Two centuries of biodiversity discovery and loss in Singapore.

There is an urgent need for reliable data on the impacts of deforestation on tropical biodiversity. The city-state of Singapore has one of the most detailed biodiversity records in the tropics, dating back to the turn of the 19th century. In 1819, Singapore was almost entirely covered in primary forest, but this has since been largely cleared. We compiled more than 200 y of records for 10 major taxonomic groups in Singapore (>50,000 individual records; >3,000 species), and we estimated extinction rates using recently developed and novel statistical models that account for "dark extinctions," i.e., extinctions of undiscovered species. The estimated overall extinction rate was 37% (95% CI [31 to 42%]). Extrapolating our Singapore observations to a future business-as-usual deforestation scenario for Southeast Asia suggests that 18% (95% CI [16 to 22%]) of species will be lost regionally by 2100. Our extinction estimates for Singapore and Southeast Asia are a factor of two lower than previous estimates that also attempted to account for dark extinctions. However, we caution that particular groups such as large mammals, forest-dependent birds, orchids, and butterflies are disproportionately vulnerable.

Ancestral social environments plus nonlinear benefits can explain cooperation in human societies.

Human cooperation (paying a cost to benefit others) is puzzling from a Darwinian perspective, particularly in groups with strangers who cannot repay nor are family members. The beneficial effects of cooperation typically increase nonlinearly with the number of cooperators, e.g., increasing returns when cooperation is low and diminishing returns when cooperation is high. Such nonlinearity can allow cooperation between strangers to persist evolutionarily if a large enough proportion of the population are already cooperators. However, if a lone cooperator faces a conflict between the group's and its own interests (a social dilemma), that raises the question of how cooperation arose in the first place. We use a mathematically tractable evolutionary model to formalise a chronological narrative that has previously only been investigated verbally: given that ancient humans interacted mostly with family members (genetic homophily), cooperation evolved first by kin selection, and then persisted in situations with nonlinear benefits as homophily declined or even if interactions with strangers became the norm. The model also predicts the coexistence of cooperators and defectors observed in the human population (polymorphism), and may explain why cooperators in behavioural experiments prefer to condition their contribution on the contributions of others (conditional cooperation in public goods games).

Ecological network assembly: How the regional metaweb influences local food webs.

Local food webs result from a sequence of colonisations and extinctions by species from the regional pool or metaweb, that is, the assembly process. Assembly is theorised to be a selective process: whether or not certain species or network structures can persist is partly determined by local processes including habitat filtering and dynamical constraints. Consequently, local food web structure should reflect these processes. The goal of this study was to test evidence for these selective processes by comparing the structural properties of real food webs to the expected distribution given the metaweb. We were particularly interested in ecological dynamics; if the network properties commonly associated with dynamical stability are indeed the result of stability constraints, then they should deviate from expectation in the direction predicted by theory. To create a null expectation, we used the novel approach of randomly assembling model webs by drawing species and interactions from the empirical metaweb. The assembly model permitted colonisation and extinction, and required a consumer species to have at least one prey, but had no habitat type nor population dynamical constraints. Three datasets were used: (a) the marine Antarctic metaweb, with two local food webs; (b) the 50 lakes of the Adirondacks; and (c) the arthropod community from Florida Keys' classic defaunation experiment. Contrary to our expectations, we found that there were almost no differences between empirical webs and those resulting from the null assembly model. Few empirical food webs showed significant differences with network properties, motif representations and topological roles. Network properties associated with stability did not deviate from expectation in the direction predicted by theory. Our results suggest that-for the commonly used metrics we considered-local food web structure is not strongly influenced by dynamical nor habitat restrictions. Instead, the structure is inherited from the metaweb. This suggests that the network properties typically attributed as causes or consequences of ecological stability are instead a by-product of the assembly process (i.e. spandrels), and may potentially be too coarse to detect the true signal of dynamical constraint.

Las redes tróficas locales son el resultado de una secuencia de eventos de colonización y extinción de especies que provienen del conjunto regional de especies o metaweb, es decir, el proceso de ensamble. Se teoriza que el ensamble es un proceso selectivo: si ciertas especies o estructuras de red pueden persistir o no está determinado en parte por procesos locales, incluido el filtrado del hábitat y las limitaciones dinámicas. En consecuencia, la estructura de la red trófica local debe reflejar estos procesos. El objetivo de este estudio fue analizar la evidencia de estos procesos selectivos comparando las propiedades estructurales de redes tróficas empíricas (locales) con la distribución esperada dada la metaweb. Nuestro particular interés fue la dinámica ecológica: si las propiedades de la red comúnmente asociadas con la estabilidad dinámica son el resultado de restricciones de estabilidad, entonces deberían desviarse de las expectativas en la dirección predicha por la teoría. Para crear una expectativa nula utilizamos el enfoque novedoso de ensamblar redes utilizando un modelo estocástico o 'modelo de ensamble nulo' que extrae especies e interacciones de una metaweb empírica. El modelo de ensamble incluye colonización y extinción, y requiere que los consumidores tengan al menos una presa, pero no tiene restricciones acerca del tipo de hábitat ni de dinámica de poblaciones. Se utilizaron tres conjuntos de datos: (a) la metaweb antártica marina, con 2 redes tróficas locales; (b) los 50 lagos de Adirondacks; y (c) la comunidad de artrópodos del experimento clásico de defaunación de los Cayos de la Florida. Contrariamente a lo esperado, encontramos que casi no hubo diferencias entre las redes empíricas (locales) y las resultantes del modelo de ensamble nulo. Pocas redes tróficas empíricas mostraron diferencias significativas con las propiedades de la red, los motivos y los roles topológicos. Las propiedades de la red asociadas con la estabilidad no se desviaron de las expectativas en la dirección predicha por la teoría. Nuestros resultados sugieren que-para las métricas de uso común que consideramos-la estructura de la red trófica local no está fuertemente influenciada por restricciones dinámicas o de hábitat. En cambio, la estructura se hereda de la metaweb. Esto sugiere que las propiedades de la red típicamente atribuidas como causas o consecuencias de la estabilidad ecológica son, en cambio, un subproducto del proceso de ensamble y podrían no tener la suficiente potencia para detectar la verdadera señal de restricción dinámica.

Having a stake in the future and perceived population density influence intergenerational cooperation.

Intergenerational common-pool resource games represent a new experimental paradigm in which the current generation's decision to cooperate or defect influences future generations who cannot reciprocate, providing key insights for sustainability science. We combine experimental and theoretical approaches to assess the roles of having a stake in the future (50% chance to pass the resource on to themselves in the next generation) and reminders of the presence of others (exposure to people-chatting sounds) on intergenerational cooperation. We find that, as expected, having a stake in the future increases cooperation with future generations, except when participants are also exposed to people-chatting sounds. We hypothesize that this interaction effect occurs because people-chatting sounds trigger a perception of large group size, which reduces the chance of individuals and their descendants benefiting from the pool in the future, thus reducing cooperation. Our results highlight the context-dependent effect of having a future stake on intergenerational cooperation for resource sustainability, and suggest an area of future work for environmental messaging.

Extinction rate of discovered and undiscovered plants in Singapore.

Extinction is a key issue in the assessment of global biodiversity. However, many extinction rate measures do not account for species that went extinct before they could be discovered. The highly developed island city-state of Singapore has one of the best-documented tropical floras in the world. This allowed us to estimate the total rate of floristic extinctions in Singapore since 1822 after accounting for sampling effort and crypto extinctions by collating herbaria records. Our database comprised 34,224 specimens from 2076 native species, of which 464 species (22%) were considered nationally extinct. We assumed that undiscovered species had the same annual per-species extinction rates as discovered species and that no undiscovered species remained extant. With classical and Bayesian algorithms, we estimated that 304 (95% confidence interval, 213-414) and 412 (95% credible interval, 313-534) additional species went extinct before they could be discovered, respectively; corresponding total extinction rate estimates were 32% and 35% (range 30-38%). We detected violations of our 2 assumptions that could cause our extinction estimates, particularly the absolute numbers, to be biased downward. Thus, our estimates should be treated as lower bounds. Our results illustrate the possible magnitudes of plant extirpations that can be expected in the tropics as development continues.

Tasa de Extinción de Plantas Descubiertas y No Descubiertas en Singapur Resumen La extinción es un tema importante para la valoración de la biodiversidad global. Sin embargo, muchas medidas de la tasa de extinción no consideran a las especies que se extinguieron antes de que pudieran ser descubiertas. Singapur, la ciudad-estado isleña altamente desarrollada, tiene una de las floras mejor documentadas del mundo. Esto nos permitió estimar la tasa total de las extinciones florísticas en Singapur desde 1822 después de considerar el esfuerzo de muestreo y las criptoextinciones cuando recopilamos los registros de herbarios. Nuestra base de datos incluyó 34,224 especímenes de unas 2,076 especies nativas, de las cuales 464 especies (22%) estaban consideradas como extintas a nivel nacional. Asumimos que las especies no descubiertas tuvieron la misma tasa anual de extinción por especie que las especies descubiertas y que ninguna especie no descubierta permanecía en existencia. Con algoritmos clásicos y bayesianos, respectivamente, estimamos que 304 (95% IC 213-414) y 412 (95% IC 313-534) especies adicionales se extinguieron antes de que fueran descubiertas; las estimaciones correspondientes de la tasa de extinción total fueron 32% y 35% (rango de 30-38%). Detectamos violaciones en nuestras dos suposiciones que podrían causar que nuestras estimaciones de extinción, particularmente los números absolutos, tuvieran un sesgo hacia abajo. Por lo tanto, nuestras estimaciones deberían ser tratadas como límites inferiores. Nuestros resultados ilustran las magnitudes posibles de las extirpaciones de plantas que pueden esperarse en los trópicos conforme el desarrollo continúa.

Inferring stratified parasitoid dispersal mechanisms and parameters from coarse data using mathematical and Bayesian methods.

Biological invasions have movement at the core of their success. However, due to difficulties in collecting data, medium- and long-distance dispersal of small insects has long been poorly understood and likely to be underestimated. The agricultural release of parasitic hymenoptera, a group of wasps that are critical for biological pest control, represents a rare opportunity to study the spread of insects on multiple spatial scales. As these insects are typically less than 1 mm in size and are challenging to track individually, a first-time biocontrol release will provide a known spatial position and time of initial release for all individuals that are subsequently collected. In this paper, we develop and validate a new mathematical model for parasitoid wasp dispersal from point release, as in the case of biocontrol. The model is derived from underlying stochastic processes but is fully deterministic and admits an analytical solution. Using a Bayesian framework, we then fit the model to an Australian dataset describing the multi-scale wind-borne dispersal pattern of Eretmocerus hayati Zolnerowich & Rose (Hymenoptera: Aphelinidae). Our results confirm that both local movements and long-distance wind dispersal are significant to the movement of parasitoids. The model results also suggest that low velocity winds are the primary indicator of dispersal direction on the field scale shortly after release, and that average wind data may be insufficient to resolve long-distance movement given inherent nonlinearities and heterogeneities in atmospheric flows. The results highlight the importance of collecting wind data when developing models to predict the spread of parasitoids and other tiny organisms.

Wind-borne dispersal of a parasitoid: the process, the model, and its validation.

The aphelinid parasitoid Eretmocerus hayati Zolnerowich & Rose (Hymenoptera: Aphelinidae) was recently released in Australia as a biocontrol agent against the crop pest Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae). It was found that the parasitoid can spread over several kilometers in a single generation and continue laying eggs for over a fortnight. A simple wind-advection model was fitted to emergence data from a first release between Fassifern and Kalbar, Queensland, and its predictive ability was tested against the second release near Carnarvon, Western Australia. The fitting of the model was used to develop several hypotheses about the dispersal of E. hayati, which were validated by the second release: E. hayati flies in the same direction as the wind to a distance proportional to the wind speed; this wind-borne flight takes place at any time during daylight hours; a flight is attempted every day after emergence unless there are high wind conditions during that day; and the high wind condition that will delay flight is wind speeds in excess of ≍2 m/s. This model of E. hayati dispersal may be contrasted with previous models fitted for Eretmocerus species, for which dispersal was dominated by diffusion processes, and parasitoid spread was constrained to the scales of tens and hundreds of meters.

The initial dispersal and spread of an intentional invader at three spatial scales.

The way an invasion progresses through space is a theme of interest common to invasion ecology and biological pest control. Models and mark-release studies of arthropods have been used extensively to extend and inform invasion processes of establishment and spread. However, the extremely common single-scale approach of monitoring initial spread leads to misinterpretation of rate and mode. Using the intentional release of a novel biological control agent (a parasitic hymenoptera, Eretmocerus hayati Zolnerowich & Rose (Hymenoptera: Aphelinidae), we studied its initial dispersal and spread at three different spatial scales, the local scale (tens of metres), field scale (hundreds of metres) and landscape scale (kilometres) around the release point. We fit models to each observed spread pattern at each spatial scale. We show that E. hayati exhibits stratified dispersal; moving further, faster and by a different mechanism than would have been concluded with a single local-scale post-release sampling design. In fact, interpretation of each scale independent of other scales gave three different models of dispersal, and three different impressions of the dominant dispersal mechanisms. Our findings demonstrate that using a single-scale approach may lead to quite erroneous conclusions, hence the necessity of using a multiple-scale hierarchical sampling design for inferring spread and the dominant dispersal mechanism of either human intended or unintended invasions.

Permanence does not predict the commonly measured food web structural attributes.

Food web assembly algorithms show great promise for investigating issues involving the dynamics of whole webs, such as succession, rehabilitation, and invasibility. Permanence, which requires that all species densities remain positive and finite, has been suggested as a good stability constraint. This study tests the validity of the permanence constraint by comparing real webs and model webs from the literature to the predictions of three assembly algorithms: one constrained by permanence and feasibility, one constrained by feasibility alone, and one with no dynamical constraint. It is found that the addition of the permanence constraint does not improve the predictive ability of the algorithm. Its main effect is to increase the efficiency of species selected for the web. Dynamically constrained webs have lower connectance and indistinct trophic levels compared to real webs and webs from other models, which is a consequence of omitting species' physiology. Although webs are less likely to be permanent if they have high omnivory and cycling, the web-building process circumvents this constraint. The challenges of testing and justifying system-level hypotheses, including isolating and detecting their effects, are discussed.