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(Loranthaceae). 293 BLUMEA 38 (1993) 65-126 Conspectus of the genera Amylotheca, Cyne, Decaisnina, Lampas, Lepeostegeres, and Loxanthera (Loranthaceae) Bryan+A. Barlow Australian National Herbarium,CSIRO Division of Plant Industry, P.O. Box 1600, Canberra, ACT 2601, Australia Summary The of Australasian/Malesian group genera Amylotheca, Cyne, Decaisnina,Lampas, Lepeostegeres and Loxanthera is reviewed. Particular attention is given to the species of the Malesian region, as a ofLoranthaceae for Flora Malesiana. The of floristic ele- precursor to a treatment genera are part a ment of Gondwanan derivation which has diversified in the southeast Asian and western Malesian region, and has reached the Papuasian/Australasian region from the west following the establish- of Charles’s Line. The six Decaisnina ment genera comprise 46 species (Amylotheca 4, Cyne 6, 25, Loxanthera with Lampas 1, Lepeostegeres9, 1). Along the Australian genus Lysiana (8 species) natural with tribal of richness in they form a group or subtribal status. Major centres species are Borneo, the Philippines, and New Guinea. Revised and necessitate number of nomenclatural genus species circumscriptions a changes to earlier regional treatments. Three species are described as new, viz., Cyne baetorta, C. monotrias and C. Decaisnina longipes. There are 15 new combinations,viz., Cyne papuana, perfoliata, Decaisnina D. aherniana,D. amplexicaulis, D. angustata, D. celebica, D. confertiflora, crassilimba, D. cumin- gii, D. miniata, D. ovatifolia, D. revoluta, D. sumbawensis, D. viridis, and D. zollingeri. Introduction One in this is of objective preparing conspectus to provide a precursor to a treatment the family Loranthaceae forFlora Malesiana. In the Malesian region the family com- prises 21-23 genera and more than 200 species. For a family of this size, publica- tion of a comprehensive revision in close conjunction with a flora treatment would involve considerable duplication of information, and inflationof publication costs. of In any case, the work of a previous student the family, B.H. Danser, provides a good basis for revisionary study (see below). For these reasons, it was concluded that publication of material precursive to a flora treatment could conveniently take the of form of a series of updating accounts of selected genera or groups genera. They will deal in suitable detail with those topics relevant to a critical study of the family, but concise flora and not appropriate to a treatment, including biogeography puta- tive phylogeny, nomenclaturalissues and the rationale for the taxonomic decisions made. Critical studies of the Loranthaceae relevant to the Malesian region were made by Danser, the major works being revisions for the Netherlands Indies (Danser, 1931) 66 BLUMEA Vol. 38, No. 1, 1993 and the Philippines (Danser, 1935) in widely circulated journals. Revisions and flora treatments by the present author (Barlow, 1966, 1974, 1984) are also accessible. In in these this conspectus the very extensive synonymy provided earlier revisions has not been repeated where it remains unchanged, and wherever possible reference has been made to previous species descriptions. In this paper the six generaAmylotheca, Cyne, Decaisnina, Lampas, Lepeostege- res, and Loxantheraare dealt with, and are collectively referred to as the 'decaisninoid' the of and Decaisnina, of the Australianand group. In cases Amylotheca treatments New Guinean/Pacific species have been published previously (Barlow, 1966, 1974, for of is 1983), but now require updating, and each a conspectus the entire genus therefore presented. This allows a synthesis for each genus of its biogeographic his- and of and and host differen- tory patterns speciation, dispersal ecological preference tiation. However, for those species which are taxonomically unchanged and for which there is no significant additional information, only brief summaries are pro- vided, withreference to previous published work. in lists of Specimen label datahave been accumulated computer files, and speci- will be where mens examined only published they represent significant new findings or newly recognized taxa. Extracts from these files can be provided on request, and National reference copies relevant to this paper have been lodged at the Australian Herbarium. the contributionin this series For additionalintroductory notes, see first (Barlow, 1991b). Species circumscription The information sources for the study of the Malesian taxa have been herbarium specimens and earlier taxonomic concepts, notably those of Danser. Conclusions about of entities in different islands have also taken of conspecificity account geo- and with Danser's taxonomic physical palaeoclimatic history, concepts being re- viewed against this background. For a fulleraccount of the systematic methodology employed, see the conspectus of Amyema in this series (Barlow, 1992). AMYLOTHECA, CYNE, DECAISNINA, LAMPAS, LEPEOSTEGERES, LOXANTHERA The six treated here natural number genera comprise an apparently group, sharing a of morphological features ofprimary taxonomic importance, and having comparable habitatrequirements and geographic distributions. Although not circumscribed in the same way by previous authors, they have usually been kept together in infrafamilial classifications of the family. Infrafamilial classification of Loranthaceaeis based main- and but is in need of revision. As the ly on ovary seedling characters, seriously con- stituents of the informalgroup L 21-26 'Amylothecae' (Barlow et al., 1989), the ge- neric groupis characterized by a partially differentiatedovary with a placental column bearing rudimentary ovules (Maheshwari et al., 1957; Bhatnagar & Johri, 1983), co- tyledons emerging on germination, and relatively large chromosomes (x = 12). Most of the readily observable external characters which together are diagnostic for the group are derived from inflorescence and flower structure. B. A. Barlow: Amylotheca, Cyne, Decaisnina,Lampas, Lepeostegeres, Loxanthera 67 Diagnostic features is di- The basic inflorescence unit(uniflorescence) in the decaisninoids a simple bract each flower. The triads chasium (triad) with a simple subtending are aggregated into conflorescences, and the plesiomorphic state for the group is an axillary raceme of uniformly spaced opposite pairs of triads. Dichasial uniflorescences in a racemose Notanthera and arrangement are characteristic of genera such as Gaiadendron, Nuyt- for because sia, which are logical choices within the family outgroup comparisons of their plesiomorphic characters which show links with other families of Santalales (Barlow & Wiens, 1973). indicate that inflorescences The same outgroup comparisons capitate represent a derived state. The condensation of the inflorescence into a capitulum is apparently an adaptive trend which has also occurred convergently in other loranth groups in the 1966, 1974, and Macrosolen region, such as the amyemoid genera (Barlow, 1990) and its allies. Along with the crowding ofmore or less sessile flowers in the inflores- of floral bracts into in- cence axis, there is a trend towards the development outer an sometimes fruit volucrumwhich encloses the flowers during development, even until involucre from tissue. In ripening. In one genus, Cyne, an is derived stem periderm all such derivedcapitate inflorescences, the basic triad uniflorescencein a racemose The in loranths be in the arrangement is still visible. changes seem to more frequent wet tropics, and may contribute to narrowing the guilds of animals which visit the plants as potential pollinators or predators. Simpler inflorescences are probably derived by reduction. There is a trend for reduction of triads to single flowers, usually along with reduction in inflorescence elaboration, and this has also occurred convergently in other generic groups in the region. The transformationseries postulated for Amyema (Barlow, 1966) illustrates the reductions which may occur. In Amylotheca, for example, the inflorescence of decussate in is often a simple raceme flowers, and Lysiana (see below) it is usual- ly a 2-flowered simple umbel and may be a solitary flower. In these cases the basic triad unit is no longer apparent, but vestigial structures (especially supernumerary bracts) and occasional aberrant developments indicatethe direction ofchange (Barlow, 1992). of decaisninoids The flowers the are hermaphrodite, mostly 6-merous, and usu- ally regular. Outgroup comparisons indicate that the plesiomorphic corolla state is choripetalous (Barlow, 1983), although it occurs in very few decaisninoid species. least Even in the apparently specialized genus,Decaisnina, the petals in most species in the lower after anthesis. remain weakly coherent part The short corolla tube so formed is sometimes dilated to form a distinct nectar chamber. Similar corolla char- acters are found in Cyne. In all other genera in the group the corollais gamopetalous. Parallel trends in Loranthaceae from flowers open choripetalous to tubular gamo- petalous ones are found in all continents, and are correlatedwith shifts from entomo- phily to ornithophily. The epipetalous stamens have basifixed immobile introrse anthers with simple filaments which in some cases are very short so that the anther is subsessile. In one the anther dorsifixed of a basal genus,Loxanthera, appears owing to development of anther. The is articulate above the distinct spur the style usually base, leaving a nipple on the fruit. 68 BLUMEA Vol. 38, No. 1, 1993 Vegetative features are relatively homogeneous.
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