Mature Colonies Research Articles

Biofilm dispersal of Neisseria subflava and other phylogenetically diverse oral bacteria.

Polystyrene petri dishes containing liquid medium were inoculated with single-cell suspensions of a fresh clinical isolate of Neisseria subflava and were incubated under conditions of low vibration. N. subflava colonies grew firmly attached to the surface of the dish, while the broth remained clear. Growing colonies released cells into the medium, resulting in the appearance of 10(2) to 10(4) small satellite colonies attached to the surface of the dish in an area adjacent to each mature colony after 24 h. Satellite colonies grew in patterns of streamers shaped like jets and flares emanating from mature colonies and pointing toward the center of the dish. This dispersal pattern evidently resulted from the surface translocation of detached biofilm cells by buoyancy-driven convection currents that were generated due to slight temperature gradients in the medium. Streamers of satellite colonies ranged from 2 to >40 mm in length. Satellite colonies in very long streamers were relatively uniform in size regardless of their distance from the mature colony, suggesting that mature colonies released single cells or small clusters of cells into the medium and that the detachment, surface translocation, and subsequent surface reattachment of released cells were a transitory process. Incubation of N. subflava single cells in a perfused biofilm fermentor resulted in a large spike of the number of CFU in the perfusate after 9.5 h of growth, consistent with a rapid release of cells into the medium. Biofilm colonies of several other phylogenetically diverse oral bacteria, including Actinobacillus actinomycetemcomitans, Haemophilus aphrophilus, Streptococcus mitis, and a prevalent but previously uncultured oral Streptococcus sp., exhibited similar temperature-dependent dispersal patterns in broth culture. This in vitro spreading phenotype could be a useful tool for studying biofilm dispersal in these and other nonflagellated bacteria and may have physiological relevance to biofilm dispersal in the oral cavity.

Meiotic differentiation during colony maturation in Saccharomyces cerevisiae.

As yeast colonies ceased growth, cells at the edge of these colonies transited from the cell division cycle into meiosis at high efficiency. This transition occurred remarkably synchronously and only at late stages of colony maturation. The transition occurred on medium containing acetate or low concentrations of glucose, but not on medium containing high glucose. The repression by high glucose was overcome when IME1 was overexpressed from a plasmid. Experiments with different growth media imply that meiosis in colonies is triggered by changes in the nutrient environment as colonies mature. HAP2 is required to sporulate in any carbon source, whereas GRR1 is required for glucose repression of sporulation. CLN3 is required to repress meiosis in colonies but not in liquid cultures, indicating that the regulators that mediate the transition to meiosis in colonies are not identical to the regulators that mediate this transition in liquid cultures.

Soldier Production Strategy in Lower Termites: From Young Instars or Old Instars?

We constructed a mathematical model to explain from which instars soldiers should be produced to maximize the growth rate of a termite colony. The model is based on the demography of the lower termite's colony: many of them feed inside the nest. The model predicts the following: (1) When the colony is young (it still has enough food and needs a high ratio of soldiers to workers) it will produce soldiers from young instars. (2) When the colony is old (it does not have enough food any more and needs a lower ratio of soldiers to workers) it will produce soldiers from old instars. This prediction fits well with the fact investigated by past empirical researches. With the samples of colonies of Neotermes koshunensis, we measured the antennal joint number of soldiers of each colony. We plotted the antennal joint number (1) of the average and (2) of the maximum one for each colony, against the total number of colony members. For both in each colony, we confirmed the consistency between the model and the samples: as the colony matures, it produces soldiers from older instars. The model also explains why higher termites produce soldiers from fixed instars. Their life style, in which they go outside to collect food, does not require a change in their soldier production strategy.

Identifying the transition between single and multiple mating of queens in fungus-growing ants.

Obligate mating of females (queens) with multiple males has evolved only rarely in social Hymenoptera (ants, social bees, social wasps) and for reasons that are fundamentally different from those underlying multiple mating in other animals. The monophyletic tribe of ('attine') fungus-growing ants is known to include evolutionarily derived genera with obligate multiple mating (the Acromyrmex and Atta leafcutter ants) as well as phylogenetically basal genera with exclusively single mating (e.g. Apterostigma, Cyphomyrmex, Myrmicocrypta). All attine genera share the unique characteristic of obligate dependence on symbiotic fungus gardens for food, but the sophistication of this symbiosis differs considerably across genera. The lower attine genera generally have small, short-lived colonies and relatively non-specialized fungal symbionts (capable of living independently of their ant hosts), whereas the four evolutionarily derived higher attine genera have highly specialized, long-term clonal symbionts. In this paper, we investigate whether the transition from single to multiple mating occurred relatively recently in the evolution of the attine ants, in conjunction with the novel herbivorous 'leafcutter' niche acquired by the common ancestor of Acromyrmex and Atta, or earlier, at the transition to rearing specialized long-term clonal fungi in the common ancestor of the larger group of higher attines that also includes the genera Trachymyrmex and Sericomyrmex. We use DNA microsatellite analysis to provide unambiguous evidence for a single, late and abrupt evolutionary transition from exclusively single to obligatory multiple mating. This transition is historically correlated with other evolutionary innovations, including the extensive use of fresh vegetation as substrate for the fungus garden, a massive increase in mature colony size and morphological differentiation of the worker caste.

COMPETITION–COLONIZATION TRADE-OFFS IN A GUILD OF AFRICAN ACACIA-ANTS

In systems where disturbance creates colonization opportunities, competition–colonization trade-offs can lead to coexistence between competitors sharing a single, limiting resource. In upland East Africa, four species of obligate acacia-ants compete for possession of Acacia drepanolobium host trees but still coexist at fine spatial scales. Previous experiments revealed a hierarchy of competitive dominance among mature ant colonies in battles over occupied trees (Crematogaster sjostedti > C. mimosae > C. nigriceps > Tetraponera penzigi), and the pattern of ant species turnover on single host trees over time is consistent with this hierarchy. Here we present evidence that competition–colonization trade-offs are likely to facilitate coexistence within this guild. The population of host trees is highly dynamic. Although most suitable host trees are occupied by ants, mature A. drepanolobium trees may be vacated after fire, extensive damage, or drought. Smaller host trees with limited nesting space become available when they recruit into the population as saplings or coppices. These unoccupied trees may be colonized by expansion of nearby, mature colonies as well as by foundress queens attempting to initiate new colonies. We identified competition–colonization trade-offs between two stages of colony development: competitive colony expansion onto nearby trees, and competition among foundress queens for nest-initiation sites. All three Crematogaster species readily occupy nearby, empty trees by colony expansion. In contrast, T. penzigi colonizes empty trees almost exclusively by dispersive foundress queens. Colonies of C. sjostedti are competitively dominant over those of all other species, but foundress queens of this species rarely attempt to start independent colonies. Foundress queens of the secondary competitive dominant (C. mimosae) are restricted mostly to low-quality, small saplings. In contrast, competitively subordinate colonies of T. penzigi and C. nigriceps produce foundress queens that disproportionately colonize empty mature trees, the highest quality host plants that are available. Competition between foundress queens is intense. Queens are significantly hyper-dispersed among potential nest-initiation sites, indicating that the presence of resident foundresses deters subsequent colonization attempts, and competitive dominance among foundresses who fight during colony founding is the reverse of that seen among mature colonies. Coexistence within this guild may therefore be promoted by ontogenetic reversals in relative competitive ability between early and late stages of colony development, a pattern that is consistent with the niche hypothesis for competition–colonization trade-offs.

The Natural History of the Arboreal Ant, Crematogaster ashmeadi

The arboreal ant, Crematogaster ashmeadi Emery (Hymenoptera: Formicidae), is the most dominant arboreal ant in the pine forests of the coastal plain of northern Florida. The majority of pine trees harbor a colony of these ants. The colonies inhabit multiple chambers abandoned by bark-mining caterpillars, especially those of the family Cossidae, in the outer bark of living pines. They also inhabit ground level termite galleries in the bark, often locating the queen in galleries. The density of chambers and ants is highest in the base of the tree and drops sharply with height on the trunk. Because chambers are formed in the inner layer of bark, they gradually move outward as more bark layers are laid down, eventually sloughing off the tree's outer surface. Chambers have a mean lifetime of about 25 yr. The abundant chambers in pine bark are excavated by a small population of caterpillars and accumulate over decades. Ant colonies also inhabit abandoned galleries of woodboring beetles in dead branches in the crowns of pines. Because newly mated queens found colonies in abandoned woodboring beetle galleries in the first dead branches that form on pine saplings, C. ashmeadi is dependent on cavities made by other insects throughout its life cycle, and does little if any excavation of its own. Mature colonies nest preferentially in chambers greater than 10 cm2 in area, a relatively rare chamber size. In natural pine forests, this does not seem to limit the ant's populations.Founding queens containabout 50% fat and lose about half of their dry weight during the claustral period, converting approximately half of this lost weight into progeny. The claustral period is about 40 to 50 days at 27°C. Mature colonies contain several tens of thousands of workers (est. up to 80,000), and have a life expectancy of 10 to 15 years. Each colony occupies an entire tree, and sometimes two trees if they are close together. Within a colony, there is a single queen capable of laying up to 450 eggs/day during the warm season. Such queens weigh 12 to 18 mg, have 50 to 60 active ovarioles and 120 to 600 vitellogenic oocytes in their ovaries.Mature colonies begin producing sexual brood in late April or early May. Sexual adults are present from late May through June. Mating flights commence in June and most sexuals have left their natal nests by late July. Female sexuals are an especially large investment; the energetic content of a single, flight-ready female sexual is almost 20 times that of a worker. The newly mated queen sequesters a mean of 2.64 × 106 sperm in her spermatheca, a supply that should last her for 16 years at the observed reproductive rates.

Allometry in the giant tropical ant, Paraponera clavata

Workers of the ponerine ant, Paraponera clavata, grow allometrically, with the head disproportionately larger in larger workers. The anterior (clypeus)-posterior (occiput) dimension of the head capsule is allometric with the lateral (interocular) dimension, so that larger workers have elongate heads. The degree of head capsule allometry varies among colonies; this could result either from differences in colony maturity or from variation in developmental patterns among colonies. These data resolve a previously controversial issue concerning caste in Paraponera clavata.

Origin of male-biased sex allocation in orphaned colonies of the termite, Coptotermes lacteus

In eusocial insects, sex allocation often constitutes a ground for intracolonial conflicts. This occurrence provides ideal opportunities to test kin-selection theory. A vast literature on this topic is available for social Hymenoptera, but the same field remains almost untouched in termites. A preeminent case is that of some species of Coptotermes, where the sex-allocation ratio in nymphs shifts from near equity to all-male when the primary reproductives are replaced by neotenics. To shed light on the developmental origin of this shift, we compared the sex ratio of the various castes and instars in primary- and neotenic-headed mature colonies of Coptotermes lacteus. The male-biased sex allocation in the latter type of colony results from two concurrent events: first, the sex ratio of the youngest instars (larvae) is male-biased by a 3:1 ratio; and second, all female larvae become workers, while a large fraction of the male larvae proceed to the nymphal and alate stages. Colony-founding experiments showed that inbreeding by itself cannot account for the male bias at hatching. We suggest that both genetic factors, due to the reproductive behaviour of neotenics, and environmental factors (colony condition and resource availability) may influence this process. Their exact nature and respective impact have not yet been clarified.

The natural history of the arboreal ant, Crematogaster ashmeadi.

The arboreal ant, Crematogaster ashmeadi Emery (Hymenoptera: Formicidae), is the most dominant arboreal ant in the pine forests of the coastal plain of northern Florida. The majority of pine trees harbor a colony of these ants. The colonies inhabit multiple chambers abandoned by bark-mining caterpillars, especially those of the family Cossidae, in the outer bark of living pines. They also inhabit ground level termite galleries in the bark, often locating the queen in galleries. The density of chambers and ants is highest in the base of the tree and drops sharply with height on the trunk. Because chambers are formed in the inner layer of bark, they gradually move outward as more bark layers are laid down, eventually sloughing off the tree's outer surface. Chambers have a mean lifetime of about 25 yr. The abundant chambers in pine bark are excavated by a small population of caterpillars and accumulate over decades. Ant colonies also inhabit abandoned galleries of woodboring beetles in dead branches in the crowns of pines. Because newly mated queens found colonies in abandoned woodboring beetle galleries in the first dead branches that form on pine saplings, C. ashmeadi is dependent on cavities made by other insects throughout its life cycle, and does little if any excavation of its own. Mature colonies nest preferentially in chambers greater than 10 cm(2) in area, a relatively rare chamber size. In natural pine forests, this does not seem to limit the ant's populations. Founding queens contain about 50% fat and lose about half of their dry weight during the claustral period, converting approximately half of this lost weight into progeny. The claustral period is about 40 to 50 days at 27 degrees C. Mature colonies contain several tens of thousands of workers (est. up to 80,000), and have a life expectancy of 10 to 15 years. Each colony occupies an entire tree, and sometimes two trees if they are close together. Within a colony, there is a single queen capable of laying up to 450 eggs/day during the warm season. Such queens weigh 12 to 18 mg, have 50 to 60 active ovarioles and 120 to 600 vitellogenic oocytes in their ovaries. Mature colonies begin producing sexual brood in late April or early May. Sexual adults are present from late May through June. Mating flights commence in June and most sexuals have left their natal nests by late July. Female sexuals are an especially large investment; the energetic content of a single, flight-ready female sexual is almost 20 times that of a worker. The newly mated queen sequesters a mean of 2.64 x 10(6) sperm in her spermatheca, a supply that should last her for 16 years at the observed reproductive rates.

Behavioral Repertoire, Age Polyethism and Adult Transport in Ectatomma opaciventre (Formicidae: Ponerinae)

This study describes the behavioral repertoire of a mature colony of Ectatomma opaciventre Roger. Twenty-eight different behavioral acts were recorded from 4538 individual observations. There was evidence of age polyethism, with young workers performing significantly more brood care and allogrooming than older workers. Experiments were also performed to investigate the behavioral organization of nest relocation in E. opaciventre. This complex social behavior was regulated by a few workers which specialized in transporting nestmates, including other workers, winged gynes, males, and the queen. Winged gynes were also observed performing adult transport. The carrying posture utilized in E. opaciventre is similar to that in other ponerines, although some idiosyncratic features were detected.

Characterization of a New Subgroup of Rhizoctonia solani Anastomosis Group 1 (AG-1-ID), Causal Agent of a Necrotic Leaf Spot on Coffee

ABSTRACT A new foliar disease on coffee leaves was observed in Mindanao, Philippines, in 1996. The symptoms appeared as large circular or irregularly shaped necrotic areas with small circular necrotic spots (1 mm or less in diameter) usually found around the periphery of the large necrotic areas. Rhizoctonia solani was consistently isolated from these diseased coffee leaves. Isolates obtained were multinucleate (3 to 12 nuclei per hyphal cell), had an optimum temperature for hyphal growth at 25 degrees C, prototrophic for thiamine, and anastomosed with tester isolates belonging to R. solani anastomosis group 1 (AG-1). Mature cultures on potato dextrose agar (PDA) were light to dark brown. Sclerotia, light brown to brown, were formed on the surface of PDA and covered the whole mature colony culture. Individual sclerotia often aggregated into large clumps (3 to 8 mm in diameter) and their color was brown to dark brown. In pathogenicity tests, isolates from coffee caused necrotic symptoms on coffee leaves, whereas isolates of AG-1-IA (not isolated from coffee), 1-IB, and 1-IC did not. The results of analyses of restriction fragment length polymorphism of ribosomal DNA internal transcribed spacer, random amplified polymorphism DNA, and fatty acid profiles showed that R. solani isolates from coffee are a population of AG-1 different from AG-1-IA, 1-IB, and 1-IC. These results suggest that R. solani isolates from coffee represent a new subgroup distinct from AG-1-IA, 1-IB, and 1-IC. A new subgroup ID (AG-1-ID) is proposed.

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal.

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999. Using a blocked, paired plot design we found that removal of colonies increased new colony recruitment in some areas of the site, but not others. Spatial variation in the importance of established colonies to recruitment was consistent across years; blocks in which density dependence was important in one year exhibited density dependent recruitment in following years. We estimated that in the blocks where recruitment was affected by established colonies, they accounted for less than 10% of the mortality of foundress queens. The increase in the number of new recruits (on average two additional new colonies) was considerably less than the number of colonies removed; average colony density in the removal plots was 14 colonies per 0.25ha plot. The consistent lack of importance of established colonies to recruitment in one block and the relatively small response to colony removal in the other blocks suggests that the number of new colonies in a year may not be equivalent to the number of deaths of established colonies in that year. Space limitation is an important influence on recruitment in P. occidentalis, but the magnitude of the limitation varies spatially.

Evaluation of green lacewings, Chrysoperla plorabunda (Fitch) (Neurop., Chrysopidae), for augmentative release against Toxoptera citricida (Hom., Aphididae) in citrus

Larvae of the lacewing Chrysoperla plorabunda (Fitch) were evaluated in laboratory and field tests for potential to control the brown citrus aphid, Toxoptera citricida (Kirkaldy) in inundative releases. Larvae surviving to pupation consumed an average of 1676, 1297, 392, 165 and 130 1st–4th instar T. citricida nymphs and apterous adults, respectively, and the mean developmental time was 27.0, 23.0, 16.5, 13.8 and 18.3 days, respectively. The average pupation rate was 37.5%, but only 6.3% of pupae yielded adults. Developmental time and survival to pupation varied with the life stage of aphids fed to larvae, 4th instars yielding the highest survival and shortest developmental time. Development of C. plorabunda was compared on diets of T. citricida and Aphis spiraecola. Only 6.3% of larvae completed development on T. citricida, whereas 37.5% yielded adults on A. spiraecola. Larvae consumed more of the brown T. citricida than the green A. spiraecola in a choice test on a white background, but no preference was evident when aphids were offered on a black background. Two separate field trials were performed in citrus groves with T. citricida infestations in which an average of 275 and 116 C. plorabunda larvae/tree were released, respectively. The rate of T. citricida colony maturation (= survival to alate production) was very low in the first trial, and relatively high in the second trial, but did not differ between control and release trees in either trial. The most apparent cause of aphid colony mortality was predation by the coccinellids Cycloneda sanguinea L. and Harmonia axyridis Pallas. Despite acceptance of T. citricida in the laboratory, very few C. plorabunda larvae were observed feeding on aphid colonies on release trees.

Reproductive skew and queen relatedness in an ant with primary polygyny

Queens of the neotropical ponerine ant Pachycondyla cf. 'inversa' may cooperate during colony founding. Pleometrosis leads to permanent primary polygyny, with queens still co-existing in the same nest when the colony has become mature. Using multilocus DNA-fingerprinting we examined the genetic structure of colonies. Nestmate queens are typically not close relatives. Excepting one pair of queens, which appeared to be closely related, the band-sharing coefficients of co-occurring queens were similar to the background similarity among individuals from different nests. In mature colonies, queens typically contribute equally to workers and sexuals. Only in one of seven colonies did one queen produce significantly more workers than the other. Polygynous ants with unrelated queens might provide an interesting system to test predictions from kin selection theory.

Polyandry and colony genetic structure in the primitive ant Nothomyrmecia macrops

Abstract The Australian endemic ant Nothomyrmecia macrops is considered one of the most ‘primitive’ among living ants. We investigated the genetic structure of colonies to determine queen mating frequencies and nestmate relatedness. An average of 18.8 individuals from each of 32 colonies, and sperm extracted from 34 foraging queens, were genotyped using five highly variable microsatellite markers. Queens were typically singly (65%) or doubly mated (30%), but triple mating (5%) also occurred. The mean effective number of male mates for queens was 1.37. No relationship between colony size and queen mate number was found. Nestmate workers were related by b=0.61 ± 0.03, significantly above the threshold under Hamilton’s rule over which, all else being equal, altruistic behaviour persists, but queens and their mates were unrelated. In 25% of the colonies we detected a few workers that could not have been produced by the resident queen, although there was no evidence for worker reproduction. Polyandry is for the first time recorded in a species with very small mature colonies, which is inconsistent with the sperm-limitation hypothesis for the mediation of polyandry levels. Facultative polyandry is therefore not confined to the highly advanced ant genera, but may have arisen at an early stage in ant social evolution.

Self-sustaining mars colonies utilizing the north polar cap and the martian atmosphere

A revolutionary new concept for the early establishment of robust, self-sustaining Martian colonies is described. The colonies would be located on the North Polar Cap of Mars and utilize readily available water ice and the CO 2 Martian atmosphere as raw materials to produce all of the propellants, fuel, air, water, plastics, food, and other supplies needed by the colony. The colonists would live in thermally insulated large, comfortable habitats under the ice surface, fully shielded from cosmic rays. The habitats and supplies would be produced by a compact, lightweight (∼4 metric tons) nuclear powered robotic unit termed ALPH (Atomic Liberation of Propellant and Habitat), which would land 2 years before the colonists arrived. Using a compact, lightweight 5 MW (th) nuclear reactor/steam turbine (1 MW(e)) power source and small process units (e.g., H 2O electrolyzer, H 2 and O 2 liquefiers, methanator, plastic polymerizer, food producer, etc.) ALPH would stockpile many hundreds of tons of supplies in melt cavities under the ice, plus insulated habitats, to be in place and ready for use when the colonists landed. With the stockpiled supplies, the colonists would construct and operate rovers and flyers to explore the surface of Mars, ALPH greatly reduces the amount of Earth supplied material needed and enables large permanent colonies on Mars. It also greatly reduces human and mission risks and vastly increases the capability not only for exploration of the surrounding Martian surface, but also the ice cap itself. The North Polar Cap is at the center of the vast ancient ocean that covered much of the Martian Northern Hemisphere. Small, nuclear heated robotic probes would travel deep (1 km or more) inside the ice cap, collecting data on its internal structure, the composition and properties of the ancient Martian atmosphere, and possible evidence of ancient life forms (microfossils, traces of DNA, etc.) that were deposited either by wind or as remnants of the ancient ocean. Details of the ALPH system, which is based on existing technology, are presented. ALPH units could be developed and demonstrated on Earth ice sheets within a few years. An Earth-Mars space transport architecture is described, in which Mars produced propellant and supplies for return journeys to Earth would be lifted with relatively low ΔV to Mars orbit, and from there transported back to Earth orbit, enabling faster and lower cost trips from Earth to Mars. The exploration capability and quality of life in a mature Martian colony of 500 persons located on the North Polar Cap is outlined.

Why are leaf‐cutting ants more common in early secondary forests than in old‐growth tropical forests? An evaluation of the palatable forage hypothesis

I evaluated the hypothesis that leaf‐cutting ants are more common in early successional forests than in old‐growth forests because pioneer species, which dominate in early successional habitats, appear more susceptible to leafcutters than shade‐tolerant species, which dominate primary forests (palatable forage hypothesis). The relative importance of pioneer and shade‐tolerant species as plant resources for leaf‐cutting ant colonies was evaluated (1) by literature review of leaf‐cutting ants’ diet, and (2) experimentally, using field assays to determine leafcutter's selectivity. Pioneer species were harvested three times more frequently than shade‐tolerant species and made up the largest component of the diet in all the studies reviewed. The amount harvested was not correlated with the plant species abundance. In addition, leaves from pioneer plants were selected eight times more than leaves from shade‐tolerant species in the field assays. These results support the palatable forage hypothesis. Leafcutters probably select pioneer leaves because of their low level of chemical defenses and high nutrient content. The high availability of pioneer species in early successional forest probably decreases the cost to locate palatable resources. Therefore, early successional habitats support more ant colonies than old‐growth forests. On the other hand, the effective defense mechanisms of mature plant species and the high dispersion of palatable plants could explain the low density of leaf‐cutting ant colonies in old‐growth forests. The palatable forage hypothesis is compared with other hypotheses that explain leaf‐cutting ant density. The preference of foundress queens for forest clearings, the dependence of small colonies on herbs, and the importance of pioneer plant species for mature colonies (palatable forage hypothesis) can be considered complementary, because they focus on different stages of the colony's life history. Consequently, the availability of pioneer plants appears to be one of the most influential factors determining mature leaf‐cutting ant nest densities in Neotropical forests.

Queen mating frequency and relatedness in young Atta sexdens colonies

Queen mating frequency is correlated with mature colony size across monogynous ant species as a whole. However, recent comparisons between Atta and Acromyrmex leafcutter ants found an opposite trend. We tested whether this result could be reproduced for Atta sexdens, using a new Atta colombica microsatellite marker to analyse queens and worker offspring from six young colonies. We found that all queens had mated multiply and used sperm from at least 2-3 males. This result is comparable to previous findings for Atta colombica and seems to confirm that queen mating frequency in Atta is lower than in Acromyrmex.

Mathematical analysis of growth and interaction dynamics of streptomycetes and a bacteriophage in soil.

We observed the infection cycle of the temperate actinophage KC301 in relation to the growth of its host Streptomyces lividans TK24 in sterile soil microcosms. Despite a large increase in phage population following germination of host spores, there was no observable impact on host population numbers as measured by direct plate counts. The only change in the host population following infection was the establishment of a small subpopulation of KC301 lysogens. The interaction of S. lividans and KC301 in soil was analyzed with a population-dynamic mathematical model to determine the underlying mechanisms of this low susceptibility to phage attack relative to aquatic environments. This analysis suggests that the soil environment is a highly significant component of the phage-host interaction, an idea consistent with earlier observations on the importance of the environment in determining host growth and phage-host dynamics. Our results demonstrate that the accepted phage-host interaction and host life cycle, as determined from agar plate studies and liquid culture, is sufficient for quantitative agreement with observations in soil, using soil-determined rates. There are four significant effects of the soil environment: (i) newly germinated spores are more susceptible to phage lysis than are hyphae of developed mycelia, (ii) substrate mycelia in mature colonies adsorb about 98% of the total phage protecting susceptible young hyphae from infection, (iii) the burst size of KC301 is large in soil (>150, 90% confidence) relative to that observed in liquid culture (120, standard error of the mean [SEM], 6), and (iv) there is no measurable impact on the host in terms of reduced growth by the phage. We hypothesize that spatial heterogeneity is the principal cause of these effects and is the primary determinant in bacterial escape of phage lysis in soil.

Maintaining an ant-plant symbiosis: secondary polygyny in the Macaranga triloba-Crematogaster sp. association.

Ant-plant associations are common and important elements of tropical forest ecosystems. Their relationships vary from loosely facultative to obligate symbiosis (for an overview see Davidson and McKey 1993). In SE Asia many species of the pioneer tree genus Macaranga are myrmecophytes that provide their ant-partners with nutrients and nesting space in exchange for herbivoreprotection by the ants. The majority of the associated ant species belong to Decacrema, a subgenus of the myrmicine Crematogaster (Fiala et al. 1999) and they are obligatorily and specifically associated with their Macaranga plant-partners. Ants and plants are highly dependent on each other in this myrmecophytic complex. In most Macaranga species even very young saplings develop stem domatia, which are colonised by queens throughout the year. It is very rare to find more than one founding queen in one domatium but frequently several queens occupy different domatia of the same young plant. After claustral colony foundation and the emergence of the first workers, however, only one monogynous colony survives per host tree (Fiala and Maschwitz 1990). Generally it is supposed that the monogynous status is maintained throughout the entire existence of the symbiosis. The death of either partner – ant colony or host tree – can result in severe damage or even the death of the other partner. In the case of the death of the queen the colony dies out and the tree is highly endangered by herbivores (Fiala et al. 1989; Heil et al. 1997). Small trees may be colonised again by their specific ant-partners and then suffer only little damage during the development of the new colony which will soon be large enough to patrol the whole tree effectively. In mature trees, however, with their much larger leaf area, the establishment of a worker force would take too long. Do we find other mechanisms in ant colonies on mature trees that might enable the ants to further protect the tree against herbivoredamage as well as safeguard its territory and resources against competing ant species? Whereas much information now exists on the founding and growth of colonies on saplings (Fiala and Maschwitz 1990), no survey has ever been conducted on the development of mature ant colonies on larger Macaranga trees. We therefore studied queen numbers and colony development in maturing trees in the common myrmecophyte Macaranga triloba (Bl.) Muell. Arg., that is obligatorily associated with Crematogaster msp. 4 (Fiala et al. 1999).