The island biogeography of human population size.

For decades, biogeographers have sought a better understanding of how organisms are distributed among islands. However, the island biogeography of humans remains largely unknown. Here, we investigate how human population size varies among 486 islands at two spatial scales. At a global scale, we tested whether population size increases with island area and declines with island elevation and nearest mainland, as is common in non-human species, or whether humans escape such biogeographic constraints. At a regional scale, we tested whether population sizes vary among islands within archipelagos according to the positioning of different cultural source pools. Results illustrate that on a global scale, human populations increased in size with island area, similar to non-human species, yet they did not decline in size with elevation and distance to nearest mainland. At a regional scale, human population size often varied among islands within archipelagos relative to the location of different cultural source pools. Despite broad-scale similarities in the geographical distribution of human and non-human species among islands, results from this study indicate that the island biogeography of humans may also be influenced by archipelago-specific social, political and historical circumstances.

Minimal models provide maximally parsimonious explanations.

We are delighted that Diniz-Filho et al. agree with the main premise of our paper, and we welcome their critique, as constructive debate will help foster a better understanding of size evolution on islands. Our perspective on each of their criticisms is discussed in greater detail below.

A simple null model predicts the island rule.

The island rule is a putative pattern in island evolution, where small species become larger on islands and large species become smaller. Despite decades of study, a mechanistic explanation for why some taxonomic groups obey the island rule, while others do not, has yet to be identified. Here, we explore whether the island rule might result from evolutionary drift. We derived a simulation model that predicts evolutionary size changes on islands based on random evolutionary trajectories along bounded trait domains. The model consistently predicted the island rule and could account for its occurrence in plants inhabiting islands in the Southwest Pacific. When support for the island rule was not detected, insular gigantism was often observed, suggesting that natural selection was at work. Overall results indicate that evolutionary drift can provide a parsimonious explanation for the island rule, suggesting future work should focus on circumstances where it does not occur.

Parasite-offspring competition for female resources can explain male-biased parasitism in plants.

Male-biased susceptibility to parasites is common in dioecious plants. However, why males have higher parasite loads than females is unclear. Unlike males, females must subsidize post-fertilization costs of reproduction (e.g. seed and fruit development). As a result, females may have smaller pools of resources potentially available to parasites, thus leading to lower parasite loads. We tested this prediction in New Zealand's largest native moth ( Aenetus virescens: Lepidoptera), whose larvae parasitize Aristotelia serrata (Elaeocarpaceae), an endemic species of dioecious tree. We measured parasite loads in male and female trees, as well as annual seed set in females. We then derived a technique to equate the energetic cost of seed set in females to an equivalent number of parasitic larvae. Our results showed evidence for male-biased parasitism: male trees harboured more larval parasites than female trees. However, when parasite loads in males were compared with parasite loads in females, plus the energetic cost of seed production calculated as an equivalent number of parasitic larvae, differences in parasitism between the sexes disappeared. We conclude that male-biased parasitism in plants could arise from parasite-offspring (i.e. herbivore-seed) competition for female resources.

Can you teach an old parrot new tricks? Cognitive development in wild kaka (Nestor meridionalis).

Despite recent efforts to characterize innovative individuals within a species, we still know very little about the ontogeny of innovation ability. A number of studies have found that innovation rates are correlated with personality traits, such as neophilia and exploration. Juvenile birds are frequently more neophilic and explorative, yet few studies have found evidence of age-related differences in innovative problem-solving success. Here, we show consistently higher innovation efficiency in juveniles of a wild, omnivorous parrot species across a variety of tasks and contexts. We tested 104 kaka (Nestor meridionalis), ranging in age from four months to 13 years. Twenty-four individuals participated in all three of our problem-solving tasks, two of which involved a familiar feeder and one an entirely novel apparatus. Juveniles were the most efficient problem-solvers in all three tasks. By contrast, the adults' success was context dependent and limited to the novel apparatus, which did not require modification of a pre-learned behavioural response. This suggests greater behavioural flexibility in the juvenile birds, who also showed higher persistence and exploratory diversity than adults. These traits may enable young kaka to discover efficient foraging techniques, which are then maintained throughout adulthood.

A Theory of Island Biogeography for Exotic Species.

The theory of island biogeography has played a pivotal role in the way ecologists view communities. However, it does not account for exotic species explicitly, which limits its use as a conservation tool. Here, I present the results of a long-term study of plant communities inhabiting an archipelago of small islands off the coast of New Zealand and derive a modified version of the theory of island biogeography to predict differences in the turnover and diversity of native and exotic species. Empirical results showed that, although species richness of both native and exotic plant species increased with island area, native species consistently outnumbered exotic species. Species turnover increased with species richness in both groups. However, opposite to species-area patterns, turnover increased more rapidly with species richness in exotic species. Empirical results were consistent with the modified version of the theory of island biogeography, which distinguishes exotic species from native species by decoupling extinction rates of exotic species from island area, because they are represented by only small populations at the initial stages of invasion. Overall results illustrate how the theory of island biogeography can be modified to reflect the dynamics of exotic species as they invade archipelagos, expanding its use as a conservation tool.

The repeated evolution of large seeds on islands.

Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous species often evolve to become woody and species frequently evolve larger leaves, regardless of growth form. However, our understanding of how seed sizes might evolve on islands lags far behind other plant traits. Here, we conduct the first test for macroevolutionary patterns of seed size on islands. We tested for differences in seed size between 40 island-mainland taxonomic pairings from four island groups surrounding New Zealand. Seed size data were collected in the field and then augmented by published seed descriptions to produce a more comprehensive dataset. Seed sizes of insular plants were consistently larger than mainland relatives, even after accounting for differences in growth form, dispersal mode and evolutionary history. Selection may favour seed size increases on islands to reduce dispersibility, as long-distance dispersal may result in propagule mortality at sea. Alternatively, larger seeds tend to generate larger seedlings, which are more likely to establish and outcompete neighbours. Our results indicate there is a general tendency for the evolution of large seeds on islands, but the mechanisms responsible for this evolutionary pathway have yet to be fully resolved.

Wild robins (Petroica longipes) respond to human gaze.

Gaze following and awareness of attentional cues are hallmarks of human and non-human social intelligence. Here, we show that the North Island robin (Petroica longipes), a food-hoarding songbird endemic to New Zealand, responds to human eyes. Robins were presented with six different conditions, in which two human experimenters altered the orientation or visibility of their body, head or eyes in relation to mealworm prey. One experimenter had visual access to the prey, and the second experimenter did not. Robins were then given the opportunity to 'steal' one of two mealworms presented by each experimenter. Robins responded by preferentially choosing the mealworm in front of the experimenter who could not see, in all conditions but one. Robins failed to discriminate between experimenters who were facing the mealworm and those who had their head turned 90° to the side. This may suggest that robins do not make decisions using the same eye visibility cues that primates and corvids evince, whether for ecological, experiential or evolutionary reasons.

Large quantity discrimination by North Island robins (Petroica longipes).

While numerosity-representation and enumeration of different numbers of objects-and quantity discrimination in particular have been studied in a wide range of species, very little is known about the numerical abilities of animals in the wild. This study examined spontaneous relative quantity judgments (RQJs) by wild North Island robins (Petroica longipes) of New Zealand. In Experiment 1, robins were tested on a range of numerical values of up to 14 versus 16 items, which were sequentially presented and hidden. In Experiment 2, the same numerical contrasts were tested on a different group of subjects but quantities were presented as whole visible sets. Experiment 3 involved whole visible sets that comprised of exceedingly large quantities of up to 56 versus 64 items. While robins shared with other species a ratio-based representation system for representing very large values, they also appeared to have developed an object indexing system with an extended upper limit (well beyond 4) that may be an evolutionary response to ecological challenges faced by scatter-hoarding birds. These results suggest that cognitive mechanism influencing an understanding of physical quantity may be deployed more flexibly in some contexts than previously thought, and are discussed in light of findings across other mammalian and avian species.

On the selective advantage of coloniality in staghorn ferns (Platycerium bifurcatum, Polypodiaceae).

The staghorn fern (Platycerium bifurcatum, Polypodiaceae) is an epiphyte from Australasia that displays many life history characteristics commonly associated with eusocial animals. Here, I hypothesize about the selective advantage of living in cooperative groups by comparing the morphological characteristics of colonies to their solitary congeners.

Scientists' warning - The outstanding biodiversity of islands is in peril.

Despite islands contributing only 6.7% of land surface area, they harbor ~20% of the Earth's biodiversity, but unfortunately also ~50% of the threatened species and 75% of the known extinctions since the European expansion around the globe. Due to their geological and geographic history and characteristics, islands act simultaneously as cradles of evolutionary diversity and museums of formerly widespread lineages-elements that permit islands to achieve an outstanding endemicity. Nevertheless, the majority of these endemic species are inherently vulnerable due to genetic and demographic factors linked with the way islands are colonized. Here, we stress the great variation of islands in their physical geography (area, isolation, altitude, latitude) and history (age, human colonization, human density). We provide examples of some of the most species rich and iconic insular radiations. Next, we analyze the natural vulnerability of the insular biota, linked to genetic and demographic factors as a result of founder events as well as the typically small population sizes of many island species. We note that, whereas evolution toward island syndromes (including size shifts, derived insular woodiness, altered dispersal ability, loss of defense traits, reduction in clutch size) might have improved the ability of species to thrive under natural conditions on islands, it has simultaneously made island biota disproportionately vulnerable to anthropogenic pressures such as habitat loss, overexploitation, invasive species, and climate change. This has led to the documented extinction of at least 800 insular species in the past 500 years, in addition to the many that had already gone extinct following the arrival of first human colonists on islands in prehistoric times. Finally, we summarize current scientific knowledge on the ongoing biodiversity loss on islands worldwide and express our serious concern that the current trajectory will continue to decimate the unique and irreplaceable natural heritage of the world's islands. We conclude that drastic actions are urgently needed to bend the curve of the alarming rates of island biodiversity loss.

Geographic patterns in fruit colour diversity: do leaves constrain the colour of fleshy fruits?

We tested for geographic patterns in fruit colour diversity. Fruit colours are thought to promote detection by seed dispersers. Because seed dispersers differ in their spectral sensitivities, we predicted that fruit colour diversity would be higher in regions with higher seed disperser diversity (i.e. the tropics). We collected reflectance data on 232 fruiting plant species and their natural backgrounds in seven localities in Europe, North and South America, and analysed fruit colour diversity according to the visual system of birds-the primary consumer types of these fruits. We found no evidence that fruit colours are either more conspicuous or more diverse in tropical areas characterised by higher seed disperser diversity. Instead, fruit colour diversity was lowest in central Brazil, suggesting that fruit colours may be more diverse in temperate regions. Although we found little evidence for geographic variation in fruit hues, the spectral properties of fruits were positively associated with the spectral properties of backgrounds. This result implies that fruit colours may be influenced by selection on the reflectance properties of leaves, thus constraining the evolution of fruit colour. Overall, the results suggest that fruit colours in the tropics are neither more diverse nor more conspicuous than temperate fruits, and that fruit colours may be influenced by correlated selection on leaf reflectance properties.

Memory Performance Influences Male Reproductive Success in a Wild Bird.

Despite decades of comparative research, how selection shapes the evolution of cognitive traits remains poorly understood [1-3]. Several lines of evidence suggest that natural selection acts on spatial memory in food-caching species [3-6]. However, a link between reproductive fitness and spatial memory ability has yet to be demonstrated in any caching species [1, 3, 6]. Here, we show that memory performance influences reproductive success differentially for males and females in a caching songbird, the New Zealand robin (Petroica longipes). Males' memory performance in a spatial task during winter influenced their subsequent breeding success; individuals with more accurate performance produced more fledglings and independent offspring per nesting attempt. Males with superior memory performance also provided an increased proportion of large prey items to chicks in the nest and spent less time flying while foraging and provisioning. No such effects were found for females. Previous research reveals that trade-offs may constrain selection and act to maintain variation in cognitive traits [7]. The gender dimorphism in the reproductive benefits of robin memory performance suggests an additional role for divergent selection between the sexes in constraining runaway selection on male memory ability [8], ultimately maintaining variation in this cognitive trait.

Phenotypic trait matching predicts the topology of an insular plant-bird pollination network.

Conceptualizing species interactions as networks has broadened our understanding of ecological communities. However, the factors shaping interaction patterns among species and, therefore, network structure remain unclear. One potentially important factor is the matching of phenotypic traits. Here, we tested for trait matching in a bird-flower visitation network from New Zealand. We first quantified the overall network structure and tested whether flower size could account for differences in the visitation rates of flowering plants. We then explored the relationship between the flower size and bill size. The results showed that the interaction network is nested. Plant species with large flowers received more visits from birds than plant species with small flowers. Moreover, plant species with large flowers were visited more frequently by birds with large bills, while species with smaller flowers were visited more frequently by birds with small bills. Overall, the interaction patterns between birds and flowering plants could be predicted by their morphology, suggesting that phenotypic trait matching is an important predictor of network structure.

Male New Zealand robins (Petroica longipes) cater to their mate's desire when sharing food in the wild.

In many species that have bi-parental care, food-sharing males provide vital nutritional resources to their mates during reproduction. However, it is currently unknown whether females can signal specific desires to their mates, or if males can cater to female desire in the wild. Here we investigate whether and how wild male North Island robins (Petroica longipes) respond to changes in their mates' desires and nutritional need when sharing food. We demonstrate that wild female robins' desire for particular foods changes over short time periods; when given the choice between two types of insect larvae, females prefer the type they have not recently eaten. In our experiments, wild male robins preferentially shared the larvae type that their mate was most likely to desire and also increased the quantity of food shared if she had begun incubating. Males catered to their mates' desire when female behaviour was the only cue available to guide their choices. This is the first evidence that females may behaviourally communicate their specific food desires to their mates, enabling males to cater to fine-scale changes in their mates' nutritional requirements in the wild. Such a simple behaviour-reading mechanism has the potential to be widespread among other food-sharing species.

Drivers of aggregation in a novel arboreal parasite: the influence of host size and infra-populations.

As a novel arboreal parasite, New Zealand's largest endemic moth, Aenetus virescens, is a biological oddity. With arguably the most unusual lepidopteran life history on earth, larvae grow to 100mm, spending ∼6 years as wood-boring parasites feeding on host tree phloem. Parasite fitness is a product of host suitability. Parasite discrimination between heterogeneous hosts in fragmented populations shapes parasite aggregation. We investigated whether A. virescens aggregation among hosts occurs randomly (target area effect), or if larvae select hosts based on host quality (ideal free distribution). Using long-term larval growth as an indicator of energy intake, we examined A. virescens aggregation in relation to host size and infra-population. Using a generalised linear model, the relationship between parasite intensity and host tree size was analysed. Reduced major axis regression was used to evaluate A. virescens growth after 1 year. Linear mixed-effects models inferred the influence of parasite infra-population on parasite growth, with host tree as a random factor. Results indicate parasite intensity scaled positively with host size. Furthermore, parasite growth remained consistent throughout ontogeny regardless of host size or parasite infra-population. Aenetus virescens aggregation among hosts violates the ideal free distribution hypothesis, occurring instead as a result of host size, supporting the target area effect.

Evaluating Frugivore-fruit Interactions Using Avian Eye Modelling.

Fruit phenotypes are often hypothesised to be affected by selection by frugivores. Here, we tested two hypotheses concerning frugivore-fruit interactions from the perspective of fruit colours. We measured the spectral properties of 26 fruits and the associated leaves of plants from 2 islands in New Zealand. Visual observations were also performed to record the birds that fed on the fruits. First, we tested the fruit-foliage hypothesis, where fruit colours are assumed to be evolutionarily constrained by their own leaf colour to maximise colour contrast and fruit conspicuousness. We ran a null model analysis comparing fruit colour contrast using an avian eye model. Second, we tested the frugivore specificity hypothesis, where specific fruit colours are thought to be connected with a specific bird frugivore. We performed a regression on the number of bird visits against the fruit colour in tetrahedral colour space based on an avian eye calculation using Mantel's test. The results show that fruit colours are not constrained by their own leaf colours. There is also no relationship or pattern suggesting a link between a specific fruit colour and specific bird visitors. We suggest that although fruit colour is one of the most highly discussed components, it is not the most important single deciding factor in frugivore fruit selection.

The ability of North Island Robins to discriminate between humans is related to their behavioural type.

Animals are able to learn to identify persistent threats to themselves and their offspring. For example, birds are able to quickly learn to discriminate between humans that have previously threatened their nests from humans with whom they have had no prior experience. However, no study has yet examined whether a bird's ability to discriminate between humans is related to the bird's underlying behavioural type. In this study, we examined whether there were differences among North Island (NI) robins (Petroica longipes), based on their underlying behavioural type, in their abilities to discriminate between familiar and novel human observers. Using a simple feeding experiment, we timed how long it took birds to attack a food item placed next to an observer on each of 7 days. On the eighth day, a different observer timed the birds. We found that birds could be split into two behaviour types based on their attack behaviour: fast attackers (latencies <20 sec) and slow attackers (latencies >20 secs). Interestingly, the fast birds did not increase their attack latency in response to the novel observer whereas the slow attackers did. This result, for the first time, demonstrates that a bird's ability to discriminate between humans can vary among birds based on their behavioural type.

Is crypsis a common defensive strategy in plants? Speculation on signal deception in the New Zealand flora.

Color is a common feature of animal defense. Herbivorous insects are often colored in shades of green similar to their preferred food plants, making them difficult for predators to locate. Other insects advertise their presence with bright colors after they sequester enough toxins from their food plants to make them unpalatable. Some insects even switch between cryptic and aposomatic coloration during development. Although common in animals, quantitative evidence for color-based defense in plants is rare. After all, the primary function of plant leaves is to absorb light for photosynthesis, rather than reflect light in ways that alter their appearance to herbivores. However, recent research is beginning to challenge the notion that color-based defence is restricted to animals.