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1969 Cytotaxonomy of the in Louisiana. James Howard Brooks Louisiana State University and Agricultural & Mechanical College

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Recommended Citation Brooks, James Howard, "Cytotaxonomy of the Genus Echinochloa in Louisiana." (1969). LSU Historical Dissertations and Theses. 1640. https://digitalcommons.lsu.edu/gradschool_disstheses/1640

This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 70-9040 BROOKS, James Howard, 1932- CYTOTAXONOMY OF THE GENUS ECHINOCHLOA i IN LOUISIANA. j The Louisiana State University and Agricultural and Mechanical College, Ph.D., 1969 Agronomy

University Microfilms, Inc., Ann Arbor, Michigan CYTOTAXONOMY OF THE GENUS ECHINOCHLOA IN LOUISIANA

A Dissertation

Submitted to the Graduate Faculty of the Louisiana State University Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy

in

The Department of Botany and Pathology

by James Howard Brooks B.S., Stephen F. Austin State College, 1957 M .S., Stephen F. Austin State College, 1964 ACKNOWLEDGEMENTS

The author wishes to express his sincere appreciation to Dr.

Clair A. Brown for serving as chairman of his committee, for aid

in selecting the problem, guidance throughout the study, and helpful

suggestions for improving the manuscript and companionship during

some of the collecting trips. Thanks are due to the other members

of his committee, Dr. St. John P. Chilton, Dr. Irwin Forbes, Dr.

M. T. Henderson, Dr. B. Exner, and Dr. J. B. Baker, for their

encouragement and helpful criticism in the preparation of this

dissertation. The author is grateful to the Department of Botany

and for cooperation and assistance in respect to

utilization of equipment, materials and greenhouse space.

The author also wishes to thank the curators and staff of all the herbaria who supplied their herbarium specimens and to thank

other persons who supplied and seed for this investigation.

Thanks are due also to Mrs. Denise Sylvester for securing several interlibrary loans.

He is grateful to Mrs. Jackie Lockwood for typing the final

draft of this manuscript and her invaluable assistance during the

past three years.

Finally, he wishes to express his deep appreciation to his wife,

Vera, for encouragement at all times and limitless patience in typing this dissertation. TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS...... ii

TABLE OF CONTENTS ...... iii

LIST OF TABLES . . . v

LIST OF FIG U RES...... vi

ABSTRACT...... vii

INTRODUCTION...... 1

HISTORICAL ACCOUNT...... 2

MATERIALS AND M ETHODS...... 8

CYTOLOGY...... 11

Chromos ome s ...... 11

Method of Reproduction...... 17

Correlation of Pollen Size and Chromosome Number .... 18

ANATOMY...... 22

Leaf Epidermis...... ' ...... 22

Embryo-Caryopsis ...... 24

ANALYSIS OF HERBARIUM SPECIM ENS...... 30

Discussion of Taxonomic Characters ...... 34

TAXONOMIC TREATMENT...... 38

SUMMARY...... 69

iii Page

LITERATURE CITED...... ' ...... 71

APPENDIX...... 75

VITA...... 84

iv LIST OF TABLES

TABLE Page

I. Concepts of Echinochloa taxa correlated with Weigand (1921)...... 6

II. Mean diameter of pollen grains in different taxa of Echinochloa ...... 21

III. The difference in caryopsis length and embryo le n g th ...... 29

IV. Mean values of some quantitative characters of Echinochloa taxa...... 32

V. Ranges of some quantitative characters of Echinochloa ta x a ...... 33

v LIST OF FIGURES

FIGURE Page

1-7. Chromosomes of Echinochloa taxa at diakinesis. . . . 15

8. Photomicrograph of a meiotic anaphase I of E. colonum (n = 2 7 ) ...... 16

9. Photomicrograph of an endosperm nucleus showing approximately 81 chromosomes (3n) and 54 chromosomes (3 n ) ...... 19

10-19. Abaxial epidermis of leaves and sheaths of grasses of various Echinochloa ta x a ...... 26

20. Caryopsis of various taxa of the genus Echinochloa . showing a longitudinal cross-sectional view ...... 27

21. Caryopsis of various taxa of the genus Echinochloa, showing the dorsal view ...... 28

22. Polygons of Echinochloa ta x a ...... 31

23. Spikelets of various taxa of the genus Echinochloa. showing the nature of the spikelet and relative size. Each spikelet is shown in two views, dorsal and ventral...... 37

24. Maps showing the distribution of E. crusgalli. E. colonum and E. colonum form JL in Louisiana. . . . 6F

25. Maps showing the distribution of E_. muricata. E.. muricata var. microstachya and E. occidentalis in Louisiana...... 67

26. Maps showing the distribution of E. zelayensis. E. walteri and four forms in Louisiana...... 68

vi ABSTRACT

The present investigation was undertaken to determine the taxonomic relationship of various taxa of the genus Echinochloa in

Louisiana. The investigation included studies of anatomical, cyto- logical and morphological characters of each taxon within the genus.

The characters used for delimitation of the tasa were, anther, caryopsis, fertile floret, embryo, palea of lower floret, sheath, spikelet, sterile lemma, panicle, pustulate based setae on the branches of panicle, length of panicle branches, length and width of blades, ligule, and presence or absence of stamens in the lower floret. Based on the above characters, twelve taxa of the genus

Echinochloa were recognized in Louisiana, namely, E. colonum

(n = 27), E. colonum form JL (n = 27), E. crusgalli (n = 27), E. muricata (n = 18), E. muricata var. microstachya (n = 18), E_. occidentalis (n = 27), E. walteri (n = 18), E. walteri form 1_ (n = 18),

E. walteri form_2 (n = 18), E. walteri form 3^ (n = 18), 13. walteri form A (n = 18) and 12. zelayensis (n = 18). A key is provided for the above taxa, based on material from specimens taken from

L o u isiana.

v ii INTRODUCTION

Echinochloa. the sixth largest grass genus in the tribe

(Hitchcock, 1951), is one of the less economically important genera.

Most of the taxa are grazed by stock but are not usually utilized as forage due to the sparse growth habit. Some of the cultivated varie­ ties of E. colonum and E. crusgalli are used as a source of food in tropical Asia and tropical . In Louisiana, E. colonum and E. crusgalli are weeds in the fields as well as other agronomic crops. Echinochloa is one of the important sources of food in the coastal marsh for the water fowl.

The taxa of Echinochloa are widely distributed throughout the tropical and temperate regions of the world. There is no agreement as to the exact number of polymorphic represented in this genus. A total of twenty-one species has been described in North

America alone. Yabuno (1962) reported about twenty species world w id e .

The most recent treatment of the genus Echinochloa in North

America was by Hitchcock (1951). It was evident from the use of this treatment that an extensive investigation of the genus Echinochloa was necessary.

The present study, utilizing an anatomical, cytological and morphological approach, was made to attempt to offer a better delimitation of the taxa in Louisiana. HISTORICAL ACCOUNT

Palisot de Beauvois (1812) proposed and described the genus

Echinochloa, based on crusgalli L.; numerable changes and revisions have been made since.

Early investigation in the Echinochloa group of grasses, placed it in the genus Panicum. Linnaeus (1753) described Panicum crusgalli and a variety B, his description, "gromen panicum, spica divisu, aristis longis armata, " list of synonymy, and literature of the histori­ cal account leaves little doubt that he was aware of both forms of awned and awnless Panicum crusgalli L.

Andre Michaux (1803) described Panicum muricatum and labelled the type specimen "Lac Champlain. " His description was accurate and fits the accounts of muricata Beauv. (1812) and Echinochloa pungens Poir, (1816). was proposed by M. L.

Fernald (1915) based on Michaux's Panicum muricatum. Femald stated,

"This indigenous plant of eastern America. . . well matches Michaux's description of his Panicum muricatum. . . and is clearly indicated by the present writer's notes, made in 1903, after examination of the

Michaux type."

Rydberg (1931) and Fairbrothers (1952) changed E. muricata

(Michx.) Fern, to E. pungens (Poir.) Rydb. Based on the homonym rule P. muricatum Michx. became invalid because of the earlier £. muricatum Retz. in 1786. Fernald (1935), unaware that his epithet was valid, later accepted E. pungens because of the homonym rule.

Hitchcock (1951) listed Setaria muricata Beauv. as a synonym of E. crusgalli (L.) Beauv. Shinners (1954) reported if the homonym rule is strictly adhered to, the basonym must be designated as

Oplismenus muricatus Kunth. Fairbrothers (1956) reported that

Palisot de Beauvois had transferred Panicum muricatus Michx. to the genus Setaria in 1812 and stated on page 118, "P. muricatum

Michx. equals Setaria. 11

Link (1833) transferred P. colonum L. to the genus Echinochloa, based on the description by Linnaeus in 1759. Wiegand (1921), in reference to the use of colona, pointed out that, "Hitchcock, follow­ ing Greene, has called attention to the fact that the name colonum is not an adjective and hence should not be declined." Ramakreshman

(1960) distinguished two distinct ecotypes of JE. colonum, a tall form and a short form. Yabuno (1962) reported two distinct strains of native E.. colonum and three distinct cultivated varieties.

Steyermark (1963) recognized two forms and one variety in his treatment of E. crusgalli, while Ali (1968) recognized three varieties.

Radford, Ahles and Bell (1968) described E. crusgalli (L.) Beauv. as a polymorphic species in which many characters were used to delimit the taxa overlap. Hitchcock (1951) reported E. crus-pavonis (H.B.K.) Schult. as occurring in Virginia, Alabama, Louisiana and southern Texas. Ali

(1968) reported E. crus-pavonis as indigenous to California.

Hitchcock (1920) (1951) reported E. polvstachva (H.B.K.)

Hitchc. for Louisiana. Wiegand (1921) did not report this species for Louisiana, and it was not confirmed in the present study.

Hitchcock's report of E. polvstachya in Louisiana was of a single collection which was sterile; fide John Lynch, collector.

Wiegand (1921) described E. muricata var. occidentalis. Since then it has been elevated to species rank by Rydberg (1931) and included as a synonym by Hitchcock (1951) and Ali (1968). Rydberg

(1931) stated, "This form was recognized by me forty years ago when

I regarded it as a native species and distinct from E_. crusgalli."

Hitchcock (1950), Farwell (1941) and Ali (1968) treated E. zelavensis (H.B.K.) Schult. as a variety of E. crusgalli, however,

Weigand (1921) and Rydberg (1941) included it as a distinct species.

Wiegand (1921) used the citation E. walteri (Pursh) Nash and in his footnote stated, "Heller in. . . listed E. walteri (Pursh) but with no description or synonymy. Not withstanding that Pursh's name is in parenthesis, the reference is too vague to warrant the acceptance of this as a valid publication of the combination. "

Since 1803 at least twenty-one specific epithets and numerous varieties and forms have been applied to the genus Echinochloa. A complete synonymy is listed for each taxon in the section on taxonomic tre a tm e n t.

Comparisons of selected authors' concepts of the taxa are

shown in Table I. The taxa recognized by the various authors are correlated with Wiegand's (1921) concept which is listed first. The most recent treatment is usually shown when an author has published more than one. TABLE I

Concepts of Echinochloa taxa correlated with Wiegand (1921).

Wiegand (1921) Hitchcock (1935) Small (1933) Rydberg (1941)

(1) colonum colonum colonum colona var. zonalis (2) zelay en sis z elay e n sis var. macera var. subaristata (3) frumentacea crusgalli edulis frumentacea (4) crusgalli crusgalli crusgalli crus-galli (5) m uricata pungens var. ludoviciana var. occidentalis occidentalis var. microstachya microstachya var. multiflora (6) ech inata crus-pavonis var. decipens (7) w alteri w alteri ; w alteri w alteri forma laeviqate (8 ) oplismenoide s oplismenoides (9) holciformis holciformis (10) polystachya polystachya (11) guadenloupensis pyramidalis (12) paludiqena paludiqena paludiqena var. soluta TABLE I (continued)

Fernald (1950) Hitchcock (1951) Gleason (1963) Steyermark (1963)

(1) colonum colonum colonum forma colonum (2) cru sg alli var. zelayensis (3) frum entacea cru sg alli cru sg alli var. frumentacea var. frumentacea (4) cru sg alli cru sg alli cru sg alli cru sg alli var. crusgalli forma longiseta (5) pungens m uricata muricata var. muricata var. ludoviciana var. ludoviciana var. weigandii occidentalis var. occidentalis var. microstachya crusgalli var. mites microstachya var. microstachya var. multiflora var. coarctata (6) crus-pavonis (7) w alteri w alteri w alteri forma laevigata forma laevigata forma breviseta forma breviseta (8 ) (9) (10) polystachya (11) (12) paludigena MATERIALS AND METHODS

Field trips were made through the State of Louisiana to collect plants from natural growing populations of Echinochloa. Data were obtained from examination of herbarium specimens as well as from greenhouse grown plants. These included herbarium specimens from all the hprbaria in the State of Louisiana. Voucher specimens were deposited at the Louisiana State University Herbarium. Duplicate specimens were deposited at the McNeese State College Herbarium and the United States National Herbarium.

Immature inflorescences collected in the field or from the greenhouse grown plants were killed in 3:1 absolute alcohol-glacial acetic acid solution and the anthers were squashed in aceto-carmine

(Sass 1958). The fixed plant material was stored in 70 percent alcohol for periods up to six months. The microsporocytes were examined and chromosome counts were usually made at diakinesis in pollen mother cells. Chromosome counts were documented by photomicrographs of microsporocytes with good chromosome distribution. Abbe camera lucida drawings were also made of many specimens.

The somatic chromosome numbers were determined from root-tip squashes. The roots were pretreated in a 0.002M. solution of 8-hydro- xyquinoline for 2-3 hours, stained and hydrolyzed in a 9:1 solution of two percent aceto-orcein and IN HC1, and squashed in a one percent

aceto-orcein solution.

The endosperm in several collections were studied in an effect to

determine if functional embryo sacs were sexual. Florets were selected

in which the ovary had begun to enlarge but had not yet formed large

quantities of starch for the determination of chromosome numbers in

endosperm cells. Isolated ovules were stained and hydrolyzed in a

warm 9:1 solution of two percent aceto-orcein (Emery 1957).

Seeds collected in the field were germinated by placing them

on moist paper in a petri dish. Seedlings were potted in six inch

clay pots in a mixture of equal parts of sand, clay loam and peat

moss, watered and fertilized to promote growth. Four seedlings from

each sample were grown to maturity. The plants grown from seed were observed at regular intervals to determine whether there were morpho­ logical differences of taxonomic significance among them and whether they differed morphologically from the original collections. Pressed

specimens were made of those plants that showed different morpho­ logical characters when compared with the original field collections

(Appendix A). Primary roots collected from germinated seed and young seedlings were exceedingly small. It was difficult to obtain chromo­

some counts from these. Culms of plants were cut at the base or below a node and placed in water. Adventitious roots larger in diameter than primary roots developed within several days. These 10 were collected according to the previously described method and the plants were pressed for voucher specimens.

Anatomical studies of the leaf epidermis were made from fresh as well as dry material following Stewart's (1965) procedure. Epi­ dermal peels were also made using cellulose acetate; it was spread on the leaf surface and allowed to harden. The strips were removed with forceps and mounted dry on a slide.

Pollen measurements were taken following the procedure by

Gould (1957). Measurements of pollen grains were made using a calibrated ocular micrometer on a compound microscope at 430x.

Spikelets, anthers, caryopsis and embryos were measured with an ocular micrometer in a lOx eyepiece of a stereoscopic microscope with lx and 2x objectives. CYTOLOGY

The importance of chromosome counts has been well established as an integral part of grass . Brown (1950) suggested that the basic chromosome numbdr as x = 9 for the genus Echinochloa. All the species thus far counted have been reported as tetraploid (2n = 36), hexaploid (2n = 54) and octoploid (2n = 72). The earlier chromosome counts recorded on a world wide basis were listed by Darlington and

Wylie (1955) and Carnahan and Hill (1961).

A summary of chromosome counts reported in the literature are as follows:

2n Chromosome Taxon Number Region Recorded by colonum (L.) Link 36 . United States Brown, 1950

54 South Africa de Wet, 1954b

54 United States Gould, 1968

48 India Kirshnaswamy & Raman, 1949

54 T anzania Tateoka, 1965

var. frumentacea (Roxb.)) 54 54 Japan Yabuno, 1962 crusgalli (L.) Beauv. 54 United States Brown, 1948

42 United States Church, 1929b

48 India Rau, 1929

11 12

54 T anzania Tateoka, 1954

42 United States H e is e r & Whitaker, 1948

var. crusgalli 72 United States Gould, 1958

72 United States Gould, 1966

54 United States A li, 1968 m uricata

var. microstachya 36 United States A li, 1968 frumentana (Roxb.) Link. 36 C anada Hunter, 1934

56 T ropicana Church, 1929b

Pungens (Poir.) Rydb. 36 United States Fair brothers, spp. pungens Fairbr. 1952

spp. microstachya Fairbr. 36 United States Fairbrothers, 1952

var. zelayensis (H.B.K.) (H itch. 36 United States Brown, 1948

36 United States A li, 1968 walteri (Pursh) Heller 36 United States Gould, 1968

Aneuploid counts have been reported by Kiirshnaswamy and Raman

(1949), Heiser and Whitaker (1948), Church (1929b), and Mulligan

(1961b). Church also reported a chromosome number of 2n = 56 for

E. frumentacea (L.) Link. Brown (1950) suggested that Church's count of 2n = 28 for E. frumentacea was in error by one chromosome. 13

Only tetraploid and hexoploid counts were made and meiosls appeared normal in all pollen mother cells studied (Fig. 8). The dif­ ferences in chromosome size and morphology within or between taxa were slight. Photomicrographs of chromosomes of each species are shown in Figures 1 to 8.

Chromosome counts have been made for the following collections.

Voucher specimens for both gametic and somatic counts of these col­ lections were deposited in the Louisiana State University Herbarium.

E. colonum. n = 27. (Fig. 1). Bossier: 278; Calcasieu: 261, 302.

Catahoula: 285; East Baton Rouge: 155, 225; Lafourche: 199;

Ouachita: 298; Rapides: 220; St. James: 241; St. Mary: 234;

Terrebonne: 239.

E. occidentalis. n = 27. (Fig. 2). East Baton Rouge: 269;

Evangeline: 266; Jefferson Davis: 328; St. Mary: 231, 249, 250;

Tangipahoa: 207; West Baton Rouge: 246.

E. crusgalli. n = 27. (Fig. 3). Acadia: 135; Allen: 263; Calcasieu:

262; East Baton Rouge: 216, 217, 245; East Feliciana: 308;

. Jefferson Davis: 256, 257, 258; Sabine: 273; St. Landry: 131,

131F1; Tangipahoa: 210, 213; Terrebonne: 238; Vermillion: 254;

W est Baton Rouge: 182.

E. muricata. n = 18. (Fig. 4). Caddo: 276; Lafayette: 344;

Ouachita: 296; Red River: 282; Sabine: 272; Tensas: 309;

West Baton Rouge: 247. muricata var. microstachya. n=18. (Fig. 5). East Baton Rouge

342. zelayensis. n= 18. (Fig. 6). Bossier: 279; Iberville: 187,

187F1; Madison: 314; St. Mary: 236. w a lte ri. n=18. (Fig. 7). Acadia: 138; Calcasieu: 303; East

Baton Rouge: 179, 244; Jefferson Davis: 330, 331; Lafayette:

143; St. James: 242; St. Mary: 248. Fig. 1-7. Chromosomes of Echinochloa species at diakinensis.

Fig. 1. Echinochloa colonum n = 27

Fig. 2. Echinochloa occidentalis n = 2 7

Fig. 3. Echinochloa crusgalli _n = 2 7

Fig. 4. Echinochloa muricata n = 18

Fig. 5. Echinochloa muricata var. microstachya n = 18

Fig. 6. Echinochloa zelayensis n = 18

Fig. 7. Echinochloa walteri n = 18. 15

' ** Fig. 8. Photomicrograph of a meiotic anaphase I of E. colonum (n = 27).

A. Two lagging chromosomes.

B. Normal anaphase I. 16

B 17

Method of Reproduction

Recent investigations in some taxa of Echinochloa have revealed the normal eight-nucleate embryo sac. While reproduction in grasses is usually sexual many generaialso reproduce asexually. Brown and

Emery (1958) reported the frequent occurrence of apomixis in the sub­ family , however, they observed that E. colonum. E_. crusqalli and E. w alterlhave normal eight-nucleate embryo sac develop­ ment. Ali (1968) reported the mature embryo sac in E. muricata as having the normal eight-nucleate development. There have been only a few studies of reproductive behavior of the Echinochloa taxa. Most of the studies of reproduction have involved embryology , megasporo- genesis and embryo sac development. Emery (1957) reported the chromosome counts of endosperm tissue as related to reproduction.

Endosperm in the sexual embryo sacs of most grasses is triploid by the fusion of two reduced polar nuclei with a reduced male gamete

(In + In + In). In most apomicts the endosperm results from the fusion of two unreduced polar nuclei (2n + 2n) or by triple fusion of two unreduced polar nuclei and a reduced male gamete (2n + 2n + In). A

4n or 5n endosperm constitution would indicate a pseudogamous development of an unreduced embryo sac.

These studies show fertilization within the eight-nucleate sexual embryo sac appears to be effected in a normal manner. Examination of 18

endosperm smears from ovules of E.. colonum, E. crusqalli. E.. muricata.

12. walteri. J2. walteri form 2. and zelayensis revealed a 3n chromo­

some constitution (Fig. 9).

Emery (1957) reported the frequent presence of three anthers and

a complete stigma impressed in the surface of the enlarging ovary,

indicating either pseudogamous or cleistogamous development in the

florets. Youngman and Roy (1923) in their work on pollination methods

of some reported E. crusqalli var. frumentaceus as a self­

fertilized taxon. Only one case was found where the three anthers and complete stigma were impressed in the surface of the enlarged ovary of several hundred florets. However, a panicle in an enclosed

sheath from _E. muricata was collected from greenhouse grown plants with a mature caryopsis. This indicated cleistogamous development.

Correlation of Pollen Size and Chromosome Number

Pollen size has been shown to be correlated with chromosome number in several grass genera. Positive correlation between pollen diameter and chromosome number of Poa pratensis was found by Akerbery

(1942). Gould (1957) also reported a correlation, however, the study did show some pollen size differences due both to genetical differences between populations with the same chromosome numbers and to seasonal environmental changes. Bell (1959) found considerable variation in pollen size resulting from different levels of mineral nutrition. Fig. 9. Photomicrograph of an endosperm nucleus showing approximately 81 chromosomes (3n) and 54 chromosomes (3n).

A. E. colonum. n = 27.

B. E. m u ric ata . n = 18. 19

B 20

Schoch-Bodmer (1940) reported a positive correlation between avail­

able habitat moisture and pollen size in Lythrum.

Pollen samples were taken from seven taxa in a total of twenty-

one populations, three of each taxa (Table II). Pollen samples were

taken from fresh material, except for only two herbarium specimens

of E.. muricata var. microstachya. Size measurements were made from

only the populations which chromosome counts had been made pre­

viously. One hundred pollen grains mounted in water were measured

from macerated anthers from several florets on one panicle.

This investigation indicates that a correlation does exist

between the taxa of different ploidy levels. However, it also indi­

cates the variation among the populations within a tax is of a magni­ tude greater than the variation among the ploidy levels. Two

populations of tetraploids, E_. muricata var. microstachya and

E. zelayensis, show a mean diameter of 37.4ju and 35.4^, which

is equal to the hexaploid populations. The coefficient of variation

indicates a wide range at the 95 percent level. This variation may be

correlated to available moisture. Echinochloa is a semi-aauatic / sewer ditch grass that can be found growing in water or in a dryer

h a b ita t. TABLE II

Mean diameter of pollen grains in different taxa of Echinochloa.

Mean diameter of 100 pollen Taxa grains per plant (in microns) Range Coefficient ______n = 18______n = 27 ______of variation

E. colonum 32.9 29.4-36.7 2 .0 E . colonum 37.1 32.0-41.6 5.4 E. colonum 3 2 .2 28.8-38.4 9.1 E. occidentalis 3 2 .6 2 8 .8 -3 8 .4 2 .3 E. occidentalis 3 3 .4 3 0 .4 -4 0 .0 2 .1 E. occidentalis 3 3 :6 3 0 .4 -4 0 .0 2 .0 E. crusqalli 3 2 .5 2 8 .8 -3 6 .8 4 .8 E. crusgalli 3 5 .1 3 2 .0 -3 8 .4 3 .4 E. crusqalli 37.8 3 5 .2 -4 0 .0 3 .0 E. zelayensis 2 9 .4 2 5 .6 -3 6 .8 7 .4 E. zelayensis 3 5 .4 2 8 .8 -4 1 .6 6 .6 E. zelayensis . 2 8 .3 2 5 .6 -3 2 .0 4 .7 E. muricata 3 1 .2 2 8 .8 -3 6 .8 4 .5 E. muricata 32.4 2 7 .2 -3 6 .8 5 .4 E. muricata 3 4 .2 2 7 .2 -4 0 .0 8 .0 E. muricata var. microstachya 2 9 .4 2 7 .2 -3 5 .2 5 .4 microstachya 3 2 .9 30.4-36.8 5.7 microstachya 37.4 3 2 .0 -4 1 .6 5 .2 E. w alteri 2 9 .2 24.5-36.7 7.8 E. w a lte ri 2 9 .4 2 5 .6 -3 5 .2 6 .0 E. w a lte ri 2 9 .7 2 5 .6 -3 3 .6 5 .8 ANATOMY

The importance of anatomical characters in grass systematis has been shown to be useful in differentiating taxa at higher ranks than at the species level. The principal recent work on the anatomy of grass is Brown (1958), Metcalfe (1960) and Stewart (1965). Stewart (1965) used epidermal characters to delimit various grass genera and pointed

out there was a certain amount of intraspecific variation in the charac­ ters cf the epidermis. Metcalfe (1960) gave a detailed account of the

characteristics of the epidermis of grasses. The present investigation utilized epidermal characters based on Metcalfe (1960) and Stewart

(1965) work.

Leaf Epidermis

The abaxial surface of the leaf was used because in most litera­ ture reports, it bears a greater diversity of characters and preparations were easier. The surface is divided longitudinally into alternate zones, the costal zone, that occurs over the vascular bundles, and the inter­

costal zones. The cells of the epidermis are of two main types, long cells and short cells. The long cells vary in depth of sinuous cell walls from slightly sinuous to distinctly sinuous (Fig. 10). The short cells occur mostly over the vascular bundles in rows of three or more

(Fig. 13, 14). 22 23

Silica cells occur over the veins in various shapes, cross, dumb-bell and nodular. There was no significant difference in the taxa, except for some varietion of the ratio of cross-shaped versus nodular.

The cross-shape was more numerous in E. walteri (Fig. 13), E_. muricata and J3. zelayensis. Some of the leaves examined showed the nodular shape silica cells almost exclusively (Fig. 10, 19), while another leaf from the same taxon had many dumb-bell shaped.

Macrohairs were consistently found on a few of the taxa, with several specialized cells at the base as shown in Figure 16 of E_. walteri. E. crusqalli produced several papillose macrohairs along the blade margin near the sheath in some populations. E. occidentalis. typically found in sewer ditches, produced macrohairs scattered over th@ epidermal surface.

Microhairs are usually very abundant, delicate and occur mostly in the intercoastal zone of all the taxa. They are bicellular and the anterior cell usually breaks off (Fig. 12).

Prickle hairs were found on all taxa; they were more numerous in all but one form of E. walteri (Fig. 11), E. muricata and E.. zelayensis.

E. crusqalli had strong prickle hairs only on the major veins. Weak and scattered prickle hairs were present in the intercostal zones of E., zelayensis.

A papilla or swollen point is present on almost all epidermal cells. These vary in the degree of thickness at the ends. They were 24 less distinct in E. walteri and E_. muricata, probably due to the abundant prickle hairs.

The stomatal pattern varies in the number of rows and frequency as shown in Figures 11 and 15. In most of the taxa the stomata are low dome shaped in fresh leaf peels. The subsidiary cells are bordered by one or two long cells. The length of the stomatal opening varies from plant to plant with a rather wide range of 22.4)1 in E. zelaVensis to 54.4u in E. walteri.

S heath

The sheath has been found to be a significant taxonomic character i in delimiting various taxa. E. muricata and.E. muricata Var. micro­ stachya in Louisiana were found to have scabrous sheaths (Fig. 18).

E. walteri (Fig. 15) exhibited a wide range of conditions, but always consistent in all forms except form JL. In form _1 the scabrous condition was sometime indistinct and not rough to the touch. This same condi­ tion of only slightly scabrous was found in one population of E.. zelayensis. while the rest were scabrous. E_. colonum. E. crusqalli.

E. occidentalis and E.. walteri form 3_ had glabrous sheaths.

C ary o p sis

Differences in size and shape of caryopsis and embryo length compared with caryopsis length has been recognized by Martin (1946) and Reeder (1957) as taxonomically significant in grass systematics. The caryopsis was cut longitudinally and stained in the present investigation. Each taxon was examined to determine the difference in length of the embryo and caryopsis (Fig. 20). E. muricata (Fig. 20/

I-J) and E. imuricata var. microstachya (Fig. 20, K-L) were found to have a difference in embryo and caryopsis length that was quite dif­ ferent from the other taxa. E. walteri form 2 (Fig. 21, O-P) produced a larger endosperm than 12. walteri while the embryo, length of both remained about the same. The ranges of the embryo length and caryopsis length were variable but the differences in caryopsis- embryo length were quite stable for all the samples of any given taxon. The differences in caryopsis length and embryo length is given in Table I . A dorsal view of the caryopsis of various taxa is presented in Figure 22. Fig. 10-19. Abaxial epidermis of leaves and sheaths of grasses of various Echinochloa taxa.

Fig. 10. E. crusqalli (lOOx)

Fig. 11, 13. 16. E. walteri (lOOx)

Fig. 12’, 17. E. occidentalis (lOOx)

Fig. 14, 18. E. muricata (lOOx)

Fig. 15. E. walteri form 4 (lOx)

Fig. 19. E. colonum (430x)

Silica bodies: cs, cross-shaped db, dumb-bell shaped n, nodular Macrohairs: pb, papillose based se, specialized epidermal cells Microhairs: b, bicellular Prickle Hairs: u , u n ic e llu la r lm, leaf margin Miscellaneous: p, papillae lc, long cells sc, short cells

Fig. 20. Caryopsis of various taxa of the genus Echinochloa. Showing a longitudinal and cross-sectional view. (15x)

A - B. E_. colonum

C-D. E. occidentalis

E-F. E_. crusgalli

G-H. E_. zelayensis

I - J . E_. m uricata

K-L. E.. muricata var. microstachya

M-N. E_. walteri

O P . E. w a lte ri form 2 27

mm■i-MM

00$ m w f.wftiAvivs-g iw ■r.

m

m & m Fig .21. Caryopsis of various taxa of the genus Echinochloa. Showing the dorsal view. (15x)

A. E.. muricata

B. E_. c ru sg a lli - -•

C. E. muricata var. microstachya

D. E. occidentalis

E. E. zelavensis

F. E. colonum

G. E. walteri

H. E. walteri form 2 28 TABLE III

The difference in caryopsis length and embryo length (mm).

Embryo M ean C aryo psis Mean Caryopsis-Embryo Taxa Range Value Range V alue______D ifference

E. colonum 0 .9 - 1 .2 1.05 1 .3 - 1 .5 1.43 0 .3 8

form _1 1. 0- 1.3 1.18 1 .4 - 1 .7 1.52 0 .3 4

E.. crusgalli 1 .4 - 1 .9 1 .74 1 .4 - 2 .3 2 .0 8 0 .3 4

E.. muricata 1 .7 - 2 .0 1.93 1 .4 - 2 .1 2.00 0 .0 7 E. muricata var. microstachya 1 .3 - 1 .5 1.40 1. 2- 1.8 1.50 0.10

E.. occidentalis 1 .3 - 1 .6 1.45 1 .4 - 2 .0 1.85 0 .4 0

JJ. walteri 1. 0- 1.4 1.21 1 .4 - 1 .9 1 .6 0 0 .3 9

form X 1 .3 - 1 .5 1 .4 0 1 .5 - 1 .8 1.70 0 .3 0

form X 1 .3 - 1 .5 1 .38 1 .9 -2 .1 1 .9 7 0 .5 9

form 3^ 1. 2- 1.4 1 .2 6 1 .5 - 1 .7 1.66 0 .4 0

form 4^ 0. 8- 1.2 1.00 1 .3 - 1 .6 1 .4 4 0 .4 4

E.. zelayensis 1. 1- 1.2 1.15 1 .3 - 1 .5 1.42 0 .2 7 ANALYSIS OF HERBARIUM SPECIMENS

Observations were made of the pressed specimens collected in

1966, 1967 and 1968 and specimens borrowed from the following herbaria: GH, Gray Herbarium; LSU, Louisiana State University; LPI,

Louisiana Polytechnic Institute; NO, Tulane University; NSC,

Northwestern State College; NEC, Northeastern State College; SSC,

Southeastern State College; USL, University of Southwestern Louisiana.

Morphological characters for 323 specimens for this investigation were evaluated by measurements. Characters that appeared to be most significant are spikelet length, seed length and width, anther length, i caryopsis length and width, caryopsis-embryo length ratio, sheath surface, nature of first glume. Spikelet length measurements were made from primary s pikelets from the terminal recemes. Three to five spikelets per plant were used in computing average sizes. Anther lengths were taken from dried anthers. Leaf blade length and width, v. height of plant, racemes were measured to the nearest millimeter.

The data were evaluated by preparing and studying frequency distribution. The polygonal graph method as outlined by Davidson

(1947) was used for correlating six quantitative characters. The mean values for the six characters of each species obtained from previous data were used (Table IV). The characters and scale used for the

30 22. Polygons of Echinochloa taxa.

1. E. colonum. 2. E. colonum form 1

3. E. occidentalis. 4. E. crusgalli.

5. E_. walteri form j4. 6. E.. muricata.

7. E. muricata var. microstachya.

8. E.. zelayensis. 9. E. w a lte ri

10. E. walteri form 1. 11. E. walteri form 3.

12. E. walteri form 2. 31

ipikiut a i t k i r lll|lk(M ) im ik m

Hint liiftk (■■) Hint ilktk

Seal* TABLE IV

Mean values of some quantitative characters of Echinochloa taxa (mm).

F ertile F ertile F loret F loret C ary o p sis C ary o p sis Taxa Snikelet Anther L enath W idth Lencrth W idth

E. colonum 2 .3 4 0 .6 9 2 .0 0 1 .3 4 1.43 1.10 form .1 2 .5 6 0 .7 3 2 .1 4 1.48 1.52 1 .2 8

E. crusgalli 3 .3 0 0 .8 5 2 .8 2 1 .6 4 2 .0 8 1.37

E. muricata 4 .0 0 0 .8 0 3.30 1.80 2.00 1.50

E. muricata var. . microstachya 3 .0 9 0 .7 1 2 .6 5 1 .7 0 1 .5 0 1 .3 7

E. occidentals 2 .8 4 0 .6 0 2.41 1.56 1.85 1.21

E. walteri 3.55 1.03 3.10 1.22 1 .6 0 1 .0 6

form _1 3 .7 3 0 .8 3 3 .3 0 1 .2 4 1 .7 0 1 .12

form 2_ 4.70 0 .9 5 3 .9 6 1 .2 6 1 .9 7 1.1 5

form 3_ 3 .4 4 1 .0 8 3 .0 3 1.20 1 .6 6 1.05

form A 3 .5 5 1 .0 4 3 .0 4 1.22 1 .4 4 0 .9 4

E. zelayensis 2.90 0.90 2.30 1.20 1 .42 1.11 TABLE V

Ranges of some quantitative characters of Echinochloa taxa (mm).

Num ber of F ertile F ertile Specim ens S p ik elet A nther F loret Floret C ary o p sis Caryopsis Taxa Exam ined Lenoth Lenoth Lenoth W idth Lenoth W idth

E . colonum 59 1 .8 - 2 .8 0.5-0.9 1.7-2.3 1.1-1.7 1.3-1.5 0.9-1.2 form _1 15 2.3-3.1 0.5-1.0 2 .0 - 2 .3 1 .2 - 1 .7 1 .4 - 1 .7 1 .2 - 1 .4

E. crusaalli 54 2.8-4.1 0.7-1.1 2.1-3.4 1 .3 - 1 .9 1 .4 - 2 .3 1 .1 - 1 .6

E. muricata 45 . 3 .5 - 4 .5 0.6-1.2 2.8-4.1 1.6-2.2 1.4-2.1 1 .3 - 1 .9

E. muricata var. microstachva 22 2.5-3.6 0.6-0.9 2.4-3.0 1.5-2.0 1.2-1.8 1.1-1.7

E. occidentalis 25 2.5-3.3 0 .4 - 0 .7 ‘2.2-2.7 1.4-1.7 1.4-2.0 1.1-1.4

E. walteri 43 3.0-4.5 0.7-1.4 2.7-3.6 1.0-1.4 1.4-1.9 0 .9 - 1 .2 form JL 14 3 .0 - 4 .5 0 .7 - 1 .1 2 .8 - 3 .5 1.2-1.3 1.5-1.8 1 .0 - 1 .2

form 2_ 3 4 .0 - 5 .2 0 .9 - 1 .0 3 .5 - 4 .5 1.1-1.4 1.9-2.1 1 .0 - 1 .3

form 3_ 12 3 .0 - 4 .0 0.8-1.2 2.8-3.5 1.1-1.3 1.5-1.7 0.9-1.1

form _4 6 3.0-4.2 0.9-1.2 2.7-3.6 1.1-1.3 1 .3 - 1 .6 0 .9 - 1 .0

Ev zelavensis 6 2 .6 - 3 .3 0.8-1.1 2.1-2.7 1.1-1.3 1.3-1.5" l."0 -1 .2 34

polygons are shown in Figure 22. The range of the six characters are

presented in Table V.

Discussion of Taxonomic Characters

Spikelet. (Fig. 23). The nature of spikelet has been an important

character in previous taxonomic treatments of grasses. These studie.s

show spikelet size was not a dependable character to delimit all the taxa due to the variation within each taxon. J E [ . colonum has a uniform

size with 85 percent occurring between 2.1-2. 6 mm. E. muricata and

E. muricata var. microstachya also have a dependable spikelet size.

The range variation in both taxa was 10 mm, with only a slight overlap of spikelet size.

WidgcSftd (1921), Fairbrothers (1952) and others have used the nature of spikelet hairs to separate the Echinochloa taxa. The vari­ ability of the hair length and number was great and found to be only of secondary importance.

The back of the sterile lemma was found to exhibit a coriaceous spot near the middle of spikelet and was consistently recorded for one ta x o n .

Caryopsis length - width. (Fig. 21). The shape of the caryopsis has taxonomic importance in this genus. E. muricata, E. muricata var. microstachya, E.. crusqalli, E. colonum and JE. occidentalis varies from 35 broadly oblong to nearly orbicular, while the other taxa have mostly narrowly oblong caryopsis.

Fertile floret length and width. The fertile floret has been used by some authors as importance in delimiting some taxa. The character is relatively consistent within all taxa. This character was therefore, fairly important in delimiting two major taxa.

Anther. The anther was used by Weigand (1921)- as an important taxonomic character. It was found that color as well as size range was important. However, due to the range of variation, size was important only in delimiting E. occidentalis from the other taxa. E. colonum has purple anthers, this corroborates Yubano's (1962) findings.

Sheath. Hitchcock (1920; 1951) and Wiegand (1921) used the nature of the sheath in the taxonomic treatment of Echinochloa. The sheath has been found to exhibit three distinct conditions in forms examined from Louisiana, glabrous, scabrous and papillose-hispid.

The characters were consistent in each of the seven taxa and five forms that were examined from field and greenhouse specimens.

Awn length. The presence or absence of an awn was variable in most of the taxa. The variation was conspicuous within all taxa except

E. colonum which has not been recorded as having awns. The presence and absence of awns were conspicuously variable even between spikelets

i on the same inflorescences as well as tillering branches in all taxa except E.. colonum and E. occidentalis. 36

First Glume. The first glume has been recognized in some taxo­ nomic treatments as of secondary importance. The first glume in

E. muricata and E. muricata var. microstachya can be consistently delimited from the other taxa based on the presences of strong setae on five nerves. The first glume was always loose and saccate.

Setae on the panicle branches. The pustulate based setae variable in length and frequency on the branches of the panicle occurs in most of the taxa. The absence of pustulate based setae was significant in delimiting E. zelayensis from the other taxa. Fig. 23. Spikelets of various taxa of the genus Echinochloa showing the nature of the spikelet and relative size. Each spikelet is shown in two views, dorsal and ventral. (7X).

A-B. E. muricata

C-D. E. crusgalli

E-F. E. muricata var. microstachya

G-H. E. zelayensis

I - J . JL colonum

K-L . J2. o c c id e n ta lis

M-N. E. walteri

O-P. E. walteri form 2 ISWSi&n&ttli TAXONOMIC TREATMENT

I believe it is best to recognize Echinochloa as a genus with highly variable polymorphic taxa as shown by the evidence in the preceding section. I have chosen to separate subunits of E_. colonum and E. walteri by form number rather than form name. It is my opinion that additional names in this group at this time would only add to the confusion. E_. muricata var. microstachya was treated as a variety, however, in the preliminary investigation it was considered as only a form of E. mucricata.

Echinochloa Beauv.

Yellowish green to green purplish annuals with tillering culms,

prostrate, spreading ascending, or erect, up to 2.5 meters tall;

panicle terminal and axillary; racemes 1.5-21 cm long, simple or com­

pound; spikelets paired or crowded along the raceme, mostly plano­

convex, strongly mucronate to coarsely pubescent, sometimes purple

spotted green to brownish, elliptic to orbicular; first and second glume

distinct; fertile floret variable with spikelet length, smooth, shining

and coriaceous; leaf blades ascending to spreading, glabrous to

puberulent to papillose-hirsute; ligule none to minute white hairs; leaf

sheaths glabrous, scabrous to strongly papillose-hispid, the lower ones

sometimes purplish.

38 39

Its greatest diversity of form in Louisiana occurs in the southern

part, mostly along the coastal marsh region.

Key to Species, Varieties and Forms of Echinochloa in .Louisiana

Sheath glabrous or papillose-hispid; palea of lower floret well developed;

first glume rarely 5-nerved, appressed to sterile lemma.

Fertile floret length less than twice the width; caryopsis ovate or

oblong, not ellipsoid; sheath glabrous.

Sterile lemma of lower floret with smooth coriaceous spot on most

spikelets; anther yellow or purple., mostly 0.6 mm long(0.4-.7 mm);

lower floret often staminate.

6. E.. occidentalis

Sterile lemma of flower floret with microhairs and spinules on mid­

rib; anther yellow or purple, mostly 0.7-.9 mm long, lower

floret rarely, if every staminate.

Spikelets 2.8-4.1 mm long, awn length variable; anther 0.7-

1 .1 mm lo n g , y e llo w . 3. E. crusgalli

Spikelets rarely more than 2.8 mm long, awnless, spinules not

distinct; anther purple, ca. 0\7-0.8 mm long.

Panicle loose mostly spreading, branches simple, often on one

side; spikelets mostly 2.0-2.5 mm long.

I.E. colonum 40

Panicle tight, erect, green, usually substended by the two

upper blades; spikelets mostly 2.4-2.9 mm long.

2. E. colonum form JL

Fertile floret length greater than twice the width; caryopsis ellipsoid;

sheath glabrous, or papillose-hispid.

Sheath papillose-hispid, sometimes only marginal.

Caryopsis more than 1.8 mm long; spikelet 4.0-5.2 mm long,

long awned, awn 9-13 cm long; embryo ca. 0.5 mm less than

caropsis length.

9. E. walteri form 2_

Caropsis less than 1.8 mm long; spikelet 3.0-4.5 mm long,

usually awned; embryo ca. 0.4 mm less than caryopsis

le n g th .

Panicle loose nodding, branches usually descending, green or

purple-ginged; lower sheaths papillose-hispid.

7. E_. walteri

Panicle tight, branches ascending, only slightly nodding, green

with purple awn; sheaths papillose-hispid up to panicle;

ligule a row of white hairs.

11. E. walteri form 4

Sheath not papillose-hispid.

Sheath glabrous. 41

Panicle tight, mostly erect; branches ascending; spikelet

green with purple awns or awn points; culms slender;

ligule a row of white hairs. 8. E. walteri form 1

Panicle loose, branches mostly descending, nodding;

spikelets green to purplish-tinged; culms usually strong

at the base; no ligule. 10. E. walteri form 3

Sheath scabrous, pubescent or sometimes only slightly scabrous at the

top; first glume 3 or 5-nerved; palea of lower floret present or

a b s e n t.

Palea of lower floret well developed; pustulate based setae present

on branches, sometime few.

Fertile floret length less than twice the width; first glume 5-nerved

with strong spinules; spikelets with weak to distinct pustulate

based spinules.

Spikelet 3.5 mm long or more to base of awn; anther ca. 0.8 mm

long; pustulate based spinules distinct.

4. E_. muricata

Spikelet 3.5 mm long or less to base of awn; anther ca. 0.7 mm

long; pustulate based spinules few and weak.

5. E_. muricata var.

microstachya Fertile floret length greater than twice the width; first glume

mostly 3-nerved.

8. E. walteri form 1 42

Palea of lower floret absence or rudimentary; pustulate based setae

absent on branches.

12. E_. zelayensis

1(. Echinochloa colonum (L.) Link, Hort. Berol. 2: 209. 1833.

Panicum colonum L ., Syst. Nat. ed. 10. 2: 870. 1759.

Milium colonum Moench, Meth. PI. 202. 1794.

Qplismenus colonum H.B.K., Nov. Gen. etSp. 1; 108. 1815.

Panicum sonale G uss., FI. Sic. Prodr. 1: 62. 1827.

Qplismenus repens Presl., Rel. Haenk, 1: 321. 1830.

Qplismenus colonum var. zonalis Schrad., Linnaea 12: 429. 1838.

Panicum incertum Bose. exSteud., Nom. Bot. ed. 2. 2: 258. 1841.

Echinochloa zonalis Pari.. FI. Panorm. 1: 119. 1845.

Panicum prorepens Steud.. Syn PI. Glum. 1:46. 1854.

Qplismenus crusgalli var. colonum Coss. and Dur., Expl. Sic.

Alger. 2:28. 1854.

Panicum colonum var. sonale L. H. Dewey, U. S. Nat. Herb. cont.

2: 502. 1894.

Echinochloa colonum var. zonalis Woot. and Standi., N. Mex. Cd.

Agr. Bui. 81: 45. 1915.

Echinochloa crusgalli subsp. colonum Honda. Bot. Mag. (Tokyo)

37: 122. 1923.

Panicum crusgalli subsp. colonum Makino and Nemoto. FI. Jap.

1470. 1925. Plants annual. Culms decumbent to suberect, weak, sometimes rooting on lower nodes. Panicle loose, erect, branches simplq, from

1.2-3.5 mm long, mostly on one side of panicle axis. Nodes glabrous t and usually swollen toward~the base. Sheath glabrous. Ligule absent.

Blade 0.5-1.5 cm broad, 7-38 cm long. Spikelets planoconvex, 1.8-

2 .8 mm long, awnless or with awn point, green to purple colored, usually in four rows along each branch of the panicle, spinules of spikelet minute. Palea of lower floret well developed and subequal to palea of fertile floret. Sterile lemma and first glume sqbequal. Anther

0.5-0.9 mm long, purple, rarely yellow. Fertile floret 1.7-2.3 mm long, 1.1-1.7 mm broad. Caryopsis 1.3-1.5 long, 0.9-1.2 mm broad, brownish colored. Embryo 0.9-1.2 mm long, 0.38 mm less than caryopsis length. Flowering from June to November.

Type: Tamacia, Browne, location of type specimen unknown.

Distribution: Found abundantly throughout Louisiana, as a weedy grass

in usually damp cultivated fields and waste places.

Discussion: This taxon is native and cosmopolitan in Louisiana. It is

recognized by the thin slender culms, simple branches on a short

green or purple panicle. One specimen, B-290. was collected with

purple bands, a form that was recognized by Wiegand (1921). The

variability of purple pigment in leaves of this taxon was considered

i to be of no taxonomic value. 44

Representative specimens:

LOUISIANA: Parish. Acadia, Gooch 14 (USL). Assumption,

B-229. Avoyelles, B-289. B-290. Beauregard, Thieret 29438

(USL). Bossier. Robinette 214 (LPI), B-280, Shinners 21192

(GH). Calcasieu, Thieret 28001 (USL), B-261, B-301. Caldwell,

B-292. Cameron, Thieret 9613 (USL), Valentine 6084 (USL).

Catahoula, Moore 6471 (LPI. LSU), B-285, B-2870 East Baton

Rouge, Cain and Heath, December 12, 1925 (LSU), B-153, B-155,

B-158, B-159, B-167, B-176, B-201. B-204, Thieret 27777 (USL).

East Carroll, B-319, Thieret 29997 (USL). East Feliciana,

Thieret 21290 (USL). Evangeline, B-268. Franklin, Thieret

30060. 30070 (USL), Iberia, B-304, Thieret 15924 (USL)..

Iberville, B-184. B-186. Jefferson, B-329, Guidry 14 (LSU).

Jefferson Davis, Dickinson, Sept. 15, 1918. Lafayette, B-142,

Claycomb (?) (LSU, USL), Correll 9365 (LPI), Haynes and Hinton

2526 (GH, USL), Lasseigne 195 (USL), Thieret 8649 (USL),

Reynolds 61 (NSC). Lafourche, Lasseigne 656 (USL), B-198.

Lincoln, B-326. Madison, B-316, Thieret 28231. Natchitoches,

Reynolds 71 (NSC), Montgomery 68 (NSC), Barrett 56 (NSC), Rabb

63, (NSC). Orleans, Bank, Aug. 2L, 1940 (NO), Ewan 18800 (NO,

GH), Morrison, Tune 30, 1929 (NO). Ouachita, B-297, Thieret

20713 (USL), Thomas 722, 3781, 10980 (NEC). Plaquemines,

• Ewan 17986 (NO, GH), Red River, Thieret 20652 (USL). Sabine, '45

Shinners 26544 (GH). St. Bernard, Lemaire 847, 923 (LSU,

USL). St. Charles, Griffinq, Oct. 1, 1913 (LSU), Montz 153

(LSU). St. Tames. B-196, B-197, B-241. St. Landry, B-129.

B-130. St. Martin, Russ 358 (USI), Kidder 178 (USL). St..

Mary, B-191, B-230, B-231. B-233, Correll 9343 (LPI, GH),

Ewan 18953 (NO). St. Tammany, Rockett 6. Tangipahoa. B-208, t B-209, B-211, Brown 19896 (LSU), Smith 51 (LSU). Tensas,

B-310, B-311. Terrebonne. B-239, B-240, Arceneaux 212 (LSU),

Ingram and Haynes 172 (LSU). W est Carroll. Thieret 28219 (USL).

Winn, Brown 9586B (LSU). Parish unknown, Hale 365 (GH).

MEXICO: Llano Caguihui, Wiggins 15526A (GH).

2. Echinochloa colonum form 1_.

Plants annual. Culms erect, not rooting at the nodes. Panicle tight, branches, ascending. Upper blades mostly erect projecting.above the end of panicle. Sheath glabrous. Nodes usually pubescent, short appressed hairs, some glabrous. Spikelet 2.3-3.1 mm long, green, awnless, scarcely echinate, sterile lemma and first glume subequal, sterile palea white, well developed, subequal to palea of fertile floret.

Anther 0.5-1.0 mm long, purple to pink colored. Fertile floret 2.0-2.3 mm long, 1.2-1.7 mm broad, ovoid. Caryopsis 1.4-1=. 7 mm long, 1.2-

1.4 mm broad, brownish colored. Embryo 1.0-1.3 mm long, 0.34 mm

. » less than caryopsis length. 46

Distribution: Scattered throughout Louisiana, occurring mostly in

the rice fields or associated with cultivated crops.

Discussion: This form is recognized or distinct from the typical

E. colonum by the somewhat larger seed, erect culms and the

tendency of the upper two blades to somewhat enclose the green

compact panicle.

Representative specimens:

LOUISIANA: Parish. Calcasieu, B-302. Catahoula. Moore

6471 (GH). Concordia. B-288. East Baton Rouge. B-223. East

Carroll, B-320, B-321. Iberia, B-305. Lafayette, B-337.

Lincoln, B-325. Madison, B-317. St. Mary, B-234. Vermilion,

Lass eigne 436 (USL). W est Carroll, B-323. West Feliciana,

Thieret 21894 (USL).

3. Echinochloa crusgalli (L.) Beauv., Ess. Agrost. 53, 161. 1812.

Panicum crusgalli L., Sp. P. 56. 1756.

Panicum crusgalli var. A L ., Sp. PI. 56. 1753.

Panicum crusgalli var. B L ., Sp. PI. 56. 1753.

Milium crusgalli Moench, Meth. PI. 202. 1794.

Panicum grossum Salish., Prod. Stirp. 18. 1796.

Panicum crusgalli (L.) aristatum Pursh FI. Amer. Sept. 66. 1814.

Pennisetum crusgalli Baumg., Enum. Stirp. Transsilv. 3: 277. 1816.

Echinochloa crusgalli var. aristata S. F. Gray, Nat. Arr. Brit. PI.

2: 158. 1821. 47

Qplismenus crusgalli Dum., Obs. Gram. Belg. 138. 1823. i Orthopoqon crusgalli Spreng., Syst. Veg. 1: 307. 1825.

Panicum crusgalli (L.) var. lonqisetum Trin. Sp. Gramin. ii. t.

162. 1829.

Echinochloa crusgalli (L.) Veauv. forma vittate Hubb., Rhodora

2 3 2 . 1916.

Echinochloa crusgalli (L.) Beauv. forma longiseta (Trin.) Farwell,

Rep. Mich. Acad. Sci. ssi. 349. 1919.

Plants annual, to 1.5 meters .tall, erect to decumbent and rooting

on the lower nodes. Culms slender, nodes glabrous, slightly, swollen.

Sheath glabrous, often purple at the g$se. Ligule absent. Blade 0.5-

2.1 cm broad, 9-48 cm long, often pubescent with a few pustulate based

hairs along the margin at the base of the blade. Panicle 5.5-24 cm long, with mostly spreading or sometime appressed branches. Spikelets

panoconvex, 2.8-4.1 mm long to the base of the awn or awn point.

Glumes mucronate and hispid, the first glume only minutely mucronate

o$ the nerves, s'pinules of second glume and sterile lemma of spikelet without swollen base, sometime only slightly. Sterile palea well

developed, whitish or purplish. Anther 0.7-1.1 mm long, yellow at anthesis. Fertile floret 2.1-3.4 mm long, 1.3-1.9 mm broad, smooth

coriaceous, white to dark gray. Caryopsis 1.4-2.3 mm long, 1.1-1.6

mm broad ovate brownish colored, embryo 1.4-1.9 mm. Flowering from June to November. 48

Type: Canada, Kalm (?), in the Linnean Herbarium. The type has not

been examined.

Distribution: E. crusgalli (L.) Beauv. is distributed throughout i Louisiana. It is more common in cultivated fields and sewer

d itc h e s .

Discussion: The size of the spikelet varies from 2.8-4.1 mm long.

The spinules are shorter in general than E_. muricata or E. walteri.

The presence or absence of an awn was quite distinct and variable,

and unimportant as a taxonomic character. Field plot specimens

B-203; and B-217 were examples of this variability. The older

panicles bear spikelet that were distinctly awnless, while the

panicle produced later in the growing season develops spikelets

with awns. Hitchcock (1920) recognized only the one taxon, JB.

crusgalli (L.) Beauv. and several varieties in the United States

as being difficult to separate from the European specimens.

Weigand (1921) recognized two taxa as distinct, one indigenous,

E. muricata (Beauv.) Fernald and E. crusgalli (L.) Beauv. as

introduced.

Representative specimens:

LOUISIANA. Parish. Acadia. B-133, B-134, B-135, B-136.

B-137. Allen. B-263. B-264. Bienville, Ware 66 (NSC).

Calcasieu, B-262, B-300, Thieret 20634, 23507, 28000 (USL). i Caldwell, B-294. Cameron, Haynes 85 (U&L)/ Thieret 8781 (USL). Claiborne, Teansonne. Tune 24. 1938 (LSU). East Baton Rouge,

B-203, B-217, B-224. B-245, B-270, Brown 5086 (LSU).

Burnett 8. (LSU). East Feliciana, B-308, Thomas 11530 (NEC)»•

Evangeline, B-267. Iberville, B-183. Jackson, Hammons 16

(NO). Jefferson Davis, B-255, B-256, B-257, B-258, Thieret

26356 (USL). Lafayette. Clavcomb 361, 364 (USL), B-141.

Reese 1456 (USL, GH), 3855 (USL). Lafourche, B-199. LaSalle,

B-284. Lincoln, Woodword 638. 729 (LPI). Livingston, Rockett

35. (LSU). Natchitoches. Ware 72 (NSC)i; Ouachita. B-299.

Orleans, McArthur. Tuly 2. 1928 (NO). Sabine, B-273. St.

Bernard, Lemaire 281 (USL). St. Landry, B-131. B-132. St.

Mary, B-235. Lemmon 1219 (LSU). Tangipahoa, B-215. B-226 .

Thomas 11654 (NEC).. . Wascorn 182 (LSU). Terrebonne, B-238.

Lasseigne 871 (USL). Vermilion, B-251, B-253. B-254. Hansen

20885 (USL), Lasseigne 430 (USL). West Baton Rouge, B-182.

Winn, Brown 9586A (LSU).

Echinochloa muricata (Beauv.) Fern. Rhodora 17: 106. 1915

Panicum muricatum M ichx., FI. Bor. Amer. 1: 47. 1803. Not

P. muricatum Retz. 1786.*

Setaria muricata Beauv., Ess. Agrost. 51, 170, 178. 1812.

Panicum .pungens Poir. in Lemark'd Encycl. Meth. Bot. Sup. 4:

273. 1816.

Qplismenus muricatus (Beauv.) Kunth, Rev. Gram. 1: 45. 1829. 50

Echinochloa crusgalli (L.) Beauv. var. muricata Farwell, Mich.

Acad. Sci. Rept. 21: 350. 1920. I Echinochloa crusgalli var. michauxii House, N. Y. State Mus.

i Bull. 243-244: 42. 1923.

Echinochloa pungens (Poir.) Rydb., Brittonia 1: 81. 1931.

Echinochloa pungens var. coarctata Fern, and G risc., Rhodora 37:

136, pi. 336, fig. 1, 2. 1935.

Plants annual. Culms suberect to erect, strong. Panicle 10-33 cm long, branches 3.0-11 cm, mostly widespread along panicle axis, with few to no pustulate based setae along the bfanch. Sheath often purplish, scabrous. Nodes glabrous. Spikelets 3.5 to 4.5 mm long, usually awned, rarely awn pointed, pustulate based setae often strong, thick at base, usually purplisri. green, crowded along panicle branches, palea well developed, usually purplish. First glume 5-nerved, with distinct setae on nerves, saccate or loose around spikelet. Fertile floret 2.8-4.1 mm long, 1,6-2.2 mm broad. Caryopsis 1.4-2.1 mm long, 1.3-1.9 mm broad. Anther 0.6-1.2 mm long, usually yellowish to tan. Caryopsis 0.07 mm longer than embryo. Flowering from July to November.

Type: CANADA. Lake Champlain Michaux. The type has not been

exam in ed.

Distribution: E. muricata is widespread in the northern part of Louisiana, ■ i South to the Coastal Plain marsh. 51

Discussion: Fernald (1915), Wiegand (1921)/ Fassett (1949), arid

others treated this taxon as native to North America. Examina­

tion of field collections and herbarium specimens show only

slight variations. Some specimens were found to have a green

inflorescence while others were purplish. The strong pustulate

based setae was prominent on all spikelets examined.

Representative specimens:

LOUISIANA: Parish. Bossier. Robinette 171 (LSU), Thieret

26085 (USL), Penn, Tulv 24, 1941 (USL), B-276. Caldwell,

B-291. Catahoula, B-286. DeSoto, Thieret 2Q742 (USL). East ..

Baton Rouge, Lasseigne 1149 (USL). East Carroll, B-318,

Thomas 3316, 3336 (NEC). East Feliciana, B-309. Franklin,

Thieret 30058 (USL). Lafayette, Lasseigne 222 (USL). LaSalle,

Thieret 25043 (USL). Lincoln, Dike 14 (LPI), Moore 5458 (LPI), '

Moore 5400, 5458. Poland 278. Madison, B-312, Lasseigne

840 (USL). Morehouse, Thieret 28211, 29920, 20433 (USL)'.

Natchitoches, Barrett 53 (NSC), Harris 67 (NSC), Montgomery

59. (NSC), Rabb 75 (NSC), Ware 69, 73 (NSC). Orleans, Cocks

Tune 1900. Ouachita, Thieret 20828, 28164 (USL), Thomas 20,

3565, 11404 (LPI). Red River, Thieret 20307 (USL). Richland,

Thieret 20928 (USL). Sabine, B-272. Tangipahoa, E. A. Bessey

t May 12, 1909 (LSU). Union, Moore 6345 (IPl). West Baton

Rouge, B-247. West Carroll, B-322, Ricks JL3 (LSU). TEXAS: Shelby Co. B-274.

Echinochloa muricata var. microstachya Weiqand Rhodora 23: 58. 1921

Panicum crusgalli var. mite Prush, FI. Amer. 1814.

Panicurr crusgalli var. purpureum Pursh. FI. Amer. 1814. i Panicum crusgalli var. mutlcum E ll.. Bot. S. C. and Ga. 1: 114.

1816.

Echinochloa crusgalli var. mitis (Pursh) Peterm., FI. Lips. 82.

1838.

Panicum sciendens Ness ex steud. . Svn. PI. Glum. 1: 47. 1854.

Qplismenus crusgalli var. muticus Wood. Amer. Bot. and Flor.

pt. 2: 393. 1871.

Panicum crusgalli normale var. mite forma hispidum Kuntze. Rev.

Gen. PI. 2: 783. 1891.

Echinochloa crusgalli var. mutica Rvdb. . Colo. Agr. Col. Bui.

100: 21. 1906.

Echinochloa crusgalli forma purpurpea Farwell. Mich. Acad. Sci.

Rpt. 21: 349. 1920.

Echinochloa muricata var. multiflora W eigand. Rhodora 23: 59.

1921.

Echinochloa muricata var. ludoviciana Wiegand, Rhodora 23: 58.

1921. \ Echinochloa microstachya Rvdb.. Brittonia 1: 82. 1931. j. 53

Echinochloa pungens (Poir'.) Rydb.# var. microstachya

(Wiegand) Fern, and G risc.# Rhodora 37: 137. 1935.

Echinochloa pungens var. multiflora Fern, and G risc., Rhodora i 37: 137. 1935.

Echinochloa crusgalli var. microstachya (Wiegand) Shinners,

Rhodora 56: 33. 1954.

Plants annual. Culms erect, usually strong. Sheath often

purplish, scabrous. Blade 0.9-2.0 cm broad by 20-62 cm long.

Panicle to 25 cm long, branches 3.0-9.0 cm long often crowned along

panicle axis. Spikelets 2.5-3.6 mm long, purplish or purple tinged.

The awns absent in most forms. Spikelets with or without pustulate

based setae. First glume 5-nerved, with setae on nerves, saccate or

loose around spikelet. Fertile floret 2.4-3.0 mm long, 1.5-2.0 mm

broad. Caryopsis 1.2-1.8 mm long, 1.1-1.7 mm broad. Anther

0.6-0.9 mm long. Caryopsis 0.10 longer than embryo. Flowering from July to November.

Type: NEW YORK. Ithaca jE. L. Palmer 97 (GH). The type has not been

e x a m in e d .

Distribution: Scattered throughout the northern central and southeastern

part of Louisiana.

Discussion: E. muricata var. microstachya is a variable taxon as

Wiegand (1921) recognized. His treatment of E. muricata included

four varieties, ludoviciana, occidentalis, microstachya and 54

multiflora. E. muricata var. ludoviciana, Billing 14 (GH),

Baton Rouge, Louisiana, has been examined and is equal to

E. muricata var. microstachya specimens collected from the

same area help to confirm it is not synonompus with

E. muricata. The spikelets are smaller, anther smaller,

panicle shorter, tight and more erect.

Representative specimens:

LOUISIANA: Parish. Bienville. Gouedy 724 (LPI). Caddo,

Thieret 20980 (USL). East Baton Rouge, B-154, B-342 (LSU),

Billings 14 (GH). Jackson, Posher 638 (LPI), Freeland 693. »*» Jefferson, Ewan 17430 (NO). Lafayette, Delahoussage 195

(USL). LaSalle, Hollis 226 (LPI). Lincoln, Hansford 84, 115

(LPI). Livingston. Thieret 24771 (USL). Natchitoches,

Robbertson 62. 74, 70 (NSC). Ouachita, Krai 8054 (NEC)..

St. Mary, Delahoussage, 208 (USL). Tensas, Thieret 21560.

28250 (USL). .

ARKANSAS: Scott Co., Moore 520859 (LPI).

NEBRASKA: Lancaster C o., Maples 286 (LPI).

6. Echinochloa occidentalis (Wleg.) Rydb. Brittonia 1: 82. 1931.

Echinochloa muricata var. occidentalis Wieg. Rhodora 23: 58.

1921.

Echinochloa pungens var. occidentalis Fern, and G risc.,

Rhodora 37: 137. 1935.

Echinochloa pungens var. wlegandii, Rhodora 51: 2. 1949. 55

Plants annual# to 1 meter tall. Culms suberect# slender. Nodes glabrous, slightly swollen. Sheath glabrous to slightly scabrous rarely purple at the base. Ligule absent. Blade 6-11 mm broad#

10-30 cm long# usually smooth. Panicle 8-23 cm, green# erect# slightly spreading but ascending# branches with many pustulate setae# usually extending beyond spikelets. Spikelets planoconvex, 2.5-3.3 mm long# spinules short# only slightly swollen at base# lower floret often staminate# first glume ca. one-half length of second glume# sterile lemma with a smooth slightly coriaceous spot along the mid- nerve# spikelets green to only slightly purple tinged# awnless to short awn points. Anther 0.4-0.7 mm long# average 6.5 mm# yellowish to purple-tinged. Sterile palea well developed. Fertile floret ovate#

2.2-2.7 mm long# 1.4-1.7 mm broad# smooth coriaceous, brownish gray. Caryopsis 1.4-1. 7 mm long# 1.1-1.4 mm broad# brownish colored. Embryo 1.3-1.6 mm long# 0.4 mm less than caryopsis length. Flowering from June to November.

Type: Grand Tower. 111.. Gleason 1720 (GH).

Distribution: Scattered throughout most of Louisiana# on rich soil#

sewer ditches and damp waste places.

Discussion: E. occidentalis Rydb. has been recognized at the species

level by Rydberg (1931) and Gleason (1963). This investigation

has found ■ that morphological differences do exist between all

other tasia and E. occidentalis. Wiegand (1921) included it as 56

a variety of E. muricata primarily on the slightly swollen spinule

base, anthers 0.4-0.6 mm long. The smooth coriaceous spot on

the sterile lemma, lower floret often staminate, and the n = 27

chromosome constitution has been the basis for using Rydberg

taxonomic treatment of this taxon.

Representative specimens:

LOUISIANA: Parish. Acadia, B-259. Allen, B-265. Caddo

B-277. Calcasieu, B-260. Caldwell, B-293. East Baton Rouge,

B-224, B-269. Evangeline, B-266. Grant, Thieret 23212 (USL).

Tefferson, B-328. LaSalle, B-283. Livingston, Brown 19595

(LSU). Madison. B-313, B-315. Morehouse, Thieret 29921

(USL). Natchitoches, B-271. Ouachita, B-295, B-296, B-298.

Rapides. B-221. St. Charles, Montz 205 (LSU). St. Mary,

B-249, B-250. Tahgipahoa, B-207. West Baton Rouge, B-246.

TEXAS: Shelby Co. . B-275.

7. Echinochloa walteri (Pursh) Nash, Britton's Manual 78. 1901.

Panicum hirtellum W alt., FI. Carol. 72. 1788.

Panicum walteri Pursh, FI. Amer. Sept. 66. 1814.

Echinochloa longearistata Nash in Small, FI. Southeast U. S. 84.

1903.

Panicum crusgalli var. walteri Farwell, Mich. Acad. Sci. Rept.

6; 202. 1904. j

Echinochloa walteri forma breviseta Fern, and G risc., Rhodora

37: 137. 1935. Plants annual. Culms strong, up to 2.5 meters tall. Panicle

15-61 cm long, branches spreading, loose, usually compound with many to no pustulate based setae. Nodes glabrous to slightly pre- i bescent. Sheath papillose-hispid, usually only on the lower sheaths, if on upper, only marginal, no ligule. Spikelets 3.0-4.5 mm long, variable in color, green to purple, setae strong, pustulate base not always distinct, awns variable up to about 3.5 cm long. Sterile palea usually white, from well developed to only one-half length of fertile floret, often bifid at apex. Anther 0.7-1.4 mm long, usually brownish yellow. Fertile floret 2 .7-3.6 mm long, 1.0 -1.4 mm broad.

Caryopsis 1.4-1.9 mm long, 0.9-1.2 mm broad, brownish. Embryo

0.39 mm less than caryopsis. Flowering from June to November.

Type: SOUTH CAROLINA. W alter. Type has not been examined.

Distribution: Abundant from Cameron Parish, South along the Coastal

Plain to St. Bernard Parish and throughout the Florida parishes.

Scattered throughout the northern part of Louisiana mostly al ong

the rivers and bayous.

Discussion: E. walteri (Pursh) Nash, was recognized by Wiegand (1921)

on.the basis of a short-awned glume. Hitchcock (1951) separated

E. walteri on the nature of the sheath. The spikelet size varies

from 3.0 to 4.5 mm and the awns are almost as variable. The . > first glume may be awned or awnless, while the second glume is

more consistently awn pointed or awned. The sheath is 58

consistently papillose-hispid, if only on the margin, generally

on the lower sheaths.

Representative specimens:

LOUISIANA: Parish. Caddo. Thieret 20978 (USL). Cameron,

B-334. B-340, Brown 5901 (LSU), Chamberlain. Aug. 10. 1955

(LSU), Chamberlain. Aug. 10, 1955 (LSU). DeSoto, Thieret

20681 (USL). East Baton Rouge, B-222, B-225. B-243, Cain,

Oct. 12. 1925 (LSU), Thieret 27785. Iberia, Correll 9555 (LPI).

Iberville, B-185. Jefferson, 6^227, B-228, B-331, Lambremont

7. (NO). Lafayette, Claycomb 367 (USL). Lafourche, B-200.

Livingston, B-206. Morehouse, Thieret 20312 (USL).

Natchitoches. Ware 64, 65 (NSC), Barrett 55 (NSC), Robinson

76. (NSC). Orleans, Lake Pontchartrain (LSU), Cocks 1928 (NO).

Ouachita, Thieret 28188, 20832 (USL). Plaquemines, Keller

Tuly 12, 1917 (NO). St. Charles, Lambremont 10 (NO). St.

Tammany, Thieret 23407 (USL). Tangipahoa, Wascom 270,

246 (LSU), B-214. Brown 19897, 19902 (LSU). Tensas, Thieret

21524 (USL). Terrebonne, Lasseigne 869, 582 (USL), Wurzlow.

Tulv 27. 1912 (NO).

8. Echinochloa walteri form 1.

Plant annual. Culms slender, tillering from central base. Sheath scabrous, ligule present as a row of short hairs up to ca. 0.8 mm long.

Blades 0.6-1.2 cm broad, 18-65 cm long. Nodes glabrous to only 39

slightly pubescent, panicle tight nodding only slightly. Spikelet

3.0-4.5 mm long, strong setae on margins of second glume and sterile lemma, pustulate base not distinct, green with purple awn, awn variable in length. Anther 0.7-1.1 mm long yellowish color. Fertile floret 2.8-3.5 mm long, 1.2-1.3 mm broad. Caryopsis 1.5-1.8 mm long, 1.0=1.2 mm broad, brownish colored. Embryo ca. 0.3 mm less than caryopsis length. Flowering from July to November.

Distribution: Scattered along the southern part of Louisiana, from

Cameron to St. Bernard Parish.

Discussion: This form is recognized by the slender culms, purple

coloration, mostly on the lower sheaths, tight panicle and

ligule a row of short white hairs. Sheath is consistently

Scabrous. A specimen examined from University of Southwestern

Louisiana, Reese 1452. reported the roots appeared to be perennial,

this could not be determined from the specimen at USL, however,

the specimen from Gray Herbarium did have the distinct appear­

ance of having rhizomes.

Representative specimens:

LOUISIANA; Parish. Acadia, B-138. Cameron, B-327,

Thieret 8678 (USL). Lafayette, B-143. Haynes and Hinton 2539

(GH), Cemaiie 1718. =947 (NO). Orleans, Hall. Tulv 1935 (NO).

St. Martin, Reese 3869, 5903 (USL) (GH). St. Mary, B-248.

Vermilion. B-139, Reese 1452 (USL). 60

9. Echinochloa walteri form .2

Plants annual. Culms strong and thick at the base. Panicle i

spreading and nodding, green to purple tinged, turning brown with i age. Sheath pubescent with weak and indistinct papillose based hairs.

Ligule with a few short hairs, some indistinct. Blades ca. 2 cm broad. » Spikelet 4.0-5.2 mm long, with second glume and sterile lemma long awned up to 13.5 cm long. Anther 0.9-1.0 mm long, brownish yellow.

Fertile floret 3.5-4.5 mm long, 1.1-1.4 mm broad. Caryopsis 1.9-2.1

mm long, 1.0-1.3 mm broad. Embryo 1.3-1.5 mm long, ca. 0.59 mm

less than caryopsis, usually brownish colored.

Type: The type specimen Panicum longisetum Torr. and E.. longearistata

Nash are both in the herbarium at Columbia University. Wiegand

(1921) reported these two types as having been inappropriately

named and gave them the epithet E . walteri forma laevigata.

Distribution: The only collections were reported from St. Mary Parish.

The third specimen was from Louisiana but no location was given.

Discussion: This form was recognized by the long awns and spikelets

that measure 4.0-5.2 mm long. One other distinguishing character

was the large caryopsis, 1.9-2.1 mm long.

Wiegand (1921) placed all the E. walteri that had glabrous

sheaths into 12. walteri forma laevigata, regardless of awn,

spikelet or caryopsis length. This present investigation recog­

nizes a difference in that taxon with glabrous sheaths and the taxon with pubescent, scabrous or papillose-hispid sheaths.

The specimen from Gray Herbarium was only the upper one-half i of the plant, it was glabrous while the two specimens from St. t Mary Parish are distinctly pubescent and weakly papillose-

h is p id .

Representative specimens: *

LOUISIANA: Parish. St. Mary. Brown 18926. 19843 (LSU).

Parish unknown (GH).

10. Echinochloa walteri form 3

Plants annual. Culms strong, thick and usually purple at the \ base. Panicle spreading and nodding , branches mostly compound and

up to 9 cm long. Sheath glabrous, ligule none, nodes usually swollen

and glabrous. Spikelets green to purple-tinged, 3.0-4.0 mm long, awn variable from an awn point to ca. 3.0 cm long, mostly purple.

Sterile palea well developed, white. Anther 0.8-1.2 mm long,

brownish colored when dry. Fertile floret 2.8-3.5 mm long, 1.1-1.3

mm broad, apex subacute. Caryopsis 1.5-1.7 mm long, 0.9-1.1 mm

broad, brownish colored. Embryo 1.2-1.4 mm long, 0.4 mm less than

caryopsis length. Flowering from July to November.

Distribution: Scattered through the Florida parishes and South from

St. Bernard along the coastal marsh to Cameron Parish.

Discussion: Wiegand (1921), Hitchcock (1951) separate this form on i the basis of glabrous sheath. Both authors recognize E.. 62

lonqisetum Torr. as a synonym of E. walteri forma laevigata but

Hitchcock listed jE. lonqearistata Nash as a synonym of JE.

walteri. The main delimiting character was the glabrous sheath.

The specimen examined in this investigation from Gray Herbarium,

labeled 13. lonqisetum and annotated by Wiegand in 1919 was dis­

tinctly glabrous. While this specimen has long awns, the

pubescent lower sheaths can not be a certain Uue to the lack of

these parts.

Representative specimens:

LOUISIANA: Parish. Cameron, Thieret 9615a (USL). East

Baton Rouge, B-179, B-242, Cappaqe (?) (LSU). Jefferson,

B-327. Orleans, T. _F. Hall, Jr. (?) (NO), Penfound (?) (NO),

Sidney (?) (NO). St. Mary, B-188, B-232. St. Tammany,

Thieret 27165 (USL). Tangipahoa, Wascom 180 (LSU, SSC).

11. Echinochloa walteri form 4 •1 1 1 i ~ Plants annual. Culms strong but not as thick at the base as typical E. walteri. Panicle tight to only slightly spreading, purple only on the awns, branches compound. Sheath with papillose-hispid condition on all sheaths, the papillose base usually purple. Nodes were very hairy appressed pubescent, ligule with short hairs to 4.2 cm in a row of stiff white hairs. Spikelets 3.0-4.2 mm long, with awns up.to 3 cm long. Anther 0.9-1.2 mm long. Sterile palea usually one- half the length of fertile floret, white. Fertile floret 2.7-3.6 mm long, 63 1.1-1.3 mm broad. Caryopsis 1.3-1.6 mm long/ 0.9-1.0 mm broad, brownish colored. Embryo 0.8-1.2 mm long, 0.44 mm less than the caryopsis length. Flowering from'July to November.

Distribution: Restricted to the coastal .marsh region of Louisiana,

usually growing in water or at. the margin.

Discussion: This form has never been reported for any part of the

United States. The distinct hairy condition of the sheath and

stiff rows of ligulated hairs. E_. walteri is a semi-aquatic

taxon while this form has been found in the marsh where the

water remains throughout the year.

Representative specimens:

LOUISIANA: Parish. Calcasieu, B-303. Cameron, Thieret

8782 (USL). Jefferson Davis, B-330. St. Charles, Brown 20263

(LSU). Terrebonne, Charbreck 37 (LSU), Palmisano 23 (LSU).

12. Echinochloa zelayensis (H.B.K.) Schult. Mantissa 2: 269. 1824.

O'plismenus zelayensis H.B.K., Nov. Gen. etSp. 1: 108. 1815.

Panicum zelayense Steud., Nom. Bot. ed. 2. 2: 265. 1841.

Echinochloa crusgalli var. zelayensis (H.B.K.) Hitchc., U. S.

Dept. Agr. Bui. 772: 238. 1920.

Echinochloa crusgalli forma zelayensis Farwell, Mich. Head.

Sci. Papers. 26: 4. 1941.

Plants annual. Culms strong, slender, erect, producing only a few tillers. Panicle tight, erect, branches ascending and without 64 pustulate based hairs along the branches. Nodes glabrous, usually purple. Sheath strongly scabrous to only slightly. A young plant or a young sheath tends to be on minutely scabrous. Spikelet 2.6-3.3 mm long, green to purple tinged at the apex, sometimes short awned.

Anther 0.8-1.1 mm long, brownish colored. Sterile palea absent in most spikelets rudimentary in only a few. Fertile floret 2.1-2.7 mm long, 1.1-1.3 mm broad. Caryopsis 1.3-1.5 mm long, 1.0-1.2 mm broad, brownish colored. Embryo 1.1-1.2 mm long, ca. 0.27 mm less than caryopsis length. Flowering from July to November.

Type: Location unknown; based on zelayensis H.B.K.,

Nov. Gen. et Sp. 1: 108. 1815, Zelaya, Mexico.

Distribution: Sparsely scattered throughout Louisiana.

Discussion: Wiegand (1921) delimited E. zelayensis on the basis of

coriaceous subacute lemma, lower palea often wanting and nodal

and branch-setae of panicle much reduced or wanting. Hitchcock

(1951) and Ali (1968) placed it as a variety of E.. crusgalli. The

data gathered in the present investigation provides evidence that

E. zelayensis (H.B.K.) Schult., (n = 18) was a valid species

and not a variety of E_. crusgalli (L.) Beauv., (n = 27), but was

more closely related to E.. walteri (Pursh) Nash. The polygon

graph (Fig. 22) shows a closer relation to E. walteri than E.

crusgalli. The embryo-caryopsis (Fig. 20, Table III) ratio gives

a good indication that_E_. zelayensis was not a variety of E..

crusgalli. Representative specimens:

LOUISIANA: Parish. Bossier, B-279. Iberville, B-187.

Madison, B-314, Thieret 28247 (USL). St. Mary, B-236. i Tangipahoa, Wascorn 198 (SSC, LSU).

Distribution maps for the above taxa in Louisiana are shown as follows

Fig. 24. E. colonum

jS. colonum form JL.

2±* crusgalli

Fig. 25. E. occidentalis

E_. muricata

E_. muricata var. microstachva

Fig. 26. E. walteri

E.. walteri form _1

JE. w alteri form 2_

jS. walteri form 3,

E_. walteri form 4.

E.. zelayensis Fig. 24. Maps showing the distribution of E. colonum. E. colonum form JL and E_. crusgalli in L ouisiana. • E. colonum (o r i Fig. 25. Maps showing the distribution of E. muricata, E,. muricata var. microstachya and JS. occidentalis in Louisiana.

I 67

I • E. m u ric a ta i • E. m u ric a ta var. m lc ro sta c lya Fig. 26. Maps showing the distribution of E. zelayensis, E. walteri and four forms in Louisiana. 68

• E. w a lte ri E. w a lte ri (arm m E. w a lte ri to rn

E. w a iter) form

E. w a lteri form SUMMARY

The present investigation was undertaken to determine the taxonomic relationships of various taxa and forms throughout Louisiana.

The investigation included studies of anatomical, cytological, morpho­ logical and reproductive characters of the genus. A detailed investi­ gation of natural growing populations, greenhouse, fieldplot and herbarium specimens provided characters that were used for the delimitation of the taxa. These characters were anther, caryopsis, fertile floret, embryo, palea of lower floret, sheath, spikelet, and sterile lemma. Some other characters utilized to differentiate the taxa were, the nature of panicle, the presence or absence of pustulate based setae on the branches of panicle, length of panicle branches, length and width of blades, ligule, and presence or absence of stamens in the lower floret. On the basis of some of the above, characters, twelve taxa of the genus Echinochloa were recognized in Louisiana. .

Cytological studies confirmed the previous reports of two ploidy levels, the tetraploid, 2n = 36, and the hexaploid, 2n = 54, in all the taxa. Gould (1958) reported an octoploid (n = 36) for E^. crusgalli var. crusgalli on page. 766, and a hexploid (n = 27) for the same species on page 764, the latter of which is apparently in error. An octoploid

69 70 count has been confirmed. The study of the reproduction behavior indicated normal,sexual reproduction to all the taxa examined. The development of a mature caryopsis in an enclosed sheath indicated cliestogamy. The correlation of pollen size between ploidy levels was of a lesser magnitude than between populations of one taxon.

Analysis of pollen samples indicated that the variation between the two ploidy levels was small, while among each polymorphic taxon the variation was distinct. Echinochloa was found growing in wet and dryer habitats that may contribute to some degree the variation in pollen size.

Polymorphism among the taxa was also suggested by the diversity of characters of the abaxial surface of the leaves, while the characters of the sheath were found to be taxonomically signifi­ cant in all the taxa.

The caryopsis length compared with the embryo length was found to be significant in delimiting 13. muricata and E.. muricata var. microstachya from the other taxa. LITERATURE CITED

Akerberg, E. 1942. Cytogenetic studies in Poa pratensis and its hybrid with Poa alpina. Hereditas 28: 1-126.

Ali; Md. Arshad. 1968. The Echinochloa crusgalli complex in the United States. Ph.D. Dissertation, Texas A and M Univ., College Station, Texas.

Bell, C. R., 1959. Mineral nutrition and flower to flower pollen size variation. Amer. Jour. Bot. 46(9): 621-624.

Brown, W. V., 1950. A cytological study of some Texas grasses. Bull. Torrey Bot. Club 77: 63-76.

Brown, W. V. 1958. Leaf anatomy in grass systematics. Bot. Gaz. 119: 170-178.

Carnahan, H. L., and H. D. Hill. 1961. Cytology and genetics of forage grasses. Bot. Rev. 27: 1-162.

Church, George L. 1929b. Meiotic phenomena in certain Gramineae. II. Paniceae and Andropogoneae. Bot. Gax. 88: 63-84.

Darlington, C. D., and A. P. Wylie. 1955. Chromosome atlas of flowering plants. 2nd ed. Allen and Unwin, Ltd., London.

Davidson, J. F. 1947. The polygonal graph for simultaneous portrayal of several variables in population analysis. Madrono 9: 105-110. de Wet, J. M. J. 1954b. Chromosome numbers of a few South African g ra sse s. Cytologia 19: 97-103.

Emery, W. H. P. 1957. A study of reproduction in Setaria macrostachya and its relatives in the southwestern United States and norhtern Mexico. Bull. Torrey Bot. Club 84: 106-121.

Fairbrothers, D. E. 1952 . A cytotaxonomic investigation within the genus Echinochloa. M. S. Thesis, Cornell Univ., Ithaca, N. Y.

Fairbrothers, D. E. 1956. Nomenclatural change in the grass genus ‘Echinochloa. Rhodora 58: 48.

71 72

• Fassett, Norman C. 1949. Some notes on Echinochloa. Rhodora 51: 1-3.

Fernald, M. L. 1915. Michaux's Panicum Muricatum. Rhodora 17: 105-107.

Fernald, M. L. 1950. Gray's Manual of Botany. American Book Company, New York. 1632.

Gleason, H. A ., and A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada . Van Nostrand Co.

Gould, F. W ., 1957. Pollen size as related to polyploidy and speci- ation in the Andropoqon saccharoides - A. barbinodis complex. Brittonia 9: 71-75.

Gould, F. W. 1958. Chromosome numbers in southwestern grasses. Amer. J. Bot. 45: 757-767.

Gould, F. W. 1966. Chromosome numbers of some Mexican grasses. Canad. J. Bot. 44: 1683-1696.

Gould, F. W. 1968. Chromosome numbers of Texas grasses. Canad. J. Bot. 46: 1315-1325.

Heiser, C. B., Jr., andT. W. Whitaker. 1948. Chromosome number, polyploidy and growth habit in California weeds. Amer. J. Bot. 35: 179-186.

Hitchcock, A. S. 1920. The North American Species of Echinochloa. Contr. U. S. Nat. Herb. xxii. pt. 3, 133-153.

Hitchcock, A. S. 1935. Manual of the grasses of the United States. U. S. Dept. Agric. Misc. Publ. 200. 1040 pp.

' Hitchcock, A. S. 1951. (Revised by Agnes Chase) Manual of the Grasses of the United States. U. S. D. A. Misc. Pub. #200. 1051 pp.

Hunter, A. W. S. 1934. A karyosystematic investigation in the Gramineae. Canad. J. Res. 11: 213-241.

Kamakrishnan, P. S. 1960. Ecology of Echinochloa colonum (L.) Link. Proc. Indian Acad. Sci. Sect. B. 52(3): 73-90. 73

Krishnaswamy, N ., and V. S. Raman. 1949. A note on the chromo-- some numbers of some economic plants of India. Curr. Sci. 18: 376-378. 1

Link, H. F. 1833. Hort. Berol. 2: 209. (Cited from Hitchcock 1951). i Linnaeus, C. 1753. Species plantarum .. 1st ed. Vol. I. page 56. Bartholomew Press, Dorking, Great Britain.

Martin, A. C. 1946. The comparative internal morphology of seeds. Amer. Midi. Nat. 36: 513-660. (Gramineae, pp. 536-542.)

Metcalfe, C. R. 1960. Anatomy of the . Vol. I. The Gramineae. Oxford, Claredon Press.

Michaux, Andre. 1803. Flora Boreali-Americana. Vol. I. Paris, page 47.

Mulligan, G. A. 1961b. Chromosome numbers of Canadian weeds. III. Canad. J. Bot. 39: 1057-1066.

Palisot de Beauvous, A. M. F. J. 1812. Flore D'Oware Et De Benin EnAfriquell. Paris. Pages 14, 80, plate 58, fig. 1.

Poiret, J. L. M. 1816. Lamark's Encyclopedie Methodique Botanique. Sup. Tome IV. Paris. Page 273.

Pursh, S. 1814. Flora of North America. Vol. I. White, Cochrane, and Company, London. Pages 66-87.

Radford, A. E., H. E.. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill.

Rau, N. S. 1929. On the chromosome numbers of some cultivated plants of South India. Jour. Indian Bot. Soc. 8: 126-128.

Reeder, J. R. 1957. The embryo in grass systematics. Amer. J. Bot. 44: 756-768.

Rydberg, P. A. 1931. Taxonomic notes of the flora of the prairies and plains of central North America. Brittonia 1: 81-82.

Sass, J. E. 1958. Botanical Microtechnique. 3rd ed. Iowa State College Press, Ames, Iowa. 74

Schoch-Bodmer# Helen. 1940. The influence of nutrition upon pollen grain size in Lvthrum salicarla. J. Genetics 40: 393-402.

i Shinners, L. H. 1954. Notes on North Texas grasses. Rhodora 56: 33-34. i

Stewart# D. R. M. 1965. The epidermal characters of grasses# with special reference to East African plains species. Bot. Jb. 84(1): 63-116.

Tateoka# T. 1965. Chromosome numbers of some East African grasses. Amer. J. Bot. 52: 864-869.

Wiegand# K. M. 1921. The genus Echinochloa in North America. Rhodora 23: 49-65.

Yabuno# T. 1962. Cytotaxonomic studies on the two cultivated species and wild relatives in the genus Echinochloa. Cytologia 27: 296-305.

Youngman# W.# and S. C. Roy. 1923. Pollination methods amongst the lesser millets. Agr. Jour. India. 18: 580-583. APPENDIX 76

APPENDIX A

Plants grown in field plots and greenhouse.

Taxa Collection Number

E.. colonum 129, 130, 155, 158, 184, 203, 204, 208, 231, 241, 278, 345, 348, 349

E. colonum form j;, 287, 302, 337

JE. crusgalli 131, 133, 134, 135, 137, 140, 183, 212, 217, 253, 255, 256, 257, 258, 263, 267, 273, 308, 270

E_. muricata 247, 281, 286, 309

E_. muricata var. microstachya 247A, 342

JL* occidentalis 207, 246, 249, 250, 259, 260, 263, 265, 271, 283

JE. walteri 178, 237, 243

form JL. 138, 248

form 2. Brown 18926

form 3. 179

E.. zelayensis 187, 236, 279, 314 77

APPENDIX B

Plants examined for leaf anatomy.

Taxa Abaxial Surface

E.. colonum 191, 204, 209, 211, 219/ 234, 249

E.. colonum form ^ 287

E_. crusgalli 215, 216, 257, 258, 273

E_. muricata 272, 281, 286

muricata var. microstachya 247A, 342

E.. occidentalis 207, 231, 271

E_. walteri 210, 232

form 1_ 143, 248

form 2_ Brown 18926

form 3. 179

form 4, 303

E,. zelayensis 314, 236 APPENDIX C, NO. 1

Frequency distribution of taxa. Spikelet lengths in percentages.

Spikelet Length (mm) 1.9- 2.1- 2.3- 2.5- 2.7- 2.9- 3.1- 3.3- 3.5- 3.7- 3.9- 4.1-- 4.3-• 4.5- 4.7- 4.9- 5.1 Taxa 2.0 2.2 2.4 2.6 2.8 3.0 3.2 3.4 3.6 3.8 4.0 4.2 4.4 4.6 4.8 5.0 5.2 E. colonum 9.3 27.5 32.6 24.65.9 1.3

E. colonum form 1. 21.3 57.413.1 3.2 4.9

E. crusgalli 3.8 10.3 19.7 15.9 29.6 16.33.2 1.1

E. muricata 29.7 27.624.8 9.0 5.4 3.5

E. muricata var. microstachva 3.612.3 30.2 29.4 17.0 7.5

E. occidentalis 15.6 35.936.7 9.6 1.7 ' E. walteri 5.06.721.0 37.8 13.410.0 2.5 1.7 1.7

form 1. 0 .2 0 .4 12.4 29.0 12.720.8 6.4 6.2 0.2

form 1_ 9 .0 9 .0 13.5 22.6 9.0 31.8 4.5 form 3. 8 .5 10.6 10.6 46.7 14.8 10.6

form A 3 .8 19.3 61.4 3.8 3.8 7.6

E. zelayen sis 3 .7 25.9 33.333.3 3.7 APPENDIX C, NO. 2

Frequency distribution of taxa. Anther lengths in percentages.

Anther Length (mm) * Taxa 0.4 0.5 0.6 0.7 0.8 0.9 1.0 i=.l 1.2 1.3 1.4

E. colonum .5 25.6 40.2 19.5 6.1

E. colonum form _1 20.0 36.0 40.0 4.0

E. crusgalli 5.6 12.7 23.9 21.1 23.9 12.7

-E. muricata 3.1 9 .6 26.7 22.3 13.8 18.1 4.2 2.1

E. muricata microstachya 28.0 32.0 24.0 16.0

E. occidentals 3.2 35.5 32.2 29.0

E. w alteri 3.1 9.5 7.9 31.1 28.5 15.8 1.6 1.6

form _1 27.2 54.5 9.1 9.1 -

form 2_ 50.0 50.0

form 3. 17.5 6.3 12.5 25.0 43.7

form A 26.6 . 20.0 40.0 13.3

E. zelayensis 20.0 53.3 13.3 13.3 APPENDIX C, NO. 3

Frequency'distribution of taxa. Fertile floret lengths in percentages.

Fertile Floret Length (mm) 1 .8 - 2 .0 - 2 .2 - 2 .4 - 2 .6 - 2 .8 - 3 .0 - . 3.2- 3.4- 3.6- 3 .8 - 4.0- 4.2- 4.4- Taxa 1.9 2.1 2.3 2.5 2.7 2.9 3.1 3.3 3.5 3.7 3.9 4.1 4.3 4.5 E. colonum 32.8 51.8 15.4

E. colonum form _1 57.2 42.8

E. crusgalli 3.3 3.3 6.5 31.1 13.1 24.5 14.7 3.3

E. muricata 9.6 22.6 22.6 22.6 16.1 3.3 3.3 E. muricata var. microstachya 28.0 30.0 24.0 8.0

E. occidentalis 32.0 41.0 26.0

E. w alteri 8.0 40.0 32.0 12.0 4.0 4.0

form JL. 12.5 31.2 25.0 13.8 12.4

form 2_ 20.0 20.0 20.0 20.0 20.0 form 3. 37.5 43.8 12.4 6.2

form 4. 17.5 25.0 50.0 12.5

E. zelayensis 18.2 45.4 18.2 18.2 APPENDIX C, NO. 4

Frequency distribution of taxa. Fertile floret width in percentages.

Fertile Floret Width (mm) Taxa 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 2 .0 2.1 2.2

E . colonum 1.7 22.4 32.7 25.9 13.8 1.7 i .7

E. colonum form _1 7.1 21.4 14.3 14.3 28.5 14.3

E. crusgalli 1.6 12.7 7.9 25.4 20.6 22.3 9.5 E. muricata 9.625.8 22.616.112.9 9 .6 3.3

E. muricata var. microstachya 15.4 30.7 23.0 19.2 7.7 3.9

E. occidentalis 16.0 24.0 44.0 16.0

E. w alteri 4.5 22.7 22.7 31.8 18.2

form _1 55.5 44.4

form 2_ 20.020.0 40.0 20.0

form 3_ 33.341.7 25.0 form 4_ 20.0 40.0 40.0

E. zelayensis 18.1 36.4 45.4 APPENDIX C , NO. 5

Frequency distribution of taxa, Caryopsis length in percentages.

Caryopsis Lencrth (mm) Taxa 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 2 .0 2.1 E . colonum - 4.1 58.3 37.5

E. colonum form 1_ 8.1 54.6 9.1 18.1

E. crusgalli 4.6 4.6 18.2 13.6 22.7 22.7 13.6

E. muricata 4.2 29.1 8.3 16.7 8.3 16.7 12.5 4.2

E. muricata var. microstachya 9.1 4.5 27.2 31.8 9.1 13.6 4.5

E. occidentalis 27.3 36.3 18.2 18.2

E. w alteri 18.4 28.9 26.4 21.1 2 .6 2 .6 form 1_ 12.5 12.5 50.0 25.0 form 2_ 37.5 50.0 12.5

form 3_ 8.3 25.0 66.6

form 4. 11.1 41.4 33.3 11.1

E. zelayensis 11.2 55.5 33.3 APPENDIX C, NO. 6

Frequency distribution of taxa. Caryopsis width in percentages.

Caryopsis Width (mm) Taxa 0.9 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 2.0 2.1

E. colonum 4.1 25.0 41.7 29.2 E. colonum form JL ■ 36.4 45.4 18.2

E. crusgalli 9.1 9.1 27.2 22.7 22.7 9.1

E. muricata 16.7 20.8 29.1 20.8 4.2 4.2 4.2

E. muricata var. microstachya • 4.5 50.1 22.7 13.6 4.5 4.5

E. occidentalis 27.3 36.3 27.3

E. w alteri 2 .6 50.1 31.5 15.8

form JL. 12.5 50.0 37.5

form 2_ 25.0 25.0 25.0 25.0

form 3_ 8.3 33.3 58.3

form A 55.5 44.4

E. zelayensis 11.2 66.6 22.2 James Howard Brooks was born in Shelby County, Texas in 1932,

attended Timpson High School and was graduated in 1950. He entered

the University of Texas on an athletic scholarship and played football

for three years. He volunteered for the Army after receiving an injury

playing football. He spent one year of his Army service in Germany.

He entered Stephen F. Austin State College after his release from the service and finished his B.S. degree in 1957. Prior to finishing his B.S. degree, he married and presently has two boys. He taught

biology and chemistry at Bridge City High School in Bridge City,

Texas and after teaching there for two years, received a summer

NSF fellowship at Stephen F. Austin State College to work toward his Master's degree. In 1961, he accepted "a position at Stark High

School in Orange, Texas and taught biology, and in the summer of

1964 finished his Master's degree. He accepted an appointment at

Stephen F. Austin State College for September, 1965 and the early part of 1966 he accepted a Graduate Assistantship from Louisiana

State University to work toward a Ph.D. degree. In September,

1968 he accepted an appointment at McNeese State College where he is presently employed. EXAMINATION AND THESIS REPORT

Candidate: James Howard Brooks

Major Field: Botany

Title of Thesis: Cytotaxonomy of the Genus Echinochloa in Louisiana

Approved:

Cd/MOi $ Major Professor and Chairman

Dean oftho Graduate School

EXAMINING COMMITTEE:

Date of Examination:

July 24, 1969