South-East Asian Tropical Rainforest Research Articles

Energy Balance Closure Problem over a Tropical Seasonal Rainforest in Xishuangbanna, Southwest China: Role of Latent Heat Flux

The unresolved energy-unclosed problem in micrometeorology refers to the fact that the sum of turbulent fluxes (sensible and latent heat fluxes, Hs and LE) monitored by eddy covariance (EC) methods tends to be lower than the available energy (net radiation (Rn), soil heat flux (G), and heat storage (S)). The lack of energy balance closure (EBC) increases evapotranspiration-measurement uncertainty. Using EC data from Xishuangbanna, a Southeast Asian tropical seasonal rainforest, we analyzed the energy distribution and closure based on micrometeorological features. We found that: (1) the EBC in the rainy season exceeds that in other seasons and that the seasonal moisture content, frictional wind velocity (u*), and LE contribute to the high seasonal variability in EBC; (2) the annual closure is approximately 65%, and energy non-closure is influenced by turbulence intensity and atmospheric stability. When the atmospheric state is unstable to near neutral, u* is greatest, and EBC can reach nearly 80%. (3) energy is mainly allocated to LE, and energy non-closure leads to LE underestimation, especially in the foggy-cool and hot-dry seasons. (4) Heat storage and large time-scale flux effects on EBC were excluded. The causes of energy non-closure in the tropical calm zone need further investigation.

Sensitivity to high temperature and water stress in recalcitrant Baccaurea ramiflora seeds.

Southeast Asia experiences one of the highest rates of deforestation in the tropics due to agricultural expansion, logging, habitat fragmentation and urbanization. As tropical rainforests harbour abundant recalcitrant-seeded species, it is important to understand how recalcitrant seeds respond to deforestation and fragmentation. Baccaurea ramiflora is a recalcitrant-seeded species, widely distributed in Southeast Asian tropical rainforest. In this study, B. ramiflora seeds were sown in three plots, one in a nature reserve and two in disturbed holy hill forests, to investigate seed germination and seedling establishment in the field, while laboratory experiments were conducted to investigate the effects of high temperature and water stress on germination. It was found that seed germination and seedling establishment in B. ramiflora were clearly reduced in holy hills compared to the nature reserve, although the seeds were only moderately to minimally recalcitrant. This was potentially caused by increased temperature and decreased moisture in holy hills, for laboratory experiments showed that seed germination was greatly inhibited by temperatures ≥35°C or water potentials ≤-0.5MPa, and depressed by heat treatment at 40°C when the continuous heating period lasted for 240h or daily periodic heating exceeded 10h. Unlike orthodox seeds, which can endure much higher temperatures in the air-dried state than in the imbibed state, both blotted and immersed B. ramiflora seeds lost viability within a narrow temperature range between 50 and 60°C. As recalcitrant seeds can be neither air-dried nor heated, species producing recalcitrant seeds will suffer more than those producing orthodox seeds in germination and seedling establishment from increased temperature and decreased moisture in fragmented rainforests, which results in sensitivity of recalcitrant-seeded species to rainforest fragmentation.

Occurrence of stomatal patchiness and its spatial scale in leaves from various sizes of trees distributed in a South-east Asian tropical rainforest in Peninsular Malaysia.

In this study, we demonstrated the occurrence of stomatal patchiness and its spatial scale in leaves from various sizes of trees grown in a lowland dipterocarp forest in Peninsular Malaysia. To evaluate the patterns of stomatal behavior, we used three techniques simultaneously to analyze heterobaric or homobaric leaves from five tree species ranging from 0.6 to 31 m in height: (i) diurnal changes in chlorophyll fluorescence imaging, (ii) observation and simulation of leaf gas-exchange rates and (iii) a pressure-infiltration method. Measurements were performed in situ with 1000 or 500 μmol m(-2) s(-1) photosynthetic photon flux density. Diurnal patterns in the spatial distribution of photosynthetic electron transport rate (J) mapped from chlorophyll fluorescence images, a comparison of observed and simulated leaf gas-exchange rates, and the spatial distribution of stomatal apertures obtained from the acid-fuchsin-infiltrated area showed that patchy stomatal closure coupled with severe midday depression of photosynthesis occurred in Neobalanocarpus heimii (King) Ashton, a higher canopy tree with heterobaric leaves due to the higher leaf temperature and vapor pressure deficit. However, subcanopy or understory trees showed uniform stomatal behavior throughout the day, although they also have heterobaric leaves. These results suggest that the occurrence of stomatal patchiness is determined by tree size and/or environmental conditions. The analysis of spatial scale by chlorophyll fluorescence imaging showed that several adjacent anatomical patches (lamina areas bounded by bundle-sheath extensions within the lamina) may co-operate for the distributed patterns of J and stomatal apertures.

Linking biogenic hydrocarbons to biogenic aerosol in the Borneo rainforest

Abstract. Emissions of biogenic volatile organic compounds are though to contribute significantly to secondary organic aerosol formation in the tropics, but understanding these transformation processes has proved difficult, due to the complexity of the chemistry involved and very low concentrations. Aerosols from above a Southeast Asian tropical rainforest in Borneo were characterised using liquid chromatography–ion trap mass spectrometry, high-resolution aerosol mass spectrometry and Fourier transform ion cyclotron resonance mass spectrometry (FTICRMS) techniques. Oxygenated compounds were identified in ambient organic aerosol that could be directly traced back to isoprene, monoterpenes and sesquiterpene emissions, by combining field data on chemical structures with mass spectral data generated from synthetically produced products created in a simulation chamber. Eighteen oxygenated species of biogenic origin were identified in the rainforest aerosol from the precursors isoprene, α-pinene, limonene, α-terpinene and β-caryophyllene. The observations provide the unambiguous field detection of monoterpene and sesquiterpene oxidation products in SOA above a pristine tropical rainforest. The presence of 2-methyl tetrol organosulfates and an associated sulfated dimer provides direct evidence that isoprene in the presence of sulfate aerosol can make a contribution to biogenic organic aerosol above tropical forests. High-resolution mass spectrometry indicates that sulfur can also be incorporated into oxidation products arising from monoterpene precursors in tropical aerosol.

OH reactivity in a South East Asian tropical rainforest during the Oxidant and Particle Photochemical Processes (OP3) project

Abstract. OH (hydroxyl radical) reactivity, the inverse of the chemical lifetime of the hydroxyl radical, was measured for 12 days in April 2008 within a tropical rainforest on Borneo as part of the OP3 (Oxidant and Particle Photochemical Processes) project. The maximum observed value was 83.8 ± 26.0 s−1 with the campaign averaged noontime maximum being 29.1 ± 8.5 s−1. The maximum OH reactivity calculated using the diurnally averaged concentrations of observed sinks was ~ 18 s−1, significantly less than the observations, consistent with other studies in similar environments. OH reactivity was dominated by reaction with isoprene (~ 30%). Numerical simulations of isoprene oxidation using the Master Chemical Mechanism (v3.2) in a highly simplified physical and chemical environment show that the steady state OH reactivity is a linear function of the OH reactivity due to isoprene alone, with a maximum multiplier, to account for the OH reactivity of the isoprene oxidation products, being equal to the number of isoprene OH attackable bonds (10). Thus the emission of isoprene constitutes a significantly larger emission of reactivity than is offered by the primary reaction with isoprene alone, with significant scope for the secondary oxidation products of isoprene to constitute the observed missing OH reactivity. A physically and chemically more sophisticated simulation (including physical loss, photolysis, and other oxidants) showed that the calculated OH reactivity is reduced by the removal of the OH attackable bonds by other oxidants and photolysis, and by physical loss (mixing and deposition). The calculated OH reactivity is increased by peroxide cycling, and by the OH concentration itself. Notable in these calculations is that the accumulated OH reactivity from isoprene, defined as the total OH reactivity of an emitted isoprene molecule and all of its oxidation products, is significantly larger than the reactivity due to isoprene itself and critically depends on the chemical and physical lifetimes of intermediate species. When constrained to the observed diurnally averaged concentrations of primary VOCs (volatile organic compounds), O3, NOx and other parameters, the model underestimated the observed diurnal mean OH reactivity by 30%. However, it was found that (1) the short lifetimes of isoprene and OH, compared to those of the isoprene oxidation products, lead to a large variability in their concentrations and so significant variation in the calculated OH reactivity; (2) uncertainties in the OH chemistry in these high isoprene environments can lead to an underestimate of the OH reactivity; (3) the physical loss of species that react with OH plays a significant role in the calculated OH reactivity; and (4) a missing primary source of reactive carbon would have to be emitted at a rate equivalent to 50% that of isoprene to account for the missing OH sink. Although the presence of unmeasured primary emitted VOCs contributing to the measured OH reactivity is likely, evidence that these primary species account for a significant fraction of the unmeasured reactivity is not found. Thus the development of techniques for the measurement of secondary multifunctional carbon compounds is needed to close the OH reactivity budget.

半島マレーシア熱帯雨林はどのようにして気候変動に対して熱・水・二酸化炭素フラックスの平衡を保っているか？

半島マレーシア熱帯雨林はどのようにして気候変動に対して熱・水・二酸化炭素フラックスの平衡を保っているか？

DOAS measurements of formaldehyde and glyoxal above a south-east Asian tropical rainforest

Abstract. Tropical rainforests act as a huge contributor to the global emissions of biogenic volatile organic compounds (BVOCs). Measurements of their oxidation products, such as formaldehyde (HCHO) and glyoxal (CHOCHO), provide useful indicators of fast photochemistry occurring in the lower troposphere. However, measurements of these species in tropical forest locations are extremely limited. To redress this, HCHO and CHOCHO were measured using the long-path (LP) and multi-axis (MAX) differential optical absorption spectroscopy (DOAS) techniques above the rainforest canopy in Borneo during two campaigns in spring and summer 2008, as part of the Oxidant and Particle Photochemical Processes above a south-east Asian tropical rainforest (OP3) project. The results were compared with concurrent measurements of hydroxyl radical (OH), isoprene (C5H8) (which was the dominant organic species emitted in this forest environment), and various meteorological parameters. Formaldehyde was observed at a maximum concentration of 4.5 ppb and glyoxal at a maximum of 1.6 ppb, significantly higher than previous measurements in rural locations. A 1-D chemistry model was then used to assess the diurnal evolution of formaldehyde and glyoxal throughout the boundary layer. The results, which compare well with the LP-DOAS and MAX-DOAS observations, suggest that the majority of the glyoxal and formaldehyde is confined to the first 500 m of the boundary layer, and that the measured ratio of these species is reproduced using currently accepted product yields for the oxidation of isoprene by OH. An important conclusion is that the measured levels of glyoxal are consistent with the surprisingly high concentrations of OH measured in this environment.

Vertical variation in leaf gas exchange parameters for a Southeast Asian tropical rainforest in Peninsular Malaysia

Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.

A nocturnal Provespa wasp species as the probable pollinator of epiphytic orchid Coelogyne fimbriata

AbstractMany vespid wasps visit flowers to forage nectar. These hymenopterans sometimes contribute to flower pollination. However, none of the nocturnal wasp species is a known pollinator. We collected individuals of light‐attracted Provespa nocturna workers in a montane rainforest on Peninsular Malaysia: some wasps collected bore orchid pollinia on their thoraxes. Among 114 trapped individuals, four bore pollinaria and nine bore only viscidia, suggesting that pollinia had been successfully transported. Molecular barcoding of the pollinia (based on their ITS sequences) assigned the orchid to a species in Coelogyne fimbriata complex. These findings and our other analyses suggest that this nocturnal wasp contributes to pollination of an epiphytic nectarless orchid that probably releases olfactory attractants. This discovery sheds light on the importance of mutualistic relationships between the nocturnal social wasps and epiphytic orchids in Southeast Asian tropical rainforest canopies.

In situ measurements of isoprene and monoterpenes within a south-east Asian tropical rainforest

Abstract. Biogenic volatile organic compounds (BVOCs) emitted from tropical rainforests comprise a substantial fraction of global atmospheric VOC emissions, however there are only relatively limited measurements of these species in tropical rainforest regions. We present observations of isoprene, α-pinene, camphene, Δ-3-carene, γ-terpinene and limonene, as well as oxygenated VOCs (OVOCs) of biogenic origin such as methacrolein, in ambient air above a tropical rainforest in Malaysian Borneo during the Oxidant and Particle Photochemical Processes above a south-east Asian tropical rainforest (OP3) project in 2008. Daytime composition was dominated by isoprene, with an average mixing ratio of the order of ~1 ppb. γ-terpinene, limonene and camphene were the most abundant monoterpenes, with average daytime mixing ratios of 102, 71 and 66 ppt respectively, and with an average monoterpene toisoprene ratio of 0.3 during sunlit hours, compared to 2.0 at night. Limonene and camphene abundances were seen to be related to both temperature and light conditions. In contrast, γ-terpinene emission continued into the late afternoon/evening, under relatively low temperature and light conditions. The contributions of isoprene, monoterpenes and other classes of VOC to the volatile carbon budget and OH reactivity have been summarised for this rainforest location. We observe good agreement between surface and aircraft measurements of boundary layer isoprene and methacrolein above the natural rainforest, suggesting that the ground-level observations are broadly representative of isoprene emissions from this region.

Isoprene oxidation mechanisms: measurements and modelling of OH and HO<sub>2</sub> over a South-East Asian tropical rainforest during the OP3 field campaign

Abstract. Forests are the dominant source of volatile organic compounds into the atmosphere, with isoprene being the most significant species. The oxidation chemistry of these compounds is a significant driver of local, regional and global atmospheric composition. Observations made over Borneo during the OP3 project in 2008, together with an observationally constrained box model are used to assess our understanding of this oxidation chemistry. In line with previous work in tropical forests, we find that the standard model based on MCM chemistry significantly underestimates the observed OH concentrations. Geometric mean observed to modelled ratios of OH and HO2 in airmasses impacted with isoprene are 5.32−4.43+3.68 and 1.18−0.30+0.30 respectively, with 68 % of the observations being within the specified variation. We implement a variety of mechanistic changes into the model, including epoxide formation and unimolecular decomposition of isoprene peroxy radicals, and assess their impact on the model success. We conclude that none of the current suggestions can simultaneously remove the bias from both OH and HO2 simulations and believe that detailed laboratory studies are now needed to resolve this issue.

JENUFA GEN. NOV.: A NEW GENUS OF COCCOID GREEN ALGAE (CHLOROPHYCEAE, INCERTAE SEDIS) PREVIOUSLY RECORDED BY ENVIRONMENTAL SEQUENCING(1).

The diversity of eukaryotic microorganisms is far from fully described, as indicated by the vast number of unassigned genotypes retrieved by environmental sequencing or metagenomics. We isolated several strains of unicellular green algae from algal biofilms growing on tree bark in a Southeast Asian tropical rainforest and determined them to be relatives of an unidentified lineage of environmental 18S rDNA sequences, thus uncovering its cellular identity. Light, confocal, and electron microscope observations and sequencing the 18S rRNA gene revealed that the strains represent two different species within an apparently new genus, described here as Jenufa gen. nov. Both species formed minute coccoid cells with an irregular globular outline, a smooth cell wall, and a single parietal chloroplast without a pyrenoid. The two species, described herein as J. perforata and J. minuta, differed in chloroplast morphology and cell wall structure. Phylogenetic analyses of 18S rRNA gene sequences showed a firm relationship between the two species and placed the Jenufa lineage in an unresolved position within the CS clade (Chlamydomonadales + Sphaeropleales) of the class Chlorophyceae, although possible affinities to the genus Golenkinia were suggested both by maximum-likelihood (ML) and Bayesian methods. Furthermore, two almost identical environmental 18S rDNA sequences from an endolithic microbial community occurring in dolomite rock in the central Alps turned out to be specifically related to, yet apparently distinct from, the sequence of J. minuta, indicating the existence of an undescribed Jenufa species occurring in the temperate zone.

Fluxes and concentrations of volatile organic compounds from a South-East Asian tropical rainforest

Abstract. As part of the OP3 field study of rainforest atmospheric chemistry, above-canopy fluxes of isoprene, monoterpenes and oxygenated volatile organic compounds were made by virtual disjunct eddy covariance from a South-East Asian tropical rainforest in Malaysia. Approximately 500 hours of flux data were collected over 48 days in April–May and June–July 2008. Isoprene was the dominant non-methane hydrocarbon emitted from the forest, accounting for 80% (as carbon) of the measured emission of reactive carbon fluxes. Total monoterpene emissions accounted for 18% of the measured reactive carbon flux. There was no evidence for nocturnal monoterpene emissions and during the day their flux rate was dependent on both light and temperature. The oxygenated compounds, including methanol, acetone and acetaldehyde, contributed less than 2% of the total measured reactive carbon flux. The sum of the VOC fluxes measured represents a 0.4% loss of daytime assimilated carbon by the canopy, but atmospheric chemistry box modelling suggests that most (90%) of this reactive carbon is returned back to the canopy by wet and dry deposition following chemical transformation. The emission rates of isoprene and monoterpenes, normalised to 30 °C and 1000 μmol m−2 s−1 PAR, were 1.6 mg m−2 h−1 and 0.46mg m−2 h−1 respectively, which was 4 and 1.8 times lower respectively than the default value for tropical forests in the widely-used MEGAN model of biogenic VOC emissions. This highlights the need for more direct canopy-scale flux measurements of VOCs from the world's tropical forests.

Simulating atmospheric composition over a South-East Asian tropical rainforest: performance of a chemistry box model

Abstract. Atmospheric composition and chemistry above tropical rainforests is currently not well established, particularly for south-east Asia. In order to examine our understanding of chemical processes in this region, the performance of a box model of atmospheric boundary layer chemistry is tested against measurements made at the top of the rainforest canopy near Danum Valley, Malaysian Borneo. Multi-variate optimisation against ambient concentration measurements was used to estimate average canopy-scale emissions for isoprene, total monoterpenes and nitric oxide. The excellent agreement between estimated values and measured fluxes of isoprene and total monoterpenes provides confidence in the overall modelling strategy, and suggests that this method may be applied where measured fluxes are not available, assuming that the local chemistry and mixing are adequately understood. The largest contributors to the optimisation cost function at the point of best-fit are OH (29%), NO (22%) and total peroxy radicals (27%). Several factors affect the modelled VOC chemistry. In particular concentrations of methacrolein (MACR) and methyl-vinyl ketone (MVK) are substantially overestimated, and the hydroxyl radical (OH) concentration is substantially underestimated; as has been seen before in tropical rainforest studies. It is shown that inclusion of dry deposition of MACR and MVK and wet deposition of species with high Henry's Law values substantially improves the fit of these oxidised species, whilst also substantially decreasing the OH sink. Increasing OH production arbitrarily, through a simple OH recycling mechanism , adversely affects the model fit for volatile organic compounds (VOCs). Given the constraints on isoprene flux provided by measurements, a substantial decrease in the rate of reaction of VOCs with OH is the only remaining option to explain the measurement/model discrepancy for OH. A reduction in the isoprene+OH rate constant of 50%, in conjunction with increased deposition of intermediates and some modest OH recycling, is able to produce both isoprene and OH concentrations within error of those measured. Whilst we cannot rule out an important role for missing chemistry, particularly in areas of higher isoprene flux, this study demonstrates that the inadequacies apparent in box and global model studies of tropical VOC chemistry may be more strongly influenced by representation of detailed physical and micrometeorological effects than errors in the chemical scheme.

Overview: oxidant and particle photochemical processes above a south-east Asian tropical rainforest (the OP3 project): introduction, rationale, location characteristics and tools

Abstract. In April–July 2008, intensive measurements were made of atmospheric composition and chemistry in Sabah, Malaysia, as part of the "Oxidant and particle photochemical processes above a South-East Asian tropical rainforest" (OP3) project. Fluxes and concentrations of trace gases and particles were made from and above the rainforest canopy at the Bukit Atur Global Atmosphere Watch station and at the nearby Sabahmas oil palm plantation, using both ground-based and airborne measurements. Here, the measurement and modelling strategies used, the characteristics of the sites and an overview of data obtained are described. Composition measurements show that the rainforest site was not significantly impacted by anthropogenic pollution, and this is confirmed by satellite retrievals of NO2 and HCHO. The dominant modulators of atmospheric chemistry at the rainforest site were therefore emissions of BVOCs and soil emissions of reactive nitrogen oxides. At the observed BVOC:NOx volume mixing ratio (~100 pptv/pptv), current chemical models suggest that daytime maximum OH concentrations should be ca. 105 radicals cm−3, but observed OH concentrations were an order of magnitude greater than this. We confirm, therefore, previous measurements that suggest that an unexplained source of OH must exist above tropical rainforest and we continue to interrogate the data to find explanations for this.

Pollen foraging and resource partitioning of stingless bees in relation to flowering dynamics in a Southeast Asian tropical rainforest

We used microscopic pollen analysis to investigate the diversity and similarity of pollen diets of six colonies of stingless bees (Apidae; Meliponini) located within one monospecific (three colonies of Trigona collina) and one mixed nesting aggregation (one colony of T. collina, and one colony of each of the close relatives T. melina and T. melanocephala) in lowland tropical rain forest in Sabah, Malaysia. Samples of 20 corbicular loads, collected six times over a period of three months from each colony, contained a total of 74 different morphotypes of pollen grains with an average between 4.7 to 8.5 per sample for the different colonies. In an analysis on total diet composition intraspecific similarity was much greater than interspecific similarity. The focal colony of Trigona collina from the mixed aggregation distinctly clustered according to species rather than nest location, suggesting that some interspecific resource partitioning occurs. The sampling period was accompanied by a drastic increase in flowering activity as evidenced by data from a flower phenology transect. At times of limited flowering similarity of pollen diets was generally low, both within and between species. It is hypothesized that this is so because bees are forced to forage from scattered subsets of flower patches spread out over a large foraging range. In times of increased flowering pollen diet similarity significantly increased between colonies of the monospecific aggregation, presumably because colonies concentrated on more profitable sources in closer proximity. In contrast, similarity remained low within the mixed aggregation, suggesting that innate differences in foraging preferences precluded any effect of diet convergence.

Use of multitemporal ERS-2 SAR images for identification of burned scars in south-east Asian tropical rainforest

Evaluation of an area severely affected by fires in 1998 using a multitemporal series of ERS-2 Synthetic Aperture Radar (SAR) images showed that fire induced changes of the vegetation cover strongly affected C-band radar backscatter. We investigated the changes in radar backscatter over a period of ten months in areas of interest that represented different land-cover types at a study site in East Kalimantan, Indonesia. The impact of fire was found to cause a strong decrease in backscatter (2-5 dB) for all land-cover classes while areas not affected by fire showed only slight variations in backscatter (maximum 0.5 dB). Ground and aerial evidence suggests that the marked decrease in backscatter can be attributed to the removal of the vegetation cover and subsequently higher contribution of backscatter from dry soil. After the onset of rain the radar backscatter increased to 5.5 dB in areas severely affected by fire while in unburned forests it returned to values similar to those before the drought. Burned scars could be identified visually in multitemporal principal component analysis-enhanced ERS SAR colour composites.

The influence of ants on patterns of colonization and establishment within a set of coexisting lycaenid butterflies in a south-east Asian tropical rain forest.

In Peninsular Malaysia ten species of lycaenid butterflies use leaf flushes or inflorescences of the legume tree Saraca thaipingensis as larval hostplant. Resource partitioning among these species is regulated by a complex mixture of patterns of interaction with ants. Females of obligately myrmecophilous species lay their eggs exclusively on trees colonized by their specific host ants. On trees colonized by weaver ants, only specialist mutualists adapted to these territorial ants are able to survive, while larvae of other species are killed. The formicine ant Cladomyrma petalae, which inhabits hollow twigs of the myrmecophytic hostplant, likewise precludes oviposition by female butterflies. Lycaenid larvae confronted with this ant species never survive, but one concealed feeding species (Jamides caeruleus) escapes removal due to the cryptic life-habits of the larvae. Two facultative myrmecophiles associate in a mutualistic way with a wide and largely overlapping range of ant genera which forage at the extrafloral nectaries of leaf flushes. One species (Cheritra freja) is not myrmecophilous, but is tolerated by all but the most territorial ants. Ant-dependent hostplant selection and egg-clustering characterize the obligate mutualists, whereas facultative myrmecophiles and the non-myrmecophile distribute their eggs singly over appropriate hostplants. Signals mediating caterpillar-ant communication are highly specialized in one obligate myrmecophile (Drupadia theda), but rather unspecific in four other species tested. Altogether our observations indicate that colonization and establishment of lycaenid butterflies on S. thaipingensis trees are governed by specializations as well as opportunistic use of resources (ants and hostplant parts). Therefore, the diversity of this species assemblage is maintained by deterministic as well as stochastic factors.

Egg Clustering and the Southern Hemisphere Lycaenids: Comments on a Paper by N. E. Stamp

Previous articleNext article No AccessNotes and CommentsEgg Clustering and the Southern Hemisphere Lycaenids: Comments on a Paper by N. E. StampR. L. KitchingR. L. Kitching Search for more articles by this author PDFPDF PLUS Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinkedInRedditEmail SectionsMoreDetailsFiguresReferencesCited by The American Naturalist Volume 118, Number 3Sep., 1981 Published for The American Society of Naturalists Article DOIhttps://doi.org/10.1086/283833 Views: 2Total views on this site Citations: 13Citations are reported from Crossref Copyright 1981 The University of ChicagoPDF download Crossref reports the following articles citing this article:Daniela Rodrigues, André V. L. Freitas Contrasting egg and larval performances help explain polyphagy in a florivorous butterfly, Arthropod-Plant Interactions 7, no.22 (Oct 2012): 159–167.https://doi.org/10.1007/s11829-012-9230-3Hans Van Dyck, Sofie Regniers Egg spreading in the ant-parasitic butterfly, Maculinea alcon: from individual behaviour to egg distribution pattern, Animal Behaviour 80, no.44 (Oct 2010): 621–627.https://doi.org/10.1016/j.anbehav.2010.06.021Rod Eastwood, Ann M. Fraser Associations between lycaenid butterflies and ants in Australia, Austral Ecology 24, no.55 (Oct 1999): 503–537.https://doi.org/10.1046/j.1440-169x.1999.01000.xPeter Seufert, Konrad Fiedler The influence of ants on patterns of colonization and establishment within a set of coexisting lycaenid butterflies in a south-east Asian tropical rain forest, Oecologia 106, no.11 (Apr 1996): 127–136.https://doi.org/10.1007/BF00334414MERRILL A. PETERSON Unpredictability in the facultative association between larvae of Euphilotes enoptes (Lepidoptera: Lycaenidae) and ants, Biological Journal of the Linnean Society 55, no.33 (Jan 2008): 209–223.https://doi.org/10.1111/j.1095-8312.1995.tb01060.xRonald No�, Peter Hammerstein Biological markets: supply and demand determine the effect of partner choice in cooperation, mutualism and mating, Behavioral Ecology and Sociobiology 35, no.11 (Jul 1994): 1–11.https://doi.org/10.1007/BF00167053P. R. SAMSON MORPHOLOGY AND BIOLOGY OF ACRODZPSAS ZLLZDGEZ (WATERHOUSE AND LYELL), A MYRMECOPHAGOUS LYCAENID (LEPIDOPTERA: LYCAENIDAE: THECLINAE), Australian Journal of Entomology 28, no.33 (Aug 1989): 161–168.https://doi.org/10.1111/j.1440-6055.1989.tb00874.xN. E. Pierce, R. L. Kitching, R. C. Buckley, M. F. J. Taylor, K. F. Benbow The costs and benefits of cooperation between the Australian lycaenid butterfly, Jalmenus evagoras, and its attendant ants, Behavioral Ecology and Sociobiology 21, no.44 (Oct 1987): 237–248.https://doi.org/10.1007/BF00292505R. E. Jones Behavioural evolution in the cabbage butterfly (Pieris rapae), Oecologia 72, no.11 (Apr 1987): 69–76.https://doi.org/10.1007/BF00385047R. Buckley Ant-Plant-Homopteran Interactions, (Jan 1987): 53–85.https://doi.org/10.1016/S0065-2504(08)60087-2Naomi E. Pierce, Mark A. Elgar The influence of ants on host plant selection by Jalmenus evagoras, a myrmecophilous lycaenid butterfly, Behavioral Ecology and Sociobiology 16, no.33 (Mar 1985): 209–222.https://doi.org/10.1007/BF00310983 Steven P. Courtney The Evolution of Egg Clustering by Butterflies and Other Insects, The American Naturalist 123, no.22 (Oct 2015): 276–281.https://doi.org/10.1086/284202C. B. COTTRELL Aphytophagy in butterflies: its relationship to myrmecophily, Zoological Journal of the Linnean Society 80, no.11 (Jun 2008): 1–57.https://doi.org/10.1111/j.1096-3642.1984.tb02318.x