Mature Colonies Research Articles

SIRT1 deficiency compromises mouse embryonic stem cell hematopoietic differentiation, and embryonic and adult hematopoiesis in the mouse

AbstractSIRT1 is a founding member of a sirtuin family of 7 proteins and histone deacetylases. It is involved in cellular resistance to stress, metabolism, differentiation, aging, and tumor suppression. SIRT1−/− mice demonstrate embryonic and postnatal development defects. We examined hematopoietic and endothelial cell differentiation of SIRT1−/− mouse embryonic stem cells (ESCs) in vitro, and hematopoietic progenitors in SIRT1+/++/−, and −/− mice. SIRT1−/− ESCs formed fewer mature blast cell colonies. Replated SIRT1−/− blast colony-forming cells demonstrated defective hematopoietic potential. Endothelial cell production was unaltered, but there were defects in formation of a primitive vascular network from SIRT1−/−-derived embryoid bodies. Development of primitive and definitive progenitors derived from SIRT1−/− ESCs were also delayed and/or defective. Differentiation delay/defects were associated with delayed capacity to switch off Oct4, Nanog and Fgf5 expression, decreased β-H1 globin, β-major globin, and Scl gene expression, and reduced activation of Erk1/2. Ectopic expression of SIRT1 rescued SIRT1−/− ESC differentiation deficiencies. SIRT1−/− yolk sacs manifested fewer primitive erythroid precursors. SIRT1−/− and SIRT1+/− adult marrow had decreased numbers and cycling of hematopoietic progenitors, effects more apparent at 5%, than at 20%, oxygen tension, and these progenitors survived less well in vitro under conditions of delayed growth factor addition. This suggests a role for SIRT1 in ESC differentiation and mouse hematopoiesis.

Strict monandry in the ponerine army ant genus Simopelta suggests that colony size and complexity drive mating system evolution in social insects

Altruism in social insects has evolved between closely related full-siblings. It is therefore of considerable interest why some groups have secondarily evolved low within-colony relatedness, which in turn affects the relatedness incentives of within-colony cooperation and conflict. The highest queen mating frequencies, and therefore among the lowest degrees of colony relatedness, occur in Apis honeybees and army ants of the subfamilies Aenictinae, Ecitoninae, and Dorylinae, suggesting that common life history features such as reproduction by colony fission and male biased numerical sex-ratios have convergently shaped these mating systems. Here we show that ponerine army ants of the genus Simopelta, which are distantly related but similar in general biology to other army ants, have strictly monandrous queens. Preliminary data suggest that workers reproduce in queenright colonies, which is in sharp contrast to other army ants. We hypothesize that differences in mature colony size and social complexity may explain these striking discrepancies.

Recruitment Strategies and Colony Size in Ants

Ants use a great variety of recruitment methods to forage for food or find new nests, including tandem running, group recruitment and scent trails. It has been known for some time that there is a loose correlation across many taxa between species-specific mature colony size and recruitment method. Very small colonies tend to use solitary foraging; small to medium sized colonies use tandem running or group recruitment whereas larger colonies use pheromone recruitment trails. Until now, explanations for this correlation have focused on the ants' ecology, such as food resource distribution. However, many species have colonies with a single queen and workforces that grow over several orders of magnitude, and little is known about how a colony's organization, including recruitment methods, may change during its growth. After all, recruitment involves interactions between ants, and hence the size of the colony itself may influence which recruitment method is used—even if the ants' behavioural repertoire remains unchanged. Here we show using mathematical models that the observed correlation can also be explained by recognizing that failure rates in recruitment depend differently on colony size in various recruitment strategies. Our models focus on the build up of recruiter numbers inside colonies and are not based on optimality arguments, such as maximizing food yield. We predict that ant colonies of a certain size should use only one recruitment method (and always the same one) rather than a mix of two or more. These results highlight the importance of the organization of recruitment and how it is affected by colony size. Hence these results should also expand our understanding of ant ecology.

Effects of Selfing on Offspring Survival and Reproduction in a Colonial Simultaneous Hermaphrodite (Bugula stolonifera, Bryozoa)

Understanding the consequences of selfing in simultaneous hermaphrodites requires investigating potential deleterious effects on fitness at all stages of life. In this study, I examined the effects of selfing throughout the life cycle of the marine bryozoan Bugula stolonifera, a colonial simultaneous hermaphrodite. In 2008, larvae from field-collected colonies were cultured through metamorphosis to reproductively mature colonies either in the presence of one other colony, the paired treatment, or alone, the solitary treatment. Results demonstrated that selfing in this species is possible, in that colonies in the solitary treatment produced viable larvae that successfully completed metamorphosis. On average, however, these colonies released significantly fewer larvae, which experienced reduced rates of metamorphic initiation and completion compared to the paired treatment. These experiments were extended in 2009, when metamorphs from colonies reared in the solitary (n = 58) and paired (n = 61) treatments were transferred to the field for growth to reproductive maturity and then brought back to the laboratory for larval collection. Results revealed additional deleterious effects associated with selfing, as no viable larvae were recovered from colonies deriving from the solitary treatment. In contrast, offspring from the paired treatment released 1030 larvae and 99% initiated metamorphosis, 97% of which completed metamorphosis. Overall, selfed larvae not only had significantly decreased chances of survival, but those that did survive did not successfully reproduce.

Patriline shifting leads to apparent genetic caste determination in harvester ants

The harvester ant, Pogonomyrmex occidentalis, is characterized by high levels of intracolonial genetic diversity resulting from multiple mating by the queen. Within reproductively mature colonies, the relative frequency of different male genotypes (patrilines) is not stable. The difference between samples increases with time, nearing an asymptote after a year. Patriline distributions in gynes and workers display similar patterns of change. A consequence of changing patriline distributions is that workers and gynes appear to have different fathers. However, apparent genetic differences between castes are caused by changing paternity among all females. Temporal variation in the relative frequency of patrilines may be a consequence of processes that reflect sexual conflict, such as sperm clumping. Recent work documenting genotype differences between physical castes (workers and gynes; major and minor workers) in several species of ants has been interpreted as evidence of genetic caste determination. Reanalysis of these studies found little support for this hypothesis. Apparent caste determination may result from temporal variation in sperm use, rather than from fertilization bias among male ejaculates.

Reproductive Synchrony inAcroporaAssemblages on Reefs of New Caledonia

Despite a recent expansion in the geographical focus of studies on coral reproduction, there remain many regions in the Indo-Pacific, such as Melanesia, where research is limited. For example, although New Caledonia in southern Melanesia is home to the world's second largest barrier reef, which has recently been given UNESCO World Heritage listing, almost nothing is known of the reproductive biology of the coral fauna there, in particular the timing of spawning. In this study we sampled Acropora assemblages in November 2004 to test for reproductive synchrony at eight sites in New Caledonia separated by up to 200 km. In total, 80% of 1,055 Acropora colonies sampled contained mature oocytes, and 34 (92%) of 37 species sampled had at least one mature colony. These data demonstrate that reproduction in Acropora is highly synchronous over a large scale in New Caledonia and suggest a multispecies spawning event following the full moon in November coincident with the mass spawning period on the Great Barrier Reef, Australia. The high synchrony of reproductive effort implies that even a brief halt of activities that threaten fertilization and early development of coral propagules, such as discharge of liquid waste from ore processing, could have a major mitigating effect on the potential damage to these globally valued reefs.

Worker Allometry in Relation to Colony Size and Social form in the Fire AntSolenopsis invicta

Workers of the polymorphic fire ant Solenopsis invicta Buren (Hymenoptera: Formicidae) show modest changes of shape with increases in body size. These shape changes (allometries) have been described only for workers taken from mature colonies of the monogyne social form. For the study reported here, workers were collected from small and large monogyne and large polygyne colonies for tests of the effects of colony size and social form on allometry. The differential growth of body parts in relation to total body growth was determined by measurement of all major body parts and regression of the logs of these measurements, or their ratios, on the log of the body size. The slopes of these regressions defined the allometric relationships, and the slopes for these three types of colonies were compared for determination of the influence of colony size and social form on allometric rules. Most allometric constants did not differ with colony size or social form, but head shape, relative antennal size, and alinotum shape did. For a given worker size, heads of workers from small monogyne colonies or from polygyne colonies were narrower above the eyes. Antennae of workers from large monogyne colonies were relatively shorter than those from small monogyne or polygyne colonies (which did not differ). Alinotum heights of small workers from small monogyne colonies were greater than those from large monogyne or polygyne colonies (which were isometric and did not differ). These observed differences in allometric constants suggest that the relative growth rules are not completely determined by worker body size but are affected by colony size and social form. These differences are discussed in light of the growth of imaginai discs under conditions of fixed resources.

FOG1 requires NuRD to promote hematopoiesis and maintain lineage fidelity within the megakaryocytic-erythroid compartment

AbstractNuclear factors regulate the development of complex tissues by promoting the formation of one cell lineage over another. The cofactor FOG1 interacts with transcription factors GATA1 and GATA2 to control erythroid and megakaryocyte (MK) differentiation. In contrast, FOG1 antagonizes the ability of GATA factors to promote mast cell (MC) development. Normal FOG1 function in late-stage erythroid cells and MK requires interaction with the chromatin remodeling complex NuRD. Here, we report that mice in which the FOG1/NuRD interaction is disrupted (Fogki/ki) produce MK-erythroid progenitors that give rise to significantly fewer and less mature MK and erythroid colonies in vitro while retaining multilineage capacity, capable of generating MCs and other myeloid lineage cells. Gene expression profiling of Fogki/ki MK-erythroid progenitors revealed inappropriate expression of several MC-specific genes. Strikingly, aberrant MC gene expression persisted in mature Fogki/ki MK and erythroid progeny. Using a GATA1-dependent committed erythroid cell line, select MC genes were found to be occupied by NuRD, suggesting a direct mechanism of repression. Together, these observations suggest that a simple heritable silencing mechanism is insufficient to permanently repress MC genes. Instead, the continuous presence of GATA1, FOG1, and NuRD is required to maintain lineage fidelity throughout MK-erythroid ontogeny.

Diploid male production in a leaf-cutting ant

1. In haplodiploid social insects where males are haploid and females are diploid, inbreeding depression is expressed as the production of diploid males when homozygosity at the sex-determining locus results in the production of diploid individuals with a male phenotype. Diploid males are often assumed to have reduced fitness compared with their haploid brothers. 2. While studying the reproductive biology of a leaf-cutting ant, Atta sexdens, in Gamboa, Republic of Panama, we detected the presence of a larger male morph. Using microsatellite markers we were able to confirm that the large male morph was diploid in 87% of cases. 3. We infer that the Gamboa population of A. sexdens experiences inbreeding depression because diploid males were found in three out of five mature colonies. However, their frequencies were relatively low because queens were multiply mated and our estimates suggest that many diploid male larvae may not survive to adulthood. 4. We measured two traits potentially linked to male reproductive success: sperm length and sperm number, and showed that diploid males produced fewer but longer sperm. These results provide indirect evidence that diploid male reproductive success would be reduced compared with haploid males if they were able to copulate. 5. We conclude that diploid male production is likely to affect the fitness of A. sexdens queens with a matched mating, as these males are produced at the cost of workers and, if the colony survives to reach mature size, also gynes.

Colony fusion causes within-colony variation in a parthenogenetic ant

The evolutionary stability of cooperation and altruism in colonies of social insects requires that nestmates be to some extent related. An efficient system of discrimination against non-nestmates protects the nest against unrelated conspecifics, which might exploit or parasitize the colony. The co-occurrence of unrelated individuals in mature colonies therefore is a rare event that deserves more attention. Here, we report on the relatively common incidence of colony fusion in the ant Platythyrea punctata. Workers of this ant can produce genetically identical female offspring from unfertilized eggs through thelytokous parthenogenesis. Consequently, the majority of colonies has a “clonal structure” and consists of individuals with identical multilocus genotypes. Nevertheless, field observations indicate that a surprisingly large percentage of colonies contain workers belonging to two or more different genetic lineages. Much of this genetic heterogeneity is incompatible with eventual recombination or mutation events, but instead appears to result from colony fusion or the adoption of unrelated individuals. Indeed, colonies of P. punctata from the Dominican Republic and Barbados readily merged in the laboratory and, after elimination of one of the two reproductive workers, formed stable, genetically heterogeneous colonies. We discuss the possible causes and benefits of colony fusion in natural populations.

Synchronous reproduction of corals in the Red Sea

Multi-species synchronous spawning was first described on reefs off the east and west coast of Australia. In contrast, locally abundant species in the northern Red Sea and the central Pacific have little overlap in the time of reproduction. Consequently, the idea developed that high levels of spawning synchrony both within and among species was largely confined to Australian reefs. Here, we show that gamete maturity in colonies of the genus Acropora was highly synchronous in the Red Sea. In early April 2008, at two locations separated by 300 km, 13 of 24 species sampled had mature colonies, and a further 9 species had immature colonies. In late April–early May 2008, all colonies sampled had no oocytes, indicating colonies had spawned a few days after the full moon of 20 April 2008. Similarly, in 2009, 99% of colonies from 17 species at Hurghada were mature in late April, and all were empty in early May. Spawn slicks suggested many of these colonies had released gametes three night prior to the full moon on 8 May 2009. This level of synchrony in gamete maturity is among the highest ever recorded and similar to that typically recorded in Acropora assemblages on Australian reefs. While further work is required to document the night of gamete release, these data strongly suggest that high levels of spawning synchrony are a regular feature of these Red Sea coral assemblages and that multi-species spawning occurs on or around the full moon in April and/or May.

Colony growth and reproductive ability of feral nests of the introduced bumblebee Bombus terrestris in northern Japan

The introduced Bombus terrestris has been blamed for the decline of native bumblebee populations in Japan. To control this invasive species, it is necessary to understand its ecological traits in its newly established range. Here, we investigated the colony growth pattern and reproductive ability of feral nests of B. terrestris in northern Japan from 2003 to 2006. Nests collected at various times showed initially slow growth followed by rapid development. This pattern is consistent with findings in previous studies using laboratory-raised colonies. Our results also suggested that protandrous colonies tend to specialize in male production, whereas protogynous colonies specialize in gyne production, producing a split sex ratio in feral colonies of B. terrestris. The numerical population sex ratio was 1.40 gynes per male, calculated from the numbers of pre-emergence cocoons for gynes and males, showing a female-biased sex ratio at the population level. Mature colonies produced a mean of 376.5 cocoons and 90.2 gyne cocoons (22.1% of the total). The proportion of gynes produced by B. terrestris nests exceeded both those of conspecifics observed in other countries and those of consubgeneric native Japanese species. The propagule pressure hypothesis appears to explain the probability of establishment of this invasive species. Suitable nest sites for B. terrestris queens appeared to be in short supply, and B. terrestris may win usurpation contests against native species due to its large size, resulting in the decline of native bumblebee species.

Prudent sperm use by leaf-cutter ant queens

In many species, females store sperm between copulation and egg fertilization, but the consequences of sperm storage and patterns of sperm use for female life history and reproductive success have not been investigated in great detail. In hymenopteran insect societies (ants, bees, wasps), reproduction is usually monopolized by one or relatively few queens, who mate only during a brief period early in life and store sperm for later use. The queens of some ants are particularly long-lived and have the potential to produce millions of offspring during their life. To do so, queens store many sperm cells, and this sperm must remain viable throughout the years of storage. Queens should also be under strong selection to use stored sperm prudently when fertilizing eggs. We used the leaf-cutter ant Atta colombica to investigate the dynamics of sperm use during egg fertilization. We show that queens are able to fertilize close to 100 per cent of the eggs and that the average sperm use per egg is very low, but increases with queen age. The robustness of stored sperm was found to decrease with years of storage, signifying that senescence affects sperm either directly or indirectly via the declining glandular secretions or deteriorating sperm-storage organs. We evaluate our findings with a heuristic model, which suggests that the average queen has sperm for almost 9 years of normal colony development. We discuss the extent to which leaf-cutter ant queens have been able to optimize their sperm expenditure and infer that our observed averages of sperm number, sperm robustness and sperm use are consistent with sperm depletion being a significant cause of mortality of mature colonies of Atta leaf-cutter ants.

UNEXPLAINED SPLIT SEX RATIOS IN THE NEOTROPICAL PLANT-ANT,ALLOMERUS OCTOARTICULATUSVAR.DEMERARAE(MYRMICINAE): A TEST OF HYPOTHESES

We investigated sex allocation in the Neotropical ant Allomerus octoarticulatus var. demerarae. Because Allomerus is a plant symbiont, we could make geographically extensive collections of complete colonies and of foundresses in saplings, allowing us to estimate not only population- and colony-level sex allocation but also colony resource levels and the relatedness of competing ant foundresses. This species exhibits a strongly split sex ratio, with 80% of mature colonies producing >or=90% of one sex or the other. Our genetic analyses (DNA microsatellites) reveal that Allomerus has a breeding system characterized by almost complete monogyny and a low frequency of polyandry. Contrary to theoretical explanations, we find no difference in worker relatedness asymmetries between female- and male-specialist colonies. Furthermore, no clear link was found between colony sex allocation and life history traits such as the number of mates per queen, or colony size, resource level, or fecundity. We also failed to find significant support for male production by workers, infection by Wolbachia, local resource competition, or local mate competition. We are left with the possibility that Allomerus exhibits split sex ratios because of the evolution of alternative biasing strategies in queens or workers, as recently proposed in the literature.

Effect of exogenous cytokinins on dynamics of development and differentiation of infectious structures of the pathogen of wheat powdery mildew

The infectious structures for the development and differentiation of Erysiphe graminis DC. f. sp. tritici March., the pathogen of wheat powdery mildew, under the effects of exogenous zeatin was studied by methods of light and electron scanning microscopy. It was shown for the first time that physiologically active substances, specifically phytohormones of the cytokinin type, can affect dimensions of the halo revealed in the pathogen penetration site at cytochemical staining. Treatment with zeatin affected conidia germination and pathogen growth in the ectophytic stage. The concentration curve of the action of zeatin for the number of mature pathogen colonies (6 days after infection) was represented by a multiphase curve with two maxima (1 and 3 μM) and one minimum (1.5 μM). Similar curves have been obtained for the number of normal appressoria and for large halo diameters, which possibly indicates the existence of the factors affecting these both parameters, as well as the final number of pathogen colonies. The obtained data indicate that the origin of the multiphase dose response curve for effect of cytokinins on the development of powdery mildew pathogen is connected with factors that are active at the early stages of pathogenesis.

Phase-Unknown Linkage Mapping in Ants: Figure 1.

To create a genetic linkage map of an ant genome, the phase of a marker (i.e., which marker/allele came from the grandmother and which came from the grandfather of the individuals used for linkage mapping) must be known. However, field colonies contain only two generations: the queen(s) and their offspring. Normally, virgin queens disperse from their nest, mate with one or multiple males who die after mating, and start a new colony. A new reproductive brood (i.e., virgin queens and males) is usually produced after colony establishment, which can take from 6 mo to 3-5 yr. Hence, determining the phase of markers from field samples is impossible because the grandparents of a reproductive queen are no longer available for genetic analysis. Ants raised in the laboratory are likewise unsuitable for such analyses, because most ants cannot be bred regularly in the laboratory, and attempts to inseminate ant queens artificially have not been successful. However, three features facilitate "phase-unknown" linkage mapping for any ant (or other social hymenopteran) species, which is described here: (1) Mature colonies usually produce many reproductive offspring during a short period of time, which allows for the collection of a good-sized mapping population (>100 males) from a field colony; (2) Ants have a haplodiploid sex determination system (i.e., males are haploid and females diploid); (3) Queens produce males parthenogenetically. Thus, because they have no father, only those markers for which the queen is heterozygous are expected to segregate 1:1 in her male offspring.

Fungus gardens of the leafcutter ant Atta colombica function as egg nurseries for the snake Leptodeira annulata

Attine ants are well known for their mutualistic symbiosis with fungus gardens, but many other symbionts and commensals have been described. Here, we report the discovery of two clusters of large snake eggs in neighboring fungus gardens of a mature Atta colombica colony. The eggs were completely embedded within the fungus garden and were ignored by the host ants, even when we placed them into another, freshly excavated fungus garden of the same colony. All five eggs contained embryos and two snakes eventually hatched, which we identified as being banded cat eyed snakes Leptodeira annulata L. Ant fungus gardens are likely to provide ideal climatic conditions for developing snake eggs and almost complete protection from egg predation. Our observations therefore indicate that mature banded cat eyed snakes are able to enter and oviposit in large and well defended Atta colonies without being attacked by ant soldiers and that also newly hatched snakes manage to avoid ant attacks when they leaving their host colony. We speculate that L. annulata might use Atta and Acromyrmex leafcutter ant colonies as egg nurseries by some form of chemical insignificance, but more work is needed to understand the details of this interaction.

Vasa and the germ line lineage in a colonial urochordate

Germ cell sequestering in Animalia is enlightened by either, launching true germ line along epigenetic or preformistic modes of development, or by somatic embryogenesis, where no true germ line is set aside. The research on germ line-somatic tissue segregation is of special relevancy to colonial organisms like botryllid ascidians that reconstruct, on a weekly basis, completely new sets of male and female gonads in newly formed somatic tissues. By sequencing and evaluating expression patterns of BS-Vasa, the Botryllus schlosseri orthologue of Vasa, in sexually mature and asexual colonies during blastogenesis, we have demonstrated that the BS-Vasa mRNA and protein are not expressed exclusively in germ cell lineages, but appeared in cells repeatedly emerging de novo in the colony, independently of its sexual state. In addition, we recorded an immediate Vasa response to cellular stress (UV irradiation) indicating additional functions to its germ line assignments. To confirm germ lineage exclusivity, we examined the expression of three more stem cell markers (BS-Pl10, Bl-piwi and Oct4). Vasa co-expression with Pl10 and Oct4 was detected in germ line derivatives and with Bl-piwi in somatic tissues. Presumptive primordial germ cells (PGC-like cells), that are Vasa+/Pl10+/Oct4+ and 6–12 μm in diameter, were first detected in wrapped-tail embryos, in oozooids, in sexual/asexual colonies, within a newly identified PGC niche termed as ‘budlet niche’, and in circulating blood borne cells, indicating epigenetic embryogenesis. Alternatively, BS-Vasa co-expression with piwi orthologue, an omnipresent bona fide stemness flag, in non germ line cell populations, may indicate germ cell neogenesis (somatic embryogenesis) in B. schlosseri. Both alternatives are not necessarily mutually exclusive.

EPHEMERAL WINDOWS OF OPPORTUNITY FOR HORIZONTAL TRANSMISSION OF FUNGAL SYMBIONTS IN LEAF-CUTTING ANTS

Evolutionary theory predicts that hosts are selected to prevent mixing of genetically different symbionts when competition among lineages reduces the productivity of a mutualism. The symbionts themselves may also defend their interests: recent studies of Acromyrmex leaf-cutting ants showed that somatic incompatibility enforces single-clone gardens within mature colonies, thereby constraining horizontal transmission of fungal symbionts. However, phylogenetic analyses indicate that symbiont switches occur frequently enough to remove most signs of host-symbiont cocladogenesis. Here we resolve this paradox by showing that transmission among newly founded Acromyrmex colonies is not constrained. All tested queens of sympatric A. octospinosus and A. echinatior offered a novel fragment of fungus garden accepted the new symbiont. The outcome was unaffected by genetic distance between the novel and the original symbiont, and by the ant species the novel symbiont came from. The colony founding stage may thus provide an efficient but transient window for horizontal transmission, in which the fungus is unable to actively defend its partnership position before the host feeds on it, so that host fecal droplets remain compatible with alternative strains during the early stage of colony founding. We discuss how brief stages of low commitment between partners may increase the evolutionary stability of ancient coevolved mutualisms.

MODELING THE MAINTENANCE OF A DEPENDENT LINEAGE SYSTEM: THE INFLUENCE OF POSITIVE FREQUENCY-DEPENDENT SELECTION ON SEX RATIO

In insect societies, worker versus queen development (reproductive caste) is typically governed by environmental factors, but some Pogonomyrmex seed-harvester ants exhibit strict genetic caste determination, resulting in an obligate mutualism between two reproductively isolated lineages. Queens mate randomly with multiple males from each lineage and intralineage crosses produce new queens, whereas interlineage crosses produce workers. Early colony survival is negatively frequency dependent; when lineage frequencies are unequal, queens from the rarer lineage benefit because they acquire more interlineage sperm, and produce more workers. Here we examine theoretically and empirically the effect of relative lineage frequency on sex ratio. We predict that the ratio of inter- to intralineage sperm acquired by queens of each lineage will affect the sex ratio produced at colony maturity. Consistent with model predictions, we found that gyne production in mature colonies was positively frequency dependent, increasing significantly with increasing lineage frequency across 15 populations. Unequal lineage frequencies are common and likely maintained by a complex interplay between an ecological advantage specific to one lineage, and opposing frequency-dependent selection pressures experienced throughout the colonies life-cycle; rare lineage colonies benefit during early colony growth, and common lineage colonies benefit at reproductive maturity.