Investing in a nest egg: intraspecific variation in the timing of egg laying across a latitudinal gradient.

Avian reproductive strategies vary widely, and many studies of life-history variation have focused on the incubation and hatching stages of nesting. Birds make proximate decisions regarding reproductive investment during the laying stage, and these decisions likely constrain and tradeoff with other traits and subsequent behaviors. However, we know relatively little about egg-laying stage behaviors given the difficulty of locating and monitoring nest sites from the onset of laying. We used non-invasive continuous video recording to quantify variation in the egg-laying behaviors of burrowing owls (Athene cunicularia) along a 1400-km latitudinal gradient in western North America. Burrowing owls laid eggs disproportionately in the morning hours, and that tendency was strongest among first eggs in a clutch. However, selection appeared to act more strongly on laying intervals (the time between laying of consecutive eggs) than on the diel time of laying, and laying intervals varied widely among and within clutches. Laying intervals declined seasonally and with increasing clutch size but increased with increasing burrow temperature and as a function of laying stage nest attentiveness, which may be a strategy to preserve egg viability. Laying interval was positively correlated with the duration of hatching intervals, suggesting that laying interval duration is one mechanism (along with timing of incubation onset) that generates variation in hatching asynchrony. Our results lend support to two general hypotheses to explain laying schedules; selection favors laying eggs in the morning, but other selective pressures may override that pattern. These conclusions indicate that allocation decisions during laying are an important part of avian life-history strategies which are subject to energetic constraints and tradeoffs with other traits.

Ophthalmic Parameters and Ophthalmoscopy of Burrowing Owls (Athene cunicularia).

Twelve adult burrowing owls (Athene cunicularia) maintained in a managed environment underwent complete bilateral ophthalmic examinations to assess ocular parameters and, if present, describe lesions (n = 24 eyes). Tear production was measured with a Schirmer tear test (STT), and intraocular pressure (IOP) was measured with rebound tonometry using established calibration settings (D = dog/cat, P = other species). Retinography was performed for all birds after application of topical rocuronium bromide, and corneal diameter was measured. Menace response was absent bilaterally in 7 of 12 (58.3%) owls; however, this did not appear to be related to the presence of fundic lesions. Ocular lesions were visualized in 6 of 12 (50%) owls. The most common ophthalmic abnormality noted was mild multifocal fundic pigment clumping, suggestive of chorioretinal scarring. Other ocular lesions included 1 retinal tear and 1 incipient cataract. Mean tear production was 6.1 ± 3.0 mm/min. Mean IOPs were 11.6 ± 1.8 mm Hg and 7.1 ± 1.3 mm Hg for the D and P settings, respectively, and these were significantly different (P < 0.001). The IOP results did not differ significantly based on patient age or between the right and left eyes, but a higher mean was obtained from males versus females using the D setting (P < 0.039; male mean 12.1 ± 1.9 mm Hg; female mean 10.9 ± 1.2 mm Hg). Measurements obtained from the STT were not affected by either age or sex. Corneal height was 11 mm and width was 12 mm, regardless of age or sex. The rebound tonometer D setting is recommended for measuring IOP values in this species. Burrowing owls had inconsistent mydriasis following topical rocuronium bromide application to the eye; however, a complete fundic examination was possible with or without complete mydriasis.

Drivers of the Ectoparasite Community and Co-Infection Patterns in Rural and Urban Burrowing Owls.

Urbanization creates new ecological conditions that can affect biodiversity at all levels, including the diversity and prevalence of parasites of species that may occupy these environments. However, few studies have compared bird-ectoparasite interactions between urban and rural individuals. Here, we analyze the ectoparasite community and co-infection patterns of urban and rural burrowing owls, Athene cunicularia, to assess the influence of host traits (i.e., sex, age, and weight), and environmental factors (i.e., number of conspecifics per nest, habitat type and aridity) on its composition. Ectoparasites of burrowing owls included two lice, one flea, and one mite. The overall prevalence for mites, lice and fleas was 1.75%, 8.76% and 3.50%, respectively. A clear pattern of co-infection was detected between mites and fleas and, to less extent, between mites and lice. Adult owls harbored fewer ectoparasites than nestlings, and adult females harbored more lice than males. Our results also show that mite and flea numbers were higher when more conspecifics cohabited the same burrow, while lice showed the opposite pattern. Rural individuals showed higher flea parasitism and lower mite parasitism than urban birds. Moreover, mite numbers were negatively correlated with aridity and host weight. Although the ectoparasitic load of burrowing owls appears to be influenced by individual age, sex, number of conspecifics per nest, and habitat characteristics, the pattern of co-infection found among ectoparasites could also be mediated by unexplored factors such as host immune response, which deserves further research.

Revisión del conocimiento actual y conservación de la lechuza de los arenales Athene cunicularia (Molina, 1782) en el Perú / A review of the knowledge and conservation of the Burrowing Owl Athene cunicularia in Peru

Resumen La lechuza de los arenales Athene cunicularia (Aves: Strigidae) se distribuye desde Canadá hasta Tierra del Fuego en América. Ocupa una gran variedad de hábitats naturales, ambientes urbanorurales y agroecosistemas. En el Perú residen tres subespecies A. c. nanodes, (Berlepsch y Stolzmann, 1892), A. c. juninensis (Berlepsch y Stolzmann, 1902) y A. c. cunicularia (Moliln, 1782). El presente trabajo, constituye una revisión de la bibliografía complementada con datos propios, con la finalidad de identificar el estado actual del conocimiento de la biología y estado de conservación de A. cunicularia con especial énfasis en las poblaciones que habitan en el Perú.

Abstract The Burrowing Owl, Athene cunicularia, is distributed from Canada to Tierra del Fuego in America. It occupies a wide variety of natural habitats, urban-rural environments, and agro-ecosystems. Three subspecies reside in Peru: A. c. nanodes (Berlepsch and Stolzmann, 1892), A. c. juninensis (Berlepsch & Stolzmann, 1902) and A. c. cunicularia (Molina, 1782). The present work constitutes a bibliography review, supplemented with our own data, to identify the current state of knowledge about the biology and conservation status of A. cunicularia; with special emphasis on the populations that inhabit Peru.

Agriculture creates subtle genetic structure among migratory and nonmigratory populations of burrowing owls throughout North America.

Population structure across a species distribution primarily reflects historical, ecological, and evolutionary processes. However, large-scale contemporaneous changes in land use have the potential to create changes in habitat quality and thereby cause changes in gene flow, population structure, and distributions. As such, land-use changes in one portion of a species range may explain declines in other portions of their range. For example, many burrowing owl populations have declined or become extirpated near the northern edge of the species' breeding distribution during the second half of the 20th century. In the same period, large extensions of thornscrub were converted to irrigated agriculture in northwestern Mexico. These irrigated areas may now support the highest densities of burrowing owls in North America. We tested the hypothesis that burrowing owls that colonized this recently created owl habitat in northwestern Mexico originated from declining migratory populations from the northern portion of the species' range (migration-driven breeding dispersal whereby long-distance migrants from Canada and the United States became year-round residents in the newly created irrigated agriculture areas in Mexico). We used 10 novel microsatellite markers to genotype 1,560 owls from 36 study locations in Canada, Mexico, and the United States. We found that burrowing owl populations are practically panmictic throughout the entire North American breeding range. However, an analysis of molecular variance provided some evidence that burrowing owl populations in northwestern Mexico and Canada together are more genetically differentiated from the rest of the populations in the breeding range, lending some support to our migration-driven breeding dispersal hypothesis. We found evidence of subtle genetic differentiation associated with irrigated agricultural areas in southern Sonora and Sinaloa in northwestern Mexico. Our results suggest that land use can produce location-specific population dynamics leading to subtle genetic structure even in the absence of dispersal barriers.

Burrowing owls eavesdrop on southern lapwings' alarm calls to enhance their antipredatory behaviour.

Eavesdropping is a widespread behaviour among animals, providing the receiver with valuable information to assess the habitat, resources or threats. This kind of behaviour has been reported for the burrowing owl (Athene cunicularia), which in its northern range lives in close association with fossorial mammals and eavesdrops on their alarm calls as indicators of risk. In their southernmost range, burrowing owls do not associate with mammals, but they are often found sharing foraging and nesting patches with the southern lapwing (Vanellus chilensis), a noisy, territorial and aggressive plover species. We designed a field experimental study aimed at determining if burrowing owls are able to use lapwing calls as indicator of potential risk. We exposed focal owls to a sequence of sounds including lapwing alarm calls, and biological and non-biological controls, and registered their response as alert or relax behaviours. Linear mixed modeling showed that owls increased their alert behaviour in response to lapwing alarm calls but not in response to control treatments. In addition, owls' response was consistent between habitats (rural and urban) and seasons (breeding and non-breeding). Our results suggest that eavesdropping is a generalized strategy of burrowing owls to acquire environmental information throughout its distribution range.

Ecología trófica del búho terrestre Athene cunicularia punensis (Strigiformes: Strigidae) en el archipiélago de Jambelí, provincia de El Oro, suroeste de Ecuador / Feeding ecology of the Burrowing Owl Athene cunicularia punensis (Strigiformes: Strigidae) in the Jambelí archipelago, El Oro province, southwestern Ecuador

Estudiamos la dieta del búho terrestre Athene cunicularia punensis en cuatro territorios dentro del archipiélago de Jambelí, provincia de El Oro, suroeste de Ecuador, entre noviembre de 2015 y abril de 2016, mediante la recolección y análisis de 182 egagrópilas. Los artrópodos constituyeron el principal grupo-presa en frecuencia de aparición (86.5%), seguido por los mamíferos (12%) y las aves (1.5%). En cuanto a biomasa consumida, los mamíferos (Mus musculus y Rattus rattus) aportaron 58.5%, los artrópodos 33.2% y las aves 8.2%. No encontramos diferencias significativas de la dieta entre las cuatro parejas estudiadas. En este trabajo se confirmó el rol de Athene cunicularia como un importante depredador de plagas como roedores introducidos y artrópodos potencialmente dañinos.

The diet of Burrowing Owl (Athene cunicularia punensis) was studied in four territories in the Jambelí archipelago, El Oro, southwestern Ecuador, from November 2015 to April 2016, collecting 182 pellets. Arthropods were the main prey group in frequency of occurrence (86.5%), followed by mammals (12%) and birds (1.5%), but mammals (Mus musculus and Rattus rattus) contributed 58.5% of the biomass, arthropods 33.2% and birds 8.2%. No significant differences were found in the diet among the four pairs studied. Our study confirmed the important role of Athene cunicularia as predator of pests like introduced rodents and potentially harmful arthropods.

LOW PREVALENCE OF WEST NILE VIRUS ANTIBODIES IN SELECT NORTHERN CALIFORNIA OWL SPECIES (2007-2014).

The objective of this study was to determine evidence of previous West Nile virus (WNV) infection in northern California owls. Owl serum samples were collected from birds presenting to a veterinary medical teaching hospital between 2007 and 2014 and were screened for the presence of WNV antibodies by an indirect enzyme immunoassay (EIA). Only one of 71 samples (1.41%) tested was positive by EIA and confirmed by a plaque reduction neutralization test; it was the most recent sample collected. The reason for the low prevalence of WNV in these California owls despite a high prevalence in sympatric avian species in the same region is unknown and should be a topic for further research.

Differential risk perception of rural and urban Burrowing Owls exposed to humans and dogs.

Urban areas expose wildlife to an array of novel predators, amongst which, humans and dogs are highly frequent. Thus, wild animals living in urban areas are forced to invest more time and energy in defence behaviours, which depend on how the risk is perceived and assessed. We experimentally tested whether Burrowing owls coming from rural and urban habitats showed differences in behavioural responses when facing humans and domestic dogs. We measured flight initiation distances (FIDs), nest returning, and aggressiveness level when owls faced a human and a human with a dog walking towards them. Our results showed that urban owls recognise a human with a dog as a greater threat than a human alone, thus indicating that fear of domestic animals should be considered as affecting owls' settlement in cities and towns. On the other hand, rural owls perceived human and dogs as similar threats, but showed higher FIDs, less aggressiveness, and lower tendency to return to the nest than urban owls in both treatments. These findings emphasize the importance of modified habitats in modelling the response of urban and rural owls to predators and represent another step in the explanation of how wild animals assess and respond to threats associated with living in urbanized environments.

Rapid warming and drought negatively impact population size and reproductive dynamics of an avian predator in the arid southwest.

Avian communities of arid ecosystems may be particularly vulnerable to global climate change due to the magnitude of projected change for desert regions and the inherent challenges for species residing in resource limited ecosystems. How arid-zone birds will be affected by rapid increases in air temperature and increased drought frequency and severity is poorly understood because avian responses to climate change have primarily been studied in the relatively mesic northern temperate regions. We studied the effects of increasing air temperature and aridity on a Burrowing Owl (Athene cunicularia) population in the southwestern United States from 1998 to 2013. Over 16 years, the breeding population declined 98.1%, from 52 pairs to 1 pair, and nest success and fledgling output also declined significantly. These trends were strongly associated with the combined effects of decreased precipitation and increased air temperature. Arrival on the breeding grounds, pair formation, nest initiation, and hatch dates all showed significant delays ranging from 9.4 to 25.1 days over 9 years, which have negative effects on reproduction. Adult and juvenile body mass decreased significantly over time, with a loss of 7.9% mass in adult males and 10.9% mass in adult females over 16 years, and a loss of 20.0% mass in nestlings over 8 years. Taken together, these population and reproductive trends have serious implications for local population persistence. The southwestern United States has been identified as a climate change hotspot, with projections of warmer temperatures, less winter precipitation, and an increase in frequency and severity of extreme events including drought and heat waves. An increasingly warm and dry climate may contribute to this species' decline and may already be a driving force of their apparent decline in the desert southwest.

Detecting Burrowing Owl Bloodmeals in Pulex irritans (Siphonaptera: Pulicidae).

Pulex irritans L. is a cosmopolitan flea species that infests a wide variety of hosts. In North America it generally parasitizes large wild mammals, but in the Pacific Northwest an association has emerged between P. irritans and the western burrowing owl (Athene cunicularia hypugaea). While investigators have recognized this association for decades, it has not been clear if P. irritans feeds on burrowing owls, or if the owls serve exclusively as phoretic hosts. Here we describe using a real-time assay that was originally developed to identify bloodmeals in Ugandan cat fleas (Ctenocephalides felis Bouché) to detect burrowing owl DNA in P. irritans collected from burrowing owls in southern Idaho. Of 50 fleas tested, 12 had no detectable vertebrate bloodmeal. The remaining 38 (76%) contained burrowing owl DNA. The assay did not detect vertebrate DNA in unfed fleas exposed to owl or mouse pelts and is therefore unlikely to detect DNA in fleas from vertebrates that have served exclusively as phoretic hosts. We conclude that P. irritans feeds on burrowing owls. We discuss the potential implications of this finding for burrowing owl conservation and enzootic plague dynamics.

Burrowing Owls, Pulex irritans, and Plague.

Western Burrowing Owls (Athene cunicularia hypugaea) are small, ground-dwelling owls of western North America that frequent prairie dog (Cynomys spp.) towns and other grasslands. Because they rely on rodent prey and occupy burrows once or concurrently inhabited by fossorial mammals, the owls often harbor fleas. We examined the potential role of fleas found on burrowing owls in plague dynamics by evaluating prevalence of Yersinia pestis in fleas collected from burrowing owls and in owl blood. During 2012-2013, fleas and blood were collected from burrowing owls in portions of five states with endemic plague-Idaho, Oregon, Washington, Colorado, and South Dakota. Fleas were enumerated, taxonomically identified, pooled by nest, and assayed for Y. pestis using culturing and molecular (PCR) approaches. Owl blood underwent serological analysis for plague antibodies and nested PCR for detection of Y. pestis. Of more than 4750 fleas collected from owls, Pulex irritans, a known plague vector in portions of its range, comprised more than 99.4%. However, diagnostic tests for Y. pestis of flea pools (culturing and PCR) and owl blood (PCR and serology) were negative. Thus, even though fleas were prevalent on burrowing owls and the potential for a relationship with burrowing owls as a phoretic host of infected fleas exists, we found no evidence of Y. pestis in sampled fleas or in owls that harbored them. We suggest that studies similar to those reported here during plague epizootics will be especially useful for confirming these results.

New record of Pelecitus sp. (Nematoda, Onchocercidae) as a parasite of Athene cunicularia (Strigiformes, Strigidae) in southeastern Brazil / Novo registro de Pelecitus sp. (Nematoda, Onchocercidae) como parasita de Athene cunicularia (Strigiformes, Strigidae) no Sudeste do Brasil

The aim of this study was to report the burrowing owl Athene cunicularia as a new host for the filarid nematode Pelecitus sp. in southeastern Brazil for the first time, as well as reporting the occurrence of this nematode species in the body cavity, near the cervical air sac and lung region. This study contributes towards knowledge of parasitism in Brazilian wild birds and an anatomical region of the host as an infection site for Pelecitus sp.

O objetivo do estudo foi relatar, pela primeira vez, a coruja buraqueira Athene cunicularia como um novo hospedeiro para o nematóide filarídeo Pelecitus sp. do Sudeste do Brasil, bem como a ocorrência desta espécie de nematóides na cavidade corporal, próximos à região dos sacos aéreos cervicais e pulmões. Este estudo contribui para o conhecimento do parasitismo em aves silvestres brasileiras e uma região anatômica do hospedeiro como sítio de infecção para Pelecitus sp.