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Djvu Document BULL. BOT. .SURV. INDIA Vol. 7, Nos. 1-4, pp. 73-88, 1965 THE TRANSFER OF CYRTANDROMOEA FROM GESNERIACEAE TO SCROPHULARIACEAE, WITH NOTES ON THE CLASSIFICATIONOF THAT FAMILY B. L. BURTT Royal Botanic Garden, Edinburgh 1. THE STRUCTURE AND AFFINITIES OF the Matang plant. The seedline raised had cotyledons CYRTANDROMOEA which were equal in size on germination and remained so throughout. Thus the doubts already felt about the In a recent attempt to classify and give keys to position of Cyrtandromoea were immediately confirmed, the Old World genera of Gesneriaceae, Cyrtandromoea .for no member of Cyrtandroideae is known to have was ten~~tivelyassigned a position in Cyrtandroideae- isocotylous seedlings (cf. Burtt, 1963, 205). Loxonieae, "for want of a better place". At the same The plants of Cyrtandromoea raised from Sarawak time attention was drawn to its resemblances to Brookea seed produced flowers in the late autumn of 1963 and Benth. and Rehmannia Libosch., two genera whose from one of them serial sections of the ovary were exclusion from Gesneriaceae was thdught justified. A prepared. The illustration (fig. 1) shows a selection thorough re-investigation of Cyrtandromoea was of somewhat diagrammatic cross-sections at different advocated (Burtt, 1963, 210, 217). levels of the ovary ; it had been taken from a flower The opportunity, one might say incentive, for, such that had been hand-pollinated but had fallen without a study has arisen from a visit paid to Malaya and fruit development. Pollen tubes are not visible in any Sarawak in the summer of 1962-after the above-quoted of the sections. The ovary is clearly bilocular through- paper was written. Cyrtandromoea grandis was found out its length ; the placentae are lamellar in form and in the Ulu Gombak near Kuala Lumpur, and another ovuliferous over the whole surface, species (possibly C. subsessilis) on Mt. Matang, near Examination of young fruits of Cyrtandromoea Kuching, Sarawak. gmndis, preserved in spirit, showed that the fruit wall is The genus has been introduced $to cultivation at thin while the placentae are fleshy (Plate I, H). These Edinburgh from a few seeds found in a broken fruit on characters' recall such genera as Fieldia (Gesnerioideae- Fig. 1 Cyrtmdromoea subsessilis (Miq.) B. L. Burtt ? (B. 1944) ; series of trans- verse sections of ovary from apex (1) to base (12) X 59. 74 BULLETIN OF THE BOTANICAL SURVEY OF INDIA [VO~.7 Mitrarieae), though there the' ovary is unilocular. The seed structure of Cyrtandromoea also needed Although the possession of a unilocular ovary and elucidation. Endosperm is definitely present and the parietal placentae is usually given as a diagnostic feature inner and side walls of the testa have laminate thicken- of Gesneriaceae, a bilocular ovary does not absolutely ings (Plate I, J) : the outer wall remains thin and exclude a genus from thii family-it is, in fact, quite easily becomes broken, so that the testa appears to have remarkable how, many illustrations of Gesneriaceae a reticulate pitting. show, without comment, a bilocular ovary. Before considering .possible relationships between Plate I Cyrtandromoea grandis Ridl. : A, basal portion of stem X 3 ; B, apical portion of atem X 4 ; C, corolla, dissected X 9 ;.D, gynoecium X 4 ; E, anthem, posterior and anterior views X3 4 ; F, pollen grain X 650 ; G, ovary in longitudinal &on XI* ; H, young fruit in transverse section X 3 ; 1, seed X 63 ; J, seed in section X 63. Cyrtandromoea subsessilis ? K, seedling X 4. (A-J from BUM & Woods, B 1616; K from Burtt & Woods, B 1944). 19651 BURTT : TRANSFER OF CYRTANDROMOEA FROM QESNERIACEAE TO SCROPHULARUCEAE 75 Cyrtandromoea and groups outside Gesneriaceae it dorsal corolla lobes are external in aestivation and the seemed desirable to find out whether there was an leaves opposite), and the convex lower lips of the affinity with any genera of the subfamily Gesnerioideae, bilabiate spurless corolla and the cymose inflorescence which has constantly isocotylous seedlings and seeds lead directly to the tribe Cheloneae. with endosperm. Ridley has inadvertently provided a Within Cheloneae Wettstein's key asks first oi all pointer in this direction, for the plant he described from whether the fruit is a capsule or berry and then, if a Sarawak as Cyrtandromoea minor proves to be the capsule, whether it is septicidal or loculicidal. This is Americag Chysothemis pulchella (Don) Decne., somewhat less than satisfactory ; for it separates which must have been cultivated in Kwhing. The Paulownia from Wightia, Uroskinnera from Brookea, resemblances between these two genera lie in the general Hemichaena from Leucocarpus. Cyrtandromoea can leaf shape, the tubular winged shortly toothed calyx have a more or less indehiscent thin walled berry or a and the external form of the gynoecium. The differ- weakly loculicidal capsule. If the first it is to be placed ences are manifold : Chrysothemis has bent filaments next to Leucocarpus D. Don, if the second next to and anthers cohering by their tips, a large unilateral Hemichaena Benth. Both genera are Central American nectary, bifid parietal placentae and spirally striate and monotypic, and both have been figured in Curtis's seeds with a funicular food-body. Even the resemblance Botanical Magazine : Leucocarpus in 1831 (t. 3067 as in the calyx is superficial, for the wings or ridges on Mimulus) and Hemichaena in 1875 (t. 6164). The the calyx of Chrysothemis run to the sinuses whereas best account of Leucocarpus is, however, David Don's, in Cyrtandromoea they are median to the calyx-teeth. published by Sweet (D. Don, 1831). If Cyrtandromoea belongs to Gesnerioideae, it The fruit of Leucocarpus is a white berry, but the should be in the affinity of Besleria with which it has skin is rather thin and most of the substance of the the annular disc in common. Now Besleria is a large fruit is due to the fleshy placenta. It is interesting and varied genus of some 140 species, of which very that Don made comparison with the fruit of Fieldia*. few are in cultivation, and of which even the herbarium The calyx of Leucocarpus is tubular and shortly toothed, material available to me is rather scanty. Fortunately but is not so long as in Cyrtandromoea and is not there is a fine revision of the genus by C. V. Morton accrescent after flowering, so that the fruit is not in- (1939). It appears that constant generic features of cluded within it. The ovary, I and especially the Besleria are that the anther-thecae are confluent at placentation (cf. Hartl, 1956: 204, Abb. 12), the style- their tips, that the two parietal placentae are ovuli- and bilamellate stigma are essentially the same in both ferous only on their inner faces, that the stigma is genera. In the stamens of Leucocarpus (Plate 11, E) bilobed or stomatomorphic and that the fleshy berry the anther thecae are widely divergent, and confluent has a thick verrucose exocarp and spirally striate seeds. on dehiscence, whereas they are subparallel in Cyrtan- All these features are in sharp disagreement with the dromoea and only slightly divergent at the base. The characters of Cyrtandromoea and the two genera are exserted fruit and the anthers provided the best dis- clearly not closely related. tinction between Leucocarpus and Cyrtandromoea. The only possible affinity remaining within Ges- The seeds of Leucocarpus are similar to those of nerioideae is with the tribe Mitrarieae. Here we find Cyrtandromoea in general appearance and have about a resemblance in fruit characters, for this group also the same quantity of endosperm. In sections, however, has a thin fruit wall and fleshy placenta. However they do not show the' distinctive laminated thickening the ovary of Mitrarieae is unilocular and the placentae of the inner and side walls characteristic of Cyrtandro- are parietal and bifid. In flowers, leaves and habit moea. It has not been possible to make wide enough there is little resemblance and Mitrarieae are an austral comparative studies to assess the value of this charac- pup, in contrast to Cyrtandromoea which is tropical. ter : in this affinity Schultes ( 1941) refers to. the seeds It is concluded that Cyrtandromoea has no close of Uroskinnera being enclosed in a thin membrane- afhity with other members of Gesneriaceae. * In reading this article on Leucocarpus I learned for bilafular OvV and other of Cyrtan- the fint be Don hd wi&bwn his p~viou~~ypm- &ornoen to saggat that an affinity is most likely be fa~lyMdymoearpOCCoc (bn, sa9ng found for it in the large polymorphic family Scrophu- accumte examination has fully convinced me that they must lariaceae ; in fact there is a notable superficial resem- be united to the Gcsnsriaceae, which again are hardly dis- blance to same cultivated species of ~j~~l~~.using tinguishable by MY tangible character from Scrophularinae." Thin implies, that Don recognized the family Gesncriaceat Wettstein's (1895) cla~ificationCyrtandromoea should in 182& ia accopkd publiertion only fmm 1821 be placed in subfamily Antirrhinoideae (because the (c., ~~x~~,viii, 173 : 1959). BULLETIN OF THE BOTANICAL SURVEY OF INDIA Plate I1 Leucocarpus alatus D. Don : A, habit X 3 ; B, calyx X ; C, corolla, dissected X 3 ; D, gynoecium X 8; E, anthers, posterior and anterior views X 2d ; F, pollen grain X 650 ; G, fruit in longitudinal sectiqn X % ; H, fruit in transverse section X 4 ; I, seed X 40. (A, B, C, G & H from British Flower Garden. ser. 2(2) : t. 124, 1831). The map indicates the general arsas of Cyrtandromoea (cross-hatched) and Leucocarpus (qtraight lines). In the inset X marks districts where Cyrtandromoea is known to occur, not individual records. this may well be the thin outer walls of the cells of lobes and full yellow flower, red-spotted in the throat the testa.
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