Operculum development in theEudesmieae B eucalypts andEucalyptus caesia (Myrtaceae)

In theEudesmieae B eucalypts and inEucalyptus caesia, the perianth of the mature flower consists of a single, anatomically continuous, opercular structure that is crowned by the tips of the original free and separate calycine and corolline whorls. Ontogenetic and comparative evidence supports the hypothesis that this operculum is mostly corolline in composition, and that the calycine parts have been elevated distally onto the dorsal surface. In theEudesmieae B eucalypts this condition appears to be due to precocious initiation of the corolline primordia, followed by expansion and continuity of their growth centres which incorporates the areas at or below the base of the still differentiating calycine whorl. InEucalyptus caesia the corolline primordia are not precocious, but a similar situation is effected by a seemingly retarded increase in receptacle diameter relative to lateral expansion of the corolline growth centre(s). In these two examples the same final perianth form apparently derives from two different sets of growth processes which, nonetheless, result in the same allometric relationships within the developing flower. Although identical at maturity, the operculum form in these two taxa is best described in terms of convergence, rather than homology.     

Perianth development inAngophora and the bloodwood Eucalypts (Myrtaceae)

The petals ofAngophora flowers are compound structures consisting of two morphologically distinct components that develop along separate morphogenetic pathways. These two components are also evident in the corolline parts of the bloodwood eucalypts. In occasional flowers ofAngophora and some bloodwoods, several adjacent corolline primordia may become continuous due to interprimordial growth, but the petals are mostly free at anthesis. In other bloodwood eucalypt species all the primordia in the corolline whorl become continuous at some stage in development, resulting in an operculum that is anatomically unresolvable into its original petaline parts. The varying degrees of this continuity that are evident within individual trees (and even within single flowers) suggests that operculum formation is an epigenetic event that is determined by morphogenetic processes within the flower. It is suggested that these may relate to differing rates of growth in different regions of the bud.     

Operculum development inEucalyptus clöeziana andEucalyptus informal subg.Monocalyptus (Myrtaceae)

Flowers ofEucalyptus clöeziana have two clearly distinct perianth whorls. The small free parts of the outer (calycine) whorl cease growth early and are lost from the flower; the parts of the inner (corolline) whorl become continuous laterally by confluence of growth centres and form an operculum in the mature flower. The stamens are inserted on a circumfloral buttress (staminophore) that is homologous to the adaxial corolline component inAngophora and the bloodwood andEudesmia eucalypts. Flowers ofMonocalyptus have only one perianth whorl, which is opercular. The stamens are similarly inserted on a circumfloral buttress. Developmental study does not provide conclusive evidence for either a calycine or corolline determination of theMonocalyptus operculum, but comparison with other eucalypt groups, includingE. clöeziana (the sister taxon), predicts an essentially corolline composition.     

Problems of species and genera inEucalyptus (Myrtaceae)

Eucalyptus is recognised as containing somewhat more than 450 species, but this number could reasonably be either decreased or increased considerably by slight variations of species concept. Recognition of taxa, at least in the wild, is less difficult than supposed but their ranking depends on degree and nature of discontinuities or steepened gradients of correlated variation, in this group of diploid sexual outbreeding trees of the dominant stratum. Some 8–10 rather distinct lines marked by different character constellations are discernible inEucalyptus s. lat. (incl.Angophora), and it is clear that the floral operculum is of diverse nature and multiple origin. The advisability of recognising several distinct genera depends on the wideness of polyphylesis in relation to other Myrtaceous genera, and (unfortunately) on practical convenience.Presented at the symposium Speciation and the Species Concept during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Aspects of floral structure and phenology in the genusConophytum (Mesembryanthemaceae)

The complex floral structure in the southern African genusConophytum (Mesembryanthemaceae; 77 spp.) is described in detail and assigned to three basic floral types, two of which can be divided into two subtypes. Correlations between structural features and phenological patterns are demonstrated and discussed in the family context as well as in relation to the systematic subdivision of the genus.     

Evolutionary trends in the androecium of theOrchidaceae

The evolution of the androecium in theOrchidaceae shows three major trends. There is a progressive trend in the degree of fusion of the filament(s) and staminode(s) to the gynoecium. Secondly, there is a reduction in the number of fertile anthers. Finally, there is a progressive change in the position of the base of the anther relative to the apex of the stigma; in the more primitive orchids the apex of the stigma is always higher than the base of the anther (this position is reversed in the higher orchids). All three trends reflect variation in the evolution of pollen dispersal and pollen reception mechanisms in theOrchidaceae. Trends in the evolution of the orchid anther(s) tend to parallel trends in the evolution of their pollinaria.     

Early floral development and androecium organization in Fouquieriaceae (Ericales)

Early floral development with focus on the androecium was studied with the help of scanning electron microscopy and serial microtome sectioning in Fouquieria columnaris and F. splendens. Perianth organs appear in a spiral pattern on the floral apex. The spiral may be a clockwise or anti-clockwise. The androecium is best interpreted as two-whorled with all the stamens arranged in a single series. In F. splendens, two or more of the five epipetalous stamen positions are doubled, i.e. they are occupied by stamen pairs. Unusual features in the floral development of Fouquieriaceae include (1) a strong spiral component even in whorled organ categories and (2) a pronouncedly asymmetric floral apex during an early phase of floral development. From a phylogenetic point of view, it seems plausible that the common ancestor of Fouquieriaceae and its sister family Polemoniaceae was characterized by two alternating, pentamerous stamen-whorls.     

A phylogenetic and allozyme approach to understanding rarity in three “green ash” eucalypts (Myrtaceae)

Phylogenetic relationships among the 12 species of the green ash group of eucalypts were examined using allozyme data, to investigate the causes of rarity in three localized endemics of the group. The relationships suggested by the allozyme data showed both similarities to and differences from those estimated from morphological data byLadiges and coworkers. The phylogenetic relationships suggest that rarity inEucalyptus burgessiana may be related to recent divergence, whileE. paliformis andE. rupicola are relatively old species, more likely to be relicts, and/or restricted to a rare habitat.     

Corolla wilting facilitates delayed autonomous self-pollination in Pedicularis dunniana (Orobanchaceae)

Structural changes associated with corolla wilting may serve as a mechanism for effecting self-pollination. Low pollinator visitation, high seed production and a corolla that persists after anthesis indicates that Pedicularis dunniana is autogamous. Delayed autonomous self-pollination is facilitated by corolla wilting. Wilting of the upper lip (galea) brought the pollen laden anthers into contact with the stigma resulting in the deposition of self pollen on the stigma. The seed set of flowers either emasculated, or with restrained galeae thus preventing anthers brushing against the stigma, was significantly lower than that of open-pollinated flowers. This demonstrates that autogamy occurs in this species through corolla wilting. Germination experiments indicated that outcross seedlings were more vigorous than selfed seedlings as a result of inbreeding depression. It is likely that autogamy provides reproductive assurance for P. dunniana under conditions of pollinator scarcity.     

New species ofBeaufortia andChamaelaucium (Myrtaceae),Drosera (Droseraceae) andPultenaea (Fabaceae) from SW. Australia

Four new species of the generaBeaufortia, Chamaelaucium, Drosera, andPultenaea are described based on the author's collections from SW. Australia. Their affinities are discussed and full illustrations are provided.Dedicated to Hofrat Univ.-Prof.K. H. Rechinger on the occasion of his 80th birthday.     

The effect of crossing distances and population subdivision on floral morphology inScleranthus annuus (Caryophyllaceae), a selfing annual

I used a discontinuous population ofScleranthus annuus (Caryophyllaceae) to study the effect of crossing distances on flower morphology of the progeny. Four types of progeny were produced by artificial selfing, crossing with pollen-donors from the same patch in the population, crossing with pollen from donors from other patches in the population and inter-population crosses. The size of gynoecium parts and 12 sepal characters in this petal-lacking species were significantly influenced by the type of cross and the patch in the population from where the seed-parents originated. All comparisons of progeny types except selfed vs progeny produced by within-patch crosses were significantly separated from each other, while all four seed-parent patches used were significantly separated in a multidimensional space.     

Observations on pollination inSarcostemma (Asclepiadaceae)

Field observations on pollination in New World species of the genusSarcostemma R. Br. sensuHolm are reported. Morphological and anatomical comparisons of pollinated flowers ofSarcostemma subg.Ceramanthus Kuntze (New World) andSarcostemma subg.Sarcostemma (Old World) are presented.     

Floral anatomy ofHypseocharis (Oxalidaceae) with a discussion on its systematic position

The floral anatomy of threeHypseocharis spp. has been studied. The genus resemblesOxalidaceae as well asMonsonia andSarcocaulon of theGeraniaceae. As it is closer toGeraniaceae than toOxalidaceae, it perhaps serves as a connecting link between them.     

Floral morphology and phylogenetic analysis inCrossostylis (Rhizophoraceae)

Floral morphology in all ten species ofCrossostylis, one of the inland genera of Rhizophoraceae and is distributed in the South Pacific Islands, was studied to increase our knowledge on floral features of individual species as well as on relationships among the species. Flowers ofCrossostylis, unlike those of the other Rhizophoraceae, always have semi-inferior ovaries and entire petals, but are diversified concerning the number and arrangement of stamens and carpels, the presence or absence of staminodia, sexuality and the structure of nectaries. Despite some doubt of the presence of apomorphies restricted to the whole genus, we tentatively definedCrossostylis by a combination of the presence of the semi-inferior ovary, entire petals, and arillate seeds, and then performed cladistic analysis on the basis of 24 floral and other morphological characters and withCarallia andGynotroches as outgroups. Our phylogenetic analysis suggested that the species ofCrossostylis are divided into two monophyletic groups: one comprising six species distributed in the Solomon Islands, Vanuatu and the Fiji Islands, and the other comprising four species distributed in New Caledonia and Polynesia.     

Flower and fruit development in the Macaronesian endemicCeballosia fruticosa (syn.Messerschmidia fruticosa,Boraginaceae, Heliotropioideae)

The ontogeny of the flower and the fruit of the Macaronesian endemicCeballosia were investigated morphologically and anatomically by SEM and LM. The fruit does not break into four mericarps, but splits into two two-seeded carpids. Exo- and mesocarp wither after fruit-ripening and the endocarp constitutes the remaining outer wall. Within the stony endocarp tubular parenchymatic isles develop which are linked with the mesocarp. Subsequent disintegration pretends additional locules in the mature fruit. Similar pericarp formations are also found in someHeliotropium species but result from a different ontogeny. Therefore, although a close relationship ofCeballosia toHeliotropium is obvious, the taxon should be treated as a separate genus.Carpological investigations in theHeliotropioideae (Boraginaceae) 2. For part 1 seeHilger (1987).     

A comparative study of the ontogenetic development of the cupules inCastanea andLithocarpus (Fagaceae)

Two contradicting theories have been proposed for the morphological nature of fagaceous cupules; the intercalary growth theory and the higher order dichasial branch theory. All the previous ontogenetic studies insist on the latter one, but the genera investigated have been rather restricted and have not covered all the cupule types. A comparative study of the ontogenetic development of cupules inCastanea crenata andLithocarpus edulis, which are representatives of fundamentally different cupule types, revealed that both the theories are incomplete. InL. edulis, the higher order dichasial branches contribute to cupule formation along the anterior portions of the lateral flowers. However, along the adaxial portion of the central flower, the cupule develops as an intercalary growth, represented by rapid increase of tangentially oblong epidermal cells. InCastanea, intercalary growth is not clearly observable, for presumably, the flowers are surrounded by a well-developed partial inflorescence mound from the beginning of development.Contributions from the Osaka Museum of Natural History No. 305.     

Hair morphology and systematics ofPhysalis (Solanaceae)

Hair morphology was exammed on developing seedlings and mature plants of ten species from five of seven series inPhysalis. The taxonomic importance of hair development and the distribution of hair types then was evaluated with respect to other comparative data. Two classes of hair types were observed, as found in earlier investigations ofSolanum andRhododendron. The first hair class comprises a sequence from unbranched hairs to dendroid-stelliform hairs. Hairs of this class show considerable variability among taxa in size, presence or absence of glandular tips, and presence and degree of branching, and so were useful in making taxonomic comparisons. The second hair class consists of short hairs, each tipped with a multicellular gland. This second hair class was not useful taxonomically because of its lack of variability in morphology and its often rare occurrence on a plant. The taxa studied exhibit distinctive trichome features which serve to distinguish series and, in some cases together with other data, provided new insight into previously enigmatic species relationships.