Ecología de Interacciones. MCB Unidad 3.

Trabajo de investigación. Especie – “hormiga”

Título original

Introducción especie de estudio

Ants, which represent the family Formicidae, have a stinging apparatus and so belong to
the aculeate suborder within the insect order Hymenoptera (some evolved taxa later lost
the ability to sting). The oldest ant fossils date from ≈100 mya, meaning that ancestral ants
most likely appeared during the early Cretaceous Period (144-65 mya)

The family Formicidae is extremely diverse with 12,651 known ant species [7], and an
estimated 3,000 to 9,000 additional species as yet unknown to science.

Importancia en el ecosistema que se desarrolla

Ants as eco-system engineers modify the habitat structure and change the nutrient
content of the soil that may increase the density of important decomposers and thereby
also possibly indirectly affect their predators.
Predation by ants could therefore lead to a reduction in density of other arthropods at all
trophic levels or potentially lead to a classic top-down trophic cascade whereby some
functional groups may increase because the density of their natural enemies is reduced
The bottom-up effects of ants include the modification of physical and chemical soil
parameters by bioturbation and the accumulation of organic material.
Because of the building of below-ground galleries, mounding and material mixing, the soil
of ant nests is characterized by the impeded formation of soil horizons, increased porosity,
drainage and aeration, reduced bulk density and modified texture and structure. Increased
content of organic matter, P, N and K.
Ecosystem engineers are defined as “organisms that directly or indirectly affect the availability of
resources for other organisms through modifications of the physical environment”

Ants are an important functional component of most terrestrial ecosystems. They reach
high abundances and occupy numerous niches both above and below ground. Because of
their high biomass density they dominate many ecosystems, comprising between 20 and
52% of animal biomass in the tropics (Stork, 1996; Dial et al., 2006). For example, there are
estimated to be 8.6 million ants per hectare in Amazonian rainforest (Beck, 1971). High
densities of ants have also been recorded in temperate regions with ants surpassing other
macroinvertebrate groups in biomass (King, Warren & Bradford, 2013) (Fig. 1). Some ant
species create extremely large colonies, which can be interconnected and cooperate in a
phenomenon known as a supercolony. This can allow ants to reach even higher densities:
Japanese grasslands can support 1.13 million ants per hectare of a single species, Formica
yessensis (Higashi & Yamauchi, 1979). Ants interact with many organisms in diverse roles.
They are effective predators of a wide range of animals (Hölldobler & Wilson, 1990). The
predation pressure ants exert on arthropod communities is of great importance, and they
can shape entire insect communities (Floren, Biun & Linsenmair, 2002), and increase plant
growth by reducing herbivore numbers (Schmitz, Hambäck & Beckerman, 2017). Many
ants are involved in mutualisms. Ants tend honeydew-producing insects on plants,
protecting them against predators and feeding on their carbohydrate-rich excretions
(Styrsky & Eubanks, 2007). Many ants also form mutualisms directly with plants, trading
protection against herbivores and plant competitors for housing space inside the plant,
and consuming plant-provided food (Rico-Gray & Oliveira, 2010). Through mutualisms with
cellulose-decomposing fungi (Agaricales: Leucocoprinus), leafcutter ants (tribe Attini) are
also able to use plant materials as a food source, thus acting locally as effective and often
selective herbivores that consume up to 17% of annual tree leaf production in Neotropical
forests (Vasconcelos & Cherrett, 1997). Ants are also important seed dispersers for an
estimated 4.5% of angiosperm plant species globally (Lengyel et al., 2009). Because of
their high abundances, ants may have important effects on the environment. Recent
studies show support for ants being the main scavengers in some ecosystems, particularly
in the tropics (Fayle et al., 2011; Tan & Corlett, 2012) with ants being responsible for 61%
of all invertebrate-removed food items on the rainforest floor (Griffiths et al., 2018). As a
result of this, ants indirectly accelerate the decomposition of dead organic matter and the
redistribution of nutrients (Frouz, Santruckova & Kalcik, 1997; Frouz & Jilková, 2008). Ants
build large, mainly underground nests. During the building process they turn over vast
quantities of soil – a process called soil bioturbation. The amount of soil moved is
estimated at 1–5 tons per hectare per year, but could reach 5–50 tons per hectare per year
(Wilkinson, Richards & Humphreys, 2009). Ant bioturbation affects the distribution of soil
colloids and soil organic matter, stimulates microbial activity, and creates soil pores, which
increases aeration and water infiltration, thus influencing overall soil health (Gabet,
Reichman & Seabloom, 2003; Meysman, Middelburg & Heip, 2006). As a result, ants have
positive impacts on plant growth in both natural ecosystems and in agricultural areas
(Evans et al., 2011).
-Ant-termite interactions: an important but under-explored ecological linkage

Especies que interactúan en ese hábitat nombre tipo de interacción de cada caso,
intensidad
Types of interaction between ants and termites

Because of their ecological significance, numerous interactions with other organisms,

relative ubiquity and common co-occurence, understanding the relationships between
ants and termites is important. However, these interactions are still very poorly
documented (see online supporting information, Fig. S1). This is despite the two groups
being estimated as each having as much global biomass as all other terrestrial arthropods,
and nearly an order of magnitude more biomass than all wild vertebrates (Fig. 1). Ants and
termites interact in a number of ways, including living together commensally,
mutualistically, competing for nesting space, and probably most importantly as
predators/prey (see Table S1; Hölldobler & Wilson, 1990). Co-habitation involves ants
living inside a termite nest taking advantage of termitarium structures. This arrangement is
usually beneficial for ants, when cleptobiosis (theft of food or another item of value from
another animal) or lestobiosis (cleptobiosis but with the thieving species nesting in or near
the chambers of the host species) is involved, and can range from detrimental to beneficial
for termites. Co-habitation has been best studied for the minority of termite species that
build externally visible mound structures. These structures represent protected spatial and
functional niches in the environment and consequently, the termite mound is often used
as a nesting site by numerous ant species (Holt & Greenslade, 1980), even while still
inhabited by termites. Wheeler (1936) recorded 198 ant species inhabiting termite nests.
Although these inquiline ants interact with host termites in various ways, there is little
information on the nature of these interactions. Ants can either inhabit the parts of
termite nest where termites do not occur, or ants can exclude termites from a certain part
of the nest (Lubin & Montgomery, 1981). It is likely that inquiline ants living in termite
nests feed opportunistically on termite brood or adults (Jaffe, Ramos & Issa, 1995), as well
as on other inquiline arthropods present in the termitaria (Gallego Ropero & Feitosa,
2014). The relationship between inquiline ants and termites can also shift from
commensal, in which only the ants benefit and the fitness impacts on termites are
minimal, to more mutualistically beneficial interactions (Jaffe, Ramos & Issa, 1995). Where
the relationship is mutualistic, ants can benefit from the use of a nesting site in the termite
nest, while termites can consume ant food remnants, which are rich in nitrogen, and even
benefit from ant protection (Hölldobler & Wilson, 1990; Diehl, Junqueira & Berti-Filho,
2005). Jaffe, Ramos & Issa (1995) observed a common protective reaction of ants and
termites living in the same nest against other attacking ant species. In this case, ants and
termites were not physically separated in the nest. Sometimes the protective burden falls
on the ant partner, with the ant Camponotus sp. effectively protecting nests shared with
termites against intrusion of the regular termite predator ant Iridomyrmex sanguineus
(Higashi & Ito, 1989). Similarly, nesting in the base of the mound of the termite
Odontotermes latericius, the ant Pheidole megacephala was observed to attack predatory
Megaponera analis workers when they attempted to raid the termite colony in African
savanna (Sheppe, 1970). Note that P. megacephala is globally invasive, although this
behaviour was observed in Zambia, which might be part of its native range
[www.antmaps.org (Economo & Guénard, 2016)]. However all these instances of ‘defence’
could be by-products of ants protecting their own nest or territory. It is unclear whether
any of these ant–termite interactions have progressed beyond being ‘by-product’
mutualisms (De Jaegher, 2017) to a stage where there is reciprocal altruism between
partners. Furthermore, the degree to which termites benefit from co-habiting with ants
appears to vary among both ant and termite species. Ants can also inhabit the same niche
as termites and hence potentially compete for nesting space. Deadwood is an important
nesting site for both ants and termites and they compete for this resource (Kimber &
Eggleton, 2018). Termites furthermore not only inhabit (and defend) pieces and logs of
deadwood, but they also consume wood and thus remove this nesting site from the
environment. Another example comes from standing, living trees. Colonies of the two
groups inhabit the tree Cecropia pachystachya in Brazil, being spatially and mechanically
(termites build protected foraging galleries from fibrous material) segregated from each
other, without any apparent direct antagonistic interactions (Neves, Bernardo & Santos,
2014). Similarly, ants build physical barriers from fibrous material at territory boundaries in
this system (Quinet, Tekule & de Biseau, 2005). Termites can also use vibrations to detect
or avoid ants, or even mimic ant vibrational signals to prevent direct confrontation (Oberst
et al., 2017). Something similar is observed in epiphytic bird’s nest ferns (Asplenium spp.),
in which ant colonies and termite colonies are able to co-exist in the root mass of larger
ferns, but smaller ferns support colonies of either ants or termites with the two groups not
co-occurring (Ellwood, Jones & Foster, 2002). Presumably, ants and termites compete for
nesting space in the ferns. The behaviour and feeding nature of ants is also of importance.
Ants predating arthropods can lower the activity of foraging arboreal termites, while non-
predatory ants had no such effect in Brazilian rainforest (Conçalves et al., 2005). This
suggests that predation but not competition (for space) is a limiting factor for termite
activity. Because termites represent an abundant food source, and many ant species are at
least partly predatory, predation of termites by ants is probably the most common type of
interaction (Table S1) and this forms the focus of the remainder of this review. Ants have
been described as being the most significant and regular predators of termites (Deligne,
Quennedey & Blum, 1981; Abe & Darlington, 1985; Hölldobler & Wilson, 1990). Since the
global-scale distributions of ants and termites overlap (Fig. 2), the two groups are often
found in the same habitat, and many ants are often predate a broad range of insects, it is
expected that predation of ants on termites should also be widespread. Since termites are
mainly detritivores, predation of ants by termites is unlikely to occur. The only evidence for
any consumption of animal-derived food by termites is keratophagy (consumption of skin)
on mammal carcasses in the African savannah (Freymann et al., 2007), feeding on
vertebrate carcasses by Nasutitermes termites in Panama (Thorne & Kimsey, 1983) and
feeding on rat carrion by Rynchotermes nasutissimus in Brazil (Prestes et al., 2014). An
anecdotal case of termites foraging for ant bodies was recorded when the termite
Nasutitermes corniger harvested Azteca sp. ants, freshly killed during defence of their nest
(Jaffe, Ramos & Issa, 1995). However, the latter might be a case of hygienic behaviour, as
termites often clean up dead nestmates, or consume them as they are rich in nitrogen,
which is a scarce nutrient in wood-feeding termites (Shelton & Grace, 1996; Neoh et al.,
2012; Sun, Haynes & Zhou, 2013). Since there is no evidence of termite predation on ants,
we here discuss only cases in which ants directly predate termites.
Ant-termite interactions: an important but under-explored ecological linkage

Interacciones negativas:

Many interactions between species have winners and losers: one individual benefits, and
the other one suffers. These are called negative species interactions. For
example, predation is a type of species interaction in which one organism (a predator)
eats another organism (the prey)—this is good for the predator but very bad for the prey!
Different ant species can eat many different things, and some ants are important
predators. Predatory ants often eat other insects like termites and caterpillars, while other
ants eat only plants or fungus. Megaponera ants have only one food—termites—and
these ants organize huge hunts in which worker ants infiltrate termite colonies, capture as
many termites as they can carry, and bring the termites back to the ant nest to feed to
their larvae [3] (Figure 2). But even though many ants are predators, they can also be prey
for other animals. Lots of animals like to eat ants: birds, ant eaters, even humans in some
parts of the world! An animal that eats ants is called a myrmecophagous animal. The
authors of this paper have eaten ant larvae that were specially prepared, and we can
confirm that they are both delicious and nutritious.

Another type of negative interaction is called parasitism. Just like predation, parasitism
has winners and losers, but the losers typically survive the interaction. Parasitism occurs
when an organism steals a resource from another organism. For example, some ant
species called slave-making ants parasitize neighboring ant colonies by stealing their
larvae. The slave-making ants raise the stolen young ants to work in their colony. Many
other organisms parasitize ant colonies, including some very special butterflies! These
butterflies trick ants into thinking that the butterfly caterpillar is an ant larva, so that ants
will bring the caterpillar into the ant nest and feed the caterpillar as if it were an ant.
Because the butterfly caterpillar benefits by taking food that would otherwise be given to
the ants’ young, this is an example of a parasitic interaction [4].

Competition is another type of negative species interaction. Competition occurs when two
organisms fight for the same resource, such as food, territory, or mates. Ants are great
competitors, and even though they are small they can compete with organisms much
bigger than they are. Some ants in the savannahs of East Africa do not build underground
nests, but instead live in Vachellia trees. These trees are a popular food source for large
animals like elephants and giraffes, meaning that the ants must protect their homes from
being eaten. Who do you think would win in a competition between an ant and an
elephant? It is easy to think that an ant is no match for these large animals, but when an
elephant starts feeding on a tree where ants have made their home, the ants will respond
by ferociously attacking the elephant, even climbing inside the elephant’s trunk to bite
and sting it.

Interacciones positivas

Unlike predation, parasitism, and competition, some species interactions do not have
winners and losers, they have winners and winners. When both species benefit from an
interaction this is considered a positive species interaction, often called a mutualism. One
of the most amazing things about ants is that they have so many mutualisms with other
organisms. Some of these mutualisms involve other insects, like aphids. Aphids are small,
soft, and slow insects that feed on plant juices, and even though aphids might seem like
they would be easy and tasty prey for ants, most ants do not prey on aphids. Instead, the
ants guard and protect aphids from other predators, while feeding on the nutritious
sugary liquid that aphids excrete. Because the ants get food and the aphids get protection,
both species benefit from the interaction.

Ants also form mutualisms with plants, for example by protecting plants from herbivores
(like elephants), cleaning parasitic fungi off of plants’ leaves, or dispersing plant seeds. The
seeds of bloodroot plants have special structures called elaiosomes that ants like to eat.
When an ant finds a bloodroot seed it will take the seed back to the nest, remove the
elaiosome from the seed, and feed the elaiosome to the ant larvae. The ant will then
typically discard the seed underground, where it has a good chance of sprouting (Figure
3). By taking seeds back to the ant nest, the ants transport seeds far away from parent
plants, so when new seedlings sprout, they do not compete with their parent plants for
light or nutrients. This is beneficial for the plants as well as for the ants. Many plants rely
on ants to transport their seeds; in fact, seed dispersal by ants is so common it has its own
name, myrmechochory.

Other ants engage in mutualisms with microorganisms like fungi and bacteria. Leafcutter
ants live in jungles and are well-known for cutting up small pieces of leaves, which they
carry back to their nests (Figure 4). But leafcutter ants do not actually eat leaves. Instead,
they use the leaves to build underground compost piles on which they grow a special type
of fungus, and they feed this fungus to their young. The ants benefit from this interaction
because the fungus provides food for the colony, and the fungus benefits because the ants
bring it fresh leaves to grow on, keep it free from pests, and carry the fungus along when
they establish new colonies [5].

Whitaker M and Stolzmann B (2019) Species Interactions and Ants. Front. Young Minds.
7:53. doi: 10.3389/frym.2019.00053

Describir interacciones existentes (nombre, intensidad, tipo)

Discutir los efectos directos e indirectos de los interactuantes salud/éxito de la población

Elaborar esquema

Interacciones:
Mutualistas:
Hempiteran artrópodo mutualista a parásito
Mariposa artrópodo mutualista a parásito
Nectario extrafloral planta
elaiosome-based dispersal by ants
domatium-based symbioses
cosecha de plantas (plant farming)
cosecha de hongos attine-fungus farming
bacteria endosimbiosis y parásita
bacteria simbiosis intestinos gut
hongos simbiosis gut

Parasítica:
Interacciones entre especies hormiga/hormiga mutualista-parabiosis, parásita-parasitismo
social
Artrópodos parásito:
Molusco
Díptera
Araña
Escarabajo
Avispa
Pececillo de plata

Depredación:
Depredación de semillas planta

Domacio
estructuras huecas formadas por las plantas, que sirven de abrigo a pequeños animales
con los cuales viven en simbiosis.
A los domacios de hormigas se los denomina mirmecodomacios
Overall, nearly 700 plant species form symbioses with ants, offering nesting sites in
modified stems or leaves, in return for protection and/or nutrition

Ant/plant
P. nagasau workers cultivate multiplant colonies of Squamellaria epiphytes that can
contain 50 or more individuals whose plant-formed cavities in modified stems (domatia)
house one queen and ∼250,000 workers (11, 18). In this mutualism, the ants control
dispersal, fertilization, and defense of the epiphytes. They do this by actively
collecting Squamellaria seeds, planting them under the branch bark of their host tree, and
subsequently protecting both seedlings and adults from herbivory (11, 18). The ants also
fertilize these nutrient-limited (soilless) epiphytes by defecating on specialized and highly
absorptive warts within a plant’s tuberous domatium, which is also the ants’ obligatory
nest site (11, 18). In return, the ants feed on sugar- and amino acid-rich food rewards
produced by the flowers’ nectaries. The food rewards and the domatia, which contain a
complex network of interconnected cavities.

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