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BLUMEA 26 403-410 (1980) A note on Heterophylly and Branching patterns in the Amyema complex Job of University (Loranthaceae) Kuijt Lethbridge, Canada Summary in Attention is drawn to the unusual distribution of flowers and inflorescences to certain far has of branching and peculiarities escaped notice, involving cataphylls. A similar The genus branching Amyema Ignoring the eight or dealing with the Danser regarded Amyema’. several of these complex the to to genera in Van Sogerianthe both the system the but (1931, 1933) as An by Barlow). been context encircling in wood) between peculiar a the stem and words, inflorescences feature, 1974; a of including which internodes Valeton ( stem to also are for more very Rubiaceae; Winkler prolific Amyema creator (1966, 1974). Tetradyas, Pa- polymorphous detailed investigations a was in the has followed not nomenclaturally for Amyema has narrow showing of more and Rhizomonanthes are even to regard. workers important and Barlow the latter genus is sense an several types which at a inflorescence survey of Loranthaceae generally. features of Amyema the axillary region of found may be a an found in those 1931). lies in the fact that, in the merely in the axils of zone the often foliage leaves. In other (often profusely, especially anatomical The like the curiosity extreme are flowers found on paid in expressions species (Dicymanthes leafy completely leaf pair. Little attention has been constitutes much not floriferous transverse (Merr.) Danser)where inflorescences themselves, the most regard reliable generic limits and develop endogenously. inflorescence proliferation and D. seriata with whole. A discussion of these types will follow an narrow, on a most infrageneric organization proposed. the members of inflorescences appear constitute the 'extreme forms of the more so intercalary Helicanthes. of Loranthaceae. Barlow's work striking morphological rather of minute Diplatia, Distrianthes, great majority of species, the inflorescences emerge organs in mostly by two diverse genus in its inflorescence structure, One of the heterophylly which World, especially need, however, achieve separate genera (Barlow later date within the number of species, and a pair a undoubtedly Old Tieghem, Dicymanthes, in Loranthaceae as unique is evident generally although Dicymanthes knowledge, my exceedingly are as the have been Danser genera overlooked apparently not, names He stressed the Danser in these views maintained early disappearance of relatives nearest a explained by nine additional genera proposed, generic placement accurate are Much differenceof opinion has existed in this latter Rhizomonanthes and puanthes, genus its and The latter flower distribution and pattern and generic circumscription. of Loranthaceous the formation Loranthaceae in complex of difficult phyllotaxy. older to that this the of such suluana Danser irregularly along of Stichianthus the minutiflorus 404 BLUMEA Plate Sogerianthe versicolor, I. between intercalary cataphylls enlarged. The 3. Fig. — of young prominence lower next flower Brass 3227, two successive zone. node Fig. — 26, L. Fig. — of pairs from 4. No. VOL. 2, 1. Young leaves. foliage 1, Fig. Emergence 1980 branch from Cataphyll of showing disappearance of four flowers of tip showing position 2. Fig. — Fig. 1, and cataphylls from the flower zone, all in approximately prophyllar positions. In number of a from epicortical endogenous inflorescences Amyema species 1 roots In these ). of flowers origin or inflorescences, number of quite unrelated angiospermic species strictly showing endogenous origin, is nature taken referred are numerous nor both genera show the species buds axillary covered the covering what also, all. (As axillary endogeny Old it is very as appears be Worlds. from internodes there is of these plants. and but Hooker as In a It needs The implication at the be can of all this find an of in flaps be cannot of it in traces of the species some to but that able was as well which the cushion hard, protective that in in fact may endogeny). in the young Amyema, be this not understand how the true of matting together a proper anatomical a show that to through is, therefore, nodes flowering anything emphasis we tissue to a conspicuous hairy covering no such for inflorescences (1846) caplike a axillary region, drying endogeny roots usually of a those In Misodendrum (Misodendraceae), , several mentioned above, however, it is difficult enough, in by irregular flaps phenomenon a general, pronounced secondarily by over in the area. to Even emergence. Such bud eventually emerges. In Cladocolea there is evidence of perhaps glandular hairs from known evidence some ruptured during and New phenomenon, same were the example, (Kuijt 1975). innovations vegetative been in Loranthaceae in rare in is doubt of the no 1969). involved may be invested This apparent for be 'chaffy scales' by Barlow (1974), but the implied leafy to as neotropical Cladocolea, as apparently is it there (Kuijt directly may emerge can phenomenon a parasites flowering structures appropriate. not face value, at nodal the which have corky tissue, Sogerianthe as flowers or latter instances there at flowering Interestingly flowering study zones is needed to elucidate these features. In the course of general evident that the overall inadequately explored; relationships to understanding complex, present studies of the inflorescence of Loranthaceaeit has become structure I of the members of the referring especially am the position of inflorescences. of this there is no doubt that it will on be an herbarium key My original attention present was focussed neither of which seem on to successive than ') at See, pairs of foliage perpendicular planes for example, A. leaves as My are been Rhizomonanthes) also the inflorescences of the above three leaves. and A. can a Danser seen and be rather where two misleading. puzzling the we features literature. The each percurrent stem, however, A. in the from on the same plants hastifolium Barlow. Flowers both when which, recorded in expected dilatipes species) are only always aligned would be curvifolium (Krause) Danser's constituting are an of the material branches of these plants frequently bifurcate. On two that pretend essential tool for this purpose. Sogerianthe, have cannot and their systematic analysis to concerned with the position of branches particularly, are has been Amyema complex branching patterns While I is the morphological aspect contribution is based to in plane (Ridl.) Danser Distrianthes epicortical rather with decussate (together (and perhaps roots and in the axils of Job Kuijt: Heterophylly and branching patterns in Amyema 405\I 406 BLUMEA phyllotaxy (Plate I). Secondly, from the (often apparently the foliage leaves, nearest alternate with the a VOL. often than more flowering zone) in their axils not if especially somewhat only one of the 2, 1980 not the innovations placed are In other innovations two with the at a node obvious in to such branches the members of pressed material; its orientation is often altered develops, internode words, between always not arising perpendicularly foliage leaves, arising nearest aligned be to No. (Plate II). As mentionedearlier, this feature is pair. 26, and below, it may be to appear percurrent. accounting for both A solution of particularly good specimen which has young Islands, L) that, an on a of these aberrant features in the process of twigs percurrent branch, successive two with presented itself The fact is elongating (Plate I). of pairs foliage leaves separated by are extremely inconspicuous, caducous pair of intercalary cataphylls which for only seen a very brief period. Each and about 0.5 dorsally, the expanded material in the flowering node before cited above there all in the It be can in does, leave these that these fact, exist but successive pairs Innovations The observed exists in several 1959), of pairs the intercalary novations, although been of the first material. I at or a few P. with the calls these and correct. cataphyll in in flowering assuming prophyllar, all above the I have flowering as not a to axils of and scarce I axillary positions but in some observed any In seem P. as Phoradendron, to subtend in- crassifolium (DC.) intercalary cataphylls. No Sogerianthe. is zone The next flowers difficult they seem to and such be to foliage appear tempting too designation a regular pattern to It is petiole (fig. 4). some petiolar base, to leaves. foliage parallel Trel. never such species single flowering the same far to observe leaves designate the side for this may therefore in the further in the certainly are to are not development of be the zone. Danser did same in mind the (H.B.K.) persistent, mature. common of caducous pair a nearest that the flowers in the axils of to a which It is of considerable interest that another the single (Kuijt elaborate immediately flanking positions not at a one piperoides are more least among the first those found along genus Phoradendron of Viscaceae immediately common in am In the collection Regular decussate phyllotaxy frequently most position neotropical cataphylls seen The sequence of pressed of the swelling leaves intercalation of the Eichl. inflorescences regularly stand in the axils of flowers have fully corky leaves find themselves in the foliage originate seem to phenomenon species especially are leaves below foliage scars. foliage keeled of the cataphylls provide intercalary of and thus alternate in cataphylls, however, discernible no successive ten leaves foliage production because of the considerable plane (on percurrent branches) through cataphylls. the be can scale, plane. therefore, seen, before the for both of the above mentioned features. explanation brownish minute, between it and the node of the cataphylls least at are same a fallen, possibly through zone the flowering, Even long (fig. 2). have is cataphyll Both because of the cork and (fig. 3). shoot, mm cataphylls a versicolor Danser (Brass 3227, Solomon Sogerianthe not consider closely branching pattern cannot but, certainly crescent-shaped a of it is to (monotypic) Sogerianthe described above. be certain that all most the members of as related branching (Plate III). Young leaf-pair. The first flower leaf scar) is in a is as or genus, Helicanthes, which Amyema, precisely described for Sogerianthe above, innovations arise in (only shows Since material of Helicanthes is a scar in positions fig 10, directly primary axillary position, but between above the subsequent ones JOB Kuijt: Sogerianthesogerensis. Plate II. Flower positions indicated pattern, branching as seen petiole, those of Sogerianthe. 2 ), one branching 5. Cruciate Fig. — black dots. Amyema patterns branching in pattern. luzonensis. — — not 6 Fig. 8. Figs. 7, Since flowers supposed, in affinity even a series of flowers Helicanthes, between the if the two geographical or again as seen from Variations been able to ascertain the scars in as genera in above. cruciate very much like Sogerianthe might not distributionof the and Eastern New Guinea and island groups to the east, distinct. I have 407\II Amyema from above. wonders if the closer than Danser (India, by and and older nodes show flank the solitary Heterophylly occurrence of , are be much two respectively) genera is quite intercalary cataphylls in Helicanthes. 2 ) The statement The inflorescence (Johri, Agrawal, probably referred and to Garg a leafy 1957, p. branch is a raceme 352) with is not a with about fifteen sessile only self-contradictory nodal whorl of flowers. flowers but arranged also in error; in the a whorl' authors 408\III BLUMEA VOL. 26, No. 2, 1980 Job Kuijt: While all pattern, numerous species of lateral branches in Australian the be of for in this Amyema species. lack cataphylls type there This is A. maidenii. As would be There quite separate. Amyema complex, the thus are two is, In those together. occur leaves. are My impression with normal, percurrent system, a decussating foliage example of 'cruciate' Similarly, cataphylls. always 409 Amyema have to appear all clearly and patterns in several predicted in such seen cushions from which inflorescences emerge axillary two ramification in the which branching the axils confluent but not seen for true Amyema representatives, species also, are not branching either we would thus expect showing not Sogerianthe this is that cruciate however, species of species branching and Heterophylly at node a distinct patterns of very of which needs systematic significance to explored. The In habit of forking the of majority which forms only The entire place. aborts we which are many this cases one decussate with the is effect, area on lateral innovations only a single and leaves, own two minute scales in placed are can field a or each of apex takes internode long. Where cushion that future flowers hairy develop, before abortion of its which sometimes foliage brownish hairs. It is from this abortion of the shoot apex. early on an two foliage leaves, in innovation, very blunt see a that means of pair is based species an be apex seen, of short, inflorescences originate. In least at axils of the of species one leaves foliage the fourinnovations is present above the species, such even on the same inflorescences; the as at luzonensis, cruciate two two foliage individual, is the quartet of branches (fig. a these Strictly speaking, A. Amyema, may join compound an apparent whorl of Also common of four foliage in this leaves under 8). any rate, the transition with normal innovations in the Thus (fig. 7). occurrence to one is not already be considered leafy great. In several members of inflorescences of this type have evolved, Distrianthes, Diplatia,and Papuanthes. Amyema species leaves lateral innovations may Amyema complex, indeed, specialized in two ones. In turn, it is possible that the umbels may have inflorescences of derivation, involving the reduction of all leafy organs on such a numerous comparable branches. ACKNOWLEDGEMENTS I should like to acknowledge working from the National Science and facilities at the Research Engineering Rijksherbarium, Leiden, Council, and financial support Canada. REFERENCES BARLOW, B. A. 1966. A revision of the Loranthaceae of Australia and New Zealand. Austral. J. Bot. 14: 421-499. 1974. A revision Bot. DANSER, of the Loranthaceae of New Guinea and the South-western Pacific. Austral. J 22: 531-621. B. H. 1931. The Loranthaceae of the Netherlands Indies. Bull. Jard. Bot. Buitenzorg, Ser. 3, 11 233-519. Plate III. innovations axillary Helicanthes (i); flower one scar elastica, Wight 35, leaf removed. (s) above a leaf — Fig. scar, L. 10. and — Fig. 9. Cruciate Flowering node two flowers (f). branching, showing showing two older two innovations young (i), an BLUMEA 410 . 1933. A new Old World 29: Sect., HOOKER, JOHRI, B. J. D. M., system for the species JOB. No. genera of Loranthaceae subfamily. 1946. Flora Antarctica. Verh. Kon. Part II. J. S. AGRAWAL, and S. GARG. 1959. A Phoradendron . 26, 1980 2, Loranthoideae, with Akad. Wetens. a nomenclator for the Amsterdam, Afd. Natuurk., 2d 1-128. Loranthaceae.-I. KUIJT, of this VOL. 1969. The Helicanthes study of elastica of Loranthaceae, pp. Morphological (Desr.) heterophylly (Loranthaceae). biology 1957. and Danser. 8: — 302. embryological studies Phytomorphology inflorescence Acta Bot. Neerl. 289 and 506 — parasitic flowering plants. University structure- in 7: 336 — in the family 354. Dendrophthora and 546. of California Press, Berkeley and Los Angeles. . 1975. The WINKLER, H. genus Cladocolea 1931. Ueber die Valeton. Planta 13: (Loranthaceae). J. Arn. Arb. eigenartigeStellungderBlUten 85—101. 56: 265 — 335. bei der Rubiacee Stichianthus minutiflorus