John Swaddle

Assessing the long-term macroevolutionary consequences of sexual selection has been hampered by the difficulty of studying this process in the fossil record. Cytheroid ostracodes offer an excellent system to explore sexual selection in the fossil record because their readily fossilized carapaces are sexually dimorphic. Specifically, males are relatively more elongate than females in this superfamily. This sexual shape difference is thought to arise so that males carapaces can accommodate their very large copulatory apparatus, which can account for up to one-third of body volume. Here we test this widely held explanation for sexual dimorphism in cytheroid ostracodes by correlating investment in male genitalia, a trait in which sexual selection is seen as the main evolutionary driver, with sexual dimorphism of carapace in the genus Cyprideis. We analyzed specimens collected in the field (C. sale-brosa, USA; C. torosa, UK) and from collections of the National Museum of Natural History, Washington, DC (C. mexicana). We digitized valve outlines in lateral view to obtain measures of size (valve area) and shape (elongation, measured as length to height ratio), and obtained several dimensions from two components of the hemipenis: the muscular basal capsule, which functions as a sperm pump, and the section that includes the intromittent organ (terminal extension). In addition to the assessment of this primary sexual trait, we also quantified two dimensions of the male secondary sexual trait—where the transformed right walking leg functions as a clasping organ during mating. We also measured linear dimensions from four limbs as indicators of overall (soft-part) body size, and assessed allometry of the soft anatomy. We observed significant correlations in males between valve size, but not elongation, and distinct structural parts of the hemipenis, even after accounting for their shared correlation with overall body size. We also found weak but significant positive correlation between valve elongation and the degree of sexual dimorphism of the walking leg, but only in C. torosa. The correlation between the hemipenis parts, especially basal capsule size and male valve size dimorphism suggests that sexual selection on sperm size, quantity,

Synopsis Each year, billions of birds collide with large human-made structures, such as building, towers, and turbines, causing substantial mortality. Such bird-strike, which is projected to increase, poses risks to populations of birds and causes significant economic costs to many industries. Mitigation technologies have been deployed in an attempt to reduce bird-strike, but have been met with limited success. One reason for bird-strike may be that birds fail to pay adequate attention to the space directly in front of them when in level, cruising flight. A warning signal projected in front of a potential strike surface might attract visual attention and reduce the risks of collision. We tested this idea in captive zebra finches (Taeniopygia guttata) that were trained to fly down a long corridor and through an open wooden frame. Once birds were trained, they each experienced three treatments at unpredictable times and in a randomized order: a loud sound field projected immediately in front of the open wooden frame; a mist net (i.e., a benign strike surface) placed inside the wooden frame; and both the loud sound and the mist net. We found that birds slowed their flight approximately 20% more when the sound field was projected in front of the mist net compared with when the mist net was presented alone. This reduction in velocity would equate to a substantial reduction in the force of any collision. In addition to slowing down, birds increased the angle of attack of their body and tail, potentially allowing for more maneuverable flight. Concomitantly, the only cases where birds avoided the mist net occurred in the sound-augmented treatment. Interestingly, the sound field by itself did not demonstrably alter flight. Although our study was conducted in a limited setting, the alterations of flight associated with our sound field has implications for reducing bird-strike in nature and we encourage researchers to test our ideas in field trials.

Recently, numerous studies have observed changes in bird vocalizations—especially song—in urban habitats. These changes are often interpreted as adaptive, since they increase the active space of the signal in its environment. However, the proximate mechanisms driving cross-generational changes in song are still unknown. We performed a captive experiment to identify whether noise experienced during development affects song learning and the development of song-control brain regions. Zebra finches (Taeniopygia guttata) were bred while exposed, or not exposed, to recorded traffic urban noise (Study 1) or pink noise (Study 2). We recorded the songs of male offspring and compared these to fathers' songs. We also measured baseline corticosterone and measured the size of song-control brain regions when the males reached adulthood (Study 1 only). While male zebra finches tended to copy syllables accurately from tutors regardless of noise environment, syntax (the ordering of syllables within songs) was incorrectly copied affected by juveniles exposed to noise. Noise did not affect baseline corticosterone, but did affect the size of brain regions associated with song learning: these regions were smaller in males that had been had been exposed to recorded traffic urban noise in early development. These findings provide a possible mechanism by which noise affects behaviour, leading to potential population differences between wild animals occupying noisier urban environments compared with those in quieter habitats. INTRODUCTION Research into the effects of urban noise on the behaviour of wild animals has increased over the past decade. We now know that several species of birds change their song in association with anthropogenic noise. Specifically, birds appear to alter the frequency (Brumm,

Mercury contamination from mining and fossil fuel combustion causes damage to humans and animals worldwide. Mercury exposure has been implicated in mammalian hearing impairment, but its effect on avian hearing is unknown. In this study, we examined whether lifetime dietary mercury exposure affected hearing in domestic zebra finches (Taeniopygia guttata) by studying their auditory brainstem responses (ABRs). Zebra finches exposed to mercury exhibited elevated hearing thresholds, decreased amplitudes, and longer latencies in the ABR, the first evidence of mercury-induced hearing impairment in birds. Birds are a more appropriate model for the human auditory spectrum than most mammals because of similarities in frequency discrimination , vocal learning, and communication behavior. When mercury is considered in combination with other anthropogenic stressors such as noise pollution and habitat alteration, the hearing impairments we document here could substantially degrade avian auditory communication in wild birds.

Environmental stressors can negatively affect avian cognitive abilities, potentially reducing fitness, for example by altering response to predators, display to mates, or memory of locations of food. We expand on current knowledge by investigating the effects of dietary mercury, a ubiquitous environmental pollutant and known neurotoxin, on avian cognition. Zebra finches Taeniopygia gut-tata were dosed for their entire lives with sub-lethal levels of mercury, at the environmentally relevant dose of 1.2 parts per million. In our first study, we compared the dosed birds with controls of the same age using tests of three cognitive abilities: spatial memory, inhibitory control, and color association. In the spatial memory assay, birds were tested on their ability to learn and remember the location of hidden food in their cage. The inhibitory control assay measured their ability to ignore visible but inaccessible food in favor of a learned behavior that provided the same reward. Finally, the color association task tested each bird's ability to associate a specific color with the presence of hidden food. Dietary mercury negatively affected spatial memory ability but not inhibi-tory control or color association. Our second study focused on three behavioral assays not tied to a specific skill or problem-solving: activity level, neophobia, and social dominance. Zebra finches exposed to dietary mercury throughout their lives were subordinate to, and more active than, control birds. We found no evidence that mercury exposure influenced our metric of neophobia. Together, these results suggest that sub-lethal exposure to environmental mercury selectively harms neurological pathways that control different cognitive abilities, with complex effects on behavior and fitness.

The fields of behavioral ecology, conservation science, and environmental toxicology individually aim to protect and manage the conservation of wildlife in response to anthropogenic stressors, including widespread anthropogenic pollution. Although great emphasis in the field of toxicology has been placed on understanding how single pollutants affect survival, a comprehensive, interdis-ciplinary approach that includes behavioral ecology is essential to address how anthropogenic compounds are a risk for the survival of species and populations in an increasingly polluted world. We provide an integrative framework for behavioral ecotoxicology using Tinbergen's four postulates (causation and mechanism, development and ontogeny, function and fitness, and evolutionary history and phylogenetic patterns). The aims of this review are: 1) to promote an integrative view and redefine the field of integrative behavioral ecotoxicology; 2) to demonstrate how studying ecotoxicology can promote behavior research; and 3) to identify areas of behavioral ecotoxicology that require further attention to promote the integration and growth of the field.

Mercury is a ubiquitous metal contaminant that negatively impacts reproduction of wildlife and has many other sub-lethal effects. Songbirds are sensitive bioindicators of mercury toxicity and may suffer population declines as a result of mercury pollution. Current predictions of mercury accumulation and biomagnification often overlook possible genetic variation in mercury uptake and elimination within species and the potential for evolution in affected populations. We conducted a study of dietary mercury exposure in a model songbird species , maintaining a breeding population of zebra finches (Taeniopygia guttata) on standardized diets ranging from 0.0–2.4 μg/g methylmercury. We applied a quantitative genetics approach to examine patterns of variation and heritability of mercury accumulation within dietary treatments using a method of mixed effects modeling known as the 'animal model'. Significant variation in blood mercury accumulation existed within each treatment for birds exposed at the same dietary level; moreover, this variation was highly repeatable for individuals. We observed substantial genetic variation in blood mercury accumulation for birds exposed at intermediate dietary concentrations. Taken together, this is evidence that genetic variation for factors affecting blood mercury accumulation could be acted on by selection. If similar heritability for mercury accumulation exists in wild populations, selection could result in genetic differentiation for populations in contaminated locations, with possible consequences for mercury biomagnification in food webs.

Collisions between birds and aircraft cause billions of dollars of damages annually to civil, commercial, and military aviation. Yet technology to reduce bird strike is not generally effective, especially over longer time periods. Previous information from our lab indicated that filling an area with acoustic noise, which masks important communication channels for birds, can displace European Starlings (Sturnus vulgaris) from food sources. Here we deployed a spatially controlled noise (termed a “sonic net”), designed to overlap with the frequency range of bird vocalizations, at an airfield. By conducting point counts, we monitored the presence of birds for four weeks before deployment of
our sonic net, and for four weeks during deployment. We found an 82% reduction in bird presence in the sonic net area compared with change in the reference areas. This effect was as strong in the fourth week of exposure as in the first week. We also calculated the
potential costs avoided resulting from this exclusion. We propose that spatially controlled acoustic manipulations that mask auditory communication for birds may be an effective long term and fairly benign way of excluding problem birds from areas of socioeconomic importance, such as airfields, agricultural sites, and commercial properties.

Vocal responses to anthropogenic noise have been documented in several species of songbird. However, only a few studies have investigated whether these adjustments are made in ‘‘real time’’ or are longer-term responses to particular
soundscapes. Furthermore, increased ambient noise often is accompanied by structural changes to the habitat, including the introduction of noisy roadways and the removal of native vegetation. To date, no studies have simultaneously investigated the impact of both acoustic and structural disturbance on the same species. The relevance of each of
these variables must be quantified if we wish to refine our understanding of the ways in which human activities influence
avian communication. In this study, we quantified both among-male and within-male adjustments of song in response to
ambient noise, and also investigated whether anthropogenic modifications of the habitat explained variations in songs’
parameters. Recordings of songs were collected from male, breeding eastern bluebirds (Sialia sialis) residing in a network
of nestboxes distributed across a gradient of anthropogenic disturbance. Levels of ambient noise were associated both
with the average song-parameters of each male and with the change in a male’s song-parameters between the loudest and
quietest periods at his nest box. Males’ song parameters were also related to habitat structure, as assessed using geographic
information systems techniques. Males in noisier sites produced both higher-pitched and louder songs than did
birds in quieter areas. Likewise, individual males demonstrated immediate adjustments to disturbance by noise, increasing
the amplitude of their song between periods of quiet and loud ambient noise. Both spectral and temporal aspects of a
male’s song were related to whether his habitat was more ‘‘natural’’ or ‘‘anthropogenic.’’ Our results indicate that males’
adjustments of song may represent simultaneous responses to multiple modifications of the habitat by humans. However,
we also conclude that biotic noise remains an important influence on avian signals even in anthropogenic areas.
We suggest that human habitats provide an ideal setting in which to perform experiments on communication strategies,
with resulting data poised to reveal underlying evolutionary processes while also informing conservation and management.

ABSTRACT.—Although phylogenetic systematics is used to reconstruct evolutionary relationships, undergraduates have a difficult time mastering its fundamental concepts. Because it is a key part of the mainstream professional thinking, we explored in what ways students misread cladograms, which are the abstract and synthetic diagrams of phylogenetic systematics.

We investigated the effects of the plumage changes associated with moult on the anti-predator take-off performance of European Starlings Sturnus vulgaris. By altering the plumage to simulate moult, we have isolated the biomechanical consequences of changes in wingform from the underlying physiological and metabolic changes that may occur during natural moult. Previous analyses of avian take-off performance have relied on descriptive observations of wingtip kinematics or dual measures of take-off speed and angle.

Male-female pair bonds are common to most bird species, and these bonds affect fundamental aspects of mating systems and the strength of selection, for example, by limiting extrapair paternity. Therefore, understanding factors that strengthen and erode pair bonds are important in elucidating the selection pressures that avian populations will experience.

Abstract Moult is a costly and extremely important stage of the annual cycle for most birds. It also often delimits periods of breeding and migration and so factors that influence moult parameters have important life-history implications. Although the direct energetic and physiological costs of moult have been well-studied, there has been relatively little consideration or formalisation of the biomechanical costs of moult and feather replacement.

The chapter introduces fluctuating asymmetry and shows why it is relevant to behavioral ecologists. It illustrates where the knowledge about fluctuating asymmetry is lacking in important areas, and to offer suggestions that will help behavioral ecologists fill those gaps. The chapter reviews relationships between asymmetry and fitness. These relationships appear to be taxon and trait specific. The specificity of such relationships probably arises from the weak correlation of fluctuating asymmetry with developmental instability. The chapter discusses common methodological problems of studying fluctuating asymmetry. A hypothesis is presented that fluctuating asymmetry may have significant genetic redundancy and could be viewed as a neutral character. Many behavioral ecologists have quantified fluctuating asymmetry in secondary sexual characters and investigated whether such asymmetry plays a role in sexual selection processes. The chapter concludes that a link is established between behavioral studies of fluctuating asymmetry and broader evolutionary questions, and it is suggested that researchers adopt a more integrated approach that brings together a proximate understanding of the impact of asymmetry with more long-term studies of the origins of asymmetry and evolutionary consequences of selection against fluctuating asymmetry.