Leaf hopper and meat ant relationship advice

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leaf hopper and meat ant relationship advice

been appreciated that these facultative relationships can be mutualistic. P. candidus system suggests that plant-spider associations, like other facultative Australia (Maslin and Hopper ), reaching 5 m in advice and help, and to D. Phillips for advice on analyzing . seeds and meat ants Iridomyrmex viridiaeneus. The leafhopper populations in two almond witches'-broom phytoplasma (AlmWB) prove a definite vector relationship, the technique is useful in narrowing the search .. and presence of aggressive ants) and leafhopper richness and abundance. .. of salivary sheath termini in plant tissue generated from insect stylet tips. Meat ants, also known as gravel ants are an omnivorous species of ants found throughout Australia. They live in vast underground nests and are very hostile to .

Ants contaminated with tergal gland secretions in these episodes usually displayed an aversive reaction, releasing the grip on the beetles and grooming and wiping their mouthparts and antennae on the substrate. But the beetles then had to escape quickly, because other ants nearby became alerted and rapidly converged on the scene, apparently in response to the alarm pheromones of their nestmates.

Even with this element, however, isolated tergal gland contents of P. Apparently the repellent effect of the quinones, the major components of the tergal gland secretions, is stronger than the attractant effect of undecane.

When the ants' antennae were directly contaminated with the beetles' tergal gland secretions, they hung almost motionless, and the ants were disoriented for several minutes. In early spring, when most of the Pella adults are close to the entrance of the ants' nest, they feign death when attacked by ants. The ants then either ignore the motionless beetles or carry them around and finally discard them on the refuse piles. Later in the season, when the activity of ants and beetles is much higher, the beetles employ a different appeasement technique.

Each time they encounter ants they flex their abdomen forward and point the abdominal tip toward the head of their adversaries. Usually the ants respond by antennating the tip and licking it briefly Figure This ordinarily damps the ants' aggressions, allowing the beetle to escape.

When on occasion the ants become persistent, the beetle extrudes a white, viscous droplet from the abdominal tip, which the ants eagerly lick up. It is not yet possible to assign one specific gland to the appeasement function. Kitchen middens and peripheral nest chambers. We have described the genus Pella in some detail as representative of an early evolutionary grade in the myrmecophilous evolution of the aleocharine staphylinids.

Species of Pella are specialized predators and scavengers, but they are less advanced in their myrmecophilic relationships than aleocharine species specialized on the kitchen middens, peripheral nest chambers, and brood chambers of the host ants' nests.

A representative of the latter, more advanced evolutionary grade is Dinarda. The myrmecophilous habits of this genus have been known since Wasmann's first account published inand the genus has since been recorded with many species of Formica Bernard, Kistner points out that the species-level taxonomy is still in poor condition, so that the best studied form, Dinarda dentata, might in fact comprise several species.

The following account is based mostly on the form or species of Dinarda dentata that lives in nests of the red slavemaking ant Formica sanguinea of Europe Figure The larvae of Dinarda are concentrated in the kitchen middens of the Formica hosts, where they feed on dead ants and debris. When they encounter worker ants they exhibit the same appeasingly defensive behavior described for Pella larvae Figure Not surprisingly, they also possess a similar complex glandular structure in the tip of their abdomens.

Some adult Dinarda beetles are found in the kitchen middens, where they exist as scavengers. Most, however, patrol through the peripheral nest chambers, where they feed on arthropod prey brought in by the host workers. Wasmann reported that Dinarda also eats mites and occasionally even ant eggs and larvae. This parasitism occurs mostly in the peripheral chambers of the nest, where the bulk of regurgitation occurs between newly returning foragers and nest workers.

The Dinarda beetles sometimes insinuate themselves between two workers exchanging food and literally snatch the droplet of food from the donor's mouth. They also use a simple begging behavior in order to obtain food directly from the returning foragers.

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The beetle approaches an ant and furtively touches its labium, causing the ant to regurgitate a small droplet of food Figure The ant, however, soon recognizes the beetle as an alien and commences to attack it. At the first sign of hostility the beetle raises its abdomen and offers the ant the appeasement secretions at the abdominal tip. The abdominal tip is quickly licked by the ant, and almost instantly the attack ceases. During this brief interlude the beetle makes its escape. Dinarda also possesses a well-developed tergal gland, but its repellent secretions are applied only as a last resort against ant attacks.

It is noteworthy that a myrmecophile belonging to a radically different taxonomic group obtains food in a similar manner to Dinarda. Janet discovered that the thysanuran Atelura formicaria, which is a very common symbiont of many European ant genera Bernard,occasionally snatches food during food exchanges between worker ants.

Like the Dinarda beetles, Atelura favors the peripheral chambers of the host nests. Another example of a myrmecophile adapted especially to kitchen middens and peripheral nest chambers is the histerid beetle genus Hetaerius Figure As Wasmannfirst reported, the European species Hetaerius ferrugineus is most frequently found in nests of Formica fuscawhere it feeds on dead and wounded ants.

Occasionally it also consumes ant larvae and pupae. Most of the time the ants pay no attention to the Hetaerius. When they do attack an adult, the beetle exhibits death-feigning behavior by holding perfectly still with its legs closely appressed to its body.

The ants frequently react to this nonviolent resistance by carrying the beetle around, licking it and finally releasing it. It has been suggested that Hetaerius has special trichome glands opening on the margins of the thorax Escherich and Emery, ; Wasmann, ; Wheeler, c. However, no histological investigations have been performed to confirm this assumption. Wheeler c observed that adults of the North American Hetaerius brunneipennis solicit regurgitated food from the host ants.

The beetle sometimes waves its forelegs toward passing ants, and by this action appears to attract their attention. A very similar behavior has been more recently recorded in H. The soliciting beetle takes an upright position, stretching its forelegs widely apart and waving them slightly towards the approaching ant. The ant antennates and licks the beetle with her mandibles usually held relatively closed Figure Thus, contrary to a suggestion originally made by Wheeler, Hetaerius brunneipennis does not represent a more advanced evolutionary state over H.

The brood chambers constitute the optimal niche within an ant nest for a social food-flow parasite, because there the food of the highest quality is concentrated to be fed to the developing larvae, callow workers and queen s. Moreover, the immature ant stages housed in the brood chambers provide the most valuable prey for specialized ant predators. The brood chambers are nevertheless difficult for parasites to penetrate, because they are fiercely defended by the ants. Very special adaptations are thus needed for myrmecophiles to exploit these sites as an ecological niche.

This has been accomplished by some of the evolutionarily most advanced myrmecophilous staphylinids, of which the aleocharine beetles Lomechusa strumosa and several species of the genus Atemeles are the premier examples.

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Lomechusa Figure lives with Formica sanguinea in Europe. Atemeles pubicollis Figurealso a European species, is normally found with the mound-building wood ant Formica polyctena during the summer.

But in the winter it inhabits the nests of ants of the myrmicine genus Myrmica. We know from the observations of Wasmann of more than 70 years ago that these beetles are both fed and reared by their host ants. The behavioral patterns of the larvae of these beetles are similar for Lomechusa and the various Atemeles species; in particular the larvae prey to a certain extent on their host ants' larvae.

It is therefore astonishing that nurse ants not only tolerate these predators but also feed and protect them as readily as they do their own brood Figure The beetle larvae show a characteristic begging behavior toward their host ants. As soon as they are touched by an ant, they rear upward and try to make contact with the ant's head.

If they succeed, they tap the ant's labium with their own mouthparts. This apparently releases regurgitation of food by the ants. The ant larvae beg for food in much the same way but usually less intensively than the beetle parasites.

With the aid of experiments using food mixed with radioactive sodium phosphate it has been possible to measure the social exchange of food in a colony.

The results show that myrmecophilous beetle larvae present in the brood chamber obtain a proportionately greater share of the food than do the host ant larvae. The presence of ant larvae does not effect the food flow to the beetle larvae, whereas ant larvae always receive less food when they compete with beetle larvae.

This disparity suggests that the releasing signals presented by the beetle larvae to the nurse ants may be more effective than those presented by the ant larvae themselves. The Lomechusa and Atemeles larvae are also frequently and intensively groomed by the brood-keeping ants. Thus it appears probable that chemical signals are also involved in the interspecific relationship.

The transfer of substances from the larvae to the ants was detected with the aid of radioactive tracers. These substances are probably secreted by glandular cells that occur beneath the integument of the dorsolateral area of each segment. The biological significance of the secretions was elucidated by the following experiments. Beetle larvae were first completely covered with shellac to prevent the release of the secretion. They were then placed outside the nest entrance, together with freshly killed but otherwise untreated control larvae.

The ants quickly carried the control animals into the brood chamber. The shellac-covered larvae, on the other hand, were either ignored or carried to the garbage dump. It was found that for adoption to be successful, at least one segment of the larva must be free of shellac. Furthermore, after all the secretions have been extracted with solvents, the larvae are no longer attractive.

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When the extracted larvae are then contaminated with secretions from normal larvae, they regain their attractiveness. Even filter paper dummies soaked in the extract are carried into the brood chambers. To summarize, the experiments show that the adoption of the Lomechusa and Atemeles larvae and their care within the ant colony depend on chemical signals.

It may be that the beetle larvae imitate a pheromone which the ant larvae themselves use to release brood-tending behavior in the adult ants.

This inference is supported by the fact that the beetle larvae and host ant larvae can be transplanted from the host species to closely related Formica species. But the ant species that do not accept the larvae of the ant hosts also reject the beetle larvae.

The question next arises as to how the ant colony manages to survive the intensive predation and food parasitism by the beetle larvae.

Meat ant and Leaf Hopper

The answer appears to be very simple. The beetle larvae are cannibalistic, and this factor alone appears to be effective in limiting the number of beetle larvae in the brood chambers at any time. The larvae of both genera pupate in the summer, and they eclose as adult beetles at the beginning of autumn. The young Lomechusa adults leave the ant nest and after a short period of migration seek adoption in another nest of the same host ant species.

Atemeles adults, on the other hand, emigrate from the Formica nest, where they have been raised, to nests of the ant genus Myrmica. Formica polyctena nests normally occur in woodland, while those of Myrmica are found in grassland around the woods. Experiments have revealed that when Atemeles leave the Formica nest they show high locomotor and flight activity and orient toward light.

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This may well explain how they manage to reach the relatively open Myrmica habitat. Once they reach the grassland, the beetles must distinguish the Myrmica ants from among the other species present and locate their nests.

Choice experiments in the laboratory have revealed that they identify the Myrmica nests innately by specific odors. Windborne species-specific odors are equally important in the springtime movement back to Formica nests. Having found the hosts, the beetles must then secure their own adoption.

The process comprises the four sequential steps depicted in Figure First, the beetle taps the ant lightly with its antennae and raises the tip of the abdomen toward it. The latter structure contains the appeasement glands, the secretions of which are immediately licked up by the ant and appear to suppress aggressive behavior. The ant is then attracted by a second series of glands along the lateral margins of the abdomen.

The beetle lowers its abdomen in order to permit the ant to approach this part of its body. The glandular openings are surrounded by bristles, which are grasped by the ant and used to carry the beetle into the brood chambers. Before leaving the Formica nest the Atemeles beetle must obtain enough food to enable it to survive the trek to the Myrmica nest. This it obtains by vigorous solicitation from its hosts.

The begging behavior is essentially the same toward both Formica and Myrmica Figure ; see also Figure The beetle attracts the ant's attention by rapidly drumming on it with its antennae. Using its maxillae and forelegs it taps the mouthparts of the ant, thus inducing regurgitation.

As noted previously, the ants themselves employ a similar mechanical stimulation of the mouthparts to obtain food from one another. It is thus clear that Atemeles is able to obtain food by imitating these very basic tactile food-begging signals. What is the significance of the seasonal changes in hosts in Atemeles? There are good reasons for believing that Atemeles first evolved myrmecophilic relationships with Formica rather than with Myrmica. It seems likely that the ancestral Atemeles beetles hatched in Formica nests in the autumn and then dispersed, returning to other Formica nests only to overwinter.

However, in the Formica nest the immature stages disappear during the winter, and consequently social food flow is reduced. In contrast, the Myrmica colony maintains brood throughout the winter. Thus, in Myrmica nests, larvae and nutrients from the social food flow are both available as high-grade food sources to the myrmecophiles. These circumstances, coupled with the fact that the beetles are sexually immature when they hatch, suggest why it is advantageous for the beetle to overwinter in Myrmica nests.

While there the Atemeles complete gametogenesis, so that when spring comes the beetles are sexually mature. They then return to the Formica nest to mate and lay their eggs. At this time the Formica are just beginning to raise their own larvae and the social food flow is again maximized. The life cycle and behavior of Atemeles is thus synchronized with that of its host ants in such a manner as to take greatest advantage of the social life of each of the two species in turn. The North American staphylinid myrmecophiles of Xenodusa seem to have a similar life history as Atemeles.

The larvae are found in Formica nests and the adults overwinter in the nests of the carpenter ants of the genus Camponotus Wheeler, It is undoubtedly significant that Camponotuslike Myrmicamaintains larvae throughout the winter. It may well be that the host-changing behavior of Xenodusa has the same significance as that inferred for Atemeles.

Wasmann considered Lomechusa strumosa to be the most evolutionarily advanced of the myrmecophilous staphylinids. Lomechusa also appears to have at least one migratory phase in spring, when, shortly after overwintering as an adult inside the Formica sanguinea nest, it travels to another nest of the same host species, where mating takes place Figure The adoption procedure into the new host ants' nest is even more complicated than that of Atemeles.

Like other aleocharine myrmecophiles, Lomechusa is equipped with well-developed tergal glands that produce defensive substances.

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The secretion consists of benzoquinone, methyl-benzoquinone, ethyl-benzoquinone and n-tridecane, with the latter compound accounting for more than 80 percent of the volatiles Blum et al. This powerful repellent mixture, however, is normally employed only when the beetle is attacked by non-host ants. When approached by host ants Lomechusa behaves much more gently. It first presents the trichome structures on its legs, particularly the femora.

The beetle then slowly circles around on the spot, with its legs widely extended so that the femoral trichomes are easily accessible. Simultaneously the beetle antennates the ants, bending its body sideways or backwards in order to reach the ants with its antennae. Next it slowly points its abdominal tip at the ants. Lomechusa possesses a battery of exocrine glands at the abdominal tip, and during the early adoption phase it frequently extrudes a whitish, viscous droplet which is eagerly licked up by the ants.

The secretion is proteinaceous and contains no appreciable amounts of carbohydrates. The ants that feed on the material seem to grow calmer in the process. The Lomechusa now permit the ants access to trichomes along the lateral margins of the abdomen that project above the adoption glands. The ants lick and grasp the trichome bristles and finally carry or drag the beetle into the brood chambers of their nests. Inside the nest the Lomechusa continues to be tended by its host ants, despite the fact that it preys on the ants' brood Figure In addition the adult Lomechusa, like the larva, receives food directly from the nurse ants.

Its soliciting behavior is markedly different from that of adult Atemeles. Lomechusa possess trichomes at the labrum, which are frequently licked by the ants. During this procedure the beetle uses its maxillae to stimulate the mouthparts of the ants, an action that seems to elicit the regurgitation of liquid food.

In contrast to Atemeles, however, Lomechusa does not use its forelegs when begging for food. One gets the impression, as Wasmann first suggested, that the ants' feeding behavior toward adult Lomechusa resembles more the nursing behavior toward ant larvae than the trophallactic food exchange between adult ants. It is possible, but not yet proved, that Lomechusa is fed primarily with high-grade secretions from the postpharyngeal and labial glands of nurse workers--in other words, the same food given the larvae.

In short, Lomechusa strumosa presents its host ants with a complex variety of chemical signals, including imitations of brood-tending pheromones and other kinds of pseudopheromones. The species is thereby superbly well adapted to live with Formica sanguineaespecially in the brood chambers where it finds both food and a protected and regulated environment.

Most of the more casual observations of regurgitation between host and symbionts suggest that the exchange is exploitative, meaning that the liquid flows exclusively from the host to the symbiont. However, an exception has been reported in the case of Amorphocephalus coronatus.

Adults of this brenthid beetle live with Camponotus in southern Europe Figure According to Le Masne and Torossian they receive food from some of the host workers and regurgitate part of it back to other host workers. This is the first reported example that could be construed as altruistic behavior on part of the symbiont beetles Wilson, However, quantitative investigations are needed to see how much of the received food the beetle shares with other host ants.

Other adaptations of predators in the ants' brood chambers. Entomologists generally agree that a select few of the aleocharine staphylinids, including Atemeles and Lomechusa, have reached the evolutionary pinnacle of myrmecophilous adaptations. In mimicking the ants' important communication signals they are able to integrate themselves into the ant societies and exploit the hosts' brood chambers.

There nevertheless exists a diversity of other arthropods that also preys on ant brood but uses a very different set of sophisticated techniques to invade the ants' nests. An especially instructive example is provided by the scarabaeid beetle genus Cremastocheilus Figure Cremastocheilus is a North American genus comprising about 50 species Krikken,various of which have been studied by Wheeler bCazier and MortensonAlpert and Ritcherand Kloft et al.

Most recently Alpert has provided an insightful review of the ecology, behavior and evolution of the genus. Various species of Cremastocheilus, the greatest number of which are found in the deserts of the southwestern United States and Mexico, use the formicine ants FormicaMyrmecocystusLasius and Camponotusand the myrmicines PogonomyrmexVeromessor and Aphaenogaster as hosts.

Ant nests in this arid area provide a high concentration of food resources and a refuge from predators and severe climatic change.

Cremastocheilus larvae, like other scarabaeid larvae, feed primarily on decaying vegetable material. They develop within piles of vegetable matter inside ant nests, despite the apparent lack of any morphological adaptations for defense from their hosts.

Alpert's field observations revealed few interactions between beetle larvae and ants. Experiments showed that the larvae are generally ignored by ant workers, while unrelated scarabaeid larvae of similar size used as controls were attacked immediately. Alpert concluded that while Cremastocheilus larvae are not nutritionally dependent on ants they can be raised in complete isolationthey gain protection from predators and desiccation simply by virtue of their residency in ant nests.

Cazier and Statham suggested that the adult beetles are first brought into the ant nests as booty by the foraging ants. The beetles, however, are so well protected by their heavily sclerotized cuticle that the ants are unable to kill them. Once inside the nest, the beetles themselves prey on the ant larvae Cazier and Mortenson, This scenario has been basically confirmed by Alpert's field and laboratory observations. Although Cremastocheilus adults are capable of feeding on any species of ant larvae, only one or a few ant species are selected as hosts under natural conditions.

This specificity appears to be a product of the host-searching process. It was originally believed that secretions from glandular trichomes, located on prominent pronotal angles in all species of Cremastocheilus, attracted ants to beetles outside the nest and induced adoption Cazier and Mortenson, ; but experiments conducted by Alpert did not confirm this assumption. Alpert released dead Cremastocheilus and other scarabaeid beetles with live C.

There was no significant preference by the ants for live C. Nor did glandular extracts elicit special attraction on the part of the ants. All the evidence supports the hypothesis that certain Cremastocheilus species are mistaken for prey items and thereby gain access to the nest. In fact, Cremastocheilus beetles commonly feign death when approached by ants, retracting their antennae into protective grooves and sticking out their heavily sclerotized legs.

While some species gain entry into the host ants' nest by mimicking prey items, other species are able to march directly through the nest entrance or burrow through thatch piles into the interior. Often the beetles are attacked by ants inside the nest. Occasionally they are forcibly ejected, only to be carried in again by other ants. Attacks usually persist for several minutes, until the ants give up and the Cremastocheilus move to concealed corners in chambers or passageways.

In the end the beetles move slowly toward the brood chambers, where they are completely ignored by the ants. Although pronotal angles were specifically licked, most of the dorsal surface of the beetles was also involved. Alpert concludes that while adults of Cremastocheilus feed on ant brood, trichome secretions may distract workers, reduce aggression, and prevent workers from evacuating brood, while other glands serve to absorb colony odors. They claimed that by transferring Cremastocheilus trichome substances to other workers, the entire colony could be appeased.

By carefully sectioning Camponotus castaneus and members representing most of the other species groups, Alpert has shown that this conclusion is incorrect. The prosternal apophysis and propygidial spiracles are not glandular in the genus Cremastocheilus. Alpert nevertheless did find many new areas that bear exocrine glands.

In a series of experiments with radioactive tracers, he further discounted the claim by Kloft et al. Entomologists continue to be intrigued by the possible evolutionary pathways that led to the specialized predaceous behavior of Cremastocheilus. One hypothesis is that Cremastocheilus has changed from what was almost certainly a herbivorous diet to become a fully carnivorous scarabaeid Wilson, ; Kistner, This route might have started with the adaptation of larvae for development in ant nests.

In typical scarabaeid fashion Cremastocheilus larvae eat vegetable matter in the soil, and because of the concentration of vegetation in ant nests and protection from predators, ant colonies became the new oviposition site. Such behavior is known in cetonine scarabaeids, such as Euphoria inda, which develops in Formica obscuripes nests in North America Windsor,and the European species Cetonia curprea, the larvae of which also develop in Formica nests Donisthorpe, Alperton the other hand, argues that the adults of Ephoria and Cetonia are not preadapted for survival in ant colonies and did not evolve in this direction.

Cremastocheilus adults, in contrast, have many morphological adaptations for integration into ant nests and an examination of other genera of Cremastocheilini gives the best understanding of how the genus Cremastocheilus might have evolved. Genuchinus ineptus develops in plants of the liliaeceous genus Dasylirion of Mexico and the southwestern United States, and any association with ants is entirely accidental.

The adults are general predators, feeding on many different soft bodied insects in the laboratory. Dasylirion plants are inundated with fly larvae, the most probable source of food for Genuchinus ineptus in the field.

Genuchinus ineptus, in this interpretation, is preadapted for myrmecophilous existence as a predator on ant brood. The predaceous mouthparts of Genuchinus ineptus adults are almost identical to those of Cremastocheilus. There are other morphological features common to Genuchinus and Cremastocheilus which are apparently adaptations for predation. These include the flattened body shape, hard, pockmarked integument, retractable antennae, and protective mentum. The larvae of Genuchinus ineptus develops in leaf compost of Dasylirion plants, and the pupae are sheltered by protective cases.

The Genuchinus differ from Cremastocheilus by remaining inside their cases until spring. According to Alpert's reconstruction, the major evolutionary step taken by the genus Cremastocheilus was to specialize on ant brood. The development of pronotal angles and trichome hairs is a consequence of this specialization. Other genera of New World Cremastocheilini, such as Centrocheilus Krikken,appear to occupy an intermediate level leading to this stage. Other species of Genuchinus from South America occur commonly in banana leaves and bromeliads.

Ant nests are also frequently found at these sites, and beetles may have moved from the leaf litter into the ant colonies to promote their larval development.

A study of the life history of Paracyclidius bennetti, which is found with ants on bromeliads in Trinidad, might provide insight into this evolutionary stage. The genus Cremastocheilus adaptively radiated into ant colonies inhabiting the southwestern deserts of North America. Today members from all five subgenera are found in Arizona, New Mexico, and Colorado. All species groups have at least one member from the Southwest, except for the castaneus group which is strictly eastern in distribution.

A major question remaining is whether after speciation and adaptive radiation species of Cremastocheilus are now at different evolutionary stages along a pathway toward full integration. Alternatively, differences in behavior and morphology might simply reflect adaptive radiation to the behavioral ecology of different species of ants.

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Alpert argues and we agree that the latter hypothesis is more likely, since all species of Cremastocheilus have the same basic relationship with ants. Most species of the beetle family Pselaphidae are carnivorous, with members of the subfamilies Pselaphinae and Clavigerinae being additionally myrmecophilous. According to Park all of the approximately species of the clavigerines are especially adapted to live in ant nests.

All observations to date indicate that these clavigerine beetles live among the brood of their host ants, either preying on it or exploiting it in some other manner. Kistner has provided a table listing all the behavioral acts thus far observed in Adranes and Claviger. From these data we learn that Claviger testaceus Figurea frequent myrmecophile of the formicine genus Lasiuspreys on ant eggs, larvae and pupae while at the same time receiving food from larvae and adult ants by means of trophallaxis Figure The beetles occasionally eat dead ants or booty brought into the brood nest by their hosts.

Finally, Claviger adults are also frequently groomed by the ants. The host workers lick secretions from the numerous trichomes and other exocrine glands, the anatomy of which has been carefully studied by Cammaerts The ants rarely treat Claviger aggressively.

When transplanted experimentally to new nests the beetles are accepted by a diversity of ant species--even those genera with which they have never been found to occur naturally Donisthorpe, ; K. When attacks do occur, almost invariably during the initial phase of adoption, the Claviger usually show death-feigning behavior.

The ants often respond by grasping the beetles on their slender thoraces and carrying them around for a short while before releasing them again, usually in the midst of the ant brood. Claviger rarely suffer any harm from this treatment.

Cammaerts observed that Lasius workers regurgitate liquid food to Claviger testaceus after they have licked them intensively. He noted a similarity of the movements with those occasionally observed when ants lick prey objects before they place them as food among the ant larvae.

It is noteworthy in this connection that Akre and Hill observed that ant larvae lick the trichomes of the clavigerine Adranes taylori, although in contrast to the species of Claviger the Adranes do not eat ant larvae. Instead they depend on food regurgitated to them by the larvae. In both cases, however, the mimicking of prey signals would be an efficient way to gain access to the brood chambers.

One other category of myrmecophilous adaptation by brood predators deserves special mention. The larvae of the syrphid fly genus Microdon, which develop in ant nests, have unusual shapes Figure They resemble slugs or limpets and were originally described as mollusks, later as coccids, before they were correctly identified as syrphid larvae for recent reviews of the literature see Duffield,and Kistner, Also, the larvae of at least two other nest parasites are convergent in this slug or limpet-like form: The genus Microdon is cosmopolitan and contains about species, most of which have been recorded from nests of such diverse ant genera as CamponotusFormicaLasiusPolyergusAphaenogasterCrematogasterMonomoriumPheidoleIridomyrmexand Tapinoma.

Microdon larvae either feed on detritus Akre et al. Duffield reports that the first instar larvae of M. Second and third instar larvae, on the other hand, consume small ant larvae but never pupae. Frequently the adult ants pull their larvae away from the Microdon.

Successful Microdon larvae move up and over the ant larvae, piercing the larval skin, emptying the body contents, and discarding the empty shell. Duffield observed third instar larvae consuming ant larvae each during various minute periods of observation. How are the Microdon larvae able to survive inside the ants' brood chambers? The fly larvae are usually bulky and have a heavily reticulated skin, which appears to confer mechanical protection from ant attacks.

Recently Garnett et al. These investigators studied three Nearctic species, Microdon albicomatus and M. The larvae are obligate predators on the brood of their hosts. However, they appear to feed almost exclusively on cocoon occupants, that is, on larvae already encased by a cocoon, or prepupae, or pupae.

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Attacks by ant workers on the Microdon larvae were extremely rare, but obvious acts of aggression did occur when second and third instars of M.

This result indicates the presence of a pseudopheromone that mimics the colony odor of the hosts. They observed when an ant nest is exposed to sunlight Microdon larvae compress their bodies laterally into the rough shape of an ant cocoon.

They are then quickly picked up by workers and transported into undisturbed parts of the nest along with real cocoons.

Third instar larvae are evidently too large and bulky to exhibit curling behavior, and they are never transported. If the first instar of Microdon gains access to a defenseless pupa in cocoon, it will not only find nourishment and protection from attack, but an opportunity to acquire recognition chemicals characteristic of the host.

Upon its exit from the cocoon the first instar has gained the necessary chemicals to be accepted and even transported as brood. The possible dilution of such chemicals by growth of the larvae is partially compensated by the second instar continuing to attack and enter cocoons. The eyes are semicircular and positioned around the midpoint of the ants' head capsule. The frontal carinae a keel-shaped ridge or structure are convex and the antennal scapes extend beyond the head's posterior margin by two or three times the diameter.

Erect setae are found all over the antennal scape and noticeably prominent on the clypeal margin a shield-like plate at the front of an insect. The mandibles are elongated and triangular, with long curved setae around the head capsule. These setae are mostly short and bristly.

The mesonotum is sinuous meaning it has many curvesand, like the pronotum, has 12 or more mesonotal setae. The mesothoracic spiracles are very small and the propodeal dorsum is smooth or convex. There are also a number of propodeal setae. There are both non-marginal and marginal setae present on the first gastral tergite around the gaster.

Queens are easily distinguishable from workers by their black colour and larger size, measuring The antennae and legs are ferruginous rust-like colourthe head is fusco-ferruginous, and the sides beneath the face and mandibles are ferruginous. The head is wider than the thorax and emarginate. There is an impressed line that runs from the anterior nearer to the front of the body stemma to the base of the clypeus. The wings are subhyalineexhibiting a glassy appearance. The wings are yellowish along the anterior margin of the superior pair and also around the base; the nervures the veins of the wings are rufo -fuscous.

Males are bright violet, and the antennae except for the first joint and tarsi are ferruginous. Like the queen, the wings are subhyaline imperfectly hyaline and the nervures are rufo-fuscous. The abdomen shows a bright green tinge when seen under certain light.

The head and anus are ventral. The integument is covered in spinules that are either isolated from each other or in short rows on the posterior somite and on the ventral surface. The body hairs are very short, measuring 0.

These spinules are either isolated or seen in near parallel rows. Several head hairs are present but they are small at 0. Each lobe has spinules and three sensilla simple sensory receptors around the anterior surface.

The ventral border only has two sensilla and a number of spinules, and on the posterior surface, there are several rows of spinules and three sensilla.

The mandibles have a central apical most distal plate or appendage from the body tooth which is clearly noticeable and sharp. The maxillae have lobes, and the labial palps sensory structures on the labium are knob-shaped. Aside from colour differentiation that was a key morphological character to distinguish I. For example, those that are found in very hot regions tend to be larger, whereas those found in regions of high humidity tend to be smaller than average.

For example, populations restricted to the coasts of Western Australia usually have pale setae, compared to most colonies throughout the country, which have the common blackish setae. The variation of the iridescence is, however, a consistent pattern found in other Iridomyrmex species with little distinction, making it a subtle character.

Shattuck further notes that populations found throughout the Northern Territory and South Australia have reduced pubescence on the first gastral tergite, but this is different elsewhere.

Its isolation has also allowed meat ants to form associations with neighbouring nests of the same species. In Queenslandthey are frequently encountered in the eastern regions, whereas their abundance is limited around the north and central parts. In the Northern Territory, specimens have been collected in the north and south regions but compared to other jurisdictions the ant is uncommon.

The meat ant shares its distribution with many other animals and insects, some of which may cause harm to the ant or rival it, such as the banded sugar ant Camponotus consobrinus. Meat ants are able to survive in dry areas if there is a rich supply of water and food resources such as honeydew and arthropod preyespecially along river banks, station properties and irrigated areas.

In the south coast of New South Wales, meat ants are mainly found in heath shrubland, but are absent from heavily timbered slopes and cannot build nests in quartz. Other areas where the ants do not occur include dense pastures, dense bushes, tropical rainforests and treeless areas. Their populations would later flourish and nests became numerous around houses after shrubs and trees were planted.

On the surface on the nest, workers clear the area of vegetation and cover the mound with gravel, but may use other materials that are available, including sand, pebbles, dead vegetation, eucalyptus fruits and twig fragments. The ant is a polydomous species, meaning that they live in more than one nest.

Hence, nests can be very old as suggested in one study. The regrowth of vegetation which shades the nest, soil damage or even a disease may wipe out a colony and leave the nest site completely abandoned. Satellite nests may diverge from their parent nests to become independent, as suggested by the antagonism of worker ants from different nests or when others are uninhabitable by insecticide treatment. After the eradication of a nest, satellite nests emerge nearby, and may sever their connections with the parent nest.

As most satellite nests have 11 holes and accept a queen of their own, a satellite nest may easily develop maturity in one year. Beneath the surface, there are widened circular vertical shafts which are 1. Below these shafts, the tunnels turn into irregular galleries with paths going outward and downward which form more galleries.

Almost all of these galleries are clustered together 15 to 20 centimetres 5. However, there is no known physical connection. Each gallery has a flat floor, a domed roof and is irregularly oval-shaped. A gallery is typically 1. Below the galleries are a small number of shafts in undisturbed soil with large, yet scattered chambers where the population remains during the winter.

Overall, a nest may dwell extremely deep beneath the soil as excavated nests are as deep as 3 metres 9. However, some ants such as the green-head ant Rhytidoponera metallica are not affected by the presence of meat ants and are still successful in finding food sources. They heavily rely on any food source and the impossibility of successfully defending it from other ants may have led to its peaceful coexistence with dominant species, including meat ants. This means green-head ants avoid conflict with meat ants.

In particular, Monomorium ants have been observed occupying baits regardless of the presence or absence of the meat ant. However, the presence of meats ants does not affect abundant species, and most of the time it is only Iridomyrmex ants that increase their foraging rate.

If present, meat ants rely on rapidly discovering food sources and retain their dominance so other ants cannot collect them, as well as exploitation and interference which helps displace other ants. The habitat meat ants live in may affect their dominance among the fauna. They are less successful in complex habitats and more successful in open areas, allowing workers to forage efficiently; for example, workers forage around rocks and collect food sources more successfully in contrast to those in vegetation.