Acacia Collinsii
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''Vachellia collinsii'', previously ''Acacia collinsii'', is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
native to
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
and parts of
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
.


Distribution

''Vachellia collinsii'' is native to Central America and parts of Africa. In southern Central America, where there are seasonally dry ecosystems, this tree grows in
secondary succession Secondary succession is the secondary ecological succession of a plant's life. As opposed to the first, primary succession, secondary succession is a process started by an event (e.g. forest fire, harvesting, hurricane, etc.) that reduces an a ...
, preferring
Savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
-like
climate Climate is the long-term weather pattern in a region, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteoro ...
s. Acacias like full sun and are rarely found in the trophic
understory In forestry and ecology, understory (American English), or understorey (English in the Commonwealth of Nations, Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the Canopy (biology), forest ca ...
of many
jungle jungle is land covered with dense forest and tangled vegetation, usually in tropical climates. Application of the term has varied greatly during the past century. Etymology The word ''jungle'' originates from the Sanskrit word ''jaṅgala'' ...
s. Acacias can thrive in climates with higher humidity, above 70% humidity. The ''Vachellia collinsii'' has a wide distribution across the world compared to other acacias, as well as a wide ecological distribution, considering it can grow from
sea level Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...
to 1000 meters elevation.


Description

The ''Vachellia collinsii'' can grow upwards of 40 feet tall. The tree grows relatively straight with thorns generously distributed across the branches. The small, pinnate leaves grow opposite from each other similar to a
Mimosa ''Mimosa'' is a genus of about 600 species of herbs and shrubs, in the mimosoid clade of the legume family Fabaceae. Species are native to the Americas, from North Dakota to northern Argentina, and to eastern Africa (Tanzania, Mozambique, and ...
. Since this species has a diverse geological and ecological distribution, it has a wider range of morphological traits. Nonetheless, differences between other ant-acacias can be seen through elongated cylindrical
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
s, somewhat small stipular spines, terete—cylindrical—spines that wrap around a cross section, 3-5 often dome-shaped petiolar glands, a lack of
rachis In biology, a rachis (from the [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this case the ''rachi ...
glands, and leaflets with lateral
veins Veins () are blood vessels in the circulatory system of humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are those of the pulmonary and fetal c ...
. On the petioles, are green bumps called
extrafloral nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
that create sugar for
symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s. From top to bottom of these trees, there is less suppression of lateral growth, thus, allowing for more numerous and full branches near the base of the tree, creating a pyramid shape with a strong central trunk. While most tree species typically have
auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essent ...
that grow downward, suppressing branch growth on the sides, this is not the case for ''Vachellia Collinsii,'' as it lacks a strong
apical meristem In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of stem cells, known as meristematic cells, which are undifferentiated cells capable of continuous cellular division. These meristematic ce ...
. This results in branching throughout the entire length of the tree. They are extrafloral because the true, yellow flowers produce a different kind of nectar. The tips of leaflets may produce
Beltian bodies A Beltian body is a detachable tip found on the pinnules of some species of ''Acacia'' and closely related genera. Beltian bodies, named after Thomas Belt, are rich in lipids, sugars and proteins and often red in colour. They are believed to have ...
, food bodies which are rich in protein and lipids. These are also created as part of their symbiotic relationship with ants.


Ecology

''Vachellia collinsii'' exhibits a symbiotic relationship with several species of ants. Some noted species include Pseudomyrmex spinicola and
Pseudomyrmex ferruginea The acacia ant (''Pseudomyrmex ferruginea'') is a species of ant of the genus ''Pseudomyrmex''. These arboreal, wasp-like ants have an orange-brown body around 3 mm in length and very large eyes. The acacia ant is best known and named for ...
. The ant-Vachellia system involving this species has been studied by ecologists like Daniel Janzen in Palo Verde National Park and
Santa Rosa National Park Santa Rosa National Park (), is a national park, in Guanacaste Province, northwestern Costa Rica, it was created in 1966 by decree 3694. Geography The main entrance of Santa Rosa National Park is north of Liberia, Costa Rica, Liberia on National ...
, which are both located in
Guanacaste Province Guanacaste () is a Provinces of Costa Rica, province of Costa Rica located in the northwestern region of the country, along the coast of the Pacific Ocean. It is bordered by Nicaragua to the north, Alajuela Province to the east, and Puntarenas Pro ...
,
Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...
. The ants chew holes in the tips of the hollow stipular thorns, known as
domatia A domatium (plural: domatia, from the Latin "domus", meaning home) is a tiny chamber that houses arthropods, produced by a plant. Ideally domatia differ from galls in that they are produced by the plant rather than being induced by their inhabi ...
, so that they can enter and create their colony inside. A single ant colony may span several ''V. collinsii'' trees. Medium sized herbivores are often deterred by the thorns alone, but the ants help protect the trees from other potentially threatening animals. Smaller animals such as caterpillars have no issue avoiding the thorns, and larger animals like elephants are less affected by these thorns. When a predator brushes and shakes the plant’s thorns in an attempt to feed, the ants will become disturbed, run outside, and release alarm
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s to alert other ants. All ants that come in contact with the alarm pheromones become aggressive and attack the animal by biting and stinging. Beyond defending the trees from herbivores, ants occupying ''V. collinsii'' trees may even cut down surrounding vegetation and trim the encroaching branches of other plants. This provides ''V. collinsii'' with valuable space and unimpacted access to sunlight, allowing the trees to thrive. In exchange, ''V. collinsii'' not only provides the ants with hollow thorns in which to live, but also produces lipid- and protein-rich food bodies, known as
Beltian bodies A Beltian body is a detachable tip found on the pinnules of some species of ''Acacia'' and closely related genera. Beltian bodies, named after Thomas Belt, are rich in lipids, sugars and proteins and often red in colour. They are believed to have ...
, on the tips of new leaflets. These bodies are consumed by the ants, providing critical nutrients for the colony. When starved of nutrients, V. collinsii produces more Beltian bodies to encourage the presence of ants. When the tree has enough extra nutrients, it will create less Beltian bodies. This behaviour suggests a
feedback loop Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause and effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handle ...
. ''Vachellia collinsii'' also provides the ants with sugar-rich nectar from
extrafloral nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
located at the leaf petiole. Since there are several species of ants that may occupy a ''V. collinsii'', there has been observations of intraspecific interactions between these species of ants, especially between Pseudomyrmex spinicola and
Crematogaster brevispinosa ''Crematogaster brevispinosa'' is a species of ant in tribe Crematogastrini. It was described by Mayr Mayr is a German surname. Notable people with the surname include: * Andrea Mayr (born 1979), Austrian female long-distance runner * Ernst May ...
. C. brevispinosa may take over trees occupied by '' Pseudomyrmex spinicola'' or
Pseudomyrmex nigrocinctus
'. C. brevispinosa will also occupy trees that are dying or heavily damaged, or trees that were previously inhabited. Furthermore, a hypothesized benefit to hosting ant colonies is that the acacia may have receptors within the
domatia A domatium (plural: domatia, from the Latin "domus", meaning home) is a tiny chamber that houses arthropods, produced by a plant. Ideally domatia differ from galls in that they are produced by the plant rather than being induced by their inhabi ...
for ant feces that triggers absorption pathways for additional nutrient uptake at the extremities of the plant's stem tissue. When ants defecate in the domatia, their feces contain nutrients from their food which could be good for the plant. This allows the plant to obtain additional nutrients and in a shorter period of time as the nutrients do not need to travel all the way up starting from the roots.


Chemical Ecology

The symbiotic ants living on this tree become alarmed at tissue disruption of the plant’s leaves by herbivores like the scarab, ''Pelidnota punctulate''. This beetle only feeds on ant-acacias and is protected by its heavy cuticle. Tissue disruption of the tree leaves releases ''trans''-2-hexenal, a compound that ants detect as a kairomone. On exposure to test samples of this compound the ants become alarmed and displayed alarm behavior. This compound is not found in the mandibular gland secretions of Costa Rican acacia ants, that are likely to be the source of the ant’s alarm pheromones.


Development

Diving deeper into why the acacia collinsii produce traits of the “swollen thorn syndrome,” the mechanism and pathways are still unknown but there have been experiments and strong evidence related to a change in
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
expression of miR156/miR157 and SPL
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription (genetics), transcription of genetics, genetic information from DNA to messenger RNA, by binding t ...
s, in different environmental conditions. The production of food bodies that are high in proteins and lipids as well as extrafloral nectaries is very costly so the plant must have some indication of when to start producing those traits. Generally, an acacia will not produce these traits immediately after germinating so it is age dependent and the extrafloral nectaries will be produced first around 50–75 days of age, then swollen and elongated stipules, followed by full beltian bodies. Along with the production of these traits, are declines in miR156/miR157 genes. The miR156-SPL pathway has been known in many plants to coordinate when the plant flowers as well as plant development combined with stress tolerance. In an
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organ ...
, the miR156 will keep the plant juvenile, then become suppressed when it is in the right conditions in order to further develop adult traits. When put in low light conditions, there is higher miR156/miR157 as well as a delay in swollen thorn syndrome traits. The other way around is also true in that when put in well lit conditions such as the natural environment, there is a low expression of miR156/miR157 genes when the plant is producing extrafloral nectaries, swollen stipules, and beltian bodies. Although it is unknown how the expression of the traits are linked to the miR156/miR157 genes, hypotheses include temporal coordination or regulation where the miR156/157 turn on and off in a pattern in order to turn on those genes for the swollen thorn syndrome(which are still unknown). There is also evidence that these traits are part of their
defense mechanism In psychoanalytic theory, defence mechanisms are unconscious psychological processes that protect the self from anxiety-producing thoughts and feelings related to internal conflicts and external stressors. According to this theory, healthy ...
and that nectar secretion from the extrafloral nectaries depend on
jasmonic acid Jasmonic acid (JA) is an organic compound found in several plants including jasmine. The molecule is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway. It was first isolated ...
but the mechanism is unknown as jasmonic acid could have a different function here than in a typical plant. The developmental timing of the “Swollen Thorn Syndrome” can also be influenced by natural selection as well. More research needs to be done on what developmental constraints and factors that may have influenced the later development of these traits.


References

{{DEFAULTSORT:Vachellia collinsii collinsii Flora of Central America