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New paper on spatial synchrony of forest tree population dynamics published in Proceedings of the Royal Society B

Light red meranti (Shorea parvifolia) is a canopy tree species native to tropical forests in Southeast Asia that can grow to over 65 m. In the two decades to 2005, the abundance of light red meranti decreased by 60% in a ForestGEO research plot in Pasoh, Malaysia. In a ForestGEO plot in Lambir, Malaysia, over 1000 km away, the same species decreased by 25% over a similar period. This example prompts the question of whether forest tree population dynamics are in general synchronised across space. In a paper just published in Proceedings of the Royal Society B, we found that the answer is yes.

We collated forest inventory data at three different scales—local, regional and global—and found signals of synchrony up to scales well beyond 100 km. For the Pasoh and Lambir forests, in particular, there was clear evidence of synchrony for larger trees (greater than 10 cm diameter) although not for smaller trees. A technical challenge in our analyses was developing statistical methods to produce an aggregate estimate of synchrony for each pair of forest sites while accounting for the noise inherent in datasets with large numbers of rare species. Although existing methods are available to estimate synchrony for individual species, these were not appropriate for our forest data because of the short duration of our datasets (a few decades at most) relative to the typical tree generation time. Our novel methods make use of copulas, which are used to model multivariate statistical phenomena in quantitative finance.

We attributed the observed synchrony in tree population dynamics to synchronised environmental drivers, such as climate, although we lacked the statistical power to identify particular drivers. The observed synchrony suggests greater risk of extinction for tree species that are more spatially constrained, especially those with range dimensions less than ~100 km. On the other hand, species with larger ranges may be buffered from extinction by the lack of synchrony in their population dynamics across space.

Forest tree population dynamics are synchronised across space, with synchrony detectable even beyond 100 km. Dark blue, light blue and green curves indicate the results of local, regional and global analyses. Line types indicate the spatial grain of each analysis: synchrony is lower for smaller spatial grains because of immigration.

Chisholm, R. A., T. Fung, K. J. Anderson-Teixeira, N. A. Bourg, W. Y. Brockelman, S. Bunyave- jchewin, C.-H. Chang-Yang, Y.-Y. Chen, G. B. Chuyong, R. Condit, H. S. Dattaraja, S. J. Davies, S. Ediriweera, C. E. N. Ewango, E. S. Fernando, I. A. U. N. Gunatilleke, C. V. S. Gunatilleke, Z. Hao, R. W. Howe, D. Kenfack, T. L. Yao, J.-R. Makana, S. M. McMahon, X. Mi, M. Bt. Mohamad, J. A. Myers, A. Nathalang, Á. J. Pérez, S. Phumsathan, N. Pongpattananurak, H. Ren, L. J. V. Rodriguez, R. Sukumar, I.-F. Sun, H. S. Suresh, D. W. Thomas, J. Thompson, M. Uriarte, R. Valencia, X. Wang, A. T. Wolf, and J. K. Zimmerman (2024) Assessing the spatial scale of synchrony in forest tree population dynamics. Proceeding of the Royal Society B 291: 20240486

Fanhua’s paper on tree diversity in gaps in tropical forests published in Ecology

Kong Fanhua was a visiting PhD student in our lab in 2022–2023, during which time she worked on a study of tropical forest gap diversity that has just been published in Ecology. Forest gaps are created when large trees fall. Gaps experience greater below-canopy light levels than non-gaps and thus see increased recruitment of young trees. It has long been hypothesised that forest gaps have higher tree diversity than non-gaps, but surprisingly until now there has not been a satisfactory statistical test of the hypothesis. We focussed our analyses on the 50 ha forest plot on Barro Colorado Island in Panama, leveraging regular forest census data going back decades, as well as canopy-height survey data that facilitated gap identification.

Our main finding was that although the average number of tree species in any given gap was only slightly higher than a same-sized non-gap site, collections of gaps had many more species than similar collections of non-gaps. We identified 124 newly formed gaps (each 25 m2) in 2003 and when these were seven years old they contained 149 tree species in total compared to only 109 tree species in a comparable collection of non-gap sites. Qualitatively similar, but somewhat weaker, results were observed when the gaps were two and 12 years old. These notable differences between gaps and non-gaps strongly indicate an important role for gaps in the maintenance of plot-level tree diversity.

Fanhua now holds a post-doctoral research position at the Nanjing Institute of Environmental Sciences.

Kong, F., H. Fangliang, and R. A. Chisholm. (in press) High beta diversity of gaps contributes to plot-level tree diversity in a tropical forest. Ecology

A tropical forest gap (photo credit: Rachel Lim)

Sean’s paper on conserving Southeast Asian trees published in Nature Sustainability

Sean Pang, who completed his PhD in our lab in 2022, has successfully published a chapter from this thesis about conserving Southeast Asian tree species in the face of both climate and land-use change. He modelled nearly 1,500 tree species distributions and explored how they would fare under four future climate-change scenarios involving different assumptions about human mitigation efforts. As expected, the tree species overall fared best under the most sustainable climate pathway (SSP1-2.6), but surprisingly the intermediate climate pathways (SSP2-4.5 and SSP3-7.0) were worse for biodiversity than the least-sustainable climate pathway (SSP5-8.5). The main reason was that the intermediate climate pathways involved greater land-use change for biofuel production to support a partial transition away from fossil fuels. The results demonstrate how halfway efforts to prevent climate change can potentially have detrimental effects on biodiversity. The paper is just out in the journal Nature Sustainability.

Sean was originally a student in Edward Webb’s lab, but completed the last year of his PhD in our lab after Webb moved to the University of Helsinki. Sean is now a post-doctoral research fellow at Aarhus University in Denmark.

Pang, S. E. H., J. W. F. Slik, R. A. Chisholm, and E. L. Webb. Conserving Southeast Asian trees requires mitigating both climate and land-use change. Nature Sustainability

Sankar wins best poster in the multi-taxa category at the World Congress of Herpetology in Kuching, Sarawak

From 5th to 9th August 2024, Sankar attended the 10th World Congress of Herpetology, held in Kuching, Sarawak. This is a meeting of global herpetologists held every four years. Sankar presented a poster on his work tracking squamate (lizard and snake) extirpations in Singapore using the SEUX and MODGEE models developed in our lab.

The SEUX model takes in the first and last years of detection for the species in a group and estimates the rates of known extinctions and dark extinctions (i.e., extinction prior to discovery) that occur within that group. The MODGEE model takes advantage of the entire detection time series and can additionally estimate the extinction probability of individual species and the time of extirpation. A recent comprehensive compilation of herpetofaunal records in Singapore has made it possible to use MODGEE to obtain these estimates for squamates (snakes and lizards). The resulting assessments of which species are extinct can be valuable in planning species reintroductions in the future.

The conference was well attended (~1500 delegates) and there were many opportunities to engage and network with other herpetologists from around the world working on conservation translocations, capture-mark-recapture monitoring and population viability analysis.

At the end of the conference, Sankar was awarded the best poster in the multi-taxa category. Congratulations, Sankar!

Tak attends the Society for Mathematical Biology meeting in Seoul

In the first week of July, Tak attended the Joint Annual Meeting of the Korean Society for Mathematical Biology and the Society for Mathematical Biology, held in Seoul. During the conference, he presented some new results on deriving probability distributions of extinction time for a population model with a growth rate that varies stochastically due to environmental fluctuations. This is an important topic for conservation because many species are in danger of extinction and it is unclear how environmental changes over time affect their extinction risk.

Specifically, the environmental fluctuations in the model are assumed to follow a random process that is called 1/f noise, following empirical evidence that many environmental variables in reality follow such a random process. Analyses of the model show that the mean extinction time should increase with the autocorrelation time of environmental fluctuations, because of non-linear averaging of the proportional growth rate over different sets of environmental conditions (technically, this is an application of Jensen’s inequality to the proportional growth rate). Because environmental variables in marine ecosystems tend to have longer autocorrelation times than in terrestrial systems, our model predicts that the mean extinction time would generally be greater for marine versus terrestrial populations, provided that the populations are comparable in other aspects of their biology.

In addition, Tak engaged with numerous other researchers and exchanged thoughts on different areas in mathematical biology, including but not limited to ecology and evolution. In particular, he met up with Nao Takashina from the University of Tokyo, who collaborated with Tak and Ryan on a recent paper, published in the Journal of Ecology, on mechanistic partitioning of species richness in tropical forest tree communities.

The conference was held at the Seoul campus of Konkuk University (photo used under a CC BY-SA 4.0 license, with credit to Konkukeditor).

Ryan gives an online seminar at Alphabet about our ecological modelling work

As part of a talk series on modelling physical, biological and social phenomena run by Alphabet (Google’s parent company), Ryan presented an online seminar about our lab’s work on ecological modelling. He talked about our Singapore extinctions project, our research programme on the maintenance of diversity, and potential novel applications of optimised individual-based ecological models to video games and virtual reality applications.

Special Programme in Science students win best presentation award for their Integrated Science Project

Over the last six months, three undergraduate Special Programme in Science students, Chin Zhen Jie, Andrew Teoh, and Cheng Jun Yuan (pictured left to right, below), have been conducting a project in our lab for their Integrated Science Project. They built a computational model to explore the joint effect of stage structure and conspecific negative density dependence on forest tree biodiversity. Their main result was that stage structure enhanced the diversity generated by density dependence. They recently completed their report, underwent a viva voce, gave a poster presentation, and gave a slide presentation at the Special Programme in Science student congress. For the latter, they won the best presentation award. Congratulations, guys!

The Special Programme in Science was established in 1996 at NUS to nurture enthusiastic students with a passion for science. The students take six tailor-made courses over the first two years of their degree, including the Integrated Science Project in the second year.

New modelling paper on impact of non-pharmaceutical interventions on total disease burden published in the Journal of Theoretical Biology

Infectious diseases exert a large burden on the well-being of human societies, as vividly illustrated by the recent COVID-19 pandemic. In response to an infectious disease, governments enact a variety of non-pharmaceutical interventions (NPIs) such as social-distancing guidelines, mask-wearing mandates and lockdowns. These NPIs reduce spread of the virus causing the target disease, and also reduce spread of other viruses that cause non-target diseases. For example, NPIs enacted during the COVID-19 pandemic reduced the spread of non-target respiratory diseases caused by influenza and respiratory syncytial virus (RSV). However, due to lack of pharmaceutical interventions aimed at these non-target diseases (such as vaccinations), they often rebounded to levels higher than pre-pandemic levels after the NPIs were relaxed. Thus, the net effect of NPIs on the long-term total disease burden of non-target diseases is unclear.

This knowledge gap was addressed in a new study by Tak, Ryan and Jonah Goh (former Honours student at Chisholm Lab), recently published in the Journal of Theoretical Biology. We considered a scenario where NPIs were enacted for a year and then completely removed. Under this scenario, we analysed a suite of four epidemiological models of varying generality and complexity, to quantify the net effect of the NPIs on the long-term total disease burden of a seasonal, non-target respiratory disease.

The simplest model of the four was a SIR model with seasonal disease outbreaks and temporary immunity of recovered individuals. For this model, we performed a rigorous mathematical analysis to show that the net effect of NPIs was always to reduce the long-term total disease burden. The number of susceptible individuals increased during the year when NPIs were enacted and this led to the number of infections rebounding to high levels after the NPIs were removed (Fig. 1). But this was more than offset by the number of infections prevented by the NPIs when they were being applied (Fig. 2). We found that the net reduction in the number of infections depended critically on the rate at which immunity was lost. In the extreme case of near-permanent immunity, the pool of susceptible individuals remained small and resulted in small outbreaks, such that the NPIs produced a small net reduction in the number of infections. However, as immunity was lost more quickly, the size of outbreaks increased commensurately and the NPIs produced increasingly larger net reductions in the number of infections, with the net reduction reaching large values of 70-100% of population size when immunity loss occurred quickly (on the order of months).

These key results from the simplest model were supported by simulations of the three more-complex models, which were parameterised for specific locations including Singapore and China. Overall, our study highlights a hitherto under-appreciated role of NPIs in reducing the long-term total disease burden of non-target diseases, which should be factored into cost-benefit assessments of NPIs in public health management.

Fung T., J. Goh, R. A. Chisholm. 2024. Long-term effects of non-pharmaceutical interventions on total disease burden in parsimonious epidemiological models.
Journal of Theoretical Biology 587:111817

Fig. 1. Time-series of the daily number of infections of a seasonal non-target respiratory disease, produced from a simple epidemiological model under (i) a scenario with no NPIs and (ii) a scenario with NPIs imposed for the first year and then subsequently removed. In this example, half of the recovered individuals at the end of each year are assumed to lose their immunity. The photo shows an influenza virus under a transmission electron microscope (photo from the public domain by the Public Health Image Library (PHIL) of the Centers for Disease Control and Prevention, under identification number #8160).

Fig. 2. Time-series of the cumulative number of infections of a seasonal non-target respiratory disease, produced from the simple epidemiological model under the two scenarios described in the caption for Fig. 1.

New coauthored Nature paper on the impacts of conspecific neighbours on forest tree survival

To explain high tropical forest tree species diversity, ecologists have hypothesised that pests, pathogens and resource limitation keep the abundances of common tree species in check, thereby allowing rare tree species to persist. If this hypothesis is true then individual trees surrounded by more neighbours of the same species should have lower survival. This is known as conspecific negative density dependence (CNDD). Past tests of the CNDD hypothesis have been criticised for statistical flaws, mainly arising from the use of static data (see here and here). In a study led by Lisa Hülsmann of the University of Bayreuth and just published in Nature, we conducted a comprehensive test of the hypothesis using a dynamic data set comprising repeated censuses of 23 forest plots from the global ForestGEO network.

We found that average CNDD across tree species is generally weak and exhibits little relationship to latitude, contrary to the hypothesis that CNDD explains high tropical tree diversity. Moreover, CNDD was extremely variable across species within a forest, which may limit its ability to maintain diversity. However, CNDD was higher for rare than common species in the tropics, suggesting that CNDD plays a stronger role in structuring abundances in the tropics. Our results paint a more nuanced picture of the role of CNDD in forest community ecology, and point to future experimental and theoretical studies that will be needed to clarify this role further.

The idea for this study arose when Ryan visited Lisa Hülsmann and Florian Hartig at the University of Regensburg, Germany in 2018. The paper is a collaboration between 52 authors, primarily associated with the different ForestGEO plots around the world.

Hülsmann, L., R. A. Chisholm, L. Comita, M. D. Visser, M. de Souza Leite, S. Aguilar, K. J. Anderson-Teixeira, N. A. Bourg, W. Y. Brockelman, S. Bunyavejchewin, N. Castaño, C.-H. Chang-Yang, G. B. Chuyong, K. Clay, S. J. Davies, A. Duque, S. Ediriweera, C. Ewango, G. S. Gilbert, J. Holík, R. W. Howe, S. P. Hubbell, A. Itoh, D. J. Johnson, D. Kenfack, K. Král, A. J. Larson, J. A. Lutz, J.-R. Makana, Y. Malhi, S. M. McMahon, W. J. McShea, M. Mohamad, M. Nasardin, A. Nathalang, N. Norden, A. A. Oliveira, R. Parmigiani, R. Perez, R. P. Phillips, N. Pongpattananurak, I.-F. Sun, M. E. Swanson, S. Tan, D. Thomas, J. Thompson, M. Uriarte, A. T. Wolf, T. L. Yao, J. K. Zimmerman, D. Zuleta, and F. Hartig. 2024. Latitudinal patterns in stabilizing density dependence of forest communities. Nature https://doi.org/10.1038/s41586-024-07118-4

Pasoh Forest Reserve, in Peninsular Malaysia, is the site of one of the 23 ForestGEO plots used in our analyses. Photo credit: ForestGEO

Aloysius’s paper on litterfall in forests in Singapore published in Journal of Tropical Ecology

Aloysius finished his PhD in our lab several years ago, and another chapter from his thesis has just been published. The paper reports on work he did assessing leaf litter turnover and nutrient dynamics in forests in Singapore, looking at differences between old-growth forests and novel forests. The novel forests have arisen in the last few decades from cleared farmland and villages. His main findings were that leaf litter decays much faster in novel forests than in old-growth forests, resulting in a litter pool only one third the size, and that phosphorus is highly elevated in the litter of novel forests. The differences may be attributable to dominance of exotic tree species in novel forests. The reduced litter pools in novel forests have negative consequences for ecosystem carbon balance and climate change mitigation. This may have broad relevance because of the increasing prevalence of novel forests across the tropics.

Teo, A, T. A. Evans, and R. A. Chisholm. Elevated litterfall phosphorus reduces litter and soil organic matter pools in exotic-dominated novel forests in Singapore. Journal of Tropical Ecology. 2024;40:e4