Flora of North America North of Mexico

FNANM Comp Guide, 2000/06/22, p. 1 of 32

Special Guidelines

Contributors of Treatments of Composites

In most respects, happily, most manuscripts of treatments for Flora of North America North of Mexico (FNANM) are prepared according to requests and admonitions given in the primary Guide for Contributors. Some frequent deviations are noted here, as reminders, before the special guidelines for contributors of treatments of Composites.

Descriptions should be parallel as to structure and content (Guide 5.1) for all taxa of a given rank within a single more inclusive taxon. For example: all varieties of a particular species; all species of a particular section; all genera within a particular subfamily; etc. All traits used in keys should be treated in descriptions at appropriate rank (also in section 5.1). If ovary position is used in keying genera of Floraceae, then each generic description for Floraceae should include indication of ovary position. If key to the species of genus Xxxxx has pollen color as a discriminator, then pollen color should appear in all descriptions of species of genus Xxxxx. See also 9.1.3. If all species of genus Xxxxx have red corollas (and corolla color is important), include corolla color in description of genus Xxxxx and do not state “corollas red” for each species of genus Xxxxx.

Size and number. In descriptions and in keys, adjectives of scale such as small, short, medium, large, and tall and adjectives of number such as few, many, numerous, and several are not very informative. In Guide 12.2, we are admonished to avoid such adjectives and we have examples to guide us: 1--2 mm, not “small” 75--100, not “many” ca. 200, not “numerous” In one context, “small” leaves may be 3--5 cm and in another context, leaves 3--5 cm may be “large.” Disc florets 20--30 may equate to “disc florets few” in one genus and to “disc florets many” in another. And so on. In keys and descriptions, use measurements and use numerical ranges or estimates of numbers rather than adjectives such as small, teensy, many, slew, middlin’, humongous, beaucoups, etc.

In descriptions of plants, “upper” and “lower,” “above” and “below,” and “dorsal” and “ventral” sometimes mean different things to different botanists. For FNANM, use proximal/proximally and distal/distally, base/basal/basally and apex/apical/apically, or abaxial/abaxially and adaxial/adaxially, as appropriate. See Guide 12.4.2.

Synonymy. With few exceptions, only basionyms and names used as accepted names in a listed flora should be included in synonymies in FNANM (see Guide 8.6). A name of a forma should not be included in a synonymy unless it is the basionym of the accepted name of the taxon. FNANM Comp Guide, 2000/06/22, p. 2 of 32

In keys (Guide 7.1) with unequal branches, place the lead with fewer subordinate couplets before the lead with more subordinate couplets.

Compositae Particulars

In preparing contributions for FNANM, contributors of treatments of composites are asked to follow the recommendations given here, in addition to those given in the primary Guide for Contributors. Both sets of recommendations are intended to promote consistency in use of descriptive terms and in ordering of elements in treatments in general and in descriptions in particular. In addition to the following, see also: description and discussion of the family Compositae and the Sample Treatment for Compositae.

Draft descriptions provided to contributors by Taxon Editors should be checked for accuracy; corrections of errors are to be made by contributors. Some draft descriptions may include superfluous material (which should be removed) or may lack essential material (which should be added). Additions and deletions are to be made by contributors.

Maintain distinctions between inflorescences and capitulescences. The inflorescence of a composite is a head (or capitulum). Where deemed desirable or necessary to refer to the overall arrangement of heads on a plant, the term capitulescence is used in conjunction with terms such as corymbiform, paniculiform, spiciform, etc.

For FNANM, follow conventions used by George Bentham and others (cf. Bentham 1873, p. 363 ff.) in morphological interpretation of florets and corollas. A typical disc corolla comprises a proximal tube and a distal limb; the tube is that part of the corolla proximal to the insertion of the filaments and the limb is that part distal to insertion of the filaments (cf. Bentham 1873, p. 365). The limb of a disc corolla comprises a proximal throat and (3--)5 distal lobes. The distinction between tube and limb is maintained regardless of the external form of the corolla (i.e., the corolla may be dilated distal to, at, or proximal to the point of insertion of the filaments; the region of the dilation, if any, does not always correspond to the interface of tube and throat). A ray corolla comprises a cylindrical tube and a flattened lamina; the lamina of a ray corolla has 0--3(--4) terminal lobes or teeth. The corolla of a ligulate floret (a floret from a liguliflorous head; i.e., a member of Lactuceae) comprises a tube and a ligule; a ligule has 5 terminal lobes or teeth. The length of any floret includes its ovary and is greater than the length of the corolla of that floret. Ovaries are usually white to ochroleucous at anthesis; therefore, corollas, not florets, are described as yellow, orange, red, purple, etc. FNANM Comp Guide, 2000/06/22, p. 3 of 32

Descriptions of genera and/or species of composites may include (in addition to elements usual for all flowering plants) description of expressions of the characters listed following. Not all of the following need be included in each description; so far as practicable, contributors should keep descriptions short and diagnostic.

Capitulescences. Whether spiciform, paniculiform, corymbiform, etc.

Peduncles. Length, indument, . . . .

Involucres. Shape and overall dimensions, or, at least, length or diam.

Phyllaries. Number (as "20--35" rather than "few", "several", "many", etc.). Number of series, i.e., 1, 1--3, 3--5+, etc. Color, shape, dimensions, texture, indument.

Receptacles. Shape, dimensions (if useful), indument and/or “ornamentation” of surface (if any).

Paleae (receptacular bracts). Duration, color, shape, dimensions, texture, indument, . . . .

Ray florets. Number of series, if more than one. Number in each series or total number (as numerical range such as 3--5, 20--30+, etc., not as "many", "numerous", etc.). Indication as to pistillate and fertile, styliferous and sterile, or neuter. Corolla color. Tube length. Lamina dimensions.

Disc florets. Number (20--50, 50--150+, etc.; not "few", "many", etc.). Indication as to bisexual or functionally staminate. Corolla color. Tube length; indument. Throat shape and dimensions; indument. Lobe number, shape, and dimensions; indument.

Stamens. Color, dimensions, bases/tails, apical appendages, etc.

Styles (of fertile disc florets). Stigmas, appendages, etc.

Cypselae. Color, shape, dimensions, indument and/or “ornamentation.”

Pappi. Color, number(s) of elements, and form(s)/shapes of elements, dimensions.

* * * * * * FNANM Comp Guide, 2000/06/22, p. 4 of 32

Contributors are encouraged to append brief notes, as appropriate, for genera or species that are significant economically (e.g., as sources of allergens, as noxious weeds, as toxic plants, as plants of special horticultural interest, etc.).

* * * * * *

An artificial key to genera plus a synoptical key to tribes and keys to subtribes and genera within tribes will be included in the treatment of Compositae. For the artificial key, 15 "groups" are recognized (copy herewith). Contributors are asked to verify that all appropriate (and no inappropriate) groups are listed for each genus they treat (see draft descriptions supplied by Taxon Editor). Copies of the keys will be made available to any contributor who wishes to "test" the keys.

References. Bentham, G. 1873. Notes on the classification, history, and geographical distribution of Compositae. J. Linn. Soc. Bot. 13: 335--582. Bremer, K. 1994. Asteraceae: Cladistics and classification. Portland, Ore.: Timber Press. Heywood, V. H., J. B. Harborne, and B. L. Turner, eds. 1977 [1978]. The biology and chemistry of the Compositae. London: Academic Press.

Characterizations of 15 artificial groups of FNANM comps.

Group 1. Heads liguliflorous (Lactuceae).

Group 2. "Ray" corollas or all corollas distinctly bilabiate (most of Mutisieae).

Group 3. Receptacles paleate; heads radiate; pappi none or nearly so.

Group 4. Receptacles paleate; heads radiate; pappi wholly of bristles.

Group 5. Receptacles paleate; heads radiate; pappi wholly, or partially, of scales or awns.

Group 6. Receptacles paleate; heads eradiate; pappi none or nearly so.

Group 7. Receptacles paleate; heads eradiate; pappi wholly of bristles.

Group 8. Receptacles paleate; heads eradiate; pappi wholly, or partially, of scales or awns.

Group 9. Receptacles epaleate; heads radiate; pappi none or nearly so.

Group 10. Receptacles epaleate; heads radiate; corollas of ray florets white, pink, or purple; pappi wholly of bristles.

Group 11. Receptacles epaleate; heads radiate; corollas of ray florets yellow, orange, red, or brown; pappi wholly of bristles.

Group 12. Receptacles epaleate; heads radiate; pappi wholly, or partially, of scales or awns.

Group 13. Receptacles epaleate; heads eradiate; pappi none or nearly so.

Group 14. Receptacles epaleate; heads eradiate; pappi wholly of bristles.

Group 15. Receptacles epaleate; heads eradiate; pappi wholly, or partially, of scales or awns. FNANM Comp Guide, 2000/06/22, p. 5 of 32

Description and Discussion of the Family

Asteraceae Link, nom. alt. [ Compositae Giseke, name conserved] · Composite Family

Theodore M. Barkley

Luc Brouillet

John L. Strother

Herbs (annual, biennial, or perennial), shrubs, vines, or trees. Leaves opposite or alternate, sometimes in basal rosettes, rarely in whorls, rarely stipulate, usually petiolate, sometimes sessile, sometimes with bases decurrent on the stems; blades usually simple, often pinnatifid or palmatifid, rarely compound. Inflorescences: indeterminate heads (also called capitula); each head comprising a surrounding involucre of phyllaries (involucral bracts), a receptacle, and (1--)5--200+ florets; individual heads sessile or each borne on a peduncle; heads borne singly or variously arranged in usually determinate, rarely indeterminate, capitulescences. Involucres sometimes subtended by calyculi (sing. calyculus, collective term for bractlets immediately subtending the phyllaries proper). Phyllaries borne in 1 or more series proximal to (i.e., outside of or abaxial to) the florets. Receptacles usually flat to convex, sometimes conical or columnar, either paleate (bearing paleae or receptacular bracts that individually subtend some or all of the florets) or epaleate (lacking paleae); epaleate receptacles sometimes bristly or hairy or bearing subulate enations among the florets. Florets bisexual, pistillate, functionally staminate, or neuter (also called neutral); calyx none, instead of sepals each ovary bears (usually) a pappus (pl. pappi) of bristles, awns, or scales (sometimes in combination in a single pappus); petals united, corollas (3--4--)5-merous, actinomorphic or zygomorphic (one or both kinds in a single head, see descriptions of radiate, discoid, liguliflorous, disciform, and radiant following); stamens (4--)5, alternate with corolla lobes, filaments inserted on corollas, usually distinct, anthers introrse, usually connate and forming a tube around the style (rarely filaments connate and anthers distinct; e.g., Ambrosiinae); ovaries inferior, 2-carpellate, and 1-locular with 1, basally attached, anatropus ovule that develops into an exalbuminous seed with a straight embryo; styles 1 in each bisexual, functionally staminate, or pistillate floret; each style usually ringed at base by a nectary, distally 2-branched with stigmatic papillae borne on adaxial face of each branch in 2, separate or contiguous bands or in 1, continuous band (in functionally staminate florets, styles often not branched), styles variously truncate or appendaged beyond the stigmas, usually papillate to hirsute distally on abaxial faces. Fruits (technically cypselae, historically called achenes) usually dry with relatively thick, tough, pericarps, sometimes rostrate (beaked) or alate (winged), often dispersed with aid from pappi. Genera ca. 1500, species ca. 23,000 (XXX genera and XXXX species in the flora): nearly worldwide, especially rich in numbers of species and/or in numbers of plants in arid and semiarid regions of subtropical and lower to middle temperate latitudes.

SELECTED REFERENCES: Bentham, G. 1873. Notes on the classification, history, and geographical distribution of Compositae. J. Linn. Soc. Bot. 13: 335--582. Bremer, K. 1994. Asteraceae: Cladistics and classification. Portland, Oregon. Heywood, V. H., J. B. Harborne, and B. L. Turner. [eds.] 1977[1978]. The biology and chemistry of the Compositae. London.

Inflorescences of composites are called heads (or capitula, sing. capitulum). Arrays of heads are called capitulescences; the arrays of heads on composites correspond to arrays of individual flowers (inflorescences) on plants of other families. Terms for kinds of capitulescences parallel terms for kinds of inflorescences: corymbiform, paniculiform, racemiform, spiciform, etc. In radiate heads, peripheral florets (ray florets) in one or more series have corollas with zygomorphic limbs and may be pistillate, or styliferous and sterile, or neuter; the central florets (disc florets) in radiate heads have actinomorphic corollas and may be bisexual or functionally staminate. In liguliflorous heads, all florets are bisexual and fertile and have zygomorphic corollas (ligulate florets); liguliflorous heads are characteristic of Lactuceae and FNANM Comp Guide, 2000/06/22, p. 6 of 32 are found in no other composites. In discoid heads, all florets have actinomorphic corollas and all florets are either bisexual and fertile or all are either functionally staminate or pistillate (in monoecious or dioecious taxa). In disciform heads, all florets have actinomorphic corollas and peripheral florets (in one or more series) are pistillate and, usually, have relatively slender corollas; the central florets of disciform heads are usually bisexual, sometimes functionally staminate. In radiant heads, all florets have actinomorphic corollas and the peripheral florets have much enlarged corollas and may be bisexual, pistillate, or neuter; the central florets of radiant heads are usually bisexual. Some composites have peripheral, bisexual florets with slightly to strongly zygomorphic corollas (e.g., some members of Chaenactis, Lessingia, Thymophylla, et al.); heads of such plants do not quite conform to any of the five types just described. Heads with all florets of one sexual form (bisexual, pistillate, or functionally staminate) are called homogamous (discoid, liguliflorous, and some radiant heads) and heads with florets of two or more sexual forms are called heterogamous (radiate, disciform, and some radiant heads). Receptacles may bear paleae (i.e., some or all florets are individually subtended by a bractlet called a palea or receptacular bract). Collectively paleae have been called “chaff” and paleate receptacles have been described as “chaffy.” Some receptacles that lack paleae (epaleate heads) may bear subulate enations (e.g., Gaillardia), or stiff bristles or subulate to linear scales (e.g., some Cardueae), or fine hairs (e.g., some Anthemideae). After fruit-fall, epaleate receptacles, or paleate receptacles after paleae have fallen, may be smooth or variously pitted (alveolate, foveolate, etc.). The terms tube, throat, and limb have been variously used in descriptions of corollas of composites. In FNANM, in actinomorphic corollas of bisexual and functionally staminate disc florets, the tube is the part of the corolla proximal to the insertion of the staminal filaments and the limb is the part that is distal to insertion of the filaments. The limb comprises, proximally, the throat and, distally, the lobes. The distinction between tube and throat hinges on insertion of filaments, not on external morphology. The relatively flat portion of a corolla of a ligulate floret from a liguliflorous head is called a ligule; it terminates in 5 teeth or lobes. The relatively flat portion of a corolla of a ray floret is called a lamina; it terminates in 0--3(--4) teeth or lobes. More or less bilabiate corollas are characteristic of some members of Mutisieae and are seldom found in members of other tribes. Fruits of composites have been called "achenes" because they resemble true achenes. Achenes are dry, hard, single-seeded fruits derived from unicarpellate, superior ovaries. Ovaries of composites are bicarpellate and inferior. Fruits derived from ovaries of composites are called cypselae (sing. cypsela). Morphology of an ovary of a composite at anthesis is often markedly different from the morphology of the mature fruit (cypsela) derived from that ovary. References to cypselae in keys and descriptions in FNANM refer to mature fruits, not to ovaries at anthesis. Shapes of cypselae have been used in distinguishing among species, genera, and even subtribes of composites. In most composites, cypselae are ± isodiametric in cross section. In some composites, cypselae are characteristically ± lenticular to elliptic in cross section. Such cypselae are said to be compressed or laterally flattened if the longer axis of the cross section is ± parallel to a radius of the head (e.g., Verbesina spp.). Cypselae are said to be obcompressed or radially flattened if the shorter axis of the cross section is ± parallel to a radius of the head (e.g., Coreopsis spp.). Pappi show a great range of diversity and are often diagnostic for recognition of taxa, especially at rank of genus and below. The various forms of pappus elements intergrade. For keys and descriptions in FNANM, the following arbitrary distinctions are made: Cross sections of bristles and awns are ± circular or polygonal and have the longer diameter no more than 3 times the shorter diameter. Pappus elements with "flatter" cross sections (i.e., longer diameter more than 3 times the shorter diameter) are called scales, regardless of relative overall lengths and widths of the elements. As used here, "subulate scale" means much the same as "flattened bristle" of some authors. Pliable to stiff pappus bristles with diameters less than 50 m are called fine bristles; pliable to stiff bristles with diameters greater than 50 m are called coarse bristles. Rigid pappus elements with ± circular or polygonal cross sections greater than 100 m in diameter are called awns. Bristles, awns, and scales may be smooth or variously finely to coarsely barbed or plumose. A scale of a pappus may terminate in one or more bristle-like or awn-like appendages; such scales are said to be aristate. References to pappi in keys and descriptions in FNANM refer to pappi of mature fruits, not to pappi of ovaries at anthesis. FNANM Comp Guide, 2000/06/22, p. 7 of 32

Sample Treatment for Compositae

XXX.4.23 Compositae - Heliantheae - Pectidinae

George Spelvin

Pectidinae ('Pectideae') Lessing, Linnaea 5: 134. 1830

Tageteae Cassini, J. Phys. Chim. Hist. Nat. Arts 88: 162. 1819

Herbs (annual or perennial), subshrubs, or shrubs. Leaves opposite or alternate, usually simple and entire, toothed, or pinnatifid, sometimes pinnate, the blades or lobes usually bearing pellucid oil-glands (cavities

0.1--1.5 mm diam. filled with strong-scented, essential oils).

Capitulescences corymbiform or heads borne singly. Involucres narrowly cylindric to hemispheric, often subtended by calyculi (each calyculus comprising 1--10+ bracts immediately subtending the phyllaries proper).

Phyllaries 5--21 in 1--2 series, all free or the adjacent weakly to strongly connate, usually some or all bearing pellucid oil-glands. Receptacles flat to convex, sometimes hairy, minutely setose, or foveolate; paleae none. Ray florets 0 or 3--20+, mostly 5, 8, or 13, pistillate, fertile. Disc florets mostly 10--80+, bisexual, fertile; anther sacs truncate to weakly sagittate at base; style branches linear or reduced to knobs (in Pectis), variously truncate, appendaged (penicillate to subulate); stigmatic papillae in 2 bands. Cypselae narrowly prismatic to cylindric or clavate, lengths usually

2--4(--12) times widths, faces commonly ribbed or striate, glabrous or variously hairy; pappi of scales, bristles (these all free or variously connate basally in fascicles of 3--10 bristles), or awns, or combinations thereof, sometimes none or reduced to coronas.

Genera 19, species ca. 200 (9 genera, ??? species in the flora): all native to New World, mostly North America, especially Mexico. FNANM Comp Guide, 2000/06/22, p. 8 of 32

Pectidinae is sometimes treated as a distinct tribe, Tageteae.

Regardless of rank, the group is clearly coherent and distinct from other groups of composites. Schizogenous glands that contain strong-scented essential oils (terpenoids) are found in the leaves and/or phyllaries of all members of the group; such oil-glands are not known to occur in other composites.

SELECTED REFERENCE Strother, J. L. 1977[1978]. Tageteae---systematic review. In: V. H. Heywood, J. Harborne, and B. L. Turner, eds. The biology and chemistry of the Compositae. London. Chap. 27, pp. 769--783.

1. Phyllaries weakly to strongly connate 1/3 or more their

lengths.

2. Involucres mostly fusiform or cylindric, rarely

campanulate; pappi of 10 or fewer elements, 1--2(0--5)

elements linear to subulate or awnlike, the others ±

erose, often some or all ± connate...... xx8. Tagetes, p. xxx.

2. Involucres mostly obconic to campanulate or hemispheric;

pappi of 8--22, free elements (8--12, erose, lacerate, or

1--3-aristate scales and/or 10--20 scales each comprised

of 3--11, basally connate bristles), all or the alternating

elements similar.

3. Stems mostly 3--30 cm; involucres 3--7 mm; phyllaries

strongly connate 3/4+ their lengths, margins of the

outer seldom free more than 1/2 their lengths; receptacles

smooth or shallowly alveolate...... xx7. Thymophylla, p. xxx.

3. Stems mostly 3--8+ dm; involucres 7--20 mm; phyllaries

weakly to strongly connate 1/3--3/4+ their lengths,

often separating in age, margins of the outer usually FNANM Comp Guide, 2000/06/22, p. 9 of 32

free to base; receptacles fimbrillate to setose.

4. Leaf blades mostly pinnate or pinnatifid (if not divided,

then oblanceolate to ovate), oil-glands marginal and

subterminal at bases and tips of lobes; lobes of

disc corollas lance-linear to subulate.

...... xx5. Adenophyllum, p. xxx.

4. Leaf blades not divided, linear, toothed, oil-glands

scattered along both sides of midveins; lobes of disc

corollas lance-deltate...... xx6. Dysodiopsis, p. xxx.

1. Phyllaries free to base or nearly so.

5. Leaves opposite, not divided, bristly-ciliate at base; style

branches very short, knob-like...... xx9. Pectis, p. xxx.

5. Leaves opposite or alternate, usually divided, if not, then

little, if at all, bristly-ciliate at base; style branches

linear.

6. Involucres calyculate; pappi partially or wholly of

scales or fascicles of bristles, each fascicle comprising

5--15, ± free or ± connate bristles.

7. Ray corollas yellow to orange; pappi of 15--20,

similar fascicles, each comprising 5--10, basally

connate bristles...... xx1. Dyssodia, p. xxx.

7. Ray corollas white to pink or magenta; pappi of

5--6 fascicles (each of 7--15, ± free bristles) plus

5--6, lanceolate, 1-aristate scales.

...... xx2. Nicolletia, p. xxx.

6. Involucres ecalyculate; pappi of fine to coarse, free

bristles. FNANM Comp Guide, 2000/06/22, p. 10 of 32

8. Phyllaries 8--14, strongly carinate; heads radiate;

pappi of 30--40 bristles in 1 series.

...... xx3. Chrysactinia, p. xxx.

8. Phyllaries 5--10, weakly, if at all, carinate; heads

discoid; pappi of 25--100+ bristles in 1 or more

series...... xx4. Porophyllum, p. xxx.

[Only four of the nine genera of Pectidinae that occur in the flora are included in this Sample Treatment.] FNANM Comp Guide, 2000/06/22, p. 11 of 32 xx1. DYSSODIA Cavanilles, Descr. Pl. 202. 1802 · [Greek dysodia, a bad odor]

John L. Strother

Herbs, annual [perennial]. Stems erect to decumbent, 1--3(--9) dm.

Leaves all opposite or the distal alternate, blades pinnatisect with linear to linear-cuneate lobes, little, if at all, bristly at base, on teeth, or at tips of lobes, oil-glands submarginal. Capitulescences: heads borne singly.

Calyculi of 1--9, deltate to linear bractlets. Involucres ± campanulate,

5--10 mm. Phyllaries 6--12, free to base or nearly so, bearing oil-glands.

Receptacles convex, fimbrillate to minutely setose. Ray florets pistillate, fertile, corollas yellow to orange. Disc florets bisexual, fertile; corollas yellow to orange, cylindric, lobes 5, deltate to lance-deltate; style branches linear, apically truncate to deltate, papillate. Cypselae obpyramidal to obconic; pappi of 15--20, unequal to subequal fascicles, each comprising 5--10, basally connate bristles. x = 13.

Species 4 (1 in the flora): United States, Mexico, Guatemala.

SELECTED REFERENCES: Strother, J. L. 1969. Systematics of Dyssodia

Cavanilles (Compositae: Tageteae). Univ. Calif. Publ. Bot. 48: 1--88.

Strother, J. L. 1986. Renovation of Dyssodia (Compositae: Tageteae). Sida

11: 371--378.

1. Dyssodia papposa (Ventenat) A. Hitchcock, Trans. Acad. Sci. St. Louis 5:

503. 1891 · Dogweed

Tagetes papposa Ventenat, Descr. Pl. Nouv. plate 36 + text. 1801

Stems erect to decumbent, 1--7 dm. Leaves 15--50 

10--40 mm, lobes 11--15, linear to linear-cuneate, 5--20  1--3 mm, glabrous or sparsely hairy, dotted with oil-glands. Peduncles 1--5(--10) mm.

Involucres obconic to campanulate, 6--10 mm. Phyllaries 6--12, oval to oblanceolate, each bearing 1--7 oil-glands. Ray florets 5--8; corollas FNANM Comp Guide, 2000/06/22, p. 12 of 32 yellow-orange, tubes 2--3 mm, laminae 1.5--2.5  1--2 mm. Disc florets 12--50; corollas pale yellow, ca. 3 mm. Cypselae 3--3.5 mm; pappi of ca. 20, unequal fascicles 1--3 mm, each comprising 5--10, basally connate bristles. 2n = 26

[Strother 1969].

Flowering summer--fall. Grasslands, often ruderal, in fields and along roadways; 0--2000 m; Ont.; Ariz., Ark., Colo., Idaho, Ill., Ind., Iowa,

Kans., Ky., Maine, Mich., Minn., Mo., Mont., Nebr., N.H., N.Mex., N.Y.,

N.Dak., Ohio, Okla., Pa., S.Dak., Tenn., Tex., Utah, Vt., Wis., Wyo.; Mexico.

Dyssodia papposa is (or was) introduced in California. It is locally naturalized in South America and, perhaps, naturalized in much of its range within the flora. FNANM Comp Guide, 2000/06/22, p. 13 of 32

XX5. ADENOPHYLLUM Persoon, Syn. Pl. 458. 1807 · [Greek adeno, gland, and phyllon, leaf]

John L. Strother

Herbs (annual or perennial), subshrubs. Stems erect, (2--)3--7+ dm, variously branched. Leaves all opposite or the distal alternate, blades usually pinnate or pinnatifid, bases, rachises, and teeth usually bristly- ciliate, oil-glands often borne at bases of lobes and subterminally in tips.

Capitulescences: heads borne singly. Calyculi of 12--16+, often bristle- tipped, bractlets. Involucres campanulate to hemispheric, 7--20 mm.

Phyllaries 8--20+, weakly connate 1/3--3/4 their lengths (separating in age), margins of the outer free to base or nearly so, bearing oil-glands.

Receptacles convex, fimbrillate to setose; paleae none. Ray florets pistillate, fertile, corollas yellow to orange. Disc florets bisexual, fertile; corollas yellow to orange, cylindric, lobes 5, lance-linear to subulate; style branches linear, apically conic and papillate or subulate and hispidulous. Cypselae obpyramidal; pappi of 8--20 scales, each scale muticous or comprising 3--11, basally connate bristles. x = 7, 13.

Species 10 (3 in the flora): southwestern United States, Mexico, and

Central America.

11: 371--378.

1. Annuals; bractlets of calyculi eglandular; outer pappus

scales erose, the inner each unequally 3-aristate... 3. A. wrightii

1. Perennials; bractlets of calyculi bearing oil-glands; each

pappus element comprising 5--11, basally connate bristles.

2. Leaves divided into 3--5, linear to cuneate or FNANM Comp Guide, 2000/06/22, p. 14 of 32

oblanceolate lobes; disc florets 25--40; pappi of 8--12

fascicles of bristles...... 1. A. porophylloides

2. Leaves not divided, blades ovate to oblanceolate; disc

florets 50--80; pappi of 15--20 fascicles of bristles.

...... 2. A. cooperi

1. Adenophyllum porophylloides (A. Gray) Strother, Sida 11: 377.

Dyssodia porophylloides A. Gray, Mem. Amer. Acad. Arts, ser. 2.

5:322. 1854

Subshrubs. Stems erect, 2--5+ dm. Leaves divided, 15--40 mm, lobes

3--5, linear to cuneate or oblanceolate with oil-glands at base of each and subterminal in tips. Peduncles 2--8 cm. Calyculi of 12--16, subulate, gland-bearing bractlets 3--8 mm. Involucres obconic, 10--15 mm. Phyllaries

12--20, lanceolate, separating in fruit. Ray florets 10--14; corollas yellow, becoming red-orange, tubes ca. 2 mm, laminae ca. 6  2 mm. Disc florets 25--40; corollas yellow-orange, 7--8 mm. Cypselae ca. 5 mm; pappi of

8--12 fascicles 7--8 mm, each comprising 7--11, basally connate bristles. 2n

= 26 [Turner and Flyr 1966].

Flowering spring and fall. Alluvial fans and rocky slopes; 0--1200 m;

Ariz., Calif., Nev.; Mexico (Baja California and Sonora).

2. Adenophyllum cooperi (A. Gray) Strother, Sida 11: 377. 1986 · E

Dyssodia cooperi A. Gray, Proc. Amer. Acad. Arts 9: 201. 1874

Subshrubs. Stems erect, 3--5+ dm. Leaves sessile, not divided, blades oblanceolate to ovate, 8--25  4--8 mm, sometimes toothed or obscurely lobed at base, bearing 1--2 pairs of oil-glands near base and 1 gland near tip.

Peduncles 6--15 cm. Calyculi of 12--22, linear-attenuate, gland-bearing bractlets 5--8 mm. Involucres obconic to campanulate, 15--18 mm. Phyllaries ca. 20, linear to lanceolate, separating in fruit. Ray florets usually FNANM Comp Guide, 2000/06/22, p. 15 of 32

7--13, rarely none; corollas yellow-orange, becoming red-orange, tubes ca. 5 mm, laminae 8--10  2.5--4 mm. Disc florets 50--80+; corollas yellow, 8--10 mm. Cypselae 5--7 mm; pappi of 15--20 fascicles 7--10 mm, each comprising

5--9, basally connate bristles. 2n = 26 [Strother 1969].

Flowering spring--fall. Sandy and gravelly soils of washes and alluvial fans; 0--1300 m; Ariz., Calif., Nev.

3. Adenophyllum wrightii A. Gray, Smithsonian Contr. Knowl. 5(6): 92. 1853

Dyssodia neomexicana (A. Gray) B. L. Robinson

Varieties 2 (1 in the flora): southwestern United States and Mexico.

3a. Adenophyllum wrightii A. Gray var. wrightii

Annuals. Stems erect, to 7 dm. Leaves divided, 25--35 mm, lobes 5--9, linear, each bearing an oil-gland at base. Peduncles 3--6 cm. Calyculi of

3--5, subulate, eglandular bractlets 3--5 mm. Involucres obconic to broadly campanulate, 7--8 mm. Phyllaries 9--13, narrowly to broadly oblanceolate, remaining connate in fruit. Ray florets ca. 8; corollas yellow-orange, tubes ca. 3 mm, laminae ca. 2  1 mm. Disc florets 30--50; corollas pale yellow, often tipped with crimson, 4--5 mm. Cypselae 3.5--4.5 mm; pappi of 10, outer, erose scales 0.7--1 mm and 10, inner, unequally 3-aristate scales

5--6.5 mm. 2n = 14 [Turner and Flyr 1966].

Flowering fall. Apparently in wet places in otherwise arid areas;

1500--2600 m; Ariz., N.Mex.; Mexico (Chihuahua).

[Reference for database: Turner, B. L. and L. D. Flyr. 1966. Chromosome numbers in the Compositae. X. North American species. Amer. J. Bot. 53:

24--33.] FNANM Comp Guide, 2000/06/22, p. 16 of 32

XX6. DYSODIOPSIS (A. Gray) Rydberg in N. L. Britton et al., N. Amer. Fl. 34:

171. 1915 · [Greek dysodia + opsis, indicating similarity to Dyssodia

Cavanilles] |E|

Hymenatherum Cassini sect. Dysodiopsis A. Gray, Smithsonian Contr.

Knowl. 3(5): 116. 1852

John L. Strother

Herbs, annuals, short-lived perennials. Stems erect, 4--8+ dm, distally branched. Leaves all opposite or the distal alternate; blades not divided, linear, coarsely toothed, obscurely bristly-ciliate at base, oil- glands scattered along both sides of midveins. Capitulescences: loosely corymbiform. Calyculi of 5--8, subulate or pinnatisect bractlets. Involucres campanulate, 9--12 mm. Phyllaries 10--12, strongly connate 3/4+ their lengths, margins of the outer free to base, bearing submarginal oil-glands.

Receptacles convex, fimbrillate or foveolate. Ray florets pistillate, fertile, corollas lemony to greenish yellow. Disc florets bisexual, fertile; corollas dull yellow, cylindro-funnelform, lobes 5, lance-deltate; style branches linear, apically truncate, papillate, each with a fine, fragile, setaceous appendage. Cypselae obpyramidal; pappi of 10--12, 1(--3)-aristate scales. x = 13.

Species 1.

11: 371--378.

1. Dysodiopsis tagetoides (Torrey & A. Gray) Rydberg in N. L. Britton et al.,

N. Amer. Fl. 34: 171. 1915 · |E|

Dyssodia tagetoides Torrey & A. Gray, Fl. N. Amer. 2: 361. 1842 FNANM Comp Guide, 2000/06/22, p. 17 of 32

Stems erect, 4--8+ dm. Leaves 4--9 cm  2--6 mm with subopposite teeth, glabrous. Peduncles 3--5 cm. Calyculi of 5--8 gland-bearing bractlets 8--12 mm. Involucres obconic-cylindric, 9--12 mm. Phyllaries 10--12, lance- linear. Ray florets 7--12; corolla tubes ca. 3 mm, laminae linear-elliptic,

10--15  3--6 mm. Disc florets 20--40+, corollas ca. 4 mm. Cypselae 3--3.5 mm; pappus scales narrowly lanceolate, ± aristate, 1--2.5 mm. 2n = 26

[Strother 1969].

Flowering late spring--summer. Limestone outcrops and derived soils;

100--700 m; Okla., Tex. FNANM Comp Guide, 2000/06/22, p. 18 of 32 xx7. THYMOPHYLLA Lagasca, Gen. Sp. Nov. 25. 1816 · [Greek thymon, thyme, and phyllon, leaf]

Hymenatherum Cassini  Dyssodia Cavanilles sect. Hymenatherum (Cassini)

Strother; D. Cavanilles sect. Gnaphalopsis (A. P. de Candolle) Strother

John L. Strother

Herbs (annuals, short-lived perennials), shrubs. Stems erect to spreading or decumbent, mostly 5--30 cm, variously branched. Leaves opposite or alternate, simple or divided, the blades or lobes spatulate to linear or filiform, little, if at all, bristly-ciliate at base, oil-glands scattered in lamina or submarginal. Capitulescences: heads borne singly. Calyculi none or of 1--8, deltate to linear bractlets. Involucres campanulate to obconic,

3--7 mm. Phyllaries 8--13(--22), strongly connate 3/4+ their lengths, seldom with outer margins free more than 1/2 their lengths (exceptions: T. aurea, T. pentachaeta var. belenidium), bearing oil-glands. Receptacles convex, smooth or shallowly alveolate. Ray florets pistillate, fertile, corollas usually yellow to orange, rarely white. Disc florets bisexual, fertile; corollas yellow to orange, cylindric, lobes 5, deltate; style branches linear, apically deltate or conic, papillate. Cypselae obpyramidal, obconic, or cylindro-clavate; pappi of 10(--20), free scales, each scale erose or 1--5- aristate or comprising a fascicle of 5--9 basally connate bristles. x = 8.

Species 13 (8 in the flora): mostly southwestern United States and

Mexico (some adventive beyond the flora and Mexico).

SELECTED REFERENCES Strother, J. L. 1969. Systematics of Dyssodia

Cavanilles (Compositae: Tageteae). Univ. Calif. Publ. Bot. 48: 1--88;

11: 371--378; Strother, J. L. 1989. Chromosome numbers in Thymophylla

(Compositae: Tageteae). Sida 13: 351--358.

1. Leaf blades entire or toothed or distally trifid, not divided into FNANM Comp Guide, 2000/06/22, p. 19 of 32

3--13+ linear lobes; bractlets of calyculi usually more than

1/2 as long as phyllaries.

2. Plants green, usually puberulent, sometimes glabrescent,

rarely glabrous; leaves mostly opposite...... 4. T. acerosa

2. Plants ashy-white, arachnose, lanate, floccose, or

tomentose; leaves mostly alternate.

3. Stems spreading, often prostrate; leaf blades spatulate.

...... 2. T. micropoides

3. Stems erect; leaf blades linear to filiform, sometimes

trifid distally...... 3. T. tephroleuca

1. Leaf blades divided into 3--13+ linear to filiform lobes

(blades linear, entire in T. tenuiloba var. wrightii); bractlets

of calyculi none or less than 1/2 as long as phyllaries.

4. Short-lived perennials (sometimes flowering first year);

leaves mostly opposite.

5. Plants green, usually puberulent to canescent, sometimes

glabrescent or glabrous; calyculi of 1--5, deltate

bractlets...... 5. T. pentachaeta

5. Plants ashy-white, tomentose; calyculi of 1--3

subulate bractlets...... 1. T. setifolia

4. Annuals (rarely persisting); leaves mostly alternate

(sometimes opposite at 1--3 proximal nodes).

6. Calyculi of 3--8 bractlets; disc florets 50--100+.

...... 6. T. tenuiloba

6. Calyculi none or of 1--2 bractlets; disc florets

25--45.

7. Margins of outer phyllaries free nearly to their FNANM Comp Guide, 2000/06/22, p. 20 of 32

bases...... 7. T. aurea

7. Margins of outer phyllaries free less than 1/2+ their

lengths...... 8. T. concinna

1. Thymophylla setifolia Lagasca, Gen. Sp. Nov. 25. 1816

Dyssodia setifolia (Lagasca) B. L. Robinson

Varieties 2 (1 in the flora): southwestern United States and adjacent

Mexico.

1a. Thymophylla setifolia Lagasca var. greggii (A. Gray) Strother, ined.[to be published prior to use in flora]

Thymophylla greggii A. Gray var. greggii [created by Thymophylla greggii

A. Gray var. radiata A. Gray, Smithsonian Contr. Knowl. 3(5): 119. 1852];

Dyssodia setifolia (Lagasca) B. L. Robinson var. radiata (A. Gray) Strother;

D. greggii (A. Gray) B. L. Robinson

Short-lived perennials, ashy-white, tomentose. Stems spreading, mat- forming, to 15 cm. Leaves opposite, 6--12 mm, blades of the proximal divided into 3--7, stiff, filiform lobes from near the base, the distal entire, acerose. Peduncles 3--4 cm, glabrous or nearly so. Calyculi of 1--3, subulate bractlets less than 1/2 as long as phyllaries. Involucres obconic to campanulate, 3.5--4 mm, glabrous. Phyllaries 9--15, margins of the outer free less than 1/3 their lengths. Ray florets 7--10; corollas bright yellow, tubes ca. 1 mm, laminae 2--3  0.8--1.2 mm. Disc florets 20--40; corollas dull yellow, 2--2.5 mm. Cypselae 1.5--2.2 mm; pappi coronas of connate scales 0.3--0.5 mm.

Flowering summer. Rocky, calcareous outcrops and derived soils;

1200--1500+ m; N.Mex., Tex.; Mexico (Chihuahua and Coahuila).

2. Thymophylla micropoides (A. P. de Candolle) Strother, Sida 11: 377. 1986

Gnaphalopsis micropoides A. P. de Candolle, Prodr. 7: 258. 1838;

Dyssodia micropoides (A. P. de Candolle) Loesener FNANM Comp Guide, 2000/06/22, p. 21 of 32

Short-lived perennials, ashy-white, arachnose to floccose. Stems spreading, often prostrate, to 15+ cm. Leaves alternate, not divided, blades spatulate, 10--25  3--6 mm. Peduncles 0--5+ mm, tomentose. Calyculi of

3--5, linear bractlets more than 1/2 as long as phyllaries. Involucres obconic, 6--7 mm, tomentose. Phyllaries 12--14, margins of the outer free less than 1/5 their lengths. Ray florets 10--15; corollas bright yellow, becoming greenish, tubes 3 mm, laminae 5  3 mm. Disc florets ca. 60; corollas yellow, 3 mm. Cypselae 2.3--3 mm; pappi of 5, erose scales less than 1 mm alternating with 5, lanceolate, 1-aristate scales to 3 mm. 2n = 16

[Strother 1969].

Flowering through the year, following rains. Calcareous outcrops and derived soils; 50--200+ m; Tex.; Mexico (Coahuila, Nuevo León, and

Tamaulipas).

3. Thymophylla tephroleuca (S. F. Blake) Strother, Sida 11: 378.

Dyssodia tephroleuca S. F. Blake, J. Wash. Acad. Sci. 25: 320. 1935

Perennials, ashy-white, lanate to tomentose. Stems erect, 1--3 dm.

Leaves all or mostly alternate, blades not divided, linear to filiform, usually entire, sometimes trifid distally, 10--15  0.3--1 mm. Peduncles 1--3 cm, tomentose. Calyculi of 3--4, linear bractlets more than 1/2 as long as phyllaries. Involucres campanulate, 5--7 mm, tomentose. Phyllaries 12--13, margins of the outer free less than 1/4 their lengths. Ray florets 12--13; corollas golden yellow, tubes 2 mm, laminae 6--8  3--4 mm. Disc florets ca.

30; corollas yellow, 4.5--5 mm. Cypselae 3--4 mm; pappi of 10--11, subequal scales 4--5 mm, each 3--5-aristate. 2n = 16 [Strother 1969].

Flowering early spring and late summer, following rains. Light, sandy soils; 50--100 m; Tex. [probably in adjacent Mexico].

4. Thymophylla acerosa (A. P. de Candolle) Strother, Sida 11: 376. 1986 FNANM Comp Guide, 2000/06/22, p. 22 of 32

Dyssodia acerosa A. P. de Candolle, Prodr. 5: 641. 1836

Shrublets, green, puberulent, glabrescent, or glabrous. Stems erect, branched from base, to 25 cm. Leaves mostly opposite, blades undivided, linear to acerose, 10--18 mm. Peduncles 0--10 mm, puberulent or glabrous.

Calyculi of 5, lance-linear bractlets more than 1/2 as long as phyllaries.

Involucres campanulate to cylindric, 5--7 mm, puberulent or glabrous.

Phyllaries ca. 13, margins of the outer free less than 1/6 their lengths.

Ray florets 7--8; corollas lemon-yellow, tubes 2.5 mm, laminae 5--6  2--3 mm.

Disc florets 18--25+; corollas pale yellow, 3--4 mm. Cypselae 3--3.5 mm; pappi of ca. 20 fascicles, each fascicle 3--5-aristate, 3--4 mm. 2n = 16, 24

[Strother 1989].

Flowering through the year, mostly summer--fall. Calcareous outcrops and derived soils; 1000--2000+ m; Ariz., Nev., N.Mex., Tex., Utah; Mexico.

5. Thymophylla pentachaeta (A. P. de Candolle) Small, Fl. S.E. U.S. 1295. 1903

Hymenatherum pentachaetum A. P. de Candolle, Prodr. 5: 642. 1836;

Dyssodia pentachaeta (A. P. de Candolle) B. L. Robinson

Short-lived perennials, green, usually puberulent to canescent, sometimes glabrescent or glabrous. Stems erect or spreading, to 15(--25) cm.

Leaves opposite, 6--28 mm, blades divided into 3--11, linear to filiform, usually stiff, setiform lobes. Peduncles 2--10 cm, puberulent or glabrous.

Calyculi none or of 1--5, deltate bractlets less than 1/2 as long as phyllaries. Involucres obconic to campanulate or hemispheric, 4--6 mm, puberulent or glabrous. Phyllaries 12--21, margins of the outer ones free

1/5 to nearly all their lengths. Ray florets (8--)12--21; corollas yellow to orange-yellow, tubes 1.5--2 mm, laminae 2--6(--8)  1--3 mm. Disc florets

16--40 or 50--80; corollas yellow, 2--4 mm (tending to zygomorphy in peripheral florets in some plants). Cypselae 2--3 mm; pappi of 10, erose and/or aristate scales to 3 mm. FNANM Comp Guide, 2000/06/22, p. 23 of 32

Mexico; adventive in Argentina.

1. Leaves with terminal lobes longer than the laterals;

involucres cylindric, less than 3.5 mm diam.; disc florets

16--40...... 5d. Thymophylla pentachaeta var. hartwegii

1. Leaves with all lobes subequal; involucres obconic to

hemispheric, mostly greater than 3.5 mm diam.; disc florets

50--80.

2. Peduncles 2--5 cm; margins of outer phyllaries free

almost to base..... 5a. Thymophylla pentachaeta var. belenidium

2. Peduncles (4--)5--10 cm; margins of outer phyllaries

free less than 2/3 their lengths.

3. Leaves mostly 9--11-lobed; involucres glabrous or

nearly so...... 5b. Thymophylla pentachaeta var. pentachaeta

3. Leaves mostly 5--7(--10)-lobed; involucres densely

puberulent...... 5c. Thymophylla pentachaeta var. puberula

5a. Thymophylla pentachaeta (A. P. de Candolle) Strother var. belenidium (A.

P. de Candolle) Strother, Sida 11: 377. 1986

Dyssodia belenidium A. P. de Candolle, Prodr. 7: 292. 1838; D. pentachaeta (A. P. de Candolle) B. L. Robinson var. belenidium (A. P. de

Candolle) Strother; D. thurberi (A. Gray) B. L. Robinson

Leaves 10--20 mm, lobes 3--7, subequal. Peduncles 2--5 cm. Calyculi of 3--5 bractlets. Involucres obconic to campanulate, 4.5--6  4--5 mm, glabrous but for the ciliolate margins of the phyllaries. Phyllaries ca. 13, margins of the outer free almost to their bases. Disc florets 50--70. Pappi usually of 10, 3-aristate scales, sometimes of 5, muticous scales alternating with 5, 3-aristate ones. FNANM Comp Guide, 2000/06/22, p. 24 of 32

Flowering spring--summer. Sandy or rocky soils of washes and fans, commonly with limestone, often ruderal along roadways; 500--2500+ m; Ariz.,

Calif., Nev., N.Mex., Tex., Utah; Mexico; Argentina.

5b. Thymophylla pentachaeta (A. P. de Candolle) Small var. pentachaeta

Leaves 6--25 mm, lobes (3--)9--11, subequal. Peduncles 6--10 cm.

Calyculi of 3--5 bractlets. Involucres obconic to campanulate, 4.3--6  4--5 mm, usually glabrescent or glabrous, rarely hairy. Phyllaries 12--21, margins of the outer free ca. 1/2 their lengths. Disc florets 50--80. Pappi of 5, erose scales alternating with 5, 1--3-aristate scales. 2n = 32

[Strother 1989].

Flowering early spring and late summer, following rains. Calcareous outcrops and derived soils; 50--700+ m; Tex.; Mexico.

5c. Thymophylla pentachaeta (A. P. de Candolle) Small var. puberula (Rydberg)

Strother, Sida 11: 377. 1986

Thymophylla puberula Rydberg in N. L. Britton et al., N. Amer. Fl. 34:

177. 1915; Dyssodia puberula (Rydberg) Standley; D. pentachaeta (A. P. de

Candolle) B. L. Robinson var. puberula (Rydberg) Strother

Leaves 10--28 mm, lobes 3--7(--10), subequal. Peduncles 4--10 cm.

Calyculi none or of 1--3 bractlets. Involucres campanulate to hemispheric,

4.5--5.5  4--5 mm, densely puberulent. Phyllaries 12--17, margins of the outer free ca. 1/3 their lengths. Disc florets 50--70. Pappi of 5, erose scales alternating with 5, 1--3-aristate scales.

Flowering early spring and late summer following rains. Calcareous bluffs and slopes; 350--2200 m; Tex.; Mexico.

5d. Thymophylla pentachaeta (A. P. de Candolle) Small var. hartwegii (A.

Gray) Strother, Sida 11: 377. 1986 FNANM Comp Guide, 2000/06/22, p. 25 of 32

Hymenatherum hartwegii A. Gray, Smithsonian Contr. Knowl. 3(5): 117.

1852; Dyssodia hartwegii (A. Gray) B. L. Robinson; D. pentachaeta (A. P. de

Candolle) B. L. Robinson var. hartwegii (A. Gray) Strother

Leaves 12--34 mm, lobes 3--7, unequal (terminal lobe longer than laterals in each leaf). Peduncles 2--5 cm. Calyculi of 3--5 bractlets.

Involucres cylindric, 3--5  2--3.5 mm, sparsely puberulous, often glabrescent. Phyllaries ca. 13, margins of the outer free 1/3 or less their lengths. Disc florets 16--40. Pappi of 5, erose scales alternating with 5,

1--3-aristate scales.

Flowering early spring and late summer following rains. Calcareous outcrops and bluffs; 1500--2500 m; Ariz., N.Mex., Tex.; Mexico.

6. Thymophylla tenuiloba (A. P. de Candolle) Small, Fl. S.E. U.S. 1295. 1903

Hymenatherum tenuilobum A. P. de Candolle, Prodr. 5: 462. 1836;

Dyssodia tenuiloba (A. P. de Candolle) B. L. Robinson

Annuals (rarely persisting), green, glabrous or sparsely hirtellous.

Stems decumbent to erect, to 3 dm. Leaves mostly alternate (sometimes opposite at 1--3 proximal nodes), 12--28 mm, blades entire or divided into

7--15, linear to filiform, pliable lobes. Peduncles 3--8 cm, glabrous or hirtellous. Calyculi of 3--8 bractlets less than 1/2 as long as phyllaries.

Involucres obconic, 5--7 mm, glabrous or hirtellous. Phyllaries 12--22, margins of the outer free less than 1/5 their lengths. Ray florets 10--21; corollas yellow-orange, tubes ca. 2 mm, laminae 4--10  1.5--3 mm. Disc florets 50--100; corollas yellow, 2.5--4.5 mm (± zygomorphic in some plants).

Cypselae 2--3.5 mm; pappi of erose and/or aristate scales.

Varieties 4 (4 in the flora): all in Texas, 3 in Mexico.

1. Leaf blades usually entire, linear, rarely with 3--5, tooth-like

lobes...... 6d. Thymophylla tenuiloba var. wrightii

1. Leaf blades divided into 7--15, linear lobes. FNANM Comp Guide, 2000/06/22, p. 26 of 32

2. Pappus scales all 3--5-aristate.

...... 6a. Thymophylla tenuiloba var. tenuiloba

2. Pappus scales truncate to lanceolate or 1-aristate.

3. Pappi of 5, erose scales alternating with 5, 1-aristate

scales...... 6b. Thymophylla tenuiloba var. treculii

3. Pappi of 5--10, lanceolate to muticous scales, all

all erose or 1--3 each bearing a single, delicate

arista...... 6c. Thymophylla tenuiloba var. texana

6a. Thymophylla tenuiloba (A. P. de Candolle) Small var. tenuiloba ·

Dahlberg Daisy, Golden Fleece

Dyssodia tenuiloba (A. P. de Candolle) B. L. Robinson var. tenuiloba

Leaves 7--15-lobed. Pappi of 10--12, 3--5-aristate scales 2--3.5 mm.

2n = 16, 24, 32, 40 [Strother 1989].

Flowering early spring--late fall. Heavy soils or loams or sands, often with limestone, frequently ruderal along roadways and in other disturbed places; 0--300 m; Tex.; Mexico (Tamaulipas). Adventive in Ala.,

Fla., La., Miss., Bahamas, Cuba, Africa, Asia.

The names Dahlberg Daisy and Golden Fleece are recent coinages in the horticultural trade.

6b. Thymophylla tenuiloba (A. P. de Candolle) Small var. treculii (A. Gray)

Hymenatherum treculii A. Gray, Proc. Amer. Acad. Arts 19: 42. 1883;

Dyssodia treculii (A. Gray) B. L. Robinson; D. tenuiloba (A. P. de Candolle)

B. L. Robinson var. treculii (A. Gray) Strother

Leaves 7--15-lobed. Pappi of 5, oblanceolate, erose scales 0.8--1.1 mm alternating with 5, unequally aristate scales 2.5--3 mm. 2n = 32, 40

[Strother 1989]. FNANM Comp Guide, 2000/06/22, p. 27 of 32

Flowering early spring and late fall. Mostly sandy soils; 0--300 m;

Tex.; Mexico (Coahuila, Nuevo León, and Tamaulipas).

6c. Thymophylla tenuiloba (A. P. de Candolle) Small var. texana (Cory)

Dyssodia texana Cory, Rhodora 49: 162. 1947; D. tenuiloba (A. P. de

Candolle) B. L. Robinson var. texana (Cory) Strother

Leaves 7--15-lobed. Pappi of 5--10, lanceolate to muticous scales

0.7--1.2 mm, all erose or 1--3 each bearing a delicate arista 0.5--1 mm. 2n

= 16 [Strother 1969].

Flowering spring and fall. Limestone outcrops and derived soils in grasslands; 800--900 m; Tex.; Mexico (Coahuila and Nuevo León).

6d. Thymophylla tenuiloba (A. P. de Candolle) Small var. wrightii (A. Gray)

Strother, Sida 11: 378. 1986 · |E|

Hymenatherum wrightii A. Gray, Mem. Amer. Acad. Arts, ser. 2. 4: 89.

1849; Dyssodia wrightii (A. Gray) B. L. Robinson; D. tenuiloba (A. P. de

Candolle) B. L. Robinson var. wrightii (A. Gray) Strother

Leaves linear to spatulate, usually entire, rarely with 1--5, lateral, tooth-like lobes. Pappi of 10--12, unequally aristate scales to 3 mm. 2n =

16 [Strother 1989].

Flowering spring. Sandy soils; ca. 0--25 m; Tex.

7. Thymophylla aurea (A. Gray) E. Greene in N. L. Britton and A. Brown,

Illustr. Fl. n. U.S. 3: 435. 1898

Lowellia aurea A. Gray, Mem. Amer. Acad. Arts, ser. 2. 4: 91. 1849;

Dyssodia aurea (A. Gray) A. Nelson

Annuals, green, glabrous or puberulent. Stems branched from base, to

30 cm. Leaves alternate, 15--40 mm, blades divided into 5--13, linear lobes.

Peduncles 1--7 cm, glabrous or puberulent. Calyculi none or of 1--2, subulate bractlets less than 1/2 as long as phyllaries. Involucres obconic to campanulate, 5--6 mm, glabrous or puberulent. Phyllaries 12--15, margins FNANM Comp Guide, 2000/06/22, p. 28 of 32 of the outer free almost to base. Ray florets 8--12; corollas bright yellow, tubes 1.5 mm, laminae 4--6  2--3 mm. Disc florets 30--45; corollas yellow, 3 mm. Cypselae 3 mm; pappi of erose or 3--5-aristate scales.

Mexico.

The two varieties of Thymophylla aurea are sometimes found in mixed populations, which may include intermediate plants.

1. Pappi of 8--10, erose scales...... 7b. Thymophylla aurea var. aurea

1. Pappi of 18--20, 3--5-aristate scales.

...... 7a. Thymophylla aurea var. polychaeta

7a. Thymophylla aurea (A. Gray) E. Greene var. polychaeta (A. Gray) Strother,

Sida 11: 376. 1986

Hymenatherum polychaetum A. Gray, Smithsonian Contr. Knowl. 3(5): 116.

1852; Thymophylla polychaeta (A. Gray) Small; Dyssodia polychaeta (A. Gray)

B. L. Robinson; D. aurea (A. Gray) Nelson var. polychaeta (A. Gray) M. C.

Johnston

Plants to 3 dm. Peduncles 2--7 cm. Pappi of 18--20,

3--5-aristate scales 2--3 mm. 2n = 16 [Strother 1969].

Flowering late summer. Grasslands and Larrea-desert; 1000--1500+ m; N.Mex., Tex.; Mexico (Chihuahua, Coahuila, and Durango).

7b. Thymophylla aurea (A. Gray) E. Greene var. aurea

Dyssodia aurea (A. Gray) A. Nelson var. aurea

Plants to 2 dm. Peduncles 1--3 cm. Pappi of 8--10, erose scales

0.3--0.6 mm. 2n = 16 [Powell in Strother 1969].

Flowering summer. Swales in grasslands; 1500--2000 m; Colo., Kans.,

N.Mex., Tex.; Mexico (Chihuahua).

8. Thymophylla concinna (A. Gray) Strother, Sida 11: 376. 1986 FNANM Comp Guide, 2000/06/22, p. 29 of 32

Hymenatherum concinnum A. Gray, Synop. Fl. N. Amer. 1(2): 453. 1886;

Dyssodia concinna (A. Gray) B. L. Robinson

Annuals, green, glabrous. Stems spreading to ascending or erect,

3--12+ cm. Leaves alternate, 10--15 mm, blades divided into (3--)5--9, linear to filiform lobes. Peduncles 1--5+ mm, glabrous. Calyculi none or of

1--2, subulate bractlets less than 1/2 as long as phyllaries. Involucres obconic to campanulate, 5--6.5 mm, glabrous. Phyllaries 8--16, margins of the outer free less than 1/2 their lengths. Ray florets 9--12; corollas white to pale yellow, tubes 1.5--2.5 mm, laminae 3--4  1--3.5 mm. Disc florets ca. 25; corollas pale yellow, 3--4.5 mm. Cypselae 2.5--3 mm; pappi of 10--15 fascicles, each comprising 5--9, basally connate bristles 1--2.5 mm. 2n = 16 [Strother 1969].

Flowering spring, following rains. Sandy washes and flats; 50--100 m;

Ariz.; Mexico (Sonora).