.—Nearly acaulescent, perennial herbs or shrubby plants, with tufts of very numerous short stems coming from a hard and thick root-stock containing many scaly stipules; stems and leaves covered with silky and finely appressed hairs, or smoothish ; leaves pinnate; leaflets linear; flowers racemose or spicate, rather large and elongated, purple, violet, or sometimes white ; stamens dia- delphous; keel tipped with a sharp projecting point. This is one of the loco weeds, poisonous to cattle. Distribution—Western Minnesota, western Iowa, and Missouri to Texas, and New Mexico, north to British Columbia, and northwest territory. _Extermination.—It seldom gives much trouble in cultivated fields. DESCRIPTIVE MANUAL 195 Fig 112. Purple Loco Weed (Oxytropis lamberti). A weed poisonous to cattle. (Drawn by F. C. Collins.) : Fig. 112-A. Distribution of Purple Loco Weed. 196 WEED FLORA OF IOWA Wild Liquorice (Glycyrrhiza lepidota (Nutt.) Pursh.). Description.—A branching perennial 2-3 feet high, leaves com- pound of 15-19 oblong-lanceolate leaflets with mucronate points; young leaflets sprinkled with a resinous material ; flowers in spikes; short peduncled; whitish; pods oblong, covered with hooked prickles, resembling a cocklebur. Distribution—Common in the west, western lowa to Rocky mountains, Utah, New Mexico, and Montana to Canada, also re- bp YY TY Fig. 113. Wild Liquorice (Glycyrrhiza lepidota.) Occurs in western and cen- tral parts of the state. DESCRIPTIVE MANUAL 197 Z a ‘ JANN Sane ERELERERES = Fig. 113-A. Distribution of Wild Liquorice. ported from the Great Lakes; commonly reported from Story, Greene, Pottawattamie, Harrison, Monona, Fremont, and Decatur counties. Extermination—tThis perennial plant whose burs resemble the cocklebur is easily exterminated by cultivation. Do not permit the seeds to mature. Cut off the plants at the surface of the ground to prevent flowering, if they occur in the pasture or meadow. Where the land can be cultivated, give the ordinary plowing, fol- lowed with harrowing to bring the roots to the surface so they are exposed to the sun. A few days’ exposure to the sun will kill them. Common Vetch (Victa satiwa L.). Description.—A smooth or slightly pubescent annual from 1-21 ft. high with simple stem; leaflets 5-7 pairs, obovate-oblong to lin- ear, notched or mucronate at the tip; 1 or 2 nearly sessile flowers borne in the axils of the leaves, corolla violet-purple; pod linear, several-seeded, seeds black. Distribution —This weed has long been known as troublesome in the grain fields of Europe, and in the northern states. It is particu- larly abundant in northeastern and northwestern Jowa and in some of the grain growing sections of the southern part of the state. Extermination.—Clean seed sown in clean soil is the only method of displacing the weed. 198 WEED FLORA OF IOWA Fig. 114. Common Vetch (Vicia sativa). Common in grain fields, especially wheat fields, frequently found in screenings from flour mills. (Photographed by Colburn.) ENE AE G Fig. 114-A. Distribution of Common Vetch. DESCRIPTIVE MANUAL 199 Chemical Composiiion.—The chemical composition of vetch (Vicia sativa) according to U. S. Dept. Agr., 1880; p. 152, is as follows: FRESH OR AIR DRY MATERIAL. i=} ~~ He # 2S = He = 4 ay ie ur opahae a Guiees in’ full bloom’ 2.2. | 86.20 | 1.60 | 4.14 | 2.11 | 5.34 | 0.61 WATER FREE SUBSTANCE. | 11.6 3.00 | 15.3 | 38.7 | re Wild Bean (Strophostyles helvola (L.) Britton). Description.—An annual, with prostrate stem; compound leaves ovate to oblong-obovate, with a prominent rounded lobe at the base; corolla greenish white and purplish; pod 4-8-seeded, large, usually - pubescent. Distribution—Common in sandy places in northern United States; from Wisconsin, Minnesota, and Texas east to Massachu- setts ; common in gravel pits on Muscatine Island, along Des Moines, Cedar and Iowa rivers. Extermination.—Kasily exterminated by cultivation. OXALIDACEAE, WOOD SORREL FAMILY. This family contains the cultivated yellow and purple-flowered oxalis. Few of the plants are economic. 200 WEED FLORA OF IOWA Fig. 115. Wild Bean (Strophostyles helvola). Common in sandy or gravelly soil, sandy river bottom. (Photographed by Colburn.) Fig. 115-A. Distribution of Wild Bean. DESCRIPTIVE MANUAL 201 Field Sorrel (Oxalis stricta L.). Description—aA pale green pubescent annual or perennial ; leaves compound, with evident stipules; flowers pale yellow, cymose, 1-4, at length deflexed ; in fruit columnar, short-pointed. Distribution.—Common in fields and waste places from New Eng- land to Dakota; common in Iowa probably over the entire state. Extermmation.—Persistent cultivation and crop rotation will usually exterminate the weed. Fig. 116. Yellow Field Sorrel (Ozalis stricta). Common in fields. a, Trichome or plant hair. (a drawn by Charlotte M. King. The whole plant photographed by Hart.) 202 WEED FLORA OF IOWA HALAL ames ee Boe oe US SPENSER wee Fig. 116-A. Distribution of Yellow Field Sorrel. Field Sorrel (Oxalis cormculata L.). Description.—An erect or decumbent perennial herb, spreading by numerous slender, pale runners; leaflets 3; flowers in cymose clusters, yellow; peduncle ascending and sparingly pubescent. Fig. 117. Field Sorrel (Owalis corniculata). Fields, gardens, etc. (Photographed by Colburn. 1. Drawn by Charlotte M. King.) DESCRIPTIVE MANUAL 203 Fig. 117-A. Distribution of Field Sorrel. Distribution—-A very common weed in dry or moist soil through- out eastern North America. Extermination. —Apply the same method as to the preceding species. ZYGOPHYLLACHAH, CALTROP FAMILY. A small family. Caltrop (Tribulus terrestris L.). Description.—The caltrop is a hairy, procumbent annual, branch- ing from the base, producing a stem which is a foot or more long, branches bear numerous small, compound leaves with short pe- duncles and small stipules at the base; each compound leaf has 4-8 pairs of short-stalked leaves; small, yellow, axillary flowers about 1 in. across with peduncle much shorter than leaves; fruit very spiny and divided into two nearly equal parts, each part consists of 2 long spines, 2 shorter and a row of very short ones, forming a crest on the back; 5-angled, spiny fruit splits into 3-5 divisions. Distribution.—Introduced from the Old World and oceurs from the Atlantic states to Nebraska and Kansas; in Iowa has been found only on Muscatine Island. Extermination.—Caltrop is disseminated by wool, and hence the waste of woolen mills should not be thrown in fields. The weed is easily destroyed by cultivation. 204 WEED FLORA OF IOWA ao YS SR Y FSS 4 NY te Q My “oN = Ae QZ ett aS 87000 Sy VS Sh =) SS SSH) NINN sn LOG» \W Wy Zz \ ( NA Wy y NO Sk iB AWS BIOT Ee N NAW a ss XA SU) aaa S KG) p ¥ 1) SW_/ Nv WINE KZ HY) rN agoeg We pa Ground Bur-nut (Tribulus terrestris). waste scil, Sandy Caltrop, Fig. 118. Muscatine Island. (Drawing by Charlotte M. King.) * places, DESCRIPTIVE MANUAL | 205 Fig. 118-A, Distribution of Caltrop. ASA EUPHORBIACEAE, SPURGE FAMILY. Many of the plants of this family contain an irritating milky juice. Few are of economic importance. The poinsettia commonly cultivated in greenhouses, snow-on-the-mountain in gardens, and eastor-oil bean belong to this family. Three-seeded Mercury (Acalypha virginica L.). Description—A smoothish or hairy annual from 1-2 ft. high, turning purple especially in the autumn; leaves ovate or oblong- ovate, sparingly serrate, long-petioled; sterile spike, few-flowered, pistillate flowers 1-3 at the base of staminate peduncle surrounded by a large leaf-like bract; capsule 3-lobed, subglobular, 2-valved earpels. . Distribution—F rom Nova Scotia to Texas and northward to Min- nesota. Common everywhere in Iowa along roadsides and in fields. Especially noticeable in the fall on account of the purple bracts. Extermination.—Three-seeded Mercury is not a difficult weed to exterminate. The small, reddish, striate seeds are expelled from the plant to some little distance in a manner similar to the dispersal of the eastor-oil bean. Thorough cultivation by preventing the for- mation of seed will eradicate the weed. 206 Fig. 119. WEED FLORA OF IOWA’ Three-seeded Mercury (Acalypha virginica). Common in many parts of the state. (Drawing by Charlotte M. King.) DESCRIPTIVE MANUAL 207 Fig. 119-A. Distribution of Three-seeded Mercury. Spurge (Euphorbia presli Guss.). Description—An annual from 1-144 ft. high; erect or ascending ; leaves oblique at the base, ovate, oblong, or sometimes oblong-linear, ala Ct mA, X y lp EER : ty ay SS My | Sic = My | i 4 | Y, \ Fig. 120. Spotted Spurge (Huwphorbia preslii). Common everywhere in the state. r, (Drawing by Charlotte M.. King.) 208 WEED FLORA OF IOWA Fig. 120-A. Distribution of Spotted Spurge. frequently falcate, serrate, generally with a conspicuous red spot, or margin; flowers pedunculate in terminal cymes; appendages en- tire; pod glabrous; seeds ovate sometimes wrinkled. Distribution.—In loose soils and fields from New England and Canada to Wisconsin, Minnesota and Nebraska, also southward. Extermination—tThis annual weed is easily exterminated by cul- tivation; therefore practice rotation of crops and give thorough cul- tivation. Seeds of several species of this genus retain their vitality for some length of time. Milkweed or Creeping Spurge (Euphorbia maculata L.). Description—Slightly pubescent or hairy annual, with prostrate stems; leaves oblong-linear, oblique at the base; pubescent or some- times nearly smooth, usually with a brown-red spot in the center; serrulate above, stipules lanceolate; flowers pedunculate in lateral clusters; glands of the involucre minute; appendages usually red; pods acute-angled; seed sharply 4-angled with 4 shallow grooves across the sides ; trichomes several-celled, gradually tapering to apex. Distribution.—Found in sandy fields, or generally in fields from New England and Canada westward; common in every part of Towa. Extermination.—May be exterminated in the same way as the preceding species. ‘ DESCRIPTIVE MANUAL 209 Fig. 121. Milkweed or Prostrate Spurge (Euphorbia maculata). Sandy fields. (Photographed by Quade. a, drawn by Charlotte M. King.) y . Kossuth OBrien ee 5 4¢ Bucte ‘ car LG ° 5 ies lda 3 + reene| +- |e) © 5 7 ae + : +e Clinton 5 bY ° ei ones} + < Zes| Moi +? ~ 2 . Keokuk npc watid mie] . ES a a . + . . Fig. 121-A. Distribution of Prostrate Spurge. 14 210 WEED FLORA OF IOWA Snow-on-the-mountain (Huphorbia marginata Pursh.). Description.—Stems stout, high, erect, hairy or smoothish ; annual, from 2-3 ft. high; leaves sessile ovate-oblong, acute; uppermost leaves white, petal-like margins. The flowering spurge (Huphorbia corollata L.) with white flowers in forked umbels; long peduncles; involucres showy, white append- ages appearing like petals; deep perennial root; is common in gray- elly and sandy soils in many parts of the state, and it is often weedy. Distribution—Snow-on-the-mountain occurs from Minnesota to Missouri and Colorado; also reported eastward to Ohio and South Carolina. A frequent escape from gardens in Iowa; common only in the western part. Fig. 122. Snow-on-the-Mountain (Huphorbia marginata). A frequent escape from gardens. a, Whole plant, one-third natural size; b, seed capsule, natural size. DESCRIPTIVE MANUAL 211 Fig. 123. Flowering Spurge (Huphorbia corollata). Common in sandy fields, gravel Knolls, and roadsides. (Drawing by Lois Pammel.) Extermination.—This annual weed is easily exterminated by cul- tivation. The young plant should be cut off below the surface of the ground. 212 WEED FLORA OF IOWA Cypress Spurge (Huphorbia cyparissias L.). Description—Plant with perennial running root-stocks and densely clustered stems from 6 in.-1 ft. high; linear, crowded leaves; many-rayed umbel with glands crescent-shaped and granular pods. Distribution.—Common westward from New England to Nebraska usually in the vicinity of gardens and cemeteries. Extermination.—This perennial weed because of its running root- stocks is often difficult to exterminate. The ground should be given a shallow plowing and the root-stocks exposed to the sun. It may be necessary to repeat this process two or three times during the summer. In addition to its propagation by the running root-stocks it also propagates by its seeds. Fig. 124. Cypress Spurge (Euphorbia cyparissias). Yellow ‘flowered’ plant with milky juice and narrow leaves. Escaped from gardens to roadsides. (Photographed by Colburn.) DESCRIPTIVE MANUAL 213 >) NM = SN (Ce Sia ate oe z ee L is Perk - La Se 9 PETELEBRRY Fig. 124-A. “pisteanution of Cypress Spurge. ANACARDIACEA®, SUMACH FAMILY. Many of the plants are poisonous. “Many, like sumach, contain tannin. Some, as the smoke tree, are cultivated for ornamental purposes. Poison Ivy (Rhus toxicodendron L.). Description.—A climbing or trailing shrub sometimes erect, cling- ing to trees or other objects by aerial rootlets; 3 leaflets ; inconspicu- ous flowers; waxy fruit, frequently remaining on plant until late winter or early spring. It is often mistaken for Virginia creeper (Psedera quinquefolia (L.) Greene) which, however, has 5 leaflets. Many persons are sensitive to poisoning from this plant, every part of which contains the poisonous principle. The usual remedy for in- fection from it is to wash the skin with a solution of sugar of lead. Distribution.—Poison ivy is abundant throughout eastern North America and the Rocky mountains. It is common everywhere in Iowa, in hedge rows, thickets or woods. Extermination —Poison ivy is not easily destroyed: because in most cases it is troublesome in wood lots along fences and in yards. It is difficult to destroy except by giving thorough cultivation. If persistently cut off below the surface of the ground it can be de- stroyed. Sodium arsenite at the rate of one and one-half to two pounds to 52 gallons of water will help to destroy the weed. It is not, however, safe to use this since it is a strong poison. MALVACEAE, MALLOW FAMILY. Cotton, hollyhock and okra are well known plants of the family. 214 WEED FLORA OF IOWA Fig. 125. Poison Ivy (Rhus toxicodendron). Common in woods and along fences. - (Photographed by Colburn.) Fig. 125-A. Distribution of Poison Ivy.- DESCRIPTIVE MANUAL 215 Fig. 125-B. Poison Ivy (Rhus toxicodendron). a, spray showing rootlets; ; b, fruit. ; (After Chesnut, U. S. Dept. Agr.) Indian Mallow or Velvet-leaf, Butter-print (Abutilon theophrasti Medice.). Description.—A usually tall annual from 2-4 ft. high; plant with strong odor; leaves velvety, roundish heart-shaped, taper-pointed ; peduncles shorter than the petioles; corolla yellowish ; carpels 12-15, hairy-beaked seeds rough, rather large and blackish. Distribution.—Common in waste places, corn fields, vacant lots, barnyards, ete. Common throughout eastern North America, naturalized from tropical regions, probably India. Extermination.—This plant propagates only by its seed, which retains its vitality for some length of time, having been known to germinate after a period of 60 years. The young plants are easily exterminated. The plant should be pulled up before it begins to 216 WEED FLORA OF IOWA Fig. 126. Indian Mallow, Velvet Leaf, or Butter-print (Abutilon theophrasti). Common in carn fields, waste places, barnyards, (Photographed by Colburn.) flower. Wallace’s Farmer suggests the following treatment: ‘‘It has been seriously thought of by some persons as a substitute for manilla or sisal in the manufacture of binder twine. It is an an- nual, and if not allowed to go to seed, the farm can in time be cleared of it. But it will be a long time. The seeds have a most astonishing vitality. We have known cases where it has been pulled up for fifteen years, not a plant allowed to go to seed, and yet it makes its appearance every spring. Fortunately, it grows only on rich land, and is therefore found in evidence largely in hog yards and feed lots and other places where the land is exceedingly rich. In fact, in sections of the country where it is being introduced, it may be found in almost every farmyard. Why farmers allow it to mature seed passes our comprehension. It goes variously by the DESCRIPTIVE MANUAL 217 Fig. 126-C. Pigweed, Foxtail, and Velvet Leaf, in a potato patch. Too many such garden patches in Iowa. (Photographed by Pammel.) zoe Ve =e mAs = SEER Fig. 126-B. Distribution of Indian Mallow. 218 WEED FLORA 'OF IOWA names of velvet weed, butter print and in the locality of one of our farms, Davis weed, from the fact that it was introduced many years ago by a man named Davis, who regarded it as a rather good orna- mental plant.’’ Sida (Sida spinosa L.). Description.—An annual from 10-20 in. high; frequently much- branched; leaves ovate-lanceolate; serrate with a long petiole, pe- duncles in axils of leaves, 1-flowered; flowers small, yellow; 5 car- pels, each 2-beaked. Distribution—Common in the southern states, as far north as Massachusetts to southern Iowa and Kansas. Extermination.—This weed propagates entirely by its seeds which retain their vitality for a considerable length of time, as the seed coat is hard. The growing plant is, however, easily destroyed by pulling the weed or by cultivation. Fig. 127. Sida (Sida spinosa). Common in fields in southern Iowa. (Photographed by Hart.) DHSCRIPTIVH MANUAL 219 Fig. 127-A. Distribution of Sida. Cheeses or Common Mallow (M alva rotundifolia L.). Description.—A procumbent biennial ; leaves round, heart-shaped on long petioles, crenate ; flowers white, petals longer than the calyx ; a 3-leaved involucre at the base of the calyx; carpels pubescent. Distribution.—A widely distributed weed in eastern North Amer- ica, native to Europe, common in eastern and central Iowa in door- yards, barn lots, etc., in Story, Marshall, Polk, Marion, Linn, Clin- ton, Winneshiek and Allamakee counties. Extermination.—This weed is easily exterminated by cultivation. Do not permit any of the seeds to mature. The seeds retain their vitality for a considerable length of time. 220 WEED FLORA OF IOWA Fig. 127-B. Common Mallow or Cheeses (Malva rotundifolia). Common in door- yards, barn lots, ete. (Mich. Agr. Exp. Sta.) DESCRIPTIVE MANUAL 221 Fig. 127-C. Distribution of Common Mallow. Shoo-fly (Hibiscus trionum L.). Description—A low, rather hairy annual from 1-2 ft. high ; upper leaves 3-parted with 3 lanceolate divisions, the middle longest ; calyx inflated in fruit, membranous, 5-winged, with numerous dark nerves ; flowers sulphur-yellow with a blackish eye, ephemeral. Distribution —Common in fields and waste grounds in the south- ern states and eastward ; abundant in some places in Iowa, especially in gardens where it has become naturalized from cultivation. Extermination.—A. growing plant is not difficult to exterminate. A thorough cultivation and exposing the roots to the sun will de- stroy the plant. The seeds, however, retain their vitality for a con- siderable length of time. A correspondent of southeastern Iowa stated that this weed kept coming up in spite of constant and thor- ough cultivation. This was owing to the prolonged vitality of the seed. HYPERICACEHAH, ST. JOHN’S-WORT FAMILY. This small family contains a few ornamental plants with yellow flowers, commonly found in northern states. 222 WEED FLORA OF IOWA Fig. 127-D. Bladder Ketmia or Shoo-fly, (Hibiscus trionum). Gardens and corn fields. Large white flower with a dark spot at the base of each petal. a and b trichomes or plant hairs. (a and b drawn by Charlotte M. King. The general aspect photographed by Colburn. ) Fig. 127-E. Distribution of Shoo-fly. DESCRIPTIVE MANUAL 223 St. John’s-wort (Hypericum perforatum L.). Description.—A branched perennial, 114-2 ft. high with runners; leaves elliptical, or linear-oblong, with pellucid dots; flowers numerous in cymes, petals deep yellow, black-dotted, twice the length of the lanceolate sepals. Distribution.—Common in eastern North America in clay soils. Abundant in Jowa only in eastern counties, especially northeast- ward. Extermination—This weed spreads both by seeds and by runners. It is difficult to exterminate. Clark and Fletcher give the follow- ing methods: ‘‘Close cutting several times during the summer will reduce it in pastures. An application of salt—a small handful to Fig. 128. St. John’s-wort (Hypericum perforatwm). Old fields and woodland i pastures, eastern and northeastern Iowa. (Photographed by Quade.) 224 WEED FLORA OF IOWA y TRS Fig. 128-A. Distribution of St. John's-wort. each plant after close cutting in hot dry weather—will kill it and may be practicable where the pest is not abundant and the land cannot be brought under cultivation. Prevent it from going to seed. St. John’s-wort is easily suppressed on land that cart be cultivated under a systematic rotation of crops. Where it is established, it would be well not to seed to grass until it is suppressed. ’’ ONAGRACEAE, EVENING PRIMROSE FAMILY. A small family, some plants with showy flowers, a few cultivated for ornamental purposes. Evening Primrose (Oenothera biennis L.). Description.—aA stout, erect, pubescent or hirsute perennial, 3-5 feet high, sparingly branched; leaves lanceolate, or rarely ovate- lanceolate, denticulate, acute, bracts shorter or as long as the cap- sule; flowers yellow, petals obovate, stigma lobes linear, capsule sub- cylindrical; seeds small, brownish. Distribution—Common everywhere in eastern North America, Rocky mountains and Utah. Occurs in every county in Iowa. Extermination.Spreads by seed. This plant is not difficult to exterminate. Cut off the young plants a few inches below the sur- face of the ground. UMBELLIFERAE, CARROT FAMILY. Carrot, celery, parsnips and caraway are members of this family. Kt includes also many poisonous plants. DESCRIPTIVE MANUAL 225 Tig. 129. Evening Primrose (Oenothera biennis). Fences, gardens, meadows and pastures. (Photographed by Hart. a, drawn by Charlotte M. King.) \e tchell 2 . Fig. 129-A. Distribution of Evening Primrose. 15 226 WEED FLORA OF IOWA Caraway (Carum carvi L.). Description—A smooth, erect, slender herb 1-214 feet high with fusiform roots; leaves pinnate with filiform divisions; flowers in umbels, white; calyx teeth small; fruit ovate, or oblong with fili- form ribs. Distribution—Common in eastern North America, the Rocky mountains, Utah and scattered in places in Iowa. Extermination.—Easily exterminated; cut off the plants below the surface of the ground. Fig. 130. Caraway (Carum carvi). Commonly escaped from gardens. Flowers white. (Photographed by Quade.) DESCRIPTIVE MANUAL 227 Fig. 130-A. Distribution of Caraway. Wild Carrot (Daucus carota L.). Description.—A bristly, hirsute biennial from 2-214 feet high; leaves pinnately decompound; involucral bracts foliaceous; flow- Fig. 131. Wild Carrot (Daucus carota). Common in clover meadows. Flowers in white umbels. (Photographed by Colburn.) 228 WEED FLORA OF IOWA Fig. 131-A. Distribution of Wild Carrot. ers in compound umbels, white, or occasionally pink; fruit oblong, flattened dorsally, carpel with 5 slender bristly primary ribs, and 4 winged secondary ones, each of these bearing a single row of barbed prickles. canna ; 0 pean ' Distribution—Common in eastern North America, especially in dry fields. Becoming common in Iowa clover fields in Scott, Story, Polk, Franklin, Linn, Clinton, Cerro Gordo, Boone, Webster, Sac and Clay counties. Extermination.—Plant clean seed in a clean field. Easily killed by cutting off the plant a few inches below the surface of the eround. Parsnip (Pastinaca sativa L.). Description.—A tall, stout, glabrous biennial with grooved stem; leaves pinnately compound, cut-toothed; flowers yellow, small; calyx teeth obsolete; fruit oval, flattened dorsally, the lateral ribs with broad wings. Distribution—Common in eastern North America, Rocky moun- tains and the Pacific coast.. Common on roadsides in every part of Towa. Extermination.—Propagated by seeds. Easily exterminated by cultivation ; cut off the young plants a few inches below the surface of the ground. DESCRIPTIVE MANUAL ; 229 Fig. 132. Wild Parsnip (Pastinaca sativa). Common along roads. (Mich. Agr, Exp. Sta.) 230 WEED FLORA OF IOWA Fig. 132-A. Distribution of Wild Parsnip. Cowbane (Cicuta maculata L.). Description.—A smooth, marsh perennial, 2-5 ft. high, with pin- nately compound leaves, 2 or 3 times pinnate; leaves with long petioles; coarsely serrate leaflets lanceolate to oblong-lanceolate, 1-5 in. long; stalks of umbellets numerous and unequal; flowers white; fruit broadly ovate to oval, small, 1144 in. long. The plant grows in marshes and in low grounds, the stems springing from thick fleshy underground roots tapering at the lower end, usually numbering from 3-8 although single specimens are also met with. On cutting the roots there is given off a sharp pungent odor, which becomes intensified on boiling. Distribution.—Common throughout the northern states, south- west to Louisiana, Rocky mountains and Utah. Very common in low grounds and swales in northern Jowa, less common in southern Towa. Extermination—The field where cowbane occurs needs drain- age; plow the field, break.up the sod, and expose the fascicled roots to the sun. This will soon destroy the weed. DESCRIPTIVE MANUAL 231 Wig. 133. Cowbane (Cicuta maculata). In low moist meadows and roadsides. (Photographed by Colburn.) Fig. 133-A. Distribution of Cowbane. 23 Fig. WEED FLORA OF IOWA 133-B. Cowbane (Cicuta maculata). (Photographed by Caughey.) In low woods. DESCRIPTIVE MANUAL 233 Fig. 133-C. Cowbane roots (Cicuta maculata). In low places. - (Photographed by Gardner.) 234 WEED FLORA OF IOWA Cow Parsnip (Heracleum lanatum Michx.). Description.—A stout, hairy, pubescent perennial 4-8 ft. high; leaflets broad and large, irregularly cut-toothed; flowers white, in broad umbels. ig. 134. Cow Parsnip (Heraclewm lanatum). Common in woodland pastures. . (Photographed by Charlotte M. King.) Distribution—From the Atlantic coast, Newfoundland, through the northern states and Allegheny mountains to California. mon in the Rocky mountains. Com- APOCYNACEAE, DOGBANE FAMILY. In the tropics several species are important rubber producing ‘ plants. They are commonly called milk-weeds in Iowa. DESCRIPTIVE MANUAL 235 Spreading Dogbane (Apocynum androsaemifolium L.). Description.—Root-stock horizontal, smooth, or rarely soft-tomen- tose, branched above, spreading, leaves ovate, petioled, cymes loose, . spreading, both terminal and axillary; the latter pale rose color, open, bell-shaped; calyx segments shorter than the tubes of the corolla. Distribution—Common along borders of thickets from eastern Canada to British Columbia to Arizona and Georgia. Abundant in Iowa in grain fields near thickets. Fig. 135. Distribution of Spreading Dogbane (Apocynum androsaemifolium). Extermination.—Spreading dogbhane produces long, creeping roots, which are quite tenacious of life. In order to destroy the weed, the field should be given a shallow plowing after the grain is harvested, followed by a disking in a week or ten days, depending on the character of the weather. This should be followed by a harrow. If the fall is dry dragging will probably get the larger number of these weeds. Indian Hemp (Apocynum cannabinum L.). Description —Glabrous or more or less softly pubescent; 2-3 ft. high, smooth, terminated by an erect, close, many-flowered cyme; corolla lobes nearly erect, the tube not longer than the lanceolate segments of the calyx, greenish white; appears in July and August. Distribution.—Common species eastward and troublesome as a weed in northern Mississippi valley. Common in small-grain fields and pastures. 236 WEED FLORA OF IOWA Figure 136 Figure 1364 Fig. 136. Indian Hemp (Apocynum cannabinum). Fig. 136-A. Apocynum cannabinum hypericifolium. A low growing variety of the above, but with leaves broader at base and more abruptly pointed at apex. (Schuyler Mathews in Mich. Agr. Exp. Sta. Bull.) DESCRIPTIVE MANUAL 237 136-B. Distribution of Indian Hemp. Extermination—This weed should be treated like spreading dogbane. ASCLEPIADACEAE, MILKWEED FAMILY. A few plants, only, are of economic importance. Some are cul- tivated for ornamental purposes. In Jowa they are commonly ealled milkweeds. Showy Milkweed (Asclepias speciosa Torr.). Description.—A perennial 1-4 ft. high, white-tomentose or ca- nescent; leaves thick, broadly ovate or oval, petioled; pedicel elabrate above; flowers borne in dense umbels or rarely solitary, the pedicels stout; corolla purplish green, large, follicle erect or spreading on the recurved pedicels. Distribution—Showy milkweed is common from Minnesota to southern Iowa, Kansas, the Rocky mountains and Utah. In Iowa it is abundant in Emmet, Palo Alto and Dickinson counties. Extermination.—This weed should be treated like common milk- weed. 238 WEED FLORA OF IOWA Fig. 137. Showy Milkweed (Asclepias speciosa). In grain fields, meadows and roadsides, northern Iowa. (Photographed by Colburn.) Fig. 137-A. Distribution of Showy Milkweed. DESCRIPTIVE MANUAL 239 Milkweed (Asclepias syriaca L.). Description—A perennial herb with a stout stalk 2-5 ft. high, finely, softly pubescent or tomentose; leaves oblong, oval or ovate, obtuse or roundish at the base, the young leaf somewhat pubescent above, soon becoming glabrate; petioles stout, flowers from a few to many, borne in umbels; peduncles pubescent or tomentose; co- rolla dull purple or greenish purple, occasionally pale in color; fruit a follicle and borne on erect pedicels; trichomes multicellular from a single cell, somewhat floccose. Distribution —This milkweed is common from New England to North Carolina and Kansas. In Iowa it is abundant in oat fields, on highways and in gardens throughout the state. Fig. 1838. Common Milkweed (Asclepias syriaca). Fields, meadows, roadsides, ete. a trichomes or plant hairs. (Drawing by Charlotte M. King; general aspect photographed by Colburn.) 240 WEED FLORA OF IOWA jaa INaeeal 4 \ +, z eles [ee MaeMN. | + ‘ Kossuth » ee * fore hE a ae || tae : . . . ens . 5 . beni) OSE ee Fa serey antl reene| ore ® a + ss Sea ies 7 | ? FE L ‘ ‘ \oaes| Morbes Be ciali, | elas ye ° > aR OE LY ezell| T>0ni| So Wesabe e: hea | : {eel S SLE | Salaam Fig. 138-A. Distribution of Common Milkweed. Extermination.—Both milkweeds have the same habit of growth. The weed is perhaps known by its long roots which are frequently 10-15 ft. in length; another important point is that it produces ad- ventitious buds at frequent intervals from which new shoots arise. Fig. 138-B. Common Milkweed (Asclepias syriaca). Grain fields, waste places. (Photographed by Charlotte M. King.) DESCRIPTIVE .MANUAL 241 Wallace’s Farmer says concerning this weed: ‘‘We would ad- vise our correspondent to plow this field as scon as_ possible and prepare his seed bed for winter wheat. By plowing it again next August he will undoubtedly weaken the stand. He will fail, however, unless in working his corn he uses surface cultiva- tion. These weeds have no doubt been distributed through the field during the three years it was in corn by using a shovel cul- tivator, which takes up the roots and carries them over the field in the same way that many northern farmers are now seeding their fields with quack grass and damaging them to the extent of from five to twenty dollars per acre. By giving these two thorough August plowings and taking care of the roots that may be thrown up, then preparing the seed bed very thoroughly for corn and giving it as far as practical surface cultivation, he will probably get rid of these noxious weeds.’’ Climbing Milkweed (Gonolobus laevis Michx.). Description.—A climbing perennial; leaves oblong, cordate with a deep, narrow sinus; flowers borne in axillary umbel-like cymes, 5-10 flowered, large greenish flowers; calyx 5-parted; corolla 5- parted, wheel-shaped, the lobes narrowly linear-lanceolate, obtuse, larger than the calyx; anthers horizontal under the flattened stigmas ; pollen masses 5 pairs, follicles with soft warty projections. Distribution.—Troublesome in woods and fields in the southern states. It is reported as troublesome from a few counties in south- ern Iowa. Extermination.—This perennial weed is as difficult to destroy as common milkweed. Give thorough cultivation. If this will not suf- fice get the field into a meadow. CONVOLVULACEAE, MORNING-GLORY FAMILY. Few economic plants. Sweet potato and cultivated morning glory are representatives. 16 242 Pig. WEED FLORA OF IOWA 139. Climbing Milkweed (Gonolobus laevis). southern Iowa. (Drawing by Ada Hayden.) Troublesome in fields in DESCRIPTIVE MANUAL 243 rere] S y Bren iS Masset a N yi ES \ Baer \ ae h Wy phim oy 1] i S x ee AC IN {| Fig. 139-A. Distribution of Climbing Milkweed. Morning-glory (Ipomoea hederacea Jacq.). Description— Stems retrorsely hairy; leaves heart-shaped, 3-lobed, the lobes usually acute; peduncle variable in length; 1-3 Fig. 140. Blue Field Morning-glory (Ipomoea hederacea). Fields, common southern Iowa and Missouri. (Photographed by Colburn.) 244 WEED FLORA OF IOWA FSI DSRS EPR Fig. 140-A. Distribution of Morning-glory. flowers; calyx densely hairy below; corolla funnel-form, white and purple or pale blue; lobes of stigma and cells 3. Fig. 141. Annual Morning-glory (Ipomoea purpurea). A frequent escape from gardens into fields, (Photographed by Colburn.) DESCRIPTIVE MANUAL 245» Fig. 141-A. Distribution of Annual Morning-glory. The I. purpurea has heart-shaped leaves with retrorsely hairy stem; peduncles long, umbellately 3-5 flowered, purple to white. Distribution.—Common in waste and cultivated grounds from: New England southwestward; abundant from Missouri southward; native from tropical America; rare in Iowa. The J. purpurea is. commonly cultivated and is a frequent escape from cultivation in. Hardin, Story, Marshall, Wapello and Marion counties. Extermination—It is an annual and is easily destroyed; how- ever, the seeds retain their vitality for some time, frequently springing up for several seasons after the most thorough ecultiva- tion. ) Hedge Bindweed or Morning-glory (Convolvulus sepium L.). Description—Smooth, occasionally, however, pubescent, twining around supports or trailing; leaves triangular, halberd or arrow shaped, the tip acute, or pointed, the basal lobes obliquely trun- cate or sinuate lobed; flowering peduncles 4-angled with 2 leaf-like bracts which are commonly acute; corolla white or tinged with rose purple. DistributionHedge bindweed or morning-glory is common in the northern states and in the southwest from Texas to Canada, also in the Great Basin country. A form of it is also found in Europe and Asia. Common in every section of Iowa not only in corn fields, but along highways and in small-grain fields. 246 WEED FLORA OF IOWA Fig. 142. Morning-glory (Convolvulus sepiwm). Common in grain, corn fields and meadows. (Photographed by Colburn.) Fig. 142-A. Distribution of Hedge Bindweed or Morning-glory. DESCRIPTIVE MANUAL 247 Extermination —The morning-glory must be treated like horse nettle since it is a perennial. In addition to the usual methods of cultivation sheep have been recommended to destroy the weed. This method is certainly applicable where the weed occurs in pas- tures. In Wallace’s Farmer, Mr. L. C. Greene’s experience in killing morning-glory is given as follows: ‘‘A large farmer had 145 acres of corn. One piece of twenty acres, fall plowed, on a south slope, was planted to corn the first of May, and by the time the plowing and planting were all done it was near the last of May. The early planted field was thick with-morning-glories and had received no cultivation since they commenced to grow. By the time the corn was four inches high the morning-glories were eight or more inches tall, growing in mats on the ground hunting for something to climb upon. The proprietor viewed the field, and instead of sending out the cultivator sent out three stirring plows and the planter soon followed. In two days the field was plowed and planted again and a fine crop of corn was raised with very little bother from the vines, and even the following year the vines bothered but little. ‘‘Some years ago I fall plowed a small field that was badly in- fected with morning-glory vines and smartweeds. The 24th of the next May I was ready for that field, but from a little distance it looked as if a mowing machine and a rake would be the proper tools to use. I plowed rather deep to do a good job, the planter im- mediately followed, and in four days after the planter some corn could be seen, and it was eight inches high when the cultivator got to it. It was just a matter of stirring the soil all season, for there were no large weeds to kill.’’ The Prairie Farmer makes these suggestions concerning the eradication of morning-glory: ‘‘ Another way to fight the morning- glory is to grow two or three pasture crops a year on the land for sheep. One of them ought to be a cultivated crop. The morning- elory would not be able to hold out long against such treatment. The strong point in favor of this method is the profitable character of the work.’’ Wallace’s Farmer says concerning its destruction: “‘They do not spread rapidly except under cultivation, as they grow mostly from the roots, and these are distributed over the fields: by cultivators. If when the farmer first discovers a patch out of cultivation for a year or two he plows it shallow and frequently 248 WEED FLORA OF IOWA and harrows he can get rid of them. If he cultivates the plants with the rest of his field it is only a short’ time until he will find these weeds scattered all over his field. A great many ways have been suggested to get rid of this troublesome weed. Special at- tachments have been invented for the use on corn cultivators known as the morning-glory blades. These are designed to shave off the plants just below the surface of the ground.”’ ‘“We certainly would fence up this pasture, or part of it, and would sow a mixture of grains that would furnish hog feed, and LA while we were at it would sow clover and timothy, and when the grains were three or four inches high turn in the hogs. The only trouble is that there are not enough hogs to go around the whole tract. We had a field in that condition twenty years ago. We made a hog pasture of it, and while the morning-glories are yet to be seen in the road alongside, there are none of them in the f , pasture, and have not been since the first year.”’ European Bindweed or Morning-glory (Convolvulus arvensis L.) Description.—The European bindweed or morning-glory is a deep-rooting perennial; stem procumbent, twining or creeping. Like the horse nettle, this species propagates freely by under- ground root-stocks; leaves 1-2 in. long, ovate, oblong, arrow-shaped, lobes at the base running to a point; flowers borne in 1-flowered peduncles with very small leaf-like bracts some distance from the flowers; flowers an inch or less long, short, broadly funnel-shaped, white or commonly of a rose tinge. Distribution.—European bindweed is a troublesome weed in Eur- ope and in eastern North America. It occurs also in the southern states and on the Pacific coast. It is scattered in many parts of Towa in small patches. Extermination.—A short rotation of crops should be practiced, including late sown roots or other cultivated crops: rape is useful for this purpose. Frequent use of a broad-shared cultivator will destroy new growths and exhaust the vitality of the plants. Sow no crop seeds containing those of field bindweed. Applications of salt or lime, sometimes recommended to kill this weed, are useless unless appled in large quantities. SS DESCRIPTIVE MANUAL 249 ra mae > a \ Fig. 143. European Morning-glory or Bindweed (Convolvulus arvensis). Com- mon in gardens and becoming frequent in Iowa. (Photographed by Colburn.) Fig. 143-A. Distribution of European Morning-glory. 250 WEED FLORA OF IOWA Prof. H. A. Hitchcock in Farmer’s Review, says: ‘‘The black bind-weed or perennial morning-glory (Convolvulus arvensis), which I suppose is the kind meant, is a great pest and difficult to eradicate. If a patch is not too large, heavy mulching is the best way to destroy it. Then watch the patch closely and eut off im- mediately any stray shoot that may appear above its surface. Nothing but persistent watching and the careful cutting off of all parts above the ground will eradicate this weed.’’ Prof. Ten Eyck, quoted in Wallaces’ Farmer, states that the only method of culture applicable to large areas which promises any great degree of control or destruction of the pest is very late fall or winter plowing. The plots which were plowed in November (no plowing was done later than November 20th) showed a very scat- tering and feeble growth of bindweed on April 26th, the date of inspection. The weeds were thinner and more feeble also on the unplowed land which produced a crop of sowed cane or sowed kaffir last season, than they were on any of the lots cultivated in intertilled crops. Clover Dodder (Cuscuta epithymum Murr.). Description.—A spreading, climbing plant; yellowish or reddish with a few minute scales in place of leaves; flowers whitish or pink- ish in heads, small, globular, urn-shaped; cylindrical tube longer than the nearly erect, acute sepals; scales large-toothed; stigma elongated; style longer than the ovary; stamens exserted, fruit in capsules. Distribution—This weed has long been known as troublesome in Europe and has been more or less common in the Rocky moun- tains on clover and alfalfa. For some years also frequent in the east; becoming more abundant on clover and alfalfa. Extermination—Where the plant occurs cut down the clover at once and burn. Sow the patch or field to another crop, preferably to small grain or with corn. Clark and Fletcher recommend the following treatment: ‘‘As soon as the pest is noticed, the infected patches should be at once mown with a scythe and the refuse removed and destroyed. Fields DESCRIPTIVE MANUAL 251 Fig. 144. Clover Dodder (Cuscuta epithymum). . S49 dt Ea eo ; ; Kossuth = ~ state C © OBrien ; : Sees 50 ° ary eee 0, 2 ° “4 ely * fe 0 + ° . . i) +H * |e s + 04 Loy AUS; SAIRSSSI o wll *,\ 728k | * PS s L sige ly A. Solo lol eae 5 ‘ SG * qubuque + "af e Ma, a ae a . . t 2. : e OOO 2g rene Pies ° : 4 Gt foe fl oP te | AP Remedy oe | 6 iy [ e| ¢ irs . > > 2 = = . © Clinton + + he ° . ° ry ° onsen ° te s ° 0 © \oces| Motes © e? . 5 Tataranine| ‘ ave : “|eokuk | 5 wattdrne) 7 ¢ * ‘ee e ‘ : 5 5 . ° . A A . 5 ° , . ° 0 Fig. 153-A. Distribution of Hoary Vervain. a, drawn by Charlotte M. King.) 266 WEED FLORA OF IOWA AA SESS sani Fig. 153-B. White Vervain, Nettle-leaved Vervain (Verbena urticaefolia). (Mich. Agr. Exp. Sta.) DESCRIPTIVE MANUAL 267. taba Ee. Cue hey Fig. 153-C. Distribution of White Vervain. Common Vervain (Verbena bracteosa Michx.). Description.—A widely spreading, hairy annual; leaves wedge- lanceolate, cut-pinnatifid or sometimes 3-cleft; flowers in spikes, i ‘ oe d ‘ : 7 2 os EES ee i Fig. 154. Prostrate Vervain (Verbena bracteosa). a, Plant hair or trichome. Common along roadsides, streets, and gravelly places; small, blue flowers, — (General aspect photographed by Quade. a drawn by Charlotte M. King.) 268 WEED FLORA OF IOWA Basse et Hit bets Bee oe: ee GRE PSSese AREER C/A are: Fig. 154-A. Distribution of Prostrate Vervain. with large bracts, small, purple; trichomes few-celled from a broad several-celled base. Distribution.Common in waste places, roadsides, walks, gravel- ly and sandy fields from Virginia to Wisconsin and Minnesota and southward. Eaxtermination.—It is easily exterminated by cultivation. LABIATAE, MINT FAMILY. The common pepperment, scarlet sage, catnip, pennyroyal, thyme and basil belong to this family. All are aromatic plants. Mint (Mentha arvensis L.).. Description.—This perennial weed has freely branching stems 1-114 ft. high, retrorsely pubescent, leaves oblong to ovate, rounded at the base, minutely pubescent, closely serrate, petioled or nearly sessile; flowers white, pink or violet. The varity canadensis has lanceolate to oblong-lanceolate, pubescent leaves and is the common form in Iowa in low grounds. Several other species occur in Iowa, namely: peppermint (Mentha piperita) along brooks, a smooth and pungent-tasting herb with ovate-oblong leaves and running root-stocks, and spearmint (Mentha spicata) with oblong or ovate- lanceolate unequally-serrate leaves. Distribution.—This Eurasian species occurs from Newfoundland to Nebraska and the Pacific coast. Extermination.—Easily exterminated by giving thorough culti- vation and dragging the soil to bring the root-stocks to the surface of the ground. DESCRIPTIVE MANUAL - 269 Fig. 155. Mint (Mentha arvensis var. canadensis). Common in low meadows; whitish flowers. Plant with odor of peppermint. (Photographed by Colburn.) Fig. 155-A. Distribution of Mint. 270 WEED FLORA OF IOWA Fig. 155-B. Peppermint (Mentha piperita). In some gardens, (Photographed by Colburn.) Carpenter Weed, Self-heal (Prunella vulgaris L.). Description.—A low, perennial weed with ovate-oblong, entire or toothed leaves, hairy or smooth; flowers collected in heads of 3-flowered clusters, corolla violet or flesh-color, or rarely pale in color, longer than the purplish calyx which is tubular bell-shaped. Distribution—Widely distributed in northern United States, west across the continent in clay soils and woods, also from New- foundland to Florida. Extermination—This weed is not difficult to exterminate by giving cultivation with a cultivator and hoe. The weed is mainly spread by seed. The seed of this is not uncommon in clover seed. DESCRIPTIVE MANUAL 271 Fig. 156. Self-heal (Prunella vulgaris). Common along roadsides, wooded pastures. (Photographed by Quade.) Fig. 156-A. Distribution of Self-heal. ae id 272 WEED FLORA OF IOWA American Germander (Teucrium canadense L.). Description —A_ perennial, downy, erect herb 1-3 ft. high with running root-stocks; leaves pubescent, short-petioled, downy be- neath, ovate, lanceolate, serrate with a rounded base; floral leaves small; flowers in ample wandlike spikes; calyx 5-toothed, the up- per lobes obtuse; corolla purple, rose or whitish. Distribution—From New England to Mexico and northwest to Manitoba. Common in Iowa alluvial grounds in pastures, mead- ows, grain fields, ete. Extermination.—A very troublesome weed in the north. It is, however, an excellent bee-plant. Should have the same treatment as quack grass. The root-stocks should be exposed and allowed to dry. After plowing the field follow with a dise and harrow once a week after the small grain crop is removed. Fig. 157. Germander or Wood Sage (Teuchriwmn canadense). Common in fields and woods. (Photographed by Colburn.) DESCRIPTIVE MANUAL 273 Fig. 157-A. Distribution of Germander. L. H. Pammel in the Weekly Register says: ‘‘Germander is a troublesome weed found in many parts of northern Iowa. It pro- duces root-stock very much like mint and quack grass. Each sev- ered portion produces a new plant, and for this reason it is some- what difficult to destroy. The only way to exterminate this weed is by thorough cultivation. Plowing in the fall during the dry season, then plowing again in the spring and giving thorough cul- tivation during the growing season should destroy the weed with- out difficulty.’’ Catnip (Nepeta cataria L.). Description.—A perennial, erect herb, 1-3 ft. high; leaves ovate, cordate, coarsely serrate, petiolate, whitish, downy underneath; flowers in cymose clusters; corolla whitish, dotted with purple; trichomes several-celled, rough, thick-walled. Distribution—Native to Europe; widely naturalized in the northern states. Extermination.—Give the same treatment as to motherwort. Chemical Composition.—According to the University of Min- nesota*, it is as follows :— k Nitrogen free Dry Matter Crude Protein Ether Extract Extract and Ash Fiber 94.30 22.25 | 2.66 | 63.07 12.77 *Snyder: Bull. Univ. Minn. Agr. Exp., 101. 18 274 WEED FIORA OF IOWA Fig. 158. Catnip (Nepeta cataria). Trichome or plant hair at the right. Com- mon in waste places. (Photographed by Colburn. Drawing of hair by Charlotte M. King.) + Jared \ . 4 AUS Figure 158A Figure 158B Fig. 158-A. Plant hair or trichome from leaf of catnip. (Drawing by Charlotte M. King.) Fig. 158-B. Distribution of Catnip. DESCRIPTIVE MANUAL 275 Ground Ivy (Nepeta hederacea (L.) Trevisan). Description.—A creeping, trailing perennial, with leaves all alike, petioled, round, kidney-shaped; crenate, smooth, green on both sides; flowers light blue in axillary whorls of about 6, ap- pearing in early spring and summer. Distribution.—Native to Europe, widely naturalized in the northern states, especially in shady places. Common everywhere in Jowa. Extermination—Somewhat difficult to exterminate in lawns, but thorough cultivation will destroy the weed. Fig. 159. Ground Ivy or Creeping Charley (Nepeta hederacea). Common in some gardens. (Photographed by Colburn.) 276 WEED FLORA OF IOWA eee si Fig. 159-A. Distribution of Ground Ivy. Motherwort (Leonurus cardiaca L.). Description.—Tall perennial herb with erect stem, 2-6 ft. high; leaves long-pointed, the lower round and palmately lobed, the up- ig 160. Motherwort (Leonurus cardiaca). A common weed in waste places and gardens. (Photographed by Colburn.) DESCRIPTIVE MANUAL 277 eee Fig. 160-A. Distribution of Motherwort. per crenate at the base, 3-cleft; flowers pale purple, in close axil- lary whorls; corolla bearded. Distribution.—Native to Europe, widely naturalized in the north- ern States. Extermmation.—This perennial weed can be exterminated by plowing the field with frequent cultivation subsequently. Blue Sage (Salvia lanceaefolia Poir.). Description—A_ slightly pubescent or nearly smooth annual, 1-214 ft. high; leaves petiolate, lanceolate serrate or nearly entire; flowers in interrupted, erect spikelike clusters; calyx bilabiate, upper entire, the lower 2-cleft; corolla, blue, slightly exserted; stamens with long connective, attached to a sterile anther which blocks the throat of the corolla; pistil 1, deeply 4-lobed. Plant related to the cultivated garden salvias. Distribution.—Chiefly west of Missouri river, Kansas, Nebraska, Texas, and Arizona, Missouri and Indiana, introduced in Ohio. Common in Pottawattamie, Fremont, Monona and Woodbury coun- ties, less common in Story, Boone, Polk and Muscatine counties. Extermination—tThis weed is easily exterminated by cultivation. Do not allow the plant to produce seeds. SOLANACEAE, NIGHTSHADE FAMILY. Potato, tomato, tobacco, ground cherry, jimson weed, flowering tobacco and black henbane, belong to this family. 278 WEED FLORA OF IOWA Fic. 161. Lance-leaved Salvia, Blue Sage (Salvia lanceaefohe). DESCRIPTIVE MANUAL 279 Wig. 161-A. Distribution of Lance-leaved Salvia. Ground Cherry (Physalis lanceolata Michx.). Description —aA hirsute perennial with short, stiff hairs, some- times nearly smooth; forms short and stout underground stems; leaves oblong-ovate to lanceolate, sparingly angulate-toothed or more often entire; flowers in axillary peduncles, calyx hirsute, co- rolla yellowish with a dark eye; berry reddish. The P. virginiana Mill., is also an erect perennial with narrowly ovate acutish leaves or acutish or rounded teeth, corolla pale yellow. The P. subglab- rata MacKenzie and Bush, has ovate or ovate-oblong leaves, oblique at the base, entire or repand, and brownish corolla; berry, large, reddish or purple. The P. heterophylla is perennial, leaves obtuse repand, or obtusely toothed; trichomes several-celled, glandular and non-glandular. Distribution—The P. lanceolata in southern Iowa and south- ward, the P. virginiana common everywhere in Iowa, in dry grav- elly soil, from Connecticut to Jowa and southward; the P. sub- glabrata from Rhode Island to Minnesota and southward. Extermination —All of the perennial ground cherries are diffi- cult to destroy because of the strong underground root-stocks which freely spread the weed. These plants are also scattered by seed. To exterminate the perennial weeds plow shallowly and expose the plants to the sun; give frequent cultivation. 280 WEED FLORA OF IOWA Wig. 162. Ground Cherry (Physalis lanceolata). Common in gravelly and sandy soils; waste places and gardens. 1, leaf; 2, flower. (Photographed by Quade. Drawing by Charlotte M. King.) Fig. 162A Figure 162B Fig. 162-A. Trichome or plant hair from leaf of Ground Cherry (Physalis heterophylla). (Drawing by Charlotte M. King.) Fig. 162-B. Distribution of Ground Cherry. DESCRIPTIVE MANUAL 281 Fig. 162-C. Distribution of Virginia Ground Cherry (Physalis virginiana). Common Nightshade or Stubbleberry (Solanum nigrum L.). Description.—Annual, low-branched and often spreading; glab-_ rous or hairy, hairs roughened on the angles; leaves ovate, petioled, Fig. 163. Black Nightshade (Solanum nigrum). Shady places, gardens and fields. (Photographed by Colburn.) 282 WEED FLORA OF. IOWA + 2a am 7 Sel S| é 83 ws ZN + Kossuth SZ OBrien i Wray tr). \ «, Me == Uy Yo biowe erty ie ie /da . = . en eeCne = . 4 : Ame: . o ey ere Cinte oruradow " Des) Mor 5 od ; Heohktih oe rate Bz e AN Y \ AS SN AWAY) Fig. 168-B. Black Nightshade (Solanum nigrum). DESCRIPTIVE MANUAL 283 flowers white in small, umbel-like, drooping, lateral clusters; calyx spreading, the lobes obtuse, much shorter than the white corolla; = glabrous, globose, black; occasionally large. - Distribution—Found in northern United States; abundant everywhere in Towa in shady grounds and fields. A cosmopolitan: weed. Horse Nettle (Solanwm carolinense L:). Description—aA deep-rooting perennial, propagating freely by its underground root-stocks, the running roots often being 3 ft. long; stem 1-2 ft. high, somewhat straggling, half shrubby at the base; stem hairy or merely roughish with minute hairs which are star-shaped, also armed with numerous stout, subulate, yellowish prickles; leaves oblong or sometimes ovate, obtusely sinuate-toothed or lobed, or deeply cut, 2-4 in. long; flowers borne in one-sided racemes; calyx consists of slender lobes, ¢orolla light blue or white, an inch or less in diameter ; flowers followed by yellow globose ber- ries 14-84 in. in diameter; small yellowish seeds, a little less than 1/,, in. long, minutely foudhened: Flowers and yellow berries resemble those of the potato. The spiny character of the leayes and the further resemblance of the flower to the potato should render it easy of, detection. Distribution.—Horse nettle is indigenous to thé southern states, but now oceurs from Connecticut to northern Iowa. This weed is most abundant in southern Iowa and has been reported from Floyd, Story, Boone and Linn counties; in the north half of the state, however, it is a recent introduction. ead ew ee aiinction —There are two ae ne ne ondedt Oi: one by seeds, the other by perennial roots. It is so exceedingly tenacious a weed, thatit is almost impossible to remove it when fully estab- lished. ‘The following very suggestive methods have ‘been given. Smothering. This is an effective method of removing the plant. For this purpose probably rape or sorghum is the most suitable crop. If the soil is not already rich, a liberal dressing of barnyard manure should be applied during the winter or spring. The soil should be harrowed or cultivated frequently until the time of seed- ing, which may be any time during the months of May or June. This cultivation will prevent the weed growth, and will also assist 284 WEED FLORA OF IOWA in the retention of moisture. If the rape is sown in drills about two pounds of seed per acre is sufficient quantity and three pounds if sown broadcast. When the crop has attained a rank growth it may be pastured or removed and fed to stock. Where land is lack- ing in vegetable matter it is good practice to plow the crop under when it is properly manured. The latter is not necessary when the object is to destroy the nettle, as the rank growth of the crop is very effective in completely smothering the weed. Hoed crops. Planting of corn or roots is a method much in ~ vogue for the destruction of this vile intruder. As in the previous method the plant should be kept down before seeding time. When the crop appears above the ground the use of horse and hand hoe should not be sparing. When the welfare of the crop prohibits the use of the horse hoe, the hand hoe should be used at intervals until the crop is removed; even after this, it is sometimes necessary to give attention to the pest. There is no question about this mode of treatment being effective if properly carried out, but often failure results from negligence during the latter part of the season. The Iowa Homestead suggests the following treatment: ‘‘Corn land that has grown up to horse nettles this year should be burned over, if possible, next spring, as this will destroy many of the seeds. Afterwards the land should be plowed lightly and kept cul- tivated at intervals until somewhat late in the season. A surface cultivator will be all that is necessary, and this need not be run deeper than two or three inches, just enough to effectively cut the plants off below the surface. By checking the growth several times before planting corn the root system becomes much weakened, so that ordinary cultivation the remaining part of the season will often keep them completely in check. Where nettles have been going to seed for a number of years it may require considerable time to free the land because these will germinate as they are brought near the surface by the various plowings. However, it should be kept in mind that any perennial root system may be killed outright in one season if it is not allowed to develop roots or stems.’’ Buffalo Bur (Solanum rostratum Dunal.). Description Herbaceous; woody when old, somewhat hoary or yellowish, 8 in.-2 ft. high, covered with copious stellate pubescence ; branches and main stems, when it begins to branch, covered with sharp yellow prickles; leaves somewhat melon-like, 1-3 times pin- Fig. 164. Horse Nettle (Solanum carolinense). sometimes extend 3-4 feet in the soil. the potato. DESCRIPTIVE MANUAL Common in southern Iowa. (Photographed by Colburn.) assuth | : S OBrren 1%. ° 2 . Her ° Tou Oy ubugue 57 loa 5 . 58 2 © reene oa © = mes =+ = . +> + ” < ° thie he onsen eel a? a id VF el aes} < . OM Ieee aS SS [reokuk |» ge a yatrawatidmie| / O é e hte 2 5 . : 5 + + : selene E ota Page °, \aecah ; - . we |. jecatur + sO Olorailn ) E 3 il | ° o 8 Fig. 164-A. Distribution of Horse Nettle. - 285 Deep rooted perennial, roots Flowers and berries somewhat like 286 WEED. FLORA OF IOWA Fig. 165. Buffalo Bur (Solanum .rostratum).. Pastures, gardens, railways, ete. ‘ (Photographed by Colburn.) : CRETE Fig. 165-A. Distribution of Buffalo Bur. DESCRIPTIVE MANUAL 287 aatifid, lobes roundish or obtuse and repand, covered with soft pubescence; hairs stellate, flowers yellow, corolla gamopetalous, about an inch in diameter, nearly regular, the sharp lobes of the corolla broadly ovate; stamens 5, declined, anthers tapering up- ward, linear-lanceolate, dissimilar, the lowest much larger and longer, with an incurved beak, hence the technical name rostratum ; style much declined; fruit a berry but enclosed by the close-fitting or prickly calyx, which has suggested the common name buffalo bur; pedicels in fruit erect; seeds thick, irregular, round or some- what longer than broad, wrinkled, showing numerous small pits and surrounded by a gelatinous substance. Fig. 165-B. Buffalo Bur (Solanum rostratum); a, general sketch showing habit of plant; b, flower; d, seed natural size; c, seed enlarged. (After Dewey, U. S. Dept. of Agr.) 288 WEED FLORA OF IOWA Distribution.—Buffalo bur is originally native of the plains region, between the Missouri river and the Rocky mountains. It has spread eastward in the northern states and extensively in ' Texas, Mississippi and Arkansas. It is not infrequent from Tennessee to New York. It has been found in Germany. ‘This weed is widely scattered in small patches in many parts of the state of Iowa. Extermination—Inasmuch as this weed is an annual, it can be easily exterminated by cutting off the young plants below the ground and this should be done before the pods are formed. If plants are older they should be cut off and burned. The Iowa Homestead says concerning this weed: ‘‘On account of the fact that the buffalo bur is an annual its destruction or eradication is simply a matter of preventing it from maturing its seed. Corn fields that are badly infested may need but little at- tention after the regular time for laying the corn by, for which purpose the one horse cultivator may be pressed -into use.’’ Dr. C. E. Bessey in Breeder’s Gazette recommends as follows: ‘“To get rid of it the best thing is first not to allow the weed to get a good start, as its deep roots are hard to get out. Second, if it has a good start, the plants must be cut down frequently so as to prevent their seeding and thus starting new plants. In the third place, the deep roots must be killed by digging out or by smother- ing. This can be done by using a very heavy dense crop like some of the sorghums, or by covering the patch with wet manure. Of course constant stirring of the soil will kill them. It will pay to watch this weed wherever it appears.’’ Purple Thorn-apple, Purple Stramonium, or Jimson Weed (Datura tatula L.). Description—A glabrous annual from a few inches to 5 feet high ; stem purplish ; leaves thin, ovate, acute or acuminate; flowers consisting of a 5-toothed calyx and a 5-lobed funnel form corolla, with stamens included; filiform filaments inserted below the mid- dle of the corolla tube; capsule globular, prickly, 4-valved and 2-celled. The common thorn-apple (Datura stramonium) is a glab- rous annual with green stem, sinuate-toothed leaves and white corolla. DESCRIPTIVE MANUAL 289 Fig. 166. Purple Jimson Weed (Datura tatula). Barnyards, roadsides. (Photographed by Colburn.) er ee ———— a Mitchell Kossuth OBrien SS utlor-| u ou Cy i Ida weene id 4 | dames- < —— = . . Q Des) Moines . Poitawalta mre) 7 * Page ature lima S Fig. 166-A. Distribution of Jimson Weed. 19 290 ; WEED FLORA OF IOWA Distribution.—Both species abundant in field and waste places from New England to North Dakota and Texas and naturalized from Europe; originally native to India. Fig. 166-B. Jimson Weed or Thorn-apple (Datura stramonium). a, flowering spray; b, fruiting capsule. (U. S. Dept. Agr.) RASS PACE th ny fis ml Sie val ee Fig. 166-C. Distribution of Thorn-apple. DESCRIPTIVE MANUAL 291 Extermination.—Both of the jimson weeds are easily destroyed by cultivation. They produce an enormous amount of seed which probably retains its vitality for a considerable length of time. How long, however, has not been determined. SCROPHULARIACEAH, MULLEIN FAMILY. Snapdragon and Simpson honey plant belong to this family. Mullein (Verbascum thapsus L.). Description.—A tall, densely woolly annual or biennial herb 2-6 ft. high; leaves oblong, thick, covered with branched hairs, the basal leaves margined, petioled; flowers in long dense spikes; corolla Fig. 167. Mullein (Verbascum thapsus). A hairy biennial, common road- side weed; gravel hills along Mississippi river, old fields. (Photographed by Colburn.) 292 WEED FLORA OF IOWA Seem oe Figure 167A Figure 167B Fig. 167-A. Glandular trichome from viscid pod of Moth Mullein (Verbascum blattaria). (Drawing L. H. Pammel and Charlotte M. King.) Fig. 167-B. Distribution of Mullein. rotate, yellow or rarely white; stamens unequal, the 3 upper shorter, woolly, with short anthers; the 2 lower smooth with larger anthers; trichomes many-celled, branched with central axis. Distribution—From Nova Scotia west across the continent. Southwest to Missouri and Kansas and Utah. Common in waste places, especially in eastern Iowa. Extermination.—Mullein is easily destroyed by cutting the plant off a few inches below the surface of the ground. This may be done in the autumn after the appearance of the root leaves, or in the second season when the plant shoots up. Toadflax (Linarva vulgaris Hill.). 4 Description.—Persistent, deep-rooted perennial, 114-214 ft. high, with erect, slender stem; leaves smooth, sessile, crowded, alternate- linear, somewhat fieshy ; flowers in racemes, showy, pale yellow and orange lips; corolla 2-lobed, closed; seeds small, dark brown to black and roughened; flowers from June to October. Distribution.—Introduced from Europe first as a cultivated plant from whence it has spread to roadsides, meadows, and waste places. Somewhat widely distributed in this state, but particularly common in Clayton, Allamakee and Winneshiek counties; local in Story county. DESCRIPTIVE MANUAL 293 Roadsides, gardens, Fig. 168.. Toadflax or Butter and Eggs (Linaria vulgaris). etc. ier (After Clark and Fletcher.) 294 WEED FLORA OF IOWA Fig. 168-A. Distribution of Toadflax. Extermination.—F letcher and Clark recommend as follows: ‘“‘Short rotation of crops with deep, thorough cultivation in spring and fall will suppress it. Hand-pulling when the soil is wet is effective in pasture lands that cannot be cultivated. Badly in- fested meadows or pasture lands should be brought under cultiva- tion by plowing in July, summer-fallowing until autumn, and planting with hoed crop the following spring.’’ Simpson Honey Plant (Scrophularia marilandica L.). Description—aA glabrous, somewhat glandular, pubescent peren- nial, 3-5 ft. high; stems 4-angled; leaves thin, ovate, or ovate-lan- ceolate, sharply serrate; flowers cymose; calyx lobes ovate, about the length of the tube, corolla brownish purple; capsule subglob- ular; seeds small, numerous. Distribution—Common in woods and thickets from Maine to the Rocky mountains. Abundant in woods and adjacent fields. Extermination.—Simpson honey plant produces a large number of small seeds. However, but little is known of their vitality.. The weed is easily killed by cultivation and easily crowded out by clover and small cereals. The Scrophularia leporella Bicknell, is similar as regards the foliage, the rudimentary stamen is, however, yellowish-green in- stead of brownish-purple as in the 8. marilandica. DESCRIPTIVE MANUAL 296 Fig. 169. Simpson Honey Plant (Scrophularia marilandica). waste places. (Photographed by Colburn.) Woods and Vwirchel eel | aL ( = “S C} \ IS Buller. aN = SINE ida J S yk i ¢ | Forrawattd mie Fig. 169-A. Distribution of Simpson Honey Plant. 296, WEED FLORA OF IOWA Purslane Speedwell, Neckweed (Veronica peregrina L.). Description.—A glabrous, glandular, or nearly smooth, branch- ing annual 4-9 in. high; leaves petioled, upper oblong, linear and entire; floral leaves like those of the stem but reduced; flowers axillary and solitary, white; capsule orbicular. Distribution—A common weed in fields in Iowa and in eastern North America from Nova Scotia southward; also west to Texas and the Pacific coast. Found in South America and in Europe; cos- mopolitan. ; Extermination.—The seeds are produced abundantly, but young as well as older plants are easily killed by cultivation. Vig. 170. Speedwell (Veronica peregrina). Common in gardens and fields in early spring. (Photographed by Quade.) PLANTAGINACEAE, PLANTAIN FAMILY. The family contains few plants of economic importance. The seeds of a few species are used as medicine. DESCRIPTIVE MANUAL - 297 Fig. 170-A. Distribution of Speedwell. Common Plantain (Plantago major L.). - Description. —A smooth, glabrous perennial with short root- stocks; leaves with a long channeled petiole, ovate, oblong or oval; spike long, linear, cylindrical, capsule circumscissile near the mid- dle; flowers proterogynous; seed smooth, angled, reticulated : tri- chomes short, several-celled, from a broad base.. - Distribution—Common plantain is widely distributed in North America from the Atlantic to the Pacific. Perhaps native far northward; probably naturalized in Iowa, It is found in every county in the state; frequent in dooryards, fields and pastures. Extermination.—Usually not a difficult weed to exterminate in cultivated fields. Clark and Fletcher recommend the following treatment: ‘‘Hoed crops every four years will keep this weed in ‘check. Working with a broad-shared cultivator, followed by a. harrow, to drag the plants with their fibrous roots to the surface, is recommended for spring cultivation. Plantain in lawns may be weeded out when the soil is firm by forcing a small implement like a chisel, with a half-round blade having a point like the tip of a spoon, between the soil and the fleshy crown of the weed to a depth sufficient ‘to break the plant away from its fibrous roots without disfiguring the turf. northwestern and western Iowa. (Photographed by Pammel.) Extermination —Correspondents sending this weed frequently refer it to Canada thistle. It occurs not only in pastures and mead- 376 WEED FLORA OF IOWA ows, but also in corn and grain fields. It grows in patches like the Canada thistle. These patches increase in size from year to year. It was described as a doubtful perennial by early botanical writers and so far as I have been able to determine, it is a perennial. It may be exterminated by thorough cultivation, plowing well and then following with the cultivator. Prairie Thistle (Cirsium discolor (Muhl.) Spreng.). Description.—Tall, branching, leafy biennial, 5-7 ft. high, with heads larger than in Canada thistle; stem striate, slightly hirsute; leaves radical, 12-14 in. long, deeply pinnatifid, the divisions fre- quently divided, prickly-toothed, the upper surface smoothish, and the lower white ; woolly single heads terminating the branches, with Fie, 221. Prairie or Wood Thistle (Cirsiwm discolor). Common border of woods, ete. (Photographed by Colburn.) DESCRIPTIVE MANUAL 377 Fig. 221A Figure 221B Fig, 221-A. Trichomes or plant hairs of wood thistle. Fic. 221-B. Distribution of Prairie Thistle. purple flowers; heads 1-11% in. long: bracts of the globose involucre somewhat suppressed, slightly arachnoid, lower bracts ovate with a broad base and a weak prickly recurved bristle, slight dorsal ‘gland, inner linear-lanceolate with a nearly colorless entire appendage; flowers purple, tube of the corolla 11-12 lines long, lobes of the corolla terminating in clavate tips; anther tips acute, filaments pubescent; bristles of pappus plumose; achene 22 lines long, ‘smooth, upper part yellow. ee Distribution—Common in many portions of Iowa; Marshall, Johnson, Winnebago, Lee, Winneshiek, Allamakee, Greene, and Emmet counties; at Keokuk, Muscatine, Ames, Cedar Rapids, Car- roll; Des Moines, Polk City, Steamboat Rock, Mason: City, Belle Plaine and Iowa City. Extermination.—This field thistle should be treated like all other biennial weeds. The flattened masses in the spring should be cut off below the ground and none of the plants allowed to go to seed. We have received numerous inquiries in regard to this weed from western and northwestern Lowa. Iowa Thistle (Curstwm towense (Pammel) Fernald). Description.—Biennial with downy, branching stem; leaves roughly hairy above but white-woolly beneath, oblong-ovate to nar- rowly lanceolate, sinuate-toothed, or somewhat pinnatifid, lobes or 378 WEED FLORA OF IOWA Fig. 222. Iowa Thistle (Cirsium iowense). Common in pastures and meadows. (Photographed by Colburn.) Fic. 222-A. Distribution of Iowa Thistle. DESCRIPTIVE MANUAL 379 teeth with weak prickles; rather large heads; involucre 1-114 in. long; bracts with broad glandular back, the inner with a some- what attenuated colorless tip. Distribution—Common in borders of woods and in fields. From Iowa to South Dakota and Kansas. In Story, Emmet, Kossuth, Marshall, Boone, Linn, Clinton, Webster and Carroll counties in prairie meadows. Extermination.—This biennial is readily destroyed by cutting the plants off below the surface of the ground. When left to flower it dies but in meadows where cut off above the surface of the ground it acts like a perennial. Canada Thistle (Cirsium arvense (li.) Scop.). Description.—Smooth perennial, spreading by roots and root- stocks, 1-3 ft. high, corymbosely branched at the top; stem smooth ; leaves lanceolate, sessile, and deeply pinnatifid, lobes and margins of leaf with spiny teeth; heads small, 34-1 in. high, bracts ap- pressed, the outer with a broad base, inner narrow, all with an acute, never spiny, tip; somewhat arachnoid flowers purple, di- oecious; in staminate plant, flowers exserted with abortive pistils, in pistillate less so, scarcely exceeding the bracts; tube of the corolla 6 lines long; stamens with abortive anthers, anther tips acute, filaments minutely pubescent; young achene pubescent; all of the bristles of the pappus plumose; trichomes simple, long, floccose. Distribution —This European weed is widely distributed in Can- ada to the Pacific coast; found in Jowa in many counties, more common in northern counties than in the southern. It is more or less abundant in Hardin, Pocahontas, Clinton, and Worth coun- ties, frequently in clover meadows and in pastures. Extermination.—The Canada thistle can be treated with sodium arsenite. No other chemicals, so far as our experiments extend, will entirely destroy this weed. Carbolic acid only partially de- stroys the roots and the plants shoot up again from below the point of injury, but by repeating the process the Canada thistle can ulti- mately be exterminated. A good method of eradicating the weed is to plow shallow and cultivate frequently during the summer. The roots of the Canada thistle extend deeply down into the soil, hence for this reason deep cultivation will be of no avail. After 380 WEED FLORA OF IOWA Fig, 223. Canada Thistle (Cirsium arvense). Common and widely distributed in Iowa. (After Clark and Fletcher.) DESCRIPTIVE MANUAL 381 Figure 223A Figure 223B Fic. 223- A. Trichome or plant hair from leaf of Canada Thistle. -(Drawing by Charlotte M. King.) Fic. 223-B. Distribution of Canada Thistle. plowing, the soil should be dragged and the roots exposed to the sun and removed, when possible. It may be necessary to run over the field with a hoe to cut off the stray plants which appear. This method was tried on a patch several years ago and no Canada thistles have since made their appearance in this place. Various crops, such as clover and sorghum, are said to be ef- fective in subduing the thistles. Of the various chemicals which have been used to exterminate Canada thistles none are more effective than sodium arsenite. It is apphed at the rate of 114-2 pounds to 52 gallows of water. Carbolic acid at the rate of one part to one part water destroys the root when it comes in contact with the mixture, and for a little distance beyond. This is not an effective method, as the roots sprout out from below. In response to circulars of inquiry sent out by the Iowa Experi- ment Station, the majority of correspondents recommend shallow plowing, disking, and harrowing; and continuing cultivation and hoeing as long as the thistles make their appearance. Some report successful treatment with galt when scattered thickly about the thistles, especially if cattle or sheep are given access to it. Some report success with carbolic acid when it is applied directly to the stem. Tarred paper in a few cases gave success, as did also the method where the thistle was covered thickly with straw or ma- nure. The depth of covering was not, however, given. 382 WEED FLORA OF IOWA Clark and Fletcher recommend the following treatment: ‘“The chief safeguard against the Canada thistle and all similar deep-rooted perennials is undoubtedly a regular short rotation. A three-year rotation, including two cuttings of early red clover for the first year, followed by a deep fall plowing for hoed crops with clean cultivation, and a cereal crop for the third year, will sup- press it.”’ A correspondent in The Prairie Farmer on the resisting quali- ties of Canada thistle says: ‘‘I have been debating in my mind and endeavoring to determine whether or not root-stocks or por- tions thereof have to some extent the hibernating qualities of the seed. For instance, take two seeds of the cocklebur—one hiber- nates for one season and the other for two seasons, with growth always present. Now the question is may not a Canada thistle with its root-stock or a portion of it lie dormant for a season or more? If this be true, then some of the best known methods of extermination are useless.’’ To this Prof. R. A. Moore says: ‘‘In regard to Canada thistle roots hibernating and retaining their vitality for several years, will say, that I think your correspondent’s version seems reason- able, and it is quite conclusive from this fact that the eradication of the pest is all the more difficult. It seems that many of the seeds and plants are given this power of remaining dormant when subjected to adverse conditions. In the lower order of plants, many of the species of bacteria when subjected to adverse environ- ments are transformed into resistant spores and will not vegetate until the conditions are favorable.’’ Wallace’s Farmer says concerning its eradication: ‘‘We believe if we had a quarter of an acre of Canada thistles we would let them alone until August, when the thistle will put forth its utmost efforts to produce seed. While the thistles are in full blossom we would mow them, rake them up and burn them, and then plow the ground about eight inches deep, throwing the furrow flat. Letting them put forth their full strength to produce seed and thwarting that by mowing and burning would weaken the roots materially. Then by plowing them under eight inches deep, if possible, you would attack them at their weakest point. DESCRIPTIVE MANUAL 383 Knapweed (Centaurea solstitcalis L.). Description.—Annual erect, branched, cottony stem; lower leaves’ lyrate, deeply pinnatifid, upper leaves, linear, entire or nearly so, decurrent wings on the stem; outer bracts, with long spreading spines, those at base few and smaller; flowers yellow, with soft pappus. Itc. 224. Knapweed, Star Thistle (Centaurea solstitialis). Im alfalfa fields. ; (Mich. Agr. Exp. Sta.) 384 WEED FLORA OF IOWA Fic. 224-A., Distribution of Knapweed. Distribution —Massachusetts to Ontario and Iowa; introduced with alfalfa seed into other western states. Extermination.—This annual weed has been introduced largely with alfalfa seed. Sow clean seed. Use seed coming from the Rocky mountains or Kansas where the weed is not common. It is an extremely objectionable weed because of ane spiny heads. It is easily killed by cultivation. Chicory or Succory (Cichoriwm intybus L.). Description—aA branching perennial with deep roots, alternate leaves; blue flowers, or sometimes pink or purple; basal leaves spreading on the ground; stem leaves oblong or lanceolate, partly clasping. Distribution.—It is common along roadsides in fields and waste places from New England to Canada to Nebraska, especially com- mon where chicory has been cultivated; it has become a trouble- some weed in Wisconsin and Minnesota. It is allied to endive, cul- tivated as a salad plant. Extermination.—Chicory is not difficult to destroy where ro- tation of crops is practiced. Though a perennial, the roots are easily killed by repeated cultivation. Clark and Fletcher recommend the following treatment: ‘‘A short rotation of crops will soon suppress it. Chicory is not often seen in good farming districts except as a wayside weed. Individual plants may be destroyed by close cutting and applying salt to the root in hot, dry weather.’’ DESCRIPTIVE MANUAL | 385, Piate 64 ’ CHIGORY - ay (Cichoriam infybus- 4) Fic. 225. Chicory (Cichorium intybus). Im clover and alfalfa fields, some- times in waste places. (After Clark and Fletcher.) 25 386 WEED FLORA OF IOWA WoC a owe TER SE, EEE RS, SSR N VT STS SRS sy ae UU hkl RSS Fic. 225-A. Distribution of Chicory. Dandelion (Taraxacum officinale Weber). Description.—A smooth, or at first pubescent, biennial or per- ennial; the many-flowered head borne on a slender hollow scape; root leaves pinnatifid or runcinate; involucre double, the outer of Fic. 226. Dandelion (Taraxacum officinale). Long root, sometimes to a depth of three and one-half feet in the soil. (Photographed by Gardner.) DESCRIPTIVE MANUAL tues 387 Fig. 226-A. Distribution of Dandelion. short scales, the inner of long, linear, erect scales in a single row; after flowering the inner involucre closes, the fruit is ripened close to the ground, and when ripe the hollow scape elongates and the whole involucre is reflexed permitting the wind to scatter the “seeds ;’’ “‘seeds’’ oblong, long-beaked, the beak being 2 or 3 times as long as the remainder of the achene, bearing at the end the pappus. j Fig. 226-B. A patch of Dandelion in a lawn, early in June. (Photographed by Charlotte M. King.) 388 WEED FLORA OF IOWA Distribution—The dandelion is cosmopolitan. It is quite as common in Europe as in the United States, being common and abundant at high elevations, as in the Rocky mountains. Found everywhere in lowa on lawns and pastures. Fic. 227. Common Dandelion (Taraxacum officinale). A weed common in lawns, pastures, etc. (Photographed by Colburn.) Extermination.—The dandelions are not difficult to exterminate in cultivated fields but in garden crops it is more difficult to do so, especially in strawberry beds. As the dandelions are perennial herbs, seed formation should be prevented. Where they occur in small patches a spud may be used successfully. Where they are abundant in lawns it is only necessary to keep the lawn closely cropped and dig them up with a small spud to prevent seeding. Blue grass and clover, especially the latter, will crowd them out. It is rare that dandelions give trouble after the middle of June. DESCRIPTIVE MANUAL 389 W RAN 7 Oi? S NS Ne ny AR Fie. 227-A. Common Dandelion (Taraxacum officinale). 1, head; 2, single flower; 3, achene; 4, receptacle and seed with pappus. Experiments have been made with herbicides and the one most commonly used is iron sulphate which is usually applied at the rate of 100 pounds to one barrel of water. The weed where prop- erly sprayed will be destroyed, but owing to the fact that the dan- delion possesses a long perennial root, sometimes 4 feet long, it will sprout again. The spraying must be kept up until fall. Chemical Composition.—According to the report of the Bussey Institution the chemical composition of the dandelion is as follows: FRESH OR AIR DRY MATERIAL. . Z Nitrogen free Water | Ash | Protein | Fiber | Extract Fat 85.54 | 1.99 | 2.81 | 1.52 745 | 0.69 WATER FREE SUBSTANCE. | 13.8 | 19.4 | 10.5 51.5 4.8 *Bull. 1877. Compiled by Jenkins and Winton; Bull. 11, Off. Exp. Sta., U. S. Dept. Agr. 390 WEED FLORA OF IOWA Red-seeded Dandelion (Taraxacum erythrospermum Andyz.). Description—A perennial weed with long root; leaves deeply runcinate-pinnatifid or pinnately divided into narrow segments; heads somewhat smaller than in common dandelion, sulphur yel- low; involucre glaucous, the inner bracts corniculate, appendaged at tip; the outer short, spreading or ascending; achene reddish, tuberculate above. Distribution.—Red-seeded dandelion is a much more recent in- troduction than common dandelion. It occurs from Maine to Kansas. Naturalized from Europe, in similar situations to com- mon dandelion in Iowa; sometimes less abundant and sometimes more abundant than the latter. Extermination—May be exterminated in the same way as the common dandelion. Fic. 228. Red-seeded dandelion (Taraxacum erythrospermum). Common in lawns. (Photographed by Colburn.) Clark and Fletcher recommend the following treatment: ‘‘It is important to prevent dandelions from seeding in lands adjacent to lawns. Though entailing much labor, the most satis- factory way to deal with them, when deeply rooted in lawns, is to loosen the soil with a digging fork and pull them up. The use of DESCRIPTIVE MANUAL 391 the fork may not be necessary in some wet and soft soils. The ap- plication of kerosene to the crown of the plant, in the centre of the rosette of leaves, is recommended. Sulphate of ammonia or sulphate of iron in excess is also recommended. A small table- spoonful of salt applied in the morning of a hot day, when the soil is dry, will kill them. Annual Sow-thistle (Sonchus oleraceus L.). Description.—Annual, succulent herb with leafy, smooth stems, and pale yellow flowers in corymbose or umbellate clusters; leaves of stem dentate, runcinate-pinnatifid, terminal with a large seg- ment; heads numerous; blossoms in late summer and fall. Distribution.—This weed is common in fields and waste places throughout North America, especially in the north. It is common in Europe and also occurs in Mexico and South America. Extermination.—Sow thistle is easily exterminated by cultiva- tion. It can also be exterminated by the use of iron sulphate at the rate of 100 pounds to a barrel of water; where it is abundant it may be necessary to make two or three applications. Clark and Fletcher recommend as follows: ‘‘Prevent them from seeding in waste places by cleaning them up and seeding them to permanent, vigorous grasses. This annual weed, with its rela- tively small, pale yellow flowers, when compared with perennial species is not difficult to control by ordinary methods of cultivation and alternation of crops. Sheep, if sufficient in numbers, will pre- vent sow thistles from seeding in pasture lands.’’ The Iowa Homestead says concerning its eradication: ‘‘We have seen the sow thistle take complete possession of a soil, growing so thickly that other crops were entirely choked out. Nec- essarily the best way to destroy it is to cultivate freely. If it makes its appearance in stubble ground we would advise plowing as early as possible after harvest and the cultivation of surface at intervals during the late summer and fall in order to keep the thistles below the ground. 392 WEED FLORA OF IOWA Fic. 229. Common Sow Thistle (Sonchus oleraceus). Waste places. (After Clark and Fletcher.) DESCRIPTIVE MANUAL 393 Fic. 229-A. Distribution of Annual Sow Thistle. Sow Thistle (Sonchus asper (.) Hill). : Description.—Like the preceding species except that stem leaves are less divided and more spiny-toothed, with auricles of the Fic, 230. Sow Thistle (Sonchus aspen), Waste places, yellow flowers and milky juice. (Photographed by Colburn.) 394 WEED FLORA OF IOWA oo asi aes Fic. 230-A. Distribution of Sow Thistle. clasping base rounded; achenes 3-nerved on each side and margined, smooth. Distribution—Common with the preceding species, in waste places. Extermination—This annual is not difficult to exterminate by giving thorough cultivation. Perennial Sow Thistle (Sonchus arvensis L.). Description.—A perennial with creeping root-stock, and milky juice; leaves runcinate, pinnatifid and spiny toothed, heart-shaped base; flowers yellow; peduncle and involucre bristly; achenes ob- compressed, wrinkled on the ribs. Distribution—Common from Nova Scotia west to Saskatchewan, North Dakota, Minnesota, occasional in Iowa to New Jersey and New England. : Extermination.—A most difficult weed to exterminate and should be treated like the Canada thistle. Summer fallow with frequent cultivation is the only successful method. Clark and Fletcher of Canada, recommend the following: “Small patches may be eradicated by digging out the roots as thoroughly as possible and destroying them. This may have to be done several times during a season. Great care must be taken not to distribute pieces of the rootstocks over the fields by harrows or other implements. To exterminate Perennial Sow Thistle, some system must be adopted which will prevent the development: of leaves for a period sufficiently long to kill the roots by smothering DESCRIPTIVE MANUAL 395 Fic. 231. Perennial Sow Thistle (Sonchus arvensis). Plants with bright yel- low flowers, milky juice. Not common in Iowa. A difficult weed to exter- minate. (Photographed by Colburn.) Fic. 231-A. Distribution of Perennial Sow Thistle. 396 WEED FLORA OF IOWA them. When a field is badly infested it requires special treatment for a season and close’ attention for a number of years. One of the most effective methods is to plow lightly immediately after the hay or grain crop is removed and follow with frequent use of a broad-shared cultivator. Late in the fall plow again, somewhat deeper. In the spring give frequent cultivation, so as to prevent the development of any leaves and thus weaken the roots to the greatest possible extent. About the middle of June or first of July sow rape in drills at the rate of about 114 lbs. per acre. Cultivate between the drills as soon as possible and repeat at short intervals until the rape completely covers the ground. Some hand hoeing may be necessary to keep all the thistles down. This should pretty well exterminate the pest but if some plants still remain when the rape is cut or pastured off, the field may be fall plowed and put into hoed crop the next season, when special attention can be given to any small patches that may appear. Buckwheat is sometimes used instead of rape for a smothering crop.’’ Prickly Lettuce (Lactuca scariola L. var. integrata Gren. & Godr.). Description.—Tall, erect herbs, glaucous, green, 2-5 ft. high, simple or branched except the lower part of stem which has stiff bristles; leaves glaucous, green, smooth except the midrib which is beset with weak prickles lanceolate to oblong in outline, with spinulose, denticulate margins, occasionally sinuate-toothed; flow- ers pale yellow. The leaves of ZL. scariola are pinnatifid and more prickly ; trichomes multicellular. Distribution.—Prickly lettuce was introduced into Massachu- setts about 1863 from Europe. It is quite widely distributed in northern Africa and Europe and has become frequent throughout the northern Mississippi valley to the Pacific coast. It is com- mon everywhere in Iowa particularly along roadsides, highways and in gardens. The ZL. scartola is becoming more frequent in Iowa, in Ames, Des Moines, Boone, etc. Extermination—The weed is easily exterminated from culti- vated fields and in waste places by cutting off young plants below the ground. Where the stem is cut off below the surface of the ground it will give no trouble, but in meadows and lawns where the plants are cut off above the ground the weed will continually reappear, producing from 3-6 branches. The following excellent suggestions are made by L. H. Dewey: DESCRIPTIVE MANUAL 397 a PRICKLY LETTUCE fuCR SEGMIGI G fier rica: A Ge Fic. 232. Prickly Lettuce (Lactuca scariola var. integrata). Common road- sides and gardens. (After Clark and Fletcher.) 398 WEED FLORA OF IOWA SetMERMneS e BUYS ee iM ase NE Ane = aa ‘et Figure 232A Figure 232B Fic. 232-A. Bristles on Prickly Lettuce (integrata). (Drawing by Charlotte M. King.) Fic, 232-B. Distribution of Prickly Lettuce. ‘“Sheep and sometimes cattle will eat the young prickly lettuce, and in some localities their services have been found very effective in keeping it down, especially in recently cleared land where culti- vation is impossible. Repeatedly mowing the plants as they first begin to blossom will prevent seeding and eventually subdue them. Thorough cultivation with a hoed crop, by means of which the seed in the soil may be induced to germinate, will be found most effective. The plowing should be shallow so as not to bury the seeds too deep. Under no circumstances should the mature seed- bearing plants be plowed under, as that would only fill the soil Fic. 232-C. Distribution of Prickly Lettuce (Lactuca scariola). DESCRIPTIVE MANUAL 399 with seeds buried at different depths to be brought under condi- tions favorable for germination at intervals for several years. Ma- ture plants should be mowed and burned before plowing. The seed appears as an impurity in clover, millet and the heavier grass seeds, and the plant is doubtless most frequently introduced by this means. As the seeds may be carried a long distance by the wind, the plants must be cleared out of the fence rows, waste land and roadsides.”’ Clark and Fletcher recommend as follows: ‘‘The seed is short-lived and if the plant is kept closely cut and prevented from seeding in waste places for two or three years it will soon disappear from cultivated areas. Clean waste lands and seed to permanent vigorous grasses. Ordinary methods of cultiva- tion will suppress it in the fields.’’ Wild Lettuce (Lactuca canadensis L.). DescriptionStem leafy, glabrous or nearly so, glaucous; lower leaves sinuate, pinnatifid, upper entire; heads numerous, in a rather long, open panicle; flowers yellow. Distribution.—Nova Scotia to Ontario, Ohio and westward; fre- quently found in Iowa. ; Extermination This perennial is not difficult to destroy by cultivation. . 400 WEED FLORA OF IOWA Fic. 233. Wild or Canadian Lettuce (Lactuca canadensis). Fields, roadsides and meadows. (Photographed by Quade.) DESCRIPTIVE MANUAL 401 Fic, 233-A. Wild or Canadian Lettuce (Lactuca canadensis). Plant with milky juice and yellow flowers. (Photographed by Quade. ) Fic. 233-B. Distribution of Wild Lettuce. 26 402 WEED FLORA OF IOWA Blue Lettuce (Hactuca pulchella (Pursh.) DC.). Description.—Perennial plant, deep-rooted, pale or glaucous; stem simple, about 1 foot high; leaves sessile, oblong or linear- lanceolate, glabrous, entire, or lower leaves somewhat pinnatifid; racemose heads large, erect; peduncles with scaly bracts; bracts of involucre imbricated in 3-4 ranks; flowers blue. Distribution.—Northern Michigan and Ontario southward; re- ported from several localities in Iowa. Extermination.—Prevent from distributing seed by continued cutting before flowering. If it becomes established in a field, try thorough summer fallow with deep cultivation so as to check growth of long rootstocks. Fig. 234. Blue Lettuce (Lactuca pulchella). Blue flowered lettuce. Plant with milky juice. Common in western Iowa. (Photographed by Quade.) DESCRIPTIVE MANUAL 403 Fic. 234-A. Distribution of Blue Lettuce. Rushlike Lygodesmia (Lygodesmia juncea (Pursh.) D. Don.). Description —A tufted, smooth, frequently glaucous perennial a foot or more high, coming from a thick woody root, with copious Fic. 235. Lygodesmia or Skeleton Weed (Lygodesmia juncea). A deep rooted perennial with a yellowish milky juice. (Photographed by Colburn.) 404 WEED FLORA OF IOWA Fie. 235-A. Distribution of Lygodesmia. yellowish juice; lower leaves rigid, linear-lanceolate, small, entire, the upper scalelike; flowers purple in erect heads; achenes nar- row-ribbed, pappus light brown. This weed is sometimes called skeleton weed. Distribution.—This weed is common from Missouri river to west- ern Montana and east to St. Croix river in Wisconsin. A some- what troublesome weed in northwestern Iowa; easily recognized by the yellowish juice and rushlike stems. Lixtermination.—This plant produces a long root which enables the plant to spread rapidly. It also produces a large number of ““seeds.’? Where the weed is common, plough the field after harvest and disk the plants. It is a good plan to follow up with a hoe, cutting off the plants. CHAPTER II. THE GENERAL CHARACTERS OF SEEDS By L. H. PAMMEL and CHARLOTTE M. KING CHAPTER II. CYPERACEAE, SEDGE FAMILY. Sedge (Carex vulpinoidea Mx.). Achene enclosed by an utricle called the perigynium, hardly 3- angled, somewhat flattened, tipped by lanceolate 2-toothed beak, the persistent base of the flower-style; achene flask-shaped, about Figure 236A Figure 236B Fic. 236. Forms of Sedges. a, Hleocharis palustris; b, Carex vulpinoidea. (After Gray.) one-sixteenth of an inch in length, light brown; surface incon- spicuously nerved; whitish, projecting scar. Spike Rush (Eleocharis palustris R. Br.). Achenes slightly flattened, somewhat lenticular, with 2 or 3 ob- tuse angles one-twelfth to one-tenth in. long, brown, smooth, shin- ing, minutely cross-striated ; seed with persistent tubercle from tip ; tubercle conical, triangular, constricted; bristles pale, longer than 408 WEED FLORA OF IOWA Fic. 237. Achenes, “seeds,’’ of common forms of Sedges (Carer). (After Hillman, Bull. Nevada Agr. Exp. Sta.) achene; seed may appear with or without tubercle and bristles. Seed occasionally found in seed of alsike and red clover from wet erounds. GRAMINEAH, GRASS FAMILY. Johnson Grass (Sorghum halepense (li.) Pers.). Sessile spikelet broadly lanceolate, acute, 4-6 mm.* long, be- coming dark at maturity ; callus small, obtuse, shortly and sparsely barbate; first glume coriaceous, slightly pubescent on the flattened back, 5-7 nerved; second similar, equal to first, with hyaline, ciliate, Fic. 238. Spikelet of Johnson Grass (Sorghum halepense). (Drawn by Charlotte M. King.) inflexed margins; the third glume shorter, membranous, faintly 2-nerved, with ciliate, infolded margins; fourth glume broad, ob- tuse, shorter than second, 2-lobed at apex, ciliate, awned; awn 10-16 mm. long; palea shorter than glumes, without nerves, ciliate. Smooth Crab Grass (Digitaria humifusa Pers.). Spikelets lanceolate or elliptical, one-twelfth in. in length, acut- ish; glumes usually present, first glume wanting, sometimes rudi- mentary, 3-nerved; the first and second, hairy on the margins; *1 mm. = 0.0394 inch, or nearly one-twenty-fifth inch. THE GHNHRAL CHARACTERS OF SEEDS 409 Fig. 239. Seeds of Smooth Crab Grass (Digitaria humifusa). A and B, spike- lets; A showing the second glume, B showing the third glume. C, D and #, florets; D, bearing a portion of the second glume, H, the inner face, showing the edges of the flowering glume. F, a spikelet of Digitaria filiformis, show- ing the shorter second glume. G, the natural size of both of these species. (After Hillman, Bull. Nevada Agr. Exp. Sta.) the third, 7-nerved; the fourth, dark purplish brown. Occurrence very frequent in alfalfa clovers and commercial grass seed. Crab Grass (Digitaria sanguinalis (L.) Scop.). Spikelets one-seventh in. in length with usually persistent scale- like glumes, lanceolate, pedicellate; second glume usually ciliate on margins, short; fourth glume silky-villous along marginal nerves, 5-nerved, color pale. Frequently occurs in red and alsike clover, as well as in timothy. FF a co fe He Fig. 240B Figure 240C Fic. 240. Common Crab Grass (Digitaria sanguinalis). A, an enlarged view of Crab Grass. B, with small glume attached. C. A, spikelets showing the second glume, floret, and edges of the third glume. B, the opposite face, showing the minute first glume and third glume. D and H, the two faces of the floret. F, seeds, natural size. (A and B, Drawings by C. M. King; C, after Hillman, Bull. Nevada Agr. Exp. Sta.) 410 WEED FLORA OF IOWA Tickle or Hair Grass (Panicum capillare L.). Spikelets small, ovate or acute, one-fifteenth to one-twelfth in. long, acuminate-pointed, smooth, shining; sterile glumes usually absent, when present not shining; first glume 1-3-nerved, obtuse to Figure 241A Figure 241A Figure 241B Fic. 241. A. Tickle or Hair Grass (Panicum capillare)., At the left a spikelet opened, at the right various views. Fig. 365A i! 1 \ . Fig. 3650 Fic. 365. Achenial hairs of Beggar-ticks. A. Bidens frondosa. B. Bidens cernua. C. Bidens aristosa. (Drawings by Mary A. Nichols.) Sneezeweed (Helenium autumnale L.). Achenes straw-colored, with several longitudinal ribs, length 1 mm., breadth 0.3 mm. at apex, toward which the achene widens; surface bears scattered appressed hairs; pappus several chaffy points on margin of truncate apex. Fetid Marigold (Dyssodia papposa (Vent.) Hitche.). Achene slender, 4-angled; length 3 mm., width at truncate apex .75 ram.; black, with numerous, scattered, appressed black hairs; pappus a row of chaffy scales, dividing into numerous, rough, bristly hairs; 3 mm. long. According to Mr. Fracker, the achenial hairs are simple and duplex, the tip in duplex hairs deeply cleft. THE GENERAL CHARACTERS OF SEEDS 491 Fig. 366A Fig. 366B Fig. 366C Fic. 366. Seeds and achenial hairs of Sneezeweed and Fetid Marigold.’ A. Sneezeweed (Helenium autumnale). B. Fetid Marigold (Dyssodia papposa). C. Achenial hairs of Fetid Marigold. (A and B, drawings by Charlotte M. King; C, from drawing by Mary A. Nichols.) Yarrow (Achillea millefolium L.). Achenes one-twelfth to one-tenth in. long, oblong to obovate, somewhat compressed; light on the margin, the remainder some- what brownish; sometimes slightly curved, base with prominent scar with slightly raised border; apex larger with a notch in the center ‘and a projecting knob; surface of the achene marked with fine lines ; pappus absent. A Fig. 367A Fig. 367B Fic. 367. Seeds of Yarrow and Mayweed.! A, Yarrow (Achillea millefolium) ; a seeds (achenes) in side view, the two at the left showing the minute, cir- cular sear; b, group showing the natural size. B. Mayweed (Anthemis cotula); a, a group of seeds showing the prevailing forms; b, a group showing the natural size. (After Hillman, Bull. Nev. Agr. Exp. Sta.) Mayweed (Anthemis cotula L.). Achenes one-twentieth to one-sixteenth in. long, oblong with preminent tubercled ribs, or occasionally smoothish curved, the base tipped with smooth nipple-like projections; pappus absent; straw- colored to light brown; bearing a projection scar; base with round, light-colored sear. Common in clover and grass seeds. 492 WEED FLORA OF IOWA Ox-eye Daisy (Chrysanthemum leucanthemum L.). Achenes flattened, club-shaped, straight or slightly curved, one- twentieth to one-tenth in. long, oblong; angles white, with brown interstices ; 5-10-ribbed ; small scar at basal end; pappus wanting. Fic. 368. Seeds of Ox-eye Daisy (Chrysanthemum leucanthemum). A, pre- vailing forms of seeds (achenes); B, one in cross section; C, a group showing the natural size. ‘ (After Hillman, Bull. Nev. Agr. Exp. Sta.) Wormwood (Artemisia biennis Willd.). Achenes brown, smooth, from 3-4 flattened faces, angled between longitudinally ; length .£8 mm., width .3 mm., broader at apex than at base. 2 x a . oe ~ ~ ~ eo 4 = . . a 4 b b Fig. 369A Fig. 369B Fic. 369. Seeds of Wormwood and Fireweed. A. Wormwood (Artemisia biennis) ; a, achenes, b, natural size. B. Fireweed (Hrechtites hieracifolia) ; a, achenes, b, natural size. (After Hillman, Bull. Ney. Agr. Exp. Sta.) Fireweed (Hrechtites hieractfolia (L.) Raf.). Achene linear-oblong, straight, or curved, prominently striate, pubescent, beakless, one-sixth in. in length; upper end with a white ring, within the ring a slight depression; scar at the lower end with whitish ring, and a small depressed opening; pappus of numerous white soft, capillary bristles. THE GENERAL CHARACTERS OF SEEDS 493 Indian Plantain (Cacalia tuberosa Nutt.). Achenes oblong, straight, or slightly curved, one-fifth in. in length; brown; prominently ribbed, the ribs minutely roughened ; the apex of the seed with a slightly projecting rim or border to which the pappus is attached; the base with a circular ring; pappus with numerous fine, white, capillary bristles. Fig. 370A Fig. 370B Fic. 370. Seeds of Indian Plantain and Groundsel. A. Indian Plantain (Cacalia tuberosa). B. Groundsel (Senecio vulgaris). (Drawings by Charlotte M. King.) Groundsel (Senecio vulgaris L.). Achenes teretish or those of marginal flowers compressed, narrow, eylindrical; 5 or 10-ribbed, pubescent; one-half in. long; upper part expanded, extending beyond narrow portion of upper part of achene; lower portion with a depression; color reddish; pappus of fine, white, capillary bristles. Burdock (Arctium lappa L.). Achenes 3-5-ridged, upper portion truncate, one-fifth to one- fourth in. long, compressed or oblong, nearly straight to slightly eurved, 3-angled; surface mottled in appearance, due to the small serrulate scales with projecting tips of ridges beyond the border; scar surrounded by a circular lighter colored area; scar at base lighter in color; pappus of numerous short scales. In commercial seeds, occasionally. 494 WEED FLORA OF IOWA 74 Fic. 871. Seeds of Burdock (Arctium lappa). a, a side view of one of the inner achenes of a bur; b, showing the inner surface of a curved, outer achene, and exhibiting the character of the apex, both enlarged; c, a group show- ing natural size. (After Hillman, Bull. Nev. Agr. Exp. Sta.) Tall Thistle (Cirsium altissimum Willd.). Achenes lanceolate-obovate, tapering toward lower end, and some- what narrowed toward the apex, wider in the middle, one-eighth in. in length; width one-twelfth in.; dull brown or grayish excepting light colored ring at upper end; numerous fine ridges; apex con- cave, with the remnant of corolla tube projecting from the center; color uniform; not marked by light and dark areas as the bull thistle. Fig. 372A Fig. 373B Fic. 372, Seeds of Thistles. A. Different views of Canada Thistle (Cirsium arvense). B. Bull Thistle (Cirsium lanceolatum). (Drawings by Charlotte M. King.) Canada Thistle (Cirsium arvense (L.) Scop.). Achene lanceolate, narrowed at lower end, tapering from some- what thickened top, one-twelfth to one-eighth in. long; the eup- shaped top with a projecting conical portion, straight or slightly curved; marked with longitudinal dark lines or furrows; apex with a light colored border, giving appearance of a ring. Found in seed of small grains, clovers and grasses. THE GENERAL CHARACTERS OF SEEDS 495 Field Thistle (Oirsiwm discolor (Muhl.) Spreng.). Achenes obovate, inner edge nearly straight, outer curved, convex ; one-seventh to one-sixth in. long, tapering from narrowed base to apex, upper part longitudinally striated, slightly pigmented ; Fic. 373. Seeds of Thistle. 3. Cirsium altissimum. 4. Cirsium discolor. 5. Cirsium undulatum. 6. Cirsium ioense. (Drawing by Charlotte M. King.) grayish, upper part yellowish for one-third length of the achene. Found in seed of alfalfa and red clover. Towa Thistle (Cirsium ioense (Pammel) Fernald). Achenes obovate, lanceolate, one side of seed straight, outer side slightly convex, one-fifth to one-fourth in. long, one-twelfth to, one- sixteenth in. in width, gradually tapering from lower end toward upper end, prominently widened just below the apex, marked by longitudinal striations and dark areas, upper part yellowish, promi- nent rim glossy; apex concave, with the prominent projecting rem- nant of flower center. Bull Thistle (Cirsium lanceolatum (.) Hill). Achene lanceolate, curved, tapering, in many cases somewhat angular, one-eighth to one-sixth in. long; brownish, not darkly striated, marked with definite grooves, upper part lighter, ringed, also showing at center of concave apex; apex with projecting point. Found in red clover and in alfalfa seed. Waivy-leaved Thistle (Cirsium undulatum (Nutt.) Spreng.). Achene in general outline lanceolate, tapering from base, inner edge slightly convex, several prominent ridges, light brown, not pigmented; upper part yellowish; apex with prominent rim and prominent remnant of corolla tube; achene slightly thicker than that of C. discolor. Found in seeds of red clover and alfalfia. 496 WEED FLORA OF IOWA Chicory (Cichorium intybus L.). - Achenes oblong, ribbed, 5-angled, spotted, grayish or straw- colored, with darker spots, one-twelfth to one-eighth in. in length apex with the base of the pappus scales extending beyond the scar; Fic. 374. Chicory (Cichorium intybus). A, B, two views of seeds; C, seeds, natural size. (After Hillman, Bull. Mich. Agr. Exp. Sta.) minutely, transversely roughened ; base of seed lighter in color, with small depressed scar; achenes from the inner part of the flower more slender and straight than from outer part. Found with various commercial seeds. Corn-flower (Centaurea cyanus L.). Achene oblong or obovoid, compressed, one-sixth to one-fifth in. long; smooth, shining; with inconspicuous veins or nerves; lower part of achene oblique; ivory-white; the elliptical or somewhat cir- cular scar at base with whitish rimmed border, depressed, made up of soft tissue; pappus of several series of scaly bristles; brownish in color. The achenial trichomes, according to Mr. Fracker, are long and simple. Barnaby’s Thistle (Centaurea solstitialis L.). - Involucre ovoid, 1.5 cm. in diameter, with stout straw-colored spines, widely spreading ; achene cream or pale brown after matted; length 2 mm., scar of attachment in a notch above rounded base; apex truncate, bearing tubercle in the center. THE GENERAL CHARACTERS OF SEEDS 497 Fig. 375A Fig. 375B é Fig. 375C Wie. 375. Seeds of Corn-flower and Knapweed. A. SBarnaby’s Thistle or Knapweed (Centaurea solstitialis); a, achene with pappus, ¢c, pappus re- moved, b and d, achenes, natural size. B. Corn-flower (Centaurea cyanus). C. Achenial hairs of Corn-flower. (A, after Hillman, Bull. Nev. Agr. Exp. Sta.; B, after Nobbe; C, drawing by S. B. Fracker.) Bristly Ox-tongue (Picris echioides L.). Achene elliptical, narrowed at base, with projection at tip ; one- tenth in. in length; light brownish red; wrinkled transversely, espe- cially toward apical end. Found in seed of alfalfa and clover. Oyster Plant, Salsify (Tragopogon porrifolius L.). Achenes linear, terete, beaked or long, covered with scalelike tubercles on the ribs, or merely roughened, light straw-colored or darker pappus of numerous plumose bristles; small scar at apex, whitish with a depression scar at base with an oval depression. Reported as a “‘weed of alfalfa meadows.”’ Meadow Salsify (Tragopogon pratensis L.). Achenes linear, terete, one-half in. or little more long, exclusive of beak; scar at base, whitish, with an oval depression; beak one- third in. long; scar at end of beak whitish, with a depression, striate, smooth or slightly roughened, light straw-colored or darker ; pappus of numerous soft, brownish, plumose bristles. : 382 498 WEED FLORA OF IOWA es. — SS === SOR ae Eo ee ee . aos St Cant saa x ENS et a Raabe eit Fig. 376A Fig. 376A1 Fig. 376B Fig. 376C Fic. 376. Seeds of Picris and Tragopogon. A and Al. Bristly Ox-tongue (Picris echioides). B. Oyster Plant (Tragopogon porrifolius) ; a, achenes enlarged, b, achenes natural size. C. Meadow Salsify (Tragopogon pratensis). (A, drawings by L. R. Collins; B, after Hillman, Bull. Mich. Agr. Exp. Sta. ; C, drawing by C. M. King.) Dark-seeded Dandelion (Taraxacum erythrospermum Andrz.). Achenes one-sixth to one-fifth in. long; lance-shaped, or spindle- shaped, 5 longitudinal ridges, upper end with rough tubercles, per- sistent long beak about two-fifths in. long, brownish, somewhat pointed, prominently ribbed; pappus of numerous capillary, fragile, white bristles ; color reddish ; toothed at apex; beak and pappus both shorter; achenes more prominently tubercled than in 7’. officinale. Sor >, =< i Fig. 377A Fig. 377B Fic. 377. Dandelion seeds. A. Red Seeded Dandelion (Taraxacum erythros- permum) ; a, seed enlarged, b and c, seed, natural size, c, with pappus. B. Common Dandelion (Taraxacum officinale).- A, an enlarged view of one of the two similar faces of an achene, in which no attempt is made to show the minute surface-scales which are not evident under the ordinary lens. B, a group of seeds, natural size. b, one bearing the beak and pappus. C, a group showing common variation in the form of the achenes. (After Hillman: A, Bull. Mich. Agr. Exp. Sta.; B, Bull. Nev. Agr. Exp. Sta.) THE GENERAL CHARACTERS OF SHEDS 499 Dandelion (Taraxacum officinale Weber). Achenes two-fifths in. long including persistent beak, fusiform in shape, prominent ribs, and projecting teeth at ribs, especially at upper end; beak four-fifths to one in. in length; achenes light gray in color, otherwise like preceding; pappus capillary, whitish, fragile. With grass seeds. Field Sow Thistle (Sonchus arvensis L.). Achenes dark reddish brown, dull, ends slightly truncate, length 2.5-3 mm., width 0.8 mm., somewhat flattened, with 4 strong ribs or angles, between which lie smaller ridges ; numerous transverse ridges on the ribs. Fig. 378C Fic. 378. Seeds of Sow Thistles. A. Field or Perennial Sow Thistle (Sonchus arvensis) ; B, natural size, showing pappus on one achene, B. Sonchus oleraceus, different views of achenes. C. Sonchus asper, A and B, different views of achenes, C, achenes, natural size. (After Hillman, Bull. Mich. Agr. Exp. Sta.) Spiny Sow Thistle (Sonchus asper Vill.). Achenes broadly oblong or lance-shaped, similar faces, many- ribbed, one-tenth in. long, slightly pubescent, prominent ridges minutely roughened; base with a minute sear, brownish; the apex with projecting point and fine, capillary, white, bristles. Found with grass seeds. 500 WEED FLORA OF IOWA Sow Thistle (Sonchus oleraceus L.). Achenes light brown, flattened, ribbed, the prominent ribs rough- ened; one-eighth in: long; pappus of white capillary bristles, much like the preceding. ‘‘Apt to occur among grass seeds.’’ Wild Lettuce (Lactuca canadensis L.). Achene three-twentieths in. in length, straight or curved, sides somewhat unequal, transversely wrinkled, blackish brown, beak one- twentieth in. long, shorter than in L. floridana; one faint rib on each side of prominent midrib; decidedly convex on each side of midrib to the flattened margin ; scar with whitish ring and small depression ; pappus white, capillary. Much like ZL. floridana, beak smaller and shorter. With commercial seed. Fig. 379A Fig. 379B Fic. 379. Seeds of Lettuce. A. Lactuca canadensis; a, enlarged achene, b, natural size. B. Lactuca floridana. (A, after Hillman, Bull. Mich. Agr. Exp. Sta.; B, drawing by Charlotte M. King.) False Lettuce (Lactuca floridana (l.) Gaertn.). Achene brown, transversely wrinkled, three-twentieths to seven- fortieths in. in length, straight or slightly curved; beak light brown, about one-fifth in. long, generally persistent, convex; ribs rather faint, 1 on each side of prominent midrib, strongly convex on each side of midrib, margin flattened; pappus white, capillary. Prickly Lettuce (Lactuca scariola var. integrata Gren. & Godr.). Achenes brownish, margin somewhat lighter, surface roughened, one-ninth to one-sixth in. long; beak one-tenth to one-eighth in. long, lance-shaped, straight or slightly curved, upper end tapering THE GENERAL CHARACTERS OF SEEDS fe 501 v BS Dp ~ ( Fig. 380A Fig. 380B Fic. 380. Lettuce Seeds. A. Lactuca scariola; A, side view of an achene, en- larged. B, a group of the same, natural size. C, an achene bearing its beak and fragile pappus. B. Prickly Lettuce (Lactuca scariola var. integrata.) (A, after Hillman, Bull. Nev. Agr. Exp. Sta.; B, drawing by L. R. Collins.) toward the beak, somewhat flattened, on one side margined, faces convex, with 5-7 longitudinal nerves; scar circular, with a small depression. Rattlesnake Root (Prenanthes alba L.). . Achene oblong or columnar, truncate, somewhat flattened, 4-5 angled, brownish; upper part with a projecting ring to which the bristles of pappus are attached; scar at the base whitish, not prom- inent; one-eighth in. in length; pappus tawny. Fig. 381A Fig. 381B Fic. 381. Seeds of Rattlesnake Weed and False Calais. A. Rattlesnake Weed (Prenanthes alba). B. False Calais (Agoseris cuspidata). (Drawings by Charlotte M. King.) 502 WEED FLORA OF IOWA False Calais (Agoseris cuspidata (Pursh.) Steud.). Achenes fusiform, slightly contracted at the apex, with 10 prominent ribs; one-half in. long, or little longer; pappus of soft, white, capillary bristles; scar at base, whitish, with a small opening. Orange Hawkweed (Hieraciwm aurantiacum L.). Achene oblong, columnar, one-tenth in. long; blackish or dark brown, marked with prominent longitudinal ridges, minutely roughened; base of achene with small circular raised border, lighter than remainder of achene; pappus of numerous somewhat brown- ish bristles, frequently breaking away near the top of the achene, then showing short bristles. Fig. 382A Fig. 382A1 Fig. 382B Fic. 382. Seeds of Hawkweeds. A and Al, Orange Hawkweed (Hieracium aurantiacum). B, Hawkweed (Hieracium canadense). (A, after Hillman, Bull. Mich. Agr. Exp. Sta.; Al, drawing by L. R. Collins; B, drawing by Charlotte M. King.) Hawkweed (Hieracium canadense Mx.). Acheres one-tenth to one-eighth in. long, oblong, columnar, 10-15-ribbed; blackish or reddish; base with prominent disklike area marked by circular ring; apex with somewhat fine tawny- colored capillary bristles; below point of attachment is an area bulging outward; capillary bristles have tendency to break. CHAPTER III. MICROSCOPIC STRUCTURE OF SOME WEED SEEDS By L. H. PAMMEL and CHARLOTTE M. KING CHAPTER II. -GRAMINEAE,* GRASS FAMILY. Smooth Crab Grass (Digitaria humifusa Pers.). A cross section of seed shows but slight development of the pericarp and testa. The epidermal cells of the former are smaller than the underlying rows of cells of the wall of the ovary. The testa is reduced to a single layer of cells, longer than broad. The aleurone layer is filled with protein grains. The starch cells of the endosperm are larger and densely packed with polygonal grains. Fic. 383. Structure of the grain fruits of some Grasses (Gramineae). I. Wild Barley (Hordeum jubatum). II. Timothy (Phleum pratense). III. Smooth Crab Grass (Digitaria humifusa). IV. Sandbur (Cenchrus tribu- loides). V. Millet (Setaria italica). VI. Corn (Zea mays). VII. Barn- yard Grass (Echinochloa crusgalli). t=testa. al=aleurone. en=endosperm. n=nucellus. (Drawings by L. H. Pammel.) ae, *The material for this family has been taken from the accounts by Pammel and Winton. 506 WEED FLORA OF IOWA Broomecorn Millet (Panicum miliaceum L.). The walls of the ovary are similar to those of Digitaria humifusa except that they are wider. The testa is much compressed and consists of several layers of small cells. The cells of the aleurone layer are small, somewhat longer than broad. The cells of the starch layer are similar to those of D. humrfusa. Barnyard Grass (Echinochloa crusgalli (l.) Beauv.). The adherent glumes in this species consist of several rows of parenchyma cells, the inner portion, of one row of thick-walled sclerotic cells with pore canals. The cells of the pericarp and testa are much as in the other species, thin-walled and compressed. The protective features are preserved in the coriaceous glumes. The starch and aleurone layers are similar to those of Digitaria humifusa. Hungarian Grass or Millet (Setaria italica (L.) Beauv.). The colorless smooth pericarp is but slightly thickened and con- sists of three or four rows of elongated cells. These in colored seeds contain the pigment. The testa is but slightly developed. The cells of the aleurone layer are not much longer than broad, and are densely filled with protein grains. Green Foxtail (Setaria viridis (L.) Beauyv.). Dr. A. L. Winton describes the microscopic structure as follows: ~ Empty Glumes and Glume of Sterile Flower. The lower empty glume is three-nerved and less than 1 mm. long; the upper empty glume and the glume of the staminate flower are five-nerved and 2 mm. long. In microscopic structure the three are practically identical. 1. Outer Epidermts. Characteristic of this layer are the elongated cells with sinuous side walls and longitudinal rows of pits so arranged that one pit occurs in each concave bend of the wall. On the middle portion of the mature glume each of these pits is so large that it fills completely the bend of the wall, and in addition has a thickened border, half of which coincides with the cell wall, thus giving the tissue a lacelike appearance. This structure is optically delusive, the pit borders often appearing to be the cell walls, but is resolved by careful focusing and comparison with the tissue in earlier stages of growth. In addition to these MICROSCOPIC STRUCTURE OF WEED SEEDS 507 Fic. 384. Green Foxtail (Setaria viridis). I, spikelet with ripe fruit. g', lower empty glume; g%, upper empty glume; gf, glume, and p', palet of the staminate flower; ef?, glume, and p?, palet of fertile flower; c, caryopsis; b, bristles. II and III. caryopsis enclosed by flowering glume and palet. X 8. (After Winton, Conn. Agr. Exp. Sta.) elongated cells, pairs of short cells, one isodiametric, probably a hair-scar, the other more or less crescent-shaped, occur here and there, and less frequently stomata and thin-walled one to three- jointed hairs. 2. Mesophyl. Only about the nerves and the basal Fig. 385A. Fig. 385B Fic. 385. Green Foxtail. A. Outer epidermis of the staminate flower; I, at the edge; II, in the middle. X 300. B. Outer epidermis of the glume of the fertile flower, showing the wrinkled central portion and the smooth edge. (After Winton, Conn, Agr. Exp. Sta.) 508 WEED FLORA OF IOWA portions of the glumes is this coat evident. It has no diagnostic importance. 3. The Inner Epidermis is composed of elongated cells with straight walls. Palet of Staminate Flower. Within the glume of the staminate flower is the palet, a hyaline scale only 1 mm. or less long with a notch at the end. In general structure, it is much the same as the other thin envelopes, but the cell walls are thinner. 1. Outer Epiderms. The narrow, elongated cells are wavy in out- line, but pits are lacking or are indistinct. Isodiametric cells are thin-walled; jointed hairs also occur. 2. Inner Epidermis. Except at the base, where traces of mesophyl are sometimes evident, the inner epidermis immediately underlies the outer epidermis. Glumes and Palet of Perfect Flower. Both the glume and the palet of the fertile flower closely envelop the grain at maturity, the former being strongly convex, the latter flat except on the edges, which clasp about the caryopsis. At the time of flowering these envelopes are thin and of a green color, but at maturity they are coriaceous, silicified and of a brown or mottled color. Under the lens, numerous transverse wrinkles are evident on the glume and on the middle or flat portion of the palet, the lateral portions of the latter which clasp the caryopsis being smooth and shining. 1. Outer Epidermis. Throughout the glume and on the middle portion of the palet, the cells are isodiametric or moderately elon- gated and are arranged not only in longitudinal rows but also in irregular transverse rows, the wrinkles being formed by the out- bending of the cells at the end walls and the inward bending half way between. At the time of flowering, it may be seen that at the outer surface the end walls are sinuous and the side walls are compoundly sinuous, but farther inward the end walls are nearly straight and the side walls are simply, not compoundly sinuous. At the end of each cell nearest the apex of the envelope, a cuticular wart bearing a group of pits is usually evident, par- ticularly on the palet. About these warts the adjoining end walls are more or less curved and the side walls are not so deeply sinuous. At maturity the cell cavity beneath the wart is conspicuous (on the palet nearly circular), but at the other end of the cell is nar- row or not evident at all owing to the encroachment of the strongly thickened walls. The cell contents during the early stages of devel- opment are colorless, but later on usually become dark brown. The epidermal cells on the lateral or smooth portions of the palet which MICROSCOPIC STRUCTURE OF WEED SEEDS 509 Fic. 386. Green Foxtail. Outer epidermis from the middle of the glume of the fertile flower. I, outer surface, and II, inner surface soon after blooming. III, outer surface when in fruit. X 300. (After Winton, Conn. Agr. Exp. Sta.) elasp about the caryopsis are longer, narrower, and less complex than those already described. At maturity the wrinkles are usually from 0.03 to 0.06 mm. apart. 2. The Hypodermal Fibers may be readily isolated by treatment on the slide with caustic alkali. They vary in length up to 0.6 mm. and are often toothed at margin. 3. Mesophyl. Rectangular parenchyma cells without intercellular spaces make up this layer. Numerous chlorophyl granules are present at the time of flowering. 4. The Inner Epidermis is com- 510 WEED FLORA OF IOWA posed of rectangular cells resembling those of the mesophyl. Both of these layers become more or less obliterated at maturity and are of no diagnostic importance. Pericarp. The ventral side is flat and has a darker colored spot, the remains of the hilum, near the base. Extending half way from the base to the apex on the dorsal side is a groove, which marks the position of the embryo. Vogl describes minutely the histology of the caryopsis of common millet (Panicum miliacewm L.) and Fic. 387. Green Foxtail. Outer epidermis from the edge of the glume of the fertile flower. X 300. (After Winton, Conn. Agr. Exp. Sta.) states that German millet (Setaria panis Jessen) has practically the same structure. I find that his description applies also to the caryopsis of both green and yellow foxtail. 1. Epidermis. As in the outer epidermal layers of the floral envelopes the cells are elongated and wavy in outline. On the dark colored spot already referred to, the epidermal cells are more or less rectangular. 2. The Cross-Cells are similar to the tube-cells in form but are usually shorter, broader, and more irregular in shape. 3. Tube- Cells. These are 0.002 to 0.004 mm. wide and often reach the length of 0.3 mm. a. Nucellar or Hyaline Layer. After treatment with alkali, this layer is clearly seen in surface view. The cells are of large size and have beaded walls. b. Endosperm. 1. Alewrone Layer. The cells vary in diameter from 0.01 to 0.02 mm. MICROSCOPIC STRUCTURE OF WEED SEEDS 511 2. Starch-Cells. Polygonal starch granules with conspicuous hilums fill the parenchyma cells of the endosperm. In the outer layers they are from 0.004 to 0.008 mm. in diameter but farther inward they reach the maximum diameter of 0.018 mm. After dissolving the starch with potash, there remains a network of threads con- taining conspicuous granules. In this respect, however, this fruit cannot be distinguished from the fruits of S. glauca Beauv., S. panis Jessen, Panicum miliaceum L. (see Vogl) and all the other species of Panicum which I have examined. Detection m Powder Form. The membranous glumes with pores in the bends of Fic. 388. Green Foxtail. Transverse section of caryopsis. F, pericarp con- sisting of the epidermis ep and the tube-cells sch; N, nucellar layer; H, endosperm consisting of the aleurone-cells al and the starch cells s. X 300. (After Winton, Conn. Agr. Exp. Sta.) the walls and the coriaceous, transversely wrinkled, more or less spotted, envelopes of the fertile flower with compoundly sinuous, thickened cell walls are highly characteristic of both green and yellow foxtail. These tissues are usually present in all stages of development. The fruit elements are like those of common millet and German millet. Treatment with caustic alkali brings out the structure of the fruit coats and nucellar layer, and serves to dis- tinguish this fruit from the common cereals. The starch is hardly distinguishable from the starch of bindweed. Dr. A. L. Winton describes the microscopic character of yellow foxtail (Setarta glauca Beauv.) as follows: The fruit of this species is larger than that of green foxtail, the envelopes are also proportionately larger (with the’ exception of the upper empty glume which is but half the length of the 512 WEED FLORA OF IOWA spikelet) and the wrinkles on the glume of the fertile flower are more pronounced. In microscopic structure the fruits of the two species are identical. The floral envelopes are also much alike, the only distinction being in the distance apart of the wrinkles on the mature flowering glumes. In green foxtail this distance is usually from 0.03 to 0.06 mm., but in yellow foxtail it is often from 0.08 to 0.12 mm. Since this distinction does not apply to the mature glumes and since the wrinkles on the palets of the two species are practically the same, it is often difficult to identify the species in ground mixtures. Fortunately, identification of the genus is all that is usually required. Fic. 389. Green Foxtail. Caryopsis in surface view. Significance of letters. same as in Fig. 388. (After Winton, Conn. Agr. Exp. Sta.) Broom Corn (Andropogon sorghum var. technicus Koern). Dr. A. L. Winton describes the microscopic structure of the pericarp and testa of broom corn as follows: “As Empty Glumes.—Both glumes are from 4 to 6 mm. long, equal- ing and closely enveloping the fruit. They vary in color from yellow-brown to red-brown. The soft hairs, which nearly cover the outer surface, are loosely attached and most of them are re- moved during the threshing and cleaning of the seed, leaving the glumes smooth and shining. 1. The Outer Epidermis consists of strongly sclerenchymatized cells several times as long as they are broad, with wavy contour, interspersed here and there with iso- diametric hair-scars, each accompanied by a crescent-shaped cell MICROSCOPIC STRUCTURE OF WEED SEEDS 513 Fig. 390. Broom-corn. Fruit with chaff. r, two staminate spikelets; g1, lower empty glume; g2, upper empty glume; g%, glume of rudimentary flower; gf, flowering glume with awn; p, palet; ec, caryopsis. X 4. (After Winton, Conn. Agr. Exp. Sta.) with granular contents. The hairs, which are almost invariably detached in preparing the mount, if not in cleaning the seed, are often 1.0 mm. long and 0.012 mm. broad in the middle but taper toward both ends. Invariably the lumen is much broader than the walls. 2. The Hypoderm Fibers, of which there are several layers, have thick walls and narrow cavities. They vary in length Kaan pry ADwe™ ) y b ~. ECR ae eras Seca Vie oe Lh iso, en am Ores RUN ere se aw £Y5 rey pe iep.---- eae ue ep. ---+ SER hy ae Joie mes. --- os Blue Qeaas Seis sere) ah zl tp | Stee sheen orcad a |. sch- CRY 3 S~-~__ te rea) A al Sc seas z Fic. 391. Broom-corn. Transverse section of caryopsis and an empty glume. Sp, empty glume, consisting of the outer epidermis aep, the fiber layer f, the spongy parenchyma p, and the inner epidermis iep; g, bundle; sto, stoma; Fs, pericarp, consisting of the epidermis ep with the cuticle ec, the hypoderm hy, the starchy mesocarp mes, the cross-cells q, and the tube-cells sch; N, nucellar or hyaline layer with swollen inner walls s; H, endosperm, consisting of the aleurone layer al and the starch-cells with starch granules st and proteid network a. X 160. : (After Winton, Conn. Agr. Exp. Sta.) 33 514 WEED FLORA OF IOWA up to 0.5 mm. or more. 3. Spongy Parenchyma. As seen in sur- face view, the cells of this layer are more or less rectangular with circular intercellular spaces and resemble those of rice and barley glumes. 4. Inner Epidermis. In cross section this layer is not readily studied since the radial walls are usually collapsed; but in surface preparations, the large elongated cells, often 0.15 mm. long and 0.05 mm. wide, interspersed with stomata and hairs, are clearly displayed. Thin Glume.—Within the lower or first glume and nearly equal- ing it in length, is the third or thin glume, the remnant of an abortive flower. This glume is exceedingly thin and membranous and bears numerous hairs, particularly in the margin. 1. Outer Epidermis. In general form the cells are similar to those of the outer epidermis of the thick glumes, but are narrower and much thinner-walled. The marginal hairs are long (often 0.5 mm.) single-celled and pointed; but on the surface shorter hairs, with two or three joints and blunt ends, also occur. Both of these Fig, 392A Fig. 392B Fic. 392. Broom-corn. A. aep, outer epidermis and f, fiber of an empty glume in surface view. X 300. B. p, spongy parenchyma and iep, inner epidermis of an empty glume in surface view; sto, stoma; h, hair. X 300. (After Winton, Conn. Agr. Exp. Sta.) MICROSCOPIC STRUCTURE OF WEED SEEDS 515 forms have exceedingly thin walls. 2. The Inner Epidermis is dis- tinguished from the outer by the straight walls and almost entire absence of hairs. Flowering Glume.—The fourth or flowering glume, situated be- tween the upper or second glume and the grain, is also mem- branous and bears an upwardly barbed awn 5 to 7 mm. long. This awn, with the larger part of the flowering glume, being readily de- tached by threshing, it is seldom found in the grain on the market. Fic. 393. Broom-corn. Glume of rudimentary flower in surface view. aep, outer epidermis with h, one-celled hair and h, two-celled hair; iep, inner epidermis. X 300. (After Winton, Conn. Agr. Exp. Sta.) Palet—tThis is membranous and hairy like the third or ite glume, but is much smaller. Pericarp.—The grain or caryopsis is about 5 mm. long and from 2 to 3 mm. wide, tapering to a blunt point at both ends. It varies in color from yellow-brown to red-brown. Harz, Hassack and particularly Mittlacher have described so fully the histological elements of the caryopsis, that only a brief description, essential for a clear understanding, need here be given. 1. Eyidermis. The cells are longitudinally extended and have thick wavy side walls, with more or less distinct pores. Hassack has noted that the cuticle is of uneven thickness, due to minute granules or crystals, which may be seen in either section or sur- 516 WEED FLORA OF IOWA FIG. 394. Broom-corn. Layers of the pericarp in surface view. Significance of letters same as in Fig. 391. X 160. (After Winton, Conn. Agr. Exp. Sta.) face view. 2. The Hypoderm consists of from one to three layers of cells, with walls somewhat thinner than those of the epidermis. 3. Starchy Mesocarp. Several layers of thin-walled parenchyma cells, filled usually with small round or rounded polygonal starch granules seldom over 0.006 mm. in diameter, make up this coat. In all the varieties here described the starch appears during the early stages of growth and persists until the fruit nearly or quite - reaches full maturity. As the caryopsis, even when nearly mature, is intensely green owing to chlorophyll grains in the outermost layers of the mesocarp,. it may be inferred that this starch is a direct product of photosynthesis in the pericarp. So far as I have observed, the presence or absence of a starchy mesocarp in the erain at the time of harvest is not a definite varietal peculiarity, but is dependent on the ripeness of the fruit or other conditions. Some kernels of the same variety may possess it, while others show only empty, obliterated cells. Whether or not the starch is present in a given seed may often be determined by careful scraping and observation with the naked eye. 4. Cross-Cells. These cells are usually long and narrow, being distinguished from the tube-cells only by their transverse arrangement. Near the extremities of the seed they are, however, shorter and of more irregular shape. 5. Tube-Cells (sch). The cells of this layer lie at right angles to the _ eross-cells. They are about 0.005 mm. wide and often reach a length of 0.20 mm. a. Nucellar or Hyaline Layer. This layer is frequently 0.05 mm. thick. The outer radial walls are thin, but the inner wall is greatly swollen. In surface view the large cells MICROSCOPIC STRUCTURE OF WHED SEEDS 517 are conspicuous, not only because of their size, but because of their yellow or brown color. b. Endosperm. 1. Aleurone Layer (al). The individual cells of this layer are characterized by their great variation in size (0.01 to 0.04 mm. in diameter) and form. 2. Starch-Cells (st). In the outer layers the starch granules, if present, are much smaller than in the interior of the seed, where they sometimes reach a diameter of 0.083 mm. They are usually sharply polygonal, with a distinct hilum and radiating fissures. The starch is surrounded by small protein granules, forming a network which is especially evident after removing the starch by reagents. In some specimens, one or more of the outer cell layers are filled with these protein granules to the complete exclusion of the starch. ) Sandbur (Cenchrus tribuloides L.). The pericarp is divided into two portions. The cells of the outer portion are thick-walled and short; of the inner portion, elongated, thick-walled and fusiform. The testa consists of one to two rows of thin-walled cells, much compressed. The aleurone cells are much larger than in Panicum and Setaria, thick-walled, densely filled with protein grains. Walls of the starch cells thick- ened; starch grains larger and loosely arranged. Timothy (Phlewm pratense L.). The testa and pericarp are dark colored. The epidermal cells are thin-walled, elongated, sometimes slightly irregular. The testa consists of several rows of thick-walled dark brown cells much longer than broad. The aleurone layer consists of a single row of cells relatively thin-walled, somewhat variable in size, solidly packed with aleurone grains. The nucellus is very evident as a remnant in some places. The cells of this layer are thick- walled, clear and colorless. The starch cells are much larger than the aleurone, and contain angular starch grains. Cheat or Chess (Bromus secalinus L.). Dr. A. L. Winton describes the microscopic structure as follows: Flowering Glume.—The structure throughout is much the same as in darnel, but the cells of the outer epidermis are much more conspicuously thick-walled, and the wavy-walled cells throughout much longer than broad. The circular cells also have wavy walls. The cells on the margins, interspersed with lance-shaped hairs, are the same as in darnel. 518 WEED FLORA OF IOWA Fic. 395. Chess (Bromus secalinus). Outer epidermis of flowering glume in surface view. X 160. (After Winton, Conn. Agr. Exp. Sta.) Palet—The flowering glume and palet of chess are very similar in structure, but the outer epidermis of the latter is barbed on the keel, the stiff hairs often reaching 45“ in length. Pericarp (F).—The pericarp consists of two layers with rudi- ments of another layer in parts. 1. The Epidermal Cells (ep) are large, elongated-polygonal, and have thin, non-porous walls. 2. Mesocarp. As a rule, the cross-cells immediately underlie the Fic. 396. Chess. Transverse section of fruit. F, pericarp consisting of epidermis ep, and cross-cells q; S, testa; N, perisperm; H, endosperm con- sisting of aleurone layer al, and starch-parenchyma st. X 160. (After Winton, Conn. Agr. Exp. Sta.) epidermis, but occasionally traces of the mesocarp are evident. 3. Cross-Cells (q). Whether this layer corresponds with the cross- cells of the tube-cells of other grasses, I have been unable to decide. MICROSCOPIC STRUCTURE OF WEED SEEDS 519 The tissue is made up of irregular spongy-parenchyma cells, usually transversely elongated with large, round or elongated inter- cellular spaces. The Testa (S).—Consists of one layer of elongated brown cells 10-20 wide. Perisperm (N).—This layer is enormously developed. As may be seen in cross section, the cells are 40“ thick, but the walls are so swollen as to almost entirely obliterate the cavity. After soaking for some time in 1 per cent soda solution they are evident in sur- face view. Endosperm.—l1. The Alewrone Layer (al) is not of especial in- terest. 2. The Starch-Parenchyma (st) is remarkable for the thick- ness of the cell walls (often 10 thick) and the elliptical starch grains 3-204 in diameter. With proper illumination each grain may be seen to have an elliptical hilum. Fic. 397. Chess. Elements of fruit in surface view. Significance of letters same as in Fig. 396. (After Winton, Conn. Agr. Exp. Sta.) Wild Barley or Squirrel-tail Grass (Hordeum jubatum L.). The grain is adherent to the palet. The epidermis consists of thick-walled, tangentially elongated ‘cells, most of which are longer than broad. In part the cells are developed into short conical trichomes. The underlying cells are thick-walled with prominent pore canals. The remainder of the adhering palet consists of thin- walled cells much larger than broad. The pericarp as well as the 520 WEED FLORA OF IOWA testa is but slightly developed. In some cases the underlying parenchyma cells are not clearly defined. The blackish pigment is found in the internal part of the palet; some also occurs in peri- carps and the aleurone layer. The pericarp consists of one or two rows of rather thin-walled, tangentially elongated cells. The testa is reduced to a single layer of cells longer than broad. The nucellus is nearly absent except in the groove. It consists of a single row of thin-walled, colorless, compressed cells. In the groove several rows of cells occur. The aleurone in the specimens studied is made up of a single row of cells. The cells of the starch layer are larger than those of the aleurone. Darnel (Lolium temulentum L.). Dr. A. L. Winton describes its microscopic structure as follows: The Flowering Glume is 6-8 mm. long, equaling or exceeding the caryopsis. It is obscurely five-nerved, lobed at the end, and bears an upwardly barbed awn often 15 mm. long. Like the glumes of barley, oats, and other cereals, it consists of four coats, some of which, however, are lacking on the margins and at the end. 1. The Outer Epidermis differs greatly in structure in different parts of the glume. At the margins (as is clearly shown in Fig. 1 by Moeller*), it consists of straight-walled, elongated cells inter- spersed here and there with short lance-shaped hairs. On the ereater part of the surface, however, the cells, as in barley and some other cereals, are of three kinds: first, cells of wavy outline, lic. 398. Darnel (Lolium temulentwm). Margin of flowering glume showing lance-shaped hairs. X 300. (After Winton, Conn. Agr. Exp. Sta.) MICROSCOPIC STRUCTURE OF WEED SEEDS 521 into which the straight-walled cells at the margin pass; second, circular cells corresponding to the conical hair-cell of barley ; third, exceedingly short, more or less crescent-shaped cells. Near the ‘ margins and on the veins, where they alternate with stomata, the cells of wavy outline are elongated; but in other parts they are very short, often being broader than long. Although thick-walled, the walls are transparent, whereas the middle lamella is con- spicuous, giving the impression of thin-walled cells. Pores are few and inconspicuous. Near the margin the circular cells are small and are usually accompanied by crescent-shaped cells which often exceed them in size. On the greater part of the glume, however, the circular cells are much larger, often being 70# in diameter. Numerous pores are conspicuous, both in the radial and tangential walls. Often one, sometimes two, crescent-shaped cells accompany a circular cell. Characteristic of this coat are the short, wavy Fic. 399. Darnel. Middle portion of flowering glume. X 160. (After Winton, Conn. Agr. Exp. Sta.) cells and the numerous circular cells, the latter frequently exceed- ing in area the former. 2. Hypoderm. The fibers in this layer are much the same as in cereals. Fibers of similar structure also make up the ground-tissue of the awn. 3. Spongy-Parenchyma. The elements are more or less rectangular in shape, like those of the corresponding layer of barley, and are readily distinguished from the star-shaped elements of oats. 4. Inner Epidermis. This layer is made up of thin-walled cells and stomata, and is of no diagnostic importance. 522 WEED FLORA OF IOWA Palet.—The two-keeled palet is about the same size as the flower- ing glume, but is of a thinner texture, owing to the absence of a well developed hypoderm layer. The Outer Epidermis is much the same as that of the flowering glume, except that it is barbed on ~ the keels with rigid thornlike hairs 150“ or less in length, as is shown in Fig. 3 by Moeller. Fie. 400. Darnel. Keel of palet showing outer epidermis with hair h, and hypoderm fibers f. X 160. (After Winton, Conn. Agr. Exp. Sta.) The Pericarp (F').—Consists of four coats, of which only two, the epidermis and cross-cells, are fully developed. 1. Epidermis (ep). Cross sections of the mature seed show that this layer consists of collapsed, moderately thick-walled cells, which are best studied after heating with potash. Seen in surface view, the cells at the apex of the seed are nearly isodiametric, but at other parts are elongated. The walls are indistinctly beaded. 2. The Mesocarp (m) is not developed on all parts of the seed, but is conspicuous on the angles. The cells vary greatly in shape and size, some being ir- regularly isodiametric, others transversely elongated, resembling the cells of the next layer. 3. Cross-Cells (q). Especially striking are the cells of this layer, which resemble the cross-cells of barley. As has been noted by Moeller, the radial walls appear indistinctly beaded, but this is evident only under favorable conditions. 4. Tube-Cells, spongy-parenchyma, and various intermediate forms, make up the interrupted inner layer of the pericarp. Testa (S).—The cells are for the most part elongated and are often diagonally arranged with reference to the axis of the fruit. In MICROSCOPIC STRUCTURE OF WEED SEEDS 523 42m” 4 Fic. 401. Darnel. Transverse section of fruit. F, pericarp consisting of epidermis ep, mesocarp m, cross-cells q, and tube-cells sch; S, testa con- sisting of outer layer a. and inner layer i; N, perisperm; f, fungus layer; HE, endosperm consisting of aleurone layer al, and starch-parenchyma st. x 160. (After Winton, Conn, Agr. Exp. Sta.) transverse sections this coat often separates from the pericarp on the one hand and the perisperm on the other. Examined in water, only one cell layer (the inner) is evident; but successive treat- ments with 5 per cent potash, dilute acetic acid and chlorzine iodine, bring out two layers. 1. The Outer Layer (a) is made up of thin-walled cells with cuticularized outer walls. Treated as above described, the cuticle is colored yellow-brown, the radial and inner walls, blue. 2. The Inner Layer (1) is not only thicker than the outer, but the cells are thicker-walled and, in addition, swell greatly with potash. These swollen walls are stained deep blue by chlorzine iodine, thus differentiating them from the yellow-brown cuticle on the inner wall. Perisperm (N).—Characteristic of this seed is the nucellar-coat, consisting usually of two cell layers. In cross section these cells are rectangular with swollen walls; in surface view, as maybe seen after soaking for a long time in dilute potash, they are:ir- regularly polygonal or more or less elongated. Shines Fungus Layer (F').—In most specimens a layer of fungus-threads 20 thick is present between the perisperm and the aleurone layer. So commonly is this fungus present in darnel grown in Europe, that it is of no little value in identifying the grain; but it remains to be determined whether in California, where the plant is a pest 524 WEED FLORA OF IOWA in wheat fields, the fungus is also a common accompaniment. After treatment with potash this layer is stained bright yellow by zine chloride of iodine. Endosperm.—1. The Aleurone Cells (al) vary from less than 20 to 40“ in diameter. 2. Starch-Parenchyma (st). The thin-walled cells contain small polygonal grains 3 to 7# in diameter. The in- dividual starch grains are not distinguishable from the grains of rice and oats, and like the latter, often occur in aggregates of various sizes. : Nez Ps ne : Fic. 402. Darnel. Elements of fruit in surface view. Significance of letters same as in Fig. 401. X 160. (After Winton, Conn. Agr. Exp. Sta.) URTICACEAE, NETTLE FAMILY. Hemp (Cannabis sativa L.). The smooth crustaceous achene consists of an outer epidermal layer of thick-walled cells with wavy outline, the thickness of the radial walls varying somewhat. This is followed by a layer of loosely arranged parenchyma cells, and a layer of cells with brown contents. A layer of small cells spoken of as the dwarf cells by Winton, may also be made out in some sections. The palisade layer consists of thick-walled cells with pore canals, the walls having a wavy outline. The cell cavity is very much reduced. The testa is very thin and consists of thin-walled elongated paren- chyma cells, the second layer of spongy parenchyma. The com- MICROSCOPIC STRUCTURE OF WEED SEEDS 525 pressed nucellus follows. The endosperm consists mostly of a single layer of cells, the embryo of small epidermal cells and the elongated palisade parenchyma on the upper face of the cotyledon; cells of embryo and endosperm contain fat and protein. Nettle (Urtica gracilis Ait.). The pericarp of the small achenes consists of an outer epidermal layer of rather large cells with exterior walls thickened, and underneath several layers of loosely arranged parenchyma cells. The testa and nucellus are compressed; cells of the embryo squarish, containing fat and protein grains. Fic. 403. Microscopic structure of the seeds of the Nettle family. (Urticaceae). I. Hemp (Cannabis sativa). II. Common nettle (Urtica gracilis). ep=epidermis, thick-walled short cells, underneath followed by a pigment layer, thinner-walled parenchyma cells and a layer of thick-walled cells. pal= palisade cells. p=parenchyma cells with the pigment. t=testa. n= nucellus. em=embryo. (Drawings by L. H. Pammel and Charlotte M. King.) POLYGONACEAH,* BUCKWHEAT FAMILY. Miss E. Sirrine finds, ‘‘In the seed coats of the family, the palisade portion constitutes the outer part of the achenium; this is followed in most cases by the testa consisting of several layers of eells varying, however, in some cases; they are quite regular in form and in some instances are of a dark color. In the mature palisade cell, the cavity is present; this varies greatly in the dif- ferent genera; in some cases occupying nearly the entire cell, in others small and irregular.’’ *A study of the microscopic structure of the achenes is largely based on Miss Emma Sirrine’s work and upon that of Dr. Winton. 526 WEED FLORA OF IOWA Sorrel or Sour Clover (Rumesx acetosa L.). This species has very small palisade cells, rectangular in shape and with a small cell cavity which occupies only a small portion of the lower end of the palisade cell. No canals or irregularities of cell cavity.. The cell is light in color while canal is darker. The sub-palisade portion is composed of small round cells of a variable number. The endosperm is composed of irregularly ar- ranged cells. Measurements: whole seed coats, 38.34; palisade cells, 23"; sub-palisade, 13.2. HULL ee rt mat OE =| unum: UNE } TUM pee twin mM mn Fic. 408. Black Bindweed. Seed in surface view. Significance of letters same as in Fig. 406. (After Winton, Conn. Agr. Exp. Sta.) Detection in Powder Form.—Characteristic of this fruit are the papillae on the outer epidermis of the calyx and the epicarp with sinuous cell walls and rows of warts. The outer epidermal cells of the testa are sinuous in outline, like those of buckwheat, but, unlike the latter, are commonly elongated. Although the cross- cells are morphologically the same as the spongy parenchyma of buckwheat, they resemble more nearly in structure the tube-cells of the cereals.. The starch granules are not characteristic and the network obtained after treatment with caustic alkali serves merely as an indication that the seed belongs to a Polygonaceous plant. Buckwheat (Fagopyrum esculentum Moench.). The achenium consists of elongated epidermal cells with thick- ened walls, underneath similarly elongated thick-walled sclerotic parenchyma cells with pore canals. This layer contains the pig- ment. The testa follows the pericarp and is differentiated into an epidermal layer of yellowish brown walls, followed by smaller thin-walled, parenchyma cells. The testa is much compressed; the albumen consists of an outer aleurone layer of small cells followed by larger cells. The albumen cells contain compound starch grains. 532 WEED FLORA OF IOWA 4) b) Fic. 409. Black Bindweed. Surface view of the pericarp from below. X 160. Significance of letters same as in Fig. 406. (After Winton, Conn. Agr. Exp. Sta.) CHENOPODIACHAH, GOOSEFOOT FAMILY. Goosefoot or Lamb’s Quarters (Chenopodium album L.). Harz has given an account of the structure of Beta and Spinacia. The structure of the seed of lamb’s quarters is somewhat different. The thin utricle consists of an epidermal layer of somewhat wavy eells and an indistinct layer underneath. The testa contains the brownish pigment. The cells are thick-walled; the nucellus is com- pressed. The cells of the endosperm are large and contain an abundance of small starch grains. Russian Thistle (Salsola kali (.) var. tenutfolia G. W. F. Meyer). The seed is without endosperm; the embryo fills the seed, and is coiled in a conical spiral. The seed coat consists of two parts, a layer of three rows of elongated cells followed by a layer of three rows of irregular parenchyma cells. One side of the seed coat is wider than the other and these cells contain an abundance of calcium oxalate crystals. The cells of the outer layer of the MICROSCOPIC STRUCTURE OF WHEHD SEEDS 533 embryo are somewhat elongated; the remainder are nearly as broad as long; these contain no starch but albuminoids and fat. Fic. 410. Microscopic structure of seeds of the Chenopod family. (Chenopodia- ceae). I. Russian Thistle (Salsola kali var. tenwifolia). II. Lamb’s Quar- ters (Chenopodium album). us ep=epidermis. p=parenchyma cells. t=testa. n=nucellus. en=endosperm. (Drawings by L. H. Pammel and Charlotte M. King.) NYCTAGINACEAE, FOUR-O’CLOCK FAMILY. Wild Four-o’clock or Umbrella Plant (Oxybaphus nyctagineus (Mx.) Sweet). The nutlike fruit of the wild four-o’clock or umbrella plant is somewhat hairy; the outer portion of the pericarp is made up of thick black-brownish cells from 6 to 9 rows. The epidermal cells are smaller, some of the cells elongated into one-celled thick-walled trichomes. Adjacent to the testa are bundles of thick-walled, sclerotic cells, the outer layer of cells of the testa consisting of mucilaginous cells with colorless walls. This is followed by a second layer of thin-walled parenchyma cells. The nucellus con- sists of an indefinite granular mass, followed by the endosperm, and the thin-walled cells of the embryo. | CARYOPHYLLACHAH, PINK FAMILY. Soapwort or Bouncing Bet (Saponaria officinalis L.). The kidney-shaped, black, roughish seeds consist of tangentially elongated, thick, black, rough cells, the outer layer being brownish colored, while the cell cavity is red or blackish brown. . The inner walls of the testa are thinner, the cells are elongated and the inner seed coat is much compressed. The nucellus is indistinguishable from the inner seed coat. This is followed by the endosperm, 534 WEED FLORA OF IOWA Fig. 411. Wild Four-o’clock (Oxybaphus nyctagineus). The sclerotic paren- chyma, parenchyma and epidermis belong to the pericarp. ep=epidermis. p=parenchyma. scl=sclerotic parenchyma. t=testa. t=tri- chome. en=endosperm. r=ribs. (Drawing by L. H. Pammel and Charlotte M. King.) consisting of an outer tangential layer with granular contents. The aleurone layer and remaining portion of the endosperm cells consist of large rather thin-walled cells with compound starch grains and protein. Slender Catchfly (Silene antirrhina L.). The small kidney-shaped seeds are rough and brownish in color. The outer epidermal walls are thick, brownish black in color. The underlying parenchyma cells are tangentially elongated with near- ly colorless walls, This is followed by an indefinite layer con- MICROSCOPIC STRUCTURE OF WEED SEEDS 535 sisting of an inner seed coat and a nucellus. The endosperm con- sists of thin-walled parenchyma cells with granular contents con- sisting of starch grains and protein. Zi Fic. 412. Microscopic structure of some seeds of the Pink family (Cary- ophyllaceae). I. Bouncing Bet (Saponaria officinalis). II. Catchfly (Silene antirrhina). III. Virginia Catchfly (Silene virginica). ep=epidermis. p=parenchyma. n=nucellus. h=hypoderm cells. en=endo- sperm. (Drawings by L. H. Pammel.) The seeds of Virginia Catchfly (Silene virginica) are irregular. The outer walls are greatly thickened, irregular, the underlying parenchyma cells being slightly elongated. This layer is followed by the nucellus and the aleurone layer and remaining endosperm cells containing starch grains and protein material. Silene inflata is described and figured by Harz*. *Samenkunde 2 :10-79. 536 WEED FLORA OF IOWA RANUNCULACEAE, CROWFOOT FAMILY. Common Buttercup (Ranunculus abortivus L.). The greenish achenes of the buttercup consist of three distinct layers. The outer epidermal is of elongated, thickish black cells. The cells containing the brown pigment are yellowish brown. The underlying layers of three or four rows of seeds are similar but larger and paler in color. This layer is followed by a thick- walled sclerotic layer containing three or four layers of thick- walled cells with pore canals. The testa consists of an outer layer of thin-walled parenchyma cells; the inner layer consisting of sclerotic parenchyma cells filled with pore canals. The endosperm cells are large or thin-walled and contain protein material. Harz has given an account of the structure of R. arvensis*. ic. 413. Microscopic structure of the seed of Buttercup (Ranunculus aborti- VUS). ep=epidermis. p=parenchyma. scl=sclerotic parenchyma. en=endosperm. (Drawing by L. H. Pammel and Charlotte M. King.) PAPAVERACEAE, POPPY FAMILY. Common Poppy (Papaver somniferum L.). Prof. A. L. Winton describes the seeds of this plant as follows: Testa (S).—Cross sections are prepared after soaking the seed in water that may be cleared with chloral or alkali. After soaking the whole seed for about twenty-four hours in 1 per cent sodium hydrate solution, the first four layers readily separate from the *Landw. Samekunde 2 :10-64. MICROSCOPIC STRUCTURE OF WEED SEEDS 537 fifth. Subsequent treatment with hydrochloric acid dissolves out the calcium oxalate, and staining with chlorzine iodine or safranin renders the outer layers more distinct. 1. The Hpidermal Cells (ep) are polygonal and of enormous size, corresponding to the net- work on the seed. As appears in cross section, the cells are col- lapsed except in the neighborhood of the radial walls. In surface view the radial walls are sinuous and thin, what are often con- sidered the thick dark walls of this layer being not the walls at all, but the ribs formed by the thickening of the second and third layers. This conclusion is consistent with Meyer’s and Hanausek’s figures of cross sections, also with Meyer’s drawings and Mach’s photomicrographs of surface preparations. The statement of Tschirch and Oesterle that the epidermis consists of elongated cells situated over the ribs, with large polygonal cells between, has since been corrected by the authors themselves. Doubtless they mistook some of the cells of the second layer for epidermis. Hanausek’s surface view, on the other hand, might convey the impression that the ribs were the cell walls, but his description and cross section clearly show their true nature. 2. Crystal Layer (k). On the Fic. 414. Poppy Seed (Papaver somniferum). Transverse section. s, testa consisting of epidermis ep, crystal layer k, fiber layer f, cross-cells q and netted-cells n; e, endosperm containing aleurone grains al. X 160. (After Winton, Conn. Agr. Exp. Sta.) ribs, the cells of this layer are more or less tangentially elongated, but between the ribs are isodiametric and polygonal, the elongated cells having longer radial walls than the others, thus contributing to the formation of the ribs. They contain fine, granular crystals ‘of calcium oxalate. Meyer has demonstrated that the blue color of the poppy seed is due, not to a blue pigment, but to the inter- ference of light by the crystals over the brown cells in the back- ground, and is the same phenomenon that causes the apparent blue color of the sky and the iris of the eye. As soon as these erystals are dissolved in hydrochloric acid, the seed appears brown. 538 WEED FLORA OF IOWA 3. Fiber Layer (£). The fibers of this layer are 15-40" broad and are parallel to the curved axis of the seed. Seen in cross section, this layer is thickest in the ribs, the walls throughout being distinctly thickened and stratified. In surface view they are rendered more distinct by chlorzine iodine. 4. Cross-Cells (q). The fourth layer consists of moderately thick-walled, transversely elongated, pointed cells arranged side by side in rows. The walls _ are impregnated with brown material. 5. Netted-Cells (n). Ow- ing to the netted-veined, colorless walls and the presence of deep brown contents, these cells are particularly striking. They are arranged transversely and often side by side in rows. The cell contents are insoluble in alkali and do not give the tannin re- action. Some authors designate the cells of this layer ‘‘Pigment cells,’ notwithstanding the fact that in the white poppy they do not contain pigment. Meyer, Tschirch and Oesterle, Vogl, and Hanausek describe an inner layer of thin-walled cells, but I am unable to find such a layer except in the vicinity of the hilum. Fic. 415. Poppy seed. Testa in surface view. Significance of letters same as in Fig 414. pig, pigment. X 160. (After Winton, Conn. Agr. Exp. Sta.) The Endosperm (E) contains aleurone grains up to 34 in the outer layers and 7 in the inner layers, each grain containing several globoids and crystaloids. Embryo.—In the cotyledons there is only one layer of palisade- cells and these cells are only slightly elongated. The aleurone grains are like those of the endosperm. MICROSCOPIC STRUCTURE OF WEED SHEDS 539 Prickly Poppy (Argemone intermedia Sweet). The blackish pitted seeds are difficult to study. They consist of nearly colorless epidermal cells, with thick outer walls and cuticle. The cells of the underlying layer are thick-walled, walls and contents blackish, difficult to make out on account of dense Fic. 416. Rocky Mountain Poppy (Argemone intermedia). ep=epidermis with thick outer walls. pi=pigment layer, n=nucellus, en=en- dosperm. : (Drawing by L. H. Pammel and Charlotte M. King.) pigment. The nucellus is compressed, colorless. The endosperm of large cells contains oil in large amounts, and protein. The structure seems to differ in a marked degree from Papaver as described by Harz and G. Kraus.. CRUCIFERAH, MUSTARD FAMILY. Virginia Peppergrass (Lepidium virginicum L.). The seed coats consist of three well defined layers. The outer or epidermal cells are tabulated, somewhat compressed. The cuticle forms a continuous layer over these. On the addition of water the epidermal cells elongate and form a mucilaginous mass, showing stratified layers. These are not difficult to make out when the specimen is mounted in water. The cell cavity is very much re- duced; that portion of the cell wall in contact with the cell cavity is differentiated from the outer cell wall substance. Long con- tinued addition of water causes the cuticle to break and the exterior becomes very irregular. The second layer is colored brown, the cell walls are considerably thickened laterally and project upwardly in the shape of cones. A section made through the ends of these seeds shows that the second layer is considerably more developed and there are evidences 540 WEED FLORA OF IOWA here of an indistinct layer between the first and second. The layer following this consists of thin-walled parenchyma cells, in some cases considerably elongated but in others short. The third layer is followed by the endosperm which consists of a layer of rather thick-walled parenchyma cells. These carry granular protein grains. This is followed by one or more layers of elongated cells, in which the cell cavity is very much reduced. These cells reach their highest development between the folds of the caulicle and cotyledon. The cells of the first layer of the embryo are smaller, quite uni- form in size and filled with protein grains and oil. Small Peppergrass (Lepidium apetalum Willd.). The cuticle forms a continuous layer over the epidermal cells, which are larger than in LD. virginicwm. On the addition of water the cell wall rapidly elongates, emitting a copious mucilage. The cell cavity is very much reduced but longer than in L. vwirginicum. It is surrounded by a denser, more or less differentiated part of the cell wall which is more yellow in color than the remainder of the cell wall. The second layer is of a yellow straw color and consists of very minute cells with small cell cavities. The cell walls of the third layer are strongly thickened, brown, and serve the same purpose as in the other species. The endosperm consists of thick-walled parenchyma cells. In the first layer of cells, the cell walls are very much larger and packed with protein grains. The other layers of endosperm consist of smal] elongated thick-walled cells with a small cell cavity. These attain their greatest development between the caulicle and cotyledon. In the embryo, the cells of the first row are isodiametric filled with protein grains and oil. The outer cells are elongated larger, and also densely packed with the same material. Shepherd’s Purse (Capsella bursa-pastoris (L.) Medic.). The seed coats attain their maximum development in the region of the caulicle. Cuticle covers the epidermal cells, the latter tabular, compressed but in the addition of water elongating, be- coming mucilaginous and showing stratification. The second and third layers are brown with thick cell walls. Fourth layer consists of endosperm, one layer of isodiametric cells filled with protein grains, followed by thick-walled cells reaching MICROSCOPIC STRUCTURE OF WHED SEEDS 541 Fic. 417. Microscopic structure of seeds of Mustard family (Cruciferae). I. Hedge Mustard (Sisymbriwm canescens). II. Small Peppergrass (Lepidium apetalum). To the right an enlarged epidermal cell with mucilaginous walls. ep=epidermis. p=parenchyma. em embryo. (Drawing by L. H. Pammel.) their greatest development between the cotyledon and caulicle. First row of cells of embryo. nearly isodiametric, filled with oil and protein grains. Others somewhat larger contain the same substances. Cotyledons incumbent. Central part of caulicle sepa- rated from the rest. Cells of caulicle very much larger than cells of cotyledons. Black Mustard (Brassica nigra Koch.). The cuticle covers the epidermal cells as a continuous layer; when mounted in alcohol the outer layer is very much compressed and shows very slight stratification; the cell walls expand and after it has been moist for a considerable time the cuticle breaks. Stratification is very evident on the addition of water. The second layer consists of rather thin-walled parenchyma. The cells of 542 WEED FLORA OF IOWA Fic. 418. Microscopic structure of some Cruciferous seeds (Cruciferae). I. False Flax (Cameélina sativa). II. Black Mustard (Brassica nigra). III. Common Mustard (B. arvense). IV. Winter Cress (Barbarea vulgaris). V. jPeppergrass (Lepidium virginicum). VI. Tumbling Mustard (Sisym- brium altissimum). VII. Shepherd’s Purse (Capsella bursa-pastoris). t=testa. p=parenchyma cells. n=nucellus. em=embryo. I, II, III, and IV. Hlongated palisade cells below the epidermal layer and parenchyma cells. To the right of I, epidermal cells after the addition of water. (Drawings by L. H. Pammel.) this layer differ greatly with reference to their size, being scarcely at all developed in places, in others nearly as large as the cells of the outer layer. MICROSCOPIC STRUCTURE OF WEED SEEDS 543 The third layer consists of thick-walled parenchyma cells, densely packed, radially elongated, sides presenting a cone-shaped appear- ance. Underneath this is a layer of thick-walled parenchyma cells which contain some coloring matter. The endosperm follows this layer. The first layer consists of thick-walled cells, densely packed with albuminous matter. The remaining cells vary in number, are much elongated, thick-walled with a small cavity; these cells extend down between the contiguous portions of the cotyledon or caulicle. The Embryo—tThe cells of the first layer surrounding the cotyledon or caulicle are smaller, filled with fat and protein grains. The remaining cells are larger, also filled with fat and protein grains. The central part of the caulicle shows a differentiation of the embryonic vascular portion, consisting of small cells. Charlock or Common Mustard (Brassica arvensis (L.) Ktze.). The outer layer of cells is compressed, tabular, with stratification not evident, and cuticle well developed, and forms a continuous layer over the outer cells; on the addition of water, the cell walls become mucilaginous, elongate, stratification becomes evident, the cuticle breaks, and an irregular surface is formed. The second layer is but slightly developed, made up of thin-walled parenchyma cells. The cells of the third layer are elongated and thickened laterally. These cells are much longer than in B. nigra and brown in color. The fourth layer consists of one to two rows of rather thin-walled cells carrying pigment. Endosperm consists of several rows of cells; first row nearly isodiametric, filled with protein erains. The three or four layers of cells following are thick-walled with a small cell cavity. Embryo.—First layer of cells nearly isodiametric, those follow- ing somewhat larger, filled with protein and fat grains. Fiedge Mustard (Sisymbrium. officinale Scop.). Cuticle covering the epidermal cells, the latter tabular, much compressed. On the addition of water the cell walls become mucilaginous with evident stratification. The cells of the second layer are brown and thin-walled, much compressed. On addition of choloral hydrate they expand. Third layer much darker than the second, thick-walled, followed by endosperm, cells elongated, filled with protein grains, followed by elongated thick-walled cells 544 WEED FLORA OF IOWA with a small cavity. These reach their highest development between cotyledons and caulicle. First row of cells of the embryo nearly isodiametric, filled with protein grains and oil. Tumbling Mustard (Stsymbrium altissimum L.). On the addition of water the cell wall of the outer seed coat becomes mucilaginous. Outer epidermal layer covered with cuticle, cells elongated, on the addition of water, walls become mucilaginous and show stratification. Cell walls of second layer thick, light brown, followed by endosperm of two layers of cells, first elongated, thick-walled. Cells of embryo as in S. officinale. Hairy Hedge Mustard (Sisymbriwm canescens Nutt.). The testa of the small brownish cells consists of an outer epidermal layer with thick walls. These cell walls become mucil- aginous on the addition of water. This is followed by a layer of several rows of brownish cells and a compressed layer, the nucellus. The cells of the outer row of the embryo are smaller than the underlying cells. Common Winter Cress or Yellow Rocket (Barbarea vulgaris R. Br.). First layer of outer seed coat not well developed, cells elongated in the direction of the seed. Cuticle covers the epidermal cells. On the addition of water a slight mucilaginous modification takes place. Second layer with thick lateral walls and quite large cell cavities, colored brown. Third layer of rather thick-walled parenchyma cells also colored brown, followed by endosperm as is usual in cruciferous seeds. False Flax (Camelina sativa (Li.) Crantz.). Seed coats consisting of four layers. The outer epidermal cells not much longer than wide, on the addition of water become mucilaginous and well stratified. On the addition of chloral hydrate stratification is more evident. The cell walls are differentiated into several layers. The second layer is not always developed. Cells of third layer with thick walls and brown pigment, followed by a narrow layer of thick-walled brown cells. The first row of cells of endosperm, rather thick-walled, filled with protein grains, the other layers of unequal development, cells elongated, thick- walled; followed by cells of embryo; these contain protein grains and fat. MICROSCOPIC STRUCTURE OF WEED SEEDS 545 ROSACHAH, ROSH FAMILY. White Avens (Geum canadense Jacq.). A greater part of the so-called seed is made up of the pericarp consisting of a layer of small epidermal cells with trichomes. The underlying parenchyma cells are large with numerous intercellular spaces. -The testa is thin, consisting of an outer pigment layer followed by several rows of thick-walled, colorless cells. Fic. 419. Microscopic structure of fruit of Avens (Geum canadense). ep=epidermis. p=parenchyma. per=pericarp. t=trichome. t'!=testa. (Drawing by L. H. Pammel and Charlotte M. King.) LEGUMINOSAE,* PULSE FAMILY. Rattlebox (Crotalaria sagittalts L.). The testa not strongly developed; endosperm 196” in thickness. From Nadelmann’s studies it appears that in Crotalaria verrucosa the horny endosperm is well developed, being four and one-half times as wide as the testa. The aleurone layer contains fat. and aleurone grains. The cells of the embryo contain protein and fat but no starch. *The descriptions here given are taken for the most part from a paper by L. H. Pammel, Trans. Acad. Sci. St. Louis, 9. 30 546 WEED FLORA OF IOWA Malpighian.—The cells are prismatic in surface view, 9.84 across, with five to six canals. In cross sections they are 84# long. The cuticle forms a continuous layer, with longitudinal canals pro- jecting into the cells; these extend down through the upper part of the cell wall and the cuticularized substance; this layer is lighter in color than the rest of the cell wall, and separates from the re- mainder of the cell in the form of a band. The narrow light line occurs close under the cuticularized layer. The cell cavity is narrow and gradually tapers upward; it contains some protein matter. The cell wall consists of cellulose. Osteosclerid.—The walls are thickened. Cells wide in the lower part and narrowing upwards, with large intercellular spaces. Cells contain protein. Nutrient— Consists of radially elongated cells, which are slightly compressed. Walls of medium ‘thickness, shghtly colored. Fic. 420. Microscopic structure of the testa of Rattle-box (Crotalaria sagittalis). ll=light line. m=malpighian cells. n=nutrient layer. o=osteosclerid. (Drawing by L. H. Pammel.) ‘Endosperm—Not strongly developed. The cells of the aleurone layer large, containing protein. Two layers follow this, the cells very much compressed, and somewhat radially elongated, of thick- walled cells. All of the cells contain protein and fat. Embryo.—Cells of outer row smaller; those adjoining the en- dosperm thicker walled, the inner part with thinner walls. The re- maining cells of the embryo larger. All of the cells filled with protein. Starch is absent. Black Medick (Medicago lupulina L.). Testa and endosperm well developed, 245“ in thickness on sides. More than half of this thickness consists of endosperm. MICROSCOPIC STRUCTURE OF WEED SEEDS 547 Malpighian.—Cells 40-424 in length. Cuticle slightly irregular; underneath the cuticle a light colored area with conical projec- tions, as in Melilotus, but somewhat more prominent. With chlor- iodide of zine this rapidly colors blue. It corresponds to the mucilaginous ‘‘membrana interna’’ of Mattirolo and Buscalioni, and, as shown by Schips, is chemically differentiated from the cuticle and remainder of the cell wall. The conical layer is highly refractive. The light line occurs below the conical layer and colors blue soon after the addition of chlor-iodide of zine. The cell cavity is broadest at the base, gradually tapering upward. A large chromatophore occurs at the base or near the middle of the cell cavity. In colored seed some pigment occurs in the cavity as well as considerable amounts in the walls. Small pore-canals occur in the upper part of the cell wall. The cell walls color blue more slowly with chlor-iodide of zine than the cuticularized layer. Osteosclerid—Cells broad at the base, with conspicuous longi- tudinal pores; intercellular spaces below the Malpighian cells tri- angular; walls colored brownish; cells containing pigment and tannin. ri ; Nutrient.—This layer is much compressed, and differentiated into two parts; cells elongated, rather thin-walled; those in the lower portion carry a great deal of pigment, and are much more compressed than the upper portion. Endosperm.—The endosperm is of unequal development, later- ally as much as 150% in thickness. Harz gives the thickness as 250". Cells of the aleurone layer rectangular, thick-walled, and filled with fat and protein grains. This layer is followed by re- serve cellulose. The primary wall persists when treated with weak solvents. The walls, except the primary, color blue with chlor- iodide of zinc. The inner portion of the endosperm consists of . thick-walled, elongated cells. Embryo.—Cells of the first row smaller than those below; ex- terior walls thickened more than the lateral; all of the cell walls consist of cellulose. Cells contain fat and protein grains; starch grains do not occur, though Harz says they are usually abundant. In several specimens examined starch was not found even when potassium hydrate or weak sulphuric acid was used with the iodine. 548 WEED FLORA OF IOWA Fe Fic. 421. Microscopic structures of seeds of some leguminous weeds. I. and V. Yellow Sweet Clover (Melilotus officinalis). 1. White Sweet Clover (Melilotus alba). III. Black Medick (Medicago lupulina). IV. Bur Clover (Medicago denticulata). m=malpighian cells. ll=light line of the same. o=osteosclerid. em=embrya t=testa. en=endosperm. p=parenchyma. n=nucellus, .(Drawings by L. H. Pammel.) MICROSCOPIC STRUCTURE OF WEED SEEDS 549 Bur Clover (Medicago denticulata Willd.). The seeds of this species agree with those of M. lupulina. Malpighian cells 35-384 long; the narrow light line occurs be- low the conical layer; the chromatophores are absent. Osteosclerids 16-184 long; longitudinal striae well marked. Cross sections show beyond a doubt that these striae are canals. The nutrient layer is much compressed. Aleurone layer of endosperm as in M. lupulina. The mucil- aginous reserve cellulose not so strongly developed as in the last species. Treatment with iodine gives no reaction for starch; nor do blue grains appear when treated with weak sulphurie acid and iodine, or potash and iodine. An abundance of fat and protein grains occurs in the cells. Walls of the reserve cellulose color light blue. Malpighian cells a darker blue. Sweet Clover (Melilotus alba Lam.). Testa and endosperm vary in thickness, average 75". Malpighian cells as long as the thickness of the endosperm and remainder of testa. : Malpighian.—Cuticle wavy and well developed; the cuticularized layer below with small, conical projections, those of two adjoining cells meeting at the middle lamella of the lateral walls, giving the layer the appearance of consisting of conelike projections. These cones are also connected with the small pore-canals. This cuticular- ized layer is highly refractive. The lhght line consists of a nar- row but distinct refractive zone below the conical layer. The refractive zone colors blue with chlor-iodide of zinc. The whole upper part is more or less refractive. The remainder of the cell wall contains pigment and is colored blue with chlor-iodide of zine; the cuticularized layer as well as the conical layer colors blue. Small canals project into the walls, and in some cases extend beyond the light line. The chromatophores are irregularly dis- tributed in the cell cavity, some near the base, others in the center. Osteosclerid.—Cells with a broad base and a small triangular intercellular space above; longitudinal pore-canals in the upper - part of the cell, but these do not extend its entire length. Nutrient—This layer is much compressed; consists of thin- walled cells, divided into two parts; cell walls of lower part thicker. Both layers contain pigment and tannin, the upper more than the lower. Cell walls consist of cellulose. 550 WEED FLORA OF IOWA - Endosperm.—The aleurone layer is quite distinct; the cells are rectangular; cell walls made up of cellulose. The walls of the re- mainder of the endosperm, except where it joins the embryo, are thick, consisting of mucilaginous reserve cellulose. Cells of the internal layer of the endosperm thick-walled, clon sey containing some protein grains and fat. Embryo.—Cells of the exterior walls of first row thickened; smaller than those below. All of the cells contain fat, protein grains, and small starch grains. Procambial vessels well developed. Yellow Sweet Clover (Melilotus officinalis Lam.). Testa with endosperm varying from 260-300 in thickness. The Malpighian cells of this species are longer than in WM. alba, and also more abundantly supplied with pigment. The conical pro- jections are longer. The osteosclerids are longer and nearly as wide above as below. The longitudinal canals are as conspicuous and well developed as in that species. Cells of the nutrient layer, especially in the lower part, are abundantly supplied with pig- ment and some tannin. The walls of the aleurone cells are thick; the mucilaginous reserve cellulose and the thick-walled, elongated cells are not essentially different from the last species. It also agrees with it with respect to the embryo. Hairy Prairie Clover (Dalea alopecuroides Willd.) (Parosela dalea (l.) Britt.). Testa and endosperm from 150-265 in thickness. Variation is mostly due to endosperm, which reaches its greatest development laterally. Malpighian.—Cells are 36.4" in length. Cuticle prominent; cuticularized layer not conspicuous; narrow light line near the cuticle; pores prominent, extending into the walls beyond the light line. Cell cavity broad at the base, containing protein erains. Osteosclerid.—Cells thick-walled, lighter in color than the Mal- pighian layer. They contain pigment, tannin and protein. Nutrient—This layer is compressed and the cells are elongated. Walls color blue with chlor-iodide of zinc. Brown pigment abundant in the vascular region. Endosperm.—Aleurone cells nearly isodiametric, containing fat, and protein; most of the endosperm consisting of reserve cellulose MICROSCOPIC STRUCTURE OF WEED SEEDS 551 Fie. 422. Microscopic structure of leguminous seeds (Leguminosae). I. Silky Sophora (Sophora sericea). II. Parosela (Dalea alopecuroides). III. Common Vetch (Vicia sativa). IV. Stemless Locoweed (Oxvytropis lam- verti). V. Wild Liquorice (Glycyrrhiza lepidota). m=malpighian cells. l=light line. em=embryo. o=osteosclerid. p=parenchy- ma. al=aleurone layer. en=endosperm. en=endosperm reserve cellulose cells. tr=tracheae. (Drawings by L. H. Pammel.) 552 WEED FLORA OF IOWA with prominent pore-canals; the internal layer consisting of elongated thick-walled cells, with cell cavity much reduced. Embryo—First row of cells of embryo smaller than underlying, with thickened outer walls. Cells below with small intercellular spaces. Reserve material consists of fat and protein grains; starch is absent. Stemless Loco Weed (Oxytropis lamberti Pursh.). Testa and endosperm 170-1754. This, the so-called loco weed, is said to cause disturbances in animals, but alkaloids have not been found in the seed or any other part of the plant. Malpighian.—Cells 40-424 long. Cuticle somewhat uneven; the narrow well marked cuticularized layer colors blue with chlor- iodide of zinc; the light line occurs below the cuticularized layer, and this also colors blue; the remainder of the cell wall takes on a darker blue color. Cells contain an abundance of pigment, tannin, and some plastic material. Osteosclerid.—Cell walls thickened, not prominently I-shaped, but with an elongated intercellular space. Nutrient.—Layer consists of thin-walled elongated cells from ten to twelve rows. Pigment more abundant in lower than in upper part. Walls color blue with chlor-iodide of zine. Endosperm.—The aleurone layer consists of thick-walled cells; the underlying thick-walled ‘cells of the reserve cellulose become mucilaginous on the addition of water. The internal part consists of thick-walled, elongated cells. The cells contain protein. Embryo.—Cells of the first row smaller, with thick exterior walls; cells below not so compact atid with thinner walls. Cell walls color blue with chlor-iodide of zine. Starch is absent but cells contain fat and protein. Bush Clover (Lespedeza capitata Mx.). Testa and endosperm 90-100 thick. Light line occurs close under the cuticle. A large spherical chromatophore occurs in the pigmented Malphigian cells which is variable as to its position in the cell. The long pores extend to the middle of the cell. The os- teosclerids are short. The nutrient layer is compressed, containing much pigment. Endosperm as in D. stuvei; the aleurone cells are thick-walled, and the reserve cellulose is mucilaginous. Embryo as in the other species containing fat and protein but no starch. MICROSCOPIC STRUCTURE OF WHED SEEDS 553° Common Vetch (Vicia sativa L.). This species has been studied by Harz, Tschirch and Oesterle, Beck, and Sempolowski. Testa irregular, with small projections, 126 thick. Endosperm reduced to a single layer. The presence of endosperm has been indicated by the above writers. Beck speaks of an aleurone spot: (Aleurone fleck) in the epidermal cells of the cotyledons of this and other species of the genus Vicia. Malpighian.Cells 72-75" long, pointed at the upper end; cuticle very irregular because of the projections; cuticularized layer most prominent in the depressions; pores project into the walls below the light line, and partly connect with the cell cavity; _the upper part of the cell is not pigmented, or very little. The light line oceurs just above the pigmented part of the cell. Cell cavity is large at the base, narrows upward, becoming much con- stricted below the light line, and above widens again. Small lateral projections or pores extend into the wall at right angles to the cavity. A large chromatophore, some pigment, and small granules occur in the cavity. The walls in lower part of cell are colored bluish brown. Osteosclerid.—Cells are thick-walled, 13-16.8- long, longitudinally striated. Upper and lower ecross-bars nearly equal; the inter- cellular spaces elongated. Tschirch and Oesterle state that this layer is not very strongly developed, but in specimens which I have examined it is well developed. These cells are more or less variable, as indicated by Harz, who states that they are from 11-13" long. Nutrient —This layer is differentiated into two parts; the upper consists of thin-walled, elongated cells with a yellowish pigment; the cells of the lower part are larger, thin-walled and elongated, containing a brown pigment. Nucellus.——This consists of a narrow zone of compressed cells. Endosperm.—Ocecurs in the form of thick-walled elongated cells with a narrow eell cavity. Usually only one or two rows of cells. _ Hmbryo.—The outer row of cells of the cotyledon is continuous. The exterior walls are thickened; cells below are more loosely ar- ranged; small intercellular spaces in the angles of the cells; the epidermal cells contain fat and protein, the others in addition an abundance of spherical or elliptical starch grains measuring 25 x22.5" to 50x25". Palisade cells wanting. 554 WEED FLORA OF IOWA Wild Liquorice (Glycyrrhiza lemdota (Nutt.) Pursh.). Testa and endosperm 490-500" thick. Endosperm variable in different parts of the seed, but usually well developed. Malpighian—Cells 70-75" long. Cuticle somewhat irregular; the light colored cuticularized layer is followed by a narrow but sharply marked zone, the light line; cell cavity is large at the base, gradually tapering upward. Pore-canals extend into wall beyond the ight line. Cells contain pigment and some plastic material. Osteosclerid.—The I-shaped cells are thick-walled, with small projections somewhat similar to those shown for Ervuwm lens by Mattirolo and Buscalioni. The intercellular space is elongated. The cells attain their greatest development in the hilar region. All of the cells carry some pigment and plastic material. Nutrient.—This layer is much compressed and thin-walled; cells number from four to six rows. Pigment is most abundant in the lower part of the layer. Endosperm.—Aleurone layer consists of nearly isodiametric thick-walled cells. The mucilaginous reserve cellulose is variable in quantity. Cell walls differentiated into primary, secondary and tertiary. Internal part of the endosperm consists of thick-walled, elongated cells. All of the cells contain protein grains. Embryo.—Cells of the outer row smaller than those within; ex- terior walls thickened, those below more loosely arranged than the -epidermal; more compact and with thicker walls than those of Astragalus meaxicanus. Cells contain fat and protein grains but no starch. LINACEAE, FLAX FAMILY. Common Flax (Linwm usitatissimum L.). The shining, brownish seeds consist of an outer epidermal layer of thick-walled cells; walls colorless and stratified; these become mucilaginous on the addition of water. The cell cavity is very small, the underlying layer consists of yellowish parenchyma cells. The third, called the fiber layer by Winton, consists of sclerotic parenchyma cells with pore-canals, the sclerotic parenchyma fol- lowed by a layer of colorless cross-cells with thin walls. The parenchyma cells of the pigment layer follow; these cells are squarish, pigment yellowish brown. The endosperm follows the MICROSCOPIC STRUCTURE OF WEED SEEDS 555 Fic. 428. Microscopic structure of the seed of common Flax (Linum usitatis- simum). : ep=epidermis. p=parenchyma, underneath thick-walled sclerenchyma cells and the pigment layer. pi=pigment layer. en=endosperm. (Drawing by L. H. Pammel and Charlotte M. King.) pigment layer and consists of 2-6 layers of cells, the walls being thicker than those of the embryo, and containing fat and aleurone grains. The epidermal cells of the embryo are squarish, the cells” underneath on the upper face of the cotyledon are palisade-like. All of the cells contain fat and protein. ' GERANIACEAE, GHRANIUM FAMILY. Carolina Cranesbill (Geranium carolinianum L.). The testa of the smooth small seeds consists of an outer epider- mal layer, the cell walls not greatly thickened; underlying it is a single row of elongated parenchyma cells followed by the Mal- phigian cells; the light line is narrow and occurs in the middle of the cell; this is followed by a layer of much larger cells with thick walls. The inner seed coat is much compressed and consists of several rows of small cells and a single row of large cells. The walls in both cases are not greatly thickened. The nucellus is much compressed. The cells of the endosperm are not much longer than broad. The cells of the outer row or the aleurone layer are much smaller than those of the second layer. 556 WEED FLORA OF IOWA pal Fe wes Se ee € 9Gn. A Z ee en aa 3Be Fic. 424. Microscopic structure of the seed of common Geranium (Geranium carolinianum). p=parenchyma, malpighian cells below. ll=light line of mal- ep=epidermis. pal=palisade cells. it=inner testa. n=nucellus. en=en- pighian cells. dosperm. (Drawing by L. H. Pammel and Charlotte M. King.) EUPHORBICEAH,* SPURGH FAMILY. Flowering Spurge (Huphorbia corollata L.). The outer layer of the seed coat is mucilaginous; the walls of the cells are thickened and colorless; showing stratification upon addition of water; the cell contents are dark in color. The granu- lar layer beneath the mucilaginous cells is well developed; the con- tents give a blue reaction to iodine. Underneath the granular layer is a row of slightly elongated thin-walled parenchyma cells; beneath these are the long palisade-like cells, in which are pores of less prominence than in HL. marginata and others. Next lie two compressed layers of thin-walled parenchyma cells. Spotted Spurge (Huphorbia preslii Guss.). The outer layer of cells are darkened; these cells are not mucilaginous. The palisade-like cells are present, as in all the species. *The descriptions here given are from a paper by L. H. Pammel, Trans. Acad. Sci. St. Louis. 5:543. The literature will be found in this paper. MICROSCOPIC STRUCTURE OF WEED SEEDS 557 Prostrate Spurge (Huphorbia maculata L.). The walls of the outer cells are dark in color; these cells are but slightly mucilaginous. The granular layer is not pronounced. The palisade-like cells resemble those of the other species. Fie. 425. Microscopic structure of the seeds of the Spurge family (Huphor- : biaceae). I. Cypress Spurge (Huphorbia cyparissias). II. Flowering Spurge (Huphor- bia corollata). ep=epidermis, in figure II cell-walls mucilaginous. pal=palisade cells. p= parenchyma cells. ; (Drawing by L. H. Pammel.) Yellow Spurge (Huphorbia cyparisstas L.). The seed shows the usual palisade-like cells with the overlying and underlying thin-walled parenchyma cells. MALYVACEAH, MALLOW FAMILY. Velvet Weed, Butterprint (Abutilon theophrasti Medic.).* The outer layer a, of the first integument is transformed into a strongly refractive layer. The second layer is composed of radially elongated cells. The seed hairs arise from a single cell and are large and conspicuous. The hairs are spindle-shaped and thin- walled; they occur mostly at the ends of the seed and are more or less pressed to it. There is little or no coloring matter in this in- *From Rolfs, P. H., Bot. Gaz. 1892 :33-39. 558 WEED FLORA OF IOWA tegument excepting in the base of the hair cells. The palisade cells, c, are narrow for their length. The cell cavity is not promi- nent and the nodosity is inconspicuous. The light line is narrow and occurs near the outer end of the palisade layer. The sub- palisade portion, d, is made up of two layers of light brown cells. They are symmetrical and elongated tangentially. Measurements, seed coats, 1474; outer integument, 134; palisade layer, 96"; subpalisade, 38. Shoo-fly (Hibiscus trionum L.). P. H. Rolfs has made a study of H. militaris, the microscopic structure of which species closely resembles that:of H. trionwm. The dark grayish seeds are roughened with tubercular processes, which contain the ‘‘seed hairs.’? The seed hairs consist of cells somewhat longer than broad beginning with a broad base, ex- tending into a several-celled trichome, the terminal portion larger than remainder of cell. These hairs contain a yellow pigment. The Malpighian cells occupy about one-half the thickness of the testa, the cuticle is well developed, the hght line is near the upper portion of the cell. The cell cavity is spindle-shaped and near the middle of the cell. The remainder of the testa consists of a subpalisade portion of parenchyma cells carrying a brownish pig- ment; underneath it, larger thin-walled cells; walls brownish, a compressed narrow, brownish layer, the nucellus, n. This is fol- lowed by the squarish cells of the endosperm. Common Mallow (Malva sylvestris L.). The surface of M. sylvestris is rough in appearance. The second layer, b, of the outer integument, a, has been compressed into a thin layer and seems to have no definite arrangement. The outer layer, a, has been elongated radially. In places, these elongated ‘eells have divided forming a double layer of cells. There is no brown coloring matter in this integument nor is there any between the integuments. The palisade cells, c, are clear; the walls thick. The cell cavity occupies about one-third the length of the cells, the lower end reaching to the middle. The nodosity is promi- nent. Below the cavity the cells are clear, almost transparent. The subpalisade portion, d, is usually made up of two layers, at some places only one, of large dark brown eells. MICROSCOPIC STRUCTURE OF WEED SEEDS 559 Measurements, seed coats, 122”; outer integument, 274; outer layer of same, 22”; inner layer of same, 5“; palisade layer, 70#; subpalisade, 254. (P. H. Rolfs.) Fie. 426. Microscopic structure of the seeds of some Malvaceous weeds. I. Shoo-fly (Hibiscus trionum). II. Indian Mallow or Butterprint (Abutilon theophrasti). III. Mallow (Malva sylvestris). IV. Sida (Sida spinosa). V. Cheeses or common Mallow (Malva rotundifolia). ep=epidermal cells. m=malpighian cells. ll=light line. pi=pigment layer. D= parenchyma cells. pal=palisade or malpighian cells. n=nucellus. en=en- dosperm. h=trichome. t=plant hair or trichome surface view (Drawings by P. H. Rolfs.) Cheeses or Mallow (Malva rotundifolia L.). The roughened, somewhat velvety, dark grayish seeds consist of an outer row of rather large, rather thick-walled cells, walls colorless. This layer is followed by the very long Malpighian ‘cells, which are longer than the cells of the rest of the testa, light line near the upper part of the cell, lower portion of cell slightly yellowish. Cavity spindle-shaped near the middle. The dark 560 WEED FLORA OF IOWA brownish pigment cells of two or three rows. Adjacent to the pigment layer is the colorless compressed nucellus and the squarish endosperm cells. Sida (Sida spinosa L.). The brownish seeds are minutely roughened. The Malpighian cells occupy more than one-half of the thickness of the testa, the cuticle is well developed, the light line occurs near the upper end of the cells. The cell cavity is spindle-shaped near the lower end of the cell. The subepidermal layer contains the brownish pig- ment, the walls of these cells are thickened; the parenchyma cells of the layer underneath are colorless, the walls are less thickened; a narrow pigment layer follows. The cells of the endosperm are thick-walled containing protein grains. ONAGRACHAH, EVENING PRIMROSE FAMILY. Evening Primrose (Oenothera biennis L.). The small, irregular, winged seeds are rough. The epidermal cells, ep, of the testa are small, thick-walled with minute pore canals; the underlying parenchyma cells, p, are large, thick-walled and with small pore canals; these cells reach their greatest develop- ment where the wings occur. The remaining portion of the testa consists of four or five layers of cells more or less rectangular. The nucellus and the endosperm much compressed, outer epidermal cells of the embryo, em, thin-walled, a little longer than wide. Bez Fic. 427. Microscopic structure of the aie. of Evening Primrose (Oenothera biennis). ep=epidermis of thick-walled cells. p=thick-walled parenchyma underneath thinner-walled parenchyma. n=nucellus and below a few layers of cells of the endosperm. (Drawing by Charlotte M. King and L. H. Pammel.) MICROSCOPIC STRUCTURE OF WEED SEEDS 561 UMBELLIFERAH, PARSLEY FAMILY. Sweet Cicely (Osmorrhiza longistylis (Torr.) D.C.). The epidermal cells are longer than broad, the outer walls are thickened, five to six rows of thin-walled parenchyma cells occur underneath the epidermal layer; the ribs contain the vascular ele- ments, thick-walled sclerenchyma cells; the testa consists of a layer of thin-walled. parenchyma eells, p, followed by a layer tan- gentially elongated and-another of large parenchyma cells, p*, and the compressed nucellus, n.. The endosperm consists of large thin- walled cells. Fic. 428. Microscopic structure of some Umbelliferous weeds. I. Sweet Cicely (Osmorhiza longistylis). II. Cow Parsnip (Heracleum lana- tum). III. Wild Carrot (Daucus carota). Underneath, parenchyma cells, thick-walled sclerotic parenchyma, sectional view to the right. n=nucellus. p=parenchyma. t=testa. en=endosperm. ep=epidermis. o=oil duct. (Drawings by L. H. Pammel.) 36 562 WEED FLORA OF IOWA Wild Carrot (Daucus carota L.). The cremocarp consists of an outer epidermis of cells some- what longer than broad; the underlying cells are of similar struc- ture except the cells near the inner epidermis, which are much longer than wide; a large oil duct occurs in each rib. The testa consists of a single layer of parenchyma cells and a few rows of compressed elements, probably the nucellus. The endosperm of thin-walled parenchyma cells contains protein. Cow Parsnip (Heraclewm lanatum). An outer epidermis with outer walls thickened. The underlying parenchyma cells similar, followed by a layer of thick-walled sclerenchyma cells. The testa of single layer of cells. The en- dosperm cells contain protein. ASCLEPIADACEAE, MILKWEED FAMILY. Milkweed (Asclepias syriaca L.). The flattened reddish brown seeds consist of an epidermis of isodiametric cells; the outer walls wavy, and thickened; each cell with a projecting point, outer walls colorless, contents brownish; the epidermis is followed by 10-14 rows of thin-walled parenchyma cells containing a brownish pigment. This layer is followed by oP Fic. 429. Microscopie structure of the seed of Milkweed (Asclepias syriaca). ep=epidermis. p=parenchyma. n=nucellus. em=embryo. (Drawing by L. H. Pammel and Charlotte M. King.) MICROSCOPIC STRUCTURE OF WEED SEEDS 563 the compressed perisperm, brownish yellow in color. The endo- sperm consists of thickish-walled, colorless parenchyma cells, the outer row smaller than the underlying, the inner layer of smaller elongated cells. The contents consist of protein and fat. The inner epidermal cells of the embryo are smaller than the endo- sperm cells; they contain protein and fat. CONVOLVULACHAE, CONVOLVULUS FAMILY. European Bindweed (Convolvulus arvensis L.). The testa consists of an outer row of short or elongated cells p with thick walls and brownish contents. This is followed by a layer of small cells with colorless walls. Contents brownish. The Malphighian cells m are situated underneath. This is followed by a layer of parenchyma cells consisting of 8-12 rows of cells. The endosperm consists of thick-walled colorless cells. The walls in part mucilaginous. Fic. 430. Microscopic structure of the seeds of Convolvulaceae. I. Morning-glory (Convolvulus sepium). II. Cultivated Morning-glory (Ipomoea purpurea). III. Dodder (Cuscuta epithymum). IV. Bindweed (Convol- vulus arvensis). ep=epidermis. m=malpighian cells. ll=light line. p=parenchyma. n=nucellus. en=endosperm. a8 (Drawings by L. H. Pammel and Charlotte M. King.) 564 WEED FLORA OF IOWA Wild Morning Glory (Convolvulus sepium L.). The testa of the seed of the common morning glory consists of an outer layer of elongated cells containing the blackish pigment followed by the Malpighian cells with small cell cavity and promi- nent light line. The colorless parenchyma cells are thick-walled followed by a compressed indistinct layer, the compressed nucellus and the endosperm cells. Cultivated Morning Glory (Ipomoea purpurea (.) Roth.). The black-brownish seeds are rough. The testa. consists of a superficial layer of cells with granular brownish contents. The Malpighian cells m occur underneath; the light line is near the upper part of the cell, cell cavity small. The cells of the paren- chyma layer are compressed. The cells of the endosperm are thick- walled and somewhat mucilaginous. Dodder (Cuscuta epithymum Maurr.). The small seeds are minutely roughened and velvety. The outer layer of the testa consists of large cells with yellowish contents, the following layer consists of thick-walled cubical or prismatic cells p, the colorless Malpighian cells m follow. They are elongated with a small cell cavity. The layer following consists of com- pressed cells, tangentially elongated. The endosperm layer con- sists of an outer aleurone layer of nearly square cells. The cells below are irregular. The endosperm cells contain compound or simple starch grains and protein. BORAGINACHAEH, BORAGE FAMILY. Wild Comfrey (Cynoglossum virginianum L.). The rough fruits are covered with hooked appendages, the sides roughened with small colorless points; the epidermal layer is ir- regular and prominent, in some cells developed into short trichomes, whose walls are greatly thickened. The epidermal walls and con- tents are blackish, the underlying sclerotic parenchyma cells have. thick walls and are blackish. The parenchyma cells of the testa are thinner walled and colorless, followed by the compressed layer of the nucellus n. The endosperm cells are longer than broad, colorless, contain protein and fat. MICROSCOPIC STRUCTURE OF WEED SEEDS 565 Fic. 431. Microscopic structure of some fruits of the Borage Family (Boragin- aceae). I. Stickseed (Lappula echinata) part of fruit or bur. II. Stickseed (Lap- pula echinata). Enlarged view of epidermal cells showing irregularities in cell wall and cavity. III. Wild Comfrey (Cynoglossum virginianum). ep=epidermal cells. n=nucellus. pi=pigment layer. en=endosperm. t=tri- chomes. ; (Drawings by L. H. Pammel and C. M. King.) Stickseed (Lappula echinata Gilibert). The fruits are rough and provided with hooked appendages, the cells of the appendages are elongated, thick-walled, the walls colorless; the cells of the epidermis toward the exterior are very . irregular with projecting rounded or sharp points, walls colorless. The epidermal cells are elongated with central cell cavity and prominent pore canals at right angles with the cell cavity, these having the appearance of a series of cells one above the other. The epidermal cells contain a brownish pigment, the underlying parenchyma cells are thinner walled and also contain a brownish 566 WEED FI.ORA OF IOWA pigment. The testa is thin, consisting of thin-walled parenchyma cells with granular contents. VERBENACEAE, VERVAIN FAMILY. Hoary Vervain (Verbena stricta Vent.). Epidermal cells thick-walled; ‘outer cells colorless. Underneath this lie several layers of pigmented cells, slightly longer than broad. This layer is followed by rather thick-walled, colorless cells, with small cell cavity; one or two rows toward the middle are thick- walled and smaller. Next follow the cells of the embryo. Fic. 432. Miscroscopic structure of seed and fruit of Vervain (Verbena stricta). ep=epidermis, thick-walled. p=parenchyma layer. pi=pigment layer of small parenchyma cells. The two inner layers of cells of the testa. (Drawings by L. H. Pammel and Charlotte M. King.) LABIATAE, MINT FAMILY. Catnip (Nepeta cataria L.). The small dull brown seeds are finely roughened. The pericarp consists of an epidermal layer with thin cuticle, on the addition of water becoming mucilaginous. This layer is followed by under- lying parenchyma cells, then the Malpighian cells with conspicuous light line. The cells are colored yellowish brown; underlying this, MICROSCOPIC STRUCTURE OF WEED SEEDS 567 the testa, consisting of a single layer of cells, thin-walled. The nucellus of whitish, elongated, rather thick-walled cells, the en- dosperm differentiated into an outer layer of larger parenchyma and a compressed inner layer of thick-walled cells. The epidermal cells of the embryo nearly isodiametric, contain protein and fat. The epidermal cells of the upper epidermis nearly like the lower, palisade parenchyma underneath. Fic. 433. Microscopie structure of seeds of the Mint Family (Labiatae). I. Catnip (Nepeta cataria). II. Giant Hyssop (Agastache scrophulariaefolia). ep=epidermis, underneath, parenchyma cells in I and sclerotic parenchyma in II, the epidermal cells irregular showing some trichomes. scl=sclerotic parenchyma. en=endosperm. m=malpighian cells. n=nucellus. ll=light line. p=parenchyma. em=embryo. (Drawings by L. H. Pammel and C. M. King.) Giant Hyssop (Agastache scrophulariaefolia (Willd.) Ktze.). The small pubescent nutlet consists of the epidermal cells of the pericarp with small unicellular thickened trichomes; cells con- tain a brownish pigment; the cells of: the layer underneath are thick-walled, cell wall and cavity contain a brownish pigment. The Malpighian layer follows, the cells being longer than wide, the light line near the upper part of the cell. These cells are yellowish. The testa is compressed and consists of elongated cells, brownish in color. The endosperm varies in thickness, the cells are in some cases elongated, in others squarish, and contain fat and protein material. 568 WEED FLORA OF IOWA SOLANACHAE, NIGHTSHADE FAMILY. Horse Nettle (Solanum carolinense L.). The testa consists of an outer row of cells whose walls are mucilaginous ; the underlying cells are thick-walled with interven- ing air spaces broad below and narrow above. The cells of this layer contain the yellow pigment. The underlying parenchyma layer is compressed, composed of colorless cells which are longer than wide; the layer adjacent to the endosperm is compressed and not clearly defined. The cells of the inner portion of the endosperm are smaller and contain protein. Cells of embryo are small and contain protein. Fig. 434. Microscopic structure of the seeds of some Solanaceous weeds. I. Jimson Weed (Datura stramoniwm). II. Black Nightshade (Solanum ni- grum). III. Buffalo Bur (Solanwm rostratum). IV. Horse Nettle (Sola- num carolinense). V. Ground Cherry (Physalis pubescens). ep=epidermis. en=endosperm. pal=palisade cells. em=embryo. p=parenchyma. p!= elongated parenchyma cells. (Drawings by L. H. Pammel and C. M. King.) MICROSCOPIC STRUCTURE OF WEED SEEDS | 569 Buffalo Bur or Spiny Nightshade (Solanum rostratuwm Dunal). The black seeds are irregular on the surface, which is mucil- aginous; the underlying layer consists of thick-walled cells with large cavities into which thickened processes extend; the third layer is also pigmented and consists of squarish cells or elongated, compressed elements. The endosperm is composed of thick-walled cells, the outer nearly square, the others elongated. Starch ab- sent; protein present. The cells of the embryo are smaller, thin- walled and contain protein and fat. Black Nightshade (Solanum nigrum L.). The surface of the yellowish seeds is slightly irregular. The superficial layers consist of mucilaginous, parenchyma, cells, fol- lowed by large parenchyma cells with thickened folds. These cells are, however, variable as to structure and are sometimes short with folds not evident. The elongated, thick-walled portion of the testa contains the yellowish pigment. The cell cavity is tri- angular. The second layer of parenchyma cells is compressed. Jimson Weed (Datura stramonum L.). The blackish seeds are rough, surface irregular, cells and walls of epidermis ep thick; the underlying thinner-walled cells » are small, longer than broad, followed by an indistinct layer p’, whose walls are not clearly defined. The layer adjacent to the endosperm consists of somewhat larger cells also thin-walled. The endosperm cells e are large, containing protein. Ground Cherry (Physalis pubescens L.). The epidermal cells are thick-walled, the outer walls colorless. These cells contain the pigment. The parenchyma cells of the second layer are elongated; an indistinct compressed layer follows. The outer row of the endosperm cells are smaller, containing protein. A discussion of the seed coats of this family is to be found in Harz, Samen- kunde, 2 996-1025. 570 WEED FLORA OF IOWA SCROPHULARIACEAE, FIGWORT FAMILY. Mullein (Verbascum thapsus L.). The minute, roughened seeds are irregular. The epidermal cells are dark colored, longer than broad and thick-walled. The epider- mal layer is followed by one or more layers of parenchyma eells, more numerous in the micropilar region; an indistinct and com- pressed layer with large intercellular spaces follows, these corre- sponding to the irregularities on the surface, smaller intercellular Fic. 435. Microscopic structure of the seeds of the Figwort family (Scrophul- ariaceae). I. Moth Mullein (Verbascum blattaria). II. Common Mullein (Verbascum thapsus). ep=epidermis. p=parenchyma. al=aleurone layer. en=endosperm. em=em- bryo. (Drawings by L. H. Pammel and Charlotte M. King.) spaces occurring between the larger. The nucellus is compressed, showing remnants of cell walls. The endosperm consists of an outer aleurone layer, the walls with pore canals; this is followed . by cells of the same character. The cells of the embryo are smaller. The testa of Verbascum blattaria in an undeveloped seed consists of the epidermal cells with chlorophyll and the SEES much larger cells also with chlorophyll. MICROSCOPIC STRUCTURE OF WEED SEEDS 571 PLANTAGINACEAH, PLANTAIN FAMILY. Common Plantain (Plantago major 1.). The small, yellowish brown seeds consist of an epidermal layer of thick black-brownish seeds. The walls on the addition of water become mucilaginous and expand. The cell cavity is small. The epidermal layer is followed by several layers and small parenchyma cells. These cells are usually somewhat compressed and brownish in color. The pigment layer and endosperm consist of an outer layer of aleurone cells smaller than the underlying eells. These cells contain protein grains and starch; the walls have small: pore canals. Bre en ' t ' 1 wn : Fig. 486. Microscopic structure of the seeds of Plantain family. I. Bracted Plantain (Plantago aristata). II. Common Plantain (Plantago major). III. Buckhorn (Plantago lanceolata). 1l=epidermal cells showing stratification. c=cell cavity. ep=epidermis. pi=un- derlying parenchyma cells. 2=underlying parenchyma cells of the testa. em=embryo. n=nucellus. p=parenchyma cells. en=endosperm. (Drawings by L. H. Pammel and C. M. King.) 572 WEED FLORA OF IOWA _ Buckhorn (Plantago lanceolata L.). The smooth, brownish, elongated seeds consist of epidermal cells with a small cavity and thick walls; the outer walls become mucil- aginous on the addition of water. The underlying cells are thin, blackish, elongated, followed by a brown pigment layer as in the last species. The aleurone cells and remaining endosperm cells are of about the same shape and size; the walls are provided with small pore canals. Bracted Plantain (Plantago aristata Mx.). The seeds of this are similar in structure to those of P. lance- olata. The walls of the epidermal cells are mucilaginous. The underlying parenchyma cells and the underlying pigment layer are followed by the endosperm and embryo. The cells of the nucellus are narrow, elongated, and thick-walled. The eells of the embryo are isodiametric and are thin-walled. Several species of the genus have been studied by Harz*. Numerous earlier papers and references will be found in Pammel’s article in Transactions St. Louis Academy of Science, 9:91. RUBIACHAH, MADDER FAMILY. Bedstraw (Galium aparine L.). The dry fruit of the common bedstraw is provided with hooked prickles. The epidermal cells are somewhat elongated, brownish. The trichomes consist of single hooked cells with pitted walls. The underlying thin-walled parenchyma cells are elongated. Harz ob- served mucilaginous cells in this portion of the- pericarp. The testa is very much reduced, consisting of several layers of thin- walled cells with granular contents. The endosperm consists of an aleurone layer, thick-walled cells followed by a thick-walled irregular layer of cells with pore canals. Contents consist of pro- tein grains. *Samenkunde 2; 983. MICROSCOPIC STRUCTURE OF WEED SEEDS 573 Fic. 437. Microscopic structure of fruit of Bedstraw (Galium aparine). ep=epidermis. t=testa. t! =trichome. en=endosperm. (Drawing by L. H. Pammel and Charlotte M. King.) CUCURBITACEAE, GOURD FAMILY. Wild Cucumber (Sicyos angulatus L.). The testa of the brownish seeds consists of elongated epidermal cells with narrow cell cavities; walls thickened, with pore canals. The underlying portion consists of a layer of one or two rows of cells with thin walls, and protein contents; the remaining portion of the testa is composed of thin-walled cells with large inter- cellular spaces. The related species, Echinocystis lobata, contains a thick, pigment layer of brownish sclerotic parenchyma followed by a thinner layer of thick-walled cells with blackish brown pig- ment. In this species, as in Sicyos angulatus, frequently portions of the fruit adhere to the surface. The seed coats of this family have been described-by Harz. 574 WEED FLORA OF IOWA Fic. 438. Microscopic structure of seeds of some cultivated weeds. I. Wild Cucumber or Wild Balsam Apple (Echinocystis lobata). II. Bur Cucumber (Sicyos angulatus). f ep=epidermis. ll=light line. p=parenchyma. m=malpighian layer. scl=scler- otic cells. (Drawings by L. H. Pammel and C. M. King.) COMPOSITAE, COMPOSITE FAMILY. Large Ragweed (Ambrosia trifida IL.). The so-called ‘‘seed’’ consists of an inyolucre of rather thick- walled sclerotic parenchyma cells, occurring underneath the epidermis; some of these cells are radially elongated, others are spherical in cross section, showing numerous pore canals. The internal layer of the involucre is composed of nearly isodiametric, thick-walled, sclerotic cells. The testa consists of a layer of brown- ish colored cells followed by a layer containing black pigment. Next within lies a layer of thick-walled, small, nearly colorless cells, then the compressed layer of the nucellus, followed by the nearly square cells of the embryo. MICROSCOPIC STRUCTURE OF WEED SEEDS > 575 Common Sunflower (Helianthus annuus L.). The microscopic structure of the fruit and seed has been studied by Hanausek, Harz, Winton and Moeller. The obovoid achenes are more or less four-sided. The pericarp in some varieties is nearly black, in others it is striped with white and black. The pericarp consists of thin porous walls which are dark colored in the black seeded varieties, although in the varieties with striped seeds only a part of the cells are colored. Some of the cells are elongated, forming duplex hairs, which are attached to what Hanausek called the ‘‘foot cell.’? The underlymg hypodermal cells of 4-6 rows of cells are thick-walled and porous, the cells ar- ranged in rows. These contain the blackish pigment, pitchlike in character. The third layer consists of thick-walled sclerotic parenchyma cells. These cells are more or less isodiametric. This layer contains the fibrovascular bundles which occur adjacent to the thin-walled parenchyma eells. Extending into the sclerotic parenchyma are radial rows of thinner-walled parenchyma cells. This layer is followed by large thin-walled parenchyma cells. The testa consists of thin-walled loose parenchyma cells. The outer or epidermal cells are roundish and have obscurely beaded walls; the spongy parenchyma follows and contains the fibrovascular bundles. The spongy parenchyma is followed by the rectangular cells of the inner epidermis. The endosperm consists of one or two rows of aleurone cells. The epidermal layer of the embryo consists of small, rather thin-walled cells of the cotyledons and underneath! this on the upper surface are several rows of palisade cells. These cells contain irregular spherical aleurone grains larg- er than those in the epidermal cells. Crownbeard, (Verbesina helianthoides Mx.). The microscopic structure of the achene includes a series of small, rather thick-walled epidermal cells, followed by elongated or short, rather thick-walled parenchyma cells. The pigment layer is com- posed of thick-walled cells, whose walls contain a blackish pigment. The testa consists of two layers; an outer of elongated, thick-walled cells and an inner layer of shorter cells also colorless. The cells of the embryo are much larger and contain protein grains. Boot-jack, Spanish Needle (Bidens discoidea (T. & G.) Britton). The pericarp consists of an outer epidermal layer underlaid by a similar layer of elongated, thick-walled cells; between which are 576 WEED FLORA OF IOWA thinner-walled parenchyma cells. The outer layer of the testa is composed of nearly isodiametric cells, followed by larger thick- walled parenchyma cells; cells of the lower portion compressed. The endosperm is much reduced, of elongated cells; embryo with row of outer cells longer than broad. cK J > Z ‘ o> 7. AWA ) A haces (i Bra \e < Ay Wy ry Co 1 . é / Fig. 439A Fig. 439 B Fic. 439. Sunflower (Helianthus annuus). A. Cross section of outer layers of pericarp. 0, epicarp with h, .hairs; K, hypoderm; H, fiber bundles separated by m, parenchyma; p, parenchyma with g, fibro-vascular bundles. X 160. B. Epicarp with h, twin hairs, in surface view. After Winton. Burdock (Arctium lappa L.). The brownish mottled fruits consist of a thick-walled pericarp, the outer epidermal layer of thick-walled cells, walls colorless or but slightly colored; the underlying layer of six or eight rows of thick-walled cells, not as thick, however, as the epidermal cells. The first layer carries the brownish or blackish brown pigment. MICROSCOPIC STRUCTURE OF WEED SEEDS 577 7? OG= rar par c yo) em, SIS® SEY Fic. 439A. Miscroscopic structure of some weeds of the Sunflower family (Compositae). I. Small Ragweed (Ambrosia artemisiaefolia). II. Verbesina (Verbesina hel- ianthoides). III. Spanish Needle (Bidens discoidea). IV. Burdock (Arctium lappa). ep=epidermis. scl=sclerotic parenchyma, t=testa. n=nucellus. en=endosperm. em=embryo. pal=palisade cells. (Drawings by L. H. Pammel and Charlotte M. King.) The pigment layer is followed by elongated sclerotic parenchyma with a narrow cell cavity; cells with granular contents. The testa is thin, cells elongated, cell wall thickened slightly, tinged with yellow. The testa is followed by remnants of the nucellus. The parenchyma cells of embryo follow the nucellus. Wood or Field Thistle (Cirstum discolor (Muhl.) Spreng.). The microscopic structure of the yellowish gray seeds shows a elear relationship to Arcttwm; the outer epidermal layer of the pericarp consists of thick-walled colorless cells, the walls bright and lustrous; the pigment layer underneath the walls, not nearly 37 578 WEED FLORA OF IOWA as thick-walled as the epidermal, pigment yellowish. The elong- ated sclerotic parenchyma cells are yellowish white or nearly color- less; this layer is followed by the parenchyma cells of the testa and remnants of the nucellus. The parenchyma cells of the embryo with numerous small intercellular spaces contain protein and fat. The fibrovascular bundles are located one at each end of the achene. Fic. 440. Microscopic structure of the seeds of Thistles (Cirsiwm). I. and II. Common Wood Thistle (Cirsiwm discolor). III. Iowa Thistle (Cir- sium iense). ep=thick-walled epidermal cells. pi=pigment layer. scl=sclerotic palisade cells with narrow cell-cavity. p=parenchyma cells of testa. t=testa. n=nucellus. em=embryo. fv=fibro-vascular bundles. (Drawings by L. H. Pammel and Charlotte M. King.) Iowa Thistle (Cirstum ioense (Pammel) Fernald). The microscopic structure of the seeds is similar to that of pre- ceding, comprising a thick-walled epidermal layer with bright colored walls and underlying pigment layer, an elongated sclerotic parenchyma and the testa of small parenchyma cells. MICROSCOPIC STRUCTURE OF WEED SEEDS 579 Chicory (Cichorium intybus L.). The microscopic structure of the pericarp and seed of chicory has been given by Harz and Lavialle. The pericarp consists of epidermal cells whose outer walls are irregular and cuticularized. This is followed by a variable number of rows, usually 10-15, of sclerotic parenchyma, some of the cells of which as observed by eee e 7 Broce Iria xt, oa Bay (fer Fic. 441. Microscopie structure of the seed of Chicory (Cichorium intybus). ep=epidermis. scl=sclerotic parenchyma. per=pericarp. t=testa. al=aleu- , rone layer. en=endosperm. (Drawing by Charlotte M. King and L. H. Pammel.) Kraus, contain crystals of calcium oxalate. Beneath are 6-8 rows of thin-walled, elongated parenchyma. The testa consists of a layer of large epidermal cells, followed by smaller thin-walled parenchy- ma, the inner portion much compressed. The aleurone layer is of elongated cells containing aleurone grains. Dandelion (Taraxacum officinale Weber). - Harz has given an account of the structure of the pericarp and seed. The epidermal cells are irregular, many of the cells pro- longed into one-celled trichomes, the ribs forming tubercular processes. The underlying portion containing the thinner-walled parenchyma and the sclerotic parenchyma cells of the ribs. This is followed by several rows of compressed, elongated, thin-walled parenchyma cells. The testa consists of the epidermal layer of elongated, spirally thickened walls, followed by a compressed layer of parenchyma cells. The endosperm of one or two rows of aleurone cells. . WEED FLORA OF IOWA 580 Saag, Rees Sam SSN x, ile C nN () Dandelion (Taraxacum officinale). Fic. 442. BIBLIOGRAPHY. ANATOMICAL STUDY OF SEED COATS. GENERAL. » Buscaloni, L. Contribuzione allo studio della membrana cellulare. Malpighia 6 :3-40, 217-228; pl. 1-2. 7:105-162; pl. 1-2. 8:3-13; pl. 1-3. 1875. Anatomisch-physiologisch Untersuchung tber d. Dahmen, Max. Funiculus d. Samen. Prings. Jahrb. f. wiss. Bot. 23 :441-478; pl. 20-22. Fluckiger, F. A., and Hanbury, D. Pharmacographia: a history of the principal drugs of vegetable origin met with in Great Britain and British India. Ed. 2, 803. London, 1879. Frank, A. B. Ueber die anatomische Bedeutung und die Entste- hung der vegetabilischen Schleime. Pringsh. Jhrb. f. wiss. Bot. 5:161; pl. 15-16. 1886. MICROSCOPIC STRUCTURE OF WEED SEEDS 581 Godfrin, H. Etude histologique sur les téguments sémineaux des angiosperms. Bull. Soc. des Sci. de Nancy. 5:109-219. Godfrin, J. Recherches sur l’anatomie comparée des cotylédons et de l’albumen. Ann. Sci. Nat. Bot. Soc. des Sci. de Nancy. 59:109-219; pl. 1-5. 1880. Guignard, L. 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Ueber Pflanzenschlerme und seine Entstehung in der Samen epidermis von Plantago maritima und Lepidium satiwwum. Flora. 58 :173-200, 209-216; pl. 6. Winton, A. L. Anatomy of oil seeds among Cruciferae. Microscopy of vegetable products. 174-193. EUPHORBIACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:825-841. Pammel, L. H. On the seed coats of the genus Euphorbia. Contr. Shaw School of Bot. 8:5438-568. Trans. St. Louis Acad. Sci. 5 :543-568. 584 WEED FLORA OF IOWA GENTIANACEAE. Guerin, . Développement et structure anatomique du tégument séminal des Gentianées. Compt. Ren. Acad. des Sci. de Paris. vi: 136. GRAMINEAE. Bessey, C. EK. Structure of the wheat grain. Bull. Neb. Agr. Exp. Sta. 32 Hanausek, T. F. Vorlaufige Mittheilung tber der von Vogl. in der Frucht von Lolium temulentwm entdeckten Pilz. Ber. Deutsch. Bot. Ges. 16:203. 1898. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:1145-1349. Hillman, F. H. The seeds of fescue grass and chess. Bull. U. S. Dept. Agr., Bur. Pl. Ind. 25:4. Holm, Theo. A study of some anatomical characters of N. A. Graminee. Bot. Gaz. 16 :166-171, 219-225, 275-281. 17 :358-363. 20 :362-365. 21 :357-360. 22 :403-406. Jumelle. Sur le constitution des fruit des Graminées. Compt. Rend. 107 :285. 1888. Krutizky. On some peculiarities in the structure of the caryopsis of wheat. Just’s Bot. Jahresb. 21: (Abt. I) 571. 1893. Moeller, J. Die Mikroskopie der Cerealien. Pharm. Centralh. 25: 507. 1884. Pammel, L. H. Comparative anatomy of the corn caryopsis. Trans. Ja. Acad. Sei. 1897. 5. The histology of the caryopsis and endosperm of some grasses. Trans. St. Louis Acad. Sei. 1898 :199-220. Pammel, L. H., Weems, J. B., and Lamson Scribner, F. The Grasses of Iowa. Bul. Ia. Geol. Survey. 1:525. Winton, A. L. The anatomy and microscopic identification of the fruits of darnel and chess. Rept. Conn. Agr. Exp. Sta. 1903. 165-174. Anatomy of the seed of green foxtail. Rept. Conn. Agr. Exp. Sta. 1902. 352-358. Anatomy of seed of yellow foxtail. Ibid. 1902: 358. Ueber amerikanische Weizenausreuter. Ztschr. Unters. Nahr-Genussm. 6:433. LABIATAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :866-870. MICROSCOPIC STRUCTURE OF WHED SEEDS 585 LEGUMINOSAE. Beck, G. Die Samenkunde einiger Leguminosen. Sitzb. K. K. Akad. Zu. Wien. 1878. 79. : Vergleichende Anatomie d. Samen von Vicia und Ervum. Sitz. d. k. Akad. d. Vissench. Wien. 76 :335-352; 2 pl. Chalon, J. La graine des légumineuses. I. Cellule de la carapace. II. Albumen. Soc. des Arts, et des lettres du Hainaut. 1875. 66; pl. 1-3. Haberlandt, G. Ueber die Entwicklungsgeschichte und den Bau d. Samenschule bei der Gattung Phaseolus. Sitz. d. k. Akad. d. Wissensch. Wien. 75:33-47; 2 pl. 1877. Harz, C. D. See Landwirthschaftliche Samenkunde. 2; 562-736. Mattirolo, O. Ricerche anatomo-fisiologische sui tegumenti seminali delle Papilianacee. Nota preventiva. Atti. R. Accad. delle sei. di Toniro. 24. 1889. Ricerche anatomo-fisiologische sui tegumenti seminali delle Papilionacee. I. Anatomia. II. Stor ia di sviluppo dei tegumenti seminali. JII. Contribuzioni allo studio della fisiol- ogia del seminali. IV. Bibliographia. Reale Accad. delle Sci. di. Torino II, 42 :223-318, 359, 445; pl. 1-5. 1892. | Nadelmann, Hugo. Ueber d. Schleimendosperme d. Leguminosen. Prings. Jahrb. Wisensch. Bot. 21:609-691; pl. 16-78. 1890. Pammel, L. H. Anatomical character of seeds of Leguminose. Trans. St. Louis Acad. Sci. 9:91-263; Tables A-G; pl. 7-35. The seed coats of Crotalaria sagitalis and Astragalus mollis- simus. Bienn. Rept. Ia. St. Agr. Coll. and Farm. 13 :17-24; pl. 59-58. | On the structure of the testa of several leguminous seeds. Bull. Torr. Bot. Club. 13 :17-24; pl. 52-58. Schips, K. Ueber die Cuticula und die Auskleidung der Intereel- lularen in den Samenschalen der Papilionaceen. Ber. Deutsche. Bot. Ges. 11:311. 1893. Schively, Adeline F. Contributions to the life history of Amphi- carpaea monoica. Cont. Bot. Lab. Univ. Penn. 1:270-363; pl. 19-36. Schleiden, M. J. Beitrage zur Entwickelungsgeschichte d. Bluthen- theile bei d. Leguminosen. Nov.-Oct. K. Leopold-Carol. Akad. Naturf. 19 :59-84; pl. 9-11. p Winton, A. L. Anatomy of the seeds of Vicia sativa. Winton’s Microscopy of vegetable products. 251-252. 586 WEED FLORA OF IOWA MALVACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:736-750. Rolfs, P. H. The seed coats of Malvacex. Bot. Gaz. 17 :33-39. ONAGRACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :875-878. PHYTOLACCACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:1088-1031. PLANTAGINACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :982-987. Winton, A. L. Anatomy of plantain seeds. Microscopy of vegeta- ble products. 163. Uloth, W. Ueber Pflanzenschleime und seine Entstehung in der Samenepidermis von Plantago maritima und Lepidum sativum. Flora. 58 :193-200, 209-216; pl. 6. 1885. POLYGONACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:1101-1113. Sirrine, Emma. Structure of the seed coats of Polygonacez. Proc. la. Acad. Sei. 2:128-134; pl. 7-9. Winton, A. L. Anatomy of seed of black bindweed. Rept. Conn. Agr. Exp. Sta. 1902 :340-352. Winton’s Microscopy of vege- table products. 157-158. RANUNCULACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :1063-1074. Winton, A. L. Anatomy of Ranwnculus arvensis. Microscopy of vegetable products. 152-154. SAPINDACEAE. Guérin, M. P. Développement de la graine et in particulier du tégument séminal de quelques Sapindacées. Jour. de Bot. 15; 336-348. SCROPHULARIACEAE. Bachmann, E. Die Entwicklungsgeschichte und der Bau der ., Samenschalen der Scrophularinéen. 179. Halle, 1880. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :966-973. SOLANANCEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2 :997-1025. MICROSCOPIC STRUCTURE OF WEED SEEDS 587 UMBELLIFERAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:1030-1063. Winton, A. L. Anatomy of the seeds of wild carrot. Microscopy of vegetable products. 158-159. URTICACEAE. Harz, C. D. See Landwirthschaftliche Samenkunde. 2:978-982. Winton, A. L. The anatomy of certain oil seeds with especial reference to the microscopic examination of cattle foods. Conn. Agr. Exp. Sta. 1903. 175-180. Microscopy of vegetable products. 212-217. CHAPTER IV. MORPHOLOGY OF LEAVES AND FLOWERS By J. N. MARTIN CHAPTER IV. LEAVES. INTRODUCTORY STATEMENT. Leaves are distinguishable into primary and secondary. The pri- mary leaves arise directly from the first cells produced by the division of the fertilized egg and in seed plants are called cotyle- dons. They are usually transient, and not rarely are so distorted by acting as storage places for reserve food that they do not function as foliage leaves at all. The secondary leaves arise upon the sides of the stem and are the ordinary foliage leaves of the plant. They are very important organs in connection with the work of nutrition. PARTS OF A LEAF. In the typical foliage leaf there are three parts—the expanded portion which is called blade or lamina, the leaf stalk (petiole), and a pair of appendages at the base of the petiole known as stipules. Fic. 443. Apple leaf; b, blade; p, petiole; s, stipules. (After Thomé.) 592 WEED FLORA OF IOWA In some cases, as in the Hare’s-ear mustard, (Conringia orient- alis) and in the upper leaves of Canada thistle (Cirsium arvense) the petiole is absent and the blade is directly attached to the stem. Such leaves are designated as sessile. Fie. 444. Sessile leaf of Thistle. (After Thomé.) The stipules are small leaf-like structures which appear at the place where the leaf is attached to the stem. They are very often absent but are conspicuous in the cinquefoils, vetches and other members of the rose and pulse families. VEINING. The blade of the leaf is traversed by a frame work of fibro- vascular bundles known as veins. In the leaves of grasses, sedges, and rushes, the veins run more or less parallel from the base to the tip of the leaf. These leaves are the parallel-veined type. In the leaves of most of our common weeds, the veins are branched so as to form a network. These are the netted-veined type. Palmate and pinnate veining.—Netted-veined leaves are palmately veined when the primary ribs radiate from the base of the petiole as in the great ragweed. If there is only one midrib from which smaller ribs extend both ways, as in dandelion, dock, goldenrod, etc., the veining is said to be pinnate (meaning featherlike). LEAF ARRANGEMENT. Sometimes as in dandelion and evening primrose, the stem does not appear above ground or is late in appearing and the leaves at the surface of the ground are called radical leaves in distinction to the stem or cauline leaves. MORPHOLOGY OF LEAVES AND FLOWERS 593 Fig. 4451 Fig. 44511 Fic, 445. I. Vetch. (Lathyrus aphaca), showing opposite leaves; r, tendril; b, flower; f, fruit. Stipules performing the function of leaves. II. Grass type of leaf; L, leaf blade; G, leaf sheath; Lig, ligule. ‘ (After Thomé.) Leaves are usually arranged so as to secure the best exposure to the light. In the milkweed there is a pair of leaves at each node and the two leaves are on opposite sides of the stem. Here the leaves are said to be opposite. Fig. 446. The clinging stem of the Bindweed, showing the alternate leaves. (After Thomé.) 38 594 WEED FLORA OF IOWA In the asters, ironweed, goldenrods, lamb’s quarter, ete., there is only one leaf at each node, and they are spoken of as alternate. In some cases several leaves appear at each node in a whorl. Such examples of whorled or verticillate leaves are found in the bed- straws and Joe Pye weed. BRANCHING. The outline of a blade is extremely various. When the general outline is completely filled out and the margin represents an even line, the leaf is said to be entire. Examples of such leaves are found in water pepper (Polygonum hydropiper), mild water pepper (P. hydropiperoides), Pennsylvania smartweed (P. pennsylvani- cum), ete. : | Fic. 447. Leaf of the Privet plant showing entire margin. To designate the amount and character of the branching, the following terms are used: Fig. 4481 Fig. 44811 Fig. 448'II Fig. 4481V Fic. 448. I. Triangular, lobed leaf of the Notch-weed. II. The reniform or kidney-shaped leaf of Ground Ivy, illustrating wavy margin. III. Arrow- shaped leaf of Bindweed. IV. Spear-shaped leaf of Sorrel. (After Thomé.) MORPHOLOGY OF LEAVES AND FLOWERS 595 Wavy margin, when the margin forms a wavy line bending slightly inward and outward in succession, as in the bitter dock (Rumex obtusifolius). Fig. 449 T Fig. 449 II Fig. 449 III Fig. 449 1V Fig. 449 V - Fic. 449. I. Leaf of Daisy, spatulate in shape with a serrate margin. San Spiny-pointed, serrulate leaflet of alfalfa. III. Wavy-margined leaf of Pigweed (Amaranthus). IV. Ovate, dentate leaf of Snowball. V. Serrate leaf of Hen-bit. ; (After Thomé.) Toothed or dentate, when the margin is cut into sharp teeth and the teeth point out, as in the lower leaves of the daisy fleabane (Erigeron annuus). Fic. 450. Cleft and pinnatifid leaves of Wild Lettuce. (After Thomé.) 596 WEED FLORA OF IOWA Serrate, when the teeth point forward, as in the common sun- flower (Helianthus annuus). Serrulate, when the margin is finely serrate as in milk purslane (Euphorbia maculata). Crenate, when the teeth are broad and rounded as in the common mallow (Malva rotundifolia). 100 ONONN HHH Fig. 451 1 Fig. 451 II Fig. 451 It Fic. 451. Compound leaves. I. Leaf of Clover with three leaflets. I1. Pedately divided leaf of Dragon Root. III. Pinnate leaf of Locust with 19 leaflets. (After Thomé.) Lobed, when the leaf is deeply cut, as in the great ragweed. The projecting portions are then called lobes. When the incisions are sharp the term cleft is often used; the leaf is pinnatifid when the incision extends almost to the midrib, as in the prickly lettuce (Lactuca scariola) ; it is pinnate when the incisions have extended to the midrib and each separated portion takes the character of a leaf. Each of the smaller portions is a leaflet, and the leaf is now considered compound. The spotted cowbane (Cicuta maculata), spring vetch (Vicia sativa) and cinquefoil (Potentiula monspelien- sis) are good examples. MORPHOLOGY OF LEAVES AND FLOWERS 597 Fig. 452 I Fig. 452 II Fig. 452. I. Wedge-shaped leaflet of Horse Chestnut. II. Pointed leaf of the Wall Pellitory. (After Thomé.) All of the above marginal characters may characterize the lobes of a simple leaf or the leaflets of a compound leaf. LEAF STRUCTURE. Before considering the work of the leaf it will be necessary to become acquainted with its structure. The leaf is covered with an epidermis which is composed of com- pact layers of cells, so modified as to protect the more delicate inner parts. The epidermis may be peeled off as a delicate trans- parent skin. A microscope shows that this transparent skin is made up of many cells, so closely fitted together as to make a contin- uous sheet or covering. Many slitlike openings between two cres- cent-shaped cells (guard cells) appear quite evenly distributed in the epidermis. The opening and guard cells constitute the stoma (plural stomata) which really means mouth. These numerous openings are passage ways into the interior of the leaf and per- mit interchange of gases between outside air and the air in the leaf interior. The guard cells can change their shape and so vary the size of the opening. In horizontal leaves the stomata are chiefly and sometimes exclusively on the lower surface, a fair average number being about 62,500 to the square inch. . 598 WEED FLORA OF IOWA O) pOwCOEDS f aSnases avaly d p Fic. 453. I. A cross section of a leaf of Peppergrass (Lepidium) showing the upper epidermis (e), the lower epidermis (é), stoma (s), the chlor- enchyma (c) consisting of closely placed palisade cells (p) and more loosely placed spongy tissue (f), and a vascular or conductive tract (v) With bundle sheath (b), hadrome or xylem (h), and leptome or phloem (1). II. Surface view of stoma from Haster Lily; g, the kidney-shaped guard cells en- closing the stomatal aperture (Ss); b, the subsidiary cells. III. Cross section of stoma; g, guard cell; s, central slit; 0, outer slit; i, inner vestibule; c, stomatal cavity ; b, subsidiary cell. IV. Surface view of a grass stoma (Poa pratensis) showing the guard cells (g), With their dumb-bell-shaped lumina; b, subsidiary cells with prominent nucellus (n). V. Median cross section and cross section through end of stoma of Poa annua; g, guard cell lumina; b, lumina of subsidiary cells. VI. | ee Yellow foxtail .. £22 2 0|Se 28 22u)_ 2 ceo eee ee Green: foxtall: 5a ae a eee 098) |e 1785 Pennsylvania smartweed _~--------- 15 38 65 49° De See eee Mustard) 222222 93° Niue S ee ee ee ‘Horseweed: ] 2 eeeeesases| 222 a/b 9 10 11 Secribner’s panic grass_|_-_-_--_|-----_-- +4 |. =-2-- |. eee Tansy mustard —__--~-_|_-------|-------- Fae -Yanens emer fete SS |e Ny Cteleay- Sesame 6 as 2 Sree | Se eee § _|-23 eae Yellow dodke2222)). | |h eae ———E—E————ee +158 Large spottedtspurce..|22 = Se ee eee eee 129 Old: witchworasse: 2-2) 22 ee ee ee ee 2 Wild fourocloelk 22.222) ee ee 12 Selif-he ail eee | ee ee 2 *In sheltered place on the south side of the railroad grade. Weeds were not all counted on April 27th, but there were hundreds of small ragweeds and smartweeds, and probably more than a thousand green foxtail plants. ++Winter annual. +Perennial. These tables reveal some interesting facts. The first weeds to appear in March were the winter annuals, like peppergrass and shepherd’s purse, certain perennial weeds, and in one ease horse- weed (Erigeron canadensis) in a timothy meadow. Two weeks later smartweed (Polygonum) was abundant. Spurge (Huphor- bia), an annual, was abundant in the latter part of April and early May. Foxtails in some cases surpassed all other weeds in abundance early in May. The smartweeds (Polygonum) and spurges increased enormously. These weeds were entirely removed from the plots. Fall plowing and clean cultivation certainly in- dicate a smaller number of weeds. Long, who measured off a square yard of ground in* Great Britain, roughly grouping its plant life therein into species, found on this square yard 1,050 seedlings or 5,082,000 per acre. There were 654 buttercup seedlings, 107 of annual meadow grass, 60 of dock, 26 of goosefoot, 25 of groundsel, 15 of shepherd’s purse, 14 of *Trans. of Highland Agri. Soc. Scotland V. 28: 52. NUMBER AND KINDS OF WEEDS ON DIFFERENT SOILS 661 Fic. 515. A weedy cornfield, mostly Foxtail and Smartweeds. (Photographed by Colburn.) annual sow thistle, and 10 of chickweed, besides 139 of other species. He states that Korsmo’s investigation revealed the pres- ence of an even larger number of seeds having the power of ger- mination, the seeds per square yard to a depth of 9.8 inches being as follows: Fallow field, 8,682 weed seeds (over 42,000,000 per acre) ; field for spring grain bearing the same crop for four suc- cessive years, 28,213 weed seeds (over 136,000,000 per acre) ; fallow field, 1,474 weed seeds (over 7,000,000 per acre). Mr. Long gives the following species of weeds found in Great Britain within an area 100 feet square: 662 Fic. 516. WEED FLORA OF IOWA ChE we ay ay, N WY " SING 2 ny’ V ¢ gy. S| \V a i Shepherd’s Purse, a common winter annual. (After Vasey, U. S. Dept. Agr.) NUMBER AND KINDS OF WEEDS ON DIFFERENT SOILS 663 Fig. 517. A cornfield from which Quack Grass was removed by hoeing and cultivation. In adjacent check it formed a sod. The check was given the ordinary methods of cultivation. (Photographed by Colburn. ) *Convolvulus arvensis (Euro- pean bindweed.) *Polygonum convolvulus (Black bindweed ) *Polygonum aviculare (Knot- grass) *Rumex sp. (Dock) *Tussilago farfara ‘(Coltsfoot) *Mentha arvensis (Pepper- mint) *Sinapis arvensis (Common mustard ) *Sonchus arvensis (Field sow thistle) *Stellaria media (Chickweed) Papaver sp. (Poppy) *Ranunculus arvensis (Corn crowfoot) *Viola sp. (Violet) Potentilla anserina (Silver weed) Aethusa cynapium (Fool’s parsley) ; Scandia pecten-veneris (Ven- us’ comb) Senecio vulgaris (Groundsel) *Galium aparine (Cleavers) Victa sativa (Vetch) 664 WEED FLORA OF IOWA *Matricaria wmodora (Wild *Agrostis sp. (Bent grass) chamomile) Agropyron repens (Quack Plantago major (Common grass) plantain) Poa annua (Low spear grass) Lychnis alba (White cam- *Veronica sp. (Speedwell) pion) , | Myosotis sp. (Forget-me-not) “Euphorbia exigua (Spurge) Alchemilla arvensis (Parsley- “Alopecurus agrestis (Foxtail piert) erass) Most of these species are troublesome weeds, and it may be added that those marked with * were abundant. For an Iowa cultivated field on June 2, 1903, the following weeds were found in one square rod. Name of weed. No. per sq. rod. Smartweed (Polygonum pennsylvanicum) ........0..0.e ee eeeeee 40,324 Hedge mustard (Sisymbrium officinale)..............5++-+- to 3,060 Black beggar-ticks (Bidens frondosd)...............0ceceee cere 476 Prickly lettuce (Lactuca scariola var. integrata)..........+--- os 204 Pigweed \(Ohewepon1un .al 1m) sane yee eee ee 340 Horseweed (Hrigeron CANAdENSIS) .... sce eee e ween cere cece ceneee 7,412 Dandelion (Tarawacum Officinale) .......e cece reece eee eee eceeee 68 Moxtail (CS€t@rig) sce c2 cdiemacaies oe Satie olgae oda aier gees mee eine 136,000 WEEDS AND CONTINUOUS CROPPING. Everyone has observed that continuous cropping increases the number of weeds; not only is this true for many. parts of Iowa but it is equally true for the older sections of the United States and of Europe. A. D. Hall who reported on the crops grown in grain land at Rothannstad* said: After continuous cropping for forty-seven years uel weeds of all descriptions occupy considerably more space after continuous crop- ping than before. The relative proportion they bear to the grasses and clover has increased from year to year. Such weeds as barnet, hawkbit and black knapweed became abundant. Mr. Long gives the following list of worst weeds made by four expert Scotch agriculturists : Arable Land.—Charlock, runch, chickweed, spurrey, docks, thistle, groundsel, coltsfoot, day nettle, red-shank, annual meadow grass, bulbous oat grass (pearl grass), couch grass, fine bent grass or black couch, and wild oats. *Jour. Roy Agr. Soc. 64:-88. NUMBER AND KINDS OF WEEDS ON DIFFERENT SOILS 665 Fic. 518. Green Foxtail (Setaria viridis). Common in gardens and fields. ‘ (Photographed by Colburn.) ' Grass Land.—Buttercups, self-heal, docks, ragwort, daisy, thistles, ribwort, plantain, creeping soft grass, common bent grass, York- shire fog or woolly soft grass, moss. There are, however, many other weeds which are given by him which are serious pests to the agriculturist of that country. Of the weeds of the arable land he lists buttercups, poppies, fumitory, charlock (Brassica arvensis), runch (Raphanus raphamstrum) , shep- herd’s purse, corn cockle, spurrey, silver weed, cleavers, thistle (Canada thistle), sow thistle, coltsfoot, groundsel, bindweeds (Poly- gonum convolvulus), (Convoluulus arvensis), dodder, buckhorn, plantain, broom rape, corn or field mint, hemp and dead nettle, smartweed, or red-shank (Plygonum persicaria), knotweed, docks, goosefoot, quack grass, fine bent grass, pearl grass (Arrenatherum avenaceum), slender foxtail (Alopecurus agrestis), wild oats and 666 WEED FLORA OF IOWA Fic. 519. Needle Grass (Stipa spartea). In pastures. (Photographed by Charlotte M. King.) Fie. 520. Purple Cone Flower (Brauneria purpurea). In a prairie pasture. (Photographed by Pammel.) NUMBER AND KINDS OF WEEDS ON DIFFERENT SOILS 667 horsetail. Of grass land weeds there are listed (Colchicum autum- nale), buttercups, leguminous weeds, gorse (Ulex) broom, rest har- row (Ononts spinosa), dyer’s green weed (Gemsta tinctoria), wild carrot, burdock, knapweed (Centaurea nigra), thistles (Cirsiwm arvense, C. acaule, C. lanceolatum, C. palustris), cotton thistle (Onopordum acanthium), daisy (Bellis perennis), ox-eye daisy, rag- wort (Senecio jacobaea), plantains, yellow rattle, self-heal (Pru- nella vulgaris), docks, sorrel (Rumex acetosa, R. acetosella), sting- ing nettle (Urtica dioica) ; grasses: bent grass (Agrostis alba) tus- sock grass (Aira caespitosa), Yorkshire fog (Holcus lanatus), creeping soft grass (H. mollis), quaking grass (Briza media), bar- ley grass (Hordeum pratense), and the bracken (Pieris aquilina), horsetail, and mosses. \ Fic. 521. Common Porcupine or Needle Grass (Stipa spartea). A weed native to gravel soil, soon succumbs to cropping. a, single spikelet; b, fruit with sharp pointed callus. (Lamson-Scribner, U. S. Dept. Agr.) CHAPTER VIII. INJURIOUSNESS OF WEEDS CHAPTER VIII. INJURIOUSNESS OF WEEDS. Weeds are injurious to man in the following ways: 1. They crowd out the growing crop. 2. They consume the moisture necessary for a crop. 3. They consume the mineral or other food elements essential to a crop. 4, They pull down the crop. 5. They are injurious because the seeds are difficult to remove. 6. They are injurious because they harbor insects. 7. They are injurious because they harbor parasitic fungi. 8. They prevent the proper cultivation of the soil. 9. They may cause conditions which breed disease. 10. They may poison the soil. 11. They stop drains. 12. They poison animals and man. 1. WEEDS CROWD OUT GROWING CROPS. When weeds are abundant they crowd out growing crops. Every plant requires a certain amount of space to bring forth a bountiful erop. Two plants cannot grow together in the same place; sooner or later, one plant will crowd out the other. If the weed is a more vigorous grower, like the greater ragweed, it will prevent the oat plant from maturing a crop. It prevents the proper amount of light coming to the plant, and so the food neecssary for a crop can- not be made. 2. WEEDS CONSUME THE MOISTURE. To produce a crop all weeds need moisture, which should go to a more desirable crop. They transpire water just as cultivated plants do. Long, in ‘‘Common Weeds of the Farm and Garden,’’ says: 29 Weeds also absorb from the soil and ‘‘transpire,’’ or pass off into the atmosphere, large quantities of moisture which would be of great service to the growing crop. For example, a maize plant has been observed to transpire in the 16 weeks between May 22d and September 4th as much as 36 times its own weight. A large oak 672 WEED FLORA OF IOWA Fic. 522. Ragweeds (Ambrosia trifida) consume an enormous amount of mois- ture. INJURIOUSNESS OF WEEDS . 673 tree is also stated to transpire 10 to 20 gallons of water in a day; while barley, beans, and clover were found to transpire, during five months of their growth, over 200 times their dry weight of water. Experiments conducted at the agricultural experiment sta- tion of Cornell university showed that during the growth of a 60- bushel crop of maize the plants pumped from the soil, and tran- spired into the air through the leaves, upwards of 900 tons of water. A 25-bushel crop of wheat similarly disposed of 500 tons of water. Weeds also transpire, and if the ground be covered with weeds it is certain that much of the moisture which would be of value to the crop will be lost in the manner indicated. Weeds are especially harmful in this way in a hot summer, and the loss is most felt by the cultivated crop on light sandy soils. 3. WEEDS CONSUME MINERAL AND FOOD ELEMENTS. A weed needs not only the carbon dioxide of the air to make food, but the nitrogenous and mineral elements of the soil to make plant food, all of which should go to the crop. Long gives the fol- lowing in his book, ‘‘Common Weeds of the Farm and Garden’’: Some analyses made at Konigsberg, and lately reported by Profes- sor Stutzer and L. Seidler, show that the amounts of nitrogen, phos- phorie acid, potash, and lime which are removed are deserving of serious consideration. A number of weeds without their roots were collected from oat fields, the soil of which was fairly heavy and poor in humus. In the case of the Wild Radish or White Char- lock the plants had already formed many seed-pods, but the other weeds were in full bloom. The table following shows the percentage of ingredients in the dry matter. These figures indicate in a gen- eral way the amount of the chief plant foods required by weeds. The nitrogren in the Persicaria nearly equaled 20 per cent, and that in the Sow Thistle nearly 15 per cent of albuminoids in the dry mat- ter. Phosphoric acid was chiefly taken up by Spurrey and Persi- earia; potash by the Sow Thistle and Spurrey; and lime by Per- sicaria, Yarrow, and Cornflower. 43 674 WEED FLORA OF IOWA PERCENTAGE CONTENTS OF DRY MATTER. 1 ] f=) (oho) e ee Zz ° E 34 = 2 © 2 iE za GZ ce ou = nN io) Sow thistle (Sonchus oler- ACCUS) 0 MSE eR ai there ans eek 2.39 0.88 4.77 1.94 2.16 14.95 Cornflower (Centaurea cy- CNUS) A Vda oe eich eee 2.30 0.78 1.94 3.13 1.07 8.12 Spurrey (Spergula arven- SUS)! Sasa Oe Ress oe eee 2.36 1.08 4.21 1.52 1.91 10.12 Wild Radish (Raphanus raphanistrum) .......... 1.85 0.78 1.30 1.81 0.71 5.22 Persicaria or red-shank (Polygonum persicaria).'| 3.12 1.16 3.12 4.93 2.53 10.58 Yarrow (Achillea millefol- AULT) ©. ctsisec ee ROOT ue 2.30 0.93 3.15 3.84 ali 9.61 Average of six weeds..... 2.38 0.93 3.08 2.86 1.59 9.76 4. WEEDS PULL DOWN THE CROPS. Weeds like morning-glory, bindweed, wild buckwheat, and others, pull down a cultivated plant and then prevent the formation of a good erop. 5. WEEDS ARE INJURIOUS BECAUSE IT IS DIFFICULT TO RE- MOVE THE SEEDS. It is difficult in many cases to remove weed seeds from seed of various kinds. Buckhorn can be removed with difficulty from clover seed; peppergrass with difficulty from timothy seed; greater ragweed from wheat; wild oats from oats; quack grass from brome grass; cockle from wheat. These impurities often greatly reduce the quality of the grain or seed. 6. WEEDS HARBOR INSECTS. Long gives the following insects found on weeds: Uf ZA ENGIN LAWS Ez . . i) Fic. 523. Morning-glory pulls down the corn and other crops. (After Vasey, U. S. Dept. of Agriculture. ) 676 WEED FLORA OF IOWA SS Fic. 524. Impurities found in Red Clover: 1, Bull Thistle; 2, Canada Thistle; 3, Green Foxtail (Setaria viridis) ; 4, common Plantain (Plantago rugelit) ; 5, Peppergrass (also found in Timothy); 6, Chicory; 7, Pigeon Grass (Setaria glauca) ; 8, Crabgrass (Digitaria sanguinalis) ; 9, Old Witch Grass ; 10, Timothy; 11, Wild Carrot; 12, Pigweed (Amaranthus retroflexus) ; 13, Smartweed (Polygonum persicaria) ; 14, Lamb’s quarters; 15, Dropseed Grass (Muhlenbergia). (Drawings by Charlotte M. King.) Weed “host.” Insect. Turnip Flea Beetle or “Fly” (Phyl- Charlock (Sinapis arvensis) and lotreta (Haltica) nemorum) similar Crucifers Cabbage and Turnip Gall Weevil - (Ceutorhynchus sutlcicollis) Goosefoot (Chenopodium album) Thistles (Cirsium sp) Sow Thistle (Sonchus) . Mangold Fly (Pegomyia betae) Dandelion (Taraxacum) and appar- ently Docks (Rumez) INJURIOUSNESS OF WEEDS 677 Charlock and other Crucifers Diamond-Back Moth (Plutella mac- Prickly saltwort (Salsola kali) ulipennis) Thistles, and the Cotton Thistle (Onopordon Acanthium) } Celery Fly (Acidia heraclei) Hops, Yorkshire Fog, Poa annua, aN Daisy, Shepherd’s Purse, Spurrey, : Buttercup, Cornflower, Sow This- + See Merete devas tle, Black Bindweed (Polygonum Convolvulus) and Plantain Nightshades, Henbane, Hedge, cana Colorado Beetle (Doryphora decem- tard, Thistles, Goosefoot, and lineata) many other plants Shepherd’s Purse, Winter Cress, | Cabbage Root Fly (Phorbia bras- Hedge Mustard sicae) Various Grasses } Frit Fly (Oscinis frit) Docks, Goosefoot, and some ee weeds Bean Aphis (Aphis rumicis) Ghost or Otter Moth (Hepialus Docks, Thistles, Burdock lupuli) 7. WEEDS ARE INJURIOUS BECAUSE THEY HARBOR PARA- SITIC FUNGI. ' Long and Percival, in ‘‘Common Weeds of the Farm and Gar- den,’’ have the following table of fungus diseases that affect va- rious weeds: Weed “host.” Disease. Finger-and-toe of turnips (Plasmodio- phora brassicae Wor.) Peronospora parasitica De Bary Charlock and other Crucifers White Root-rot (Rosellinia necatria Prill.) Sclerotium Disease (Sclerotinia scleroti- orum Mass.) Many species inum) sporium gramineum ESSE) Shepherd’s Purse and other Crucifers White Rust of Cabbages (Cystopus can- didus Lev.) Many species a. Root-rot (Rhizoctonia violacea Agrostis canina L., and other grasses Reed-mace Fungus (EHpichloe typhina Wild Barley (Hordeum mur: u Blindness in Barley and Oats (Helmintho- i } Tul.) 678 WEED FLORA OF IOWA Barberry { Rust of wheat (Puccinia graminis Pers.) Aakers Gooseberry-leaf Cluster-cup (Puccinia Sedges (Carex) \ pringsheimiana Kleb.) Goosefoot t Peronospora effusa Rab., on spinach. Docks, Sorrel, and many) Rosellinia radiciperda Mass. A white others Root-rot in New Zealand. \ Chrysanthemum Rust (Puccinia hieracii Hawkweed (Hieracium) Mart.) Groundsel, Ragwort, and) Pine Cluster-cups (Peridermium pini other species of Senecio Waillr.) Many wild grasses t Ergot of rye (Claviceps purpurea Tul.) SOD : (Sexes QOS (Tee, - ) RS Ce e Ets WSs an CD @ cea ) JOOS ss aes KS 52000) ) (EE Ce lo = il D0. Solavacedow Nie Mismadesmamily 55 2. Seams eiee es 5 ee: 9 NE NUNLO CLO Cracmmam leven almiliy so... 5. ln me eran rq eee ows. a aa ae alt Benes UTULCOCEUe wNGtUleMU aM Yin 020. ct le cnt ge) Meee etch ses + oom « 5) jon. Valeranacedos Valerian Wamily.. 5 i205 ee oon Sens oe i pia, Verbenaceaeverbenaub amily : 022 2-38 leis sare oie e'o es it Dm) a ViLOLACCRC MNO ley MCA eh es «Sie ike «aw date ty Apricorn cte «14 il The weeds of the Pacific coast are somewhat different from those of the Atlantic coast. Eugene Hilgard, in a series of interesting articles in Garden and Forest, some years ago, S. B. Parish, J. Burtt Davy and other California botanists, have contributed notes on the introduction of these weeds of the coast. They enumerated the more important weeds, among which mention may be made of black mustard: (Brassica nigra), catchfly (Silene gallica), storksbill (Erodium cicutariwm), musk erodium (Hrodium moschatum), small-flowered mallow (Malva borealis), black medick (Medicago lupulina), bur clover (M. denticulata), white sweet clover (Meli- lotus alba), and yellow sweet clover (M. officinalis), star thistle (Centaurea cyanus), knapweed (C. solstitialis), brown knapweed (C. jacea), black knapweed (C. nigra), wild carrot (Daucus car- ota), beet (Beta vulgaris), spinach (Spinacia oleracea), lady’s thistle (Silybum marianum), cotton thistle (Onopordon acanthiwm), dodder (Cuscuta epithymum). The writer recently noted the following weeds in the San Fran- cisco Bay region, Oakland, and Sacramento, many of our eastern weeds being common among them: Hedge mustard (Sisymbrium officinale), Raphanus sativus, Marrubium vulgare, dock (Rumex crispus), cheeses (Malva rotundifolia), mallow (M. crispa), bull thistle (Cirsium lanceolatum), sow thistle (Sonchus oleraceus), eat’s-ear (Hypochaeris radicata), knotweed (Polygonum erectum), black medick (Medicago lupulina), Brassica campestris, lamb’s quarters (Chenosodium album), chickweed (Stellaria media), Poa annua, Alfilaria (Hrodwm cicutaria), pineapple weed (Matricaria suaveolens), common groundsel (Senecio vulgarts), burdock (Arc- tium lappa), Silybum marianum, Centaurea melitensis, prickly let- tuce (Lactuca scariola), mustard (Brassica campestris), also such native weeds as Amsinckia and Escholtzia. 726 WEED FLORA OF IOWA Fig. 549A 1 Fig. 549A IT Fig. 549A III Fie. 549-A. Seeds of weeds scattered with agricultural seeds. I. Dodder in clover and alfalfa seed. II. White Sweet Clover in alfalfa seed. III. Bitter Dock (Rumex obtusifolius) in red clover seed. Many of these weeds have made their way into California by way of South America. Others are indigenous and widely scat- tered on the Pacfiic coast. The Pacific northwest has in common with the rest of the Pacific coast many European weeds, but also many that are indigenous to the country. Many years ago Cusick called attention to the predominance of the Northern fireweed (Epilobium angustifolium), which, after the removal of the forest and the burning of the slashings, comes up in great abundance. The writer in another connection* discussed the weeds commonly *The Problem of Weeds in the West; Proc. Ia. Acad. Se. 15: 34. WEED MIGRATION 727 Fie. 550. Curled Dock (Rumex crispus). .A common European weed now occurring across the continent. (After Chesnut, U. S. Dept. Agr.) found in the northwest. A few of the abundant species may be mentioned. Russian thistle (Salsola kali var. tenutfolia) is abun- dant in the drier regions, especially east of the Cascades. In the Puget Sound country are found such common European weeds as Canada thistle (Cirsium arvense), bull thistle (C. lanceolatum), batchelor’s button (Centaurea cyanus), chicory (Cichorium inty- bus), common mustard (Brassica arvensis), mullein (Verbascum thapsus) and also V. blattaria, and toadflax (Linaria vulgaris). It is not uncommon to find in clearings of the forest the small Kenilworth ivy (Linaria cymbalaria), the common pansy (Viola tricolor), the petunia (Petunia grandiflora), the dead nettles (La- mum amplexicaule and L. album), ox-eye daisy (Chrysanthemum leucanthemum), hemp nettle (Galeopsis tetrahit), foxglove (Digi- talis purpurea), European bindweed (Convolvulus arvensis). The 728 WEED FLORA OF IOWA eats’-ear (Hypochaeris radicata) is one of the most common weeds in lawns. There is a common belief that it was introduced from Chile. In the Great Basin country, especially in Utah, a large number of the weeds are of Huropean origin. Of the most common of these mention may be made of the brome grasses (Bromus brizae- formis and B. tectorum), bouncing Betty (Saponaria vaccaria), black medick (Medicago lupulina), dodders (Cuscuta arvensis and C. epithymum), poison hemlock (Coniwm maculatum), moth mul- lein (Verbascum blattaria), mint (Mentha viridis), Russian thistle (Salsola kali var. tenuifolia), horehound (Marrubium vulgare), Fie. 551. Awned Brome Grass (Bromus tectorum).. Common in the Great Basin country and California, occasionally in Iowa. WEED MIGRATION 729 prickly lettuce (Lactuca scariola) and storksbill (Hrodiwm cicu- tartwm). There are also such native weeds as bee-plant (Cleome serrulata), wild liquorice (Glycyrrhiza lepidota), the squirrel-tail grasses (Hordeum jubatum and H. caespitosum), prickly lettuce _(Lactuca pulchella), pine-apple weed (Matricaria suaveolens) and prickly poppy (Argemone platyceras). In the Rocky mountain re- gion, especially Colorado, many of the more recent introductions are from Europe. Among these are the sweet clovers (Melilotus alba and M. officinalis), black medick (Medicago denticulata), purslane (Portulaca oleracea), dodder (Cuscuta arvensis and C. 2: b Fig. 551A I Fig. 551A IL G @ oy 92 Fig. 551A IIL : Fig. 551A IV Fic. 551-A. Seeds scattered with agricultural seeds... I. Barnyard Grass (Echinochloa crusgalli). 11. Medicago denticulata, common in alfalfa seed. II. Horehound (Marrubium vulgare), a weed commonly scattered with alfalfa seed. Common in the Great Basin country. IV. Wild Buck- wheat or Bindweed (Polygonum convolvulus), commonly scattered with grain seed. (All after Hillman.) epithymum), foxtail (Setaria viridis), storksbill (Erodiwm cicu- tarium), Russian thistle (Salsola kali var. tenuifolia), vegetable oyster (Tragopogon porrifolius), goat’s beard (T. pratensis) and lamb’s quarters (Chenopodium album). A large number of native plants have become weedy, such as marsh elder (Iva xanthifolia), 730 WEED FLORA OF IOWA (Franseria discolor), the two annual sunflowers (Helianthus an- nuus, H. petiolaris), small ragweed (Ambrosia artemisiaefolia), buffalo bur (Solanum rostratum), Rocky Mountain bee plant (Cleome serrulata), gumweed (Grindelia squarrosa) and squirrel- tail grass (Hordewm jubatum). Fig. 551B IIT Fig. 551B IV Fic. 551-B. Seeds of immigrant weeds. I. Squirreltail (Hordeum jubatum) ; widely scattered with hay and stock trains from the western plains. II. Spinage (Spinacia oleracea) ; a common weed of the Great Basin country. Seattered from cultivated plants. III. Russian Thistle (Salsola kali var. tenuifolia) ; brought to Dakota with flax seed and grain seed; now scat- tered in Iowa with alfalfa seed. IV. Winged Pigweed (Cycloma atri- plicifolium). (II, drawing, L. R. Collins; the others after Hillman.) Cosmopolitan Weeds.—A great many weeds, originally, had a very wide distribution, although many of these so-called weeds were probably introduced by man. In many eases it is almost impossible to tell how and whence they came to the places in which they occur. . ; Who can trace the immigration of such weeds as common purs- lane, or charlock, or barnyard grass? In the first: place, these WEED MIGRATION So tetod plants immigrated when little was known about the species of plants. What was said about them by the early botanical writers was often very indefinite. In the second place, many of the early writers did not take pains to leave statistics concerning the intro- duction of the plants. The future records, however, will be more accurate as the adventive plants are being recorded by a host of botanical writers the world over. The notes in such floras as Britton’s Manual, Robinson and Fernald-Gray’s Manual, Bentham and Hooker’s Handbook of the British Flora, Garcke’s Flora of Germany, Acloque’s Flora of France, Moore’s Handbook of the Flora of New South Wales, Arcangeli’s Flora of Italy, Baron Fer- dinand von Mueller’s Systematic Census of Australian Plants, Grisebach’s Flora of the British West Indies, Millspaugh’s Flora of Yucatan, Hemsley’s Botany of Central America, Urban’s papers on the Flora of the West Indies give more or less detailed infor- mation on introduced weeds. Fie. 552. Crabgrass (Digitaria sanguinalis). A cosmopolitan weed. The following weeds are more or less cosmopolitan: crab grass (Digitaria sanguinalis), found in North and South America, Eu- rope, Asia, Africa, Australia and New Zealand; barnyard grass (Echinochloa crusgallt), in Europe, Asia, Africa, North and South America and Australia; green foxtail (Setaria viridis), in Europe, 732 WEED FLORA OF IOWA Fig. 553. Foxtail Grass (Alopecurus geniculatus). Widely distributed. Cos- mopolitan grass. (U. S. Dept. of Agr.) Asia, Africa, Australia, North and South America; pigeon grass (Setaria glauca), in Europe, Asia, Africa, North and South Amer- ica; whorled millet (Setaria verticillata), in Europe, Asia, Africa, North and South America; Johnson grass (Sorghum halepense), in Europe, Asia, Australia and North America; foxtail grass (Alopecurus genculatus), im Asia, New Zealand, Australia, North America; hair grass (Agrostis hyemalis), in Australia, New Zealand, North America; Bermuda grass (Cynodon dactylon), a valuable forage plant, but, in cultivated fields, a weed, Europe, Asia, Africa, Australia, New Zealand, North and South America; southern spear grass (Hragrostis pilosa), Europe, Asia, Africa, Australia, New Zealand, North and South America; crowfoot grass WHED MIGRATION 733 (Eleusine indica), Europe, Asia, Africa, Australia, New Zealand, North and South America; cheat (Bromus secalinus), Europe, Asia, North America, a weed; brome grass (Bromus tectorum), Europe, Asia, Africa, North America; soft chess (Bromus arven- sis), Europe, Asia, Africa, North America... It is singular that only one species of Bromus is given by Moore and Ferdinand von Mueller, the Bromus arenarius. Reed (Phragmites communis), though not regarded as a weed in the United States, is a cosmo- politan plant-found in Europe, Asia, Africa, Australia, New Zealand, Papua, North America (Canada, United States and Mexico). Yy if \ y f \\ J 7 Fic. 554. Tumbling Mustard (Sisymbrium altissimum). Introduced with grain in the Dakctas. (Dewey, U. S. Dept. Agr.) Cress (Barbarea vulgaris) occurs in Europe, Asia, Australia, New Zealand and North America. None of the most common North American weeds among the remaining members of the mus- 734 WEED FLORA OF IOWA tard family occur in Australia or New Zealand, although shep- herd’s purse (Capsella bursa-pastoris), common mustard (Brassica arvensis), the hedge mustards (Sisymbrium officinale and S. altis- stmum) and peppergrass (Lepidiwm apetalum) are common in Europe and Asia. Of the pulse family the bird’s-foot trefoil (Lotus corniculatus) — occurs in Europe, Asia, Africa and Australia; the indigo plant (Indigofera hirsuta) in Africa, Australia, Papua and Asia. The absence of black medick (Medicago lupulina) and hop clover (Tri- folium agrartum) from Australia is striking. Of the geranium family the common European, African and Asiatic species of storksbill (Hrodium cicutariwm) are absent in Australia and New Zealand and the little yellow flowered sorrel (Oxalis cormculata) is the only representative in New Zealand, Australia and Papua. This species also occurs in Europe, Asia, Africa and America. The only malvaceous weeds in Australia common also to the United States are sida (Sida spinosa) and butter-print (Abutilon theophrasti), both of tropical origin. The former occurs in Europe, Asia, Africa, North and South America, Australia and Papua. The pigweed (Amaranthus retroflexus) of southern North America is common in Europe but has not reached Australia. None of our troublesome weedy species of dock, which are cosmopolitan, occurs in Australia. Of the smartweeds there are two species, the water pepper (Polygonum hydropiper) (Europe, Asia, North and South America) and slender smartweed (P. lapathifolium) of Europe, Asia, North ‘and South America. Silky cinquefoil (Potentilla anserina), of the rose family, is com- mon in the west and here and there in northern Iowa; it is found in Europe, Asia, Africa, Australia, New Zealand and North America. Feverfew (Agrimonia striata) is found in Europe, Africa and North America. None of our Oenotheras is cosmo- politan; however the primrose willow (Jussiaea suffruticosa), a sub-tropical plant, is widely distributed in Asia, Africa, America and Papua. Very few of the Umbelliferae are cosmopolitan. Moore and Mueller record water parsnip (Siwm latefolwm) for New Zealand and Australia. The Mexican Ageratum conyzoides of the sunflower family, found in Africa, Asia, Europe, North and South America, is often weedy. The Spanish needle (Bidens bipinnata) is found in Asia, Europe, North and South America. The small number of plants of this family found in Australia.and New Zealand is remarkable. None WEED MIGRATION "735 of our plantains is of common occurrence. There is no morning- glory. It may be of interest to compare the noxious weeds of Ger- many with those of the United States. Dr. A. Thaer of the Uni- versity of Giessen in 1881 published a small book on the agricul- tural weeds of that country listing the following: (Those printed in small capitals are also weedy in Iowa.) Corn poppy (Papaver rhoeas), mustard (BRASSICA ARVENSIS), charlock (RAPHANUS RAPHANISTRUM), Canada thistle (CiRstUM ARVENSE), corn cockle (AGROSTEMMA GITHAGO), chickweed (STELLARIA MEDIA), sheep sorrel (Rumex acetosella), vetch (Vicia hirsuta), colt’s-foot (Tussilago farfara), corn chamomile (Anthemis arvensis), corn marigold Fic. 555. Cypress Spurge (Euphorbia cyparissias). Common in Europe and waste places in Iowa. In many cases started in the vicinity of cemeteries. (After Strasburger, Noll, Schenck and Karsten.) (Chrysanthemum segetum), groundsel (Senecio vernalis), corn- flower (Centaurea cyanus), sow thistle (SoNCHUS OLERACEUS), small bindweed (CoONVOLVULUS ARVENSIS), clover dodder (CuSCUTA EPITHYMUM), broom rape (Orobanche ramosa), garden orach 736 WEED FLORA OF IOWA (Atriplex hortense), cock’s comb (Rhinanthus crista-galli major and minor), smartweed (PoLYGONUM PERSICARIA), cypress spurge ( EUPHORBIA CYPARISSIAS) , meadow saffron ( Colchicum autumnale), silky bent-grass (Agrostis spica-ventr), wild oats (AVENA FATUA), chess (BROMUS SECATINUS), quack grass (AGROPYRON REPENS), horse-tail (EQUISETUM ARVENSE). L. Danger in his work on weeds and parasites of Germany, pub- lished in 1887, divides the weeds of Germany into two classes: A. Root weeds, B. Seed weeds. A. Root weeds—Quack grass (Agropyron repens), reed grass (Phragmites communis), bindweed (Convolvulus arvensis), deadly nightshade (Atropa belladonna), goatweed (Aegopodium roda- graria), sheep sorrel (Rumex acetosella), sour dock (R. crispus), meadow saffron Colchicum autumnale), marsh marigold (Caltha palustris), mint (Mentha arvensis), thistle (Cirsium arvense, C. alteraceum, C. acaule, C. heterophyllum, C. palustre, C. lan- ceolatum), (Carduus crispus, C. lanceolatus, C. natams), (Onopor- don acanthium), sow thistle (Sonchus arvensis, 8. palustre, S. asper, S. oleraceus), colt’s-foot (Tussilago farfara, T. hybrida, T. peta- sites), yarrow (Achillea millefolium), (Leontodon autumnalis) dandelion (Taraxacum officinale), plantain (Plantago coronopus, P. arenaria, P. major, P. media, P. maritima, P. lanceolata), horse- tails (Hquisetum arvense, E. palustre). B. Seed weeds—Chess (Bromus secalinus), (Alopecurus agrestis), wild oats (Avena fatua), (Agrostis spica-venti), darnel (Lolium temulentum), black nightshade (Solanum nigrum), black henbane (Hyoscyamus niger), Jimson weed (Datura stramonium), fool’s parsley (Aethusa cynapium), hemlock (Coniwm maculatum), cowbane (Crcuta virosa), Venus’ comb (Scandia pecten-veneris), buckwheat (Fagopyrum esculentum), smartweed (Polygonum persicaria), dooryard knotweed (Polygonum aviculare), corn cockle (Agrostemma githago), buttercups (Ranunculus septentrionalis, k. flammula, R. arvensis), nettle (Lamtum), charlock (Raphanus raphamstrum), foxglove (Digitalis purpurea), mustard (Brassica arvensis), mayweed (Anthemis arvensis and A. cotula), groundsel (Senecio vulgaris), horseweed (Erigeron canadensis), field mari- gold (Chrysanthemum segetum), Frenchweed (Galinsoga parvi- flora), spurge (Euphorbia cyparissias), orach (Atriplex hortense), nettles (Urtica wrens, U. dioica). WHED MIGRATION 737 AMERICAN WEEDS INTRODUCED INTO EUROPE. Every year more weeds are introduced into Europe from Amer- ica; however, it is very noticeable that the more abundant of these are from the warmer regions of America, rather than from the colder regions. A good many of these are weeds of the open country of North America. I have looked rather hastily through a few of the floras of Great Britain, Italy, Austria and Germany, in addition to a number of recent references on the adventive flora of Europe, from which these data have been compiled. Apple of Peru (Nicandra physaloides) Germany (Garcke). Aster (Aster novi-belau) Italy (Arcangeli), Germany (Garcke), France (Acloque). Aster (Aster parviflorus) Germany (Garcke). Aster (Aster salicifolius) Germany (Garcke). Bug seed (Corispermum hyssopifolium) Germany (Garcke), Italy (Arcangeli). Bur cucumber (Sicyos anglinus) Germany (Garcke), Italy (Areangell). Clotbur (Xanthiwm spinosum) Italy (Arecangeli), Germany (Garcke, Bitter, Kiefer), Austria (Neilreich), France (Acloque). Collomia (Collomia grandiflora) Germany (Garcke). Cone flower (Rudbeckia hirta) Germany (Garcke). Cone flower (Rudbeckia lacimata) Germany (Garcke, Bitter; Lehmann). Cone flower (Lepachys pinnata) Germany (Bitter). Kvening primrose (Oenothera sinuata) Germany (Bitter). Fleabane (Erigeron annwus) France (Acloque), Italy (Ar- eangeli). Fleabane horseweed (Hrigeron canadensis) France (Acloque), Austria (Neilreich), Germany (Garcke, Lehmann), England (Hooker), Italy (Areangeli). Galinsoga (Galinsoga parviflora) Italy (Arcangeli), Germany (Garcke). Pigweed (Amaranthus retroflexus) Italy (Arcangeli), Germany (Garcke, Alpers), France (Acloque), Austria (Neilreich). Goldenrod (Solidago canadensis) Germany (Garcke, Wegelin), Austria (Neilreich). - Goldenrod (Solidago lanceolata) Germany (Gareke), England (Hooker). 47 738 WEED FLORA OF IOWA Fic. 556. Goldenrod (Solidago rigida). Common in Iowa. (Photographed by Gardner.) Goldenrod (Solidago serotina) Italy (Arcangeli), Germany (Garcke). Mexican tea (Chenopodium ambrosioides) Germany (Garcke, Alpers), Italy (Arcangeli). Nightshade, buffalo bur (Solanum rostratum) Germany (Garcke, Alpers). Nightshade, buffalo bur (Solanum heterodoxum) Germany (Garcke). Nightshade, Three-flowered (Solanum triflorum) Germany | (Garcke, Alpers). Pellitory (Parietaria pennsylvanica) Germany (Garckane Peppergrass (Lepidiwm virginicum) Germany (Garcke), France (Tourlet).. Phacelia (Phacelia tanacetifolia) Gemede (Garcke). Pigweed, Tumbleweed (Amaranthus graecizans) Italy (Arean- geli), Germany (Bitter). Pigweed (Amaranthus retroflexus) Italy (Arecangeli), eine (Garcke, Alpers), France (Acloque), Austria (Neilreich). WEED MIGRATION 739 Fig. 557. Rice Cut Grass (Leersia oryzoides). Common in low grounds, Iowa; not, however, regarded as a weed. Common in Italy. (Lamson-Scribner, U. S. Dept. of Agr.) 740 Sat WEED FLORA OF IOWA Pigweed, winged (Cycloloma platyphyllum) Italy (Arcangeli). Plantain (Plantago aristata) Germany (Bitter). Ragweed, small (Ambrosia artemisiifolia) Germany (Garcke, Laubert). Rice cut-grass (Leersia oryzoides) Italy (Arcangeli), Germany (Garcke,. Buchenau), France (Acloque), Austria (Neilreich), England (Hooker). Sida (Sida spinosa) Germany (Alpers). Spanish needle (Bidens bipinnata) Italy (Areangeli), Germany (Garcke, Boute), France (Acloque). Spanish needle (Bidens frondosa) Italy (Arcangeli). Spanish needle (Bidens leucanthus) Germany (Garcke). Speargerass (Hragrostis pilosa) Italy (Arcangeli), Germany (Garcke), France (Acloque). Speedwell (Veronica peregrina) Italy (Arcangeli), Austria (Neilreich). Sunflower, artichoke (Helianthus tuberosus) Italy (Areangeli), Germany (Garcke). Sunflower, common (Helianthus annuus) Germany (Garcke), Austria (Neilreich). _ Tickle grass (Panicum capillare) Germany (Alpers, Care France, Italy (Arecangeli), Austria (Neilreich). Tickseed (Coreopsis tinctoria) Germany (Bitter). Waterweed (Elodea canadensis) Germany (Garcke, Scherer, Weshoff, Bitter), England (Hooker), France (Acloque). INDIGENOUS WEEDS COMMON TO THE NORTHERN HEMISPHERE. In all probability a few of the weeds here listed have been intro- duced, at any rate their origin is in doubt; some occur in Hurope and North America only; those found in Asia are marked (As.). Achillea millefolium (yarrow) (As.). Alopecurus geniculatus (marsh foxtail). Anagallis arvensis (pimpernel) (As.). Anaphalis margaritacea (everlasting). Atriplex patula var. hastata (orach). Barbarea vulgaris (yellow rocket). Bidens cernua (sticktight). Capsella bursa-pastoris (shepherd’s purse). Cerastiwm arvense (mouse-ear chickweed). Convolvulus sepium (morning-glory) (As.). WEHED MIGRATION 741 Sem CAN WP UNAS Fic. 558. Yarrow (Achillea millefolium). Common in the northern hemis- phere. (Drawing by Charlotte M. King.) Cyperus esculentus (northern nut grass) (As.). Epilobium angustifolium (fireweed) (As.). Erigeron acris (fleabane). © Erysimum chewranthoides (treacle mustard). Galwum trifidum (bedstraw) (As.). Galium aparine (cleavers) (As.). Gnaphalium uliginosum (cudweed). Hierochloe borealis (holy grass)... Juncus tenuis (wire-grass). Lepidium apetalum (peppergrass). Mentha arvensis (mint) (As.). Myosurus minimus (mouse-tail). Oxalis corniculata (field sorrel). 742 WEED FLORA OF IOWA Plantago major (common plantain) (As.). Polygonum aviculare (dooryard knotweed) (As.). Polygonum hydropiper (smartweed). Polygonum lapathifolium (smartweed). Potentilla anserma (silvery cinquefoil). Potentilla monspeliensis (five-finger) (As.). Fic. 559. Holy or Vanilla Grass (Hierochloe borealis). Common in the northern hemisphere, mountain regions. In northern Iowa. (Lamson-Scribner, U. S. Dept. Agr.) Prunella vulgaris (self-heal). Radicula palustris (marsh cress) (As.). Ranunculus cymbalaria (erowfoot). Ranunculus pennsylvanicus (crowfoot) (As.). Ranunculus repens (crowfoot). Rhinanthus crista-galli (yellow rattle). WEED MIGRATION 743 Senecio palustris (ragwort). Stachys palustris (woundwort) (As.). Taraxacum officinale (dandelion). This species occurs every- where in the United States and Europe, even on the highest moun- tains. Fic. 560. Dandelion (Taraxacum officinale). Common in the northern hemis- phere, across the continent. INTRODUCTION OF SOME WEEDS INTO THE UNITED STATES AND INTO IOWA. © CHENOPODIACEAEH, GOOSEFOOT FAMILY. Russian Thistle (Salsola kali var. tenwifolia). This plant has long been known as a troublesome weed. Henfrey, in his work, “‘The Vegetation of Europe, its Condition and Causes,”’ published in 1852, notes its abundance in Russia, east of the Volga. 744 WEED FLORA OF IOWA Linnaeus, the Swedish botanist, seems to have known the plant as it occurred in eastern Europe. Prof. L. H. Dewey has given us a good account of the introduction of this weed in the United States.* It seems to have been first observed in Scotland, Bonhomme county, South Dakota, in 1873 or 1874, the seed having been brought in with flax seed. In 1877, it was reported from Yankton county; five years later in the counties to the north and west of Bonhomme. By 1888 it had infested many of the counties east of Missouri river and two years later practically all of the counties of South Dakota and southern North Dakota. About the same time it invaded Fic. 561. Russian Thistle (Salsola kali var. tenuifolia). Common in eastern Europe. Introduced into the Dakotas, 1873 or 1874. Now common in northern United States, particularly in the west. (Drawing by Charlotte M. King.) *Bull. Div. of Bot., U. S. Dept. of Agr. 15:12. 1894. WEED MIGRATION 745 northwestern Iowa, northeastern Nebraska and western Minnesota. In 1898 it was reported from Colorado, Wisconsin, Illinois and other western states. In 1894 Pammel gave the following account of its distribution: Prof. Dewey’s map indicates that the badly infested area extends from the east bank of the Missouri river at Bismarck to Jamestown and Moorehead in North Dakota; south to Sioux City, in Iowa, it also occurs in many isolated places in Minnesota, Wisconsin, Iowa, and at Denver, Colorado. The localities at which he found it in Towa are Rock Rapids, Emmetsburg, Spencer and west to the Missouri river, Council Bluffs, and on the Missouri river opposite Nebraska City. To these we must add Edna, Ames, Little Rock (Ball), Calumet (Louthan), Missouri Valley, Mason City, Hagle Grove, and in all probability it occurs along our great trunk lines across the state. Last season Mr. G. W. Carver found a small specimen of what undoubtedly was Russian thistle along the Chi- cago & Northwestern railroad; the place was revisited this year, and an abundance of the weed was found. he saw the importance of making careful examination for the im- purities of various grass and clover seeds. In 1876 appeared his classical book, Handbuch der Samenkunde. In addition to this work he was the author of many other important papers on via- bility of seeds and other physiological seed problems. Among other important contributions along this line, we may mention the papers and work of Kraft, Luhn, and Harz. The work of Harz summarizes not only the facts pertaining to the anatomy of seeds but also to many other important topics; the literature bearing on the subject of seeds is also given. The work of Burchard on the adulteration of seed with special reference to their origin is particularly noteworthy. In his book he has published statistical records showing the origin of clover seed and the weed seeds found in the same from Middle Europe, Eastern Europe, Southern Europe and North and South America. He sometimes missed the important weed species that are found in our American clover seed, but on the whole it is true that the character of the weed seeds found in our clover and other seeds will enable one to tell where they were grown. Burchard, in his account published in a contribution from the Seed Control Station of Hamburg, states that a large number of seeds investigated by him had impurities. Of two hundred and — eighty seeds tested, one hundred and sixty-seven belonged to the pulse family, seventy-nine to the grass family, twenty-three to forest seeds, and eleven were miscellaneous. He found the mini- mum purity of the clover seed was 54.2 per cent. The highest percentage of purity was found in timothy, 99.42. The lowest germinative energy was found in fescue grass, 0.17 per cent, and *The more important references here given upon the history of seed investi- gation are taken from bulletins 88, 99 and 105, issued by the Iowa State Agri- cultural Experiment Station, which are more completely listed in the bibliography. 828 WEED FLORA OF IOWA the highest in clover, 97.25 per cent. Of the sixty samples of red clover examined for dodder, twenty-eight were free. The most common species was clover dodder, although others were also found. One lot of alfalfa from South America contained the Chilian dodder. In this investigation as in others, he emphasizes the im- portance of determining the kinds of weeds found in the seed, thus: In Australian orchard grass the hairy brome grass was common, and in North American seed, orchard grass, timothy, blue grass and tickle grass. The work of Settegast treats extensively the subject of agricul- tural seeds and seed testing, especially with reference to vitality and seed production. Vandevelde’s work treats of the morphology and physiology of germination and includes a splendid bibliography. Attention should be called to the excellent contributions of Wollny whose reports of his splendid work on seeds and the care of agricultural crops often give considerable detail on the ger- mination and viability of various agricultural seeds. Kienitz gives a detailed account of the methods to be followed in the study of germination of seeds. Fruwirth carried out a study on the color and speeie gravity of clover seeds. He found that the dark violet seeds were heavier than the light colored ones of the same head, and perhaps had greater germinative energy. Samek carried on an experiment, testing seeds for a period of eleven years, showing the results of germination after the first and eleventh years. Hiltner in a somewhat recent paper discusses the limitations of seed testing and the importance of obtaining pure seed. H. von Guttenberg gives the description of five species of Cuscata and a key for their identification. Von Degen notes the germination of dodder under varying conditions. Kinzel has made a study of the effect of freezing seed and its bearing upon agri- cultural weeds. Pieper discusses the method of testing the ger- mination of seeds and presents a new method of determining the germination of grass seed. Dorph-Petersen gives a summary of seed tests carried on by the Danish Seed Control Station, during which 9,024 samples of clover, grass, and other seeds were analyzed. The paper gives a list of weed seeds found and includes notes on germination and purity. F. G. Stebler, of the Zurich Seed Control Station, Switzerland, gives the results of tests made of 10,335 HISTORY AND BIBLIOGRAPHY 829 seeds examined in 1908, with reference to adulteration, purity, vitality, and the presence of Cuscuta. Vilke gives an account of the presence of weed seeds found in Swedish seeds. J. Paczoski notes the important weeds found in the Cherson Government. A. Malzew reports on the more important weeds found in Russia; black bindweed (Polygonum convolvulus) and corn cockle (Agros- temma githago) were common in wheat; oats had, in addition, wild oats (Avena fatua, glabrata), and Neshia panculata. In rye, the black bindweed (Polygonum convolvulus), lamb’s quarters (Cheno- podium album), stickweed (EHchinospermum lappula), and Neslia paniculata were found. Lesage made a study upon the effect of solutions of common salt and alcohol on the germination of garden eress. O. Munerati and T. V. Zapparoli of Italy have made a study of weed seeds with hard coats. F. Johnson and R. Hensman discuss the source of the alien flora of Ireland through weed im- purities found in agricultural seed. A. J. Ewart made tests of various grass and garden seeds with reference to the vitality of seeds after a sea voyage. In a lengthy report on the prolonged vitality of seeds it was found that quite a per cent of seeds ger- minated after half a century. Rees made a study of hard seeds with reference to the cuticular layer. This layer repelled the water, and for this reason seeds retained their vitality. A recent paper by E. Lehmann on the temperature and lght relations and the germination of seeds, gives both an exhaustive summary of the subject and a great deal of original matter. It is difficult to say under what conditions seed will germinate. Older investigators regarded light as having no definite action on the germination of seeds. Crocker found that the structure of the seed coats retarded germination in the case of Xanthium. Shull, Crocker, and others found that oxygen hastened the process. In a recent paper by H. Lehmann, attention is called to the action of light and temperature on the germination of seeds, both of which influence this action. The seeds of Whitlavia grandtflora indicated that germination is retarded at higher temperatures but not at lower. He designates some seeds, such as H'pilobium rosewm, E. hir- sutum (Lehmann), and Veronica peregrina (Heinricher) as light germinating seeds. Dark germinating seeds are represented by Phlox drummondii. Tehmann found that 4 per cent germinate in light and 31 per cent in darkness. Maturity also influences ger- mination of seed. When wheat is ripened at a lower temperature, it requires a longer optimum temperature for germination than seed 830 WEED FLORA OF IOWA ripened at a higher temperature. The seeds of blue grass when planted soon after maturity did not germinate in darkness but under the influence of light 88 per cent germinated. In spring, however, the germination was equally good in darkness and light. Cieslar found that light influenced germination. It would seem from this and other investigations that structure, the oxygen, sub- stratum moisture, temperature, light, in short, the ecology and structure as well as physiology play an important part in germin- ation. There are many other contributions, but most of these papers will be found recorded in the literature cited by Vandevelde, Harz and Nobbe. American work.—Connecticut.—The pioneer work in this coun- try was done by Profs. E. H. Jenkins and Warneke, of the Con- necticut Agricultural Experiment Station. Miss Mary G. Jagger and E. H. Jenkins report the results of analyses for 1908-1909-1910-1911, and note the character of the im- purities of seed found in the Connecticut market. North Carolina——Gerald McCarthy, of the North Carolina Agri- cultural Experiment Station, published an elaborate paper on the subject in which he gave the details of an extensive investigation carried on in North Carolina on germination of seeds. Michigan.—Some excellent work was done by Prof. W. J. Beal who reported results of germination of clover seeds furnished by seedsmen. From seeds grown on moist paper the results of the germination showed that large red clover had a germination of 88 per cent, medium red clover 88 per cent, white clover 84 per cent and alsike clover 64 per cent. Prof. Beal called attention to the difference in germination of seeds when grown in sand and in blotting paper. From seeds grown in sand the following results were obtained: Medium clover had a germination of 76 per cent, white clover 92 per cent, red clover (two separate lots) 70 and 56 per cent. : A later report of Prof. Beal gives two tests of twelve years old clover seeds in both of which the germination was 35.8 per cent. Subsequent reports on the vitality of clover and other seeds were also made. Prof. Beal also reports tests on seeds sent by farmers. Fifty- eight samples ranged from 25 to 97 per cent, seventeen being above 90 per cent. He early called attention to the presence of rib grass in clover seed. Recently he has published an exhaustive bulletin on the seeds of Michigan weeds with excellent figures made by HISTORY AND BIBLIOGRAPHY 831 F. H. Hillman. Dr. R. Zceuw has published the results on the comparative viability of seeds, fungi, and bacteria, when subjected to various chemical agents. Ernest Bessey gives the results of seed analysis for 1911 and 1912 from which it appears that buckhorn and plantain were frequent. The paper gives the number of weed seeds in a pound. Mr. Parsons in 1893 made some interesting tabulations in a summary of American seed vitality tests. In the results offered by him we find the following: Alsike 72.7 per cent, crimson clover 59 per cent, red clover 84.8 per cent, mammoth clover 82.5 per cent, white clover 72.1 per cent, alfalfa 61.6 per cent. Pennsylvania.—Prof. Butz studied many hundreds of samples of seeds, chiefly with respect to their germinative power. In many cases the percentage of germination was very low. Thus we find recorded the following: Alfalfa 52 per cent, alsike clover 61 per cent, Japan clover 69 per cent and seradella 13 per cent. Ohio.—Mr. Devol early recognized and emphasized the im- portance of the experiment station in studying the viability of seeds. In tests conducted by him clover showed a germination of 93 per cent. ; Later Selby and Hicks made a study of fifty-two samples of clover and alfalfa seeds sold in Ohio. They found not only that the seeds had a low vitality but that they contained considerable impurities. Nevada.—Several very important papers have been published by Mr. Hillman on clover seeds and their impurities. One of his earlier bulletins deals with the descriptions of weed seeds and their distribution, together with an incidental account of the occurrence of these seeds in commerce. In a later publication he considers the weed seeds found as impurities in various types of seeds, in- cluding alfalfa, red clover, white clover, alsike clover, crimson clover, Japan clover, Bokhara clover, yellow trefoil and esparcette. In his investigations a large number of samples was examined. The paper therefore gives a fair estimate of the impurities gen- erally found in the various clover seeds offered for sale in this country. ; Kansas.—Prof. Roberts and Mr. Freeman carried on an extensive investigation of alfalfa seed, showing adulteration, substitutes and impurities; and the methods of detecting the latter. They found some adulteration in alfalfa seed. The yellow trefoil (Medicago lupulina) was most frequent though there were occa- 832 WEED FLORA OF IOWA sional instances in which burr clover (Medicago denticulata) and sweet clover (Melilotus alba) occurred. The most noxious weed seeds found were the docks and the English plantain. The average germination of alfalfa seed was 83 per cent. The subject is also briefly discussed by Ten Eyck. Prof. Roberts and Mr. Freeman also made quite an exhaustive study of the grass seeds commonly sold in Kansas. The dodder may prove to be a destructive clover parasite in the United States. Mr. Brown, in a paper on legal and customary weights per bushel of seeds, has also brought together much important matter on the subject of the weight of commercial seeds. Maine.—Prof. Harvey and other members of the staff of the Maine Experiment Station investigated the vitality and the im- purity found in the agricultural seeds offered for sale in that state. The results of Prof. Harvey’s investigations disclosed the advisability of having a law to regulate the sale of seeds. Such a law was passed by the state of Maine and the work of carrying on this law was placed in the hands of the director of the Station. The results of the tests and regulations concerning the seed testing for Maine were published by Charles D. Woods, director of the Station. In a subsequent bulletin he discusses seed inspec- tion and lists 79 kinds of weed seed. With Hammond the require- ments of the law are given as well as results of seed testing. Minnesota.—W. L. Oswald, Bulletin 127 of the Minnesota Station gives the results of purity and germination tests for 1275 samples of commercial seed. North Carolina—McCarthy in an early bulletin of the North Carolina Station reported on the analyses of seed in 1912. O. I. Tillman reported on the analyses of commercial seed in accordance with the North Carolina pure seed act. Kentucky.—Prof. Garman and others, of the Kentucky Experi- ment Station, have investigated the impurities in grass and other forage plants sold in Kentucky. It was found that thirty-six of the five hundred samples examined were adulterated; among these were red clover, blue grass, timothy and orchard grass. They con- cluded that the greatest fraud perpetrated is in the sale of certain varieties under an assumed name. In several publications Prof. Garman discussed the impurities and adulterations of grass seed sold in Kentucky and recommended that no field seeds should be sold in Kentucky containing more than 5 per cent weed seeds. He also gave a comprehensive account of seed testing apparatus HISTORY AND BIBLIOGRAPHY 833 and the conditions under which germination tests are made, with special reference to work in Kentucky. Kentucky now has-in consequence of this work an admirable law forbidding adulteration or misbranding of the clovers and timothy sold for seed in the state. Its effects have been most salutary and have been studied with profit by other common- wealths. Vermont.—Mr. L. W. Barton, under the direction of Profs. Jones and Orton of the Vermont Experiment Station, made an examination of thirty-four samples of clover seed in Vermont. He reports the total percentage of impurities in red clover as being from .3 to 5.3, with an average of 1.8 per cent; alfalfa as having a maximum of 7.1 per cent and a minimum of .6 per cent. Sorrel was found in 60 per cent of the red clover samples and wild carrot in a few, the rib plantain occurred in 77 per cent, dodder in 5 per cent, and Canada thistle in 5 per cent. Dodder did not occur in the alfalfa, but in 8 per cent of the samples Canada thistle was found. Massachusetts.—G. E. Stone reports on seed tests made in 1908 and 1910. Smith, Chapman, and Stone describe fourteen weed seeds most commonly found in grass seed and cattle foods, with a brief account of the weeds of the state. Maryland.—E. I. Oswald made a series of experiments to deter- mine the vitality of seeds when placed in manure under different conditions. It was found that weed seeds when left for six months in manure had lost their vitality completely. When left there for one month under conditions usually followed by dairymen, the seeds of ribgrass, horse nettle, dock, and a few others were still firm. Mention of weed seeds is made in the treatise by J. B. S. Norton on Maryland weeds. The same author gives the result of seed analysis of various commercial seed. Nebraska.—E. M. Wilcox in a paper on dodder in alfalfa seed discusses the frequency of the impurities and means of eradicating the weed. New Hampshire.—F.. W. Taylor in reports of seed tests for 1910 and 1911 discusses the New Hampshire law giving results of purity and germination tests. New York—F. C. Stewart under the head of alfalfa troubles discussed dodder in alfalfa seed and subsequently gave a method of screening dodder out of alfalfa seed. In 1910 G. T. French re- 53 834 WEED FLORA OF IOWA ported on an examination of several hundred samples of seed with special reference to the occurrence of dodder, Russian thistle, and Centaurea repens; an earlier report by the same author reported on seed tests made at the station. Further discussion is found in the New York Agricultural Experiment Station Circular 8:1-4, Dodder in Alfalfa, by F. C. Stewart and G. T. French. Mr. M. T. Munn in Bulletin 362 gives the seed tests made in 1912, giving results of examinations made for purity of commercial samples. North Dakota.—H. L. Bolley discussed the pure seed law of North Dakota and seed work ending December 31, 1909, with an account of weed seeds found. Wisconsin.—A. L. Stone in a circular of information discussed the seed inspection law of the state, adding some general informa- tion. With G. T. Moore he also discussed the question of the eradi- cation of farm weeds which have been introduced by means of im- pure seed. Mr. George T. Harrington discussed the worst weeds in connec- tion with grass and clover seeds. D. L. Beach made a study of commercial feeds with reference to the germination of weed seeds found in these feeds. Feed subjected to steaming before feeding contained no germinable weed seeds. Texas.—According to O. M. Ball alfalfa seed sold in Texas con- tains the following chief impurities: Russian thistle, ribgrass, tumbleweed, pigweed, two kinds of dodder, green foxtail, curled dock, bur clover, and sweet clover. The vitality varied from 49.5 to 96.5 per cent. A subsequent report also on alfalfa seed was made. Arizona.—Prof. Thornber made an examination of alfalfa seed sold in Arizona. This showed a high percentage of germination. Iowa.—The Iowa Agricultural Experiment Station has produced several publications relating to weed seeds. Ball discussed im- purities in grass seed; Pammel, Buchanan and King have pub- hshed a bulletin covering impurities of commercial seed. Two bulletins by I. H. Pammel and Charlotte M. King, ‘‘Results of seed investigations for 1907’’ and ‘‘Results of seed investiga- tions for 1908 and 1909,’’ give the results of analyses of seed sent by farmers to the Station. The results of the 1907 investigation indicated that the weed seeds were different in many cases from those reported in a previous publication; dropseed grass was common, occurring in 3.3 per cent of the samples; Canada thistle seed was also less frequent. The investigations for 1908-1909 in- HISTORY AND BIBLIOGRAPHY 835 dicated that the seed sold in the state has improved, field tests of germination were lower than laboratory tests. A method of deter- mining the vitality is also given. The subject of delayed germina- tion is discussed by H. S. Fawcett, who found that a relationship exists between the hardness of the seed coat and the dormant period. L. H. Pammel states that different conditions influence the germination of seeds, such as hard coats and freezing of seeds. The delayed vitality of weed seeds is discussed by L. H. Pammel and Charlotte M. King. Weed seeds after freezing germinate more readily when frozen. Work of the United States Government.—The National Govern- ment began a serious investigation of agricultural seeds and their impurities in 1893. Early articles on the subject appeared under the head of ‘‘Pure Seed Investigations.’’ Mr. Hicks called attention to the abuses in the seed trade. The matters that were mentioned by him have evidently not been remedied since 1894. Since then the work of the Department has been immeasurably increased, especially by such contributions as have been made by Mr. Duvel on the vitality of buried seeds and the storage and germination of wild rice; and the paper by Mr. Pieters on ‘‘The Farmer’s Interest in Good Seed.’’ Among the notes on seed testing in 1897, Prof. Hicks and Mr. Sothoron Key published an account of the germination of several forage plants and flower garden seeds. A good table on the percentage of germination sanders re- quired of seeds will be found in a paper by Mr..Pieters. These seed standards were also published in the Year Book of 1896. Later Mr. Pieters discussed the presence of clover dodder and other im- purities in clover seed. In a circular by Prof. Dewey of the same division the dodders found in clover were discussed. The quality of the seed, especially its germinative energy, de- pends somewhat on the manner in which seeds are kept. Two ar- ticles on this subject are of special interest in this connection, one by Pieters, ‘‘Agricultural Seeds, Where Grown and How to Handle,’’ and an article by Pieters and Brown, ‘‘Kentucky Blue Grass Seed, Harvesting, Curing and Cleaning.’’ The United States Department of Agriculture has published rules and described apparatus for seed testing. These were adopted by the standing committee on seed testing of the Association of American Agricultural Colleges and Experiment Stations. 836 WEED FLORA OF IOWA In reports of more recent work, A. F. Woods found dodder seed frequent in alfalfa and red clover seed. The adulteration and mis- branding of alfalfa, red clover and grain seeds was discussed by W. A. Taylor and B. T. Galloway. F. H. Hillman in several papers on the impurities, adulteration, etc., of clover and forage plant seeds and vetches, presented admirable figures and good descrip- tions of weed seeds found as impurities. Westgate, McKee, Evans, and Vinall gave a brief account of impurities found in alfalfa seed and sweet clover. The subject of low grade clover seed was con- sidered by Edgar Brown and Miss Crosby. In a paper by Brown and Hillman, ‘‘The Seed of Red Clover and its Impurities,’ the more important impurities found in Eu- ropean and American grown clover seed are given. Attention is called to the introduction of bad seeds from Chile. In 1905 two hundred and seventy-five thousand pounds of Chilean red clover seeds were imported into the United States, and this clover seed contained Chilean clover dodder seed. Canadian work.—In 1892 Prof. J. H. Panton called attention to the importance of making an investigation of seed purity and of carrying on a campaign for better and cleaner seed. He thought a large number of the weeds on the Ontario farms were introduced in clover seed. He found the number of weed seeds varied greatly, all the way from one to four thousand five hundred and forty per half ounce. Among the weed seeds he reported several that have become trouble- some in the Mississippi valley; these include ribgrass and chicory. Saunders in several reports of the Experiment Farms has dis- cussed the vitality of grass and clover seeds. In 1903 there were tested one hundred and eighty-six samples of clover. The highest percentage of germination was low, the lowest 17 and the average 76.3. A large number of grass seeds were also tested. The work has been continued by Saunders and Grisdale (1910) relative to determining the climatic conditions favorable to high vitality of seeds. Since 1903 the Department of Agriculture of the Dominion of Canada has created a seed division with Mr. G. H. Clark in charge of the work in purity and vitality tests. A very notable publication on the subject of weeds and weed seeds has been contributed by Clark and Fletcher who have given colored illustrations of the more important impurities found in clover and other seeds. HISTORY AND BIBLIOGRAPHY 837 A. B. McCready of the Ontario Agricultural College gives the results of germination tests of alfalfa and clover in which the superiority of bright, clean seed is shown. Australia.—A. J. Ewart made tests of various grass and garden seeds with reference to the vitality of seeds after a sea voyage. In a lengthy report on the prolonged vitality of seeds it was found that quite a per cent of seeds germinated after half a century. Rees made a study of hard seeds with reference to the cuticular layer. This layer repelled the water and for the same reason seeds retain their vitality. BIBLIOGRAPHY. CHEMICAL COMPOSITION OF WEEDS. Bailey, EH. H. S., & Sayre, L. E. On the presence of barium in the ash extraet of certain Kansas weeds. Trans. Kans. Acad. Sci. 23-24 :194-198. Cassidy, James, & O’Brine, D. Some Colorado grasses and their chemical analysis. Bull. Col. Agr. Exp. Sta. 12:151. Goessman, C. A. Analyses of weeds. Bull. Mass. Agr. Exp. Sta. i 5-Ge Jenkins, E. H., & Winton, A. L. A compilation of analyses of American feeding stuffs. Bull. Exp. Sta. U. 8S. Dept. Agr. JL lays, Knight, H. G., Hepner, F. F., Nelson, Aven. Wyoming forage plants and their chemical composition. Bulls. Wyo. Agr. Exp. Sta. 60:52; 70:75; 76:119-. 87 :153. Pammel, L. H. The Russian Thistle. Bull. Ia. Agr. Exp. Sta. 38 :24. . Patrick, P. E. Chemical analysis of Russian thistle. Bull. Ia. Agr. Exp. Sta. 26 :26-29. Peter, A. M. Kentucky Forage Plants—Analyses of some Ken- tucky grasses. Bull. Ky. Agr. Exp. Sta. 87 :111-122. Weems, J. B., & Heileman, W. H. The chemical composition of squirreltail grass. Bull. Ia. Agr. Exp. Sta. 30:320-321. Stutzer, A., & Seidler, L. The content of important food ingre- dients in some common weeds. Fiihling’s Landw. Ztg. 57: 429-430. 1908. DISTRIBUTION OF WEEDS. Acloque, A. Flore de France. 816. Paris, 1896, Flores régional de la France. 1904. 838 WEED FLORA OF IOWA Anonymous. Corn Cockle. Jour. Bd. Agr. London, 1910: 38-45. Note in Exp. Sta. Rec. 23 :340. Distribution ; burdock attached to advertisement. See Na- ture 84 :547. The dodder pest. Natal. Agric. Jour. 9 :959. Pasture weeds; their prevention and eradication. Bull. Kan. Acr Exp sta. 119-oere ball. 113: Weeds. List and description prepared by Office of Experi- ment Stations. U. S. Dept. Agr. Bull. Office Exp. Sta. 15 :356-376. . Arcangeli, Giovanni. Compendia della Flora Italiana ossia manu- ale per la determinazione delle piante che trovansi selvatiche od inselvatischite nell Italia e nelle isole adiacenti. 889. Rome, 1882. Arthur, J. C. Wild or prickly lettuce. Bull. Ind. Agr. Exp. Sta. 52 :83-112. Bailey, F. M. The weeds and suspected poisonous plants of Queens- land. 245. Brisbane, 1906. : Beal, W. J. Michigan Weeds. Bull. Mich. Agr. Exp. Sta. 267 :281-458. Beck, L. C. Botany of the northern and midland states, or a description of the plants found in the United States, north of Virginia, arranged according to the natural system; with a synopsis of the genera according to the Linnaean system, a sketch of the rudiments of botany and a glossary of terms. 471. Albany, 1833. 2nd ed. 185. Bentham, G., and Hooker, J. D. Handbook of the British Flora. 584. London, 1904. Bergen, Joseph W. Foundations of Botany. 257. Boston, 1902. Blankenship, J. W. The loco and other poisonous plants of Montana. Bull. Mont. Agr. Exp. Sta. 45:75-104. Notes on weeds (Dodder, Coreopsis and Dandelion). Rept. Mont. Agr. Exp. Sta. 1902:70-71. Weeds of Montana. Bull. Mont. Agr. Exp. Sta. 30:7. Blatchley, W. S. The Indiana weed book. 191. Indianapolis, 1912. Bogue, E. E. An annotated catalog of the ferns and flowering plants of Oklahoma. Bull. Ok. Agr. Exp. Sta. 45. 48. Weeds of Oklahoma. Bull. Ok. Agr. Exp. Sta. 41:12. Bohmer, G. R. Commentatio de plantis segeti infestis; accedit commentatio de plantis auctoritate publica exstirpandis, cus- todiendis, et e foro proscribendis. 128. Wittenberg, 1785. HISTORY AND BIBLIOGRAPHY 839 Bolley, H. Iv. Distribution of weed. seeds by winter winds. Bull. ; -N.°D. Agr. Exp. Sta. 17 :1038-105. Bolley, H. L., & Waldron, L. R. A preliminary list of the sperma- tophyta eed ‘bearing plants of North Dakota. Bull. N. D. Agr. Exp. Sta. 46 :593-675. Bruhl, P. Recent plant immigrants. Jour. and Proc. Asiatic Soe. Bengal, 4:603-656. : Buchenau, F. Die Verbreitung von Oryza: clandestina A. Braun. Bot. Zeit. 52:83-96. 1894. Buckhout, W. A. On troublesome weeds in Pennsylvania. Report of Botanist Penn. Agr. Exp. Sta. 1903 :587-590. Weeds in general ; two newcomers into Pennsylvania [ Allium carinata and Scabiosa australis}. Bull. Pa. Agr. Exp. Sta. boc. | 1902. Budd, J. L. The Russian thistle in its natal home. Bull. Ia. Agr. Exp. Sta. 26 :30-33. Buettner, R. Flora advena mordica. ay Verh. Botan. Ver. Bran- denberg, 25:1-59. 1883. Bull, C. P. Weeds in flax fields. Cy. of Am. Agr. 2:299. Burtt-Davy, Joseph. Dodder in luzerne. Transvaal Agric. Jour. 5 :677. Four noxious weeds [Burweed, cocklebur, Mexican poppy, dodder]. Far. Bull. Transvaal Dept. Agr. 54:13. Some injurious weeds. Trans. Agr. Jour. 3:291-299. Abs. in Exp. Sta. Rec. 16:1091. Cates, J. S., and Cox, H. R. The weed factor in the cultivation of conne bull We Se Dept: Aor, Bur Pll Inds 257 235. Clark, G. H., & Fletcher, Jas. Farm weeds of Canada. With il- lustrations by Norman Criddle. 97; pl 56. Ottawa, W906. Claypole, E. W. On the migration of plants from Europe to Aus- tralia, with an attempt to explain certain phenomena connected therewith. Rep. Montreal Hort. Soc. and Fruit Growers’ Ass. of the Province of Quebee. 3:70-91. 1877. Clinton, G. P. The Russian thistle and some plants that are mis- taken for it. Bull. Ill. Agr. Exp. Sta. 39 :87-118. Cockerel, T. D. A. Some western weeds and alien weeds in the west. Bot. Gaz. 20:503-504. 1895. Coit, J. E., and Packard, W. E. Imperial Valley settlers’ crop manual. Bull. Cal. Agr. Exp. Sta. 210:139-253. 840 WEED FLORA OF IOWA Cotta, H. Forstrath in Tharand—auf der kleinen Zillbach einen Hennebergschen Forsthause am 30. Oct. 1763. 25 p. Tharand 1844. Cook, L. C. Questions concerning weed distribution. Bull. Vt. Bot. Club. 3 :14-15. Covile, F. V. Noxious weeds. Rept. U. S. Dept. Agr. 1890: 388-392. Craig, Moses. The farmers’ foes. (Thistles.) Bull. Ore. Agr. Exp. Sta. 32 :99-115. Some Oregon weeds and how to destroy them. Bull. 19:21. Crosby, J. O. Incursion of weeds into our gardens and farms. Trans. Ia. St. Hort. Soe. 42 :153-154. Cryer, J): Hawkweeds. Naturalist. 1909: 143-144. Cusick, Wm. Forest Fires in Oregon. | Refers to introduction of Epilobium spicatum or ‘‘Elkweed.’’| Bot. Gaz. 8:176. Darlington, Wm. American weeds and useful plants. Rev. Ed. 460. New York, 1859. Dewey, L. H. Distribution of prickly lettuce in the United States. Bot. Gaz. 21 :34-35. Migration of weeds. Yrbk. U. S. Dept. Agr. 1896: 263-285. Table of two hundred weeds. Yrbk. 1895 :594-611. Twenty-five most harmful weeds. Yrbk. 1897 :641-645. Dodson, W. R. Rice weeds in Louisiana. Bull. La. Agr. Exp. Sta. 77 :394-429. Note in Ex. Sta. Ree. 16:168. Ehrhart, F. Beitrage zur Naturkunde und den damit verwandten Wissenschaften, hesonders der Botanik, Chemie, Haus und Landwirtschaft, Arzneigelehrtheit und Apothekerkunst. 7 pt. 1293. Hannover und Innsbruck, 1787-1792. Oekonomische Pflanzenhistorie nebst dem Kern der Land- wirthschaft, Garten-u. Arzneikunst. 12. Ulm and Mem- mingen 1753-1762. Ewart, A. J., and Tovey, J. R. Californian Stinkweed, or sheep- weed (Glia squarrosa H. & A.). Jour. Dept. Agr. Victoria. VI:592. The Thorn Apple, Ib. 11:672. Ewart, A. J. The proclaimed plants of Victoria. Jour. Dept. Agr. Victoria. 5:28, 104, 230, 276, 336, 438, 498, 540, 606, 680, 720. Abs. in Just’s Bot. Jahrb. 22:610. Fernald, M. L. Some recently introduced weeds. ‘Trans. Mass. Hort. Soe. 1905: 11-22. ? HISTORY AND BIBLIOGRAPHY 841 French, C. An introduced weed (Hypericum perforatum). Jour. Dept. Agr. Victoria. 1903: 65-67. Garcke, August. Illustrierte Flora von Deutschland. 795. Ber- lin, 1903. Gleditsch, J. G. Systematische Einleitung in die neue Forstwissen- schaft. 544. Berlin, 1775. Gmelin. J. F. Abhandlungen von dem Arten des Unkrauts auf den Aeckern in Schwaben. 408. Lubeck, 1779.. Goff, E. S. Noxious weeds. Bull. Wis. Agr. Exp. Sta. 39:38. Noxious weeds of Wisconsin. Bull. 76:52. The Russian thistle. Bull. 37:13. Gray, Asa. Burs in the Borage family. Am. Nat. 10:4. Pertinacity and predominance of weeds. Am. Jour. Sci. 18 :161-167. Also in Scientific Papers. 2:234-242. Greene, E. L. Is Rhus glabra in Canada? Ottawa Nat. 22:179-181. Greif, O. Queckenvertilgung. Mitt. Deutsch. Landw. Ges. 1906: 339. Abs. in Just’s Bot. Jahrb. 9 :482. Haglund, E. Characteristic weeds on cultivated marsh soils. See Svenska Mosskultur for Tidskr. 25: 238-245. Note in Exp. Sta. Ree. 26 :538. ; Halstead, B. D. American and English weeds compared. PI. World. 3:171-173. Our worst weeds.