Long-Term Study Reveals Fluctuations in Birds’ Nesting Success

AUK-17-189 D Janus

Long-term data on Song Sparrows reveals factors that affect their nesting success. Photo credit: D. Janus

Understanding the factors that affect a bird species’ nesting success can be crucial for planning effective conservation efforts. However, many studies of nesting birds last only a few years—and that means they can miss the effects of long-term variation and rare events. A new study from The Auk: Ornithological Advances demonstrates this with nearly four decades of data from Song Sparrows in British Columbia.

The University of British Columbia’s Merle Crombie and Peter Arcese used 39 years of data from an island population of Song Sparrows to examine how the factors influencing their nesting success changed over long periods of time. Over almost 3,000 nesting attempts, 64% of which were successful, a number of patterns emerged. Some, such as the fact that older female birds were less successful, remained consistent over time. However, others, such as the effects of rainfall, population density, and nest parasitism, interacted with each other in complex ways that caused their importance to wax and wane over the decades, and inbreeding only became a significant negative factor when it increased sharply during the middle portion of the study. Unpredictable, rare fluctuations such as this can have large effects that shorter-term studies rarely capture.

“Researchers have been learning about the Song Sparrow population on Mandarte Island since 1960, and monitoring the population continuously since 1975,” says Arcese. “Because the population is semi-isolated, small, and resident year-round, we band all birds in the nest and have genotyped all nestlings since 1991.” A close focus on individuals, fitness, and relatedness in the Mandarte Song Sparrow population has allowed researchers to report the most precise demographic and population genetic parameters yet estimated in wild populations.

“Most studies of plant and animal populations in nature last three to five years, but ecological processes are often dramatically affected by climate and community change, which plays out over decades,” he continues. “Long-term studies like ours provide an invaluable record of change in population processes, which can help interpret the results of short-term studies of species not as easily studied as Song Sparrows.”

Temporal variation in the effects of individual and environmental factors on nest success is available at http://www.bioone.org/doi/full/10.1642/AUK-17-189.1.

About the journal: The Auk: Ornithological Advances is a peer-reviewed, international journal of ornithology that began in 1884 as the official publication of the American Ornithologists’ Union, which merged with the Cooper Ornithological Society in 2016 to become the American Ornithological Society. In 2009, The Auk was honored as one of the 100 most influential journals of biology and medicine over the past 100 years.

AUTHOR BLOG: Are All Eggs Created Equal? Saltmarsh Sparrows Support Gender Equality

Bri Benvenuti and Adrienne Kovach

Linked paper: Annual variation in the offspring sex ratio of Saltmarsh Sparrows supports Fisher’s hypothesis by B. Benvenuti, J. Walsh, K.M. O’Brien, M.J. Ducey, and A.I. Kovach, The Auk: Ornithological Advances 135:2, April 2018.

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Three Saltmarsh Sparrow chicks banded as part of a study on offspring sex ratios. Photo: B. Benvenuti

In birds, females have the ability to control the sex of individual eggs; therefore, a mother may be able to choose whether she prefers each egg laid to be a male or female. This means that offspring sex ratios are not usually left to chance. From an evolutionary standpoint, this can be very beneficial, as different circumstances may favor the success of sons versus daughters.

But how does one know if it would be better to have more sons or daughters? Evolutionary theory suggests that if the potential benefits of raising one sex over the other vary in relation to environmental or maternal conditions, then females should favor the production of that sex. Typically, high quality sons are more beneficial to mothers, because they have the potential to produce far more grandchildren than daughters can (males can mate many times, but females are limited by how many eggs they can produce, incubate, and raise to fledging). More grandchildren = greater lifelong success.  Still, there is a risk to biasing offspring production toward sons; if the son is low quality (competitively inferior), he may not reproduce at all.  On the other hand, daughters tend to be “cheaper” to raise than sons. They need less resources to reach maturity, and if they survive, they almost always reproduce. With this in mind, one could logically say that producing daughters represents the “safe bet”—you might get a smaller payout (in terms of numbers of grandchildren), but you know you’ll get something.

Armed with this information, we chose to investigate whether Saltmarsh Sparrows manipulate the sex of their offspring based on environmental or maternal conditions as we would expect based on evolutionary theory. These tidal marsh specialist birds live a stressful life—they build nests in the marsh grasses just inches above the marsh surface that is regularly subjected to tidal flooding. Nests are more likely to escape these flooding events and successfully fledge offspring if they are timed to fledge within the 28 day lunar tidal cycle. Saltmarsh Sparrows are also one of the world’s most promiscuous birds, with almost every egg in a nest having a different father thanks to the scramble of competition among males for access to females. These characteristics provide interesting hypotheses in the context of evolutionary theory, so we asked, would Saltmarsh Sparrow mothers produce more sons, who would be larger and a) more likely to survive a nest flooding event and b) have the ability to produce more offspring through multiple matings? Or would they take the “safe bet” and produce more females?

To test our hypotheses, we collected nesting data from Saltmarsh Sparrow breeding locations in New England marshes over five years. We used DNA analysis to determine the sex of Saltmarsh Sparrow chicks and calculated the offspring sex ratio for our four study sites and across the whole study population. We then used a modeling approach to determine if there was an influence of environmental conditions (year, tidal flooding, precipitation), temporal effects (nest initiation in relation to flood tides, timing within the breeding season), or maternal condition on offspring sex ratios.

Surprisingly, we found an even offspring sex ratio of 1.03:1 (males to females) across all years and sites, and offspring sex ratios did not vary as a function of the environment, tidal flooding risk, or female condition. What we did find was an interesting pattern of annual variation between male and female bias that mirrored the adult sex ratio in the preceding year.

While numerous studies have provided evidence that female birds may have the ability to adjust offspring sex ratios in an adaptive way, we found no evidence for adaptive sex ratio manipulation in Saltmarsh Sparrows in relation to our hypotheses. Instead, the observed time-lagged relationship between offspring and adult sex ratio meets expectations of frequency-dependent selection, whereby females respond to higher frequencies of one sex by increasing production of the rarer sex, which would have a temporary fitness advantage. Our findings overall show support for balanced offspring sex ratios at a population level over time.

Scientists Remind Their Peers: Female Birds Sing, Too


Northern Cardinals are among the familiar North American bird species in which females sing in addition to males.

When North American ornithologists hear a bird singing, they’re likely to assume it’s a male. But in many species, the females sing too—and a new commentary in The Auk: Ornithological Advances argues that a better understanding of these unappreciated female songs could lead to advances in many aspects of bird biology.

Authors Karan Odom of Cornell University and Lauryn Benedict of the University of Northern Colorado both discovered the world of female birdsong through their own research. “I started studying California towhees 17 years ago, and I was fascinated by the duet vocalization given by females and males,” says Benedict. “That led me to start looking for female song in other North American bird species, and I was surprised to learn that it was much more common than I expected. The reports of female song are buried in odd corners of the literature, but when you put them all together, you start to see some interesting patterns.”

Though singing females were likely the norm among the ancestors of today’s songbirds, female song today is understudied and is underrepresented in collections of bird sound recordings. This, say Odom and Benedict, may be result of bias toward the world’s temperate regions—though more widespread in temperate species than many ornithologists appreciate, female song is most common among tropical birds. They argue that better documentation of which species female song is present in and more detailed descriptions of female song structure and output could improve our understanding of birds’ comparative physiology, neurobiology, behavioral ecology, evolution, and even conservation. Birds of conservation concern are often located and identified by song during surveys, and assumptions that all singing birds are male could mislead wildlife managers about the state of populations.

Odom and Benedict urge their fellow ornithologists to spread the word that female birds sing, to share resources, and to disseminate their findings. You don’t need to be a professional ornithologist in order to help expand our knowledge of female song, either—Odom has created a website where any birdwatcher can upload their observations. “If you hear a bird singing, do not assume it’s a male,” she says. “If you observe a female bird singing, document it by uploading field notes, audio, or video to the collections on our website, femalebirdsong.org. Make sure to indicate how you recognized the bird was female.”

“Odom and Benedict have written an excellent appeal to document and record more female bird song,” adds Leiden University’s Katharina Riebel, a former collaborator of Odom’s. “They rightly point out that the extent of female bird song has been starkly underestimated, as almost by default we assume that a singing bird must be the male of the species. As a consequence, we might have missed out many aspects and the dynamics of male and female vocal signaling in songbirds—clearly, there is still lots to discover! I am confident that ornithologists in the field can make substantial contributions toward these questions by sharing their observations and recordings, as I very much hope this article will encourage them to do.”

A call to document female bird songs: Applications for diverse fields is available at http://www.bioone.org/doi/full/10.1642/AUK-17-183.1.

About the journal: The Auk: Ornithological Advances is a peer-reviewed, international journal of ornithology that began in 1884 as the official publication of the American Ornithologists’ Union, which merged with the Cooper Ornithological Society in 2016 to become the American Ornithological Society. In 2009, The Auk was honored as one of the 100 most influential journals of biology and medicine over the past 100 years.

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AUTHOR BLOG: Recognizing the Importance of Female Birdsong

Karan Odom & Lauryn Benedict

Linked paper: A call to document female bird songs: Applications for diverse fields by K.J. Odom and L. Benedict, The Auk: Ornithological Advances 135:2, April 2018.


House Wrens are among the familiar birds in which females sing. Photo credit: J. Hudgins/USFWS

Can you name ten North American or European bird species in which females sing? Can you name twenty? Fifty? That may seem like a lot, but in fact it’s only a small percentage of temperate-zone bird species with female song. There are at least 144 North American passerine species with female song, and many more non-passerines with elaborate vocalizations that could be classified as song (defining “song” is a topic we won’t even go into here!). Across all avian species, approximately 64% have female song, but these estimates are rough. The true numbers could be much higher. Why are the estimates so rough? Because documentation and reports of female song are lacking. We highlight this problem in our paper “A call to document female bird songs: Applications for diverse fields.” We ask all of you to help us address the deficit.

We know that the data are out there; we regularly have conversations with ornithologists and citizen scientists who tell us that they have observed singing females in myriad species. Our response: Publish it! Archive it! We’ve chatted with many senior researchers who have years of data including observations and recordings of female song in their study populations, but who haven’t published these data because female song is rare or is not their main research focus. We’ve also heard from students working as field assistants whose cohorts regularly observe singing females, but those observations are seldom documented. On field projects with many technicians, word-of-mouth data can be extensive and highly informative, but staff turnover means that known population traits never get put down on paper (or audio).

Citizen scientists frequently tell us about singing females, and many of them have taken the next step to document their observations through The Female Bird Song Project. Contributors have recorded female song in species as diverse as the Mexican Sheartail (Doricha eliza), Black-goggled Tanager (Trichothraupis melanops), Saffron Finch (Sicalis flaveola), and Cerulean Warbler (Setophaga cerulea), all of which seem to be the first documentation of female song in their species!

Each of these contributions plays a role in understanding the distribution of species with female song – data that researchers can use to address a wide range of biological questions. A complete picture of when and how female birds sing will offer insights into the biological mechanisms, evolution, and applications of avian vocal signals. Neurobiologists can ask how bird brains perceive and produce these variable signals, and whether that differs by sex. Evolutionary ecologists can ask why songs differ among species with different ecology and life-history traits. Conservation biologists can use songs to census and monitor the presence of males and females across populations.

What can you do? Don’t assume that a singing bird is a male. Look, listen, and document without bias. Teach your students to do the same. In 73% of all bird species we lack enough published information to even determine whether females sing. We are confident, however, that in some of those species females do sing because we have talked to colleagues about them. Common knowledge suggests that female song is rare, but our experiences make us question that: if all ornithologists talked to each other about female song the way that they talk to us, then that assumption would change. Your random observations of a female warbler singing can probably be backed up by the observations of many others. Females of temperate-breeding species may not sing as often as males, but when we pool all our knowledge and observations it’s likely that we’ll find more parity than we expect.

AUTHOR BLOG: Latitudinal Gradient in Bird Survival

Gonçalo Ferraz

Linked paper: Age effects on survival of Amazon forest birds and the latitudinal gradient in bird survival by A.P. Muñoz, M. Kéry, P.V. Martins, and G. Ferraz, The Auk: Ornithological Advances 135:2, April 2018.


A Guianan Warbling Antbird, Hypocnemis cantator, banded and ready for release. Photo credit: E. Johnson

Bold, recurring patterns are fascinating, and more so if they are unexplained. For biologists living in the tropics, perhaps the most striking examples are the various latitudinal gradients in community and life history metrics. Science excites our curiosity in the search for universal rules, but large-scale latitudinal patterns hint that some things may be fundamentally different in different parts of the world. Could there be really such a thing as “tropical biology”?

One of the best-documented of these latitudinal gradients is the tendency for avian clutch size to increase with increasing latitude. Since there is no evidence that high-latitude populations grow faster than equatorial ones, it’s been widely assumed that some other life history trait must also change with latitude to compensate for the variation in clutch size. Thus, throughout the second half of the 20th century, the idea that adult tropical birds live longer than their temperate counterparts became widely accepted among ornithologists.

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A Rufous-throated Antbird, Gymnopithys rufigula. Photo credit: E. Johnson

The acceptance of a latitudinal gradient in survival, however, was more based on common sense than hard data—so much so that when Karr et al. (1990, The American Naturalist 136: 277-291) challenged the idea in a paper subtitled “Will the dogma survive?” there were not many voices sounding in defense of the “dogma.” Indeed, two decades later, Corey Tarwater and colleagues suggested that, if there is no latitudinal trend in the survival of adult birds, there could be a trend in the survival of juveniles (2011, Ecology 92: 1271-1281). Their work drew on detailed analysis of age-dependent survival in one Central American passerine species and proposed that tropical juveniles survive more than temperate juveniles. If adult survival shows no relationship with latitude this implies relatively little difference between adult and juvenile survival in the tropics.

For a paper just published in The Auk, Alejandra P. Muñoz, myself, and our colleagues used bird banding data to tackle the problem of the latitudinal gradient in bird survival on two fronts. First, we quantified the effect of age on survival for forty species of Amazon forest passerines, showing that adults at our tropical site had substantially higher survival probabilities than juveniles. Second, we compared our adult survival estimates with estimates from 175 species from Peru to Alaska and found that survival does decrease with increasing latitude. This latitudinal effect persists even after accounting for effects of migration mode, phylogeny, and time of data collection, and we conclude that the latitudinal gradient in survival is a fact after all, at least as seen among New World forest passerines.

Our work benefited a great deal from two recent advances. First, there is an ongoing transformation in how tropical ornithologists assess bird age. The Wolfe-Ryder-Pyle molt-cycle system, which we employed, is extremely useful for aging birds in populations that have poorly delimited breeding periods, which is typical of tropical regions. This was central for quantifying the effect of age on survival. Second, we tapped into the phenomenal database of Vital Rates of North American Birds made available by the Institute of Bird Populations (IBP), enabling us to include data from as far north as Alaska. The IBP estimates, combined with a wealth of tropical passerine survival estimates published since 1990, made it possible to take up Karr et al.’s (1990) challenge anew.

From the analytical perspective, we used a multi-species Cormack-Jolly-Seber (CJS) approach, a standard method for estimating apparent survival from capture-recapture data on open populations, to model our Amazon data. We treated each of the forty species in our dataset as a random draw from a wider distribution of species, and the inference about age effects was made at the level of this wider distribution, strengthening the generality of the conclusions. The CJS accounts for the possibility of capture failure, and a “mixture” component of the model, developed by co-author Marc Kéry, makes it possible to incorporate birds of unknown age in the analysis. Our work thus addresses a variety of sources of uncertainty before reaching its final conclusions.

The observation of a latitudinal trend in survival probability helps explain the maintenance of a latitudinal trend in clutch size, but it need not be the only explanation. It’s also possible, for example, that the number of clutches laid each year also varies with latitude in association with the length of the breeding season. With or without latitudinal change in number of clutches, one cannot tell whether the variation in survival we documented is a sufficient explanation for the current variation in clutch size. What’s more, even if one explains the maintenance of current variation, there is still the question of how that variation evolved. There is certainly still much to explore in the latitudinal variation of bird life history traits.

Long Incubation Times May Defend Birds Against Parasites

Some tropical birds have longer egg incubation times than their temperate cousins, even though their habitat is teeming with egg-eating predators. The reason why has long been a mystery, but a new study from The Auk: Ornithological Advances applies new methods to confirm the evidence for an old hypothesis—that a longer development period leads to a stronger, more efficient immune system.

The University of Missouri-St. Louis’s Robert Ricklefs first studied this relationship in the early 1990s, using data from microscopic examination of avian blood samples for the presence of parasites, primarily those that cause malaria. He found that the longer a species’ incubation period, the lower its prevalence of parasites. However, Ricklefs remained concerned that especially low parasite loads could have been missed during microscope examination, affecting parasite prevalence estimates.

Advances in DNA sequencing offered a new solution. For their new study, Ricklefs and his colleagues collected blood samples from birds in the eastern U.S. and several Neotropical countries and checked for the presence of parasite DNA, tabulating how many individuals from various families of birds were sampled at each site and how many were infected with Haemoproteus or Plasmodium parasites. About 22% of individual birds in both temperate and tropical regions had parasite DNA in their blood. While incubation time varies little among temperate species, it does vary among tropical species, and in tropical birds parasite prevalence was significantly lower in species with longer incubation times.

These results confirm those of the old blood smear analysis. While there is still no direct evidence for the hypothesis that a longer incubation time promotes a stronger immune system, this correlation provides a strong hint that this could indeed be the solution to the mystery of why the embryos of some tropical birds take so long to develop. “My interest in blood parasites was stimulated by a former graduate student, Victor Apanius, primarily in the context of community ecology. However, I had been working on the diversification of avian life histories, particularly embryo and chick growth rates, and I couldn’t help but notice a connection between the two,” says Ricklefs. “I wasn’t surprised that the new results confirm the old ones so well, really, because the two techniques estimate the same attribute. However, more detailed studies of the avian immune response and how variation in host defense is related to development certainly are warranted.”

“This paper is a nice follow up the 1992 study that showed an inverse relationship between parasite prevalence and egg incubation period. Since that time, PCR methods have been developed that detect more infections than microscopy, and the work is important because it verifies the previous results with revised methods,” according to San Francisco State University’s Ravinder Sehgal, an expert on avian blood parasites who was not involved in the study. “Moreover, it renews interest in a phenomenon that has gone largely unexplored. It will be now be important to test the work in an experimental system, to study the parasitology and explore the tradeoffs between embryo growth rate and immune function.”

Duration of embryo development and the prevalence of haematozoan blood parasites in birds is available at http://www.bioone.org/doi/full/10.1642/AUK-17-123.1.

About the journal: The Auk: Ornithological Advances is a peer-reviewed, international journal of ornithology that began in 1884 as the official publication of the American Ornithologists’ Union, which merged with the Cooper Ornithological Society in 2016 to become the American Ornithological Society. In 2009, The Auk was honored as one of the 100 most influential journals of biology and medicine over the past 100 years.

Are Flamingos Returning to Florida?

CONDOR-17-187 J Patterson

American Flamingos should be considered native to Florida, argue the authors of a new review. Photo credit: J. Patterson

Flamingos are a Florida cultural icon, and sightings of American Flamingos in the state have been on the rise in recent decades. However, whether they’re truly native to the U.S. or only arrive via escape from captivity has long been subject to debate, making developing a plan for managing Florida’s flamingo population challenging. A new study from The Condor: Ornithological Applications reviews the evidence and provides a fresh argument that the birds should be considered part of the Sunshine State’s native fauna.

Zoo Miami’s Steven Whitfield, along with colleagues from Audubon Florida’s Everglades Science Center, the National Park Service, Big Cypress National Preserve, and the Rookery Bay Estuarine Research Reserve, reexamined the historical evidence of flamingos in Florida and evaluated the likely origins of birds seen in recent years. Overall, they conclude, the evidence from both narrative accounts and museum records suggests that American Flamingos once occurred naturally in large flocks in Florida and probably even nested there before being eliminated by hunting around 1900. From 1950 to the present, however, birdwatchers have reported almost 500 new observations of flamingos in Florida, with both flock size and the frequency of observations increasing over time. While it’s plausible that some of these individuals could be escapees from captive flocks, there is also strong evidence for dispersal from wild populations in Mexico and the Caribbean.

The population history Whitfield and his coauthors describe is consistent with that of some native species already protected by state and federal endangered species laws, and they hope that their study will lead to a better plan for managing wild flamingos in Florida. “Living in Florida, you see flamingos everywhere—in advertising, in place names, even on the logo for the state lottery—but as an actual organism, as a species, there was essentially no information available on the biology of flamingos,” says Whitfield. “Some biologists considered them native birds that were extirpated during the plume trade of the late 1800s, and urged for population recovery measures, while others considered the rare flamingos seen around Florida to be escapes from captive colonies. We often say that in south Florida we have just two types of species, introduced and endangered, but a species can’t be both at the same time.”

“This article finally sheds welcome light on status of these iconic birds in Florida. The authors meticulously researched historic records and compiled more recent sightings to reconstruct the history and population trends of flamingos in Florida,” adds the American Museum of Natural History’s Felicity Arengo, a flamingo conservation expert who was not involved in the study. “Flamingo numbers have increased notably since the 1950s due to protections to species and habitats in Florida and throughout the Caribbean. The authors are cautious and recognize the limitations of the data in their study, but they provide ample evidence that Florida was the northernmost extent of the American Flamingo prior to the early 1900s and that populations have been recovering.”

Status and trends of American Flamingos (Phoenicopterus ruber) in Florida, USA is available at http://www.bioone.org/doi/full/10.1650/CONDOR-17-187.1.

About the journal: The Condor: Ornithological Applications is a peer-reviewed, international journal of ornithology. It began in 1899 as the journal of the Cooper Ornithological Club, a group of ornithologists in California that became the Cooper Ornithological Society, which merged with the American Ornithologists’ Union in 2016 to become the American Ornithological Society. In 2016, The Condor had the number one impact factor among 24 ornithology journals.

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