Paul M. Nolan

Sex, size, and plumage redness predict house finch survival in an epidemic

Mycoplasma gallisepticum is a well–known disease of poultry but until 1994 had not been observed in passerine birds. From 1994 to 1996, tens of millions of house finches (Carpodacus mexicanus) are believed to have died in an epidemic of mycoplasmal conjunctivitis, similar to ‘pinkeye’ in humans. The outbreak of Mycoplasma gallisepticum affected finches of both sexes but disproportionately killed males, shifting the sex ratio from male–biased to female–biased. This differential male mortality is consistent with a cost of testosterone, which is a key prediction of the immunocompetence handicap hypothesis. Males and females that survived the epidemic weighed significantly less and had significantly shorter wing chords, tarsi, and bills than did individuals before the epidemic. Male survivors also had significantly redder plumage than males that did not survive, supporting the idea that plumage brightness serves as an indicator of condition, as proposed by the honest advertisement model of sexual selection.

YOU CAN'T JUDGE A PIGMENT BY ITS COLOR: CAROTENOID AND MELANIN CONTENT OF YELLOW AND BROWN FEATHERS IN SWALLOWS, BLUEBIRDS, PENGUINS, AND DOMESTIC CHICKENS

Abstract The two main pigment types in bird feathers are the red, orange, and yellow carotenoids and the black, gray, and brown melanins. Reports conflict, however, regarding the potential for melanins to produce yellow colors or for carotenoids to produce brown plumages. We used high-performance liquid chromatography to analyze carotenoids and melanins present in the yellow and brown feathers of five avian species: Eastern Bluebirds (Sialia sialis), Barn Swallows (Hirundo rustica), King Penguins (Aptenodytes patagonicus), Macaroni Penguins (Eudyptes chrysolophus), and neonatal chickens (Gallus domesticus). In none of these species did we detect carotenoid pigments in feathers. Although carotenoids are reportedly contained in the ventral plumage of European Barn Swallows (Hirundo rustica rustica), we instead found high concentrations of both eumelanins and phaeomelanins in North American Barn Swallows (H. r. erythrogaster). We believe we have detected a new form of plumage pigment that gives penguin and domestic- chick feathers their yellow appearance. No Puedes Juzgar un Pigmento por su Color: Contenido de Carotenoide y Melanina de Plumas Amarillas y Marrones en Golondrinas, Azulejos, Pingüinos y Gallinas Domésticas Resumen. Los dos tipos principales de pigmentos que las aves incorporan en sus plumas son carotenoides, para desarrollar plumajes rojo, naranja o amarillo, y melaninas, para adquirir coloración negra, marrón, gris o tonalidades color tierra. Sin embargo, existe información conflictiva sobre la potencial coloración de plumas amarillas basadas en melanina y la presencia de caroteniodes en el plumaje marrón de ciertas especies. En este estudio, usamos cromatografía líquida de alto rendimiento para analizar los tipos y cantidades de carotenoides y melaninas presentes en las plumas amarillas y marrones de cinco especies de aves: el azulejo Sialia sialis y la golondrina Hirundo rustica, los pingüinos Aptenodytes patagonicus y Eudyptes chrysolophus y el plumón natal amarillo de la gallina doméstica Gallus domesticus. En ninguna de estas especies detectamos pigmentos carotenoides en las plumas. A pesar de que los carotenoides han sido encontrados en el plumaje ventral de la golondrina Hirundo rustica rustica, nosotros en cambio encontramos altas concentraciones de eumelaninas y feomelaninas en H. r. erythrogaster y en azulejos que variaron entre individuos y regiones de plumaje. Creemos que hemos detectado una nueva forma de pigmento de plumaje que le da a las plumas de pingüinos y pollos domésticos su apariencia amarilla.

THE EVOLUTION OF SEXUAL SIZE DIMORPHISM IN THE HOUSE FINCH. II. POPULATION DIVERGENCE IN RELATION TO LOCAL SELECTION

Recent colonization of ecologically distinct areas in North America by the house finch (Carpodacus mexicanus) was accompanied by strong population divergence in sexual size dimorphism. Here we examined whether this divergence was produced by population differences in local selection pressures acting on each sex. In a long‐term study of recently established populations in Alabama, Michigan, and Montana, we examined three selection episodes for each sex: selection for pairing success, overwinter survival, and within‐season fecundity. Populations varied in intensity of these selection episodes, the contribution of each episode to the net selection, and in the targets of selection. Direction and intensity of selection strongly differed between sexes, and different selection episodes often favored opposite changes in morphological traits. In each population, current net selection for sexual dimorphism was highly concordant with observed sexual dimorphism—in each population, selection for dimorphism was the strongest on the most dimorphic traits. Strong directional selection on sexually dimorphic traits, and similar intensities of selection in both sexes, suggest that in each of the recently established populations, both males and females are far from their local fitness optimum, and that sexual dimorphism has arisen from adaptive responses in both sexes. Population differences in patterns of selection on dimorphism, combined with both low levels of ontogenetic integration in heritable sexually dimorphic traits and sexual dimorphism in growth patterns, may account for the close correspondence between dimorphism in selection and observed dimorphism in morphology across house finch populations.

Female choice for song characteristics in the house finch

Song has been proposed to function in mate choice, and such a role has been demonstrated experimentally in a number of species. We presented captive female house finches, Carpodacus mexicanus, with a choice of two songs, each of which was identical except for a single parameter that had been either accentuated or minimized. To ensure that our playback tapes offered choices within the normal range of variation in house finch song, we recorded wild male finches, then created playback tapes using values for the song parameters under consideration that were above or below the mean for the local population, but within the natural range of variation. We presented females that had not previously mated with choices between songs that (1) were long or short, (2) were given at fast or slow rates, and (3) included a large or small repertoire of unique song elements. Females showed significant preferences for long songs and for songs presented at a faster rate, but there was no significant preference for large or small repertoires of unique song elements. Song length and rate each seem likely to indicate a males energy reserves, and thus could be important sources of information for females choosing mates.

ULTRAVIOLET BEAK SPOTS IN KING AND EMPEROR PENGUINS

Abstract In seabirds, colors of feathers and external tissues have only recently been studied, and ultraviolet (UV) color has not yet been detected. Using live individuals as well as museum skins, we found UV peaks of reflectance in two large Aptenodytes species, King (A. patagonicus) and Emperor (A. forsteri) Penguins. UV reflectance did not occur on the feathers, claws, or skin of these species, nor did we find UV reflectance in five other genera of penguins (11 species). UV peaks overlapped with spots of color on the lower beak that appeared orange for human observers, and beak spots differed slightly in location between the two species. Adults of both sexes possessed these UV markings, but they were lacking in juveniles, as was the orange color of the beak spot, and auricular patches used for selecting mates. Finally, measurements of free-ranging King Penguins showed that recently paired birds had higher UV reflectance than courting ones, suggesting possible roles of UV beak spots in pairing and as an indicator of sexual maturity. Manchas Ultravioleta en el Pico de los Pingüinos Aptenodytes patagonicus y A. forsteri Resumen. En las aves marinas, los colores de las plumas y los tejidos externos sólo han sido estudiados recientemente, y el color ultravioleta (UV) todavía no se ha detectado. En individuos vivos así como en pieles de museo, nosotros encontramos picos de reflectancia UV en dos especies de pingüinos, Aptenodytes patagonicus y A. forsteri. El color UV no se encontró en las plumas, las garras o la piel de estas especies, ni encontramos color UV en otros cinco géneros de pingüinos (11 especies). Los puntos UV se encontraban superpuestos con manchas de color ubicadas en la parte baja del pico que parecían anaranjadas para los observadores humanos. Las manchas del pico difirieron levemente en forma y localización entre las dos especies. Los adultos de ambos sexos presentaron las manchas UV, pero éstas no estaban presentes en los juveniles, al igual que el color anaranjado de la mancha del pico y los parches auriculares empleados en la selección de parejas. Medidas tomadas en individuos libres de la especie A. patagonicus demostraron que los que se habían apareado recientemente tenían presentaban reflectancias de UV mayores que las de aquellos que aún estaban cortejando, lo que sugiere un posible papel de las manchas UV del pico en el apareamiento y como indicadoras de la madurez sexual.

The Number of Provisioning Visits by House Finches Predicts the Mass of Food Delivered

Abstract One classic means of assessing variation in avian foraging success and provisioning effort, counting the number of trips to the nest, assumes that parents bring equal amounts of food during each trip. We tested this assumption in male House Finches (Carpodacus mexicanus) by using video cameras to record both the number of nest visits and the mass of food delivered as measured by an electronic balance. We compared the number of feeding visits and mass of food delivered at each of three stages in the nest cycle: incubation, young nestlings, and older nestlings. The number of provisioning trips was significantly correlated with the mass of food provided by a male to his mate or to their offspring during each stage of the nesting cycle. Furthermore, this correlation became stronger as the breeding cycle progressed. These observations support the assumption that, for this species and perhaps others that carry food in their crop, the number of provisioning visits to the nest is a reasonable predictor of the mass of food provided. El Número de las Visitas del Aprovisionamiento de Carpodacus mexicanus Predice la Masa del Alimento Entregada Resumen. Un método clásico para establecer la variación en el éxito de forrajeo y esfuerzo de aprovisionamiento en aves, que consiste en contar el número de viajes hacia el nido, asume que los padres llevan igual cantidad de alimento durante cada viaje. Probamos este supuesto en machos de Carpodacus mexicanus mediante cámaras de video que registraron tanto el número de visitas al nido como el peso del alimento entregado, este último medido mediante una balanza electrónica. Comparamos el número de visitas al nido y el peso del alimento entregado en cada uno de los tres estados del ciclo de nidificación: incubación, pichones jóvenes y pichones tardíos. El número de viajes de aprovisionamiento se correlacionó positivamente con el peso del alimento provisto por un macho a su pareja o a su progenie en cada estado del ciclo de nidificación. Además, esta correlación se hizo más fuerte a medida que el ciclo de nidificación progresó. Estas observaciones apoyan el supuesto que, al menos para esta especie y otras que transportan alimento en el buche, el número de visitas de aprovisionamiento al nido predice razonablemente bien la masa de alimento provista.

Characterization of mycoplasma gallisepticum infection in captive house finches (Carpodacus mexicanus) in 1998.

Since 1995, the epidemic of mycoplasmal conjunctivitis in eastern house finches has affected the Auburn, AL, house finch population. To better characterize the current status of this host-parasite interaction, we established a captive flock of 38 seronegative, healthy finches in fall 1998. After a minimum quarantine period of 4 wk, two Mycoplasma gallisepticum (MG)-infected house finches were introduced into this flock. Over a 12-wk period, the flock was captured every 2 wk and each bird was observed for conjunctivitis. Blood and choanal swabs were collected from each bird for serologic analysis and for the detection of MG by polymerase chain reaction. The infection spread rapidly through the flock just as it had in a similar study performed in 1996 at the height of the epidemic. Unlike the earlier study in which birds remained chronically infected, most of the birds in our study recovered rapidly, and only three of the birds died during the study. Two patterns of host response to infection with MG were observed. Twenty-seven birds (73%) experienced an acute conjunctivitis that resolved, and the birds appeared to clear the infection. Ten birds (27%) suffered prolonged clinical disease, and MG could be detected in these birds intermittently throughout the experiment. These results, in conjunction with our surveys of MG in the wild population, suggest an evolving host-parasite interaction.

CHARACTERIZATION OF THE MYCOPLASMAL CONJUNCTIVITIS EPIZOOTIC IN A HOUSE FINCH POPULATION IN THE SOUTHEASTERN USA

An epidemiological study of the prevalence of mycoplasmal conjunctivitis in the house finch (Carpodacus mexicanus) was conducted in Auburn (Alabama, USA) between March 1998 and February 1999. Clinical disease was observed in 4% of the 1,214 finches trapped and examined. This rate is comparable to the average annual prevalence observed in this population since 1996, although the prevalence of clinical disease observed in the peak months of September through November was lower than in previous years. Clinically ill birds were observed in all months of the study. To estimate the prevalence of recovering and asymptomatic, infected birds, we tested a subset of 334 house finches serologically for exposure to Mycoplasma gallisepticum (MG) using the serum plate agglutination (SPA) assay. The prevalence of clinical disease in this subsample was slightly higher (7%) than in the entire sample, reflecting the fact that the serological survey was initiated in the late summer when the prevalence of MG infection peaks in our study population and a sampling bias for symptomatic birds. The serological survey indicated that 13% of this subpopulation had been exposed to MG. We also tested 46 of 334 finches by polymerase chain reaction (PCR) to detect MG in seropositive, asymptomatic birds. Use of the PCR in conjunction with the SPA detected six asymptomatic, infected birds that may represent potential carriers or birds in the early stages of infection. The decreasing prevalence of clinical disease observed during the peak months suggests a changing host-parasite relationship. Continued surveillance of this population, employing both clinical observation and serological analysis will be useful in characterizing these changes over time.

PARENTAL CARE IN RELATION TO BROOD SIZE IN THE HOUSE FINCH

Abstract Studies that quantify parental care in birds are often faced with the confounding effects of variation in brood size. That is, nestlings from broods of varying sizes may receive different quantities of food for reasons not entirely related to the parental quality of the adults. To control for variation in brood size, researchers often divide feeding visitation rates by brood size to yield a per-nestling feeding rate. This presents problems, however, if adults adjust food load size in response to variation in brood size. We examined the relationship between brood size and parental care in the herbivorous House Finch (Carpodacus mexicanus) by considering not only visitation rates but also food load sizes. As brood size increased, the overall visitation rate increased but both per-nestling visitation rate and per-nestling food load rate decreased. The relationship between brood size and per-nestling care was similar regardless of whether we considered visitation rate or actual food loads, suggesting that in the House Finch per-nestling visitation rate serves as a reasonable index of total mass of food received by the nestlings. However, we urge caution in assuming that per-nestling visitation rate is an adequate measure of parental care in other species.

Effects of Mycoplasma gallisepticum on Reproductive Success in House Finches

Long known as a pathogen of poultry, Mycoplasma gallisepticum (MG) was first detected in house finches in 1994. The disease rapidly spread throughout the eastern United States and Canada and was associated with debilitating disease and high mortality in house finches. However, in the late 1990s, the proportion of infected finches dying as a result of infection with MG decreased, and asymptomatic infection was more common among wild birds than in the past. We documented MG infections in breeding house finches and concluded that adults of both sexes transmit the infection to dependent young, probably after hatch. MG infections of breeding adults occurred late in the breeding season and were found in birds completing significantly more nests than birds that never tested positive for MG, implying that higher rates of reproduction carry a cost in the form of increased risk of infection. We found evidence of an MG-induced delay in dispersal of nestlings from their natal area and demonstrated a significant impact of infection on nestling growth.