Myrmecophily in Staphylinidae
Many species of
Taxonomy
Staphylinidae is currently considered to be the largest family of beetles, with over 58,000 species described. As such, many myrmecophilous species are unknown. The majority of studied myrmecophilous rove beetles belong to the subfamily
Ecology and behavior
Due to their large number and diversity, myrmecophilous rove beetles occupy an array of behaviors. Myrmecophilous interactions can be generalized into categories, in three of which staphylinids can be found. The synecthrans, or “persecuted guests,” the synoeketes, or “tolerated guests,” and the symphiles, or “true guests.”[4]
Synecthrans
Synecthran staphylinids live on the periphery of the host colony and are not accepted into the colony. The majority are in the subfamily Aleocharinae, having defensive glands on their terminal abdominal segments. Species in this group, such as those in the genus Pella, commonly lay their eggs in the refuse heaps of their host ant, where the larvae feed on the discarded carcasses of ants. If detected by the host ant, the larvae enters a typical defense position, facing the ant with its abdomen tip raised. Usually this behavior results in the ant palpating the beetle's abdomen and stopping the attack.[5] At least two members of Pella, P. funestus and P. humeralis, produce several of their host ant's alarm pheromones to avoid aggression.[6] Other species, like members of Myrmechusa, Aenictonia, and Anommatochara, prey on raiding columns or on the nests of driver ants (Dorylus).[7] Myrmechusa species attack the ants from the rear before pulling them from the main column. If pursued, they rapidly wave their abdomens through the air, releasing a noticeable scent. Ants encountering this scent seem disoriented and take almost 20 minutes to recover.[8]
Synoeketes
Synoeketetic staphylinids live in close contact with their host ants but are not integrated into the colony. These species may be further categorized as neutral, mimetic, loricate, and symphiloid synoeketes. Neutral synoeketes ignore and are ignored by their host, but feed on refuse. There are few staphylinid neutral synoeketes, but some are found in the genus Athetini, which live in the debris and fungal chambers of leaf cutter ants (
Symphiles
Symphilic staphylinids have been fully integrated into the host ant's society. Symphilic species have undergone complex morphological adaptations, many becoming myrmecoid. Most have developed trichomes, which secrete appeasement pheromones. The most extreme adaptations, found in members of tribe Clavigerini, include the reduction of mouthparts for
Chemical mimicry
Chemical mimicry refers to the production of one species’ chemical signals by another species. Many myrmecophilous staphylinids have evolved chemical mimicry to deter or placate ants. Synecthrans, as non-tolerated guests, primarily produce defensive secretions. Pella species produce two compounds found in their host ant Lasius fuliginosus, undecane and sulcatone, which elicit aggressive and panic reactions respectively. Although it seems counterintuitive to release an aggressive alarm pheromone as a defense, the presence of sulcatone stops the aggression response to undecane. Ants exposed to the defensive secretion act less aggressively and avoid the odor.
For staphylinids accepted into the host colony, chemical mimicry is used more for camouflage. The majority of the chemical signals used are cuticular hydrocarbons, which are produced in the cuticle of the host ant at certain concentrations and are palpated to determine the identity of an ant. Species in close contact with their host ants are able to pick up the host's hydrocarbons and imitate the ant's hydrocarbon pattern, thus appearing in scent at least to be the same species as the host ant. As hydrocarbon patterns are specific to an individual colony, the rove beetles are generally restricted to one nest. The production of a new hydrocarbon pattern takes time, during which the beetle is vulnerable to detection and attack.[6] Some species, such as Zyras comes, produce volatile pheromones as well as cuticular hydrocarbons, which may provide it more protection than contact based pheromones while traveling with its host in foraging trails.[13]
See also
References
- ^ Lapeva-Gjonova, Albena (April 2013). “Ant-Associated Beetle Fauna in Bulgaria: A Review and New Data”. Psyche (Cairo: Hindawi Pub. Co.) 2013: 1-14
- ^ O'Keefe, Sean (2000). "Ant-Like Stone Beetles, Ants, and Their Associations (Coleoptera: Scydmaenidae; Hymenoptera: Formicidae; Isoptera)". ‘’J. N. Y. Entomol. Soc.’’ (NYC: N. Y. Entomol. Soc.) 108 (3/4): 273–303.
- ^ Parker, Joe; Eldredge, Taro (10 Dec 2012), "The Bizarre, Beetle-Biased World of Social Insect Exploitation", ‘’Scientific American’’
- ^ Mynhardt, Glené (2013). "Declassifying Myrmecophily in the Coleoptera to Promote the Study of Ant-Beetle Symbioses". ‘’Psyche’’ (Cairo: Hindawi pub co) 2013: 1–8.
- ^ Holldobler, B.; Moglich, M.; Maschwitz, U. (1981). "Myrmecophilic Relationship Of Pella (coleoptera: Staphylinidae) To Lasius fuliginosus (hymenoptera: Formicidae)". Psyche(Cairo: Hindawi Pub Co.) 88: 347–374
- ^ a b Stoeffler, Michael; S. Maier, Tanja; Till, Tolasch; L. M. Steidle, Johannes (2007). "Foreign-language Skills in Rove-Beetles? Evidence for Chemical Mimicry of Ant Alarm Pheromones in Myrmecophilous Pella Beetles (Coleoptera: Staphylinidae)". ‘’J Chem Ecol’’ (NYC: Springer Pub) 33: 1382–1392
- ^ . Koblick, Tonya; H. Kistner, David. "A Revision of the Species of the Genus Myrmechusa from Tropical Africa with Notes on Their Behavior and Their Relationship to the Pygostenini (Coleoptera: Staphylinidae)". “Ann. Entomol. Soc. Am” 58 (1): 28–44
- ^ M. Wheeler, William (1960). ‘’Ants: Their Structure, Development, and Behavior.’’ 9. NYC: Columbia Univ Press.
- ^ Cammaerts, R. (1992). "Stimuli inducing the regurgitation of the workers of Lasius flavus (Formicidae) upon the myrmecophilous beetle Claviger testaceus (Pselaphidae)". “Behavioural Processes” (ScienceDirect) 28 (1-2): 81–95
- ^ W. Rettenmeyer, Carl; D. Akre, Roger (1968). "Trail-Following by Guests of Army Ants (Hymenoptera: Formicidae: Ecitonini)". J. Kans. Entomol. Soc. (Kans. Entomol. Soc.) 41 (2): 165–174
- ^ a b Parker, Joseph (2016). "Myrmecophily in beetles (Coleoptera): evolutionary patterns and biological mechanisms" (PDF). Myrmecological News. 201, 22: 65–108 – via https://authors.library.caltech.edu/.
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- ^ Akino, Toshiharu (2002). "Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae)". Chemoecology (NYC: Springer pub) 12 (2): 83–89