Natural Populations Of Vertebrates Research Articles

Contrasting epidemic histories reveal pathogen-mediated balancing selection on class II MHC diversity in a wild songbird.

The extent to which pathogens maintain the extraordinary polymorphism at vertebrate Major Histocompatibility Complex (MHC) genes via balancing selection has intrigued evolutionary biologists for over half a century, but direct tests remain challenging. Here we examine whether a well-characterized epidemic of Mycoplasmal conjunctivitis resulted in balancing selection on class II MHC in a wild songbird host, the house finch (Carpodacus mexicanus). First, we confirmed the potential for pathogen-mediated balancing selection by experimentally demonstrating that house finches with intermediate to high multi-locus MHC diversity are more resistant to challenge with Mycoplasma gallisepticum. Second, we documented sequence and diversity-based signatures of pathogen-mediated balancing selection at class II MHC in exposed host populations that were absent in unexposed, control populations across an equivalent time period. Multi-locus MHC diversity significantly increased in exposed host populations following the epidemic despite initial compromised diversity levels from a recent introduction bottleneck in the exposed host range. We did not observe equivalent changes in allelic diversity or heterozygosity across eight neutral microsatellite loci, suggesting that the observations reflect selection rather than neutral demographic processes. Our results indicate that a virulent pathogen can exert sufficient balancing selection on class II MHC to rescue compromised levels of genetic variation for host resistance in a recently bottlenecked population. These results provide evidence for Haldane's long-standing hypothesis that pathogens directly contribute to the maintenance of the tremendous levels of genetic variation detected in natural populations of vertebrates.

Senescence and age‐specific trade‐offs between reproduction and survival in female Asian elephants

Although studies on laboratory species and natural populations of vertebrates have shown reproduction to impair later performance, little is known of the age-specific associations between reproduction and survival, and how such findings apply to the ageing of large, long-lived species. Herein we develop a framework to examine population-level patterns of reproduction and survival across lifespan in long-lived organisms, and decompose those changes into individual-level effects, and the effects of age-specific trade-offs between fitness components. We apply this to an extensive longitudinal dataset on female semi-captive Asian timber elephants (Elephas maximus) and report the first evidence of age-specific fitness declines that are driven by age-specific associations between fitness components in a long-lived mammal. Associations between reproduction and survival are positive in early life, but negative in later life with up to 71% of later-life survival declines associated with investing in the production of offspring within this population of this critically endangered species.

Extreme natal philopatry in female Antarctic fur seals ( Arctocephalus gazella)

Natal philopatry is an important component of mammalian behaviour but is difficult to study in natural vertebrate populations due to the requirement for long-term individual-based spatial observations. Consequently, we quantified fine-scale patterns of natal philopatry in an intensively studied colony of Antarctic fur seals ( Arctocephalus gazella), where a scaffold walkway allows individual locations to be measured to the nearest metre. Using subcutaneous PIT tags, we tracked the early life histories of 335 females born within the colony, of which 38 were resighted as breeding adults. We found that individual females returned to as little as one body length (2 m) of their birth locations. Moreover, distances between natal and pupping sites were not correlated with female age, but instead tended to decrease with the number of seasons an individual was sighted ashore. This suggests that breeding experience may be a better predictor than age of the ability of females to occupy preferred sites within fur seal colonies.

DNA EVIDENCE FOR NONHYBRID ORIGINS OF PARTHENOGENESIS IN NATURAL POPULATIONS OF VERTEBRATES

Naturally occurring unisexual reproduction has been documented in less than 0.1% of all vertebrate species. Among vertebrates, true parthenogenesis is known only in squamate reptiles. In all vertebrate cases that have been carefully studied, the clonal or hemiclonal taxa have originated through hybridization between closely related sexual species. In contrast, parthenogenetic reproduction has arisen in invertebrates by a variety of mechanisms, including likely cases of "spontaneous" (nonhybrid) origin, a situation not currently documented in natural populations of vertebrates. Here, we present molecular data from the Neotropical night lizard genus Lepidophyma that provides evidence of independent nonhybrid origins for diploid unisexual populations of two species from Costa Rica and Panama. Our mitochondrial and nuclear phylogenies are congruent with respect to the unisexual taxa. Based on 14 microsatellite loci, heterozygosity (expected from a hybrid origin) is low in Lepidophyma reticulatum and completely absent in unisexual L. flavimaculatum. The unique value of this system will allow direct comparative studies between parthenogenetic and sexual lineages in vertebrates, with an enormous potential for this species to be a model system for understanding the mechanisms of nonhybrid parthenogenesis.

Immunomodulatory parasites and toll-like receptor-mediated tumour necrosis factor alpha responsiveness in wild mammals

BackgroundImmunological analyses of wild populations can increase our understanding of how vertebrate immune systems respond to 'natural' levels of exposure to diverse infections. A major recent advance in immunology has been the recognition of the central role of phylogenetically conserved toll-like receptors in triggering innate immunity and the subsequent recruitment of adaptive response programmes. We studied the cross-sectional associations between individual levels of systemic toll-like receptor-mediated tumour necrosis factor alpha responsiveness and macro- and microparasite infections in a natural wood mouse (Apodemus sylvaticus) population.ResultsAmongst a diverse group of macroparasites, only levels of the nematode Heligmosomoides polygyrus and the louse Polyplax serrata were correlated (negatively) with innate immune responsiveness (measured by splenocyte tumour necrosis factor alpha responses to a panel of toll-like receptor agonists). Polyplax serrata infection explained a strikingly high proportion of the total variation in innate responses. Contrastingly, faecal oocyst count in microparasitic Eimeria spp. was positively associated with innate immune responsiveness, most significantly for the endosomal receptors TLR7 and TLR9.ConclusionAnalogy with relevant laboratory models suggests the underlying causality for the observed patterns may be parasite-driven immunomodulatory effects on the host. A subset of immunomodulatory parasite species could thus have a key role in structuring other infections in natural vertebrate populations by affecting the 'upstream' innate mediators, like toll-like receptors, that are important in initiating immunity. Furthermore, the magnitude of the present result suggests that populations free from immunosuppressive parasites may exist at 'unnaturally' elevated levels of innate immune activation, perhaps leading to an increased risk of immunopathology.

EVOLUTIONARY POTENTIAL OF A LARGE MARINE VERTEBRATE: QUANTITATIVE GENETIC PARAMETERS IN A WILD POPULATION

Estimating quantitative genetic parameters ideally takes place in natural populations, but relatively few studies have overcome the inherent logistical difficulties. For this reason, no estimates currently exist for the genetic basis of life-history traits in natural populations of large marine vertebrates. And yet such estimates are likely to be important given the exposure of this taxon to changing selection pressures, and the relevance of life-history traits to population productivity. We report such estimates from a long-term (1995-2007) study of lemon sharks (Negaprion brevirostris) conducted at Bimini, Bahamas. We obtained these estimates by genetically reconstructing a population pedigree (117 dams, 487 sires, and 1351 offspring) and then using an "animal model" approach to estimate quantitative genetic parameters. We find significant additive genetic (co)variance, and hence moderate heritability, for juvenile length and mass. We also find substantial maternal effects for these traits at age-0, but not age-1, confirming that genotype-phenotype interactions between mother and offspring are strongest at birth; although these effects could not be parsed into their genetic and nongenetic components. Our results suggest that human-imposed selection pressures (e.g., size-selective harvesting) might impose noteworthy evolutionary change even in large marine vertebrates. We therefore use our findings to explain how maternal effects may sometimes promote maladaptive juvenile traits, and how lemon sharks at different nursery sites may show "constrained local adaptation." We also show how single-generation pedigrees, and even simple marker-based regression methods, can provide accurate estimates of quantitative genetic parameters in at least some natural systems.

GEOGRAPHIC VARIATION IN MATERNAL INVESTMENT: ACIDITY AFFECTS EGG SIZE AND FECUNDITY IN RANA ARVALIS

Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.

Sex differences in ageing in natural populations of vertebrates

In many long-lived vertebrates (including humans), adult males have shorter lifespans than adult females, partly as a result of higher annual rates of mortality in males and partly owing to sex differences in the rate of ageing. A probable explanation of the evolution of sex differences in ageing is that, in polygynous species, intense intrasexual competition between males restricts the number of seasons for which individual males are able to breed successfully, weakening selection pressures favouring adult longevity in males relative to females. If this is the case, sex differences in adult longevity and in the onset and rate of senescence should be greater in polygynous species than in monogamous ones and their magnitude should be related to the duration of effective breeding males compared with females. Here, we use data from longitudinal studies of vertebrates to show that reduced longevity in adult males (relative to females) is commonly associated with a more rapid decline in male than female survival with increasing age and is largely confined to polygynous species. The magnitude of sex differences in adult longevity in different species is consistently related to the magnitude of sex differences in the duration of effective breeding, calculated across surviving adults. Our results are consistent with the suggestion that sex differences in senescence in polygynous species are a consequence of weaker selection for longevity in males than females.

Molecular and cellular studies in evolutionary physiology of natural vertebrate populations: influences of individual variation and genetic components on sampling and measurements

Studies combining ecological, genetic and physiological approaches are needed in evolutionary biology. Although the combination of approaches has been emphasized, such studies have been rare with regard to molecular and cellular studies on natural vertebrate populations. The major reasons for this are that the generation time of vertebrates is long and it is difficult to find a molecular or cell physiological measurement that is both relevant for the fitness of the population and can be repeated an adequate number of times to enable estimations of individual variability. The paucity of suitable physiological parameters is partly due to the fact that most physiological studies have not been directed towards understanding the behaviour of populations but towards understanding the basic mechanisms of the function of individuals. Also, physiological measurements that appear most relevant from the point of view of evolutionary studies are often integrative functions, composed of the function of many genes. When dissecting the integrative functions into components, it is often observed that the same integrative response can be achieved via different routes, i.e. changes in the responses of different genes. To enable cellular and molecular physiological studies to be increasingly combined with ecological and genetic studies, it is important that such studies include and report individual variability and that the sample size is increased. In addition, more sophisticated statistical methods should be used than is traditionally done, and when the function of most genes in the integrative response are not known, techniques such as QTL mapping should be used. Hitherto in vertebrates, the methodology has mainly been used in production biology (e.g. meat or milk production). With regard to combining genomic and physiological studies, one must bear in mind that the massive datasets associated with genomic studies need to be further enlarged to enable estimates of individual variation. It is also important to remember that microarray and proteomic data give the levels of mRNA and proteins, respectively. Since the function of the protein can be regulated independently of its transcription or its level in the cell, direct physiological measurements are also needed if estimations of protein activity in the individuals of a population are wanted.

Admixture and patterns of linkage disequilibrium in a free‐living vertebrate population

Linkage disequilibrium (LD), a measure of nonrandom association of alleles at different loci, is of great interest to evolutionary geneticists as it can be used to help identify loci that explain phenotypic variation. Surveys of the extent of LD across genomes have been carried out in a number of systems, most notably humans and model organisms. However, studies of natural populations of vertebrates have rarely been performed. Here, we describe an investigation of LD in a free-living island population of red deer Cervus elaphus. Relatively high levels of LD extended several tens of centimorgans, and significant LD was frequently detected between unlinked markers. The magnitude of LD varied depending on how the population was sampled. It also varied across different chromosomes, and was shown to be a function of sample size, intermarker distance and marker heterozygosity. A recent admixture event in the population led to an ephemeral increase in LD. Association mapping may be possible in this population, although a high 'baseline' level of LD could lead to false positive associations between marker loci and a trait of interest.

Fishing for the secrets of vertebrate evolution in threespine sticklebacks

The threespine stickleback (Gasterosteus aculeatus) is rapidly emerging as a new model genetic system to study questions at the interface of evolution and development. The relatively rapid and recent diversification of this small teleost fish, combined with the development of genetic and genomic tools for this fish, provides an unprecedented opportunity to identify the genetic and molecular basis of morphological variation in natural populations of vertebrates. Recently, the genes underlying two different adaptive morphological traits in stickleback have been identified. This work has provided answers to four longstanding questions in the field of evolution and development: (1) How many genes underlie morphological variation in natural populations? (2) What are the genes that underlie morphological variation in natural populations? (3) Do coding or regulatory mutations underlie morphological evolution? (4) What is the molecular and genetic basis of parallel morphological evolution? Because stickleback populations also display natural variation in morphology, life history, physiology, and behavior, extending the approaches used to identify the genetic basis of morphological variation in sticklebacks to other phenotypes is sure to yield further important insights into the genetic and developmental basis of diversity in natural populations.

An Mhc class I allele associated to the expression of T-dependent immune response in the house sparrow

The major histocompatibility complex (Mhc) encodes for highly variable molecules, responsible for foreign antigen recognition and subsequent activation of immune responses in hosts. Mhc polymorphism should hence be related to pathogen resistance and immune activity, with individuals that carry either a higher diversity of Mhc alleles or one specific Mhc allele exhibiting a stronger immune response to a given antigen. Links between Mhc alleles and immune activity have never been explored in natural populations of vertebrates. To fill this gap, we challenged house sparrows (Passer domesticus) with two T-dependent antigens (phytohemagglutinin and sheep red blood cells) and examined both primary and secondary immune responses in relation to their Mhc class I genotypes. The total number of Mhc alleles had no influence on either primary or secondary response to the two antigens. One particular Mhc allele, however, was associated with an increased response to both antigens. Our results point toward a contribution of the Mhc, or of other genes in linkage disequilibrium with the Mhc, in the regulation of immune responses in a wild animal species.

Geographic and within-population structure in variable resistance to parasite species and strains in a vertebrate host

Host resistance to parasites and parasite infectivity may be subject to significant genetically determined variation within species. However, relatively little is known of how this variability is structured in natural vertebrate populations and their macroparasites. A laboratory experiment on host susceptibility–parasite infectivity variation in a wildlife host–parasite system (subspecies of the anuran X. laevis and their polystome flatworms), including 33 pairwise allopatric and sympatric host–parasite combinations (three parasite geographical isolates×11 host full-sibling families, n=600), revealed a complex pattern of infection success. Results amongst host sibships from different localities suggested that infection success was subject to a highly significant locality×parasite isolate interaction. Within localities, a highly significant sibship×isolate interaction also occurred in one of two groups of sibships examined. The existence of such interactions suggests a potential for frequency-dependent, Red Queen-like selection. Interaction between locality and isolate was partly due to higher infection levels in sympatric combinations, consistent with a general pattern of host-specific adaptation. However, some allopatric combinations produced unpredictably high infection levels, resulting in very asymmetrical cross-infectivity patterns (where the reciprocal cross-infections produced negligible infection). This phylogeographically structured host–parasite system may, therefore, sometimes generate local parasite strains with high infectivity to allopatric hosts. Secondary contact between populations could thus result in significant, and unequal, transfer of parasites.

Demographic and genetic estimates of effective population size (Ne) reveals genetic compensation in steelhead trout.

Estimates of effective population size (Ne) are required to predict the impacts of genetic drift and inbreeding on the evolutionary dynamics of populations. How the ratio of Ne to the number of sexually mature adults (N) varies in natural vertebrate populations has not been addressed. We examined the sensitivity of Ne/N to fluctuations of N and determined the major variables responsible for changing the ratio over a period of 17 years in a population of steelhead trout (Oncorhynchus mykiss) from Washington State. Demographic and genetic methods were used to estimate Ne. Genetic estimates of Ne were gained via temporal and linkage disequilibrium methods using data from eight microsatellite loci. DNA for genetic analysis was amplified from archived smolt scales. The Ne/N from 1977 to 1994, estimated using the temporal method, was 0.73 and the comprehensive demographic estimate of Ne/N over the same time period was 0.53. Demographic estimates of Ne indicated that variance in reproductive success had the most substantial impact on reducing Ne in this population, followed by fluctuations in population size. We found increased Ne/N ratios at low N, which we identified as genetic compensation. Combining the information from the demographic and genetic methods of estimating Ne allowed us to determine that a reduction in variance in reproductive success must be responsible for this compensation effect. Understanding genetic compensation in natural populations will be valuable for predicting the effects of changes in N (i.e. periods of high population density and bottlenecks) on the fitness and genetic variation of natural populations.

Expression of P-glycoprotein and cytochrome P450 1A in intertidal fish ( Anoplarchus purpurescens) exposed to environmental contaminants

Whether P-glycoproteins (P-gps) like those which confer multidrug resistance in tumor cell lines are important in adaptation to chemicals in natural populations of vertebrates exposed to contaminant mixtures is the focus of this study. P-gp expression was examined in the intertidal fish high cockscomb blenny ( Anoplarchus purpurescens) exposed to crude oil or pulp mill effluent. The relationship between P-gp expression and cytochrome P450 1A (CYP1A) induction also was investigated. Immunohistochemical (IHC) analysis revealed that levels of P-gp expression in the bile canaliculi were three- to five-fold greater in oil exposed fish than in control fish. Levels of P-gp expression were highly correlated with hepatic CYP1A levels previously measured in these fish. In fish from sites near pulp mills, P-gp expression in freshly caught fish did not correlate with proximity to pulp mills. However, hepatic P-gp expression levels in freshly caught fish were 14-fold higher than in fish from those sites that were depurated in clean water for 6 weeks. CYP1A levels were also elevated in liver of freshly caught as compared with depurated fish. Expression of neither CYP1A nor P-gp was elevated in depurated fish exposed to sediment and food from within the original pulp mill effluent stream. Depurated fish, which were injected with the aryl hydrocarbon receptor (AHR) agonist ß-naphthoflavone (BNF) showed an expected induction of CYP1A but no induction of P-gp. These results suggest that in blennies, unlike CYP1A, P-gp expression is not regulated by the AHR pathway; although P-gp and CYP1A both may be induced by some compounds in petroleum and unidentified xenobiotics at field sites. While our data indicate that CYP1A and P-gp are not coordinately regulated, these proteins may play complementary roles in cellular detoxification. Thus the elevation of P-gp activity may be an important mechanism of multixenobiotic resistance for organisms, such as intertidal fish, which are commonly exposed to anthropogenic contaminants and naturally occurring toxins.

The molecular basis of an avian plumage polymorphism in the wild: a melanocortin-1-receptor point mutation is perfectly associated with the melanic plumage morph of the bananaquit, Coereba flaveola.

Evolution depends on natural selection acting on phenotypic variation, but the genes responsible for phenotypic variation in natural populations of vertebrates are rarely known. The molecular genetic basis for plumage color variation has not been described in any wild bird. Bananaquits (Coereba flaveola) are small passerine birds that occur as two main plumage variants, a widespread yellow morph with dark back and yellow breast and a virtually all black melanic morph. A candidate gene for this color difference is the melanocortin-1 receptor (MC1R), a key regulator of melanin synthesis in feather melanocytes. We sequenced the MC1R gene from four Caribbean populations of the bananaquit; two populations of the yellow morph and two populations containing both the yellow morph and the melanic morph. A point mutation resulting in the replacement of glutamate with lysine was present in at least one allele of the MC1R gene in all melanic birds and was absent in all yellow morph birds. This substitution probably causes the color variation, as the same substitution is responsible for melanism in domestic chickens and mice. The evolutionary relationships among the MC1R haplotypes show that the melanic alleles on Grenada and St. Vincent had a single origin. The low prevalence of nonsynonymous substitutions among yellow haplotypes suggests that they have been under stabilizing selection, whereas strong selective constraint on melanic haplotypes is absent. We conclude that a mutation in the MC1R is responsible for the plumage polymorphism in a wild bird population and that the melanic MC1R alleles in Grenada and St. Vincent bananaquit populations have a single evolutionary origin from a yellow allele.

Similarity is Relative: Hierarchy of Genetic Similarities in the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) in Hawaii

Termites (Isoptera) have a strong hierarchical structure due to their social organization. To interpret genetic differences within and among termite populations it is first necessary to create a quantitative scale to determine the meaning of “different.” To accomplish this, the entire hierarchy of genetic similarities in a Hawaiian population of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) was defined by multilocus DNA fingerprinting using the oligonucleotide (GTG)5 as a probe. The genetic background similarity in the population, based on the genetic similarity between unrelated termites, was 0.21 and falls in the range of other natural invertebrate and vertebrate populations. The mean genetic similarity of termites collected from the same collection site was 0.64 (range, 0.58–0.72). This equals an average relatedness of 0.54 within collection sites. The genetic similarity among the offspring of an artificially outbred laboratory colony lies in the same magnitude (0.65). However, genetic similarity within field colonies was significantly lower than the genetic similarity within laboratory colonies derived from pairs of siblings (mean 0.79). This indicates moderate inbreeding within most of the field colonies. This wide range of genetic similarities defines the basis for a detailed description of the population structure as well as the identification of individual colonies of C. formosanus.

Cadmium toxicity among wildlife in the Colorado Rocky Mountains.

Cadmium is known to be both extremely toxic and ubiquitous in natural environments. It occurs in almost all soils, surface waters and plants, and it is readily mobilized by human activities such as mining. As a result, cadmium has been named as a potential health threat to wildlife species; however, because it exists most commonly in the environment as a trace constituent, reported incidences of cadmium toxicity are rare. Here we have measured trace metals in the food web and tissues of white-tailed ptarmigan (Lagopus leucurus) in Colorado. Our results suggest that cadmium toxicity may be more common among natural populations of vertebrates than has been appreciated to date and that cadmium toxicity may often go undetected or unrecognized. In addition, our research shows that ingestion of even trace quantities of cadmium can influence not only the physiology and health of individual organisms, but also the demographics and the distribution of species.

PARASITE-MEDIATED SELECTION AGAINST INBRED SOAY SHEEP IN A FREE-LIVING ISLAND POPULATON.

Parasites are thought to provide a selective force capable of promoting genetic variation in natural populations. One rarely considered pathway for this action is via parasite-mediated selection against inbreeding. If parasites impose a fitness cost on their host and the offspring of close relatives have greater susceptibility to parasites due to the increased homozygosity that results from inbreeding, then parasite-mediated mortality may select against inbred individuals. This hypothesis has not yet been tested within a natural vertebrate population. Here we show that relatively inbred Soay sheep (Ovis aries), as assessed by microsatellite heterozygosity, are more susceptible to parasitism by gastrointestinal nematodes, with interactions indicating greatest susceptibility among adult sheep at high population density. During periods of high overwinter mortality on the island of Hirta, St. Kilda, Scotland, highly parasitised individuals were less likely to survive. More inbred individuals were also less likely to survive, which is due to their increased susceptibility to parasitism, because survival was random with respect to inbreeding among sheep that were experimentally cleared of their gastrointestinal parasite burden by anthelminthic treatment. As a consequence of this selection, average microsatellite heterozygosity increases with age in St. Kildan Soay sheep. We suggest that parasite-mediated selection acts to maintain genetic variation in this small island population by removing less heterozygous individuals.

Molecular advances in understanding social insect population structure.

Social insects present many phenomena seen in all organisms but in more extreme forms and with larger sample sizes than those observable in most natural populations of vertebrates. Microsatellites are proving very much more informative than allozymes for the analysis of population biological problems, and prolifically polymorphic markers are fairly readily developed. In addition, the male-haploid genetic system of many social insects facilitates genetic analysis. The ability to amplify DNA from sperm stored in a female's sperm storage device enables the determination of mating types long after the death of the short-lived males, in addition to information on the degree of mixing of sperm from different males. Mitochondrial (mt) DNA sequences are also proving important, not only in phylogenetic studies but also in molecular population genetics, as a tracer of female movements. Mitochondrial markers have definitively shown the movement of females between colonies, challenging models giving exclusive primacy to kin selection as the explanation for multiqueen colonies, in Australian meat ants, Iridomyrmex purpureus, and the aridzone queenless ant Rhytidoponera sp. 12. Microsatellite and mtDNA variation are being studied in Camponotus consobrinus sugar ants, showing an unexpected diversity of complexity in colony structure, and microsatellites have shown that transfer of ants between nests of the weaver ant Polyrhachis doddi must be slight, despite an apparent lack of hostility.