Floral development and systematics ofCistaceae

The floral development of representatives of six genera ofCistaceae has been studied. Calyx development involves the formation of a ring primordium in several taxa. Androecium development in species with intermediate or higher stamen numbers starts with the formation of a ring meristem on which the stamens are initiated in a centrifugal direction. In many taxa five alternipetalous leading stamen primordia can be observed. In the apetalous (cleistogamous) flowers ofTuberaria inconspicua androecium development appears to be unordered; this is probably due to the lack of petals. InLechea intermedia (also cleistogamous) the corolla is trimerous and three complex stamen primordia are produced, which give rise either to one or three stamens. Relationships withinCistaceae are discussed. Floral development inCistaceae is compared with that in otherMalvanae. Among the eight families ofMalvanae from which information on floral development is availableCochlospermaceae andBixaceae exhibit the greatest similarities toCistaceae. InCistaceae the leading stamen primordia are alternipetalous. InBixa the same condition seems to be present. InMalvales s. str. mostTiliaceae also show earliest stamen initiation in alternipetalous sectors, whereas the stamens of the innermost alternipetalous position are retarded early or even suppressed inSterculiaceae, Bombacaceae, andMalvaceae. WithinMalvales s. str. the diversity of androecial developmental patterns seems to decrease inBombacaceae andMalvaceae due to increasing synorganization in the mature androecium. The derivation of polyandry inMalvanae from diplo- or obdiplostemony is discussed by comparison with the sister clades ofMalvanae as shown in recentrbcL studies (i.e.Sapindales, Rutales, the glucosinolate producing clade, andMyrtales).     

Ovule and seed ontogeny inGnetum gnemon L.

The ovule ofGnetum gnemon has three envelopes around the nucellus. The outer one forms two clear swellings at the lateral sides during the early developmental stages. The middle envelope also shows two swellings in many cases arranged decussately with respect to those of the outer one. All these swellings become obscure or disappear later. The inner envelope arises as an annular primordium and forms several manifest lobes at the stage of pollination. it develops two proliferating structures from its middle portion, viz., a flange and a micropyle-closing tissue. The three envelopes differentiatiate into the fleshy outer, sclerenchymatic middle and compressed inner layers of the seed coat, respectively. The inner one, however, remains restricted to the apical part of the seed owing to endochalazal growth. The outer envelope is derived from both dermal and subdermal cells of the ovule primordium and, therefore, is of dual origin. The middle and inner envelopes are subdermal in origin. The present study has cleared up some conflicting reports in the previous publications.     

Seed and fruit anatomy ofCocculus hirsutus (Menispermaceae)

The ovules ofCocculus hirsutus are anatropous, bitegmic and, crassinucellate. The fruit is drupaceous, black purple, and laterally compressed and has a pericarp demarcated into exocarp and endocarp. The seeds are curved around the basal bodies. Testa and tegmen cells are thin-walled and unspecialized. In the ripe seed the inner epidermis of the tegmen persists, whereas the entire testa and the outer layer of the tegmen degenerate. Relationships of theMenispermaceae toRanunculaceae, Berberidaceae, andLardizabalaceae are supported.     

Seed morphology and anatomy in someAnnonaceae

Twelve species ofAnnonaceae, namelyAncana sp.,A. stenopetala, Annona cherimola, A. montana, A. muricata, A. squamosa, Bocagea sp.,Bocageopsis canescens, two species ofUnonopsis, Xylopia aromatica, andX. emarginata, were investigated with respect to the morphology and anatomy of the seed. They show the basic structural pattern characteristic of annonaceous seeds: perichalaza, fibrous mesotesta, and rumination developed by both integuments. However, several differential characters, some of them never previously described, were found: All taxa exhibit an additional endotestal mechanical layer which forms the micropylar plug except inAncana. InAncana the plug is built by the inner integument. Aril, sarcotesta and pits on the seed surface are present in some taxa and show different origin and structure. Special anatomical adaptations possibly functioning during germination are described. The results obtained from the investigated taxa are discussed and compared with published data on seed structure inAnnonaceae.     

Amino acids,polyamines and proteins during seed germination of two species of Dipterocarpaceae

Summary Amino acid, polyamine and protein concentrations in seeds and their evolution during seed germination of two dipterocarp species, Hopea odorata and Dipterocarpus alatus, were determined with the help of a multianalytical system. Glutamic acid and glutamine were the major amino compounds present. Hopea seeds also contain high levels of aspartic acid/asparagine, serine, threonine, arginine and alanine, while those of Dipterocarpus contain high levels of alanine, arginine and threonine. These species were quite different in their germination behavior and thus in their protein and amine metabolism rates. In Hopea, polyamines increased during the first 3 days of germination and reached a maximum by the 3rd day, 1 day before maximum germination rate. In Dipterocarpus polyamines reached their maximum at the 6th day while maximum germination rate is observed by the 7th day. This suggests that polyamine compounds could play a role in the early part of the germination process in Hopea and Dipterocarpus seeds. The possibility that control of polyamine biosynthesis could be used for the establishment of biochemical methods to improve seed storage and to control germination of these recalcitrant seeds is discussed.     

Comparative wood anatomy ofCoriaria

Wood anatomy ofCoriaria was surveyed to clarify generic features on the basis of 14 species collected from various regions of the World to cover the whole range of geographic distribution and habitual variation. Wood anatomy ofcoriaria is considerably uniform, and the species share a combination of the following features: 1) pores are thin-walled, polygonal in outline and mostly in multiples; 2) vessel elements and libriform fibers are very short; 3) perforation plates are exclusively simple; 4) intervessel pits are alternate; 5) vascular tracheids are present; 6) wood parenchyma is vasicentric and sometimes confluent; 7) rays are heterogeneous and large. Its species differ in several characters, such as distinctness of growth rings, pore size, pore patterns, type and abundance of wood parenchyma, and distinctness of storied structure. Comparisons among species indicate that the species of the Northern Hemisphere show a tendency toward having semi-ring porosity, while those of the Southern and Western Hemisphere have diffuse porosity. The other infrageneric variations appear to be related to different habits of the species rather than to geographic distribution. Small trees mostly have confluent and vasicentric parenchyma composed of fusiform cells and distinctly storied tissues, while shrubs and herbs have less abundant parenchyma which is vasicentric and comprises strands of two to four cells and indistinctly storied tissues.     

Floral anatomy and systematic position ofVivianiaceae

The floral anatomy of four species ofViviania has been studied. In the basic floral plan and essential floral anatomical featuresViviana closely resembles theGeraniaceae. Evidence from vegetative and floral anatomy, ultrastructural studies on phloem as well as phytochemistry supports geranialean affinity ofViviania; the placement within thePittosporales sensuHutchinson being unnatural.     

Masting,seed dispersal and seed predation in the cycad Macrozamia communis

Summary Little is known about the adaptive value of mast seeding, a common phenomenon in temperate trees and shrubs. Masting is likely to affect both seed dispersal and seed predation. In systems where similar taxa of animals are involved in these two processes, the consequences of mast seeding are likely to be particularly complicated. This study examined the effects of mast seeding in a cycad, Macrozamia communis, on the dispersal of seeds, the pattern of dispersion of seeds and post-dispersal predation on seeds. Dispersal of seeds by possums was poorer from source plants in a masting population than from source plants in an adjacent, non-masting population. This resulted in fewer seeds per seeding female plant in the masting plot being dispersed to favourable sites. We conclude that this is caused by the feeding behaviour and movements of possums in the masting site. The abundance of seeds in this site did not satiate the post-dispersal predators, native rats. In fact, more seeds in this site were eaten than in the nonmasting site. We suggest that the mast seeding observed in M. communis may not be adaptive, but is more likely a consequence of other factors which synchromize flowering within local populations.     

Ribosomal RNA integrity and rate of seed germination

The integrity of ribosomal RNA (the percentage of complete, un-nicked molecules) in seeds was studied by electrophoresis under denaturing conditions. Two batches of carrot seed, harvested at different stages of maturity, and four batches ofNicotiana seed stored for various times were used. Within each species, there was a correlation between the integrity of the rRNA of the dry seed and the rate of germination of that seed. In carrot seed, there was extensive degradation of existing rRNA in both the embryo and endosperm during the first two days of imbibition.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Regeneration of Cyperaceae, with particular reference to seed ecology and seed banks

Cyperaceae (sedges) are an important component of many ecosystems. To understand better their regeneration, we examined seed ecology, including dispersal, seed characteristics, and germination behavior that relate to seed bank development and persistence. We also evaluated sedge seed banks from 104 studies, representing a wide array of habitats. Sedge seed bank development and persistence were associated with germination and dormancy traits, namely: dormancy level, seasonal dormancy patterns, and requirement for light, alternating temperatures, and aerobic conditions. Interplay of traits appears to have resulted in low-risk germination strategies adapted to exploit infrequent occurrence of gaps and facilitate formation of persistent seed banks. A variety of dispersal modes and morphological adaptations occurred, but many species had no apparent specialized structures. The main dispersal vectors were water and then animals. About 216 species, in 21 genera, were recorded in the seed bank survey. High densities (>50,000 m⁻²) occurred occasionally in wet habitats, but generally values were low (<500 m⁻² in 70% of entries). Species richness was also generally low (mean 4.8 species study⁻¹), but ranged from 10 to 33 species in certain wetlands. Our studies showed varied reproductive strategies within habitats, persistence, and ability of many species to colonize disturbed habitats. Overall, seed banks tended to be persistent (>1 year). Maximum longevity ranged between 10 and 295 years, but for certain species viability was lost in <3 years. Seeds of many sedges occurred in deeper soil layers to depths >1 m. Seed production, low in rhizomatous species, ranged between 0 and 345,000 seeds m⁻² year⁻¹. Amphicarpy or pseudo-viviparous plantlets occurred in limited numbers of species. The relation between seed production, seed rain, and seed bank is largely obscure and awaits further investigation. For successful restoration and species conservation projects, seed banks (or a source of seeds) are necessary, combined with suitable germination and establishment conditions. Future seed bank studies are considered.     

Accumulation of seed storage proteins and the taxonomy ofPoaceae

The accumulation of specific seed proteins is a taxonomically valuable feature and can be used to additionally characterize plant taxa. To date, mainly crop proteins have been analysed in thePoaceae. In this investigation seed proteins from 147 species were screened with emphasis on legumin-like proteins and prolamins. The groups resulting from evaluation of the protein profiles correspond with well-known subfamilies and tribes.Panicoideae are clearly separated fromPooideae. WithinPooideae, theBromeae plusTriticeae tribes revealed obvious similarities.Lolium, Festuca andVulpia, generally included in the tribeFestuceae, revealed a protein profile similar to the profile of theBromeae/Triticeae. Legumin-like proteins are accumulated abundantly inBambusoideae andPooideae exceptBromeae/Triticeae, however, only the species included in theAveninae subtribe produce soluble (globulin-type) legumins as already known fromAvena sativa. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Morphological studies on the genusClematis linn. VIII. Floral and inflorescence anatomy inClematis patens with eight-sepaled flowers

The vascular anatomy of inflorescence axes and flowers ofClematis patens have been studied. The species shows a unique behaviour of the vascular bundles in the transition node from vegetative stem to pedicel: stelar bundles increase in number from six to eight as they ascend through the transition node so that the number of vascular bundles coincides with that of sepals. In the pedicel stele the resulting eight bundles are disposed opposite to eight sepals. respectively; each sepal receives its vascular supply from the bundle facing it. Morphological and anatomical evidence suggests that the calyx of eight sepals in this species should be interpreted as having consisted originally of four pairs of opposite organs, rather than as having been derived secondarily through chorisis of sepals from a calyx of four sepals as seen in most other species ofClematis.     

Seed coat anatomy ofCrossostylis (Rhizophoraceae): its evolutionary and systematic implications

The seed coat structure of all 13 species ofCrossostylis was studied to contribute to an understanding of species delimitation and relationships within the genus. The mature seed coat is relatively uniform and consistently constructed mainly by a well-developed exotesta and a well-developed fibrous exotegmen. The species differ in the thickness of the exotesta and exotegmen, the anatomy of exotestal cells, the presence and absence of persistent mesotesta, and so forth. On the basis of comparisons of these characters, close relationships are suggested in the species groups such as:Crossostylis banksiana andC. cominsii; C. biflora, C. raiateensis andC. multiflora; C. gandiflora, C. sebertii andC. imera; and five species in the Fiji Islands. These relationships except for those of Fijian five species are also supported by cladistics as their common characters are evaluated as synapomorphy. Species-level separation ofC. banksiana, C. pedunculata andC. raiateensis each from the closest species is doubted based on the results of seed coat structure.     

Chronic herbivory negatively impacts cone and seed production,seed quality and seedling growth of susceptible pinyon pines

Although herbivory often reduces the reproduction of attacked trees, few studies have examined how naturally occurring insect-resistant and susceptible trees differ in their reproduction, nor have these effects been experimentally examined through long-term herbivore removals. In addition, few studies have examined the effects of herbivory on the quality of seeds produced and the implications of reduced seed quality on seedling establishment. We evaluated the impact of chronic herbivory by the stem-boring moth, Dioryctria albovittella, on cone and seed production of the pinyon pine (Pinus edulis) during two mast years. Three patterns emerged. First, moth herbivory was associated with reductions in cone production, viable seed production and seed mass. Specifically, pinyons susceptible to moth attack had 93–95% lower cone production, and surviving cones produced 31–37% fewer viable seeds, resulting in a 96–97% reduction in whole tree viable seed production. In addition, surviving seeds from susceptible trees had 18% lower mass than resistant trees. Second, long-term experimental removal of the herbivore resulted in increased rates of cone and seed production and quality, indicating that moth herbivory was the driver of these reductions. Third, seed size was positively associated with seed germination and seedling biomass and height, suggesting that trees suffering chronic herbivory produce poorer quality offspring. Thus, the resistance traits of pinyons can affect the quality of offspring, which in turn may affect subsequent seedling establishment and population dynamics.     

Wood anatomy ofTrimeniaceae

Four collections of three species ofTrimenia and one collection ofPiptocalyx were studied; early-formed and later-formed wood was analyzed for oneTrimenia. Liquid-preserved material permitted analysis of mucilage and starch storage in wood ofT. neocaledonica andP. moorei. BecausePiptocalyx is scandent whereasTrimenia is arborescent, wood differences relative to evolution of a climbing habit could be examined.Piptocalyx contrasts withTrimenia in having wider vessels, more numerous per mm², resulting in a conductive area five times greater per unit area than that of theTrimenia woods averaged.Piptocalyx has appreciably fewer bars per perforation plate and thus much greater conductive area per perforation plate than have the species ofTrimenia. Rays inPiptocalyx are much taller and wider than those ofTrimenia. Wood ofTrimeniaceae is highly primitive in its scalariform perforation plates, scalariform lateral wall pitting on vessels, relatively long vessels elements, and heterocellular rays. Imperforate tracheary elements are septate nucleate fibertracheids (or even libriform fibers) rather than tracheids, but loss of borders on pits (and thus lowered conductive function of the imperforate tracheary elements) can be explained by the development of these elements into starchstoring cells. Some fiber-tracheids inT. neocaledonica are enlarged mucilagecontaining cells. Details of vessel structure inTrimeniaceae are similar to those ofMonimiaceae (s. s.), but similarity to some other lauralean (annonalean) families may be found: in mucilage presence,Trimeniaceae resembleLauraceae rather thanMonimiaceae. Wood ofTrimeniaceae may be regarded as highly mesomorphic, corresponding to the moist habitats in which all of the species occur.     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

Studies in the floral morphology and anatomy of nymphaeales

Floral morphology ofBrasenia schreberi Gmel. andCabomba caroliniana A. Gray was observed chiefly from an anatomical point of view. The receptacle ofB. schreberi is rather flat and a vascular plexus is observable in the mature flower. The vasculature in this plexus is so complex taht it is not easy to trace its structure in detail. by observation on small buds, it can be seen that the receptacular vasculature consists of a girdling bundle in the basal area and usually nine receptacular strands from which traces to the petals and stamens branch off. The vasculature in the receptacle is reconstructed and diagramatically shown as though split longitudinally and spread out in one plane. Floral vasculature inCabomba caroliniana is simpler, and is probably related to the smaller number of stamens and carpels. It also has a girdling bundle at the bottom of receptacle and this vasculature is suggested to be derived by simplification from aBrasenia-type vasculature. Evidence from floral anatomy suggests that these two genera are closely related. InNymphaea, a vascular plexus in the receptacle is also observed (Moseley, 1961; Ito 1983). The plexus ofBrasenia andNymphaea are not the same in their construction. Nevertheless, their fundamental floral vasculature is comparable and it is preferable to place them in the same family or same order.     

Wood anatomy of theSarraceniaceae; ecological and evolutionary implications

The wood of theSarraceniaceae has a considerable number of primitive features including scalariform perforation plates, long and oblique end walls, scalariform lateral wall pitting, solitary vessels, tracheids with scalariform pitting, and diffuse axial parenchyma. Vessel elements in the genusHeliamphora have the greatest number of primitive features, while vessel elements inDarlingtonia andSarracenia appear to have modifications relating to temperate climates. The wood anatomy suggests thatHeliamphora is growing in a habitat more similar to the original habitat for the family thanDarlingtonia andSarracenia. The wood of theSarraceniaceae is similar to the wood of theTheales.     

Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.