Life-history strategies affect aphid preference for yellowing leaves

According to the nutrient-translocation hypothesis, yellowing tree leaves are colonized by aphids at the end of the growing season owing to improved availability of nutrients in the phloem sap after chlorophyll degradation. We measured aphid densities on potted Betula pendula seedlings in a field site where a small proportion of foliage rapidly turned yellow before normal autumn coloration as a consequence of root anoxia. The number of adults and nymphs of the birch-feeding specialist aphids Euceraphis betulae, Betulaphis brevipilosa and Callipterinella tuberculata were counted from leaves on each of the 222 plants. Aphids were detected on 19 per cent of green leaves and on 41 per cent of yellow leaves. There was no indication of aphid avoidance of yellow leaves, and the number of winged (alate) viviparous E. betulae adults and their nymphs were significantly higher on yellow leaves than on green leaves, while the numbers of apterous B. brevipilosa and C. tuberculata did not differ between the leaf colour types. Our result suggests that only aphid species with alate generation during colour change can take advantage of yellowing leaves. This may explain the exceptional abundance of E. betulae compared with other aphid species on birches.     

Behavioral interactions between three birch aphid species and Adalia bipunctata larvae

First and fourth instars of three birch aphid species were exposed to first and fourth instars of Adalia bipunctata (L.), the most common aphid predator on silver birch, Betula pendula Roth, in northern California. Defensive behavior differed by aphid species. Euceraphis betulae (Koch) (Eb), the most successful escapee, was highly mobile and frequently walked away from coccinellid larvae. Betulaphis brevipilosa Börner (Bb), a flat, sessile species, was the least successful aphid at actively escaping from A. bipunctata larvae, but could passively escape detection when coccinellid larvae walked over nymphs and did not perceive them. Active escape behavior was much safer for aphids than passive avoidance of detection. Both instars of Eb and fourth instars of Callipterinella calliptera (Hartig) (Cc) escaped from coccinellid larvae more frequently when approached from the front, apparently using vision for pre-contact detection of A. bipunctata. These aphids avoided physical contact with larger predators more often than with smaller predators. Level of predation by A. bipunctata on these three aphid species is dependent upon types of aphid defense.

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Résumé Des larves de premier et troisième stades de 3 espèces de pucerons du bouleau: Betulaphis brevipilosa, Callipterinella calliptera et Euceraphis betulae ont été exposées aux attaques des larves du premier et du quatrième stades d'Adalia bipunctata, prédateur le plus fréquent de pucerons sur Betula pendula en Californie du Nord. Le comportement défensif dépend beaucoup de l'espèce de puceron. E. betulae, qui s'échappe avec le plus de succès, est très mobile et s'écarte fréquemment des larves agressives de coccinelles. B. brevipilosa, espèce plate, sessile, a le moins de succès dans la protection active contre les attaques des larves de A. bipunctata, mais elle peut échapper passivement à la détection des larves de coccinelles qui ne peuvent pas les discerner lorsqu'elles circulent parmi les larves de pucerons. E. betulae (aux deux stades) et C. calliptera (au quatrième) échappent le plus aux larves de coccinelles qui attaquent de front, ce qui laisse supposer que la vision permet de détecter A. bipunctata avant le contact. Le succès des larves de coccinelles varie selon les stades du puceron et de la coccinelle. Les possibilités d'obtention de niveaux de prédation élevés de ces 3 pucerons par A. bipunctata dépend du type de protection du puceron.

     

Fungi associated with the aerial parts of Malaysian mangrove plants

The leaf surface fungi associated with nine species of mangrove plants includingAvicennia alba, A. officinalis, Bruguiera parviflora, Ceriops tagal, Rhizophora apiculata, R. mucronata, Sesuvium portulacastrum, Sonneratia alba, andXylocarpus mollucensis were studied using direct observation techniques and leaf washings. Over 40 fungal taxa were isolated from the leaf washings. Of these, species ofAspergillus, Choanephora, Cladosporium, Curvularia, Fusarium, Nigrospora, Penicillium, Pestalotiopsis, Trichoderma, andZygosporium were frequently encountered in the washings of all nine mangrove plants. Fewer species of fungi includingCladosporium oxysporum, Corynespora cassiicola, Fusarium, Penicillium, Pestalotiopsis, andZygosporium were capable of growth on the washed leaves. The major phylloplane fungus on plants with higher leaf tannin content (e.g.B. parviflora, C. tagal, Rhizophora spp., andX. mollucensis) wasPestalotiopsis. Leaves with relatively lower amounts of tannin supported the proliferation ofFusarium as the major fungus. Fungi were present on the plumule and cotyledonous sheath even before the leaves opened.Pestalotiopsis persisted throughout the development and growth of the leaves. Many of the fungi encountered on senescent leaves have been reported in earlier studies to be the primary colonisers of submerged decaying leaves.     

Diet divergence in two sympatric congeneric butterflies: Community or species level phenomenon?

Summary Two species ofEuphydryas butterflies were studied in California, USA, and showed considerable diet overlap at the species level. They utilize many of the same plant genera for oviposition. However,E. editha is less likely to use woody perennials than isE. chalcedona.Both butterfly species are known to specialize on different host plants in different populations, so species level divergence may not be a good predictor of community level divergence. Within five communities,E. editha andE. chalcedona showed no dietary overlap. A major component of the niche ofE. editha in one community was occupied byE. chalcedona in a second community, even though both butterfly species occupied both communities. These resource use patterns indicate that community level interactions may affect diet divergence. The degree to which divergence within communities is greater (or less) than expected from a species level comparison may be used to provide a measure of community organization. Equations are given in the Appendix for calculating overlap probabilities from presence/absence types of data; in this study, presence is oviposition on a particular plant species, absence is no oviposition on that plant species. Given the various assumptions of the model,E. editha andE. chalcedona show significant community level components of their dietary divergence in the areas studied. However, in some other communitiesE. editha andE. chalcedona do share host plant species. Therefore, we could not demonstrate community level divergence conclusively, nor has this been demonstrated for any other pair of insect herbivore species. We do not know whether this is because the phenomenon is truly rare or just very hard to demonstrate.     

Habitat preferences of four chironomid species associated with aquatic macrophytes in an irrigation reservoir

Habitat preferences of four chironomid species associated with aquatic macrophytes were studied during the vegetation season from April to October at an irrigation reservoir located in Nagoya, Japan.The two smaller sized species,Corynoneura cuspis Tokunaga andPentapedilum tigrinum Hashimoto, were found to be abundant in two floating-leaved plants,Nymphoides indica O. Kuntze andTrapa japonica Flerov. Phytophages,C. cuspis larvae were observed freely moving on the surfaces of leaves and stems within both of these floating-leaved plant communities. Although found within both plant communities,P. tigrinum larvae, as facultative leaf miners, were more concentrated on the surfaces and within the leaves ofN. indica, which have a thin epidermis and thick mesophyll layer.The two larger sized species,Glyptotendipes viridis (Macquart) andPentapedilum sordens (van der Wulp), were found to be living principally within the stems ofN. indica and the petioles ofT. japonica. These parts of the plants, with thin epidermis and many lacunae in the parenchyma, are favorable for miners.     

Karyological and taxonomical notes on three species of the genus<Emphasis Type="Italic">Epipactis (Neottioideae,Orchidaceae)</Emphasis> in the central-western Iberian Peninsula

Studies on the karyotypes and chromosome numbers of species ofEpipactis from the central-western Iberian Peninsula show that the species harbour enormous chromosome variability, have very asymmetric karyotypes and possess extraordinary diversity of aneuploidy. This paper provides the first report of a chromosome number forE. fageticola (2n=36, 40 + 0–2 B), as well as the first counts for Portuguese populations ofE. helleborine (2n=18, 32, 38) and first counts for Iberian populations ofE. tremolsii (n=20, 30, 2n=16, 24, 32, 34, 36, 38 + 1B, 40 + 1B, 52, 60). Among populations ofE. tremolsii there is a significant differentiation in ecology and somatic chromosome number, suggesting that there may be two different taxa in the region studied. Chromosomes are large to small, ranging in length from 10.8 μm to 1.8 μm. Karyotype asymmetry is of type 3C inE. fageticola andE. tremolsii and 2C inE. helleborine andE. tremolsii.     

Allozyme diversity and the evolution ofCrepidiastrum (Compositae) on the Bonin (Ogasawara) Islands

The genusCrepidiastrum is distributed in East Asia and includes 7 species. In the Bonin Islands, three species ofCrepidiastrum occur, and all of them are endemic to the islands. For detecting the origin and speciation of these endemic species, electrophoretic studies have been done in three endemic species of the Bonin Islands as well as in the remaining four species ofCrepidiastrum, andYoungia denticulata which is considered to be closely related toCrepidiastrum. A total of 386 individuals were sampled from 14 populations. As a result, 17 loci of 10 enzyme systems were resolved and gene frequencies for each population were calculated. The genetic variability was low in island species, as reported in some oceanic island plants. Four groups were recognized in the dendrogram generated by the UPGMA method. The Bonin endemics were clustered together, suggesting a monophyletic origin.C. ameristophyllum andC. linguaefolium were found to be genetically very similar, and this may suggest recent and rapid speciation within the islands.     

Ecological differentiation ofEuphorbia lasiocaula andE. sinanensis (Euphorbiaceae) I plant height,phenology and allocation to stems and leaves

Resource allocation and the seasonal change of stem length inEuphorbia lasiocaula Boiss. andE. sinanensis (Hurusawa) T. Kurosawa et H. Ohashi were examined in 10 populations on hills in Miyagi Prefecture, northern Japan. Differences were found in the diameter of stem, stem/leaf ratio of dry weight, vegetative dry weight/leaf area, and the beginning, end and duration of stem growth.Euphorbia lasiocaula has a thicker stem, a larger stem/ leaf ratio, a larger vegetative dry weight/leaf area, a later beginning and end of stem growth and a longer period of stem growth thanE. sinanensis. These differences support the relationships among plant height, resource allocation and phenology predicted by the mathematical models of Givnish (1982) and Sakai (1991, 1994). The tall and thick stems ofE. lasiocaula are considered to be favorable for capturing sunlight in grassy places, causing it to allocate much of its resources to the stems. On the other hand,E. sinanensis is considered to be adapted to deciduous forest floors or forest margins because it completes growing before it is shaded by canopy trees or by tall herbs, which is enabled by the larger allocation of resources to the leaves.     

Low genetic differentiation betweenAdiantum capillusveneris L. andA. ogasawarense Tagawa (Adiantaceae), an endemic pteriodophyte of the bonin (ogasawara) islands

Genetic differentiation was examined betweenAdiantum capillus-veneris L. andA. ogasawarense Tagawa. A total of 136 individuals from two populations ofA. ogasawarense and three populations ofA. capillus-veneris were sampled. Allozymes encoded by 19 putative loci for nine enzyme systems were observed. The populations ofA. ogasawarense are genetically differentiated from those ofA. capillus-veneris; conspecific populations cluster together. However, genetic identity between two species was estimated to be 0.815, and indicates that the degree of allozyme differentiation is lower than those between endemic flowering plants and their progenitors. It is possible that the low genetic differentiation observed betweenA. ogasawarense andA. capillus-veneris results from a recent origin of the endemic species,A. ogasawarense.     

Fungi associated with the aerial parts of Malaysian mangrove plants

The leaf surface fungi associated with nine species of mangrove plants includingAvicennia alba, A. officinalis, Bruguiera parviflora, Ceriops tagal, Rhizophora apiculata, R. mucronata, Sesuvium portulacastrum, Sonneratia alba, andXylocarpus mollucensis were studied using direct observation techniques and leaf washings. Over 40 fungal taxa were isolated from the leaf washings. Of these, species ofAspergillus, Choanephora, Cladosporium, Curvularia, Fusarium, Nigrospora, Penicillium, Pestalotiopsis, Trichoderma, andZygosporium were frequently encountered in the washings of all nine mangrove plants. Fewer species of fungi includingCladosporium oxysporum, Corynespora cassiicola, Fusarium, Penicillium, Pestalotiopsis, andZygosporium were capable of growth on the washed leaves. The major phylloplane fungus on plants with higher leaf tannin content (e.g.B. parviflora, C. tagal, Rhizophora spp., andX. mollucensis) wasPestalotiopsis. Leaves with relatively lower amounts of tannin supported the proliferation ofFusarium as the major fungus. Fungi were present on the plumule and cotyledonous sheath even before the leaves opened.Pestalotiopsis persisted throughout the development and growth of the leaves. Many of the fungi encountered on senescent leaves have been reported in earlier studies to be the primary colonisers of submerged decaying leaves.     

Application of soft X-ray microradiography to observation of cystoliths in the leaves of various higher plants

Soft X-ray microradiography was applied to observation of the cystoliths, calcified bodies of higher plants, in the leaves ofMorus bombycis, Humulus scandens, Ficus elastica, F. retusa (Moraceae),Boehmeria platanifolia, Pilea viridissima (Urticaceae) andMomordica charantia (Cucurbitaceae). It was proved that this technique is useful for examination of the shape, size, distribution and number of cystoliths in fresh leaves. The microradiographs revealed large cigar-shaped cystoliths in the leaf ofP. viridissima, and neighbor-cystoliths in somewhat restricted areas of the leaves ofM. bombycis andH. scandens, and two to seven radially arranged cystoliths in the leaf ofM. charantia. The number of cystoliths per unit area of leaf (nos./cm²) was estimated to be from 1,090 to 3,900 by means of the microradiographs, varying from species to species. The CaCO₃ content of the leaf calculated from the volume and number of cystoliths was approximately 0.4 mg/cm² in all species exceptF. retusa. InF. retusa, it was about 1.06 mg/cm², the highest value among all species tested. Hand-sections of the leaves showed that the lithocysts were localized in the upper and/or lower epidermis, and they were associated with many photosynthetic cells in all species, suggesting some relationship between CaCO₃ deposition in cystoliths and photosynthesis. This paper is dedicated to Professor Kurazo Furuya, Department of Biology, Tokyo Gakugei University, on the occasion of his retirement (1986).     

Taxonomic relationships in East AsianVicia species with unijugate leaves based on random amplified polymorphic DNA markers

Random amplified polymorphic DNA(RAPD) markers were investigated to clarify the taxonomic positions ofVicia linearifolia andV. bifolia, and to assess the genomic diversity among the 9 populations ofV. unijuga, each of which represents a geographical variation or infraspecific taxa in southern Korea. These species are characterized by unijugate leaves in East Asia and have been controversial as to infra-or interspecific classification. The polymorphic markers among the populations examined were observed for fifteen decamer primers. The degree of band sharing was used to calculate genetic similarity between populations, and a phenogram using UPGMA cluster analysis was generated based on the Dice similarity coefficient. The taxa studied were divided into two main groups and the populations ofV. unijuga were all grouped together in the phenogram. The genetic similarities ofV. unijuga were very high among the populations and did not show distinctions between the infraspecific taxa, although the populations of Mt. Odae and adjacent areas in eastern Korea were different from others of the species.V. linearifolia fell within the range of the genomic variation among the populations ofV. unijuga, whileV. bifolia was grouped withV. venosa var.cuspidata having multijugate leaves rather thanV. unijuga. The result from studying RAPD markers suggested thatV. linearifolia should be integrated intoV. unijuga and that species with unijugate leaves ofV. bifolia andV. unijuga are polyphyletic.     

COMPARATIVE PHOTOSYNTHESIS OF TWO SPECIES OF INTERTIDAL EPIPHYTIC MACROALGAE ON MANGROVE ROOTS DURING SUBMERSION AND EMERSION

Photosynthesis and dark respiration rates were measured in water and in air, and the capacity to recover photosynthetic activity from emersion stress was examined for two species of intertidal, epiphytic macroalgae—Bostrychia calliptera (Montagne) Montagne and Caloglossa leprieurii (Montagne) J. Agardh—collected on prop roots of the red mangrove Rhizophora mangle L. in Buenaventura Bay, Pacific coast of Colombia. In both species, net photosynthetic rates were significantly higher under submersed conditions. Maximum photosynthetic rates (P_max) in water and in air were highest in B. calliptera, 126 ± 4 versus 52 ± 9 μmol O₂·mg chl a⁻¹·h⁻¹, respectively. In C. leprieurii, P_max of submerged plants in water and in air were 98 ± 9 versus 30 ± 11 μmol O₂·mg chla⁻¹·h⁻¹. The photoinhibition model of Platt et al. (1980) was used to fit the experimental data in both water and air for both species. Photoinhibition occurred at irradiance as low as 200 μmol·m⁻²·s⁻¹. The photosynthesis–light response curves demonstrated an adaptation to shaded habitats for both species, as light compensation points in water and air for both species were below 17 ± 5 μmol·m⁻²·s⁻¹. The rate of dehydration was significantly lower in thalli of B. calliptera compared to C. leprieurii. An increase of photosynthetic activity in B. calliptera was evident between 5% and 15% water loss, but rates decreased thereafter with declining water content. In C. leprieurii, desiccation negatively influenced photosynthetic rates that significantly decreased linearly with declining water content. In B. calliptera, net photosynthesis reached zero only at a water content between 29% and 35%, whereas in C. leprieurii no net photosynthesis occurred in plants containing less than about 50% of their relative water content. Resubmerged plants ofB. calliptera exhibited 100% photosynthetic recovery after 45 min, whereas C. leprieurii recovered 100% at about 120 min. On the basis of the comparison of rates of light-saturated net photosynthesis for B. calliptera in air versus in water, aerial photosynthetic activity ranged from 35% to 42% of that in water, whereas the emersed photosynthetic capacity of C. leprieurii ranged from 24% to 29% of that in water. Using tidal predictions and the emersed photosynthetic rates, a carbon balance model was constructed for both species over a single daylight period. The calculations indicated that emersed photosynthesis increased average daily carbon production of B. calliptera by 17% and C. leprieuri by 12%. The physiological responses to desiccation stress and the photosynthetic recovery capacities between species correlated with, and may determine, their vertical distribution in the mangrove habitats of Buenaventura Bay.     

Epizootiology of virus diseases in three lepidopterous insect species of alfalfa

Summary The incidence of virus infections in three lepidopterous insect species was studied from 1965 to 1968 in alfalfa fields in California. The insects were the alfalfa caterpillar,Colias eurytheme; the beet armyworm,Spodoptera exigua; and the alfalfa looper,Autographa californica. InC. eurytheme, the major virus was a nuclear polyhedrosis virus (NPV); inS. exigua, a granulosis virus (GV) and an NPV; inA. californica, a GV. Virus epizootics did not develop in very high densities ofC. eurytheme. Virus epizootics occurred in low host densities of the three insect species, especially in populations ofA. californica. The virus acted as a density-dependent factor in the regulation of the populations ofS. exigua andA. californica. Temperature, humidity and rainfall had no marked effect on the incidence of virus infections.     

Distribution and sporulation phenology of myxomycetes in the Sonoran Desert of Arizona

All pith samples from 68 dead saguaro cacti in 3 plots and 11 isolated dead plants in Saguaro National Monument, Arizona, produced at least one species of myxomycete upon incubation at 20 or 30°C. Three species,Badhamia gracilis (Macbr.) Macbr.,Physarum straminipes Lister, andDidymium eremophilum M. Blackwell et Gilbertson, developed at high frequencies on the substrates in moist chamber culture.Perichaena corticalis (Batsch) Rost, andProtophysarum phloiogenum M. Blackwell et Alexopoulos were also present. Although previous literature reports [9] indicated that Myxomycetes grow best at low pH, these species all tolerated substrates of pH 8.7–10.4.Didymium eremophilum andP. phloiogenum had peaks in sporulation within 6 days; other species were slower. There was no difference in time of sporulation ofB. gracilis orD. eremophilum at 20 and 30°C; however, sporulation ofP. straminipes was significantly later at 30°C. Reduced spore germination and slower buildup of critically sized amoebal populations ofP. straminipes at 30°C may be a factor.     

Responses of four arid zone grass species from varying habitats to drought stress

The effects of 4 or 8 drought cycles on four grass species,Cenchrus pennisetiformis, Leptochloa fusca, Panicum turgidum, andPennisetum divisum were assessed in a pot experiment. There were significant differences between the species in biomass production under water stress.C. pennisetiformis andP. turgidum produced significantly greater fresh and dry matter thanP. divisum and especially thanL. fusca. L. fusca had the lowest andP. divisum highest osmotic potentials compared with the other species after the completion of 4 or 8 drought cycles. Osmotic adjustment (difference between osmotic potential of droughted/rehydrated plants and control plants) was highest inL. fusca. The stomatal conductance was significantly decreased with increased drought stress inC. pennisetiformis. The elasticity ofC. pennisetiformis, P. turgidum andP. divisum increased with increase in number of drought cycles, whereas that ofL. fusca remained unchanged.L. fusca andP. turgidum had the lowest leaf hydration of all species after 8 drought cycles. The chlorophyllsa andb in all species remained unaffected by drought treatments. The proline content ofC. pennisetiformis andL. fusca increased significantly with increased drought stress, whereas that ofP. turgidum remained unaffected after 4 or 8 drought cycles.L. fusca synthesized great amount of leaf soluble proteins during 8 drought cycles, whereasP. divisum had low protein content after 4 drought cycles. The protein contents ofC. pennisetiformis andP. turgidum remained unaffected after 8 drought cycles. The leaf epicuticular wax ofL. fusca increased consistently with increased drought stress, but leaf wax ofP. divisum increased only at the highest drought stress and that ofC. pennisetiformis andP. turgidum increased after 4 drought cycles. On the basis of these results it was established thatC. pennisetiformis andP. turgidum were the most tolerant,P. divisum intermediate, andL. fusca the most sensitive to drought stress. The osmotic adjustment did not positively correlate with the degree of drought resistance.     

A biotechnology strategy for medium- and long-term conservation of cycads

It has been possible to regenerate a few cycad species in vitro by somatic embryogenesis, either from zygotic embryos (Ceratozamia hildae, C. mexicana, Encephalartos cycadifolius,E. dyerianus, E. natalensis, Zamia fischeri, Z. furfuracea, andZ. pumila) or from leaves of mature phase trees (C. euryphyllidia, Ceratozamia hildae, andC. mexicana). This strategy has great potential for the commercial vegetative propagation of certain highly endangered species (e.g.,C. euryphyllidia) and should indirectly protect wild populations of these species by discouraging collection in situ. Embryogenic cultures of several cycad species have grown vigorously and are highly morphogenic more than 11 years after induction. The long-term conservation of cycad genetic resources can also be addressed for species that can be regenerated by somatic embryogenesis. Preliminary studies indicate that embryogenic cultures that have been pretreated on plant growth medium containing 0.75 M sucrose for two days, encapsulated in sodium alginate, and desiccated for six hours can survive immersion in liquid nitrogen (−196°C).     

A survey of the socially parasitic ant generaEpimyrma Emery, 1915 andChalepoxenus Menozzi, 1922 in Italy (Hymenoptera,Formicidae, Myrmicinae)

Summary An up-to-date synthesis is presented of the available faunistic and biological data concerning the Italian species belonging to the socially parasitic ant generaEpimyrma Emery andChalepoxenus Menozzi.The first known populations ofE. corsica (Emery) andE. stumperi (Kutter) in Italy were discovered in the Lucretili mountains (Latium) and in Alto Adige, the former species being previously recorded only in Corsica and Dalmatia (Yugoslavia), the latter in the French and Swiss Alps.The first records ofE. ravouxi (André),E. kraussei (Emery) andC. muellerianus (Finzi) in central Italy are presented. Since no species of either genera has ever been collected in Italy in the area between the Po valley and Calabria, these new records are of great interest.Maps, showing the currently known distribution of each taxon in Italy, are provided.     

Shoot structure and dynamics of saplings and canopies of three deciduous broad-leaved trees of a coppice forest in central Japan

Three deciduous broad-leaved trees, Quercus serrata, Castanea crenata and Carpinus laxiflora, were the main constituents of a coppice forest in central Japan. The shoot elongation and leaf emergence modes of both saplings and the canopy of the three species were investigated. The shoot elongation modes of Q. serrata and C. crenata were the same in saplings and the upper layer of the canopy. The second shoots of these two species were formed after the first shoots were elongated. C. laxiflora was different between saplings and the upper layer of the canopy. In saplings, only the first shoots took a long time to elongate. In the upper canopy layer, higher order shoots were formed in the same way as in the other two species. In the lower layer of the canopy, all three species showed the same shoot elongation mode, in which only the first shoot and its duration of elongation was short. Leaf longevity, individual leaf area, leaf mass per unit leaf area and the stem mass per unit stem length of C. laxiflora were significantly shorter or significantly smaller than those of Q. serrata and C. crenata. The length of the stem per unit leaf area of C. laxiflora was three times that of Q. serrata and five times that of C. crenata. The elongation growth of C. laxiflora was highly efficient as it occurred with a small leaf area. The shoot dynamics and the shoot structure of C. laxiflora are more suitable for elongation growth than in Q. serrata and C. crenata. Furthermore, the shoot structures of the three species were compared and ecological characteristics of the three species are discussed. Received: 29 September 1998 / Accepted: 17 September 1999