This dissertation has been 65—3903 microfilmed exactly as received

PINKAVA, Donald John, 1933- BXDSYSTEMATIC STUDY OF GENUS DC. (COMPOS1TAE).

The Ohio State University, Ph.D., 1964 Botany

University Microfilms, Inc., Ann Arbor, Michigan BIOSYSTEMATIC STUDY OF OENUS BERLANDIERA DC. (COMPOSITAE)

DISSERTATION

Presented in Partial Fulfillment of tha Requirement* for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

DONALD JOHN PINKAVA, B.Sc.t M.Sc.

The Ohio State University 1964

Approved by

Adviser Department of Botany and Pathology PLEASE NOTE Figure pages are not original copy. They tend to "curl". Filmed in the best possible way. University Microfilms, Inc. ACKNOWLEDGMENTS

I wish to express my sincere gratitude to Dr* T.

Richard Fisher, my adviser, who not only suggested thia problem, but moat effectively guided it to ita completion*

I am thankful alao to Drs* Clara G. Weiahaupt, Emanuel 0.

Rudolph, Dale A. Ray and Carroll A* Swanson for reading thia dissertation and for their helpful criticisms and suggestions*

Special recognition is extended to John M* Speer*

Unless otherwise cited, the photography is to be credited to his talents so unselfishly shared*

I am deeply indebted to Dr* Ray for statistical assistance; to the curators of the herbaria for loaned specimens and/or photographs; and to my colleagues for

their many useful suggestions*

Financial assistance was provided by The Ohio State

Research Foundation and the National Science Foundation*

ii TABLE OF CONTENTS

Pag*

ACKNOWLEDGMENTS...... 11

LIST OF T A B I E S ...... It

LIST OF ILLUSTRATIONS...... Yl

INTRODUCTION...... 1

GENUS D E S C R I P T I O N ...... 6

HISTORY OF G E N U S ...... 12

ECONOMIC I M P O R T A N C E ...... 16

ARTIFICIAL KEY TO HERLANDIERA T A X A ...... 17

DESCRIPTION AND DISCUSSION OF T A X A ...... 19

PROBLEMS OF NOMENCLATURE...... 66

EXPERIMENTAL METHODS AND MATERIALS...... 68

CYTOGENETICS ...... 75

CROSSABILITY...... 10A

GERMINATION AND SURVIVAL...... 113

ANALYSIS OF PARENTAL EXPERIMENTAL P L A N T S ...... 117

KEY TO AND ANALYSIS OF HYBRID EXPERIMENTAL . . . 129

FIELD COLLECTION ANALYSIS (EAST) ...... l46

FIELD COLIECTION ANALYSIS (WEST)...... 157

DISTRIBUTION M A P S ...... 168

SUMMARY ...... 172

BIBLIOGRAPHY ...... 17^

AUTOBIOGRAPHY...... 178

111 LIST OF TABLES

1 Summary of chromosome count* reported in lltormturo ......

2 Chromosome data of current s t u d y ...... •

3 Pollen atainability data from fisld oollsctlons......

4 Genetic data supporting tstraploidy hypothesis ......

5 Chi-square tsst of gsnstic data supporting tstraploidy hypothssis ••*•••••••

6 Sslfing achene-set data ......

7 Summary of achsns-sst data from artificial crosses ......

8 Germination and ons-ysar survival records of artificial crosses ...... ••••

9 Germination data of field-collected fruits of eastern Berlandiera taxa ......

10 Morphological analysis of parental experi­ mental plants ......

11 Morphological analysis of artificial hybrid plants ••••»•• ......

12 List of herbaria specimens used in hybrid index analysis of eastern taxa complex • • .

13 Hybrid index analysis of eastern taxa complex ......

14 Analysis of variance between populations of eastern taxa complex

15 Analysis of variance between species-hybrid groups of eastern taxa complex ••••••

iv LIST OF TABLES (Continued) Table Page

16 Liat of herbaria specimens used in hybrid index analysis of western taxa complex . • . 158

17 Hybrid index analysis of western taxa complex • • 160

v LIST OF ILLUSTRATIONS Figure Page

1 DC. (Holotype illustration)...... 7

2 Berlandieralyrata Benth. var. lyrata (Holotype ...... 21

3 var. macrophylla 6ray (Holotype, Sheet 1) ...... 32

Berlandiera lyrata var. macrophylla Oray (Holotype, Sheet 2) ...... 32

5 Berlandiera aubacaulia Nutt. (Holotype) .... 37

6 Berlandiera x humilia (Small) comb. nov. Tfeolotype of B. humilia Small)...... kz

7 Berlandiera x humilia (Small) comb. nov. (Ieotype of B. humilia Small) ...... kz

8 Berlyidiera pumila (Michx.) Nutt. (Holotype) ...... kk

9 (Michx.) Nutt. (Holotype of caroliniana Poiret) ...... kk

10 Berlandiera pumila (Michx.) Nutt. (Holotype and paratypea of B. tomentosa var. deal­ ba t a T. & 0.) . 7 ...... k6

11 Berlandiera pumila (Michx.) Nutt. (Paratypea of B. tomentosa var. dealbata T. & G.) . . .

12 Berlandiera texana DC. (Iaotype) ...... 5^

13 Berlandiera texana DC. (Holotype of B. longifolia Nutt.) ...... 51*

l*f Berlandiera x betonicifolia (Hooker) comb. nov. (Syntype of betonicifolium Hooker) . . . . . 7 7 ...... *1

vi LIST OF ILLUSTRATIONS (Continued)

15 Berlandiera x betonicifolia (Hooker) comb, nov. (Syntype of Silohium betonicifolium Hooker)

16 Microsporogenesis I ......

17 Microsporogenesis II ......

18 Microsporogenesis III ......

19 Microaporogenesia I V ...... • .

20 Microaporogeneaie V ......

21 Microsporogenesis VI ......

22 Heteromorphic Pollen ......

25 Corolla Colora ......

2*+ Leaf Shapea ......

25 Pubescence Types I ......

26 Pubescence Types II ......

27 13. lyrata var. lyrata 1 0 1 0 ......

28 B. texana 1002 ...•• ......

29 B. x betonicifolia 655 ......

30 B. x betonicifolia 657 ......

31 — * Puaila 525 ......

32

33 lyrata var. lyrata 35-*+......

B. lyrata var. lyrata x subacaulls 5-5 . • • LIST OF ILLUSTRATIONS (Continued) Figure Page

35 B. subacaulis 11-2 ...... 131

36 B. texana 660-1 ...... , 13 2

37 B. lyrata var. lyrata x texana 40-2 ...... 132

38 B. lyrata var. lyrata x oumila 255-1 ..... 132

39 B. texana x oumila 197-3 ...... 134 40 B. texana x subacaulis 2-1 ...... 134

41 B. oumila x subacaulis 170-2 ...... 134

42 B. lyrata var. lyrata x B. x betonicifolia i i 8 - i ...... 135

43 B. texana x B. x betonicifolia 176-1 ...... 135

44 B. x betonicifolia x B. pumila 242-1 ...... 135

45 B. x betonicifolia x B. subacaulis 144-1 . . . . 136

1*6 offspring of B. lyrata var. lyrata x F2 oumila 31-1 ...... 136

offspring of B. subacaulis x texana 469-1 . . 156 1*7 F2 48 Histograms of hybrid index values of eastern and western taxa complexes ...... 163

49 Distribution map of eastern t a x a ...... 168

50 Distribution map of western taxa ...... 170

viii INTRODUCTION

The genus Berlandiera was described by Augustin-Pyrasue

De Candolle in 1828 in honor of Jean Louis (or Luis) Berlandier who had served as his eaissary to the then relatively unknown

Texas region and as a scientist of the Mexican Boundary

Coaaission (1827-1829)* Although born into a poor faaily near

Fort-de-l'£cluse, France, Berlandier's initiative and keen interest in natural sciences enabled hia, as a young aan in his early twenties, to be the first to collect extensively and explore the boundary region* According to Qeiser (1937) and McKelvey (1933)* the travels of Berlandier extended froa

Mexico City to the Trinity River via Laredo and San Fernando de Bejar along the Old Bejar Road* The expedition's objective was Nacogdoches, but Berlandier and others of the party contracted aalaria* The scientific staff never crossed the

Trinity River but were forced back to Bejar by June, 1828*

During this tiae Berlandier collected a series of speciaens that later were naaed in his honor, Berlandiera texana DC*

With Berlandier ill and his speciaens daaaged again and again by the incleaent weather, the elder De Candolle's hopes of a large and valuable collection froa a virgin land faded*

Subsequent sojourns were aade later in 1828 and again the following year but none included the type locality of B* texana* After the Commission had completed its task, Berlandier remained in , married and settled at Matamoras. He made still other excursions, particularly one to Goliad and Bejar in l$3k. He continued his investigations until his death in 1851 when he drowned in an attempt to cross the San Fernando River south of his home.

As to his scientific accomplishments, McKelvey (1955) lists his "Memories de la Comleion de Limites. Historia Natura

por El General Teran y L. Berlandier" (1832) which contains descriptions of eleven newly proposed species and four new genera; his handwritten Expedition (in U.S. National Museum); co-authorship with Raphael Chovel in 1850 of Diario de viage de la comision de limites aue puso el gobierno de la republica. ba.io la direccion del Bxmo: his collections distributed among various herbaria, particularly Gray at Harvard University,

New fork Botanical Garden, Paris, London, Geneva, Leipzig and

Vienna; his manuscripts at Yale, Library of Congress, Library of Smithsonian Institution, University of Texas, and Gray

Herbarium Library; his zoological specimens; and his meteoro­ logical and topographic observations.

Geiser in 1937 defends the reputation of Berlandier from severe criticisms of A.P. De Candolle and Alphonse De Candolle

(published by the latter in 1862) and of Asa Gray (in 1863).

Alphonse De Candolle's criticisms of Berlandier for poor speci­ mens of insufficient quantity and quality to the value of but one-quarter what was expected — all due to neglect and misconduct — la difficult to Interpret since Alphonse Do

Candolle received tho complete assortment of 2320 "numbers" including 55,07? plant specimens plua aooda, insects, blrda, aolluaoa and other aatoriala preserved in alcohol (Ooiaor, 1937)*

McKelvey (1955) eoaaonta that "in light of tho roeorda concerned with hia aeooapliahaonta, Borlandior'a r$le appoara to have boon aoro croditablo than tho ono played by tho ronownod Do Candolle."

Sarly workora of tho gonua woro eoncornod primarily with

doaeriptiro aorphology, largoly of vegetative charaetora, as in

original doaeription, rodofinition of taxa, and differentiation

aaong taxa via "keys." In hia descriptive work, Qray (1842)

roeogniaoa internediacy in charaotora among plants of tho

"betonicifolia-texana" complex* Small (1903) separates out

from B* oumila an intermediate to B* subacaulis*

Some time later, working with Texas species, Shinners (1951)

and Turner and Johnston (195&) recognise intermediacy, using

overlapping characters in their respective keys* Shinner's

observations on the "betonioifolia-texana" complex are based on

the number of leaves below the inflorescence, the length of the

petiole, the leaf shape, and the pubescence as distinctive

characterst Turner and Johnston employ the number of leaves in

the upper ten centimeters of stem from below peduncles of

flowering heads, the length of petiole, and the type of pubes­

cence to delineate plants of the "texana-betonicifolia-pumila

(including the variety dealbata)11 complex* Dyar (1959) performs a hybrid index analysis of ths

"pualla-texana" complex vis harbaria specimens, delegating significance to laaf blada width-length ratio, intarnoda langth above laaf, patiola langth, and basa angla of laaf -- all aaaauraaants froa tha aiddla cauline laaf of aaeh spaciaan. Tha rasult is a histogram with two paaks corresponding rathar wall to tha two spacias in quastion, but linked by many intermediates*

No one oharaetar, she continues, could be used to single out tha variety betonicifolia. postulated product of hybridisation of tha two spacias with subsequent intrograssion* She accounts for tha variability as possibly due to ecological influence. No

"perfect" spaciaan of either spacias was found.

Chroaosoae counts are reported by Turner and Johnston

(1956)t Dyar (1959)* Turner, Powell and Xing (1962); and Da Jong and Longpre (19&5)* Chroaosoae aorphology was briefly consider­ ed by Turner and Johnston (1956). To ay knowledge tha only hybridisation program carried out involving Berlandiera v^as by

Fisher (unpublished).

Tha major objective of this three-year study is to attaapt interpretation of natural populations collected in tha field by naans of analyses of artificial crossing prograa, cytology, pollan stainability, and comparative aorphology. Consideration of anatoaical study, chroaotography, and pollan growth rates are in future plans, as is additional study of B. lyrata var. macrophylla. Herbaria specimen citations follow abbreviations standardized by Lanjouw and Stafleu (1964): British Museum (BM);

Chicago Natural History Museum (F); Gray Herbarium of Harvard

University (OH); KEW (K); Herbarium of State University

(LSU)| Botanical Garden (M0){ Michigan State University

Herbarium (MSC); Tulane University Herbarium (NO); The New York

Botanical Garden (NY)t Herbarium of University of

(NCU)f The Ohio State University (OS); Muslum National d'Bistoire

Naturelle, Paris (P); Philadelphia Academy of Natural Sciences

(PH); and Herbarium of Southern Methodist University (SMU). An asterisk (*) precedes abbreviations if specimen(s) cited have intermediate characters* DESCRIPTION OF GENUS

Berlandiera De Candolle, A.P., 1836. Prodromus. Part V: 517* Illustrated by Heyland in leones Selaectae Plantarum by Benj. De Lessert, 1839* IV: 11, Tab. 26. (Figure 1)

Berlandiera is a New World genus of plants belonging to the Melampodinae of the Helianthae, tribe Tubuliflorae, family

Compositae. Closest relationship is to Silnhium. less affinity to Melampodia. Engelmannia. Parthenium. and Polymnia. Its distribution extends along the coastal plain from through Texas, southward into Peninsula and northward into the Ouachitas and the Ozarks; westward through into the Chiracahuas in southeastern and on into the mountainous regions of northern Mexico* There is a definite scarcity of plants in most of , Mississippi, and eastern

Louisiana, interrupting the otherwise continuous range. This gap in distribution, as well as the northern limits of eastern taxa, may be explained, in part, on the basis of a map of southern forests by Hedlund and Janssen (1963)* See Figures *+9* 30 on pages 169* 171* All taxa, except B. lyrata. are usually asso­ ciated with mixed pine, pine-oak, or oak-hickory forests but not with either bottomland oak-gum-hickory forests or with pine forests without slash or longleaf pines, singly or in combination.

The pinnatifid-leaved taxa usually grow in exposed areas, often ( (Hi *• - S l H H H M M lit*- likafi.

Figure 1. Berlandiera texana D.C. Copy of Holotype Illustration by Jean- Christophe Heyland (Plantarum, 1839), Scale: 0.64x. — — ^ along roadsides and In lawns; the others usually border woodland areas in light shade. All are rather restricted to dry or well- drained soils.

Roots. All taxa have a perennial tap root system, becoming woody with age« from which develop one to several shoots. Roots grow in soils ranging from sands to dry hard rocky limestone types. Vegetative reproduction from any portion of plants appears to be rare or absent in native habitats* although attempts to propagate stem cuttings of members of various taxa and inter­ specific hybrids under greenhouse conditions have been successful with and without the use of rooting hormone.

Stems. Stems are terete* solid, and have resin ducts. They range in height from but a few inches in some species (or dwarf forms of others) to three or four feet. They are herbaceous to suffrutescent, decumbent to erect, branched or unbranched, scapose to leafy throughout. The inflorescences are solitary to paniculate-corymbose.

Leaves. Leaves are highly variable, even on the same plant, yet leaves still serve to distinguish taxa. They are alternately arranged, rarely opposite. Leaves vary in shape from ovate to obovate to deltoid to spatulate; their bases, from cuneate to truncate to cordate; their apices, from obtuse to acute; their petioles, from short (even absent) to long; their margins, from dentate to doubly serrate to sinuate-

(alternately) to lyrate-pinnatifid. Blade width-length ratios range from 20 to l*tO percent. Palisade tissue, at least in cortain taxa, ia of one to two colls thick; stomates aro on both surfaces (Wallace Weber, unpublished).

Heads. Individual heads are 1.2-2.7 cm wide, of many flowers. Phyllaries are spreading, leaf-like, with margins crenate, serrate or nearly entire; in three series, becoming progressively larger toward the summit. The lowest phyllary may arise from the peduncle as a scale. The lowest 3-4 phyllaries are oblong to oval; the middle 4-3, usually obovate; the upper ones (really paleae), as many as the ray flowers, obovate to rhomboid. Phyllary texture is membranous, becoming chartaceous and reticulate with drying and age. The receptacle is small, somewhat turbinate, with a flattened to depressed upper surface.

Upon abscission of phyllaries, string-like appendages are retained about the receptacle. Ligulate flowers are monosporangiate, ovulate, fertile, in a single series of 5-10, usually 8, inserted at the axils of the uppermost phyllaries (paleae). Rays are pale to bright yellow, the latter becoming orange upon drying. The strap-like portion distal to the short tubular corolla base atop the achenes measures 4-15 mm wide, 7-24 mm long and is notched or sometimes cleft or tubular. Nine to twelve anastomosing veins on the under surfaces of rays are either green or red to maroon.

Colors often change with drying and hence are unsafe to determine from herbarium specimens. Amber to reddish resin dots appear on the drying leaves, stems, and flower parts. The style is two- or sometimes three-parted, lined, pubescent on outer surface.

Disc flowers are monosporangiate, staminate, their tubular 10 corollas 3-4 mm long, five-toothed, either yellow or red to maroon* Five blackish coherent anthers surround a yellow style which is pubescent toward its summit* The pappus is an incon­ spicuous crown of aristate teeth. Paleae progressively narrow toward center of head. Adherent to the ripening achene and ab­ scising with it are two disc flowers and their abaxile enclosing paleae. The degree of enclosure about the disc flowers also pro­ gressively decreases toward center of head. Paleae have dentate to eroae apices.

Achenes. The achenes are 2.8-4.8 mm wide, 4.0-7.5 sun long; obcompressed with a pubescent inner surface. At the achene apex the pubescence tufts about the inconspicuous pappus crown of teeth which are sometimes pubescent, and about additional short awns if present. The achenes are truncate or shallowly notched at the apex, but are not winged. Germination occurs in 2-30 days under greenhouse conditions, without requirement of cold treat­ ment.

There are two cotyledons (or falsely three via splitting) which are fleshy, entire, oval in outline. The first several leaves of the seedling usually form a basal rosette and have shapes often much different from leaves developing later. The first leaves of pinnatifid-leaved taxa are only oval to spatu- late with crenate to dentate margins.

Pubescence throughout is of multicellular, jointed hairs which vary in thickness, length, density and amount of purple pigment (Figures 25* 26 on pages 121, 122). 11

The common name of the genua la Green Eyea (Small, 1933) becauae after disc flowera fall, the center of the head appeara green (paleae) aurrounded by the peraiating yellow raya. HISTORY OF GENUS

The genus Berlandiera was originated by A.P. De Candolle who, in his Prodromus. assigns to it only the type species, B, texana.

In l84l, Thomas Nut tall transfers three eastern U.S. flllphium taxa to genus Berlandiera. namely £. pumilum of Michaux

(1805), Pursh's &• tomentosa (l8l4), and his own £• subacaule

(1821). Curiously enough, Nuttall's original description does not clearly delineate "subacaule," the second word therein, from the remainder of the passage. This led John Torrey in 1826 to assign to what he considered an anonymous taxon the epithet

"nutt&llianum," In his observations, Torrey mentions: "First discovered in Florida by Mr. Ware, and described but not named by Nuttall." The comments appear in an account of plants collected in the west by E.P. James, and Torrey considers both the western species "lyrata" and its eastern counterpart

"subacaule" as the same taxon because in 1842 he and Asa Gray create another name, "incisa," to include the western plant of

James but not the Florida plant of Ware which they assign to £. subacaulis (Nutt.) T. & G. Thus they place jS. nuttallianum. in part, in synonomy to B. Incisa. and remark that it has more cauline leaves than has B. lyrata of Bentham (1839)* Neverthe­ less, Gray places B, incisa in synonomy to B. lyrata in 1884.

12 13

Thus, the Florida plant collected by Ware belongs to B. suba-

caulis Nutt.

In 1842, Torrey and Qray introduced two new valid varieties

to established species of the genus. The first, a western taxon

with a fine white tomentum, is assigned to _B. tomentosa as the

variety dealbata. differing by being larger, leafier, and with

more heads than the variety tomentosa. The holotype was collect­

ed by Nattall along the "Arkansa", but with it is mounted para-

types, similar plants collected in Texas by Drummond. Qray

(1884) places £. pumila in synonomy to B. tomentosa. but Trelease

(1891) completely reverses this decision since publication of B.

pumila antedates that of B. tomentosa by eleven years. There­

fore "dealbata" became a variety of B. pumila until Small (1903)

raised it to specific status, B. dealbata (T. & G.) Small.

Turner and Johnston (1936) consider as possible "isotype" of

tomentosa var. dealbata a typical specimen of B. pumila and

accordingly assign it there. The plant considered is not an iso­

type, but as they stated, is "fairly typical" of B. pumila.

However, they annotated the paratype of B. tomentosa var. deal­ bata as B. texana var. betonicifolia.

The second variety named by Torrey and Qray (1842) is B.

texana var. betonicifolia. formerly Hooker*s Sllphium betonic1-

folium (1833) based on syntypes having petiolate leaves with

deeply and coarsely crenate margins and with peduncles "clothed with beautifully jointed, purplish hairs." According to herb­ arium labels, these syntypes were collected by Drummond in New Orleans, but this locale was queatioaed by Torrey and Qray

(1842)• At tha aaaa tiae thay aantlon that plants eollaotad by

Dr. Hala in Lottisiana "have tha padunelas and nppar part of tha stas elothad with purplish hairs (oolorad by tha daposition of a rssiaous natter), just as Hooker daseribas his Silohiua beton- ielfoliuni but tha uppar leaves ara all sessile, as in Da

Candolle's plant|jj. texanaj..." Bara in reeognition of intor-

■adiata speciaens. In 1903, B. taxana var. batonicifolia was elevated to B. batonicifolia by 8sall, only to ba raturnad to its rariatal position by Turnar and Johnston (1936)* Huttall

(1841) daseribas a naar ally of typical B. taxana as B. loniti- folia. Tha holotypa fros tha plains of tha Bad River, , is about two fast tall with erowdad ooryabs, short-patiolata leaves, and "padunelas and stas olothad with dense, long and soft hairs, but not oansscent or toaentose." It has since bean assigned to B. taxana by Qray (1884) and to its variety baton­ icifolia by Turner and Johnston (1936)*

A third variety introduced by Torray and Qray (1842) is B. subacaulis var. £., described but not naaad and hanca as yet in­ valid. A possible isotype (collected by Leavenworth in Florida) and tha original description suggest characters intersedlate between B. nusila and B. subacaulis. but tending sostly toward tha latter. This is interesting because in 1903 Snail recog­ nises another intarsadiata between these same two taxa as B. hunills. based on a plant collected by Mary Reynolds at St. 15

Augustine, Florida. This plant appsars to approach B. punila.

Anothar spacias is raprasantad in far wastarn U. S. and northarn Hazico. It has velvety lyrata-pinnatifid leaves, not scabrous as In aastarn B. subacaulis. Hookar (1839) thus introdueas B. lyrata. In l88*», Gray nanas a variety (B. lyrata var. nacrophylla) basad on a plant Dr. Lannon collacted in southarn Arisona. Its basal leayas ara as nuch as a foot long and ara oranata to pinnatifid at tha basa. Tha rarlaty was alavatad to spaeific status by Marcus £. Jonas (1908) but has sines bssn ra turnsd by Tumar and Johnston (1958).

According to tha aost racant works on Barlandiara by

Turnar and Johnston (1956) and Dyar (1959)1 tha ganus contains four spacias (B. taxana. B. punila. B. subacaulis. and B. lyrata) and varieties (B. taxana var. betonicifolia and B. lyrata var. nacrophylla). Tha ganus has not baan monographed in its entirety. ECONOMIC IMPORTANCE

All tax* of Borlandiora aro weedy and prosont problems In lawns and along roadways. Plants of B. subacaulia porslst In lawns, oven if nowod regularly, thus deserving its coaaon nano,

Florida Dandelion (Baker, 1926). •

Only ono easo is known to no of attoapts to cultivate a aoabor of this genus. Dr. David Fairchild roports on an herbarium labol (B. aubacaulis spocimon froa Dado County,

Florida, Fairchild an, 28 May 1932, NT)* "Mrs. Fairchild has doaosticatod a fow plants in Qoraan Post soil in shady local­ ity and has planted sono soods to soo if oho can raiso it froa seed."

According to horbariua labels (Rio Mayo explorations of

Howard Scott Sentry, 2304, 30 Aug 193&t F» MO, GH), the roots of B. lyrata var. aacrophylla aro reported to bo sold in local drugstores in the vicinity of Coronilla, Mexico, as a decoc­

tion or infusion for stoaach troubles.

Castetter (1935) records Indians using B. lyrata»

"Noahityini ayaahityi aaskawi wawa, bustard aedicine. The

flowers are aixed with sausage by the Acoaa and Laguna for

the purpose of seasoning."

16 ARTIFICIAL ESI TO BEBLAMDIEBA TAXA

Diao oorollaa and raina on under aurfaoea of raja rad to aaroon ; plaata with paraiating baaal roaette, deeuabent laafj branehaai paduaolaa aeabroua to aubacabroua; laaraa ralraty abore, uaually at laaat aoaa lyrata- pinnatifid* Taxaa Panhandla and Paooa Rirer wa at ward ...... (A) B. lyrata

a) Bladaa (except aoaatiaaa upperaoat and/or lowaraoat) Irregularly lyrata-pinnatifid •..... (B) var* lyrata

Bladaa irragularly toothad or only alightly pinnatifid at baae, aa nuch aa a foot long} intargradaa to (B)«*«(C) var. nacrophyll<

Disc oorollaa yallow or rad to aaroon and raina on undaraurfacaa of raya green; plaata with or without baaal roaattaa« daouabant to araot branehaai plaata not with both aoabroua to aub- acabroua padunelaa and valraty uppar laaf aur- faoaai laaf ahapaa rariabla ......

Diae oorollaa yallow; raina on undar aurfacaa of raya graani laaraa (axoapt upperaoat and/or lowaraoat) alternately ainuate-pinnatifidt laaraa and padunelaa aoabroua to aubaoabroua; uaually one to few flowera froa a aoapoaa roaottet Florida...... (D) B« aubacaulia

Diao oorollaa rad to aaroon, ray raina beneath green; ataaa araot, l e a f y ...... 3

Laaraa unaranly diatributad along ataa, oluatarad below; aoaa laaraa uaually lyrata-pinnatifid, uppar aurfaeea finely hirauta; pedunclea with long fine haira uaually nixed with atout abort onea, intargradaa to (D) and (0) •••••••••••••••• (B) B. x huailia

Laaraa aranly diatributad along ataa; not lyrata- pinnatifid (or if ao, than laaraa ralraty); intargradaa to (B) ...... k

17 18

Padunelaa with Ions fiat hairs, netted; bladaa velvety, uaually ovate, about aa broad aa long; S. C. to .... (F) B. punila

k Peduncles with atoutar araet, spreading or ourliag hairat bladaa final? hirauta to subseabroua above, ovate to triaagulata, laaa broad than l o n g ...... 5

5 Bladaa hirauta to subseabroua, triangulate, about two-thirds as long aa broad or least peduncles densely long hirauta ...... (Q) B. texana

5 Bladaa finely to coarsely hirauta, one-half to one and one-quarter tinea aa broad aa longt peduncles with spreading, curling to araot hirauta hairst intargradaa to (F) and (G)t east-central Texas, La...... (H) B. x betonicifolia DESCRIPTION AND DISCUSSION OF TAXA

(A) Barlandiara lyrata Benth* PI* Hartw* 17* 1839*

Barlandiara incisa T. & 0* A FI. of N.A. Vol. II, Part lit EET. 1842.

Perennial barbs with decuabent branchad leafy staas up to thraa faat high, rising froa a basal rosatta or a cluatar of ahortanad leafy branehas* Leaves velvety, lowar onas long-pat- iolad* Padunclas long, scabrous to subscabrous. Haads 1*3-1.? on wida. Says daap yallow to yallow-oranga; 10-14 aa long, 5*5-

8*0 aa wida; with rad to aaroon rains banaath (soaa rays with antira lowar surfacas rad to aaroon)* Achanas snail, alongata,

4.5-6.0 an long, 2.7-3*7 aa wida, quick-aaturing* Dry rocky liaastona soils, roadsidas*

Typei MEXICO. "Aguas Caliantas" (city or state of Aguas- caliantas), Hartwag 120, 1839* (Holotypa K, photograph MSClt isotypas NT1 QHI). Figure 2.

(B) Barlandiara lyrata rar• lyrata

Blades, except soaatiaas the upperaoat and/or lowaraoat, lyrata-pinnatifid, the terainal lobe cranata to irregularly incised* The nuaber of lobes par laaf varies widely between and within populations* The typical variety intargradaa to B. lyrata var* aacronhylla.

19 Specimens examined: MEXICO. Aguaacalientes: Aguas Cali- entes, Hartweg 120, 1839* NY Sc OH (isotypes). : Haci­ enda Mariposa in open brush land near Puerto Santa Anna, Muzquiz,

F. Lyle Wynd & C.H. Mueller 272, 24 Je 1936, OH,NY,MO{ El

Berrendo near Muzquiz at 4000', Stephan S. White 1873, 13-16

Ju 1939* GH; Muzquiz Palm Canyon, E.Q. Marsh, Jr. 993* 19 Sept

1936, F, OH; Muzquiz, E.Q. Marsh, Jr. 1146, Apr 1938, F,QH;

Sierra de Santa Bosa s of Muzquiz, E.Q. Marsh, Jr. 1534, 27 Ju

1938, QH,F; Sierra de Hechiceros in vicinity of Rancho El Tule near the Chihuahuan boundary in silty flat below old springs,

I.M. Johnston & C.H. Mueller 1321, 17-21 Sept 1940, QH; Canyon del Indio Felipe, ca. lat 28® 33' in a deep wooded canyon with running water in igneous Sierra Hechiceros near Chihuahuan boundary along banks of stream, Bobert M. Stewart 121, 27-29

Sept 1940, QH; On sandy flat in vicinity of Rancho El Tule ca.

24 km due n of Castillon and near Chihuahuan boundary, B.M.

Stewart 496, 13 Je 1941, QH (3 sheets); Qrassy flat near mouth of Cafton de San Enrique on e side of Sierra de la Encantado 3 km w of Bancho Buena Vista, B.M. Stewart 1364, 4 Sept 1941,

QH. : Meoqui, Harde Le Sueur 48, 6 Aug 1936, MO,PH,

QH; Chihuahua, Harde Le Sueur 48, 3 Sept 1933* F; Sandy hill­ side at broad valley 4 km s of Rancho de Encinillas, R.M.

Stewart 720, 7 Ju 1941, QH; Twelve km s of Escobillas on silty flat, B.M. Stewart 2337* 22 Sept 1942, QH; Bachimba Canyon,

C.Q. Pringle 844, 2 Apr 1886, NY,F,NY; Five mi e of Jimenez, 21

Figure 2. Berlandlera lyrata Benth. var. lyrata. Copy of holotype photo­ graph(TS5T, courtesy of J.H. Beaman.

Stephan S. White 2125* 31 Ju 1939* QH; Vicinity of Chihuahua at ca. 1300 m, £. Palmer 98, 8-27 Apr 1908, NY,GH; Dry plain

se of Chihuahua, Dr. Gregg (?) 536, 23-28 Apr 19^7, MO,NY;

Gallejo Springs in Northern Chihuahua s of Paso, A. WislizenuB

129, 21 Aug 18A6, QH,MO; Colonia Juarez on rocky slopes of

Sierra Madre Occidentale, at 1500-1700 m, Francis V/. Pennell

19057* 21 Sept 193^* NY,PH; El Paso, Geo. Thurber 73*N Aug

1852, GH; St. Diego at 6000', C.V. Hartman 617* 20 Apr 1891,

GH; Four mi n of Tucabaya on grassy plains road from Mestenas n« to Ojinagfc at 4700', I.M. Johnston 7966, 25 Sept 1938, QH.

Durangot City linita of along roadaida on opan sandy gravel, R.M. King 3730, 16 Aug I960, NT ("Chroaosoae nunbar dataraination froa aaiotio natarial aa n * 15* B.L. Turnar,

W.L. filliaon, R.M. King, I960"); Dry baaaltie rook naar

Carpintaro and 28-29 ka b of Durango at 2000-2100 a, F.W.

Pannall 18207* 25 Aug 193*», NT,PH; Durango City, £.W. Nalaon

4582, 1 Aug 1898, QH{ Naar Ignacio Allanda at 6300', Forraat

Shreve 9133* 24 Aug 1939* GH; Tapahuanaa at 6000', Gao. L.

Fiaher 44233, 28 Ju 19****, NT,MO,GH,SMU; Tapahuanaa, £. Painer

17, 25 Mar - 16 Apr 1906, NI,QH,M0,Fi In bottoa lands of City of Durango and vicinity, £• Painer 15*** Je 1896, QH,GH, and

Apr-Nov 1896, MO*F,MO,MO,MO,NT; Fiva ai aw of Quadalupa Victoria naar highway aaid Qaaaa graaaland with Opuntia in ahallow coarsa browning clay loan over calicha at ± 2000 a* Howard

Scott Qantry 8427, 23 Sept 1948* QH. ; Batwaan Huejuq- uilla and Maaquitic, J.N. Rosa 2573* 25 Aug 1897, NT,NT*M0.

San Lula Potosi; Minaa da San Rafaal Buana Vista, C.A. Purpus

5145* Ju 1911* F,GH,M0,NT;'Chareaa, C.L. Lundall 5096, Ju-Aug

1934, NT. Sonorai San Pedro, C.V. Hartaan 832, 14 Sept 1890,

GH; Batwaan San Pedro and Frontaras, C.V. Hartaan 930, 20-24

Sapt 1890, GH; Thraa ai a of Agua Priata along road to Colonia

Moraloa, Rio da Baviape at 3900', Stephan S. White 3821, 7 Aug

1941, GH; Along roadaida batwaan Sta Cruz and £1 Paso, and Chihuahua, A. Schott 42013, 8 Oct 1858, F. ;

Zacatecas, C.A. Purpua 462, Aug 1903, MO. . ARIZONA: Cochise: Three mi se Dos

Cabezos, Bassett Maguire 11155* 3 Hay 1935* HI; Four mi ns

Dragoon on w slopes with gravelly soilt Bassett Maguire lll82t

6 May 1935* NY j Sandy wash 5 oi s of Hilltop in Chiraoahua

Mts., Bassett Maguire 11056, 1 May 1935* PH,NY,SMU; Along

wash bed 3 ni n of Paradise, Bassett Maguire 11107, 2 May 1935*

NY,GH,M0, and 11107a, M0,GH,NY; Barren spur of mts at Stark

Ranch n of mouth of Pinery Canyon 5200-5500' in ChiracahuaB,

Witmer Stone 923* 30 Ju 1919* PH; Desert between the Chiracahuas

and the Southern Pacific RR 4 mi s of Dos Cabezos, Witmer Stone

464, 2 Ju 1919* PH| Seven mi s of Coohise at 4200', Forrest

Shreve 4454, 26 Sept 1914, MO; Near Douglas at 4000', Forrest

Shreve sn, 30 Apr 1933* F; Fort Bowie in Chiracahuas, Lemmon

Herbarium of Oakland, Cal. sn, Aug l88l, F,F; Ft. Bowie, Lemon

an, Aug l88l, F; Chiracahuas, J.W. Tourney an, 26 Ju 1894, NY,

QH; Chiracahua Mts., M.E. Jones 23617, 22 Sept 1931* MO; Qraasy

bottoms and dry ponds at Douglas, Leslie N. Goodding 2258, 22

May 1907* OH,MO; Ten mi n of Douglas, O.J. Goodman & C.L.

Hitchcock 1231, 19 Je 1930, MO; In grasslands 6 mi nw of

Chricahua at 5000', F.W. Gould & L.M. Pultz 3167, 18 May 1945,

NY; Roadside gravel 12 mi w of Douglas, K.M. & M.C. Wiegand

2398* 13 Apr 1935* GH. Cochlse-Pima: Mesas around Mustang Mts.,

C.G. Pringle sn, 27 Je 1884, F,MO,NY,F,NY,F,F. Pima: Foothills

of Mt. Rincon near Sienega Tucson, Lemmon 120, GH. Santa Cruz:

Two mi n of Vaughn, J.F. Arnold sn, 27 May 1938, GH; West of

Sonoita on road to Fairbank at 4700', Mrs. F.M. Stone 36, 22 Apr 1934, NT; Sonoita to Elgin at 4850', R.H. Peebles &

M. J. Fulton 11478, 5 May 1935, F. : Baca: Sand canyon 25 *1 a of Pritchatt naar Okla. lina at 4100*, C.L.

Portar 4264, 11 Ju 194?* QH; Sandy soil 8 il i of Caapo at

3900', C.W.T. Panland & J.B. Hartwell 5285, 31 May 1946, MT.

KANSAS: Morton: Stony hilla, A.S. Hitchcock 253, 6 Aug 1895*

NT,MO. NEW MEXICO: Barnallilo: Albuquarqua, C.L. Harriok an,

6 Sapt 1894, NT; Albuquarqua. M.S. Jonas 4898, 4 Sept 1884, F,

PH,GH; Rocky plains naar Albuquarqua, E.J. Palaer 31218, 21

Ja 1926, PH} On aesa ca. 2 ai a of Albuquarqua at 5000',

Alfred L. Kaaaarar 51• 1915* PH,M0,MT; Vicinity of Albuquerque at 5000-7000', D. Daaaraa & £. J. Palaer 388, 20 Ja 1926,

UARK, Chaves: Arroyo Ranch near Roswell, David Griffiths

4280, 4-9 May 1903* MO; Twenty ai s of Roswell on high plains,

F.S. Earle 28?, 9 Aug 1900, MO, and with E.S. Earle, MO,OS,NT.

Chaves-Eddy; Roadside Roswell to Carlsbad, Avan Nelson 11345*

22 May 1931* MO,NY,OH. Donna Ana: Organ Mts. at 4500', E.O.

Wooton sn, 17 Aug 1895* NT; Organ Mts. at 4700', E.O. Wooton

143, 15 Ju 1897* NT,MO. Eddy: In rocky sandy clay in valley of Black R. 22 ai aw of Carlsbad on Hwy 62, U.T. Waterfall

3747, 14 Aug 1942, GH; Forty-five ai sw of Carlsbad in Dark

Canon along Rocky Arroyo Road at 5200-6400' in valley or botton of canon, Carol Grassl 30, 8 Ju 1930, F; Washington Ranch,

B.E. McKechnie 555* 1 Sept 1959* MO, About 8 ai above the aouth of Dark Canyon of Guadelupe Mts., Hans Wilkens 1567, 22 Apr

1932, PH; Top of low ridge near Three Forks of Rooky Arroyo in 25 Guadelupe Hans Wilkens 1710, 30 Apr 1932, PH; Pasturs of Carson Shssp Ranger Station in Guadelupe Mts., Hans Wilksns

2269S, 4 Ju 1932, PH; Foothills of Quadslups Mts., Ladislaus

Cutak A Arthur Christ 47, 18 Sspt 1935• NO. Grant* Naar

Silver Citj, E.L* Qrssns 12613, 11 May 1880, NT,MO, and sn, QH;

Sandy loan of w slopss at tank #3 at Had Bock Caap, Bassstt

Maguire 11414a, 16 May 1935* NT; Sandy rocky soil on nvr slopss

1 ai sw of Oils Caap along road, Bassstt Magulrs & B.L. Richards,

Jr. & Thao. Moeller 11997, 14 Ja 1935* NT,PH; Sandy flats 12 ai n of Silver City, Bassstt Maguirs, B.L. Richards, Jr., & Thso.

Moeller 11597, 22 May 1935, NT; Oog Spring, E.A. Mearns 136,

27 May 1892, *NT; Hills at Silver City, Alice Eastwood 8364,

28 Apr 1919, GH; Silver City to Tyrone at 5500', Mrs. F.M.

Stone 16, 17 Apr 1934, NT; Gila R. near Silver City, M.E.

Jones 25937, 4 May 1930, MO; At Mangas Springs 18 ai nw of

Silver City at 4770', O.B. Metcalfe 4?, 6 May 1903, GH, NT,

MO; Gravelly banks at Mangas Springs, Henry H. Rusby 243,

May 1880, MO,NT,F. Quadslups * Cuervo, J.T. Rothrock 98, Js

1874, NT,F,GH; Halfway Anton Chico to Sta. Rosa at 1600 a,

Bro. G. Arsens & Bro. A. Benedict 16687, 2 Aug 1926, F,PH.

Lincoln* Rocky canyon wall at 7300* 15 ai w of Lincoln,

C.L. Hitchcock, R.V. Rethke & R. van Raadshooven 4267, 26

Ju 1933, GH; Near Gray at 6000', Miss Josephine Skehan 33,

15 Ju 1898, NT,MO,MO,NT,F,OS,GH; Gray at 6000-6500', F.S. &

Esther Earle sn, 22 Ju 1900, NT. Mora* Between Wagon Mound and

Las Vegas at 6500', Susan Delano McKelvey 4904, 4 Je 1934, QH. Otero* Saeraaento Mte. in Juniper belt, F.S. Earle 555,

8-19 May 1902, NT| High Holla and vicinity n hillside at 6000',

H.L. Viereck sn, 21-28 May 1902, PH* Quay* Qrassy plains at

San Jon site 10 ni s of San Jon, Kirk Bryan sn, 26 Aug 19^2,

OH* Mesa Rica near Paloaas, Leslie Hubricht, Cora Shoop &

Dorothy B. Heines Bl*t60, 14 Ju 19??, MO. San Miguel: Las Vegas

Aqua Zarca at 2000 a, Arsens 18601, 1? May 1927, 7* Langinjuela

Las Vegas at 2000 a, Bro* 0. Arsens 18966, 7 Apr 1927, NY| Las

Vegas Hot Springs at 6500', W.C. Sturgis sn, 12 May 1902, NT,

OH| Las Vegas, O.M. Kellogg sn, Spring 1890, QH{ San Miguel Co.,

T.S. Brandegee 12058, 1879, MO. Santa 7e: On hills 10 ai n of

Santa 7e at 5800*, A.A. & E. Gertrude Heller 3662, 5 Ju 1897,

GH,NT,NT,NI,M0. Sierra* In and around the s end of Black

Hangs Kingston at ca* 6600* in dry gravelly soil, O.B. Metcalfe

932, 25 May 1904, M0,NT,GH,7. Socorro* Socorro, F.H. Snow sn,

Aug l88l, MO. * Beaver * Prairie 10 ai s of Liberal,

Kansas, G.W. Stevens 381, 10 May 1913, GH,M0. Ciaarron* Valley near butte near Kenton, G.W. Stevens 501, Ik May 1913, MO,NY,

GH* Black Mesa at Kenton, D. Deaaree 133^1, 28 Ju 1938, NT,QH|

Near base of Black Mesa near Kenton, G.W. Stevens ^78, Ik May

1913, GH* Rocky soil 11 ai n of Boise City, Robert Stratton

^52,^53, 22 Aug 1927, MO,M0,SM1I; Dry hills 5 ai e of Kenton,

George J. Goodaan ZkOkt 21 Apr 1935, NY,MO. On plains 15 ai n of Boise City, George J. Goodaan 6007, ^ Sept 195^, GH{ On top of the Black Mesa in black volcanic rock and soil derived froa it on dry zeric plateau 2 ai n of Kenton, M. Hopkins A M. Van Valkenburgh 5795# 17 Apr 1941# NT* Harmon*Jackson:

Eldorado# D. Demaree 12203# 18 Apr 1956# HO. TEXASt Austin:

Industry# E.H. Boyd sn# no dats, F. Ballsy; Eight mi nw of

Hulsshos In silt at roadside, Lloyd H. Shinners 8341# 28 Je

1945# SHU,OH,MO. Baylor: Sandly plains# J. Beverchon 480,

Sept 1879# SHU,HO, and 20# NT. Brewster: Ungrazed grassland between BB and Hwy in igneous area 3 mi ne of Alpine# Bogers

McVaugh A A.M. Harvill, Jr. 7878, 9 Apr 1947# SMU; In grassland on rocky Igneous slopes at Sul Boss Hill at Alpine, Beginald

Bose-Innes & Bruneele Moon 1249# 19 Je 1941, F,GH; Vacant lot with rocks (limestone) soil in Marathon, L.H. Shinners 8719#

2 Aug 1946# SMU; Agua Fria Mts., B.C. Tharp 3820# Aug 1925#

PH; At Alpine at 4450*# Geo. L. Fisher sn, 24 Aug 1932# NO;

Bocky hillside at Alpine at 4600*# T.L. Steiger 991# Aug 1952#

NT; Bocky slope 8 mi e of Alpine at ca. 4500', F.L. Steiger

1956, Apr 1952, NT; Chisos Mts., C.H. Mueller 8219# 13 Ju 1931#

M0,GH,KT; Dry rocky plains near Alpine# E.J. Palmer 30581,

7 Oct 1926, MO; Stony hillside college grounds at Alpine, K.M.

Sc M.C. Wiegand 2397# 7 Apr 1935# OH; Pulliam Bluff in Chisos

Mts., no collector cited 745# 1 J* 1937# GH; At Alpine, Barton

H. Warnock 21773# 7 Jc 1940, NT; Between Alpine and Fort Davis,

Susan D. McKelvey 1951# 22 Apr 1931# GH. Crosby: Walker Tank near Blanco B., C.O* Erlanson 1088, 21 Je 1925# SMU; Three mi e of Crosbyton on gravelly slopes w side of ,

L.H. Shinners 8368, 28 Je 1945# SMU. Culberson: Canon Guade­ loupe, V. Hanard sn, Oct l88l, F. Along gully running through Larrea dlvericata near He Vey Tourist Courts in Van Horn, L.

Waterfall *+135, 20 Aug 19*+2, NT,GH,MO; In silt of vacant lot in

Van Horn, L.H* Shinners 8895, 11 Aug 19*+6, SMU; Sands in Van

Horn, Reverchon sn, 10 Je 1903, SMU. Dallam; Texline, David

Griffiths 5635• 26-28 Aug 1903, MO. Dawson-Lvnn: Plains between

Lamesa and Tahoka, J.K. Small & Edgar T. Wherry 121 *+0, 29 Apr

1923, SMU,NT. Deaf Smith: East side of Hereford on gravelly bluffs in park on Tierra Blanca Creek, L.H. Shinners 8320, 27

Je 19*+5, SMU; One mi e of Glen Rio, N* Mex., by roadside, V.L.

Cory 50385* 19 Oct 19*+5, SMU. El Paso; El Paso, G.R. Vasey sn, l88l, F. Hale: Two mi ne of Hale Center along RR right-of-way,

L.H* Shinners 18640, 28 Apr 195*+, SMU. Hardeman: Eight mi nw of

Quanah in pasture land, V.L. Cory 50123, 13 Oct 19*+5, SMU,NT.

Hartley: Fifteen mi se of Dalhart in tight hard "B" horizon of scraped and regraded roadside, L.H. Shinners 8170, 2*+ Je 19*+5,

SMU. Haskell: Along fencerow 8 .*t mi s of Haskell, L.H. Shinners

23737, SMU. Hudspeth: In arroyo on SP RR b mi w of Sierra

Blanca, Roxana S. Ferris & Carl D. Duncan 2*+79, k Ju 1921, MO,

NT; Igneous slopes of Eagle Mts. e of Eagle Peak, U.T. Waterfall

7790, 16 Aug 19*+7« GH; One and three-quarters mi nw of Allamore,

V.L. Cory 53009, 15 May 19*+6, SMU, Hutchinson: Three mi n of

Stinnet on slope above draw in pasture land, V.L. Cory 50325,

18 Oct 19*+5, SMU. Jeff Davis: Goat Canyon in Davis Mts. at

5500*, L.C. Hinkley 2*tl, 31 Ju 1935, NT,NT,F, and near Mt.

Livermore, Je 1936, GH; on Verhalen clay 2 mi sw of Chispa,

U.T. Waterfall 4?23, 26 Je 19*+3, NY,SMU,GH,MO; Plain w of Ft. Ft. Davis, J.K. Small & Edgar T. Wherry 12071, 23 Apr 1925. NY;

Ft. Davis in Davis Mts., Roxana S. Ferris & Carl D. Duncan 2655,

9-12 Ju 1921, MO; On shoulders of hwy 11 % mi s of Toyahvale,

V.L. Cory 52051* 5 May 1946, SMU, Jones: Near Phantom Hill,

Sutton Hayes 4l4, 15 May 1858, NY,F. Kent: Hills s of Delaware

Mts. near Kent, M.S. Young sn, 4 Aug 1916, MO; Kent, M.S. Jones

26422, 11 Apr 1930, MO. Kimble: Sandy plains, J. Reverchon

480, May 1885* F. Knox: One mi n of Qoree, H. Sumanth 262, 14

Sept 1954, SMU. Lubbock: Near Lubbock, D. Demaree 74-60, 1950,

¥ MO; Lubbock, D. Demaree 7476, 11 Apr 1930, UARK,GH; Lubbock, D.

Demaree 7708, 26 May 1930, GH,NY,F,M0,M0,UARK,UARK,N0; Canyons and plains escarpement "" ("Cap-Rock") in short- grass prairie on eroded slopes and rocky gullies and bottom of

"Yellow House Canyon" (Canyon of the Mountain Fork of the

Brazos R) 15 mi ese of Lubbock 0.5 mi s of Acuff at 2700-5200',

H.H. Iltis, D.M. Moore & H. Barkley 738, 28 Apr 1954, UARK,F.

McCullouch: Highway 190, 5 mi s of Rochelle, V.L. Cory 58512,

28 Sept 1930, SMU. Martta: Stanton, H. Eggert sn, 14 Je 1900,

MO. Midland: Nine and three-tenths mi ne of Midland, V.L.

Cory 5553* 18 Apr 1930, GH.GH. Mitchell: Along RR n of

Colorado* H. Eggert sn, 9 Je 1900, MO; Along Hackberry Creek

22 mi saw of Colorado City sec 5 4 SP RR Block 17 on dry rock ledge, Richard W. Pohl 5178, 16 Aug 1945, SMU; In creek bottoms, aw K sec. 40 T & P RR Block 25* 7 mi ene of Colorado

City, Richard W. Pohl 4978, 17 Je 1945* SMU. Moore: Level grassland at Cactus Ordnance Works at Dumas at 5600', Beryl Si Holger Jaspersen, 10 Ja 1945, NO,SMU,NY,MO. Nolans Sweet­ water, B.C. Tharp an, 9 Ju 1941* SMU,MO; Dry plains sandy and gypsous soil naar Sweetwater, £.J. Palaar 33968, 13 May 1928,

NY,PH,MO| Dry opan ground of gypsous soil at Swaatwatar, £.J.

Palaer 1376?, 27 May 1918, MO. Pacoss Ft. Stockton on Coaan- cha Craak, Patrick Duffy sn, i860, NY; Along Hwy 3 ai a of Ft.

Stockton in liaastona soil, Barton H. Warnock 46162, 19 Apr

1946, UARK,MO; Liaey clayish gravel scrubland and naar draw

13 ai w of Ft. Stcokton, Frad A. Barkley 14T857* 2 Oct 1944,

MO. Potters Prairie at Aaarillo, J. fieverchon sn, 29 May 1902,

MO| Hillside 16 ai a of Aaarillo, Qaorga J. Goodaan 21f27, 22

Apr 1933* MOf Aaong rocks in canyon of Short Craak 3 ai sw of

Fritch naar Canadian R, Rogers McVaugh 7239* 23 Ja 1945, SMU.

Presidios In liaastona in Pinto Canyon on a side of Chinati

Mts. at 3200*, Jin Scuddy 16, 26 Apr 1932, SMU; Vicinity of

Marfa at 4688', D. Daaaraa A E.J. Palaar 230, 18 Ja 1926,

UARE; Race track at football field in Marfa at 4700', L.C.

Hinklay 2702, 1 Ja 1943* SMU; Twenty-two ai s of Marfa, Forrest

Shrove 8113* 2 Aug 1937* F; Along dry craak 17 ai n of Shaftar,

Hugh C. Cutler 1941, 1 Ja 1938, OH,MO; Marfa at 1420 a, W.W.

Eggleston 17329, 18 Sapt 1920, NY. Randalls ,

R.R. Innas A Brunelle Moon 1134, 10 Ja 1941, QH; Canyon, H.C.

Benke 3002, 23 Apr 1929* GH,F; Dry opan ground, plaina at

Canyon, E.J. Palaar 14080, 13 Ja 1918, MO; Canyon City, H.

Eggert sn, 14 Aug 1900, MO,MO. Randall-Arastronas Dry slopes in Palo Duro Canyon region, M.S. Young sn, 9 Sapt 1917, 31

MO. Reagan: Waste ground at tourist court at Big Lake, V.L*

Cory 53^6^, 2k Apr 19^7, SMU,NY. Reeves: Balmorhea, Aven &

Ruth A. Nelson 3010, 31 Mar 19^2, GH,MD; Balmorhea, M.S. Jones

29513* 5 Apr 1932, MO; Along hwy 6 ml e of Balmorhea In lime­

stone soil, Barton H. Warnock 99^6, 22 Apr 1951, SHU. Swisher:

In sand plains 7*3 ®i e of Tulia Rd near Tule Canyon, Eula

Whitehouse 9975, 18 Je 1 9^5, SMU. Taylor: Camp Barkley In

grassland on rocky soil, W.L. Tolstead 6950, 8 Apr 19^3, NY,

MO,SMUi Sandy plains, J* Reverchon sn, Apr 1882, NY,F; Sandy

ground n of Abilene, H. Eggert sn, 7 Je 1900, MO; Camp Barkley,

W.L. Tolstead 7678, 1 Oct 19^3, GH. Tom Green: Dry hills in

calcareous soil at San Angelo, E.J. Palmer 11399, 20 Mar 1917,

MO; Sandy prairies at San Angelo, J. Reverchon sn, 19 May 1903,

MO; Tom Green Co., Mary Lovland (?) sn, 1883, MO; Tom Green

Co., Frank Tweedy sn, 1879, NY; Twelve me nw of San Angelo

on dry slopes and moist areas at base of slope with limestone

base, Charles Smith 101 and 133, 22 Je 19*tl, NY,NY. Ward:

Roadside 1 mi e of Monahans, H.R. Totten 8296, 16 Je 1930, NCU.

(C) Berlandiera lyrata Benth. var. macronhylla Gray. Syn. tfl. ft. Am. Vol. I, Part II: 2^3. 1884.

Berlandiera macrophylla (Gray) Jones. Contr. West. Bot. 12: kT. 1908.

Basal leaves measuring as much as a foot long, either

merely crenate or pinnatifid at the base.

Type: ARIZONA: Southern Arizona, J.G. Lemmon sn, l88l

(Holotype on two sheets, GH1). Annotation label mentions

Apache Pass, Arizona* See Figures 3 and U. Figure 3* Berlandiera lyrata var. Figure Berlandiera lyrata yar. macrophylla Gray. Holotype, aheet 1 (GH) macrophylla Gray. Holotype, aheet 2 f Scale: 0.28x. large specinena left (GH). Scale; 0.28x. This taxon la retained pending furthsr experimentation.

This entity ie segregated froa B. lyrata var* lyrata using size of parts and degree of lobing of the blades. The lobes on one leaf of this holotype outnuaber those on certain leaves of the holotype of the typical variety. With the large-leaved specimens are several saall unlobed-leaved plants of another oolleotion. Most taxonoaists have considered all as belonging to variety aacronhylla. This taxon is not considered in the remainder of this paper.

Specimens examined: MEXICO. Chihuahua: Dwarf fora froa vicinity of Madera at ca. 2250 a, £. Palaer 273* 2? May-

3 Je 1908t GH,M0,F,NTj Near Colonia Juarez in the Sierra

Madre, £.W. Nelson 6035* Je-Ju 1899* GHt £1 Ciaa, Harde Le

Sueur 1037* 29 Je 1936, GH,F* Dwarf fora froa open Mesa de

Oonselea above Arroyo Ancho, District of Guerrero, Tnes Mexia

26l8, 8 Je 1929* F,NT; In Sierra Madras near Colonia Garcia at 8000', C.H. Townsend & C.M. Barber 21, 23 Je 1899, MO,GH,F

NT; Guayanopa Canyon at 5000* in Sierra Madras, Marcus £.

Jones sn, 23 ^ept 1903* GH,NY* Pine plains at base of Sierra

Madras, C.G. Pringle 1311, 26 Sept 1887, GH,PH,F,NY,NY,{

Sierra Canelo, Bio Mayo JSxpl. of Howard Gentry 250*+, 30 Aug

1936, M0,G,GH. Nuevo Leon: Hacienda Pablillo, Galeana,

Mary Taylor 16, 3 Ju 1936, *F,*M0,NY| Sierra Madre Oriental

San Francicso Canyon ca. 15 ai aw of Pueblo Galeana at 7500-

8000*, C.H. & M.T. Mueller 336, l*t May 193^, *GH,GH*Fj

Shrubby meadow in lower canyon of Canon Denuncio, Rancho La Bolla, Municipio de Villa Santiago, C.H. Mueller 2025* 2? Je

1935, *GH,F; "Encinal," Fabillo se of Galeana in reddish soil among bushes at 2000-2100 a, F.W. Pennell 17003* 26-30 Je 1934,

•PHi On plain Arroyo Hondo* Kacianda San Jose de Raices, Munici­ pio de Galeana, C.H. Mueller 2318, 1 Aug 1933* MO; Hacienda

Cieneguillas on Cerro Ptosi on dry open stony slopes in oak chaparral at 8000', Municipio de Galeana, Mex. Biol. Expedition of U. of Illinois 989* 11 Aug 1938, GH,NY. : Three ai n of Micquihuana in pine forest, S.M. Labor, L.R. Stanford St

L.A. Taylor 2409. 12 Ju 1949, SMU,NY(MQ; Atop broad flat at. 22 km se of Miquihuana at 2300 m at 23* 40' N, 99* 41' W, L.R.

Stanford & K.L. Retherford Sc R.D. Northcroft 873, 12 Aug 1941,

NY,MO.

UNITED STATES. ARIZONA: Cochise: North Fork of Pinery

Canyon 2-4 mi above jet with South Fork at 6000-7000* in Chira­ cahuas, Witmer Stone 134, 24 May 1919* FH; Ft. Huachuca, T.E.

Wilcox sn, Ju 1893, OS, and dwarf form, Je 1892, NY. Santa Cruz:

Sonoita, R. Darrow sn, 22 May 1937, GH; Harslow Camp Patagonia

District, J.G. Lemmon 121, l88l, GH. Apache Pass, J.G. Lemmon sn, l88l, GH (holotype). Santa Crus - Pima: Foothills of Santa

Rita Mts., C.G. Pringle 1, 23 Je 1882, GH. NEW MEXICO: Eddy:

In, canyons in Guadelupe Mts. at 4000', A.L. Hershey 3031* 23 May

1944, *GH. OKLAHOMA: Cimarron: In arroya running up the nw slopes of Black Mesa 4 mi n of Kenton, U.T. Waterfall 7433, 9 Ju

1947, *SMU, TEXAS: Brewster: Rare near top of Baldy Peak of

Glass Mts. on limestone ridges, Barton H. 'Warnock 233* 13 Ju 1940, 35

•QH. Culberson: Among boulders in dry stream bed of Upper

McKittrick Canyon of Quadelupe Mts. at 6000', Fred 0. & Lillian

E. Meyer 2176, 21 Je 19^7* MO,NY; South slope of Pine Top Mt. of

Guadelupes on cliff faces at ca. 7000', Qrady L. Webster **576,

28-29 Ju 1952, SMU; In sand in bottom of S. McKittrick Canyon at

5^00', L.C. Hinkley 12 Ju 19*+&t SMU; Guadelupe Mts. on open rocky ridge above McKittrick Canyon at 2500 m, J.A. Moore & J.A.

Steyermark 3630, 25 Ju 1931» PH,MO. Pecos: Dwarf forms from mountain side 25 mi s of Ft. Stockton, Herbert C. Hanson 667, 18

Apr 1919, MO,NY.

(D) Berlandiera subacaulia Nutt. Trans. Am. Phil. Soc. Vol. Ill n.s.: 543. " m i .

Silphium subacaule Nutt. Am. Journ. Sci. V: 301. 1822. Silphium nuttallianum Torrey. Ann. Lyc. NY. II: 216.

Perennial herbs with very short stems up to A—5 inches high.

Long leafless or nearly leafless scapes typically bearing a single head 1.6-1.9 cm wide. Leaves are petiolate to Bessile, scabrous to subseabroua, alternately sinuate-pinnatifid, some­ times lyrate. Hays 10.0-12.6 mm long, 5.0-7.8 mm wide, lemon yellow with veins beneath green. Disc corollas yellow. Achenes

5 .2-7.0 mm long, 3*0-3.8 mm wide. Plants of sandy lawns and road­ sides. Common name is Florida Dandelion.

Type: Florida: Collected in east Florida by Mr. Ware

(Holotype BM; Photograph MSCI; possible isotype PHI). See

Figure 5. Specimens examinedt ALABAMA. Locality uncertain: Alabama,

S.B. Buckley sn, no date, MO. FLORIDA, Alachua: Alachua Co., A.S. Hitchcock. ant Je-Ju 1898, F,M0. Brevard: Indian R, E.

Palmer 268, 1874, MO; Scrub-oak land, Okechobee region, A.

Fredholm 5736, 25 Mar 1903, OH} Dry roadside at Rockledge, E.B.

Bartram an, 8-18 Mar 1916, PH; Eau Oillie at Indian R, A.H.

Curtiss 5699, 11 Ju 1896, MO,SMU, GH, NY, Broward: Hammocks from

Fort Lauderdale to Miami, J.K. and O.K. Small 2nd J.J. Carter sn

Feb 1911, NI. Citrus: Two mi s of Rte 4l at Dunnellon, F.H.

Sargent 6064, 4 May 1952, SMU. Columbia: Lake City, P.H.

Rolfs 307, 1891, MOtF. Dade: Plnelands about Humbugus Prairie,

J.K. Small & C.A. Mosier 5547, 28 Feb 1915, NCU,NY,OH} Along hwy in Redland region west of Perrine on Peters Rd, D. Fairchild

sn, Je 1932, NY} Near Camp Longview, J.K. Small & J.J. Carter

993, 9*12 Nov 1903, NY,PH{ Pinelands near Long Prairie, N.L.

Britton 185, 24 Mar 1904, F,NY; In plnelands near Homestead Rd

between Cutler and Longview Camp, J.K. Small & J.J. Carter sn,

9-12 Nov 1903, F, and 1460, NY} Between RR and river at Miami,

A.A. Eaton 141, 8 Nov 1903, *F} Miami, A.P. Oarber sn, May 1877

NY} Sandy cutover lands at Miami, Delzie Demaree 10246, 18 Feb

1933, MOf Buena Vista, H.N. Moldenke 485, 21 Jan 1930, NY,MO;

Plnelands s of Miami R., J.K. & O.K. Small sn, 26 Nov-20 Dec

1913, MO, and 4764, NY{ Plnelands w of Miami, J.K. and O.K.

Small and J.J. Carter 3247, Feb 1911, NY, and 7, PH. Gilchrist:

RR right-of-way at Wilcox, R.K. Godfrey 5543, 3 Mar 1957, OH,

NY. Hardee: Sandhills w of Avon Park (then De Soto Co.),

J.K. Small & J.B. Winkeler 9086, 1 May 1919, *NY. Highlands:

Along Rte 8 s of Sebring, J.A. Steyermark 63310, 15 Mar 1946, 37

N r k i

Figure 5* Berlandiera eubacaulis Nutt. Copy of photograph (MSC) of holotype (upper left specimen), courtesy of J.H. Beaman.

F; Be Soto City, L.J. Brass 15218, 25 May 19^5* QH. Hills­ borough : Tampa, N.L. and E.O. Britton and J.A. Shafer 7*+«

7-8 Mar 1903* NY; Tampa, A.M. Ferguson sn, Je-Aug 1898, MO;

Sand barrens about Tampa, J.R, Churchill sn, 11 Apr 1923* OH,

MO,F. Lake: Sandy woods n of Mineola, H.N. Moldenke 1971

5 Bee 1929t NY,*MO; Sandy woods at Mount Bora, F.W. Hunnewell

17929* 18 Feb 1 9^1 OH; Bustis, L.M. Underwood sn, Jan-Apr,

1891, NY; Eustis, P.H. Rolfs 385. Ju 189?, MO,F; Vicinity of Eustia, A.S. Hitchcock an, Je-Ju 1894, M0,F,F; Vicinity of

Euatis, Oeo. V. Naeh 808, 16-31 May 1894, NY,OH,MO, and 34,

12-31 Mar 1894, PH,GH,MO,NY, and 1920, 28 May-13 J« 1895,NY,

NCU. Lae: Flatwooda at (Ft.) Meyera, A,S. Hitchcock 134, Ju-

Aug, 1900, MO,OH,NY,F. L e w ; One mile n of county (Citrua) line, O.K. Cooley, C.E. Wood, Jr. 8c K.A. Wilaon 5958, 23 Apr

1958, NY. Manatee; Open aandy woods, J.T. Kothrock an, Mar

1887, F* "Braidentown," S,M. Tracy 6906, 16 May 1900, M0,F,NY.

Marion; Cedar landing on Ocklawaha K, E. Perot Walker 1601,

19 Mar 1940, PH; Ocala, J.R. Churchill an, 1 Apr 1897, MO;

Sandy open ground near Ocala, E.J. Palmer 38297, 31 Mar 1931,

QH,M0. Orange; Scrub-oak land, A. Fredholm 5406, 14 Ju 1902,

OH; Near Winter Park, A.M. Huger 16, 29 Mar 1900, MO; Roadaide

13 mi e of Winter Garden on State Rte 24, D.E. 8c M.S. Eylea

8047, 12 Apr 1942, OH; Forest City, F.L. Lewton sn, 20 Ju 1894,

NY; Orlando, P.H. Rolfs 399, 23 Feb 1894, F; Rock Springs, collector's name not legible sn, 18 Feb 1933, F. Osceola:

Kissimmee, A.W. Bitting 326, 3 Mar 1892, F. Palm Beach: Palm

Beach, M. Canby sn, Mar 1902, NY. Pasco: High pine land at

Jessamine, J.H. Barnhart 2507, 11 Mar 1898, NY; Pine wood clearings at Jessamine, Mrs. H.M. Platt 2748, 23 Feb 1897,

NY: Dry plnelands at St. Leo, Rev. H. O'Neill sn, 30 Apr 1927,

MO, and 28 Mar 1926, MO. Pinellas: Near St. Petersburg,

Mrs. C.C. Dean 4010, 16 Mar 1908, QH,F; St. Petersburg, Mary

H. Williams sn, 13 Feb 1926, PH; Clearwater, M.S. Bebb sn,

1894, F (unusually hairy specimens with some heads with ten rays); Clearr/ater, A. Brown sn, 26 Mar 1901, NY; Clearwater 39

Harbor, no collector cited an, Oct 1875* MO; Clearwater Harbor,

H.A. Pilsby an, Mar 1904, PH; Dunedin, 5.M. Tracy 6916, 14 Apr

1900, M0,GH,NY,F. Polk: Winter Haven, J.B. McFarlin an, 20 May

1931, NO; Fort Meade, Dr. & Mra* Q.M. Armatrong an, 11 Ju 1922,

MO; Sandy place, P.O. Schallert an, 1 May 1941, NY; Sandy aoil

at Mountain Lake, F.W. Hunnewell 9051« 10 Mar 1924, QH; Bartow,

Jno A. Allen, Mar 1890, QH. Putnam: Palatka, A. MacElwee an,

18 Apr 1897, PH; Creacent City, Dr. Q. Martin an, 4 Mar 1880, PH,

MO; McMeekin, A.L. Pennock an, Mar 1890, PH; Palatka, C.S.

Williamson sn, Apr, PH,NY; Palatka, Wm.M. Canby sn, Apr l869t NY,

NY,GH; Open pine barrens at '.Velaka, C. Mohr sn, Mar 1882, F;

Palatka, H.C. Benke 3846, 10 Mar 1924, F; Welaka, B.F. Leeds sn,

15 Mar 1892, F; Palatka, Geo. Smith sn, 17 Mar 1871, F, and Mar

1872, NY; Sandy soil in high pine lands at Johnson, Rev. J.W.

Barnhart 1331, 14 Mar 1896, NY, and 2172, 28 Apr-3 May 1897* NY.

Sarasota: South Creek at Osprey, B.H. Smith sn, Mar 1904, PH;

Near Osprey, Dr. Anne E. Perkins sn, 18 Dec 1943, GH. Seminole:

Collected in sand at Sanford, S. Rapp sn, Feb 1918, LSU; Alamonte

Springs, P.O. Schallert 6106, 1 Ju 1959. NCU and 25 Mar 1956,

SMU. Taylor: Scrub-oak-palmetto along Fla. 361 e of Adams

Beach, A.E. Radford 8319. 9 Apr 1955. NCU. Volusia: Open pine

woods 1 mi se of Lake Bereaford, E. Perot Walker 1729. 3 Apr 1940,

PH; Pierson, no collector cited 7029, 20 Mar 1926, NY; Waste

places near Lake Helen, Mrs. Stella Deam 1760, 26 Apr 1906, GH,

MO; Edge of N. Amelia Ave. of De Land, E.C. Prichard 732, 17 Apr

1954, GH; De Land, Janet R. Perkins sn, Mar-Apr 1918, GH; Pinland 40 at Orange City, S.C. Hood an, 19 Feb 19H« MO,MO; Dry pine barrens at Mosquito Lagoon, A.H. Curtiss 1393* J«» *NY,UARK,F,NY,GH,MO,

NY. Wakula: Dry oak barrens near Shell Point, Ludlow Oriscom

21606, 9 Apr 192*+! GHf M collefl oinetonun prone Shelloolnt.

F. Rugel sn, Ju 1843, NY. East Florida: East Fla., Ware, no date, PH (possible isotype). . Fulton - De Kalb (?):

Atlanta, S.B. Buckley sn, no date, MO (Buckley's specimens from

Oeorgia and Alabama are doubtfully cited). Without locality:

Georgia, LC (LeConte) sn, no date, NY.

(E) Berlandiera x humilis (Small) comb. nov. (* Berlandiera subalcaulis x pumila).

Berlandiera humilis Small. FI. SE U.S. 1246. 1903. n^otype" W r Tsotype FI)

Perennial herbs with erect stems 1-3 dm tall, commonly branching. Peduncles v/ith long fine hairs usually intermixed with short, stout ones. Leaves crov/ded toward root; blades lyrate-pinnatifid or unlobed, slightly hirsute above. Heads

1.6-2.2 cm wide. Rays 12-1? mm long, 5*5-7*8 mm wide, deep yellow to yellow-orange, veins beneath green. Disc corollas red to maroon. Achenes obovate, 5-6 mm long.

Type: FLORIDA: St. Augustine, Mary C. Reynolds sn, Je-Dec l8?5« Figures 6 and 7*

Putative backcross progeny much resemble parental types.

Intermediate specimens between B. pumila and B. x humilis bear either some lyrate-pinnatifid leaves or narrowed ones or both.

Otherwise the progeny are within the variation pattern of B. pumila. Most are found in a narrow coastal band from eastern Alabama to South Carolina across northern Florida. Putative B. subacaulis backcross progeny differ from parental type by bearing leaves with fewer lobes on more elongate, sprawling stems. Such plants are found scattered in northern Florida and along the east coast. The center of hybridization is apparently the Jackson- ville-St, Augustine area.

Specimens examinedi FLORIDA. Clay: Hibernia, Wm.M. Canby sn. Mar 1869, NY,MO,NY. Duval: Dry pine barrens near Jackson­ ville, A.H. Curtiss 1392, May l877( F^F.NY^Y.GH.MO; Pine barrens, J.D. Smith sn, 16 Mar 1882, F,QH; South Jacksonville,

J.R. Churchill sn, 7 Apr 1897, MO; Dry pine barrens near Jackson­ ville, A.H. Curtiss 7 May 1894, GH,F,NY; Near Jacksonville,

A.H. Curtiss 4477« H Apr 1893t MO,NY; Jacksonville, Nathaniel

Thayer Kidder sn, 7 Apr 1885, GH; Jacksonville and vicinity,

Thos. Hogg sn, 1873, NY; St. Nicolas, Rev. L.H. Lighthipe 78,

19 Mar 1896, NY; Jacksonville, H.C. Benke 3845, 1 Apr 1924, F.

Hillsborough: Tampa Bay, Dr. Burrows sn, no date, NY. Nassau:

St. Marys R., Read 3399* no date, MO. St. Johns: Dry pine barrens at St. Augustine, Mary C. Reynolds sn, Je-Dec 1873, NY,F

(holotype and isotype of jJ. humilis Small respectively). Suwanee

Hammock along Suwanee R. near Withlacoochee Bridge, J.K. Small,

J.B. DeV/inkeler & C.A. Hosier 11182, 21 Apr 1924, NY. Volusia:

Pine woods at Ormond, G.G. Kennedy sn, 29 Mar 1897, GH. SOUTH

CAROLINA. Barnwell: "Scrub" on sandy uplands near V/illiston,

S.J. Palmer 39865» 15Apr 1932, GH. Lexington: Dry sandy pine- lands s of Columbia, C.A. V/eatherby 6119, 27 Apr 1932, GH,NY. Figure 6. Berlandiera x humilis (Small) Figure 7* Berlandiera x hmailia (Small) comb. nov. Holotype (GH) of B. hunilia comb. nov. Isotype (lover left specimen, F) Small. Scale: 0.28x. of B. humilie Small. Scale: 0.2ox. ■p- ro 43

(F) Berlandiera jjumila (Michx.) Nutt. Trane. Am. Phil. Soc. VII n.s.: 342-343. l84l.

Silphium puailum Michx. FI. Am. Bor. 2: 146. 1803* l&olotype P, photograph MSC!) Silphium tomentoeun Purah. FI. Am. Sept. 2: 379* l8l4. Berlan&era tomentoaa (Pureh) Nutt. Trane. Am. Phil, doc. VII n.e.t 3*3. 1841. Berlandiera tomentoaa var. dealbata T. & 0. A FI. of fcl.A. Vol. II, Part II: ISIt 1&42. (Holotype NY! Paratypes NY! OH! 0H1) Berlandiera pumlla var. dealbata (T. & 0.) Trel. Ann. kep. fleol. Surv. of Ark. for 1888. IV: 193* 1891. Berlandiera dealbata (T. & 0.) Small. FI. SE U.S. HUT. T903. Polymnia caroliniana Poiret. Ency. Suppl. V: 303* 1804. (Holotype P,' photograph 01)

Herbaceous to suffrutescent perennials, erect, usually branching. Leaves well-distributed along stem; without a persis­

tent basal rosette. Central leaves ovate, usually three-fourths

to one and one-fourth times as broad as long, velvety, with

crenate margins, unlobed (except intergrades to B. x humilis), often long-petioled. Peduncles with dense, matted, fine long hairs. Heads 1.4-1.8 cm wide. Bays 12-20 mm long, 6-9 mm wide, deep yellow to yellow-orange, with green veins beneath.

Disc corollas red to maroon. Achenes 4.3-6.0 mm long, 3*0-4.0 mm wide. Along roadsides and grazed areas, woodland borders, and open woodlots.

Type: FLOBIDA; Collected in Florida, A. Micheaux 10, no date (Holotype P, photograph MSC1). Figure 8. Silphium

tomentosum holotype was collected in west Florida near Talla­ hassee. Type not seen. Polymnia carolinlna holotype was

collected by Fraser in Carolina (Figure 9). Type sheet has

two specimens of B. pumlla plus fragments of another plant, i f to

...... 4 f \. * Figure 8. Berlandiera pmaila Figure 9* Berlandiera pumila (Hiehr.) Nutt, Copy of holotype (Michx.) Nutt. Copy of photograph photograph (MSC), courtesy of (GH) of holotype (upper two speci­ J.H. Beaman. mens) of Polrmnia caroliniana Poiret courtesy of Gray Herbarium. probably Polymnia. In his description, Poiret excludes the fragment, and states that all material was of such poor con­ dition that a more accurate description was not possible. B. tomentosa var. dealbata was collected along the "Arkansa"

(Arkansas River) by T. Nuttall. Torrey and Qray's description cites paratypes collected in Texas by Drummond, 220. Figure

10 .

B. pumlla has hybridized with B. subacaulis in eastern

U.S. and with B. texana in the eastern Texas region to the extent that "typical" material is not common. Interbreeding in the west has produced a complicated system of hybrids dis­ cussed under B. x betoniclfolia.

The velvety texture of the leaves is responsible for its common name Woolly-leaf (Wells, 1932).

Specimens examined (East of Mississippi River): ALABAMA:

Baldwin: Qateswood, S.M. Tracy 8557* 6 May 1903* *M0,*F,NY,

*GH,*M0. Escambia: Escambia Exp. Forest, Section 27, S.M.

Qarnes 219* 13 Apr 19**9, NY. FLORIDA. Alachua: Two mi e of

Alachua, K.M. Wiegand & V/.E. Manning 333**, 30 Ju 1927, OH.

Calhoun: Eight ml n of Wewahitchka, E.S. Ford 3329* 19 Ju

195**» OH. Clay: Oreen Cove, Dr. G. Martin sn, 23 Apr 1880,

PH; Tacoi, E. Palmer 267, 16 Apr 18?**, *M0,*NY. Columbia:

At Lake City, Geo.V. Nash 2215, 11-19 Ju 1895, •0H,0S(M0,NYtF;

At Lake City, R.H. Rolfs 629, 23 M a y 1895, M0,F; At Lake City,

Geo.V. Nash 129**, 11-19 Ju 1895, *F; Dry sandy woods no of

High Springs, H.M. Moldenke 1101, 1 May 1930, NY,M0; Lake City, Figure 10. Berlandiera punila Figure 11. Berlandiera puaila (Michx.) Nutt. Holotype' (right), (Michx.) Nutt. Paratypee of B. toaent- two paratypee (left) of B. toaent* osa var. dealbata T. & G. (off). osa var. dealbata T. & gT (ite). Scale: 0.29x. Scale: 0.2Bx. A. S. Hitchcock sn, Je-Ju 1898, F{ Dry Black Jack ridge at Lake

City, R.H. Rolfs 368, F. Duvali Dry pine barrens near Jackson­ ville, A.H. Curtiss 1392, Kay 1877, *PH,NY,*GH,*M0; Pine barrens,

J.D. Smith sn, 16 Mar 1882, *F; South Jacksonville, J.R. Church­ ill sn, 7 Apr 1897, *F,*GH; Dry copses of Jacksonville, A.H.

Curtiss 6183, 4 Je 1898, *NCU; Jacksonville, A.H. Curtiss sn,

Apr, *0S; Dry pine barren, A. Fredholm 50l8, 2 Apr 1902, *GH;

One mi s of Tisonia, K.K. Wiegand & W.E. Manning 3333, 26 Ju

1927, *GH; In high pineland at Jacksonville, Hugh O'Neill 6003,

12 Oct 1929, *NY; Jacksonville, Alexander MacElwee Bn, 13 Apr

1897, *PHj Jacksonville, 1 S T . Y / . Calkins sn, 27 Dec 1886, *F and

24 Dec 1886, *F; Jacksonville, H.A. Lang sn, 9 Aug 1909, PH,PH;

Jacksonville, C.S. 7/illiamson sn, Aug 1894, *PH; Jacksonville,

A.S. Hitchcock sn, Ju-Aug 1900, *F. Escambia; Open field in

Pensacola, R. Brinker 6l, 21 Je 1941, MO; West shore of Bayou

Texar at Pensacola, R. Brinker 82, 23 Je 1941, MO. Gadsden;

One mi s of River Junction, E.S. Ford 3601, 23 Ju 1934, *GH;

Pine barrens near Havana, Ludlow Griscom 21604, 13 Apr 1934, *GK;

Quincy 12 mi s of Rte 267, F.H. Sargent 6163, 27 Apr 1932, *SMU;

Chatahoochee, A.H. Curtiss sn, May 1882, *GH. Gulf; Dry pine barrens at Wewahitchka, no collector cited 6l6a, 12 Aug 1896, GH,

NY; Wewahitchka, no collector cited 2447, Aug 1896, MO. Jeffer­ son; Jefferson Co., A.S. Hitchcock sn, Je-Ju 1898, F. Lafayette

Slash pine flatwoods 3 ml nw of Branford, M. Knott 4203, 17 Mar

1937, *SMU. Leon; Open sandy field 8 mi w of Tallahassee, R.

Krai 2435, 27 May 1956, •SMU,*NCU; At Bellair, Geo. V. Nash 2549, 48

3 Sept 1893, MO,NY,GH; Near Tallahassee, N.K. Berg ant Summer,

NY; Sandy open ground near Tallahassee, E.J. Palmer 33218, 9 Apr

1929* GH; Ad collea in pinetls inter Tallahassee jet St, Harks

(Leon-Wakulla Counties) Rugel sn, Mai 1843, NY,MO. Liberty:

Bristol, no collector cited sn, no date, HO; Open oak woods near

Torreya State Pk, R. Krai 1662, 14 Oct 1955. *GH. Nassau:

Fernandina, A. Ruth sn, Apr 1893, *MO,*OS. Polk: Polk Co., L.B.

Ohlinger sn, May 1894, *F. Santa Rosa: Oak-pine woods near US

90, 8 mi w of Milton, A.E. Radford 7915. 18 Apr 54, NCU. St.

Johns: Dry sandy woods near Moultrie, H.N. Moldenke 1?6, 27 Nov

1929. *NY,*MO; Dry pine barrens at St. Augustine, Mary C. Rey­ nolds sn, Je-Dec 1875. *NY,*F. Volusia: Sandy soil in flatwoods by Volusia Rd at Ormond, W.A. Purdie sn, 17 Apr 1903* *GH.

Walton: De Funiak Springs, 0. Clyde Fisher sn, Apr 1909. *NY“;

Dry pine barrens at De Funiak Springs, Chas Mohr sn, 11 May 1892,

*M0. Washington: Turkey-oak woodland 3 mi n of post offics at

Crystal Lake, E.S. Ford 3754, 29 Ju 1954, *GH. GEORGIA. Charl­ ton: Old field on oak ridge 4 mi w of Folkston, Jean S. Harper

812, 22 Mar 1936, PH; Pine barrens 4 mi w of Folkston, Arthur N.

Leeds 2563. 14 Apr 1936, *P; Trail ridge 1 mi w of Low Ford on oak ridge along canal, Molly & Francis Harper 322, Ju 1931. PH;

Gopher Ridge near Starling Branch, Jean S. Harper 417, 14 Apr

1932, PH; Open pine lands in sandy soil at Camp Cornelia, Wilbur

H. Duncan 7429. 27 Apr 1947. *M0. Dougherty: Vicinity of

Albany, C.L. Pollard & W.R. Maxon 519. 13 Aug 1900, NY; Along the Flint R. at Albany, J.K. Small sn, 24-28 Kay 1895. NY. Decatur: Bainbridge, Alphonso Wood ant no date, *NY; Flint R.

Swamp, J.K. Small an, 4 Je 1895* F,NY. : In sandy border

of woods ca* 1)5 ml se of Wellston, 83*36' Long. 32*3' Lat, Y/alter

Ainsworth 44507*3, 7 Hay 19^, PH* Sumter: Near Amerlcus, A.W.

Huger sn, Mar 1897, *NY; Dry soil se of Amerlcus, Roland M. Harper

1018, 8 Ju 1901, NY,F,*GH,*M0. Taylor: Open sandy woods several

mi n of Butler, Arthur Cronquist 5259* 23 Hay 1948, •PH,OH,NY,

SMU,M0. Toombs: Coarse sands dominated by turkey oaks ca. 2 mi

se of Lyons, R.L. Wilbur 3097, 26 Apr 1953, SMU. SOUTH CAROLINA.

Aiken: Aiken Co., Wm.M. Canby sn, May 1869, NY; Turkey oak-wire-

grass sandhills 4 mi sw of Aiken near US 1, A.S. Radford 6761,

23 Apr 1952, NCU; Highway n of Aiken, Wm.L. Hunt sn, 19 Apr 1939,

*NCU; Entrance to Aiken State Pk, T.R. Fisher 1750, 7 Aug 1958,

‘private collection; Sandy ground n of Graniteville, H. Eggert

sn, 4 Aug 1898, MO; SC 23, 3 mi e of New Ellenton, J.A. Duke 635,

16 May 1958, *NC; Five mi w of Aiken on US 1, J.A. Duke 1284, 19

Je 1958, NY; Aiken, H.Y/. Ravenel sn, no date, GH; Aiken, W.C.

Ferguson sn, 1 Apr 1922, *NY; Aiken, A. & W.V. Veldin sn, Apr

1882, *M0; Aiken, Hook sn, 24 Aug 1866, MO; Aiken, W.W. Eggleston

5057, 12-15 Sept 1909, MO,NY,NY,F,PH. Barnwell: "Scrub" on

sandy uplands near 'Williston, E.J. Palmer 39865, 15 Apr 1932,

*GH,M0. Chesterfield: Oak-hickory forest 0.9 mi s of Patrick near US 1, A.E. Radford 12423, 5 Je 1956, NCU; Pine forest 8 mi

e of Patrick, A.E. Radford 15804, 11 Aug 1956, NCU; Camp forest,

L.F. Swails, Jr. 37, 5 Je 1956, NCU. Calhoun: US 21 near the

Franklin Co. (?) line, Oliver M. Freeman 56175, 16 Hay 1956, •NCU; Sandhill 0.1 mi n of Calhoun-Lexington Co. line on US 21 wnw of St. Matthews, Harry S. Ahles & J.Q. Haesloop 25642, 19

May 1957, NCU. Darlington: Along SC 102, 6.5 ni b of Patrick,

J.A. Duke 1521, 18 Ju 195$, NCU; In sandhills 4 mi n of Harts- villa, W.C. Coker sn, 1 May 1910, •NC,*NC; Sandy soil along e side of Crowley*s Branch 1 mi e of Hartsville, B.E. Smith 6l8,

5 Je 1941, *NCU{ Sandhills n of Hartsville near Chesterfield Co. line, B.E. Smith 887, 20 Ju 1941, •NCU; Dry sandhill by Crowley’s

Spring in Hartsville, W.C. Coker 8290, 17 Je 1912, NCU; Society

Hill, no collector cited sn, no date, NY; Society Hill, Curtis sn, no date, QH; Society Hill, V/m.M. Canby sn, 5 Ju 1878, F,F,F.

Edgefield: Sandhill near US 25, 9 nti ssw of Trenton, A.E. Radford

22494, 12 May 1957, *NCU; Sandhills, A. Cuthbert 59, 24 Je 1901,

•NY. Kershaw: Sandhill 7*5 nii nnw of Bethune, A.E. Radford

25702, 4 Je 1957, NCU. Lexington: RR right-of-way by US 1, 5 mi w of West Columbia, V/.B. Fox & S.G. Boyce 4574, 22 Apr 1951,

QH; Roadside near US 178, Zh mi w of Pelion, A.E. Radford 20558,

5 Apr 1957, NCU; Sandhill near SC 602, 6 mi se of Lexington, A.E.

Radford 25525, 27 May 1937, NCU; Scrub oak and long leaf pine in sandhills 6 mi s of Columbia, R.K. Godfrey & R.M. Tyron, Jr. 1248,

7 Aug 1959, NY,GH. Orangeburg: Sandy oaks woods Zh mi sw of

Neeces on SC 4, H.E. Ahles & J.G. Haesloop 25280, 18 May 1957,

NCU,NCU. Richland: SC 12, 8 mi e of Ft. Bragg entrance, J.A.

Duke 572, 15 May 1938, NCU; Sandy open woods near Columbia, E.J.

Palmer 42405, 19 May 1954, GH,M0. Saluda: Sandhill near SC 592,

2 mi s of Ridge Spring, A.E. Radford 25156, 26 May 1957, *NCU. Specimens examined (West of Mississippi River): LOUISIANA.

Grant: Along BR bank 10 mi nw of Alexandria, Fred Corbin 195,

30 Aug 1952, SMU. TEXAS. Anderson: Along BR, H. Eggert sn, 8

Je l899t MO; In sandy upland nw of Montalba, C.L. & A.A. Lundell

1191^t H Sept 19^2, *SMU. Bastrop: In sandy soil w of Bastrop

State Pk, Eula Whitehouse 10620, 27 May 19^2, *SMU. Cass: Sandy open woods and open ground 7# n of Atlanta, iL.H. Shinners 7882,

13 Je 19^5• *SMU. Cherokee: Dry woods at Larissa, E.J. Palmer

7836, 3 Je 1915* M0,*NY. Colorado: Roadside clearing in post­ oak woods 1 mi sw of New Ulm on Rd 109 in deep sandy soil, n=15»

B.L. Turner A C.J. Marshall, 5^329, 10 Apr 195^, *SMU. :

In sand at Dallas, B.F. Bush 693* 10 May 1900, MO; Dallas, J.

Reverchon ^79* Ju 1876, *F, and V?9, 1880, *M0 , and sn, Sept

1876, *GH, and 4?9, Je 1885, *F, and sn, Ju 1879, *F, and sn, Je

1876, *NY; Dallas, G.W. Letterman sn, no date, *M0. Freestone:

Open woods in sand of hillside 15 mi s of Fairfield, L.H.

Shinners 7111, 9 Apr 19^5, SMU. Gonzales: Cottonwood Springs,

H.B. Parks 7692, 27 Oct 1933, OH. Grimes: Navasota, G.L. Fisher

3290, 11 Je 1932, *F. Harris: Hockley, W.F. Thurrow sn, 1890,

*F; Very dry open waste field near Lake Huston, rd to n shore development ca. 200 m e of US 39, 20 m s of road at 63' elev.,

Alfred Traverse 997, 7 Apr 1959, *M0; Sand w side of US 59 ca.

0.2 mi s of bridge over San Jacinto R., 23 mi n of Houston, A.

Traverse 165, 3 Ju 1956, *GH. Montgomery: Sandy open woods near Conroe, E.J. Palmer 33332, 15 Apr 1928, GH. Robertson:

In sandy soil between hwy and HR kYi mi n of Benchley, V.L. Cory 55019i 26 Mar 19**9* SMU. Smith: Roadside near Kernal's Lake,

Camp Fannin 8 mi ne of Tyler, H.E. Moore, Jr. 7*+6, 23 Apr 19*+*+»

QH; In open woods at Tyler State Pk, V.L. Cory 53391* 13 Apr

19*+7, SMU. Ugshur: A colony in sandy open woods *+.*+ mi w of

Rosewood, V.L. Cory 57696, 8 Aug 1950, SMU; Sands at Big Sandy,

J. Reverchon sn, 29 May, MO. Van 2andt: Qrand Saline, J.

Reverchon sn, 19 Oct, *M0; Sandy upland prairie-like open area

2 mi w of Qrand Saline, L.H. Shinners 7535* *+ May 19*+5» SMU.

Waller: Sandy prairie at Hempstead, Elihu Hall 32*+, 1 May 1872,

•MO,*QH,*F,*NY,*NY,*NY. Washington: At 350' alt. at Brenham,

Q.L. Fisher 132, 17 Ju 1930, *F. Wood: In pine land at Lake

Ellis 5 mi ne of Crow, C.L. & A.A. Lundell 11736, 2 Sept 19*+2,

•SMU. Uncertain locations: Arkansa, Nuttall sn, no date, *NY;

Texas, Drummond 220, no date, *NY,*GH,*GH (holotype and paratypes respectively of B. tomentoaa var. dealbata).

(Q) Berlandiera texana DC. Prodr. V: 517* 1836.

Berlandiera longifolia Nutt. Trans. Am. Phil. Soc. Vol. VII n.s.: 3^+2. l84l, (Holotype BM, photograph MSCI).

Suffrutescent herbs up to four feet high, often much branched in inflorescence. Peduncles short, densely long hirsute, bearing heads 1.8-2.? cm wide. Central cauline leaves short-petioled, about one-half to two-thirds as broad as long, wider leaves on plants found toward eastern portion of range; triangulate to ovate with serrate to partially doubly serrate margins; siffly hirsute to subscabrous above. Leaves well-distributed or crowded near summit; no persistent basal rosette. Rays 10.5-17.0 mm long, 6 .5-9*0 mm wide, deep yellow to yellow-orange, with green 53 veins beneath. Disc corollas red to maroon. Achenes large, ovate, 4.5-6.0 mm long, 3*0-4.8 mm wide. Plants commonly found at margins of wooded areas or in open or grazed woodlots, on hillsides or along lightly-shaded river banks; in sandy soils.

Type: TEXAS: Between the Trinity River and Bejar (San

Antonio), J.L. Berlandier 2044, Je 1828. Holotype not seen but is presumably in Museum National d'Histoire Naturelle (Paris)

(P); isotypes NY! MO! F!; topotype, J.L. Berlandier 1751 = 431,

Je 1828 GH!. His later collections (2474 * 1044) along the

Medina River in June, 1834, clearly belong to the same taxon.

Figure 12.

Central and western Oklahoma and northwestern Texas speci­ mens have very scabrous leaves and peduncles short hirsute to subscabrous. Missouri, Arkansas, and south-coastal Texas specimens have wider leaves with more obtuse species. The pubescence is less erect and is often matted on lower portion of stem. This latter group is consistent with holotype and description of Nuttall'a B. longifolla. Figure 13*

Specimens examined: ARKANSAS. Madison: Huntsville, D.M.

Moore 430201, 14 Aug 1943. UARK,UARK. Marion: Yellville, D.M.

Moore 330259* 11 1930, UARK. Polk: Fields, dry shaly banks with pine, post oak-hickory woods ca. 1# mi sse of Hatfield near

Six Mile Creek at 900-1000', Aileen McWilliam sn, 17 Oct 1954,

SMU. Sharp: Hardy P.O. at 700* in foothills, Delzie Demaree

26845, 20 Je 1948, SMU. LOUISIANA. Caddo: Highlands near

Shreveport, J. Gregg sn, 9 Sept 1847, MO,PH. Rapides: Pine woods at Rapides, illegible name of collector sn, Je, *M0. f

Figure 12. Berlandiera texana Figure 13. Berlandiera texana DC. DC. Isotype (NO). Scale: 0.30x Copy of holotype pkotograpk (kSC) of B. longifolia Nutt., courtesy of J.H. Beaaan. MISSOURI. Barrg: Cassville, B.F. Bush 302, 29 Sspt 1896, NY,

MO) Fislds at Cassvilla, K.K. Mackenzie an, 29 Sept 1896, MO;

Prairies at Purdy, B.F. Bush 3266, 17 Aug 1905t MO; Dry soil e of Monett along S.L. & S.F. RR, F.W, Pennell 5355i 2 Sept 1913,

NY. Carter: Dry open woods at Van Buren, E.J. Palmer 6203, 6

Ju 1914, MO; North-facing cherty slopes along Big Barren Creek T

25 N, R1W, Sect 10, 5 ni aw of Eastwood in rocky open field in valley, J.A. Steyermark 72313, 5 Aug 1951, F. Dade; Openings along Hwy 160 e of Greenfield, J.A. Steyermark 65225, 26 Sept

1947, F. Dent: Cherty limestone open rocky slopes above lime­ stone bluffs along Current R. nw of Cedar Grove sw of Jadwin near

Shannon Co. line, J.A. Steyermark 19408, 3 Aug 1935, MO. Douglas

Rocky hillsides in open woods near Ava, E.J. Palmer 34699, 26 Je

1928, MO. Greene: Rocky upland prairies and barrens near

Springfield, E.J. Palmer 28977, 2 Oct 1925, GH,M0; Roadside near

Strafford and open thickets along rd ne of Strafford, J.A.

Steyermark 24003, 31 Ju 1937, MO,OS,NY,F; Brookline, J.H. Kellogg

sn, 19 Sept 1908, MO; Greene Co., J.W. Blankinstrop sn, 5 Aug

1887, MO; Open woods w of Springfield, H. Eggert sn, 1882, MO;

Springfield, F.W. Dewart 79, 31 Ju 1892, MO; Bolivar Rd, Wm.

Trelease 566, 28 Ju.l897, MO; Willard, Blankinstrop sn, 6 Aug

1887, PH. Howell: Cherty limestone slopes in "County Hollow"

ca. 5 mi ne of Peace Valley, J.A. Steyermark 14490, 12 Aug 1934,

MO (two sheets),GH. jjasger: Forest Mill in rich bluff open woods, E.J. Palmer 2316, 23 Je 1909, MO,MO; Rocky banks at bor­

ders of woods near Sarcoxie, E.J. Palmer 32210, 17 Oct 1926, GH,

MO,PH; Upland prairies at Sarcoxie, E.J. Palmer 6259, 15 Ju 1914, 56

MO. Lawrencet Near Freistatt, C.H. Demetrio 108, 18 Aug 1897,

OB; Openings along road H n of Monett, J.A. Steyermark 65216, 26

Sept 1947, F. Me Donald: Dry ground, B.F. Bush 165, 1 Sept

1895, OH,NY; Me Donald Co., B.F. Bush sn, 24 Ju 1892, MO,MO;

Newton: Rocky hills near Neosho, H. Eggert sn, Aug 1882, MO,MO.

Oregon: Open thicket bordering field 1 mi s of Qreer 9 »i n of

Alton, J.A. Steyermark 7059, 23 Ju 1932, MO; Rocky open woods and hillsides "Irish Wilderness" w of Turner Mill Spring, E.J.

Palmer & J.A. Steyermark 41742, 14 Ju 1933, MO. Ozark: Rocky glades near Pontiac, E.J. Palmer 33105, 11 Oct 1927, NY,OH,MO;

Cherty open wooded slopes along Turkey Creek, T 23 N, R 15 W,

Sect 17, l1/6 mi ne of Hammond, J.A. Steyermark 22815, 1 Ju 1937,

MO. Phelps: Rocky prairies, J.H. Kellogg 15349, 4 Aug 1930,

MO; Cherty ravine tributary to Mill Creek, T 35 N, R 9 Sect

4, 3Y* mi (by air) n of Flat, J.A. Steyermark 74569, 1 Sept 1952,

F; Limestone glade above bluffs along Big Piney R. between

Spring Creek and Ousley Spring, J.A. Steyermark 19425, 10 Aug

1935, MO. Pulaski: Limestone glade above bluffs along Big Piney

R., Sect 32, k mi se of Tribune, J.A. Steyermark 25507, 28 Aug

1937, F,M0; Hillside above Roubidoux Spring at Waynesville,

Francis Drouet 16, 3 Ju 1932, UARK,MO; Pulaski Co., E. Hall sn,

Aug 187?, NY. Shannon: Shannon Co., B.F. Bush sn, 12 Sept 1888,

F,M0; Rocky sw-facing lime bluffs of Paint Rock Bluff along Cur­ rent R. 8 mi s of Exchange and 20 mi se of Eminence, J.A. Steyer­ mark 12304, 26 Ju 1936, MO,MO; Montier in woods, B.F. Bush 6103,

5 Aug 1910, QH,NY; High rocky hills at Montier, B.F. Bush 11433, 5 Aug 1927f MO; Rocky open woods in sandy soil near Montier, S.

J. Palmer 19276 and 13022, 3 Oct 1920, QH; In woods at Montier,

B.F. Bush 223* 2 Aug 1899* NY; Shannon Co., B.F. Bush l6l, 30 Je

189*+, NY; Rocky banks, B.F. Bush sn, 17 Ju 1891, NY. Stone:

Along Rte 86, 3*5 mi e of Viola, J.A. Steyermark 69933* 15 Ju

1950, F; Rocky hillsides at Galena, E.J. Palmer 4643, 13 Oct

1913, MO; Central Stone Co., Wm. Trelease sn, 10 Sept 1898, MO.

OKLAHOMA. Adairt Wooded valley 3 mi s of Kansas, U.T. Waterfall

9582, 7 Ju 1950, SMU. Beaver: Forgan, Robert Stratton 406, 20

Aug 1927* SMU,MO; Sand hills along small tributary of North

Canadian R. near Beaver, E.J. Palmer 41896, 24 Ju 1933* MO.

Cleveland: Ten mi flat at bottoms of S. Canadian R. at Norman,

D. Demaree 13083, 22 Je 1936, F,H0,GH; Norman, W.H. Emig 514, 21

Sept 1914, MO. Ellis: Sandy soil 5 mi e of Arnett, C.C. Smith

603* 6 Ju 1938, MO; Stabilized sand sunes 8j£ mi e of Harmon, U.T.

Waterfall 7128, 18 Je 194?, PH; In grassy draw near Shattuck, R.

L. Clifton 3200-V, 10 Je 1914, NY,GH. Greer: Grassy roadside near Granite, G.W. Stevens 1037* 17 Je 1913* M0,GH. Logan: Sandy grass prairie near Guthrie, G.W. Stevens 3264, 14 Je 1914, GH.

Me Clain: Loam soil 4 mi s of Purcell, P.R. Harding, Jr. 570,

28 Aug 1940, F. Me Curtain: In open woods near Shawneetown,

H.W. Houghton 3892, 28 May 1916, M0,GH,NY. Payne: Sandy soil near Ripley, C.D. Learn sn, 10 Ju 1916, SMU; Cimarron R. near

Perkins, Waugh 288, 14 Je 1893* MO. Roger Mills: Sandy Bhin oak area 10 mi sw of Cheyenne, W.C. Sooter sn, 14 Aug 1947, SMU.

Woodward: Woodward Co., P.J. White 20, 13 Ju 1900, MO. TEXAS. Austin: Industry (?), F. Lindheimer 253* Aug 1844, MO. Arm­

strong: One mi w of Ashtola in sandy loam ditch, Eula V/hitehouse

19150, 13 Ju 19^7* SMU. Bandera: Silt and limestone gravel

roadside 10.3 mi wnw of Bandera, L.H. Shinners 16882, 4 Nov 1953*

SMU. Bee: Beeville, M.E. Jones 29516, 30 Mar 1932, MO. Bexar:

In sandy loam bank of Cibolo 3. near Bracken, B.H.A. Groth 109*

10 Ju 1903* NY,GH,F; , Mr. & Mrs. J. Clemens 1008, 30

Je 1911* MO,NY. Blanco: Scattered in limestone soil along hvry

w of Dripping Springs, B.H. Warnock 46477, 26 Je 1946, UARK,M0;

Open calcareous banks along river near Blanco, E.J. Palmer 33950,

15 May 1928, PH,OH, Comal: Braunfels, F. Lindheimer 423 exsicc.,

May 1846, MO,MO,SMU,SMU,*PH,NY,GH,GH; Comanche Spring, New Braun­

fels, etc., F. Lindheimer 886, Je 1850, NY,F,PH,GH,MO. Cooke:

Tyler Bluff in rocky soil at western edge of county, D.S. & H.B.

Correll 12992, 2 Ju 1946, SMU. De Witt: Yorktown, H.B. Parks

17394, 28 Oct 1935* GH; Western De VJitt Co., Marguerite Hiedel

sn, 24 Apr 1942, MO. Donley: Red-brown sandy silt 11.8 mi nw

of Claredon, L.H. Shinners 15205* 19 Je 1953* SMU. Gillespie:

Near Fredicksburg, B.C. Tharp 285* Je 1939* MO, and sn, KY;

Along Hwy 16 between Kerrville and Fredericksburg, C.L. & A.A.

Lundell 13840, 8 Je 1945* SMU; Two mi e of Stonewall, V.L. Cory

4l8l8, 30 May 1943, GH; Gillespie Co., G. Jermy sn, no date, MO.

Goliad: Along 33 bank, Rev. C.B. Williams 59, Mar 1927* PH.

Hemphill: Open sandy soil at Canadian, E.J. Palmer 14101, 17 Je

1918, MO,MO; Deep sands in dune area at Gene Howe Wildlife

Management Area 5 mi e of Canadian, Chester Rowell 4056, 9 Je 59

1955, SMU; At Canadian on sandhills a of rirer, Hans Wilkens

2463, 17 Ja 1932i PH. Karnes: Dry sandy loan roadsids 3 ii & of Harass City, San Antonio River crossing, Stats Hwy 123, J.C.

Johnson 871, 28 Js 1952, SMU; Dry loany sand in fallow field 2,7

■i as of Karnes City, J.C. Johnson 1159, 15 Mar 1953* SMU. Kerr*

About Xerrville at 1600-2000', A.A. Ha liar 1874, 12-19 Js 1894,

MO,SMU,QH,NT,PH. Lira Oak: Live Oak Co., E.D. Schull 38-3 8 , 13

May 1938, F. Mason* Nsar Texas Rte 29» 10 ni w of Llano Co. lins, C.M. Rogers, C.C. Albars A Jos Barksdale 6893* 2 Ju 1949,

PH. Montague * Sandy roadside 11.2 ai se of Nocona, L.H.

Shinners 16387* 26 Sept 1953» SMU; Sandy post-oak woods 3*3 si e of Nooona on Hwy 23, Eula Whitehouse 10497, 22 Ju 1945, SMU.

Oldhaat Sandy hills 1 si w of Canadian S. on Dalhart Rd, R.S.

Ferris A C.D. Duncan 3518, 28 Aug 1921, M,NT. Pecos: Low ground near Leon Springs (7), Sister Mary Clare Mets sn, 16 Not 1931,

NY. Refugio: Refugio, H.C. Benke 5443, 12 Apr 1930, GH,F. San

Patricio: In sand section in clay bluff overlooking Nueces Bay,

8 si s of Taft, Fred B. Jones 131, Apr 1950, SMU. Trawls:

"Heill Sides," Austin, Ellhu Hall 325, 15 May 1872, F,NT,NT,MO,

QH; On linestone outcrop 5 si s of Austin, C.L. A A.A. Lundell

8919t 16 May 1940, NY,SMU,QH; Moist woods, M.S. Toung 149, 4

May 1918, MO. Victoria: Victoria, E.J. Palner 9161, 13 Mar

1916, MO. Wise: On clacareous prairie 8 si w of Slidell, V.

Newton 3, 22 Je 1939, SMU. Uncertain locations: Rio de Medina.

L. Berlandier 24?4 - 1044, Je 1834, PH,M0,QH, and 2474, NT, and

834 (7), MO. Bntre el rio de la Trinida 2 Bejar. L. Berlandier 60

2044, Junio 1026, K0,F,NI (all isotypes) and 431 (topotype).

(H) Beglandiera x betonicifolia (Hook. comb. nor. (■ Berlandi- •ra taxana x pumila)

Silphium betonioifolium Hooker. Coap. Bot. Mag. I: 99* 1835* (Syntypes BM, BM, photographs BMI BM1) Berlandiera texana var. botoaloifolla (Hook.) T. & 0. i n. of N.A. Vol. II,Part lit 281. 1842. Borlandlora botoaloifolia (Hook.) Small. FI. SE U.S. 1963. Horbaooous to suffrutooeont perennials with well-distributed loavost axis ofton branching. Loavos variable with triangulate to ovate leaves, one-half to one and one-quarter tines as broad as long} with dentate to serrate to deeply and doubly crenate margins; finely hirsute to nearly subacabrous above. Peduncles with fine curly long somewhat spreading to erect hairs, or sparse stout hirsute pubesoence. Heads 1.2-1.8 cm wide. Rays 9*18 nn long, 5.4-9.0 mm wide; deep yellow to yellow-orange, with green veins beneath. Disc corollas red to maroon. Achenea 2.5-4.0 mm wide, 4,0-5.5 nn long.

Type: This description is enlarged greatly over Hooker's original description in that it includes Drunnond's syntype collection fron New Orleans (?) in 1833 (Figures 14 and 15).

Most characters are intermediate of the parents in varying degrees and combinations over the range of east-central Texas and Louisiana. Northeast and east-central Texas specimens most nearly approach experimentally produced F^ progeny resulting from crossing parental extreme types. The hybrids look much more like B. pumlla than B. tcxana. Peduncle pubescence is of long rather fine curling hairs. Leaves slightly hirsute Figure 14. Berlandiera z betonicifolia Figure 15. Berlandiera z beton­ (Hook.) comb. nov. Copy of photograph and re icifolia (Hook.; comb. now. Copy of drawn tracings of ayntype (BM) of Silphium photograph of syntype (BM) of Silphium botonicifoliua Hooker, courtesy of British betonicifoliun Hooker, courtesy of Museum. British Museum. above. Middle cauline leaves ovate, nearly as broad or broader

than long, crenate to dentate. North and south of this group,

hybrids intergrade gradually toward the other parent, B. texana.

Drummond's syntypes more closely approach B. texana and have

been assigned to it as a variety by Torrey and Qray, who commented

that other specimens fitted neither varietal description complete­

ly. Characters used by Hooker (1835) in contradistinction to J3.

texana include longer petioles, less woody stems, hirsute leaves more ovate, of membranous texture; and from B. pumila by the

coarser and more deeply crenate margins, and by peduncles with

"beautifully jointed, purplish hairs." Peduncle pubescence of

13. texana is similar but longer and much more dense. On no

plant of any taxon have I found under surfaces with stellate hairs, a character commented upon by Hooker. The photographs

of the types do no t permit me to be certain of the character.

In south-central Texas, west of the "betonicifolia" com­

plex is a group of plants formerly called J3. dealbata which

Intergrade gradually into "typical" B. pumila northward. The

shorter petioles and the greater stem woodiness indicate possi­

ble similarity to B. texana. but other characters are of 13.

pumila to which I shall assign it. Experimentally, this taxon has been poorly investigated.

Specimens examined; TEXAS. Angelina: Diane Island, J.

Reverchon 2505* 5 May 1903* HO. Austin: Industry, F. Lind­ heimer sn, May 184A, *GH,*NY, and sn, Je 1844, *M0. Bastrop:

In pine land of Bastrop-Buescher State Pk, C.L. & A.A. Lundell

8994, 18 Hay 1940, *GH. Cass: Bevins, Ozette He Qinnis sn, 12 May 1941, *GH,*MO; Dry open fields 4 mi e of Hughes Springs,

D.S. & H.B. Correll 12490, 6 Je 1946, SMU. Colorado: Dry

prairies w Sabine R., H. Eggert sn, 28 Aug 1898, *M0 { Sandy

roadsides at Gladewater, L.H. Shinners 24269* 30 Ju 1956, *N0;

Longview, Q.W. Letterman sn, 19 Aug 1880, *M0. Hardin: In

open woods Zh mi w of Silsbee, V.L. Cory 52793* 9 Apr 1947*

NY,SMU; On grounds of Richard Collier Camp on Village Creek

7*4 mi nw of Silsbee, then 2 mi s, V.L. Cory 36664, 9 Jue 1949,

SMU; Sandy open ground at Fletcher, E.J. Palmer 9555, 25 Apr

1916, MO. Harrison: Dry sandy woods at Marshall, E.J. Palmer

7916, 8 J« 1915* *NY,*MO; Brushy cutover slope in sandy soil

4# mi ne of Marshall, L.H. Shinners 7568, 9 May 1945, SMU;

Roadside in soft sand ca. 8 mi nw of Caddo Lake State Pk on

Farm Rd 134, Karnack to Jefferson, Anne E. Orr 186, 9 May 1948,

•SMU; upland pine-hardwood forest at s edge of Marshall, D.S.

Correll 13238, 15 Ju 1946, SMU. Houston; Deep sand in open places of pine-oak woods near Grapeland, Eula Whitehouse 10631,

29 Apr 1931, SMU; Grapeland, B.C. Tharp 944, 8 Je 1920, *NY.

Jasper: Open oak woods in sandy ground 6 mi n of Jasper, L.H.

ShinnerB 76^6, 11 May 1945* SMU; In pine woods 2 mi n of

Jasper, H.R. Reed 966, 24 Apr 1950, SMU; In pine land off US

96 near Kirbyville, C.L. & A.A. Lundell 10545, 8 May 1941,

SMU. Marlon: Railroad near Jefferson, H. Eggert sn, Je 1899,

MO. Montgomery: In dry soil near JGN RR tracks, Royal A.

Dixon 469* 18-21 Ju 1909, *GH,*NY,*F; Deep sand and dry non­ calcar eous s red clay at 'iVillis, S.R. '.Varner sn, 12 Apr, •MO| Willisf R.A. Dixon 253, 20-25 Je 1908, *F. Newton; Flat pine lands with open secondgrowth stands 4 mi se of Newton, L.H.

Shinners 7857, 12 May 1945* SHU, Nacogdoches; Fern Lake, H.B.

Parks, Jr. 2346, *0.233, and 41234, 18 Ju 1940, MO,QH,QH respec­ tively. Polk: Indian Reservation, E. Girvin 285, 12 Apr 1940,

•GH; Deep loose sandy soil covered with duff beneath large old stand of Pinus oalustris at Fisher's Rd 4 mi ne of Camden, B.C.

Tharp, B.L. Turner & M.C. Johnston 280-5, 31 May 1954, SMU.

Rusk; Sandy upland oak-pine woods 4.6 mi sw of Henderson, L.H.

Shinners 14030, 6 Apr 1953, SMU. San Augustine; Sandy open ground at San Augustine, E.J. Palmer 9513, 20 Apr 1916, PH.

Shelby; Border of upland pine woods in sandy soil 16 mi se of

Center, L.H. Shinners 7628, 11 May 1945, SMU; Eight mi n of San

Augustine in Sabine Nat'l Forest, Eula Y/hitehouse 16619, 8 Sept

1946, SMU. Smith; In open pine woods at 'Western Tyler n of Pine

Burr Rd, V.L. Cory 56159, 20 Hay 1949, *SMU. Tyler; Sandy soil along Tex 287, 6.7 mi s of Y/oodville, J.F. Hennen 290, 16 Apr

1949, SMU; In open woods around pitcher-plant Hyatt Bog 2 mi s of Warren, V.L. Cory 57143, lS Apr 1950, SMU. Upshur; Clay

fencerow by partly cleared pine-oak woods 3 mi n of Ore City,

L.H. Shinners 18903, 24 Je 1954, SMU; Sandy clay soil between hwy and RR 7*8 mi se of Big Sandy, L.H. Shinners 15491, 27 Ju

1953, *SMU. Walker; Huntsville, R.A. Dixon 44, 3-12 Je 1908,

*F; Huntsville, on open sandy banks, E.J. Palmer 13386, 19 Apr

1918, *M0; In an old convict grave yard in vicinity of Hunts­ ville, R.A. Dixon 334, 9-12 Ju 1909, *F,*NY,*GH. Uncertain 65 localities: Florida (?)f "Dr. Torrey mis 1898," *K0; "Ark in

La" and Louisiana, Leavenworth sn, no date, *NY,*GH ("syntypes" of unnamed B. tomentoaa var. £ )* PROBLEMS OF NOMENCLATURE

I n * of uncertain atatua. Silphlua retloulatua Pur ah. FI*

Aa. Sept. 2: 579* 1814. Purah's daaerlption la ahort: "Sollla altarnaa oyato-laneeolatia oordatia aarratia obtuaiuaculua villoaluaculua* Harb. Banks» aaa. In Florida.H. ▼.a. in Harb.

Banka.11 Froa thia alona it belongs to or la affiliated with B. pualla. Ita taxonoaic aaaignaant auat raaain uncartain*

Silphlua patulua ? Thia taxon is listad in aynonoay by

Purah (1814) to S. puailua Purah. It ia liatad aa in "Harb*

Banka*'1 but with no othar data*

Tha Britiah Muaaua houaing tha Harbariua Bankaiana haa baan unabla to locate type apecimens of either taxon*

Orthoicraphical errora* Barlandiara aubacaula Nutt, Trana*

An* Phil* Soc* VII n*s*t 3^3* 1841, Aa Nuttall transferred tha nana froa Silphiun to Barlandiara* ha retained "aubacaule" instead of ita correct fora "aubacaulia•" Thia lad to Torray and Oray'a

B. aubacaulia (Nutt.) T. & 0. (1842). Cray (1884) finally corr­ ected the nana to B. aubacaulia Nutt.

Barlandiara texana DC. var.^batonicifolia T. & Q. FI* N*

Aa. lit 28l 1842* In tha tranafar of Silphiun botonicifoliua

Hooker (1833) to varietal status, "i" waa converted to Maa*M

Illaxitiaata nanaa. Silphlua puailua Purah* FI. Aa. Sept*

2 1 378-379* I8l4. Thia ia a later hoaonyn; "puailua" waa uaad 66 in the same genua by Michaux in 180J and therefore nust ba rejected under Article 6^ of tha Intarnational Coda of Botanical

Nomenclature (1961). Wood (1861) tranafara thia illegitimate nana to Silphiun aatariacua L. aa var. pumilun.

Unnamed varietiea. In 18^2 in their A flora of Worth

America, Torray and Oray introduce two unnamed variation: B. tonantoaa var. / baaed on a plant collected by Or. Leavenworth in waatarn Louiaiana (poaaibla ayntypaa Nil QHI){ and B. aub­ acaulia var. baaed on collection by Dr. Leavenworth in Florida

(poaaible isotype NYU. Tha latter variety ia characterized primarily by having leaves with oblong undivided terminal lobes. EXPERIMENTAL METHODS AND MATERIALS

Genus Berlandiera poses several taxonomic difficulties, notably intergradation among taxa and much variability within and between certain populations.

Analyses of morphology of herbaria specimens has been under* taken by Shinners (1951)* Turner and Johnston (1956) and Dyar

(1959)* As a result of her researches, Dyar proposes introgres* sive hybridization among plants of the Htexana-pumila,f complex.

It was also known that at least certain taxa could Interbreed via artificial crossing. The objectives of this study are to learn whether all taxa hybridize and how readily, whether there are any external or internal barriers to hybridization among taxa, and whether there is clinal variation.

Several biosystematic techniques are employed, but details will be discussed separately under respective investigations.

To conserve space two systems of abbreviations will be used. The taxa names will be replaced by the first three letters of the specific epithet, namely LYR, TEX, BET, PUM, HUM, and SUB. In addition, particularly in crossing data, genomic formulae con­ sisting of the first initial of the specific epithet will be used to indicate ancestry. Thus progeny of B. texana x B. pumila become TP, the first letter always representing the ovule parent

(TT); the second, the pollen parent (PP). Indentification numbers

68 69 following abbreviations rofor olthor to collection numbers as in

TT 1004 or to artificial cross numbers as in SS10-4, in which

case the plant is the fourth progeny of the tenth cross* In

addition, abbreviations will be used for collectors names used

repeatedly! SWP « John H. Speer, Wallace K. Weber, and D*J.

Pinkava; P ■ D.J* Plnkava.

The larger the population studied, the greater is the prob­ ability of the worker's learning of its overall variability, and

lesser is the chance of his being misled by the unusual plant

that caught the collector's eye. Specimens collected by the same person from the same locality at the same time, when compiled

froa several herbaria, still usually permit study of but a small

sample of the population* Where possible, large "herbaria popu­

lations" and mass field collections are used. Mass collections

and living plants were obtained throughout the range except from

Mexico. The following experimental plants were already placed

into cultivation under the care of Dr. Fisher when I began the

study: TEXAS: Hartley: Sand banks 35 mi sw Dalhart on Rte 54,

R.C. Jackson sn (1001, 1002, 1004,) no date cited, TEX. Lubbock:

Thirteen mi ese of Lubbock or 3 mi s of Acuff at 2700-3200*, no

collector cited sn (1009, 1010), no date cited, LYR. FLORIDA:

Pasco: Near Lacoochee, D. Smith sn (1011, 1013), 1 Apr 1961,

SUB. SOUTH CAROLINA: Aiken: Near entrance to Aiken State

Park, T.R. Fisher 750 (1008), 7 Aug 1958, PUM.

From crosses among certain of these experimental plants, Dr.

Fisher had obtained offspring: 13. texana x lyrata var. lyrata 70

(1000, 1015, 1016) and B. puaila x lyrata var. lyrata (1017, 1018, 1021, 1022).

A s*ri*s of four collecting trips rosultod in obtaining tha

following: ALABAMA: Baldwin: Along Ala. fit* 112, 5 si aw of

Perido R in dry sandy eut-ov*r pin*-oak woodlands, J.M. Sp**r,

W.R. W*b*r & D.J. Pinkava (-SWP) (1197), 30 Aug 1963, PUM; Ia

scraped ar*as and at *dg*s of oak-pin*-dogwood wood, along Ala.

Rt* 112, 1.9 ai uw of Perido R., SWP (1198), 30 Aug 1963, POM.

ARIZONA: Coohle*: Dry stony olay roadsid* of Rt* 666 at a* corner of jet 666 and rd w to Courtland, Y.J., Agn*s & D.J.

Pinkara (924-928), 28 Dec 1962, LIR. Santa Crus: Dry hard road­ sid* loam on rd froa Elgin to Rt* 82, 1.9 ai s of that jet with

Rt* 82, just w of Mustang Mta. and near Cochioe-Santa-Crus Co. line, Y.J., Agn*s & D.J. Pinkava (920-923), 28 Dec 1962, LYR.

FLORIDA: Alachua: Along Fla. Rte 236, 2 mi froa jet with Fla.

Rt* 237 and *n* of Alachua, SWP (1234), 1 Sept 1963, PUM. Cal­ houn: Hillsid* along grassy roadsid* of Fla. Rte 2?4S, 0.8 mi

from jet with Fla. Rte 6 9 , SWP (1224), 31 Aug 1963, PUM. Duval:

In churchyard lawn n*ar Beach Blvd. and Palm Lane off US Rt* 90,

3.7 ai • of Jacksonville and 2.9 mi e of Littl* Petersburg Creek,

SWP (1241), 2 Sept 1963, PUw. Escambia: Edg*s of pin*-oak woods along US Rt* 29 n froa Cantonaent between jet with Rte 63 and with Rt* 63 and with Fla. Rt* 1 9 6 , SWP (1199), 30 Aug 1963, PUM.

Oadad*n: In d**p sand along pin* woods, US Rt* 90 and off aid*

rd ca. 7 ai • of Chattahoochee, SWP (1227), 1 Sept 1963, PUM; In

scrub oak woods in sand 3*5 ai »• of Gadsden-Liberty Co. lin* on 71

Fla. Rte 267 and ca. 11 mi s of Quincy, D.J. Pinkava (»P) (524-

540), 6 Oct 1962, PUM. Holmes; In deop, dry sandy soil in pine- oak woodland along US Rte 90, 9*0 mi w of Big Reedy Creek and e of Argyle, SWP (1212), 31 Aug 1963* PUM; Roadside, 0.13 mi e of

Big Sandy Creek on US Rte 90 and 0.43 mi e of jet with Fla. Rte s-lOA, SWP (1216), 31 Aug 1963, PUM. Jackson: Sandy roadside of Fla. Rte 69» 2.6 mi n of Jackson-Calhoun Co. line and 6.1 mi s of jet with US Rte 90, SWP (1225), 31 Aug 1963, PUM; In sand wash in roadside ditches and banks on Jim Woodruff State Pk Rd

1.2 mi n of jet with US Rte 90, this 3*7 mi e of Grand Ridge e limits, P (541-551), 7 Oct 1962, PUM. Nassau; Along US Rte 1,

23, 0.5 mi s of St. Mary's R. and 0.15 mi s of Ga. State line,

SWP (1243), 2 Sept 1963, PUM* Okaloosa: In pine-oak woods along US Rte 90, 1.3 mi e of Holt, SWP (1206), 31 Aug 1963, PUM;

In sandy ditch wash and pine-oak woodland along US Rte 90, 7.1 mi w of Walton Co. line, S7/P (1208), 31 Aug 1963, PUM. Pasco:

In sand along Rte 19, 0.2-0.3 mi n of Hudson, Stephan, Armstrong,

Arnholt & Pinkava (1014, 1025), Mar 1962, SUB. Putnam: In lawn near US Geodetic Survey Marker along US Rte 20, 11.4 mi e of jet with S315, near Palatka, SY/P (1235), 2 Sept 1963, SUB. Santa

Rosa: Ditch bordering oak-pine woods along Fla. Rte 196, 0.8 mi e of Escambia R., SWP (1202), 30 Aug 1963, PUM; Along scrub oak- pine woods border along US Rte 90, 2.5 mi e of Milton City and

Blackwater R., SWP (1203), 31 Aug 1963, PUM; On grassy roadside of US Rte 90, 7.8 mi w of Holt, SY/P (1205), 31 Aug 1963, PUM.

St. Johns: Roadside, lawn, and woodland border along Fla. Rte 207* 3.1 mi ne of Elkton w limits, SWP (1238), 2 Sept 1963t PUM-

HUM. Walton: Along RR near US Rte 90, 13*2 mi e of Y/alton Co. line, SWP (1210), 31 Aug 1963, PUM. Washington; Along rd and pine woodlot 2.4 mi e of Holmea-Y/ashington Co. line, SWP (1217),

31 Aug 1963* PUM. GEORGIA: Charlton; Oak-pine woodlot at 2nd

St. and Bay St. off US Rte 1, 23 near Folkton, SWP (1246), 2

Sept 1963, PUM. Lee: In sand of fresh roadcut along Rte 118,

3*4 mi e of Smithville e limits, P (306-323). Houston: Front lawn in sand along US Rte 341, 2.7 ml sc of Peach Co. line, SWP

(1262), 4 Sept 1963, PUM. Taylor; Georgia Rte 96 w of Reynolds,

4.6 mi e of Butler e limits, SY/P (1264), 4 Sept 1963, PUM; Along

Ga, Rte 96, 3.4 ml e of Butler e limits, SWP (1266), 4 Sept 1963,

PUM; Along Ga. Rte 96, 6.6 mi w of Butler e limits, S'WP (1267),

4 Sept 1963, PUM. Toombs: Along unnamed "dirt" rd 0.3 mi from jet with US Rte 280 and 0.8 mi e of Lyons se limits in deep sand of hillsides in very dry oak woods, S'WP (1253), 4 Sept 1963, PUM.

MISSOURI: Barry; Stony roadcut among few oaks along Mo. Rte 37,

2.15 mi s of Butterfield s limits, P (894-901), 20 Oct 1962, TEX.

Lawrence: Foncerow in red rocky soil along Rte H off US Rte 60,

0.5 mi n of Monott n limits, P (902-905), 20 Oct 1962, TEX. NEW

MEXICO: Chaves; Roadside along US Rte 285, 14.8 mi s of Roswell s limits, P (839-844), 17 Oct 1962, LYR. Eddy; Roadside along

US Rte 285, 16.4 mi s of Artesia s limits, P (836-838), 17 Oct

1962, LYR. SOUTH CAROLINA; Aiken; Sand 1-2-feet thick atop light-red sandy clay along S.C. Rte 215, 5*7 mi w of entrance to

Aiken State Pk, SWP (1291), 5 Sept 1963, PUM; Deep sand in dry 73 oak woods along roadsid# at #ntranc# to Alk#n Stats Pk, SWP

(1294), 3 S#pt 1963» PUM, Orangeburg: Oesp sand n#ar oak wood­ land along S.C. fit# 4, 2.2 ai w of N##o#s w limits, SWP (1297-

1298), 5 Sopt 1963* PUM. TEXAS: Band#ra: Along T#x. fit# 16,

20.6 ai w of jot with T#x. fit# **6 , P (714-730), 14 Oct 1962, TEX;

With grass in hard, dry clay soils along T#x. fit# 16 ca. 100 yds s# of M#dina s limits and roadsid# pk, P (733-743), 14 Oct 1962,

TEX. Br#wst#r: Along T#x. fit# 118, 18.1 ai s of Alpin# s limit#, P (730-751), 16 Oct 1962, LYR1 Along T#x. fit# 118, 15.35 ai s of Alpin# s limits (s##ds only), P, 16 Oct 1962, LYR; Along

T#x. fit# 118, 0.45 ai s of Alpin# s limits, P (767-776), 16 Oct

1962, LYfi. Cass: Roadsid# opposit# roadsid# pk on T#x. Rt# 11,

2* ai • of Hughe# Springs, P (584-595), 10 Oct 1962, PUM.

CulbTSon: Roadsid# along US fit# 8 0 , 0.05 ai • of Van Horn • limits, P (822-833), 16 Oct 1962, LYfi; Pin# Top Mt. at Pin#

Springs on T#x. Rt# 54, P (834-839A), 16 Oct 1962, LYR. Fayette:

Roadsid# #abankn#nt of red clay along T#x. fit# 71, 9*1 ai • of

La Grang# • limits, P (700-712), 13 Oct 1962, TEX. Qr#gg-

Ppahur: In sand of roadcut, US Rte 80, 0.75 ai w of Glad#- water, P (621-630), 11 Oct 1962, BET. Hardin: Cl«ar#d pine- oak woodlot in sand along T«x. Rt# 327 w of Silab## and 1.3 ai

• of Village Cr##k, P (680-692), 12 Oct 1962, BET. Harrison:

Along spars# pin# stand at roadsid# near f#nc#row, T#x. Rt#

43, 8.9 ai sw of jet with Farm Rd 134 at Karnak, P (631-642),

11 Oct 1962, BET. Hemphill: Along US Rt# 60, 0.05 ai sw of

Glazier sw limits, P (865-866), 18 Oct 1962, TEX. J#ff Davis: 74

Roadaido of Tox. Rt* 118, 13 ai • of Ft. Davie • liaito, P.

(777-788), 16 Oet 1962, LYR; Boadaldo of Tox. Rt* 17, 3.6 ai a of Ft. Davia a lialta, P (789-799)* 16 Oot 1962, LYR; Boad- alda of Tox. Bto 17, 13.3 ai a of jet with US 290, P (800),

16 Oet 1962, LYR; fioadaido of Tax. Bto 17, 6.23 ai a of jet with US 290, P (801-810), 16 Oet 1962, LIB. Lipaooabt 11on®

US Bto 60, 1.1 ai aw of Higgina aw liaita, P (868-867), 18 Oet

1962, TEX. Mowtoa1 fioadaido under pin* and extending up paaturo

hillaido, Tox. Bto 6 3 , 4.4 ai w of Tox.-La. lino and 5*0 ai o

of Burkvill* o liaita, P (643-656), 12 Oet 1962, BET; In aand

banka noar edge of wooda, Tox. Bto 87, 0.07 ai n of Newton n liaita, P (668-679) (637), 12 Oet 1962, BET. Palaort Roadsid* pk in rod loaay aoil, US Bto 60, 1.45 ai aw of Bovina w liaita,

P (8^5 -850), 18 Oot 1962, LYR. Pocoa-Browater1 Very dry liaey

rocky roadaido of US Rte 385, 21.3 ni no of jet with US Rte 90

between Marathon and Ft. Stoekton, P (744-746), 15 Oct 1962,

LTR. Randall1 In rod aoil along Tox. Rte 217 e of Santa Fo

RR ovorpaaa noar 11th St. and 3rd Av* in Canyon, P (851-864),

18 Oot 1962, LTR. Roovoat Roadaido of US Rt* 290, 14.3 ai oao

of Kent o liaita, P (811-821), 16 Oct 1962, LYR. Upahurt Sandy

aoil of roadeut of Tox. Rt* 154, 7.4 ai o of jet with Tox. Rte

155 at Qilaor, P (596-616), 11 Oet 1962, PUM; On wound of aand

along Tox. Rte 155 ea. 500 ft no of jet with Fara Rd 1404 and

6.65 ai no of Big Sandy no liaita, P (617-620), 11 Oct 1962,

PUM-BET. CYTOGENETICS

Various workers have made chromosome counts of plants of

western Berlandlera taxa which are summarized in Table 1 under

their original identifications in the literature. My study

concerns chromosome number and behavior in both the parental

experimental plants and their F1 hybrid progeny (Table 2).

Buds of greenhouse-cultured plants were killed in a

solution of six parts chloroform, three parts absolute ethanol

and one part propionic acid and maintained at approximately

3° C until examinations were made. Nearly all counts and

behavior observations cited in Table 2 and Figures 17-21 are

from pollen mother cells squashed from fixed anthers stained

with propio-orecln. Exceptions include a tetraploid tapetal

cell chromosome count (Figure 19c) and a mitotic root tip

smear prepared by Y/ilbur Settle (unpublished) using TBA-Fuelgen

technique (Figure l8h).

All parental and hybrid plants examined had n * 15, con­

sistent v/ith previously cited data. However, many of the plants have "abnormalities." Stickinoss of chromosomes is

troublesome because it results in formation of false bridges

in mitotic and neiotic figures, as many as four such "bridges"

in one cell (Figure 20c).

Laggard chromosomes were commonly encountered, particu­

larly in B. lyrata plants and their hybrid offspring (Figure 21).

75 76

TABLE 1

SUMMARY OF CHROMOSOME COUNTS REPORTED IN LITERATURE

Taxon Description n

B. lgSfita var. lyrata. Grown in Austin from soed collected in Alpine, Brewster County, Texas. Johnston 5*11032. (Turner & Johnston, 1956) 15

B, lyrata. City limits of Durango. Durango. ~~ King 3750. (Turner, Powell & King, 1962) 15

B. lyrata. Along highway to Buenaventura, 3*6 mi _ ------Rj_car

B. texana var. texana. Two mi w of Bandera, Bandera County, Texas. Turner 3841. (Turner & Johnston, 1956) 15

B. texana var. betonicifolia. Nineteen mi w of *” Giddings, fiastrop bounty, Texas. Turner & Johnston 5**357. (Turner & Johnston, 1956) 15

B. texanavar. betonicifolia. Two mi e of Fulshear, ~ Fort Bend County. Texas. Turner3819. (Turner St Johnston, 1956) 15

B. texana var. betonicifolia,. Fourteen mi s of fairchild. Turner 3814 (this specimen ap­ proaching B. pumila). Leon County, Texas. (Turner Sc Johnston, 1956) 15

B. pumila. One mi sw of New Ulm, Colorado County, ” Texas. Turner Sc Johnston 5^329* (Turner St Johnston, 1956) 15

B, pumila. Three mi n of Belmont, Gonzales County, ~ Texas. Thompson Sc Turner sn. (Turner 8c Johnston, 1956) 15

B. pumila. Grown in Columbus, Ohio} from entrance ~~ to Aiken State Park, Aiken County, South Carolina. Fisher 1750 (Dyar, 1959) 15 77

TABLE 2

CHROMOSOME DATA OF CURRENT STUDY Phaaaa Taxon n Raoordad Additional Coauaanta

LL 1010 15 M*. D, M 2 Micronuolaii thraa apin< dlaa at A 2 j laggarda

LL 838 Maioaia aa uaual 15 «1 TT 1004 15 M (Mitoaia) Tatraploid tapatal call with 60 ohromoaomaa, two nuclaoli*

TT 1002 Laggarda at A thraa 15 *1 \ "bridgae" at A. Cona call) 1

TT 1001 15 D Laggard at A^

TT 901 15 D Two "quadriralants" and 11 bivalanta at D| "bridga"

TT 730 15 D

BB 680 15 D t M1

PP 595-2 15 D Thraa nuclaoli of thraa aizaa in ona call

PP 526 15 0

SS 11-5 15 Four "bridgaa" at A. in S ’ M i ’ ona call 2 TL 240-7 15 M 1 LT 22-5 - - Laggarda at M 2

LT 29-6 With laggarda 15 M 2

* Symbols of phaaaa: M « metaphaaa; A * anaphasa; D * diaklnaaio; aubacripta 1 , 2 rafar to firat and aacond maiotic diviaiona raapactivaly. 78

TABUS 2 (Continued)

Pbaaoa Taxon a Additional Comaanta Recorded

LT 10X6 15 Laggard at A2 ; micro- nuelait inveraion bridge ♦ fragment croae apin- dlaa at A2

BL 740-1, "Bridge" at A of mitosia in root tip call, TBA + Fualgan preparation (Settle, unpub*)

LB 118-1 - m Laggard at A? ; "quadri­ valent" at

PL 255-1 15

SL 8-8 - - Laggarda at

SL 8-2 15 D, M2 Moet A. cella with lag­ garda i micronuclei; atickineae

LS 6-1 15 D

SL 5-5 - - Laggarda; "bridges"; micronuclei

TP 197-2 15

BT 175-6 15 D

PB 172-4 15 D Many cella with linked bivalenta at D and

TS 1-4 15 D, M x 11 Quadrivalent a": 7 IV, 1 II; 5 IV, 5 11; moat with 15 II

SB-144-1 15

PS 230-1 15 D Figure 16• Microaporogeneaia I* Heioaia I diakinesia and metapha.ee configurational all n * 15. a. SS 11-?. b. PP 526* c. LL 1010. d. TT 1001.

79 % 80

f 1

MU

lu

t C \ • * # * MU

y < Figure 17* Microsporogenesie II. Meiosis I diakinesis and metaphase figuresf all n ■ 1$. a. SS 11-5. b. BS 144-1. c. SL 6-1. d. LT 240-7. e. TS 1-4. f. LP 255-1* nucleolus stippled. Scale: ca. 1500x.

81 4 a * • v . ;

o

f t

^ %s i 4 > N \ • ?

■* * < j

• Figure 17 Figure 18, Hieroaporogenesis III* All figures n * 15* a. LL 838, M . b. TT 730, D. c. BB 680, D. d. TP 197-3, M-. e. BT 175-6, M2 . f. BP 172-^, M,. g. "Tetrad” of five microsporea with five micronuclei (solid) and four nucleoli (stippled), LL 1010. h. "Bridge," root tip mitosis, BL 7^0-1, preparation by Wilbur Settle (unpub.)* Scale: ca* 1250x.

83 r *

\

A i

9 r'm

Figure 18 Figure 19* Microsporogenesis IV. a. Persistent Anaphase I inversion bridge cross spindles plus fragment, LT 1016. b. Diakinesis configuration with evidence for sets of synapsed pairs (tetraploidy)| n ■ 15, PP 595-2. c. Tetraploid tapetal cell (60 chromosomes), mitosis, TT 1004, Scale: ca. 1250x.

85 L

86

t • j * / , * » §& y *

Mu

If *;

o v * i * ^

MU

% * £ >

Figure 19 Figure 20. Microsporogenesis V. Meiotic irregularities, a. "Quadrivalentsf" Met. 1, TT 901. b. False bridges* Ana* I, TT 1002* c. Four false bridges* Ana. I* SS 11-5. d. Seven "quadrivalenta" and one bivalent, Met. I* TS 1-4.

87 88

«•

•

MU

MO

•IS i t - • * &4

MU

d }1U Figure 20 Figure 21. Microsporogeneaia VI* a-k. Meioaia I and II irregular!tiea, LS 8-2. Scale} ca. 1250x.

89 & %

A % * t ;

x © $>

Figure 21 8 91

The cause of laggards la unknown, but nay rosult from late sepa- ration of chromosomes bald togather by stickiness.

Laggards baeona variously oriantad in tha call. Thay nay ba inoorporatad into nioroaporaa containing nuolai with full complement, giving risa to aaeropollan. Or, tha laggards nay baoona tha only nuolai (nicronuclai) of tha nierosporas, giving risa to nicropollan. Hataronorphic pollan grains nay ba obsarvad sida by sida in praparations fron anthers of tha sane flower

(Figure 22).

In a study of LL 1010, it was obsarvad that tha nunber of nicronuclai par intact tetrad of miorospores ranged fron none

to six, and that 62 percent of such tetrads had at least one micronucleus. At tha tine miorospores of tha tetrad separata nearly 98 percent are without nicronuclai. Thus, tha micro­ nuclei are either incorporated or disintegrate, probably tha latter since in this plant macropollen has not bean found. In either oasa, incorporation or disintegration (unless reconsti­

tution is exact), there is an unequal distribution of nuclear

■aterlal. Ninety-two percent of tha mature pollan of this plant stains positively with analine blue. This figure compares

favorably with tha ninety-eight percent micronuclaus-frea micro- spores cited above. Some micropollen stains positivelyf most does not.

The presence of heteromorphic pollen in counts made in the

field fron fresh material (Table 3) la evidence that hetero- morphy of pollen was not induced by greenhouse conditions. 92 During this investigation evidence was found that Borland-

iera taxa are tetraploid. Usually chromosomes synapseaas

fifteen bivalents during Meiosis I. Occasionally, however,

chromosomes are found groups of four (possible quadrivalents)•

One anther smear preparation from B. toxana x subacaulis (TS

1-40 contained two cells in each of which were one bivalent

and seven "quadrivalents" (Figure 20d); one cell with five

bivalents and five "quadrivalents"; and several cells with

fifteen bivalents.

A cell at early diakinesia (Figure 19b) contained synapsed

pairs of chromosomes in two morphological sets. Clearly the

two pairs of longest chromosomes are similar and the two pairs

of "doughnut"-shaped pairs are similar. Ten other pairs could

be matched but with less certainty. The remaining unmatched

pair was of the same "V" shape as the lone bivalent found in

TS 1-4- described above.

Additional evidence of tetraploidy is the association of

two similarly shaped synapsed pairs of chromosomes with the

single nucleolus, presumably the product of fusion of nucleolar

material of the two "homologous" pairs of chromosomes of the

tetraploid set. See Figures l8b, c; 19b; 21a.

Even though very few backcrosses and F^ plants were suc­

cessfully raised to maturity, of those that were, certain

characters are better explained based on tetraploid ratios than

on diploid ones. Vein color on under surfaces of ray flowers

and color of disc corollas (Figure 23) are controlled apparently 93

TABUS 3

POLISH STAIHABILITT DATA FROM FT«tji nnr.TJfiTTnwg

Plant Total 8talnablo Non-ataln- 96 Stain- Micro- Macro- Ho. Pollon Pollon ablo Pollon ablo Pollon Pollon

Bacaabia County. Florida r-ii22

2 912 863 49 94.63 0 0 if 700 695 5 99.29 0 0 7 674 662 12 98.22 2 2 15 652 597 55 91.56 1 0 17 llif9 726 423 63.19 1 2 19 780 752 28 96.41 0 0 21 609 552 57 90.64 2 2 23 873 872 1 99.89 0 1 2if 767 759 8 98.96 0 1

9 7116 6478 638 92.53 6 8 0.0896 0.1156

Santa Soaa County. Florida - 1205

1 518 505 13 97.49 1 0 2 6ifl 350 291 54.60 1 1 3 861 845 16 98.14 0 0 if 8if8 726 122 85.61 1 0 5 if76 469 7 98.53 0 1 6 725 705 20 97.24 0 0 7 692 689 3 99.57 0 0 8 84l 823 18 97.86 1 0 9 1123 1114 9 99.20 0 0 10 1226 1207 19 98.45 1 0 11 836 826 10 98.80 0 0 12 8ifl 837 4 99.52 1 0 13 536 520 16 97.01 0 0

13 10l6if 9616 548 94.00 6 2 0.0696 0.0296

Walton County. Florida -> 1210

1 546 528 18 96.70 0 0 2 809 796 13 98.39 2 4 3 732 721 11 98.50 0 0 if 1089 1059 30 97.25 0 0 94 T A B U 3 (Continued)

Plant Total Stalnablo Non-stain- % Stain- Micro- Macro- No* Pollon Pollon abl* Pollon ablo pollon pollon

6 513 494 19 96.30 0 0 7 571 550 21 96.32 0 1 8 746 734 12 98.39 0 0 9 1564 1553 11 99-30 1 4 10 509 502 7 99-01 1 0 11 532 525 7 98.68 1 0 12 622 619 3 99-52 0 4

11 8233 8081 152 98.03 5 13 0.0696 0.1696

Holaoa County. Florida - 1212

5 670 640 30 95.52 1 0 6 622 584 38 93.89 0 0 9 1136 1113 23 97.98 4 2 10 518 503 15 97.10 0 0 11 1124 1074 50 95.55 1 0 12 797 775 22 97.24 0 1 13 643 588 55 91.45 0 1

7 5510 5277 233 95.53 6 4 0.1196 0.0796

Putnam County. Florida - J £ 2 2 1 989 978 11 98.89 1 1 2 669 665 4 99.40 0 2 3 569 564 5 99.12 0 0 if 840 832 8 99.05 0 0 5 804 795 9 98.88 0 0 7 772 761 11 98.58 0 0 9 628 623 5 99.20 0 0 10 770 759 11 98.57 0 1 12 941 930 11 98.83 0 0

9 6982 6907 75 98.95 1 4 0.0196 0.0696

St. Johns County,. Florida - 1238

1 854 850 4 99.53 0 0 4 534 528 6 98.88 0 0 5 654 629 25 96.18 6 0 6 645 638 7 98.91 2 0 8 IO69 1059 10 99.06 0 0 10 788 785 3 99.62 0 0 12 l48l 1459 22 98.51 0 0 TABLE 3 (Continued)

Total Stainable Non-stain- % Stain- Micro- «» Pollen Pollen able Pollen able pollen _ *______

Nassau County. Florida - 1243

1 876 858 18 97.94 0 3 1189 1120 69 94.20 2 5 1039 1025 14 98.65 0 7 799 774 25 96.8? 0 8 691 666 25 96.38 0 9 816 750 66 91.91 0 11 1161 1123 38 96.73 0 12 861 855 6 99.30 0 13 685 679 4 99.41 2 14 649 642 7 98.91 0

10 8764 8492 272 97.03 4 0.0596

Taylor County. Georgia - 1264

1 513 510 3 99.42 0 6 734 719 15 97.96 0 8 533 507 26 95.12 0 9 750 741 9 98.80 2 10 630 626 4 99.37 0 13 904 893 11 98.78 0

6 4064 3996 68 98.24 2 0.0596

Orangeburg County. South Carolina - 1297

1 1052 1050 2 99.81 1 2 563 549 14 97.51 0

2 1615 1599 16 98.66 1 0.0696 Figure 22. Heteromorphic pollen. SL 8-7 Only the macronucleus (center) stained posi­ tively. Note oil droplets in field. Scale: ca. 650x. by two pairs of simple dominant-recessive gene loci linked on the sane chromosome. Some dilution effect has been found in certain plants of various taxa and is as yet unexplained. In ray-vein color, red to maroon (R^) is dominant to green (r^); in disc- corolla color, red to maroon (Rg) is dominant to yellow (r^).

Thus postulated genotype for B. lvrata is R^R^R^R^R^R^R^R^} for

"texana-pumila" complex, ririR2R2rlrlH2R2 * **** for — * eut>ac r^r^r^r^r^r^rgrg* In the following data, it has been assumed that there is diploidization of the chromosomes (absence or near absence of quadrivalents) and that, consistent with data so far obtained, linkage is tight enough that cross-over between the two loci in question is virtually non-existent. All these assump­ tions are vulnerable to change with additional data. Under this hypothesis, segregating backcrosses should yield for either character a 3:1 ratio if tetraploidt a 1:1 ratio if diploid.

Segregating progeny should yield a 15*1 ratio if tetraploid; 97 3*1 ratio if diploid. Bven though the sample ia emailf data tabulated below are not inconsistent with tetraploid hypothesis.

Chi-square test is much more in accord with the tetraploid hypo­ thesis than with the diploid hypothesis (Tables 5).

Berlandiera taxa very closely resemble taxa of the genus

Silphiumt in fact, early-named taxa were called members of genus

Silphium. The genus Berlandiera differs from Bllphimn in that the former has but a single series of fertile ray flowers, ach- enes that are wingless and not distinctly toothed atop, and two disc flowers and their enclosing paleae which adhere to the ach- ene and abscise with it. Many parts of Berlandiera taxa are smaller than or are fewer in number than those of Sllphium taxa.

The pubescence (except specialized forms) of both genera are of the same basic construction, but much more dense in Berlandiera species. Stipitate glands are found in some Silphium taxa but not in Berlandiera taxa. Multicellular "glands" along the axis of the anther appear in all Berlandiera plants investigated, none are known to exist in Silphium taxa.

Although the chromosomes of Silphium are about ten times the size of those of Berlandiera. their behavior and morphology are similar if not identical, so far am one can judge from present observations. Stickiness of chromosomes and false bridges

(Cruden, i960) are found in both genera. Silphium has x = 7, and doubling this number would yield n = 14, one less than is found in Berlandiera. One might still postulate Silphium or a

Silphium-like prototype ancestry if one permits the acceptance 98 of another theory. Silphium genomes contain one very large chro­ mosome each. Some Silphium plants have an additional chromosome pair of very small slse (Fisher and Cruden, 1962). One might reason that Silphium was originally x ■ 8 ; but that due to an un­ equal translocationv one chromosome became very large* the other very small* so small that genetic material and cross-over frequen­ cy are greatly reduced, possibly to the extent that the small chromosomes remain on the metaphase plane of Meiosls 1* lagging behind and becoming lost from the chromosome complement with no severe effect on the phenotype. If the genetic value of the lost chromosome is small* or at least non-vital, the hypothesis of re­ duction of chromosome number is reasonable. Berlandiera*s 2n ■

30 might, then, arise by union of gametes with 7 and 8 chromo­ somes and subsequent doubling, or by union of unreduced gametes of 14 and 16 chromosomes; or by union of 8- and 8-chromosome gametes and doubling or union of 16- and 16-chromosome gametes, in both latter cases then with a loss of a pair of chromosomes

(aneuploidy).

The proof of the ancestry lies in successful crossings among Berlandiera and Silphium taxa with resulting triploids which could be easily detected by chromosome number. All crosses to Silphium taxa have failed thus far.

Evidence against tetraploidy includes (l) the occurrence of velvety-leaved taxa not known in Silphium; and (2) the reduction of many parts, not previously known for tetraploids, though some 99 tetraploida, for example Tradeacantla ohioenala (Dean, 1959), are Indiatinguishable from dlplolda. 100

Figure 23. Corolla Color*• Say and diec corolla colors. Top group of five heads, from left and from topi BB 615* LL 851* TT 1004 (center)* PP 526* SS 10*4. Center group of five heads: Same sequence, heads overturned exhibiting ray-vein color beneath. Bottom group of two heads exhibiting segrega­ tion of ray-vein color, siblings, SSxLP 24-1 (left), 24-6 (right). Scale: ca. 0.7*. TABLE 4

. GENETIC DATA SUPPORTING TETRAPLOIDY HYPOTHESIS

Ray-vein and Disc-Cor- Expected Expected Taxon olla Colorsv Linksd Ratio if Ratio if R-R a-R R-Y 0-1 Diploid Tetraploid Backcrosses

LT x SS 13 (1015x1014) 1 1 SS x LT 14 (1014x1015) 3 1 SS x LT 27 (1011x1016) 2 1 LT x SS 28 (1016x1011) 0 1 - -

Subtotal 6 4 - 5:5:0:0 7.5:2.5*0:0

SS x LP 16 (1014x1021) 8 0 SS x LP 24 (1013x1018) 2 4 - -

Subtotal 10 4 - 7:7:0:0 10.5:3.5:0:0

SS x SL 253 (10-4x 5-6) 3 1 S3 x SL 387 (10-4x 5-6) 4 2 SL x SS 388 (5-6 xlO-4) 1 - - 0

Subtotal 8 - - 3 5.5:0:0:5.5 8.25:0:0:2.75

ST x SS 366 (2-1 xlO-4) 6 1 S3 x ST 488 (11-lx 2-1) - 2 - 3

Subtotal - 8 - 4 0:6:0:6 0:9:0:3

F 2 Progeny TS x ST 355 102 TABLE 4 (Continued)

Ray-vein and Disc-Cor- Expected Expected Taxon olla Colorst Linked Ratio if Ratio if R-R Q-R R-Y 0-Y Diploid Tetraploid

ST x TS 356 (2-1 x 1-3) - 1 1 ST x ST 1*69 (2-5 x 2-3) - 2 0

Subtotal - 6 0:5.25:0:1.75 0:6.56:0:0.1*4

LP x LP 31 (1021x1022) 2 0 LP x LP 32 (1022x1021) 3 0

Subtotal 3.75:1.25:0:0 4.67:0.35:0:0 103

TABLE 3 M 1 0 0 Class Observed (o) Expected (c) o-c c

Baclccrosses - For ¥etraploidy 3*i

Dominant 32.00 35.25 - 3.25 0 .3 00

Recessive 13.00 11.75 + 3,25 0.899 if7 .66 If?.0 0 6.06 1 .1 69

Backcrosses - For Diploidy 1:1

Dominant 32.00 23.50 ♦ 8.50 3.07^f

Rscsssive 15.00 23,50 - 8.50 47.00 W . w O .60

Fg Progeny - For Tetraploidy 15*1

Dominant 11.00 11.25 - 0.25 0 . 0 0 6

Rscsssive 1.00 0,75 ♦ 0.25 0 . 0 8 3 11.00 2 .o6 6.60 X 5358?t

Fg Progeny - For Diploidy 3:1

Dominant 11.00 9.00 + 2.00 0.444

Recessive 1.00 3.00 - 2 . 0 0 TSt55 12.00 0.00 ----- 1— 8T at 0.3 level. 10*t

CROSSABILITY

The criterion of crossability among taxa in this study is artificial crossing achene-set. Achenes that appear plump are considered to have viable seeds} those that are flattened are considered to have non-viable seeds* Germination studies are consistent with this hypothesis. Crossability in the field has not been investigated except by casual observations made while collecting fruits that achene-set was frequently less than 50 percent*

Pollination was effected by manipulating staminate disc flowers with a pair of forceps in such a way that the pollen­ laden anthers were brushed against the inner surface of each stigma lobe of each ray flower until pollen could be seen ad­ hering to the stigma. In those flowers which set achenes, the stigmas were found to wither soon after pollination. Both staminate and ovulate flowers used in a cross were always from heads bagged long before maturity with glycine sacks held fast to the peduncle by a stringed tag.

In no case were achenes set if the bags were not removed.

Nor were achenes set in "brush-pollinated" flowers in the same head or different heads of the same plant (Table 6). Since

Berlandiera plants are completely self-sterile, emasculation is unnecessary. 105

TAOltbTABLE o6

SEIZING ACHENE-SET DATA

PARENTAGE SEIZING ACHENES PERCENT ATTEMPTS SET ACHENES-SET

Among Different Heads of Sams Plant

SS 16 0 0.0 TT 8 0 0.0 PL 16 0 0.0

Within Hoad of Same Plant

SS 119 0 0.0 PP 118 0 0.0 BB 79 0 0.0 TT 130 0 0.0 LL 626 0 0.0

TL or LT 648 0 0.0 PL or LP 290 0 0.0 ST or TS 115 0 0.0 SL or LS 6l4 0 0.0 SB or BS 15 0 0.0 BP or PB 8 0 0.0 PT or TP 16 0 0.0

PLxPL 56 0 0.0 TSxST 24 0 0.0

LLxPL or PLxLL 231 0 0.0 LLxLT or LTxLL 12 0 0.0 SSxSL or SLxSS 93 0 0.0 SSxST or STxSS 57 0 0.0

LTxSS or SSxLT 165 0 0.0 PLxSS or SSxPL 231 0 0.0 The overall achene-set among all parental stock Investi­ gated (Table 7) la 74.2 percent. B. subacaulls is most fertile;

B. lvrata var. lyrata and B. pumila. the least fertile. All produced vigorous, healthy F^ progeny. In all cases* however,

progeny had both a low fertility (achene-set) and a low pollen stainability. The few plants are slightly less vigorous than F^ progeny, but are more fertile. Also in every case the F^ plant crossed much more easily back to cither parent (ovule parent) than did the parent to the F^ plant

(reciprocal cross) or the F^ plant to its kind. Progeny result­

ing from crosses with B. lyrata var. lvrata as one of the parents had a markedly low fertility and a correspondingly low pollen stainability (Table 10, page 126)* The low fertility may be due, in part, to the meiotic lagging chromosome behavior.

This great sterility may help account for the absence (or near absence) of hybrids between B. texana and B. lyrata var. lyrata in the field.

Other types of sterility occur. Siblings developing from

seeds of a head of a B. texana plant collected in the field are

completely intersterile, yet easily cross with members of other populations of that species. Progeny resulting from a cross between B. subacaulls and B. subacaulls are interfertile.

Progeny from the reciprocal cross are also interfertile. How­

ever, no achenes are set from crosses between the two sets of

progeny ("reciprocal sterility"); hence the low value of 28.4 107

TABLE ?

SUMMARY OF ACHENE-SET DATA FROM ARTIFICIAL CROSSES o v r a T " ' c r o s s i t o ■— i w ~ — HBCTff ' PARENT PARENT ATTEMPTS SET ACHENE-SET

Intra- and Interspecific Crosses

SS SS 24 24 100.0 SS PP k7 44 93.6 SS BB 63 59 93.7 SS TT 6a 47 75.8 SS LL 96.9 &

PP SS 48 34 70.8 PP PP 24 18 75.0 PP BB 53 40 75.5 PP TT 131 65 49.6 PP LL 64.3 $ 62.6

BB SS 64 57 89.1 BB PP 73 65 89.0 BB BB 135 113 83.7 BB TT 145 98 67.6 BB LL 68 70.8 1 $ *+01 TT SS 24 22 91.7 TT PP 132 84 63.6 TT BB 106 79 74.5 TT* TT 225 179 79.6 TT LL 80.0 #

LL SS 31 27 87.1 LL PP 63 33 52.4 LL BB 91 66 72.5 LL TT 67 54 80.6 LL LL 204 135 66.2 456 lit

7 Does not include sibling intersterility. io8

TABLE 7 (Continued) OVULE polisat ■ '■ z m s m — " i t m u —PBB5HF" PARENT PARENT ATTEMPTS SET ACHENE-SET

Qeneration Crosses F1 SS SS 95 27 28.4

8P SF 48 3 6.3

SB SB 52 6 11.5 SB BS 64 13 20.3 BS SB 63 17 27.0 BS BS $ * tH ST ST 71 0 0.0 ST TS 48 5 10.4 TS ST 32 4 12.3 TS TS $ 8 m

SL SL 306 3 1.0 SL LS 114 1 0.9 LS SL 158 2 1.3 LS LS 107 1 ■585 7 ■ft* PP PP 8 5 62.5

PB PB 28 10 35.7 PB BP 30 8 26.7 BP PB 21 5 23.8 BP BP 0 0.0 -# TS 2C5 LL LL 56 0 0.0

LT LT 32 0 0.0 LT TL 40 0 0.0 TL LT 45 0 0.0 TL TL 77 6 T94 "8

LP PL 7 0 0.0 109

OVULE POLLEN CROSSING ACHENES PERCENT PARENT PARENT ATTEMPTS SET ACHENE-SET

Generation

TSxST TSxST 15 20.0 PLxPL PLxPL 16 12.' "IT

Generation Backcroesee

S3 SP or PS 16 12 75.0 PP SP or PS 5 k 80.0 SP or PS SS 15 5 33.3 SP or PS PP 15 8 53.3

SS SB or BS 2k 15 62.5 BB SB or BS kO 30 75.0 SB or BS SS 2k 1 k.2 SB or BS BB 15 6 *f0.0

SS ST or TS k$ 29 6k A TT ST or TS 32 20 62.5 ST or TS - S3 7 k 57.1 ST or TS TT 72 31 *3.1

SS SL or LS 121 37 30.6 LL SL or LS 16 6 37.5 SL or LS SS 121 1 0.8 SL or LS LL 30 k 13.3

PP PT or TP 8 75.0 TT PT or TP PT or TP PP 7 0 0.0 PT or TP TT 7 2 28.6

LL PL or LP 82 22 26.8 PP PL or LP 8 1 12.5 PL or LP LL 76 10 13.2 PL or LP PP 38 2 5.3 LL BL or LB 11 6 5^.5 BB BL or LB 16 9 56.3 BL or LB LL BL or LB BB 8 12.5 110

TABLE 7 (Continued) OVULE POLLEN CROSSING ACHENES PERCENT PARENT PARENT ATTEMPTS SET ACHENE-SET

LL LT or TL 82 25 30.5 TT IT or TL 14 6 42.9 LT or TL LL 16 1 6.3 IT or TL TT 53 2 3.8

Intergeneric Groesee

Berlandiera Silnhium 201 0 Berlandiera Polrnnia 16 0 Berlandiera Engel- mannia 0 olo olo o o olo olo o o "3 • I* • • Ill percent in Table 7* This sterility among closely related off­

spring is unexplained as yet in view of present data* This

sterility increases the significance of obligate outcrossing, beyond mere self-sterility*

These restrictions on fertility increase the significance of introgressive hybridization. It appears that both B. x humilia and B. x betonlclfolia have resulted from introgression*

Neither complex has a complete gene exchange with resulting swamping, nor is there just a "trickle" of genes since many In­

tergrades are found. The value lies somewhere between these extremes* More comment in this regard will be made later in regard to field populations.

Limitations in using achene-set data are evident. Arti­

ficial pollination may have been attempted when the stigmas were no longer receptive or, less likely, when pollen was not mature. There is opportunity for contamination by insects

(insect pollination is suspected because of the oil-laden pollen and the presence of insects about the flowers), particularly when heads are unbagged while making crosses. Fortunately,

contamination in interspecific crosses are detectible because of the different morphologies of the F1 offspring. The exper­

imental plants used represent but a small sample of the total

species gene pool.

Crosses involving three and four species result in off­

spring with usually very low fertility. Those of higher fer­

tilities always have B. subacaulis as one of the parents with but one exception* Several intergeneric crosses among Silghium spp. (n ■ 7) (Fisher and Cruden, 1962), Engelmannia oinnatifida

(n - 9), Polymnia canadensis (n ■ 15) (Wells, 1963), and Ber­ landiera spp* and hybrids (n * 15) were attempted. All were unsuccessful. GERMINATION AND SURVIVAL

In an effort to learn whether germination or survival of young plants may be factors in delimiting taxa, achenes obtained from the various crosses were placed on wetted filter papers in

Petri plates and stored at room temperatures. Germination occurs from two to thirty days. When the roots were at least one-quarter inch long, the fruit walls and seed coats were re­ moved from the young embryos, exposing the paired cotyledons.

Transfer was made to four-inch clay pots filled with moistened mixture of three parts sand and one part loam. Petri plate lids were loosely arranged above the young plants to maintain high humidity. Plants were later transferred to six-inch pots.

The plant cycle from seed to seed may take as little as 157 days.

Both germination percentages and one-year survival percent­ ages (Table 6) are subequal for all taxa with one noteworthy exception. Germination percentages of the "texana-betonicifolla- pumila" complex are lower than those of other Berlandiera taxa.

Germination of field-collected achenes under treatment described above is favorably comparable to that of achenes collected from greenhouse-cultured plants (Table 9)•

113 114

TABLE 8

GERMINATION AND ONE-YSAR SURVIVAL RECORDS OF ARTIFICIAL CROSSES

Taxa Fruits Fruits Parcant Offspring Parcant Sat Garai- Garai* Surviving Surviving a»tlng natlnx Ganaration

LL x SS 16 14 87.50 14 100.00 LL x PP 11 5 45.45 * 4 80.00 LL x BB 15 9 60.00 8 88.89 LL x TT 44 36 8l.8l 22 61.11 LL x LL 43 42 97.84 23 54.76

TT x SS 8 8 100.00 8 100.00 TT x PP 27 21 77.78 17 80.95 TT x BB 11 3 27.27 2 66.67 TT x TT 46 10 21.74 7 70.00 TT x LL 37 36 97.30 24 66.67

BB x SS 7 6 85.71 5 83.33 BB x PP 21 11 52.38 8 72.72 BB x BB .------BB x TT 17 10 58.82 9 90.00 BB x LL 17 11 64.71 4 36.36

PP x SS 17 12 70.59 12 100.00 PP x PP 12 9 75.00 5 55.56 PP x BB 14 7 50.00 7 100.00 PP x TT 11 4 36.36 2 50.00 PP x LL 15 15 100.00 9 60.00

SS x SS 16 15 93.75 13 86.67 SS x PP 23 17 73.91 14 82.35 SS x BB 8 7 87.50 7 100.00 SS x TT 9 7 77.78 7 100.00 SS x LL 16 16 100.00 14 87.50 F2 Ganaration TS x ST 4 4 2 100.00 3 75.00 ST x TS 5 5 100.00 3 60.00 ST x ST 4 2 50.00 2 100.00 LS x LS 2 0 00.00 ------LP x LP 17 14 82.32 7 50.00 115

TABLE 8 (Continued)

Tax* Fruits Fruits Percent Offspring Percent Sst Germi- Germi­ Surviving Surviving natinx nating Backcrosses

LL x LP 19 18 94.74 8 44.44 LP x LL 8 8 100.00 3 37.50 LL x LT 4 4 100.00 1 25.00 LT x LL 4 3 75.00 2 66.67 LS x LL 4 4 100.00 2 50.00 SL x SS 1 1 100.00 1 100.00 SS x SL 14 12 85.71 11 91.67 SS x ST 8 6 75.00 5 83.33 ST x SS 8 7 87.50 7 100.00 iThree-species Crosses SS x LP 15 15 100.00 15 100.00 LP x SS 2 2 100.00 0 00.00 SS x LT 4 4 100.00 1 25.00 ST x SL 2 2 100.00 1 50.00 LP x LT 3 2 66.67 0 00.00 LT x LP 2 2 100.00 0 00.00 Summary Crosses

LL x Others 129 106 82.64 71 66.98 TT x Others 129 78 64.16 58 74.36 BB x Others 62 38 61.29 26 68.42 PP x Others 69 4? 68.12 35 74.47 SS x Others 72 62 86.11 55 88.71

F^ Crosses 461 331 71.80 245 74.02

Crosses 32 25 78.13 15 60.00

Backcrosses 70 63 90.00 40 63.49

Three-species Crosses 36 35 97.22 20 57.14 116

TABLE 9

GERMINATION DATA OF FIELD-COLLECTED FRUITS OF EASTERN BER1ANPIERA ______TAXA______

Population Fruits Consi­ Fruits Percent Number dered Viable Germinating Germinating

Hybrid Populations 1197 10 9 90.00 1199 15 11 73.33 1202 21 11 52.38 1203 29 26 89.66 1205 32 26 81.25 1206 30 25 83.33 1208 16 Ik 87.50 1210 3 2 66.67

Pumila Populations 12^6 9 9 100.00 1262 5 k 80.00 126*t 20 20 100.00 129^ k k 100.00

Total (12) 19k 161 82.99

Neither germination for first-year survival appears to be significant as barriers to crossability among taxa, including hybrids^ unless field conditions are much different from the experimental conditions. MORPHOLOGICAL ANALYSIS 07 PARENTAL EXPERIMENTAL PLANTS

Analysis of morphology of living plants used in ths

brssding program is mads according to msthods dsscribsd below

bassd primarily upon distinct!vs eharactsrs ussd in original

dsscriptions and in ksys. All plants ars cultursd undsr Ths

Ohio Stats Univsrsity grssnhouss conditions. To promots

continuous flowering, thsss plants ars exposed year-around

to a minimum of twslvs hours of light psr day. Thsy ars pottsd in a mixture of three parts sand to one part loam.

Liquid fertiliser and water are supplied regularly. Measure­ ments are made from living plants where possible. The plants

listed in Table 10 are arranged according to taxa and place

of collection. Growing conditions and measurement methods also apply to the hybrid experimental plants*

Pollen etainabillty. Pollen is obtained by scraping sur­

faces of anthers and protruding style of disc flowers at anthe-

sis, under bagged conditions only. A pollen grain is considered

stainable, and presumably viable, if at least some portion of

the content stains deep blue with the addition of analine blue.

This stain is prepared from one gram of analine blue crystals,

25 grams of phenol crystals, 25 ml each of glycerine, lactic acid and distilled water. Pollen counts are based upon a minimum of 500 grains each. 117 Stem measurements. The height of the plant, is that distance from 0.5 cm above the base of the stem (to eliminate juvenile rosette) to the base of the uppermost head of the uppermost branch* Percentage of leaves along the upper half of stem is based on the total number of leaves on portion of stem used for height measurement just described* Peduncle length is measured from the uppermost non-scale-like leaf to the uppermost head of the uppermost branch. Internode length is that distance from the middle cauline leaf (if two middle leaves, the lower one is used) petiole attachment to the one immediately below.

Leaf measurements. A leaf is considered to be lobed if a sinus extends to at least half the distance to the midrib* All intact leaves are counted in calculating the percentage value of leaves lobed* The remaining measurements are based on values obtained from the middle three cauline leaves; or if there is an even number of leaves, then based on the two middle leaves and the one immediately below. The number of lobes (average) includes those on the left side of leaf only; the blade-width

(actually one-half width) measurement is also only from the left side of the leaf. Length of blade is measured from apex to level of lowest serration, cremation, etc., either side; or if blade is cordate-based, then measurement is made to the junction of lowest major vein to the midrib. Petiole length is the remaining length of the leaf to the lowest point of attach­ ment. Terminal lobe extends from apex to the lowest serration, 119 crenation, etc., of the longest unlobed terminal portion of the leaf, either aide. Blade length equals terminal lobe length in unlobed leaves. Percentage values are obtained by dividing sum values of blade width, petiole length, and terminal lobe length by sum value of total blade length. Figure 24.

Head width. This value is the maximum width across top of head from phyllary tip to phyllary tip without applying pressure.

Flower colors. Although there is a dilution effect in some plants, the disc corolla is either yellow or red to maroon; the ray veins beneath, either green or red to maroon (Figure 23)•

Ray measurements. Average maximum lengths and widths are from all rays of uppermost head of uppermost branch. The length of the ray is measured from apex to short tubular portion atop the achene. Notch length varied widely, even on the same head and was subsequently abandoned.

Pubescence (vesture). All pubescence is of multicellular, jointed hairs varying in length, width, density, and base form.

This character is subjectively evaluated and assigned scaled values as listed below. For comparisons, see Figures 23 and 26.

Leaves Peduncles

3. Scabrous 3* Scabrous or subscabrous 4. Subscabrous 4. Dense, erect, long hirsute 3. Hirsute 3* Sparse, spreading, long 2. Finely hirsute hirsute 1. Velvety 2. Spreading or curling, long, fine 1. Hatted, dense, long, fine 120

Figure Zk. Leaf shapes of parental and hybrid plants. Figure 25. Pubescence Types I.Peduncle vesture. Top row from left: SS 1235 SP 170-1i PP 550; PP 590. Bottom row from left: TP 197-2; HB 6?6; TT 708. Scale: 1 mm: ca. 10 microns. All peduncles hairs are from dried specimens and are mounted directly in balsam. 122

t

t > %

Figure 26* Pubescence Types II. Top row, from left: TT 903, peduncle; LL 750, peduncle. Bottom: Specialized hairs from along anthers, TT 901. Scale: of peduncle hairs, 1 mm: ca. 10 microns; of specialized hairs, 1 mm: 1.3 microns. 123 Berlandiera taxa aa described earlier were distinguished in part on basis of these measurements and will not now receive further comment. See Figures 27-32. 455

i 1 Figure 2?. j*. lyrata Figure 28. B, texana 1002. ' Figure 29* B. x beto«*ci- var. lyrata ToiO. Scale: 0.26x7 folia 655* Scales 0.32x. Scale: 0.233c. 657 525 Figure 30. B. x betonicifolia Figure 31. B. pumila 657- Scale: 0,lf2x. 525. Scale: 0.37z. 126

TABLE 10

M0BPH0L0QICAL ANALYSIS OF PARENTAL EXPERIMENTAL PLANTS • TmrTiiwrwnar TSL7- ~ w . u#pfii m u ■TJ5T NUMBER 8TAIN8 COROLLA VEIN cm LEAVES LOBED LOBE£ porcont COLOR COLOR porcont norcont SUB •1611 99.$ follow drttn — ------SUB •1013 99.7 Yallow Orton — ooowm v -- SUB 1014 9a .3 Yollow Orton 22 20.00 100.00 2.5 SUB 1025 98.8 Yollow Orton 38 56.25 100.00 1.0 FUK-SUB 1205 99.8 Rod Oroon 30 33.33 30.77 0.7 PUM 1199 99.6 Rod Orton 21 25.00 0.00 0.0 PUM 526 84.9 Rod Oroon 42 36.36 0.00 0.0 PUM 525 98.9 Rod Oroon 39 42.86 0.00 0.0 PUM 507 98.3 Rod Oroon 23 22.22 0.00 0.0 PUM 506 98.8 Rod Oroon 18 46.15 0.00 0.0 PUM *595-1 98.1 Rod Oroon 27 44.44 0.00 0.0 PUM *595-2 89.0 Rod Oroon 32 55.56 0.00 0.0 PUM-BET 616 99.8 Rod Oroon 41 57.14 0.00 0.0 BET 641 98.9 Rod Oroon 43 65.52 0.00 0.0 BET 642 98.3 Rod Oroon 36 31.25 0.00 0.0 BET 655 99.5 Rod Oroon 18 64.29 0.00 0.0 BET 656 98.2 Rod Oroon 19 70.59 0.00 0.0 BET-TEX 657 98.7 Rod Oroon 16 46.67 0.00 0.0 BET-TEX 680 99.7 Rod Oroon 24 64.71 0.00 0.0 TEX 710 96.5 Rod Oroon 40 76.92 0.00 0.0 TBX 729 99.1 Rod Rod 28 61.11 0.00 0.0 TEX 730 98.9 Rod Oroon 48 83.78 0.00 0.0 TEX 743 98.8 Rod Oroon 46 71.43 0.00 0.0 TEX 900 98.5 Rod Oroon 26 66.67 0.00 0.0 TEX 901 99.8 Rod Oroon 27 24.00 0.00 0.0 TEX •1001 99.6 Rod Oroon 24 54.55 0.00 0.0 TEX 1002 99.8 Rod Oroon 37 60.00 0.00 0.0 TEX 1004 99.6 Rod Oroon 32 71.43 0.00 0.0 LYR 746 96.6 Rod Rod 64 38.89 100.00 2.7 89.4 LYR 751 Rod Rod 71+ 23.08 90.91 2.0 LYR 799-1 99.6 Rod Rod 26+ 16.67 100.00 4,0 LYR 799-2 99.8 Rod Rod 54 60.00 100.00 1.3 LYR 821 99.5 Rod Rod 67 64.71 100.00 2.7 LYR 838 99.8 Rod Rod 86 62.50 100.00 5.3 LYR 851 98.7 Rod Rod 51 75.00 73.33 0.3 LYR 920 99.5 Rod Rod -- ..... — LYR 921 98.9 Rod Rod 129 37.84 100.00 4.0 LYR 922 99.8 Rod Rod 116 50.00 100.00 3.3 LYR 923 99.8 Rod Rod 45 28.57 89.47 3.7 LYR 1009 98.7 Rod Rod 50 50.00 81.82 2.0 LYR 1010 92.5 Rod Rod 42 53.85 87.50 2.0 * Data i’rora dried voucher spocimens. 127

TA.BUS 10 (Continued) turnm . beab"m m - m m —m —bumt tolpr ism. NUMBER L WIDTH NODE L W W/L L W LOBE cm cm L cm ma mm percent cm cm L cm "T3TI— ~ — — — —

1014 10.4 1.9 0.2 12.6 7.8 61.90 16.8 3.9 11.6 1025 5.3 1.4 2.3 9.0 5.6 62.22 20.7 3.8 13*7 1205 1.6 1.4 0.7 16.0 8.0 50.00 17.4 4.7 16.7 1199 2.7 1.8 0.9 16.4 8.0 48.78 16.8 3.8 16.8 526 1.8 1.8 0.3 20.2 9.0 44.55 16.7 5.5 16.7 525 0.8 1.8 2.6 17.4 7.9 45.40 18.4 5.3 18.4 507 6.3 1.5 1.6 15.5 8.3 53.55 13.1 5.2 13.1 506 1.4 1.4 2.8 12.1 6.5 53.72 10.7 4.4 10.7 595-1 ... 1.5 ...... 18.5 5.4 18.5 595-2 ...... 2.8 ...... 18.6 6.3 18.6 616 1.7 1.5 2.1 14.1 7.4 52.48 14.0 5.5 14.0 641 1.0 1.7 0.9 15.4 7.7 50.00 10.3 4.1 10.3 642 1.9 1.7 2.0 15.1 5.4 35.80 16.5 4.0 16.5 655 1.2 1.5 1.3 17.1 9.0 52.63 7.6 2.5 7.6 656 0.7 1.2 0.8 11.0 6.9 62.73 4.3 2.6 4.3 657 2.1 1.8 1.2 16.4 6.6 40.24 12.5 4.6 12.5 680 ... 1.5 1.3 9.2 5.2 56.56 710 4.1 1.9 1.6 16.9 9.6 56.80 16.7 5.4 16.7 729 1.8 2.3 2.5 15.4 8.7 56.49 17.5 5.1 17.5 730 0.9 2.0 0.7 15.0 6.9 46.00 11.2 3.1 11.2 743 2.0 1.7 1.4 14.4 6.4 44.44 20.4 5.0 20.4 900 1.1 2.0 2.3 16.4 9.0 54.88 25.0 6.5 25.0 901 2.0 1.5 0.4 13.3 7.2 54.14 12.7 3.4 12.7 1001 1.0 ... 3.0 ...... ------15.7 4.3 15.7 1002 2.8 2.4 3.7 13.3 8.5 63.91 31.0 7.8 31.0 1004 0.6 2.4 1.8 10.7 6.8 63.55 20.4 4.1 20.4 746 3.7 1.7 1.9 10.4 5.9 56.73 28.7 5.3 14.5 751 16.0 1.3 0.4 10.9 5.7 52.29 21.8 4.1 15.4 799-1 3.7 1.8 0.2 15.0 7.7 51.33 29.9 5.2 16.9 799-2 5.8 1.5 4.7 10.7 5.5 56.40 14.6 4.2 10.8 821 2.7 1.6 4.2 12.3 6.5 52.85 19.3 3.2 12.5 838 4.2 1.7 4.2 13.0 5.8 44.62 16.8 4.5 4.6 851 4.1 1.7 0.8 13.8 6.8 59.28 10.7 2.8 9.9 920 ...... ------921 7.7 1.7 1.2 10.0 6.0 60.00 23.7 3.6 6.3 922 8.2 1.4 4.2 11.5 6.7 58.26 17.4 4.3 8.4 923 6.6 1.5 1.9 13.8 6.9 50.00 16.9 3.0 7.8 1009 6.4 1.7 8.2 11.0 6.0 54.54 25.2 7.3 15.1 1010 6.3 1.3 8.7 14.0 7.9 56.43 18.7 5.0 11.7 128

TABU 10 (Contiauod) “TO T * m : — szzra—v r n n m — m r iy v b b w bb— NUMBER L W/L I/BLADE L BLADE L FED. LEAF on porcont porcont porcont (ABOVE) T B i r —— ------— 1013 1014 ” 4 23.21 69.05 58.10 5 5 1025 5.6 18.36 66.18 17.59 5- 5 1205 6.3 27.10 95.98 56.21 1+ 1 1199 4.1 22.62 100.00 24.40 1+ 1 526 10.2 52.95 100.00 61.08 1 1 525 4.3 28.80 100.00 25.57 U 1 507 5.2 59.69 100.00 24.43 14- 1 506 4.6 40.19 100.00 42.99 14- 1 595-1 5.8 29.19 100.00 20.54 1 1 595-2 4.6 55.87 100.00 24.73 1 1 616 2.0 59.28 100.00 14.29 1 2 641 2.2 59.81 100.00 21.36 2 3- 642 1.8 24.24 100.00 10.91 2 3- 655 1.2 52.89 100.00 15.79 2 3- 656 0.5 60.47 100.00 11.63 3- 3 657 2.5 36.80 100.00 20.00 3 3+ 680 > It 710 0.8 52.34 100.00 4.79 4 5+ 729 1.9 29.14 100.00 10.86 4 3* 730 0.4 27.68 100.00 5.57 44* 4 743 0.8 24.51 100.00 5.92 4 3+ 900 2.5 26.00 100.00 10.00 4 4 901 1.9 26.77 100.00 14.96 4 4 1001 5.6 27.39 100.00 22.92 44- 4 1002 5.1 25.16 100.00 10.00 44. 4 1004 1.2 20.10 100.00 5.88 4+ 4 746 14.0 27.18 50.52 48.78 5 1 751 15.2 18 .81 70.64 60.55 5 1 799-1 4.7 17.59 56.52 15.72 5 1 799-2 7.5 28.77 75.97 51.37 5 1 821 4.0 16.58 64.77 20.73 5 1 838 1.8 26.78 27.58 10.71 5 1 851 6.6 26.17 92.52 61.68 5 1 1 920 3c X 921 4?8 15.17 26.58 20.25 5 1 922 1.9 24.71 48.28 10.92 5 1 923 5.8 17.75 46.15 22.49 5 1 1009 8.0 28.97 59.92 31.75 5 1 1010 8.8 26.74 62.57 47.06 5 1 KEY TO AND ANALYSIS OF HYBRID EXPERIMENTAL PLANTS

First-generation hybrids bar* been obtained froa all eoabiaatloas of tho four species* Progeny from rooiprooal oroaaoa art similarf la tho koy aad doseriptloast aeroly for convenience, hybrids art listod with tho wostoraaost taxa first ia tho formulae* Tho koy bolow is supportod by subsoquoat doooriptioas, photographs of living plaats (Figures 33-4-7) aad tabulated morphology data (Tablo 11)* Tho koy is basod 00 hybrids obtaiaod froa paroats without (as much as possible) iatergradiag characters*

1 Rays with veins boaoath rod to aarooa • •..... 2

2 Middle cauliae leaves with 5 or aoro lateral lobes, tho toraiaal lobe loos thaa one-half blade loagth ••••• (A) B* lyrata var• lyrata z aubacaulis

2 Middle cauliae loaves with 0-3 lateral loboe, tho toraiaal lobe aoro thaa three-quarters blade loagth ,...... 3

3 Loaves velvety •••• (B) B* lyrata var* lyrata x pualla

3 Leaves hirsute • ••• (C) B* lyrata v a r . lyrata x toxaaa

1 Rays with veins beneath green ...... 4

4 Blades unlobedf leaves evenly distributed{ peduncle with long fine hairs but not m a t t e d ...... (D) B. texana x puaila

4 Some blades lobed (or if unlobed, not with both long fine hair on the peduncle and even distribution of leaves) ...... 3

129 130

3 Peduncles hirsute to sub- scabrous (E) B. texana x subacaulls

5 Peduncles with long fine hairs (F) B. pumlla x subacaulls

(A) Berlandiera lyrata var. lyrata x subacaulls

Herbs with, few to several sprawling leafy stems 10-30 cm high| with many heads. Persisting basal rosette. Leaves all lobed (except sometimes uppermost and/or lowermost); terminal lobe and petiole length reduced by Increase of lateral lobest greater In average number than In either parent. Leaf pub­ escence Intermediate of parents. Heads 1.7-2.0 cm wide. Disc corollas and ray veins beneath red to maroon. Figure 34.

(B) Berlandiera lgrata var. lyrata x texana

Herbs with one to few erect stems, 24-85 cm high. Blades of middle cauline leaves either unlobed or having 1-3 lateral lobes; hirsute above, Intermediate of parental types. Heads

1.4-2.2 cm wide. Disc corollas and ray veins beneath red to maroon. Figure 37.

(C) Berlandiera lyrata var. lyrata x pumlla

Herbs with one to few erect stems, 24-48 cm high. Blades of middle cauline leaves unlobed or having 1-3 lateral lobes, velvety. Heads small, 1.2-1.6 cm wide. Disc corollas and ray veins beneath red to maroon. Figure 38 .

(D) Berlandiera texana x pumlla

Herbs with one to few erect stems 28-72 cm high. Leaves unlobed, finely hirsute above. Peduncle with long fine spreading 5*5 IM

Figure 33* B. lyrata Figure Jk, B. lyrata Figure 35* B. suba- var* lyrata 35-^. Scale: var* lyrata x subacaulls caulis 11-2* Scale: O.^lx. 0 .26x. 5-5 * Scale: 0 .32x« 4 0 4

Figure 36. JJ. texana Figure 37- J3. lyrata Figure 3d. B. lyrata 660-1. Scale: 0.?3x. var. lyrata x texana fro-2. var. lyrata x pumila 2^5-1 . Scale: 0.23x. Scale: 0.23x.

V* ru 133 or curling hairs* Hoads l.Jf-1.8 cm wide* Disc corollas red to maroon} ray veins beneath green* Other discussion appears under description of B. x betgnicifgilia, p. 66* Figure 39*

(E) Berlandiera texana x subacaulis

Suffrutescent herbs with one to few leafy erect stems, 19-

39 cm high. Middle cauline leaves unlobed or having 1-3 lateral lobes, hirsute to subscabrous above. Heads large, 2.1-2.6 cm wide. Disc corollas red to maroon. Ray veins beneath green.

Figure kO.

(F) Berlandiera pumila x subacaulis

This taxon is described on page *f3 as B. x humilis. These plants duplicate holotype of B. humilis Small. Figure *tl.

In Tables 10 and 11 are presented morphology data for comparison of F^, F^, and backcross progeny.

In all cases red to maroon color was dominant to green in ray veins and to yellow of disc corollas. Other characters used are apparently quantitatively inherited. Never did lobed- leaved offspring result from any crossing combination of the

"texana-pumila" complex. In more than 90 percent of crosses observed, crosses of unlobed-leaved taxa to lobed-leaved taxa

(and reciprocally) result in offspring with at least some leaves that are lobed. Most of the exceptions (22 of Zk) occur when B. lyrata var. lyrata is a parent. In the hybrids, the number of lobes is usually but two or three at the blade base, the remaining portion of the blade accounting for 80-95 percent IfM J l

Figure 39. B. texana x Figure 40. B. texana Figure klm B. puaila puaiila 197-3. Scale: 0.19x. x subacaulis 2-1 •”* Scale: x subacaulis 1 7 0 - ? . Scale: 0.23x1 0.23x.

vx -p- IM i m i Figure 1*2. B. lyrata Figure 1*3. texana Figure kb, B. x beton­ var. lyrata x IJ. x beton- x B. x betonicifolia 176-1. icifolia x B. pmiiTi» 21*2-1. Iclfolia 118-1 . Scale: Scale: 0.21x. Scale: 0.23x. 0 .21x. M VX NJI JH

Figure 45. B. x beton- Figure 46. F2 off­ Figure 47. F_ offspring ici folia x 13. subacaulis spring of B. lyrata var. of B. subacaulis x texana 144-1. ScaTe: O.zSx. lyrata x pumila 31-1 - 469-1. Scale: 0.23x. VJ4cr\ Scale: 0 .233c. of the blade length as compared to 20-75 percent in lobed-leaved parental plants. There is an additive effect in number of lateral

lobes in offspring from crosses of the two lobed-leaved taxa,

B. subacaulis and B. lyrata var. lyrata. For comparisons of leaf

shape, see Figure 24, page 120.

In general, the most variable characters within a taxon are

those of the stem; namely internode length, total height of plant, and peduncle length. Hence, stem characters are not good key characters. Head width, petiole length, pubescence types and the percentage of leaves in the upper half of the stem are

better characters to distinguish among taxa because character

ranges of variability are such that no overlap or but slight

overlap occurs between sympatric taxa. 138

TABLE 11

MORPHOLOGICAL ANALYSIS OF ARTIFICIAL HYBRID PLANTS ------m m '6 vpl5~rm - 1 ------CROSS TAXON PARENT PARENT VIDU- STAINS COROLLA VEIN HEIGHT ALS percent COLOR COLOR cm K Generation 10 S3 X SS 1013 1014 6 83.2rll.5 Yellow Green 18 £ 3.7 11 SS X SS 1014 1013 6 78.5-13.5 Yellow Green 17 £4.1

170 SS X PP 1014 507 1 96*4--— Red Green 21 ---- 229 SS X PP 1025 526 4 98.9 .-1.1 Red Green 28 ;6.7 230 PP X SS 526 1025 3 87.4tl0.9 Red Green 16 £4.6

351 PP X PP 526 507 2 99.2 £1.2 Red Green 35 78.5 352 PP X PP 507 526 2 Red Green 29 £0.0

156 BB X PP 641 595-1 3 98.8 £l.O Red Green 49 -3.5 171 PP X BB 595-1 641 1 99.6 -— Red Green 6 7 --- 172 BB X PP 641 595-1 2 99.3 -0.5 Red Green 59-12.7 196 BB X PP 657 507 1 99.6 -— Red Green 30 7 — zkz PP X BB 595-2 680 5 99.7 ^0.4 Red Green 40 £5.9

144 SS X BB 1014 642 7 91.4 £7.8 Red Green 29 £7.4 145 BB X SS 642 1014 5 97.6 £0.9 Red Green 35 i2.3

1 TT X SS 1001 1014 8 79.6J10.0 Red Green 29£l0.3 2 SS X TT 1014 1011 7 81.5-12.9 Red Green 29 -1.7

194 PP X TT 526 1002 1 99.6.--- Red Green 37.--- 197 TT X PP 710 526 2 6l.?£30.5 Red Green 50£21.2

176 TT X BB 743 641 1 9 9 . 0 --- Red Green 5 2 ---

659 TT X TT 730 1002 3 99.6 £0.3 Red Green 35 £9.3 660 TT X TT 1002 730 1 9 9 . 5 --- Red Green 3 0 ---

5 SS X LL 1014 1009 6 26.3£j0.3 Red Red 17 73.8 6 LL X SS 1009 1014 7 25.3 j7.4 Red Red 16 £2.7 7 LL X SS 1010 1011 6 17.1 76.8 Red Red 23 7 4 .O 8 SS X LL 1011 1010 8 10.9 £3.8 Red Red 20 £2.7

255 PP X LL 525 1010 3 76.7£31.3 Red Red 40 £6.7 390 PP X LL 507 1010 2 ...... Red Red 31 £0.0 624 PP X LL 526 1010 1 49.4 -— Red Red 30 7-- 651 LL X PP 851 526 2 98.9 -0.3 Red Red 28 £4.2

118 LL X BB 1009 642 1 9 1 . 1 --- Red Red 4 3 --- TABLE 11 (Continued) ------w u w 'w g T B g - m m im- — m z ------CROSS TAXON PARENT PARENT VIDU- STAINS COROLLA VEIN HEIGHT ALB percent COLOR COLOR cn

9 LL X TT 1010 1004 3 35.8 ±8.7 Red Red 32 ±5.0 22 LL X TT 1010 1004 2 40.5 r5.2 Red Red 28 ±0.0 29 LL X TT 1010 1001 6 35.8 ±6.3 Red Red 29 *3.6 37 TT X LL 1001 1010 4 27.9 r2.0 Red Red 27 *3.4 38 LL X TT 1010 1001 2 29.7 rl.9 Red Red 29 *0.0 40 TT X LL 1002 1010 5 51.9.-1.6 Red Red 27 *3.3 41 LL X TT 1010 1002 5 49.5*17.1 Red Red 27 *2.2 240 TT X LL 900 1009 4 78.8-10.1 Red Red 43 *9.2 241 LL X TT 1009 900 1 8 6 . 4 --- Red Red 26 --- 577 TT X LL 1002 920 2 33.2* 21.1 Red Red 38 ± 2.8

15 LL X LL , 1010 1009 1 76.5 z— Red Red 28 ---- 35 LL X LL 1010 1009 5 96.2-3.3 Red Red 22 ±8.7 36 LL X LL 1009 1010 4 88.3*10.4 Red Red 28 ±8.2 179 LL X LL 923 1010 3 Red Red 41 ±7.6

Generation

355 TS X ST 1-3 2-1 3 76.4±l8.7 Both Green 29±12.5 356 ST X TS 2-1 1-3 2 84.5 *7.4 Both Green 34 *0.7 469 ST X ST 2-5 2-3 2 98.3 *0.5 Red Green 22 ± 9.2

31 PL X PL 1021 1022 1 7 3 . 0 --- Red Red 24 ---- 32 PL X PL 1022 1021 2 49.3*29.6 Red Red 18 ±1.4

Backcrosses

366 ST X SS 2-1 10-4 7 a Both Green 30 ±8.0 488 SS X ST 11-1 2-1 5 94.0-10.3 Both Green 23 *5.2

472 LS X LL 6-8 1009 2 Red Red 15 *0.7

253 SS X SL 10-4 5-6 4 94.4-4.0 Both Both 25 *7.8 387 SS X SL 10-4 5-6 6 84.1±15.2 Both Both 22 ±8.4 388 SL X SS 5-6 10-4 1 9 8 . 5 --- Red Red 1 6 ---

3 LL X PL 1010 1021 2 18.6 -3.7 Red Red 20 ±3.5 18 LL X PL 1009 1018 1 34.7 .--- Red Red 29 T— 19 PL X LL 1018 1009 3 85.5*15.4 Red Red 28 ±1.0 33 LL X PL 1010 1021 3 49.6-18.2 Red Red 22 ±3.5 39 LL X PL 1009 1022 1 9 6 . 7 --- Red Red 2 4 ---

25 LL X LT 1009 1015 1 7 4 . 9 --- Red Red 3 2 --- 26 LT X LL 1015 1009 1 4 1 . 2 --- Red Red 2 3 --- 120 LT X LL 1015 1009 1 Red Red 2 9 ---- TABLE 11 (Continued)

UPPER LOBED LOBE PEDUNCLE HEAD INTERNODE CROSS LEAVES LEAVES NUMBER LENGTH WIDTH LENGTH PTctnt percent ca cm cm Generation

10 20.0±13.* 95.* ±5.8 *•0*Jl.8 5.9±2.8 1.8±0.3 1 .5±1.2 11 20.i±i3 .* 100.0 ±0.0 3.3--0.5 5.0±3.* 1.7±0.3 0 .7±0.8

170 33.3 r— 8 1 . 8 --- 1.7 2 . 8 -- 1 .8—- 2.1 — 229 *0.8 ±*.2 *6.0±31.* 0.9 ±1.1 3.*±1.* 2 .0±0.2 3.6±2.5 230 39.3 -6.5 60.9-28.8 1.5 ±0.7 1 .8±0.8 1.7±0.1 3 .2±o.*

351 **.3x22.2 0.0 ±0.0 0.0 ±0.0 1.2±0.9 1 .6± 0.2 *.3±0.1 352 53.9-10.2 0.0 -0.0 0.0 -0.0 0 .9±0.0 1 .5±0.l 3.3±9.*

156 57.* -6.7 0.0 ±0.0 0.0 ±0.0 1.0±0.5 1.5-1.5- 3 .1±0.1 171 60.7 z— 0.0 ---- 0.0 2 ,2—- 2 . 0 -- 1.9.— 172 57 .* -0.6 0.0 -0.0 0.0 ±0.0 1.0±0.9 1.6±0.3 2 .5±0 .* 196 *7.1 r— 0.0 ---- 0.0 2 . 6 -- 1.5+ — 3.*— - 2*2 60.1 -2.9 0.0 -0.0 0.0 ±0.0 l.*±0.7 1 .6±0 ,* 2.7±0.7

1** 39.3 ±9-9 32.2±l*.l 0.7 ±0 .* 3 .8± 1.8 1 .8±0.2 2.8±0.7 1*5 *6.3 -8.7 25.9-19.0 0.3 ±0 .* 3.1±0.9 1 .8± 0.2 2.7±0.9

1 38.2 ±*.2 69.7J 21.* 0.8 ±0.3 3 .*±1.0 2 .3±0.2 3.3±1.1 2 *5.2 -7.7 70 .8-28.1 0.9 ±0.2 2 .6±0.6 2 .3± 0.2 3.*±1.1

19* *7.* 1-- 0.0 ---- 0.0 0.9+— l.*+--- 2.2 — 197 61.8 ±2.6 0.0 ±0.0 0.0 ±0.0 2 .0±0.6 1 .7±0.1 3 .2± 1.6

176 6 2 . 5 --- 0 , 0 --- 0.0 --- 0 . 7 -- 1 .* -- 1.8 —

659 71.0 ±7.1 0.0 ±0.0 0.0 ±0.0 1.1±0.3 1 .9-0.1 1.1±0.7 660 70 . 0 - 0 . 0 --- 0.0 1 . 0 -- 1.9 — 1 . 7 --

5 2*.0 ±*.* 9*.9 ±2.7 7.0 ±1.9 5.2±1,9 5.2±1.* 2 .0±0,1 6 23.5 ±3.7 96.5 ±5.1 5.* ±2.3 *.8±1.8 1 .9±0.2 1 .1±0.8 7 33.*±1*.0 97.9 ±3.5 *.8 ± 1.2 *.3±2.1 1 .7±0.2 2.5±2.5 8 30 .2-11.1 98.6 ±2.6 5.6 ±1.7 3.7±1.7 1.7±0.3 2.6±1.9

255 *0.8 ±7.6 *9.9±**.0 0.7 ±0 .* l.*±0.3 1.5±o.l *.8±0.9 390 *2.3 -0.8 13.1 ±5.1 0.2 ±0.2 1 .8±0.2 1 .3±0.1 *.*±0.6 62* 30.7 T-- 5 2 . * --- 1.0 3.7+-- 1 .*— 2 . 6 -- 651 37.5 ±5.9 6l.8± 5*.0 1 .* ±0.5 1 .2±0.6 l.*±0.0 3 .*±2.0

118 2 7 . 8 --- * . 6 --- 0.0 .... 2 . 3 -- 2 . 0 -- 2 . 5 -- TABLE 11 (Contiautd) Q $ 3 SB. o A 55 3 as° 338 b © a 2 * «Q s - „ O 09 4» m o 2 • d a •

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? + M o 14 iHO (VJ O M ( • • Ov •g tv o 11 T 11 1 +1+1 +1 +1+1 +T +1+1 + + + 1+1+i +1 + 1 (Vi O0 m CO 00 -4- Cv -4- -4- Cv Ov (M-4 (vjao v o m m-4- m r m m irv o h i l cv cv •• • • • • • • • • + l - - V r-l r-i r-l r-ir-l (Vir-i(VJ I r-l H r-i Ol rvj CO O ( vj i o +i +i +? + + 1 j -4 -4 -4 vO tv vOv Ov r-l (VJ m m m lTv-4 m r-l r-i h h ov ov h - • • • • • (vi m m v rj I

+

rl rl 0 C O m o m (i v | J J | i cv (vi i ojm (vj » 1 •1 • 1 CV (VJ l + l 1CM O 1Ov O • • • 1 (vji 1 m t 1CvvO h • • • • • +1 +1 + 1 + ♦ ♦ 1+1 « H H Q r-l Q h i • + + mm (vi m evi m (vi h Ov ov H i i •

cv rl O o m * * * • i 1 i i 1 (vj 1 1 1 1 1 • • 5 m (vj m 5 O(VI O H cvj m m-4 o o v m o o o m o o m mvo m o m i i i i i i i I I I I l l i I i I i I t I I I I I I I I i i i i i i i i i i i i i • • • • •• • 4 - \o \ r-i « I h • •

142 TABU! 11 (Contlimed)

RAY RAY HAY WIDTH/ BLADE BLADE CROSS LENGTH WIDTH LENGTH LENGTH WIDTH mm mm v«rc«nt cm cm F 1 Aentration 10 10.3x1*4 6.1;0.7 59.2 i7.2 21.7x4.8 3 .8J0.6 11 11.0-2.3 6 .011.0 54.5-10.2 21.9-3.8 3.9-0.7

170 10.9.— 6 . 5 -- 59.6 -— 24.6.-- 6 . 2 -- 229 17.5x3.0 8.5x0.6 49.5 x7*2 20.5x5.5 4.7x6.9 230 17.213.2 8 .110.6 48.0 i6.0 l?.6i4.9 3.8io.9

351 14.5x5.2 6 .3x0.5 45.5il3.0 13.3x4.2 5 .oil.8 352 13.5-0.2 6 .4-0.6 47.4 I 5.5 13 .612.3 4.7-0.6

156 13.2-2.0 6 .6-0.4 50.otll.2 15.2io.9 6 .210.6 171 19.5.— 8 .5+— 43.6 -— 11.4 — 5.3+ --- 172 14.110.2 7.2-0.9 51.1 15.3 l4.8io.O 5.9io.i 196 15.1.-- 8.1 — 53.6 -— 16.8 — 6.5+- ~ 242 13.2-1.6 7.010.9 53.0 i6.2 l4.8il.? 5.010.7

144 15.0jl.9 7.011.3 46.7 x 2 ‘7 17.6J2.4 4.6il.O 145 13.9-1.9 6.811.3 48.9 i4.0 I6.2t6.5 5 .oio.8

1 15.3x2.3 9.3jl.l 60.8 J5.5 25.3x3.7 5.?io.7 2 14.412.0 8 .110.8 56.3 13.7 24.oi2.6 5 .510.5

194 13.0.-- 6 .9+— 53.1 r— 19 .4+ — 6.0 — 197 17.6-1.4 8 .2il.l 46.8 il.5 l6.8io.4 5.2io.9

176 8 . 3 -- 4.7 — 56.6 13.7 — 4.8 —

659 14.6-1.4 8.6io.5 58.9— 2.7 13.9il.9 4.113.0 660 1 4 . 4 -- 8.1 — 5 6 . 3 --- 15.2 — 5.0 —

5 14.7x2.3 8 .811.8 59.9 x7.4 29.7x3.0 5.9x0.4 6 14.2J2.5 8 .6J2 .O 60.6 ;5.4 25.0J3.3 4.4io.7 7 15.2jl.6 7.oii.o 46.1 i5.5 26.2J 5.2 6 .1J1.2 8 15.812.2 7.410.8 46.8 i5.4 26.7-3.2 5.9-0.7

255 14.7x1.9 7.2J0.5 49.3 i3.7 21.3x2.4 6 .7x1.2 390 11.512.3 5.712.3 48.6il0.2 15.9-0.4 5 .2io.o 624 12.9 .-- 6.1 — 47.3 x — 18 • 5 . — 5 . 2 -- 651 13.8-1.4 5.910.4 42.5 i l .8 I8.8i2.5 4.8io.6

118 2 0 . 0 -- 8.5 — 4 2 . 5 --- 15.6 — 7.3 — K\

TABUS 11 (Continifd) asa 09 o i + i+f+i-ti+a+ ♦ •♦T+i+i+itT+i+l +7+WW+TV7+1V? '+

cm cm 1 t-o vo i m tv-oo v o m + - * - - * - * - * - m m 1 + T +

knoo i 1 -t - i o v m + + rl O H i 1+1 + Cv. - - cm i +i i +

cm I 1 + m CM 1

m o o r - + m m ^ -+ m o o r - «-i h m oo r oo m h Cv-vO Cv»sO m m m m CM H CM CM OvCM CM -T N K H v O O cm c m mcM + - v t ov m o m v r+ m m i+i+i+i • • •I N \ I I M I vo -T OO -T vo I *+?+

cm vo m cm • « •

r +1 + ■rl * *» o +• r-H-i +• o a * * 9 . 5 ° .*0 ** M • • • CM +

+ + + 1 + 1 + + 1 + vo 1 vo Cv- -* -* vO -* i m | l r h H l rH r m v - vo O cv.-r - 1 C- H rH m Ov Cv-00 O CM Cv-OO m m 1 CM i m n CM 1 m m CM l r i t 1 •1 1 • • • • • • • i • i • • • m 30 CM cm l r 1 •

M n m 4 o o • • # + + + 1 1+1 + T I+ + + vo m vo-* VO OV VO Cv- oo m -r m OV -* vo CM vOOO vO tv- m o o [V-T CM

+i +1 +? +- r - m Q m c m m rH Cv- CM I Cv- O o v m tv- - r * - m OvrH * - l r \ K rl l r l r l r cm o oo a m ov o ov c m m o o v c m i 1 + 1 • • • • • • • • • • • • • • I • • • • • • + 1 +

m c O 1 O 1 l r mvo m m i • • m » cm O i r o h i i i i i i i i i • •

+

Ov O cv- m m Ov m m • i + + 1 - 1 r-l m m • 1

m r o v Ov * - * - rm-rm m cm rHOO m CM -T -T Ov o v m rH OO Cv- cv-co ^ O OV C^O ov - j m CM CM CM o v m cm i t i i i i i i i i i i i i i i i i i I I I I I II I I I i i i i i i i i i i i i 1 1 1 1 i 1 1 1 1 • • • • • • • • • • • • • • • • • • •

mvo v m h i i m M C O l r

TABLE 11 (Continued) it— mrm— rons--- TiBH m i m 'M CROSS LOBE LENGTH W/L LOBK/L i/ b l a d e v e s t u r e cm cm percent BUDS LENGTH PED.LF. Generation fl 10 8 .822.8 3 .8* 1.6 17.5*5.9 40.6'■J9.1 17.5 *5.1 5 5 11 9.9*1.3 4.2*2.8 17.8*1.4 45.2 *5.8 19.2*10.3 5 5

170 2 2 . 4 -- 8 .8A— 25.2^— 91.1, 35.8.--- 2+ > 229 17.8*3.9 6 .4*4.4 23.9*4.4 87.9*10.1 46.7*23.5 2- 1+ 230 14.2*3.4 4.0* 1 .4 22.2*4.4 81.4 *5.1 22.4 * 2.0 2- 1+

351 13.3*4.2 2 .9*1.9 37.6*1.7 100.0 *0.0 21.8 *8.0 U 1 352 13.6*2.3 2 .8*0.6 34.6*1.0 100.0 *0.0 20.6 *7.7 1 1

156 15.2*0.9 1 .4*1.0 40.8*5.6 100.0 *0.0 9.2*22.3 1+ 2 171 11.4 — 1 . 1 -- 46.5. 100.0 9.7 r— 1+ 2 172 14.8*0.0 1 .4*0.1 39.9*1.0 100.0 *0.0 9.4 *1.0 1+ 2 196 16.6 — 4 . 2 -- 38.7.— 100.0 25.0 -— 1+ 2 242 14.8*1.7 1 .1*0 .4 33.8*3.5 100.0 *0.0 7.4 *3.1 1+ 2

144 15.5*2.3 4.5*2.2 26.1*5.6 88.1 *7.6 25.6*14.6 3 4 145 15.5*2.2 3.9*1.4 34.0*1.3 95.7 *5.5 24.1 *8.9 3 4

1 23.6*3.4 4.2*1 .4 22.5*2.0 93.3 * 2 .! 16.6 *4.2 4+ 4 2 22.6*2.2 3 .0* 1.1 22.9*2.6 94.2 *1.9 12.5 *3.7 4+ 4

194 19-4.-- 3 . 2 -- 30.9.-- 100.0 16.5 r— 1+ 2- 197 16.8*0.4 1 .6*0 .3 30.7*6.2 100.0 *0.0 9.6 *1.9 1+ 2

176 1 3 . 7 -- 1.4 — 3 5 . 0 -- 100.0 — 1 0 , 2 --- 2+ 3

659 13.9*1.9 1 .3*0.4 29.5*2.6 100.0 *0.0 9.9 *3.9 4 4 660 1 5 . 2 -- 1.5 — 3 2 . 9 -- 100.0 ---- 9 . 9 --- 4 4

5 9 .6*1.1 9.5*3.3 19.9*2.8 32.3 * 5.6 32.0* 11.6 5 3 6 10.1*1.8 8 .1*1.6 17.6*1.4 40.4 *6.7 32.4 *6.9 5 3 7 11.7*2.3 6 .7*2.1 23.3*0.6 44.7 *5.1 25.6 *8.9 5 3 8 10.2*1.2 6 .0* 3.2 22.1* 2.1 38.2 * 6.8 22.5*10.8 5 3

255 18.3*3.6 9.9*1.8 31.5*2.1 85.3 *8.1 46.5 *4.8 1+ 1 390 15.5*0.5 8.7*2.3 32.7*0.3 97.2 *4.0 54.8*16.8 1+ 1 624 15.3.-- 3 . 6 -- 2 8 . 1 -- 82.7 ...» 19.5 ---- 1+ 1 651 14.8*1.4 4.9*1.3 25.6*0.4 78.9 *3.2 25.9 *3.4 1+ 1

118 15.6 — 6.9 — 46.8 — 100.0 .... 4 4 . 2 --- 3 3 145

TABLE 11 (Continued) TERMINAL BLADE TERMINAL CROSSLOBE LENGTH W/L LOBE/L i/ b l a d e v e s t u r e cm cm nercent BLADE LENGTH PED.LF.

9 22.0-1.9 9.6*4.4 21.5 *2.4 89.9 *1.2 38.7*15.8 5 2 22 18.2-0.2 8.3*0.1 25.6 *4.0 89.7*10.2 40.9-*4.9 5 2 29 20,1-3*4 5.8*2.8 26.2 *2.4 98.3 *1*8 29.6*15.9 5 2+ 37 16.9*0.7 6.1*0.7 27.1 *3.4 89.9 *2.6 32.5 *3.6 5 2+ 38 14.9*1.4 5.9*0.2 34.0 *3.4 97.4 *3.0 37.5 *6.1 5 2+ 40 20.9-1*2 8.7*4.5 27.2 *2.8 96.3*10.5 40.1*19.0 5 2+ 41 20.7-2.2 10.6*6.4 24.8 *2.7 96.7 x 2*-8 49.5*25.0 5 2+ 240 24.1-3.2 6.4*0.7 32.6 *3.1 93.1 *2.6 25.0 *3.0 5- 3 241 18.6a-- 4.4 — 30.5 x— 9 4 . 4 --- 22.3 x -- 5- 3 577 19.9-0.1 7.9*1.1 28.7 *1.4 91.6*11.9 35.9 *0.3 5 2+

15 1 2 , 2 -- 15 .2^-- 20.6 x-- 62.9 x — 78.4 5 1 35 14.3x1.6 14.6*2.5 22.3 x1*8 65.0 *2.2 66.4*14.2 5 1 15.8*4.2 1 36 13.6*3.1 23-S & * 0 61.8*13.7 71.8*16.7 5 179 13.5-2.4 8 .8*4.0 21.8 *4.7 55.9 * 2.0 35.1 *8.4 5 1

F 2 Generation 355 16.7*3.1 3.4*3.0 28.3 *5.6 92.8 * 6.6 18.9*12.4 5- 4 356 16.3*3.3 3.2*0.5 28.6 *0.2 100.0 *0.0 20.1 * 7.2 5 4 469 17.0-3.9 2 .2*0.1 20.0 *4.5 94.4 *1.7 12.2 *4.9 5- 4

31 18.5.-- 6.9.-- 27.5+x-- 87.7 x— 3 2 . 7 --- 5 1 32 13.4-1.9 8 .2*4.2 26.9*12.0 100.0 *0.0 64,4-41.0 5- 1

Backcroases

366 22.1*6.8 5 .6* 2.0 18.0 *3.5 86.3 *8.6 21.9 * 6.2 5- 5 488 17.9-5.1 4.3*2.0 16.2 *2.5 78.2*14.4 18.8*14.2 5 5-

472 13.3-0.0 7.9*1.1 23.6 *3.9 75.4 *8.7 44.9*11.2 5- 1+

253 16.3j 5.8 7.4*2.3 17.5 *2.7 64.9 *9.5 26.7 *9.5 5 5- 387 14.9-2.2 7 .4*2.4 18.3 *2.4 58.0* 10.0 28.8* 12.2 5 5- 388 18.8 — 1 1 . 3 -- 2 0 . 2 --- 6 0 . 3 --- 3 6 . 2 --- 5 5

3 16.1*1.1 14.6*3.6 22.1 *1.5 72.5 * 2.1 65.8*15.3 5 2 18 16.5*— 8 . 1 -- 24.9 x-- 80.5 x— 3 9 . 5 --- 4 1 19 17.1*2.9 12.4*0.3 23.3 *1.6 78.1 *7.5 56.6* 11.0 5- 1+ 33 16.1*3.1 10.6*3.6 27.9 *1.3 84.7*13.6 55.8*17.4 5 1+ 39 14.8 — 1 9 . 0 --- 1 6 . 5 --- 6 1 . 2 --- 7 8 . 5 --- 5- 1

25 17.3 — 1 0 . 7 -- 2 2 . 0 ---- 8 0 . 8 ---- 50.O --- 5- 1+ 26 17.3 — 7 . 2 --- 2 1 . 9 ---- 9 4 . 5 ---- 3 9 . 3 ---- 5 1+ 120 2 4 . 3 --- 1 1 . 2 --- 2 9 . 7 ---- 73 .O ---- 3 3 . 6 ---- 5 1+ FIELD COLLECTION ANALYSIS (EAST)

This analysis is mads to investigate the possibility of lntrogressive hybridization between B. pumila and B. subacaulis in the field in view of the fact that we already know that they can be induced to interbreed via artificial pollination under greenhouse conditions. The basic methods of Anderson (19^9) are here expanded to populations as units rather than individ­ ual plants. Sixty-eight populations are considered. Thirty- two are mass collections made on successive trips to the region; thirty-six are compilations of specimens from various herbaria. Selected were large herbaria specimens compilations and those, regardless of size, from areas otherwise poorly represented.

Five measurements identical to those in experimental plant analyses were selected. As a result of the analyses, hybrid index values were established and scaled in such a way that

"ideal" B. pumila should have a hybrid index value of 5t

"ideal" j). subacaulis. 25* The values are scaled thus because

the F^ hybrids are not exactly intermediate*in all characters.

The population data of Tables 12 and 13 are arranged geographically. 1V7

TABLE 12

LIST OF HERBARIA SPECIMENS USED IN HYBRID INDEX ANALYSIS OF ______EASTERN TAXA COMPLEX______top. fl.i.

TABU 12 (Continued) POP. H.X. COUNTY STATE DATE NAME NUMBER COLLECTEDCOLUCTOR(S)

D 11 Duval Fla — May 1877 Curtiss, A.H. .... E 9 Duval Fla 7 Apr 1897 Churchill, J.R. F 10 Duval Fla 16 Mar 1882 Smith, J.D. ---- 1238 11 St. Johns Fla 2 Sept 1963 Speer,Weber&P. P1238

East Coast,, North 1to South

R 15 St. Johns Fla Je-Dec 1875 Reynolds, Mary C' • -- C 16 Volusia Fla 29 Mar 1897 Kennedy, 0.0. — B 19 Volusia Fla 30 Je Curtiss, A.H. 1393 A 23 Volusia Fla 26 Apr 1906 Deam, Stella 1760 FF 23 Brevard Fla 11 Ju 1896 Curtiss, A.H. 5699 DD 2k Highlands Fla 15 Mar 19V6 Steyermark, J.A.63310 EE 25 Palm Beach Fla — Mar 1902 Canby, Wm.M. ... 00 2k Dade Fla 28 Feb 1915 Small & Mosier 55V7 L 23 Dade Fla 2k Mar 190V Britton, N.L. ... K 2k Dade Fla 21 Jan 1930 Moldenke, H.N. V85 J 23 Dade Fla 9+Nov 1903 Small & Carter ...

West Coast and Inland Florida, South to North

JJ 21 Manatee Fla 16 May 1900 Tracy, S.M. 6906 II 23 Hillborough Fla 11 Apr 1923 Churchill, J.R. 30 X 2k Pinellas Fla IV Apr 1900 Tracy, S.M. 6916 0 22 Pasco Fla — Mar 1962 Armstrong,Arnholt, S t ephan&Pinkava PO BB 2k Lake Fla 12+Mar 189V Nash, Geo. V. 33 AA 2k Lake Fla 28+ May 1895 Nash, Geo. V. 1020 Z 2k Lake Fla 16+May I89V Nash, Geo. V. 808 EE 2k Gilchrist Fla 3 Mar 1957 Godfrey, R.K. 55V33 T 23 Taylor Fla 9 Apr 1935 Radford, A.E. 8319 1235 2k Putnam Fla 2 Sept 1963 Speer,Weber&P. P1235 CC 23 Putnam Fla — Apr 1869 Canby, Wm.M. TABLE 13

HYBRID INDEX ANALYSIS 07 EASTERN U.S. TAXA 1 raw 'mm ~ ■ 'izm mm 1 ~ POP. H.I. r X x x t* Bt 0Yt x t 8t cvt

126? 6 1 44.4 6.7 ——- 0.0 0.7 ------1266 6 2 50.0 7.1 0.0 0.0 0.0 0.7 0.0 0.0 1264 7 9 40.3 6.3 1.0 15.0 0.0 0.7 0.0 0.0 H 7 5 47.9 6.9 0.8 11.1 12.2 2.6 2.8 107.5 1 7 11 36.2 6.0 0.7 11.3 0.0 0.7 0.0 0.0 1262 6 3 43.5 6.6 0.8 11.6 0.0 0.7 0.0 0.0 1253 6 9 50.5 7.1 0.8 11.1 0.9 1.0 0.8 78.6 S 6 5 44.4 6.7 0.3 5.1 0.0 0.7 0.0 0.0 1291 5 1 56.3 7.5 ---— 0.0 0.7 ------1294 6 10 49.8 7.1 0.7 10.6 0.0 0.7 0.0 0.0 T 9 3 33.6 5.8 0.8 13.4 14.3 2.7 3.4 127.1 1297 6 3 51.1 7.2 0.1 1.7 0.0 0.0 0.0 0.0 V 8 2 37.5 6.1 1.5 24.0 5.0 2.0 1.8 90.6 W 8 2 39.7 6.3 0.7 11.5 0.0 0.7 0.0 0.0 U 8 3 42.0 6.5 0.5 8.1 0.0 0.7 0.0 0.0 1197 7 25 42.1 6.5 0.9 13.2 2.5 1.1 1.4 122.9 1198 10 13 37.4 6.1 0.8 12.6 11.7 2.7 2.3 86.7 I 9 9 36.2 6.0 0.9 15.7 2.0 1.1 1.2 108.3 1199 8 21 41.6 6.5 0.7 10.1 3.5 1.4 1.5 107.5 1202 8 18 47.2 6.9 0.8 11.3 22.5 3.5 3.4 98.1 1203 9 12 37.6 6.1 1.1 18.1 7.1 1.9 2.1 113.4 1205 10 12 38.2 6.2 0.9 14.9 8.4 2.0 2.3 118.3 1206 7 17 49.3 7.0 0.6 8.9 1.8 1.1 1.1 102.2 1208 6 12 42.7 6.5 0.7 10.5 0.7 0.9 0.6 62.8 1210 7 10 41.7 6.5 0.8 12.6 4.7 1.5 1.8 117.3 1212 9 13 41,0 6.4 1.1 17.5 16.0 2.7 3.1 113.5 1216 7 1 55.6 7.5 — ---- 0.0 0.7 — ---- 1217 8 11 34.3 5.7 1.5 26.6 12.1 2.6 2.5 95.8 1224 7 3 39.2 6.3 0.8 12.8 0.0 0.7 0.0 0.0 1225 7 4 44.2 6.7 0.5 6.8 8.3 2.2 2.4 108.8 M 7 2 48.1 7.0 0.2 2.9 0.0 0.7 0.0 0.0 2 9 11 39.4 6.2 1.1 17.0 9.4 2.1 2.4 115.6 122? 7 13 42.9 6.5 1.0 15.1 6.0 1.9 1.8 99.6 0 7 3 36.9 6.1 0.2 3.5 0.0 0.7 0.0 0.0 Q 6 2 60.8 7.8 0.0 0*0 20.6 4.6 0.4 8.3 P 7 2 53.6 7.4 0.3 4.6 4.6 1.9 1.7 88.7 1234 6 1 48.0 7.0 ------0.0 0.7 ------N 6 4 53.5 7.3 0.4 5.7 0.0 0.7 0.0 0.0 0 8 6 33.2 5.8 0.7 12.5 2.4 1.2 1.3 103.9 1246 7 13 38.3 6.1 1.3 21.2 2.7 1.2 1.3 108.4 * Data aarktd "t" baa«d on tranaforaation x ■ x+)i 150

TABLE 13 (Continifd) _UPPZR LEAVES — u > m m m — 4» POP. H.I. r X X *t Bt CVt Bt ovt

1243 7 13 35.3 5.9 1.0 16.5 1.1 1.0 0.9 92.0 1241 10 9 40.7 6.3 1.2 19.0 14.8 3.0 2.6 87.5 0 11 11 39.6 6.3 1.0 16.0 28.3 5.0 2.0 40.8 6 9 3 37.9 6.2 0.4 6.4 20.0 3.1 4.1 133.^ 1238 11 12 42.8 6.5 0.9 13.5 23.1 3.7 3.3 90.7 r 10 3 29.9 5.5 0.7 12.8 16.1 4.4 4.1 93.4

R 13 4 34.9 5.9 1.0 16.5 38.0 4.7 4.6 98.2 C 16 2 27.9 5.3 0.1 1.4 77.1 8.8 1.4 15.8 B 19 11 40.7 6.3 1.1 18.0 79.0 8.9 0.7 7.6 A 23 5 4.0 1.5 1.7 116.2 100.0 10.0 0.0 0.0 FT 23 4 26.7 5.1 1.4 28.0 97.2 9.9 0.3 2.9 DD 24 2 9.5 3.2 0.1 3.2 95.5 9.8 0.3 3.2 EE 25 3 0.0 0.7 0.0 0.0 100.0 10.0 0.0 0.0 00 24 3 11.9 3.1 2.1 67.3 91.7 9.6 0.8 8.0 L 23 4 9.1 2.5 2.1 84.2 87.7 9.4 0.4 4.6 K 24 2 6.3 2.2 2.1 94.9 93.8 9.7 0.5 4.6 J 23 10 3.8 1.4 1.6 109.8 84.9 9.2 0.6 6.9

JJ 21 3 16.7 3.6 2.5 70.8 97.8 9.9 0.2 2.0 II 25 7 5.3 1.8 1.8 101.4 95.8 9.8 0.3 2.7 X 24 6 0.0 0.7 0.0 0.0 97.6 9.9 0.3 3.1 0 22 2 3.6 1.7 1.5 83.5 95.5 9.8 0.4 3.5 BB 24 5 0.0 0.7 0.0 0.0 88.6 9.4 0.4 4.2 AA 24 2 10.0 2.6 2.7 103.1 100.0 10.0 0.0 0.0 Z 24 3 7.4 2.1 2.3 113.7 100.0 10.0 0.0 0.0 HH 24 6 1.5 1.1 1.0 88.3 97.2 9.9 0.4 3.6 T 23 3 7.4 2.1 2.3 113.7 100.0 10.0 0.0 0.0 1235 24 10 8.3 2.7 1.4 53.2 90.1 9.5 0.7 6.9 CC 25 3 2.0 1.3 1.1 131.5 97.8 9.9 0.2 2.0 TO 0*0 V0 0*0 0*0 0*0 0*01 6*66 VOT 9*0 V 5 8*2S 5*721 8**7l 1*0 8*0 1*0 5*0 1*0 0*01 5*66 V 6 5*0 9*S V 05 9+72T 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 S**r! 8*o v s 8*82 0 0*0 0*0 V 0 0*0 0*0 0*0 0*01 0*001 6*8 5*0 5*5 5*62 N ------VO 0*0 ----— 0*0T 0*001 ... 1*9 2*9f *7521 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 0*6 5*0 1*5 V 52 d 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*00T s**l 5*0 8*S 9*25 b 0*0 0*0 VO 0*0 0*0 0*0 0*0T 0*001 2*8 5*0 8*5 8*25 0 2*5l 1*0 8*0 VO 5*0 1*0 0*01 5*66 2*2 1*0 9 ‘S V 62 £221 6*9£ 5*0 V0 2*0 2*0 0*0 0*01 9*86 5*01 5*0 2*5 8*92 2 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 0*0 0*0 2*5 1*92 W 0*0 0*0 v 0 0*0 0*0 0*0 0*01 0*001 5*£ *•0 6*5 S * tf 5221 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 0*21 V 0 8*S 5*55 *?221 5*02 2*0 8*0 1*0 2*2 2*0 0*01 V 8 6 5*51 V 0 v s 5*62 £121 ... ---- VO 0*0 ------0*01 0*001 ... VS 8*82 9121 5*6l 1*0 8*0 1*0 V I 2*0 0*01 1*66 1*21 9*0 2*5 V 92 2121 0*0 0*0 V o 0*0 0*0 0*0 0*01 0*001 V 8 5*0 5*5 1*05 0121 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 6*51 8*0 5*5 V 05 8021 9**? 1*0 0*1 0*0 5*0 VO 0*01 V 66 5*01 9*0 VS 9*82 9021 9*£ 1*0 Vo 0*0 5*0 0*0 0*01 8*66 8**1 8*0 **5 V 82 5021 8*61 2*0 8*0 1*0 0*1 0*1 0*01 5*66 8**?t V o 0*5 6**72 5021 6*15 vo 8*0 2*0 6*1 2*0 6*6 0*86 1*21 9*0 2*5 2*^2 2021 o *6t v o VO 0*0 0*0 0*0 0*01 0*001 8*8 5*0 2*5 6*92 6611 0*0 0*0 V 0 0*0 0*0 0*0 0*01 0*001 8** 2*0 V * 1*12 I 9*1 0*0 8*0 v o 5*1 1*0 6*6 2*86 V £T VO 6** 6*52 8611 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 8*8 5*0 2*5 V 92 £611 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 6*8 5*0 5*5 5*42 n 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 9*£ **0 5*5 £*£2 M 0*0 0*0 V o 0*0 0*0 0*0 0*01 0*001 1*02 0*1 1*5 A 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 8*f 2*0 V 5 v is £621 1*22 2*0 8*0 2*0 0*1 v o 0*01 0*66 **9 5*0 1*5 8*52 0*0 0*0 V 0 0*0 0*0 0*0 0*01 0*001 V* 2*0 6*5 5**r5 *7621 ------V o 0*0 ---- o*ot 0*001 ...... 8*5 V 25 1621 0*0 0*0 V o 0*0 0*0 0*0 0*01 0*001 2*5 5*0 9*S 6*05 8 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 1*£ **0 0*9 2*95 5521 0*0 0*0 V0 0*0 0*0 0*0 0*01 0*001 V 5 5*0 9*9 9*2*7 2921 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 0*8 5*0 6*5 S** t5 1 5*22 2*0 8*0 1*0 5*0 l* o 0*01 5*66 2*6 5*0 5*5 5*05 H 0*0 0*0 V o 0*0 0*0 0*0 0*01 0*001 **8 5*0 0*9 V 55 *7921 0*0 0*0 VO 0*0 0*0 0*0 0*01 0*001 5*21 VO 6*5 V*r5 9921 »* V 0 0*0 ““ “ “““ 0*01 0*001 ••• V 9 V*7*7 £921

1 % % *A0 X AO 0 *x X *d0d *• *A0 *• *x X* HSHHfllfSHO* 1 n m g / j m o t •hSm ___i / » s a n e ~ ^•ntrnuoo) tx ffiari

151 152

TABU 13 (Continued) ------w a i w — msrijSwmsrL — eobttober— pop. ; ;t .t c»t x ;t «t cvt s ;t »t cvt

124l 23.6 4.9 0.3 5.4 97.2 9.9 0.2 2.3 0.1 0.8 0.1 17.1 D 22.7 4.8 0.6 11.6 96.7 9.9 0.2 1.7 0.3 0.9 0.1 25.1 & 24.9 5.0 0.2 3.2 97.9 9.9 0.2 1.7 0.2 0.8 0.1 26.4 F 22.3 4.8 0.2 4.5 98.3 9.9 0.1 0.9 0.1 0.8 0.1 17.4 1238 23.1 4.8 0.5 11.3 96.2 9.8 0.2 2.2 0.4 0.9 0.3 26.8

R 21.8 4.7 0.3 7.1 92.8 9.7 0.5 4.8 0.6 1.0 0.4 35.5 C 20.9 4.6 0.4 8.0 87.1 9.4 0.3 2.8 0.9 1.2 0.1 8.6 B 17.9 4.3 0.4 9.6 59.9 7.7 1.0 13.0 1.7 1.5 0.3 16.3 A 14.7 3.9 0.5 13.6 49.5 7.0 1.0 14.0 3.0 1.9 0.2 12.2 FF 13.5 3.7 0.2 4.3 23.0 4.8 0.4 8.1 4.4 2.2 0.3 15.0 DD 12.2 3.6 0.2 4.4 61.1 7.1 1.4 19.0 3.0 1.8 0.6 32.4 EE 10.7 3.4 0.1 3.2 29.7 5.5 0.7 12.0 4.0 2.1 0.4 16.5 OG 10.7 3.4 0.1 4.2 22.4 4.8 0.5 9.7 4.5 2.2 0.3 15.4 L 14.5 3.9 0.4 10.5 22.4 4.7 0.8 17.2 5.0 2.3 0.2 7.4 K 12.8 3.6 0.5 12.8 23.3 4.9 0.2 3.2 4.2 2.1 0.4 17.8 J 14.8 3.9 0.3 7.7 41.1 6.4 1.0 15.2 2.9 1.8 0.3 18.3

JJ 18.5 4.4 0.4 8.4 59.9 7.7 1.0 12.3 1.9 1.6 0.1 9.1 II 12.9 3.6 0.4 11.1 34.4 5.9 0.9 15.1 3.7 2.0 0.1 6.6 X 16.8 3.3 0.4 10.9 24,1 4.9 1.1 22.4 3.5 2.2 0.4 16.0 0 16.0 4.0 0.6 14.6 34.7 5.9 0.0 0.5 3.0 1.9 0.0 2.1 BB 11.5 3.4 0.5 14.9 41.8 6.3 1.9 29.4 3.8 2.1 0.2 10.2 AA 15.5 4.0 0.0 0.0 21.9 4.7 0.3 5.8 4.0 2.1 0.2 11.1 Z 14.0 3.8 0.5 13.0 34.6 5.8 1.4 24.3 3.4 2.0 0.4 18.0 HH 14.1 3.8 0.2 5.8 27.7 5.2 1.0 19.7 3.7 2.1 0.2 10.2 Y 20.4 4.6 0.5 11.0 31.2 5.6 0.4 6.9 3.5 2.0 0.3 15.9 1235 13.6 3.6 0.3 8.9 24.5 4.9 1.2 23.6 4.1 2.1 0.3 133.3 CC 9.9 3.2 0.3 8.4 27.4 5.2 1.0 19.6 4.3 2.2 0.3 12.6 153

Hybrid Percent Percent Blade Terminal Lobe Index Upper Leaves W/L Lobe I/Blade Number Value Leaves Lobed Ratio L Ratio

1 56* 0-5 29* 96.5-100.0 0.00-0.25 2 42-56 5-35 24-29 89.5- 96.5 0.25-0.75 3 26-42 35-65 19-24 75.5- 89.5 0.75-1.50 4 12-26 65-95 14-19 47.0- 75.5 1.50-2.50 5 0-12 95-100 9-14 0.0- 47.0 2.50f

The histogram drawn from these data (Figure 48a) indieataa two major groups corresponding to the two apaciaa9 B. subacau- li£ and B. £ggila; plus two populations centering about the 7^ plant hybrid index value (H.I. 15). The B. humilla holotype and iaotype belong to the population with H.I. 13. Both species groups of the histogram have been skewed toward one another indicating gene flow from one species to the other, but restricted to the extent that swamping has not occurred. It is known that

F^ plants easily backcross to the parents, but I have not yet observed the offspring. If these should fall into the skewed

"bulges" of the species groups on the histogram, I feel confident that introgression is occurring, and slowly. The center of hybridisation is apparently the Jacksonville-St. Augustine,

Florida, region; this is confirmed by the clinal variation of all characters (deviations from H.I. 13 increases in all directions from the center). This is also the collection site of B* humilis types. F^ plants have been collected in distant regions, for example at Tampa Bay, and cannot be yet explained unless they are from local contamination by transported specimens. Analyses of variance indicat* in all characters tested significant variation between populations and between species- hybrid groups (Tables 14 and 15). This great variation also is indicative of hybridisation. 155

TABLE 14

ANALYSIS OF VARIANCE BETWEEN POPULATIONS OF BERLANDIERA EASTERN TAIA COMPLEX Sourcs of 1 1 V Values Variation df 3S MSF 0.05 0.1 Percentage Upper Leaves

Among 67 1502.90 22.43 20.77* *1.43 <1.32 Within 390 419.96 1.08 Total 457 1922.86

Percentage Leaves Lobed

Among 67 5034.08 75.13 22.70* Within 390 1290.52 3.31 Total 457 6324.60

Blade Width-Length Ratio

Among 67 242.96 3.63 13.58* Within 390 104.08 0.27 Total 457 347.04

Terminal Lobe Length - Blade Length Ratio

Among 67 1397.38 20.86 99.31* Within 390 82.01 0.21 Total 457 1479.39

Number of Lobes

Among 67 116.94 1.75 45.32* Within 390 15.03 0.04 Total 457 131.97

* Values are significant atthe 0.05 0.1 levels against the hypothesis that thsrs are no differences between populations due to location* 156

TABLE 15

ANALYSIS OF VARIANCE BETWEEN SPECIES-HYBRID GROUPS OF BBRLAND- IXRA EASTERN TAXA COMPLEX

Source of F Values Variation df SS MS F 0.05 0.1 Percentage Upper Leaves Groups 2 1164.18 582.09 538.97* <3.07 <2.35 Among 65 338.72 5.21 4.84* < 1.43 < 1.32 Within 390 419.96 1.08 Total 457 1922.86

Percentage Leaves Lobed

Groups 2 4625.17 2312.59 698.67* Among 65 408.91 6.29 1.90* Within 390 1290.52 3.31 Total <*57 6324.60

Blade Width-Length Ratio

Groups 2 182.99 91.50 338.89- Among 65 59.97 0.92 3.46- Within 390 104.08 0.27 Total 457 347.04

Terminal Lobe Length - Blade Length Ratio

Groups 2 1253.96 626.98 2985.62- Among 65 143.42 2.21 10.50* Within 390 82.01 0.21 Total 457 1479*39

Number of Lobes

Groups 2 112.90 56.45 1411.25* Among 65 4.04 0.06 1 .61- Within 390 15.03 0.04 Total 457 131.97

9 V.*' ______1 1 A A — — A Mb m m 1 the hypothesis that there are no differences between species- hybrid groups due to location (upper value of each pair) and no differences between populations due to location (lower value of each pair). FIELD COLLECTION ANALYSIS (WEST)

This analysis is of taxa centering about sastsrn Texas.

Techniques used for eastern taxa also apply to western taxa except that the distinguishing characters between groups are different. Collectors ordinarily do not collect the eniire specimen (even from above the root) of these larger, coarser plants} thus the total height is unknown in most cases.

This, unfortunately, makes it impossible for data to correspond directly to parental and hybrid eaqterimental plant analyses.

The characters used are: the number of leaves along stem ten centimeters from below the lowest branch of the inflores­ cence; the blade width-length ratio and petiole length-blade length ratio as previously described but based on sum values of the three leaves immediately below the inflorescence rather than on central cauline ones; head width as described; and peduncle vesture. Peduncle vesture scale is altered as follows:

5 - dense, erect, long hirsute; - sparse, spreading, erect hirsute; 3 - spreading or curling, long fine hairs; 2 - long fine hairs, not matted; and 1 - long fine hairs, matted.

Hybrid index values cited below are scaled according to experimental data. The arrangement of these is that "ideal"

B. pumila should receive a hybrid index value (H.I.) of 3;

"ideal" B. texana. a value of 23* Tables 16 and 17 are

157 158

TABLE 16

LIST OF HERBARIA SPECIMENS USED IN HYBRID INDEX ANALYSIS OF WESTERN TAXA COMPLEX t o . t . t . — m m — B r o s — b u s------bibb m m COLIJBCTEP COLLECTOR(S)

15 24 Bandera Tex 14 Oct 1963 Pinkava, D.J. P15 V 24 Bandera Tex 4 Nov 1953 Shinners, L.H. 16882 14 20 Bandera Tex 14 Oct 1963 Pinkava, D.J. Pl4 w 23 Kerr Tex 12+Je 1894 Heller, A.A. 1874 X 21 Qllleeple Tex — Je 1939 Tharp, B.C. 285 AA 23 Blanco Tex 15 May 1928 Palmer, E.J. 33950 Z S3 Comal Tex -- Je 1850 Lindhelmer, F. 886 Y 20 Bexar Tex 30 Je 1911 Clemens, Mr & Mrs 1008 U 23 Travis Tex 16 May 1940 Lundell, C.L. 8c A . 8919 cc 21 Refugio Tex 12 Apr 1930 Benke, B.C. 5443 12 21 Fayette Tex 13 Oct 1963 Pinkava, D.J. P12 11 13 Bardin Tex 12 Oct 1963 Pinkava, D.J. PU 10 12 Newton Tex 12 Oct 1963 Pinkava, D.J. P10 9 13 Newton Tex 12 Oct 1963 Pinkava, D.J. P9 Q 9 Austin Tex ■* May 1844 Lindhelmer, F. --- p 11 Waller Tex 1 May 1872 Hall, Elihu 324 R 11 Montgomery Tex l8+Ju 1909 Dixon, Royal A. 469 S 14 Walker Tex 9+Ju 1909 Dixon, Royal A. 334 DD 9 Cherokee Tex 3 Je 1915 Palmer, E.J, 7856 EE 13 Harrison Tex 8 Je 1915 Palmer, E.J. 7916 8 11 Harrison Tex 11 Oct 1963 Pinkava, D.J. P8 7 11 Gregg-Up, Tex 11 Oct 1963 Pinkava, D.J. P7 5 11 Upshur Tex 11 Oct 1963 Pinkava, D.J. P5 6 9 Upshur Tex 11 Oct 1963 Pinkava, D.J. P6 it 11 Cams Tex 10 Oct 1963 Pinkava, D.J. P4 3* 11 Jackson Fla 7 Oct 1963 Pinkava, D.J. P3 1262* 8 Houston Ga 4 Sept 1963 Speer, Weber & P. P1262 1253* 7 Toombs Ga 3 Sept 1963 Speer, Weber 8e P. P1253 T 12 Case Tex 12 May 1941 Me Ginnis, 0. mmrnrn M 14 Madison Ark 14 Aug 1943 Moore, D.M. 1430201 I 20 Me Donald Mo 1 Sept 1893 Bush, B.F. 163 J 18 Me Donald Mo 24 Ju 1892 Bush, B.F. —- H 22 Jasper Mo 33 Je 1909 Palmer, E.J. 2316 35 18 Lawrence Mo 20 Oct 1963 Pinkava, D.J. P35 E 19 Greene Mo 2 Oct 1925 Palmer, E.J. 28977 F 20 Greene Mo 31 Ju 1937 Steyermark, J.A. 24003 K 20 Ozark Mo 11 Oct 1927 Palmer, E.J.O 33105 a 20 Howell Mo 12 Aug 1934 Steyermark, J.A. 14490 N 20 Shannon Mo 12 Sept 1888 Bush, B.F. 0 22 Shannon Mo 26 Ju 1936 Steyermark, J.A. 12304 * Eastern extension of complex* 159

TABU 16 (Continued) WF.TTT.—m m — stub— bub------ms sweet COLLECTED COLLECTORS)

L 18 Pul*ski Mo 28 Aug 1937 Steyermark, J.A. 25507 A 22 Me Curtain Okla 28 May 1916 Houghton, H.W. 3892 C 23 Cleveland Okla 22 Je 1936 Demaree, Delete 13083 B 23 Oreer Okla 17 Je 1913 Stevens, Q.ff. 1037 BB 22 Hemphill Tex 17 Je 1918 Palmer, E.J. 14101 TABLE 17

HTBBID INDEX ANALYSIS 0T WESTERN U.S. TAXA

At BLADE W/L BtPET. VBLADS l FOP. rA B x ■ cv X 8 cv 15 16 27.1 4.5 16.5 2.9 2.6 67.8 3" V l 26.6 —. .... 1.1 .... 1 14 5 30.3 5.2 17.2 4.0 2.0 50.5 7 w 5 29.3 0.9 3.1 2.7 1.7 64.3 5 X 2 29.9 6.6 22.2 2.3 0.1 4.4 2 AA 3 26,5 1.1 4.3 1.5 0.3 21.3 2 Z 1 27.1 2.2 1.4 63.9 2 y 1 30.4 -- --- 5.1 .... 1 u 3 27.6 2.9 10.6 1.2 1.2 98.4 3 cc 1 23.8 ...... 3.6 .... 1 12 7 31.3 4.5 14.5 2.3 0.3 13.7 7 11 8 35.9 6.9 19.3 10.3 5.0 48.4 8 10 7 37.0 4.8 13.0 12.0 4.8 40.0 7 9 11 33.1 6.2 18.7 12.8 6.4 50.0 .1 Q 2 49.3 4.4 8.9 8.5 3.6 42.0 2 P 5 36.6 1.9 5.3 7.5 3.6 47.7 5 R 2 37.8 0.4 1.1 5.7 1.3 22.6 2 S 2 28.9 1.7 5.7 9.3 2.0 21.9 2 DD 3 36.1 4.9 13.5 6.1 4.3 71.4 3 EE 2 34.2 3.5 10.3 1.9 2.7 142.2 2 8 8 40.4 4.6 11.5 7.2 6.5 90.9 8 7 9 32.6 4.8 14.6 11.1 3.4 30.2 9 5 19 36.9 4.3 11.6 11.6 5.6 48.5 9 6 4 33.3 3.9 11.6 12.7 4.0 31.2 4 10 36.3 4.7 13.0 9.9 3.0 29.8 .0 3 10 35.5 5.4 15.2 7.0 4.4 63.2 ,0 1262 3 43.3 3.2 7.3 14.0 2.9 20.6 3 1253 6 36.6 3.7 10.2 22.8 10.7 47.0 6 T 1 34.4 .... 4.6 — .... 1 M 2 35.5 1.2 3.3 1.5 0.2 11.2 2 T 1 24.4 ... 2.4 ... 1 J 2 33.8 4.2 12.5 1.7 0.5 26.6 2 E 1 24.3 ...... 3.6 ...... 1 35 3 35.3 4.8 13.7 2.4 0.1 4.2 3 E 3 26.8 4.2 15.5 3.0 0.9 29.9 3 F 3 27.8 8.9 31.8 1.0 0.2 17.9 3 K 2 23.1 0.3 1.1 3.2 2.0 62.2 2 0 1 22.0 — —— 3.1 ...... 1 »

161

TABLS 17 (Coatlnuod) h .i . POP. At BLADE V/L PA rB x m CT z • OT i 20 N 3 24.1 5.3 21.8 2.8 0.5 17.7 4 22 0 2 25.7 0.4 1.4 2.5 1.7 65.9 2 18 L 2 28.1 5.2 18.4 2.0 1.2 62.0 2 22 A 2 18.0 2.8 15.4 1.7 0.2 9.8 2 23 C 3 27.5 1.3 4.7 2.8 1.1 38.5 3 25 B 2 24.9 4.4 17.8 2.7 0.8 29.1 2 22 BB 2 30.6 2.3 7.6 1.9 0.5 26.3 2 162

T A B U 17 (Continued)

POP. r. Ct LV8. BSLOW r. r. Si PSD. VSSTUBB C D S X • «TX 0 CTX ■ CT

15 10 8.A 8.5 3A.6 10 2.0 0.2 8.5 10 A.9 0.3 6.5 V 1 6.0 ---.... 1 2.3 —- . . . . 1 5.0 --- lA 10 5.6 1.6 28.2 9 2.1 0.2 11. A 10 5.0 0.0 0.0 V 5 6.0 1.9 31.2 5 2.3 0.3 13.1 5 5.0 0.0 0.0 X 2 5.0 A.2 8A.8 2 2.1 0.1 6.7 2 5.0 0.0 0.0 AA 2 6.5 2.1 32.6 3 2.1 0.2 9.5 3 5.0 0.0 0.0 Z A 5.3 1.5 28.6 2 2.3 0.2 10.0 A 5.0 0.0 0.0 I 1 A.O — 2 2.7 0.6 23.7 2 5.0 0.0 0.0 u 3 8.0 1.0 12.5 3 1.8 0.2 13.3 3 5.0 0.0 0.0 cc 2 A.5 0.7 15*8 2 2.2 0.3 12.7 2 5.0 0.0 0.0 12 7 5.3 1.0 18.0 8 2.5 0.2 9.2 8 A.9 O.A 7.A 11 9 A.l 1.5 35.3 9 1.9 0.1 5.3 9 3.7 0.5 13.6 10 7 A.O 0.6 lA.3 8 2.0 0.2 12.0 9 3.8 O.A 11.6 9 12 A.7 2.2 A7.8 12 1.8 0.2 11.1 12 3.5 0.5 1A .9 Q 2 3.0 0.0 0.0 2 1.9 O.A 19*0 2 1.0 0.0 0.0 P 5 A.O 0.7 17.8 5 1.9 0.3 13.7 5 1.0 0.0 0.0 s 3 A.7 2.1 AA.? 3 1.7 0.1 5.9 3 1.7 0.6 3 A .6 8 3 A.O 1.0 25.0 3 2.1 0.2 8.1 3 2.0 0.0 0.0 DD 3 3.0 1.0 33.3 3 1.7 0.1 5.9 3 1.0 0.0 0.0 SS 2 3.0 0.0 0.0 2 1.9 0.0 0.0 2 2.0 0.0 0.0 8 9 A.7 l.A 30.2 7 1.8 0.1 7.8 9 2.A 0.5 21.7 7 10 A.A l.A 32.5 10 1.7 0.2 11.8 10 2.0 0.0 0.0 5 18 3.6 0.6 17.A 19 1.9 0.2 12.6 19 1.9 0 .5 2A .3 6 A 3.5 1.7 A9.A A 1.7 0.1 2.9 A 2.0 0.0 0.0 A 10 3.A 1.1 31.8 10 2.1 0.3 lA.3 10 2.0 0.0 0.0 3 10 2.7 0.8 30 .A 10 2.1 0.3 12.A 10 1.0 0.0 0.0 1262 3 2.3 0.6 2A.5 2 1.8 0.1 5.6 3 2.0 0.0 0.0 1253 6 A.2 1.7 A1.3 1 ' 1.1 --- .... 6 1.2 O.A 35.0 T 2 5.0 2.8 56.6 2 1.6 0.1 8.8 2 1.0 0.0 0.0 M 2 3.5 0.7 20.3 2 1.5 0.5 33.3 2 5.0 0.0 0.0 Z 2 A.5 2.1 A7.1 2 1.8 0.0 0.0 2 5.0 0.0 0.0 J 1 3.0 .... 2 2.1 O.A 20.0 2 5.0 0.0 0.0 H 2 7.5 2.1 28.3 2 1.9 0.2 12.1 2 5.0 0.0 0.0 35 3 A.O 1.7 A3.3 3 2.0 0.3 15.0 3 5.0 0.0 0.0 S 3 5.0 1.0 20.0 3 1.6 0.1 6.3 3 5.0 0.0 0.0 T 3 5.0 1.0 25.0 2 1.8 0.1 3.9 3 5.0 0.0 0.0 K 2 5.5 2.1 38.6 1 1.8 .... 2 5.0 0.0 0.0 a 2 A.5 0.7 15.8 2 2.0 0.1 3.5 2 5.0 0.0 0.0 N A A.5 0.6 12.7 A 1.8 0.1 27.8 A 5.0 0.0 0.0 0 2 9.0 0.0 0.0 1 1.7 ... ----- 2 5.0 0.0 0.0 163

T A B U 17 (Continued)

POP. rQ Ct LVS. BELOW rD rfi S: PSD. VESTURE ______I I CV______X I CT______X a_____ CT

L 2 5.5 2.1 38.6 2 1.5 0.1 4.7 2 5.0 0.0 0.0 A 2 11.5 2.1 18.4 2 1.7 0.0 0.0 2 5.0 0.0 0.0 C 3 6.0 1.0 16.7 3 2.1 0.7 33.8 3 5.0 0.0 0.0 B 2 8.0 0.0 0.0 2 2.3 0.6 24.8 2 5.0 0.0 0.0 BB 2 6.5 0.7 10.9 2 2.1 0.4 20.0 2 5.0 0.0 0.0 164 arranged geographically. This study includes 45 populations of which 15 are field mass collections, 30 are herbaria specimens compilations.

Hybrid Blade Petiole V Leaves Head Peduncle Index W/L Blade L 10 Cm Width Vesture Value Ratio Ratio Below

1 37+ 12+ 4- 1 .6- 1 .0 -1 .5 2 34-37 9-12 4-5 1 .6-1.8 1.5-2.5 3 31-34 6 -9 5-6 1 .8-2.0 2.5-3*5 4 28-31 3 -6 6-7 2 .0 -2.2 s.s-'t.s 5 28- - -3 7+ 2 .21- **■•5-5*0

The resulting histogram (Figure 48b) has three major groups and other subgroups. Similar results were obtained in a study by Dyar (1939) in which she used different characters and specimens, and individuals rather than populations.

The group with hybrid index values 7-9 (-11) are B. pumila and "dealbata" specimens; group H.I. 11-14 are hybrid populations with variously combined characters and key to

B. x betonicifolla: group H.I. (14-) 18-23 nre B. texana specimens Missouri, Arkansas, and south-coastal Texas, the

"longifolia" group; group H.I. 20-23 are B. texana specimens from mid-northern Texas, the and Oklahoma.

Detailed descriptions of taxa appeared earlier.

No population scored as "ideal" F^ hybrid (H.I. 15)» but the hybrid plants group is shifted somewhat toward B. pumila.

One could interpret that there is a gene flow to both parental types, but a greater flow to B. pumila. The "longifolia" Figure *t8 « Histograms of Hybrid Index values of eastern and western taxa complexes* a* Eastern taxa* b* Western taxa*

165 NUMBER a Figure 48 Figure YRD NC VLI TIX T VALUIS INDCX HYBRID 166 group night be the result of different ecology* but plants taken from the field and placed under greenhouse conditions did not change markedly. The variability is tentatively interpreted as introgression with a freer gene flow among taxa of western complex than among taxa of the eastern complex; i.e. there appear to be less effective internal and/or external barriers to hybridization in the western group than in the eastern one.

The hybridization pattern is complex and a more nearly complete analysis is not justifiable until more data are available• Figure 49. Distribution asp of sastsrn taxa. Ecological zones redrawn from map by Hedlund and Janssen (1963)« courtesy of USDA, U.S. Government Printing Office.

168 Figure . Distribution map of eastern taxa. Ecological zones redrawn from map by Hedlund and Janssen (1963 ), courtesy of USDA, U.S. Qovernment Printing Office.

168 169

Figure ^9 Figure 50. Distribution map of western taxa. Ecological zones redrawn from map by Kedlund and Jassen (1963)i courtesy of USDA, U.S. Government Printing Office. For explanation of zones, see Figure 49.

170 r— ;

X

Figure 50 SUMMARY

The genus Berlandiera is tentatively revised to include

four species: B. texana. B. lyrata. B. pumila, and B. sub- acaulis: one variety of these, B. lyrata var. macrophylla; and

two new hybrid complexes, B. x humilis and j3. x betonicifolla.

Biosystematically, it becomes one comparium of one distinct ecospecies and three less distinct ecospecies. Variety macrophylla was not investigated experimentally.

All species are highly interfertile. Internal barriers include partial F^ and F^ generation sterility, notably those in which B. lyrata var, lyrata is a parent, probably due in part to meiotic irregularities. In addition to self-sterility of all members of all taxa, there is partial or complete inter­ sibling and "reciprocal offspring sterility" in certain taxa combinations.

Introgressive hybridization occurs among sympatric species, more commonly in the western "texana-pumila" complex than in the eastern "subacaulis-pumila" complex.

External barriers to hybridization are apparently in­ effective except for (1) ecological separation of high dry limestone soils habitats of B, lyrata var. lyrata from lowland sandy habitats of adjacent J3. texana; and (2) geographic iso­ lation between eastern and western segments of B. pumila ani among all taxa that are not sympatric.

172 173

Evidence ia presented favoring the hypothesis of origin of Berlandiera from Silphium or Silphium-like prototype via tetraploidy. Its origin is probably recent with as yet poorly developed barriers among most taxa.

Experimental investigations of B. lyrata var. macrophylla and of all backcrosses are necessary before complete taxonomic revision can be made. BIBLIOGRAPHY

Anderson, Edgar* 19^9* Introgressive Hybridisation. J. Wiley, N.Y.

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Clausen, Jens. 1951* Stages in the Evolution of Plant Species, Cornell University Press, N.Y.

Cruden, Robert William. I960. Biosystematic studies in the genus Silphium: the perfoliate taxa. Unpub. M.S. thesis. The Ohio State University. Columbus.

Dean, Donald S. 1959* Distribution of Tetraploid and Diploid Tradescantia ohioensls in Michigan and Adjoining Areas. American Midland Naturalist 6l(l): 20^-209.

De Candolle, Augustin-Pyramus. 1 8 3 6 . Prodromus Systematis Naturalis Regni Vegetabilis. ars V: 517* Paris.

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De Lessert, Benj. 1 8 3 9 . Icones Selaectae Plantarurn. Quas in Prodromo Systematis Universalis, ex Herbariis Parisiensibus Prasertim ex Lessertiano. Vol. IV. Exhibens Compositas. Paris. 17*+ 175

Dyar, Mary Hapner. 1959* A Biosystematic Study of the Genua Berlandiera. Unpub. M.S. theaia. The Ohio State University. Coltuabua.

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Fiaher, T. Bichard and Bobert C. Cruden. 1962. Chromosome Numbera and Obaervationa in the Genua Silphium. Ohio J. Sci. 62(5): 258-259.

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Geiser, Samuel Wood. 1937* Naturalists of the Frontier Chapter 3: In Defense of Jean Louis Berlandier. University Press, Dallas.

Gray, Asa. 1884. Synoptic Flora of North America. Vol. I, Part II: 24-3* July. Ivison, Biakeman, Taylor and Co., N.Y.

Hedlund, Arnold and Paul Janssen. 1963* Major Forest Types in the South. Map. Forest Service, USDA. Washington, D.C.

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Lanjouw, J. and F.A. Stafleu. eds. 1964. Regnum Vegitabile Vol. 31* Part I: Index Herbariorum. The Herbaria of the World. Fourth Edition. The International Bur. for Plant and Nomen. Utrecht, Netherlands.

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Ii Donald John Pinkava, was born on August 29* 1933* in

Cleveland, Ohio. The majority of my life unfolded in nearby

Orange Village, where I attended Orange Elementary, Junior High, and High Schools, receiving a diploma in June, 1951* The fall of the same year I was enrolled at the Ohio State University where I shall have earned all three degrees: B.Sc. in education in 1933, H.Sc. in botany in 1961, and Ph.D. in botany in 1964.

This study was interrupted by a teaching position held at Solon

High School (Cuyahoga County, Ohio) from 1933 to I960.

At The Ohio State University I was granted a Du Pont

Fellowship, a National Science Foundation Co-operative Fellow­ ship, a graduate assistantship and an instruetorship in botany, the last at the Lakewood Branch.

This fall (1964) I shall join the Botany Staff at Arizona

State University as Assistant Professor.

178