The origins and relatedness of multiple reproductives in colonies of the termite Nasutitermes corniger

Colonies of the termite Nasutitermes corniger often contain multiple reproductive queens and kings. We used double-strand conformation polymorphism (DSCP) analysis of mitochondrial DNA (mtDNA) to determine the probable origins of co-occurring reproductives. Colonies differed in queen and king number, in the number of nests containing reproductives, and in the genetic relationships among reproductives. Most of the 44 colonies contained a single pair of maternally unrelated reproductives. In the two single-nest colonies with a pair of queens, the two queens differed in mtDNA haplotype, suggesting nest-founding by unrelated queens. In the seven single-nest colonies with larger numbers of reproductives (11–49), all reproductives shared the same haplotype, a pattern consistent with replacement of a single pair by several offspring. As predicted by theory, the number of coexisting queens was greater for replacement reproductives than for co-foundresses. Several complex colonies contained multiple queens of two or more haplotypes distributed among several interconnected nests. This indicates that several matrilines can persist within a colony through one or more generations of budding and replacement, a hypothesis confirmed by orphaning experiments. The various modes of termite colony formation rival the diversity seen in ant species and demonstrate the remarkable convergence of behaviours between the two groups.     

Mitochondrial DNA distributions indicate colony propagation by single matri-lineages in the social spider Stegodyphus dumicola (Eresidae)

Colony-dwelling social spiders of the genus Stegodyphus are characterized by high colony turnover, within-colony mating, inbreeding and skewed sex ratios. These phenomena may purge genetic variation from the entire species gene pool. Social Stegodyphus have previously been discussed as ecologically unstable and evolutionary dead ends. We investigated the distribution and age (sequence divergence) of mitochondrial DNA variation for inferences of colony propagation, colony discreteness and maintenance of genetic variation in the social spider S. dumicola . In contrast to our expectations, we found abundant mtDNA variation, consisting of 15 haplotypes belonging to four haplotype lineages. Lineage divergence ranged between 2.75 and 6% for the gene ND1. Nearly all colonies (86%) were monomorphic and even neighbour colonies showed fixed differences. Simulations show that genetic drift in multifounder colonies cannot alone explain monomorphism within colonies. Haplotypes in polymorphic colonies and from neighbouring colonies were always genealogically similar. Monomorphism and the genealogical pattern among colonies suggest 'clonal' colony propagation involving single matrilineages. The divergence of haplotype lineages and distribution of haplotypes imply that colony turnover is not high enough to purge genetic variation in the species gene pool, and that S. dumicola as a species is old enough to question the instability (in ecological time) of a social spider.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 591–600.     

Micro- and macrogeographical genetic structure of colonies of naked mole-rats Heterocephalus glaber

Patterns of genetic structure in eusocial naked mole-rat populations were quantified within and among geographically distant populations using multilocus DNA fingerprinting and mitochondrial DNA (mtDNA) sequence analysis. Individuals within colonies were genetically almost monomorphic, sharing the same mtDNA control region haplotype and having coefficients of band sharing estimated from DNA fingerprints ranging from 0.93 to 0.99. Family analysis of a hybrid captive colony of naked mole-rats with increased levels of genetic variability using multilocus DNA fingerprinting gave results consistent with Mendelian inheritance, and has revealed for the first time that multiple paternity can occur. In a survey of wild colonies from Ethiopia, Somalia and locations in northern and southern Kenya, we have examined mtDNA control region sequence variation in 42 individuals from 15 colonies, and together with multilocus DNA fingerprinting and mtDNA cytochrome- b sequence analysis in selected individuals have shown that these populations show considerable genetic divergence. Most of the variance in sequence divergence was found to be between geographical locations (Φ_ct= 0.68) and there was a significant correlation between sequence divergence and geographical separation of haplotypes. Six colonies from Mtito Andei in southern Kenya shared the same control region haplotype, suggesting a recent common maternal ancestor. In contrast, out of four colonies at Lerata in north Kenya, three haplotypes were identified, and phylogenetic analysis suggests that this area may be a zone where two distinct lineages are in close proximity. Genetic distances were maximal between Ethiopian and southern Kenyan populations at 5.8% for cytochrome- b , and are approaching interspecific values seen between other Bathyergids.     

Unrelated queens coexist in colonies of the termite Macrotermes michaelseni

Relatedness increases the likelihood of cooperation within colonies of social insects. Polygyny, the coexistence of numerous reproductive females (queens) in a colony, is common in mature colonies of the termite Macrotermes michaelseni. In this species, polygyny results from pleometrosis and from several female alates that jointly found a new colony. To explain this phenomenon, it was suggested that only related females cooperate and survive during maturation of colonies. Using multilocus fingerprints as well as microsatellites, we showed that nestmate queens in mature colonies are unrelated. Furthermore, we found that all nestmate queens contributed to the production of steriles. Even in mature colonies, several matrilines of steriles coexist within a colony. Although genetic diversity within colonies may increase the likelihood of conflicts, high genetic diversity may be important for foraging, colony growth, and resistance to disease and parasites.     

Restricted effective queen dispersal at a microgeographic scale in polygynous populations of the ant Formica exsecta

Ecological constraints on effective dispersal have been suggested to be a key factor influencing social evolution in animal societies as well as the shift from single queen colonies (monogyny) to multiple queen colonies (polygyny) in ants. However, little is known about the effective dispersal patterns of ant queens. Here we investigate the microgeographic genetic structure of mitochondrial haplotypes in polygynous populations of the ant Formica exsecta, both between pastures and among nests within pastures. An analysis of molecular variance revealed a very high genetic differentiation (phiST = 0.72) between pastures, indicating that queens rarely disperse successfully between pastures, despite the fact that pastures were sometimes as close as 1 km. Most of the pastures contained only a single haplotype, and haplotypes were frequently distinct between nearby pastures and even between groups of nests within the same pasture. In the three pastures that contained several haplotypes, haplotypes were not randomly distributed, the genetic differentiation between nests being phiST = 0.17, 0.52, and 0.69. This indicates that most queens are recruited within their parental colonies. However, a large proportion of nests contained more than one haplotype, demonstrating that colonies will sometimes accept foreign queens. The relatedness of mitochondrial genes among nestmates varied between 0.62 and 0.75 when relatedness was measured within each pasture and ranged between 0.72 and 1.0 when relatedness was assessed with all pastures as a reference population. Neighboring nests were more genetically similar than distant ones, and there was significant isolation by distance. This pattern may be due to new nests being formed by budding or by limited effective queen dispersal, probably on foot between neighboring nests. These results show that effective queen dispersal is extremely restricted even at a small geographical scale, a pattern consistent with the idea that ecological constraints are an important selective force leading to the evolution and maintenance of polygyny.     

High within-population mitochondrial DNA variation due to microvicariance and population mixing in the land snail Euhadra quaesita (Pulmonata: Bradybaenidae)

A high level of geographical variation at an exceptionally fine scale was detected in the mitochondrial 16S ribosomal RNA genes of a land snail species Euhadra quaesita from the Kanto region of Japan. In total, 50 haplotypes were detected from 27 populations, with most sample sites possessing private alleles. In some individual populations the different haplotypes do not fall as a monophyletic group, so that some of the haplotypes are phylogenetically distant, differing from each other by > 10%. In contrast, phylogenetically similar haplotypes were found in separate sites at long distances from their main distribution. Together, these findings strongly suggest that contraction and expansion of populations has occurred repeatedly in the past. The subsequent expansion of populations and migration from different areas may have mixed distant populations. This repetition of isolation and mixing has resulted in an exceptionally fine scale of geographical variation, and the accumulation of high genetic diversity within and between populations of this species.     

Mitochondrial DNA (mtDNA) reveals that female Bechstein's bats live in closed societies

We present a microgeographic analysis of mitochondrial DNA (mtDNA) in Bechstein's bats using three sources of control region sequence variability, including a novel mtDNA microsatellite, to assess individual relatedness both within and among 10 maternity colonies. Comparison of marker variability among 268 adult females revealed little genetic variability within each colony. However, most colonies were clearly distinguished by colony-specific mitochondrial haplotypes (total n = 28). Low intracolony variability and strong haplotype segregation among colonies, was reflected by an extraordinary high FST of 0.68, indicating a very low intercolony dispersal rate of approximately one female in five generations. Haplotype distribution among 18 solitary males showed that males frequently disperse between colony locations, indicating the absence of dispersal barriers. Bechstein's bat maternity colonies are thus closed groups that comprise 20-40 females probably belonging to only one or, at most, two matrilines. The genetic population structure of Bechstein's bats is in agreement with the hypothesis that females seek familiar and, at least, partially related cooperation partners for raising their young. Alternatively strong philopatry might reflect the importance of profound roost or habitat knowledge for successful reproduction in female Bechstein's bats.     

The local distribution of highly divergent mitochondrial DNA haplotypes in toque macaques Macaca sinica at Polonnaruwa, Sri Lanka

Surveys of mitochondrial DNA (mtDNA) variation in macaque monkeys have revealed extremely high levels of intraspecific divergence among haplotypes. One consistent pattern that has emerged from these studies is that divergent haplotypes are geographically segregated so that sampling a few matrilines from a given region shows them to be identical, or a closely related subset of haplotypes. Geographically structured mtDNA variation has also been commonly observed in other taxa. In this study, haplotype variation and distribution are studied in detail within a local population of toque macaques. The results show that highly divergent haplotypes, differing by 3.1% in their nucleotide sequences, coexist in this population and that they may be spatially segregated even on this micro-geographic scale. Furthermore, these differences are maintained between social groups that exchange male migrants, and thus nuclear genes, frequently.     

Complex fine‐scale phylogeographical patterns in a putative refugial region for Fagus sylvatica (Fagaceae)

Broad‐scale plastid (chloroplast) DNA studies of beech (Fagus sylvatica) populations suggest the existence of glacial refugia and introgression zones in south‐eastern Europe. We choose a possible refugium of beech in northern Greece, Mt. Paggeo, which hosts a private plastid haplotype for beech, to conduct a fine‐scale genetic study. We attempt to confirm or reject the hypothesis of the existence of a small‐scale refugium and to gain an understanding of the ecological and topographical factors affecting the spatial distribution of plastid haplotypes in the area. Our results reveal a high haplotype diversity on Mt. Paggeo, but the overall distribution of haplotypes shows no significant correlation with the ecological characteristics of the beech forests. However, the private haplotype is found at high frequencies in beech forests located in or near ravines, having a high spatial overlap with a relict vegetation type occurring in ecological conditions found mainly in ravines. This result emphasizes the importance of topography in the existence of glacial refugia in the wider area. Furthermore, haplotypes originating from two more widespread beech lineages in Greece are found on Mt. Paggeo, indicating a possible mixing of populations originating from a local refugium with populations from remote refugia that possibly migrated into the area after the last glaciation. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 516–528.     

金沙江中下游圆口铜鱼遗传多样性与种群历史动态分析

以线粒体Cytb和CoⅠ基因作为分子标记,对金沙江中下游江段的圆口铜鱼进行遗传多样性和种群历史动态分析.结果显示,393尾圆口铜鱼样本共检测出91个串联基因序列单倍型,呈现较高的单倍型多样性(0.936±0.006)和较低的核苷酸多样性(0.00489±0.00009);基于单倍型构建的分子系统发育树及Median-j...     

Global mtDNA genetic structure and hypothesized invasion history of a major pest of citrus,Diaphorina citri (Hemiptera: Liviidae)

The Asian citrus psyllid Diaphorina citri Kuwayama is a key pest of citrus as the vector of the bacterium causing the “huanglongbing” disease (HLB). To assess the global mtDNA population genetic structure, and possible dispersal history of the pest, we investigated genetic variation at the COI gene collating newly collected samples with all previously published data. Our dataset consists of 356 colonies from 106 geographic sites worldwide. High haplotype diversity (H‐mean = 0.702 ± 0.017), low nucleotide diversity (π‐mean = 0.003), and significant positive selection (Ka/Ks = 32.92) were observed. Forty‐four haplotypes (Hap) were identified, clustered into two matrilines: Both occur in southeastern and southern Asia, North and South America, and Africa; lineages A and B also occur in eastern and western Asia, respectively. The most abundant haplotypes were Hap4 in lineage A (35.67%), and Hap9 in lineage B (41.29%). The haplotype network identified them as the ancestral haplotypes within their respective lineages. Analysis of molecular variance showed significant genetic structure (F_ST = 0.62, p < .0001) between the lineages, and population genetic analysis suggests geographic structuring. We hypothesize a southern and/or southeastern Asia origin, three dispersal routes, and parallel expansions of two lineages. The hypothesized first route involved the expansion of lineage B from southern Asia into North America via West Asia. The second, the expansion of some lineage A individuals from Southeast Asia into East Asia, and the third involved both lineages from Southeast Asia spreading westward into Africa and subsequently into South America. To test these hypotheses and gain a deeper understanding of the global history of D. citri, more data‐rich approaches will be necessary from the ample toolkit of next‐generation sequencing (NGS). However, this study may serve to guide such sampling and in the development of biological control programs against the global pest D. citri.     

Matrilineal genetic structure within and among populations of the cooperatively breeding common marmoset,Callithrix jacchus

Common marmosets are members of the family Callitrichidae, South American primates characterized by highly social group living and cooperative breeding. In this study we analysed 1112 base pairs (bp) of the mitochondrial control region in 59 Callithrix jacchus individuals, sampled mainly from two geographically distinct field sites in N.E. Brazil. Analysis of molecular variation revealed a highly significant genetic structuring of haplotypes between social groups and between populations. Examination of matrilineal genetic structure within social groups revealed that seven of nine recorded breeding pairs were from different maternal lineages, indicating assortative mating and outbreeding. In addition to the breeders, at least six of 10 groups contained adult individuals from different matrilines, with five haplotypes present in one social group of nine animals. Groups of mixed lineages raise questions about potential reproductive conflicts of interest, and the extent of kin-selected altruism in the evolution and maintenance of cooperative breeding in this species.     

Contrasting patterns of historical colonization in white oaks (Quercus spp.) in California and Europe

Phylogeography allows the inference of evolutionary processes that have shaped the current distribution of genealogical lineages across a landscape. In this perspective, comparative phylogeographical analyses are useful in detecting common historical patterns by either comparing different species within the same area within a continent or by comparing similar species in different areas. Here, we analyse one taxon (the white oak, genus Quercus, subgenus Quercus, section Quercus) that is widespread worldwide, and we evaluate its phylogeographical pattern on two different continents: western North America and Western Europe. The goals of the present study are: (i) to compare the chloroplast genetic diversity found in one California oak species vs. that found in the extensively studied European oak species (in France and the Iberian Peninsula); (ii) to contrast the geographical structure of haplotypes between these two taxa and test for a phylogeographical structure for the California species. For this purpose, we used the same six maternally inherited chloroplast microsatellite markers and a similar sampling strategy. The haplotype diversity within site as well as the differentiation among sites was alike in both taxa, but the Californian species has higher allelic richness with a greater number of haplotypes (39 vs. 11 in the European white oak complex). Furthermore, in California these 39 haplotypes are distributed locally in patches while in the European oaks haplotypes are distributed into lineages partitioned longitudinally. These contrasted patterns could indicate that gene movement in California oak populations have been more stable in response to past climatic and geological events, in contrast to their European counterparts.     

Genetic population structure in the yellow mongoose, Cynictis penicillata

Phylogeographic structure was determined for the yellow mongoose, Cynictis penicillata , using mtDNA RFLPs and control region sequences. The RFLP analysis revealed 13 haplotypes which showed weak geographical patterning consistent with a recent range expansion from a refugial population(s). An analysis of molecular variance ( AMOVA ) revealed no correspondence between mtDNA phylogeography and subspecies delimitation, nor between matrilines and areas characterized by a high incidence of the viverrid-type rabies, of which the yellow mongoose is the principal vector. The lack of structure was also shown by control region sequences although four of the maternal lineages shared a near-perfect 81 bp repeat. We speculate that regional hot spots of the viverrid rabies biotype reflect population density differences in the yellow mongoose that are not underscored by genetic partitioning, at least at the level of resolution provided by our analyses.     

Phylogeographical approaches to assessing demographic connectivity between breeding and overwintering regions in a Nearctic-Neotropical warbler (Wilsonia pusilla)

We characterized the pattern and magnitude of phylogeographical variation among breeding populations of a long-distance migratory bird, the Wilson's warbler (Wilsonia pusilla), and used this information to assess the utility of mtDNA markers for assaying demographic connectivity between breeding and overwintering regions. We found a complex pattern of population differentiation in mitochondrial DNA (mtDNA) variation among populations across the breeding range. Individuals from eastern North America were differentiated from western individuals and the eastern haplotypes formed a distinct, well-supported cluster. The more diverse western group contained haplotype clusters with significant geographical structuring, but there was also broad mixing of haplotype groups such that no haplotype groups were population specific and the predominance of rare haplotypes limited the utility of frequency-based assignment techniques. Nonetheless, the existence of geographically diagnosable eastern vs. western haplotypes enabled us to characterize the distribution of these two groups across 14 overwintering locations. Western haplotypes were present at much higher frequencies than eastern haplotypes at most overwintering sites. Application of this mtDNA-based method of linking breeding and overwintering populations on a finer geographical scale was precluded by the absence of population-specific markers and by insufficient haplotype sorting among western breeding populations. Our results suggest that because migratory species such as the Wilson's warbler likely experienced extensive gene flow among regional breeding populations, molecular markers will have the greatest utility for characterizing breeding-overwintering connectivity at a broad geographical scale.     

Extreme genetic mixing within colonies of the wood-dwelling termite Kalotermes flavicollis (Isoptera,Kalotermitidae)

The existence of altruism in social insects is commonly attributed to altruistic individuals gaining indirect fitness through kin selection. However, recent studies suggest that such individuals might also gain direct fitness through reproduction. Experimental studies on primitive wood-dwelling termites revealed that colony fusion often causes the death of primary reproductives (queen and king), allowing opportunities for workers to inherit the nest by developing into replacement reproductives (neotenics). Therefore, colony fusion has been proposed as an important factor that may have favoured sociality in termites. However, whether colony fusion occurs frequently in natural populations of wood-dwelling termites remains an open question. We analysed eleven colonies of the wood-dwelling termite Kalotermes flavicollis (Kalotermitidae), using two mitochondrial and five nuclear microsatellite markers. Nine of eleven colonies (82%) were mixed families, with offspring of three or more primary reproductives. To our knowledge, this result represents the highest frequency of mixed-family colonies ever reported in termites. Moreover, genetic mixing of colonies appeared extreme in two ways. First, the number of haplotypes per colony was exceptionally high (up to nine), indicating that colonies were composed of multiple queens' offspring. Second, some mixed-family colonies included individuals belonging to two highly divergent genetic lineages. F-statistics and relatedness values suggest that mixed-family colonies most likely result from colony fusion, giving support to the accelerated nest inheritance theory. These findings raise important questions about the mode of foundation of mixed-family colonies and the evolutionary forces that maintain them within populations.     

Intraspecific differentiation of Allium wallichii (Amaryllidaceae) inferred from chloroplast DNA and internal transcribed spacer fragments

In the present study, we used two chloroplast DNA (cpDNA) fragments (trnL-F and rps16) and the nuclear ribosomal internal transcribed spacer (ITS) sequence data to examine intraspecific differentiation and phylogeographical history of Allium wallichii. Based on wide scale sampling (28 populations and 174 individuals) across the entire distribution range of this species, 33 cpDNA haplotypes and 25 ITS ribotypes were detected in our investigation. These cpDNA haplotypes were divided into three major lineages, which was further supported by the ITS phylogenetic results. High haplotype/ribotype diversity and population differentiation, together with most of the haplotypes/ribotypes being exclusive to single populations, implied restricted gene flow among populations and significant geographical isolation. Nearly all populations with high haplotype/ribotype diversity were found in the Hengduan Mountain Region (HMR), whereas the populations of the Himalayas and Nanling Mountains showed a lower level, suggesting the HMR might serve as a potential divergence center for A. wallichii. The main lineages of A. wallichii diverged from each other between Mid–Late Pliocene and Early Quaternary based on two sets of molecular markers, indicating that the Quaternary climatic fluctuation could not have contributed greatly to the divergence of the main lineages of A. wallichii. Instead, the intricate topography and heterogeneous habitats resulting from the drastic uplift of the Qinghai–Tibet Plateau from the Late Pliocene could be responsible for the intraspecific differentiation of A. wallichii. The present study further highlights the importance of geographic isolation and habitat heterogeneity in shaping and maintaining high species diversity within the HMR.     

Fine‐scale impacts on avian biodiversity due to a despotic species,the bell miner (Manorina melanophrys)

Bell miners (Manorina melanophrys; Meliphagidae) are a highly social and very aggressive honeyeater. They are despotic and cooperate in the defence of their territories against other bird species, leading to the almost complete exclusion of other avifauna from miner‐occupied regions. This study aimed to resolve some of the fine‐scale effects of bell miner aggression on avian diversity both within and adjacent to colonies to determine the true impact of a colony on local avifaunal abundance. Three areas, distributed throughout the range of the bell miner, were surveyed across both non‐breeding and breeding seasons to assess the temporal and spatial impacts of bell miner aggression on other bird species. Bell miner colonies were found to occupy very clearly defined areas and had the expected negative impact on avian diversity within their colony. The effects of bell miner colony presence on abundance and richness of avian species were found to cease at the colony boundary, with both recovering to normal levels immediately outside the bell miner colony. Whether bell miners were breeding or not, and irrespective of the amount of vegetation coverage, bell miner colonies were found to have relatively marginal impacts on avian richness and abundance. No impact of colony presence/absence was found on the richness or abundance of the avian species that dwell in the undergrowth, with some evidence that these species were actually more common at the colony edge. Our results demonstrate that the influence of bell miner colony presence upon avian biodiversity is restricted to the confines of the colony and does not radiate outwards into the surrounding habitat. Colony presence influences, therefore, have implications when considering the impact of bell miner behaviour on the diversity of insectivorous birds and processes, most notably the propagation of Bell Miner Associated Dieback.