Predatory Behavior in Dominant Arboreal Ant Species: The Case of Crematogaster sp. (Hymenoptera: Formicidae)

Crematogaster sp. is a dominant arboreal ant species that captures and retrieves very large prey. Hunting workers forage collectively thanks to short-range recruitment. They detect prey by contact, then rapidly attack, seizing small prey by the body and large prey by a leg. In this study, almost all the active prey were spread-eagled by several workers, even when small enough to permit a single worker to easily master them. While certain workers spread-eagled the prey, others deposited venom on the prey body using their spatulated sting (topical action of the venom). The well-developed arolia on the pretarsus of workers' legs have crucial importance for the success of prey capture (spread-eagling) and transport in an arboreal habitat. These results are compared with those known for other arboreal-dwelling generalist predator ant species.     

Effects of experience on food-searching behavior in the antFormica schaufussi (Hymenoptera: Formicidae)

Foragers of the ant Formica schaufussitend to return to and search at a site of a previous food find. The search tactic employed by a forager on its return trip is related to the type of food previously encountered: search is more persistent in response to carbohydrate than to protein food. Using different reinforcement schedules with carbohydrate and protein food rewards, we show that, on a short-term as well as on a long-term basis, the basic pattern of search observed in naive foragers is only slightly modified through foraging experience. Foragers do not increase their search effort or adjust their search pattern when either type of food is systematically renewed on a fixed reward schedule and thus do not seem to be able to learn to assess the food predictability. Collective responses that could compensate for this lack of individual flexibility and increase foraging efficiency at the colony level are discussed.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

Respiratory patterns in nurses of the red wood ant Formica polyctena (Hymenoptera, Formicidae)

Summary Cycles of discontinuous gas exchange (DGCs) and abdominal ventilatory movements were studied in nurses of red wood ant, Formica polyctena, using an electrolytic respirometer and an infra-red (IR) gas analyser or flow-through respirometry. Both respirometry systems were combined with an IR actographic device based on IR-emitting and IR-sensor diodes. After recovering from handling and apparatus stress, lasting 1–3 h, completely motionless intact ants displayed regular DGC. After decapitation the ants displayed DGC whose frequency was somewhat lower than that of the intact individuals (7.17 ± 0.79 mHz and 10.43 ± 01.12 mHz, respectively). In headless ants, there occurred continuous slow movements of legs. Bursts of carbon dioxide in the intact and in the headless ants always coincided with a bout of telescoping movements (contractions) of abdominal segments, which was interpreted as active ventilation. During interburst periods, the headless ants exhibited telescoping movements characterised by rapid protraction, lasting 0.07–0.09 s, followed by a slow retraction of segments, suggesting passive ventilation. The intact ants were very sensitive to the flowing air and tended to be continuously active during flow-through respirometry. The decapitated ants, on the contrary, were insensitive to the air current.Received 26 March 2003; revised 30 July 2003; accepted 6 August 2003.     

Attraction of the sexes inFormica lugubris Zett (Hymenoptera: Formicidae)

Summary Sexuals ofFormica lugubris fly to mating places, where females attract males by using a sex pheromone. Females collected on the nest surface before departing on a mating flight are much less attractive than those collected on the mating place after the mating flight, suggesting that the mating flight triggers the release of the sex pheromone. Olfactory cues are essential for males to locate females while they patrol. Males probably use visual cues to locate females once they have alighted nearby them. Males are also attracted by aggregations of other males on the ground, probably because one or several females are likely to be close to male aggregations.     

Feinstruktur des Komplexauges der Roten Waldameise,Formica polyctena (Hymenoptera,Formicidae)

Zusammenfassung Die Analyse der Feinstruktur des Komplexauges der Ameise Formica polyctena ergab, daß der dioptrische Apparat der insgesamt 750 Ommatidien aus einer lamellierten Cornealinse und einem euconen Kristallkegel besteht. Zwei Hauptpigmentzellen umgeben schalenförmig den Kristallkegel, 8 Nebenpigmentzellen schirmen das Ommatidium in seiner ganzen Länge von der Cornea bis zur Basalmembran ab. Jedes Ommatidium besteht in seinem distalen Teil aus 8 Retinulazellen, 2 gegenüberliegenden schmalen und je 3 gegenüberliegenden großen Sehzellen. Weiter proximal tritt eine 9. Retinulazelle hinzu. Die Mikrovillisäume der Sehzellen verschmelzen zu einem zentralen Rhabdom. Im distalen Teil sind die Mikrovilli in 3 Richtungen angeordnet. Es wird im besonderen die Orientierung der Mikrovilli zur Augenlängsachse und zur Ommatidien-Symmetrieachse untersucht. Auch die 9. Sehzelle bildet Mikrovilli. Das Rhabdom endet an 4 basalen Pigmentzellen.Auf den Mikrovillisaum folgt im Querschnitt des dunkel adaptierten Ommatidiums ein Kranz von großen intrazellulären Vakuolen. Die anschließende cytoplasmatische Zone der Retinulazellen enthält viele Pigmentgranula und Mitochondrien; multivesikuläre und multilamelläre Körper sowie Golgiapparate treten nur selten auf. Die funktionelle Bedeutung des Ommatidienaufbaues und die Verteilung der Organellen bei Dunkeladaptation werden diskutiert.

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The fine structure of the complex eye of the ant Formica polyctena (Hymenoptera, Formicidae)

Summary The fine structure of the compound eye of the ant Formica polyctena was investigated. The eye consists of a total of 750 ommatidia, each containing a dioptric apparatus of a lamellated cornea lens, a eucone-type crystalline cone, and 8 long pigment cells which surround the ommatidium for its total length, i.e. from the cornea to the basal membrane. Each ommatidium has in its distal portion 8 retinula cells—6 large plus 2 small ones. The retinula cells are arranged in such a way that 3 pairs of large cells, and the one pair of the small cells lie opposite each other. Further proximally, a 9th retinula cell is encountered. The fused, centrally located rhabdom is built up by the microvilli of all nine retinula cells. The rhabdom ends at 4 basal pigment cells. In dark adapted ommatidia, a ring of large intracellular vacuoles is to be seen immediately peripherally to the rhabdom. The outer, cytoplasmic zone of the retinula cells contains many pigment granules and mitochondria; multivesicular bodies, onion bodies and Golgi apparatus are of relatively rare occurrence. The functional significance of the ommatidial structure and the arrangement of the cell organelles in the dark adapted condition are discussed.

Für technische Hilfe danke ich Frau Langer und Frl. Müller, Herrn Prof. H. Markl für die Durchsicht des Manuskripts.     

Foraging Area Marking in Two Related Tetramorium Ant Species (Hymenoptera: Formicidae)

The related ants Tetramorium caespitum and T. impurum mark their foraging area in a species-specific, home range and short-lasting manner. Indeed, ants reaching a new area have a slow linear speed which increases during the marking. Conspecific ants are arrested and attracted by marked areas, while heterospecific ants are reluctant to visit them. However, when the latter do visit marked areas, they move more quickly and less sinuously than conspecific ants and do not stay on the areas. The marking is performed in about 3 min by T. caespitum and in 3 to 6 min by T. impurum. If not reinforced, the marking vanishes in the same time intervals. Neither poison gland nor last sternite extracts reproduce the activity of naturally marked areas, whereas a Dufour gland extract does exactly that. Foraging ants touch the ground with the tip of their gaster. Consequently, we can postulate that the workers mark their foraging area with the contents of this gland, which is associated with the sting apparatus, and that they deposit with the extremity of the gaster. Alien conspecific ants are seldom aggressive to one another, even on marked areas. When encountering each other on unmarked areas, heterospecific ants present some aggressive reactions. On marked areas, their aggressiveness is enhanced and intruder ants are restless, while resident ones walk freely. On ground marked by T. impurum, ants of this species are more aggressive than antagonistic T. caespitum workers. The marking of foraging areas thus induces defense against heterospecifics but not against conspecific ants.     

Relationships between the ant Brachymyrmex obscurior (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae)

Summary. In central Mexico, the ant Brachymyrmex obscurior Forel feeds on nectar produced by extrafloral nectaries of Acacia pennatula (Schlecht. & Cham.) Benth. However, no studies have determined whether the ants visitation is related to plant nectar availability and whether ants protect A. pennatula from herbivory. The objectives of this 2-yr study (2000–2001) were to assess whether seasonal changes in ant visitation coincide with extrafloral nectar productivity in A. pennatula and to determine whether ants protect the plant. At the end of the dry season (April–June) B. obscurior was the only ant species on A. pennatula and extrafloral nectar production is limited to this period. Exclusion experiments, performed at the end of the dry season showed that A. pennatula did not receive a protective benefit when visited by ants. Branches with ants and branches where ants are excluded had similar numbers of the nonmyrmecophile leafhopper Sibovia sp. which was the only herbivore observed under natural conditions.Received 24 March 2004; revised 4 September 2004; accepted 8 September 2004.     

Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Oral trophallaxis in adult leaf-cutting ants Acromyrmex subterraneus subterraneus (Hymenoptera, Formicidae)

Adult leaf-cutting ants of the subspecies Acromyrmex subterraneus subterraneus were fed with an Evans Blue dye solution, which allowed the investigation of subsequent exchange of liquids between ants by oral trophallaxis. Trophallactic behavior was filmed and the antennation patterns of donor and recipient ants were described. The ants’ crop capacity was measured following ad libitum feeding on dye solution. Ants previously fed on the dye solution (donors) were placed individually with unfed ants of the same caste (recipients) and the amount of dye solution passed from the donor to the recipient by oral trophallaxis was measured after one hour. The volume received was 0.26 ± 0.15μL (mean ± SD) and the residual volume of dye in the crop of the donors was 0.49 ± 0.23μL. There were 38 trophallactic events recorded for 50 pairs of ants. Trophallaxis was observed from 2.3 min to 21.5 min after initial exposure, with a mean latency of 8.4 ± 5.6 min. The mean duration of a trophallatic event was 2.3 ± 1.3 min. As far as we know, this is the first time that trophallaxis in leaf-cutting ants has been described and quantified. Received 2 December 2005; revised 6 April 2006; accepted 10 April 2006.     

Raiding behavior of the dulotic antChalepoxenus muellerianus (Finzi) in the field (Hymenoptera: Formicidae,Myrmicinae)

Summary Little information is available regarding the raiding behavior in nature of dulotic ants belonging to the tribeLeptothoracini. Between July 17 and 24, 1991, several raids ofChalepoxenus muellerianus (Finzi) were observed in nature near Tignale/Lago di Garda, Italy. Apparently, this species raids frequently during its summer raiding season (5 raids were observed during 7 days spent observing 8 colonies). A singleChalepoxenus colony sometimes raids more than one host colony more or less simultaneously. Observations during which oneChalepoxenus colony raided another and captured slavemaker brood indicate that intraspecific raids can occur either accidentally, or as a result of competition or territoriality when there is a sufficient dense slavemaker population.     

Integration raids in the Amazon ant Polyergus rufescens (Hymenoptera, Formicidae)

Summary. Groups of enslaved Formica fusca workers from mixed colonies of Polyergus rufescens with numerous slave workforce tend to split off and found small and almost homospecific nests around the main nest, with at least some of them connected with the latter with underground passages. Their inhabitants are able, at least temporarily, to adopt young F. fusca gynes. P. rufescens invades these satellite nests in a manner similar to the normal slave raids, and carries the slaves back to the main nest. The supposed evolutionary cause of this behaviour is to keep integrity of mixed colonies and prevent possible emancipation of slaves.Received 18 August 2004; revised 27 September 2004; accepted 11 October 2004.     

Longitudinal observation of attine (Atta cephalotes isthmicola) ants (Hymenoptera; Formicidae)

A colony of the leaf-cutting Attine (Atta cephalotes isthmicola) ants was cultured and observed in the laboratory. During the 114 month life of the colony, a maximum garden volume of 54 L. was attained. The colony used a total of 139,902 g of substrate at a maximum rate of 2,660 g per month and occupied one hundred forty-four 1.25 L. chambers. Observations on the colony ranged from behavior of the castes to biochemistry and the absence of cholesterol. The dramatic display quality of Attine colonies for zoological exhibit is discussed.     

Respective contributions of leader and trail during recruitment to food inTetramorium bicarinatum (Hymenoptera: Formicidae)

Summary This paper describes the food recruitment strategy of the antTetramorium bicarinatum, at both the individual and collective levels. The general organisation of recruitment used by this species during the exploitation of sucrose solutions shows similarities with group recruitment described for other species. However, our experiments demonstrate that inT. bicarinatum, the first trail laid by a recruiter during its return trip to the nest is more efficient than in other species using group recruitment, for exampleTetramorium impurum. Moreover, the efficiency of the first trail of aT. bicarinatum recruiter is comparable with that ofTapinoma erraticum, a species that uses mass recruitment. Despite the efficiency of the trail, choice experiments show that the recruited workers prefer to follow the leader rather than the first trail, suggesting the emission of a more attractive signal by the leader on its way back to the food. The function of the leader in this strategy is discussed in terms of collective decisions.     

Three new species of ants (genus <Emphasis Type="Italic">Aphaenogaster</Emphasis>, Hymenoptera: Formicidae) from Turkey

Three new species of the ant genus Aphaenogaster (A. aktaci sp. n., A. maculifrons sp. n., A. radchenkoi sp. n.) are described from Turkey and their taxonomic positions are discussed. Aphaenogaster aktaci sp. n. belongs to the obsidiana group, A. maculifrons sp. n. to subterranea group and A. radchenkoi sp. n. to pallida group of the Attomyrma subgenus. Key characteristics for identification of the species and data on their ecology are given.     

Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria

The present catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria is made on a base of critical reconsideration of literature (covering the period from 1892 till 2009 and part of 2010) as well as on examination of the authors' and several museum's collections. A lot of data were omitted in the previous Bulgarian monograph on ants, lots of new data were recently added and many important additions and alterations were made due to taxonomic revisions of Eurasian Formicidae during the last three decades. Two new species are reported for the country [Temnothorax graecus (Forel, 1911) and Temnothorax cf. korbi (Emery, 1924)].This catalogue contains a list of 163 ant species belonging to 40 genera of 6 subfamilies now known from Bulgaria. Synonyms and information on the previously reported names in relevant publications are given. Known localities of the species are grouped by geographic regions. Maps with concrete localities or regions for each species were prepared. The conservation status of 13 ant species is given as they are included in IUCN Red List of Threatened Species and Bulgarian Biodiversity Act. In comparison with adjacent Balkan regions the ant fauna of Bulgaria is quite rich and its core is composed of South European elements.