80. Pyrolaceae - Flora of North America

80. Pyrolaceae - Flora of North America

Hydrangeaceae05dSI (Freeman)
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HYDRANGEACEAE Dumortier
* Hydrangea Family
Craig C. Freeman
Subshrubs, shrubs, trees, or vines [herbs], evergreen or deciduous. Stems ascending to erect,
sometimes climbing or trailing. Leaves cauline, opposite or, sometimes, whorled [alternate],
simple; stipules absent; petiole present or absent; blade margins entire, denticulate, or lobed,
venation pinnate or acrodromous (Fendlera, Fendlerella, Philadelphus, and Whipplea).
Inflorescences terminal or axillary, corymbiform to paniculiform cymes; bracts present or
absent. Flowers bisexual or, sometimes, marginal ones sterile, radially symmetric (bisexual
ones) or bilaterally symmetric with enlarged petaloid sepals (sterile ones); perianth and
androecium half-epigynous to epigynous; hypanthium present; sepals 4–12, distinct or connate
proximally; petals 4–12, connate proximally or entirely, then calyptrate; nectary disc usually
present; stamens 4–200; filaments distinct or connate proximally, forming corona; anthers
dehiscencing by longitudinal slits; pistils 1, 2–12-carpellate; ovary less than ½ inferior, ½
inferior, or completely inferior, locules 1–12; placentation usually axile proximally, parietal
distally, rarely strictly axile or parietal; ovules 1–50 per locule, anatropous; styles (1–)2–12;
stigmas as many as style branches or, sometimes, 1, lobed. Fruits capsules [berries],
dehiscence septicidal or loculicidal. Seeds 2–350, funicular appendage present (Fendlerella,
Whipplea) or absent.
Genera 17, species ca. 220 (9 genera, 25 species in the flora): North America, Mexico,
Central America, South America, Eurasia, Pacific Islands.
A. Cronquist (1981) placed Hydrangeaceae among a group of woody families traditionally
allied with Saxifragaceae. Recent phylogenetic studies consistently place Hydrangeaceae in the
Cornales and sister to Loasaceae (A. L. Hempel et al. 1995; Hufford 2004; Hufford et al.
2001; D. E. Soltis et al. 1995). Within Hydrangeaceae, the western North American genera
Jamesia and Fendlera form a clade (subfamily Jamesioideae) that is sister to the rest of the
family (subfamily Hydrangeoideae) (L. Hufford et al. 2001; L. Hufford 2004). Subfamily
Hydrangeoideae comprises two tribes: Philadelpheae and Hydrangeae. North American
genera in the former are Carpenteria, Deutzia, Fendlerella, Philadelphus, and Whipplea.
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Relationships in the Hydrangeae are poorly resolved; North American genera in the tribe are
Decumaria and Hydrangea.
The Hydrangeaceae are well represented in the paleobotanical record dating back to the
Upper Cretaceous but best represented in the Tertiary (L. Hufford 2004). Some genera are
sources of popular introduced or native ornamentals, including Carpenteria, Deutzia,
Hydrangea, and Philadelphus. Some ornamentals have become established outside of
cultivation in the flora area. A few North American Hydrangeaceae have reputed medicinal
(D. E. Moerman 1998) or toxicologic (G. E. Burrows and R. J. Tyrl 2001) properties.
SELECTED REFERENCES Hufford, L. 1995. Seed morphology of Hydrangeaceae and its phylogenetic implications. Int. J.
Plant Sci. 156: 555–580. Hufford, L. 1997. A phylogenetic analysis of Hydrangeaceae based on morphological data. Int. J.
Plant Sci. 158: 652–672. Hufford, L. 1998. Early development of androecia in polystemonous Hydrangeaceae. Amer. J. Bot.
85: 1057–1067. Hufford, L. 2004. Hydrangeaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular
Plants. 7+ vols. Berlin etc. Vol. 6, pp. 202–215. Hufford, L., M. L. Moody, and D. E. Soltis. 2001. A phylogenetic analysis of
Hydrangeaceae based on sequences of the plastid gene matK and their combination with rbcL and morphological data. Int. J.
Plant Sci. 162: 835–846. Small, J. K. and P. A. Rydberg. 1905. Hydrangeaceae. In: N. L. Britton et al., eds. North American
Flora.... 1905+. New York. 47+ vols. Vol. 22, pp. 159–178. Soltis, D. E., Xiang Q. Y., and L. Hufford. 1995. Relationships
and evolution of Hydrangeaceae based on rbcL sequence data. Amer. J. Bot. 82: 504–514. Spongberg, S. A. 1972. The genera
of Saxifragaceae in the southeastern United States. J. Arnold Arbor. 53: 409–498.
1. Plants woody vines ........................................................................................................... 9. Decumaria, p. nnn
1. Plants subshrubs, shrubs, or trees.
2. Twigs with stellate and simple trichomes ........................................................................ 5. Deutzia, p. nnn
2. Twigs with simple or, sometimes, branched trichomes, never with stellate trichomes.
3. Inflorescences with both sterile and bisexual flowers, sterile flowers peripheral
.............................................................................................................. 8. Hydrangea (in part), p. nnn
3. Inflorescences with only bisexual flowers.
4. Stamens 20–200; filaments terete.
5. Styles 3–5; leaves deciduous; stamens 20–70 ....................................... 6. Philadelphus, p. nnn
5. Style 1; leaves persistent; stamens 150–200 ........................................... 7. Carpenteria, p. nnn
4. Stamens 8–12(–20); filaments flattened or terete.
6. Stems prostrate to decumbent................................................................... 4. Whipplea, p. nnn
6. Stems erect, ascending, or spreading.
7. Apices of filaments 2-lobed, lobes surpassing anthers .......................... 2. Fendlera, p. nnn
7. Apices of filaments not 2-lobed.
8. Inflorescences 100–1000-flowered............................... 8. Hydrangea (in part), p. nnn
8. Inflorescences 1–35-flowered.
[9. Moved to left margin.–Ed.]
9. Margins of leaf blades entire; blades elliptic to lanceolate or oblanceolate, obovate, or
linear-oblong .................................................................................................................... 3. Fendlerella, p. nnn
9. Margins of leaf blades crenate to dentate or, rarely, entire; blades ovate or broadly ovate
to obovate, rhombic, or suborbiculate ................................................................................... 1. Jamesia, p. nnn
1. JAMESIA Torrey & A. Gray, Fl. N. Amer. 1: 593. 1840, name conserved * Cliffbush,
waxflower [For Edwin P. James, 1797–1861, American physician and naturalist on the
Stephen H. Long expedition of 1819–1820]
Craig C. Freeman
Shrubs. Stems ascending or spreading. Bark exfoliating in sheets, strips, or strings. Branches
ascending, spreading, or descending; twigs with simple trichomes. Leaves deciduous, opposite,
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herbaceous; petiole present; blade ovate or broadly ovate to obovate, rhombic, or
suborbiculate, margins crenate to dentate or, rarely, entire, plane; venation pinnate.
Inflorescences terminal, cymose, 2–35-flowered, sometimes flowers solitary; peduncle absent
or present. Pedicels present. Flowers bisexual; perianth and androecium half-epigynous;
sepals 4–5, connate proximally into tube; floral tube hemispheric, adnate to proximal part of
ovary, without ribs in fruit, sparsely to densely strigose, canescent, or sericeous, lobes
persistent, valvate, erect, lanceolate to narrowly ovate; petals 4–5, imbricate, spreading, white
or pink, obovate or oblanceolate, base clawed, sometimes obscurely, adaxial surface hairy;
stamens 8 or 10; filaments distinct, dorsiventrally flattened, gradually or abruptly tapered
from base to narrow summit, apex not 2-lobed; anthers depressed-ovate; pistil 3–5-carpellate;
ovary to half-inferior, partially 3–5-locular initially, becoming 1-locular; placentation axile
proximally, parietal distally; styles persistent, (2–)3–5, distinct. Capsules ovoid or conic,
indurate, dehiscence basipetally septicidal to middle of fruit. Seeds 25–50 per locule, orangish
brown or tan, ellipsoid. x = 16.
Species 2 (2 in the flora): United States, n Mexico.
This treatment of Jamesia is essentially an adaptation of that by N. H. Holmgren and P. K.
Holmgren (1989).
Phylogenetic studies place Jamesia as a basal genus in Hydrangeaceae, sister either to
Fendlera (D. E. Soltis et al. 1995) or a clade comprising Fendlera, Fendlerella, and Whipplea
(L. Hufford 1997), which are, in turn, sister to the rest of the family. Fossil leaves referred to
the genus have been identified in Oligocene sediments from Colorado and Montana; the
identity of material from the latter site is ambiguous (N. H. Holmgren and P. K. Holmgren
1989).
SELECTED REFERENCE
Brittonia 41: 335–350.
1.
1.
Holmgren, N. H. and P. K. Holmgren. 1989. A taxonomic study of Jamesia (Hydrangeaceae).
Sepals 5; petals 5; stamens 10; inflorescences 5–35-flowered, cymose or, rarely, 1- or 2-flowered
on lateral branches; leaf blade margins (5–)9–51(–69)-toothed
.........................................................................................................................................................1. Jamesia americana
Sepals 4; petals 4; stamens 8; inflorescences solitary flowers or, rarely, 2- or 3-flowered; leaf
blade margins 3–13(–16)-toothed, rarely entire
........................................................................................................................................................ 2. Jamesia tetrapetala
1. Jamesia americana Torrey & A. Gray, Fl. N. Amer. 1: 593. 1840 * American cliffbush
F
Edwinia americana (Torrey & A. Gray) A. Heller
Stems 5–20(–40) dm. Bark exfoliating readily or tardily in reddish brown or blackish sheets, or gray to brown
strips or strings. Branches spreading or ascending, often stunted and straggly; twigs ascending-strigose. Leaves:
petiole (1–)2–18(–54) mm, ascending-strigose to canescent or sericeous; blade ovate or broadly ovate to obovate,
rhombic, or suborbiculate, (0.7–)1.3–8(–10) × (0.5–)1–6.3(–8.5) cm, base cuneate to obtuse or rounded, usually
asymmetric, margins crenate to dentate, (5–)9–51(–69)-toothed, apex obtuse to rounded, abaxial surface
moderately to densely canescent or sericeous, adaxial sparsely strigose to glabrescent. Inflorescences cymose, 3–
35-flowered, or, rarely, 1- or 2-flowered on lateral branches; peduncle absent or 4–30 mm, sparsely to densely
strigose. Pedicels 1.5–8 mm, sparsely to densely strigose. Flowers: sepals 5; floral tube 1.3–2 × 2.5–4.5 mm,
sparsely to densely strigose, lobes lanceolate to deltate-ovate, 1.5–7(–8) × 1.1–2 mm, margins entire, sometimes 2–
3-lobed apically, apex acute to obtuse, abaxial surface sparsely to densely strigose; petals 5, white or pink, (4–)5.5–
11(–11.5) × 3–4.5 mm, sparsely to densely strigose or canescent, especially distally; stamens 10; filaments (2–)2.7–
10 × (0.2–)0.5–1 mm; anthers 0.7–1.1 mm; styles (2–)3–5, (3–)3.5–6.5(–8) mm. Capsules 3.5–5.5(–7) × 2–3.8
mm. Seeds 0.6–1 mm. 2n = 32.
Varieties 4 (4 in the flora): United States, n Mexico.
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N. H. Holmgren and P. K. Holmgren (1989) recognized subsp. americana, the widespread eastern element,
including only var. americana, and subsp. californica (Small) A. E. Murray, the western element, with three
allopatric varieties: macrocalyx, rosea, and zionis.
The Chiricahua Apache and Mescalero Apache of the southwestern United States and northern Mexico used the
seeds as food (D. E. Moerman 1998). The species has been cultivated as an ornamental since 1862 (R. A. Vines
1960).
1. Bark exfoliating readily in reddish brown or blackish sheets; leaf blade margins crenate to dentate, teeth (13–)19–51(–69);
inflorescences
(7–)15–35-flowered;
se
Arizona,
c
Colorado,
New
Mexico,
se
Wyoming
.....................................................................................................................................................1a. Jamesia americana var. americana
1. Bark exfoliating tardily in gray or brown strips or strings; leaf blade margins dentate, teeth (5–)9–27(–33); inflorescences (1–)3–11(–
19)-flowered; s California, s Nevada, Utah.
2. Petals pink or, rarely, white; lobes of floral tubes (1.5–)1.9–4(–6) mm; s California, s Nevada
.......................................................................................................................................................1c. Jamesia americana var. rosea
2. Petals white or pinkish white; lobes of floral tubes (2.5–)3–7(–8) mm; Utah.
3. Leaf blades (0.7–)1.2–3(–3.9) × (0.5–)0.8–1.8(–3.2) cm; n Utah
.......................................................................................................................................1b. Jamesia americana var. macrocalyx
3. Leaf blades (2.5–)3–5(–5.5) × (1.3–)2–4(–4.5) cm; sw Utah
................................................................................................................................................ 1d. Jamesia americana var. zionis
1a. Jamesia americana Torrey & A. Gray var. americana
F
Bark exfoliating readily in reddish brown or blackish sheets. Leaves: petiole (1–)3–18(–54) mm; blade ovate, (1–
)2–8(–10) × (1–)1.3–6.3(–8.5) cm, margins crenate to dentate, teeth (13–)19–51(–69). Inflorescences (7–)15–35flowered. Flowers: floral tube lobes triangular-ovate, 1.5–3(–3.8) mm; petals white, 6–11 mm; styles 3.5–5(–7)
mm. 2n = 32.
Flowering May–Aug(–Oct). Streambanks, forested canyon bottoms, rocky slopes, cliffs; 1500–3000(–3800) m;
Ariz., Colo., N.Mex., Wyo.; Mexico (Chihuahua, Nuevo León).
1b. Jamesia americana Torrey & A. Gray var. macrocalyx (Small) Engler in H. G. A. Engler and K. Prantl, Nat.
Pflanzenfam. ed. 2, 18a: 196. 1930 C E F
Edwinia macrocalyx Small in N. L. Britton et al., N. Amer. Fl. 22: 176. 1905
Bark exfoliating tardily in gray strips or strings. Leaves: petiole 2–8(–12) mm; blade ovate to obovate or rhombic,
(0.7–)1.2–3(–3.9) × (0.5–)0.8–1.8(–3.2) cm, margins dentate, teeth (5–)11–21(–27). Inflorescences (2–)3–9(–13)flowered, sometimes flowers solitary. Flowers: floral tube lobes lanceolate to triangular-ovate, (2.5–)3–7 mm;
petals white or pinkish white, (6.5–)7.5–11.5 mm; styles 4–6.7 mm.
Flowering Jun–Aug. Cliffs, crevices, rocky slopes; of conservation concern; (1700–)2100–3300(–3700) m; Utah.
1c. Jamesia americana Torrey & A. Gray var. rosea Purpus ex C. K. Schneider, Ill. Handb. Laubholzk. 1: 375.
1905 E
Bark exfoliating tardily in brown or gray strips or strings. Leaves: petiole 1–6(–14) mm; blade ovate to elliptic or
suborbiculate, (0.8–)1.3–3(–5) × (0.5–)1–2.2(–2.9) cm, margins dentate, teeth (9–)11–23(–29). Inflorescences (4–
)6–10(–19)-flowered. Flowers: floral tube lobes lanceolate, (1.5–)1.9–4(–6) mm; petals pink or, rarely, white, (4–
)5.5–9.5 mm; styles 3–6.5(–7.8) mm.
Flowering Jun–Aug. Cliffs, crevices, rocky slopes, talus; 2000–3700 m; Calif., Nev.
1d. Jamesia americana Torrey & A. Gray var. zionis N. H. Holmgren & P. K. Holmgren, Brittonia 41: 344,
fig. 2G–I. 1989 C E
Bark exfoliating tardily in brown or gray strips or strings. Leaves: petiole 4–18 mm; blade ovate to broadly ovate
or suborbiculate, (2.5–)3–5(–5.5) × (1.3–)2–4(–4.5) cm, margins dentate, teeth 13–27(–33). Inflorescences 4–12flowered. Flowers: floral tube lobes lanceolate, (3.6–)4–7(–8) mm; petals white or pinkish white, 5.5–7(–11) mm;
styles (4.5–)6–8 mm.
Flowering Jun–Aug. Moist cliffs, hanging gardens; of conservation concern; 1200–2000(–2500) m; Utah.
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2. Jamesia tetrapetala N. H. Holmgren & P. K. Holmgren, Brittonia 41: 348, fig. 3J–K. 1989 * Four-petal
cliffbush C E
Stems 5–20(–40) dm. Bark exfoliating in gray, orangish gray, or brownish strips or strings. Branches spreading or
descending, often stunted and straggly; twigs densely spreading- or retrorse-pilose. Leaves: petiole 1–4(–7) mm,
ascending-strigose to canescent or sericeous; blade ovate or obovate, (0.5–)1–2(–2.7) × (0.2–)0.8–1.2(–1.6) cm,
base obtuse to rounded or, sometimes, cuneate, usually asymmetric, margins usually dentate, rarely entire, teeth 3–
13(–16), apex obtuse to rounded, abaxial surface moderately to densely canescent or sericeous, adaxial sparsely
strigose to glabrescent. Inflorescences 2(–3)–flowered, sometimes flowers solitary; peduncle 2–10 mm, canescent
or subsericeous. Pedicels 1–5 mm, canescent or subsericeous. Flowers: sepals 4; floral tube 2.5–3.5 × 1–1.5 mm,
canescent or sericeous, lobes lanceolate to deltate-ovate, (3–)5–9 × (1.5–)2–2.8 mm; margins usually entire, rarely
2-lobed apically, apex acute to obtuse, abaxial surface canescent or sericeous; petals 4, white with pinkish margin
or entirely pinkish, 5–13(–15) × 3–4.9 mm, sparsely to densely strigose or canescent, especially distally; stamens 8;
filaments (4–)5–8 × 0.8–1.4 mm; anthers 0.8–1.1 mm; styles 3–5, 4.8–7.7 mm. Capsules 3–5.5(–7) × 2.5–3.5 mm.
Seeds 0.6–0.8 mm.
Flowering Jun–Aug. Limestone cliffs, crevices, talus; of conservation concern; 2000–3300 m; Nev., Utah.
2. FENDLERA Engelmann & A. Gray, Smithsonian Contr. Knowl. 3(5): 77, plate 5. 1852
* Fendler-bush [For August Fendler, 1813–1883, German-born plant collector in North
and South America, early botanical explorer of southwestern United States]
Ronald L. McGregor
Shrubs. Stems ascending or spreading. Bark exfoliating in grayish or reddish strings or strips
or, sometimes, not exfoliating. Branches erect or arching; twigs with simple trichomes.
Leaves deciduous, opposite, sometimes appearing clustered on lateral branches; petiole absent
or present; blade linear, elliptic, lanceolate, oblong, ovate, or falcate, herbaceous or
coriaceous, margins entire, plane or strongly revolute; venation acrodromous or with single
midvein. Inflorescences terminal, solitary flowers; peduncle absent. Pedicels present. Flowers
bisexual; perianth and androecium nearly hypogynous; sepals persistent, 4, valvate, eventually
erect or strongly recurved, essentially distinct, usually triangular, sometime ovate, turbinate,
broadly campanulate, or hemispheric, glabrous or hairy; petals 4, imbricate, spreading, white,
sometimes tinged pink or red, ovate, ovate-deltate, or ovate-rhombic or, rarely, obovate or
ovate, base clawed, surfaces finely pubescent; stamens 8; filaments distinct, dorsiventrally
flattened, gradually tapered toward apex, apex 2-lobed, lobes prolonged beyond anthers;
anthers oblong; pistil 4-carpellate; ovary to half-inferior, 4-locular; placentation parietal; styles
persistent, 4, distinct.
Capsules elongate-ovoid, ovoid, or ellipsoid, cartilaginous or
subcartilaginous, dehiscence basipetally septicidal to middle of fruit. Seeds (1–)2–4(–6) per
locule, reddish brown or dark reddish brown, ellipsoid. x = 11.
Species 2 (2 in the flora): sw United States, n Mexico.
A. J. Rehder (1920) found that the species of Fendlera show great uniformity in floral
characters and may be distinguished easily by their leaves. N. H. Holmgren and P. K.
Holmgren (1997) also reported that floral characters are remarkably uniform. They
considered size, shape, and vestiture of leaves to be too variable to warrant recognition of
infraspecific taxa.
B. L. Turner (2001) recognized five species of Fendlera, including four in the flora. In
addition to the species recognized here, B. L. Turner also recognized F. falcata Thornber and
F. wrightii (Engelmann & A. Gray) A. Heller in our area; as noted later, these are best
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included within F. rupicola. The fifth species of Turner is F. tamaulipana B. L. Turner, a
taxon from Tamaulipas, Mexico; this species should be included in F. linearis.
Inflorescences of Fendlera have been described as solitary flowers, or clusters of two, three,
or five flowers, racemes, or dichasial cymes. The inflorescence consists of a single flower
terminating a branch, these flower-bearing branches sometimes aggregated. Frequently, the
lateral branches are much-reduced and consist of a relatively short axis bearing two relatively
small leaves and terminated by a flower. More rarely, the flowers appear nearly sessile in leaf
axils, subtended by scales; this phenomenon is most frequent in F. linearis.
SELECTED REFERENCE Turner, B. L. 2001. Taxonomic revision of the genus Fendlera (Hydrangeaceae). Lundellia 4: 1–11.
1.
1.
Leaf blades coriaceous, linear, margins strongly revolute, touching broad, flat midrib, abaxial
surface hidden, with 2 layers of vestiture: trichomes of lower layer dense, minute, branched;
trichomes of upper layer coarser, appressed
............................................................................................................................................................ 1. Fendlera linearis
Leaf blades herbaceous to coriaceous, elliptic, lanceolate, oblong, ovate, or falcate, rarely linear,
margins plane to strongly revolute, not touching midrib, abaxial surface visible, with 1 layer of
vestiture: trichomes sparsely to densely appressed; or with 2 layers of vestiture: trichomes of
lower layer dense, minute, branched; trichomes of upper layer moderately appressed
...........................................................................................................................................................2. Fendlera rupicola
1. Fendlera linearis Rehder, J. Arnold Arbor. 1: 205. 1920 * Narrow-leaf Fendler-bush
Fendlera rigida I. M. Johnston; F. tamaulipana B. L. Turner
Stems 5–30 dm. Branches somewhat spinescent; twigs densely hairy. Leaves: petiole absent or essentially so; blade
linear, 7–25(–30) × 1–1.5(–2) mm, coriaceous, base cuneate, margins strongly revolute, touching broad, flat
midrib, apex mucronulate to acute, abaxial surface hidden, with 2 layers of vestiture: trichomes of lower layer
dense, minute, branched; trichomes of upper layer appressed, coarser, adaxial surface glabrous or sparsely
appressed-pubescent, especially near margins. Pedicels 2–3 mm, densely canescent. Flowers: floral tube 2.5–4 ×
3.5–4.5 mm, lobes becoming reflexed, triangular, 3–4 × 3.5–4.5 mm, canescent abaxially; petals white, ovate,
round-obovate, or triangular-ovate, 3–5 × 3–5 mm, margins erose; filaments 3.5–8 × 2–2.5 mm; anthers 1–2 mm;
styles 1–1.2 mm. Capsules ovoid, 6–12 × 3–6 mm, subcartilaginous. Seeds dark reddish brown, 3–5 mm.
Flowering May–Oct. Limestone walls, slopes, rocky woodlands, chaparral, gypsum hills; 1100–1900 m; Tex.;
Mexico (Chihuahua, Coahuila, Nuevo León, Tamaulipas).
Fendlera linearis is well-marked although variable in habit and flower and fruit size. In Texas, it is known only
from Brewster and Presidio counties. It reportedly is grazed heavily by deer and other animals.
2. Fendlera rupicola Engelmann & A. Gray, Smithsonian Contr. Knowl. 3(5): 77, plate 5. 1852 * Cliff
Fendler-bush F
Stems 5–30 dm. Branches never spinescent; twigs glabrous or puberulent. Leaves: petiole absent or essentially so;
blade elliptic, lanceolate, oblong, ovate, or falcate, rarely linear, 10–38 × 1–10 mm, herbaceous to coriaceous, base
cuneate to rounded, margins plane or slightly to strongly revolute, not touching midrib, apex obtuse, acute, or
slightly attenuate, abaxial surface visible, with 1 layer of vestiture: trichomes appressed and sparsely to moderately
dense, or with 2 layers of vestiture: trichomes of lower layer dense, minute, branched, trichomes of upper layer
appressed, moderately dense, adaxial surface glabrous or sparsely to moderately strigose or hispidulous. Pedicels
1–25 mm, glabrous or slightly hairy. Flowers: floral tube 2.5–4 × 4–6 mm, lobes eventually erect or reflexed,
narrowly to broadly triangular, (3.5–)4–7(–8.5) × 1.5–3.2 mm, glabrous or moderately hairy abaxially; petals
white or tinged with pink, ovate, ovate-rhombic or, rarely, oblanceolate, obovate, or oval, 8–15 × 5–14 mm,
margins erose; filaments 3–7 × 1–2.5 mm; anthers 2–2.5 mm; styles 4.5–6 mm. Capsules ovoid or ellipsoid, 7–15
× 5–8 mm, cartilaginous. Seeds reddish brown, 4.5–5.5 mm.
Varieties 2 (2 in the flora): sw United States, n Mexico.
B. L. Turner (2001) separated Fendlera falcata from F. rupicola by leaf length, blade vestiture, and distribution
(F. falcata with leaf lengths mostly three to six times widths, blades mostly moderately to densely strigose abaxially
and variously hairy adaxially, and occurring in western Texas and westward; F. rupicola with leaf lengths three to
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four times widths, very sparsely strigose abaxially and glabrous adaxially, and occurring in central Texas).
Material agreeing with Turner’s concept of F. rupicola occurs in New Mexico and westward to the Grand Canyon
area of Arizona. Morphologically intermediate specimens from other areas suggest that F. falcata is best placed in
synonymy with F. rupicola. Vegetatively, var. rupicola is nearly impossible to distinguish from similar forms of
Philadelphus microphyllus A. Gray. I have been unable to separate them clearly.
Fendlera rupicola is reportedly a favorite browse of deer, goats, sheep, and cattle. D. E. Moerman (1998)
reported that early Native Americans used plants for drugs and as ceremonial items, and the hard wood for
making arrow shafts.
1. Leaf blades herbaceous to coriaceous, margins plane or revolute, abaxial surface with 1 layer of vestiture: trichomes appressed, sparse
to moderately dense, adaxial surface smooth, glabrous or scabrous and densely to moderately strigose
.......................................................................................................................................................... 2a. Fendlera rupicola var. rupicola
1. Leaf blades coriaceous, margins strongly revolute, abaxial surface with 2 layers of vestiture: trichomes of lower layer minute,
branched, forming dense, white tomentum; trichomes of upper layer appressed, moderately dense, adaxial surface scabrous, strigose,
and, often, hispidulous, especially near margins
............................................................................................................................................................2b. Fendlera rupicola var. wrightii
2a. Fendlera rupicola Engelmann & A. Gray var. rupicola
F
Fendlera falcata Thornber; F. rupicola var. falcata (Thornber) Rehder
Leaf blades herbaceous to coriaceous, margins plane or slightly revolute, abaxial surface with 1 layer of vestiture:
trichomes appressed, sparse to moderately dense, adaxial surface smooth, glabrous or scabrous and densely to
moderately strigose.
Flowering Apr–Jun (Aug–Oct). Limestone bluffs and ledges, rocky and brushy slopes, among boulders,
chaparral, juniper-oak woodlands; 1000–2800 m; Ariz., Colo., Nev., N.Mex., Tex., Utah; Mexico (Chihuahua,
Coahuila, Sonora).
2b. Fendlera rupicola Engelmann & A. Gray var. wrightii Engelmann & A. Gray, Smithsonian Contr. Knowl.
3(5): 77, plate 5, fig. 2, 5–14. 1852
Fendlera tomentella Thornber; F. wrightii (Engelmann & A. Gray) A. Heller
Leaf blades coriaceous, margins strongly revolute, abaxial surface with 2 layers of vestiture: trichomes of lower
layer minute, branched, forming dense, white tomentum; trichomes of upper layer appressed, moderately dense,
adaxial surface scabrous, strigose, and, often, hispidulous, especially near margins. 2n = 22.
Flowering Apr–Jul (Aug–Oct). Limestone hillsides and ledges, chaparral, oak woodland, desert grassland,
granitic soils, red-sandstone hillsides; 700–2100 m; N.Mex., Tex.; Mexico (Chihuahua).
In its extreme form, var. wrightii appears as a distinct species and was so recognized by A. A. Heller (Bull.
Torrey Bot. Club 24: 537. 1897), A. Rehder (1920), and B. L. Turner (2001). Turner cited 11 specimens from
Chihuahua and Sonora, Mexico, as more or less intermediate between Fendlera falcata and F. wrightii; he did not
map them. Other intermediates were cited for Arizona and New Mexico. Two specimens from New Mexico were
said to approach F. wrightii, the fuzzy indumentum of the abaxial leaf blade surface absent or nearly so. Although
F. wrightii has been cited for Arizona, Colorado, Nevada, New Mexico, Texas, and Utah, my examination of 195
specimens from these areas reveals that var. wrightii occurs only in New Mexico, Texas, and northern Mexico,
where the two varieties intergrade to some extent. The complex merits a modern monographic study.
3. FENDLERELLA (Greene) A. Heller, Bull. Torrey Bot. Club 25: 626. 1898 * Yerba
desierto [Genus Fendlera and Latin -ella, diminutive]
Ronald L. McGregor
Fendlera Engelmann & A. Gray sect. Fendlerella Greene, Bull. Torrey Bot. Club 8: 26. 1881
Shrubs. Stems ascending or spreading. Bark exfoliating in grayish strips or flakes. Branches
spreading; twigs with simple trichomes. Leaves deciduous, opposite, sometimes appearing
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clustered on lateral branches; petiole present or essentially absent; blade elliptic to lanceolate
or oblanceolate, obovate, or linear-oblong, coriaceous, margins entire, plane or revolute;
venation acrodromous. Inflorescences terminal, compound-cymose, (3–)5–26(–34)-flowered;
peduncle present. Pedicels present. Flowers bisexual; perianth and androecium nearly
hypogynous; sepals persistent, 4–5, valvate, erect, essentially distinct, lanceolate, turbinatecampanulate, strigose; petals (4–)5, valvate, spreading, white to yellowish, oblanceolate, base
clawed, surfaces glabrous or sericeous-pilose abaxially, strigose adaxially; stamens (8)10;
filaments distinct, dorsiventrally flattened, gradually or abruptly tapered from base to tip,
apex not 2-lobed; anthers globose; pistil (2–)3(–5)-carpellate; ovary slightly inferior, (2–)3(–5)locular; placentation strictly axile; styles persistent, (2–)3(–5), distinct. Capsules lanceoloid,
cartilaginous, dehiscence basipetally septicidal to near base of fruit. Seeds 1 per locule,
reddish brown, ellipsoid.
Species 3 (1 in the flora): sw United States, Mexico.
1. Fendlerella utahensis (S. Watson) A. Heller, Bull. Torrey Bot. Club 25: 626. 1898
F
Whipplea utahensis S. Watson, Amer. Naturalist 7: 300. 1873; Fendlerella utahensis var. cymosa (Greene ex Wooton &
Standley) Kearney & Peebles
Stems to 10 dm. Twigs reddish to orangish, strigose. Leaves: petiole 0.1–0.3 mm, or essentially absent, pilose in
axils; blade 5–20(–25) × 1–5(–6.5) mm, base cuneate, apex obtuse to acute, abaxial surface glabrous or strigose to
tomentose, adaxial glabrous or strigose. Inflorescences congested to open, bracteate; peduncle 2–10 mm, strigose.
Pedicels 0.8–3(–4.5) mm, strigose. Flowers: floral tube 0.5–2 × 0.9–1.2 mm, lobes 1–1.5 × 0.2–0.6 mm; petals 2–4
× 0.8–1 mm; filaments 1.5–3 × 0.2–0.4 mm; anthers 0.2–0.5 mm. Capsules 5–6.1 × 1.5–9 mm. Seeds 2–3 mm.
Flowering Apr–Jun (Aug–Sep). Cracks and crevices, usually in limestone but, sometimes sandstone outcrops,
sandy soil of mixed desert scrub, pine communities; 1200–2800 m; Ariz., Calif., Colo., Nev., N.Mex., Tex., Utah;
Mexico (Chihuahua, Nuevo León).
Some plants in the southern range of Fendlerella utahensis, in southern Arizona, southern New Mexico, the
Pecos of Texas, and northern Mexico, have leaves that average narrower, longer, and more acute than the leaves
on plants to the north. These have been recognized as var. cymosa. Intergradation is common; the variety is not
recognized here.
In southern Coahuila, Mexico, are plants with petals undulate and sericeous-pilose, leaf blades tomentose
abaxially, and styles three; these have been named Fendlerella lasiopetala Standley. In southern Puebla, Mexico,
plants with leaf blades white-tomentose and styles two have been named F. mexicana Brandegee.
4. WHIPPLEA Torrey in War Department [U.S.], Pacif. Railr. Rep. 4(5): 90, plate 7. 1857 *
Modesty, yerba de silva [For Lieutenant Amiel Weeks Whipple, 1816–1863, commander of
Pacific Railroad Expedition 1853–1854] E
Ronald L. McGregor
Subshrubs. Stems prostrate to decumbent. Bark exfoliating in grayish or grayish-brownish
strips. Branches erect to decumbent; twigs with simple trichomes. Leaves marcescent,
opposite; petiole absent or present; blade elliptic, ovate, ovate-elliptic, or oval, herbaceous,
margins entire or shallowly crenate-serrate, plane; venation acrodromous. Inflorescences
terminal, racemes or racemose cymes, 4–12-flowered; peduncle present. Pedicels present.
Flowers bisexual; perianth and androecium nearly hypogynous; sepals (4–)5–6, connate
proximally into tube; floral tube free from ovary, weakly 9–11-ribbed in fruit, hemispheric,
appressed-pubescent abaxially, lobes eventually deciduous, valvate, erect, narrowly oblong or
oblong-lanceolate; petals 4–6, valvate, reflexed, white, obovate or oblong, base clawed,
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surfaces glabrous; stamens 8–12; filaments distinct, dorsiventrally flattened, abruptly tapered
medially, apex not 2-lobed; anthers elliptic; pistil (2–)4–5-carpellate; ovary slightly inferior,
(3–)4–5-locular; placentation strictly axile; styles caducous, (3–)4–5, distinct or connate
basally. Capsules depressed-spheric, cartilaginous, dehiscence basipetally septicidal, carpels
separating entirely. Seeds 1 per locule, brown or reddish brown, ellipsoid, (3-gonous).
Species 1: w United States.
1. Whipplea modesta Torrey in War Department [U.S.], Pacif. Railr. Rep. 4(5): 90, plate 7. 1857
E
F
Stems to 10 dm; adventitous roots at proximal nodes. Flowering branches weak, (0.4–)0.6–1.5 dm, appressedpubescent. Leaves: petiole 0.2–3 mm, pilose, or absent; blade 10–40 × 10–30 mm, base rounded or tapered, apex
obtuse, abaxial surface strigose, adaxial with white, pustule-based trichomes. Inflorescences congested to open;
peduncle 20–50 mm, appressed-pubescent. Pedicels 0.5–2.5 mm, appressed-pubescent. Flowers: odorless; floral
tube 1.4–2 × 2.3–2.6 mm, lobes 1.5–2 × 0.2–0.5 mm; petals 2.5–4 × 1–1.5 mm; filaments 1.2 × 0.5 mm; anthers
0.4–0.6 mm; styles 1–1.2 mm. Capsules 1.5–2 × 2–2.5 mm. Seeds 1–1.5 mm.
Flowering Mar–Jun. Dry, rocky sites, open to sparsely forested areas; (30–)400–1300(–1700) m; Calif., Oreg.,
Wash.
Whipplea modesta grows on the west side of the Cascades and Coast Ranges.
5. DEUTZIA Thunberg, Nov. Gen. Pl., 19. 1781 * [For Johann van der Deutz, ca.1743–
1784, Dutch merchant and patron of Carl Peter Thunberg] I
Ronald L. McGregor
Shrubs. Stems erect to spreading. Bark exfoliating in grayish, brownish, or reddish sheets or
flakes. Branches erect, ascending, spreading, or arching; twigs with stellate and simple
trichomes. Leaves deciduous or semideciduous, opposite, herbaceous; petiole present; blade
lanceolate, elliptic, or ovate, margins serrulate to crenate-denticulate, plane; venation pinnate.
Inflorescences terminal, cymose, paniculate, racemose, or corymbose, (2–)5–60-flowered or,
rarely, flowers solitary; peduncle present or absent. Pedicels present. Flowers bisexual;
perianth and androecium epigynous; sepals 5, connate proximally into tube; floral tube
campanulate or ovoid, adnate to proximal part of ovary, without ribs in fruit, glabrous or
abaxially short-hairy or stellate-pubescent, lobes deciduous, valvate, erect, triangular to
broadly ovate; petals 5 [10 in some cultivars], valvate or imbricate, spreading, white, pink, or
purplish, elliptic to oblong, base clawed, surfaces glabrous or hairy; stamens 10 (2-seriate);
filaments distinct, dorsiventrally flattened or terete, gradually or abruptly tapered medially or
filiform, apex 2-toothed or entire; anthers globose; pistil 3–5-carpellate; ovary completely
inferior, 3–5-locular; placentation axile proximally, parietal distally, sometimes strictly
parietal; styles persistent, 3(5), distinct. Capsules hemispheric, cartilaginous, dehiscence
acropetally septicidal along septum at base of fruit and also apically. Seeds 10–20 per locule,
dark brown, ellipsoid. x = 13.
Species ca. 60 (3 in the flora): introduced; Mexico, Asia (China, Himalaya, Japan, Korea),
Pacific Islands (Philippines).
A. J. Rehder (1940) recognized 23 species (including eight named hybrids), 27 varieties, and
about 27 cultivars of Deutzia growing mostly at the Arnold Arboretum. L. H. Bailey et al.
(1976) listed taxa only in the horticultural trade and cited 34 species and 36 cultivars that had
been listed in the previous two decades. M. A. Dirr (1998) reported that Deutzia has lost
favor as an ornamental over the years. He noted that identification of species, particularly
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cultivars, borders on the impossible, and nothing is clear-cut in the world of Deutzia
identification. He listed seven species and 22 cultivars, including nine cultivars of D. scabra.
In cultivation, deutzias may spread by suckers; several species have escaped and become
marginally naturalized. M. L. Fernald (1950) reported that Deutzia gracilis and D. scabra
were beginning to spread to thickets and roadsides; there is no evidence that either species has
become widely naturalized. H. A. Gleason and A. Cronquist (1991) noted that D. scabra
rarely escapes from cultivation.
1.
1.
Inflorescences corymbose cymes; petals broadly ovate or suborbiculate, imbricate; filaments
filiform, toothless or inner ones 2-toothed at apex
.......................................................................................................................................................... 3. Deutzia parviflora
Inflorescences racemes or panicles; petals narrowly elliptic to oblong or oblong-lanceolate,
valvate; filaments narrowly oblong, toothed at apex.
2. Leaf blades glabrous or nearly so
abaxially;
inflorescences
glabrous
....................................................................................................................................................... 1. Deutzia gracilis
2. Leaf blades densely stellate-pubescent abaxially (trichomes 10–20-rayed); inflorescences
stellate-pubescent
.........................................................................................................................................................2. Deutzia scabra
1. Deutzia gracilis Siebold & Zuccarini, Fl. Jap. 1: 22, plate 8. 1835
F
I
Shrubs 5–20 dm. Branches spreading or arching. Leaves: petiole 2–8 mm, glabrous; blade lanceolate, ellipticlanceolate, oblong-lanceolate, or broadly ovate-lanceolate, 30–35 × 15–30 mm, base cuneate or rounded, margins
unevenly serrate, apex acuminate, abaxial surface light green, glabrous or nearly so, adaxial dark green, sparsely
stellate-pubescent. Inflorescences racemose or narrowly paniculate, open, (2–)10–50-flowered, sometimes flowers
solitary, 8–12 × 3–6 cm, glabrous; peduncle 5–10 mm, glabrous. Pedicels 5–10 mm, glabrous. Flowers faintly
fragrant, 15–20 mm; floral tube campanulate, 2.5–3 × 2.5–3 mm, glabrous or short-hairy, lobes triangular, 1.5–1.7
× 1–1.5 mm, apex obtuse, surfaces glabrous or nearly so; petals valvate, white, oblong or oblong-lanceolate, 10–
12(–15) × 4–6 mm, glabrous or sparsely stellate-pubescent abaxially, glabrous adaxially; outer filaments
dorsiventrally flattened, narrowly oblong, 5–6 mm, 2-toothed at apex (teeth not reaching anther), or entire,
subulate, toothless, inner filaments dorsiventrally flattened, narrowly oblong, 3–4 mm, all 2-dentate at apex; styles
3, 4–7 mm. Capsules hemispheric, 6–7 × 5–6 mm. Seeds 1.2–1.5 mm. 2n = 26 (Asia).
Flowering Apr–Jun. Waste areas, roadsides; 10–900 m; introduced; Ga., Md.; Asia (China, Japan).
2. Deutzia scabra Thunberg, Fl. Jap., 185, plate 24. 1784
I
Deutzia crenata Siebold & Zuccarini
Shrubs 10–30 dm. Branches erect to ascending. Leaves: petiole 1–3 mm, sparsely to densely stellate-pubescent;
blade ovate-lanceolate to ovate, 30–80 × 15–50 mm, base rounded or broadly cuneate, margins crenatedenticulate, apex acute to acuminate, abaxial surface light green, densely stellate-pubescent (trichomes 10–20rayed), adaxial dark green, stellate-pubescent (trichomes 4–6-rayed). Inflorescences compound racemes or
panicles, loose, 5–50-flowered, 4–10 × 2–3 cm, stellate-pubescent; peduncle absent. Pedicels 2–10 mm, sparsely
stellate-pubescent. Flowers faintly fragrant, 8–20 mm; floral tube campanulate, 2.5–5 × 3–4 mm, densely stellatepubescent, lobes triangular to ovate, 1.5–2.5 × 1–2.2 mm, apex obtuse, surfaces stellate-pubescent; petals valvate,
white or pinkish, narrowly elliptic to oblong, 7–15 × 2.5–3 mm, stellate-pubescent abaxially, glabrous or sparsely
stellate-pubescent adaxially; outer filaments dorsiventrally flattened, narrowly oblong, 7–9 mm, 2-toothed at apex
(teeth nearly horizontal, not reaching anthers), inner filaments dorsiventrally flattened, narrowly oblong, 5–6 mm,
2-toothed at apex; styles 3(–4), 5–11 mm. Capsules 3.8–5 × 4–5 mm. Seeds 1.5–2 mm. 2n = 26, 130 (Asia).
Flowering Apr–Jul. Roadsides, waste areas, homesteads, parks; 10–200 m; introduced; Conn., Fla., Ga., Ill.,
Ky., Md., Mass., N.J., N.Y., N.C., Pa., R.I., Vt., Va., W.Va.; Asia (Japan).
Deutzia crenata and D. scabra are highly variable and have long been variously interpreted. A. J. Rehder
(1920), assisted by H. O. Juel of Uppsala University, studied the three specimens at UPS that Thunberg had named
D. scabra. He selected one as the type but said it was the same as the later-described D. crenata. J. Ohwi (1965)
listed eight synonyms for D. scabra and interpreted D. crenata as consisting of three varieties, which included six
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synonyms, including “D. scabra Thunberg in part.” In view of the above, and the many cultivars that have been
developed, I was unable to separate D. crenata from D. scabra.
3. Deutzia parviflora Bunge, Enum. Pl. China Bor., 31. 1833
I
Shrubs 10–20 dm. Branches ascending to erect. Leaves: petiole 2–8 mm, glabrous; blade ovate or eliptic to ovatelanceolate, 30–90 × 20–40(–50) mm, base cuneate or rounded, margins serrulate, apex acute to acuminate, abaxial
surface green, glabrous or with 6–9-rayed trichomes, adaxial green, glabrous or with 3–5-rayed, stellate trichomes.
Inflorescences corymbose, open, 10–60-flowered, 5–10 × 4–7 cm, glabrous; peduncle 20–80 mm, villous or
stellate-pubescent. Pedicels 10–15 mm, stellate-pubescent. Flowers faintly fragrant, 9–10 mm; floral tube ovoid,
2.2–3 × 1–1.3 mm, densely stellate-pubescent, lobes ovate-deltate or broadly ovate, 0.9–1.2 × 0.3–0.4 mm, apex
obtuse, surfaces densely stellate-pubescent; petals imbricate, white to pink, broadly ovate or suborbiculate, 2–7 ×
3–5 mm, stellate-pubescent; outer filaments terete, filiform, 4–5 mm, apex toothless, inner filaments terete,
filiform, 3–4 mm, apex 2-toothed; styles 3, 2–3 mm. Capsules hemispheric, 2–3 × 2–3 mm. Seeds 0.9–1.3 mm.
2n = 26, 78 (Asia).
Flowering Apr–Jun. Disturbed areas, moist ravines; 100–200 m; introduced; Ga.; Asia (n China, Korea).
Deutzia ×lemoinei Lemoine, the hybrid between D. parviflora and D. gracilis, often has been grown in the flora
area and is considered one of the hardiest cultivars (A. J. Rehder 1940; M. A. Dirr 1998). Three varieties of D.
parviflora are recognized in China; plants in the flora area seem closest to var. parviflora.
6. PHILADELPHUS (not yet ready)
7. CARPENTERIA Torrey, Proc. Amer. Assoc. Advancem. Sci. 4: 192. 1851 * California
tree-anemone [For William Marbury Carpenter, 1811–1848, Louisiana physician and
botanist] C E
Craig C. Freeman
Shrubs. Stems erect, ascending, or spreading. Bark exfoliating in grayish sheets or strips.
Branches ascending or spreading; twigs with simple trichomes. Leaves persistent, opposite,
herbaceous to coriaceous; petiole present; blade lanceolate to narrowly elliptic or oblong,
margins entire or obscurely denticulate, usually revolute, sometimes plane; venation pinnate.
Inflorescences terminal and axillary, cymose, 3–9(–13)-flowered; peduncle present. Pedicels
present. Flowers bisexual; perianth and androecium nearly hypogynous; sepals 5–7, connate
proximally into tube; floral tube patelliform, free from base of ovary, without ribs in fruit,
sparsely to densely appressed-pubescent abaxially, lobes persistent, valvate, spreading, ovate
to ovate-lanceolate or triangular; petals 5–7(–8), imbricate, spreading to reflexed, white,
ovate, round, or depressed-elliptic, base sessile or obscurely clawed, surfaces glabrous; stamens
150–200; filaments distinct, essentially terete, not noticeably tapered from base to summit,
apex not 2-lobed; anthers depressed-ovate; pistil 5–7-carpellate; ovary nearly superior, 5–7locular; placentation axile proximally, parietal distally; style persistent, 1. Capsules conic to
depressed-spheric, corticate to coriaceous, dehiscence basipetally septicidal to near base of
fruit, delayed apically by persistent style. Seeds ca. 50 per locule, brown or reddish brown,
ellipsoid. x = 10.
Species 1: California.
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Phylogenetic analyses of morphological (L. Hufford 1997) and molecular (D. E. Soltis et al.
1995) data place Carpenteria as sister to Philadelphus. Androecium development in the two
genera is unique in Hydrangeaceae (Hufford 1998).
SELECTED REFERENCE Cheatham, N. H. 1974. Carpenteria, the mystery shrub. Fremontia 1: 3–8.
1. Carpenteria californica Torrey, Smithsonian Contr. Knowl. 6(2): 12, plate 7. 1853
C
E
F
Stems 10–30(–40) dm. Twigs glabrous or sparsely to moderately appressed-pubescent. Leaves: petiole (2–)5–10
mm, appressed-pubescent abaxially, glabrous or sparsely pubescent adaxially; blade 2.7–8.5(–10) × 0.8–2.5(–3.6)
cm, base acuminate, apex acute to rounded, abaxial surface whitish, densely appressed-tomentulose, adaxial green,
glabrous. Inflorescences open; peduncle 1.5–5 cm, glabrous or sparingly appressed-pubescent; bracts prominent,
sessile. Pedicels 1.5–5.2 mm, sparsely to densely strigose. Flowers: fragrant; floral tube 2–4 × 8–13 mm, lobes (6–
)8–14 × (3–)4–6 mm, margins entire, apex acute to acuminate, abaxial surface glabrous or sparingly pubescent
except for tomentulose tip; petals 20–40 × 20–36(–40) mm; filaments 4–9 × 0.2–0.6 mm; anthers 0.9–1.1 mm;
style 3.5–6 mm. Capsules 10–12 × 8–12 mm. Seeds 0.7–1 mm. 2n = 20.
Flowering Apr–Jun. Dry, rocky slopes in chaparral and lower ponderosa pine woodlands; of conservation
concern; 300–1500 m; Calif.
Carpenteria californica is a fire-adapted shrub endemic to the central and southern Sierra Nevada Foothills
between the Kings and San Joaquin rivers, Fresno County (R. Casamajor 1950). Rare in the wild, it has a long
history of cultivation in both Europe and the United States (including elsewhere in California) (N. H. Cheatham
1974), where it is prized for its lightly scented, showy flowers and attractive, persistent foliage. The plant does
well in sunny or partially shaded, sheltered sites with well-drained soils.
8. HYDRANGEA Linnaeus, Sp. Pl. 1: 397. 1753; Gen. Pl. ed. 5, 189. 1754 * Graybeard,
sevenbark, hortensia [Greek hydor, water, and angeion, diminutive of angos, vessel or
container, alluding to shape of mature, dehisced capsule]
Craig C. Freeman
Shrubs or trees. Stems erect, ascending, or spreading. Bark exfoliating in grayish, brown, or
reddish brown sheets. Branches erect, ascending, or spreading, sometimes arching; twigs with
simple or branched trichomes. Leaves deciduous, opposite or 3-whorled, herbaceous; petiole
present; blade ovate, elliptic-ovate, elliptic, or suborbiculate (sometimes lobed), margins
serrate to serrulate, plane; venation pinnate. Inflorescences terminal, cymose panicles, 100–
1000-flowered; peduncle present. Pedicels present. Flowers bisexual or, often, marginal ones
asexual; perianth and androecium epigynous or perigynous; sepals 5, connate proximally into
tube; floral tube campanulate or hemispheric, adnate to ovary to near its top, weakly or
strongly 7–10(–11)-ribbed in fruit, glabrous or, rarely, abaxially sparsely hairy, lobes
persistent, valvate, spreading or reflexed, deltate to shallowly triangular; petals 5, valvate,
spreading or reflexed, white to yellowish white, ovate-lanceolate, elliptic, oblong, spatulate, or
narrowly ovate to ovate, base sessile, surfaces glabrous; stamens 10; filaments distinct, terete
or flattened proximally, gradually or abruptly tapered from base to summit, apex not 2-lobed;
anthers depressed-ovoid or transversely oblong; pistil 2–4-carpellate; ovary completely inferior
or nearly so, or half-inferior, 2–4-locular; placentation axile proximally and parietal distally;
styles persistent, 2–4, distinct or connate at middle or distally. Capsules hemispheric,
suburceolate, or oblong-ovoid, coriaceous, dehiscence interstylar, creating elliptic to circular
pore at base of styles. Seeds 10–40 per locule, light brown to dark brown, fusiform or
ellipsoid. x = 18.
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Species ca. 29 (5 in the flora): United States, Mexico, Central America, w South America, e
Asia.
Hydrangea enjoys considerable esteem as an ornamental shrub, especially for its prominent
asexual flowers. North American species have been cultivated in Europe since before the mid
1700s (W. L. Stern 1978). Besides the species treated here, popular ornamentals in North
America are H. anomala D. Don, H. aspera D. Don, H. heteromalla D. Don, H. involucrata
Siebold, H. macrophylla (Thunberg) Seringe, and H. serrata (Thunberg) Seringe. Among
these, H. macrophylla may be the most widely grown; M. A. Dirr (2004) listed nearly 170
cultivars of this species. Surprisingly, it has not escaped from cultivation.
Tubercles comprising crystals of calcium carbonate often are visible on leaf trichomes (G.
W. Burkett 1932). They are observed most easily at magnifications greater than 30×, and
some taxonomic utility has been ascribed to their presence and abundance.
Toxic and medicinal properties are attributed to some native and cultivated species of
Hydrangea (J. M. Kingsbury 1964; W. L. Stern 1978; D. E. Moerman 1998; G. E. Burrows
and R. J. Tyrl 2001). These possibly are related to various alkaloids present in roots and
leaves of some species.
Phylogenetic studies of Hydrangeaceae (D. E. Soltis et al. 1995; L. Hufford 1997) have
indicated that Hydrangea as traditionally circumscribed is paraphyletic, excluding eight genera
apparently descended from an ancestor shared with Hydrangea. One of those genera,
Decumaria, occurs in the flora area.
SELECTED REFERENCES Dirr, M. A. 2004. Hydrangeas for American Gardens. Portland. McClintock, E. 1957. A
monograph of the genus Hydrangea. Proc. Calif. Acad. Sci., ser. 4, 29: 147–256. Pilatowski, R. E. 1980. A Taxonomic Study
of the Hydrangea arborescens Complex. M.S. thesis. North Carolina State University. Pilatowski, R. E. 1982. A taxonomic
study of the Hydrangea arborescens complex. Castanea 47: 84–98. St. John, H. 1921. A critical revision of Hydrangea
arborescens. Rhodora 23: 203–208. Stern, W. L. 1978. Comparative anatomy and systematics of woody Saxifragaceae.
Hydrangea. Bot. J. Linn. Soc. 76: 83–113.
1.
1.
Leaf blade margins (3–)5(–7)-lobed, lobes coarsely serrate; inflorescences ovoid to conic
....................................................................................................................................................4. Hydrangea quercifolia
Leaf blade margins unlobed, dentate, serrate, or serrulate; inflorescences dome-shaped to
hemispheric or conic to ovoid.
2. Ovaries
half-inferior;
inflorescences
conic
or,
sometimes,
ovoid
............................................................................................................................................. 5. Hydrangea paniculata
2. Ovaries completely inferior or nearly so; inflorescences dome-shaped to hemispheric.
3. Leaf blade surfaces white or grayish, densely tomentose (trichomes at 40× either
smooth,
1–3
mm,
or
sparsely
tuberculate,
0.3–1
mm)
abaxially
.......................................................................................................................................... 3. Hydrangea radiata
3. Leaf blade surfaces green, glabrous or nearly so, or grayish, uniformly velutinous,
pilose, or tomentose (trichomes at 40× conspicuously tuberculate, 0.3–1 mm).
4. Leaf blade surfaces green, glabrous or nearly so abaxially, or sparsely hirsute along
midribs
and,
sometimes,
along
lateral
veins
............................................................................................................................1. Hydrangea arborescens
4. Leaf blade surfaces grayish, uniformly velutinous, pilose, or tomentose abaxially
................................................................................................................................... 2. Hydrangea cinerea
1. Hydrangea arborescens Linnaeus, Sp. Pl. 1: 397. 1753 * Wild or smooth hydrangea
E
Hydrangea arborescens var. oblonga Torrey & A. Gray
Shrubs, 10–30 dm. Twigs strigose to hirsute, trichomes white. Leaves opposite; petiole 1.4–8.5(–11.5) cm,
glabrous or glabrous abaxially and sparsely tomentose adaxially; blade ovate, elliptic-ovate, or broadly ovate,
(2.7–)6–17.8 × (1.4–)2.5–12(–15.5) cm, base cordate, truncate, or cuneate, margins dentate to serrate, unlobed,
apex acute to acuminate, abaxial surface green, glabrous or nearly so, or sparsely hirsute along midribs and,
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sometimes, along lateral veins (trichomes at 40× conspicuously tuberculate, 0.3–1 mm), adaxial surface green,
glabrous or sparsely hirsute. Inflorescences compact, 100–500-flowered, dome-shaped to hemispheric, (3.3–)4–14
× 3.6–12 cm; peduncle 1.5–7.8 cm, sparsely tomentose. Pedicels 1–2.5 mm, glabrous or sparsely hirsute. Asexual
flowers absent or present, terminating lateral branches, white, greenish white, or yellowish white, tube 6–16 mm,
lobes 3–4(–5), obovate to broadly ovate, round, or elliptic, 3.6–15 × 2.2–14 mm. Bisexual flowers: floral tube
adnate to ovary to near its top, 0.7–1 × 0.8–1.2 mm, strongly 8–10(–11)-ribbed in fruit, glabrous abaxially, lobes
deltate to triangular, 0.2–0.5 × 0.2–0.5 mm, margins entire, apex acute to acuminate, abaxial surface glabrous;
petals caducous, white to yellowish white, elliptic to narrowly ovate, 1–1.5 × 0.6–1.1 mm; filaments 2–4.5 × 0.1–
0.2 mm; anthers 0.3–0.5 mm; pistils 2(–3)-carpellate; ovary completely inferior or nearly so; styles 2(–3), distinct,
0.9–1.2 mm. Capsules hemispheric, 1.2–2.1 × 1.7–2.5 mm. Seeds 0.3–0.6(–0.8) mm. 2n = 36.
Flowering (May–)Jun–Jul(–Aug). Moist to dry deciduous forests and woods, moist slopes, shaded bluffs, ledges,
streambanks; 70–2000 m; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ill., Ind., Ky., La., Md., Mass., Miss., Mo., N.J.,
N.Y., N.C., Ohio, Okla., Pa., S.C., Tenn., Va., W.Va.
Hydrangea arborescens has escaped from cultivation in Connecticut and Massachusetts; it is not native there.
E. McClintock (1957) circumscribed Hydrangea arborescens as comprising three partly sympatric subspecies;
subsp. arborescens, subsp. discolor, and subsp. radiata. R. E. Pilatowski (1980, 1982) concluded that these were
best treated as three species, H. arborescens, H. cinerea, and H. radiata, citing chemical, morphological,
reproductive, and geographic discontinuities among the taxa. Most herbarium specimens are easily referred to one
of these species; occasional specimens appear intermediate between H. arborescens and H. cinerea.
The Cherokee and Delaware tribes used bark and, occasionally, leaves from Hydrangea arborescens to prepare
infusions or poultices to treat various internal and external ailments. The Cherokee used peeled twigs and
branches to make tea, or cooked twigs and branches as a vegetable (D. E. Moerman 1998).
2. Hydrangea cinerea Small, Bull. Torrey Bot. Club 25: 148. 1898 * Ashy hydrangea
E
Hydrangea arborescens Linnaeus var. deamii H. St. John; H. arborescens subsp. discolor (Seringe) E. M. McClintock; H.
arborescens var. discolor Seringe
Shrubs, 10–30 dm. Twigs sparsely hirsute to pilose, trichomes white. Leaves opposite; petiole 0.8–8.6 cm,
sparsely to densely hirsute abaxially, densely hirsute adaxially; blade ovate, elliptic-ovate, or broadly ovate, (2.5–
)4.2–14.9 × (1.7–)2.1–12 cm, base cordate, truncate, or cuneate, margins dentate to serrate, unlobed, apex acute to
acuminate, abaxial surface grayish, uniformly velutinous, pilose, or tomentose (trichomes at 40× conspicuously
tuberculate, 0.3–1 mm), adaxial surface green, sparsely hirsute. Inflorescences compact, 150–500-flowered, domeshaped to hemispheric, 3.5–14.5 × 3–16.5 cm; peduncle 0.5–6 cm, sparsely to densely tomentose. Pedicels 0.8–2.8
mm, glabrous or sparsely to densely hirsute. Asexual flowers absent or present, terminating lateral branches, white
or greenish white, tube 4–21 mm, lobes 3–4(–5), obovate to broadly ovate, round, or elliptic, 6.5–15 × 2.5–14 mm.
Bisexual flowers: floral tube adnate to ovary to near its top, rarely ca. 2/3 up ovary, 0.7–1 × 0.8–1.2 mm, strongly
8–10-ribbed in fruit, glabrous, lobes deltate to triangular, 0.3–0.5 × 0.3–0.6 mm, margins entire, apex acute to
acuminate, abaxial surface glabrous; petals caducous, white to yellowish white, elliptic to narrowly ovate, 1.2–1.5
× 0.7–1.1 mm; filaments 1.8–4.5 × 0.1–0.2 mm; anthers 0.3–0.5 mm; pistils 2(–3)-carpellate; ovary completely
inferior or nearly so; styles 2(–3), distinct, 0.8–1.1 mm. Capsules hemispheric, (1.2–)1.5–2.3 × (1.2–)2–2.8 mm.
Seeds 0.3–0.6(–0.8) mm. 2n = 36.
Flowering May–Jul(–Aug). Deciduous upland and bottomland forests, shaded cliffs, ravines, and slopes,
streambeds; 100–700 m; Ala., Ark., Ga., Ill., Ind., Kans., Ky., Mass., Mo., N.C., Okla., S.C., Tenn.
Hydrangea cinerea has escaped from cultivation in Massachusetts.
Bark or roots were used by the Cherokee to prepare infusions used as an antiemetic, emetic, cathartic, and
gynecological and liver aids (D. E. Moerman 1998).
3. Hydrangea radiata Walter, Fl. Carol., 251. 1788 * Silver-leaf or snowy hydrangea
E
Hydrangea arborescens Linnaeus subsp. radiata (Walter) E. M. McClintock
Shrubs, 10–30 dm. Twigs glabrous or sparsely to densely tomentose, trichomes usually white, sometimes brown
or orangish brown. Leaves opposite; petiole (0.7–)1.1–8 cm, glabrous or glabrous abaxially and sparsely to
densely tomentose adaxially; blade narrowly to broadly ovate or elliptic, 4–15.2 × 1.3–11.2 cm, base truncate to
cuneate, margins dentate to serrate, unlobed, apex acute to acuminate, abaxial surface white or grayish, densely
tomentose (trichomes at 40× either smooth, 1–3 mm, or sparsely tuberculate, 0.3–1 mm), adaxial surface green,
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glabrous or sparsely hirsute along major veins. Inflorescences compact, (100–)200–500-flowered, dome-shaped to
hemispheric, 4–13 × 5–14 cm; peduncle 1.6–8.3 cm, sparsely to densely tomentose. Pedicels 0.8–3.2 mm, glabrous
or sparsely hirsute. Asexual flowers usually present, rarely absent, terminating lateral branches, white or greenish
white, tube 5–14 mm, lobes 3–4(–5), obovate to broadly ovate or round, 5–18 × 3–15 mm. Bisexual flowers:
floral tube adnate to ovary to near its top, 0.9–1.2 × 1–1.3 mm, strongly 8–10-ribbed in fruit, glabrous abaxially,
lobes deltate to triangular, 0.3–0.8 × 0.3–0.5 mm, margins entire, apex acute to acuminate, abaxial surface
glabrous; petals caducous, white to yellowish white, elliptic to narrowly ovate, 1.2–1.6 × 0.7–1.1 mm; filaments
1.5–4.5 × 0.1–0.2 mm; anthers 0.3–0.5 mm; pistil 2(–3)-carpellate; ovary completely inferior or nearly so; styles
2(–3), distinct, (0.6–)0.9–1.5 mm. Capsules hemispheric, (1.3–)1.7–2.8 × 1.8–3 mm. Seeds 0.3–0.6(–0.8) mm. 2n
= 36.
Flowering May–Jul. Moist, deciduous forests, ravines, banks, rocky slopes, cliffs; 200–1200 m; Ga., N.C., S.C.,
Tenn.
4. Hydrangea quercifolia W. Bartram, Travels Carolina, 382, plate 7. 1791 * Oak-leaf hydrangea
E
F
Shrubs, 10–20 dm. Twigs densely tomentose, trichomes brown or orangish brown, sometimes white. Leaves
opposite; petiole (1–)1.5–8(–12.6) cm, densely tomentose; blade suborbiculate to ovate, (2.8–)7–26.4 × (2.6–)6.5–
26.5 cm, base truncate to cuneate, margins (3–)5(–7)-lobed, lobes coarsely serrate, apex acute to acuminate,
abaxial surface grayish, pilose to tomentose (trichomes at 40× either smooth, 1–4 mm, or tuberculate, 0.4–1 mm),
adaxial green, glabrous or sparsely hirsute. Inflorescences open or compact, 500–1000-flowered, ovoid to conic,
9–32 × (6–)8–14 cm; peduncle 4.3–7.3 cm, sparsely tomentose to hirsute. Pedicels 1–3 mm, glabrous or, rarely,
sparsely hirsute. Asexual flowers present, terminating lateral branches, white, greenish white, pink, or reddish,
tube 11–31 mm, lobes 4(–5), obovate to broadly ovate or round, 6–20 × 5–20 mm. Bisexual flowers: floral tube
adnate to ovary to near its top, 0.9–1.3 × 1.2–2.5 mm, strongly (7–)8–10-ribbed in fruit, glabrous, lobes deltate to
shallowly triangular, (0.3–)0.5–0.9 × 0.5–1 mm, margins entire, apex acute to acuminate, abaxial surface glabrous;
petals caducous, white or yellowish white, elliptic, oblong, or spatulate, 1.5–2.6 × 0.7–1.4 mm; filaments 2.5–5(–6)
× 0.1–0.2 mm; anthers 0.3–0.6 mm; pistil 2–4-carpellate; ovary completely inferior; styles 2–4, distinct, 1.5–2.5
mm. Capsules hemispheric to suburceolate, 1.5–2.5 × 2–2.5 mm. Seeds 0.6–0.8 mm. 2n = 36.
Flowering (Apr–)May–Jul. Deciduous forests, pine-oak forests, ravines and slopes, ledges; 20–400 m; Ala.,
Conn., Fla., Ga., La., Miss., N.C., S.C., Tenn.
Hydrangea quercifolia is a popular ornamental in the eastern United States, prized for its distinctive leaves,
showy inflorescences, and fall color. Records of naturalized plants exist for Connecticut and North Carolina. The
brownish color often seen in the trichomes is imparted by tanninlike substances (W. L. Stern 1978). M. A. Dirr
(2004) listed nearly 40 cultivars derived from the species.
5. Hydrangea paniculata Siebold, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 14(suppl.): 691. 1829 *
Panicled hydrangea I
Shrubs or trees, 5–70 dm. Twigs appressed-pubescent, trichomes white. Leaves opposite or 3-whorled; petiole
0.6–2.4 cm, glabrous or sparsely hirsute, blade elliptic to ovate, (5–)6.4–15 × 3–6(–10) cm, base rounded to
truncate or cuneate, margins serrate to serrulate, unlobed, apex acute or, sometimes, acuminate, abaxial surface
green, sparsely hirsute, often densely hirsute along veins (trichomes at 40× smooth, 0.5–1 mm), adaxial surface
green, glabrate or sparsely hirsute, sometimes moderately hirsute along veins. Inflorescences open or compact,
250–1000-flowered, conic or, sometimes, ovoid, 7–25 × 6–11.5 cm; peduncle 2–5.5 cm, sparsely to densely hirsute.
Pedicels 1–4 mm, glabrous or sparsely hirsute proximally. Asexual flowers present, terminating lateral branches,
white or greenish white, sometimes tinged pink or blue, tube 9–16 mm, lobes 4(–5), elliptic to round, (6–)10–21 ×
(4–)8–15 mm. Bisexual flowers: floral tube adnate to ovary to near its middle, 1–1.8 × 1.2–2.2 mm, weakly 7–10ribbed in fruit, glabrous or, rarely, sparsely hairy, lobes shallowly triangular, 0.3–0.7 × 0.5–1.2 mm, margins
entire, apex acute to acuminate, abaxial surface glabrous; petals tardily deciduous, (spreading or reflexed), white
or yellowish white, elliptic, ovate, or ovate-lanceolate, 3–4 × 1.2–1.5 mm; filaments 1.8–5 × 0.1–0.3 mm; anthers
0.3–0.6 mm; pistil 2–4-carpellate; ovary half inferior; styles 2–4, connate at middle or distally, 2–3.5 mm.
Capsules oblong-ovoid, 3.5–5 × 2–3 mm. Seeds (caudate at both ends), 1.7–3 mm. 2n = 72, 108 (Japan).
Flowering Jul–Sep. Disturbed woods, roadsides, wooded swamps; 30–500 m; introduced; Conn., Maine, Mass.,
N.H., N.Y., N.C., Pa., R.I., Va., W.Va.; e Asia.
M. A. Dirr (2004) listed 26 primary cultivars derived from Hydrangea paniculata.
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9. DECUMARIA Linnaeus, Sp. Pl. ed. 2, 2: 1663. 1763 * Climbing-hydrangea, woodvamp [Latin decumae, tenths, and -aria, possessing, alluding to sometimes 10-merous
flowers]
Ronald L. McGregor
Woody vines. Stems climbing or, sometimes, trailing and forming loose, nonflowering mats.
Bark exfoliating in grayish or reddish brown sheets, strips, or strings. Branches spreading
laterally or declining; twigs glabrous or with simple trichomes. Leaves deciduous or
semideciduous, opposite; petiole present; blade ovate, elliptic, obovate, oblanceolate,
subround, or round, herbaceous, margins entire or, rarely, dentate or lobed, plane; venation
pinnate. Inflorescences terminal (on shoots of the season), corymbose, (20–)50–100-flowered;
peduncle present. Pedicels present. Flowers bisexual; perianth and androecium epigynous;
sepals 7–10(–12), connate proximally into tube; floral tube adnate to ovary to near its top,
strongly 7–12-ribbed in fruit, turbinate, glabrous, lobes persistent, valvate, erect, triangular;
petals 7–12, valvate, spreading, white, lanceolate, elliptic, or oblong, base essentially sessile,
surfaces glabrous; stamens 20–30 (inserted on rim of floral tube); filaments distinct,
dorsiventrally flattened, linear, tapering abruptly just proximal to anther, apex not 2-lobed;
anthers suborbiculate to ovate; pistil 6–12-carpellate; ovary completely inferior, 6–12-locular;
placentation axile proximally, parietal distally; style persistent, 1. Capsules turbinate,
cartilaginous, dehiscence intercostal (lateral walls separating from ribs), eventually leaving
cagelike remnants. Seeds 10–20 per locule, yellow, fusiform. x = 14.
Species 2 (1 in the flora): se United States, Asia (China).
1. Decumaria barbara Linnaeus, Sp. Pl. ed. 2, 2: 1663. 1763
E
F
Stems to 100 dm; adventitious roots usually present. Twigs glabrous. Leaves: petiole slightly winged proximally,
10–30 mm, glabrous or short-pubescent; blade 10–12 × 6–8 cm, base cuneate, truncate, cordate, or rounded, apex
acute, obtuse, rounded, or mucronate, abaxial surface light green, finely pubescent along veins, adaxial dark green,
glabrous. Inflorescences compact, congested to open, 3–8 × 4–10 cm; peduncle 2–6 cm, glabrous. Pedicels 1–6
mm, glabrous. Flowers faintly fragrant; floral tube 1.5–2.2 × 0.5–2 mm; lobes 0.2–1.3 × 0.2–0.5 mm, margins
entire, apex acute to acuminate, surfaces glabrous abaxially; petals 2.8–3.2 × 1.2–1.7 mm; filaments 3–5 × 0.2–0.3
mm; anthers 0.7–1 mm; style stout, broad at base, 0.1–2 mm, or absent; stigmatic lines 7–12, radiating. Capsules
3.2–5.5 × 3–5 mm. Seeds 1.5–3 mm. 2n = 28.
Flowering Apr–Jun; fruiting Jul–Oct. Very moist to wet woodlands and swamps mostly on Coastal Plain, rich
woodlands; 0–400 m; Ala., Ark., Del., Fla., Ga., La., Miss., N.Y., N.C., S.C., Tenn., Va.
Decumaria barbara is a rare escape from cultivation in New York.
Decumaria sinensis Oliver, of central China, differs from D. barbara in being a low-climbing vine with
semipersistent or persistent leaves, stigmas almost completely sessile, and plants growing at elevations of 600–1300
meters.