Homologies of the flower and inflorescence in the early-divergent grass Anomochloa (Poaceae)

? Premise of the study: The grass subfamily Anomochlooideae is phylogenetically significant as the sister group to all other grasses. Thus, comparison of their structure with that of other grasses could provide clues to the evolutionary origin of these characters. ? Methods: We describe the structure, embryology, and development of the flower and partial inflorescence of the monotypic Brazilian grass Anomochloa marantoidea. We compare these features with those of other early-divergent grasses such as Pharus and Streptochaeta and closely related Poales such as Ecdeiocolea. ? Key results: Anomochloa possesses several features that are characteristic of Poaceae, notably a scutellum, a solid style, reduced stamen number, and an ovary with a single ovule that develops into a single indehiscent fruit. Interpretation of floral patterning in Anomochloa is problematic because the ramification pattern of the florets places the bracts and axes in unusual positions relative to the primary inflorescence axis. Our study indicates that there is a single abaxial carpel in Anomochloa, probably due to a cryptic type of pseudomonomery in Anomochloa that resembles the pseudomonomery of other grasses. On the other hand, the Anomochloa flower differs from the "typical" grass flower in lacking lodicules and possessing four stamens, in contrast with the tristaminate condition that characterizes many other grasses. ? Conclusions: Using the median part of the innermost bract as a locator, we tentatively homologize the inner bract of the Anomochloa partial inflorescence with the palea of other grasses. In this interpretation, the pattern of monosymmetry due to stamen suppression differs from that of Ecdeiocolea.     

CENTRIFUGAL STAMEN INITIATION IN PHYTELEPHANTOID PALMS

The phytelephantoid group, considered very highly specialized among the 15 major groups of palms, includes only three dioecious genera which are distinguished by multipartite flowers. Staminate flowers of these genera have from 120 to over 900 stamens, the largest number known in palms. Developmental material shows centrifugal inception of stamens in Palandra and Phytelephas, and the course of the trunk bundles indicates centrifugal initiation of stamens in Ammandra. Stamens of all three genera are supplied by large trunk bundles that develop acropetally following the different centrifugal expansion of the floral apex in each genus. Branches of the trunk bundles form later to supply stamen primordia as they develop. Polyandric androecia occur in ten of the major groups of palms. Other multistaminate genera show expansion and change in shape of the floral apex before stamen initiation. Centrifugal development appears to represent a different method of expansion of the floral apex to accommodate increase in stamen number in this group. Centrifugal inception of stamens in monocotyledons has previously been established only for two genera of Butomaceae.     

Morphological evolution in the graminid clade: comparative floral anatomy of the grass relatives Flagellariaceae and Joinvilleaceae

New comparative data are presented on the reproductive morphology and anatomy of two genera closely related to grasses, Flagellaria and Joinvillea, in which the flowers are superficially similar, especially in stamen morphology. This investigation demonstrates some anatomical differences between the two genera. For example, both genera depart from the ‘typical’ condition of tepal vasculature (three‐traced outer tepals and one‐traced inner tepals): in Flagellaria, each tepal receives a single vascular bundle and, in Joinvillea, each tepal is supplied by three vascular bundles. Joinvillea possesses supernumerary carpel bundles, as also found in the related family Ecdeiocoleaceae, but not in Flagellaria or grasses. In the anther, the tapetum degenerates early in Flagellaria, and is relatively persistent in Joinvillea, in which the pollen grains remain closely associated with the tapetum inside the anther locule, indicating a correlation between peripheral pollen (a feature that is common in grasses) and a persistent tapetum. This study highlights the presence of a pollen‐tube transmitting tissue (PTTT) or solid style in the gynoecium of Flagellaria, as also in many Poaceae, but not in Joinvillea or Ecdeiocoleaceae. We speculate that the presence of a PTTT could represent one of the factors that facilitated the subsequent evolution of the intimately connected gynoecia that characterize grasses. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 393–404.     

Migration of endpoints of two genes relative to boundaries between regions of the plastid genome in the grass family (Poaceae)

Overlapping genes occur widely in microorganisms and in some plastid genomes, but unique properties are observed when such genes span the boundaries between single-copy and repeat regions. The termini of ndhH and ndhF, situated near opposite ends of the small single-copy region (SSC) in the plastid genomes of grasses (Poaceae), have migrated repeatedly into and out of the adjacent inverted-repeat regions (IR). The two genes are transcribed in the same direction, and the 5' terminus of ndhH extends into the IR in some species, while the 3' terminus of ndhF extends into the IR in others. When both genes extend into the IR, portions of the genes overlap and are encoded by the same nucleotide positions. Fine-scale mapping of the SSC-IR junctions across a sample of 92 grasses and outgroups, integrated into a phylogenetic analysis, indicates that the earliest grasses resembled the related taxa Joinvillea (Joinvilleaceae) and Ecdeiocolea (Ecdeiocoleaceae), with ca. 180 nucleotides of ndhH extending into the IR, and with ndhF confined to the SSC. This structure is maintained in early-diverging grass lineages and in most species of the BEP clade. In the PACMAD clade, ndhH lies completely or nearly completely within the SSC, and ca. 20 nucleotides of ndhF extend into the IR. The nucleotide substitution rate has increased in the PACMAD clade in the portion of ndhH that has migrated into the SSC.     

Reproductive morphology of the early-divergent grass Streptochaeta and its bearing on the homologies of the grass spikelet

Reproductive morphology and development are described in the Brazilian grass Streptochaeta spicata, in order to assess the homologies of the characteristic grass inflorescence, termed a spikelet, and other reproductive organs. Streptochaeta possesses some features that are commonly found in Poaceae, including a well-differentiated embryo. It also possesses some relatively unusual, presumably derived features, such as non-plumose stigmas, which indicate that it could be insect-pollinated. It shares some features with other early-divergent grasses, such as Pharus, which could represent plesiomorphic conditions for grasses. The inflorescence unit in Streptochaeta has been interpreted as a compound branching system or pseudospikelet. The present data suggest that it is a highly modified spikelet, with a modified flower borne either on a different axis to the basal bracts (glumes) or on the same axis as the basal bracts. The three bracts below the stamens are interpreted as homologous to the lodicules. The Streptochaeta spikelet could be considered as morphologically intermediate between the true spikelet of grasses and reproductive units of close grass relatives.     

杠杆状雄蕊及其进化生态学意义

对被子植物一类特化雄蕊——杠杆状雄蕊的结构多样性及其进化生态学意义进行了归纳总结。植物的花在进化过程中, 常会发生雄蕊群的改变, 包括雄蕊数目及其形态结构的变化, 同时雄蕊功能也会发生相应的适应性转变。杠杆状雄蕊是指结构特化为杠杆状或距状, 在传粉过程中具有类似杠杆功能的一类特化的雄蕊类型。目前, 已在唇形科不同亚科以及姜科6个属中发现杠杆状雄蕊, 根据其结构和形态发生方式总体上可分为2大类: 一类是唇形科中以鼠尾草属(Salvia)为代表的, 由2个可育雄蕊平行发育、药隔组织增长所形成的杠杆状雄蕊; 另一类是姜科植物中由一个可育雄蕊特化形成的带有距状附属体的雄蕊类型。在生态功能上, 两类雄蕊均能通过传粉者推动其距状下臂做杠杆运动进行传粉, 被认为是一种促进异交的传粉机制, 可通过精确传粉和花粉分发等途径影响植物的繁殖成功。杠杆状雄蕊在不同的类群中是独立起源与进化的; 仅在唇形科鼠尾草属中, 杠杆状雄蕊发生了3次独立进化, 而且它可能是触发该属物种适应性辐射的关键性状。将来需在宏观进化和微观进化两个水平深入探讨杠杆状雄蕊的进化生态学意义。     

Comparative floral development in the tribe Mentheae (Nepetoideae: Lamiaceae) and its bearing on the evolution of floral patterns in asterids

A comparative developmental study of flowers was carried out using epi-illumination light microscopy on four genera of Lamiaceae (Nepeta, Rosmarinus, Salvia, andZiziphora), representing all three subtribes of Mentheae. All species examined share unidirectional (adaxial to abaxial) sepal initiation, except Rosmarinus, which has the reverse unidirectional sequence, starting abaxially. Initiated but suppressed bracteoles were detected only in Rosmarinus. In Rosmarinus, Salvia, and Ziziphora, initiation of petals and stamens proceeds unidirectionally from the abaxial side. Floral initiation of Nepeta has bidirectional inception of petals and unidirectional stamen initiation from the adaxial side. Temporal overlap in organ initiation between petal and stamen whorls occurs in all taxa, though this feature is more prominent in Rosmarinus. Significant structural and developmental features that distinguish the four genera include: (1) polysymmetric calyx tube, highly tomentose corolla and deeply four-partitioned ovary in Nepeta; (2) monosymmetric two-lipped calyx and shallowly four-partitioned ovary in Ziziphora; and (3) suppression of adaxial stamens in Salvia and Rosmarinus. Adaxial stamens are absent from Rosmarinus, but reduced stamens remain as staminodia in Salvia. In a phylogenetic context, the late monosymmetry of Nepeta and very early monosymmetry of Rosmarinus could both be regarded as derived conditions compared with the early monosymmetry ofSalvia and Ziziphora.     

菰属系统与演化研究—胚形态

本文采用光学显微镜对全世界菰属４种２亚种及其有关属种共４属７种的胚形态进行系统研究后,获得菰属Ｚｉｚａｎｉａ Ｌ．与山涧草Ｃｈｉｋｕｓｉｃｈｌｏａ ａｐｕａｔｉｃａ Ｋｏｉｄｚ．的胚型为Ｆ＋ＦＰ,稻Ｏｒｙｚａ ｓａｔｉｖａ Ｌ．与拟菰Ｚｉｚａｎｉｏｐｓｉｓ ｍｉｌｉａｃｅａ（Ｍｉｃｈｘ．）Ｄｏｅｌｌ ＆ Ａｓｃｈｅｒｓ．的胚型为Ｆ＋ＦＰ,其中山涧草的胚型为首次报导。根据胚型,这４属均应位在禾本科Ｇ     

ANATOMY OF THE CARYOPSIS OF BRIZA MAXIMA (POACEAE)

Briza maxima (quaking grass) is a cosmopolitan grass common to Europe and North and South America. It grows in disturbed soils and on roadsides. The hemispherical caryopsis is embedded between a leaflike lemma and flattened palea. The embryo is of the festucoid type. The scutellum shows two surrounding ridges at the edge of the scutellum/endosperm boundary, and has lateral lobes. A broad epiblast extends toward the embryo apex and is continuous with the dorsal surface of the coleorhiza. The single-layered aleurone surrounds the starchy endosperm and is discontinuous around the embryo. The caryopsis coat is thin, except at the placental pad where it is thickened by the pigment strand and the nucellar projection.     

Phylogenetic relationships among Poaceae and related families as inferred from morphology, inversions in the plastid genome, and sequence data from the mitochondrial and plastid genomes

A phylogenetic analysis of the Poales was conducted to assess relationships among Poaceae and allied families. The analysis included 40 taxa, representing all families of the Poales as circumscribed by the Angiosperm Phylogeny Group (APG), plus five of the six unplaced Commelinid families in the APG system. The data matrix included 98 informative characters representing variation in morphology and chloroplast genome structure (including three inversions in the chloroplast genome), and 563 informative characters derived from rbcL and atpA nucleotide sequences. Ecdeiocolea has the 6-kilobase (kb) chloroplast genome inversion previously reported in Joinvillea and Poaceae, and like Joinvillea it lacks the trnT inversion that occurs in grasses. Analysis of the morphological data places Poaceae in an unresolved relationship relative to several other taxa, including Joinvillea and Ecdeiocolea, while analysis of the molecular and combined data resolves Ecdeiocolea as sister of Poaceae, with Joinvillea the sister of this group. Although the 6-kb and trnT inversions are non-homoplasious in the phylogenies obtained in this study, the 28-kb inversion is optimized as having originated twice (once in Restionaceae and another time in the most recent common ancestor of Ecdeiocolea, Joinvillea, and the grasses); an alternative interpretation is that it arose once and was later lost in Anarthria. Ecdeiocolea shares with Poaceae the presence of operculate, annulate pollen that lacks scrobiculi, and a dry, indehiscent fruit.     

Floral ontogeny of Annonaceae: evidence for high variability in floral form

Background and Aims

Annonaceae are one of the largest families of Magnoliales. This study investigates the comparative floral development of 15 species to understand the basis for evolutionary changes in the perianth, androecium and carpels and to provide additional characters for phylogenetic investigation.

Methods

Floral ontogeny of 15 species from 12 genera is examined and described using scanning electron microscopy.

Key Results

Initiation of the three perianth whorls is either helical or unidirectional. Merism is mostly trimerous, occasionally tetramerous and the members of the inner perianth whorl may be missing or are in double position. The androecium and the gynoecium were found to be variable in organ numbers (from highly polymerous to a fixed number, six in the androecium and one or two in the gynoecium). Initiation of the androecium starts invariably with three pairs of stamen primordia along the sides of the hexagonal floral apex. Although inner staminodes were not observed, they were reported in other genera and other families of Magnoliales, except Magnoliaceae and Myristicaceae. Initiation of further organs is centripetal. Androecia with relatively low stamen numbers have a whorled phyllotaxis throughout, while phyllotaxis becomes irregular with higher stamen numbers. The limits between stamens and carpels are unstable and carpels continue the sequence of stamens with a similar variability.

Conclusions

It was found that merism of flowers is often variable in some species with fluctuations between trimery and tetramery. Doubling of inner perianth parts is caused by (unequal) splitting of primordia, contrary to the androecium, and is independent of changes of merism. Derived features, such as a variable merism, absence of the inner perianth and inner staminodes, fixed numbers of stamen and carpels, and capitate or elongate styles are distributed in different clades and evolved independently. The evolution of the androecium is discussed in the context of basal angiosperms: paired outer stamens are the consequence of the transition between the larger perianth parts and much smaller stamens, and not the result of splitting. An increase in stamen number is correlated with their smaller size at initiation, while limits between stamens and carpels are unclear with easy transitions of one organ type into another in some genera, or the complete replacement of carpels by stamens in unisexual flowers.     

The structure and function of the scutellum of the Grarnineae

NEGBI, M., 1984. The structure and function of the scutellum of the Gramineae. Four kinds of scutella, of which only the first is universally known, can be distinguished in the Gramineae. (1) The scutellum sew stricto , the kind most commonly described in textbooks. In this scutellum the only growth activity during germination is the development of every epithelial cell into a separate elongated papilla. These papillae are involved in secretion of hydrolases, gibberellins and other hormonal factors which in their turn activate the aleurone layer; and in absorption of the mobilized endosperm reserves. (2) The kind characteristic of Auma is found in several genera. In this the scutellar tip elongates during germination, reaches the distal end of the endosperm sac and develops papillae over its whole surface. (3) The kind found in Cizuniu in which the scutellar tip elongates and extends to the far end of the caryopsis during embryo development, but not during germination. In this scutellum only the abaxial surface faces the bulk of the storage endosperm and probably only this surface becomes papillate. Several bamboo genera have the kind of scutellum characterized by Melocannu . This scutellum has evolved as a storage organ and in mature caryopses the endosperm is reduced. This kind is associated with vivipary and with the presence of storage tissue in the pericarp.
The vascularization and the structure of the scutellar epithelium, as studied mainly in a limited number of species belonging to the first kind, are related to the functions of the scutellum. The scutellum has a prime role in controlling the mobilization of endosperm reserves.     

PATTERNS OF STAMEN VASCULATURE IN THE ARACEAE

A survey of the three-dimensional organization of stamen vasculature in 100 genera and over 350 species of Araceae was made using clearings. The Araceae exhibit highly varied stamen vasculature, with three main patterns: 1) vascular bundles unbranched, 1–3 per stamen, 2) forked bundles in some or all stamens, 3) anastomosing vascular systems with several to many bundles entering a single stamen. Three major groups of taxa in the family can be recognized on the basis of their predominant pattern of stamen vasculature. Virtually all genera with bisexual flowers (most Pothoideae, Monsteroideae, Calloideae, Lasieae) have unbranched bundles, one per stamen, except two to three in some species of Holochlamys, Spathiphyllum, and Scindapsus. Forked stamen bundles are virtually restricted to and occur nearly throughout the monoecious Lasioideae, Philodendroideae, Colocasioideae and among certain Aroideae (sensu Engler), including tribes Arophyteae, Spathicarpeae (Asterostigmateae) and Protareae. No forked bundles were found in tribe Areae (Aroideae), except Theriophonum indicum or any Araceae with bisexual flowers, except two species of Cyrtosperma. Anastomosing systems are virtually limited to members of tribe Areae with larger stamens, such as Arum, Helicodiceros, Eminium and Dracunculus species. A similar pattern occurs in some Amorphophallus, but other patterns occur as well. The distributions of forked bundles and anastomosing systems in the family are notable because they are both highly congruent with Philodendroideae-Colocasioideae, and Aroideae, respectively, in Grayum's new system for the family. Virtually all of the genera with forked bundles are grouped together in the Philodendroideae-Colocasioideae. All of the genera with anastomosing systems are in the Areae, including the complex and variable Amorphophallus, which has an uncertain systematic placement.     

三白草科花部发育及其系统学意义

本研究从比较三白草科属间小花个体发育及分析花器官数量变异入手,探寻花器官在发生顺序、数目变化及排列方式等方面的演化趋势,揭示系统发育在个体发育中一定程度重现的事实及属间的进化关系。结果简述如下:首先,雄蕊和心皮发生顺序由中部优先演化到两侧优先。其次,由于远中雄蕊和心皮经历了从发育延迟、生长减缓到最终消失的历程,中部雄蕊和心皮由成对演化为单生。此外,两侧生雄蕊对由各自独立的原基发生演化到共同原基发生或减化为1枚,假银莲花属近中1枚雄蕊原基二裂成1对,蕺菜属3枚心皮发生于一环状共同原基等,都是该科花器官演化的重要事实并可归结为融合、减化和复化的结果。文章根据花器官的演化趋势及过渡类型的剖析,论述了三白草科属间的系统进化关系。     

IRREGULAR FLORAL DEVELOPMENT IN CALLA PALUSTRIS (ARACEAE) AND THE CONCEPT OF HOMEOSIS

About two-thirds of the more than 100 genera in the Araceae lack tepals and their absence is considered derived. Unlike most of these atepalate genera, Calla palustris has about twice as many stamens per flower. Using epi-illumination microscopy, we studied floral development in Calla to see if the supernumerary stamens form in positions corresponding to tepal positions in perigonate Araceae. If so, this would be an example of homeosis—in this case, the replacement of tepals with stamens—in the evolution of this genus. We found the positions of stamen primordia in many floral buds too irregular to conclude that they replace tepals positionally. However, in more regular floral buds the first formed stamens do form in what correspond to tepal sites in related genera. If the immediate ancestor to Calla had tepals, as is generally assumed, stamen positions in the more regular flowers, at least, support a homeotic interpretation. There is no evidence that the supernumerary stamens arise by dédoublement, but since morphogenesis in Calla is only partly comparable to other aroids, and the phylogeny in the family is not well understood, further studies are needed to resolve the interpretation of the flower in Calla. With regard to systematics and evolution, the absence of tepals in Calla may not be homologous with atepaly in other members of the family, as has been assumed for the past century.     

Quantitative genetics of stamen number in the selfing Scleranthus annuus (Caryophyllaceae)

Heritability of stamen fertility—different scores were given to sterile stamens developed to different degrees as well as to fertile stamens with one or two pollen sacs—was studied in Scleranthus annuus (Caryophyllaceae), a selling annual that shows extensive phenotypic variation in stamen fertility. Variation within and among 172 maternal families, derived from plants representing 20 natural populations from southern Sweden, was used to estimate heritabilities of stamen fertility for stamens/staminoids at each of the ten stamen positions in the flower. The hierarchical design of the study allowed partitioning of variation at four levels of organization using nested analysis of variance. Heritabilities ranged from 0.631 to 0.714 for stamen positions in the outer whorl of stamens and from 0.235 to 0.555 for positions in the inner whorl. When stamen fertility was pooled across all stamen positions of a flower, the heritability was 0.807. The nested ANOVA indicated that stamen positions in the outer whorl have comparatively higher proportions of among-family and among-population variation than those in the inner whorl. Furthermore, highly significant genetic correlations exist among stamen positions within the inner whorl and among positions within the outer whorl, but not so between positions from each of the two whorls.     

The Australasian grass flora in a global context

Australia's flora and fauna have long been considered unique, but whether this applies to its grasses is less known. This study characterises the Australasian grass flora biogeographically. We investigate the distribution of C₃ and C₄ grass genera across four continents and construct broad profiles of their grass flora. We use endemism to examine global patterns of specialisation, and inter-continental distributions as indicators of dispersal, using databases constructed over twenty years. We examined Australasian patterns with regard to endemicity and shared groups and categorised all of the region's genera into four age classes, from Australia's separation from Gondwana to the present. Globally, each continent presented a unique profile and C₄ grasses were more widely shared than C₃. Australasia's grasses equally comprise C₃ and C₄ genera; it shared two thirds of its C₄ types with other continents, whereas C₃ types split evenly between shared and endemic. Australasia shared relatively few genera with just one neighbour (7% C₃, 13% C₄), primarily with EurAsia. Australasian grass genera and species were either endemic or globally widespread, and 88% of C₃ and 93% of C₄ species were derived from lineages that originated elsewhere. We conclude Southeast Asia was the gateway for dispersal into Australasia, akin to rainforest taxa exchanges which increased from c12 Ma, with about 65% of Australasia's grass genera arriving in the past 3.5 Ma. The strong presence of C₄ grasses in Australasia implies they have infiltrated a wide range of ecosystems, many probably occupied by ancient taxa with which they had not co-evolved.     

CHLORANTHUS-LIKE STAMENS FROM THE UPPER CRETACEOUS OF NEW JERSEY

Fossil angiospermous stamens with in situ pollen from the Turonian (ca. 90 million years before present, Late Cretaceous) of New Jersey are described and assigned to the Chloranthaceae. The fossil stamens, which are three-parted and bear two bisporangiate thecae on the central lobe and one bisporangiate theca on each lateral lobe, are indistinguishable from stamens of several extant species of Chloranthus. The pollen is spheroidal, 13–18 μm in diameter, with a reticulate exine and apparently elongate/elliptical apertures. The pollen is similar to that in extant Chloranthus in grain size, shape, exine sculpture, and aperture structure. Like pollen of some extant species of Chloranthus, aperture number in the fossil pollen appears to be variable. Because fossil pistillate chloranthoid reproductive structures have not been found at this locality it is unknown whether the fossil stamens described here were borne on the side of the ovary, as in extant Chloranthus, or in another arrangement. The three-parted stamen of Chloranthus is unique in angiosperms and there has been considerable debate concerning the origin and evolutionary significance of the structure. Uncertainty as to whether the three-parted stamen represents a synapomorphy for the genus or a retained plesiomorphy in angiosperms is the primary reason why these fossil stamens are not assigned to the extant genus Chloranthus.     

Floral anatomy and systematics of Alliaceae with particular reference to Gilliesia, a presumed insect mimic with strongly zygomorphic flowers

The floral structure of Alliaceae is assessed in relation to the systematics of the family, especially the nature of the component parts of the remarkably insect-like flower of Gilliesia graminea. Both presence of solid styles and possession of tenuinucellate ovules represent consistent synapomorphies for Alliaceae and support the separation of Agapanthus and Themidaceae from Alliaceae. Within Alliaceae, absence of septal nectaries (i.e., complete fusion of carpel margins) is a synapomorphy for the sister genera Gilliesia and Gethyum; septal nectaries are present in all other Alliaceae. A gynobasic style and reduced ovule number are probable synapomorphies for the genus Allium. In contrast to most other Alliaceae, in Gethyum and Gilliesia only three (abaxial) stamens (A1, a1, a2) are expressed, as in the apostasioid orchid Neuwiedia, but the perianth of Gethyum is only slightly bilaterally symmetric (zygomorphic), whereas Gilliesia graminea shows bilateral symmetry in all three floral whorls: perianth (suppression of the inner adaxial tepal in most flowers), androecium (suppression of three adaxial stamens), and gynoecium (slight bilateral symmetry, evident in transverse section). The precise relationships of Miersia and Solaria, the other two genera of Alliaceae with bilaterally symmetric flowers, are unknown, but their morphology indicates a close relationship with Gilliesia and Gethyum. Appendages of tepaline origin occur in Gethyum, Gilliesia, and Miersia; their papillate epidermis suggests that they function as osmophores. Their presence in Miersia, which has six stamens, indicates that these novel structures, which develop late in floral ontogeny, evolved independently from stamen suppression in this group. Within Gilliesia graminea, the genetic mechanisms controlling tepal number and shape are apparently unstable, resulting in fluctuating asymmetry. In G. graminea the possession of insect mimicry, presence of osmophores and absence of nectar together indicate a deceitful pollination mechanism similar to that of some Orchidaceae; this would make Gilliesia highly unusual among non-orchid monocots, given that pollination by sexual deceit is normally regarded as exclusive to orchids.     

The ontogenetic basis for floral diversity in <Emphasis Type="Italic">Agonis</Emphasis>, <Emphasis Type="Italic">Leptospermum</Emphasis> and <Emphasis Type="Italic">Kunzea</Emphasis> (Myrtaceae)

Floral development in three species each of Leptospermum and Kunzea, and one species of Agonis, is described and compared. Differences in the number of stamens and their arrangement in the flower at anthesis are determined by the growth dynamics of the bud. In Leptospermum, early expansion of the bud is predominantly in the axial direction and causes the stamen primordia to be initiated in antepetalous chevrons. In Kunzea, early expansion occurs predominantly in the lateral direction and successive iterations of stamen primordia are inserted alternately at more or less the same level. In both genera, further expansion in the lateral plane spreads the stamens into a ring around the hypanthium. Agonis flexuosa is similar to Leptospermum. Other variable factors include the timing at which stamen initiation commences (earlier in Leptospermum than Kunzea), the duration of stamen initiation (hence the total number of stamens produced – varies within genera), and very late differential expansion that forces stamens into secondary antesepalous groups in A. flexuosa and L. myrsinoides.The authors thank Dr H. Toelken for kindly providing some material and the impetus for this project. This research was supported by Australian Research Council grant AS19131815.