1 Matt Johnson Lecture Notes ORNITHOLOGY (Humboldt State Univ. WILDLIFE 365) LECTURE 12 - ANNUAL CYCLE Birds operate in world punctuated by cycles of stress and opportunity. The major activities or birds -- reproduction, molt, and migration -- are guided by internal clocks so that they coincide with appropriate seasons. In general, seasonal changes in day length - called photoperiod - control gonadal development and reproductive activities. Ultimate factors such as food supplies and climate influence the evolution of activity timing. I. II. The annual cycle. A. Cycles vary considerably between species and regions.: 1. Temperate resident (e.g., a chickadee) ON BOARD 2. Tropical resident (e.g., a jacamar) ON BOARD 3. Temperate-Tropical Migrant (e.g., a migrant warbler) ON BOARD 4. The important concept of repro-survival trade-off…..ON BOARD B. Temperate can be either N or S a. birds breeding in southern hemisphere do so during the warm productive months there; Sept. - Dec. The shearwaters we saw on pelagic trip breed off of Chile and off New Zealand, they "winter" (our summer) off Pacific coast. b. Birds breeding in tropics usually do so over longer periods of time than do birds breeding in either N. or S. temperate areas. C. Varies considerably within species and regions. 1. White-crowned sparrows: OVERHEAD Gambel's migrates but Nuttall's doesn't. 2. Hermit and Swainson's Thrushes co-exist in summer in Pacific NW forests; Hermits winter along Calif. coast, but Swainson's thrushes (with whom they are sympatric in summer) winter in Central America. The rhythms. A. Circadian - having a periodicity of about 24 hr; used to regulate behavioral patterns such as feeding, sleeping, singing, etc. 1. Not always strictly 24 hrs. kept in constant dim light, Chaffinches in captivity begin to "wake up" an hour earlier 2 III. IV. each day, suggesting that their circadian rhythm is close to 23 hours. OVERHEAD 2. Discrepancies between a bird's circadian rhythm and a 24 hr cycle are adjusted, or entrained, by external cues, such as sunrise, called Zeitgebers. Indeed, sunrise and sunset have profound influences on all animals' lives, including our own behaviorally, psychologically, physiologically. 3. But why the discrepancy? Not understood, but leading hypothesis is that animals need to be flexible for day length can change with global climate changes. A rhythm shorter than 24 hrs precludes them from getting “left behind” in a warming climate, while a cooling climate would delay sunrise and allow the zeitgeber to adjust the animal’s activity. It’s an evolutionarily conservative strategy, just as it is conservative to set your alarm earlier than you actually need to get up…..”just in case.” B. Circannual - having a periodicity of about 1 year; used to regulate timing of molt, breeding, and migration. -- these cycles probably also vary in some species from precise 12 month cycles and are entrained by changing day length, but this is poorly studied. Anatomy and Integration of Environmental Cues. A. Photoperiod. William Rowan did much of this work with captive birds (e.g., juncos) in the 1920's and 30's. He found that by carefully adjusting birds' exposure to light and dark in the lab, he could get juncos in winter to increase gonad size, and her could get crows to orient to the north for what should have been winter migration. The photoperiod cycles operated even when birds' eyes were blocked or removed, so a neurological pathway was clearly involved. B. Anatomy of the whole thing. 1. Photoreceptors in the brain, the hypothalamus to be exact, are sensitive to light that penetrates the skull and brain. 2. Once stimulated, these photoreceptors signal to the pituitary to synthesize and release Follicle Stimulating and Leutenizing Hormone, that affect gonad growth and development. FSH gamete formation and LH regulates secretion of test. and estro. from gonads 3. The pathway for light to influence molt is less wellunderstood, but it in some way involves the thyroid and its hormones. Reproductive hormones, androgens and estrogens, generally inhibit molt, which is usually too energetically demanding to undertake while breeding (except in a few species). Breeding Seasons A. Most birds breed annually, but there is considerable variation. 3 V. 1. Even in birds that do breed "annually," some do so by attempting only one brood, others produce multiple broods (but undertake the general breeding activity only once per year). 2. Others actually have multiple reproductive periods per year. Some tropical species nest continually. 3. Sooty terns on Ascension Island (tropical Atlantic) nest every 9.6 month. That is, high food availability allows them to reproduce continually, but it takes time to raise young and molt. 4. Many bird species reproduce variably during a year depending on local environmental conditions. Many tropical and desert breeding species begin breeding facultatively after rains begin. B. Timing of breeding. In birds that do exhibit discrete breeding seasons, various factors influence the timing of breeding. 1. Rainfall a. Budgerigars in Australia wait, up to several years, for onset of rains, at which time they go like crazy until rains cease. b. Many others species, most species in Northern Neotropics, initiate breeding at end of dry season, so that when the chicks hatch the rains have begun and food is plentiful. This indicates that provisioning food for young is more demanding that obtaining enough food for egg production. c. Birds on Cocos Island, off Costa Rica, breed in dry season. Food is plentiful year-round, but the birds can thermoregulate better when its dry. 2. Temperature - American Robins breed across a broad latitudinal range over which the onset of egg-laying is equally variable. Robins begin nesting 3 days earlier for every degree of latitude and progressively cooler temperatures. 3. Other factors a. The link between rain and/or temp and breeding activity probably evolved so that birds could time their reproduction to coincide with abundant food. These patterns have evolved when future food is predictable based current temperature and/or rainfall. But sometimes food is not so predictable, and birds have evolved to respond to food itself. b. Pinyon Jays feed extensively and opportunistically (spring summer fall) on seeds of pinyon pine, which has patterns of abundance and shortage that are difficult to predict. Experiments in lab have indicated that just looking at green pinyon pine cones can stimulate gonadal development. Molt 4 VI. A. Complete molt is extremely energetically demanding. Feathers weigh up to about 40% of total lean dry body mass; to replace them all is very demanding. B. Because it is so costly, birds usually relegate molt to periods of the annual cycle when food availability is high but when other energetic demands are low (i.e., not during migration and reproduction). Various patterns exist: 1. The simplest pattern is (for N. temperate spp. With 2 annual molts): breedingcomplete molt (prebasic)fall migrationwinterincomplete molt (body feathers only; prealternate) breeding 2. Here's another, common in arctic nesters (which are under extreme time constraints in their short arctic breeding season): They initiate prebasic molt before breeding, arrest during incubation, and resume molt at staging (migration gathering areas) or wintering areas. ON BOARD 3. Here's another, the one for ducks: The males do their prebasic molt shortly after breeding and hold it (basic plumage) for just a few weeks (the eclipse plumage), then undergoing prealternate molt back to familiar drake plumage in fall, so that they can attract mates in winter. Females (hens) do the standard sequence; they don't have to impress anybody. 4. Here's another, one that is common in species that winter in relatively benign regions (migrant landbirds in tropics, shorebirds wintering in food-rich areas): They initiate prebasic molt before migration, arrest it during, then resume it on wintering quarters, leisurely replacing feathers over the entire winter. 5. Here's another one, a weird one: Female hornbills get "entombed" in their nests during breeding; they rapidly drop all feathers and replace them then -- might as well as long as male can feed her enough. 6. Here's another weird one: Common White-terns have all white plumage. Much feather strength is derived from pigments, which this hapless species lacks. So her poor feathers degrade quickly, she replaces them continually. That wraps up timing and physiology of annual cycle with respect to 2 activities of: molt and reproduction. Next lecture and the one after we go over one of the most fascinating and beautiful patterns in all of avian biology: migration.