Seedling development of the miombo woodland tree Julbernardia globiflora

Abstract. The development of seedlings of the miombo tree (Julbernardia globiflora) was studied for 28 months (December 1987 - April 1990) at a Zambian grassland site from which fire has been excluded for many years. Germination occurred 10–25 days after seed planting and germination rates under laboratory and field conditions did not differ significantly. Seed germinability after one year of storage declined to 38 %. Leaf production occurred between December and April with peaks in January each year. Leaf fall occurred steadily throughout the dry season but leafless seedlings were first observed in August of each year. All seedlings had shed all their leaves by the end of November during the first year while 10% of the seedlings remained leafy throughout the second year. Annual seedling mortality was about 60 % during the first year and declined to 3 6 % during the second year. Survivorship appeared to stabilise at 3 8 % when seedlings were 5 5 - 5 9 weeks old. Every seedling experienced shoot die-back during the first hot dry season (September - November) when open-pan evaporation rates averaged 5–10 mm / day. However, shoot mortality did not necessarily result in plant mortality. Because of recurrent shoot die-back there was little increase in shoot biomass and height at the end of the second and third growing seasons. Seedlings that survived shoot die-back sprouted from roots which showed a steady annual accumulation of biomass. Early shoot die-back in seedlings of Julbernardia globiflora as documented in this study was not causedby fire but most probably by drought and / or nutritional stress.     

Inoculation of pines in a nursery withPisolithus tinctorius andThelephora terrestris in Korea

Summary Mycelial inocula of the two ectomycorrhizal fungi,Pisolithus tinctorius (Pt) andThelephora terrestris (Tt) were artificially introduced into fumigated nursery soil before seed sowing. At the end of the first growing season, Pt stimulated the height growth ofPinus densiflora, P. thunbergii, P. rigida, andP. rigida×taeda by 55, 36, 69, and 37%, respectively, compared with control seedlings (with no fumigation and no inoculation). When growth stimulation was expressed by dry weight, Pt increased dry weight ofP. densiflora andP. rigida×taeda by 143% and 128%, respectively, over control seedlings. Total absorption (per plant top) of nitrogen and phosphorus of inoculatedP. densiflora seedlings was also increased by 188 and 144%, respectively, even though the concentration of these two elements in the tissue was not significantly different from that of control seedlings.Tt failed to stimulate growth of the inoculated plants possibly due to its lower level of infection (40–60% mycorrhizal tips) compared with the relatively high infection (over 80%) of Pt.Pinus koraiensis did not respond to inoculation during the first growing season, probably due to its inherent characteristics of producing a small number of short roots and its slow shoot growth during the first few years after germination.This study was supported by the International Foundation for Science in Stockholm, Sweden.     

Ectomycorrhizae in coastal miombo woodland of Tanzania

Summary Ectomycorrhizae were found in root samples of the treesAfzelia quanzensis Welw. andBrachystegia spiciformis Benth. (Caesalpiniaceae), collected in the coastal miombo type woodland 50 km west of Dar-es-Salaam, Tanzania. Root nodules with a structure resembling that of nitrogen-fixing root nodules of other leguminous plants were observed in theA. quanzensis material. The climate of the locality is rather dry, and strongly seasonal. In the tropics, ectomycorrhizae have previously been found only in humid or rain forest climate zones.     

Growth of Phragmites australis and Phalaris arundinacea in constructed wetlands for wastewater treatment in the Czech Republic

Common reed (Phragmites australis) and reed canarygrass (Phalaris arundinacea) are two most commonly used plant species in constructed wetlands for wastewater treatment in the Czech Republic. Growth characteristics of both plants (biomass, stem count, and length) have been measured in 13 horizontal sub-surface flow constructed wetlands since 1992. The results revealed that while Phalaris usually reaches its maximum biomass as early as during the second growing season, Phragmites usually reaches its maximum only after three to four growing seasons. The maximum biomass of both species varies widely among systems and the highest measured values (5070 g m⁻² for Phragmites and 1900 g m⁻² for Phalaris) are similar to those found in eutrophic natural stands. The shoot count of Phragmites decreases after the second growing season while length and weight of individual shoots increases over time due to self-thinning process. Number of Phalaris shoots is the highest during the second season and then the shoot count remains about the same. Also the shoot length remains steady over years of constructed wetland operation.     

Phenology and nutrition of miombo woodland trees in Zambia

Seasonality in nitrogen (N) and phosphorus (P) concentration in soil and shoots of five Brachystegia-Julbernardia (miombo) woodland trees was studied from September 1991 to March 1993 at two regrowth miombo sites in central Zambia. Shoot growth started in the dry season (September–November) and lasted until April during the 1991/92 season but had virtually ceased by January 1993 during the 1992/93 season. The shoot growing season was associated with low foliar N/P ratios. These ratios were much lower (<5) during the 1991/92 season than in the 1992/93 season (12–15). The increase in foliar N/P ratios after the shoot growing season was caused by a sharp drop in foliar P concentration, apparently due to reabsorption before leaf fall. There were no annual variations in biomass N concentration in contrast to P. During the 1992/93 growing season P concentrations in foliage and wood were a quarter and a third, respectively, of the 1991/ 92 levels. Since the short shoot growing season observed during the 1992/93 season is typical of savanna woodland trees in southern Africa, the high biomass P concentration and longer growing season in 1991/92 season were exceptional and may have been related to reduced competition by shallow rooting herbaceous plants caused by the severe drought of that season.     

Seasonal variation of growth and development of the roots of five second year conifer species in the nursery

Every other week over their second growing season, stem height, collar diameter, shoot and root dry masses, number of lateral roots and length of the tap root were measured on nursery grown seedlings ofAbies balsamea L. Mill.,Pinus banksiana Lamb.,Pinus resinosa Ait.,Picea mariana Mill. BSP andPicea glauca Moench Voss. Root elongation, branching and mycorrhizal development were also recorded.Given species showed distinct seasonal growth patterns. The rate and timing of maximum root growth (mg/dry weight/week) differed markedly between species.Except for the increase in height ofPinus banksiana, root and shoot growth were not negatively correlated.The results are discussed in relation to the performance of tree seedlings in the nursery.     

Stomatal conductance in a tropical xerophilous shrubland at a lava substratum

Diurnal variation in leaf stomatal conductance (g _s) of three xerophilous species (Buddleia cordata, Senecio praecox and Dodonaea viscosa) was measured over a 10-month period during the dry and wet seasons in a shrubland that is developing in a lava substratum in Mexico. Averaged stomatal conductances were 147 and 60.2 (B. cordata), 145 and 24.8 (D. viscosa) and 142.8 and 14.1 mmol m^–2 s^–1 (S. praecox) during the wet and dry season respectively. Leaf water potential () varied in a range of –0.6 to –1.2 (S. praecox), –0.6 to –1.8 (B. cordata) and –0.9 to –3.4 MPa (D. viscosa) during the same measurement periods. Stomata were more sensitive to changes in irradiance, air temperature and leaf–air vapour pressure difference in the rainy season than the dry season. Although stomatal responses to were difficult to distinguish in any season (dry or rainy), data for the entire period of measurement showed a positive correlation, stomata tending to open as increased, but there is strong evidence of isohydric behaviour in S. praecox and B. cordata. A multiplicative model relating g _s to environmental variables and to accounted for 79%–83% of the variation of g _s in three sites (pooled data); however, the performance of the model was poorer (60%–76%) for individual species from other sites not included in the pooled data.     

Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment

Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland. A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats. The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa. Not one seedling emerged from dung pats. Neither location nor the other treatments affected the density of emerging seedlings, although only 40.4 seedlings m^–2 emerged of the 200 seeds m^–2 planted. Shading dramatically increased the density of surviving seedlings. In the open, only 3 and 1.5 seedlings m^–2 remained respectively at the end of the growing season or the beginning of the next, compared to 23.3 and 19.5 seedlings m^–2 under shading for these respective times. This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall. Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline. Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years. Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings. Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated. A. karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank. This has implications for the invasion of grassland by woody species.     

Survival and recruitment favored by safe site-strategy – the case of the high alpine, non-clonal cushions of Eritrichium nanum (Boraginaceae)

In 14 permanent plots at four distant sites in the European Alps life tables, cohorts and recruitment of Eritrichium nanum were studied during a period of 5 resp. 6 years. The germination rates were tested in the Botanical Institute of the University of Basel and besides the life tables of the plots at natural habitats we followed the early stages of development in pot cultures at the University Botanical Garden in Basel. Most of the seedlings of E. nanum emerged in spring, in the pot cultures as well as under natural conditions, a behavior which confirms general findings. But the germination rates (0–30%) were markedly low. The pot cultures show that some seeds are after 4 or 5 years still viable. Premature flowering in the second summer is a rather common behavior of E. nanum seedlings, which happened in all the cohorts observed during five years, partly up to 20% of the total of living plants in the corresponding plots. It is evident that life expectancy of E. nanum plants increase rapidly with rising size, to such a degree that cushions of only 10–20 cm² surface have a survival chance of more than 70%, those of 20–30 cm² even of more than 95% during a period of 5 years. But it has to be mentioned that juvenile plants are not growing parallel to their lifetime, many of them persist during several years in the size categories I/II (0–5 cm²), while others suddenly arise from category I to III (–10 cm²) or even IV (–20 cm²), what is mostly caused by joining, a specific type of safe site strategy. Though not obligatory, juvenile plants growing close together, often add their sizes to form one larger cushion, which was found to be particularly advantageous for recruitment. This is obviously a primary strategy of E. nanum and favors, especially on bare ground, the survival of its descendants which was found higher than expected (average of 26% after 5 years’ growth at natural habitats). Focusing on the whole life cycle of E. nanum, the present studies show constraints in pollination, a small seed production and low germination rates to be well compensated by the high survivorship and successful recruitment of the seedlings.     

Emission and oxidation of methane in Equisetum fluviatile stands growing on organic sediment and sand bottoms

Methane emission and rhizospheric CH₄ oxidation were studied in stands of Equisetum fluviatile, a common cryptogam in boreal lakes. The experiment was performed in mesocosms with organic sediment or sand bottoms under natural variation of temperature and light using the light-oxic – dark-anoxic chamber (LO/DA) technique. Net CH₄ emission from the organic sediment during the growing season varied between 3.4 and 19.0 mg m^–2 h^–1, but from sand the net CH₄ emission was only 3–10% of that measured from the organic sediment. In the organic sediment net CH₄ emission was very significantly correlated with sediment temperature (r² = 0.92). In the sand mesocosms the variation of net CH₄ emission was better correlated with the shoot biomass than with sediment temperature variation during the growing season, indicating that methanogens were severely limited by substrate availability and were probably dependent on substrates produced by E. fluviatile. The proportion of the methane oxidized of the potential CH₄ emission in summer did not differ significantly between the bottom types. The net CH₄ emission during the growing season as a proportion of the seasonal maximum of the shoot biomass was significantly higher in the organic sediment mesocosms (6.5%) than in sand (1.7%). The high CH₄ emissions observed from dense well-established E. fluviatile stands in the field appear to be more related to temperature-regulated turnover of detritus in the anaerobic sediment and less to CH₄ oxidation and seasonal variation in plant growth dynamics     

Growth,chlorophyll and mineral nutrient studies on phalaris arundinaceae L. in three scottish lochs

Growth, chlorophyll and mineral nutrients studies were made in Phalaris arundinaceae L. in three Scottish lochs of varying nutrient status from March to November in 1975. The maximum shoot height and shoot dry weight attained by the plants were approximately 160 cm and 4 g respectively. Seasonal changes in the chlorophyll levels in the Phalaris leaf were studied and two peaks were found, one in April and the other in June–July. Maximum chlorophyll level attained was 9 mg g^–1 leaf dry weight. The changes in the mineral levels in the root, stem, leaf and inflorescence parts of the plants from the three lochs were also assayed throughout the growing season. A total of eight mineral elements were studied, including carbon, nitrogen, phosphorus, potassium, sodium, calcium, magnesium and iron. Variations in both the mineral concentrations and their pattern of changes during the study period among the plants from the lochs were observed and discussed.     

Nitrogen fixation in breeding lines of Phaseolus vulgaris L.

Nitrogen fixation in nine common bean (Phaseolus vulgaris L.) lines was estimated using the ¹⁵N isotope dilution method at two locations in two seasons. In the first season at one location no N₂ fixation was detected while in the second season up to 51 kg N ha^–1 were estimated. There were significant differences between lines and correlations between trials were significant for the amounts of N₂ fixed, but not for total shoot nitrogen. The plants that fixed the most nitrogen nodulated rapidly after germination. Differences in maximum nodule mass, but not specific nodule activity, were detected also.     

Effect of exposure to fluoride,nitrogen compounds and SO2 on the numbers of spruce shoot aphids on Norway spruce seedlings

Summary Development of spruce shoot aphid (Cinara pilicornis Hartig) populations was monitored in natural and artificial infestations of Norway spruce (Picea abies Karst.) seedlings, exposed to air pollutants in an experimental field. The pollutants, applied both singly and in mixtures, were gaseous sulphur dioxide, NaF (30 mg l^-1 F) and Ca(NO₃)₂ or (NH₄)₂SO₄ in aqueous solutions (200 mg l^-1 N). Aphid numbers on 10 seedlings in each treatment and two control plots were counted at 2-week intervals. At the beginning of the experiment aphid numbers did not differ between treatments. Aphid populations peaked in late June and early July. All the pollutants and their combinations significantly increased the numbers of aphids per seedling. Four apterous females were transferred to spruce seedlings which were growing in containers in the same plots. After 4–5 weeks aphid numbers were significantly higher in the fluoride treatment and in the combined treatment of fluoride, nitrogen and SO₂. The pollution treatments did not have a significant effect on shoot growth. Concentrations of F and S in needles were higher in treatments involving these pollutants. There were no significant differences in the concentrations of free amino acids in shoot stems between control and fluoride treatment. However, the relatively low concentration of arginine in the F treatment at the end of the growing season might indicate disturbances in the nitrogen metabolism of spruce seedlings.     

Survival and growth of Virola surinamensis yearlings: Water augmentation in gap and understory

Summary Factors affecting seedling Virola surinamensis (Myristicaceae) survival and growth were investigated on Barro Colorado Island, Panama. Seedlings planted 3 months after germination were monitored in treefall gaps and understory using 2.25 ha irrigated and control plots through the first dry season. During the dry season, irrigated plants in gaps increased total leaf area significantly more than did irrigated plants in the shaded understory. Over the same dry season, control plants in gaps and in the shaded understory lost similar amounts of leaf area. Seedlings in understory were suppressed in stem height and biomass in both irrigated and control plots; these measures were greater in gaps and greatest in irrigated gaps (height). Roots were similar in length in all treatments, but greater in biomass in gaps than understory due to greater proliferation of secondary roots in control and irrigated gaps than in control and irrigated understory. This experiment demonstrates both water and light limitation during the first dry season after germination. V. surinamensis seedlings are capable of survival and modest growth of leaf area in the deep shade of the understory in moist locations; they are severely disadvantaged in shaded understory subject to drought, where most seeds fall and most seedlings establish. The broken canopy of a gap allows shoot and consequently root growth that permits seedlings to survive seasonal drought.     

Response of maize and rice to 9-β-L(+) adenosine applied under different environmental conditions

The increase in growth, determined by dry weight gain, of rice (Oryza sativa L.) and maize (Zea mays L.) caused by foliar applications of 9--L(+) adenosine, a putative second messenger elicited by triacontanol, was studied under different environmental conditions. Maize seedlings cultured in the greenhouse under approximately 100 mol m^–2s^–1 of light prior to treatment with L(+) adenosine did not respond unless they received supplemental light (250–300 mol m^–2s^–1) after treatment. Exposure of rice seedlings growing for 16 h at 150 mol m^–2s^–1 to short periods of 450 mol m^–2s^–1 (< than 20 min) had no effect on the positive response of rice to L(+) adenosine; however, exposure for 60 min or more increased the growth of rice and obviated the effect of L(+) adenosine. Rice seedlings treated with L(+) adenosine at different times during the day responded only when treated 9 to 12h after initiation of the photoperiod. Normal growing temperatures under different light intensities had little or no direct effect on the response of plants to L(+) adenosine.     

The response of rice to the growth and nitrogen fixation inAzolla caroliniana andAzolla pinnata at varying rates of phosphate fertilization

The response of rice toAzolla caroliniana, newly introduced in India, was compared with the reponse to the local isolate ofAzolla pinnata at varying rates of phosphate fertilizer (4.4–8.8 kg P ha^–1) during a wet and a dry season.Fresh weight, dry weight and fixed N were more for both species 21 DAI (days after inoculation) than 14 DAI, but acetylene reduction activity (ARA) was higher 14 DAI than 21 DAI. Dry weight of Azolla and fixed N were less 14 DAI forA. caroliniana than forA. pinnata during the wet season. Twenty-one DAI, fresh weight ofA. caroliniana was 62.1 and 27.6% higher than that ofA. pinnata during the wet and dry season, respectively. However, dry weight and fixed N were more 21 DAI inA. caroliniana than inA. pinnata during only the wet season. The ARA was higher inA. caroliniana both 14 and 21 DAI, irrespective of season. The presence of either species in the rice field increased grain yield, straw yield, number of panicles m^–2, number of grains per panicle and reduced percentage sterility during both the wet and the dry season. Phosphate application significantly increased fresh weight, dry weight, ARA and fixed N for both species as well as grain and straw yields of rice. The responses to phosphate fertilizer were similar for both Azolla species and for rice grown with either one of the Azolla species.     

Hydrochory,seed banks,and regeneration dynamics along the landscape boundaries of a forested wetland

Following the environmental sieve concept, the setting in which the recruitment of Taxodium distichum occurs in, becomes increasingly restrictive from the seed to seedling stage in an impounded forested wetland. Although a wide elevational band of dispersing seed moves across the boundary of a swamp-field in the water sheet, the zone of germination is relegated to that portion of the forested wetland that draws down during the growing season. Seedling recruitment is further restricted to the uppermost zone of the winter water sheet. These patterns are likely applicable to other species of dominant swamp species, e.g., Cephalanthus occidentalis crossed the boundary of a forested wetland and abandonded field in winter flooding (November–December and November–March, respectively) in Buttonland Swamp. The elevation of the boundary was 101.3 m NGVD. While the seeds of at least 40 swamp species were dispersed across the boundary, few viable seeds were dispersed after the winter season. Kriged maps showed seeds of T. distichum and C. occidentalis dispersed in patches in the water depending on the position of the water sheet. Most species of both water- and gravity-dispersed species had a localized pattern of seed distribution (either spherical or exponential) and this indicated that seeds may not be dispersed for great distances in the swamp. Water-dispersed T. distichum and C. occidentalis had larger dispersal ranges (A ₀=225 and 195 m, respectively) than Bidens frondosa and B. discoidea (A ₀=14 and 16 m, respectively). Seed dispersal varied with season depending on the availability of seeds. In Buttonland Swamp, viable seeds typically were dispersed for T. distichum in November–June, and for C. occidentalis in November-July. Low water occurred in August 1993 and high in February 1994 (99.8 and 101.6 m NGVD, respectively). The seed banks along the landscape boundary varied in species composition according to elevation (r ² = 0.996). While the similarity of species richness between water-dispersed seeds and the seed bank at elevations that flooded (during June 1993 through May 1995) was high (10–17%), it was low between water-dispersed seeds and the seed bank at elevations that did not flood (5%). T. distichum seeds had a short germination window in that seeds germinated within a year following their production in zones that were flooded in the winter followed by drawdown during the next growing season. After 1 year, less than 5% of the T. distichum seeds remained viable on the surface of the soil. Germination of T. distichum was confined to specific elevations (above 99.3 but below 101.6 m NGVD) during this study with 4.1% of the seedlings surviving for more than 2 years at a mean of 101.4 m NGVD. All seedlings below this elevation died. To maximize natural regeneration along the boundaries of swamps in abandoned farm fields targeted for restoration, this study suggests a flood pulse regime consisting of high water in the winter to maximize dispersal of live seeds followed by low water in the summer to facilitate seed germination and seedling recruitment. Hydrologic restoration could assist in the natural recovery of damaged wetlands if a seed source exists nearby.     

Phenology and water relations of tree sprouts and seedlings in a tropical deciduous forest of South India

The phenology of sprouts (>1 year old, up to 1.5 m in height) and seedlings (<1 year old) of six woody species (four deciduous, one brevi-deciduous, and one evergreen) was examined during the dry season in a tropical deciduous forest of South India. Xylem water potential (_x), leaf relative water content (RWC; % turgid weight), and xylem specific conductivity (K _S; kg s^–1 m^–1 MPa^–1) of sprouts were measured on two occasions during the dry season. In addition, K _S of seedlings (<1 year old) of one deciduous and one evergreen species was determined to allow comparison with sprouts. _x of deciduous species was significantly higher at the second sampling date and was accompanied by a significant increase in K _S and RWC, while the brevi-deciduous and evergreen species did not show any difference in _x. Seedlings of Terminalia crenulata (deciduous) and Ixora parviflora (evergreen) had significantly lower K _S compared to sprouts, while seedlings of all four deciduous species shed their leaves much earlier in the dry season than did conspecific sprouts. More favorable water relations of sprouts compared to seedlings during the peak of the dry season may explain the lower rates of die-back and mortality of sprouts observed in dry deciduous forests of India.

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This revised version was published online in May 2005 with corrections to Received-/Accepted-dates.     

Adaptive significance of nitrogen storage in Bistorta bistortoides,an alpine herb

Summary Studies were conducted to examine the importance of nitrogen storage to seasonal aboveground growth in the alpine herb Bistorta bistortoides. Stored reserves accounted for 60% of the total nitrogen allocated to the shoot during the growing season. The stored nitrogen was equally partitioned between preformed buds of the shoot and the roots/rhizome. Reliance on stored N was similar in populations of a 105-day growing season site and of a 75-day growing season site. Contrary to our initial hypothesis, stored nitrogen reserves were not used to extend the growing season of this species into the late-spring when soils are still cold, and saturated with snow-melt water. The time at which stored nitrogen was used to initiate shoot growth coincided with the time of root initiation, rapid soil warming, and near maximum soil concentrations of NO _inf3 ^sup– and NH _inf4 ^sup+ . Thus, nitrogen demand and soil nitrogen supply were both high at the same time. The importance of nitrogen storage in this species appeared to be in satisfying the high demand of simultaneous vegetative and reproductive growth during the early-growing season after soils thawed. The initiation of rapid leaf and inflorescence growth occurred in mid-June in both sites. The maximum pool size of shoot nitrogen (maximum nitrogen demand) occurred only 12 days later in the long season site, and 28 days later in the short season site. The early-season utilization of nitrogen stores allows plants of this species to initiate reproductive allocation at the same time vegetative tissues are exhibiting maximal growth rates. By releasing vegetative and reproductive growth from competition for nitrogen, seeds could mature early in the alpine growing season, before the frost probability sharply increases in mid-August.     

Relationships between the ant Brachymyrmex obscurior (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae)

Summary. In central Mexico, the ant Brachymyrmex obscurior Forel feeds on nectar produced by extrafloral nectaries of Acacia pennatula (Schlecht. & Cham.) Benth. However, no studies have determined whether the ants visitation is related to plant nectar availability and whether ants protect A. pennatula from herbivory. The objectives of this 2-yr study (2000–2001) were to assess whether seasonal changes in ant visitation coincide with extrafloral nectar productivity in A. pennatula and to determine whether ants protect the plant. At the end of the dry season (April–June) B. obscurior was the only ant species on A. pennatula and extrafloral nectar production is limited to this period. Exclusion experiments, performed at the end of the dry season showed that A. pennatula did not receive a protective benefit when visited by ants. Branches with ants and branches where ants are excluded had similar numbers of the nonmyrmecophile leafhopper Sibovia sp. which was the only herbivore observed under natural conditions.Received 24 March 2004; revised 4 September 2004; accepted 8 September 2004.