Morphometric analysis of the genusHemerocallis L. (Liliaceae) in Korea

To better understand the patterns of variability and distributions ofHemerocallis in Korea, 53 locations were visited and measurements of 19 morphological and phenological characters were taken on plants directly from their natural habitats. For morphometric analysis, 10 plants from each of 34 populations and five herbarium specimens ofH. middendorffii were used and the data from 12 quantitative characters was analyzed using univariate analysis. Except the littoral populations of Cheju, Hong, Taehuksan, and Sohuksan Islands (H. hongdoensis M. Chung & S. Kang), three peninsular KoreanHemerocallis species can be recognized mainly in South Korea:H. hakuunensis Nakai (=H. micrantha Nakai, growing on southern, central, and northwestern Korea);H. thunbergii Baker (=H. coreana Nakai, found on southeastern and central Korea); andH. middendorffii Tr. et Mey. (central and northeastern Korea). Morphological and phenological features contributing to recognition of the three groups were; color of perianth, shape of roots, shape of inflorescence, flowering time, odor, length of inflorescence, width of the lowest bracts, length of perianth tube enclosing a ovary, width of the inner perianth lobes. Natural hybridization seems to be rare in KoreanHemerocallis. It appears that the KoreanHemerocallis species are relatively well characterized by their distribution patterns, phenology, and habitats compared with the JapaneseHemerocallis species.     

A phenological hypothesis on the thermophilous distribution of Pistacia lentiscus L.

In this study we present evidences of the importance of the phenological pattern on the distribution limits of the dioecious Pistacia lentiscus L. This species, though displaying a thermophilous distribution, has been proved to resist low freezing temperatures during winter. We try to explain this apparent paradox by studying the effects of an extreme cold event that occurred in December 2001 on a natural population of P. lentiscus in the NE of the Iberian Peninsula. In previous phenological studies conduced over 4 years, no spring branch damage was noticeable among population individuals. Female individuals of this species present a prolonged phenophase development which extends until early winter with the conclusion of fruit set. Therefore, we hypothesize that female plants would be more affected than male ones in terms of the following: (1) vegetative and reproductive organs survival; and (2) next-year vegetative growth and fruit production. To test this hypothesis, we selected 225 adult individuals (113 females and 112 males) in April 2002, and estimated their crown volume, percentage of frozen branches and reproductive buds, and the amount of 2001 fruits frozen. In June 2002 we evaluated, in the same individuals, the percentage of vegetative buds flushed into shoots and of reproductive buds producing infrutescences. Branch mortality was significantly higher in female plants and females with increased frost-damage displayed a higher amount of frozen fruits. The loss of reproductive buds caused a decrease in 2002 fruit production, while 2002 vegetative growth was unaffected by the degree of frost damage. These results verify most of the predictions of our hypothesis. Moreover, they suggest that the limited distribution of P. lentiscus in the cold areas of Mediterranean climate could be more determined by the long extent of the phenological activity of the crown than by its frost tolerance during winter.     

Genetics of osteal plate polymorphism and microevolution of threespine stickleback (Gasterosteus aculeatus L.)

Summary The results of genetical studies of osteal plate polymorphism of threespine stickleback Gasterosteus aculeatus reported to date (Munzing 1959; Hagen and Gilbertson 1973; Avise 1976) are discussed and proposed inheritance models are considered. Results of crosses between the morphs of G. aculeatus from the White Sea are presented. An attempt has been made by the author to devise a unifying model for the inheritance of plate morphs within the whole range of G. aculeatus. The role of polymorphism in the microevolutionary processes taking place within this complex species is discussed.     

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).     

Peroxisomes in intestinal and gallbladder epithelial cells of the stickleback,Gasterosteus aculeatus L. (Teleostei)

Summary The occurrence of microbodies in the epithelial cells of the intestine and gallbladder of the stickleback, Gasterosteus aculeatus L., is described. In the intestine the organelles are predominantly located in the apical and perinuclear zone of the cells and may contain small crystalline cores. In gallbladder epithelial cells the microbodies are distributed randomly. The latter organelles are characterized by the presence of large crystalloids. Cytochemical and biochemical experiments show that catalase and D-amino acid oxidase are main matrix components of the microbodies in both the intestinal and gallbladder epithelia. These organelles therefore are considered peroxisomes. In addition, in intestinal mucosa but not in gallbladder epithelium a low activity of palmitoyl CoA oxidase was detected biochemically. Urate oxidase and L- hydroxy acid oxidase activities could not be demonstrated.     

Reproductive lability in pre-dispersal biology ofRhamnus alaternus L. (Rhamnaceae)

Summary Rhamnus alaternus L. is a dioecious, fleshy-fruited shrub, typical of Mediterranean vegetation, which has been recorded from the region since the tropical Tertiary. Reproductive biology of this species has been studied in Southern Italy over a period of 4 years with both field and laboratory experiments. In all the studied populations male plants predominate over the females, have a higher flower density and flower regularly every year while females are usually alternate-bearing. A strong negative association between fruit formation and vegetative growth has been found indicating that the presence of fruits on a branch reduces the growth of new shoots. This phenomenon, together with alternate-bearing, is evidence of a strong competition between vegetative growth and reproductive activity and indicates resource limitation. Flower morphology and quantity of pollen produced are typical of insect pollinated species, and flies of the Empididae and Muscidae were trapped bearing pollen at female plants but pollen is also dispersed by wind. Field experiments on pollination success showed that the exclusion of insects as pollen vectors produces a significant decrease in fruit development. Excess pollen provided by hand-pollination increases fruit set over open-pollination and reproductive success in the field is therefore pollen limited. The variability of sex expression, male fitness, fruit production, regrowth and pollination presented here suggests that these predispersal characteristics ofR. alaternus have been subject to evolutionary change since the tropical Tertiary climate was replaced by a Mediterranean one. This points to the possibility that reproductive, as well as morphological and physiological characteristics, may be equally selected for in extreme environments such as the Mediterranean.     

Seasonal changes in components of male and female reproductive success in Raphanus sativus L. (Brassicaceae)

Summary To document seasonal changes in the reproductive behavior of the perfect-flowered, self-incompatible mustard, Raphanus sativus L., we monitored individual survival, flower and fruit production among 58 individuals in a California population over six census dates (cohorts). Population size declined dramatically and mean individual levels of fruit set changed significantly between cohorts. The frequency distribution of flower and fruit production became increasingly skewed over the first four cohorts. The phenotypic maleness of individuals, a standardized measure of phenotypic gender, oscillated during the reproductive season, peaking in the third and fourth cohorts. We calculated a simple estimate of expected male reproductive success of each plant (the number of fruits sired on conspecifics); this estimate was a function of an individual's flower production and the fruit production of its potential mates in our sampled population. Mean expected male success did not differ significantly among cohorts; expected male success per flower did, however, change significantly among cohorts. Among individuals within each cohort, maternal fruit production and expected male success were both positively correlated with flower production throughout the season. Spearman rank correlation coefficients indicate that the strength of these associations, however, changed during the season. Linear regressions of transformed variables indicated that the shape of several fitness functions also changed over time. In addition, the amount of variation in maternal or expected paternal success explained by flower production declined over the first four cohorts. If typical of wild populations, these temporal changes in these functions suggest that measurements of the intensity of phenotypic selection on flower production will depend on when and how fitness is measured in natural populations.     

Field energetics of free-living, lactating and non-lactating echidnas (Tachyglossus aculeatus)

We measured daily energy expenditure (DEE) and water turnover rates in lactating and non-lactating short beaked echidnas (Tachyglossus aculeatus) using the doubly labelled water technique during the lactation period in spring. Reproductively inactive echidnas were on average significantly heavier (median: 3354 g; range: 2929-3780 g; N=4) than lactating females (median: 2695 g; range: 2690-2715 g; N=3) during the equivalent time period. The median water flux rate of lactating echidnas (152 ml day(-1); range: 120-198 ml day(-1)) did not differ significantly from that of non-lactating females (170 ml day(-1); range: 128-227 ml day(-1)). The median DEE of echidnas that were lactating was 645 kJ day(-1) (range: 581-850 kJ day(-1)), which was not different from the median DEE of non-reproductive control females (763 kJ day(-1); range: 720-766 kJ day(-1)). Lactating females somehow compensate for the energy costs of milk production, resulting in a daily energy budget that is not different from that of non-reproductive females. At least part of their energy minimising strategy could involve the use of moderate heterothermy, allowing a greater proportion of daily energy expenditure to diverted to milk production.     

The pineal complex of the three-spined stickleback,Gasterosteus aculeatus L.

Summary The pineal complex of the three-spined stickleback (Gasterosteus aculeatus L.) was investigated by light and electron microscopy, as well as fluorescence histochemistry for demonstration of catecholamines and indolamines. The pineal complex of the stickleback consists of a pineal organ and a small parapineal organ situated on the left side of the pineal stalk. The pineal organ, including the entire stalk, is comprised mainly of ependymal-type interstitial cells and photoreceptor cells with well-developed outer segments. Both unmyelinated and myelinated nerve fibres are present in the pineal organ. Nerve tracts from the stalk enter the habenular and posterior commissures. A small bundle of nerve fibres connects the parapineal organ and the left habenular body. The presence of indolamines (5-HTP, 5-HT) was demonstrated in cell bodies of both the pineal body and the pineal stalk, and catecholaminergic nerve fibres surround the pineal complex.     

Biosystematic studies inTulipa sect.Eriostemones (Liliaceae)

Evolutionary relationships and infra- and interspecific variability within the species ofTulipa sect.Eriostemones Boissier were studied. Measurements of 35 morphological characters were used for principal component and canonical variate analyses. Crossing experiments and chromosome counts were carried out. Two axes of morphological variation were encountered within the principal component plot. Variation of subsect.Biflores was traced along one axis, the species of subsect.Australes as well asSaxatiles were found along the other axis. The nomenclature of names of subsect.Biflores was considered on the basis of their geography, specific morphological characteristics and polyploidy. Conspecificity of names of all species was tested based on both morphology and crossability.     

Flower development inSpirodela polyrrhiza (Lemnaceae)

Flowering under experimental conditions inSpirodela polyrrhiza, and the development of the spatha, stamina and pistil are described and illustrated by microphotographs. During the development of the microsporangia the existence of the middle layer between endothecium and tapetum has been documented for the first time inLemnaceae.     

Geographic variation in morphological and reproductive characters of coastal and tableland populations ofBlandfordia grandiflora (Liliaceae)

For tenBlandfordia grandiflora populations spanning about 90% of the species' range, univariate and multivariate analyses on 14 vegetative and reproductive characters separated plants into distinct coastal and tableland groups. Distinguishing characters were number of flowers and leaves, leaf length and width, and inflorescence stalk height and diameter; coastal plants were larger than tableland plants. In a transplant experiment, coastal and tableland plants retained their phenotypic distinctness, indicating that vegetative morphology was genetically determined. Coastal plants exhibited clinal variation with latitude. Compared to tableland plants, coastal plants had higher pollen: ovule ratios, and produced fewer but heavier seeds per flower. Tableland and coastal plants are phenotypically distinct, indicating that separate subspecific status is warranted.     

Expression patterns of class I <Emphasis Type="Italic">KNOX</Emphasis> and <Emphasis Type="Italic">YABBY</Emphasis> genes in <Emphasis Type="Italic">Ruscus aculeatus</Emphasis> (Asparagaceae) with implications for phylloclade homology

STM (RaSTM) and YAB2 (RaYAB2) homologues were isolated from Ruscus aculeatus (Asparagaceae, monocots), and their expressions were analyzed by real-time polymerase chain reaction (PCR) to assess hypotheses on the evolutionary origin of the phylloclade in the Asparagaceae. In young shoot buds, RaSTM is expressed in the shoot apex, while RaYAB2 is expressed in the scale leaf subtending the shoot bud. This expression pattern is shared by other angiosperms, suggesting that the expression patterns of RaSTM and RaYAB2 are useful as molecular markers to identify the shoot and leaf, respectively. RaSTM and RaYAB2 are expressed concomitantly in phylloclade primordia. These results suggest that the phylloclade is not homologous to either the shoot or leaf, but that it has a double organ identity.     

Notes on Quercus ilex L. in Liguria (NW Italy)

The climatic characteristics of the Liguria region have been outlined by means of raw data and derived indices (water balance related to potential evapotraspiration, Rivas-Martinez's index of mediterraneity and thermicity, De Martonne's index of dryness, and Emberger's pluviothermic quotient and index of summer dryness). Their interpretation suggests that Liguria is a boundary region between two different climatic areas: the Mediterranean and that of Central Europe.The distribution of Quercus ilex communities in Liguria have suffered the heavy consequences of human activity on the coastal belt of the region. There is, however, sufficient evidence to show that their presence is closely linked to climatic conditions. Quercus ilex communities are absent from the western coastal belt where the annual water balance is below –50 mm and high mean temperatures occur. They are also absent from marly limestone dominated areas where annual water balance is below 300 mm. Quercus ilex woods show a preference for water balance values between 0 and 600, with the better developed forests being found in central and eastern coastal Liguria.The Ligurian Quercus ilex woods are ascribed to Quercetum ilicis Br.-Bl. 1915. Given that the presence within their floristic composition of a group of species of Querco-Fagetea (more species in eastern stands, few in western ones) is an almost constant characteristic, the subass. fraxino-ostryetosum Mariotti 1984 can be retained for many of them. Quercus ilex is not present in the initial succession stages of Mediterranean Liguria vegetation. It can be found in later stages when the vegetation is more fully developed towards a maquis type structure. In this case it tends to become dominant and the community develops (if no fire or other human interference occurs) into a Quercus ilex forest. Frequently, this succession takes place under an old tree layer cover (generally Pinus sp.), a remanant of man's past management of the vegetation in the area.     

Heterochromatin and chromosome variation in cultivated species ofTulipa subg.Leiostemones (Liliaceae)

The chromosomes of several cultivatedTulipa species of subg.Leiostemones were examined in conventionally stained and C-banded preparations. The heterochromatin content varied from almost none to 45%. Several chromosome types were recognized with respect to chromosome morphology and heterochromatin distribution, and groups of species with common chromosome characteristics could be identified. These karyological relationships are discussed with respect to the groups formed on the basis of floral and bulb charateristics.     

Morpho-anatomical features of the leaves of the mediterranean geophyteUrginea maritima (L) baker (Liliaceae)

The morpho-anatomy and histochemistry of the hysteranthous leaf ofUrginea maritima (L.) Baker and its adaptive strategies to the Mediterranean climate were investigated. The leaf ofU. maritima is 714 μm thick and possesses moderate specific leaf mass (8.564 mg cm^-2) and low tissue density (136.5 mg cm^-3). The epidermal cells are compactly arranged and covered with cuticle. The average density of stomata in lower epidermis is higher than that of the upper one. The mesophyll cells occupy 52.96% of the total volume of the leaf, while the mesophyll intercellular spaces and the air spaces occupy 30.41%. Idioblastic cells containing raphide bundles and different phenotypes of crystalloid inclusions, embedded in polysaccharides, occur in the lower side of the mesophyll. The presence of oil droplets and lipids is evident. Bundle sheath cells are hardly visible with no chloroplasts which are a pronounced C₃ plant character. Plastids containing protein crystalloid inclusions are abundant in the protophloem sieve elements.U. maritima, a deciduous plant, possesses leaves with mesophytic characters, in order to optimize its adaptation to the seasonal fluctuation of environmental conditions of the Mediterranean climate.     

Insect and wind pollination inUrginea maritima (Liliaceae)

Three co-existing pollination mechanisms are found inUrginea maritima: insect-, wind-, and self-pollination. The flowers exhibit a typical insect-pollination syndrome; they offer abundant exposed nectar as well as pollen. Out of the many different visitors only a few could be regarded as pollinators:Apis mellifera, Polistes gallicus, andVespa orientalis. Wind pollination also occurs and generally is responsible for self-pollination.It is argued that the development of extra wind-pollination accompanied by partial self-incompatibility is an adaptation to increase pollination in an unfavourable season (August–September), when insects are scarce.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Heterochromatin and chromosome variation in cultivated species ofTulipa subg.Eriostemones (Liliaceae)

The chromosomes of several cultivatedTulipa species belonging to the subg.Eriostemones were examined using conventional staining and C-banding techniques. Most of the species have lightly banded chromosomes with heterochromatin content varying from nil to about 15%. The banding patterns of several taxa are described and discussed in regard to species relationships.