Anatomical specializations for enhanced olfactory sensitivity in kiwi, Apteryx mantelli

The ability to function in a nocturnal and ground-dwelling niche requires a unique set of sensory specializations. The New Zealand kiwi has shifted away from vision, instead relying on auditory and tactile stimuli to function in its environment and locate prey. Behavioral evidence suggests that kiwi also rely on their sense of smell, using olfactory cues in foraging and possibly also in communication and social interactions. Anatomical studies appear to support these observations: the olfactory bulbs and tubercles have been suggested to be large in the kiwi relative to other birds, although the extent of this enlargement is poorly understood. In this study, we examine the size of the olfactory bulbs in kiwi and compare them with 55 other bird species, including emus, ostriches, rheas, tinamous, and 2 extinct species of moa (Dinornithiformes). We also examine the cytoarchitecture of the olfactory bulbs and olfactory epithelium to determine if any neural specializations beyond size are present that would increase olfactory acuity. Kiwi were a clear outlier in our analysis, with olfactory bulbs that are proportionately larger than those of any other bird in this study. Emus, close relatives of the kiwi, also had a relative enlargement of the olfactory bulbs, possibly supporting a phylogenetic link to well-developed olfaction. The olfactory bulbs in kiwi are almost in direct contact with the olfactory epithelium, which is indeed well developed and complex, with olfactory receptor cells occupying a large percentage of the epithelium. The anatomy of the kiwi olfactory system supports an enhancement for olfactory sensitivities, which is undoubtedly associated with their unique nocturnal niche.

The anatomy of the bill tip of kiwi and associated somatosensory regions of the brain : comparisons with shorebirds

Three families of probe-foraging birds, Scolopacidae (sandpipers and snipes), Apterygidae (kiwi), and Threskiornithidae (ibises, including spoonbills) have independently evolved long, narrow bills containing clusters of vibration-sensitive mechanoreceptors (Herbst corpuscles) within pits in the bill-tip. These ‘bill-tip organs’ allow birds to detect buried or submerged prey via substrate-borne vibrations and/or interstitial pressure gradients. Shorebirds, kiwi and ibises are only distantly related, with the phylogenetic divide between kiwi and the other two taxa being particularly deep. We compared the bill-tip structure and associated somatosensory regions in the brains of kiwi and shorebirds to understand the degree of convergence of these systems between the two taxa. For comparison, we also included data from other taxa including waterfowl (Anatidae) and parrots (Psittaculidae and Cacatuidae), non-apterygid ratites, and other probe-foraging and non probe-foraging birds including non-scolopacid shorebirds (Charadriidae, Haematopodidae, Recurvirostridae and Sternidae). We show that the bill-tip organ structure was broadly similar between the Apterygidae and Scolopacidae, however some inter-specific variation was found in the number, shape and orientation of sensory pits between the two groups. Kiwi, scolopacid shorebirds, waterfowl and parrots all shared hypertrophy or near-hypertrophy of the principal sensory trigeminal nucleus. Hypertrophy of the nucleus basorostralis, however, occurred only in waterfowl, kiwi, three of the scolopacid species examined and a species of oystercatcher (Charadriiformes: Haematopodidae). Hypertrophy of the principal sensory trigeminal nucleus in kiwi, Scolopacidae, and other tactile specialists appears to have co-evolved alongside bill-tip specializations, whereas hypertrophy of nucleus basorostralis may be influenced to a greater extent by other sensory inputs. We suggest that similarities between kiwi and scolopacid bill-tip organs and associated somatosensory brain regions are likely a result of similar ecological selective pressures, with inter-specific variations reflecting finer-scale niche differentiation.

Is lek breeding rare in bats?

Bats (Mammalia: Chiroptera) are among the most successful mammals and likely display the widest range of mating systems within the Class. One mating system that is underrepresented in the Chiroptera is lek breeding, which is characterized by aggregations of sexually displaying males that are visited by receptive females who appraise male displays and actively choose mates, yet receive no direct benefits such as assistance in parenting. Leks are thought to form when males can defend neither resources nor females, making it more economical to establish small breeding territories and self-advertise through sexual displays. Lekking is rare in mammals, and it has been suggested that a lack in the mobility required by females to economically seek out aggregations of sexually displaying males may explain this rarity. Bats, like birds, do not suffer reduced mobility and yet out of over a thousand described species, only one has been confirmed to breed in leks. We examine the rarity of lekking in bats by providing an overview on the current state of knowledge of their mating systems and discuss the ecological and social determinants for the observed trends, contrasted with the prerequisites of lek-breeding behaviour. We use the breeding behaviour of New Zealand's lesser short-tailed bat Mystacina tuberculata, which is believed to be a lek breeder, as a case study for the examination of potential lekking behaviour in bats, and highlight the importance of such research for the development of effective conservation strategies.

Inner-ear morphology of the New Zealand Kiwi (Apteryx mantelli) suggests high-frequency specialization

The sensory systems of the New Zealand kiwi appear to be uniquely adapted to occupy a nocturnal ground-dwelling niche. In addition to well-developed tactile and olfactory systems, the auditory system shows specializations of the ear, which are maintained along the central nervous system. Here, we provide a detailed description of the auditory nerve, hair cells, and stereovillar bundle orientation of the hair cells in the North Island brown kiwi. The auditory nerve of the kiwi contained about 8,000 fibers. Using the number of hair cells and innervating nerve fibers to calculate a ratio of average innervation density showed that the afferent innervation ratio in kiwi was denser than in most other birds examined. The average diameters of cochlear afferent axons in kiwi showed the typical gradient across the tonotopic axis. The kiwi basilar papilla showed a clear differentiation of tall and short hair cells. The proportion of short hair cells was higher than in the emu and likely reflects a bias towards higher frequencies represented on the kiwi basilar papilla. The orientation of the stereovillar bundles in the kiwi basilar papilla showed a pattern similar to that in most other birds but was most similar to that of the emu. Overall, many features of the auditory nerve, hair cells, and stereovilli bundle orientation in the kiwi are typical of most birds examined. Some features of the kiwi auditory system do, however, support a high-frequency specialization, specifically the innervation density and generally small size of hair-cell somata, whereas others showed the presumed ancestral condition similar to that found in the emu.

Vocal repertoire of the New Zealand kea parrot Nestor notabilis

The unique alpine-living kea parrot Nestor notabilis has been the focus of numerous cognitive studies, but its communication system has so far been largely neglected. We examined 2,884 calls recorded in New Zealand’s Southern Alps. Based on audio and visual spectrographic differences, these calls were categorised into seven distinct call types: the non-oscillating ‘screech’ contact call and ‘mew’; and the oscillating ‘trill’, ‘chatter’, ‘warble’ and ‘whistle’; and a hybrid ‘screech-trill’. Most of these calls contained aspects that were individually unique, in addition to potentially encoding for an individual’s sex and age. Additionally, for each recording, the sender’s previous and next calls were noted, as well as any response given by conspecifics. We found that the previous and next calls made by the sender were most often of the same type, and that the next most likely preceding and/or following call type was the screech call, a contact call which sounds like the ‘kee-ah’ from which the bird’s name derives. As a social bird capable of covering large distances over visually obstructive terrain, long distance contact calls may be of considerable importance for social cohesion. Contact calls allow kea to locate conspecifics and congregate in temporary groups for social activities. The most likely response to any given call was a screech, usually followed by the same type of call as the initial call made by the sender, although responses differed depending on the age of the caller. The exception was the warble, the kea’s play call, to which the most likely response was another warble. Being the most common call type, as well as the default response to another call, it appears that the ‘contagious’ screech contact call plays a central role in kea vocal communication and social cohesion

Morphometric analysis of telencephalic structure in a variety of neognath and paleognath bird species reveals regional differences associated with specific behavioral traits

Birds exhibit a huge array of behavior, ecology and physiology, and occupy nearly every environment on earth, ranging from the desert outback of Australia to the tropical rain forests of Panama. Some birds have adopted a fully nocturnal lifestyle, such as the barn owl and kiwi, while others, such as the albatross, spend nearly their entire life flying over the ocean. Each species has evolved unique adaptations over millions of years to function in their respective niche. In order to increase processing power or network efficiency, many of these adaptations require enlargements and/or specializations of the brain as a whole or of specific brain regions. In this study, we examine the relative size and morphology of 9 telencephalic regions in a number of Paleognath and Neognath birds and relate the findings to differences in behavior and sensory ecology. We pay particular attention to those species that have undergone a relative enlargement of the telencephalon to determine whether this relative increase in telencephalic size is homogeneous across different brain regions or whether particular regions have become differentially enlarged. The analysis indicates that changes in the relative size of telencephalic regions are not homogeneous, with every species showing hypertrophy or hypotrophy of at least one of them. The three-dimensional structure of these regions in different species was also variable, in particular that of the mesopallium in kiwi. The findings from this study provide further evidence that the changes in relative brain size in birds reflect a process of mosaic evolution.

Evidence for an Auditory Fovea in the New Zealand Kiwi (Apteryx mantelli).

Kiwi are rare and strictly protected birds of iconic status in New Zealand. Yet, perhaps due to their unusual, nocturnal lifestyle, surprisingly little is known about their behaviour or physiology. In the present study, we exploited known correlations between morphology and physiology in the avian inner ear and brainstem to predict the frequency range of best hearing in the North Island brown kiwi. The mechanosensitive hair bundles of the sensory hair cells in the basilar papilla showed the typical change from tall bundles with few stereovilli to short bundles with many stereovilli along the apical-to-basal tonotopic axis. In contrast to most birds, however, the change was considerably less in the basal half of the epithelium. Dendritic lengths in the brainstem nucleus laminaris also showed the typical change along the tonotopic axis. However, as in the basilar papilla, the change was much less pronounced in the presumed high-frequency regions. Together, these morphological data suggest a fovea-like overrepresentation of a narrow high-frequency band in kiwi. Based on known correlations of hair-cell microanatomy and physiological responses in other birds, a specific prediction for the frequency representation along the basilar papilla of the kiwi was derived. The predicted overrepresentation of approximately 4-6 kHz matches potentially salient frequency bands of kiwi vocalisations and may thus be an adaptation to a nocturnal lifestyle in which auditory communication plays a dominant role.

The presence of ultrasonic harmonics in the calls of the rifleman (Acanthisitta chloris)

Ultrasonic vocalisations (frequencies > 20 kHz) have been extensively studied in the context of echolocation by bats and other mammals (Sales & Pye 1974; Wilson & Hare 2004). Ultrasonic calls have also been recorded from birds, including the blue-throated hummingbird ( Lampornis clemenciae ) (Pytte et al. 2004), where it was first thought that individuals made use of high pitch calls to avoid masking by background noise in a visually obscured environment. Similarly, city-dwelling great tits ( Parus major ) use song with a higher minimum frequency (although not ultrasonic) compared to woodland birds to communicate with conspecifics to avoid the predominantly low-frequency background noise in the city (Slabbekorn & Peet 2003). The theory that birds use ultrasound to avoid noise masking was discarded when it was discovered that there was no corresponding auditory brainstem response (i.e. sensory perception) to the ultrasonic calls in the hummingbirds producing those calls.

Evolution of brain size in the palaeognath lineage, with an emphasis on New Zealand ratites

Brain size in vertebrates varies principally with body size. Although many studies have examined the variation of brain size in birds, there is little information on Palaeognaths, which include the ratite lineage of kiwi, emu, ostrich and extinct moa, as well as the tinamous. Therefore, we set out to determine to what extent the evolution of brain size in Palaeognaths parallels that of other birds, i. e., Neognaths, by analyzing the variation in the relative sizes of the brain and cerebral hemispheres of several species of ratites and tinamous. Our results indicate that the Palaeognaths possess relatively smaller brains and cerebral hemispheres than the Neognaths, with the exception of the kiwi radiation (Apteryx spp.). The external morphology and relatively large size of the brain of Apteryx, as well as the relatively large size of its telencephalon, contrast with other Palaeognaths, including two species of historically sympatric moa, suggesting that unique selective pressures towards increasing brain size accompanied the evolution of kiwi. Indeed, the size of the cerebral hemispheres with respect to total brain size of kiwi is rivaled only by a handful of parrots and songbirds, despite a lack of evidence of any advanced behavioral/ cognitive abilities such as those reported for parrots and crows. In addition, the enlargement in brain and telencephalon size of the kiwi occurs despite the fact that this is a precocial bird. These findings form an exception to, and hence challenge, the current rules that govern changes in relative brain size in birds. Copyright (c) 2007 S. Karger AG, Basel.

Kiwi Forego Vision in the Guidance of Their Nocturnal Activities

Background In vision, there is a trade-off between sensitivity and resolution, and any eye which maximises information gain at low light levels needs to be large. This imposes exacting constraints upon vision in nocturnal flying birds. Eyes are essentially heavy, fluid-filled chambers, and in flying birds their increased size is countered by selection for both reduced body mass and the distribution of mass towards the body core. Freed from these mass constraints, it would be predicted that in flightless birds nocturnality should favour the evolution of large eyes and reliance upon visual cues for the guidance of activity. Methodology/Principal Findings We show that in Kiwi (Apterygidae), flightlessness and nocturnality have, in fact, resulted in the opposite outcome. Kiwi show minimal reliance upon vision indicated by eye structure, visual field topography, and brain structures, and increased reliance upon tactile and olfactory information. Conclusions/Significance This lack of reliance upon vision and increased reliance upon tactile and olfactory information in Kiwi is markedly similar to the situation in nocturnal mammals that exploit the forest floor. That Kiwi and mammals evolved to exploit these habitats quite independently provides evidence for convergent evolution in their sensory capacities that are tuned to a common set of perceptual challenges found in forest floor habitats at night and which cannot be met by the vertebrate visual system. We propose that the Kiwi visual system has undergone adaptive regressive evolution driven by the trade-off between the relatively low rate of gain of visual information that is possible at low light levels, and the metabolic costs of extracting that information.

A quay crane scheduling methodology for multimodal container terminals

This thesis develops an operational decision support tool for container terminal managements. The tool generates efficient schedules for shore cranes handling containers carried by mega container vessels.

Accounting for management costs in sensitivity analyses of matrix population models

Traditional sensitivity and elasticity analyses of matrix population models have been used to inform management decisions, but they ignore the economic costs of manipulating vital rates. For example, the growth rate of a population is often most sensitive to changes in adult survival rate, but this does not mean that increasing that rate is the best option for managing the population because it may be much more expensive than other options. To explore how managers should optimize their manipulation of vital rates, we incorporated the cost of changing those rates into matrix population models. We derived analytic expressions for locations in parameter space where managers should shift between management of fecundity and survival, for the balance between fecundity and survival management at those boundaries, and for the allocation of management resources to sustain that optimal balance. For simple matrices, the optimal budget allocation can often be expressed as simple functions of vital rates and the relative costs of changing them. We applied our method to management of the Helmeted Honeyeater (Lichenostomus melanops cassidix; an endangered Australian bird) and the koala (Phascolarctos cinereus) as examples. Our method showed that cost-efficient management of the Helmeted Honeyeater should focus on increasing fecundity via nest protection, whereas optimal koala management should focus on manipulating both fecundity and survival simultaneously. These findings are contrary to the cost-negligent recommendations of elasticity analysis, which would suggest focusing on managing survival in both cases. A further investigation of Helmeted Honeyeater management options, based on an individual-based model incorporating density dependence, spatial structure, and environmental stochasticity, confirmed that fecundity management was the most cost-effective strategy. Our results demonstrate that decisions that ignore economic factors will reduce management efficiency. ©2006 Society for Conservation Biology.

Regional avian species declines estimated from volunteer-collected long-term data using List Length Analysis

Long-term systematic population monitoring data sets are rare but are essential in identifying changes in species abundance. In contrast, community groups and natural history organizations have collected many species lists. These represent a large, untapped source of information on changes in abundance but are generally considered of little value. The major problem with using species lists to detect population changes is that the amount of effort used to obtain the list is often uncontrolled and usually unknown. It has been suggested that using the number of species on the list, the "list length," can be a measure of effort. This paper significantly extends the utility of Franklin's approach using Bayesian logistic regression. We demonstrate the value of List Length Analysis to model changes in species prevalence (i.e., the proportion of lists on which the species occurs) using bird lists collected by a local bird club over 40 years around Brisbane, southeast Queensland, Australia. We estimate the magnitude and certainty of change for 269 bird species and calculate the probabilities that there have been declines and increases of given magnitudes. List Length Analysis confirmed suspected species declines and increases. This method is an important complement to systematically designed intensive monitoring schemes and provides a means of utilizing data that may otherwise be deemed useless. The results of List Length Analysis can be used for targeting species of conservation concern for listing purposes or for more intensive monitoring. While Bayesian methods are not essential for List Length Analysis, they can offer more flexibility in interrogating the data and are able to provide a range of parameters that are easy to interpret and can facilitate conservation listing and prioritization. © 2010 by the Ecological Society of America.

Implicating animals: The symbolic power of scapegoating animals in marine pollution debates

Animals are often used as symbols in policy debates and media accounts of marine pollution. Images of miserable oil-soaked marine birds and mammals are prominent following high profile oil spills such as the Exxon Valdez, Prestige and Pacific Adventurer incidents. Portrayed as hapless victims, these animal actors are not only cast as powerful symbols of the effects of anthropogenic pollution but also represent an environment in crisis. Animals, like the broader environment, are seen as something which is acted upon. Less attention has been given to the ways in which animals have been cast as either the cause of marine pollution or as having the potential to actively mitigate the potential impacts of anthropogenic marine pollution. This article explores how animals are constructed with respect to vessel-sourced sewage pollution. Through a process of interpretive policy analysis, drawing on media reports and responses to an Australian regulatory review process this study found that, when defending the perceived right to pollute recreational boaters implicated animals such as dogs, fish, turtles, dolphins and seabirds in their pollution discourses. Scapegoating was an important rhetorical feature of claims-making strategies designed to avoid responsibility for changing sewage disposal practices.

Robust sensing for a 3,500 tonne field robot

The mining industry is highly suitable for the application of robotics and automation technology since the work is arduous, dangerous and often repetitive. This paper discusses a robust sensing system developed to find and trade the position of the hoist ropes of a dragline. Draglines are large `walking cranes' used in open-pit coal mining to remove the material covering the coal seam. The rope sensing system developed uses two time-of-flight laser scanners. The finding algorithm uses a novel data association and tracking strategy based on pairing rope data.