Family 33. NYCTAGINACEAE

NYCTAGINACEAE
Family 33. NYCTAGINACEAE
A.G. MILLER
Herbs or climbing shrubs. Leaves alternate or opposite, simple, exstipulate. Flowers
bracteate, in terminal or axillary cymes, sometimes whorled, umbellate or glomerulate;
actinomorphic, unisexual or bisexual. Perianths 5-lobed, tubular, petaloid, the lower
part persistent around the ovary. Stamens 1-10. Ovary superior, I-celled, I-ovulate;
style simple. Fruit (the anthocarp) a I-seeded achene enclosed by the persistent part
of the perianth.
Species of Bougainvillea are commonly cultivated as ornamentals in Arabia. They
are woody climbers with insignificant flowers concealed by large, showy bracts.
I.
+
Woody climbers or small trees; stems armed with curved spines
Herbs; stems not spiny
2.
Flowers showy, 3.5-4cm long, surrounded by a calyx-like involucre of bracts
1. Mirabilis
3
Flowers small, up to 1cm; bracts inconspicuous
+
3.
+
4. Pisonia
2
2. Boerhavia
Fruits 5-ribbed, glabrous or glandular-hairy
Fruits obscurely IO-ribbed, with conspicuous sessile or stalked, wart-like
3. Commicarpus
glands
I. MIRABILIS L.
Perennial herb. Leaves opposite. Flowers bisexual, in crowded terminal cymes, showy;
bracts united into a persistent 5-lobed and calyx-like involucre. Perianths trumpet­
shaped. Stamens 5-6, exserted. Stigma capitate, exserted. Fruit ribbed, eglandular.
143
NYCTAGINACEAE
1. M. jalapa L., Sp. pi.: 177 (1753). Illustr.: Fl. Pakistan 115: p.2 (1977).
Robust herb; stems erect or sprawling, up to lm. Leaves triangular-ovate, 5-12 x 27cm, acuminate, the base truncate or cordate; petiole l-3cm. Involucre clem long,
campanulate, divided to the middle into 5 acute teeth. Perianths red, rarely white or
yellow, 3.5-4cm; limb 5-lobed, 2-3.5cm across. Fruit subglobose, 7-8 x 5mm. Map
165, Fig. 26.
Mainly near villages and abundant in high rainfall areas or near permanent water;
(0-) 100O-2500m, the lower altitudes refer to cultivated material.
Saudi Arabia, Yemen (N & S), Kuwait. A native of tropical America, cultivated as an
ornamental throughout the tropics and widely naturalized; now rarely cultivated in
Arabia.
2. BOERHAVIA L.
Annual or perennial herbs, sometimes woody-based. Leaves opposite, entire. Flowers
bisexual in terminal or axillary umbellate heads or open panicles; bracts minute.
Perianths funnel-shaped or campanulate. Stamens 1-3, shortly exserted. Fruit 5ribbed, glabrous or glandular-hairy.
1.
+
2.
+
Stems erect; flowers solitary or rarely paired on capillary peduncles; inflor
escence a diffuse, much-branched, terminal panicle
1. B. elegans
Stems prostrate, ascending or sprawling; flowers in (2-)3-many-flowered
pedunculate clusters which are either axillary or form a loose terminal panicle
by reduction of the upper leaves
2
Stems prostrate; flowers in simple axillary pedunculate clusters; clusters shor
ter than or barely exceeding the leaves; fruits frequently crowned by the
persistent perianth
2. B. repens
Stems sprawling or ascending; flower clusters axillary or terminal, often in
panicles; peduncles exceeding the leaves; fruits not usually crowned by the
persistent perianth
3. B. diffusa
1. B. elegans Choisy in DC, Prodr. 13 (2): 453 (1849). Syn.: B. rubicunda Steud.,
Nomencl. bot., ed. 2, 1: 213 (1841) nom. nud. Type: Saudi Arabia, Schimper 744 (E,
K).
Woody-based perennial; stems erect, up to 50cm, puberulous below. Leaves nar
rowly lanceolate to oblong-ovate, 20-50(-60) x 3-10(-17)mm; petioles up to 10mm.
Flowers solitary or rarely paired, on capillary peduncles in erect, much-branched,
diffuse panicles. Perianths pink or reddish purple, 2-3mm long. Fruit clavate, 34.5 x 1-1.25mm, puberulous in the furrows. Map 166, Fig. 26.
1.
144
Inflorescence ± regularly dichotomously dividing; pedicels usually more than
1.5cm long; leaves usually narrowly lanceolate to narrowly oblong
subsp. stenophylla
2. Boerhavia
+
Inflorescence with smaller secondary branches arising from a ± continuous
primary axis; pedicels usually less than 1cm; leaves ovate to ovate-lanceolate
subsp. elegans
subsp. elegans
Dry rocky slopes and wadi-beds in semi-desert; usually on basalt; 0-650m.
Saudi Arabia, Yemen (S). Endemic.
subsp. stenophylla (Boiss.) A.G. Miller in Edinb. J. Bot. 51 (1): 40 (1994). Syn.: B.
elegans var. stenophylla Boiss., Fl. Orient. 4: 1046 (1879). Illustr.: Fl. Pakistan, 115:
p.8 (1977)as B. rubicunda; Collenette (1985 p.375);Western (1989p.44). Type: Oman,
Aucher-Eloy 5250 (G).
Open rocky slopes, wadi-beds and roadsides in semi-desert; usually on limestone;
100-900m.
Saudi Arabia, Yemen (S), Oman, UAE. Ethiopia, Sudan, Kenya, Tanzania, Uganda,
Iran, Pakistan and India.
The two varieties of B. elegans are clearly distinguished by their inflorescences: in
subsp. stenophylla the inflorescence is an open, dichotomising panicle with the fruits
borne on long, delicate pedicels giving the plants a mist-like appearance when viewed
from a distance; subsp. elegans has a more or less monochasially branching panicle
and shorter pedicels which never give the mist-like appearance so characteristic of
subsp. stenophylla. Subsp. stenophylla is widespread in the SE of the Arabian peninsula
but in the west is restricted to the dry, limestone plateaus of the interior; outside
Arabia it is distributed from Pakistan to E Africa. Subsp. elegans is endemic to Arabia
where it has a disjunct distribution being found in two small areas, apparently always
on basalt, around Jeddah in Saudi Arabia and around Aden in Yemen.
2. B. repens L., Sp. pi.: 3 (1753). Syn.: B. diandra L., Sp. pi.: 1194 (1753); B. repens L.
var. glabra Choisy in DC, Prodr. 13(2): 453 (1849). Illustr.: Fl. Pakistan, 115 p.6
(1977) as B. diandra.
Annual or perennial herb; stems prostrate, up to 75cm, glabrous or sparsely pub
erulous. Leaves ovate-triangular or oblong-ovate, 5-20x5-15mm, often whitish
beneath. Flowers in short-stalked axillary clusters; peduncles 5-8mm; clusters 1-6flowered. Perianths pale pink or white, c.lmm. Fruit 3 x 1mm, thinly pubescent and
glandular, usually crowned with the persistent perianth. Map 167, Fig. 26.
Sandy, silty and stony ground, roadsides, wadis etc.; 50-1700m.
Saudi Arabia, Yemen (N & S). Widespread in tropical Africa and Asia.
3. B. diffusa L., Sp. pi.: 3 (1753). Syn.: B. ascendens Willd., Sp. pi. 1: 19 (1797); B.
coccinea Miller, Gard. diet. ed. 8, 4 (1768); B. diffusa var. viscosa (Lagasca & Rod
riguez) Heimerl in Beitr. Syst. Nyctag.: 27 (1897); B. glutinosa sensu Deflers (1889
p.192) non Vahl; B. repens sensu Balf.f. (1888) non L.; B. repens var. diffusa (L.)
145
NYCTAGINACEAE
Fig. 26. Nyctaginaceae (1) A, Boerhavia diffusa: Aa, flowering shoot (x 1). B, B. repens: Ba, flowering shoot
(x 1); Bb, fruit (x 7). C, Mirabilis jalapa: Ca, flowering shoot (x 0.5). D, Boerhavia eleganssubsp. elegans:
Da, flowering shoot (x 0.6). E, B. elegans subsp. stenophylla: Ea, flowering shoot (x 0.6); Eb, fruit (x 10).
F, Pisonia aculeata: Fa, flowering shoot (x 0.5); Fb, fruit (x 3).
146
3. Commicarpus
Boiss., Fl. orient. 4: 1046 (1879); B. repens var. viscosa Choisy in DC, Prodr. 13(2):
453 (1849); B. repens sensu Collenette (1985) non L. Illustr.: Collenette (1985 p.375)
as B. repens.
Annual or perennial herb. Stems prostrate or sprawling, up to lm long, glabrous,
sparsely pubescent or sparsely to densely glandular, sometimes viscid. Leaves ovate
or oblong-ovate to subcircular, 10-50 x5-45mm, often paler beneath. Flowers in
long-stalked axillary clusters or (by reduction of the upper leaves) + leafless panicles;
peduncles l-3cm; clusters 3-12-flowered. Perianths usually pink, rarely magenta,
1.25-1.5mm long. Fruit 3-4 x 1.25-1.5mm, puberulous and usually glandular-viscid,
rarely + glabrous. Map 168, Fig. 26.
Found in a wide variety of habitats including sandy and gravelly plains, wadis, rocky
slopes, Acacia-Commiphora bushland, field borders and roadsides; 0-2300m.
Saudi Arabia, Yemen (N & S), Socotra, Oman, UAE. Pantropical.
A confusing complex badly in need of a world-wide revision. I have been unable to
arrive at a satisfactory treatment for the Arabian plants, and therefore this account
must be considered provisional. Within B. diffusas.l. different authors have recognized
a number of taxa at specific, subspecific and varietal level. Extreme forms are fairly
distinct, but the numerous intermediates make it impossible to define recognisable
taxa. For alternative treatments see Codd in Bothalia 9 (1): 113-121 (1966) and
Stannard in Fl. Zamb. 9 (1): 20-25 (1988).
3. COMMICARPUS Standley
Annual or perennial herbs, sometimes woody-based. Leaves opposite, entire or sinu
ously lobed. Flowers bisexual, in dense heads, umbels or whorls; bracts minute.
Perianths funnel-shaped, tubular below. Stamens 2-6, exserted. Stigma capitate,
exserted. Fruit 10-ribbed, studded with conspicuous, sessile or stalked wart-like glands.
Meikle, R.D. (1978). A key to Commicarpus. Notes Roy. Bot. Gard. Edinburgh 36:
235-249.
Measurements referring to the perianth exclude the basal anthocarp.
1.
+
Flowers in shortly pedunculate axillary umbels which are shorter than or
equalling the subtending leaves; peduncles up to 5mm
14. C. reniformis
Flowers in inflorescences longer than the subtending leaves; peduncles more
than 10mm
2
2.
Flowers numerous, sessile or shortly pedicellate, crowded into a dense head
borne on a stout peduncle
1. C. pedunculosus
+
Flowers in umbels or whorls but not in dense heads
3.
Stems, at least in the region of the inflorescence, conspicuously glandularpilose and very sticky
4
Stems glabrous, scabridulous, puberulous or clothed with a short indu
mentum of crispate hairs, not sticky
6
+
3
147
NYCTAGINACEAE
4.
+
5.
+
Buds densely covered with stalked glands; flowers usually in umbels; inflor
escence-region leafy
2. C. grandiflorus
Buds glabrous or thinly pilose; flowers in whorls or umbels; inflorescenceregion leafless
5
Flowers in whorls; stems with crisped hairs below; stamens 3
3. C. stenocarpus
Flowers in umbels or rarely in whorls; stems glabrous or scabridulous below;
stamens 3-5
8. C. heimerlii
6.
+
Stems, at least in the lower part, closely crispate-pubescent or puberulous 7
Stems glabrous, or at most sparsely scabridulous
9
7.
Leaves sinuate or lobed; flowers sessile or very shortly pedicellate
+
Leaves entire; flowers distinctly pedicellate
8.
Leaves fleshy; stems crispate-pubescent; fruit pubescent between the glands
+
Leaves not fleshy; stems sparsely puberulous; fruit glabrous between the
glands
4. C. arabicus
9.
+
Inflorescence of umbels
10
Inflorescence of whorls
11
6. C. sinuatus
8
9. C. mistus
10.
+
Inflorescence-region a leafless mass of slender, intricately branched peduncles
and pedicels; fruit fusiform, inconspicuously glandular
7. C. simonyi
Inflorescence an open panicle of umbels; fruit clavate, prominently glandular
at the tip
11.
+
Fruits clavate
Fruits fusiform
4. C. arabicus
12
14
12.
Flowers sessile or very shortly pedicellate; pedicels usually less than 4mm
(those in the lowest whorl sometimes longer); perianths up to 2.5mm long
+
Flowers distinctly pedicellate, pedicels often more than 1cm; perianths 3-
13. C. helenae
5mm long
13.
+
13
Inflorescence normally of 1—2(—3) whorls of flowers; perianths widely infun
dibuliform, without a distinct basal tube; stamens 3(-4)
10. C. boissieri
Inflorescence of 3-4(-5) whorls of flowers; perianths narrowly infun
dibuliform, with a distinct basal tube; stamens 2, rarely 3 11. C. ambiguus
14.
Perianths 2-3mm, broadly infundibuliform, without a basal tube; stamens 2
+
Perianths 7-10mm, narrowly infundibuliform, with a well-developed basal
12. C. adenensis
tube; stamens 3
15.
148
Perianths white; inflorescences mainly of whorls
15
5. C. plumbagineus
3. Commicarpus
+
Perianths mauve, pink or purple; inflorescence-region a leafless mass of
slender, intricately branched peduncles and pedicels
7. C. simonyi
1. C. pedunculosus (A. Rich.) Cuf., En. PI. Aeth.: 79 (1953). Syn.: Boerhavia pedunculosa A. Rich. (1850 p.210).
Stems finely pubescent. Leaves ovate to ovate-triangular, up to 4cm, entire. Flowers
sessile or shortly pedicellate, crowded into dense heads at the tips of stout peduncles.
Perianths magenta, c.6mm long, narrowly funnel-shaped with a well-developed basal
tube, externally thinly puberulous. Stamens 3-6. Fruit clavate, prominently glandwarted at the apex, smooth below and lacking glands. Map 169, Fig. 27.
Cliffs, stony plains and roadsides, often on volcanic rocks; 2100-2600m.
Yemen (N). Rwanda, Somalia, Sudan, Ethiopia, Uganda, Kenya and Tanzania.
All specimens examined from Arabia have 3-4 not 5-6 stamens as stated by Meikle
(op. cit.).
2. C. grandiflorus (A. Rich.) Standley in Contr. U.S. Natl. Herb. 18: 101 (1916). Syn.:
Boerhavia grandiflora A. Rich. (1850 p.209); B. plumbaginea Cav. var. grandiflora
(A.Rich.) Schweinf. (1896 p.167). Illustr.: Collenette (1985 p.375).
Stems glandular-pilose, viscid. Leaves ovate to ovate-triangular, up to 6cm, entire.
Flowers pedicellate, in umbels; pedicels 3-8mm. Perianths pink, mauve or purple, 68mm long, narrowly funnel-shaped with a well-developed basal tube, densely glandu
lar externally. Stamens 3. Fruits clavate, puberulous, prominently gland-warted over
their entire surface. Map 170, Fig. 27.
Rocky slopes and roadsides; 1100-2800m.
Saudi Arabia, Yemen (N). Chad, E and NE tropical Africa and India.
3. C. stenocarpus (Chiov.) Cuf., Enum. PI. Aeth.: 81 (1953). Syn.: Boerhavia stenocarpa
Chiov., Fl. Somala: 283 (1929).
Stems shortly glandular-pilose, viscid, with crispate hairs below. Leaves ovate or
subcircular to transversely ovate, up to 4cm, entire. Flowers shortly pedicellate, in
whorls; pedicels l-4mm. Perianths mauve or purple, (4-)7-8mm long, narrowly
infundibuliform with a well-developed basal tube, glabrous or thinly pilose externally.
Stamens 3. Fruits fusiform, with gland-warts prominent at the tip and sessile or
inconspicuous below. Map 171, Fig. 27.
Dry rocky slopes, rocky and sandy plains and wadi-beds; 0-1000m.
Yemen (S), Oman, UAE. Iran, Pakistan and Somalia.
4. C. arabicus Meikle in Kew Bull. 32: 474 (1978). Boerhavia plumbaginea Cav. var.
forskalei Schweinf. (1896 p. 167). Type: Yemen (N), Hepper 5659 (K).
Stems glabrous or sparsely scabridulous. Leaves not fleshy, ovate, up to 5cm, entire.
Flowers distinctly pedicellate, in umbels, very rarely in whorls or irregular. Perianths
mauve, pink or purple, c.5-7mm long, narrowly infundibuliform with a well149
NYCTAGINACEAE
Fig. 27. Nyctaginaceae. A, Commicarpus pedunculosus. B, C. grandifolius. C, C. stenocarpus.D, C. arabicus.
E, C. heimerlii. F, C. simonyi. G, C. mistus. H, C. sinuatus: He, leaf (x 3). I, C. plumbagineus: Ic, flower
(x 10) - a, fruits (all x 7); b, inflorescences (all x 1).
150
3. Commicarpus
developed basal tube, externally glabrous, sparsely glandular or puberulous. Stamens
3. Fruits clavate, prominently gland-warted at the tip, ± smooth and inconspicuously
glandular below. Map 172, Fig. 27.
Rocky and disturbed ground, roadsides etc.; 1500-2900m.
Saudi Arabia, Yemen (N & S). Endemic.
The records of this species from Saudi Arabia need confirmation.
5. C. plumbagineus (Cav.) Standley in Contr. U.S. Natl. Herb. 18: 101 (1916). Syn.:
Boerhavia dichotoma Vahl, Enum. pi. 1: 290 (1804); B. plumbaginea Cav., Icon. 2: 7,
t.112 (1793); B. plumbaginea Cav. var. dichotoma (Vahl) Asch. & Schweinf. (1896
p.167); B. plumbaginea Cav. var. glabrata Boiss., Fl. orient. 4: 1044 (1879); IB.
scandens Forsskal (1775) non L.; B. verticillata Poiret in Lam., Encycl. 5: 55 (1804);
Commicarpus verticillatus (Poiret) Standley in Contr. U.S. Natl. Herb. 18: 101 (1916);
Valeriana scandens sensu Forsskal (1775 p. 12) non Loefl. Illustr.: Collenette (1985
p.376).
Stems glabrous or at most sparsely scabridulous. Leaves ovate to subcircular, up
to 5cm, entire or sinuate. Flowers distinctly pedicellate, mainly in whorls. Perianths
white, 8-10mm long, narrowly infundibuliform with a well-developed basal tube,
externally puberulous. Stamens 3. Fruit narrowly fusiform, prominently gland-warted
at the tip, with scattered inconspicuous sessile glands below. Map 173, Fig. 27.
Rocky and sandy plains, wadi-beds, walls, Euphorbia shrubland and as a weed of
irrigated ground; 400-2400m.
Saudi Arabia, Yemen (N & S), Bahrain. Spain, tropical and southern Africa and
Madagascar.
6. C. sinuatus Meikle in Kew Bull. 29: 83 (1974). Syn.: Boerhavia plumbaginea Cav.
var. viscosa Boiss., FL orient. 4: 1044 (1879). Illustr.: Collenette (1985 p.376).
Stems puberulous throughout. Leaves sinuate or lobed, up to 4cm. Flowers sessile
or shortly pedicellate, in umbels. Perianths pink, 6-9mm long, narrowly infun
dibuliform with a well-developed basal tube, externally puberulous. Stamens 3-4.
Fruit clavate, puberulous and covered with sessile glands. Map 174, Fig. 27.
Dry rocky slopes, sandy wadis and disturbed ground; 100-2600m.
Saudi Arabia, Yemen (N & S). Sinai, Somalia and Ethiopia.
7. C. simonyi (Heimerl & Vierh.) Meikle in Hooker's Icon. PI. 37: t.3694 (1971). Syn.:
Boerhaviascandenssensu Balf.f. (1888) non L.; B. simonyiHeimerl & Vierh. in Oesterr.
Bot. Z. 53: 435 (1903); B. plumbaginea Cav. var. socotrana Heimerl. Type: Yemen
(Socotra), Simony (WU).
Stem glabrous, or at most very sparsely scabridulous. Leaves ovate-triangular to
subcircular, up to 5cm, entire. Flowers distinctly pedicellate, in umbels or rarely and
abnormally in whorls or irregular, forming a leafless mass of slender, intricately
branched peduncles and pedicels; pedicels up to 20mm. Perianths mauve, pink or
151
NYCTAGINACEAE
purple, 7-7.5mm long, narrowly infundibuliform with a well-developed basal tube.
Stamens 3(-5). Fruit narrowly fusiform, inconspicuously glandular. Map 175, Fig.
27.
Rocky slopes in semi-deciduous thicket on both limestone and granite; 200-850m.
Socotra. Endemic.
8. C. heimerlii (Vierh.) Meikle in Hooker's Icon. PI. 37: t.3694 (1971). Syn.: Boerhavia
heimerlii Vierh. in Oesterr. Bot. Z. 53: 435 (1903); B. ?scandens sensu Balfour (1888)
non. L. Type: Yemen (Socotra), Simony (WU).
Stem glabrous towards the base, viscid above. Leaves fleshy, ovate to subcircular,
up to 5cm, entire. Flowers distinctly pedicellate, in umbels, very seldom in whorls
or irregular; pedicels 5-7mm. Perianths pink or purple, 6-7.5mm long, narrowly
infundibuliform with a well-developed basal tube, externally glabrous. Stamens 3-5.
Fruit clavate, prominently glandular at the tip, inconspicuously glandular below.
Map 176, Fig. 27.
Rocky slopes in semi-deciduous thicket on limestone; 15-350m.
Socotra. Endemic.
9. C. mistus Thulin in Nord. J. Bot. 10(4): 405 (1990). Syn: C. squarrosus sensu auctt.
non (Heimerl) Standley. Illustr.: Collenette (1985 p.376) as C. squarrosus.
Stem crispate-pubescent towards the base. Leaves fleshy, ovate, up to 3(-5)cm,
entire. Flowers distinctly pedicellate, in umbels, very seldom in whorls or irregular;
pedicels 5-10mm. Perianths pink or purple, 4-5(-8)mm long, narrowly infun
dibuliform with a well-developed basal tube, externally puberulous. Stamens 2-3.
Fruits clavate, puberulous with prominent long-stalked glands at the tip and incon
spicuously glandular below. Map 177, Fig. 27.
Dry rocky hills, particularly on limestone; 900-2700m.
Saudi Arabia, Yemen (N & S), Oman. Ethiopia, Somalia and Kenya.
A specimen from Saudi Arabia {Collenette 6653) differs from typical plants in its
larger flowers, fruits and leaves.
10. C. boissieri (Heimerl) Cuf., Enum. PI. Aeth.: 79 (1953). Syn.: Boerhavia boissieri
Heimerl in Akad. Wiss. Wien Math.-Naturwiss. Kl., Denkschr. 71: 346 (1907).
Stems glabrous. Leaves ovate to subcircular, up to 7cm, entire or sinuate. Flowers
distinctly pedicellate, in whorls; pedicels 6-15mm. Perianths pink, purple or magenta,
3-5mm long, widely infundibuliform, with a very short, inconspicuous basal tube,
externally glabrous. Stamens usually 3(-4). Fruits clavate, prominently gland-warted
at the tip, with scattered prominent glands below. Map 178, Fig. 28.
Dry rocky hills and Euphorbia balsamifera shrubland; 0-1000m.
152
3. Commicarpus
Socotra, Oman. Pakistan and India.
11. C. ambiguus Meikle in Kew Bull. 38: 481 (1983).
Stems glabrous. Leaves broadly ovate, 3-6cm, entire or obscurely sinuate. Flowers
in whorls with long, slender pedicels up to 20mm. Perianths purple, 3-5(-8)mm long,
narrowly infundibuliform with a short, but distinct, basal tube, externally glabrous.
Stamens 2(-3). Fruits clavate, with prominent, long-stalked glands at the tip and
scattered prominent glands below. Map 179, Fig. 28.
Dry rocky and stony hills; 1220-2300m.
Saudi Arabia, Yemen (N). Ethiopia and Somalia.
12. C. adenensis A.G. Miller in Edinb. J. Bot. 51 (1): 39 (1994). Type: Yemen (S),
Miller et al 8067 (E).
Stems glabrous or occasionally minutely scabridulous. Leaves ovate or rounded,
up to 5cm, entire or sinuously lobed. Flowers shortly pedicellate (the pedicels up to
10mm in fruit), in irregular panicles of 2-3 whorls. Perianths pink or mauve, 2-3mm,
widely infundibuliform, with a short inconspicuous basal tube, externally glabrous.
Stamens 2. Fruit fusiform, prominently glandular at the tip and with inconspicuous
glands below. Map 180, Fig. 28.
Dry volcanic slopes; 0-1100m.
Yemen (S). Endemic.
13. C. helenae (Roemer & Schultes) Meikle in Hooker's Icon. PI. 37: t. 3694 (1971). Syn.:
Boerhavia helenae Schultes in Roemer & Schultes, Mant. 1: 73 (1822); B. verticillata
sensu Schwartz (1939) non Poiret; Commicarpus stellata Berhaut in Bull. Soc. Bot.
France 100: 51 (1953). Illustr.: Collenette (1985 p.376).
Stems glabrous or sparsely scabridulous. Leaves ovate-triangular, up to 5(-9)cm,
entire or sinuously lobed. Flowers sessile or shortly pedicellate, in narrow whorls;
pedicels up to 4mm. Perianths pale pink or purple, 1.5-2.5mm long, widely infun
dibuliform, with a very short, inconspicuous basal tube, externally glabrous. Stamens
usually 2. Fruits clavate, with prominent, long-stalked glands at the tip, incon
spicuously glandular below. Map 181, Fig. 28.
Stony ground, roadsides and a weed of cultivation; 10-2300m.
Saudi Arabia, Yemen (N & S), Socotra, Oman, UAE. Tropical Africa, Egypt, Palestine,
Iran and India.
An inadequate specimen (Hillcoat 395) from near Riyadh in Saudi Arabia may be
referable to this species.
14. C. reniformis (Chiov.) Cuf. in Bull. Jard. Bot. Brux. 23, Suppl.: 80 (1953).
Stems crisped-puberulous. Leaves reniform, up to 3.5 x 4cm, glabrous or thinly
puberulous. Flowers in shortly pedunculate axillary umbels; peduncles up to 5mm;
pedicels up to 9mm. Perianth mauve or purple, 8-9mm, narrowly infundibuliform
153
NYCTAGINACEAE
Fig. 28. Nyctaginaceae. A, Commicarpusboissieri. B, C. ambiguus. C, C. adenensis. D, C. helenae - a, fruits
(all x 7); b, inflorescences (all x 1); c, flowers (all x 10); d, habit ( x 1).
154
4. Pisonia
with a well-developed basal tube. Stamens 2-3. Fruits clavate, glabrous with promi­
nent long-stalked glands at the tip and inconspicuously glandular below.
Gypsum hills; I020m.
Yemen (S). Somalia.
In Arabia only collected once in an area of gypsum hills north of Mukalla.
4. PI SONIA L.
Dioecious climbing shrub or small tree; stems armed with curved spines. Leaves
opposite or alternate. Flowers in dense axillary pedunculate cymes which expand in
fruit; bracts minute. Perianths infundibuliform or campanulate. Stamens 6-8,
exserted. Stigma fimbriate, exserted. Fruit ribbed with rows of glandular hairs along
the ribs.
1. P. aculeata L., Sp. pl.: 1026 (1753). Illustr.: Fl. Zamb. 9 (1): 26 (1988).
Stems up to 8m, with opposite spreading branches. Leaves elliptic to obovate or
ovate, up to 10 x 7cm, acute or obtuse, the base obtuse to cuneate, tomentose beneath;
petiole up to 2.5cm. Flowers cream or yellow-green, 5-lobed; male flowers infundi­
buliform, c.3mm long; ovary vestigial; female flowers campanulate, c.2mm long;
stamens absent. Fruit narrowly ellipsoid, 6-10 x 2-2.5mm. Map 182, Fig. 26.
Riverine woodland; 500-1400m.
Saudi Arabia, Yemen (N). Widespread in Old and New World tropics.
155
Distribution Maps
Map 165. Mirabilis jalapa
Map 166. Boerhavia elegans
subsp. stenophylla • subsp.
elegans A
514
Map 167. B. repens
Map 168. B. diffusa
Distribution Maps
Map 169. Commicarpus
pedunculosus
Map 170. C. grandiflorus
Map 171. C. stenocarpus
Map 172. C. arabicus
Map 173. C. plumbagineus
Map 174. C. sinuatus
Map 175. C. simonyi
Map 176. C. heimerlii
Map 177. C. mistus
515
Distribution Maps
Map 178. C. boissieri
Map 179. C. ambiguus
Map 181. C. helenae
Map 182. Pisonia aculeata
516
Map 180. C. adenensis