Evolution

IIS2024

The Geography
of Evolution
Diego Tirira PhD
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Keywords for this class
Level 1 Level 2
• Geographic distribution • Wallace’s line
• Disjunct distribution • Extinction
• Biogeography • Dispersion
• Species dispersal • Vicariance
• Heredity • Phylogeography
• Ecological community
The Geography
of Evolution

Based on
Futuyma & Kirkpatrick (2017)
 Bearded pig (Sus barbatus), found in
Borneo, is one of the easternmost

The Geography of Evolution mammals limited by Wallace’s Line.

• Some of the most avidly read

books of the 18th and 19th
centuries were the tales that
explorers recounted of their
travels in exotic lands.
• One of these books was written

Futuyma and Kirkpatrick (2017)

by Alfred Russel Wallace, who
traveled unexplored rivers and
suffered tropical diseases to
contribute to the first study of
Biogeography in history.
The Geography of Evolution
• Other important travelers
were Alexander von
Humboldt, Charles Darwin,

www.biografiasyvidas.com/monografia
Friedrich Georg Weitsch
and many more.
• They told stories not only of
hardships and of encounters
with indigenous peoples, but
also of amazing animals and Alexander von Humboldt

plants, entirely distinct any

in Europe or North America.
Charles Darwin
 Diego Tirira (all)
The Geography of Evolution
• Thousand of these exotic
species from around the
world were recovered by
and deposited in natural

https://goldenagepaintings.blogspot.com
history museums, zoos,
and botanical gardens.
• All visitors to zoos and
gardens marvel at these
unique animals and
curious plants.
The Geography of Evolution
• But also, it caused a confrontation with the Bible and Noah's
Ark.

Carl Linnaeus (1758)

Naturalists in Ecuador
• The first contribution in actual Ecuador
corresponds to Pedro Cieza de León
(1553), a priest and chronicler sent by
the Kingdom of Spain.
• He documented in his Crónica del Perú

https://www.boxerlinks.eu/
all anecdotal encounters with the local
wildlife:
… «y en la comarca de Quito hay
gran cantidad de conejos, y por las
montañas algunas dantas».
Naturalists in Ecuador

• Other contribution at the end of

16th century was written by priest
José de Acosta: Historia natural y
moral delas Indias.

Public Domain
Naturalists in Ecuador
• Main naturalists that came to actual
Ecuador and recounted their travels
in exotic lands were:
• Charles Marie La Condamine
• Jorge Juan y Antonio de Ulloa
• Antonio Pineda
• Alexander Humboldt

Public Domain (all)

• Charles Darwin
• Gaetano Osculati
• Enrico Festa
The Geography of Evolution
• But the naturalists went further:
they asked why these creatures
were found only in these remote
regions?
• Why should apes be in Africa but
not South America
• Why are sloths only in tropical
America?

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The Geography of Evolution
• If European plants were found
growing near American
seaports, having been
accidentally transported across
the Atlantic, they asked:
• Why had they not already
occupied America?
• If those species could prosper

Bernard Gribble
in what proved to be a suitable
new region, why weren’t they
already there?
The Geography of Evolution

• Based both on their own travels and on

specimens brought to Europe by explorers,
naturalists started to describe the faunal and
floral differences among regions of the world.
• This was the beginning of the biogeography
(in the 18th century): the geographic

Public Domain (both)

distributions of organisms.
• All this information was compiled by Alfred
Wallace, the “father of biogeography”.
 Charles Darwin

The Geography of Evolution

• Wallace and Darwin expanded the scientific
framework for understanding geographic
distributions.
• In some instances, the geographic distribution Alfred Wallace
of a taxon may best be explained by historical
circumstances.
• But in other, ecological factors operating at the

Public Domain (both)

present time may provide the best explanation.
• Hence the field of biogeography may be roughly
divided in those two aspects: historical
biogeography and ecological biogeography.
Biogeographic Evidence
for Evolution
Biogeographic Evidence for Evolution
• The geographic distributions of organisms provided both
Darwin and Wallace with inspiration and with evidence
that evolution had occurred.
• Actually, the reasons for certain
biogeography facts are so obvious
that they hardly bear mentioning.
• If someone asks why there are
no elephants in the Hawaiian

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Islands, we will naturally answer
that elephants couldn’t get there.
Biogeographic Evidence for Evolution
• This answer assumes that elephants originated
somewhere else: perhaps on a continent.
• But in a pre-evolutionary view,
such as that of divine creation,
the answer would be different:
• “This is the work of the Creator”.
• “He, in his wisdom (sabiduría)
placed the species in the place

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he considered best.
Biogeographic Evidence for Evolution
• Darwin devoted two chapters of On the
Origin of Species to showing that many
biogeographic facts that make little sense
under the hypothesis of special creation
make a great deal of sense if a species:
• (1) has a definite site or region of origin,
• (2) reaches a broader distribution by

www.crowdfinder.be
dispersal, and
• (3) becomes modified and gives rise to
descendant species in the various regions
to which it disperses.
Biogeographic Evidence for Evolution 1
• Darwin emphasized the following points:
• First, “neither the similarity nor the
dissimilarity of the inhabitants of various
regions can be fully explained by climates
and other physical conditions.”
• Similar climates and habitats, such as

www.crowdfinder.be
deserts and rainforests, occur in both the
Old and the New World, yet the organisms
inhabiting them are unrelated.
 Biomes of the planet

Olson et al. (2001)

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Biogeographic Evidence for Evolution
• There are members of diverse plant families
adapted to deserts by convergent evolution:
• The cacti (Cactaceae) is almost restricted to the
New World, but the cactus like plants in Old

Futuyma and Kirkpatrick (2017) (all)

World deserts are members of other families. Cactaceae (Peru)

Cactaceae (Mexico) Apocynaceae (India) Euphorbiaceae (Ethiopia)

 Even their distribution
Primates they have similar
Phylogeography habitats and
diets

• Phylogeographic studies find

the phylogenetic
relationships among
populations, and can be used
to infer their history of Catarrhini
spread.

Map by Wikipedia
Strepsirrhini
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Platyrrhini
Biogeographic Evidence for Evolution 2
• Second, “barriers of any kind, or obstacles to
free migration, are related in a close and
important manner to the differences
between the productions [organisms] of
various regions.”
• Darwin noted, for instance, that marine
species on the eastern and western coasts

www.crowdfinder.be
of South America are very different.
Biogeographic Evidence for Evolution
Coypu
3
• Third, “inhabitants of the same
continent or the same sea are related,
although the species themselves
differ from place to place.”
• Darwin’s example: aquatic rodents
of South America (the coypu and
capybara), which are structurally
similar to, and related to, South

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American rodents of the
mountains and grasslands.
Capybara
 Biogeographic Evidence for Evolution
Coypu
3
• But not are related to
the aquatic rodents
(beaver, muskrat) of
the Northern
Hemisphere.

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Beaver
Wikipedia

Muskrat
Capybara
Biogeographic Evidence for Evolution
• According to Darwin, there is a deep
organic bond (vínculo), across space
and time, over the same areas of land
and water, regardless of physical
conditions.
• This link is simply heredity [i.e.,
common ancestry], that cause which
alone produces organisms closely

Public Domain
resembling each other.
Biogeographic Evidence for Evolution
• For Darwin, it was important to show
that a species had not been created in
different places, but had a single region
of origin, and had spread from there.

Public Domain
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Biogeographic Evidence for Evolution 1
• Darwin drew particularly compelling
evidence from the inhabitants of islands.
• First, remote oceanic islands generally have
precisely those kinds of organisms that are
capable of long-distance dispersal and lack
organisms that do not.

https://mapas.owje.com/maps
• For example, the only
native mammals on
many islands are bats.

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 Biogeographic Evidence for Evolution
Flightless cormorant (Galápagos)
1
• Island species with poor dispersal
ability, such as the dodo and other
flightless birds, are closely related
to strong flyers, and

https://www.magzter.com/stories
descended
from them.

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Dodo (Mauricio Island)
Biogeographic Evidence for Evolution 2
• Second, many continental species of
plants and animals have flourished on
oceanic islands to which humans have
transported them.
• Thus, said Darwin, “who admits the
doctrine of the creation of each
separate species, will have to admit
that a sufficient number of the best
adapted plants and animals were not

Shutterstock
created for oceanic islands.”
Biogeographic Evidence for Evolution 3
• Third, most of the species on
islands are clearly related to species
on the nearest mainland, implying

Heidi Snell
that was their source.
• This is the case for almost all the
birds, mammals, plants, and other
species of the Galápagos Islands.
• For example, rats of the genus

Pine et al. (2012)

Aegialomys.
Biogeographic Evidence for Evolution 3
• Island species often bear marks of their
continental ancestry.
• For example, as Darwin noted, hooks
(ganchos) on seeds are an adaptation

https://mammothmemory.net/biology
for dispersal by mammals, yet on
oceanic islands that lack mammals,
many endemic
plants nevertheless
have hooked seeds.

Jude / Flicker
 Biogeographic Evidence for Evolution 4
• Fourth, the proportion of species that are
restricted to an island (that is the Endemic
species) is particularly high when the
opportunity for dispersal to the island is low.

www.anapsid.org/chaco4figs.html
14 species

www.galapagos.org
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 Charles Darwin

Biogeographic Evidence for Evolution

• Wallace made a special study of species
distributions, especially on islands.
• He came to many of the same conclusions as
Darwin (points that hold true today, after more Alfred Wallace

than a century and a half of research).

• Greater knowledge of the fossil record and of
geological events, such as continental drift and

Public Domain (both)

sea level changes, has contributed to our
understanding, but has not invalidated any of
Darwin and Wallace's major conclusions.
Major Patterns of Distribution
Major Patterns of Distribution
• The geographic distribution of almost
every species is limited to some degree,

www.discoverlife.org/mp/20m?kind=Plethodon
and many higher taxa are likewise
restricted (endemic) to a particular
geographic region.
• For example, the salamander genus
Plethodon is limited to North America
(approx. 50 species)
• Some species are restricted to specific
areas.
 Major Patterns of Distribution Columbidae Apterygidae

• Some higher taxa are endemic

(e.g. the kiwi family, Apterygidae,
restricted to New Zealand).

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Wikipedia
• Whereas others (e.g. pigeon family,
Columbidae, are almost cosmopolitan).
Bruxaux (2018)

Wikipedia
Major Patterns of Distribution
• Wallace recognized that many higher taxa have roughly
similar distributions, and that the taxonomic composition
of the biota is more uniform Wallace’s Line

within certain regions than

https://wildlife-photographs.blogspot.com
www.britannica.com/animal/orangutan
between them.
• For example, Wallace
discovered a severe break in
the taxonomic composition
of animal species among
the islands between SE Asia
and Australia.
Major Patterns of Distribution
• As far east as Borneo,
most vertebrates
belong to Asian
families and genera,
whereas the fauna to

https://coraltriangleadventures.com
the east has
Australian affinities.
• This faunal break has
been called Wallace’s
line.
 Major Patterns of Distribution
• Wallace’s line separates
islands that, despite
their proximity and
similar climate, differ
greatly in their fauna.

https://sciencythoughts.blogspot.com
https://coraltriangleadventures.com
• The islands to the west
were connected to the
Asian mainland during
periods of low sea level;
those to the east were
not.
Major Patterns of Distribution
• Based on these observations, • Palearctic (Eurasia and
Wallace designated several northern Africa)
biogeographic realms (major • Nearctic (North America)
regions that have characteristic • Neotropical (Central and
animal and plant taxa) for South America)
terrestrial and freshwater • Ethiopian (sub-Saharan
organisms that are still widely Africa)
• Oriental (India and
recognized today:
Southeast Asia)
• These realms are more the result • Australian (Australia, New
of Earth’s history than of its current Guinea, New Zealand, and
climate or land mass distribution. nearby islands).
 Futuyma and Kirkpatrick (2017)
Biogeographic realms
Proposed por Alfred Russel Wallace (1876)
The geographical distribution of animals
Major Patterns of Distribution
• Each biogeographic realm is inhabited
by many higher taxa that are much more
diverse in that realm than elsewhere, or
are even restricted to that realm.
• For example, the endemic (or
nearly endemic) taxa of the

Futuyma and Kirkpatrick (2017)

Neotropical realm include
(anteaters and relatives,
Xenarthra), platyrrhine primates, most
hummingbirds, some suboscine birds
(flycatchers, and others), many catfishes,
and plants, as the bromeliads.
 Diego Tirira
Major Patterns of Distribution
• The borders between biogeographic realms
cannot be abruptly drawn because some taxa

BP Los Cedros
infiltrate neighboring realms to varying degrees.
• In the Nearctic realm (North America), for
instance, some species are related to, and

www.carnivorasland.com/tillandsias
have been derived from,
Neotropical stocks.
• Examples: an armadillo, an
opossum, and Spanish
moss (Tillandsia usneoides),
a bromeliad.
Disjunct distributions 20 species

• Some taxa have disjunct distributions;

that is, their distributions have gaps.
• Disjunctly distributed higher taxa
typically have different representants

Wikipedia

www.conifers.org/ar/Araucaria.php
in each area they occupy.
• Example, many taxa are represented
on two or more southern continents,
including lungfishes, marsupials,
cichlid fishes, and Araucaria pines.
Major Patterns of Distribution
• Common disjunct patterns are illustrated North America

by many taxa of animals and plants.

• The lady-slipper orchids
Eastern Asia
(Cypripedium) are found
both eastern Asia and

Futuyma and Kirkpatrick (2017)

eastern North America.
• Other examples of plants
are tulip trees (Liriodendron)
and skunk cabbages
(Symplocarpus).
Major Patterns of Distribution
• In animals, an example is alligators
(Alligator):

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• They are found in eastern North
America and in temperate eastern
Asia, but nowhere in between. Alligator mississippiensis (USA)
Alligator sinensis (China)
• Understanding how taxa
became disjunctly
distributed has long been a
preoccupation

Wikipedia
of biogeographers.
 Wikipedia (both)
Lungfishes distribution

https://www.slideserve.com/olwen/the-earth-and-the-origin-of-life-chapter-26
Historical factors affecting
geographic distributions
Historical factors affecting distributions
• The geographic distribution of a taxon is affected by both
current and historical factors.
• The limits to its distribution may be set by:
• Geological barriers that it has not crossed, or
• Current ecological conditions to which it is not adapted.
• Those patters on the historical processes that define the
current distribution of a taxon are defined by:
• (1) extinction,
• (2) dispersal, and
• (3) vicariance.
1 Extinction Horses
returned to
North America
• The distribution of: with European
• a species may have been reduced by the colonists
extinction of some populations, and
• a higher taxon by the extinction of some
constituent species.
• For example, the horses (Equidae) originated
and became diverse in North America, but
later it became extinct there.
• Only the African zebras and the Asian

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wild asses and horses have survived.
1 Extinction
• Likewise, extinction is the cause
of the disjunction between related
taxa in eastern Asia and eastern

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North America.
• During the Paleogene Alligator mississippiensis (USA)
Alligator sinensis (China)
(66–23 Mya), many plants
and animals spread
throughout the northern
regions of North America

Wikipedia
and Eurasia.
1 Extinction
• Their spread was facilitated by a
warm, moist climate and by land
connections from North America
to both Europe and Siberia.

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• Many of these taxa became extinct
in western North America in the Alligator mississippiensis (USA)
Neogene (23–5.3 Mya) as Alligator sinensis (China)
a result of mountain uplift
and a cooler, drier climate.
• Later there were

Wikipedia
extinguished in Europe by
Pleistocene glaciations.
 1 Extinction

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Alligator mississippiensis (USA)

Alligator sinensis (China)

Wikipedia
Wikipedia
2 Dispersal
• Species expand their ranges by dispersal (movement of
individuals).
• Some species of plants and animals can expand their
ranges rapidly.
• Within the last 200 years, many species of plants, and some
animals, accidentally brought from Europe by humans have
expanded across most of North America from New York and
New England.
• Some birds, such as the European starling (estornino)
(Sturnus vulgaris) and the house sparrow (Passer
domesticus), have done the same within a century.
 Futuyma and Kirkpatrick (2017)
2 Dispersal
History of
range
expansion of
the European
starling
(Sturnus
vulgaris)
following its
introduction
into New York
City in 1896
2 Dispersal
• Other species have crossed
major barriers on their own.
• The cattle egret (Bubulcus
ibis) has had the largest and
most rapid natural
expansion of all bird
species.

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2 Dispersal
• It was found only in tropical
and subtropical parts of the
Old World until about 140

FD
years ago.
• Then it arrived in South
America, apparently
unassisted by humans.
• It has since spread
throughout the warmer

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parts of the New World.
 https://dragonflyissuesinevolution13.fandom.com
3 Vicariance
• Vicarious =
“substitute” or “in
place of”.
• Ex. Vicario apostólico.
• Vicariance refers to the separation of populations of a
widespread species by barriers arising from changes in
geology, climate, or habitat.
• The separated populations diverge, and they often
become different subspecies, species, or higher taxa.
3 Vicariance
• Example, for many species of
fish, shrimps, and other marine
animal groups, the closest
relative of a species on the
Pacific side of the Isthmus of
Panama is other species on the
Caribbean side of the isthmus.

FD
3 Vicariance
• Each pair of species has
descended from a broadly
distributed ancestral species
that was separated by the rise Pacific Ocean
Caribbean Sea

Futuyma and Kirkpatrick (2017) (both)

of the isthmus of Panama
during the Pliocene (3 Mya).
• Vicariance sometimes explain
the presence of related taxa in
disjunct areas.
 Dispersion
Vicariance

Boletín Biogeograhhía SEBA (2014)

 Dispersion
Vicariance

Boletín Biogeograhhía SEBA (2014)

Historical factors affecting distributions
• In many cases, dispersal, vicariance, and
extinction together explain distributions.
• For example, during the Pleistocene
glaciations (< 2.6 Mya),
species moved their
ranges by dispersing
into new regions.
• Some northern, cold-
adapted species became

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distributed far to the south.

FD
Historical factors affecting distributions
• When the climate
became warmer,
these species
recolonized northern
regions, and
southern populations
became extinct.
• Except populations
of some species that
survived on cold
mountaintops.

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Historical factors affecting distributions
• Disjunct distribution
of a Saxifraga
cernua in northern
and mountainous
regions of the

Futuyma and Kirkpatrick (2017)

Northern
Hemisphere.
Historical factors affecting distributions
• Relict populations
have persisted at
high elevations
following the
species’ retreat

Futuyma and Kirkpatrick (2017)

from the
southern regions
that it occupied
during glacial
periods.
 Venezuela (Mérida)

Espeletia in Ecuador
Ecuador (El Ángel)

• Frailejones inhabit
continuously from

Wikipedia
western Venezuela,
Colombia and Ecuador (Llanganates)

https://laverdadnoticias.com/ecologia
northern Ecuador.
• Except for an
isolated population
in Llanganates,

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central Ecuador.
 www.goraymi.com
Historical factors affecting
distributions
• Populations or species that have been
left behind in this way may be called
relicts.
• In this case, dispersal expanded the

https://laverdadnoticias.com/ecologia
Ecuador (Llanganates)
range, and extinction of populations
in intervening habitat caused the
vicariant disjunction of populations.
Phylogeography
Phylogeography
• Phylogeography is the description and analysis of the
processes that explain the geographic distribution of
the species, especially within
closely related taxa.

https://mapofafricanew.blogspot.com
• These processes include the
dispersal of the organisms
that carry the genes.
• Provides understanding into
the past movements of
species (including humans).
Phylogeography
• Phylogeography provides insight
into the past movements of species
and the history by which dispersal
and vicariance have determined
their present distributions.
• Phylogeographic studies find the

Futuyma and Kirkpatrick (2017)

phylogenetic relationships among
populations, and can be used to
infer their history of spread.
Symphonia globulifera
Phylogeography
• For example, the tree
Symphonia globulifera
was shown to have
dispersed from West
Africa to South America
more than 15 Mya.

Futuyma and Kirkpatrick (2017)

• After which it crossed the
Andes and spread
through western South
America to Central
America.
Phylogeography

• Phylogeographic studies have

traced the expansion of species
from Pleistocene refugia after
glacial periods.
• Also shown how Pleistocene
events, such as sea level changes,
have shaped some geographic
patterns of genetic variation.

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Phylogeography
• For example, the genetic differences among populations of
many species of freshwater fishes and other species in the
SE United States show that they were separated into
western and eastern populations in the past, probably by
high sea level during interglacial episodes.

Futuyma and Kirkpatrick (2017)

Phylogeography
• These populations are thought to have been isolated and
diverged in two refugia during the Pleistocene.

Spotted sunfish Bowfin Sister species of mints

(Lepomis punctatus) (Amia calva) (genus Dicerandra)

Futuyma and Kirkpatrick (2017)

Phylogeography
• Phylogeography contributes to know the dispersion of
species and the vicariance that determine their current
distributions.
Geographic range limits:
Ecology and Evolution
Geographic range limits

• The geographic distribution of a species results not only from

the history of its ancestors, but also from current factors.
• This is major subject of ecological biogeography.
• Several difficulties can retard or prevent a species
from expanding its range. Amazon manatee

Guido Chaves
• To disperse to the new region
individuals need reproduce and to do
must find mates, which may be
difficult if there are few individuals and
the initial population density is very low.
Geographic range limits
• Self-fertilizing (selfers) plants have larger geographic ranges
than outcrossers (reproducción cruzada).
• The histogram shows the relative range sizes
of selfers compared with outcrossers
in 20 clades of flowering plants.
• Each column that extends

Futuyma and Kirkpatrick (2017)

above the horizontal axis shows
a clade in which selfers have
larger ranges than outcrossers.
• All, except one.
Geographic range limits
For a species to extend its range
and not to die in the attempt
must be able to:
• Survive physical conditions.
• Find suitable resources (food,
habitat), and
• Contend with other species,
such as competitors, predators,
and parasites.

Pinterest
Geographic patterns
of diversity
Geographic patterns of diversity
• The evolutive study of a community ecology can explain:
• Species diversity, Ecological community
• Species composition, and
• Trophic structure of assemblages
of coexisting species.
• Both ecological and historical
biogeography bear on these topics,

www.abc.com.py
since the geographic ranges of species
determine whether or not they might
coexist.
Geographic patterns of diversity
• A pattern in community ecology is called
the Latitudinal diversity gradient: the Ecological community
numbers of species (and higher taxa
such as genera and families) decline
with increasing latitude, both on land
and in the ocean.
• Most taxa of terrestrial animals and
plants are far more diverse in tropics,

www.abc.com.py
especially in lowlands with abundant
rainfall, than in extratropical regions.
Latitudinal
diversity
gradient
Vascular plants in
various regions in

Futuyma and Kirkpatrick (2017)

the Americas
drops more than
tenfold between
the Equator and
high-latitude
regions.
Latitudinal diversity gradient
• Tree frogs (Hylidae) are much less diverse
in the temperate zone than in the tropics.
• Only few lineages are adapted to the
temperate
zone,
and those

Futuyma and Kirkpatrick (2017)

only
recently.
Latitudinal diversity gradient
• The number of tree frog
species in a continental
region is strongly

Futuyma and Kirkpatrick (2017)

correlated with the time.
• Since the Hylidae first
started diversifying, as
estimated by applying a
molecular clock to the
branch points in the
phylogeny.
Resumen
 The Geography
of Evolution

Resumen
• La biogeografía, el estudio de las distribuciones
geográficas de los organismos, tiene componentes
históricos y ecológicos.
• Ciertas distribuciones son consecuencia de la historia
evolutiva a largo plazo; otras, de factores ecológicos
actuales.
• Las distribuciones geográficas de los organismos
proporcionaron a Darwin y Wallace algunas de las
pruebas más sólidas de la evolución.
 The Geography
of Evolution

Resumen
• Los procesos históricos que afectan a la distribución
de un taxón son la extinción, la dispersión y la
vicarianza (fragmentación de una distribución
continua por la aparición de una barrera).
• Estos procesos pueden verse afectados o acompañados
por cambios ambientales, de adaptación y de
especiación.
• Los procesos de dispersión o vicarianza pueden
deducirse a menudo a partir de datos filogenéticos.
 The Geography
of Evolution

Resumen
• Las distribuciones disjuntas pueden atribuirse en
algunos casos a la vicarianza, pero la dispersión
parece ser la causa más común.
• Los patrones genéticos de variación geográfica dentro de
las especies pueden proporcionar información sobre los
cambios históricos en la distribución de una especie.
• La distribución local de las especies se ve afectada por
factores ecológicos, que incluyen tanto aspectos
abióticos, del medio, como características bióticas,
como competidores y depredadores.
 The Geography
of Evolution

Resumen
• Por qué las especies no amplían indefinidamente sus
áreas de distribución, adaptándose cada vez más a
condiciones cada vez más lejanas, es una de las
principales preguntas de la biología evolutiva.
• Las pautas geográficas en el número y la diversidad de
las especies pueden deberse en parte a factores
ecológicos actuales, pero también pueden explicarse
por la historia evolutiva a largo plazo.