Intercolony aggression within and among local populations of the invasive ant, Myrmica rubra (Hymenoptera: Formicidae), in coastal Maine

Myrmica rubra L. was introduced into New England in the early 20th century, and at present, has a patchy distribution in parts of northeastern North America, including records from 31 communities in Maine. M. rubra is highly polygynous, and colonies reproduce vegetatively, forming dense local populations where conditions are favorable. Using mobile nests and baited arenas in a series of field aggression bioassays, we tested patterns of internest tolerance within and among local populations on Mt. Desert Island, ME. We found that foragers originating from fragments of the same colony or from neighboring nests retained a high level of intraspecific tolerance over several months, whereas significant intercolony aggression among workers was present between colonies within the same local patch separated by approximately 10 m. Within populations, aggression score values were found to increase linearly with internest distance within a site. Aggression was highest between colonies from spatially different populations on the island and was higher still when nests were assayed against colonies at an off-island site 70 km away in Castine, ME. These data strongly suggest a multicolonial organization within and among local populations of M. rubra in parts of its introduced range. These findings contradict the loss of intraspecific aggression and unicolonial social structure over large geographic areas that have previously been observed in other invasive ant species, particularly Linepithema humile Mayr.     

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.     

Population genetics of geographically restricted and widespread species of Myrica (Myricaceae)

Allozyme variation of 11 putative loci in five populations of the rare Myrica adenophora Hance, and four populations of its widespread congeneric species, M. rubra (Lour.) Sieb. & Zucc. was studied. Among the 21 alleles studied, no unique allele was detected for M. adenophora, whereas M. rubra had 3 alleles not found in the former species. In terms of genetic diversity, populations of the rare species contained fewer alleles per locus (1.5 versus 1.7), fewer effective number of alleles per locus (1.12 versus 1.20), fewer number of alleles per polymorphic locus (2.14 versus 2.46), lower percentage of polymorphic loci (30.9 versus 40.9), and lower expected heterozygosity (0.106 versus 0.163) than populations of the widespread species. Genetic distances within species average 0.043 for M. adenophora and 0.045 for M. rubra, and between species ranged from 0.052 to 0.177, with a mean of 0.103, which agrees with the very similar gross morphologies of these two species. Intrapopulation differentiation was similar in both species: G(ST) = 0.152 for M. adenophora, and 0.146 for M. rubra, whereas estimated gene flow based on G(ST) values were moderate in these two species (Nm = 1.39 versus 1.46). We inferred that M. rubra and M. adenophora are a progenitor-derivative species pair that emerged before migrating into Taiwan during the last glacial period. We consider the Hengchun population (Chiupeng, Hsuhai, and Chufengpi) and Taitung population (Tienkuan and Lanshan) of M. adenophora which probably arose from two subsets of the genome of M. rubra. Genetic drift was inferred to be one of the forces shaping the observed genetic structure in M. adenophora and M. rubra.     

Expression of stress proteins of family HSP70 in response to cold in Myrmica ants from different geographic populations

Expression of HSP70 is induced by stress factors, including sub-lethal chilling, but the role of HSP70 for overcoming consequenses of cold stress is not clear. If the role is positive, the level of HSP70 expression might be higher in populations from colder climates. By use of immunoblotting we investigated dynamics of HSP70 expression in response to cold stress in two Myrmica species from three localities of different latitude (50, 60 and 67 degrees N). The results showed that in the more thermophilic species Myrmica rubra expression of HSP70 after cold stress was more intense. Expression traits within both species did not show direct relationship with latitude. In M. rubra the southernmost population and in M. ruginodis the northernmost one displayed the fastest and most intense response. However, in M. rubra the two other studied populations were similar in timing and intensity of the response, while in M. ruginodis the intermediate population showed the slowest and weakest response. The data suggest that the expression of HSP70 might play the most considerable role in the adaptation to cold in the southernmost population of M. ruginodis.     

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.     

How ants find each other; temporal and spatial patterns in nuptial flights

Reproduction is a key factor in understanding population ecology and therefore species occurrence. However, patterns in reproductive behaviour for distinct ant species remain insufficiently known. In this paper strategies in mate finding are studied for six ant species (Lasius niger, Lasius umbratus, Temnothorax nylanderi, Myrmica rubra, Myrmica ruginodis, Stenamma debile) in a forest – forest edge – agricultural field gradient. Using window traps, we studied whether these species had a restricted nuptial flight season, displayed swarming behaviour, and whether the alates aggregated at the forest edge. The flight season was limited to one month or less for L. niger, T. nylanderi, M. rubra, M. ruginodis and S. debile. Swarming behaviour occurred in all but one (L. umbratus) species. Although none of the six species seemed to have highest nest density at the forest edge, three of them, M. rubra, M. ruginodis and S. debile, showed male aggregations there, indicating this to be the main reproduction site. This last finding could be due to a more suitable micro-climate, but most likely, edges are conspicuous land marks which are used by ants to meet mates. The behavioural patterns of ant sexuals at the forest edge can influence dispersal possibilities in fragmented landscapes, reproductive success and nest densities. Received 27 November 2007; revised 27 March; accepted 5 April 2008.     

Geographical variation in host–ant specificity of the parasitic butterfly Maculinea alcon in Denmark

Abstract  1. Maculinea alcon uses three different species of Myrmica host ants along a north–south gradient in Europe. Based on this geographical variation in host ant use, Elmes et al . (1994) suggested that M. alcon might consist of three or more cryptic species or host races, each using a single and different host-ant species.
2. Population-specific differences in allozyme genotypes of M. alcon in Denmark ( Gadeberg & Boomsma, 1997 ) have suggested that genetically differentiated forms may occur in a gradient across Denmark, possibly in relation to the use of different host ants.
3. It was found that two host-ant species are indeed used as hosts in Denmark, but not in a clear-cut north–south gradient. Furthermore, specificity was not complete for many M. alcon populations. Of five populations investigated in detail, one used primarily M. rubra as a host, another exclusively used M. ruginodis , while the other three populations used both ant species. No population in Denmark used M. scabrinodis as a host, although this species was present in the habitat and is known to be a host in central and southern Europe.
4. In terms of number of parasites per nest and number of nests parasitised, M. rubra seems to be a more suitable host in populations where two host species are used simultaneously. Host-ant species has an influence on caterpillar size but this varies geographically. Analyses of pupae did not, however, show size differences between M. alcon raised in M. rubra and M. ruginodis nests.
5. The geographical mosaic of host specificity and demography of M. alcon in Denmark probably reflects the co-evolution of M. alcon with two alternative host species. This system therefore provides an interesting opportunity for studying details of the evolution of parasite specificity and the dynamics of host-race formation.     

Testing the adjustable threshold model for intruder recognition on Myrmica ants in the context of a social parasite

Social insect colonies are like fortresses, well protected and rich in shared stored resources. This makes them ideal targets for exploitation by predators, parasites and competitors. Colonies of Myrmica rubra ants are sometimes exploited by the parasitic butterfly Maculinea alcon. Maculinea alcon gains access to the ants' nests by mimicking their cuticular hydrocarbon recognition cues, which allows the parasites to blend in with their host ants. Myrmica rubra may be particularly susceptible to exploitation in this fashion as it has large, polydomous colonies with many queens and a very viscous population structure. We studied the mutual aggressive behaviour of My. rubra colonies based on predictions for recognition effectiveness. Three hypotheses were tested: first, that aggression increases with distance (geographical, genetic and chemical); second, that the more queens present in a colony and therefore the less-related workers within a colony, the less aggressively they will behave; and that colonies facing parasitism will be more aggressive than colonies experiencing less parasite pressure. Our results confirm all these predictions, supporting flexible aggression behaviour in Myrmica ants depending on context.     

Highly divergent SSU rRNA genes found in the marine ciliates Myrionecta rubra and Mesodinium pulex

Myrionecta rubra and Mesodinium pulex are among the most commonly encountered planktonic ciliates in coastal marine and estuarine regions throughout the world. Despite their widespread distribution, both ciliates have received little attention by taxonomists. In order to better understand the phylogenetic position of these ciliates, we determined the SSU rRNA gene sequence from cultures of M. rubra and M. pulex. Partial sequence data were also generated from isolated cells of M. rubra from Chesapeake Bay. The M. rubra and M. pulex sequences were very divergent from all other ciliates, but shared a branch with 100% bootstrap support. Both species had numerous deletions and substitutions in their SSU rRNA gene, resulting in a long branch for the clade. This made the sequences prone to spurious phylogenetic affiliations when using simple phylogenetic methods. Maximum likelihood analysis placed M. rubra and M. pulex on the basal ciliate branch, following removal of ambiguously aligned regions. Fluorescent in situ hybridization probes were used with confocal laser scanning microscopy to confirm that these divergent sequences were both expressed in the cytoplasm and nucleolus of M. ruisra and M. pulex. We found that our sequence data matched several recently discovered unidentified eukaryotes in Genbank from diverse marine habitats, all of which had apparently been misattributed to highly divergent amoeboid organisms.     

褪黑素对模拟干旱胁迫下北美红栎幼树幼树光合性能及抗氧化酶系统的影响

为研究干旱胁迫下施加褪黑素对北美红栎幼树光合性能和抗氧化酶系统的影响,以当年生北美红栎幼树实生苗为试验材料,利用不同浓度的聚乙二醇(PEG-6000)溶液模拟干旱,研究干旱胁迫对北美红栎幼树光合性能及抗氧化酶活性的影响。结果表明:在PEG模拟的干旱胁迫下,北美红栎幼树的光合作用受到抑制并破坏了体内的氧化还原平衡,抗氧化物酶活性提高。在干旱胁迫开始前喷施100μM的褪黑素能够提高北美红栎幼树清除活性氧的能力,维持干旱胁迫条件下较高的净光合速率并能有效地缓解由于干旱胁迫对植物造成的损伤。试验结果为实践中栽培管理北美红栎幼树提供了参考。     

Musa chunii Hakkinen, a new species (Musaceae) from Yunnan,China and taxonomic identity of Musa rubra

The center of diversity of the genus Musa (Musaceae) is in Southeast Asia, a region not studied in detail and where new species and varieties continue to be reported. A new wild banana species, M. chunii Hakki-nen from Yunnan, China is described and illustrated based on observed morphological characteristics in the field. This extremely rare new species was only found in Tongbiguan Nature Reserve, Dehong District, West Yunnan. A key to M. chunii and related taxa is provided. In addition, critical notes regarding M. rubra Kurz identity are given.     

薇甘菊对香港郊野公园植物群落危害的分析

从薇甘菊(Mikania micrantha)对不同类型群落的危害,群落中各植物种类受薇甘菊危害的程度,以及薇甘菊与群落中主要乔木和灌木的种间联结性等方面,对香港郊野公园植物群落受薇甘菊危害状况进行了分析。研究样地所在的植被受薇甘菊的危害严重,草灌丛中薇甘菊的平均覆盖率高达80%,灌木群落中薇甘菊的平均覆盖率为60%左右,常绿阔叶林中的薇甘菊覆盖率平均约为30%。对香港郊野公园受薇甘菊危害的植物种类分析表明:受薇甘菊危害最大的主要是乔木和灌木,草本和藤本的受害比例和受害程度相对较小。受薇甘菊危害的乔木有30种,占群落中乔木的62.50%;受薇甘菊危害的灌木有29种,占群落中灌木的70.73%;受薇甘菊危害的草本植物有10种,占群落中草本植物的33.33%。通过无样方法采用2×2列联表的χ2检验对薇甘菊与群落中主要乔木和灌木的种间联结性分析表明:与薇甘菊呈正联结的乔木有12种,占主要乔木(28种)的42.86%,这些种大多是阳生性乔木;与薇甘菊呈负联结的乔木有16种,占主要乔木的57.16%。与薇甘菊呈正联结的灌木有11种,占主要灌木(19种)的57.89%,多是阳生性灌木;与薇甘菊呈负联结的灌木有8种,占主要灌木的42.11%,这些种大多数也是阳生性灌木,仅九节是耐阴性灌木。     

Intraspecific aggression, colony identity and foraging distances in Sudanese Microtermes spp. (Isoptera: Termitidae: Macrotermitinae)

Abstract. 1. Intraspecific aggression between termite major workers was used to obtain estimates of foraging distances for three Microtermes species in Sudan.
2. Maximum foraging distance recorded for M. sp. nr albopartitus (Sjöstedt) in Khartoum was 11.3 m, giving an estimated minimum colony area of 100 m². This is probably an underestimate. In the Tokar Delta, individuals from single colonies of M. najdensis Harris were encountered up to 42 m apart, giving a colony area of 1390 m².
3. Intraspecific aggression could not be used reliably to distinguish members of different colonies of M. lepidus Sjöstedt. Soil barrier formation between groups of workers in petri dishes may be of use as a supplementary technique, especially for species showing no clear aggression.
4. Experiments on isolated laboratory colonics of M. sp. Nr lepidus Sjöstedt, established from alates collected in Khartoum, further confirmed the value of inter-colony aggression for indicating colony identity.
5. The complications introduced into estimation of subterranean termite foraging areas by overlap and interdigitation of colonies are discussed.     

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.     

Homopterans and an invasive red ant, Myrmica rubra (L.), in Maine

Myrmica rubra (L.), is an invasive ant that is spreading across eastern North America. It is presently found in over 40 communities in Maine and areas in Vermont, New Hampshire, Massachusetts, New York, and several provinces in the Canadian Maritimes and Ontario. In addition to disrupting native ant faunas, invasive ants also have been shown to influence homopteran abundance and species composition. We conducted surveys of Homoptera in infested and noninfested sites and conducted manipulative experiments to quantify the effects of M. rubra on homopteran abundance and composition in the summers of 2003, 2006, and 2007 on Mount Desert Island, ME. In 2003, Homoptera family-level richness was higher in infested sites compared with noninfested sites with two out of three sampling methods. Homopteran abundance in infested compared with noninfested sites depended upon the site. The sites with the highest population of M. rubra were associated with significant differences in Homoptera population abundance. In 2006 and 2007, two out of three host plants sampled had significantly higher abundances of the aphids, Aphis spiraephila Patch and Prociphilus tessellatus Fitch. An ant exclusion field experiment on the native plant, meadowsweet (Spiraea alba Du Roi), resulted in higher abundances of A. spiraephila with M. rubra tending compared with native ant tending. A predator exclusion field experiment was conducted on meadowsweet using adult ladybeetles, Hippodamia convergens Guérin-Méneville, larval green lacewings, Chyrsoperla carnea Stephens, and no predators. Predator impacts on aphid populations were reduced in the presence of M. rubra with C. carnea and moderately reduced with H. convergens.     

The effect of density on aggression between a highly invasive and native fish

Population densities of invasive species fluctuate spatially and temporally, suggesting that the intensity of their aggressive interactions with native species is similarly variable. Although inter‐specific aggression is often thought to increase with population density, it is often theorized that it should be exceeded by intra‐specific aggression since conspecifics share a greater degree of resource overlap. Yet, the magnitude of intra‐specific aggression is seldom considered when examining aggressive interactions, particularly those between invasive and native species. Here, we manipulated the density of the invasive eastern mosquitofish, Gambusia holbrooki, and observed its aggressive interactions with juveniles of the native Australian bass, Macquaria novemaculeata in a laboratory setting. For both species, the magnitudes of intra‐ and inter‐specific aggression were recorded. Regardless of density, the native M. novemaculeata was more aggressive towards heterospecifics than G. holbrooki was. In addition to this, M. novemaculeata was more aggressive to G. holbrooki than towards conspecifics, at both low‐ and high‐density conditions. In contrast, G. holbrooki was similarly aggressive towards M. novemaculeata and G. holbrooki at a high density, yet at low density, displayed significantly more aggression towards conspecifics than M. novemaculeata. These findings demonstrate the importance of considering intra‐specific aggression when exploring behavioural interactions between native and invasive species.     

浙江天童国家森林公园5种常绿阔叶植物的一次和二次抽枝进程

中国东部亚热带常绿阔叶林中, 部分常绿木本物种具有明显的二次抽枝现象, 即在春天完成第一次抽枝, 间隔一定的时间后又开始新一轮抽枝。但是关于这些植物第二次抽枝的进程, 迄今鲜有报道, 限制了人们对这些植物生活史对策的理解。该文以浙江天童常绿阔叶林中有二次抽枝的5种植物光叶山矾(Symplocos lancifolia)、檵木(Loropetalum chinense)、窄基红褐柃(Eurya rubiginosa var. attenuata)、杨梅(Myrica rubra)和栲树(Castanopsis fargesii)为研究对象, 调查并比较了第一次抽枝和第二次抽枝在昆虫取食压力、展叶动态和小枝投资大小3个方面的差异, 探讨第二次抽枝动态的主要特点。结果表明: 1)昆虫取食压力上, 杨梅、栲树2种植物第二次抽枝的叶片虫食率显著高于第一次抽枝, 光叶山矾、檵木和窄基红褐柃3种植物第一、二次抽枝的叶片虫食率没有差异; 2)在现叶速率(叶片数量的增加速率)上, 同一物种第一、二次抽枝的叶片数量增长模式相同, 光叶山矾、檵木、窄基红褐柃和杨梅4个物种呈持续式增长, 栲树则呈爆发式增长, 但是完成现叶所需时间并不完全相同, 光叶山矾、窄基红褐柃和杨梅第二次抽枝完成现叶比第一次少6-9天, 檵木和栲树第一、二次抽枝完成现叶所需时间则无差异; 3)在展叶速率上, 光叶山矾、檵木和窄基红褐柃第二次抽枝显著高于第一次抽枝, 但第一次抽枝较晚的杨梅和栲树第一、二次抽枝之间没有差异; 4)调查的5种植物, 第二次抽枝的投资总量均小于第一次抽枝, 光叶山矾、窄基红褐柃、杨梅和栲树4种植物, 第二次抽枝形成的小枝在叶片数量、单叶面积、小枝长度和小枝直径上都显著小于第一次抽枝, 檵木第二次抽枝形成的小枝在叶片数量和小枝长度上没有差异, 单叶面积和小枝直径则显著小于第一次抽枝。总体上看, 所研究物种第二次抽枝中的叶片虫食率、现叶速率和展叶速率都不小于第一次抽枝, 而第二次抽枝小枝投资总量较小, 这些可能是物种对相对较大的昆虫取食压力和即将来临的冬天不利条件等的适应。     

福建省放线菌结瘤植物共生菌Frankia遗传多样性研究

[目的]了解福建省放线菌结瘤植物共生固氮菌Frankia的遗传多样性.[方法]利用16S-23SrDNA间隔区(rrn)和nifD-K基因间隔区的PCR扩增和RFLP技术,分析了福建省木麻黄、杨梅、桤木、胡颓子等共生Frankia纯培养菌株的遗传差异.[结果]17个菌株获得rrn扩增片段,2个杨梅菌株和1个胡颓子菌株扩增未成功,酶切图谱经聚类分析表明6个地点的细枝木麻黄、短枝木麻黄、粗枝木麻黄12个共生Frankia菌株同源性高,属于一个类群,2个地点的4个杨梅菌株和1个四川桤木菌株亲缘关系近,为另一类群.25个Frankia菌株的,nifD-K基因间隔区PCR-RFLP分析结果显示,7个地点的3种木麻黄14个菌株聚类为一个类群,4个地点的7个杨梅菌株、2个地点的2个四川桤木菌株以及1个台湾桤木菌株聚类为另一个类群,胡颓子菌株则为独立的类群.[结论]研究结果表明福建省共生Frankia遗传多样性丰富.     

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).