Apocephalus paraponerae

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Apocephalus paraponerae
Apocephalus paraponerae (10.3897-BDJ.8.e50943) Figure 1.jpg
Bloated, non-gravid “feeder” females of Apocephalus paraponerae Borgmeier on an injured Paraponera clavata F. ant at La Selva Biological Station, Costa Rica
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Phoridae
Genus: Apocephalus
Species:
A. paraponerae
Binomial name
Apocephalus paraponerae
Borgmeier, 1958 [1]

Apocephalus paraponerae is a species of fly in the family Phoridae discovered by Borgmeier in 1958. This species is a parasitoid of the giant tropical ant Paraponera clavata (commonly known as the bullet ant) and uses both visual and chemical cues to locate its host. A. paraponerae can locate fighting or injured ants through host-produced alarm pheromones. [2] Female flies are attracted to the ant to feed and oviposit, while males are attracted to feed and locate females for mating. There is some evidence that suggests that A. paraponerae is a cryptic species complex of at least four genetically distinct species. [3]

Contents

Description

In 2001, it was suggested that A. paraponerae may be a cryptic species complex of at least four genetically distinct, but morphologically indistinguishable species of flies that attack several different ant hosts. Evidence for this claim includes the fact that host-location cues used by A. paraponerae to locate two of the host species differ. Furthermore, these two ant species differ consistently in body size. In addition, mtDNA analysis reveals high sequence divergence between populations, but low sequence divergence within populations. [3]

Anatomy

Larvae

The larva of A. paraponerae generally resembles that of other, related phorids. However, A. paraponerae larvae possess an unusual anterior crosspiece joining the cornua of the cephalopharyngeal skeleton. [4]

Adult

Adult A. paraponerae are small phorids that are about 1.5-2.0 mm long. [5] Notably, female flies possess modified, sclerotized ovipositors which are used to deposit eggs in adult P. clavata. [5]

Distribution and habitat

The geographic range of A. paraponerae is generally coextensive with that of its host, P. clavata. They both occupy the neotropical region and are widespread in the lowland rainforests of Brazil, Nicaragua, Costa Rica, and northern Argentina. [5] [6]

Life history

The A. paraponerae life cycle begins when an adult female deposits eggs in a P. clavata worker ant. Egg hatching and larval development occur very rapidly following oviposition.

Oviposition

Upon discovering an injured ant, A. paraponerae females repeatedly probe the ant's body with their ovipositors, focusing primarily on sutures, areas of overlap between sclerites, and ruptures in sclerites. [5] When laying eggs, females completely extend their ovipositors, allowing the egg to pass through the female reproductive tract. [5]

Eggs

While most parasitic members of the Apocephalus family deposit only single eggs into their hosts, A. paraponerae females typically deposit more than one egg per host. Furthermore, several flies may deposit eggs into a single host. [4] Compared to other phorids, the A. paraponerae egg stage is extremely short, usually lasting only 6.5–7 hours. [4]

Larvae

A. paraponerae larvae develop rapidly within hosts, emerging about 48 hours after hatching. [4] Because females typically deposit several eggs into a single host, and several flies may deposit eggs into the same host, it is not uncommon for up to 20-25 larvae to develop successfully from a single host. [4]

Pupae

The pupal stage of A. paraponerae occurs outside of the host and lasts around 17–26 days, which is much longer than that of other phorids. The long pupal stage may compensate in some way for the very short larval stage. [4] After this long period of pupation, A. paraponerae ecloses as adults.

Adult

The life history of adult A. paraponerae resembles that of both a parasitoid and a scavenger. [4] Both male and female flies are attracted to injured P. clavata to feed on wounds. After oviposition, females frequently feed on fluid oozing out of oviposition wounds. [5] Males may be attracted to injured ants not only to feed but also to mate with female flies that are attracted to those ants. [5]

Food resources and parasitism

Host range

An experimental test of potential host range in A. paraponerae showed that adults were able to develop successfully in seven species in four genera: Paraponera clavata (the natural host), Pachycondyla villosa, P. apicalis, O. opaciventris, E. tuberculatum, E. ruidum, and P. obscuricornis. [7] The number of eggs that developed into pupa differed across host species. Pachycondyla apicalis had the highest percent success of puparia, and P. clavata had the second-highest percent success. The success of larval development in species other than the natural host suggests that larvae are not highly adapted to P. clavata hosts. However, ant species in the subfamilies Myrmicinae and Formicinae were not suitable for larval development, indicating a taxonomic limit to host range. [7]

Host finding

A. paraponerae utilize both visual and chemical cues to locate injured or freshly killed hosts for feeding and egg-laying. These injuries species are often the result of interspecies aggression and serve as ideal hosts for A. paraponerae to lay their egg on. [8] Studies have shown that A. paraponerae can distinguish among different ants based on body size. [9] When ant workers of varying species and sizes were treated with hexane to remove species-specific chemical signatures, A. paraponerae was found to prefer larger P. clavata over other tested species. [9] In addition to visual cues, A. paraponerae are also attracted to 4-methyl-3-heptanone and 4-methyl-3-heptanol, which are major products of the mandibular glands of the host ant P. clavata. [2] As these chemicals are released by fighting, injured, and freshly killed workers, A. paraponerae may use these pheromones to locate suitable targets for feeding or egg-laying. [2] In an experiment, A. paraponerae were found to be equally attracted to ants that were treated with hexane, but unwilling to lay eggs on them. [9] This suggests that A. paraponerae is either unwilling or unable to lay eggs on ants lacking the characteristic chemical signals of 4-methyl-3-heptanone and 4-methyl-3-heptanol. More research is necessary on additional host-finding cues, as A. paraponerae are not attracted to other ant species with the same mandibular gland components as P. clavata. [9]

Geographic variation in host-finding cues

Research conducted at Barro Colorado Island (BCI) in Panama and La Selva Biological Research Station in Costa Rica demonstrated geographic variation in host-finding cues. A. paraponerae used 4-methyl-3-heptanone and 4-methyl-3-heptanol as a host location cue at La Selva, but did not do so at BCI. [10] This is the first reported example of geographic variation in host finding cues in a phorid parasitoid.

Related Research Articles

<span class="mw-page-title-main">Fly</span> Order of insects

Flies are insects of the order Diptera, the name being derived from the Greek δι- di- "two", and πτερόν pteron "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced mechanosensory organs known as halteres, which act as high-speed sensors of rotational movement and allow dipterans to perform advanced aerobatics. Diptera is a large order containing an estimated 1,000,000 species including horse-flies, crane flies, hoverflies, mosquitoes and others, although only about 125,000 species have been described.

<span class="mw-page-title-main">Parasitoid</span> Organism that lives with its host and kills it

In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.

<span class="mw-page-title-main">Tachinidae</span> Family of insects

The Tachinidae are a large and variable family of true flies within the insect order Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in this family commonly are called tachinid flies or simply tachinids. As far as is known, they all are protelean parasitoids, or occasionally parasites, of arthropods, usually other insects. The family is known from many habitats in all zoogeographical regions and is especially diverse in South America.

<i>Paraponera clavata</i> Species of ant

Paraponera clavata, commonly known as the bullet ant, is a species of ant named for its extremely painful sting. It inhabits humid lowland rainforests in Central and South America.

<span class="mw-page-title-main">Phoridae</span> Family of flies

The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of their alternate names, scuttle fly. Another vernacular name, coffin fly, refers to Conicera tibialis. About 4,000 species are known in 230 genera. The most well-known species is cosmopolitan Megaselia scalaris. At 0.4 mm in length, the world's smallest fly is the phorid Euryplatea nanaknihali.

<i>Megaselia scalaris</i> Species of fly

The fly Megaselia scalaris is a member of the order Diptera and the family Phoridae, and it is widely distributed in warm regions of the world. The family members are commonly known as the "humpbacked fly", the "coffin fly", and the "scuttle fly". The name "scuttle fly" derives from the jerky, short bursts of running, characteristic to the adult fly. The name "coffin fly" is due to their being found in coffins, digging six feet deep in order to reach buried corpses. It is one of the more common species found within the family Phoridae; more than 370 species have been identified within North America.

<i>Jalmenus evagoras</i> Species of butterfly

Jalmenus evagoras, the imperial hairstreak, imperial blue, or common imperial blue, is a small, metallic blue butterfly of the family Lycaenidae. It is commonly found in eastern coastal regions of Australia. This species is notable for its unique mutualism with ants of the genus Iridomyrmex. The ants provide protection for juveniles and cues for adult mating behavior. They are compensated with food secreted from J. evagoras larvae. The ants greatly enhance the survival and reproductive success of the butterflies. J. evagoras lives and feeds on Acacia plants, so butterfly populations are localized to areas with preferred species of both host plants and ants.

<i>Melaloncha</i> Genus of flies

Melaloncha is a genus of phorid flies commonly referred to as "bee-killing flies". They are found almost exclusively in the Neotropical realm, although there is one record from extreme southern Texas, United States. They are small flies, usually about 2–3 millimetres (0.08–0.12 in) in length. No true fossils are known, although there are some specimens in Colombian copal, of unknown age.

<i>Apocephalus borealis</i> Species of fly

Apocephalus borealis is a species of North American parasitoid phorid fly that attacks bumblebees, honey bees, and paper wasps. This parasitoid's genus Apocephalus is best known for the "decapitating flies" that attack a variety of ant species, though A. borealis attacks and alters the behavior of bees and wasps. These flies are colloquially known as zombie flies and the bees they infect are colloquially known as zombees. Association with honey bees has so far only been documented from California, South Dakota, Oregon, Washington, British Columbia, and Vermont.

<span class="mw-page-title-main">Metopininae</span> Subfamily of flies

The Metopininae are a subfamily of flies in the family Phoridae.

Diptera is an order of winged insects commonly known as flies. Diptera, which are one of the most successful groups of organisms on Earth, are very diverse biologically. None are truly marine but they occupy virtually every terrestrial niche. Many have co-evolved in association with plants and animals. The Diptera are a very significant group in the decomposition and degeneration of plant and animal matter, are instrumental in the breakdown and release of nutrients back into the soil, and whose larvae supplement the diet of higher agrarian organisms. They are also an important component in food chains.

<i>Pseudacteon</i> Genus of flies

Pseudacteon is a genus of flies in the family Phoridae. There are over 70 described species of Pseudacteon fly. They are also known as ant-decapitating flies due to their parasitic larval stage. An egg is injected by the female fly into the shoulder joint of an ant worker. Soon after, the egg undergoes rapid inflation as it appears to absorb ant hemolymph. This first instar larva migrates into the ant head and consumes the jaw muscle and other tissues, leaving the mandibles hanging and preparing a future exit space. After about two weeks, the ant worker is termed a "zombie" because the fly larva has effectively taken control. The worker leaves the nest and dies in the leaf litter or in a crack in the soil. As it dies, the ant's head falls off, apparently because the fly larva releases an enzyme that dissolves the membrane attaching the ant's head to its body. The fly pupates in the detached head capsule, requiring a further two weeks before emerging through the ant's mouth. In tropical, subtropical areas the flies are active all year round, but in temperate regions they are active during all months except the winter months. Several Pseudacteon species were deliberately introduced to the United States to combat via biological control the invasive fire ant species Solenopsis invicta.

<i>Apocephalus</i> Genus of flies

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References

  1. Borgmeier, T. (1958). "Neue Beitraege zur Kenntnis der neotropischen Phoriden (Diptera, Phoridae)". Studia Entomologica. 1: 305–406.
  2. 1 2 3 Feener Jr., Donald H.; Jacobs, Lucia F.; Schmidt, Justin O. (January 1996). "Specialized parasitoid attracted to a pheromone of ants". Animal Behaviour. 51 (1): 61–66. doi:10.1006/anbe.1996.0005. ISSN   0003-3472.
  3. 1 2 College of Science University of Utah; Seger, Jon; Morehead, Shellee A.; Feener, Donald H.; Brown, Brian V. Evidence for a cryptic species complex in the ant parasitoid Apocephalus paraponerae (Diptera: Phoridae). Evolutionary Ecology Research. OCLC   777821970.
  4. 1 2 3 4 5 6 7 Brown, B. V.; Feener Jr., D. H. (1991-02-01). "Life history parameters and description of the larva of Apocephalus paraponerae (Diptera: Phoridae), a parasitoid of the giant tropical ant Paraponera clavata (Hymenoptera: Formicidae)". Journal of Natural History. 25 (1): 221–231. doi:10.1080/00222939100770131. ISSN   0022-2933.
  5. 1 2 3 4 5 6 7 Brown, Brian V.; Feener, Donald H. (June 1991). "Behavior and Host Location Cues of Apocephalus paraponerae (Diptera: Phoridae), a Parasitoid of the Giant Tropical Ant, Paraponera clavata (Hymenoptera: Formicidae)". Biotropica. 23 (2): 182. doi:10.2307/2388304. ISSN   0006-3606. JSTOR   2388304.
  6. "Apocephalus paraponerae Borgmeier, 1958 | The Diptera Site". diptera.myspecies.info. Retrieved 2019-10-27.
  7. 1 2 Morehead, Shellee A.; Feener, Donald H. (2000). "An experimental test of potential host range in the ant parasitoid Apocephalus paraponerae". Ecological Entomology. 25 (3): 332–340. doi:10.1046/j.1365-2311.2000.00261.x. ISSN   1365-2311.
  8. Segura, Bernardo; Brown, Brian V. (January 2014). "Inter-Colony Aggression ofCamponotus morosusAttractive to Phorid Flies". Journal of the Kansas Entomological Society. 87 (1): 84–88. doi:10.2317/0022-8567-87.1.84. ISSN   0022-8567.
  9. 1 2 3 4 Morehead, Shellee A.; Feener, Donald H. (2000-07-01). "Visual and Chemical Cues Used in Host Location and Acceptance by a Dipteran Parasitoid". Journal of Insect Behavior. 13 (4): 613–625. doi:10.1023/A:1007875921705. ISSN   1572-8889.
  10. Morehead, Shellee A. (2001). "Geographic Variation in Host Location Cues for a Dipteran Parasitoid of Paraponera clavata1". Biotropica. 33 (3): 495. doi:10.1646/0006-3606(2001)033[0495:gvihlc]2.0.co;2. ISSN   0006-3606.