Lower survival of hybrid grosbeaks, but not towhees, suggests a molt divide disfavors hybrids.

Although avian hybrid zones in the Great Plains have been studied for almost 70 years, we know surprisingly little about the fitness costs to hybrids that keep these zones narrow. We compare age ratios in grosbeaks (Pheucticus ludovicianus and P. melanocephalus) and towhees (Pipilo erythropthalums and P. maculatus), two species pairs that differ in their life histories and molt schedules, to evaluate survival between hybrids and parentals. We then contrast molt and migratory divides as possible sources of selection against hybrids. Hybrid grosbeaks had 27%-33% lower survival relative to their parentals, whereas hybrid towhees had survival rates similar to parentals. Age ratio data for hybrid grosbeaks suggest high mortality in older birds, as expected if selection operates after the first year of life. This pattern is consistent with parental species of grosbeaks having contrasting molt schedules relative to migration, suggesting high mortality costs to hybrids driven by molt biology, which are expressed later in life. Contrasts in molt schedules are absent in towhees. While migratory divides may exist for towhees and grosbeaks, the low adult survival of hybrid grosbeaks suggest that molt may be an important and underappreciated source of selection maintaining this and other narrow avian hybrid zones.

A complex genomic architecture underlies reproductive isolation in a North American oriole hybrid zone.

Natural hybrid zones provide powerful opportunities for identifying the mechanisms that facilitate and inhibit speciation. Documenting the extent of genomic admixture allows us to discern the architecture of reproductive isolation through the identification of isolating barriers. This approach is particularly powerful for characterizing the accumulation of isolating barriers in systems exhibiting varying levels of genomic divergence. Here, we use a hybrid zone between two species-the Baltimore (Icterus galbula) and Bullock's (I. bullockii) orioles-to investigate this architecture of reproductive isolation. We combine whole genome re-sequencing with data from an additional 313 individuals amplityped at ancestry-informative markers to characterize fine-scale patterns of admixture, and to quantify links between genes and the plumage traits. On a genome-wide scale, we document several putative barriers to reproduction, including elevated peaks of divergence above a generally high genomic baseline, a large putative inversion on the Z chromosome, and complex interactions between melanogenesis-pathway candidate genes. Concordant and coincident clines for these different genomic regions further suggest the coupling of pre- and post-mating barriers. Our findings of complex and coupled interactions between pre- and post-mating barriers suggest a relatively rapid accumulation of barriers between these species, and they demonstrate the complexities of the speciation process.

Revisiting the Baltimore-Bullock's Oriole hybrid zone reveals changing plumage colour in Bullock's Orioles.

Hybrid zones are dynamic areas where populations of two or more interbreeding species may change through an influx of novel genetic material resulting from hybridization or selection on standing genetic variation. Documenting changes in populations through time, however, is challenging because repeated samples are often missing or because long-term storage can affect trait morphologies, especially colour traits that may fade through time. We document a change in carotenoid-based orange breast feathers of Bullock's Orioles (Icterus bullockii) from the Great Plains hybrid zone, USA. Contemporary Bullock's Orioles are more orange than historic individuals from the same location sampled approximately 60 years ago. Spectrophotometry revealed that contemporary Bullock's Orioles resemble orange colour profiles of Baltimore Orioles (I. galbula), the species with which they hybridize. Fading or changes in diet hypotheses do not appear to explain the shift in colour we report for Bullock's Orioles. We propose that these changes in colour are facilitated through introgression with Baltimore Orioles, and favoured by females that choose brighter, more orange males. Our study highlights the long memory of natural history collections and how they offer new insights to the dynamic roll of hybrid zones in trait evolution between interacting species.

Declining growth of natural history collections fails future generations.

New additions of vertebrate specimens to natural history collections are declining in the midst of widespread and accelerating environmental change. Reversing these trends in collecting efforts is essential for addressing future unforeseen ecological issues.

Revisiting a classic hybrid zone: Movement of the northern flicker hybrid zone in contemporary times.

Natural hybrid zones have provided important insights into the evolutionary process, and their geographic dynamics over time can help to disentangle the underlying biological processes that maintain them. Here, we leverage replicated sampling of an identical transect across the hybrid zone between yellow-shafted and red-shafted flickers in the Great Plains to assess its stability over ∼60 years (1955-1957 to 2016-2018). We identify a ∼73-km westward shift in the hybrid zone center toward the range of the red-shafted flicker, but find no associated changes in width over our sampling period. In fact, the hybrid zone remains remarkably narrow, suggesting some kind of selective pressure maintains the zone. By comparing to previous work in the same geographic region, it appears likely that the movement in the hybrid zone has occurred in the years since the early 1980s. This recent movement may be related to changes in climate or land management practices that have allowed westward movement of yellow-shafted flickers into the Great Plains.

Despotic aggression in pre-moulting painted buntings.

Aggression in territorial social systems is easy to interpret because the benefits of territorial defence mostly accrue to the territorial holder. However, in non-territorial systems, high aggression seems puzzling and raises intriguing evolutionary questions. We describe extreme rates of despotism between age classes in a passerine bird, the painted bunting (Passerina ciris), during the pre-moulting period. Aggressive encounters were not associated with aggressors gaining immediate access to resources. Instead, conspecifics, and even other species, were pursued as though being harassed; this aggression generated an ideal despotic habitat distribution such that densities of adult males were higher in high-quality sites. Aggression was not a by-product of elevated testosterone carried over from the breeding season but, rather, appeared associated with dehydroepiandrosterone, a hormone that changes rates of aggression in non-breeding birds without generating the detrimental effects of high testosterone titres that control aggression in the breeding season. This extraordinary pre-moult aggression seems puzzling because individual buntings do not hold defined territories during their moult. We speculate that this high aggression evolved as a means of regulating the number of conspecifics that moulted in what were historically small habitat patches with limited food for supporting the extremely rapid moults of painted buntings.

Reconstructing avian mercury concentrations through time using museum specimens from New York State.

We examined how variation in MeHg concentrations through time is reflected in birds, a taxon commonly used as a biological indicator of ecosystem health. Using museum specimens collected from 1880 to 2016, we measured feather MeHg concentrations in six species of birds that breed in New York State and have distinct dietary and habitat preferences. We predicted that MeHg concentrations in feathers would mirror Hg emission patterns in New York State and increase through time until 1980 then decrease thereafter in response to increased regulation of anthropogenic Hg emissions. We found that MeHg concentrations increased with δ15N, and that MeHg feather concentrations for some individuals from four of the six species examined exceeded concentrations known to cause negative sublethal effects in birds. In contrast to our prediction, MeHg concentrations in feathers did not parallel global or local Hg emissions through time and varied by species, even after controlling for possible changes in diet and habitat. MeHg concentrations varied substantially within species and individual specimens, suggesting that high within-individual variation in feather MeHg concentrations caused by spatiotemporal variation in molt, environmental Hg exposure, or mobility decoupling Hg uptake from breeding sites, may obscure trends in MeHg through time. Our study provides a unique assessment of feather MeHg in six species not typically analyzed using this retrospective approach.

Geographic variation in incubation behavior of a widely distributed passerine bird.

Incubating birds must trade-off leaving the nest to forage with staying on the nest to maintain optimal temperatures for developing embryos. This trade-off is expressed through incubation behavior, which can be heavily influenced by climate, food availability, attentiveness of their mates, and nest predation risk. Comparative studies across species have shown that incubation behavior varies across latitude, but few studies have explored how incubation behavior varies across sites within species. We might expect incubation behavior to be flexible and respond to local environmental challenges; alternatively, behavior may be relatively fixed and vary little across a species' range. We explored four incubation behaviors (male feeding rate, female off-bout duration, female off-bout frequency, and the proportion of time incubating females spent on the nest) in a widespread songbird, the yellow warbler (Setophaga petechia), breeding at a temperate and subarctic site. As temperatures warmed at both sites, males fed females less often, and as male feeding rates decreased, off-bout durations and frequencies increased causing the proportion of time on the nest to decrease. While incubation behaviors changed in similar ways between sites, off-bout durations shortened with increasing male feeding rates most strongly at the temperate site. Overall, these results show flexibility in incubation behaviors in response to different environmental cues, which likely minimize costs associated with provisioning incubating parents and maintaining warm nest temperatures, and suggests that male feeding may be especially important for breeding in cold regions.

Higher Nest Predation Favors Rapid Fledging at the Cost of Plumage Quality in Nestling Birds.

High predation risk can favor rapid offspring development at the expense of offspring quality. Impacts of rapid development on phenotypic quality should be most readily expressed in traits that minimize fitness costs. We hypothesize that ephemeral traits that are replaced or repaired after a short period of life might express trade-offs in quality as a result of rapid development more strongly than traits used throughout life. We explored this idea for plumage quality in nestling body feathers, an ephemeral trait. We found a strong trade-off whereby nestlings that spend less time in the nest produced lower-quality plumage with less dense barbs relative to adults across 123 temperate and tropical species. For a subset of these species ( n=67 ), we found that variation in the risk of nest predation explained additional variation in plumage quality beyond development time. Ultimately, the fitness costs of a poor-quality ephemeral trait, such as nestling body feathers, may be outweighed by the fitness benefits of shorter development times that reduce predation risk. At the same time, reduced resource allocation to traits with small fitness costs, such as ephemeral traits, may ameliorate resource constraints from rapid development on traits with larger fitness impacts.

Primary molt in Gruiforms and simpler molt summary tables.

Molt summary tables reveal the sequence and mode of flight-feather replacement and how these feathers are divided into independent replacement series. Tables for summarizing molt are relatively new, and the rules for generating them were first formally illustrated using data from a temperate passerine. However, this first illustration failed to address (i) species with primaries divided into more than one replacement series, (ii) species with stepwise primary replacement, which almost always involves incomplete annual replacement of the primaries, and (iii) species with incomplete annual replacement within molt series characterized by single-wave replacement. Here, we review complications that arise in developing molt summary tables for such cases and we offer solutions that remove ambiguity about the direction that molt proceeds within a replacement series and about the recognition of nodal and terminal feathers that mark the beginning and end of molt series. We use these modified molt summary tables to describe the sequence of primary replacement in four groups of Gruiform birds, a group for which primary replacement has been reported to proceed from the outermost primary toward the body, unlike most other birds. Eighty molting Grey-winged Trumpeters, Psophia crepitans, and 124 molting Limpkins, Aramus guarauna, show the sequence of primary replacement is proximal in both groups; furthermore, the primaries of trumpeters are divided into two replacement series, one beginning at the outermost primary P10, and the other beginning at P3. To further evaluate the extent of this highly unusual direction of replacement in Gruiforms, we cast the data (Stresemann & Stresemann, 1966) on primary replacement in upland rails (Rallidae) and flufftails (Sarothruridae) into molt summary tables; both also replace their primaries proximally, from outermost to innermost, suggesting that this mode of primary replacement may be characteristic of Gruiformes.

Breeding and multiple waves of primary molt in common ground doves of coastal Sinaloa.

For adult Common Ground Doves from Sinaloa we demonstrate that the primaries are a single molt series, which sometimes feature two (in one case three) waves of feather replacement. Such stepwise primary replacement is found in many large birds but, at 40 g, this dove is much the smallest species reported to have multiple waves of replacement proceeding through its primaries simultaneously. Pre-breeding juvenile Common Ground Doves never feature two waves of primary replacement. Juveniles usually have more than two adjacent feathers growing simultaneously and replace their primaries in about 100 days. In contrast adults, which extensively overlap molt and breeding, usually grow just a single primary at a time, and require at least 145 days to replace their primaries. Molt arrests are thought to drive the generation of new waves of primary replacement in a diversity of large birds. For adult Common Ground Doves, we found molt arrests to be strongly associated with active crop glands, suggesting that the demands of parental care cause arrests in primary replacement in this dove. For those adults with two primary molt waves, initiation of an inner wave was most frequently observed once the outer wave had reached P10. Thus, unlike reports for large birds, Common Ground Doves usually suppress the initiation of a new wave of molt starting at P1 when the preceding wave arrests before reaching the distal primaries. This assures that relatively fresh inner primaries are not replaced redundantly, overcoming a serious flaw in stepwise molting in large birds (Rohwer, 1999).

Fluff-thieving birds sabotage seed dispersal.

Characterizing many species interactions as mutualisms can be misleading because some members of the interaction derive greater fitness benefits at the expense of other members. We provide detailed natural history data on a suspected bird-plant mutualism in South Africa where many species of birds use fluffy Eriocephalus seed material to construct their nests, potentially dispersing seeds for the plant. We focus on a common bird, Prinia maculosa, which invests heavily in gathering Eriocephalus material. Prinias spent 5 of their median 6-day nest construction period adding seed material to their nests and frequently travelled outside their territory boundary to gather Eriocephalus material. Yet, prinias gathered primarily Eriocephalus fluff and actively avoided gathering seeds. The average prinia nest contained only 6.6 seeds, but contained fluff from 579 seeds. These data suggest that prinias provide limited dispersal benefits to Eriocephalus plants. By contrast, the large amounts of Eriocephalus fluff in prinia nests, and the effort that prinias invest in gathering it, suggest that prinias benefit from constructing their nests with Eriocephalus material. We end by outlining hypotheses for possible fitness benefits that Eriocephalus material could provide prinias and other birds.

Variation in the structure of bird nests between northern Manitoba and southeastern Ontario.

Traits that converge in appearance under similar environmental conditions among phylogenetically independent lineages are thought to represent adaptations to local environments. We tested for convergence in nest morphology and composition of birds breeding in two ecologically different locations in Canada: Churchill in northern Manitoba and Elgin in southeastern Ontario. We examined nests from four families of passerine birds (Turdidae: Turdus, Parulidae: Dendroica, Emberizidae: Passerculus and Fringillidae: Carduelis) where closely related populations or species breed in both locations. Nests of American Robins, Yellow Warblers, and Carduelis finches had heavier nest masses, and tended to have thicker nest-walls, in northern Manitoba compared with conspecifics or congenerics breeding in southeastern Ontario. Together, all species showed evidence for wider internal and external nest-cup diameters in northern Manitoba, while individual species showed varying patterns for internal nest-cup and external nest depths. American Robins, Yellow Warblers, and Carduelis finches in northern Manitoba achieved heavier nest masses in different ways. American Robins increased all materials in similar proportions, and Yellow Warblers and Common Redpolls used greater amounts of select materials. While changes in nest composition vary uniquely for each species, the pattern of larger nests in northern Manitoba compared to southeastern Ontario in three of our four phylogenetically-independent comparisons suggests that birds are adapting to similar selective pressures between locations.

Whistling in caterpillars (Amorpha juglandis, Bombycoidea): sound-producing mechanism and function.

Caterpillar defenses have been researched extensively, and, although most studies focus on visually communicated signals, little is known about the role that sounds play in defense. We report on whistling, a novel form of sound production for caterpillars and rare for insects in general. The North American walnut sphinx (Amorpha juglandis) produces whistle 'trains' ranging from 44 to 2060 ms in duration and comprising one to eight whistles. Sounds were categorized into three types: broadband, pure whistles and multi-harmonic plus broadband, with mean dominant frequencies at 15 kHz, 9 kHz and 22 kHz, respectively. The mechanism of sound production was determined by selectively obstructing abdominal spiracles, monitoring air flow at different spiracles using a laser vibrometer and recording body movements associated with sound production using high-speed video. Contractions of the anterior body segments always accompanied sound production, forcing air through a pair of enlarged spiracles on the eighth abdominal segment. We tested the hypothesis that sounds function in defense using simulated attacks with blunt forceps and natural attacks with an avian predator - the yellow warbler (Dendroica petechia). In simulated attacks, 94% of caterpillars responded with whistle trains that were frequently accompanied by directed thrashing but no obvious chemical defense. In predator trials, all birds readily attacked the caterpillar, eliciting whistle trains each time. Birds responded to whistling by hesitating, jumping back or diving away from the sound source. We conclude that caterpillar whistles are defensive and propose that they function specifically as acoustic 'eye spots' to startle predators.

Migratory double breeding in Neotropical migrant birds.

Neotropical migratory songbirds typically breed in temperate regions and then travel long distances to spend the majority of the annual cycle in tropical wintering areas. Using stable-isotope methodology, we provide quantitative evidence of dual breeding ranges for 5 species of Neotropical migrants. Each is well known to have a Neotropical winter range and a breeding range in the United States and Canada. However, after their first bout of breeding in the north, many individuals migrate hundreds to thousands of kilometers south in midsummer to breed a second time during the same summer in coastal west Mexico or Baja California Sur. They then migrate further south to their final wintering areas in the Neotropics. Our discovery of dual breeding ranges in Neotropical migrants reveals a hitherto unrealized flexibility in life-history strategies for these species and underscores that demographic models and conservation plans must consider dual breeding for these migrants.

Allometry of the duration of flight feather molt in birds.

We used allometric scaling to explain why the regular replacement of the primary flight feathers requires disproportionately more time for large birds. Primary growth rate scales to mass (M) as M(0.171), whereas the summed length of the primaries scales almost twice as fast (M(0.316)). The ratio of length (mm) to rate (mm/day), which would be the time needed to replace all the primaries one by one, increases as the 0.14 power of mass (M(0.316)/M(0.171) = M(0.145)), illustrating why the time required to replace the primaries is so important to life history evolution in large birds. Smaller birds generally replace all their flight feathers annually, but larger birds that fly while renewing their primaries often extend the primary molt over two or more years. Most flying birds exhibit one of three fundamentally different modes of primary replacement, and the size distributions of birds associated with these replacement modes suggest that birds that replace their primaries in a single wave of molt cannot approach the size of the largest flying birds without first transitioning to a more complex mode of primary replacement. Finally, we propose two models that could account for the 1/6 power allometry between feather growth rate and body mass, both based on a length-to-surface relationship that transforms the linear, cylindrical growing region responsible for producing feather tissue into an essentially two-dimensional structure. These allometric relationships offer a general explanation for flight feather replacement requiring disproportionately more time for large birds.