Principios de Taxonomia
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5.12 Partially Migratory Birds – an Example of Genetic Polymorphismsj111<br />
however, se<strong>de</strong>ntary animals of the same species remain in their breeding habitat<br />
during the winter. At least in the example of long-distance migratory birds, this<br />
coexistence of migratory and se<strong>de</strong>ntary birds is not due to voluntary <strong>de</strong>cisions of the<br />
individual birds but is genetically programmed (Berthold and Querner, 1981; Sutherland,<br />
1988). The two populations of the same bird species are genotypically different.<br />
The migratory disposition of most birds, that is, their motive for leaving their<br />
breeding habitat, is not triggered by seasonal climatic changes, at least in the case of<br />
long-distance migratory birds. Only for so-called cold fugitive species<br />
( K€altefl€uchtlinge ) are <strong>de</strong>creasing temperatures or food shortage the immediate<br />
triggers for the migration. Among these species are certain European birds, which<br />
generallymigrateonlyshortormediumdistances,forexample,manyducks,geeseand<br />
swans (Anatidae) and some herons (Ar<strong>de</strong>idae). Several individuals of these species<br />
remain in their breeding habitat in the fall or pause in stopover regions on their way to<br />
thesouthwestofthebreedinghabitat.Theselatterindividualsremaininthesestopover<br />
regions until frosts and food shortages cause them to migrate further southwest.<br />
In the case of most long-distance migratory birds, however, it is not simply food<br />
shortages or the encroaching cold that causes the animals to leave their breeding<br />
habitat for southern or western climes. These migratory birds leave their breeding<br />
habitats during high or late summer, when the temperatures and food supplies are<br />
still optimal. Consi<strong>de</strong>r the European Swift (Apus apus), which leaves central Europe in<br />
mid July, when airborne insects are still available in sufficient amounts. These birds<br />
do not migrate because certain living conditions have worsened or because they are<br />
starving or cold. Instead, these migrations are controlled by the <strong>de</strong>creasing length of<br />
the day. This migration behavior has a genetic basis. The birds become restless due to<br />
an internal clock, not because of external compulsions.<br />
The genetic disposition for migratory or se<strong>de</strong>ntary behavior likely requires the<br />
cooperation of several genes. Nearly nothing is known regarding these genes (Helm,<br />
2009). However, in most cases, the genetic differences between migratory and<br />
se<strong>de</strong>ntary birds are likely not differences in structural genes but rather (1) differences<br />
within regulatory elements (enhancers) of structural genes and (2) tissue-specific<br />
activities of certain transcription factors (Chapter 4).<br />
Migratory and se<strong>de</strong>ntary birds are distinguished by a multitu<strong>de</strong> of traits. A<br />
migratory bird requires additional instincts. In migratory birds, genes are active<br />
that control the instinctive migratory behavior twice a year at the proper time; these<br />
genes are inactive in se<strong>de</strong>ntary bird populations. Migratory birds also require a<br />
different metabolism and anatomy.<br />
In contrast to se<strong>de</strong>ntary birds, migratory birds become restless in the late summer<br />
and fly continuously in a particular geographic orientation. The direction of migration<br />
and the migration distance are genetically <strong>de</strong>termined. This is known because<br />
many songbirds migrate at night without parental guidance. These birds must locate<br />
the migration paths themselves, without the help of other individuals. Small songbirds<br />
are short-lived and can hardly benefit from the experience of the previous year.<br />
Instead, the bird must know instinctively whether it must migrate in the southwest or<br />
southeast direction. The bird must also know when it has arrived in the wintering<br />
grounds. A migratory bird must be genetically programmed to know how many days