Mating flight, metrosis, and semi-claustrality in the seed-harvester ant Pogonomyrmex salinus (Hymenoptera, Formicidae)

Here we examine dispersal, metrosis, and claustrality in the seed-harvester ant Pogonomyrmex salinus at an unusually large mating aggregation. We found that mode of queen dispersal from the mating aggregation is not a function of queen mass and that wing damage among queens was relatively rare. P. salinus is haplometrotic in the field and foundress queens placed together in forced associations eventually fight to the death. While queens of Pogonomyrmex salinus can survive claustrally, producing a single minim from their body reserves in the laboratory, fed queens produce up to four significantly larger minims along with concurrent larvae and pupae during the same period. Since queens forage in the field, we interpret claustrality as a secondary reserve strategy when foraging fails, and suggest that foraging is obligate for P. salinus queens in an overdispersed and temperate environment. Thus, nest founding strategies employed by P. salinus may be environmentally determined and represent a continuum between fully claustral and obligate foraging. We discuss our results with reference to theories of pleometrosis and claustral colony founding. Received 12 November 2004; revised 12 April 2004; accepted 29 July 2005.     

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.     

Male induction and function of workers' excitability during swarming in leaf-cutting ants (Atta and Acromyrmex) (Hymenoptera,Formicidae)

Worker excitation in species of Atta and Acromyrmex prior to and during the swarming of reproductives is initiated by secretions from the male mandibular glands. During this phase, workers are highly aggressive, attacking all foreign objects. Field data suggest that this behavior is not only advantageous in protecting sexuals as they swarm, but also to prime workers to execute queens trying to initiate colonies near mature colonies following nuptial flights.     

Temporal and spatial variation in reproduction in the facultatively polygynous antMyrmica tahoensis (Hymenoptera: Formicidae)

Summary Sex allocation was measured across six seasons in colonies of the facultatively polygynous antMyrmica tahoensis. The overall proportion of colonies that produced sexuals was constant throughout the study, but population-level sex ratios varied considerably. In 1991, 1993, and 1995, a significantly greater proportion of colonies produced females than in 1990, 1992, and 1994. Sex ratios were similar across six sites within individual years, suggesting a population-wide cause of sex-ratio variation. Individual colonies tended, on average, to produce similar sex ratios in consecutive years. Within-colony genetic relatedness, while strongly correlated with sex ratios within years, did not explain the year-to-year variation. It is suggested that extrinsic factors can limit the production of female sexuals, even when genetic relatedness is high enough to favor female production.     

Acromyrmex ameliae sp. n. （Hymenoptera： Formicidae）： A new social parasite of leaf-cutting ants in Brazil

The fungus-growing ants （Tribe Attini） are a New World group of〉 200 species, all obligate symbionts with a fungus they use for food. Four attine taxa are known to be social parasites of other attines. Acromyrmex （ Pseudoatta） argentina argentina and Acromyrmex （Pseudoatta） argentina platensis （parasites of Acromyrmex lundi）, and Acromyrmex sp. （a parasite of Acromyrmex rugosus） produce no worker caste. In contrast, the recently discovered Acromyrmex insinuator （a parasite of Acromyrmex echinatior） does produce workers. Here, we describe a new species, Acromyrmex ameliae, a social parasite of Acromyrmex subterraneus subterraneus and Acromyrmex subterraneus brunneus in Minas Gerais, Brasil. Like A. insinuator, it produces workers and appears to be closely related to its hosts. Similar social parasites may be fairly common in the fungus-growing ants, but overlooked due to the close resemblance between parasite and host workers.     

Phylogeny and bionomics of Lasius austriacus (Hymenoptera,Formicidae)

Summary Phylogenetic analysis based on sequence data of the mitochondrial COI gene confirms the species status of the recently described Lasius austriacus. The five haplotypes of L. austriacus do not cluster according to their geographic origin, indicating a recent gene flow among the populations. The molecular data corroborate the morphology based hypothesis that L. austriacus belongs to the Lasius (Lasius s.str.) brunneus group. The invasive species Lasius neglectus forms a sister taxon with L. turcicus, both next related to L. austriacus. Other phylogenetic relationships within the genus Lasius are in accordance with morphological data.First data on the bionomics of L. austriacus are discussed in context with its phylogenetic position. Based on gyne and male morphology, excavations of nests, pitfall trapping and observations in formicaries, we hypothesize that L. austriacus is a mainly hypogaeic, monogynous species with nuptial flight. These are characters of the Lasius brunneus group in general, except the polygynous-polycalic, intranidally copulating L. neglectus. Aggression tests, however, revealed non-aggressive behaviour (antennation) between separated L. austriacus populations, but pronounced interspecific aggression against L. neglectus. This confirms the species status of L. austriacus and indicates a reduced level of intraspecific aggression, similar to L. neglectus.The status of L. austriacus as a native species in Central Europe is confirmed.Received 10 February 2003; revised 24 June 2003; accepted 23 July 2003.     

Labial gland disease in the genusFormica (Formicidae,Hymenoptera)

Summary Labial gland disease causes swelling of the labial glands in the pupal stage inFormica species. The resulting adults have enlarged thoraces and are called secretergates (Wasmann's pseudogynes). They infect the younger larvae, probably during feeding. The agent of the disease is unknown. It soon becomes non-infectious outside the labial glands. Diseased females were not seen to feed larvae. Yet their offspring contained secretergates. The discrepancy remains unexplained.     

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.     

Loss of mating flight and shift in the pattern of carbohydrate storage in sexuals of ants (Hymenoptera; Formicidae)

Summary In ants, energy for flying is derived from carbohydrates (glycogen and free sugars). The amount of these substrates was compared in sexuals participating or not participating in mating flights. Results show that in participating females (Lasius niger, L. flavus, Myrmica scabrinodis, Formica rufa, F. polyctena, F. lugubris), the amount of carbohydrates, especially glycogen, was higher than in non-participating females (Cataglyphis cursor, Iridomyrmex humilis). Similarly, male C. cursor and I. humilis which fly, exhibit a much higher carbohydrate content than do the non-flying females of these species. Furthermore, the quantity of carbohydrates stored was generally higher in males than in females for each species. These results are discussed with regard to the loss of the nuptial flight by some species of ants.     

Ontogenetic origin of mermithogenic Myrmica phenotypes (Hymenoptera, Formicidae)

Entomo-pathogen parasites typically induce alternative ”parasitogenic” phenotypes in ants and other insects. However, the basis of generated developmental changes is poorly understood. Parasitic mermithid nematodes also cause the formation of three discrete and aberrant morphologies within Myrmica ants. These have been called ”worker-like” (”mermithergate”), ”intermorphic” (”gynaecoid mermithergate”) and ”gyne-like” (”mermithogyne”) and their formation has been attributed to infection of worker- and queen-presumptive larvae, respectively. In order to better understand the developmental mechanisms that lead to the formation of these alternative parasitogenic phenotypes we observed allometric patterns of parasitogenic Myrmica gallienii phenotypes in comparison with uninfected workers and gynes from the same nests. It was revealed that the three discrete morphologies of parasitogenic female phenotypes did not differ significantly from each other in their scaling indicating that these were trapped in the same developmental pathway. Infected individuals scaled according to basically gyne-like allometry, however significantly differed from workers in their scaling. Based on the observed scaling patterns we herein raise an alternative explanation according to which both ”mermithergate”, ”gynaecoid mermithergate” and ”mermithogyne” Myrmica phenotypes develop from the same type of larvae, namely from the queen-presumptive larvae and their formation, therefore, is rendered as a diverging process. According to the mechanism we propose effect of nematodes may turn out to be the determining factor in the formation of alternative parasitogenic morphologies. Received 16 June 2008; revised 15 September and 7 November 2008; accepted 18 November 2008.     

Oviposition behavior of an ant-parasitizing fly,Neodohrniphora curvinervis (Diptera: Phoridae), and defense behavior by its leaf-cutting ant hostAtta cephalotes (Hymenoptera: Formicidae)

This study examines the oviposition behavior of the phorid parasitoid Neodohrniphora curvinervisand the antiparasitoid defense behavior of its leafcutting ant host Atta cephalotes. N. curvinervisfemales are diurnal sit- and- wait parasitoids that attack only outbound foragers of head width 1.6 mm or greater. Females deposit a single egg through the foramen magnum of each host successfully parasitized. Pursuit of hosts is usually initiated when an outbound forager of acceptable size passes by a parasitoid perch site. Individual foragers defend themselves against pursuing parasitoids by outrunning them along the foraging trail or by standing their ground and fending them off with their legs,antennae, and mandibles. At the colony level, susceptible foragers are protected against parasitism by a shift in the forager size distribution toward smaller unsusceptible sizes during the day when parasitoids are active and toward larger sizes at night when parasitoids are inactive. The frequency of parasitism of susceptible foragers was 15%, which is more than five times the frequency found in another system involving the phorid parasitoid Apocephalus attophilusand the leafcutting ant host Atta colombica.We offer several possible explanations for such differences in the frequency of parasitism and also examine reasons for the high incidence of superparasitism (19%) observed in the system studied.     

Mechanisms affecting load size determination inAtta cephalotes L. (Hymenoptera,Formicidae)

Summary The size of leaf-cutting ant foragers correlates more closely with the weight than the area of the leaf fragments they cut. This implies that the mechanism of load size determination is not a simple function of body geometry. Ants were found not to adjust the radius of cut to compensate for experimental changes in leaf weight during the process of cutting. However, ants changed their cutting behaviour according to the thickness of the region of the fragment being cut. Ants decreased their cut radius when cutting through leaf veins. Similarly, when cutting artificial laminae of varying thicknesses, ants reduced the cut arc radius when traversing thicker regions.Cut radius was not principally controlled by the position of the rear legs nor the overall body length but by mechanisms associated with the head and thorax of the ant.     

Colony structure of a bamboo-dwellingTetraponera sp. (Hymenoptera: Formicidae: Pseudomyrmecinae) from Malaysia

Summary The colony structure of the bamboo-inhabiting SE-Asian pseudomyrmecine antTetraponera sp. PSW-80 nearattenuata F. Smith was investigated at the Ulu Gombak Field Studies Centre, Selangor, West-Malaysia. Based on the dissection of 54 stem internodes from 12 different culms of the large bambooGigantochloa scortechinii Gamble and on the mapping of three colonies, the following demographic characteristics emerge. The colonies are monogynous but highly polydomous (at least up to 36 internodes and up to 9 stems occupied) and very populous for a pseudomyrmecine not involved in an ant-plant mutualism. One completely censused colony had 6953 adult workers and 2079 alates (adults plus pupae). The single queen suppresses gyne development in her own nest and, to a lesser extent, in other nests within the same stem. The overall numerical sex ratio was 0.961 (females:males), the investment sex ratio, 2.931, i.e., almost exactly the 31 ratio expected for a monogynous outbred hymenopteran in which the colony queen also produces all the male offspring. Brood is distributed to all other nest chambers from the queenright chamber. The symbiotic pseudococcids (Kermicus wroughtoni Newstead) are present in all inhabited internodes, with small early instar individuals prevailing numerically by far over the larger stages. The rieht well secluded honeydew supply within the internode and the efficient architectural protection provided by the internode wall (access usually only through a 2 × 3 mm-hole) allowT. sp. PSW-80 to reach an unusually large colony size without being an aggressive and protective plant mutualist like other members of its subfamily with similar demographic features.     

Sex ratio characteristics in Ixodes rubicundus (Acari: Ixodidae), the Karoo paralysis tick

The sex ratio is an important parameter which characterizes the state and dynamics of natural populations of animals. Although ixodid ticks are specialized ectoparasites, most species are bisexual and are characterized by a 1:1 sex ratio for their progeny. In natural populations and even in laboratory colonies, biased sex ratios are often observed. Ixodes rubicundus, the Karoo paralysis tick, parasitizes domestic stock and wild ungulates in South Africa. Adults quest from vegetation, can mate off or on the host and males are seldom parasitic. We hypothesized that the sex ratio for I. rubicundus would be 1:1 when observed directly in the progeny but that it would be strongly biased towards females in samples of parasitic adults. The results mostly supported the hypothesis but it was also shown that unexplained and unpredictable variations can occur. On hosts, females dominated strongly, except on adult angora goats where the sex ratio was biased in favour of the males. This disparity may be related to a greater retention of males in the coarse, curly hair of angora goats compared to the other hosts. Monthly variations in the sex ratios of the tick on hosts are believed to be related to the large fluctuations in sex ratios of questing ticks.     

Colony-level single-sex alate production inPheidole ceres andPheidole pilifera (Formicidae: Myrmicinae)

Summary Patterns of colony-level reproductive investment were estimated inPheidole ceres andP. pilifera. Both species exhibited strong degrees of colony-level, single-sex alate production, but differed in the extent to which colonies produced only one sex. The proportion of reproductive investment allocated to males was negatively correlated to the total reproductive allocation in both species.     

Raiding and Foraging Behavior of the Blood-Red Ant,Formica sanguinea Latr. (Hymenoptera,Formicidae)

In this paper we report the results of a detailed study on the behavioral ecology of slave raiding and foraging activity in the European blood-red ant, Formica sanguinea Latr. The field study was conducted over an unbroken period of 78 days, during which the activity of two dulotic colonies of this facultative slave-maker was observed for 10 h each day. It was possible to observe 26 raids distributed over 23 days, among which 18 were followed by the sacking of nests belonging to the species F. cunicularia, F. fusca, and Lasius emarginatus, whereas 8 failed. Simple, continuous, and simultaneous raids occurred. We recorded the timing, frequency, distance, and direction of slave raids, including the number of participants and the type of booty. Particular attention was devoted to the scouting behavior and raiding organization. Moreover, every day, we observed foraging and predatory behavior, during which adult insects (mainly ants), seeds, and berries were retrieved to the dulotic colonies. On the basis of our observations F. sanguinea seems to be a very efficient slave-maker and predatory species of the Raptiformica subgenus. Moreover, its dulotic behavior may be regarded as a continuation and an expansion of its foraging and predatory behavior, as predicted by Darwin's hypothesis for the origin and evolution of slavery in ants.     

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.