Energetics of newly-mated queens and colony founding in the fungus-gardening ants Cyphomyrmex rimosus and Trachymyrmex septentrionalis (Hymenoptera: Formicidae)

Abstract.  The energetics of colony founding is investigated in the fungus gardening ants (Attini) Trachymyrmex septentrionalis and Cyphomyrmex rimosus . Similar to most ants, inseminated queens of these two species found nests independently unaccompanied by workers (haplometrosis). Whereas most ant founding queens seal themselves in a chamber and do not feed when producing a brood entirely from metabolic stores (claustral founding), the majority of fungus gardening ants must forage during the founding phase (semiclaustral founding). Laboratory-reared T. septentrionalis individuals comprise 84 dealate females collected after mating flights in June 2004. Twenty are immediately killed to obtain values for queen traits and another 20 after worker emergence for queen, fungus garden and worker traits. Cyphomyrmex rimosus comprise 22 dealate females collected in June 2005; ten of which are immediately killed and similarly prepared. Newly-mated T. septentrionalis queens have 25% of their dry weight as fat; whereas newly-mated C. rimosus queens contain 11% fat. These amounts are 50–75% less than most independently founding ant species. Trachymyrmex septentrionalis queens lose merely 5% of their energetic content during colony founding, whereas the total energetic content of their brood is more than three-fold the amount lost by the queen. Incipient T. septentrionalis colonies produce approximately half as much ant biomass per gram of fungus garden as do mature colonies. Similar to most ants, T. septentrionalis produces minim workers that are approximately 40% lighter than workers from mature colonies. Regardless of their size, T. septentrionalis workers contain much lower fat than do workers of claustral species. These data indicate that fungus gardening is adaptive because colonies can produce much cheaper offspring, making colony investment much lower.     

Capital and income breeding and the evolution of colony founding strategies in ants

Ability to store resources that will be used for reproduction represents a potential life history adaptation because storage permits feeding and reproduction to be decoupled spatially and/or temporally. The two ends of a continuum involve acquiring all resources prior to reproduction (capital breeding) or acquiring all resources during the reproductive period (income breeding). Traditional life history theory examines tradeoffs between costs and benefits of such strategies, but this theory has not been integrated into life history studies of ants, even though founding queens have the analogous strategies of fully claustral (capital breeding) and semi-claustral (income breeding). This study demonstrates that facultatively semi-claustral queens of the seed-harvester ant Pogonomyrmex desertorum exhibit phenotypic plasticity during colony founding because unfed queens produced few, small minims, whereas ad libitum fed queens produced larger, heavier minims and additional brood. Fed queens also lost less mass than unfed queens despite their producing more brood. Overall, foraging provides queens with a suite of benefits that likely offset potential negative effects of foraging risk. Life history studies across a diverse array of taxa show that capital breeding is consistently associated with low availability and/or unpredictability of food, i.e., environmental conditions that favor prepackaging of reproductive resources. Such a broad and consistent pattern suggests that similar factors favored the evolution of fully claustral (capital breeding) colony founding in ants. Overall, these data suggest that ant researchers should revise their conventional view that fully claustral colony founding evolved because it eliminated the need for queens to leave the nest to forage. Instead, colony founding strategies should be examined from the perspective of environmental variation, i.e., availability and predictability of food. I also provide a functional scenario that could explain the evolution of colony founding strategies in ants. Received 16 November 2005; revised 1 March 2006; accepted 29 March 2006.     

Competition and kin discrimination in colony founding by social Hymenoptera

Summary After mating, queens of social wasp and ant species sometimes band together to start a new colony cooperatively. I assume these queens sequentially encounter potential nest sites that may or may not already contain a queen. Whether to remain at a given site or to leave in hopes of finding a better site is modelled using dynamic programming. The results suggest that discriminating competitive ability is more valuable than discriminating kinship. Wasps, which have a high survival rate in transitions between nest site encounters and in which pleometrosis seems to have a consistently high benefit, are predicted to discriminate both competitive ability and kinship of potential partners. Ants, which have lower survivorship and variable benefits, are predicted to show conditional joining behavior (sometimes based on discriminating competitive ability, but almost never based on discriminating kinship). A survey of the literature supports the model with respect to the predictions on kin discrimination in both groups and on conditional joining behavior in ants. However, whether partners are joined based on perceptions of competitive ability needs more tests.     

Dynamic changes in host–virus interactions associated with colony founding and social environment in fire ant queens (Solenopsis invicta)

The dynamics of host – parasite interactions can change dramatically over the course of a chronic infection as the internal (physiological) and external (environmental) conditions of the host change. When queens of social insects found a colony, they experience changes in both their physiological state (they develop their ovaries and begin laying eggs) and the social environment (they suddenly stop interacting with the other members of the mother colony), making this an excellent model system for examining how these factors interact with chronic infections. We investigated the dynamics of host – viral interactions in queens of Solenopsis invicta (fire ant) as they transition from mating to colony founding/brood rearing to the emergence of the first workers. We examined these dynamics in naturally infected queens in two different social environments, where queens either founded colonies as individuals or as pairs. We hypothesized that stress associated with colony founding plays an important role in the dynamics of host – parasite interactions. We also hypothesized that different viruses have different modalities of interaction with the host that can be quantified by physiological measures and genomic analysis of gene expression in the host. We found that the two most prevalent viruses, SINV‐1 and SINV‐2, are associated with different fitness costs that are mirrored by different patterns of gene expression in the host. In fact SINV‐2, the virus that imposes the significant reduction of a queen's reproductive output is also associated with larger changes of global gene expression in the host. These results show the complexity of interactions between S. invicta and two viral parasites. Our findings also show that chronic infections by viral parasites in insects are dynamic processes that may pose different challenges in the host, laying the groundwork for interesting ecological and evolutionary considerations.     

Biological constraints and colony founding in the polygynous invasive ant Lasius neglectus (Hymenoptera, Formicidae)

Summary. The polygynous invasive ant Lasius neglectus was described from Budapest, Hungary, as an unicolonial species, with no apparent colony barriers, and inferred intra-nidal mating without a nuptial flight. Here we analyze additional morphological characteristics of gynes, their physiological condition at emergence and at the time of mating and we describe the productivity of different types of colony founding in the laboratory. A low increase in dry weight and in fat content from emergence to mating indicates that gynes can only succeed in dependent colony founding. However, the queen-worker thorax volume ratio is typical of a species with independent colony founding and we were able to demonstrate independent colony founding in the laboratory (both haplo- and pleometrotic). Brood development in independent founding is rapid and the number of nanitics higher than in other Lasius species. Both colony budding and dependent colony founding could also be demonstrated in the laboratory. Worker oviposition was absent. The carbohydrate content of newly mated queens is consistent with the observed loss of mating flight of this species. However the relative wing area clearly indicates that L. neglectus queens should be able to fly. Therefore, both queen morphology and physiological state at maturity show that L. neglectus is intermediate between a monogynous, free-living, non pest ant and a multiqueened (polygynic) invasive tramp ant. This neglected ant thus offers interesting opportunities to study the origin of unicoloniality and the spread of this species in northeast Spain.     

Flexibility of first brood production in a claustral ant, Camponotus japonicus (Hymenoptera: Formicidae)

The numbers of first-batch eggs and workers (minims) in a carpenter ant Camponotus japonicus are not fixed but are regulated in response to various internal and external factors of founding queens. The initial body weight of founding queens was positively correlated with the number of minims. The amount of queen weight loss during the founding period was highly correlated with the number of minims. The numbers of first-batch eggs and minims for artificially fed queens were significantly greater than those for control group queens and for unfed queens, respectively. The number of eggs laid by a queen increased when eggs deposited in the early stage of founding were removed. In contrast, artificial addition of alien eggs reduced the number of eggs laid by a queen. The adaptive implications of this flexibility in the production of the first-batch brood in C. japonicus are discussed. Received: December 18, 2000 / Accepted: March 19, 2001     

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.     

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.     

Testing the reproductive groundplan hypothesis in ants (Hymenoptera: Formicidae)

The evolution of complex societies with obligate reproductive division of labor represents one of the major transitions in evolution. In such societies, functionally sterile individuals (workers) perform many of fitness‐relevant behaviors including allomaternal ones, without getting any direct fitness benefits. The question of how such worker division of labor has evolved remains controversial. The reproductive groundplan hypothesis (RGPH) offers a powerful proximate explanation for this evolutionary leap. The RGPH argues that the conserved genetic and endocrinological networks regulating fitness‐relevant behavior (e g. foraging and brood care) in their solitary ancestors have become decoupled from actual reproduction in the worker caste and now generate worker behavioral phenotypes. However, the empirical support for this hypothesis remains limited to a handful of species making its general validity uncertain. In this study, we combine data from the literature with targeted sampling of key species and apply phylogenetically controlled comparative analysis to investigate if the key prediction of the RGPH, namely an association between allomaternal behavior and an allomaternal physiological state holds in the largest and most species‐rich clade of social insects, the ants. Our findings clearly support the RPGH as a general framework to understand the evolution of the worker caste and shed light on one of the major transition in evolutionary history.     

Scaling of body weight and fat content in fungus-gardening ant queens: does this explain why leaf-cutting ants found claustrally?

Offspring traits are among the most important life history traits, yet we lack an adequate understanding of their role in social insect life history evolution. Colony founding in the fungus-gardening ants (Tribe Attini) is different from most other ant species because the queens forage during the founding phase. Queens of the most derived genus, Atta, are the only attines that exhibit the more typical claustral founding, where the queens seal themselves in a below-ground chamber and produce their first generation of workers with only body fat reserves. Here I report the dry weights, fat content and energetic value of newly mated queens of ten attine species. Published phylogenies were used to make inferences on the evolutionary transitions in this clade. It appears that the evolution of fungus-gardening was associated with the manufacture of smaller, leaner queens as basal taxa are characterized by small bodies that contain relatively less fat than derived taxa. Moreover, there appears to be an allometric function between fat, energetic content and dry weight, which means that for fatter and claustral queens to develop, they also must become larger.     

Phylogeny of ants (Formicidae) based on morphology and DNA sequence data

In order to reconstruct ants' phylogeny, we analysed DNA sequences for two nuclear genes, abdominal-A and Ultrabithorax, from 49 species of ants and two outgroups. As these genes control the development of the first segments of the abdomen in insects, which are very variable in ants (petiole, postpetiole, and gaster constriction), we hypothesized that the morphological variations between the subfamilies may be correlated with mutations of some abd-A or Ubx regions. Contrarily to our hypothesis, these sequences are highly conserved. The differences observed concern mainly third codon positions and present some saturation. Phylogenetic reconstructions were carried out using the genetic raw sequence data and by combining them with a set of morphological data (Total Evidence). Relations among subfamilies of ants remains poorly resolved with molecular data only, but adding these data to morphological characters confirms and reinforce the topology of : a Poneroid complex [Ponerinae, Cerapachyinae, Leptanillinae and army ants], a Formicoid complex [Dolichoderinae, Formicinae] and a Myrmecoid complex [Myrmicinae, Myrmeciinae, Pseudomyrmecinae, Nothomyrmeciinae]. Our molecular results allow resolution near the branch tips and three subfamilies (Dolichoderinae, Formicinae and Pseudomyrmecinae) always appear as monophyletic. The Formicinae and the Dolichoderinae have close relationships. The Camponotini appear as a strong clade inside the Formicinae. The Ponerinae are separated in two parts: the Ectatommini and all other tribes. The Cerapachyinae, Dorylinae, and Ecitoninae belong to the same clade, the Cerapachyinae being confirmed in their subfamily status. The Myrmicinae appears to be very heterogeneous, with the Attini forming a very stable and well-separated group.     

Haplometrosis and pleometrosis in the antAcromyrmex striatus (Hymenoptera: Formicidae)

Summary Laboratory tests with mated females of the leaf-cutting antsAcromyrmex striatus (Myrmicinae, Attini) were conducted to determine if the colony foundation is a solitary or a mutualistic process. We have also tested the effect of foundresses density and number of available areas in this process. Three bioassays were performed: with single foundresses; with paired foundresses; and with groups of 8 to 18 foundresses. The results suggest that the foundation can be by haplometrosis or pleometrosis, possibly depending on physical distances between or densities of foundresses. Foraging activity was common in haplometrotic queens and in foundress pairs, but no foraging occurred in groups of foundresses.     

Negentropy and the evolution of chemical recruitment in ants (Hymenoptera: Formicidae)

The applicability of the concept of negentropy to the study of evolution is tested by tracing the phylogeny of the information content of the communication systems used for recruitment in the Formicidae. Data from the literature is reviewed and ordered around theoretical evolutionary routes based on increasing negentropy of the chemical communication systems used. One pathway presupposes the development of chemical orientation prior to chemical attraction, i.e. signalling the presence of food; whereas the others begin with the chemical attraction of nestmates without orientation with chemical cues to the food. The results suggest that the Myrmicinae, Ponerinae, Dolichoderinae, Pseudomyrmycinae and possibly Dorylinae evolved chemical recruitment via chemical attraction, whereas the Formicinae evolved chemical recruitment via chemical orientation. These alternative patterns permit the establishment of phyletic trends based on glandular evolution for chemical recruitment. The results are compared with phylogenetic studies based on morphological and chemical characters, and it emerges that the recruitment behaviour proves to be a particularly conservative characteristic, in that it is evident in the same form in a wider range of species. The conclusion is reached that the negentropy content of living systems is a powerful tool in phylogenetic studies, even of small taxonomic groups.     

Physiological changes during colony establishment in the termite Hodotermes mossambicus (Hagen): Water balance and energy content

The changes in dry mass, wet mass and energy content were determined during the establishment of the incipient colony. The winged reproductives of H. mossambicus are weak fliers. It is postulated that the alates compensate for the large quantity of reserve material by reducing their body water. The wet mass of both males and females increased significantly (twice original mass) from emergence until the first larvae appeared which tends support to this view. Furthermore, although the termites maintained in groups imbibed water, it is also clear that the reproductives, as long as they remain social, maintain the same low body-water content. This desiccated physiological condition appears to be and adaptation which assists flight and dispersal. In contrast with sterile eggs, fertile eggs absorbed water during development. The paired females (normal and homosexual) maintained the same energy/unit wet mass from egg production until the first workers appeared. Thus the possibility exists that the hydrophobic reserves (e.g. triglycerides) were converted to the hydrophilic reserves (e.g. carbohydrates). The same applied to the eggs and offspring.The changes in dry mass, wet mass, body water and energy content of females per unit of eggs (specific mass) produced were 0.005, 8.696, 21,276 and 12.820 respectively. The small specific dry mass value can probably be ascribed to the use of small amounts of glycerides and the increase in dry mass by the conversion from relatively light and “compact” glycerides to relatively heavy and “bulky” glycogen.Females utilised far more dry mass and energy in comparison to males to survive until the first workers appeared. Wet mass, body water and energy used in rearing of the offspring shows clearly that males made a larger contribution to the water and energy requirements of the offspring.     

Colony founding in the primitively eusocial wasp, Ropalidia marginata (Hymenoptera: Vespidae)

Abstract.

• 1 In a 16-month study in Bangalore, India, about 35% of the newly founded colonies of Ropalidia marginata were single foundress colonies and the remainder were multiple foundress colonies with two to twenty-two individuals.
• 2 Larger colonies did not have a significantly higher per capita productivity, did not produce significantly heavier progeny and did not produce them significantly faster than smaller colonies did.
• 3 Predation by the hornet Vespa tropica appeared to be independent of group size.
• 4 Single foundress colonies failed more often but not often enough to make them have a lower average per capita productivity, compared to multiple foundress colonies.
• 5 Some of the advantages of multiple foundress associations came from the greater predictability of their attaining the mean per capita productivity, the relatively lower rates of usurpation experienced by them compared to single foundress colonies, and the opportunities provided by queen turnovers for workers to become replacement queens and gain direct individual fitness.

     

Mating Behavior and Colony Founding of the Slave-Making Ant Formica sanguinea (Hymenoptera: Formicidae)

The mating and postmating behavior of reproductives belonging to two sympatric dulotic colonies of the facultative slave-making ant Formica sanguinea was analyzed in the field. Our observations showed that the European blood-red ant adopts a reproductive behavior similar to the male aggregation syndrome. Newly mated females return to a dulotic colony and often wait for a raid. Following a slave raid is an advantageous strategy to locate and invade host nests and to establish a new dulotic colony. In the laboratory, the following modes of colony founding were studied: independent, adoption, alliance, usurpation, and brood raiding. Independent foundation was possible only when several females were kept together. Alliance was obtained with females of two potential slave species (F. cunicularia, F. rufibarbis). Usurpation and adoption were more frequent in the incipient than in the mature host colonies. Mixed colonies were always obtained after the sack of the host pupae. It seems likely that, rather than conspecific adoption followed by budding, F. sanguinea relies on temporary parasitism to start new colonies.     

Toxicity and repellency of borate-sucrose water baits to Argentine ants (Hymenoptera: Formicidae)

The oral toxicity of boron compounds to the Argentine ant, Linepithema humile (Mayr), was evaluated in laboratory tests. The ants were provided 25% sucrose water containing 0.5 and 1% boric acid, disodium octaborate tetrahydrate, and borax. Lethal times of these solutions were a function of the concentration of boron. In field tests, the ants showed no discrimination between disodium octaborate tetrahydrate and boric acid. There was a significant reduction in consumption of sucrose water with > 1% boric acid.     

Structure and chemistry of the Dufour gland in Pristomyrmex ants (Hymenoptera, Formicidae)

Billen, J., Ito, F., Tsuji, K., Schoeters, E., Maile, R. and Morgan, E. D. 2000. Structure and chemistry of the Dufour gland in Pristomyrmex ants (Hymenoptera, Formicidae). —Acta Zoologica (Stockholm) 81 : 159–166 All individuals of the three species of Pristomyrmex studied have a Dufour gland with a conspicuous hammer‐shaped distal part, that is connected to a thin‐walled proximal reservoir through a very narrow stalk. The secretory distal part is formed by high columnar cells that are characterized by a highly folded apical wall. The lateral cell junctions apically correspond with the crenel tops, which gives individual cells a peculiar shape with a deep apical depression. The cytoplasm of the secretory cells contains a very well developed smooth endoplasmic reticulum and Golgi apparatus, numerous mitochondria and lysosomes. Histochemical analysis indicates a positive reaction for the presence of a lipid secretion in the hammer‐like part. Gas chromatographic analysis of glands of P. pungens workers reveals the secretion to be formed of a mixture of simple volatile monoterpene hydrocarbons; α–pinene, β–pinene, limonene and camphene. Similar, but species characteristic mixtures of four monoterpenes were found in Pristomyrmex brevispinosus and Pristomyrmex sp.1. Behavioural experiments did not allow a conclusive determination of the function of the gland.