Recent from talks
Contribute something
Nothing was collected or created yet.
Dorylus
View on Wikipedia
| Dorylus | |
|---|---|
| Dorylus gribodoi | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Hymenoptera |
| Family: | Formicidae Leach, 1815 |
| Subfamily: | Dorylinae Leach, 1815 |
| Genus: | Dorylus Fabricius, 1793 |
| Type species | |
| Vespa helvola | |
| Diversity[1] | |
| 61 species | |
| Synonyms | |
|
Cosmaecetes Spinola, 1851 | |
Dorylus, also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swahili,[2] and is one of numerous similar words from regional Bantu languages used by indigenous peoples to describe various species of these ants. Unlike the New World members of the former subfamily Ecitoninae (now Dorylinae), members of this genus form temporary subterranean bivouacs in underground cavities which they excavate and inhabit - either for a few days or up to three months. Also, unlike some New World army ants, driver ants are not specialized predators of other species of ant, instead being more generalistic with a diet consisting of a diversity of arthropods. Their colonies are enormous compared to other ant species, and can contain over 20 million individuals.[3] As with their American counterparts, workers exhibit caste polymorphism with the soldiers having particularly large heads that power their scissor-like mandibles. They are capable of stinging, but very rarely do so, relying instead on their powerful shearing jaws. A large part of their diet consists of earthworms. [4] Driver ant queens are the largest living ants known, with the largest measuring between 40 - 63 millimeters (1.5 - 2.4 inches) in total body length depending on their physiological condition.[5]
Life cycle
[edit]
Seasonally, when food supplies become short, they leave the hill and form marching columns of up to 20,000,000 ants, which constitute a considerable threat to humans, though they can be easily avoided as a column can only travel about 20 meters in an hour. It is for those unable to move, or when the columns pass through homes, that there is the greatest risk.[6] The presence of a mobile column of safari ants is, conversely, beneficial to certain human communities, such as the Maasai. They perform a pest prevention service in farming communities, consuming the majority of other crop-pests, from insects to large rats.[7] For example, driver ants prey on larvae of the African sugarcane borer, a pest moth in sub-Saharan Africa.
The characteristic long columns of ants will fiercely defend themselves against anything that attacks them.[4] Columns are arranged with the smaller ants being flanked by the larger soldier ants. These instinctively take up positions as sentries, and set a perimeter corridor through which the smaller ants can run safely. Their bite is severely painful, each soldier leaving two puncture wounds when removed. Removal is difficult, however, as their jaws are extremely strong, and one can pull a soldier ant in two without it releasing its hold. Large numbers of ants can kill small or immobilized animals and strip them to husks. Such is the strength of the ant's jaws that, in East Africa, they are used as natural emergency sutures. Various East African indigenous tribal peoples (e.g. the Maasai moran), when suffering from a laceration in the wilds, will use the soldiers to stitch the wound by getting the ants to bite on both sides of the gash, then breaking off the body. This use of ants as makeshift surgical staples creates a seal that can hold for days at a time, and the procedure can be repeated, if necessary, allowing for natural healing.[8] All Dorylus species are blind, and, like most varieties of ants, communicate primarily through pheromones.[4]
In the mating season, alates (winged drones, queens of driver-ant species do not grow wings) are formed. The drones are larger than the soldiers and the queens are even larger. Driver ants do not perform a nuptial flight, but mate on the ground, and the queens go off to establish new colonies. As with most ants, workers and soldiers are sterile females and, so, do not reproduce.[4]

Male driver ants, sometimes known as "sausage flies" (a term also applied to males of New World dorylines) due to their bloated, sausage-like abdomens, are among the largest ant morphs and were originally believed to be members of a different species. Males leave the colony soon after hatching but are drawn to the scent trail left by a column of siafu once they reach sexual maturity. When a colony of driver ants encounters a male, they tear his wings off and carry him back to the nest to be mated with a recently hatched queen. As in the majority of ant species, males die shortly afterward.[4] Driver ant queens exhibit polyandry; young queens from some species with large colony sizes must mate with 10–20 males before they have gathered enough sperm for their reproductive lives.[9] Once the queen is ready, roughly half of the workers in the colony will leave with her to found a new colony.[10] Driver ant queens are the largest ants on Earth and have the greatest egg-laying capacity among insects, laying several million eggs each month.[11]
Several species in this genus carry out raids on termitaria, paralyzing or killing termites and carting them back to the nest.[12]
Colonies of driver-ant species have only one queen.[13] When she dies, the surviving workers may try to join another colony, but in other cases, when two colonies of the same driver-ant species meet, they usually change the marching directions to avoid conflicts.[citation needed]
Species
[edit]
- D. acutus Santschi, 1937
- D. aethiopicus Emery, 1895
- D. affinis Shuckard, 1840
- D. agressor Santschi, 1923
- D. alluaudi Santschi, 1914
- D. atratus Smith, 1859
- D. atriceps Shuckard, 1840
- D. attenuatus Shuckard, 1840
- D. bequaerti Forel, 1913
- D. bishyiganus (Boven, 1972)
- D. braunsi Emery, 1895
- D. brevipennis Emery, 1895
- D. brevis Santschi, 1919
- D. buyssoni Santschi, 1910
- D. congolensis Santschi, 1910
- D. conradti Emery, 1895
- D. depilis Emery, 1895
- D. diadema Gerstaecker, 1859
- D. distinctus Santschi, 1910
- D. ductor Santschi, 1939
- D. emeryi Mayr, 1896
- D. erraticus (Smith, 1865)
- D. faurei Arnold, 1946
- D. fimbriatus (Shuckard, 1840)
- D. fulvus (Westwood, 1839)
- D. funereus Emery, 1895
- D. furcatus (Gerstaecker, 1872)
- D. fuscipennis (Emery, 1892)
- D. gaudens Santschi, 1919
- D. ghanensis Boven, 1975
- D. gribodoi Emery, 1892 – includes D. gerstaeckeri Emery, 1895
- D. helvolus (Linnaeus, 1764)
- D. katanensis Stitz, 1911
- D. kohli Wasmann, 1904
- D. labiatus Shuckard, 1840
- D. laevigatus (Smith, 1857)
- Dorylus lamottei (= D. gribodoi) Bernard, 1953
- D. leo Santschi, 1919
- D. mandibularis Mayr, 1896
- D. mayri Santschi, 1912
- D. moestus Emery, 1895
- D. molestus Wheeler, 1922
- D. montanus Santschi, 1910
- D. niarembensis (Boven, 1972)
- D. nigricans Illiger, 1802
- D. ocellatus (Stitz, 1910)
- D. orientalis Westwood, 1835
- D. politus Emery, 1901
- D. rufescens Santschi, 1915
- D. savagei Emery, 1895
- D. schoutedeni Santschi, 1923
- D. spininodis Emery, 1901
- D. stadelmanni Emery, 1895
- D. stanleyi Forel, 1909
- D. staudingeri Emery, 1895
- D. striatidens Santschi, 1910
- D. termitarius Wasmann, 1911
- D. titan Santschi, 1923
- D. vishnui Wheeler, 1913
- D. westwoodii (Shuckard, 1840)
- D. wilverthi Emery, 1899
See also
[edit]References
[edit]- ^ Bolton, B. (2014). "Dorylus". AntCat. Retrieved 17 July 2014.
- ^ Swahili translation Archived July 20, 2011, at the Wayback Machine
- ^ Silcock, Lisa, ed. (1992). The Rainforests - A Celebration. San Francisco: Chronicle Books. pp. 50 (caption) and 51 (photo). ISBN 0-8118-0155-1.
- ^ a b c d e Hölldobler, Bert; Wilson, Edward O. (1990). The Ants. Belknap Press of Harvard University Press. ISBN 0-674-04075-9.
- ^ Kronauer, Daniel (2020). Army Ants Nature's Ultimate Social Hunters. Cambridge, Massachusetts: Harvard University Press. p. 237. ISBN 978-0-674-24155-8.
- ^ "NOVA - Master of the Killer Ants - PBS". www.pbs.org.
- ^ Hastings, H.** Conling, D.E., Graham, D.Y.* & (1988-03-01). "Notes on the natural host surveys and laboratory rearing of Goniozus natalensis Gordh (Hymenoptera: Bethylidae), a parasitoid of Eldana saccharina Walker (Lepidoptera: Pyralidae) larvae from Cyperus papyrus L. in Southern Africa"[permanent dead link](PDF). Journal of the Entomological Society of Southern Africa
- ^ "From Ants to Staples: History and Ideas Concerning Suturing Techniques". ResearchGate.
- ^ Kronauer, Daniel J.C.; Johnson, Robert A.; Boomsma, Jacobus J. (February 23, 2007). "The Evolution of Multiple Mating in Army Ants". Evolution. 61 (2): 413–422. doi:10.1111/j.1558-5646.2007.00040.x. PMID 17348950.
The evolution of mating systems in eusocial Hymenoptera is constrained because females mate only during a brief period early in life, whereas inseminated queens and their stored sperm may live for decades…We show that queens of Neivamyrmex and Aenictus mate with the same high numbers of males (usually ca. 10–20) as do queens of army ant species with very large colony sizes…The species of Dorylus (Anomma) and Eciton where multiple queen mating has been documented (Kronauer 2004, 2006a, 2006b; Denny et al. 2004) have extreme degrees of worker caste polymorphism and larger colonies than almost any other social insect…
- ^ Kronauer, Daniel J.C. (September 11, 2009). "Recent advances in army ant biology (Hymenoptera: Formicidae)". Myrmecological News. 12: 51–65. Retrieved 2023-07-05.
The permanent lack of wings in army ant queens has one important corollary: they do not go on mating flights. Instead, young army ant queens mate inside their natal colony with foreign males that disperse on the wing…The reproductive colony undergoes fission during which the worker force splits into two roughly equal parts…
- ^ Kronauer, Daniel J.C.; Johnson, Robert A.; Boomsma, Jacobus J. (February 23, 2007). "The Evolution of Multiple Mating in Army Ants". Evolution. 61 (2): 413–422. doi:10.1111/j.1558-5646.2007.00040.x. PMID 17348950.
Being the largest ants on Earth, African Dorylus (Anomma) queens also hold the world record in reproductive potential among the insects, with an egg-laying capacity of several millions per month…
- ^ Biotropics Archived January 24, 2009, at the Wayback Machine
- ^ Boswell, Grame P.; Franks, Nigel R.; Britton, Nicholas (19 March 2001). "Arms races and the evolution of big fierce societies" (PDF). Proceedings of the Royal Society B: Biological Sciences. 268 (1477): 1723–1730. doi:10.1098/rspb.2001.1671. PMC 1088800. PMID 11506686. Retrieved 2023-07-05.
Here we consider the largest single-queen insect societies, those of the Old World army ant Dorylus, single colonies of which can have 20 million workers.
External links
[edit]Dorylus
View on GrokipediaTaxonomy and Classification
Etymology and Naming
The genus name Dorylus derives from the Greek word dory (δόρυ), meaning "spear" or "wooden shaft," alluding to the prominent, spear-like mandibles of the soldier caste used in aggressive foraging raids.[4] This nomenclature reflects the ants' formidable predatory apparatus, which enables them to overpower and dismember prey efficiently. The full etymological breakdown ties into the subfamily Dorylinae, where the root emphasizes the weaponized morphology central to their ecology.[4] Common names for Dorylus species vary regionally and highlight their behavioral traits. In English-speaking contexts, they are widely called "driver ants" due to their swarming raids that drive other animals and insects ahead of the column, a term popularized in 19th-century accounts describing their relentless pursuit of prey.[5] They are also known as "safari ants" or "army ants," evoking images of organized military expeditions across African landscapes. In East Africa, the Swahili term "siafu" is commonly used, originating from Bantu languages to denote these invasive, nomadic predators that disrupt local communities during migrations.[6] The genus was formally established by Danish entomologist Johan Christian Fabricius in 1793, based on the type species Vespa helvola Linnaeus, 1764, originally described from a male specimen collected in South Africa.[6] Early taxonomy was complicated by the dimorphism between castes and sexes, leading to fragmented descriptions; for instance, workers and queens were often classified separately from males until connections were clarified. Subsequent revisions, including subgeneric divisions into six groups (e.g., Anomma for epigaeic raiders), have refined the classification, though molecular phylogenetic analyses indicate these subgenera are not monophyletic; comprehensive overviews provided in monographic works addressing morphological and phylogenetic ambiguities.[7][1]Phylogenetic Position
Dorylus belongs to the subfamily Dorylinae within the ant family Formicidae, where it is the sole genus classified in the tribe Dorylini. This placement reflects the monophyly of Dorylinae, a group of predatory ants characterized by specialized legionary lifestyles, as supported by molecular phylogenetic analyses that resolve Dorylus as sister to the fossil genus Aenictogiton within the Old World doryline clade.[8][9] Dorylus shares a close evolutionary relationship with the New World army ants of the subfamily Ecitoninae, both exhibiting the army ant syndrome, including nomadic colony behavior and collective raiding. Fossil-calibrated phylogenies estimate the divergence between Old World dorylines (including Dorylus) and New World ecitonines at approximately 80–100 million years ago, aligning with the Late Cretaceous radiation of ants during the angiosperm diversification.[9][10] Key synapomorphies uniting Dorylinae, including Dorylus, encompass the permanent winglessness of queens (ergatoid morphology) and extreme worker polymorphism, which facilitate the division of labor in large, nomadic colonies. These traits, conserved across army ant lineages, underscore the evolutionary stasis of the legionary syndrome, as detailed in comprehensive phylogenetic reviews.[11]Physical Description
Morphology of Workers and Soldiers
Dorylus workers exhibit pronounced polymorphism, divided into distinct subcastes such as minors, medias, majors, and soldiers, which vary significantly in size and form to fulfill specialized roles within the colony. Minor and media workers typically measure 2–5 mm in length, featuring compact bodies with mandibles that are triangular in shape and equipped with multiple denticles for efficient cutting of prey tissues and transportation of small food particles.[8] In contrast, major workers and soldiers can reach lengths of 10–15 mm or more in certain species, such as Dorylus nigricans and Dorylus arcens, with disproportionately enlarged heads that accommodate powerful musculature.[12][8] The mandibles of soldiers are a hallmark adaptation, being oversized, sickle-shaped (falcate), and often featuring smooth inner margins with minimal or no dentition, optimized for seizing and slashing larger prey during raids or repelling threats.[1] These structures provide a mechanical advantage in predation and defense, reflecting evolutionary shifts correlated with foraging strategies across Dorylus subgenera.[1] Worker polymorphism in mandible size and shape—ranging from the finely toothed versions in minors for versatile manipulation to the robust, curved blades in soldiers—enhances colony efficiency in resource acquisition and protection.[13] Like other army ants, Dorylus workers and soldiers are blind, possessing no compound eyes and only rudimentary or absent ocelli in some individuals, an adaptation suited to their predominantly subterranean or low-light foraging environments.[13] Compensation for visual limitations occurs through enhanced antennal chemoreception, where elongated antennae in epigaeic species detect volatile pheromones over distances, guiding swarm raids and maintaining trail integrity.[1] This sensory reliance underscores the caste-specific morphological specializations that enable coordinated, mass-foraging behaviors without visual input.[13]Queen and Male Characteristics
The queens of Dorylus species, also known as driver ants, are among the largest insects in the world, measuring 40–63 mm in total body length, with their size varying by species and physiological state.[6] These queens are wingless after mating, a condition typical of the dichthadiiform morphology in the genus, where they develop a massively enlarged gaster adapted for egg production.[14] The head is bulbous, featuring poorly developed, falcate mandibles that are short and lack serrations or distinct teeth, reflecting their specialized reproductive role rather than foraging.[14] They are blind, lacking compound eyes and possessing only rudimentary or absent ocelli, which aligns with their subterranean or protected lifestyle within the colony.[14] Reproductively, Dorylus queens exhibit extreme fecundity, capable of laying 3–4 million eggs per month during peak periods, contributing to colony sizes exceeding 20 million individuals over their lifespan.[6] This high output is facilitated by the enlarged gaster, which stores vast quantities of sperm from multiple matings, as queens practice polyandry, inseminating with 10–20 males or more to ensure genetic diversity and colony robustness.[15] Unlike some ant genera, Dorylus lacks gamergates—reproductive workers—relying solely on the true queen for egg production, which underscores the genus's dependence on a single, highly specialized reproductive individual.[15] Males, referred to as alates or "sausageflies" due to their distinctive elongated abdomens, measure 25–35 mm in length and are winged for nuptial flights.[16] Their morphology includes a transverse head with large compound eyes containing thousands of ommatidia and prominent ocelli, enabling navigation during mating swarms, while their mandibles are exceptionally large and falcate, far exceeding those of queens in size and adapted for brief adult functions.[14] The abdomen is sausage-like and cylindrical, housing genitalia specialized for multiple inseminations, but males have a short lifespan, typically dying within days after mating once their reproductive role is fulfilled.[14]Distribution and Habitat
Geographic Range
The genus Dorylus, known for its army ant species, ranges from North Africa through sub-Saharan Africa to Asia Minor and tropical Asia, with the majority of species occurring in central, eastern, and southern African regions.[6][17] This distribution includes countries such as Morocco, Cameroon, the Democratic Republic of the Congo, Kenya, Tanzania, and South Africa, reflecting the genus's adaptation to diverse African tropical and subtropical environments.[18] The highest species diversity within Dorylus is concentrated in equatorial Africa, particularly in forested and savanna habitats of the Congo Basin and surrounding areas, where over 50 species have been recorded.[18] In contrast, the genus is entirely absent from the Americas, where analogous army ant roles are filled by New World taxa like Eciton, and from Europe, limiting its global presence to the Old World tropics. Extensions of the Dorylus range occur in tropical Asia, with several species distributed from India through Southeast Asia to Borneo, including notable occurrences in Bangladesh, Nepal, Thailand, and Indonesia.[6] These Asian populations, comprising fewer than ten species, represent peripheral extensions of the primarily African-centered genus.[19] The broad historical spread of Dorylus is hypothesized to trace back to Gondwanan origins, with phylogenetic evidence indicating divergence between African and Asian lineages following ancient continental fragmentation around 100-160 million years ago.[20] As of assessments through 2024, no significant range expansions or invasions by Dorylus species have been documented post-2020, maintaining the established distribution patterns.[21]Environmental Preferences
Dorylus species, commonly known as driver ants or army ants, exhibit a strong preference for tropical environments in Africa and parts of Asia, thriving in rainforests, savannas, and woodlands where vegetation provides ample cover and prey resources.[22] These habitats offer the humid, resource-rich conditions essential for their subterranean foraging and raiding activities, with colonies often establishing in areas of dense undergrowth or secondary forests disturbed by human activity.[23] They actively avoid extreme desert regions, which lack the moisture and arthropod abundance required for sustained colony survival, as well as high-altitude zones above approximately 3,000 meters where temperatures drop below optimal levels.[22][24] Colonies construct temporary bivouacs that are predominantly subterranean, excavating chambers in soil, leaf litter, or pre-existing cavities such as termite mounds, which provide protection from predators and environmental fluctuations.[2] While some species may occasionally utilize arboreal sites like tree hollows in forested areas, the majority favor underground structures to maintain humidity and stability.[6] This nomadic lifestyle is closely linked to resource availability, with colonies frequently emigrating—sometimes daily—to follow prey concentrations, ensuring access to food in dynamic tropical ecosystems.[25] Dorylus ants are adapted to a temperature range of 20–30°C, characteristic of their tropical niches, where ambient warmth supports metabolic rates and brood development without exceeding thermal tolerances.[26] They demonstrate sensitivity to drought conditions, responding by burrowing deeper into the soil to access moister layers and conserve water, a behavioral adjustment that helps mitigate the impacts of seasonal dry periods in savannas and woodlands.[27]Life Cycle
Reproduction and Colony Founding
The queens of Dorylus species exhibit extreme polyandry, mating with an exceptionally high number of males—up to 17 or more—during brief swarming events that facilitate dispersal and reproduction. These alate males, often called "sausage flies" due to their distended, sperm-filled abdomens, emerge in large swarms primarily at night to locate and inseminate virgin queens near colony sites.[15] Following mating, the queens store the collected sperm in their spermatheca for lifelong use, enabling continuous egg production without remating throughout their extended lifespan. This stored sperm supports the queen's prodigious reproductive output, with Dorylus wilverthi queens capable of laying up to 3–4 million eggs per month, primarily workers, in massive synchronized batches that sustain the colony's nomadic lifestyle.[28] The highly physogastric queens, which can weigh around 2 grams and measure up to 5 cm in length, achieve this rate through their enlarged ovaries adapted for bulk egg production.[29] Sexual brood, consisting of new queens and males, is produced less frequently, approximately every 2–3 years depending on species and environmental conditions, marking a shift in the colony's reproductive cycle.[5] Unlike many ant species with independent colony founding by single mated queens, Dorylus colonies reproduce exclusively through fission, where a mature colony divides into multiple independent units during the sexual brood emergence.[15] In this process, the old queen ceases laying eggs as the colony produces thousands of males and a small number of new virgin queens; after mating, one or more new queens lead daughter colonies, each inheriting a portion of the workforce—often hundreds of thousands of workers—bypassing a vulnerable single-foundress stage. This fission strategy ensures rapid establishment of robust colonies, with excess new queens typically cannibalized by workers to maintain monogyny in each unit.[5]Developmental Stages
The developmental stages of Dorylus ants encompass the standard holometabolous progression observed in Formicidae: egg, larva, pupa, and adult. Eggs, laid by the colony queen, undergo incubation for approximately 2-3 weeks under the regulated conditions of the bivouac, where workers maintain optimal humidity and temperature through clustering and ventilation behaviors.[5] Upon hatching, the resulting larvae are legless, eruciform grubs that depend entirely on worker care for survival. Larval development spans 3-4 weeks and involves multiple instars (typically four), during which workers feed the brood via trophallaxis—regurgitating predigested liquids and solids derived from raided prey. This nutritional provisioning is critical, as differential feeding determines caste fate: well-nourished larvae develop into soldiers or reproductives, while underfed ones become smaller workers, a polyphenic response driven by larval size at the onset of pupation.[30] Larvae grow rapidly, molting several times as they consume vast quantities of protein-rich food transported back to the colony. Pupation follows larval maturity, occurring without cocoons—a trait unique to Dorylus and certain other dorylines—directly within the protective bivouac structure formed by interlocked workers. The pupal stage lasts about 2-3 weeks, during which the immobile, exarate pupae undergo metamorphosis, with eclosion yielding callow adults that harden over several days before integrating into colony activities.[8] This naked pupation exposes the brood to greater risk but aligns with the colony's mobile lifestyle. Throughout these stages, Dorylus colonies exhibit asynchronous brood development, lacking the strict synchrony seen in some New World army ants, which allows continuous growth rather than phased bursts. New colonies formed via fission, each with a new queen and portion of the workforce, expand rapidly through successive brood cycles, potentially reaching up to 20 million individuals within months as worker production accelerates foraging capacity and resource intake.[31]Behavior and Social Structure
Foraging and Raiding Patterns
Dorylus colonies exhibit highly organized foraging strategies characterized by mass raids that enable efficient resource acquisition across diverse habitats. These raids typically form as dense columns of workers extending from the colony's temporary nest, allowing the ants to exploit prey in a coordinated manner. In epigaeic species, such as those in the subgenus Anomma, raids can manifest as expansive swarms that overrun surface areas, while hypogaeic species like Dorylus laevigatus rely on subterranean columnar trails for more targeted incursions.[32] Columnar raids in Dorylus can reach lengths of up to 100 meters, advancing at speeds of approximately 20 meters per hour, facilitating the coverage of large foraging grounds. These formations consist of thousands of workers, with minor workers scouting ahead and major workers recruited to subdue larger prey upon discovery. Swarm foraging complements this by deploying broader fronts, up to several meters wide and 3 to 8 meters long, lasting 2 to 3 hours and targeting clustered arthropods in leaf litter or soil. Such patterns allow colonies to harvest resources opportunistically without fixed nest-bound trails.[32] The diet of Dorylus primarily consists of invertebrates, including earthworms, termites, beetles, and caterpillars, particularly targeting social insects such as ants and termites. Dietary preferences vary; hypogaeic species often specialize on social insects like termites and ants, while epigaeic species exhibit more generalist predation. Raids occasionally extend to vertebrate carrion, such as small mammals or experimental baits like tuna, providing supplemental protein when available. This broad opportunistic predation underscores the ants' role as generalist apex invertebrates, preying on live arthropods encountered during advances rather than pursuing specific taxa.[32] During extended raids, Dorylus workers establish temporary bivouacs—living nests formed by interlocking bodies—to house the queen, brood, and resting foragers, often in soil cavities or elevated structures. These bivouacs, measuring up to 11 cm wide, serve as mobile hubs that shift with the raid's progression, ensuring colony cohesion. Pheromone trails laid by workers' gaster tips guide subsequent recruits along exploratory and trunk paths, with stable hypogaeic systems enabling persistent foraging efficiency over distances up to 4 meters deep.[32]Communication and Defense
Dorylus ants, like other army ants, rely heavily on chemical signaling through pheromones for internal coordination. Trail pheromones are deposited by scout workers to guide foraging columns to prey sources, enabling efficient mass recruitment during raids; this mechanism is particularly prominent in large-colony species such as Dorylus, where pheromones reinforce paths and facilitate collective navigation.[33] Alarm pheromones, released from mandibular or Dufour's glands, alert nestmates to threats by triggering rapid mobilization and aggressive responses, ensuring quick defensive postures across the colony.[2] In addition to pheromones, soldiers employ stridulation—rubbing a file-like structure on their abdomen against a scraper to produce vibrational signals—as a supplementary alert mechanism, which propagates through the colony substrate to summon reinforcements without visual cues.[34] Defensive strategies in Dorylus emphasize the role of specialized soldier castes, which form compact formations to protect the colony core during raids or emigrations. These soldiers, equipped with oversized heads and powerful, serrated mandibles capable of locking onto intruders, create interlocking barriers that deter predators by inflicting severe bites; such mandibular snaps provide both offensive and defensive capabilities, often resulting in the predator's withdrawal due to pain and injury.[2] Colony-level defense is further enhanced by nestmate recognition pheromones embedded in cuticular hydrocarbons, which allow workers to distinguish familiar colony members from outsiders, thereby avoiding costly inter-colony conflicts through evasion or selective aggression.[2] This chemical discrimination promotes spatial separation between colonies, minimizing direct confrontations in shared habitats. Given their blindness, Dorylus workers navigate raids and migrations using a combination of tactile and chemical cues, with ants maintaining contact via antennae and body linkages to follow pheromone trails in dense columns.[35] Tactile feedback from neighboring ants ensures column cohesion, while chemical gradients provide directional guidance, allowing the swarm to advance blindly yet purposefully over terrain. In response to predators such as birds that opportunistically feed on flushed prey during raids or mammals like aardvarks that excavate bivouacs, the colony mounts a coordinated counterattack; soldiers rush to the threat site, locking mandibles onto the intruder to immobilize it, often forcing retreat through sheer numbers and persistence.[36] These defenses highlight the superorganism-like integration of Dorylus colonies, where individual sacrifices bolster collective survival.Ecological Interactions
Role in Ecosystems
Dorylus ants serve as top invertebrate predators in African ecosystems, exerting significant control over pest populations through their aggressive foraging raids. For instance, species such as Dorylus helvolus prey on larvae of the sugarcane borer Eldana saccharina, a major agricultural pest in sub-Saharan Africa, with serological tests confirming predation rates contributing to 33% of ant-mediated attacks on this pest in South African sugarcane fields.[37] This predatory role helps regulate herbivorous insect densities, preventing outbreaks that could otherwise devastate crops and native vegetation. Their subterranean nesting and foraging behaviors also contribute to soil aeration and structure modification. Colonies actively excavate extensive underground tunnels and nests, displacing soil and enhancing soil porosity, water infiltration, and oxygen availability in tropical forest floors. Related hypogaeic Dorylus species, such as D. laevigatus, further support this by constructing stable trail systems at depths of 8-20 cm, promoting nutrient cycling through the decomposition of consumed prey and waste deposition.[23] Dorylus ants form a critical link in food webs as prey for larger vertebrates. Chimpanzees (Pan troglodytes) across Central and East African forests actively hunt these ants using tools like sticks to extract them from nests and raids, with rare observations in Ugandan sites like Budongo Forest documenting this behavior, though ant-feeding remains uncommon in their diet.[38] Similarly, various bird species, including ant-following specialists in Afrotropical rainforests, consume Dorylus individuals disturbed during swarm raids, integrating them into avian diets while also exploiting the raids to capture fleeing arthropods.[39] By curbing herbivore populations, Dorylus indirectly supports plant health and reproductive processes, including pollination, as reduced pest pressure allows for better flower development and pollinator access in disturbed habitats. Their raids can temporarily disrupt local arthropod biodiversity by decimating prey in raided areas, yet this predation promotes overall nutrient cycling by redistributing organic matter through colony waste and prey remains, fostering soil fertility in tropical ecosystems. Primarily native to sub-Saharan Africa and parts of southern and southeastern Asia, Dorylus species exhibit no invasive tendencies, maintaining balanced ecological dynamics without expanding beyond their natural range.[1]Relationships with Humans and Other Species
Dorylus ants provide several benefits to human communities in their native African range, particularly through their role in natural pest control. Among the Maasai people of Kenya and Tanzania, these ants are valued for preying on crop-damaging insects, helping to protect agricultural fields without the need for chemical interventions.[40] In broader contexts across tropical Africa, driver ants like those in the genus Dorylus consume vast numbers of pest species during their foraging raids, contributing to biological control efforts in ecosystems where they occur.[41] Additionally, indigenous groups have long utilized the powerful mandibles of Dorylus soldier ants as natural sutures in traditional medicine; the ant is positioned to bite the edges of a wound, and its body is then removed, leaving the head clamped in place to close the injury until healing occurs.[42] Despite these advantages, interactions between Dorylus and humans often involve significant conflicts due to the ants' aggressive raiding behavior. During swarm raids, which can move at speeds of 6–14 meters per hour, columns of ants may invade homes or camps, leading to painful encounters where soldiers bite exposed skin, leaving puncture wounds that cause intense pain from the ants' alkaloid venom.[40] Such invasions have resulted in severe medical cases, including anaphylaxis, hypotension, and ulcerated lesions, as documented in a Ugandan incident where a man sleeping outdoors suffered multiple bites leading to systemic shock.[43] Encounters with red Dorylus ants, such as D. helvolus, are particularly noted during hiking and trekking activities in Uganda, where they can be aggressive if disturbed, such as when stepped on, leading to painful bites and occasional stings, as mentioned in travel guides for gorilla trekking in areas like Bwindi Impenetrable Forest.[44][45] Cultural perceptions exacerbate these tensions, with Dorylus often mythologized as "killer ants" capable of overwhelming and consuming large animals or even humans, though such accounts are greatly exaggerated—healthy individuals can easily evade swarms, and fatalities are extremely rare, typically limited to immobilized victims like infants.[46] Interspecific relationships of Dorylus extend beyond predation to include avoidance strategies and opportunistic symbioses with non-prey species. Colonies of different Dorylus species or even conspecifics typically avoid direct encounters during raids to minimize costly intercolony battles.[47] A notable symbiosis occurs with various African bird species, which follow Dorylus raids—particularly those of D. wilverthi and D. molestus—as indicator events, foraging on insects flushed by the ants; up to 56 bird species, including specialized ant-followers like the fire-crested alethe (Alethe castanea), attend these swarms, relying on them for a significant portion of their diet in rainforest habitats.[48][49] However, no mutualistic relationships between Dorylus and other ant species or plants have been documented, with interactions largely limited to predation or evasion.[41]Species Diversity
Overview of Species
The genus Dorylus comprises over 60 recognized species.[11] These species are traditionally divided into subgenera such as Anomma and Dorylus sensu stricto, though a 2016 taxonomic revision synonymized several subgenera under the main genus due to non-monophyly in some cases.[11] Over 50 species are distributed across sub-Saharan Africa, with notable endemism in the Congo Basin, while fewer than 10 occur in Asia, mainly in the Indomalayan region of Southeast Asia.[11] Taxonomy remains challenging, as many species are known only from males or workers, complicating worker-male associations.Notable Species and Variations
Dorylus nigricans, commonly known as the siafu or driver ant, is renowned for hosting some of the largest colonies among army ants, with estimates reaching up to 20 million workers per colony.[50] These massive societies are primarily distributed across West and Central Africa, where the species exhibits highly aggressive raiding behavior, forming extensive swarms that overwhelm prey through sheer numbers.[51] The dry biomass of such colonies can range from 9 to 15 kg, underscoring their ecological dominance in tropical forest understories.[7] In contrast, Dorylus helvolus is a primarily African species distributed across sub-Saharan Africa, including Uganda, with additional occurrences in Southeast Asia, and notably smaller colony sizes compared to many African counterparts.[52] This species features workers with a distinctive yellowish to reddish coloration and a broad size polymorphism, ranging from small majors to larger soldiers equipped for subterranean foraging.[52] D. helvolus exhibits aggressive behavior, with workers capable of biting or stinging if disturbed, such as when stepped on during hikes in Ugandan forests.[53] While less prone to surface raids, D. helvolus maintains the nomadic lifestyle typical of Dorylus, adapting to diverse habitats from forests to agricultural edges.[54] Intraspecific variations within the genus include notable color morphs, spanning black to reddish hues depending on geographic populations, which may reflect adaptations to local environmental conditions.[55] Size differences also occur across populations, with some exhibiting larger workers in resource-rich areas, influencing foraging efficiency. Additionally, Dorylus gribodoi is prevalent in West African regions like Benin.[56]References
- https://www.antwiki.org/wiki/The_Ants_Chapter_16
- https://www.antwiki.org/wiki/Dorylus
- https://www.antwiki.org/wiki/Dorylus_laevigatus
- https://www.antwiki.org/wiki/Dorylus_species_by_Country
- https://www.antwiki.org/wiki/Dorylus_nigricans
- https://antwiki.org/wiki/Dorylus_helvolus
- https://www.antwiki.org/wiki/Dorylus_gribodoi
