Kevin C. Burns

Adaptive numerical competency in a food-hoarding songbird

Most animals can distinguish between small quantities (less than four) innately. Many animals can also distinguish between larger quantities after extensive training. However, the adaptive significance of numerical discriminations in wild animals is almost completely unknown. We conducted a series of experiments to test whether a food-hoarding songbird, the New Zealand robin Petroica australis, uses numerical judgements when retrieving and pilfering cached food. Different numbers of mealworms were presented sequentially to wild birds in a pair of artificial cache sites, which were then obscured from view. Robins frequently chose the site containing more prey, and the accuracy of their number discriminations declined linearly with the total number of prey concealed, rising above-chance expectations in trials containing up to 12 prey items. A series of complementary experiments showed that these results could not be explained by time, volume, orientation, order or sensory confounds. Lastly, a violation of expectancy experiment, in which birds were allowed to retrieve a fraction of the prey they were originally offered, showed that birds searched for longer when they expected to retrieve more prey. Overall results indicate that New Zealand robins use a sophisticated numerical sense to retrieve and pilfer stored food, thus providing a critical link in understanding the evolution of numerical competency.

Seed dispersal by weta

Weta are giant, flightless grasshoppers that are endemic to New Zealand. In the absence of native mammals, weta are thought to perform similar ecological functions. As such, they might be expected to be important seeds dispersers. However, insects are not known to consume fleshy fruits and to disperse seeds after gut passage. We conducted a series of observations and experiments to test whether weta form mutualistic partnerships with fleshy-fruited plants as seed dispersers, similar to small mammals elsewhere in the world. Results showed that weta are indeed effective seeds dispersers, providing an example of ecological convergence between unrelated organisms.

Ontogenetic colour changes in an insular tree species: signalling to extinct browsing birds?

* Animals often use colours to hide from predators (crypsis) or advertise defences (aposematism), but there is little evidence for colour-based defence in plants. * Here, we test whether ontogenetic changes in leaf colour of lancewood (Pseudopanax crassifolius) may have been part of a defensive strategy against flightless browsing birds called moa, which were once the only large herbivores in New Zealand. We tested this hypothesis by conducting spectrographic measurements on different-sized plants grown in a common garden. We also compared these results with observations on a closely related, derived species that evolved in the absence of moa on the Chatham Islands. * Spectrographic analyses showed that birds would have difficulty distinguishing seedling leaves against a background of leaf litter. Conversely, brightly coloured tissues flanking spines on sapling leaves are highly conspicuous to birds. Once above the reach of the tallest known moa, adults produce leaves that are typical in appearance to adult leaves. The Chatham Island species lacks ontogenetic colour changes entirely. * Overall, the results indicate that P. crassifolius goes through a remarkable series of colour changes during development, from cryptically coloured seedlings to aposematically coloured saplings, which may have formed a defensive strategy to protect against giant browsing birds.

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.

Cache spacing patterns and reciprocal cache theft in New Zealand robins

We quantified cache spacing patterns, cache retrieval rates and rates of cache theft in New Zealand robins, Petroica australis. In a field experiment we presented wild birds with a superabundant supply of mealworms. Trials were conducted on competitively dominant males and subordinate females, both when birds were alone and when they were accompanied by their mate. We hypothesized that (1) dominant males would aggregate caches to facilitate their defence, whereas subordinate females would scatter their caches more widely to avoid their discovery by males, (2) sexual differences in cache spacing would be context dependent, or occur only when birds were in pairs, and (3) patterns in cache spacing would facilitate cache retrieval and reduce cache theft. Females cached food further from food sources than males when they occurred in pairs. However, both sexes cached at similar distances when they were alone. Regardless of social context, females created more cache sites than males, and both sexes created more cache sites when they were together. Cache theft was frequently observed. Female-made caches were retrieved at similar rates by both males and females whereas male-made caches were more likely to be stolen by females than retrieved by the males that made them. Therefore, the results showed no support for the hypothesis that cache spacing patterns reduce cache theft. We suggest that the apparently conflicting relations between cache spacing, recovery and theft are determined by a trade-off between winter survival tactics and mate provisioning.

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.

What causes size coupling in fruit–frugivore interaction webs?

The simplest and arguably the most ubiquitous pattern in seed dispersal mutualisms is size coupling: large frugivores tend to consume larger fruits and small frugivores tend to consume smaller fruits. Despite the simplicity of this pattern, the potential mechanisms responsible for fruit--frugivore size coupling are mechanistically divergent and poorly resolved. Size coupling could arise deterministically, if large frugivores actively seek out larger fruits to maximize their foraging efficiency. Alternatively, size coupling could also arise passively, if frugivores forage randomly, but are able to consume only those fruit species that are smaller than their gape width. I observed birds forage for fruits in a New Zealand forest reserve at approximately five-day intervals for six years to test for fruit--frugivore size coupling. I then derived a suite of network analyses to establish whether fruit--frugivore size coupling was best explained by active or passive foraging by frugivores. Results showed a strikingly strong pattern in size coupling; the average size of fruits consumed by each frugivore species increased with their maximum gape width. Simulation analyses revealed that over 70% of variation in interaction frequencies in the observed fruit-frugivore web could be explained by a size-constrained, passive, foraging model. Foraging models in which birds foraged actively for different-sized fruits to improve their foraging efficiency performed more poorly. Results were therefore consistent with the hypothesis that apparently nonrandom patterns in seed dispersal mutualisms can sometimes arise from simple stochastic processes.

Plastic heteroblasty in beach groundsel (Senecio lautus)

Abstract Senecio lautus (Asteraceae) is a heteroblastic herb that produces entire juvenile leaves and lobed adult leaves. Juveniles commonly grow from rock fissures where they are shaded and sheltered from high winds, whereas adult plants are exposed to high light and wind as they outgrow these refuges. Because the duration each plant is sheltered varies with the depth of rock fissures, I hypothesised that heteroblasty in S. lautus varies plastically in response to environmental conditions. I tested this hypothesis in a glasshouse experiment, which exposed developing plants to different light and wind conditions. Plants grown in windy conditions showed similar ontogenetic changes in leaf morphology to plants growing in a control treatment. However, the juvenile‐adult transition was slowed in shaded conditions, indicating that heteroblasty in S. lautus varies plastically in response to light. Results were therefore consistent with the hypothesis that plastic heteroblasty is favoured in species that experience unpredictable environmental gradients during ontogeny.

Mistletoe macroecology: spatial patterns in species diversity and host use across Australia

Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macroecological scales. Because of their highly unusual life history, investigating mistletoe macroecology may provide new insight into broad-scale patterns in species distributions. We collated data on the spatial distribution and host use of 65 species of Loranthaceous mistletoes across the continent of Australia, and tested two predictions. First, we predicted mistletoe diversity would be unrelated to productivity (i.e. evapotranspiration and precipitation), as the parasitic lifestyle might relax environmental constraints on their distributions. Second, we predicted that mistletoe host ranges (number of infected host species) would increase in areas with more potential host species. The basis of this prediction is that greater host generality is likely to evolve in regions with greater host diversity because of greater unpredictability in encounter rates with particular host species. Conversely, in regions with fewer potential hosts, randomly dispersing mistletoe propagules are likely to repeatedly encounter particular host species, thus favouring the evolution of host specialization. The results were generally consistent with these predictions. Mistletoe diversity across Australia was weakly associated with environmental conditions, whereas mistletoe host ranges increased significantly with total plant diversity. Macroecological patterns in mistletoes are unusual. In contrast to non-parasitic plants, mistletoe diversity is poorly correlated with productivity. Host ranges varied predictably across Australia, providing the first quantitative support for the hypothesis that mistletoes in diverse regions tend to be host generalists, whereas mistletoes in depauperate regions tend to be host specialists.

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.1 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 defense is restricted to animals.