Colony Life History Research Articles

Vasa expression in a colonial ascidian, Botrylloides violaceus

Evolution of solitary or colonial life histories in tunicates is accompanied by dramatic developmental changes that affect morphology and reproduction. We compared vasa expression in a solitary ascidian and a closely related colonial ascidian, in an effort to uncover developmental mechanisms important during the evolution of these contrasting life histories, including the ability to reproduce by budding. In this study, we explored the origin of germ cells in new buds developing by asexual reproduction in a colonial ascidian, Botrylloides violaceus and compared it to the source of germ cells in a solitary ascidian Boltenia villosa. We studied expression by in situ hybridization of vasa, a DEAD box RNA helicase gene found in germ cells across the metazoans. In B. villosa, bv-vasa mRNA was expressed in putative germ cells and oocytes of adult gonads, and was sequestered into a posterior lineage during embryogenesis. In mature colonies of the ascidian B. violaceus, bot-vasa mRNA was expressed in putative spermatogonia, in oocytes of zooids, and in some circulating cells in the zooids and differentiating buds. We propose that expression of vasa in cells other than gonadal germ cells of zooids in a colonial ascidian may serve as a source of germ-line stem cells in the colony.

Telomerase maintained in self-renewing tissues during serial regeneration of the urochordate Botryllus schlosseri

Telomerase is critical for the protection of germ line and stem cell chromosomes from fatal shortening during replication. In most organisms, telomerase activity is suppressed in progressively committed cells and falls to basal rates in terminally differentiated lineages. The colonial ascidian Botryllus schlosseri propagates asexually and sexually, presumably from pools of stem cells that self-renew throughout the 2- to 5-year colony life span. Asexual budding takes place continuously from the parental body wall. When the colony reaches a critical size, sexual reproduction commences with the generation of gonads. Here, we establish the existence of 6–15 kb telomeres on the ends of Botryllus chromosomes. We develop a real-time quantitative PCR telomeric repeat amplification protocol (TRAP) assay that reliably detects 0.2–100 TPG units in cells and tissues. We find highest levels of enzymatic activity in the gonads, developing embryos, and tissues containing the earliest asexual buds. Telomerase activity appears to be suppressed in later buds during organogenesis and falls to basal rates in mature zooids. We postulate that this pattern reflects maximum telomere restoration in somatic stem cells of early buds and suppression of telomerase activity in progenitors and terminally differentiated cells, indicative of an alternate role for stem cells as repeated body regenerators in colonial life histories.

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.

Sexual Reproduction in the Colonial Kamptozoan Barentsia hildegardae

Four aspects of sexual reproduction were examined in a northeastern Pacific population of the kamptozoan (entoproct) Barentsia hildegardae. The investigation of this colonial species revealed that (1) the population was sexually mature year-round, with a peak of sexual maturity during summer; (2) the sex ratio of the calyces in this population did not differ significantly from 1:1 male:female; (3) male colonies grown on glass slides had higher proportions of sexually mature calyces than did females and the zone of maturity extended farther toward stolon tips; (4) recruitment was much higher near female than male colonies, perhaps because brooded larvae settled near their mothers almost immediately after release. Additional key words: Entoprocta, sexual maturation, reproductive period, sex ratio, larval period Understanding the life history of a colonial animal can be complicated, because processes such as growth, reproduction, and death must be examined at multiple levels (zooid, colony, and whole genet). Moreover, determining colony (and genet) boundaries in the field is often impossible. Consequently, we know far less about the life histories of colonial animals than about their unitary counterparts. Learning even basic information about colonial life histories, however, is essential for evolutionary and ecological studies of colonial

Colony life history and demography of a swarm-founding social wasp

Colonies of social insects are sometimes viewed as superorganisms. The birth, reproduction, and death of colonies can be studied with demographic measures analogous to those normally applied to individuals, but two additional questions arise. First, how do adaptive colony demographies arise from individual behaviors? Second, since these superorganisms are made up of genetically distinct individuals, do conflicts within the colony sometimes modify and upset optima for colonies? The interplay between individual and superindividual or colony interests appears to be particularly complex in neotropical, swarm-founding, epiponine wasps such as Parachartergus colobopterus. In a long-term study of this species, we censused 286 nests to study colony-level reproduction and survivorship and evaluated individual-level factors by assessing genetic relatedness and queen production. Colony survivorship followed a negative exponential curve very closely, indicating type II survivorship. This pattern is defined by constant mortality across ages and is more characteristic of birds and other vertebrates than of insects. Individual colonies are long-lived, lasting an average of 347 days, with a maximum of over 4.5 years. The low and constant levels of colony mortality arise in part from colony initiation by swarming, nesting on protected substrates, and an unusual expandable nest structure. The ability to requeen rapidly was also important; relatedness data suggest that colonies requeen on average once every 9–12 months. We studied whether colony optima with respect to the timing of reproduction could be upset by individual worker interests. In this species, colonies are normally polygynous but new queens are produced only after a colony reaches the monogynous state, a result which is in accord with the genetic interests of workers. Therefore colony worker interests might drive colonies to reproduce whenever queen number happens to cycled down to one rather than at the season that is otherwise optimal. However, we found reproduction to be heavily concentrated in the rainy season. The number of new colonies peaked in this season as did the percentages of males and queens. Relatedness among workers reached a seasonal low of 0.21–0.27, reflecting the higher numbers of laying queens. This seasonality was achieved in part by a modest degree of synchrony in the queen reduction cycle. Worker relatedness reached peaks of around 0.4 in the dry season, reflecting a decrease to a harmonic mean queen number of about 2.5. Thus, a significant number of colonies must be approaching monogyny entering the rainy season. Coupled with polygynous colonies rearing only males (split sex ratios), this makes it possible for a colony cycle driven by selfish worker interests to be consistent with concentrating colony reproduction during a favorable season.

Population biology of bryozoans: correlates of sessile, colonial life histories in freshwater habitats

Bryozoans are common, often abundant, sessile, colonial invertebrates of freshwaters. By reviewing what is known of their general ecology and considering in further detail the population biology of specific groups, we provide evidence that 1) populations are locally ephemeral and regional persistence is attained via a dynamic equilibrium between dispersal and colonization, and 2) sex may often be infrequent and a high degree of clonality obtains at both local and regional scales. On the basis of these characteristics we discuss how the group offers exceptional opportunities for investigating a number of fundamentally important ecological and evolutionary questions.

Queen number and sociality in insects

List of contributors 1. Opportunities and pitfalls in co-operative reproduction 2. Reproductive structure and reproductive roles in colonies of eusocial insects 3. Queen - queen conflicts in polygynous societies: mutual tolerance and reproductive skew 4. Sex ratio variation in polygynous ants 5. The effects of polygny and colony life history on optimal sex investment 6. Genetic relatedness and its components in polygynous colonies of social insects 7. The maintenance of high genetic relatedness in multi-queen colonies of social wasps 8. The evolutiuon of polygyny in primitively eusocial polistine wasps with special rerference to the genus Ropalidia 9. Serial polygyny in the primitively eusocial wasp Ropalidia marginata: implications for the evolution of sociality 10. Multiple - foundress associations in sweat bees 11. Monogyny and polygyny in ponerine ants with or without queens 12. Ecological determinants of queen number in ants 13. Distribution and ecology of queen number in ants of the genus Myrmica 14. Monogyny and polygyny in Formica ants: the result of alternative dispersal tactics 15. Queen - queen interactions in polygynous ants 16. Kin discrimination and division of labour among matrilines in the polygynous carpenter ant, Camponotus planatus 17. Selective pressures on pleometrosis and secondary polygyny: a comparison of termites and ants Glossary Author index Subject index

The Adaptiveness of Worker Demography in the Attine Ant Trachymyrmex Septentrionalis

Castle theory states that worker size distributions in ant colonies have evolved for efficient division of labor and predicts that they should vary with habitat and have an effect on fitness. We tested these predictions in a comparative study of worker size variation and colony fitness in Florida and Long Island populations of Trachymyrmex septentrionalis. Colonies from the two populations were excavated shortly before the time of the mating flights, so that workers and reproductive forms could be censused. We measured worker head widths in each colony and analyzed the resulting size frequency distributions using the mean, standard deviation, skewness, and kurtoisis, and also principal components derived from these parameters. Size frequency distributions were unimodal and close to normal, with a tendency to negative skewness. Colonies in the Long Island population had both larger workers and greater size variation than those in Florida. Florida colonies followed a growth trajectory in which the mean and standard deviation of worker head width both increased with colony size, whereas in Long Island there was an inverse relationship between the mean and standard deviation that was independent of colony size. Analysis based on the principal components showed that worker size variation significantly affected fitness in both populations, and this was not a secondary effect of colony size. Population differences in colony life history strategies may account for the observed patterns: the Long Island population appears to be adapted for surviving the winter diapause, while the Florida population has experienced different selective pressures, possibly for rapid colony growth. Size variation that has evolved in the context of life history strategies may serve as a preadaptation for division of labor.

Randomly amplified polymorphic DNA analysis of clonal population structure and geographic variation in a freshwater bryozoan.

The randomly amplified polymorphic DNA (RAPD) assay was used to identify genetic polymorphisms in three clonal populations of the freshwater bryozoan, Cristatella mucedo, a species with few useful biochemical genetic markers. Of the 19 decamer oligonucleotide primers screened, 13 gave clear, reproducible RAPD profiles. Clonal population structure was evident, and one clone was dominant at each site. Cluster analysis grouped populations from more distant localities separately (Thames Valley and Norfolk), whereas populations from the Thames Valley clustered together. However, even at the regional scale a high degree of relatedness pertained. This work is one of the first RAPD studies of natural populations, and demonstrates the suitability of the technique for examining population structure and geographic variation in clonal taxa.

Nestmate recognition and the ontogeny of acceptability in the ant, Leptothorax curvispinosus

In social insects, there is often a brief period following eclosion when workers are highly acceptable in alien nests of their own or other species. This study tested for such an “acceptance period” in the facultatively polygynous ant, Leptothorax curvispinosus, and compared the duration and effectiveness of this period for conspecific and heterospecific introductions. Workers that eclosed and aged for 1–70 h or 30 days in isolation were introduced into either their parental nests (n=24), alien conspecific nests (n=265), or nests of the closely related and biologically similar species, L. longispinosus (n=341). In alien conspecific nests, acceptance was maximal for workers aged 1–12 h at introduction (67.7% not attacked, 75.8% adopted) and gradually decreased until the level of nonaggression (after 60 h) and adoption (after 36 h) were not significantly different from 30-day-old workers (5.9% not attacked, 17.6% adopted). In heterospecific nests, acceptance was maximal for workers aged 1–4 h at introduction (34.8% not attacked, 37.0% adopted) but thereafter was not significantly different from 30-day-old workers (5.6% not attacked, 8.3% adopted). In their parental nests, workers were generally accepted regardless of age (4–56 h posteclosion, 95.8% not attacked, 100% adopted); a result that is consistent with previous research on older workers (38–157 days posteclosion). This study demonstrates an acceptance period that is more effective and of longer duration within than between these species but that, under uniform laboratory conditions, is often not necessary for the integration of workers into their parental colonies. Within colonies, acceptance periods might only be important during relatively brief periods in a colony's life history when eclosing workers produce genetically based nestmate recongition cues that are not already represented in the colony and must be learned by colony members (e.g., during early colony growth or following adoption of queens), or when young workers must acquire environmentally based nestmate recognition cues to achieve and maintain acceptability.

Asexual reproduction by oceanic planktotrophic echinoderm larvae

Cloning among metazoan larvae is rare and generally restricted to a few phyla with parasitic or colonial life-histories. Echinoderms are all non-colonial animals, and although cloning is well docu-mented for some adults, its occurrence in larval stages has not been previously observed1–3. Here we describe a novel mode of cloning by fission in planktotrophic bipinnaria larvae of the sea star Luidia sp. from the Sargasso Sea and the Gulf Stream. Reproductive larvae were widely distributed, comprising about 30% of sample populations. This represents an extensive multi-plicative potential whereby the pelagic life of the genet is prolonged and the potential of recruitment into benthic adult populations can be enhanced.