Asteraceae tribe Gnaphalieae
Antennaria
Antennaire, everlasting, ladies' tobacco, pussy-toes
erect.
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).
basal and cauline; alternate;
petiolate or sessile;
blades (1–7-nerved) mostly cuneate, elliptic, lanceolate, linear, oblanceolate, or spatulate, margins entire, abaxial faces usually tomentose, adaxial glabrous or ± tomentose to sericeous or glabrescent.
staminate campanulate to hemispheric, 2–6+ mm diam.;
pistillate turbinate or campanulate to cylindric, 3–7(–9+) mm diam.
usually flat to convex, sometimes conic to ± columnar, usually epaleate, sometimes paleate (paleae sometimes enfolding pistillate florets).
flat to convex or ovoid, foveolate, epaleate.
usually 0 (sometimes peripheral florets pistillate and corollas ± zygomorphic and bearing a ± flat limb and interpreted by some as ray florets).
0.
(often 100+ in disciform heads) in 1–3+ series;
corollas (usually present) yellow or purplish to whitish, sometimes red-tipped (actinomorphic and filiform).
mostly 20–100+, (functionally) staminate or pistillate;
staminate corollas white, yellow, or red, narrowly funnelform or tubular (lobes usually 5, erect to recurved);
pistillate corollas white, yellow, or red, narrowly tubular to filiform.
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.
in 3–6+ series, usually relatively narrow, unequal (proximally papery or membranous; distally ± scarious, often black, brown, castaneous, cream, gray, green, olivaceous, pink, red, white, or yellow), apices usually acute, sometimes obtuse to ± truncate.
0.
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.
discoid (unisexual), borne singly or in corymbiform, paniculiform, racemiform, or subcapitate arrays.
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.
mostly ellipsoid to ovoid, faces usually glabrous, often papillate (stout, myxogenic twin-hairs);
pappi: falling (bristles basally connate or coherent, shed together in rings or in groups);
staminate usually of 10–20+ (usually ± clavate, sometimes capillary, barbellate to barbellulate) bristles;
pistillate usually of 12–20+ (capillary, barbellulate to smooth) bristles.
(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.
= 14.
Asteraceae tribe Gnaphalieae
Antennaria
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 45 (34 in the flora).
Some species of Antennaria, especially the stoloniferous, mat-forming species, are cultivated as rock-garden ornamentals. Among the more suitable species widely used for that purpose are A. dioica, A. microphylla, A. parvifolia, A. rosea, and A. suffrutescens. Clones with red or pink phyllaries have been selected as prized for cultivation. Some species are used in the dried-flower trade.
Phylogenetic relationships within Antennaria. Antennaria is composed of two major lineages: the Leontipes group, mostly restricted to western North America, and the Catipes group, occurring throughout the Northern Hemisphere and South America (R. J. Bayer et al. 1996). The Leontipes group consists of five smaller groups (the Geyerae, Arcuatae, Argenteae, Dimorphae, and Pulcherrimae) and comprises species that are primarily diploid (tetraploids are known only in A. dimorpha and A. pulcherrima, Bayer and G. L. Stebbins 1987, and, as far as is known, always amphimictic, sexually reproducing). Most of the species of the Leontipes group lack horizontal stoloniferous growth (except A. flagellaris and A. arcuata). Morphologically, the Leontipes group is considered primitive in the genus, based on unspecialized morphologic features such as non-stoloniferous growth, lack of extensive polyploidy, and general lack of well-developed sexual dimorphism; the Catipes group has amphimictic diploids and tetraploids. Derived from them are all of the polyploid agamic complexes (fig. 1). Most species of the Catipes group have horizontal stolons, an effective means of asexual reproduction; it is considered more specialized than the Leontipes group.
For the most part, the smaller monophyletic groups composing the Leontipes group correspond to traditionally recognized groups (R. J. Bayer 1990; Bayer et al. 1996). The Geyerae group is monotypic, consisting of Antennaria geyeri, and the tendency toward polygamodioecy in that species, along with its lack of basal leaves, makes it more similar morphologically to Anaphalis than to the remainder of Antennaria. Antennaria arcuata is the only member of the newly recognized Arcuatae group, and it was previously considered to be a portion of the Argenteae along with A. luzuloides and A. argentea (Bayer), a relationship that was always considered weak. The Argenteae group comprises A. argentea, A. luzuloides, and A. stenophylla and is sister to the A. arcuata–A. geyeri clade (Bayer et al.). The Dimorphae group, A. dimorpha and A. flagellaris, is sister to the Geyerae-Arcuatae-Argenteae clade (Bayer et al.), and the Pulcherrimae group comprises A. pulcherrima, A. anaphaloides, and A. lanata (Bayer; Bayer et al.).
The Catipes group is well supported in both morphologic and molecular phylogenetic trees (R. J. Bayer et al. 1996); support for subclades within Catipes is weak. Traditionally, members of Catipes were split into the Alpinae, distributed in tundra, with black or olivaceous phyllaries, and the Dioicae with lighter phyllaries. Based on DNA sequence data and morphology, the two groups are artificial and should be abandoned (Bayer et al.). Amphimixis, apomixis (agamospermy), and high levels of polyploidy (Bayer and T. M. Minish 1993) are prevalent among polyploid derivatives of the Catipes group, which consists of diploids and some tetraploids in which sexual dimorphism is highly evolved (Bayer 1990). Some species of the Catipes group are specialized as edaphic endemics, e.g., Antennaria virginica on Devonian-age shale barrens (Bayer and G. L. Stebbins 1987, 1993), A. suffrutescens on serpentine (Bayer and Stebbins 1993), and A. aromatica and A. densifolia on limestone talus (Bayer 1989). Five polyploid agamic complexes, A. alpina (together with the smaller A. media, A. monocephala, and A. friesiana complexes), A. howellii, A. parlinii, A. parvifolia, and A. rosea, have evolved via multiple hybridization among members of the Catipes group (Bayer 1987, 1997). The great success of the Catipes group seems to be correlated with their ability to grow in diverse habitats throughout their range across Eurasia and North America to Tierra del Fuego in South America, and to their acquisition of characteristics such as strong sexual dimorphism, polyploidy, agamospermy, and vegetative reproduction (stolons). The amphimictic taxa of the Catipes group include A. aromatica, A. corymbosa, A. densifolia, A. dioica, A. friesiana subsp. alaskana, A. friesiana subsp. neoalaskana, A. marginata, A. microphylla, A. monocephala subsp. monocephala, A. neglecta, A. plantaginifolia, A. pulchella, A. racemosa, A. rosulata, A. solitaria, A. suffrutescens, A. umbrinella, and A. virginica. Some of those have contributed to the genetic makeup of the polyploid complexes, whose morphologic variation is correlated to the number of diploid genomes contributed to the origin of the complex. Morphologic overlap between the complexes is a direct consequence of pivotal genomes recurring in some complexes. For example, the A. parlinii and A. howellii complexes share two pivotal genomes from A. plantaginifolia (PLA) and A. racemosa (RAC). Some apomictic clones (identified under the name A. howellii subsp. howellii in part) appear to bridge the morphologic gap between the two complexes. The A. parlinii complex has three diploid progenitors: A. solitaria (SOL), A. plantaginifolia (PLA), and A. racemosa (RAC); the A. howellii complex has five: A. marginata (MAR), A. neglecta (NEG), A. plantaginifolia (PLA), A. racemosa (RAC), and A. virginica (VIR). Antennaria parvifolia has three major progenitors: A. dioica (DIO), A. neglecta (NEG), and A. marginata (MAR); it is likely that high elevation segregates of the complex also contain genomic contributions from A. pulchella (PUL) and/or A. media. Antennaria rosea is morphologically the most diverse of the polyploid complexes and has as its primary progenitors: A. aromatica (ARO), A. corymbosa (COR), A. microphylla (MIC), A. racemosa (RAC), and A. umbrinella (UMB). It is likely that A. marginata (MAR), A. rosulata (ROS), and possibly A. suffrutescens (SUF) have also contributed to the origins of some A. rosea clones. The circumpolar allopolyploid A. alpina complex appears to have its origins from the amphimictic, dark-phyllaried, arctic-alpine taxa including A. aromatica (ARO), A. densifolia (DEN), A. friesiana subsp. alaskana (ALA), A. friesiana subsp. neoalaskana (NEO), A. monocephala subsp. monocephala (MON), and A. pulchella (PUL). Three polyploid complexes, A. friesiana subsp. friesiana, A. media, and A. monocephala subsp. angustata (ANG) appear to be of non-hybrid, autopolyploid origin and are direct polyploid derivatives of A. friesiana (subspp. alaskana and neoalaskana), A. monocephala subsp. monocephala, and A. pulchella, respectively; most polyploids are of multiple hybrid origin from among multiple amphimicts. Antennaria marginata has also given rise to apparent autopolyploid apomictic derivatives.
Key pivotal genomes involved in the origins of the polyploid complexes include Antennaria aromatica, A. marginata, A. neglecta, A plantaginifolia, A. pulchella, and A. racemosa; significant contributions have also been made by A. corymbosa, A. densifolia, A. dioica, A. friesiana, A. microphylla, A. monocephala, A. solitaria, A. umbrinella, and A. virginica.
Classification of Antennaria. Past practice has been to attempt to recognize each agamospecies as a distinct taxonomic entity, usually at species rank. That has led to unwieldy classifications that can be used only by experts on the group. Clearly, that method is unsatisfactory and a more reasonable scheme for classifying polyploid agamic complexes, such as the one advocated by E. Babcock and G. L. Stebbins (1938), should be adopted. R. J. Bayer and Stebbins (1982) were the first to use the Babcock and Stebbins method in Antennaria.
Because the sexual diploids are morphologically discrete, they are each recognized as species. Polyploids that are morphologically identical with sexual diploid (nonhybrid- or auto-polyploid) taxa, whether they are agamospermous or amphimictic, are treated as conspecific with their sexual diploids, e.g., tetraploid cytotypes of Antennaria virginica and some other taxa are treated as conspecific with their corresponding sexual diploids because they are morphologically (R. J. Bayer and G. L. Stebbins 1982) and, in the case of A. virginica, genetically (Bayer and D. J. Crawford 1986) inseparable from the sexual diploids. Sexual and asexual polyploids that are of hybrid origin (segmental and genomic allopolyploids) are recognized as species because their genetic composition is not attributable to any single diploid origin. For example, Bayer and Stebbins classified A. parlinii as distinct from its sexual diploid progenitors, A. plantaginifolia, A. racemosa, and A. solitaria. A. Cronquist (1945) recognized A. parlinii (sensu Bayer and Stebbins) as two varieties of A. plantaginifolia, a view Stebbins and I opposed because the polyploids, while containing genes from A. plantaginifolia in their genetic background, also have genes from A. racemosa and A. solitaria (Bayer 1985b; Bayer and Crawford). The polyploid complexes are each defined primarily by assessing their genomic composition through the use of genetic markers, as well as through morphologic studies. This philosophy and method of classification has been extended to the other polyploid agamic complexes.
Identifying Antennaria specimens. Users of this treatment should be aware that multiple details must be kept in mind when collecting and trying to identify species of Antennaria. For example, assigning specimens to species in the “mat-forming,” stoloniferous Catipes group is particularly difficult because of widespread polyploidy and apomixis. One determinative taxonomic character (whether populations are gynoecious or dioecious) may not be readily observed on herbarium specimens but is readily determined in the field by gender ratios. On herbarium specimens, assuming pistillates are always present in populations, absence of staminates could mean either that they were not collected or that they were actually absent from the population. This character comes into use in separating the infraspecific taxa within both A. monocephala and A. friesiana. If this character cannot be readily determined on herbarium material, i.e., when staminates are absent, then such specimens are best keyed to the specific level only.
Another feature of importance in identifying specimens of Antennaria is the presence or absence of well-developed stolons that root at their tips. Some Antennaria species produce stiff, semi-erect stolons that do not root at the tips, and those stolons should not be confused with the typical stolons that are more elongate and horizontal and root at their tips.
The final feature of importance in identifying specimens of Antennaria is the presence or absence of flags on tips of mid and distal cauline leaves. Flags are flat, linear, scarious appendages of the leaf tips that are similar to the tips of the phyllaries; they are not to be confused with ordinary subulate or blunt leaf tips that are essentially green and herbaceous. In keys and descriptions, leaves are referred to as flagged or not flagged.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
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 |
Key to Groups of Antennaria Species
1. Heads usually borne singly, rarely in 2s or 3s | Group 1 |
1. Heads usually (2–)3–15(–110+), rarely borne singly | → 2 |
2. Stolons none (or erect and not rooting at tips; plants not forming mats; in fruit, heights of pistillate plants ± equal to staminates) | Group 2 |
2. Stolons mostly 1–5(–18) cm (usually prostrate, sometimes ascending, usually rooting at tips; plants forming mats; in fruit, heights of pistillate plants usually greater than or equal to heights of staminates) | → 3 |
3. Basal leaves 3–5(–7)-nerved | Group 3 |
3. Basal leaves mostly 1-nerved (1–3-nerved in A. arcuata and A. marginata) | → 4 |
4. Phyllaries (proximally dark) distally mostly dark to light brown, black, or olivaceous (some times inner whitish); arctic or alpine tundra to just below treeline | Group 4 |
4. Phyllaries (proximally light) distally mostly light brown, cream, gray, green, ivory, pink, red, rose, or white; seldom arctic or alpine tundra (A. corymbosa sometimes alpine) | Group 5 |
Group 1
1. Basal leaves 3–5-nerved; se United States | A. solitaria |
1. Basal leaves 1-nerved; w North America, Arctic | → 2 |
2. Plants 0.2–1.5(–2) cm (heads subsessile among basal leaves); basal leaves silvery gray- pubescent; Arizona, Colorado, New Mexico, Utah | A. rosulata |
2. Plants 0.5–13 cm; basal leaves abaxially gray-tomentose (sometimes none at flowering, A. suffrutescens); usually not Arizona, Colorado, New Mexico, Utah (A. dimorpha sometimes Colorado, New Mexico, Utah) | → 3 |
3. Cauline leaves spatulate (apices emarginate or obtuse), adaxial faces green, gla- brous (bases of plants ± woody); nw California and sw Oregon | A. suffrutescens |
3. Cauline leaves linear or oblanceolate (apices acute to obtuse), adaxial faces green or gray, glabrous or pubescent, sericeous, tomentose, or villous; not nw California or sw Oregon | → 4 |
4. Leaves: adaxial faces green, usually glabrous, rarely villous or pubescent (flags of mid and distal cauline leaves brown); arctic and alpine tundra | A. monocephala |
4. Leaves: adaxial faces gray, sericeous or tomentose (flags of mid and distal cauline leaves none); sagebrush steppe or talus near treeline | → 5 |
5. Stolons none (plants cespitose, not forming mats); semidesert | A. dimorpha |
5. Stolons 0.5–2 or 3–10 cm (plants forming mats); semidesert or limestone talus | → 6 |
6. Gynoecious; stolons 0.5–2 cm (leafy); basal leaves spatulate, rhombic- spatulate, or cuneate; s Nevada | A. soliceps |
6. Dioecious; stolons 3–10 cm (not leafy); basal leaves linear-oblanceolate; not s Nevada | A. flagellaris |
Group 2
1. Plants 7–15 cm (mid and distal cauline leaves flagged); low and high arctic | A. friesiana |
1. Plants either 15–65 cm (low arctic or subalpine) OR 0–70 cm (desert steppe or alpine, except A. pulcherrima arctic) | → 2 |
2. Plants (3–)10–15 cm; basal leaves linear to narrowly oblanceolate (1–3 mm wide); phyl- laries distally light brown, dingy brown, or olivaceous | A. stenophylla |
2. Plants 3–70 cm; basal leaves elliptic, lanceolate, linear, oblanceolate, or spatulate (3–25 mm wide; sometimes none at flowering); phyllaries distally usually brown, cream, pink, red, or white, rarely black, dark brown, castaneous, or olivaceous | → 3 |
3. Phyllaries (scarious, glabrous) | → 4 |
3. Phyllaries (scarious distally, hairy proximally) | → 5 |
4. Basal leaves oblanceolate to elliptic; cauline leaves oblanceolate; pistillate involucres 4–5 mm; phyllaries (usually pale green proximally) distally silvery white | A. argentea |
4. Basal leaves linear to narrowly spatulate; cauline leaves narrowly oblanceolate, narrowly spatulate, or linear; pistillate involucres 3.5–6.5 mm; phyllaries (usually light brown to golden proximally) distally white (often red- or pink-flecked). | A. luzuloides |
5. Basal leaves (absent at flowering; plants woody at bases); phyllaries (densely pubescent to well distal of middle), distally usually pink to red, sometimes light brown or white | A. geyeri |
5. Basal leaves elliptic, lanceolate, oblanceolate, or spatulate; phyllaries (moderately pubescent proximal to middle) distally usually black, brown, castaneous, cream, olivaceous, or white (rarely red- or pink-flecked) | → 6 |
6. Plants 3–20 cm; phyllaries (light brown, dark brown, or olivaceous proximally) distally whitish or light brown; alpine slopes | A. lanata |
6. Plants 8–65 cm; phyllaries (outer usually each with a dark spot at base) distally black, brown, castaneous, cream, olivaceous, or white; mostly subalpine, montane, or subarctic | → 7 |
7. Pistillate involucres 4.5–7 mm; phyllaries (bases each with dark spot 0.1–1 mm) distally cream or white (apices obtuse); dry montane or steppe | A. anaphaloides |
7. Pistillate involucres 7–12 mm; phyllaries (bases each with dark spot 1–3 mm) distally black, brown, castaneous, or olivaceous (apices acute); wet sites, subalpine or subarctic (A. pulcherrima subsp. pulcherrima) or limestone near sea level (A. pulcherrima subsp. eucosma) | A. pulcherrima |
Group 3
1. Basal leaves adaxially glabrous; heads in loose racemiform or paniculiform arrays (stems proximal to heads with purple glandular hairs); w North America | A. racemosa |
1. Basal leaves adaxially gray-pubescent, floccose-glabrescent, or green-glabrescent; heads in tight, corymbiform arrays (stems proximal to heads with or without purple glandular hairs); e North America to e Manitoba, Minnesota, Missouri, Texas | → 2 |
2. Basal leaves abaxially tomentose; pistillate involucres 5–7 mm; staminate corollas 2–3.5 mm; pistillate corollas 3–4 mm (young stolons mostly ascending); Appalachians, Pied- mont, Atlantic seaboard, and driftless area of Wisconsin, Minnesota | A. plantaginifolia |
2. Basal leaves adaxially tomentose or glabrous; pistillate involucres (7–)8–13 mm; staminate corollas 3.5–5 mm; pistillate corollas 4–7 mm (young stolons mostly decumbent); e North America from Atlantic seaboard to e margin of Great Plains | A. parlinii |
Group 4
1. Stolons 0.5–2.5 cm (prostrate); basal leaves usually cuneate, cuneate-spatulate, or spatulate, sometimes oblanceolate (lengths 1–2 times widths); limestone talus, n Wyoming to Yukon and Northwest territories | → 2 |
1. Stolons 0.1–16 cm (erect or decumbent); basal leaves cuneate, linear-cuneate, oblanceolate, or spatulate (lengths 2–6+ times widths); not limestone talus, w North America, Arizona, New Mexico to circumpolar Arctic | → 3 |
2. Basal leaves mostly 5–16 × 3–10 mm; distal cauline leaves not flagged; pistillate involucres 5–7(–9) mm (living plants with odor of citronella when crushed; flowering stems, leaves, and bases of phyllaries stipitate-glandular) | A. aromatica |
2. Basal leaves mostly 3–7 × 2–5 mm; mid and distal cauline leaves flagged; pistillate involucres 4.5–7.5 mm (living plants odorless; flowering stems, leaves, and bases of phyllaries not stipitate-glandular) | A. densifolia |
3. Stolons (usually erect, slightly woody); phyllaries distally pale brown, white, or yellowish (sometimes streaked with pink or rose, usually blunt); montane, rarely above treeline. | A. umbrinella |
3. Stolons (usually decumbent, herbaceous); phyllaries distally black, brown, or olivaceous (sometimes whitish at tips, usually acute); arctic, alpine, rarely subalpine | → 4 |
4. Cauline leaves 3–11(–13) mm; staminate corollas 1.9–2.8 mm; pistillate corollas 2–3 mm (stems, leaves, and phyllaries often stipitate-glandular); Sierra Nevada (Lake Tahoe to Mt. Whitney), California, adjacent Nevada | A. pulchella |
4. Cauline leaves 4–20 mm; staminate corollas 2.5–4.5 mm; pistillate corollas 3–4.5 mm (stems, leaves, and phyllaries stipitate-glandular or not); w North America (Arizona, California, New Mexico n to Yukon, Northwest Territories, e Arctic) | → 5 |
5. Basal leaves: faces gray-pubescent; mid and distal cauline leaves mostly not flagged (sometimes flagged near heads) | A. media |
5. Basal leaves: abaxial faces tomentose, abaxial green-glabrescent to gray-pubescent; mid and distal cauline leaves flagged | → 6 |
6. Stolons 0.1–4 cm; involucres: pistillate 5.5–8 mm; corollas: staminate 2.5–3 mm, pistillate 3–4.5 mm (stems, leaves, and phyllaries stipitate-glandular, hairs purple). | A. friesiana |
6. Stolons 1–7 cm; involucres: pistillate 4–7(–10) mm; corollas: staminate 3–3.5 mm, pistillate 3.5–5 mm (stems, leaves, and phyllaries not stipitate-glandular) | A. alpina |
Group 5
1. Basal leaves adaxially usually green-glabrous, sometimes gray-pubescent; phyllaries distally brown, pink, or white | → 2 |
1. Basal leaves usually adaxially pubescent, sometimes glabrous or glabrate with age (A. neglecta); phyllaries distally black, brown, cream, green, ivory, pink, red, stramineous, white, or yellow (sometimes streaked with pink or rose) | → 5 |
2. Basal leaves 1–3-nerved, 9–12 mm wide; distal cauline leaves flagged; phyllaries distally usually light brown, sometimes white | A. howellii |
2. Basal leaves 1-nerved, 3–9 mm wide; distal cauline leaves flagged or not; phyllaries distally whitish or pink | → 3 |
3. Distal cauline leaves flagged; phyllaries distally white or cream; e, c North America | A. howellii |
3. Distal cauline leaves not flagged; phyllaries distally white or light to dark pink; sw United States, Alaska (Aleutian Islands) | → 4 |
4. Stolons (pubescent) 2–5 cm (stems not stipitate-glandular); basal leaves adaxially green-glabrous (margins not white-woolly); Alaska (Aleutian Islands) | A. dioica |
4. Stolons (densely woolly, hairs obscuring surfaces) 2–7 cm (stems sometimes stipitate-glandular, hairs white or purplish); basal leaves adaxially green-glabrous (margins white-woolly); Arizona, se California, sw Colorado, New Mexico | A. marginata |
5. Basal leaves (largest 20–65 × 6–20 mm); phyllaries distally light brown, ivory, or white (never black, dark brown, dark green, pink, or red) | → 6 |
5. Basal leaves (largest mostly 16–45 × 4–15 mm; if 20+ mm long, less than 6.5 mm wide; if 6.5+ mm wide, less than 20 mm long); phyllaries distally light to dark brown, ivory, pink, red, rose, or white | → 7 |
6. Basal leaves cuneate-oblanceolate, narrowly to broadly ovate, spatulate, or spatulate-obovate, 20–48(–65) × 2.5–20 mm, abaxially tomentose, adaxially gray-pubescent or green-glabrous; mid and distal cauline leaves mostly not flagged (sometimes flagged near heads) | A. howellii |
6. Basal leaves cuneate-oblanceolate to spatulate, 15–65 × 6–18 mm, abaxially tomentose, adaxially gray-pubescent (green-glabrescent in age); mid and distal cauline leaves flagged | A. neglecta |
7. Stolons either 4–10 cm (herbaceous, arched) or 4–16 cm (slightly woody, erect); phyllaries distally usually pale brown, sometimes whitish or yellowish (rarely white- or pink-flecked) | → 8 |
7. Stolons 1–10 cm (herbaceous, usually decumbent); phyllaries distally usually cream, gray, green, pink, red, stramineous, white (ivory to pure), or yellow (if light brown, staminate plants absent from populations) | → 9 |
8. Stolons 4–16 cm (somewhat woody, usually erect); basal leaves narrowly spatulate to cuneate, 10–17 × 2–5.4 mm (plants woody at bases); Alberta and British Columbia to California and Colorado | A. umbrinella |
8. Stolons 4–10 cm (herbaceous, arched); basal leaves narrowly to broadly spatulate or rhombic-ovate, 20–45 × 3–15 mm; Blaine County, Idaho, Elko County, Nevada, and Fremont County, Wyoming | A. arcuata |
9. Plants 2–8(–15) cm; pistillate involucres 8–10(–15) mm | A. parvifolia |
9. Plants 4–30 cm; pistillate involucres 4–10 mm | → 10 |
10. Basal leaves spatulate; phyllaries (each with chestnut brown spot near base) distally white or light brown (willow thickets, similar moist habitats, subalpine to alpine zones, Rocky Mountains and c Sierra Nevada) | A. corymbosa |
10. Basal leaves cuneate-oblanceolate, spatulate, or linear; phyllaries (uniformly or combinations of) light brown, cream, gray, green, pink, red, white, or light yellow | → 11 |
11. Gynoecious (staminate plants very rare); basal leaves linear; phyllaries distally usually (combinations of) light brown, cream, gray, green, pink, red, white, or yellow (if solid white, not Appalachian) | A. rosea |
11. Dioecious (stems sometimes stipitate-glandular distally); basal leaves cuneate-oblanceolate or spatulate; phyllaries distally stramineous, white, or light yellow | → 12 |
12. Basal leaves 10–25 × 3–9 mm, faces gray-pubescent; shale barrens, e Ohio, w Pennsylvania, Maryland, Virginia, West Virginia | A. virginica |
12. Basal leaves 6–16 × 2–6 mm, faces silvery white-pubescent (stems stipitate-glandular distally, hairs purple or white, moniliform); w North America e to Ontario | A. microphylla |
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