A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar
Author
Wassel, Anna C.
11714 Darbyshire Ct., Loveland, OH 45140, U. S. A.
Author
Applequist, Wendy L.
Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, MO 63110, USA.
wendy.applequist@mobot.org
text
Candollea
2020
2020-03-24
75
1
1
23
journal article
20351
10.15553/c2020v751a1
f25c58ff-3359-40fe-8720-ee004402f0bd
2235-3658
5724766
Homalium
sect.
Nisa
(Noronha ex Thouars) Baill. ex Warb.
in
Engl. & Prantl, Nat. Pflanzenfam. 98–99: 36. 1893
.
≡
Nisa
Noronha ex Thouars
,
Gen. Nov.
Madagasc.: 24. 1806.
≡
Homalium
[unranked as §]
Nisa
(Noronha ex Thouars) Baill.
in Bull. Mens. Soc. Linn. Paris 1: 575. 1886.
Typus
(designated by
SLEUMER, 1973: 295
):
Homalium nudiflorum
(DC.) Baill.
(≡
Nisa nudiflora
DC.
).
Stipules
opposite the petiole base and fused (in
H. intercedens
Sleumer
axillary, free), usually caducous.
Leaves
with venation more or less brochidodromous.
Inflorescences
spicate; bract and bracteoles large, usually broad, thicktextured, persistent.
Flowers
sessile (rarely subsessile); perianth 5–6(–8)-merous.
Sepals
obovate to oblanceolate (to nearly ligulate), oblanceolate-oblong or narrowly elliptic (in
H. louvelianum
H. Perrier
oblong-ovate to ovate-deltoid), spreading, accrescent (only slightly so in
H. louvelianum
); calyx tube short, broadly funnelform to cup-shaped, or in fruit hemispherical.
Petals
ovate to oblong, usually smaller than sepals, modestly accrescent and curving over fruit (in
H. louvelianum
similar to sepals in size and spreading); sepals and petals not ciliate, or ciliolate in conjunction with overall surface pubescence.
Sepal glands
large, elliptic to elongated oblong-elliptic or roughly trapezoidal.
Stamens
1 per petal, inserted between glands; anthers dorsifixed, broadly oblongelliptic with oblong-elliptic locules and a large connective, the slits of dehiscence nearly parallel. Upper surface of
ovary
often nearly flat in flower, in fruit becoming hemispherical or broadly conical; styles 3–4, fused basally (rarely fused for most of length).
Locule of fruit
subglobose or vertically compressed, internally glabrous (short-pubescent); seeds sometimes 1 per fruit, subglobose, largely filling the locule (or several small immature seeds, possibly most never maturing).
Key to the species of
Homalium sect. Nisa
1. Sepals about equal to petals at anthesis, broadly oblongovate (to ovate-deltoid), in fruit slightly accrescent, <
3.5 mm
and only modestly longer than petals; petals not curving inward over fruit ............. 5.
H. louvelianum
1a. Sepals longer than petals at anthesis, oblanceolate to narrowly oblong-oblanceolate, spatulate, narrowly elliptic, or nearly ligulate (in flower obovate to elliptic), in fruit conspicuously accrescent,>
3.5 mm
and much longer than petals; petals usually curving inward as fruit matures .... 2
2. Sepal glands densely pubescent on upper (inward-facing) surface ............................................... 10.
H. stelliferum
2a. Sepal glands glabrous or glabrate throughout .............. 3
3. Leaves lanceolate; stipules free ............ 4.
H. intercedens
3a. Leaves elliptic, oblong-elliptic, obovate or oblanceolate (rarely to ovate, very rarely a few lanceolate); stipules fused, opposite petioles ................................................ 4
4. Calyx densely pubescent to sericeous at least on cup, basal portion of abaxial sepal surface .................................... 5
4a. Calyx glabrous or sparsely pubescent, often more densely short-pubescent on cup, with trichomes confined to medial or marginal areas of one or both sepal surfaces ............. 7
5. Leaves elliptic to narrowly elliptic, obovate, or oblanceolate, (2.5–)3–6(–9) × 1.3–2.5(–3) cm, without marginal glands; bracts and bracteoles often ciliate, pubescent only along midrib or glabrous; adaxial surface of sepals glabrous; petals
1.5–2.5 mm
, densely pubescent abaxially (to sparsely at margins), densely to sparsely pubescent adaxially; filaments
0.9–1.1 mm
, anthers
0.2–0.3 mm
............ .................................................................... 11.
H. tenue
5a. Leaves elliptic to broadly or narrowly elliptic or obovate, (3.3–)5–9.5 × (1.8–)2–4.5(–5.4) cm, usually with inconspicuous marginal glands; bracts and bracteoles ciliate and usually pubescent throughout (mostly along midrib); adaxial surface of sepals short-pubescent at least basally; petals
2–5 mm
, velutinous on both surfaces; filaments (1.2–)
1.9–2.8 mm
, anthers
0.4–0.65 mm
..................... 6
6. Flowers mostly borne in pairs or closely clustered; bracts and bracteoles
3.5–5 mm
; calyx cup long-sericeous, both sepal surfaces densely pubescent; southern
Madagascar
............................................................. 2.
H. ciliolatum
6a. Flowers usually borne singly; bracts and bracteoles 1.5– 2.5(– 3) mm; calyx cup densely pubescent, sepals densely pubescent towards base of both surfaces, becoming sparsely pubescent (to glabrous) near apical margins; northeastern
Madagascar
............... 1.
H. antilahimenae
7. Petioles (10–)
15–30 mm
; inflorescences paniculate, pubescent; flowers mostly paired ...................... 9.
H. retusum
7a. Petioles
3–15 mm
; inflorescences racemose or paniculate, glabrous or seldom sparsely or patchily pubescent; flowers borne singly ................................................................. 8
8. Leaves oblanceolate, (2.7–)3.4 – 6.8 × (0.9 –)
1.1 –2 cm
, with revolute margins; inflorescences racemose, (0.5–)1–2.5(–3.3) cm; filaments
0.5–0.8 mm
, anthers ca.
0.2 mm
.............................................. 6.
H. mandenense
8a. Leaves elliptic to broadly or narrowly elliptic, oblong-elliptic, or obovate (to oblanceolate in
H. nudiflorum
), 4.9–16 ×
1.5–9 cm
, with entire (sometimes partly, not strongly revolute), crenate, or serrate to dentate margins; inflorescences racemose or paniculate, (2–)
3– 12 cm
; filaments 1.6–3.1(–3.7) mm, anthers (0.3–)0.5–0.7(–0.8) mm ..... 9
9. Leaves (5 –)6 – 16 × (1.8 –)3 – 6.2(– 7) cm with petiole 3–7(–10) mm; leaf margins usually prominently toothed
(subentire) with elongated glands in tooth apices; both surfaces of petals densely (to moderately) pubescent especially on apical half; humid forests to drier central forests, at elevations of
445–1860 m
................. 3.
H. confertum
9a. Leaves 4.9– 14.5 ×
1.5–7 cm
with petiole (3–)4–13(–15) mm; leaf margins entire to repand (crenulate, partly somewhat revolute), with glands small and round or absent; petals usually moderately (to densely) pubescent only at apex of one or both surfaces (rarely on entire apical portion), otherwise sparsely pubescent with often ciliate margins, or glabrous; humid (to littoral) forests, at elevations of
12–1220 m
............................................ 10
10. Leafing internodes (1.7 –)2– 4(–5,5) mm diam.; leaves 8.5– 14.5 ×
5 –7 cm
; inflorescences paniculate; petals in fruit
3–5.6 mm
................................. 8.
H. pachycladum
10a. Leafing internodes (0.9–)1.1–2.4(–3.2) mm diam.; leaves 4.9–9(–10.5) × 1.5–4.2(–5.4) cm; inflorescences racemose (paniculate); petals
1.5–4 mm
.............. 7.
H. nudiflorum