Microsatellite loci for Myrmica kotokui and their application in some congeneric ant species distributed in northern Japan

We developed seven polymorphic microsatellite loci for Myrmica kotokui from RAPD (rapid analysis of polymorphic DNA) fragments. These loci showed two to six alleles with expected and observed heterozygosity of 0.13–0.73 and 0.14–0.78, respectively. These loci are transferable to the congeneric species as Myrmica rubra, Myrmica luteola and Myrmica taediosa from northern Japan. These loci will allow analyses of genetic structure of Myrmica species at both the colony level and population levels.     

Social factors affecting queen fecundity in the ant Myrmica rubra

ABSTRACT. At the end of winter, normal populations of the polygyne ant Myrmica rubra L. consist of queens, workers and larvae; the latter are dormant and all in the third stadium. During experimental culture in artificial nests in the laboratory, the starting number of all three of these categories was varied and in some, food supply was restricted too. In a population of workers and larvae, queens do not interfere with each other sufficiently to reduce their fecundity; they avoid congestion by dispersing under the canopy of workers. Queens suffer if food is withheld; they probably feed themselves and are thus largely independent of the number of workers. Even in favourable conditions with full food, the distribution of egg-batch sizes is not normal but follows a hollow curve that is well described by the negative binomial series. A poor food supply accentuates the hollow shape by increasing the number of non-laying queens and reducing the number that are able to lay large batches. Larvae also suppress fecundity but only if they have sufficient food to grow actively; as their number increases, egg production by queens decreases exponentially.     

Some characteristics of the visual perception of the ant Myrmica sabuleti

Abstract. Using classical conditioning techniques, this study aims to determine what the ant Myrmica sabuleti can see and discriminate. It appears that workers see distinctly vertical as well as horizontal segments 1, 2, 3, 5, 10 and 15 mm wide and 1, 2, 3, 5, 10 and 15 cm long. They can also see distinctly horizontal segments (0.5 × 3 cm) located at 1, 2, 3, 5 and 10 cm height, but do not see such a segment situated at 15 cm height. This allowed the assessment of their maximum distance of vision. Myrmica sabuleti workers also see distinctly vertical segments (0.5 × 3 cm) sloping backwards towards the horizontal at different angles (90, 60, 45, 30, 15 and 0 °). Consequently, M. sabuleti workers see cues lying horizontally on the ground and have a stereovision of a portion of their environment. The species also see distinctly black circles of different sizes located horizontally above them. Thus, they are able to see distinctly slightly modified patterns located aside, below, above as well as before them, and have a small area of binocular vision that both eyes can look at. Myrmica sabuleti workers' maximum distance of vision is directly proportional to the square root of the surface of the object observed. Consequently, the smallest vision angular subtense of an object needed to elicit a reaction in M. sabuleti workers is evaluated as being approximately 5 angular degrees.     

The importance of daylength and queens for larval care by young workers of the ant Myrmica rubra L.

ABSTRACT. Workers whilst still less than a month from emergence rear early stage larvae better in a 16h day than in one of 14h. The callows must perceive the diel rhythm directly because there are no foragers to give them a clue, and the larvae are kept in the dark and cannot respond directly. This, and the study of egg-laying by young workers that preceded it, indicates that the extra hours daylight enable the workers to avoid the reproductive diapause that normally prepares them for hibernation. They retain active ovaries, ingest and share food with larvae either as trophic eggs or as a regurgitate from the crop.
In late summer, callows can rear sexual larvae in spring condition. Queens have a small negative effect on this, expressed as a smaller, though still gyne, pupa produced a little earlier. This weak response to queens develops into a strong negative one at a constant temperature of either 10 or 20°C. The workers then behave as they normally do in spring, and feed a watery regurgitate (dietary control) or, if this fails to arrest sexual development, physically harass the larvae by biting them in the ventral thorax (manipulative control). Large workers then result instead of gynes.     

Roles of species-preferential seed dispersal by ants and endozoochory in Melampyrum (Orobanchaceae)

Aims Melampyrum pratense and M. subalpinum are two myrmecochorous species, which possess similar habitat requirements and frequently occur together. Despite this, their population sizes differ markedly. Melampyrum pratense populations are usually very large, whereas M. subalpinum has rather small and isolated populations. We suggest that such an imbalance might be partially influenced by the difference in ant-mediated seed-removal rates. Genus Melampyrum is considered to be exclusively myrmecochorous, though to achieve the recent distribution of some Melampyrum species during the Holocene myrmecochory would be highly insufficient. We suggest that endozoochory takes place in the long-distance migration, whereas myrmecochory is important for the removal of seeds on a local scale.Methods For seed-preference analysis, M. pratense and M. subalpinum mixed seed samples were placed around Formica polyctena anthills. After a period of time, the remaining seeds of both species were counted for each sample. The results were analysed by analysis of variance and generalized linear mixed-effect model. To test myrmecochorous removal distances, M. pratense seeds were covered with fluorescent dactyloscopic powder and placed in the vicinity of a large ant trail. The area around the starting plot was searched in the dark using UV LED torchlight 7h after the beginning. The distance from the starting plot was measured for each seed found. Birds, rodents, leporine and a ruminant were fed with M. pratense seeds and fresh plants to test the possibility of endozoochorous dispersal of the species. Animal droppings were searched for intact seeds.Important findings Our field studies show that from mixed seed samples, containing both species, ants significantly preferred the seeds of M. pratense. This may be one of factors that has positive influence on M. pratense success in seed dispersal on mixed stands and consequently in the colonization of favourable sites. Experiments focusing on ant-mediated dispersal distance revealed that F. polyctena ants are able to move seeds over a distance of 36 m in only 7h. This distance is among the furthest known myrmecochorous removals of forest plant seeds. A new Melampyrum seed disperser Oligolophus tridens (Opiliones) was observed repeatedly. Our pilot study documented that Melampyrum seeds are able to pass through the digestive tract of a cow intact. This suggests that large ruminants such as deer, bison or forest-grazing livestock may function as important long-distance dispersers of Melampyrum species.     

Effects of ant behaviour and presence of extrafloral nectaries on seed dispersal of the Neotropical myrmecochore Turnera ulmifolia L. (Turneraceae)

This paper describes the myrmecochory system of Turnera ulmifolia in a coastal sand dune matorral in Mexico. Turnera ulmifolia has elaiosome‐bearing seeds and extrafloral nectaries (EFNs). In ten quadrants (4 × 15 m) ant–seed interaction was monitored, and an interaction intensity index calculated and correlated with the number of seedlings. Seed removal rates by ants were surveyed every 2 h for 24 h, the ants being observed both on and beneath the plants. The role of the elaiosome in seed removal was evaluated by offering seeds with and without elaiosomes, and elaiosomes only. Finally, the effect of ant manipulation in seed germination was evaluated. There were 25 ant species associated with seeds and/or EFNs, the most frequently recorded being Monomorium cyaneum and Forelius analis. There was a positive correlation between the intensity index and seedling number per quadrant. There was significantly higher mean seed removal during the day than during the night (19.3% and 12.3%, respectively), and from beneath than on the plant (21.9% and 9.5%, respectively). The preference for elaiosomes only was also greater during the diurnal period, and when gathered on, rather than beneath, the plant. Seed manipulation by F. analis enhanced germination by T. ulmifolia. Seed removal, dispersal distances, seed predation and germination were largely determined by ant behaviour. The presence of EFNs may be influencing seed removal on the plant by attracting a specific assemblage of omnivorous ants. Among such assemblages associated with T. ulmifolia we encountered a variety of behaviours, with ant species either good at defending plants but bad at dispersing seeds, or vice versa. We discuss the way in which these two rewards, and the processes involved (defence and dispersion), could have interacted with each other and evolved. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 67–77.     

油质体在5种蚁播植物种子散布中的作用

蚁播植物种子上常附着有蚂蚁喜食的油质体,该物体可吸引蚂蚁取食或为蚂蚁提供钳着位点从而影响种子散布。为进一步揭示油质体在种子散布中的作用,在野外研究了4属5种典型蚁播植物包括小花宽瓣黄堇(Corydalis giraldii Fedde)、假刻叶紫堇(C.pseudoincisa C.Y.Wu)、白屈菜(Chelidonium majus L.)、紫花堇菜(Viola grypoceras A.Gray)和柔毛淫羊藿(Epimedium pubescens Maxim.)其相应的搬运蚂蚁即玉米毛蚁(Lasius alienus(Foerster))和尼特纳大头蚁(Pheidole nietneri Emery)对植物完整种子(SE)、去除油质体的种子(S)和粘有人工模拟油质体的种子(S+H)的搬运行为和搬运效率。结果显示,蚂蚁对相同植物3类种子的触碰和检查次数均无显著差异。蚂蚁对种子的尝试搬运次数除玉米毛蚁对小花宽瓣黄堇、假刻叶紫堇和白屈菜去除油质体的种子(S)显著高于完整种子(SE)和粘有人工模拟油质体的种子(S+H)外,其它的均无显著差异。玉米毛蚁对小花宽瓣黄堇、假刻叶紫堇、白屈菜和紫花堇菜以及尼特纳大头蚁对小花宽瓣黄堇、假刻叶紫堇和紫花堇菜完整种子(SE)的搬运效率显著高于粘有人工模拟油质体的种子(S+H)和去除油质体的种子(S),此外,玉米毛蚁对小花宽瓣黄堇粘有人工模拟油质体的种子(S+H)的搬运效率显著高于去除油质体的种子(S)。这说明玉米毛蚁在搬运小花宽瓣黄堇种子过程中油质体不仅起到吸引作用,而且起到钳着位点作用;玉米毛蚁在搬运假刻叶紫堇、白屈菜和紫花堇菜以及尼特纳大头蚁搬运小花宽瓣黄堇、假刻叶紫堇和紫花堇菜种子过程中油质体仅起到吸引作用。研究表明油质体可通过对蚂蚁起吸引作用,或同时起吸引和钳着位点的双重作用等影响种子散布,油质体的影响作用不但取决于植物种类,也与搬运蚂蚁的种类有关。     

Queen recognition by brood-rearing workers of the ant Myrmica rubra L

The means by which queens communicate their presence to workers and so influence broodrearing, in particular caste determination, have been analysed; both chemical and topographical stimuli are used. This follows from the fact that though dead inseminated queens are effective, the following are not: dead virgins (same shape but different smell), lipid extracted mature queens (same shape but no smell), parts of inseminated queens alone or together (same smell but different shape). This also rules out a drug-like influence on workers or larvae. The chemicals are not produced in the two main head glands, the mandibular and the pharyngeal, as no loss of potency follows their excision. The abdomen is the most likely source of a substance, the emission of which must vary seasonally.     

The control of Myrmica rubra workers by queens of their own and other Myrmica species and the interaction between queenless groups of workers

ABSTRACT. Queens of two species of the ant genus Myrmica bonded to workers of the species M. rubra L. as the latter emerge from the pupal skin can use these workers nearly 6 months later to arrest gyne formation in sex-competent larvae of the same species. Queens of M. ruginodis Nylander var. microgyria (Brian & Brian, 1949) are as good at this as the natural M. rubra , but those of M. sabuleti Meinert (of a race close to M. scabrinodis) are not. Though the M. sabuleti queens induce normal aggression against sexualizing larvae, they are unable to prevent some or all of the workers feeding larvae as though they were queenless. However, queens from different colonies of M. rubra adopted by queenless populations of workers in spring, control their brood-rearing behaviour perfectly. M. rubra workers from different colonies bring gynes to maturity from female sexual larvae at different average sizes. When workers from two such sources are mixed in equal proportions, the size of gyne larva produced after a week's culture corresponds with that of one of the worker populations; it is not intermediate in size. Also, large workers can rear larger gyne-larvae than small workers of the same age. This is only true if the workers have been living with queens all the time from emergence as an imago to the moment the experiment was set. Size mixtures only achieve the same size larvae as a pure culture of small workers would. A possible reason for this is that small workers exclude the larger ones from the nursery areas of the nest.     

First report of seed dispersal by ants in Dicentra peregrina (Papaveraceae), an alpine plant in the Japanese Alps

Seed dispersal by ants, known as myrmecochory, is commonly observed among various plant taxa. The seeds of these plants have an elaiosome to attract ants. In Japan, myrmecochory has been well studied in several lowland plant species, but not in highland plant species that grow above the tree line. We investigated whether the seeds of Dicentra peregrina, known as the “queen of the alpine plants” in Japan, are carried by Formica gagatoides ants at 2510 m a.s.l. on Mt Norikura, Kita‐Alps, Japan. We observed F. gagatoides workers picking up D. peregrina seeds by grasping the elaiosome and carrying the seeds into their nests. We inferred from the observed ant behavior and the seed morphology that D. peregrina is a myrmecochorous species.     

Resource allocation in the red ant Myrmica ruginodis– an interplay of genetics and ecology

Worker‐queen conflicts over reproductive allocation (colony maintenance vs. reproduction) and sex allocation (females vs. males) were examined in two populations of the facultatively polygynous ant Myrmica ruginodis. Plasticity of social organization in the form of two co‐existing social types (microgyna and macrogyna) has a profound effect on reproductive allocation. Workers control sex allocation by biasing sex ratios towards their own interest, but local resource competition (LRC) because of restricted dispersal of microgyna females resulted in male bias in one study population. Colony sex ratios were split and followed the predictions of the split sex ratio theory: single queen colonies with higher relatedness asymmetry (RA) produced more females than multiple queen colonies with lower RA. Single and multiple queen colonies showed similar patterns in most aspects of their reproduction, and reproductive allocation could not be explained by the hypothesis tested. This suggests that reproductive allocation conflict is of minor importance in M. ruginodis.     

Isolation and characterization of microsatellite loci in the ant Myrmica scabrinodis

Five polymorphic microsatellite loci were developed for the ant Myrmica scabrinodis using a magnetic bead hybridization selection protocol. The number of alleles per locus varied between three and six. Cross‐species amplification of four of the loci yielded positive amplification products in four Myrmica species, suggesting their general suitability for microsatellite analysis within this taxonomic group.     

Behavioural activity of pure chiral 3-octanol for the ants Myrmica scabrinodis Nyl. and Myrmica rubra L.

ABSTRACT. The mandibular glands of Myrmica ants contain, among other substances, 3-octanol, a chiral substance, 90% of it being the R -enantiomer in M. scabrinodis Nyl., ruhra L. and ruginodis Nyl. (Cammaerts et al. , 1985). Pure R and S 3-octanol has been prepared and tested on M. rubra and scabrinodis , workers. Both species react specifically only to the R enantiomer; in M. rubra this constituent arrests foragers briefly, and in M. scahrinodis it attracts them and incites them to walk more quickly. It is shown that, in M. scahrinodis , the naturally produced mixture of R and S 3-octanol (proportions of 9:1, v/v) is more active than the pure R enantiomer or a mixture of R and S 3-octanol (5:5 v/v).     

Queen selection by worker groups of the ant Myrmica rubra L.

Normal queens in a colony of Myrmica rubra L. were not necessarily more attractive to the workers than strange ones grafted into the colony. Workers chose the best queens using those they already had as a standard. After winter dormancy, workers in colonies with queens were less tolerant for queen variety. One week at 20°C was enough to cause them to reject still-dormant queens, even those of their own colony, though they accepted alien queens. However, after 2–3 weeks culture queens were acceptable only if they belonged to the same colony as the natural ones or to the same colony as those queens that had been artificially implanted. To re-open the group, it was essential to remove all the resident queens; then after 2 days the workers took any queen offered, rather than remain queenless, but when offered a choice between one of their originals and an unfamiliar one, they still took the former. Three queenless weeks were required before they lost their preference for an ex-nestmate. Queens from a single colony, isolated in separate groups of workers, remained acceptable to the workers with them indefinitely, but when these queens were transferred to identical collateral groups of workers, they were rejected even after only 2 weeks' separation. Some groups of workers were less tolerant than others and some queens were more attractive than others, but their compatibility diminished with time if they lived in isolation. A group with an attractive queen was more likely to resist other queens.     

Phylogeography of the ant Myrmica rubra and its inquiline social parasite

Widely distributed Palearctic insects are ideal to study phylogeographic patterns owing to their high potential to survive in many Pleistocene refugia and-after the glaciation-to recolonize vast, continuous areas. Nevertheless, such species have received little phylogeographic attention. Here, we investigated the Pleistocene refugia and subsequent postglacial colonization of the common, abundant, and widely distributed ant Myrmica rubra over most of its Palearctic area, using mitochondrial DNA (mtDNA). The western and eastern populations of M. rubra belonged predominantly to separate haplogroups, which formed a broad secondary contact zone in Central Europe. The distribution of genetic diversity and haplogroups implied that M. rubra survived the last glaciation in multiple refugia located over an extensive area from Iberia in the west to Siberia in the east, and colonized its present areas of distribution along several routes. The matrilineal genetic structure of M. rubra was probably formed during the last glaciation and subsequent postglacial expansion. Additionally, because M. rubra has two queen morphs, the obligately socially parasitic microgyne and its macrogyne host, we tested the suggested speciation of the parasite. Locally, the parasite and host usually belonged to the same haplogroup but differed in haplotype frequencies. This indicates that genetic differentiation between the morphs is a universal pattern and thus incipient, sympatric speciation of the parasite from its host is possible. If speciation is taking place, however, it is not yet visible as lineage sorting of the mtDNA between the morphs.     

Intraspecific aggression and the colony structure of the invasive ant Myrmica rubra

1. Patterns of aggression between ants from different nests influence colony and population structure. Several species of invasive ants lack colony boundaries over large expanses, forming ‘supercolonies’ with many nests among which workers can move without encountering aggression. 2. Bioassays of aggression were used to determine the colony structure of the invasive ant Myrmica rubra (L.) at eight sites in Massachusetts, the state where the species was first discovered in North America. To improve the ability to distinguish systematic patterns from background variability in aggressiveness, a repeated‐measures design was used and replicate assays for each pair of nests were conducted. 3. Aggressive responses showed that populations at all sites consisted of multiple distinct colonies. Patterns of aggression were repeatable and transitive, with few exceptions. Colonies were identified as clusters of nests whose workers showed little to no aggression towards one another but were aggressive towards conspecifics from more distant nests. 4. The degree of aggression varied considerably among different colony pairs but did not depend in any consistent way on the distance of separation or on whether colonies were neighbours. 5. Territories of neighbouring colonies abutted, indicating that they were restricted by intraspecific competition. Mapped territories ranged in size from 0.03 to 1.2 ha, but colonies at the study sites have not undergone the enormous expansions seen in introduced populations of some other species of invasive ants, and neighbouring colonies compete locally.     

No sympatric speciation here: multiple data sources show that the ant Myrmica microrubra is not a separate species but an alternate reproductive morph of Myrmica rubra

No aspect of speciation is as controversial as the view that new species can evolve sympatrically, among populations in close physical contact. Social parasitism has been suggested to yield necessary disruptive selection for sympatric speciation. Recently, mitochondrial DNA phylogeography has shown that the ant Myrmica microrubra is closely related to its host, Myrmica rubra, leading to the suggestion that sympatric speciation has occurred. We investigated the relationships between the two ant forms using mitochondrial and nuclear DNA sequences, microsatellite genotyping and morphometrics. Molecular phylogenetic and population structure analyses showed that M. microrubra does not evolve separately to its host but rather shares a gene pool with it. Probability analysis showed that mitochondrial DNA data previously adduced in favour of sympatric speciation do not in fact do so. Morphometrically, M. microrubra is most readily interpreted as a miniature queen form of M. rubra, not a separate species. Myrmica microrubra is not an example of speciation. The large (typical M. rubra) and small (M. microrubra) queen forms are alternative reproductive strategies of the same species. Myrmica microrubraSeifert 1993 is consequently synonymized here with M. rubra Linnaeus, 1758.     

New fungal pathogens of the red ant, Myrmica rubra, from the UK and implications for ant invasions in the USA

The red ant, Myrmica rubra, is an increasingly invasive pest species in north-eastern USA, where it is known as the European fire ant. During surveys for natural enemies in part of its native range in the UK, three previously unreported fungal pathogens developed on ants when incubated in the laboratory. These are described and illustrated: Paraisaria myrmicarum sp. nov., Hirsutella stilbelliformis var. myrmicarum var. nov., and Hirsutella subramanianii var. myrmicarum var. nov. Based on analyses of the protein coding region EF-1α and LSU?rDNA, all three described taxa are shown to be affiliated with the hypocrealean family?Ophiocordycipitaceae. The implications for the management of M. rubra in its exotic North American range using classical biological control are discussed.     

Seed handling behaviours of native and invasive seed‐dispersing ants differentially influence seedling emergence in an introduced plant

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