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Sterculiacese
Cola Schott and Endlicher

Cola Schott And Endlicher, 1832, 'Meletemata Botanica,' p. 33. Masters, 1868, in Oliver, 'Flora of Tropical Africa,' I, p. 220. K. Schumann, 1895, in Engler and Prantl, 'Die Nattirl. Pflanzenfam.,' Ill, pt. 6, p. 99; 1900, 'Sterculiacea; Africans,' in Engler. 'Monogr. Afrik. Pflanzenfam.,' V, p. 110.1

Edwardia Rafinesque, 1814, Spechio delle Seienze, Palermo, I, p. 158 (not Edu.ardsia Salisbury, 1808).

Lunanea A. De Candolle, 1825, 'Prodromus Regn. Yeget.,' II, p. 92.

Trees, shrubs or bushes. Leaves entire or lobed, often polymorph, rarely digitate; glabrous, hairy or scaly. Flowers in axillary panicles or clusters, sometimes out of the old wood. Flowers through abortion unisexual or polygamous. Calyx cup-shaped or campanulate, 4- or 5-cleft. Petals absent. Staminal column sometimes very short, bearing at the top a ring of 10 to 12 anthers, disposed in one or two, regular rows; anther-cells (thecte) parallel or superposed. Ovary 3- to 10-celled, with as many styles as cells. Ovules numerous in each cell. Fruit of 4 or 5 leathery or woody oblong carpels, ultimately splitting lengthwise. Seeds numerous, obovoid, exalbuminous; cotyledons thick, sometimes deeply bifid; radicle next to the hilum. (After K. Schumann.)

This large genus is restricted to the continental part of the Ethiopian Region. About one hundred species have been described, most of which grow in that portion of Africa defined by Engler as the "Western Forest Province" and twenty-five of them occur in the Belgian Congo. A few very closely allied forms possess at the base of the leaf-blade a pair of small pouches which are occasionally inhabited by ants. In addition, these myrmecophilous species differ from their relatives in having branches and leaves covered with numerous long, stiff, erect hairs of a brown or brownish red color; the other members of the genus being glabrous.

Cola Deweyrei De Wildeman and Durand Cola Dewevrei E. De Wildeman And Th. Dttrand, 1899, Bull. Soc. Bot. Belgique, XXXVIII, 2, I>. 184; 1901, 'Reliquiae Dewevreana?,' p. 24. E. De Wildeman, 1907, 'Mission Emile Laurent,' fasc. 4, p. 406, Pi. cxxvu. Th. Andh. Durand, 1909, 'Sylloge Flor. Congol.,' p. 62. E. De Wildeman, 1920, 'Mission de Briey au Mayumbe,' p. 191.

"A low shrub, 1 to 2 m. high. Branches hollow, terete, with long pilosity. Leaves trilobate, deeply cordate at the base, abruptly acuminate at the apex, shiny on upper and under sides, pilose, especially on the veins. Secondary veins arenately anastomosing toward the margin and uniting with the reticulate finer venation, a little prominent above, more strongly so below. Petiole very long, more or less grooved above, with long pilosity, 6 to 32 cm. long. Leaves 13 to 25 cm. long and 15 to 24 cm. broad. Stipules linear-lanceolate, dropping, pilose, about 6 to 11 mm. long and 1 to 1.5 mm. broad. Flowers yellow, fasciculate, axillary, subsessile, bracteate. Calyx

According to the rules of botanical nomenclature the name Cola should be replaced, it seems, either by Edwardia or by Lun%nea, since the latter two are not preoccupied and evidently have priority.

5-lobed, ferruginous tomentose externally, brown and less pilose inside, about 11 mm. long; its lobes 2 to 3 mm. long, acute, with more or less refiexed tips. Androecium of the male flowers stipitate, smaller than the calyx, with subglabrous stipe, 4 mm. long; the anthers placed close together, parallel and united into a ring which is about 15 mm. high" (De Wildeman and Durand, 1899).

Belgian Congo: Mayombe: Lemba River (Dewevre).

It would seem from the descriptions that the later C. Laurentii De Wildeman and the earlier C. tnarsupium K. Schumann are not specifically distinct from C. Dewevrei. According to De Wildeman (1907, p. 406), the leaves of C. Dewevrei have a different shape from those of C. Laurentii, with basal lobes almost touching each other. In these Colse, however, the form of the leaves varies to such an extent even on the same plant that this character is by itself unsatisfactory for the distinction of the species. The existence of foliar pouches is not mentioned in the original description of C. Dewevrei, but De Wildeman's figures of that species published in 1907 show them distinctly.

Cola Laurentii De Wildeman

Cola Laurentii E. De Wildeman, 1907, 'Mission Emile Laurent,' fasc. 4, p. 403, fig. 68, Pis. cxxxv, cxxxvi, and cxxxvn; 1908, 'Etudes Flore Bas- et MoyenCongo,' II, 3, p. 304. Th. And H. Dui1and, 1909, 'Sylloge Flor. Congol.,' p. 63. H. Kohl, 1909, Natur u. Offenbarung, LV, p. 148. A. Engler, 1912, 'Wiss. Ergebn. Deutsch. Zentr. Afr. Exp. (1907-08),' II, p. 506. E. De Wildeman, 1919, Bull. Jard. Bot. Bruxelles, V, p. 358.

"A small tree with cylindric branches densely villose; with brownish, elongate, spreading hairs which drop late. Leaves with more or less lengthened petioles, which are cylindrical, hispid with spreading hairs, 5 to 35 cm. long. Leaf-blade 3-lobed or nearly 5-lobed, cordate at the base; the midlobe about two thirds the length of the leaf, which varies from 11 to 36 cm.; the midlobe is oblong, rather suddenly acuminate at the tip, acute; lateral lobes about of the same shape, a little shorter and narrower than the terminal lobe, which reaches a length of 23 cm. and a width of 13.5 cm. Leafblade paler on the under than the upper side or about the same color, with 7 basal veins, the lateral ones often united at the base. Leaf-blade coriaceous, glabrous, except on the veins of both sides, especially on those of the under side which are very prominent and bristling with stiff hairs. Between the midrib and the first lateral vein on each side of it there is a small pouch strongly projecting on the upper side; the two veins between which this pouch is formed are united at the base by a plate of tissue. Stipules filiform, hispid, rather dropping, about 2 cm. long, acute. Flowers fasciculate at the axils of the leaves; the rachis about 1 cm. long; the bracts linear, acuminate, hispid, about 2 mm. long; the pedicels villose, a little over 1 mm. long. Calyx campanulate, about -S mm. long; with 4 to 5 lobes one-third the length of the tube; calyx densely villose, brownish on the outer side, with more scattered hairs internally. Male flowers with an uniseriate androecium, composed of thecae a little over 2 mm. long, borne on a slender, feebly elevated androgynophore which is 3 to 4 mm. long and shorter than the calyx-tube. Female flower with a densely villose, ovoid ovary; the style shorter than the ovary, with spreading stigmata which are as long as the calyx-tube. Staminodes reduced, surrounding the base of the ovary. Fruits red, 5 to 6 cm. long including the acumen, with 4 to 5 seeds" (De Wildeman. 1907).

Belgian Congo: Lower Congo: Sabuka (M. Laurent): between Boma and Yanga (R. Verschueren). Kasai: Dibele; Kondue (Em. and M. Laurent); forest of the Sankuru (Luja). Upper Congo: Eala; Vakusu (Em. and M. Laurent); Yambinga (M. Laurent); Dundusana (F. Reygaert); Barumbu (Noveml>er 3, 1913; J. Bequaert; Coll. No. 1081). Eastern Congo Forest: Yambuya (M. Laurent); Basoko (Em. and M. Laurent); Fariala between Mawambi and Avakubi (f. inlegrifolia; Mildbraed); between Lubutu and Kirundu (village of Uchibango, February 1, 1915; .1. Bequaert; Coll. No. 6790); Stanleyville (March 1915; H. Lang).

De Wildeman classified as ''form intermedia" plants of this species in which entire and trilobed leaves are found on the same branch together with all intermediate shapes; his "form integrifoh'a" includes specimens in which all the leaves are entire, ovate-cordate at the base and as much as 35 cm. long and 18 cm. broad; in this last form pouches are also feebly developed along the midrib in the axils of the first or first and second lateral veins, above the large basal pouches.

In recording the form integrifolia, Engler (1912, p. 506) also mentions that in his opinion C. Laurentii is not specifically distinct from C. marsupium.

Cola marsupium K. Schumann

Cola marsupium K. Schumann, 1891, Ber. Deutsch. Bot. Ges., IX, pp. 68-70; 1900. 'Sterculiacea' Africans',' in Engler, 'Monogr. Afrik. Pflanzenfam.,' Y, p. 113, Pi. XII, figs. A-D. H. Kohl, 1909, Natur u. Offenbarung, LV, p. 148. "A shrub or tree, with slender, terete branches, the younger ones flattened and strongly hispid, later glabrescent. Leaves with long, terete, hispid petioles; oblong or obovate-oblong, shortly and very sharply acuminate, cordate at the base; with 7 or even 9 veins; provided with a pair of pouches forming basal swollen domatia between the midrib and the lateral veins; covered with rather long hairs on the veins on both sides, rather rigidly herbaceous. Stipules filiform, hispid, persistent for a long time. Flowers short pedicellate, axillary, fasciculate, placed either at the extremity of branches which are rather sparsely leaved below or on leafless branches. Bracts and bracteoles linear, acuminate, hispid. Calyx campanulate; its upper third split into

4 or 5 ovate, acute lobes; tomentose outside, papillate inside. Male flower: androecium uniseriate of 16 to 20 theca>, raised on a gracile, glabrous column. Female flower: ovary subglobose, pentamerous,tomentose; the style glabrous, straight, 5-lobed;

5 ovules in each cell; follicles short stipitate, fusiform.

"The shrub reaches a height of 1 to 2.5 m.; the tree as much as 10 m. The foliate, flowering branches are 3 to 3.5 mm. thick at the base and 20 to 25 cm. long; they are rough, being covered with simple, spreading, brown red or brown hairs, which are thickened into a tubercle at the base. The petiole is 1.5 to 15 cm. long and covered with the same pile. The blade has a length of (i to 30 cm. and a width of 3 to 13 cm. above the middle; in addition to the basal veins, it is crossed on each side of the midrib by only 5 to 6 stronger veins, which are a little more prominent on the under side, as is also the reticulate venation; sometimes the blade is somewhat gibbous; in life it is dark green, brownish green when dry. The basal pouches can be entered from the under side; they are not always present, but usually found on the larger leaves. The stipules are 1 to 1.5 cm. long and covered with brown hair. The bracts of the flowers are usually somewhat broader than the stipules, but otherwise similar. The yellowish green calyx is 5 to 7 mm. long. Male flower: andrcecium 1.5 mm. long, as well as the androgynophore. Female flower: calyx slightly larger; ovary 6.5 mm. l»ng, surrounded at the base by a ring of staminodes 2 mm. high. The fruit is red, but perhaps not entirely ripe" (K. Schumann, 1900).

Cameroon: Abo (Buehholz); Johann-Albrechtshohe: in the Senge Mountains (Staudt).

French Congo: Maveli Mountains near the Sibange Farm (Dinklage). It seems probable that the three forms described above, C. Dewevrei, C. Laurentii, and C. marsupium, all belong to one species, for which the name C. marsupium K. Schumann should be retained. This is, however, a question to be decided by botanists and, in order to avoid any possible confusion, I have here used the name C. Laurentii for the plants observed by me in the Belgian Congo, because the description of that species fits them most nearly.

Ecology Of Cola Laurentii

This plant is rather common in the Congo Basin, where it prefers the drier, more elevated parts of the primitive Rain Forest. It usually grows as a shrub of moderate size (1 to 2.5 m. high), more rarely as a small tree (as much as 10 m. high) and flowers in both forms. The leaves are, as mentioned above, of variable size and shape, usually elongateoval, with cordate base; the margin may be entire, or slightly or deeply lobate. The pair of basal, elongate-oval pouches on the leaves are more or less developed; wholly absent in certain cases, in others they may attain 15 mm. in length and 5 mm. in width; on the average they are 4.5 to 9 mm. long, 1.5 to 4 mm. broad and 6 mm. high. Placed at the base of the blade close to the midrib, they project on the upper side of the leaf and on the under side have a narrow slit their entire length.

The general aspect of C. Laurentii is illustrated on Plate XXVII, Figure 2, by a photograph of a branch, with flowers and fruit, made by Mr. H. Lang at Stanleyville, while the shape of the myrmecodomatia is seen in Text Figure 84. As mentioned by £m. Laurent (De Wildeman, 1907, p. 405), the pouches are only occasionally occupied by ants. They were empty on most of the many plants which I examined. On one occasion, near the village of Uchibango, between Lubutu and Kirundu (February 1915), ants belonging to the dolichoderine Engramma kohli Forel were found inside the pouches; they had closed the slit at the under side with vegetable detritus. Unidentified ants were also found in such swellings at Barumbu (November 1913). Some of the plants collected by Mr. H. Lang along the Tshopo River near Stanleyville, in March 1915, were inhabited by Plagiolepis mediorufa (Forel), an ant originally described from specimens taken by Father Kohl in a nearby locality from an unidentified myrmecophilous plant. Engramma kohli,

[graphic]

Fig. 84. Cola Laurentii De Wildeman. Lower part of a leaf seen from above, with the two pouches at ba9e of blade. Drawn from life at Barumbu, November 1913. About natural size.

like certain other members of the genus, is a frequent inhabitant of various myrmecophytes. Both Engramma and Plagiolepis are so timid and small that they could not well act as body-guards to their host plant .

Although K. Schumann (1891, pp. 68-70) describes the ascidia of Cola marsupium very fully and regards them as myrmecodomatia, he was unable to find ants on his herbarium specimens from Gaboon and Cameroon. H. Kohl (1909, p. 148) is inclined to believe that, on account of the small size, these pouches are not adapted to the use of ants but serve better as shelters for coccids or plant lice.

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