Browsing by Author "Poulsen, John R"

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    Above-ground biomass and structure of 260 African tropical forests.
    (Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2013-01) Lewis, Simon L; Sonké, Bonaventure; Sunderland, Terry; Begne, Serge K; Lopez-Gonzalez, Gabriela; van der Heijden, Geertje MF; Phillips, Oliver L; Affum-Baffoe, Kofi; Baker, Timothy R; Banin, Lindsay; Bastin, Jean-François; Beeckman, Hans; Boeckx, Pascal; Bogaert, Jan; De Cannière, Charles; Chezeaux, Eric; Clark, Connie J; Collins, Murray; Djagbletey, Gloria; Djuikouo, Marie Noël K; Droissart, Vincent; Doucet, Jean-Louis; Ewango, Cornielle EN; Fauset, Sophie; Feldpausch, Ted R; Foli, Ernest G; Gillet, Jean-François; Hamilton, Alan C; Harris, David J; Hart, Terese B; de Haulleville, Thales; Hladik, Annette; Hufkens, Koen; Huygens, Dries; Jeanmart, Philippe; Jeffery, Kathryn J; Kearsley, Elizabeth; Leal, Miguel E; Lloyd, Jon; Lovett, Jon C; Makana, Jean-Remy; Malhi, Yadvinder; Marshall, Andrew R; Ojo, Lucas; Peh, Kelvin S-H; Pickavance, Georgia; Poulsen, John R; Reitsma, Jan M; Sheil, Douglas; Simo, Murielle; Steppe, Kathy; Taedoumg, Hermann E; Talbot, Joey; Taplin, James RD; Taylor, David; Thomas, Sean C; Toirambe, Benjamin; Verbeeck, Hans; Vleminckx, Jason; White, Lee JT; Willcock, Simon; Woell, Hannsjorg; Zemagho, Lise
    We report above-ground biomass (AGB), basal area, stem density and wood mass density estimates from 260 sample plots (mean size: 1.2 ha) in intact closed-canopy tropical forests across 12 African countries. Mean AGB is 395.7 Mg dry mass ha⁻¹ (95% CI: 14.3), substantially higher than Amazonian values, with the Congo Basin and contiguous forest region attaining AGB values (429 Mg ha⁻¹) similar to those of Bornean forests, and significantly greater than East or West African forests. AGB therefore appears generally higher in palaeo- compared with neotropical forests. However, mean stem density is low (426 ± 11 stems ha⁻¹ greater than or equal to 100 mm diameter) compared with both Amazonian and Bornean forests (cf. approx. 600) and is the signature structural feature of African tropical forests. While spatial autocorrelation complicates analyses, AGB shows a positive relationship with rainfall in the driest nine months of the year, and an opposite association with the wettest three months of the year; a negative relationship with temperature; positive relationship with clay-rich soils; and negative relationships with C : N ratio (suggesting a positive soil phosphorus-AGB relationship), and soil fertility computed as the sum of base cations. The results indicate that AGB is mediated by both climate and soils, and suggest that the AGB of African closed-canopy tropical forests may be particularly sensitive to future precipitation and temperature changes.
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    African forest elephant movements depend on time scale and individual behavior.
    (Scientific reports, 2021-06-16) Beirne, Christopher; Houslay, Thomas M; Morkel, Peter; Clark, Connie J; Fay, Mike; Okouyi, Joseph; White, Lee JT; Poulsen, John R
    The critically endangered African forest elephant (Loxodonta cyclotis) plays a vital role in maintaining the structure and composition of Afrotropical forests, but basic information is lacking regarding the drivers of elephant movement and behavior at landscape scales. We use GPS location data from 96 individuals throughout Gabon to determine how five movement behaviors vary at different scales, how they are influenced by anthropogenic and environmental covariates, and to assess evidence for behavioral syndromes-elephants which share suites of similar movement traits. Elephants show some evidence of behavioral syndromes along an 'idler' to 'explorer' axis-individuals that move more have larger home ranges and engage in more 'exploratory' movements. However, within these groups, forest elephants express remarkable inter-individual variation in movement behaviours. This variation highlights that no two elephants are the same and creates challenges for practitioners aiming to design conservation initiatives.
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    Assessing the effects of elephant foraging on the structure and diversity of an Afrotropical forest
    (Biotropica, 2020-05-01) Rosin, Cooper; Beals, Kendall K; Belovitch, Michael W; Harrison, Ruby E; Pendred, Megan; Sullivan, Megan K; Yao, Nicolas; Poulsen, John R
    African forest elephants (Loxodonta cyclotis) are ecosystem engineers that browse and damage large quantities of vegetation during their foraging and movement. Though elephant trail networks and clearings are conspicuous features of many African forests, the consequences of elephant foraging for forest structure and diversity are poorly documented. In this study in northeastern Gabon, we compare stem size, stem density, proportional damage, species diversity, and species relative abundance of seedlings and saplings in the vicinity of seven tree species that produce elephant-preferred fruits (“elephant trees”) relative to control trees that do not. Across 34 survey trees, with a combined census area of 2.04 ha, we recorded data on 26,128 woody stems in three sizes classes. Compared with control trees, the area around elephant trees had the following: (a) a significantly greater proportion of damaged seedlings and a marginally greater proportion of damaged saplings (with 82% and 24% greater odds of damage, respectively); (b) no significant difference in stem density or species diversity; and (c) a significantly greater relative abundance of seedlings of elephant tree species. Increasing distance away from focal elephant trees was associated with significantly reduced sapling stem damage, significantly increased sapling stem density, and significantly increased sapling species diversity. Considered in sum, our results suggest that elephants can affect the structure and diversity of Afrotropical forests through their foraging activities, with some variation based on location and plant size class. Developing a more complete understanding of elephants’ ecological effects will require continued research, ideally with manipulative experiments. Abstract in French is available with online material.
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    Co-limitation towards lower latitudes shapes global forest diversity gradients.
    (Nature ecology & evolution, 2022-08-08) Liang, Jingjing; Gamarra, Javier GP; Picard, Nicolas; Zhou, Mo; Pijanowski, Bryan; Jacobs, Douglass F; Reich, Peter B; Crowther, Thomas W; Nabuurs, Gert-Jan; de-Miguel, Sergio; Fang, Jingyun; Woodall, Christopher W; Svenning, Jens-Christian; Jucker, Tommaso; Bastin, Jean-Francois; Wiser, Susan K; Slik, Ferry; Hérault, Bruno; Alberti, Giorgio; Keppel, Gunnar; Hengeveld, Geerten M; Ibisch, Pierre L; Silva, Carlos A; Ter Steege, Hans; Peri, Pablo L; Coomes, David A; Searle, Eric B; von Gadow, Klaus; Jaroszewicz, Bogdan; Abbasi, Akane O; Abegg, Meinrad; Yao, Yves C Adou; Aguirre-Gutiérrez, Jesús; Zambrano, Angelica M Almeyda; Altman, Jan; Alvarez-Dávila, Esteban; Álvarez-González, Juan Gabriel; Alves, Luciana F; Amani, Bienvenu HK; Amani, Christian A; Ammer, Christian; Ilondea, Bhely Angoboy; Antón-Fernández, Clara; Avitabile, Valerio; Aymard, Gerardo A; Azihou, Akomian F; Baard, Johan A; Baker, Timothy R; Balazy, Radomir; Bastian, Meredith L; Batumike, Rodrigue; Bauters, Marijn; Beeckman, Hans; Benu, Nithanel Mikael Hendrik; Bitariho, Robert; Boeckx, Pascal; Bogaert, Jan; Bongers, Frans; Bouriaud, Olivier; Brancalion, Pedro HS; Brandl, Susanne; Brearley, Francis Q; Briseno-Reyes, Jaime; Broadbent, Eben N; Bruelheide, Helge; Bulte, Erwin; Catlin, Ann Christine; Cazzolla Gatti, Roberto; César, Ricardo G; Chen, Han YH; Chisholm, Chelsea; Cienciala, Emil; Colletta, Gabriel D; Corral-Rivas, José Javier; Cuchietti, Anibal; Cuni-Sanchez, Aida; Dar, Javid A; Dayanandan, Selvadurai; de Haulleville, Thales; Decuyper, Mathieu; Delabye, Sylvain; Derroire, Géraldine; DeVries, Ben; Diisi, John; Do, Tran Van; Dolezal, Jiri; Dourdain, Aurélie; Durrheim, Graham P; Obiang, Nestor Laurier Engone; Ewango, Corneille EN; Eyre, Teresa J; Fayle, Tom M; Feunang, Lethicia Flavine N; Finér, Leena; Fischer, Markus; Fridman, Jonas; Frizzera, Lorenzo; de Gasper, André L; Gianelle, Damiano; Glick, Henry B; Gonzalez-Elizondo, Maria Socorro; Gorenstein, Lev; Habonayo, Richard; Hardy, Olivier J; Harris, David J; Hector, Andrew; Hemp, Andreas; Herold, Martin; Hillers, Annika; Hubau, Wannes; Ibanez, Thomas; Imai, Nobuo; Imani, Gerard; Jagodzinski, Andrzej M; Janecek, Stepan; Johannsen, Vivian Kvist; Joly, Carlos A; Jumbam, Blaise; Kabelong, Banoho LPR; Kahsay, Goytom Abraha; Karminov, Viktor; Kartawinata, Kuswata; Kassi, Justin N; Kearsley, Elizabeth; Kennard, Deborah K; Kepfer-Rojas, Sebastian; Khan, Mohammed Latif; Kigomo, John N; Kim, Hyun Seok; Klauberg, Carine; Klomberg, Yannick; Korjus, Henn; Kothandaraman, Subashree; Kraxner, Florian; Kumar, Amit; Kuswandi, Relawan; Lang, Mait; Lawes, Michael J; Leite, Rodrigo V; Lentner, Geoffrey; Lewis, Simon L; Libalah, Moses B; Lisingo, Janvier; López-Serrano, Pablito Marcelo; Lu, Huicui; Lukina, Natalia V; Lykke, Anne Mette; Maicher, Vincent; Maitner, Brian S; Marcon, Eric; Marshall, Andrew R; Martin, Emanuel H; Martynenko, Olga; Mbayu, Faustin M; Mbuvi, Musingo TE; Meave, Jorge A; Merow, Cory; Miscicki, Stanislaw; Moreno, Vanessa S; Morera, Albert; Mukul, Sharif A; Müller, Jörg C; Murdjoko, Agustinus; Nava-Miranda, Maria Guadalupe; Ndive, Litonga Elias; Neldner, Victor J; Nevenic, Radovan V; Nforbelie, Louis N; Ngoh, Michael L; N'Guessan, Anny E; Ngugi, Michael R; Ngute, Alain SK; Njila, Emile Narcisse N; Nyako, Melanie C; Ochuodho, Thomas O; Oleksyn, Jacek; Paquette, Alain; Parfenova, Elena I; Park, Minjee; Parren, Marc; Parthasarathy, Narayanaswamy; Pfautsch, Sebastian; Phillips, Oliver L; Piedade, Maria TF; Piotto, Daniel; Pollastrini, Martina; Poorter, Lourens; Poulsen, John R; Poulsen, Axel Dalberg; Pretzsch, Hans; Rodeghiero, Mirco; Rolim, Samir G; Rovero, Francesco; Rutishauser, Ervan; Sagheb-Talebi, Khosro; Saikia, Purabi; Sainge, Moses Nsanyi; Salas-Eljatib, Christian; Salis, Antonello; Schall, Peter; Schepaschenko, Dmitry; Scherer-Lorenzen, Michael; Schmid, Bernhard; Schöngart, Jochen; Šebeň, Vladimír; Sellan, Giacomo; Selvi, Federico; Serra-Diaz, Josep M; Sheil, Douglas; Shvidenko, Anatoly Z; Sist, Plinio; Souza, Alexandre F; Stereńczak, Krzysztof J; Sullivan, Martin JP; Sundarapandian, Somaiah; Svoboda, Miroslav; Swaine, Mike D; Targhetta, Natalia; Tchebakova, Nadja; Trethowan, Liam A; Tropek, Robert; Mukendi, John Tshibamba; Umunay, Peter Mbanda; Usoltsev, Vladimir A; Vaglio Laurin, Gaia; Valentini, Riccardo; Valladares, Fernando; van der Plas, Fons; Vega-Nieva, Daniel José; Verbeeck, Hans; Viana, Helder; Vibrans, Alexander C; Vieira, Simone A; Vleminckx, Jason; Waite, Catherine E; Wang, Hua-Feng; Wasingya, Eric Katembo; Wekesa, Chemuku; Westerlund, Bertil; Wittmann, Florian; Wortel, Verginia; Zawiła-Niedźwiecki, Tomasz; Zhang, Chunyu; Zhao, Xiuhai; Zhu, Jun; Zhu, Xiao; Zhu, Zhi-Xin; Zo-Bi, Irie C; Hui, Cang
    The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.
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    Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?
    (The New phytologist, 2013-04) Moles, Angela T; Peco, Begoña; Wallis, Ian R; Foley, William J; Poore, Alistair GB; Seabloom, Eric W; Vesk, Peter A; Bisigato, Alejandro J; Cella-Pizarro, Lucrecia; Clark, Connie J; Cohen, Philippe S; Cornwell, William K; Edwards, Will; Ejrnaes, Rasmus; Gonzales-Ojeda, Therany; Graae, Bente J; Hay, Gregory; Lumbwe, Fainess C; Magaña-Rodríguez, Benjamín; Moore, Ben D; Peri, Pablo L; Poulsen, John R; Stegen, James C; Veldtman, Ruan; von Zeipel, Hugo; Andrew, Nigel R; Boulter, Sarah L; Borer, Elizabeth T; Cornelissen, Johannes HC; Farji-Brener, Alejandro G; DeGabriel, Jane L; Jurado, Enrique; Kyhn, Line A; Low, Bill; Mulder, Christa PH; Reardon-Smith, Kathryn; Rodríguez-Velázquez, Jorge; De Fortier, An; Zheng, Zheng; Blendinger, Pedro G; Enquist, Brian J; Facelli, Jose M; Knight, Tiffany; Majer, Jonathan D; Martínez-Ramos, Miguel; McQuillan, Peter; Hui, Francis KC
    Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.
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    Deadwood stocks increase with selective logging and large tree frequency in Gabon.
    (Glob Chang Biol, 2017-04) Carlson, Ben S; Koerner, Sally E; Medjibe, Vincent P; White, Lee JT; Poulsen, John R
    Deadwood is a major component of aboveground biomass (AGB) in tropical forests and is important as habitat and for nutrient cycling and carbon storage. With deforestation and degradation taking place throughout the tropics, improved understanding of the magnitude and spatial variation in deadwood is vital for the development of regional and global carbon budgets. However, this potentially important carbon pool is poorly quantified in Afrotropical forests and the regional drivers of deadwood stocks are unknown. In the first large-scale study of deadwood in Central Africa, we quantified stocks in 47 forest sites across Gabon and evaluated the effects of disturbance (logging), forest structure variables (live AGB, wood density, abundance of large trees), and abiotic variables (temperature, precipitation, seasonality). Average deadwood stocks (measured as necromass, the biomass of deadwood) were 65 Mg ha-1 or 23% of live AGB. Deadwood stocks varied spatially with disturbance and forest structure, but not abiotic variables. Deadwood stocks increased significantly with logging (+38 Mg ha-1 ) and the abundance of large trees (+2.4 Mg ha-1 for every tree >60 cm dbh). Gabon holds 0.74 Pg C, or 21% of total aboveground carbon in deadwood, a threefold increase over previous estimates. Importantly, deadwood densities in Gabon are comparable to those in the Neotropics and respond similarly to logging, but represent a lower proportion of live AGB (median of 18% in Gabon compared to 26% in the Neotropics). In forest carbon accounting, necromass is often assumed to be a constant proportion (9%) of biomass, but in humid tropical forests this ratio varies from 2% in undisturbed forest to 300% in logged forest. Because logging significantly increases the deadwood carbon pool, estimates of tropical forest carbon should at a minimum use different ratios for logged (mean of 30%) and unlogged forests (mean of 18%).
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    Defaunation of large mammals alters understory vegetation and functional importance of invertebrates in an Afrotropical forest
    (Biological Conservation, 2020-01-01) Lamperty, Therese; Zhu, Kai; Poulsen, John R; Dunham, Amy E
    Hunting has reduced or eliminated large-bodied vertebrates in many areas across the tropics, contributing to the global process of defaunation. Elucidating the ecological consequences of hunting has important implications for managing ecosystems and for our understanding of community and ecosystem ecology. We present data collected through a combination of comparative and experimental approaches to assess how faunally-intact and heavily-hunted forests in Gabon differ in understory vegetation structure, macroinvertebrate fauna, ecological processes, and the relative importance of different taxa driving those processes. Our results show that hunted sites had denser understory vegetation and hosted approximately 170 times fewer termites compared to faunally-intact sites. While web-building spiders were positively associated with understory vegetation density, this effect did not translate to significantly higher abundances in heavily-hunted forests. Additionally, the overall rates of decomposition, insectivory, and seed predation/removal on the forest floor appeared robust to both defaunation and the associated increases in understory vegetation density. However, our exclosure experiments revealed that the contribution of invertebrates to decomposition was approximately 25% lower in hunted sites compared to faunally-intact sites. Results suggest potential resilience in this complex ecosystem such that microbial or other taxa not measured in this study may compensate for the reduced functional contribution of invertebrates to decomposition. However, while our results illustrate potential resilience, they also indicate that indirect effects following defaunation, such as increases in the density of understory vegetation, may alter invertebrate communities on the forest floor, with potential consequences for the mechanisms, and therefore the dynamics, driving critical ecosystem processes.
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    Diversity and carbon storage across the tropical forest biome.
    (Sci Rep, 2017-01-17) Sullivan, Martin JP; Talbot, Joey; Lewis, Simon L; Phillips, Oliver L; Qie, Lan; Begne, Serge K; Chave, Jerôme; Cuni-Sanchez, Aida; Hubau, Wannes; Lopez-Gonzalez, Gabriela; Miles, Lera; Monteagudo-Mendoza, Abel; Sonké, Bonaventure; Sunderland, Terry; Ter Steege, Hans; White, Lee JT; Affum-Baffoe, Kofi; Aiba, Shin-Ichiro; de Almeida, Everton Cristo; de Oliveira, Edmar Almeida; Alvarez-Loayza, Patricia; Dávila, Esteban Álvarez; Andrade, Ana; Aragão, Luiz EOC; Ashton, Peter; Aymard C, Gerardo A; Baker, Timothy R; Balinga, Michael; Banin, Lindsay F; Baraloto, Christopher; Bastin, Jean-Francois; Berry, Nicholas; Bogaert, Jan; Bonal, Damien; Bongers, Frans; Brienen, Roel; Camargo, José Luís C; Cerón, Carlos; Moscoso, Victor Chama; Chezeaux, Eric; Clark, Connie J; Pacheco, Álvaro Cogollo; Comiskey, James A; Valverde, Fernando Cornejo; Coronado, Eurídice N Honorio; Dargie, Greta; Davies, Stuart J; De Canniere, Charles; Djuikouo K, Marie Noel; Doucet, Jean-Louis; Erwin, Terry L; Espejo, Javier Silva; Ewango, Corneille EN; Fauset, Sophie; Feldpausch, Ted R; Herrera, Rafael; Gilpin, Martin; Gloor, Emanuel; Hall, Jefferson S; Harris, David J; Hart, Terese B; Kartawinata, Kuswata; Kho, Lip Khoon; Kitayama, Kanehiro; Laurance, Susan GW; Laurance, William F; Leal, Miguel E; Lovejoy, Thomas; Lovett, Jon C; Lukasu, Faustin Mpanya; Makana, Jean-Remy; Malhi, Yadvinder; Maracahipes, Leandro; Marimon, Beatriz S; Junior, Ben Hur Marimon; Marshall, Andrew R; Morandi, Paulo S; Mukendi, John Tshibamba; Mukinzi, Jaques; Nilus, Reuben; Vargas, Percy Núñez; Camacho, Nadir C Pallqui; Pardo, Guido; Peña-Claros, Marielos; Pétronelli, Pascal; Pickavance, Georgia C; Poulsen, Axel Dalberg; Poulsen, John R; Primack, Richard B; Priyadi, Hari; Quesada, Carlos A; Reitsma, Jan; Réjou-Méchain, Maxime; Restrepo, Zorayda; Rutishauser, Ervan; Salim, Kamariah Abu; Salomão, Rafael P; Samsoedin, Ismayadi; Sheil, Douglas; Sierra, Rodrigo; Silveira, Marcos; Slik, JW Ferry; Steel, Lisa; Taedoumg, Hermann; Tan, Sylvester; Terborgh, John W; Thomas, Sean C; Toledo, Marisol; Umunay, Peter M; Gamarra, Luis Valenzuela; Vieira, Ima Célia Guimarães; Vos, Vincent A; Wang, Ophelia; Willcock, Simon; Zemagho, Lise
    Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity.
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    Ecological erosion of an Afrotropical forest and potential consequences for tree recruitment and forest biomass
    (Biological Conservation, 2013) Poulsen, John R; Clark, Connie J; Palmer, Todd M
    Unprecedented rates of logging and hunting threaten to transform the remaining primary tropical forest into a degraded mosaic, emptied of wildlife. Defaunation is expected to interrupt plant-animal interactions with cascading effects for forest structure, composition, and ecosystem services. In a Central African forest first logged 35years ago, we evaluated this process of ecological erosion in 30 study sites distributed across forest disturbed by logging and hunting, logging alone, and neither logging nor hunting. Both logging and hunting tended to reduce abundances of large mammals, together shifting the relative abundance of the animal community towards squirrels and small birds. Through a series of experiments, we evaluated the effects of logging and hunting on seed dispersal, seed predation and herbivory. We demonstrate that complete defaunation is not necessary to significantly alter the strength of plant-animal interactions. Hunting reduced the mean dispersal distances of nine mammal-dispersed tree species by 22%. Rates of seed predation were similar among forest types, but hunted forest had significantly lower rates of herbivory that we attribute to the lower abundance of meso-herbivores. Hunted forest also had significantly lower above-ground biomass (301Mgha-1) than the logged only (358Mgha-1) and undisturbed (455Mgha-1) forest types, but similar numbers of tree species and individuals. Lower biomass in hunted forest is likely attributable to significantly lower wood densities at small tree size classes (<40cm). We hypothesize that over time the human-mediated modification of plant-animal interactions can alter the composition of the forest to have a higher proportion of fast-growing, low wood density tree species, diminishing the long-term potential for carbon storage. © 2013 Elsevier Ltd.
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    Expected carbon emissions from a rubber plantation in Central Africa
    (Forest Ecology and Management, 2021-01-15) Jong, Ying Wei; Beirne, Christopher; Meunier, Quentin; Mekui Biyogo, Andréana Paola; Ebang Mbélé, Alex; Stewart, Christopher G; Poulsen, John R
    The development of agriculture on degraded lands is increasingly seen as a strategy to boost food availability and economic productivity while minimizing environmental degradation and loss of forests. To understand the effects of agricultural production on forest carbon, we quantify the aboveground carbon (AGC) of a degraded forest in northeast Gabon (the Olam Rubber Gabon concession) designated for development to a rubber plantation. Combining field measurements from 19 1-ha tree plots and aerial LiDAR, we estimate forest AGC stocks and emissions under four development scenarios: no development, 30-year rubber rotation, extended rubber rotation (replanting of plantation in stages at 30 and 40 years), and 30-year oil palm rotation. On average, the degraded forest in the study area stored 123.8 Mg C ha−1, a mean AGC lower than the Gabon average (141.6 Mg C ha−1) but substantially higher than the 75 Mg C ha−1 threshold recommended by the High Carbon Stock protocol. Converting secondary forest to plantation might incur high environmental opportunity costs from lost carbon sequestration through forest succession and growth. In this study, we estimate that a rubber plantation can sequester similar amounts of AGC as secondary forest by the end of a 30-year rotation; however, the time-averaged AGC of regenerating secondary forests under no development would be 184% higher than a mature rubber plantation with a 30-year rotation, 169% higher than an extended rubber rotation, and 512% higher than a 30-year oil palm rotation. When degraded forest is developed for agriculture, measures should be taken to avoid emissions and prolong carbon retention. We specifically estimate carbon retention from extended harvest rotations and conserving high carbon value areas as set-asides and highlight recommendations from the literature such as minimizing soil disturbance and creating rubber timber products (e.g. furniture). To minimize carbon emissions from agriculture, crop plantation area should be minimized at national and regional scales in highly forested countries, and new plantations should be coupled explicitly with effective forest restoration actions, through suitable regulation and planning, to mitigate or compensate for their climate and biodiversity impacts.
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    Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment.
    (Ecology, 2012-03) Poulsen, John R; Clark, Connie J; Bolker, Benjamin M
    The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.
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    Forest elephant movement and habitat use in a tropical forest-grassland mosaic in Gabon.
    (PloS one, 2018-01) Mills, Emily C; Poulsen, John R; Fay, J Michael; Morkel, Peter; Clark, Connie J; Meier, Amelia; Beirne, Christopher; White, Lee JT
    Poaching of forest elephants (Loxodonta cyclotis) for ivory has decimated their populations in Central Africa. Studying elephant movement can provide insight into habitat and resource use to reveal where, when, and why they move and guide conservation efforts. We fitted 17 forest elephants with global positioning system (GPS) collars in 2015 and 2016 in the tropical forest-grassland mosaic of the Wonga Wongué Presidential Reserve (WW), Gabon. Using the location data, we quantified movement distances, home ranges, and habitat use to examine the environmental drivers of elephant movements and predict where elephants occur spatially and temporally. Forest elephants, on average, traveled 2,840 km annually and had home ranges of 713 km2, with males covering significantly larger home ranges than females. Forest elephants demonstrated both daily and seasonal movement patterns. Daily, they moved between forest and grassland at dawn and dusk. Seasonally, they spent proportionally more time in grassland than forest during the short-wet season when grasses recruit. Forest elephants also traveled faster during the short-wet season when fruit availability was greatest, likely reflecting long, direct movements to preferred fruiting tree species. Forest elephants tended to select areas with high tree and shrub density that afford cover and browse. When villages occurred in their home ranges elephants spent a disproportionate amount of time near them, particularly in the dry season, probably for access to agricultural crops and preferred habitat. Given the importance of the grassland habitat for elephants, maintenance of the forest-grassland matrix is a conservation priority in WW. Law enforcement, outreach, and education should focus on areas of potential human-elephant conflict near villages along the borders of the reserve. GPS-tracking should be extended into multi-use areas in the peripheries of protected areas to evaluate the effects of human disturbance on elephant movements and to maintain connectivity among elephant populations in Gabon.
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    How Bees Deter Elephants: Beehive Trials with Forest Elephants (Loxodonta africana cyclotis) in Gabon.
    (PLoS One, 2016) Ngama, Steeve; Korte, Lisa; Bindelle, Jérôme; Vermeulen, Cédric; Poulsen, John R
    In Gabon, like elsewhere in Africa, crops are often sources of conflict between humans and wildlife. Wildlife damage to crops can drastically reduce income, amplifying poverty and creating a negative perception of wild animal conservation among rural people. In this context, crop-raiding animals like elephants quickly become "problem animals". To deter elephants from raiding crops beehives have been successfully employed in East Africa; however, this method has not yet been tested in Central Africa. We experimentally examined whether the presence of Apis mellifera adansonii, the African honey bee species present in Central Africa, deters forest elephants (Loxodonta Africana cyclotis) from feeding on fruit trees. We show for the first time that the effectiveness of beehives as deterrents of elephants is related to bee activity. Empty hives and those housing colonies of low bee activity do not deter elephants all the time; but beehives with high bee activity do. Although elephant disturbance of hives does not impede honey production, there is a tradeoff between deterrence and the quantity of honey produced. To best achieve the dual goals of deterring elephants and producing honey colonies must maintain an optimum activity level of 40 to 60 bee movements per minute. Thus, beehives colonized by Apis mellifera adansonii bees can be effective elephant deterrents, but people must actively manage hives to maintain bee colonies at the optimum activity level.
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    Hunting-induced defaunation drives increased seed predation and decreased seedling establishment of commercially important tree species in an Afrotropical forest
    (Forest Ecology and Management, 2016-12-15) Rosin, Cooper; Poulsen, John R
    © 2016 Elsevier B.V. Human hunting is widespread in tropical forests and can substantially alter the plant-animal interactions that drive tree recruitment. Seed predation is a strong determinant of plant reproductive success, but it remains unclear how defaunation modifies this process. We examined the effects of hunting-induced defaunation on seed predation and seedling establishment, using replicated exclosure treatments at six sit es across a defaunation gradient in northeastern Gabon. We monitored 5580 seeds of eight commercially important tree species that varied in seed traits such as size and dispersal mode. Rodents caused the greatest seed mortality for all species, removing ∼60% of accessible seeds. In comparison, invertebrates and fungi together caused just 6% of seed mortality. With protection from rodents, more than twice as many seeds established as seedlings, demonstrating that vertebrate seed predation was a strong filter on recruitment. With increasing defaunation, the proportion of seeds removed by rodents increased significantly, and seedling establishment decreased significantly, for most species. In heavily defaunated sites, with the lowest abundances of large mammals, seed removal by rodents increased by 63% and seedling establishment decreased by 42% compared to sites with intact fauna. Diminished seedling establishment is likely to reduce the regeneration of many tree species – including some with commercial importance – in hunted forests, with detrimental economic consequences. In turn, declines in timber regeneration may increase the likelihood that selectively logged forests are converted to non-forest land uses with little conservation value. Appropriate management could preclude these outcomes, to the benefit of both wildlife and natural timber regeneration.
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    Long-term thermal sensitivity of Earth's tropical forests.
    (Science (New York, N.Y.), 2020-05-21) Sullivan, Martin JP; Lewis, Simon L; Affum-Baffoe, Kofi; Castilho, Carolina; Costa, Flávia; Sanchez, Aida Cuni; Ewango, Corneille EN; Hubau, Wannes; Marimon, Beatriz; Monteagudo-Mendoza, Abel; Qie, Lan; Sonké, Bonaventure; Martinez, Rodolfo Vasquez; Baker, Timothy R; Brienen, Roel JW; Feldpausch, Ted R; Galbraith, David; Gloor, Manuel; Malhi, Yadvinder; Aiba, Shin-Ichiro; Alexiades, Miguel N; Almeida, Everton C; de Oliveira, Edmar Almeida; Dávila, Esteban Álvarez; Loayza, Patricia Alvarez; Andrade, Ana; Vieira, Simone Aparecida; Aragão, Luiz EOC; Araujo-Murakami, Alejandro; Arets, Eric JMM; Arroyo, Luzmila; Ashton, Peter; Aymard C, Gerardo; Baccaro, Fabrício B; Banin, Lindsay F; Baraloto, Christopher; Camargo, Plínio Barbosa; Barlow, Jos; Barroso, Jorcely; Bastin, Jean-François; Batterman, Sarah A; Beeckman, Hans; Begne, Serge K; Bennett, Amy C; Berenguer, Erika; Berry, Nicholas; Blanc, Lilian; Boeckx, Pascal; Bogaert, Jan; Bonal, Damien; Bongers, Frans; Bradford, Matt; Brearley, Francis Q; Brncic, Terry; Brown, Foster; Burban, Benoit; Camargo, José Luís; Castro, Wendeson; Céron, Carlos; Ribeiro, Sabina Cerruto; Moscoso, Victor Chama; Chave, Jerôme; Chezeaux, Eric; Clark, Connie J; de Souza, Fernanda Coelho; Collins, Murray; Comiskey, James A; Valverde, Fernando Cornejo; Medina, Massiel Corrales; da Costa, Lola; Dančák, Martin; Dargie, Greta C; Davies, Stuart; Cardozo, Nallaret Davila; de Haulleville, Thales; de Medeiros, Marcelo Brilhante; Del Aguila Pasquel, Jhon; Derroire, Géraldine; Di Fiore, Anthony; Doucet, Jean-Louis; Dourdain, Aurélie; Droissart, Vincent; Duque, Luisa Fernanda; Ekoungoulou, Romeo; Elias, Fernando; Erwin, Terry; Esquivel-Muelbert, Adriane; Fauset, Sophie; Ferreira, Joice; Llampazo, Gerardo Flores; Foli, Ernest; Ford, Andrew; Gilpin, Martin; Hall, Jefferson S; Hamer, Keith C; Hamilton, Alan C; Harris, David J; Hart, Terese B; Hédl, Radim; Herault, Bruno; Herrera, Rafael; Higuchi, Niro; Hladik, Annette; Coronado, Eurídice Honorio; Huamantupa-Chuquimaco, Isau; Huasco, Walter Huaraca; Jeffery, Kathryn J; Jimenez-Rojas, Eliana; Kalamandeen, Michelle; Djuikouo, Marie Noël Kamdem; Kearsley, Elizabeth; Umetsu, Ricardo Keichi; Kho, Lip Khoon; Killeen, Timothy; Kitayama, Kanehiro; Klitgaard, Bente; Koch, Alexander; Labrière, Nicolas; Laurance, William; Laurance, Susan; Leal, Miguel E; Levesley, Aurora; Lima, Adriano JN; Lisingo, Janvier; Lopes, Aline P; Lopez-Gonzalez, Gabriela; Lovejoy, Tom; Lovett, Jon C; Lowe, Richard; Magnusson, William E; Malumbres-Olarte, Jagoba; Manzatto, Ângelo Gilberto; Marimon, Ben Hur; Marshall, Andrew R; Marthews, Toby; de Almeida Reis, Simone Matias; Maycock, Colin; Melgaço, Karina; Mendoza, Casimiro; Metali, Faizah; Mihindou, Vianet; Milliken, William; Mitchard, Edward TA; Morandi, Paulo S; Mossman, Hannah L; Nagy, Laszlo; Nascimento, Henrique; Neill, David; Nilus, Reuben; Vargas, Percy Núñez; Palacios, Walter; Camacho, Nadir Pallqui; Peacock, Julie; Pendry, Colin; Peñuela Mora, Maria Cristina; Pickavance, Georgia C; Pipoly, John; Pitman, Nigel; Playfair, Maureen; Poorter, Lourens; Poulsen, John R; Poulsen, Axel Dalberg; Preziosi, Richard; Prieto, Adriana; Primack, Richard B; Ramírez-Angulo, Hirma; Reitsma, Jan; Réjou-Méchain, Maxime; Correa, Zorayda Restrepo; de Sousa, Thaiane Rodrigues; Bayona, Lily Rodriguez; Roopsind, Anand; Rudas, Agustín; Rutishauser, Ervan; Abu Salim, Kamariah; Salomão, Rafael P; Schietti, Juliana; Sheil, Douglas; Silva, Richarlly C; Espejo, Javier Silva; Valeria, Camila Silva; Silveira, Marcos; Simo-Droissart, Murielle; Simon, Marcelo Fragomeni; Singh, James; Soto Shareva, Yahn Carlos; Stahl, Clement; Stropp, Juliana; Sukri, Rahayu; Sunderland, Terry; Svátek, Martin; Swaine, Michael D; Swamy, Varun; Taedoumg, Hermann; Talbot, Joey; Taplin, James; Taylor, David; Ter Steege, Hans; Terborgh, John; Thomas, Raquel; Thomas, Sean C; Torres-Lezama, Armando; Umunay, Peter; Gamarra, Luis Valenzuela; van der Heijden, Geertje; van der Hout, Peter; van der Meer, Peter; van Nieuwstadt, Mark; Verbeeck, Hans; Vernimmen, Ronald; Vicentini, Alberto; Vieira, Ima Célia Guimarães; Torre, Emilio Vilanova; Vleminckx, Jason; Vos, Vincent; Wang, Ophelia; White, Lee JT; Willcock, Simon; Woods, John T; Wortel, Verginia; Young, Kenneth; Zagt, Roderick; Zemagho, Lise; Zuidema, Pieter A; Zwerts, Joeri A; Phillips, Oliver L
    The sensitivity of tropical forest carbon to climate is a key uncertainty in predicting global climate change. Although short-term drying and warming are known to affect forests, it is unknown if such effects translate into long-term responses. Here, we analyze 590 permanent plots measured across the tropics to derive the equilibrium climate controls on forest carbon. Maximum temperature is the most important predictor of aboveground biomass (-9.1 megagrams of carbon per hectare per degree Celsius), primarily by reducing woody productivity, and has a greater impact per °C in the hottest forests (>32.2°C). Our results nevertheless reveal greater thermal resilience than observations of short-term variation imply. To realize the long-term climate adaptation potential of tropical forests requires both protecting them and stabilizing Earth's climate.
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    Low-intensity logging and hunting have long-term effects on seed dispersal but not fecundity in Afrotropical forests.
    (AoB PLANTS, 2019-02) Nuñez, Chase L; Clark, James S; Clark, Connie J; Poulsen, John R
    Hunting and logging, ubiquitous human disturbances in tropical forests, have the potential to alter the ecological processes that govern population recruitment and community composition. Hunting-induced declines in populations of seed-dispersing animals are expected to reduce dispersal of the tree species that rely on them, resulting in potentially greater distance- and density-dependent mortality. At the same time, selective logging may alter competitive interactions among tree species, releasing remaining trees from light, nutrient or space limitations. Taken together, these disturbances may alter the community composition of tropical forests, with implications for carbon storage, biodiversity conservation and ecosystem function. To evaluate the effects of hunting and logging on tree fecundity and seed dispersal, we use 3 years of seed rain data from a large-scale observational experiment in previously logged, hunted and protected forests in northern Republic of Congo (Brazzaville). We find that low-intensity logging had a meaningful long-term effect on species-specific seed dispersal distances, though the direction and magnitude varied and was not congruent within dispersal vector. Tree fecundity increased with tree diameter, but did not differ appreciably across disturbance regimes. The species-specific dispersal responses to logging in this study point towards the long-lasting toll of disturbance on ecological function and highlight the necessity of conserving intact forest.
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    Poaching empties critical Central African wilderness of forest elephants.
    (Curr Biol, 2017-02-20) Poulsen, John R; Koerner, Sally E; Moore, Sarah; Medjibe, Vincent P; Blake, Stephen; Clark, Connie J; Akou, Mark Ella; Fay, Michael; Meier, Amelia; Okouyi, Joseph; Rosin, Cooper; White, Lee JT
    Elephant populations are in peril everywhere, but forest elephants in Central Africa have sustained alarming losses in the last decade [1]. Large, remote protected areas are thought to best safeguard forest elephants by supporting large populations buffered from habitat fragmentation, edge effects and human pressures. One such area, the Minkébé National Park (MNP), Gabon, was created chiefly for its reputation of harboring a large elephant population. MNP held the highest densities of elephants in Central Africa at the turn of the century, and was considered a critical sanctuary for forest elephants because of its relatively large size and isolation. We assessed population change in the park and its surroundings between 2004 and 2014. Using two independent modeling approaches, we estimated a 78-81% decline in elephant numbers over ten years - a loss of more than 25,000 elephants. While poaching occurs from within Gabon, cross-border poaching largely drove the precipitous drop in elephant numbers. With nearly 50% of forest elephants in Central Africa thought to reside in Gabon [1], their loss from the park is a considerable setback for the preservation of the species.
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    Resistance of African tropical forests to an extreme climate anomaly.
    (Proceedings of the National Academy of Sciences of the United States of America, 2021-05) Bennett, Amy C; Dargie, Greta C; Cuni-Sanchez, Aida; Tshibamba Mukendi, John; Hubau, Wannes; Mukinzi, Jacques M; Phillips, Oliver L; Malhi, Yadvinder; Sullivan, Martin JP; Cooper, Declan LM; Adu-Bredu, Stephen; Affum-Baffoe, Kofi; Amani, Christian A; Banin, Lindsay F; Beeckman, Hans; Begne, Serge K; Bocko, Yannick E; Boeckx, Pascal; Bogaert, Jan; Brncic, Terry; Chezeaux, Eric; Clark, Connie J; Daniels, Armandu K; de Haulleville, Thales; Djuikouo Kamdem, Marie-Noël; Doucet, Jean-Louis; Evouna Ondo, Fidèle; Ewango, Corneille EN; Feldpausch, Ted R; Foli, Ernest G; Gonmadje, Christelle; Hall, Jefferson S; Hardy, Olivier J; Harris, David J; Ifo, Suspense A; Jeffery, Kathryn J; Kearsley, Elizabeth; Leal, Miguel; Levesley, Aurora; Makana, Jean-Remy; Mbayu Lukasu, Faustin; Medjibe, Vincent P; Mihindu, Vianet; Moore, Sam; Nssi Begone, Natacha; Pickavance, Georgia C; Poulsen, John R; Reitsma, Jan; Sonké, Bonaventure; Sunderland, Terry CH; Taedoumg, Hermann; Talbot, Joey; Tuagben, Darlington S; Umunay, Peter M; Verbeeck, Hans; Vleminckx, Jason; White, Lee JT; Woell, Hannsjoerg; Woods, John T; Zemagho, Lise; Lewis, Simon L
    The responses of tropical forests to environmental change are critical uncertainties in predicting the future impacts of climate change. The positive phase of the 2015-2016 El Niño Southern Oscillation resulted in unprecedented heat and low precipitation in the tropics with substantial impacts on the global carbon cycle. The role of African tropical forests is uncertain as their responses to short-term drought and temperature anomalies have yet to be determined using on-the-ground measurements. African tropical forests may be particularly sensitive because they exist in relatively dry conditions compared with Amazonian or Asian forests, or they may be more resistant because of an abundance of drought-adapted species. Here, we report responses of structurally intact old-growth lowland tropical forests inventoried within the African Tropical Rainforest Observatory Network (AfriTRON). We use 100 long-term inventory plots from six countries each measured at least twice prior to and once following the 2015-2016 El Niño event. These plots experienced the highest temperatures and driest conditions on record. The record temperature did not significantly reduce carbon gains from tree growth or significantly increase carbon losses from tree mortality, but the record drought did significantly decrease net carbon uptake. Overall, the long-term biomass increase of these forests was reduced due to the El Niño event, but these plots remained a live biomass carbon sink (0.51 ± 0.40 Mg C ha-1 y-1) despite extreme environmental conditions. Our analyses, while limited to African tropical forests, suggest they may be more resistant to climatic extremes than Amazonian and Asian forests.
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    Roles of seed and establishment limitation in determining patterns of afrotropical tree recruitment.
    (PloS one, 2013-01) Clark, Connie J; Poulsen, John R; Levey, Doug J
    Quantifying the relative importance of the multiple processes that limit recruitment may hold the key to understanding tropical tree diversity. Here we couple theoretical models with a large-scale, multi-species seed-sowing experiment to assess the degree to which seed and establishment limitation shape patterns of tropical tree seedling recruitment in a central African forest. Of five randomly selected species (Pancovia laurentii, Staudtia kamerunensis, Manilkara mabokeensis, Myrianthus arboreas, and Entandophragma utile), seedling establishment and survival were low (means of 16% and 6% at 3 and 24 months, respectively), and seedling density increased with seed augmentation. Seedling recruitment was best explained by species identity and the interaction of site-by-species, suggesting recruitment probabilities vary among species and sites, and supporting the role of niche-based mechanisms. Although seed augmentation enhanced initial seedling density, environmental filtering and post-establishment mortality strongly limited seedling recruitment. The relative importance of seed and establishment limitation changed with seed and seedling density and through time. The arrival of seeds most strongly affected local recruitment when seeds were nearly absent from a site (∼ 1 seed m(2)), but was also important when seeds arrived in extremely high densities, overwhelming niche-based mortality factors. The strength of seed limitation and density-independent mortality decreased significantly over time, while density-dependent mortality showed the opposite trend. The varying strengths of seed and establishment limitation as a function of juvenile density and time emphasize the need to evaluate their roles through later stages of a tree's life cycle.
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    Telemetric tracking of scatterhoarding and seed fate in a Central African forest
    (Biotropica, 2017-03-01) Rosin, Cooper; Poulsen, John R
    © 2016 The Association for Tropical Biology and Conservation In seed predation studies, removal of a seed is only the first step of a dynamic process that may result in dispersal rather than seed death. This process, termed seed fate, has received little attention in African forests, particularly in Central Africa. We experimentally assessed the initial steps of seed fate for two tree species—the large-seeded Pentaclethra macrophylla and the relatively small-seeded Gambeya lacourtiana—in northeastern Gabon. Specifically, we evaluated whether seed size and seed consumer identity are important determinants of seed fate. We established experimental stations under conspecific fruiting trees, each comprising three seeds fitted with telemetric thread tags to facilitate their recovery, and a motion-sensitive camera to identify visiting mammals. In total, animals removed 76 tagged seeds from experimental stations. Small Murid rats and mice primarily removed small Gambeya seeds, whereas large-bodied rodents and mandrills primarily removed large Pentaclethra seeds. Gambeya seeds were carried shorter distances than Pentaclethra seeds and were less likely to be cached. The two large-bodied rodents handled seeds differently: Cricetomys emini larderhoarded nearly all (N = 15 of 16) encountered Pentaclethra seeds deep in burrows, while Atherurus africanus cached all (N = 5 of 5) encountered Pentaclethra seeds singly under 1–3 cm of leaf litter and soil, at an average distance of 24.2 m and a maximum distance of 46.3 m from experimental stations. This study supports the hypothesis that seed fate varies based on seed size and seed consumer identity, and represents the first telemetric experimental evidence of larderhoarding and scatterhoarding in the region.