Baum, 1995, A
Systematic Revision of Adansonia (Bombacaceae), Ann Miss Bot Gard,
82:440-470

Adansonia za Baill., Bull.
Mens.Soc.Linn.Paris, 2: 844. 1890. TYPE: Madagascar. Mahafaly plateau, Grevi
37 (lectotype, designated by Perrier de la Bâthie (1952a), P).

Adansonia za var. boinensis H. Perrier, Notul. Syst-,
14: 304. 1952. TYPE: Madagascar. Ankara, Kamakama, Oct. 1902, Perrier de la
Bdthie 960.9 (lectotype, here designated, P).

Adansonia bozy
Jum. & H. Perrier, Ann. Mus. Colon. Marseille, 18: 447-451. 1910.
Adansonia za Baill. var. bozy (Jum. & H. Perrier) H.
Perrier, Notul. Syst., 14: 304. 1952 (= Adansonia za var. bozo).
TYPE: Madagascar. Andranomandevo, Sambirano valley,' Jan. 1909, Perrier de la
Bâthie 8159 (holotype, P).

Adansonia alba Jum. & H.
Perrier, Matieres Grasses, 1909: 1511. 1909. TYPE: Madagascar. Upper basin of
the Andranomalaza to the N of Bezofo (Bezofo), in dense forest by a waterfall
close to 500 m, on rocks (syenite), Oct. 1908, Perrier de la Bâthie 8166
(holotype, P).

Usually tall (up to 30 m), rarely stunted (less than 5
m when mature), deciduous trees with usually single, cylindrical or slightly
tapering trunks, often with irregular swellings; crowns rounded; primary
branches usually ascending and tapering. Bark gray and ± smooth. Leaves
5-8-foliolate; petiole 5-15 cm x 1-4 mm; stipules caducous; leaflets sessile
to long -petiolulate (up to 3 cm), broadly elliptic to lanceolate, apex
usually apiculate, rarely mucronate; medial leaflets up to 20 x 8 cm in the
north, usually less than 10 x 4 cm in the south, with 10-20 pairs of regularly
spaced secondary veins (more in north), usually alternating with prominent
intersecondaries, glabrous or somewhat scabrous; margins entire. Flowers
emerging simultaneously or soon after the leaves; buds erect to horizontal,
elongated-cylindrical, 15-24 cm x 1.5-2.5 cm; flower stalk 2-3 cm, green.
Calyx lobes (3-)5, reflexed and twisted at the base of the flower, 15-22 cm x
10-12 mm, green and scabrous outside, dark red villous within. Calyx tube
fitting tightly around the petal bases, with a marked annular swelling (ca. 2
mm wide). Petals yellow, sometimes with a diffuse reddish medial streak on the
adaxial surface, linear, at least 10 times as long as wide, 14-24 cm x 10-16
mm, exceeding the style and androecium. Androecium pale yellow, comprising a
long cylindrical or tapering staminal tube, 4-6.5 cm, surmounted by 100-120
free filaments, up to 12 cm long. Ovary conical to ovoid, covered in dense,
upward-pointed hairs. Style dark red, straight, 16-22 cm glabrous above,
dense, upward-pointing hairs below, usually fitting loosely in the staminal
tube and persisting after floral abscission (rarely to fruit). Stigma red, 3-5
mm diam. with irregular lobes. Fruit subglobose, oblong or ovoid, 10-30 cm
long, 6-15 cm wide, rarely curved; usually with longitudinal ridges and,
except in the north, a distinctly swollen peduncle. Pericarp thick and tough
with many longitudinal fibers, usually blackish with a sparse indumentum.
Seeds reniform, laterally flattened, variable in size, up to 12 x 11 x 8 mm.
Germination phanerocotylar.

Representative specimens examined.
MADAGASCAR. Antsiranana Province: Ambanja District: Ambodibozo near Ambanja,
alt. ca. 5 m, Mar. 1951 (1, fr), Humbert & Capuron 25500 (K, P). Mahajanga
Province: Maromandia District: Antanandava I village, ca. 11 km E of
Maromandia, isolated tree ca. 1.5 km W of village, 14°16'5, 48°08'E, 1 Nov.
1991 (1, fl), Baum 332 (MO, P, TAN); Ambondrona village (ca. 2 km E of
Ambaibo), 500 m E of village near the Andranomaiaza river, 14°13'30"S,
48°22'30"E, alt. ca. 100 m, 4 Nov. 1991 (l, fl), Baum 336 (MO, P, TAN).
Marovoay District: Ankaboka near Marovoay, Dec. 1909 (1, fl), Perrier de la
Bdthie 8189 (BR, G, P). Ambato-Boeny District: Ankara, Kamakama, Oct. 1902 (1,
fl, fr), Perrier de la Bâthie 960A & 960C (P). Kandreho District: Kariza
forest on Kelifely tablelands, Nov. 1904 (fl), Perrier de la Bâthie 960E (P).
Maevatanana District: Andriba, isolated trees at foot of Mount Andriba on
Gneiss, 17 Jan. 1942 (1, fl), Decary 17088 (P). Toliara Province: Mcrondava
District: Kirindy forest, 4.5 km E of Morondava to Belo-sur-Tsiribihina road
42 km from Morondava, 50 m from the Kirindy river bed, 20°02'S, 44°39'E, alt.
25 m, 24 Dec. 1991 (1, fl), Baum 344 (MO, P, TAN); near Analaiva, E of
Morondava, 28 Dec. 1962 (1, fl), SF (Capuron) 22141 (A, BR, G, K, bf0, P,
TEF). Mahabo District: 21 km E of Mahabo, 4 Dec. 1920 (fl), Keraudren-Aymonin
& Aymonin 25917 (P). Miandrivazo Dis trict: Sakeny River, Aug. 1910 (1,
fl), Perrier de la Bathie 8180 (P). Morombe District: Morombe, Oct. 1940 (fr),
Decary 16426 (P). Sakaraha District: Analamarina, SE of Sakaraha, alt. 500 m,
28 Dec. 1961 (l, fl), SF (Capuron) 20595 (BR, G, MO, P, TEF). Toliara
District: limestone plane of the basin of the Fiherana, Jan. 1910 (1), Perrier
de la Bâthie 8182 (P). Betioky District: near Beza Mahafaly reserve, near
Betioky, valley of the Ehazoara river, E of Sakamena, 23°40'S, 44°39'E, alt.
170 m, 29 Nov. 1987 (1, fl), P. B. Philippson 2638 (MO, P). Ampanihy District:
between Beloha and Ampanihy, 19 Feb. 1949 (1, fl), SF 455 (P, TEF). Bekily
District: Ambatofotsy, 15 Jan. 1957 (1, fl), SF 3376 (P, TEF). Ambovombe
District: Ambovombe, 24 Jan 1925 (fl, 1), Decary 3554 (BR, G, P, TAN).
Amboasary District: Fangidraty NE Androy, 21 Nov. 1931 (I, fl), Decary 9319
(K, P); Andohahela parcelle 2, NE of Amboasary near Hazafotsy, 24°50'5,
46°32'E, 100 m, 12 Dec. 1991 (l, fl), S. T. Malcomber 1141 (MO, P). s.n. (P).
MADAGASCAR. Jardin Tsimbazaza, 7 May 1941 (I), Jardin Tsimbazaza 4896 (TAN).
Immature or incomplete material with uncertain determinations. Near Radama [=
Sahamalaza], Maromandia, 11 Oct. 1922 (imm. 1), Decary 1146 (P); N of
Fiherana, dry forest of Fandrare, 40 km N of Tulear, Mar. 1934 (juv. shoot),
Humbert 14348bis (P); Besevo, Oct. 1899 (1), Perrier de la Bâthie 960 (P);
Ambolibozo, Sambirano May 1924 (seedlings), Perrier de la Bâthie 16340a (P).

Figure 6. The distribution of Adansonia za
in
Madagascar. Collection sites are marked with a circle.
When
more than one collection is made from the same
locality the number
of separate collections is
indicated.

Distribution (Fig. 6).
Adansonia za occurs in dry deciduous forest, spiny forest, savannas,
and scrubland from Andohahela and the Mandrare River in the southwest through
southern and western Madagascar to the Boina region and the Sambirano River
basin. In some deciduous forests in the south it constitutes the dominant tree
species (e.g., Analava west of Betioky and to a lesser extent Sakaraha abd the
forests east of Ankilizato). In the northwest it is less abundant,
concentrated near rivers.

Phenology. In leaf throughout the wet
season. Flowering November to February, earlier in the north, later in the
south. Ripe fruit from end of dry season.

Cytology.Miege (1974) reported 2n = 48 for specimens collected
in the southwest of Madagascar. Baum & Oginuma (1994) found 2n = 88 from
both south and north Madagascar.

Taxonomy and nomenclature. As
discussed above, there are few fixed differences between Adansonia za
and A. madagascariensis. The case for exclusivity is weaker in A.
za than it was in A. madagascariensis in view of its wide
geographical range and the fact that some individuals in the north share
features with A. madagascariensis that are absent from southern
individuals assigned to A. za. One possibility is that A. za is
a "metaspecies" (Donoghue, 1985), i.e., that the organisms constituting A.
za are not members of any exclusive species. This situation could have
arisen, for example, under the following scenario, which assumes for the sake
of illustration that A. madagascariensis and A. za are sister
groups. (1) A small subset of the common ancestral taxon became genetically
isolated. (2) This subset continued to have a small population size and/or
went through a severe bottleneck, becoming the exclusive species A.
madagascariensis. (3) The remainder of the common ancestor, constituting
the putative taxon "A. za," retained a large population size and did
not become exclusive. If this scenario were correct, the organisms in "A.
za" would not be in any basal, exclusive group, because the smallest
exclusive group that they are members of is +A. rnadagascariensis
+A. za, which contains the exclusive group A. madagascariensis.
We do not know if such a series of events occurred, and thus we cannot
determine if A. za is a species or a metaspecies. Given this
uncertainty, what is the prudent course? In the absence of guidelines for
dealing with metaspecies in a taxonomy, and in order to avoid unecessarily
destabilizing the taxonomy of Adansonia, I here advocate recognition of
A. za as a species, while flagging it as a taxon worthy of more
detailed analysis.
Perrier de la Bathie (1955) proposed an infraspecific
classification of Adansonia za, recognizing three varieties. Although
there is significant variation from the north to the south of the range of
A. za, this variation is gradual and I see no value in recognizing
subspecific taxa. The southern populations of A. za are the most
distinctive because of their long- petiolulate leaflets and ovoid-cylindrical
fruit with swollen peduncles. These characters are only gradually replaced by
sessile leaflets and normal peduncles as one goes north. Some other characters
also vary clinally, such as leaflet size (larger in north) and flowering
season (earlier in north), but again I see no use in forcing a continuum of
variation into discrete taxa.
There is some nomenclatural confusion as to
the taxon to which the name Adansonia za should apply. Hochreutiner
(1908) pointed out that the description given by Baillon (1890b) contradicts
the illustration labeled "A. za" (79C) in Histoire Naturelle des
Plantes (Baillon, 1893) and the description given by Drake del Castillo
(1902). Hochreutiner concluded that the plate predated Baillon's description
and hence had priority. However, Jumelle & Perrier de la Bâthie (1910)
pointed out that although plates 79A and 79B predate Baillon (18906), the
others, 79C to 791, were not issued until 1893, and hence the written
description has priority. Of the three plates (79C, D, and I) labeled A.
za, two (79C and I) appear to be A. suarezensis, whereas plate 79D
is an accurate representation of the fruit of A. za Baill. Thus, the
original description of A. za is Baillon (1890b). The lectotype (Greve
37) was selected by Perrier de la Bâthie (1952a), but he incorrectly gave the
collection locality as Morondava.
The first mention of Adansonia
bozy is in Jumelle & Perrier de la Bâthie (19096), but the taxon was
only recognized provisionally and hence this name was not validly published
until 1910 (Jumelle & Perrier de la Bâthie, 1910). Perrier de la Bâthie
(1952a) subsumed this taxon as a variety of A. za. However, in this
publication Perrier de la Bâthie used the spelling bozo, an error which
was corrected by Perrier de la Bâthie (1955). Only one specimen of A.
za var. bozy was collected early enough to have been consulted by
Jumelle & Perrier de la Bâthie (1910), and this is, therefore, assumed to
be the holotype. In selecting a lectotype for Adansonia za var.
boinensis I chose Perrier de la Bâthie 960A, the most complete specimen
from one of the localities mentioned by Jumelle & Perrier de la Bâthie
(1909a, 1910) that predates these articles. The putative species Adansonia
alba was named by Jumelle & Perrier de la Bâthie (1909a) to
accommodate a specimen Perrier de la Bâthie collected along the Andranomalaza
River. This species was said to be diagnosed by its white flowers and short
staminal tube. In the course of my observations and collections along the
Andranomalaza River, all the baobabs I encountered could be assigned to A.
za. Furthermore, the type specimen, though of poor quality, shows no
features that distinguish it from A. za. Thus, I believe that A.
alba is synonymous with A. za. The characters that supposedly
separate the species, white petals and short stamina] tube, were probably
incorrectly scored by Perrier de la Bâthie, being based upon fallen flowers.

Ethnobotany.

Adansonia za fruits.Image courtesy:
Dr.Pavel
Hošek, CZ, who headed a
scientific expedition to Madagascar.
Go to
Lemuria
for more images.

Adansonia za has
several common names, for example, "za" or "zabe" in the south and "bozy" or
"bozybe" in the north, and "ringy" or "boringy" in the Ambongo region (Perrier
de la Bâthie, 1955). Little is known about the human exploitation of Adansonia
za, but Jumelle & Perrier de la Bâthie (1912) reported that the seeds are
eaten and the trunk is sometimes hollowed out as a cistern for storing water.
Perrier de la Bâthie (19526) mentioned that the seedling roots are an edible
vegetable, and Miege (1974) reported that A. za is destroyed by ranchers so
that their cattle can feed on the moist wood.

Conservation. In
view of the extensive geographical range, Adansonia za is
conservationally secure, despite the fact that several local populations are
endangered by forest clearance. Adansonia za has diverse interactions
with animals: it provides nesting sites for birds, holes for carnivores and
lemurs (M. Nicoll, pers. comm.); perches for territorial surveillance and
display by birds and the sportive lemur (Lepilemur sp.); many insects
feed on the leaves, sap, nectar, and pollen; sunbirds (Nectarinia
souimanga) feed on nectar; sifaka (Propithecus verrauxi verrauxi)
feed on flower buds; and fork-marked lemurs (Phaner furcifer) feed on
exuded gum (Petter et al., 1975).

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