Ross Furbush

ESS 201
23 May 2014
Eric Steig
What Causes Mistimed Phenology in Migrating Birds
driven by Global Climate Change?
Global temperature is steadily rising (Easterling 1997) due to an
anthropomorphic increase in carbon dioxide emission (CHANGE 2001). Global
climate change affects different parts of the globe at different rates (Solomon 2007)
affecting many ecosystems and biological systems. This difference can lead to
mismatched phenology between migratory birds and their local resources, whether
at breeding grounds or wintering grounds, due to their global life history (Hegland
2009).
The ecological strategy of migratory birds is to feed productively yearlong,
particularly during breeding; this strategy comes with the acquired risk of longdistance travel. High latitude summer breeding grounds are optimal for passerines
(perching song birds: in Order Passeriformes) because of extended daylight,
particularly during the insect feeding periods of dawn and dusk; this environment,
however, is not suitable during the winter season. This migration journey has been
evolutionarily synchronized with breeding ground peak abundance. Although some
species can alter clutch size, laying interval, or incubation period to synchronize
with food peak, global climate change will make these adjustments more difficult by
extending the period between the first laid egg and peak food abundance (Cresswell
2003).
It is observed that as global temperature increases, phenological processes
advance: spring comes earlier. Although this advancement in flora flowering, insect
emergence, tree budding, avian breeding, or migration arrival (Both 2006) can be
parallel in magnitude, there is still high variation among species (Hegland 2009).

The observed variation causes a mismatch in timing between predator and peak
prey abundance, or between mutualistic partners such as migrating pollinator and
flowering plant (Hegland 2009). This phenolocial phenomenon altered by global
climate change reveals yet another negative effect of global carbon dioxide
emission.
I will explore different migratory bird ecological characteristics that have
been tested and support an effect on migrating passerines resulting in a
mismatched phenological system. These include: (1) migration cues, (2) migration
distance, (3) brood number, (4) breeding latitude, and (5) population status. I will
also explore other testable hypotheses that could potentially result in a mismatched
phenological system; these include: (1) diet, (2) geographical structures, (3) group
size, and (4) urbanization. Finally, I will consider conservation tactics that humans
could employ to facilitate natural migration synchronization with peak resource
abundance.
Migration cues can be driven by temperature, a direct effect of global climate
change, or by endogenous rhythms, which are not affected by global warming. The
pied flycatcher (Ficedula hypoleuca), for example, which nests in Europe and
migrates to Africa, relies on length of day for migration timing instead of
temperature (Both 2001). Migration nest site arrival date for these flycatchers has
not advanced, whereas the date of caterpillar peak, a temperature driven
phenomenon, has. The phenotypic plasticity of flycatcher females resulted in
advanced laying dates, but this is driven by selection not temperature (Both 2001).
If this species migration phenology relied on the same phenological cues of
caterpillars, temperature, they would not be expected to be as poorly matched.
There has been a 90% decline in the past two decades for populations that are
mistimed and a 10% decline in populations who have accurate timing with

caterpillar abundance peak (Both 2006). The pied flycatcher serves as a

representative organism and it captures population declination trends seen in
similar studies with other species (Jenni 2003, Parmesan 2006).
Migration distance of passerines can vary considerably. This species
characteristic also impacts the birds ability to adjust phenology in response to
rising global temperature. The majority of long-distance migratory passerines adjust
peak-breeding season forward, whereas short-distance migratory species actually
delay breeding ground arrival. Explanation for this trend is frankly unclear in the
literature, but it is said that areas along migration routes are more productive due
to global warming which is more favorable to short-distance migratory birds (Jenni
2003). It should also be considered that the longer, riskier pathway of long-distance
migratory birds has led to more adaptive variation in arrival date to breeding
grounds. This is driven by competition for prime nesting location on the early end
and by migration risks and complications on the delaying end. A smaller variation
for arrival date might be assumed with short-distance migratory species because
their journey is less complicated. Natural selection therefore has more early
variation to select for in long-distance migratory species.
Brood number also drives varying phenological responses to global climate
change among bird species. Those that have a fixed brood number have advanced
migration, but species with a variable brood number do not adjust their timing in
response to rising temperature (Jenni 2003). Species with variable brood number of
two or more are adapted to fail and re-nest and repeat; a flexible design that can
withstand temperature changing conditions. Those obligated to one brood will likely
show stronger selection for earlier arriving birds because mistimed, later arriving
birds will likely produce no offspring; a rarer occurrence in birds with a brood
number of 2 or more.

It is noteworthy from a conservationists prospective that species who show

population decline tend to not advance migration, whereas those with a stable or
increasing population forwardly adjust (Mller 2008). Many variables affect a species
population decline. Consider the traits of an ecological generalist and how they
might better prepare the generalist species to increase in population size in
response to many human-caused environmental changes. Fragile species to one
drastic change will likely experience low success in another, such as mistimed
phenology. These findings are important because we learn that the species of most
concern will likely have the most difficult time adjusting migration to match peak
food availability.
Communities and ecosystems at varying latitudes are affected differently by
global warming (Baldwin 2014). Higher latitudes express more sensitivity to climate
warming with respect to community composition and overall environmental quality.
It is for this reason that nesting migratory birds at these higher latitudes experience
more population declination (Mller 2008). Although it has not yet been tested, one might expect
northern migratory breeders to actually delay migration to arrive in an environment with more similar
historic temperatures.

An avian characteristic that has shown to impact migration, but not

phenology, is diet. Birds with a wintering diet primarily consisting of berries and
insects arrive earlier in migration compared to granivores, but neither diet
classification has significantly impacted adjusted phenology. In some cases the
omnivores arrived over a week before the seedeaters (Jenni 2003). Plant and insect
responses to global warming are not uniform (Both 2001) leading to a hypothesis of
phenology differentiation in the future as different diet types tackle diets reacting
diversely to global warming.
Migration flyways all encounter different geographic obstacles and physical
features (Dunn 2011). It has been shown that wintering grounds in relation to the

Sahara desert has impacted migration, but is not correlated with a species ability to
phenologically adjust to rising temperature (Jenni 2003). This brings to question the
effect of different geological features with respect to phenology timing including:
mountains, bodies of water, deserts, ecological deserts or other physical properties
such as wind and ocean currents (many of which drive particular flyways). Physical
features may impact how dissimilar a birds wintering grounds are from their
breeding grounds therefore impacting the degree of phenological difference
between each area. Populations of same or various species may also be
interspersed between said geological features with different micro-phenologies and
slightly altered migration pathways. This may lead to measurable impacts of
geological features on adjusting phenology.
No study has yet concluded that being either gregarious or non-gregarious
affects mistiming phenology. It has been noted that migration cohort size explains a
large percentage of migration timing variation (Miller-Rushing 2008). Migrating birds
that travel in larger groups may experience the most selection from this global
climate change phenomenon because a mistimed discrete arrival event would be
more detrimental than the continuous arrival of fewer individuals.
It is well understood that human ranges and urbanization has been
expanding. Humans remove the natural environment for a more uniform, built
environment. Many species have shown various population responses to these
urbanization altercations including population increases for non-native species and
those that can nest on buildings, and decrease for interior and ground-nesting
species in urban environments (Marzluff 2001). This environment is not an
ecological desert for some migrating species that take advantage higher water and
food availability (Savard 2000, Marzluff 2001), supplemented through residential
gardens, bird feeders, and sometimes garbage (Chace 2006). Phenology has shown

to advance in urbanized environments compared to surrounding natural areas (Neil

2006). Although this has not successfully been compared to effects of migrating
phenology mistiming, one would expect that populations breeding in an urban
environment would advance phenology compared to those nesting in a more
natural landscape.
Urbanization and agriculture has dramatically altered the natural
environment. Now that humans increasingly occupy many environments, it would
be of interest to explore how we may aid migrating birds with mistimed phenology,
perhaps on a mass scale. In a New York Times article, author Jim Robbins (2014)
writes about a program in California that pays farmers to flood their fields to aid
migrating shorebirds that feed in mudflats. This would aid mistimed phenology
because scientists could inform farmers to supply water at a precise time. This
would be difficult to accomplish in an urban environment, due to land ownership
fragmentation. Networks of parks and suburban residents could possibly supply
feed at feeder stations to coordinate with migratory bird arrival. This would be an
expensive endeavor and difficult to coordinate. Depending on the urgency of
population declination due to the mistimed phenology phenomenon, other
strategies may be called upon.
There is much to still learn about how climate change affects migratory birds
with respect to mistimed phenology. Many variables and ecological characteristics
have shown to advance or delay phenology in migratory birds at a different
magnitude than their diets. Birds on an endogenous rhythm, with a long migration,
constant brood size of 1, nesting at higher temperate latitude, and with a declining
population size have shown to have higher sensitivity to mistimed phenology. Other
characteristics with no recorded affects on mistimed phenology are hypothesized to
affect a birds ability to phenologically adjust due to high variation in diet,

geographic stuctures along migration route, group size, and urbanization. Continued
population declination due to this global climate change phenomenon may require
mass human action to supplement the mistimed diets.

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