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Crematogaster

Crematogaster is a genus of ants in the family Formicidae, subfamily Myrmicinae, and tribe Crematogastrini, comprising over 520 extant species and numerous subspecies worldwide.^[1] These ants are distinguished by their unique morphology, particularly the heart-shaped gaster (abdomen) formed by the postpetiole attaching to its dorsal surface, allowing workers to raise it over the thorax in a defensive posture—hence the common name "acrobat ants."^[2] The genus, first described by Johan Christian Lund in 1831 with Formica scutellaris (now C. scutellaris) as the type species, exhibits 11-segmented antennae in most species, a propodeum often armed with spines, and workers that are typically monomorphic or show gradual size variation.^[3] With a global distribution but highest diversity in tropical regions, Crematogaster species play significant ecological roles, particularly in arboreal habitats where they dominate forest canopies in the Neotropics, Africa, and Asia.^[2] Many species are arboreal nesters, hollowing out dead wood or twigs, while others occupy ground-level sites under stones or in soil; they are omnivorous foragers, preying on small arthropods, scavenging, and frequently tending hemipterans like aphids for honeydew.^[2] Some species form mutualistic associations with plants, such as in the Neotropics where they occupy domatia in myrmecophytes,^[4] or exhibit behaviors like temporary social parasitism during colony founding.^[2] The genus's evolutionary success is evident in its hyperdiversity, with ongoing taxonomic revisions revealing complex species groups and regional radiations, such as the 31 species documented in Costa Rica alone.^[2]

Taxonomy and Phylogeny

Classification

Crematogaster belongs to the subfamily Myrmicinae within the family Formicidae and is classified in the tribe Crematogastrini.^[5]^[6] The genus was first described by Danish entomologist Christian J. Lund in 1831, based on the type species Formica scutellaris (now Crematogaster scutellaris).^[6] Subsequent taxonomic work advanced understanding of its boundaries, with significant contributions in Hölldobler and Wilson's comprehensive 1990 monograph The Ants, which treated numerous proposed subgenera as junior synonyms while emphasizing the genus's ecological and morphological diversity. More recent revisions, such as Blaimer's 2012 subgeneric study, have refined this classification using molecular data to reassess phylogenetic relationships and synonymies. Phylogenetically, Crematogaster occupies a basal position within the diverse Myrmicinae subfamily, closely allied with other genera in the Crematogastrini tribe, such as Cophomyrma. Molecular phylogenies based on multilocus nuclear gene sequences indicate that the genus originated in Southeast Asia during the mid-Eocene (approximately 40–45 million years ago), with major diversification occurring through the late Oligocene and early Miocene, aligning with broader Paleogene radiations in ants driven by angiosperm proliferation.^[7] These studies highlight Crematogaster's exceptional dispersal capabilities, facilitating multiple colonization events across tropical and temperate regions. Subgeneric divisions have undergone substantial revision, reducing over a dozen historical subgenera to two primary ones: Crematogaster sensu stricto and Orthocrema. Crematogaster sensu stricto incorporates former groups like Decacrema, distinguished by traits such as a quadrate petiole with a posteriorly placed spiracle relative to the propodeal one and a postpetiole that broadly attaches to the gaster. Orthocrema encompasses other lineages, including Neocrema, characterized by a more elongate petiole and an anteriorly positioned propodeal spiracle. These divisions reflect monophyletic clades supported by morphological and genetic evidence, aiding in the genus's high species diversity worldwide.

Species Diversity and Subgenera

The genus Crematogaster encompasses more than 500 described species worldwide, representing one of the most species-rich ant genera in the family Formicidae.^[8] This diversity is particularly pronounced in tropical and subtropical regions, with the highest concentrations observed in the Indo-Australian and Neotropical realms, where these ants often dominate local arthropod communities.^[9] For instance, Asia alone accounts for approximately 145 valid species, underscoring the Indo-Australian hotspot, while the Neotropics harbor a significant portion of the genus's overall richness due to extensive adaptive radiations.^[10]^[11] Molecular phylogenetic studies have highlighted substantial undescribed diversity within Crematogaster, including numerous cryptic species that are morphologically indistinguishable but genetically distinct.^[12] These hidden lineages, often uncovered through multilocus DNA sequencing, suggest that the true species count could nearly double the current tally, especially in understudied tropical areas like Southeast Asia and the Americas.^[13]^[14] Such findings emphasize the challenges of traditional morphology-based taxonomy in this hyperdiverse group and the value of integrative approaches for revealing evolutionary patterns. A comprehensive subgeneric revision recognizes two primary subgenera: Orthocrema and Crematogaster sensu stricto, with numerous former subgenera now treated as synonyms.^[9] Orthocrema is diagnosed by a nodiform petiole (with a distinct node) and the postpetiole attached to the anterior face of the fourth abdominal tergite, encompassing species adapted to diverse Old World habitats; examples include C. (Orthocrema) viduata from Africa and various Malagasy endemics like C. (Orthocrema) tarava.^[15] In contrast, Crematogaster sensu stricto features a more variable petiole shape (often trapezoidal or rounded) and dorsal attachment of the postpetiole to the gaster, including former subgenera such as Oxygyne (noted for polymorphic queens suggestive of social parasitism) and Atopogyne (with variable propodeal spine development).^[9]^[16] Representative species in this subgenus include C. (Oxygyne) ranavalonae from Madagascar, which exhibits extreme queen polymorphism, and C. (Atopogyne) spp. with inconsistent spine morphology across populations.^[16]^[17] Conservation concerns for Crematogaster are limited but notable in certain cases, primarily driven by habitat destruction in tropical ecosystems. Crematogaster pilosa in the southeastern United States faces threats from habitat loss in coastal marshes due to development, highlighting the genus's vulnerability where species are restricted to fragile environments.^[18]

Description and Morphology

Physical Characteristics

Crematogaster ants are small to medium-sized, with workers typically measuring 2–5 mm in length and queens reaching up to 8 mm. Coloration is predominantly black or dark brown, though regional variations occur, such as reddish hues in some tropical species and lighter yellow tones in leaf-litter inhabitants.^[2]^[19] The genus features several distinctive morphological traits. The gaster is heart- or teardrop-shaped, attaching dorsally to the postpetiole, which enables elevation over the mesosoma. The propodeum bears a pair of dorsal spines, varying from short and thick to long and slender across species. The petiole is slender and node-less, often rectangular or triangular in outline with subtle sculpturing. Antennae comprise 11 segments, forming a club of 2–4 enlarged apical segments.^[3]^[2] Sexual dimorphism is pronounced, particularly in thoracic structure. Queens possess an enlarged mesosoma for flight musculature and prominent ocelli, distinguishing them from workers. Males are generally smaller, with relatively elongated antennal scapes and larger compound eyes relative to head size.^[2]

Behavioral Adaptations in Form

Crematogaster ants exhibit a distinctive defensive behavior known as the "acrobat" posture, where workers elevate their heart-shaped gaster over the thorax and head when threatened, allowing precise application of venom from the abdominal glands directly onto intruders. This elevation, often exceeding 90 degrees relative to the body axis, exposes the spatulate tip of the sting, facilitating topical smearing of the venom rather than injection, which serves as an effective threat display to deter predators and rivals. The behavior is triggered during interspecific encounters, with strong gaster flexions accompanying high aggression levels, enabling the ants to direct the secretion accurately while maintaining mobility.^[20] The tibial glands in the hind legs represent a key morphological adaptation for chemical communication in Crematogaster, consisting of class-3 exocrine structures with secretory cells and a reservoir embedded within the tibia, allowing for the production and storage of trail pheromones. These glands enable workers to lay persistent foraging trails by rubbing the hind legs against the substrate during movement, a necessity due to the ants' inability to contact the gaster tip with the ground owing to their compact form.^[21]^[22] Sensory adaptations in Crematogaster are exemplified by their compound eyes, which are relatively large for arboreal ants and optimized for detecting motion in dense foliage, aiding in predator avoidance and prey location. These eyes feature a high density of ommatidia, correlating with the foraging niche in canopy environments, where visual cues are crucial for navigation and orientation. In species like Crematogaster, eye area measurements show significant variation tied to arboreal habits, supporting behaviors such as directed aerial descent through visual targeting of landing sites.^[23]^[24]

Distribution and Habitat

Geographic Range

The genus Crematogaster exhibits a cosmopolitan distribution, with species present on all continents except Antarctica and the polar regions, primarily in tropical and subtropical zones.^[3] This widespread occurrence spans major biogeographic realms, including the Afrotropics, Neotropics, Indo-Malaya, and Australasia, where the ants have colonized diverse ecosystems through historical dispersals.^[25] While native to warmer climates, certain species have been introduced to temperate areas via human activities, expanding their range beyond original tropical strongholds.^[26] Regional hotspots for Crematogaster diversity and abundance are concentrated in the tropics, particularly the Afrotropical, Neotropical, and Indo-Malayan regions, which harbor the majority of the genus's over 520 described species.^[3]^[1] In contrast, representation is sparser in arid desert environments, where only a subset of tolerant species persist, reflecting the genus's preference for more mesic conditions.^[27] These patterns underscore the ants' adaptation to humid, vegetated landscapes over extreme xeric habitats. Phylogenetic analyses indicate that Crematogaster originated in Southeast Asia during the mid-Eocene, approximately 40–45 million years ago, with subsequent Miocene dispersals enabling colonization of Africa, the Americas, and other regions.^[25] Biogeographic patterns reveal significant endemism in island systems, notably Madagascar, where numerous species are restricted to the island's unique forests and exhibit high levels of local diversification.^[28] This endemism highlights Madagascar as a key center for Crematogaster radiation, driven by isolation and habitat heterogeneity.^[29]

Ecological Preferences

Crematogaster species primarily occupy arboreal habitats in forests, woodlands, and shrublands, where they often dominate the canopy ant fauna, though certain taxa exhibit ground-nesting behaviors in disturbed or arid landscapes. For instance, species such as C. laeviuscula and C. pinicola are characteristically arboreal, nesting in elevated vegetation, while C. lineolata and C. pilosa favor terrestrial sites in open or altered environments. This dual strategy allows adaptation to diverse ecological niches, from tropical rainforests to desert scrubs. Colonies preferentially select microhabitats offering protection and moisture retention, including tree hollows, rotten wood, dead branches, and accumulations of leaf litter. Shaded, humid sites are favored, as seen in C. ashmeadi nests within beetle galleries and logs in forested understories, or C. minutissima in hollow twigs amid moist litter. Ground-foragers like C. dentinodis utilize crevices under rocks or shrub roots in semi-arid zones, emphasizing sheltered, low-evaporation locales. Abiotic conditions significantly shape Crematogaster distributions, with optimal activity occurring at temperatures of 20–26°C, as documented for C. rogenhoferi in nest environments.^[30] Many species tolerate seasonal dryness through polydomous colony structures, distributing nests to mitigate desiccation risks. Elevational ranges vary, with lowland species below 500 m in wet climates and montane forms up to 1700 m in drier uplands. Biotic interactions, particularly co-occurrence with canopy vegetation, strongly influence nest site selection, as arboreal species like C. rifelna associate with live oaks (Quercus virginiana) for structural support and microclimate stability. Similarly, desert dwellers such as C. depilis nest near creosote bushes (Larrea tridentata), leveraging plant architecture for protection and proximity to resources. These associations underscore how vegetation structure guides habitat suitability across the genus.

Reproduction and Life Cycle

Mating and Colony Founding

In Crematogaster ants, reproduction typically begins with nuptial flights involving synchronous swarming of alate males and queens, often triggered by environmental cues such as rainfall in tropical and subtropical regions. For instance, in the Southeast Asian species C. captiosa, flights occur shortly after monsoon rains, with swarming ceasing during active precipitation but resuming on dry days following rain, facilitating mid-air mating between winged sexuals.^[31] In temperate species like C. ashmeadi, these flights are concentrated in early summer (June–July), depleting sexual brood from natal colonies by late July, while C. cerasi exhibits flights in July and August.^[32]^[33] Such synchronization enhances mating success by concentrating dispersers, though timing varies by species and locale, often aligning with the onset of rainy seasons to optimize dispersal.^[34] Following mating, dealate queens initiate colony founding primarily through independent claustral strategies in most species, where the queen seals herself in a nest site—such as abandoned beetle galleries in dead wood for C. ashmeadi—and rears the first brood using her body reserves without external foraging.^[32]^[2] However, some groups, like the acuta-group (e.g., C. montezumia), employ temporary social parasitism, with queens infiltrating host nests of related species to exploit heterospecific workers for initial brood care before assuming control.^[2] In C. scutellaris, queens may co-found colonies pleometrotically in available refugia like tree galls when nest sites are scarce, though this typically results in only one survivor dominating the mature colony. Fertilization in Crematogaster queens generally involves single mating during the nuptial flight, with spermatozoa transferred via spermatophore to the bursa copulatrix and subsequently stored immotile in the spermatheca for lifelong use, enabling egg production over a decade or more without remating.^[35] Multiple mating is rare across the genus but occurs occasionally; for example, in C. smithi, genetic analyses reveal an effective mating frequency of about 1.14, with polyandry detected in roughly 22% of colonies, while C. osakensis queens show evidence of mating with multiple males based on sperm stores and worker relatedness.^[36] Stored sperm remains viable for at least five years in C. osakensis, supporting continuous fertilization of diploid female eggs.^[35] During initial colony growth, the founding queen lays a mix of trophic (unviable) and viable eggs to nourish emerging larvae, with trophic egg production peaking around 15 days post-founding in C. ashmeadi.^[32] The claustral phase lasts 40–50 days, during which the queen loses approximately 50% of her dry weight to produce the first workers (nanitics), after which foraging begins and the colony expands.^[32] This solitary provisioning ensures the transition to a functional workforce, with initial queen weight predicting total progeny biomass.^[32]

Development Stages

The development of Crematogaster ants follows the standard holometabolous life cycle of ants, encompassing egg, larval, pupal, and adult stages, with total immature development varying from about 4 to 8 weeks depending on species, temperature, and conditions—for example, approximately 40–50 days in C. ashmeadi at 27 °C.^[32]^[3] Eggs are laid individually by the queen, primarily fertilized to produce female offspring (workers or new queens) or unfertilized to produce males; in queenless colonies, workers can also lay unfertilized eggs that develop into males. The egg stage lasts several days, during which embryonic development occurs, influenced by temperature and humidity.^[3] The larval stage consists of multiple instars and lasts several weeks, with the legless, C-shaped larvae growing rapidly while dependent on nurse workers for protection and sustenance. Larvae are nourished through trophallaxis, receiving regurgitated liquid food from adults, and caste fate is determined by nutritional input—larvae receiving abundant, high-quality provisions develop into reproductive castes or larger workers, while those fed sparingly become smaller workers.^[19] A short prepupal phase precedes pupation, during which the mature larva spins a silken cocoon for protection. The pupal stage, enclosed within this cocoon, involves internal reorganization into the adult form; the eclosing adult then chews its way out of the cocoon to emerge.^[37] Once adults, Crematogaster workers typically live 3 to 6 months, performing colony tasks until worn out, whereas queens endure 10 to 15 years, continuously producing eggs to sustain the colony.^[32]

Division of Labor

In Crematogaster colonies, workers are typically monomorphic, performing a range of tasks including brood care, nest maintenance, foraging, and defense, with specialization often influenced by age polyethism and subtle size variations.^[2] In certain species, such as those in the subgenus Orthocrema (e.g., C. smithi), workers exhibit dimorphism, with smaller workers focusing on nursing tasks and larger workers functioning as foragers, soldiers, and producers of trophic eggs—unfertilized eggs that serve as a protein-rich food source for the colony, storing nutrients from perishable prey for extended periods.^[38]^[39] Workers also display age-based polyethism, whereby younger individuals focus on internal tasks like tending larvae and older ones shift to external activities such as foraging, a pattern common across ant species including Crematogaster.^[40] The queen's primary role is egg production to sustain colony growth, with little to no participation in foraging or other labor once the nest is established; in species like C. cerasi, she mates once during nuptial flights, stores sperm, and lays eggs throughout her lifespan.^[41] Males, in contrast, are short-lived with a lifespan of a few weeks to a month, dedicated solely to mating during reproductive flights before dying shortly thereafter.^[41] Task allocation among workers is dynamically influenced by colony needs, such as resource availability or threats, and social interactions including antennal contacts that facilitate behavioral coordination; this system promotes flexibility, especially in smaller colonies where individuals often perform multiple roles to meet demands.^[42] Colony size variations can modulate the extent of such specialization, with larger nests supporting more distinct task divisions.^[43]

Colony Structure

Crematogaster colonies vary widely in size across species, typically ranging from 100 to 10,000 workers, though some can exceed 28,000 individuals in exceptional cases. Larger colonies are frequently polydomous, comprising multiple interconnected nests that enhance resource exploitation and colony resilience, particularly in resource-rich environments like tropical forests. This polydomous structure allows for spatial expansion, with nests often linked by foraging trails spanning several meters.^[44]^[45]^[3] Nest architecture in Crematogaster is predominantly arboreal, with many species constructing carton nests from masticated plant fibers mixed with soil or fungal material, forming a durable, waterproof structure. These nests exhibit a distinctive pagoda-like or spheroidal form, composed of overlapping concavo-convex chambers arranged in a conspheroidal pattern, where the concavity of each chamber faces the nest's center for internal protection. Chambers, typically 2–9 cm in diameter and 0.3–2.6 cm thick, interconnect via small passages, providing shelter for brood and workers while perched on twigs or branches 1–10 m above ground. This design optimizes defense against predators and environmental stressors like heavy rainfall.^[46] Mature Crematogaster colonies are often polygynous, housing multiple queens per nest—averaging around four in some species—which contributes to sustained reproduction and colony growth. Queens exert dominance through chemical signals, likely pheromones, that render them highly attractive to workers, prompting retinue formation and defensive behaviors such as venom spraying in response to threats. In queenless conditions, workers enforce social control via policing, aggressively eliminating eggs laid by reproductive workers to maintain colony stability and favor queen-produced offspring.^[45]^[47]^[48] Colony founding typically begins monogynously, with a single mated queen establishing an independent nest through claustral founding, where she rears the first workers without external aid. As colonies mature, they transition to polygyny and polydomy via budding, in which queens and worker groups relocate to establish satellite nests, often seasonally during resource peaks like the rainy season. This dynamic shift supports long-term colony persistence and expansion.^[45]

Foraging and Predation

Feeding Strategies

Crematogaster ants exhibit an omnivorous diet, incorporating a variety of food sources such as live insects, honeydew from hemipterans, fungi associated with wood, and opportunistic scavenging of dead arthropods.^[49]^[50] This generalist approach allows them to exploit diverse resources in their habitats, with foraging workers collecting both solid particles like plant material and prey (approximately 72.5% plant-based and 27.5% animal-based in some species) and liquid rewards such as sugary exudates, which are preferred over fats or proteins.^[51] Food sharing within Crematogaster colonies occurs primarily through trophallaxis, a mouth-to-mouth transfer of regurgitated liquids that distributes nutrients among workers, the queen, and brood.^[52] This behavior facilitates efficient circulation of carbohydrates and other resources, enabling even non-foraging colony members to access food collected externally.^[53] For storage, Crematogaster workers utilize an expandable crop to transport and temporarily hold liquid foods like honeydew, allowing repletes—distended individuals—to act as living reservoirs for the colony during periods of scarcity.^[54] While some ant species construct nest granaries for solids, evidence for this in Crematogaster is limited, with reliance more on individual crop capacity than communal storage structures. Feeding preferences in Crematogaster shift seasonally, influenced by resource availability and environmental factors like water scarcity; in wetter spring periods, colonies prioritize carbohydrate-rich sources such as sucrose solutions or nectar-like exudates, while drier summer conditions lead to increased uptake of proteinaceous foods, reflecting higher predation to meet colony protein demands.^[55] These adjustments ensure nutritional balance, with water limitation in dry seasons broadening the diet to include lower-value carbohydrates previously rejected.^[56]

Predatory Interactions

Crematogaster species typically employ group ambushes to capture arthropod prey, with workers detecting targets through contact and rapidly recruiting nestmates via short-range pheromones for collective foraging. In arboreal environments, such as tropical forests, foraging parties of up to 15 workers surround prey within 5–10 mm, seizing small arthropods by the body and larger ones by a leg to immobilize them. Venom is injected or applied topically via the ants' characteristic spatulated stinger, often from the Dufour gland, causing rapid paralysis; in species like C. striatula, this venom is emitted as a volatile vapor, allowing chemical immobilization without physical contact and causing termites such as Macrotermes bellicosus workers and soldiers to fall and become immobilized within 10 minutes.^[57]^[58] Prey primarily consists of small insects, including termites and wasps, with Crematogaster laeviuscula known to raid entire nests of Polistes exclamans, consuming eggs, larvae, and pupae to cause complete colony failure. These ants also engage in kleptoparasitism, as seen in C. limata parabiotica, which conducts nocturnal raids on Ectatomma tuberculatum nests in French Guiana, intercepting 75.2% of returning foragers and stealing their loads of sugary liquid by licking, thereby complementing their diet without direct hunting.^[59]^[60] These predatory tactics exhibit high efficiency in arboreal settings, where well-developed arolia on the ants' pretarsi aid in spread-eagling and transporting oversized prey, enabling capture of items far larger than a single worker's capacity. Chemical immobilization enhances success rates, repelling competitors and ensuring quick subdual even of agile arthropods. Notably, C. scutellaris in Mediterranean olive groves demonstrates a dual role by preying on soft scale insects (Coccidae) at high rates—significantly higher on occupied trees—while also tending them for honeydew, indirectly disrupting host plant populations and natural enemy dynamics through selective predation.^[57]^[58]^[61]

Defensive and Communication Behaviors

Defense Mechanisms

Crematogaster ants employ a unique chemical defense strategy involving the topical application of venomous secretions rather than spraying, facilitated by their characteristic spatulate sting. This reduced sting apparatus, located at the tip of the gaster, allows workers to raise their abdomen forward over the head and apply a droplet of toxic fluid directly onto the integument of intruders or predators. The secretions, primarily derived from an enlarged Dufour's gland, contain bioactive compounds such as long-chain aldehydes and acetates that act as contact poisons, deterring attackers through irritation, toxicity, or repellency. In addition to chemical application, workers bite with their mandibles to grasp and injure threats, often combining this with the venom smear for enhanced effect.^[62] Physical displays play a key role in individual defense, with workers frequently raising their heart-shaped gaster high above the body in a characteristic "acrobat" posture, which positions the sting for precise targeting and serves as a visual warning to potential predators. This elevated gaster orientation also enables the release of alarm pheromones from the sting or mandibular glands, signaling danger to nearby nestmates. Stridulation, produced by rubbing the file-like pars stridens on the gaster against a scraper on the petiole, generates vibrational signals that amplify alarm communication, particularly in species like Crematogaster scutellaris, where the intensity and pattern of stridulations vary by threat level to coordinate responses.^[63] Collectively, Crematogaster colonies mount defenses through rapid recruitment of nestmates using alarm pheromones, such as 4-methyl-3-heptanone from the Dufour's gland, which mobilizes workers to swarm and overwhelm intruders despite the lack of distinct soldier castes. In response to threats, ants may seal nest entrances with carton material constructed from masticated wood fibers, preventing access by larger predators or rival colonies, a behavior observed in arboreal species nesting in tree cavities. This cooperative sealing enhances nest fortification, particularly in exposed arboreal habitats.^[64] Antipredator adaptations in Crematogaster emphasize evasion and concealment, with many species exhibiting camouflage by nesting within tree bark or dead wood, where their dark coloration and small size blend seamlessly with the substrate to avoid detection by visually hunting predators.

Trail-Laying and Pheromones

Crematogaster ants utilize trail pheromones to establish and maintain foraging paths, primarily secreted from specialized tibial glands in their hind legs, a morphological adaptation unique to this genus. These glands produce volatile compounds that workers deposit by dragging their legs along substrates, creating chemical guides that direct nestmates to food resources with high efficiency. In Crematogaster scutellaris, for instance, the primary trail pheromone is (R)-(-)-tridecan-2-ol, an alcohol that elicits robust following behavior at concentrations as low as 0.001 ng/µL, while the Dufour's gland contributes additional components in some species for trail reinforcement.^[21]^[65]^[66] Trail formation in Crematogaster involves the continuous deposition of these pheromones, resulting in persistent paths that allow sustained access to ephemeral resources until depletion. In intricate environments like tree canopies, trails frequently branch to facilitate exploration of multiple routes, optimizing collective foraging efforts without redundant overlap. This branching pattern supports adaptive navigation in three-dimensional spaces, where physical constraints such as branch overlaps dictate trail topology.^[67] Alarm pheromones enable rapid recruitment and defensive coordination in Crematogaster, often released from the head or Dufour's gland upon disturbance. Representative components include octan-3-one as the major alarm signal in Crematogaster peringueyi, triggering agitation and attack responses among nearby workers. In species like Crematogaster castanea, 3-octanol functions similarly, with its enantiomeric composition influencing the intensity of the behavioral response. These pheromones briefly integrate with broader defensive behaviors by alerting the colony to threats, promoting swift mobilization.^[68]^[69] Beyond foraging and alarm functions, Crematogaster employs other pheromonal signals for colony regulation. Queen pheromones, characterized by distinct chemical profiles in species such as Crematogaster smithi, are involved in caste signaling and maintenance of social structure. Cuticular hydrocarbons coating the exoskeleton serve as primary cues for nestmate recognition, with colony-specific blends allowing precise discrimination of familiar individuals from non-nestmates, thereby preventing infiltration.^[70]^[71]

Ecological Role

Mutualistic Associations

Crematogaster species engage in mutualistic associations with honeydew-producing hemipterans, such as aphids and scale insects, where ants tend these insects in exchange for a carbohydrate-rich food source. Workers actively solicit and collect honeydew, a sugary excretion produced by the hemipterans as they feed on plant sap, while providing protection against predators and parasites. This tending behavior enhances hemipteran survival and reproduction, as ants remove competitors and deter natural enemies like lady beetles. For instance, Crematogaster brevispinosa tends Planococcus citri mealybugs on orchids, leading to increased mealybug populations but also heightened plant damage due to facilitated sap-feeding.^[72] Similarly, Crematogaster mimosae tends scale insects on Acacia trees, using the honeydew as a key nutritional supplement alongside extrafloral nectar.^[73] In plant-ant mutualisms, certain Crematogaster species form obligate symbioses with myrmecophytic trees, particularly in African savannas. C. mimosae occupies domatia—hollow, swollen thorns provided by host trees like Acacia drepanolobium and A. zanzibarica—which serve as secure nesting sites for colonies. In return, the ants aggressively defend the trees from herbivores, including mammalian browsers such as elephants and goats, by swarming and biting intruders in response to plant-borne vibrations signaling damage. This protection significantly reduces browsing damage; for example, trees hosting C. mimosae with associated scale insects experience 2.5 times less elephant damage over 10 months compared to those without. The trees supply year-round carbohydrates through extrafloral nectaries, supporting ant colony growth and defensive vigor.^[74]^[73] Fungal farming in Crematogaster is rare and distinct from the nutritional cultivation seen in other ant clades like the Attini. Crematogaster clariventris, an arboreal species in Central Africa, cuts small pieces of young leaves and flowers to inoculate and fertilize fungal mycelia (primarily Capnodiales) within its carton nests, creating a reinforced composite material that withstands heavy rainfall. This structural mutualism enhances nest durability without providing direct food benefits to the ants, representing a convergent evolution with fungus-growing ants but focused on architectural strength rather than provisioning. Additionally, in ant-plant systems, C. mimosae and C. nigriceps maintain distinct fungal communities within domatia, including Chaetothyriales that offer antibacterial protection and aid in nest maintenance, with alates transmitting these fungi during colony founding.^[75]^[76]^[77] These associations provide Crematogaster with essential nutrition from honeydew and nectar, stable nest sites via domatia, and structural support from fungi, while partners gain defense against herbivores, predators, and environmental stressors.^[72]^[74]^[75]

Interactions with Other Species

Crematogaster ants engage in intense competition with other ant species for resources such as nesting sites and food, often leading to territorial disputes and exclusion of rivals. In African savannas, species like Crematogaster mimosae and Crematogaster sjostedti compete aggressively with congeners and Tetraponera ants for exclusive possession of myrmecophyte trees, such as whistling thorn (Vachellia drepanolobium), where dominant colonies prune neighboring vegetation to deter intruders and secure nectar resources.^[78]^[79] In North American contexts, Crematogaster quadriformis exhibits rapid foraging and holds its own in direct confrontations with the invasive red imported fire ant (Solenopsis invicta), spraying venom to defend food baits and occasionally displacing the competitor.^[80] This resource exclusion extends to broader interspecific rivalries, where invading ants like the big-headed ant (Pheidole megacephala) outcompete Crematogaster species by disrupting their tree-based territories, reducing native ant densities by up to 90% in affected ecosystems.^[81] Predators target Crematogaster colonies at various life stages, exploiting their arboreal and ground-nesting habits. Birds such as woodpeckers and thrushes forage on worker ants and brood from exposed nests, while spiders, including orb-weavers and jumping spiders, ambush foraging Crematogaster individuals on vegetation. Parasitic organisms exploit Crematogaster colonies through direct infestation and social manipulation. Phorid flies (Pseudacteon spp.) are key parasitoids, with females ovipositing on foraging workers; the resulting larvae decapitate the host ant, emerging to pupate and reducing colony foraging efficiency in affected populations.^[41] Nematodes, including species like Diploscapter sp., are associated with Crematogaster workers and queens in tropical ant-plant associations, such as those with Macaranga trees, where they occur in the ants' habitats and may impair reproduction and longevity.^[82] Social parasites employ inquilinism, where inquiline species like certain chalcid wasps (Myrmokata sp.) integrate into Crematogaster colonies as permanent guests, relying on host workers for brood care while producing their own offspring that compete for resources.^[83] Crematogaster species pose conflicts with humans primarily through structural damage and agricultural impacts. In urban and suburban settings, acrobat ants (Crematogaster spp.) nest in moisture-damaged wood, such as roof timbers and wall voids, excavating galleries that weaken already compromised structures, though they do not consume sound wood like termites. In agriculture, species like Crematogaster cerasi exacerbate pest issues by tending honeydew-producing aphids and scale insects on crops, indirectly promoting plant damage and reducing yields in orchards and fields.^[37]^[84]

References

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Dec 1, 2023 · The genus Crematogaster Lund, 1831, includes 521 extant species, three fossil species, and 260 valid subspecies and is among the most ...
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Crematogaster ; Family: Formicidae ; Subfamily: Myrmicinae ; Tribe: Crematogastrini ; Alliance: Crematogaster genus group.Identification · Biology · Morphology · Nomenclature
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Crematogastrini - AntWiki
Aug 5, 2025 · Crematogastrini. Scientific classification. Kingdom: Animalia. Phylum: Arthropoda. Class: Insecta. Order: Hymenoptera. Family: Formicidae.
[6]
Crematogaster - AntCat
Taxonomic revision of the obligate plant-ants of the genus Crematogaster Lund (Hymenoptera: Formicidae: Myrmicinae), associated with Macaranga Thouars ...
[7]
https://doi.org/10.1016/j.ympev.2012.06.028
[8]
Multilocus phylogeny and historical biogeography of the ...
Jun 13, 2023 · The ant genus Crematogaster is a hyperdiverse group containing more than 500 species (Bolton, 2022) and has historically been subdivided ...
[9]
A subgeneric revision of Crematogaster and discussion of regional ...
Sep 13, 2012 · Crematogaster ants are diverse, widespread and abundant in tropical, subtropical and warm-temperate climatesthroughout the world. The species ...Missing: sources | Show results with:sources
[10]
(PDF) A check list of the ant genus Crematogaster in Asia ...
Jul 15, 2019 · A total of 206 names, comprising 145 species and 61 subspecies, is recognized. The list also provides information on the distribution.
[11]
Systematics and diversity of Neotropical ants
The Neotropical region has 137 genera and around 3,100 species of ants. The Neotropics seem to have been the “cradle” and “museum” of the groups of ants, which ...Missing: Indo- | Show results with:Indo-
[12]
Molecular phylogeny of Crematogaster subgenus Decacrema ants ...
To elucidate the evolution of one of the most species-rich ant–plant symbiotic systems, the association between Crematogaster (Myrmicinae) and Macaranga ...Missing: Paleogene | Show results with:Paleogene
[13]
Cryptic Diversity in the Widespread Asian Ant Crematogaster ...
The myrmicine species Crematogaster rothneyi is one of the most widely distributed ants in Asia, but it has rarely been collected in the field.Missing: undescribed | Show results with:undescribed
[14]
Integrative taxonomy reveals cryptic diversity in North American ...
Apr 8, 2022 · This case emphasizes the critical role of integrative taxonomy to detect cryptic species and identify potential biological invasions in their nascent stages.<|separator|>
[15]
Taxonomy and species-groups of the subgenus Crematogaster ...
Jun 4, 2012 · Species descriptions, images, distribution maps and identification keys based on worker and queen ants are given for all six species. A ...<|control11|><|separator|>
[16]
Untangling complex morphological variation: taxonomic revision of ...
Nov 28, 2011 · The subgenus Crematogaster (Oxygyne) shows an Old World distribution and comprises a group of ants with specialised queen morphology ...
[17]
Crematogaster (subgenus) - AntWiki
Aug 17, 2025 · Hosoishi, S. 2015. Revision of the Crematogaster ranavalonae-group in Asia, with description of two new species (Hymenoptera, Formicidae). ...
[18]
Acrobat Ant, Crematogaster ashmeadi Emery (Insecta: Hymenoptera ...
Crematogaster ashmeadi are arboreal ants, nesting in trees and rotten wood. Acrobat ants are the most dominant arboreal ant species in north Florida coastal ...
[19]
Defensive behaviour and biological activities of the abdominal ...
Aug 5, 2025 · ... Crematogaster is one of the largest ant genera, known for their raised gaster. Crematogaster species apply their venom to enemies by ...Missing: threat display
[20]
Trail pheromone identification in the ant Crematogaster scutellaris
Apr 3, 2024 · This is the case of the myrmicine genus Crematogaster, whose trail pheromone is released by the tibial glands. The genus Crematogaster ...
[21]
Occurrence and structural organization of the exocrine glands in the ...
The first exocrine gland discovered in ant legs was the tibial gland in the hindleg of Crematogaster (Leuthold, 1968). For more than two decades, however, this ...
[22]
[PDF] Vision-Linked Traits Associated With Antenna Size and Foraging ...
Sep 7, 2021 · Apposition compound eyes are com- posed of ommatidia which appear on the surface of the eye as a facet. (Cronin et al. 2014, Land and Nilsson ...
[23]
None
Nothing is retrieved...<|control11|><|separator|>
[24]
Acrobat ants go global--origin, evolution and systematics ... - PubMed
The 15 previous subgenera within Crematogaster are mostly not monophyletic. Divergence dating analyses and ancestral range reconstructions suggest that ...Missing: list | Show results with:list
[25]
Review of the Arabian Crematogaster Lund (Hymenoptera ...
Distribution map of C. antaris. Remarks. This is a very widespread and common species, which appears to be one of the most arid ...Missing: sparse | Show results with:sparse
[26]
Malagasy Region Species List - AntWiki
Aug 5, 2025 · Crematogaster agnetis Forel, 1892 (Madagascar) Endemic to this region; Crematogaster alafara Blaimer, 2013 (Madagascar) Endemic to this region ...
[27]
Journal of Biogeography | Wiley Online Library
May 16, 2021 · Centers of ant endemism are located in western dry and humid forests of northern Madagascar, eastern humid forests, and in the southern Central ...
[28]
[PDF] Determination of structural features of the nest material of ...
Dec 13, 2017 · Table 3: Differences in ambient and internal temperature ranges of Crematogaster dives nests (season) ... The optimum temperature is ±200 C to ± ...
[29]
the ant Crematogaster torosa preferentially forms new nests near ...
Nov 12, 2010 · We investigated whether colonies of Crematogaster torosa form new polydomous nests to better exploit temporally stable food resources.<|control11|><|separator|>
[30]
https://www.entomoljournal.com/archives/2018/vol6issue1/PartW/6-1-179-703.pdf
[31]
[PDF] The Natural History of the Arboreal Ant, Crematogaster ashmeadi
Jul 12, 2002 · The Natural History of the Arboreal Ant, Crematogaster ashmeadi. 15pp. Journal of Insect Science, 2:12, Available online: insectscience.org ...
[32]
Acrobat ant, Crematogaster cerasi (Fitch, 1855) - BYGL (osu.edu)
Sep 5, 2024 · Crematogaster ants can be identified using several characters: 11 segmented antennae, flattened petiole, and heart-shaped gaster that is flattened dorsally.Missing: subgenera Decacrema diagnostic
[33]
[PDF] Crematogaster pilosa Ants as Bioindicators of Ecosystem Marsh ...
Crematogaster is known to be an extremely good disperser and colonizer, which has likely facilitated their global spread. (Blaimer 2012). Methods of dispersal ...Missing: initial trophic
[34]
Journey of sperms from production by males to storage by queens in ...
Sperm are produced by males, then ejaculated with spermatophore. They are transferred to the queen's spermatheca, where they are immobilized and stored.
[35]
Egg-Laying “Intermorphs” in the Ant Crematogaster smithi neither ...
We study male parentage and between-colony variation in sex allocation and sexual production in the desert ant Crematogaster smithi, which usually has only ...
[36]
Acrobat Ant | VCE Publications - Virginia Tech
Jan 29, 2021 · Several species of ants in the genus Crematogaster are collectively known as the acrobat ants. They are native species of ants commonly found in ...
[37]
[PDF] Managing Household Ant Pests - AgriLife Extension Entomology
Worker ants develop from eggs (5 to 6 days) through several larval stages (22 to 24 days), a ... Eggs hatch in 8 to 10 days and larvae develop through four stages ...
[38]
Evolution of a soldier caste specialized to lay unfertilized eggs in the ...
Aug 7, 2025 · In Crematogaster (Orthocrema) smithi from Arizona, a third caste exists in addition to winged queens and workers; it is intermediate in size, ...
[39]
https://doi.org/10.1007/s001140050586
[40]
The Ants Chapter 8 - AntWiki
Apr 1, 2020 · Finally, the division of labor by the allocation of tasks among various castes is often referred to as polyethism, a term apparently first ...
[41]
Crematogaster cerasi | INFORMATION - Animal Diversity Web
Ants of genus Crematogaster have unique heart shaped gasters, with the gaster flattened dorsally and convex ventrally. This genus has 11 segmented antennae, ...Missing: taxonomy | Show results with:taxonomy
[42]
Social interactions promote adaptive resource defense in ants
In this study, we demonstrate that the social environment and interactions among colony members (nestmates) modulates the propensity to engage in aggressive ...<|separator|>
[43]
Caste development and evolution in ants: it's all about size
Jan 1, 2017 · Small colonies therefore produce small larvae that exclusively develop into workers. These workers contribute to colony function by nursing ...<|separator|>
[44]
Comparative analysis of the population structure of Crematogaster ...
Mar 24, 2021 · According to our calculations, the maximum colony sizes are typical for invasive species (Crematogaster subdentata, Lasius neglectus, 100–7500 ...Missing: architecture | Show results with:architecture
[45]
Dispersal strategies in the highly polygynous ant Crematogaster ...
Dispersal tactics are also thought to be associated with colony structure (number of queens) ... The mean length, width and height were then used to calculate the ...
[46]
(PDF) Architecture of Arboreal Nests of Crematogaster Ants
Aug 10, 2025 · The nests consist of a number of concavo-convex chambers arranged in a conspheroidal manner, the concavity of each being directed towards the center of the ...
[47]
Queens defense by workers in the highly polygynous ant ...
Crematogaster pygmaea is a highly polygynous ant (Quinet et al., 2009) whose queens show strong attractiveness to workers (Hamidi, 2010). Other features of its ...
[48]
Worker Reproduction in the Highly Polygynous Ant Crematogaster ...
Jul 31, 2023 · We conclude that workers strongly police the worker-produced offspring, even in the queen's absence. Probable high relatedness between queens of ...Missing: hierarchy | Show results with:hierarchy
[49]
[PDF] ants: identification & management - AustinTexas.gov
□ Eat live insects, honeydew, fungus associated with wood. □ Scavengers. □ Baits. □ Dusts, sprays. University of Minnesota. Page 16. □ One node. □ No sting.
[50]
[PDF] 3. Predation by ants on arthropods and other animals - CORE
Most ground-dwelling ant species are omnivorous, feeding on nectar or plant exudates, Hemiptera honeydew, and preying (or scavenging, see Box 1) on other ...
[51]
Nesting System and Foraging Behaviour of Crematogaster ...
Solid particles brought back to the nest were mostly from plants (72.5%) and weakly preys (27.5%). Sugary liquids were preferred over fats, carbohydrate sources ...
[52]
Emergent regulation of ant foraging frequency through a ... - eLife
Apr 17, 2023 · Trophallaxis is the main food-sharing method in many ant species (Meurville and LeBoeuf, 2021). It allows food to circulate through a complex ...
[53]
Crematogaster emeryana - Communication - Alex Wild Photography
... trophallaxis food-sharing behavior. Carrancas, Minas Gerais, Brazil.
[54]
Ants dynamically adjust liquid foraging strategies in response to ...
Sep 15, 2022 · Generally, ants either drink, and internally store liquid and share it through regurgitation and trophallaxis, or they grab liquid into a ...
[55]
How do ants go about storing up food all summer? - Quora
Dec 23, 2021 · They then share food by mutual feeding (trophallaxis) which sees one ant regurgitate food to a nest-mate or larvae, or by giving the nest-mates ...Missing: shifts | Show results with:shifts
[56]
Effect of water and resource availability on ant feeding preferences
This study provides hints on the factors affecting seasonal variations in the uptake of carbohydrates and amino acids, and more importantly, clearly shows how ...
[57]
https://link.springer.com/article/10.1023/A:1007845929801
[58]
Predatory Behavior in Dominant Arboreal Ant Species: The Case of ...
Crematogaster sp. is a dominant arboreal ant species that captures and retrieves very large prey. Hunting workers forage collectively thanks to short-range.
[59]
Paralyzing Action from a Distance in an Arboreal African Ant Species
Dec 14, 2011 · While the C. striatula venom acts from a distance both during predation and interspecific competition, in most Crematogaster species studied so ...
[60]
Parasitoids, Predators, and Group Size in the Paper Wasp, Polistes ...
Oct 1, 1981 · Parasitoids, Predators, and Group Size in the Paper Wasp, Polistes Exclamans ... Crematogaster laeviuscula (Hymenoptera:Formicidae) predation, and ...
[61]
(PDF) Sugary food robbing in ants: A case of temporal cleptobiosis
Aug 9, 2025 · Crematogaster sp. is a dominant arboreal ant species that captures and retrieves very large prey. Hunting workers forage collectively thanks to ...
[62]
help or harm in olive pest control? - Ants - ResearchGate
Crematogaster scutellaris is a common tree-nesting ant in Mediterranean olive groves, where it acts as both predator and soft scale insect tender.
[63]
Trail pheromones of ants - Royal Entomological Society - Wiley
Feb 12, 2009 · In all but Crematogaster, the pheromone is applied to the substrate by dragging the sting lance or the abdomen along the ground as the ant ...
[64]
Trail Pheromone - an overview | ScienceDirect Topics
In ants, trail pheromones are produced in the hindgut, Dufour's gland, the venom gland, and the tibial, tarsal, and abdominal glands. In some myrmecine and ...
[65]
The Dynamics of Foraging Trails in the Tropical Arboreal Ant ... - NIH
Nov 28, 2012 · Foraging trails are on small pieces of ephemeral vegetation, and persist in exactly the same place for 4–8 days, indicating that food sources may be used until ...
[66]
A distributed algorithm to maintain and repair the trail networks of ...
Jun 18, 2018 · We study how the arboreal turtle ant (Cephalotes goniodontus) solves a fundamental computing problem: maintaining a trail network and finding alternative paths.
[67]
Identification of an Alarm Pheromone in the Ant Crematogaster ...
We report the identification of octan-3-one as the major component of an alarm pheromone complex from the heads of Crematogaster peringueyi Emery. This ...
[68]
Enantiomeric composition of an alarm pheromone component of the ...
The enantiomeric composition of 3-octanol produced by the ants,Crematogaster castanea Forel andC. liengmei F. Smith, as a component of their alarm pheromon.
[69]
Chemical profiles of mated and virgin queens, egg-laying ...
Many colonies of the North American ant Crematogaster smithi contain a “third female caste” in addition to queens and workers.
[70]
Nestmate Recognition - AntWiki
Cuticular hydrocarbons (CHCs) are particularly important compounds among the substances used by ants in nest-mate recognition. Genetics and environmental ...
[71]
Ecological consequences of interactions between ants and ... - NIH
... scale insects on fig trees (Ficus vallis-choudae) in Cameroon (Dejean et al ... (Formicinae); Crematogaster sp., Pheidole sp. (Myrmicinae); Ectatoma ...
[72]
Economy of scale: third partner strengthens a keystone ant‐plant ...
Jan 12, 2018 · Crematogaster mimosae is the ecologically dominant symbiotic ant in this system, also providing the highest protection services. In addition ...
[73]
Acacia Ants Respond to Plant-Borne Vibrations Caused by ...
Mar 4, 2019 · We show that the acacia ant Crematogaster mimosae defends its host tree by exploiting plant-borne vibrations caused by browsers feeding on the tree.
[74]
An Old World leaf‐cutting, fungus‐growing ant: A case of convergent ...
Mar 15, 2023 · The African myrmicine ant Crematogaster clariventris is a territorially dominant arboreal species that constructs very hard carton nests.
[75]
Mutualistic interactions between ants and fungi: A review - PMC
Jul 28, 2023 · Ant gardens, or mutualistic associations between certain ant species and epiphytes, shelter endophytic fungi that promote the growth of the epiphytes.
[76]
Distinctive fungal communities in an obligate African ant-plant ...
Mar 15, 2017 · Third, we find that DNA extracted from female alates of Tetraponera penzigi and Crematogaster nigriceps contained matches for most of the fungal ...
[77]
Invading ant threatens unique African ecosystem | Science | AAAS
Sep 8, 2014 · The four species of native ants compete ferociously with each other to occupy each tree. Three species, in the genus Crematogaster, feast on ...
[78]
Impacts of worker density in colony‐level aggression, expansion ...
Dec 8, 2016 · In East Africa, the symbiotic ant Crematogaster mimosae is a co-dominant competitor that engages in high-mortality, intra- and interspecific ...
[79]
Ecological dominance of the red imported fire ant, Solenopsis invicta ...
Crematogaster quadriformis was one of the fastest foragers and the only ant that won an equal number of contests against S. invicta. The low presence of ...
[80]
Disruption of an ant-plant mutualism shapes interactions ... - Science
Jan 25, 2024 · Where big-headed ants encounter whistling-thorn trees, they numerically overwhelm and completely exterminate Crematogaster spp. ants, killing ...
[81]
Natural Enemies of Fire Ants - Ant Pests
Crematogaster species of acrobat ant (with tear-drop shaped abdomen) spraying venom and competing with the larger Solenopsis invicta fire ant for a slice of hot ...<|separator|>
[82]
[PDF] The guild of saprobiontic nematodes associated with ants ... - Biotaxa
Dec 29, 2016 · constituted by the various Macaranga trees inhabited by Crematogaster spp., where thousands of nematodes ... Nematode parasites and associates of ...
[83]
[PDF] Revision of the Ant Genus Crematogaster (Hymenoptera
Jan 1, 2009 · This species is in the subgenus Neocrema. The key characteristics of the Crematogaster acuta worker have very long and pointed propodeal spines.Missing: Acanthocremastogaster | Show results with:Acanthocremastogaster