A Nutritional Interpretation of Sclerophylly Based on Differences in the Chemical Composition of Sclerophyllous and Mesophytic Leaves

On the assumption that some of the characteristics of sclerophyllousleaves may have a nutritional basis, data on the leaf compositionof nineteen species from evergreen bushland in Jamaica are comparedwith published data on the leaf composition of thirty-one speciesof mesophytes from Central America. The data show that in termsof dry matter a sclerophyllous leaf is associated with a decreasein percentage protein content rather than with an increase inpercentage fibre content. There is a highly significant positivecorrelation between the protein and phosphorus contents of theleaves considered, and it is suggested that the ability to toleratelow phosphate may characterize sclerophyllous vegetation ingeneral.     

Relationships between sclerophylly,leaf biomechanical properties and leaf anatomy in some Australian heath and forest species

ABSTRACT

A previous study of 19 south-east Australian heath and forest species with a range of leaf textures showed that they varied considerably in leaf biomechanical properties. By using an index of sclerophylly derived from botanists' rankings (botanists' sclerophylly index, BSI) we determined that leaves considered by botanists to be sclerophyllous generally had both high strength and work to fracture (particularly in punching and tearing tests), both at the level of leaf and per unit leaf thickness. In the current study we have shown that leaves from the same species also varied considerably in leaf specific mass (46–251 g m^-2), neutral detergent fibre concentration (20–59% on a dry weight basis) and in leaf anatomy. Multiple regression indicated a very strong correlation between BSI and the first two components of a principal components analysis (PCA) of leaf anatomy (R ² = 0.91). In addition, there was strong correlation between the first component of a PCA of the mechanical properties (correlated with BSI) and the two axes derived from anatomical characteristics (R ² = 0.66). The anatomical properties contributing most to the significant component axes were thickness of palisade mesophyll and upper cuticle (axis 1) and percentage fibre (neutral detergent fibre) and lower epidermis thickness (axis 2). However, whether these relationships are causal, or reflect correlations with characteristics not measured in this study, such as vascularization and sclerification, is not clear. At a finer scale, however, there is evidence that there are various ways to be sclerophyllous, both in terms of anatomical and mechanical properties. This is illustrated by comparison of two of the sclerophyllous species, Eucalyptus baxteri and Banksia marginata.     

Sclerophylly: Evolutionary advantage or mere epiphenomenon?

ABSTRACT

The possible functional significance of sclerophylly (hard and coriaceous leaves) is discussed on the basis of different interpretations reported in old and recent literature. These, have been schematically divided into four groups of hypotheses, i.e. the “water conservation?d hypothesis, the “resistance to negative turgour pressures?d hypothesis, the “nutritional?d hypothesis and the “protective?d hypothesis. In particular, a section is devoted to discussing the first and second hypotheses in terms of relationships of sclerophylly to drought resistance strategies. Data reported in the literature lead to the conclusion that the sclerophyllous habitus per se seems not to be related to drought resistance of plants as previously thought. Nor do clear-cut relationships appear to exist between leaf sclerification and living leaf cells with thicker walls, leading to a high modulus of elasticity (ε), the latter known to be of importance for generating low leaf water potentials, i.e. higher driving forces for water uptake. Because leaf sclerification has proved to be related to soil nutrient deficiency, this leaf habitus can be considered as a mere epiphenomenon of nutrient stress. On the other hand, the selective advantage of sclerophylly as protection against herbivore attacks, though not demonstrated in different environments, seems to be of great interest and deserves further studies.     

Characterising sclerophylly: some mechanical properties of leaves from heath and forest

Although sclerophylly is widespread through the world and is often the dominant leaf-form in mediterranean climates, the mechanical properties of sclerophyllous leaves are poorly understood. The term ”sclerophyllous” means hard-leaved, but biologists also use terms such as tough, stiff and leathery to describe sclerophyllous leaves. The latter term has no precise definition that allows quantification. However, each of the former terms is well-defined in materials engineering, although they may be difficult or sometimes inappropriate to measure in leaves because of their size, shape or composite and anisotropic nature. Two of the most appropriate and practically applicable mechanical properties of sclerophyllous leaves are ”strength” and ”toughness”, which in this study were applied using punching, tearing and shearing tests to 19 species of tree and shrub at Wilson’s Promontory, Australia. The results of these tests were compared with leaf specific mass (LSM) and a sclerophylly index derived from botanists’ ranks. Principal components analysis was used to reduce the set of mechanical properties to major axes of variation. Component 1 correlated strongly with the botanists’ ranks. Overall, leaves ranked as sclerophyllous by botanists were both tough and strong in terms of punching and tearing tests. In addition, tough and strong leaves typically had high toughness and strength per unit leaf thickness. There was also a significant correlation between component 1 and LSM. Although more detailed surveys are required, we argue that sclerophylly should be defined in terms of properties that have precise meanings and are measurable, such as toughness and strength, and that relate directly to mechanical properties as implicit in the term. Received: 4 March 1999 / Accepted: 22 November 1999     

Phosphorus nutrition of jarrah (Eucalyptus marginata) seedlings

Summary Effects of calcium phosphate supply on plant dry matter and phosphorus concentrations of parts of jarrah (Eucalyptus marginata) seedlings grown in a lateritic topsoil from the jarrah forest were examined in two glasshouse trials. Phosphorus deficiency depressed root and shoot dry weights and severely deficient leaves were smal and purple with prominent red major veins. Phosphorus deficiency severely reduced stem phosphorus levels (0.5% to 0.02%, experiment 1). Phosphorus concentrations were higher in bark than wood and the amount of phosphorus in the bark was sensitive to stem age and phosphate supply. Phosphorus adequate plants had bark phosphorus concentrations in the range 0.2–0.9% compared to <0.1% in deficient plants (experiment 2). Jarrah leaves accumulated dry matter up to 80 days after expansion and some leaves exported phosphorus during this period. Bark analysis may therefore be preferable to leaf analysis for detecting phosphorus deficiency in this species.     

Sclerophylly in Qualea Parviflora (Vochysiaceae): Influence of Herbivory, Mineral Nutrients, and Water Status

Qualea parviflora Mart. (Vochysiaceae) is a deciduous tree, commonly observed in campo sujo, cerrado sensu stricto and cerradão vegetation types in Brazilian cerrado (savannas). In this study we investigated herbivory, nutritional, and water status effects on leaf sclerophylly of Q. parviflora. Twenty fully expanded leaves were taken from 10 plants in each vegetation type four times a year. Mean leaf concentration of N, P, K, Ca, C, Al, Si, and percentage of total phenols, herbivory and tannins were measured on a plant basis. Leaf specific mass (LSM) (g m^?2), a sclerophylly index, and pre-dawn leaf water potential (MPa) were also recorded. Soil samples below each tree were collected to quantify N–NO₃, N–NH₄, P, K, Mn, soil moisture, organic matter, Si, and Al. Qualea parviflora showed a LSM from 69 to 202 g m^?2 and leaves were younger and less sclerophyllous in November (beginning of rainy season). Q. parviflora inhabiting the cerradão had leaves with higher concentration of nutrients and lower sclerophylly while trees in campo sujo and cerrado sensu stricto did not show significant differences in leaf sclerophylly. The concentrations of N, P, K and tannins had an inverse relationship with leaf age. Concentration of phenols, Al, C, Ca, Si, C/N and Ca/K increased with leaf age. The concentrations of P and Ca/K ratio in leaves explained 60% of variation observed in leaf sclerophylly. We did not find any significant relationship between the level of sclerophylly and water potential or herbivory. Our results corroborate the hypothesis that predicts lower concentrations of essential macronutrients would be the main factors influencing higher sclerophylly in leaves of Q. parviflora plants in Cerrado.     

Leaf morphology and macronutrients in broadleaved trees in central Italy

As part of an intensive monitoring programme (MON.I.TO, Intensive Monitoring of Forests in Tuscany), a 3-year survey was carried out, which included three tree species (beech, Fagus sylvatica L.; Turkey oak, Quercus cerris L.; holm-oak, Quercus ilex L.) located at six different sites. Leaves were sampled annually and analysed for nutrient concentrations (nitrogen, sulphur, phosphorus, magnesium, calcium, potassium and sodium) and morphological parameters (leaf area, dry weight, leaf mass per area, leaf thickness and leaf density). Results indicated considerable interannual variation of all the parameters. Differences between sampling sites indicated that, of all parameters measured, leaf mass per area could explain best the differences in field performance under conditions of stress. In fact, leaf mass per area was greater in the drier sites or when sea salt deposition occurred. Nevertheless, the variation of leaf mass per area over the 3 years did not reflect the differences in rainfall. Higher leaf mass per area was accompanied by lower concentrations of phosphorus and nitrogen, which could be a dilution effect due to an increase of structural carbon compounds in sclerophyllous leaves, as revealed by the total foliar content of these elements. Leaf mass per area as a measure of sclerophylly reached very high values among mesophile vegetation. Long-range transport of sea salt from coastal areas to mountain areas was mirrored in sodium concentrations of leaves.     

A note on the nutrient and mineral composition of Leucaena leucocephala in Western Nigeria

Analysis of Leucaena leucocephala from various localities in Western Nigeria shows that young leaves, pods and seeds contain more crude protein but lower crude fibre and ether extract than mature ones. The crude fat component was higher in the seeds (4.6% and 5.5% for green and ripe seeds, respectively) than in any other edible part of the plant, while the green and brown seed coats contain the least amount of ether extract and total ash. The mineral composition of the dry matter of mature leaves is 2.8% calcium, 0.26% phosphorus, 0.37% magnesium, 1.78% potassium, 0.21% sodium and 0.12% iron.     

Sclerophylly in two contrasting tropical environments: low nutrients vs. low rainfall

The defining characteristics of sclerophylly are mechanical (e.g., hardness, toughness, stiffness), but little is known about how they vary in contrasting environments and contribute to the adaptiveness of sclerophylly. Here we investigate how the degree and nature of sclerophylly in terms of leaf mechanics differ between vegetation of two contrasting stressful environments, maquis on nutrient-deficient, moist sites and dry forest on moderate-nutrient, drier sites. We measured toughness, strength, and stiffness at the level of the whole leaf ("structural") and per unit thickness ("material"). Leaves of maquis plants were on average structurally stiffer, stronger, and tougher than those in dry forest. There was little difference in material properties between habitats, and leaf thickness was the main contributor to differences in structural mechanical properties between habitats. Flexural stiffness varied most among species and habitats, correlating strongly with leaf mass per area and thickness. We suggest that having thicker leaves allows efficient packaging of biomass to reduce branching costs in sunny but stressful environments, with subsequent impacts on structural mechanical properties. Sclerophylly is probably a complex phenomenon, however, with its mechanical constitution arising from both evolved mechanical properties that confer protection or resistance to stress and nonadaptive mechanical consequences of adaptation to stressful environments.     

Comparison of four methods for measuring osmotic potential of tree leaves

Osmotic potential (π) of tree leaves is usually determined by pressure-volume (PV) analysis, which is too time consuming for large-scale assessments such as ecological surveys and genetic screening. We investigated methods for rapid measurement of π in sclerophyllous species. The osmotic potential of expressed sap (ES), freeze-thaw leaf discs and hot water extracts – measured using a variety of techniques – were compared with PV analysis using sclerophyllous species; Eucalyptus globulus Labill. , E . camaldulensis Dehnh. , E . cladocalyx F. Muell. and non-sclerophyllous species; Populus nigra L and Alnus jorullensis H.B.K. All methods were sensitive to changes in π and produced results that were unequal but well correlated (r² 0.82–0.90). The proportion of leaf water extracted in ES was reduced by low water content and sclerophylly without affecting method accuracy. PV analysis provided a poor correction for apoplastic dilution of ES. Results of ES osmometry and PV analysis were comparable, and osmometry of ES is proposed as a suitable method for large-scale investigations of leaf π.     

Gall-forming and free-feeding herbivory along vertical gradients in a lowland tropical rainforest: the importance of leaf sclerophylly

In contrast to most insect guilds, gall-forming insects are thought to reach highest diversity on sclerophyllous vegetation, such as Neotropical savannas and Mediterranean vegetation types. The water and nutrient stress endured by meristems of canopy trees in tall wet tropical rainforests may cause leaf sclerophylly. Hence, the upper canopies of such ecosystems may represent a suitable habitat for gall-forming insects. At the San Lorenzo Protected Area, Panama, we estimated free-feeding herbivory and gall densities within five sites in 2003 and 2004, by surveying leaves in vertical and horizontal transects. In each sample, we recorded leaf density (mature and young foliage), free-feeding herbivore damage and number of galls, including the presence of live larvae, parasitoids or fungi. We surveyed 43 994 leaves, including 231 plants and 73 tree and liana species. We collected 5014 galls from 17 host-plant species, including 32 gall species of which 59% were restricted to the canopy (overall infestation rates: 2.4% in 2003, 5.5% in 2004). In 2003, 16% of the galls were occupied by live larvae, against 5% in 2004. About 17–20% of leaves surveyed suffered from free-feeding herbivory. Leaf sclerophylly increased significantly with sampling height, while free-feeding herbivory decreased inversely. Conversely, the number of live galls collected in the canopy was 13–16 times higher than in the understorey, a pattern consistent across sites and years. Hence, the probability of gall survivorship increased with increasing leaf sclerophylly as death by fungi, parasitoids or accidental chewing were greater in the understorey. Increasing harsh ecophysiological conditions towards the upper canopy appear favourable to galls-forming population maintenance, in support of the hypothesis of harsh environment. Hence, gall diversity and abundance in the upper canopy of tall tropical forests are perhaps among the highest in the world.     

Nutrient and energy content of freshwater fishes

The analyses of fresh total body homogenates of 2385 freshwater fishes from 17 species originating in natural waters and aquaculture facilities resulted in average values of 19.1–37.8% dry matter, 13.6–18.3% crude protein, 0.8–18.3% crude fat, 0.16–2.17% nitrogen-free extracts (NFE), 2.2–2.9% nitrogen and 0.39–1.02% phosphorus. The gross energy content ranged from 3.83 to 11.49 MJ/kg. With increasing fat content in the fishes, water content decreased and dry matter content increased. Through this interdependence, a significant correlation was found between dry matter and crude fat and energy content (r²=0.92–0.97). The values of crude protein, crude ash, NFE, nitrogen and phosphorus did not show any significant influence on dry matter content (r²=0.0005–0.46). Based on the significant correlation between dry matter and gross energy (gross energy [MJ/kg]=0.0253 dry matter [%]^1.6783), the energy content of freshwater fishes can easily be estimated by the dry matter content in the total body.     

Leaf specific mass confounds leaf density and thickness

Summary We explored the relationship between leaf specific mass (LSM) and its two components, leaf density and thickness. These were assessed on the leaves of (a) the moderately sclerophyllous tree Arbutus menziesii distributed along a natural nutrient/moisture gradient in California, (b) eight sclerophyllous shrub species on four substrates in south-western Australia, and (c) seedlings of two morphologically contrasting Hakea species grown under varying soil nutrient, moisture and light regimes in a glasshouse experiment. Leaf area, mass, LSM, density and thickness varied greatly between leaves on the same plant, different species, and with different nutrient, moisture and light regimes. In some cases, variations in LSM were due to changes in leaf density in particular or thickness or both, while in others, density and thickness varied without a net effect on LSM. At lower nutrient or moisture availabilities or at higher light irradiances, leaves tended to be smaller, with higher LSM, density and thickness. Under increased stress, the thickness (diameter) of needle leaves decreased despite an increase in LSM. We concluded that, while LSM is a useful measure of sclerophylly, its separation into leaf density and thickness may be more appropriate as they often vary independently and appear to be more responsive to environmental gradients than LSM.     

An evaluation of the method used in digestibility estimations of a dietary ingredient and comparisons on external and internal markers, and time of faeces collection in digestibility studies in the fish Oreochromis aureus (Steindachner)

Results of experiments carried out on the tilapiine fish Oreochromis aureus (Steindachner) to evaluate the method used in the determination of the apparent dry matter and nutrient digestibilities of an ingredient, together with validity of estimations based on day and night collection of faeces, and internal and external markers, are presented. The apparent dry matter and protein digestibilities of the test diets were linearly correlated to the percent substitution of the test ingredient, percent fibre and ash in the test diets (P less than 0.05); the correlation coefficient for the individual relationships decreased in that order. Dry matter or protein digestibilities were not correlated to the dietary protein content (P greater than 0.05). Similarly there were no statistical differences between digestibility estimations based on faecal material, voided in the day or the night; and estimated using different markers viz crude fibre and Cr2O3. The apparent dry matter and protein digestibilities of the test ingredients, leaf meal, were curvilinearly related to the percent substitution of the ingredient in the test diets. The ingredient digestibilities estimated, using crude fibre as the marker, were consistently higher than those estimated with Cr2O3 (P less than 0.05). The present study suggests that, for ingredient digestibility estimations, test diets prepared by mixing 15-20% of the ingredient to a reference diet would be more desirable.     

Biochemical aspects of parasitism by the angiosperm parasites I. Phosphate fractions in the leaves of loranthus and hosts

The leaves of Dendrophthoe falcata growing on different trees had differing dry solid content. With 1 exception, the parasite leaf contained less dry solid than host leaf.

A characteristic feature of the loranthus parasite was the accumulation of phosphate in excess of that present in host leaves. Expressed as percent of the total phosphorus, the parasite tissue contained significantly less acid-insoluble phosphate than the host leaves. When the acid-insoluble material was fractionated into phospholipid, RNA, DNA and phosphoprotein, every component was found to be present in a smaller amount in parasite than in host leaf, expressed as percent of total phosphate in leaves.

A comparative study of the phosphate fractions in the infected and uninfected branch of infected host tree revealed that, without exception, there was a demonstrable increase in the DNA content of the infected branch.

     

Untersuchungen zur Stoffproduktion bei Futterkohl

Zusammenfassung An 5 Idiotypen verschiedener Convarietäten und Varietäten vonBrassica oleracea L. wurde der Verlauf des Trockenmassen-, Rohfaser-, Rohprotein- und Reinproteingehaltes sowie der löslichen N-Fraktion während der Vegetationszeit ermittelt. Im Trockenmassen-und Rohfasergehalt treten zwischen den geprüften Formen beachtliche, bei den Proteinfraktionen geringfügige Unterschiede auf. Die Organe Stamm, Blattspreite und Blattstiel unterscheiden sich in den Gehalten beachtlich. Der Verlauf während der Vegetationszeit bei der Gesamtpflanze ist dem Trend nach für den Trockenmassengehalt steigend, Rohfasergehalt fallend, Rohproteingehalt zunächst fallend, dann steigend und Reinproteinanteil gleichbleibend. Die Sorten und Organe zeigen in einigen Fällen ein abweichendes Verhalten.Während die Gesamttrockenmasse nach Ende Oktober nicht mehr zunimmt, steigt die Rohproteinmasse weiter an. Die relative Trocken- und Rohproteinmassenbildung wird durch Bildung des rel. Proteinfaktors verglichen. Bei Wachstumsbeginn läuft die Rohproteinbildung der Trockenmassenbildung voraus. Der Umschlagspunkt liegt bei den Organen und Sorten zeitlich verschieden und fällt mit dem Beginn des Wachstumsabschnittes 3 (Stoffumlagerung) zusammen. Das Modell der Massenzunahme für die Rohproteinmasse stimmt mit dem der Trockenmasse weitgehend überein.Die Selektion trockenmassenreicher Typen (Winterfestigkeit) ist erfolgversprechend und scheint die Futterqualität kaum zu beeinflussen. Die geringe Variabilität der Proteingehalte eröffnet keine wesentlichen züchterischen Möglichkeiten zu ihrer Erhöhung. Die Ergebnisse werden diskutiert und mögliche Zusammenhänge angedeutet.

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Experiments on the production of bulk in cabbageII. Quantitative changes in some quality determining contents during the course of development

Summary In five different varieties and subvarieties ofBrassica oleracea L. dry matter, crude fibre, crude and pure protein contents as well as soluble nitrogenous fractions have been ascertained during the vegetation period. Among the various forms under study, considerable differences with respect to dry matter and crude fibre and small differences with respect to protein content were noticed. Stem, leaf blade and leaf stalk differ considerably in composition. In the course of the vegetation period dry matter content increases while crude fibre content decreases later on. Its pure protein fraction remains the same, though in some cases the varieties and organs show deviating results.While total dry matter no longer increases after October, the crude protein fraction increases continuously. The relative synthesis of dry matter and crude protein becomes equalized by the formation of the respective protein factor. At the onset of growth crude protein formation exceeds the production of dry matter. The time of change differs with the organ and varieties, and coincides with the start of the 3rd growth period. The model for mass increase of crude protein agrees with that of dry matter.Selection of types rich in dry matter (winter hardiness) promises success; it seems to have hardly any influence on fodder quality. The low variability in protein content does not lead to possibilities for improvement. The results are discussed.

     

Leaf Mechanical Properties in Sclerophyll Woodland and Shrubland on Contrasting Soils

Sclerophylly is a common feature of vegetation on infertile soils, and its adaptive significance has been linked to nutrient-use efficiency by protection of leaves to maximise carbon gain. However, there has been little investigation of how the leaf mechanical properties that contribute to the phenomenon of sclerophylly vary along nutrient gradients. In this paper, we investigate how leaf mechanical properties vary among plants on three contrasting soil types (grey sand, laterite soil, and soil overlying dolerite) in a Mediterranean climate in southwestern Australia. Most species were sclerophyllous, but there was 5-fold variation in leaf mass per unit area (LMA) and 17- to 473-fold variation in mechanical properties among species. Species growing on laterite and/or sand (low-nutrient soils) had higher punch strength, work (a measure of toughness) to punch, specific (per unit leaf thickness) work to punch, work to shear, specific work to shear, and flexural stiffness (EI _W) than those on dolerite soils (higher in nutrients). There were few differences in mean values of leaf mechanical properties between the two low-nutrient soils, possibly because the lower concentration of nutrients in the sand is balanced by the greater soil volume than the laterite soil (higher concentration of nutrients, but shallower). There were also few differences in leaf properties between plants of the same species growing on contrasting soil types. There was some variation among sclerophyllous species in their mechanical characteristics, but overall, EI _W provided the strongest contribution to sclerophylly, explaining up to 81% of the variation in LMA. There was no evidence of differences among soil types in the relationships of mechanical properties with LMA, and therefore, no evidence of variation in the mechanical constitution of sclerophylly among soil types.     

硫硒配施对茎瘤芥生长和营养效应的研究

以茎瘤芥品种‘涪杂1号’为材料,通过盆栽实验探讨不同浓度的硫(S)、硒(Se)配施处理对茎瘤芥干物质积累、矿质元素吸收及膨大茎营养品质的影响,为生产中合理施用硫、硒肥提供理论依据.结果表明:与对照(S0Se0,未施硫硒肥)相比,增施硫、硒肥处理均能显著提高茎瘤芥的根、膨大茎、叶片和单株干物质产量,并以S50Se1[S/Se=50(mg/kg)/1(mg/kg)]和S100Se1的处理效果较好,其单株干物质产量分别比对照显著增加32.3%和36.2%;不同硫、硒浓度配施处理对茎瘤芥13种矿质元素积累的影响不同,主要显著促进了茎、叶对氮、磷、钾、硫、硒的吸收积累,而对其它元素的影响不显著,其中茎、叶的硒含量以S50Se3处理最高,硫含量以S100Se1处理最高;各硫硒配施处理对膨大茎营养品质的影响不同,其中S50Se1和S50Se3处理能显著提高膨大茎有机硒、总氨基酸和粗蛋白含量,而对维生素C和可溶性糖含量无显著影响.可见,适宜的硫硒配施可以明显促进其对矿质元素的吸收,提高植株干物质积累,有效改善茎瘤芥膨大茎营养品质,且硫硒配施用量以S 50mg/kg、Se 1mg/kg为宜.     

Relationship between some chemical components of herbage, dietary preference and fresh herbage intake rate by the Japanese serow

The serow is a native Japanese mammal and has been designated as a special natural treasure. However, it has been a pest of crops since the 1970s. Since the serow is an indigenous and protected species, it must be controlled without endangering the animal itself. To do this, it is necessary to determine if the serow selects particular crops to consume, and if so, what components of the crop plants influence this choice.In Experiment 1, we measured the preference of four serows for two crop plants and three wild plants, using the cafeteria method. The plants were offered to the animals from 09.00 to 15.30. The serows preferred crop plants to wild plants. The crude protein content of crop plants (28.0 and 32.7%) was higher than that of the other plants (P<0.05). Dry matter (DM) intakes per hour correlated positively with crude protein content (P<0.01) and negatively with nitrogen free extract content (NFE) (P<0.05) and acid detergent fibre content (ADF) (P<0.01) in the offered plants. In Experiment 2, the food intakes of 17 plants consisting of 6 crops and 11 wild plants were measured. Individual 300g samples from each plant were offered for 20min. The serows ate the crops and the forbs in preference to the tree leaves. They consumed little of the conifer tree leaves and the grasses. The dry matter intake and the ADF content were negatively correlated (P<0.05). The dry matter intake and the crude protein content were positively correlated (P=0.054). A multiple regression formula was constructed as follows: Y=1.65CP+0.56NFE+0.38DM-57.45,R(2)=0.54,P=0.016The feeding characteristics suggest that crops might be selected and ingested by the serows because of their higher crude protein, NFE and the dry matter contents.     

Tree Leaf Form in Brunei: A Heath Forest and a Mixed Dipterocarp Forest Compared1

Canopy‐top leaves of the dominant tree species from two 0.96‐ha plots in Brunei, northern Borneo, were sampled for structural and chemical analysis. Thirteen species from the mixed dipterocarp forest at Andulau and 14 from the lowland heath forest at Badas were studied. The heath‐forest species had significantly thicker leaves and were lower in nitrogen and ash concentration than those from the mixed dipterocarp forest. There were no significant differences between the two species groups in leaf mass per unit area (LMA), leaf fracture toughness, carbon concentration, 8¹³C, neutral detergent fiber concentration, sclerophylly index, and stomatal density. A significant negative correlation between %C and 8¹³C was found for the species from the mixed dipterocarp forest, but not those from the heath forest. The degree of sclerophylly measured in physical terms overlapped between the two sites to a considerable degree; however, all six species tested that were present in both plots had higher leaf fracture toughness in the heath forest. The possible reasons for the marked sclerophylly in the mixed dipterocarp forest are discussed.