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cotonnière, cottonrose, cottonweed, filzkraut, fluffweed

Habit Annuals, perennials, subshrubs, or shrubs (often ± woolly annuals 1–10 cm). Annuals, 1–50(–70) cm.
Stems

1, erect, or 2–10+, ascending to prostrate.

Leaves

basal and/or cauline; usually alternate, rarely opposite;

petiolate or sessile (bases often decurrent onto stems);

blade margins usually entire, rarely denticulate (often revolute or involute, faces often tomentose or woolly and/or glandular-pubescent).

cauline; alternate;

blades subulate to obovate.

Involucres

0 or inconspicuous.

Receptacles

usually flat to convex, sometimes conic to ± columnar, usually epaleate, sometimes paleate (paleae sometimes enfolding pistillate florets).

fungiform to obovoid (heights 0.4–1.6 times diams.), glabrous.

Ray florets

usually 0 (sometimes peripheral florets pistillate and corollas ± zygomorphic and bearing a ± flat limb and interpreted by some as ray florets).

Pistillate florets

14–45+.

Peripheral (pistillate) florets

(often 100+ in disciform heads) in 1–3+ series;

corollas (usually present) yellow or purplish to whitish, sometimes red-tipped (actinomorphic and filiform).

Bisexual florets

2–10;

corolla lobes 4–5, ± equal.

Phyllaries

sometimes 0 (apparent phyllaries interpreted as outer receptacular paleae), usually persistent (often wholly or partially white or brightly colored, sometimes woolly, at least proximally and/or medially), usually (12–30+) in 3–10+ series, distinct, and unequal, sometimes in 1–2 series, distinct and subequal to equal, medially herbaceous to membranous or scarious, margins and/or apices usually notably scarious.

0, vestigial, 1–4 (unequal), or 4–6 (equal).

Calyculi

0.

Heads

usually heterogamous (usually disciform, rarely “quasi-radiate”), sometimes homogamous (discoid, sometimes pistillate or functionally staminate), usually borne in corymbiform, paniculiform, or racemiform arrays, sometimes in glomerules, sometimes aggregated into second-order heads, rarely borne singly.

usually in glomerules of 2–10(–14) in racemiform to paniculiform or dichasiiform arrays, or some [all] borne singly.

Cypselae

usually monomorphic within heads, mostly ovoid to obovoid and compressed or obcompressed, not beaked or apically attenuate, bodies smooth or ± papillate or muriculate, often 2-, 3-, or 5-ribbed (glabrous or hairy, seldom glandular, sometimes with myxogenic hairs or papillae);

pappi (rarely 0) persistent or readily falling, usually of barbellulate, sometimes plumose bristles, sometimes of scales (scales often aristate), sometimes combinations of bristles and scales.

brown, dimorphic: outer compressed to obcompressed, obovoid to ± cylindric, straight or curved, longer than inner, abaxially gibbous, faces glabrous, smooth, shiny;

inner ± terete, faces glabrous, usually papillate to muriculate, sometimes smooth, dull;

corolla scars apical to subapical;

pappi: outer pistillate 0, inner pistillate and bisexual of (11–)13–28+ bristles (visible in heads).

Disc

(inner) florets (sometimes 1–10, usually more) bisexual or functionally staminate;

corollas yellow or purplish, sometimes red-tipped, usually actinomorphic, lobes usually (4–)5, usually ± deltate, rarely lance-ovate to lanceolate;

anther bases usually ± tailed [not tailed], apical appendages mostly ovate to lance-ovate or linear (usually flat);

styles abaxially glabrous or (mostly distally) papillate, branches ± linear, adaxially stigmatic in 2 lines from bases to apices, apices truncate or truncate-penicillate, appendages essentially none.

Pistillate

paleae (except usually innermost) readily or tardily falling, erect to ascending;

bodies with 5+ nerves (nerves ± parallel, obscure), lanceolate to ovate or boat-shaped, ± saccate most of lengths (obcompressed to terete, sometimes ± galeate, each ± enclosing a floret);

wings erect to incurved (apices blunt).

Innermost

paleae usually all pistillate, in some species bisexual and pistillate, persistent, usually 5 or 8, spreading (and enlarged) in fruit, surpassing other pistillate paleae;

bodies lanceolate to ovate.

Functionally

staminate florets 0.

x

= 14.

Asteraceae tribe Gnaphalieae

Logfia

Distribution
Nearly worldwide; with centers of concentration in southern Africa and Australia; in both Old World and New World; the greater numbers of genera and species in the southern hemisphere
from USDA
North America; n Mexico; Europe; Asia; n Africa; introduced in South America; Pacific Islands; probably elsewhere
[BONAP county map]
Discussion

Genera ca. 187, species ca. 1240 (19 genera, 111 species in the flora).

Traditionally, taxa included here in Gnaphalieae have been treated in Inuleae within Gnaphaliinae (cf. A. A. Anderberg 1991; K. Bremer 1994). Gnaphalieae include everlastings and helichrysums, which have brightly colored, persistent phyllaries and are much used in dried floral arangements, and other ornamentals, e.g., species or cultivars from Anaphalis, Antennaria, and Leontopodium (edelweiss). Some species of Facelis, Gamochaeta, Gnaphalium, Helichrysum, and Pseudognaphalium are widespread as weeds.

Phyllaries in most Gnaphalieae are usually more or less herbaceous to more or less cartilaginous medially and/or proximally and membranous to scarious laterally and distally. The herbaceous to cartilagionous area of a phyllary is usually somewhat thicker than the rest and such areas are called stereomes. Stereomes may be more or less divided or not and may be more or less glandular or not. The membranous to scarious portion of a phyllary distal to the stereome is sometimes called a lamina (not to be confused with corolla laminae of other groups). The phyllaries, or laminae are often colored and may be more or less opaque or more or less hyaline.

Surfaces of cypselae of Gnaphalieae may be smooth, longitudinally ridged, or papillate (with minute bumps or projections from one or both ends of each epidermal cell; see A. A. Anderberg 1991). In addition, the cypselae may be glabrous or may bear myxogenic (producing mucilage when wetted) or non-myxogenic “twin-hairs.” The twin-hairs may be relatively long and form sericeous to strigillose induments. Very short, globose to clavate twin-hairs (lengths equaling or not much greater than diameters) are characteristic of some taxa and have sometimes been called “papillae” in descriptions of members of Gnaphalieae. Cypselae with such very short twin-hairs are described here as minutely hairy and the hairs are referred to as papilliform.

Stuartina hamata Philipson, a member of Gnaphalieae, was collected near a wool mill in South Carolina in 1957 (G. L. Nesom 2004c). It is native to Australia and may be characterized as annuals, prostrate, mostly 6–12 cm across, ± woolly, leaves cauline (crowded near heads), petioles basally dilated, blades suborbiculate, heads ca. 3 mm (borne in glomerules), phyllaries ovate to lanceolate, inner uncinate, florets 5 (outer 4 pistillate, inner bisexual), cypselae 0.8–1 mm, epappose.

Genera 97–105 below (genera following second lead 3 in key to genera, members of Filagininae in a narrow sense) have exceptionally small heads and florets (even for composites) and are closely similar in expressions of some characters. Together, as found in the flora, they may be characterized as:

Annuals, taprooted, usually arachnoid-sericeous to lanuginose throughout, sometimes glabrescent proximally on stems and/or on adaxial faces of leaves. Leaves usually sessile, sometimes obscurely petiolate (usually gradually larger and more crowded distally, sometimes again smaller among heads, where referred to as capitular leaves); blades 1-nerved, bases usually ± cuneate, sometimes rounded, margins entire. Heads disciform. Involucres absent, vestigial, or inconspicuous, often simulated by leaves or paleae. Phyllaries 0, vestigial, or 1–6. Receptacles paleate (at least peripherally), usually glabrous among paleae (bristly in Hesperevax). Florets pistillate, functionally staminate (usually referred to as staminate), or bisexual; corollas whitish, usually distally yellowish, reddish, brownish, or purplish.

Leaves of Filagininae that immediately subtend heads and/or glomerules are here called capitular leaves. Flowering branches may also immediately subtend heads or glomerules; if so, capitular leaves collectively subtend such branches and their heads/glomerules, and heads/glomerules appear to be sessile in forks of pseudo-dichotomies or -polytomies. Sometimes capitular leaves subtend only glomerules and not individual heads and individual heads may be difficult to distinguish within glomerules.

Paleae subtend all or at least some florets in members of Filagininae. They are referred to as bisexual, pistillate, or staminate paleae depending on sorts of florets subtended. Pistillate paleae persistent or shed with cypselae, usually incurved over inner (bisexual or functionally staminate) florets at flowering; margins usually thinner, ± scarious, forming wings (sometimes gradually and obscurely so); wings recurved to erect to incurved or inflexed; abaxial faces usually ± lanuginose to sericeous, sometimes glabrate or glabrous; apices rounded or obtuse to acuminate or aristate. Bisexual and/or staminate paleae usually persistent, sometimes falling in fruit, sometimes 0 (most Filago, Logfia, and Micropus, all Psilocarphus; then simulated by pistillate paleae), usually erect at flowering, incurved, erect, or spreading in fruit, sometimes enlarging as cypselae mature (then adaxially lanuginose to sericeous), shorter than or surpassing pistillate paleae; bodies ± ovate or lanceolate to spatulate (saccate in Micropsis); margins rarely forming wings; abaxial faces lanuginose to sericeous or nearly glabrous; apices usually entire, sometimes 2–3-fid, sometimes aristate to spinose (uncinate in Ancistrocarphus filagineus).

Corolla scars on cypselae of Filagininae may be offset adaxially to subapical or ± median positions and may be diagnostic for certain taxa (corolla attachments usually appear to be apical before ovaries mature).

All or outer pistillate florets of Filagininae lack pappi. Inner pistillate, bisexual, and staminate florets have pappi 0 or of 1–28+, whitish, fragile (easily broken or detached) or readily falling, ± barbellate to barbellulate or smooth (sometimes smooth only distally) bristles in 1 series. At bases of bristles, barbs are sometimes notably longer and finer, sometimes wavy or curled, and more patent than antrorse and may interweave, resulting in proximal coherence of adjacent bristles (e.g., in Logfia); cohering bristles may be shed in groups or complete rings and separate subsequently.

In descriptions of Filagininae, median refers to areas or positions about midway between a base and corresponding apex, and medial refers to areas or positions ± centered between opposing lateral edges.

As noted by A. Cronquist (1950) for Psilocarphus, the wings or apices of pistillate paleae of Filagininae are incurved during flowering, guide styles over bisexual or functionally staminate florets, and, likely, enforce nearly obligate within-head geitonogamy. Reproductive isolation created by this self-pollinating syndrome may allow interspecific or intergeneric hybrids, when fertile, to persist and become independently reproducing species among their parental taxa (J. D. Morefield 1992, 1992b).

Birds harvest shoots of Logfia, Micropus, Psilocarphus, and Stylocline species, presumably for nesting materials (neststraw is common name for Stylocline) and may be significant in shorter-distance dispersal of some taxa. The light, fluffy paleae enclosing epappose cypselae of the same genera aid in wind dispersal, as suggested by A. Cronquist (1950).

Different species and genera of Filagininae often grow together and are frequently mixedand/or misidentified on herbarium sheets. Young or stunted plants often will not fit keys and descriptions; whenever practicable in attempting identifications of members of Filagininae, use well-developed plants with at least some heads in fruit. Some diagnostic characters require careful evaluation of structures within heads, usually with magnification; for example, anther tips and corolla lobes may be similar in color and shape and may be difficult to distinguish.

Good illustrations of most North American Filagininae may be found in L. Abrams and R. S. Ferris (1923–1960, vol. 4), G. Beauverd (1913), or J. C. Hickman (1993).

Genera ca. 187, species ca. 1240 (19 genera, 111 species in the flora)

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species 12 (6 in the flora).

See discussion of Filagininae following the tribal description (p. 385).

Logfia occurs in dry open habitats of Mediterranean, semiarid, arid, and, sometimes, humid temperate climates. The three introduced species (L. arvensis, L. gallica, L. minima) do not appear to be aggressively invasive. Logfia filaginoides (formerly known as Filago californica) is somewhat weedy in the more mesic, coastal portions of its range, where it often mixes about equally with L. gallica. Some specimens of L. filaginoides, L. depressa, and L. minima may be difficult to identify without full comparison to descriptions.

Usually included in Filago in North America, Logfia (including Oglifa) has been separated for some decades in Old World treatments. Contrary to G. Wagenitz (1969), I agree with J. Holub (1976, 1998) that the two genera warrant separation. Priority of Logfia over Oglifa at generic rank is ambiguous, however, depending on whether Logfia was validated in 1819 or 1822 (J. D. Morefield 2004). Because uncertainty remains, I here preserve current usage of Logfia, pending a proposal to conserve the name.

Morphologic evidence suggests Logfia is basal in Filagininae, with ancestral, sister, and/or reticulate relationships to Filago and Stylocline (J. D. Morefield 1992). Logfia depressa appears to be transitional toward Stylocline. Ancestors of Logfia probably resembled Gnaphalium palustre Nuttall, which is frequently misidentified as a member of Filagininae. Logfia is most easily recognized by outer epappose florets subtended by saccate paleae, prominent pappi on inner pistillate and bisexual florets, and innermost paleae open, persistent, spreading.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Heads usually discoid (unisexual or nearly so, staminate or pistillate; plants unisexual or nearly so; predominantly pistillate heads rarely with 1–9 central, functionally staminate florets; predominantly staminate heads rarely with 1–4+ peripheral, pistillate florets; involucres mostly 6–10 mm)
→ 2
1. Heads usually disciform (plants not unisexual; heads mostly alike, each with 4–200+ pistillate and 1–200+ bisexual or functionally staminate florets; heads rarely discoid in Xerochrysum, which has involucres 10–30 mm and brightly colored phyllaries in 3–8+ series)
→ 3
2. Plants (0.2–)4–25(–70) cm; basal leaves usually present at flowering (withering before in A. geyeri); pappus bristles (at least pistillate) usually basally connate or coherent.
Antennaria
2. Plants mostly 20–80(–120+) cm; basal leaves usually withering before flowering; pappus bristles distinct or basally connate
Anaphalis
3. Annuals, biennials, perennials, or subshrubs; receptacles epaleate; cypselae all pappose
→ 4
3. Annuals; receptacles ± paleate (all or at least the outermost florets each subtended by a palea); cypselae (all or at least the outermost) epappose
→ 11
4. Pistillate florets fewer than bisexual
→ 5
4. Pistillate florets more numerous than bisexual
→ 6
5. Subshrubs; heads in glomerules in corymbiform arrays; involucres campanu- late, 4–8 mm
Helichrysum
5. Annuals, biennials, or perennials; heads borne singly or 2–3 in loose, corymbi- form arrays; involucres ± hemispheric, 10–30 mm
Xerochrysum
6. Annuals; pappi persistent; pappus bristles ± plumose
Facelis
6. Annuals, biennials, or perennials; pappi readily falling; pappus bristles barbellate to barbellulate
→ 7
7. Heads in spiciform or subcapitate arrays or in glomerules in continuous or interrupted, usually spiciform, sometimes paniculiform, arrays (terminal glomerules in depauperate plants); cypselae ± papillate (papillae or papilliform hairs myxogenic) or strigillose (hairs not myxogenic); pappus bristles basally connate, falling readily (in groups or rings; distinct in Omalotheca supina)
→ 8
7. Heads usually in ± capitate clusters (subtended by leafy bracts) or corymbiform or paniculiform (often bracteate) arrays; cypselae usually glabrous or minutely hairy or papillate (papillae or papilliform hairs not myxogenic), sometimes minutely roughened and/or with 4–6 longitudinal ridges; pappus bristles distinct (falling separately) or basally coherent (falling in groups or rings)
→ 9
8. Annuals or short-lived perennials; phyllaries in 3–7 series; pistillate florets 50–130+; cypselae ± papillate (papillae or papilliform hairs myxogenic)
Gamochaeta
8. Perennials; phyllaries in 2–3 series; pistillate florets 35–70+; cypselae strigillose (hairs not myxogenic)
Omalotheca
9. Involucres narrowly campanulate to cylindric; phyllaries mostly stramineous to brownish, sometimes purplish to pinkish (hyaline, stereomes not glandular)
Euchiton
9. Involucres narrowly to broadly campanulate to cylindric; phyllaries white, rosy, tawny, or brown (opaque or hyaline, stereomes usually glandular)
→ 10
10. Annuals, (1–)3–30 cm; heads usually in ± capitate clusters (in axils of leaves or bracts), sometimes in spiciform glomerules; cypselae usually glabrous, sometimes minutely hairy or papillate (hairs or papillae not myxogenic)
Gnaphalium
10. Annuals, biennials, or perennials, (4–)15–150(–200) cm; heads usually in glomerules in corymbiform or paniculiform arrays, sometimes in terminal clusters; cypselae usually smooth, sometimes papillate-roughened and/or with 4–6 longitudinal ridges, usually glabrous (papilliform hairs in P. luteoalbum)
Pseudognaphalium
11. Bisexual florets (1–)2–10(–11), pappi of (11–)13–28+ bristles visible in heads; functionally staminate florets 0
→ 12
11. Bisexual florets 0 or 2–7, pappi 0; functionally staminate florets 0 or 2–12, pappi 0 or of 1–10(–13) bristles hidden in heads
→ 13
12. Receptacles fungiform to obovoid (heights 0.4–1.6 times diams.); most pistillate paleae ± saccate, each ± enclosing a floret, apices blunt; innermost paleae spreading in fruit; cypselae dimorphic (outer longer than inner)
Logfia
12. Receptacles cylindric to clavate (heights 5–15 times diams.); most pistillate paleae open to ± folded (at most each enfolding, not enclosing a floret; apices acuminate to aristate); innermost paleae erect to ascending in fruit; cypselae mono- morphic (outer ± equaling inner)
Filago
13. Pistillate paleae open most of lengths, flat or concave to loosely folded (not enclosing florets); pappi 0
→ 14
13. Pistillate paleae saccate most of lengths (each enclosing a floret, outermost rarely open); pappi 0 or of 1–10(–13) bristles
→ 16
14. Bisexual paleae saccate, each enclosing a floret, apices 2-fid or 3-fid; cypselae (at least outer) strigose; coastal Texas
Micropsis
14. Bisexual or staminate paleae flat to concave, not enclosing florets, apices entire; cypselae glabrous; central and western North America
→ 15
15. Receptacles glabrous; pistillate paleae falling (all or the inner together); staminate (or bisexual) paleae: bodies ± spatulate (apices scarcely enlarged); central North America
Diaperia
15. Receptacles bristly; pistillate paleae persistent; staminate paleae: bodies obovate (apices enlarged); California and Oregon
Hesperevax
16. Staminate paleae 5(–7), ± spreading proximally, enlarged in fruit (apices incurved to uncinate); pistillate paleae with 3, ± parallel (prominent) nerves; cypselae: corolla scars apical
Ancistrocarphus
16. Staminate paleae 0 or 1–4, erect, not enlarged in fruit (apices erect); pistillate paleae with 5+, reticulate (and prominent) or ± parallel (and obscure) nerves; cypselae: corolla scars subapical to ± median
→ 17
17. Cauline leaves mostly opposite; phyllaries 0; pistillate paleae (nerves reticulate, prominent): wings inflexed (and ± lateral); staminate paleae 0; pappi 0.
Psilocarphus
17. Cauline leaves mostly alternate; phyllaries 0 or 1–6; pistillate paleae (nerves parallel, obscure): wings ± erect (or subapical); staminate paleae 1–4 and/or staminate pappi of (0–)1–10(–13) bristles
→ 18
18. Pistillate paleae (obcompressed to terete, not galeate): wings ± erect (and apical); receptacles cylindric to clavate (heights 2.8–8 times diams.); phyl- laries 0 or 1–4 (similar to paleae); cypselae: corolla scars subapical
Stylocline
18. Pistillate paleae (compressed, galeate): wings ± erect (and lateral) or inflexed (and subapical); receptacles depressed-spheric to obovoid (heights 0.5–1.8 times diams.); phyllaries 4–6 (unlike paleae); cypselae: corolla scars ± lateral
Micropus
1. Pistillate florets: inner (0–)1–2 pappose; branches usually leafless between proximal forks, becoming purplish to black, glabrescent (longest capitular leaves ± linear, 2–5 times head heights)
L. arizonica
1. Pistillate florets: inner (4–)8–35 pappose; branches ± leafy between proximal forks, remaining greenish to grayish or whitish, arachnoid-sericeous to lanuginose
→ 2
2. Pistillate paleae ± vertically ranked, outer inflexed 70–90° proximally, gibbous, ± galeate, bodies ± bony; leaves subulate or broader; phyllaries usually 5, equal
→ 3
2. Pistillate paleae spirally ranked, outer incurved 20–60°, somewhat gibbous, not galeate, bodies ± cartilaginous or chartaceous; leaves not subulate; phyllaries 0, vestigial, or 1–4, unequal
→ 4
3. Leaves narrowly elliptic to narrowly ovate; longest capitular leaves 0.8–1.5 times head heights; innermost paleae ± 8, spreading in 2 series; bisexual corollas 1.6–2.1 mm; pappi mostly of 13–16 bristles falling in 1s or 2s
L. minima
3. Leaves mostly subulate; longest capitular leaves 2–5 times head heights; innermost paleae ± 5, spreading in 1 series; bisexual corollas 2.2–3 mm; pappi of 18–28+ bristles falling in complete or partial rings
L. gallica
4. Outer pistillate paleae: distal 5–10% of lengths glabrous abaxially, wings obscured by indument; outer 2–4(–6) pistillate florets epappose; innermost paleae ± 8, spread- ing in 2 series
L. arvensis
4. Outer pistillate paleae: distal 15–50% of lengths glabrous abaxially; wings prominent; outer 7–13 pistillate florets epappose; innermost paleae ± 5, spreading in 1 series
→ 5
5. Outer pistillate paleae: bodies ± cartilaginous; bisexual corolla lobes mostly 4, bright reddish to purplish; inner cypselae mostly papillate, pappi of 17–23+ bristles falling in complete or partial rings; stems typically ± erect; capitular leaves mostly acute
L. filaginoides
5. Outer pistillate paleae: bodies (except midnerves) chartaceous; bisexual corolla lobes mostly 5, yellowish to brownish; inner cypselae mostly smooth, pappi of 11–15 bristles falling in 1s or 2s; stems typically ± prostrate; capitular leaves obtuse
L. depressa
Source FNA vol. 19, p. 384. Treatment authors: Theodore M. Barkley†, Luc Brouillet, John L. Strother. FNA vol. 19, p. 443. Treatment author: James D. Morefield.
Parent taxa Asteraceae Asteraceae > tribe Gnaphalieae
Subordinate taxa
Anaphalis, Ancistrocarphus, Antennaria, Diaperia, Euchiton, Facelis, Filago, Gamochaeta, Gnaphalium, Helichrysum, Hesperevax, Logfia, Micropsis, Micropus, Omalotheca, Pseudognaphalium, Psilocarphus, Stylocline, Xerochrysum
L. arizonica, L. arvensis, L. depressa, L. filaginoides, L. gallica, L. minima
Synonyms Oglifa
Name authority Cassini ex Lecoq & Juillet: Dict. Rais. Term. Bot., 296. (1831) Cassini: Bull. Sci. Soc. Philom. Paris 1819: 143. 1819 (as “genre ou sous-genre”): in F. Cuvier, Dict. Sci. Nat. ed. 2, 23: 564. (1822)
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