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Department of Systematic Botany, Aarhus University, Nordlandsvej 68, DK-8240 Risskov, Denmark; and L. H. Bailey Hortorium, 467 Mann Library, Cornell University, Ithaca, New York 14853 USA
Received for publication December 21, 1999. Accepted for publication April 27, 2000.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Key Words: Arecaceae • Aphandra natalia • floral development • monotocotyledons • palms • Phytelephantoideae • pollination • pseudopedicel
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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). Developmental studies have shown that the phytelephantoid genera have the only monopodial flower clusters in the family, a four-parted perianth otherwise known only in only one species of Chelyocarpus (Coryphoideae), and centrifugal stamen inception. Partial centrifugal stamen development is known to occur elsewhere only in the genus Eugeissona (Calamoideae) (Uhl and Moore, 1977
; Uhl and Dransfield, 1984
).
Balslev and Henderson (1987)
originally referred Aphandra natalia to Ammandra based on the prominent submarginal veins on the pinnae and the pedicellate condition of the staminate flower clusters. Monographic work on the subfamily Phytelephantoideae has shown that it is a distinct genus (Barfod, Henderson, and Balslev, 1987
; Barfod, 1991
) (Table 1) and that the structure of the floral pedicel is critical. Developmental studies of the inflorescence and flowers of Aphandra are important for eludication of the pedicel and for comparison with developmental patterns previously described for the other genera of Phytelephantoideae (Uhl and Moore, 1977
; Uhl and Dransfield, 1984
). In this study we address three issues in particular: ontogeny of the staminate flower cluster, stamen inception, and the structure of the pedicel of the staminate flower. Some observations are also presented on the development of the pistillate flower clusters and flowers.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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); developmental material is difficult to obtain since large valuable trees must be felled to get a series. Inflorescence stages were obtained from three individuals growing in semi-cultivated populations near Logroño, Province of Morona-Santiago in Ecuador, the type locality of Aphandra natalia. The young inflorescences (Figs. 1–4) (Table 2) were fixed in FAA (formalin-acetic acid-alcohol) in the field and after 2 d, transferred to storage in nine parts 70% glycerine alcohol to one part 10% glycerine. Prophylls and peduncular bracts (PD bracts) were cut open to facilitate infiltration.
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Clearings
Material for study of vasculature was cleared using a 5% solution of NaOH and left overnight in an oven at 50°C. After 24 h the NaOH was changed and we repeated the procedure until clearing was satisfactory. The material was then rinsed in distilled water and treated with commercial bleach to remove any cloudiness. Subsequently preparations were stored in glycerine alcohol. The material was stained in a basic fuchsin solution (1 g of fuchsin and 6 g of solid KOH in 100 mL water) for photographing.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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). Inflorescences form in acropetal sequence in the axils of their subtending leaves. All leaves produced during periods of flowering subtend a young inflorescence bud. The pistillate and staminate buds have a similar overall appearance while they are still included in the sheaths of the subtending leaves, but they develop differently after the peduncular bract (PD bract) breaks through the prophyll (Figs. 1–7). In the staminate bud, the PD bract continues to grow and before it splits will attain more than twice the size of the prophyll. At anthesis the entire inflorescence is exposed and pendant. The pistillate bud, in contrast, elongates only a little after the splitting of the prophyll, the PD bract reaching only about three-quarters the length of the prophyll. The rachis of the pistillate inflorescence is much shorter, and the flowers remain closely bunched. At anthesis, they are partly contained in the PD bract, which splits longitudinally along the abaxial side to expose only the tips of long strap-shaped perianth parts and stigma branches (Fig. 19C).
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3 cm long at the base of the rachis to a narrow, few millimetre wide rim near the apex.
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Organogenesis of staminate flower clusters and flowers
In the earliest developmental stages available to us, the flower cluster sites are covered by their subtending bracts. Removal of the bract exposes a lentil-shaped apical dome in each flower cluster (Fig. 16). The two bracts that subtend the lateral flowers (Fig. 16: A, B) are clearly discernible (ba and bb), whereas the subtending bracts of the posterior and anterior flowers, respectively, are not visible in distal view. The apices of the four flowers are flattened to slightly dome-shaped (Fig. 17: A, B, and D). In distal outline, they appear three- to four-lobed because of primordial bulges (Fig. 18, 19D). On the lateral flowers, a pair of subopposite bracteoles develops first, slightly displaced relative to the plane of symmetry (Figs. 16, 19: 1b, 2b, 1a, 2a). The posterior member of this pair is usually visible before the anterior one. On the posterior flower of the cluster, the first-formed pair of subopposite bracteoles can be distinguished in lateral positions (Fig. 16: 1c, 2c). The anterior floral primordium is partly hidden under the basal part of the bract that subtends the entire flower cluster.
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650 µm in diameter, the first stamens arise in an incomplete ring around the central part of the apical dome (Fig. 21). Stamen primordia are initially round but become irregularly angled by close appression as they enlarge. The sides of the expanding apical dome are indented and flattened by the appressed perianth segments (Fig. 21). Subsequent stamen primordia arise in irregular groups along these edges (Fig. 22), in more or less centrifugal order, with those primordia that are covered by perianth parts often arising and enlarging later than more exposed ones. The occurrence of a pistillode is variable. Andrew Henderson et al. collected in 1990, staminate material from a population of Aphandra natalia in Acre, Brazil (Henderson et al. 1657 [NY, AAU, BH]). The material made available to us was sampled from a bud immediately before the splitting of the PD bract. Flowers close to the base of the rachis all had large pistillodes visible to the unaided eye, consisting of rudimentary carpels extended apically in long slender stigmatic branches. In some of the young stages of the inflorescences collected at Logroño, solitary flowers near the apex produce hairy processes up to 1 cm long arising from the middle of the receptacle. These, however, appear to be staminodial in origin.
Late in staminate organogenesis, after stamen inception has stopped but immediately before splitting of the PD bract, the lower part of the floral receptacle, sheathed in adnate and connate perianth bases, undergoes zonal growth to form a stalk that we have designated as a pseudopedicel. At maturity the epidermis and one or more subepidermal layers of the pseudopedicel become split and the surface acquires a fuzzy appearance due to a covering of broken cells (Figs. 11, 12). Raphide-containing idioblasts very similar to the pollen grains in shape and size are released from the split layers in great numbers. They remain trapped in the epidermal shreds on the surface where they mix with pollen grains at anthesis (Figs. 12, 13).
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Vascular anatomy
The large number of collateral bundles supplying each flower varies with the size and developmental stage. As the floral apex expands longitudinally and in diameter the bundles curve towards the periphery and anastomose frequently (Fig. 10).
Pistillate inflorescence and flowers: structure and development
Stages of pistillate organs were incomplete but some observations were possible. The PD bract of the pistillate inflorescence bud differs from its staminate homologue by having an emarginate apex (Figs. 1, 5). The rachis of the inflorescence resembles the staminate but bears only 20 solitary, spirally arranged, pistillate flowers, each in the axil of a bract. It is noteworthy that two subopposite pairs of bracteoles are present below each flower. In fully developed flowers, the bracteoles can be distinguished from the perianth segments by being smaller and less fleshy. In Fig. 1, the youngest stage available, the subtending bract, four bracteoles, and the perianth of the pistillate flowers are already elongate. The central whitish bulges in individual flowers are the multicarpellate gynoecia. The flowers in stage number VII of our collection have free carpel primordia that are conduplicate. The carpel primordia are present before the initiation of the staminodes, which first appear as a single series of primordia surrounding the gynoecium, a second whorl eventually forming on the outside.
The development of the pistillate perianth is irregular. Our collections did not comprise the earliest stages of perianth inception, but the sizes and insertion of the segments suggest that more than one whorl is involved. Whereas the first-formed segments are uniform in shape and in size, additional segments usually form on the abaxial side of the flower. This phenomenon, possibly related to asymmetrical expansion of the apex, was also observed in Phytelephas macrocarpa R. and P. (Barfod, 1991
).
Early in organogenesis, the apices of the carpels elongate. The resulting stigmatic branches become longer than the perianth segments as can be determined from the material shown in Figs. 1 and 2. The stylar portion of the gynoecium elongates continuously until anthesis when the stigmatic branches are surrounded by the apices of the long perianth segments. At this time the style length ranges between 20 and 25 cm and the stigmatic branches are 4–5 cm long (Fig. 19C).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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), that the floral apex expands in width and height during stamen development, and that stamen development is for the most part centrifugal (Uhl and Moore, 1977
). The stalk of the staminate flower was determined to be structurally unique and designated as a pseudopedicel, which consists of an enlarged, more or less funnel-shaped floral receptacle with an adnate perianth tube. The small tips of the perianth members are free distally. The epidermis and one or more underlying layers of the pseudopedicel contain large raphide idioblasts, resembling the pollen grains in shape. The raphides are freed by disintegration of the epidermal cells and become mixed with pollen grains on the surface of the pseudopedicel at anthesis. Comparison with other genera of the Phytelephantoideae and further consideration of these findings are given below.
Staminate flower cluster
Uhl and Dransfield (1984)
suggested that in Phytelephas aequatorialis (syn. Palandra aequatorialis), the four bract-like appendages surrounding the perianth in the pistillate flowers are homologous to the four bracts that subtend the flowers of the staminate cluster and termed them "bracteoles." Aphandra differs from P. aequatorialis in that four bract-like appendages are also present below the perianth on each staminate flower. Barfod (1991)
referred to these as sepaloid bracts. The four first-formed bracteoles on each staminate floral axis and the four bracteoles, or sepaloid bracts, of the pistillate flower are homologous. Thus, the homology is between the pistillate flower and a single flower of the staminate cluster. The extra bracteoles might suggest a more branched staminate flower cluster in Aphandra.
Stamen inception
In multistaminate palms, floral receptacles expand to accommodate more stamens, the shape of the receptacle and pattern of origin of primordia varying in different groups (Uhl and Moore, 1980
). The three genera of phytelephantoid palms are distinguished by bizarre differences in receptacle shape during stamen initiation. Floral development of two species of Phytelephas has been studied. In Phytelephas aequatorialis the staminate flowers are pedicellate from early in organogenesis; the pedicels are true stalks, not pseudopedicels as in Aphandra. When the first stamen primordia originate, the receptacle in P. aequatorialis is 800 µm in diameter and relatively flat. Stamen inception occurs rapidly in two phases. During a first phase, the entire apex becomes covered by primordia except for a marginal meristematic area. Although the primordia surrounding the center are larger, Uhl and Moore (1977)
did not observe any apices with a single ring of primordia. In the second phase, the stamen primordia develop in centrifugal order in radial rows along the meristematic periphery and thereby increase the size of the apex. At this stage, a pistillode was observed in some flowers. Five to seven hundred stamens are usually produced in each staminate flower.
In Phytelephas macrocarpa, the staminate clusters and flowers are sessile, the flowers remaining sessile throughout development. At the time of inception of the first stamens, the receptacle is 550 µm wide and slightly curved with a raised apex. Two types of meristem are active, either simultanously or in succession. One produces a limited number of stamen primordia at early stages of stamen inception, apparently in a centripetal pattern (Uhl and Moore, 1977
). The remaining stamens are produced in a centrifugal sequence by the peripheral meristem, which is active throughout organogenesis. Pistillodes were not observed. The number of stamens formed in each staminate flower varies from 150 to 300.
In species of Phytelephas and in Aphandra the receptacles are shallow domes in early ontogenetic stages. The receptacle expands in diameter but only slightly in height in both species of Phytelephas. In Aphandra natalia, however, the receptacle also elongates, becoming somewhat funnel-shaped, with stamen primordia originating centrifugally along the angled and flattened sides, often partly under the perianth segments. At anthesis a naked zone remains between the level of insertion of perianth segments and the stamen-bearing part of the receptacle. Our evidence is not conclusive as to whether there is a short centripetal phase previous to centrifugal stamen inception in Aphandra. In some apices studied one to six primordia may develop later in the center of the flower as suggested by the presence of inner bulges. They do not differ from the surrounding stamen primordia in shape, only in size. The bulges observed could represent the carpel primordia, but that remains to be demonstrated. A similar developmental pattern was considered partial centrifugal development in Phytelephas macrocarpa (Uhl and Moore, 1977
, fig. 18).
In the third genus, Ammandra, the receptacle at anthesis is irregularly chunky and becomes expanded basally to cover the perianth apices on the abaxial side of the flower. The stamen filaments are much shorter, and the anthers are smaller than in the other phytelephantoid genera (Barfod, 1991
). Button-shaped pistillodes, inserted in shallow depressions and usually caducous at anthesis, are almost universally present. Anatomical sections of the fully developed flowers show that most of the major trunk bundles extend to about three-fourths the height of the flower and then curve toward the periphery and extend downward (Uhl and Moore, 1977
), suggesting centrifugal initiation. No other observations of stamen inception have been made. Up to 1200 stamens have been recorded in A. dasyneura (Burret) Barfod (Barfod, 1991
), the highest number in the family.
The pseudopedicel
Floral stalks or pedicels vary among the phytelephantoid genera. Both sessile and pedicellate flowers occur in Phytelephas. The stalked condition of the staminate flower cluster was used to separate the genus Palandra from Phytelephas (Cook, 1927
), but the character was found variable among the species of Phytelephas, and Palandra has since been placed in synonomy (Barfod, 1991
). It is noteworthy that in Phytelephas tumacana Cook, the clusters in the middle and proximal part of the inflorescence are composed of four (rarely five) flowers that are sessile to subsessile, whereas a number of solitary flowers are always present at the apex with pedicels up to 7 mm long (Barfod, 1991
). In other species of Phytelephas, as in P. macrocarpa, all flowers are sessile. Here also Ammandra differs in having stalked staminate flowers borne in clusters of up to nine on a 1.5–3 cm long branch. The floral stalk appears to be a short branch, but this requires confirmation by developmental and anatomical studies.
In Aphandra the pseudopedicel is a special structure, an elongate floral receptacle with an adnate and connate perianth tube. Several characters appear associated with the pedicellate condition. The rapid expansion of the pseudopedicel may help generate the pressure that is needed for the flower-bearing tissues to break through the PD bract. Secondly, such rapid increase in size may contribute to the heating of the bud above ambient temperature during pollination. Thirdly, the epidermis disintegrates releasing raphide idioblasts at the same time as pollen grains are shed, and the idioblasts and pollen intermingle, covering the surface of the pseudopedicel. In Ammandra raphide idioblasts are also found in great numbers, not scattered in subepidermal layers as in Aphandra, but inside blister-like structures that occur scattered on the pedicel and the receptacle as well. The blisters rupture at anthesis to release the idioblasts. As in Aphandra the idioblasts resemble the rounded pollen grains in size and shape. Such idioblasts may help to deter pollen feeding and perforation of pedicels and receptacles for ovipositing.
Pistillate flower clusters and flowers
Except for the number of flowers in the inflorescence (see Table 1), structure and development of the pistillate flower clusters and flowers in Aphrandra are like those found for Phytelephas aequatorialis where similar carpel primordia become fused laterally later in development, the ventral sutures remaining open throughout (Uhl and Dransfield, 1984
). The presence of two pairs of sepaloid bracts below the perianth has been noted. Two indistinct four-parted perianth whorls are followed by a ring of conduplicate carpel primordia. In P. aequatorialis carpels became laterally connate, but ventral sutures remain open and the single ovule of each carpel is initiated directly on the floral axis, in a position that might be considered axillary to the carpel. At maturity the gynoecium of all phytelephantoid genera has a central, cone-shaped receptacle, histologically distinguished by large parenchyma cells and scattered tannins. Similar gynoecial structure is found elsewhere in the family only in the genera of the Calamoideae, where the receptacle is different in shape and histology.
Staminodes in Aphandra originate in two whorls, the first whorl next to the carpel primorida and a second outside the first. Uhl and Moore (1977)
demonstrated the same kind of centrifugal staminodial inception in the pistillate flowers of Phytelephas aequatorialis.
Pollination
Several studies have been undertaken on the pollination of phytelephantoid palms. Barfod, Henderson, and Balslev (1987)
and Barfod (1991)
studied Phytelephas macrocarpa ssp. tenuicaulis Barfod in Amazonian Ecuador and concluded that several insect groups mediate pollen transfer. It is noteworthy that beetles of the families Staphylinidae, Nitulidae, and Curculionidae were all ovipositing in the staminate inflorescence. At the time of anthesis, they had perforated the receptacle completely. Bernal and Ervik (1996)
described the floral biology and pollination of Phytelephas seemannii Cook in Colombia. Pollination is mostly carried out by species of pollen-eating and predating staphylinids. One particular species of the genus Amazoncharis reproduces in the male inflorescences by constructing egg chambers in the fleshy receptacles of the flowers. This reproductive behavior resembles that of beetles in the closely related subtribe Gyrophaenina that feed on the spores and reproduce in fleshy mushrooms.
Aphandra natalia is pollinated mainly by Baridinae (Coleoptera) that feed on pollen and, in contrast to the situation in other phytelephantoid species, prefer the pistillate rather than staminate inflorescence for oviposition (Ervik, 1992
). The pseudopedicel may play a key role in the interaction with the visiting insects. The raphide idioblasts that are released from the hypodermal layers in great numbers could be detrimental to the larvae. Their similarity in size and shape to the pollen grains are striking (Figs. 12, 13). A simple bio-assay could be designed to demonstrate whether the pollinating insects are able to distinguish between the two.
Ervik, Tollsten, and Knudsen (in press)
have analyzed the floral scent of Ammandra decasperma, A. dasyneura, Aphandra natalia, Phytelephas aequatorialis, P. macrocarpa ssp. tenuicaulis and P. seemannii. Their results show that the major constituents of the floral scents of the three genera are of completely different biochemical origin. The floral scent of Aphandra is unusual in being dominated by a pyrazine. The presence of this compound could explain the absence in Aphandra natalia of Derelomini and Mystrops that are common visitors in all Phytelephas species studied as well as in many other palm and cyclanth species (Henderson, 1986
; Gottsberger, 1991
; Eriksson, 1994
).
Phylogenetic implications
Cladistic analyses based on morphology and restriction site fragments found the Phytelephantoideae monophyletic (Barfod, 1991
; Uhl et al., 1995
). The group is strongly supported by a number of synapomorphies, although some of the characters are found elsewhere throughout the family, such as the multicarpellate condition (some species of Attalea), seeds attached to an extension of the receptacle (Eugeissona), floral buds open (e.g., Ceroxylon), and the breaking up of the fruit mesocarp into corky processes (e.g., Manicaria, Pelagodoxa, Sommiera, and Johannesteijsmannia).
Barfod (1991)
used morphological data to infer relationships among genera of the Phytelephantoideae. An outgroup was assembled representing the major evolutionary lineages within the palms according to Uhl and Dransfield (1987)
. Floral characters were emphasized by weighting procedures. The result was ambiguous due to lack of a likely sister group, but the cladogram favored had Aphandra and Phytelephas as a clade with Ammandra as sister group. One problem of weighting in favor of floral characters is that these are often the result of coevolutionary relationships and therefore do not neccessarily reflect phylogeny. The findings of this study further corrobate an Aphandra–Phytelephas clade. The early, sessile ontogenetical stages of the staminate flowers of Aphandra natalia are very similar to similar stages in Phytelephas macrocarpa. This similarity applies to number of flowers per cluster, stamen inception, length of the flower-bearing axis, and shape and vasculature of the receptacle. Subsequent development, however, is very different.
This study further supports the circumscription of Aphandra based on several characters of the staminate flower, in particular the presence of four sepaloid bracteoles, the shape of the floral receptacle, and a structurally unique pseudopedicel. Stamen number, and the morphology and number of pistillate flowers per inflorescence, are also different (Table 1).
The relationship of the phytelephantoid genera to other palms is not yet clear. Some trees (Uhl et al., 1995
) based on morphological and molecular characters resolved the Ceroxyleae as a sister group. A chloroplast DNA (cpDNA) restriction fragment analysis that compares representatives of the three phytelephantoid genera with a joint outgroup of 12 taxa is currently underway to further seek a likely sister group. Several of the gene trees generated using simple parsimony and Nypa as a functional outgroup have Ceroxylon as sister group. Tribe Ceroxyleae in the ceroxyloid palms has a Gondwanic distribution pattern with representatives in the Juan Fernandes Islands, northern South America, Madagascar, and Australia. In general the group is less specialized than the Phytelephantoideae, but further research is necessary to establish this relationship. The Phytelephantoideae is of primary importance in understanding evolutionary trends within the palm family. The superficial resemblance of inflorescences and flowers with certain cyclanthaceous groups such as Cardulovica is noteworthy. The phytelephantoid palms share with members of this genus, strong floral dimorphy, monopodial flower clusters, pseudopedicels, a multistaminate condition, four-merous floral whorls, spicate inflorescences, and lack of vessels in the stem (Dahlgren, Clifford, and Yeo, 1985
). There are, however, numerous differences, and comparative analysis of restriction fragments (Davis, 1995
) as well as DNA sequences (Chase et al., 1995
) has clearly demonstrated that these groups are only remotely related to each other. The respective pollination syndromes of the cyclanthoid genus Cardulovica and the phytelephantoid genera are very similar. Both groups are beetle pollinated, and they share the following features: nocturnal flowering, color, scent and temperature elevation (Barfod and Henderson, 1987; Barfod, 1991
; Gottsberger, 1991
; Eriksson, 1994
). Their similarities appear to represent a striking example of convergent evolution in response to uniform selection pressures in similar habitats.
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FOOTNOTES |
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2 Author for correspondence (NWU1{at}cornell.edu
).
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Barfod, A. S. 1991 A monographic study of the subfamily Phytelephantoideae. Opera Botanica 104: 1–76
———, A. Henderson, and H. Balslev. 1987 A note on the pollination of Phytelephas microcarpa (Palmae). Biotropica 19: 191–192
Bernal, R., and F. Ervik. 1996 Floral biology of the dioecious palm Phytelephas seemannii in Colombia: an adaptation to staphylinid beetles. Biotropica 28: 682–696[CrossRef][ISI]
Borgtoft Pedersen, H., and H. Balslev. 1990 Ecuadorean palms for agroforestry. AAU reports 23: 1–122
Chase, M. W., D. W. Stevenson, P. Wilkin, and P. J. Rudall. 1995 Monocot Systematics: a combined analysis. In P. J. Rudall et al. [eds.], Monocotyledons: systematics and evolution, 685–730. Royal Botanic Gardens, Kew, Richmond, Surrey, UK
Cook, O. F. 1927 New genera and species of Ivory Palms from Colombia, Ecuador and Peru. Journal of the Washington Academy of Sciences 17: 218–230
Dahlgren, R., H. T. Clifford, and P. F. Yeo. 1985 The families of the monocotyledons: structure, evolution, and taxonomy. Springer Verlag, Berlin, Germany
Davis, J. I. 1995 A phylogenetic structure for the monocotyledons, as inferred from chloroplast dna restriction site variation and a comparison of measures of clade support. Systematic Botany 20: 503–527[CrossRef][ISI]
Eriksson, R. 1994 The remarkable weevil pollination of the neotropical Cardulovicoideae (Cyclanthaceae). Plant Systematics and Evolution 189: 75–81[CrossRef][ISI]
Ervik, F. 1992 Notes on the phenology and pollination of the dioecious palms Mauritia flexuosa (Calamoideae) and Aphandra natalia (Phytelephantoideae) in Ecuador. In W. Barthlott et al. [eds.], Animal-plant interactions in tropical environments, 7–12. Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany
Ervik, F., L. Tollsten, and J. T. Knudsen. In press Floral scent chemistry and pollination ecology in phytelephantoid palms (Arecaceae). Plant Systematics and Evolution
Gottsberger, G. 1991 Pollination of some species of the Carludovicoideae, and remarks on the origin and evolution of the Cyclanthaceae. Botanische Jahrbücher für Systematik 113: 221–235
Henderson, A. 1986 A review of pollination studies in the Palmae. Botanical Review 52: 221–259
Uhl, N. W., and J. Dransfield. 1984 Development of the inflorescence, androecium, and gynoecium with reference to palms. In R. A. White and W. C. Dickison [eds.], Contemporary problems in plant anatomy, 397–449. Academic Press, New York, New York, USA
———, and H. E. Moore, Jr. 1977 Centrifugal stamen initiation in phytelephantoid palms. American Journal of Botany 64: 1152–1161[CrossRef][ISI]
———, and ———. 1980 Androecial development in six polyandrous genera representing five major groups of palms. Annals of Botany, London 45: 57–75
———, and ———. 1987 Genera Palmarum. L.H. Bailey Hortorium and International Palm Society. Allen Press, Lawrence, Kansas, USA
———, J. I. Davis, M. A. Luckow, K. S. Hansen, and J. J. Doyle. 1995 Phylogenetic relationships among palms: cladistic analyses of morphological and chloroplast DNA restriction site variation. In P. J. Rudall et al. [eds.], Monocotyledons: systematics and evolution, 623–661. Royal Botanic Gardens, Kew, Richmond, Surrey, UK
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