1957) (.Botanical Museum, Utrecht
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Acta Botanica Neerlandica 6 (1957) 351—377 On the Position of Platycarpum Humb. et Bonpl., Henriquezia Spruce ex Bth. and Gleasonia Standl. C.E.B. Bremekamp (.Botanical Museum, Utrecht) (;received February 15th, 1957) Introduction Some time the “Institute do ago Agronomico Norte, Belem, Para, Brazil” sent me a set of specimens for identification among which I detected Bth. This a new species of Henriquezia Spruce ex discovery induced me to study once more and now in somewhat more detail the relation between this and the related genus nearly Platycarpum Humb. and the these with et Bonpl. position two genera occupy the rather regard to habitually similar Gleasonia Standi., a subject I to which had already paid some attention at an earlier occasion, and I be the on which had reported in a note which is to found at base of 16 of work “the African of Oldenlandia L p. my on Species sensu Hiern et K. Schumann” in Verh. Kon. Ned. Akad. v. Wetensch., Sect. 2, 48, no 4, 1952. By the good office of Dr Bassett Maguire I received on loan from the New York Botanical Garden the rich materials by which, as a result of the Botanical Garden’s recent these in its expeditions to Tropical America, genera are represented herbarium, and during a recent visit to England I could study also the valuable collections in the herbarium of the Royal Botanic Gardens, Kew. In the course of these studies I discovered among the Platycarpum specimens too a new species. Descriptions of the latter and of will be in my new Henriquezia found further on this paper. but of the main They are, however, secondary importance; object of this study is to ascertain as precisely as possible the taxonomical of these three rather position puzzling genera. In the note on p. 16 of my work on “the African Species of Olden- landia L sensu Hiern et K. Schumann” I stated that Gleasonia Standi, “on account of the large and flat exalbuminous seeds and also because of the of very peculiar structure the testa is to be referred to the have Henriquezieae, a tribe that will to be removed as a separate family the In of this to Tubiflorae.” support opinion I referred to Fig. c, d, and f e of Tab. V, which represent the testa of Henriquezia verticillata Spruce ex Bth., of Platycarpum orinocense Humb. et Bonpl. and of two species of Gleasonia, viz. Gl. uaupensis Ducke and Gl. macrocalyx Ducke. The study of the more abundant and better material that I had has led now at my disposition to a somewhat different standpoint. The between and the other differences Gleasonia two genera appeared be taxonomical to of far greater importance than the points of resem- 351 352 C. E. B. BREMEKAMP and shake the blance, although this more detailed study did not I had arrived with the opinion at regard to genera Platycarpum and viz. that be Henriquezia, they are to regarded as representatives of a new family Henriqueziaceae, for which on account of its affinity with the Bignoniaceae, Pedaliaceae, Martyniaceae, Thunbergiaceae, Mendonciaceae and Acanthaceae, a position will have to be found in the order Tubiflorae, I that the Gleasonia be now see genus can not regarded as a very near ally, and that it is either to be left in the Rubiaceae, where it occupies, the seeds and the on account of exalbuminous large size of the coty- ledons, an anomalous position, or else to be referred to a new family, which however, on account of the structure of flower and fruit and of the presence of colleters on the inside of the stipules, would have be in the This to placed the near vicinity of Rubiaceae. implies, of course, that the points of resemblance between Gleasonia and the and which I based former genera Platycarpum Henriquezia, on my I view, are no longer accepted by me as homologies, but that now regard them as mere analogies. The with taxonomical change in my opinion regard to the position in of Gleasonia is due to a better insight the characters in which this differsfrom the other and in in genus two ones those which it resembles them. In order facilitate I of all to a discussion, will first give an of enumeration the differences, as the latter are in this respect of more importance than the points of resemblance. Comparison between Gleasonia and the genera Platycarpum and Henriquezia. The principal differences between Gleasonia and the two other the absence in Gleosonia of the that genera are: 1) peculiar “glands” in Platycarpum and Henriquezia are found at the base of the petiole, and which in fact are found nowhere else in the whole kingdom of plants; 2) the presence of colleters or resin glands on the inside of the of character which this shares with the stipules Gleasonia, a genus great majority of the Rubiaceae, and their absence in Platycarpum and of the that of Gleasonia Henriquezia; 3) a different structure corolla, being actinomorphous with induplicate aestivation of the lobes, that of other with streak the two genera zygomorphous a longitudinal of hairs on the ventral side of the throat and with ascending aesti- which vation (cf. Fig. 1); 4) a different structure of the androecium, Fig. 1. Floral diagrams of A. Gleasonia duidana, B. Henriquezia verticillata and C. Platycarpum orinocense. ON PLATYCARPUM, HENRIQUEZIA AND GLEASONIA 353 in Gleasonia is subregular, in Platycarpum and Henriquezia distinctly zygomorphous, the filaments being inserted at unequal height, bent at the base and of unequal length; 5) a different arrangement of the ovules those of Gleasonia in each cell (cf. Fig. 2), forming ovary a doublerow which ascends to about midway the dissepiment, whereas the two to four ovules found in each of the cells of the ovary two other inserted the genera are at nearly same height; and 6) a different of the that of Gleasonia development capsule, (Fig. 9, p. 000) remaining whereas that of the other becomes either entirely inferior, two genera 11, or even almost semi-superior ( Henriquezia , Fig. p. 000) completely 10, Less of superior ( Platycarpum , Fig. p. 000). important points those difference are found in the arrangement of the pollen grains, of Gleasonia (Fig. 3) remaining united in tetrads, whereas they are free in the set two other genera, and also in their structure, those of Gleasonia those of the always being 3-colpate, other genera (Fig. 4) either 3- or 4- or 5- or 6-colpate, in the number and shape ofthe seeds, those of Gleasonia being more numerous and more or less swollen and whereas the four seeds of the other angular, two to two genera are flat, and in the structure of the testa, the cells being convex in Gleasonia, and produced into papillae in Henriquezia (Fig. 5) and Platycarpum (Fig. 6). Before proceeding to the points of resemblance, it will be well to discuss these differences, the more important ones as well as the others, in some detail. The which in and “glands” Platycarpum Henriquezia are seen at the base of the petiole on the side facing outwards, are large glabrous and shiny spots slightly protruding beyond the surrounding tissue; in black herbarium material they at once draw the attention by the colour they assume in drying. Their exact nature is unknown: it is possible that they secrete a sugary fluid, and in that case they would to the class of extra-floral but it also be belong large nectaries, may that they are bacterial nodules; without living material it will pro- be difficult settle this Howsoever this these bably to point. may be, petiolar “glands” certainly are a very characteristic feature of these like of which found nowhere else. two genera, the is The of colleters resin the inside of the presence or glands on stipules with the of Gleasonia is a character which this genus shares majority of the Rubiaceae. In shown Krause fact, they are, as was by (Ueber harzsecernierende Driisen an den Nebenblattern von Rubiaceen, in Ber. d. d. bot. Gesellsch. 27: 446-452. 1909), found in all tribes of this family with the exception of the Rubieae (Galieae). Their absence in of the the in this tribe might be an argument favour view that leaf-like of these modified supernumerary parts plants are no stipules, is as now commonly accepted, but that they are true leaves. Krause colleters would be in the According to present genus but mistake. I did Henriquezia too, this must be a Although not in- nitida vestigate the species quoted by him, the apparently very rare H. I H. Spruce ex Bth., studied three other species, viz. verticillata Spruce ex Bth., H. macrophylla Ducke and H. longisepala Brem. v. infra, and in but I addition several species ofPlatycarpum, in none of them could 354 C. E. B. BREMEKAMP therefore find the slightest trace of these structures. I suppose that the specimen studied by Krause will have been misidentified, and in other that it belonged reality to some genus. It is noteworthy that although colleters are by no means confined to the Rubiaceae (cf. Hanstein in Bot. Ztg 26: 697-713, 720-735, 744-761 and 768-787. in this of rather 1868), they are family a peculiar be structure which returns only in the Cunoniaceae, a family that may related to the Rubiaceae and in the Viola. That the distantly , genus is colleters of Gleasonia too are of this type, certainly a weighty argu- in favour of the view that this either the Rubia- ment genus belongs to else for which will have ceae themselves or to a new family a place to be found in the near vicinity of the latter.