蚂蚁参与的传粉系统——基于鸟巢兰属和对叶兰属三种兰科植物的传粉生物学研究

蚂蚁和被子植物之间存在广泛、普遍和多样的关系，但蚂蚁为被子植物传粉的报道却很少。本文通过对分布于四川省黄龙寺自然保护区黄龙沟内鸟巢兰属和对叶兰属的三种花结构十分相似的植物(高山鸟巢兰Neottia listeroides、小叶对叶兰Listera smithii以及花叶对叶兰L. puberula var. maculata)的传粉生态学研究，首次在国内报道高山鸟巢兰和花叶对叶兰中蚂蚁参与的传粉现象，并探讨了蚂蚁参与的传粉系统中蚂蚁传粉的特点、传粉效率、蚂蚁活动特点与防止自花授粉的关系、以及蚂蚁传粉的作用等问题。同时通过蚂蚁参与传粉的高山鸟巢兰和花叶对叶兰以及没有蚂蚁参与传粉的小叶对叶兰的生境对比，初步分析了蚂蚁参与传粉的生境条件。 黄龙沟内小叶对叶兰、高山鸟巢兰和花叶对叶兰居群水平的花期分别可持续27 d、40 d和50 d，大量开花期(～60%)分别在6月下旬到7月上旬、7月中下旬和8月。这3种植物都具暴露的花蜜，花蜜位于唇瓣中央不明显蜜槽内，花蜜量很小。访问小叶对叶兰的昆虫非常少，2005年和2007年均没有观察到传粉者。在2005年，姬蜂和瘿蜂为花叶对叶兰主要传粉者。在2007年，蚂蚁(包括细胸蚁和立毛蚁)分别是高山鸟巢兰和花叶对叶兰最主要的传粉者，其单花访问次数、携粉次数以及授粉次数分别在这2种植物的所有访问昆虫中是最高的。蚂蚁沿唇瓣蜜槽取食花蜜，到达唇瓣基部后，在高山鸟巢兰上，一般情况下蚂蚁头部不能接触到蕊喙先端，只有当蚂蚁头部向上抬起，才能触碰到蕊喙，花粉团通过蕊喙先端释放的粘滴粘在蚂蚁头部最顶端;而在花叶对叶兰上，只要蚂蚁在唇瓣基部活动就可以触碰到蕊喙，花粉团粘在蚂蚁头部的前额处。当小叶对叶兰、高山鸟巢兰和花叶对叶兰的蕊喙先端被触碰后，立即向下运动盖住柱头，分别约9 h、21 h和23 h重新抬起至药帽位置。蕊喙的这种运动可以避免因蚂蚁的重复访问而引起自花授粉的发生。繁育系统实验表明，这3种植物是自交亲和的，但必须依靠昆虫进行传粉。自然条件下，小叶对叶兰的自然结实率在2005和2007年分别为18.03%和14.42%，花叶对叶兰在2005和2007年分别为17.05%和43.20%，高山鸟巢兰在2007年为19.77%。 细胸蚁和立毛蚁在高山鸟巢兰和花叶对叶兰上的访问频率差异不大，前者略低于后者，但高山鸟巢兰的结实率却远低于花叶对叶兰。这主要是由于高山鸟巢兰与细胸蚁和立毛蚁之间形态上的不完全适应，而花叶对叶兰与2种蚂蚁之间形态上却较为适应的缘故。虽然2007年2种蚂蚁是高山鸟巢兰和花叶对叶兰最有效的传粉者，但这种传粉关系可能只是在特定时间和独特生态环境中出现的。生境干旱可能是导致蚂蚁访花和传粉的重要因素。我们的研究结果也初步表明当环境变化导致原有的主要传粉者缺失时，蚂蚁可以起到补充传粉的作用。

Pollination of Adenocalymma bracteatum (Bignoniaceae): Floral Biology and Visitors

Adenocalymma bracteatum is a shrub of dense foliage and yellow flowers, easily found on grasslands areas in Central Brazil. The aim of this study was to determine the reproductive biology and the flower visitors of A. bracteatum in a pasture area nearby Ivinhema city, MS (Brazil). The flowering peak occurs in winter. The flower reflects ultraviolet light. Anthesis begins at 6:30h, and pollen and nectar are the resources to visitors. We captured 1,038 floral visitors. The bees Apis mellifera (L.), Trigona sp., Trigona spinipes (Fabricius), (Hymenoptera: Apoidea) and the ant Cephalotes sp. (Hymenoptera: Formicidae) were the main visitors. The reproductive tests indicate that A. bracteatum is self compatible, justifying its expansion in altered environments; however, the largest reproductive success was dependant on cross-pollination and self-pollination, evidencing the pollinators importance. Adenocalymma bracteatum presents melittophilous syndrome and bumblebees were the main pollinators in the area. The correlations observed between the climatic variables and the main pollinator species were low or medium.

Abiotic factors shape temporal variation in the structure of an ant-plant network

Despite recognition of key biotic processes in shaping the structure of biological communities, few empirical studies have explored the influences of abiotic factors on the structural properties of mutualistic networks. We tested whether temperature and precipitation contribute to temporal variation in the nestedness of mutualistic ant-plant networks. While maintaining their nested structure, nestedness increased with mean monthly precipitation and, particularly, with monthly temperature. Moreover, some species changed their role in network structure, shifting from peripheral to core species within the nested network. We could summarize that abiotic factors affect plant species in the vegetation (e.g., phenology), meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness. While biotic factors are certainly important, greater attention needs to be given to abiotic factors as underlying determinants of the structures of ecological networks.

Effect of broadcast baiting on abundance patterns of red imported fire ants (Hymenoptera: Formicidae) and key local ant genera at long-term monitoring sites in Brisbane, Australia

In 2001, the red imported fire ant (Solenopsis invicta Buren) was identified in Brisbane, Australia. An eradication program involving broadcast bait treatment with two insect growth regulators and a metabolic inhibitor began in September of that year and is currently ongoing. To gauge the impacts of these treatments on local ant populations, we examined long-term monitoring data and quantified abundance patterns of S. invicta and common local ant genera using a linear mixed-effects model. For S. invicta, presence in pitfalls reduced over time to zero on every site. Significantly higher numbers of S. invicta workers were collected on high-density polygyne sites, which took longer to disinfest compared with monogyne and low-density polygyne sites. For local ants, nine genus groups of the 10 most common genera analyzed either increased in abundance or showed no significant trend. Five of these genus groups were significantly less abundant at the start of monitoring on high-density polygyne sites compared with monogyne and low-density polygyne sites. The genus Pheidole significantly reduced in abundance over time, suggesting that it was affected by treatment efforts. These results demonstrate that the treatment regime used at the time successfully removed S. invicta from these sites in Brisbane, and that most local ant genera were not seriously impacted by the treatment. These results have important implications for current and future prophylactic treatment efforts, and suggest that native ants remain in treated areas to provide some biological resistance to S. invicta.

Competition for pollination by the lesser short-tailed bat and its influence on the flowering phenology of some New Zealand endemics

The lesser short-tailed bat, Mystacina tuberculata, is the only native mammalian pollinator in New Zealand and the only fully temperate bat pollinator in the world. Although this unique status as a pollinator is well recognized, its reduced abundance and cryptic behaviour means little research has been undertaken to assess the contribution of the lesser short-tailed bat (hereafter ‘short-tailed bat’) to pollination in New Zealand. Accordingly, pollination by short-tailed bats has been assumed to be comparatively inconsequential, and the potential impacts of the bat's widespread extirpation have been overlooked. The recent discovery that the short-tailed bat is a major pollinator for at least some of the plants it visits emphasizes the importance of exploring this species' role as a pollinator. Here, our aim was to provide an assessment of the competition for short-tailed bat pollination through study of the temporal variation of flowering. Bats were sampled for pollen, and phenology surveys were conducted simultaneously. We found that the amount and type of pollen carried by the bats varied temporally, with one pollen type dominating samples at any given time. The two plants most consistently observed in the pollen samples flowered sequentially with little temporal overlap, suggesting that their flowering phenology may be adapted to minimize competition for the pollination services of the short-tailed bat.

Ant Colony Optimization Algorithms for Shortest Path Problems

We propose four variants of recently proposed multi-timescale algorithm in [1] for ant colony optimization and study their application on a multi-stage shortest path problem. We study the performance of the various algorithms in this framework. We observe, that one of the variants consistently outperforms the algorithm [1].

Effect of fragmentation, habitat loss and within-patch habitat characteristics on ant assemblages in semi-arid woodlands of eastern Australia.

The reliability of ants as bioindicators of ecosystem condition is dependent on the consistency of their response to localised habitat characteristics, which may be modified by larger-scale effects of habitat fragmentation and loss. We assessed the relative contribution of habitat fragmentation, habitat loss and within-patch habitat characteristics in determining ant assemblages in semi-arid woodland in Queensland, Australia. Species and functional group abundance were recorded using pitfall traps across 20 woodland patches in landscapes that exhibited a range of fragmentation states. Of fragmentation measures, changes in patch area and patch edge contrast exerted the greatest influence on species assemblages, after accounting for differences in habitat loss. However, 35% of fragmentation effects on species were confounded by the effects of habitat characteristics and habitat loss. Within-patch habitat characteristics explained more than twice the amount of species variation attributable to fragmentation and four times the variation explained by habitat loss. The study indicates that within-patch habitat characteristics are the predominant drivers of ant composition. We suggest that caution should be exercised in interpreting the independent effects of habitat fragmentation and loss on ant assemblages without jointly considering localised habitat attributes and associated joint effects.

Modelling the potential spread of Solenopsis invicta Buren (Hymenoptera: Formicidae) (red imported fire ant) in Australia

Quantifying the potential spread and density of an invading organism enables decision-makers to determine the most appropriate response to incursions. We present two linked models that estimate the spread of Solenopsis invicta Buren (red imported fire ant) in Australia based on limited data gathered after its discovery in Brisbane in 2001. A stochastic cellular automaton determines spread within a location (100 km by 100 km) and this is coupled with a model that simulates human-mediated movement of S. invicta to new locations. In the absence of any control measures, the models predict that S. invicta could cover 763 000–4 066 000 km2 by the year 2035 and be found at 200 separate locations around Australia by 2017–2027, depending on the rate of spread. These estimated rates of expansion (assuming no control efforts were in place) are higher than those experienced in the USA in the 1940s during the early invasion phases in that country. Active control efforts and quarantine controls in the USA (including a concerted eradication attempt in the 1960s) may have slowed spread. Further, milder winters, the presence of the polygynous social form, increased trade and human mobility in Australia in 2000s compared with the USA in 1940s could contribute to faster range expansion.

Demographics, pollination syndrome and conservation status of Macrozamia platyrhachis (Zamiaceae), a geographically restricted Queensland cycad

To assess the International Union for Conservation of Nature (IUCN) status of Macrozamia platyrhachis F.M.Bailey, we surveyed this central Queensland cycad for its population abundance and health and its pollinator type and pollination syndrome (thermogenesis and volatile emissions). Plants are locally abundant within the 11 discrete populations surveyed, with an estimated population of 611 315 adult plants. Plants are highly restricted to a small area of occupancy, seed dispersal is nearly non-existent and extreme fires appear to have destroyed almost all seeds and seedlings and decimated the pollinators. Of known Macrozamia pollinators, only the thrips, Cycadothrips chadwicki Mound, were found on cones, and these were found in very low numbers. The pollination syndrome for this cycad appears to be unique, based on two cone traits. For one, thermogenesis peaks in early evening, a contrast with daytime peaks of other Cycadothrips-pollinated Macrozamia, but matches that of the Tranes weevil-pollinated Macrozamia machinii. In addition, cone volatiles include both previously unreported compounds as well as those reported exclusively on either Cycadothrips- or Tranes-pollinated species. Based on its small, fragmented area of occupancy, projected population declines and the unique pollination syndrome, we recommend that M. platyrhachis retain its current status as 'Endangered'. Habitat management plans should stipulate that controlled burns be avoided during cycad coning season and that wildfires be controlled to minimise damage to seedlings and pollinators.

Parallel processor scheduling for minimizing total weighted tardiness using ant colony optimization

In the modern business environment, meeting due dates and avoiding delay penalties are very important goals that can be accomplished by minimizing total weighted tardiness. We consider a scheduling problem in a system of parallel processors with the objective of minimizing total weighted tardiness. Our aim in the present work is to develop an efficient algorithm for solving the parallel processor problem as compared to the available heuristics in the literature and we propose the ant colony optimization approach for this problem. An extensive experimentation is conducted to evaluate the performance of the ACO approach on different problem sizes with the varied tardiness factors. Our experimentation shows that the proposed ant colony optimization algorithm is giving promising results compared to the best of the available heuristics.

Revised benefits and costs of eradicating the red imported fire ant

The paper revisits estimates of cost/benefit for eradication in Australia provided in 2001 which were based largely on information about a US ecosystem. The study had two major components; spread modelling using a cellular automation model provided by Joe Scanlan and an impact analysis undertaken by the remaining authors. The revised figures provided in this study increased the damage estimate from $2.8 billion to $45 billion and the benefit-cost ratio of eradication efforts improved from 25:1 to 390:1.

Reproductive Biology of Corymbia citriodora subsp. variegata and Effective Pollination across its Native Range.

The spotted gum species complex represents a group of four eucalypt hardwood taxa that have a native range that spans the east coast of Australia, with a morphological cline from Victoria to northern Queensland. Of this group, Corymbia citriodora subsp. variegata (CCV) is widespread in south-eastern Queensland and northern New South Wales. It is currently the most commonly harvested native hardwood in Queensland. However, little basic knowledge of the reproductive biology of the species is available to inform genetic improvement and resource management programmes. Here we take an integrative approach, using both field and molecular data, to identify ecological factors important to mating patterns in native populations of CCV. Field observation of pollinator visitation and flowering phenology of 20 trees showed that foraging behaviour of pollinator guilds varies depending on flowering phenology and canopy structure. A positive effect of tree mean flowering effort was found on insect visitation, while bat visitation was predicted by tree height and by the number of trees simultaneously bearing flowers. Moreover, introduced honeybees were observed frequently, performing 73% of detected flower visits. Conversely, nectar-feeding birds and mammals were observed sporadically with lorikeets and honeyeaters each contributing to 11% of visits. Fruit bats, represented solely by the grey-headed flying fox, performed less than 2% of visits. Genotyping at six microsatellite markers in 301 seeds from 17 families sampled from four of Queensland's native forests showed that CCV displays a mixed-mating system that is mostly outcrossing (tm = 0.899 ± 0.021). Preferential effective pollination from near-neighbours was detected by means of maximum-likelihood paternity analysis with up to 16% of reproduction events resulting from selfing. Forty to 48% of fertilising pollen was also carried from longer distance (>60 m). Marked differences in foraging behaviour and visitation frequency between observed pollinator guilds suggests that the observed dichotomy of effective pollen movement in spotted gums may be due to frequent visit from introduced honeybees favouring geitonogamy and sporadic visits from honeyeaters and fruit bats resulting in potential long-distance pollinations.

Controlled pollination methods for creating Corymbia hybrids.

Inter-specific Corymbia hybrids are of increasing interest to plantation forestry, yet there is little knowledge of the most suitable controlled pollination methods for this genus. Inter-specific crosses were made between C. torelliana [CT(maternal parent)] and C. citriodora subsp. variegata (CCV), C. henryi (CH) and C. citriodora subsp. citriodora (CCC) using conventional pollination, one-stop pollination (OSP) and artificially-induced protogyny on yellow buds (AIP Y) pollination methods. Additional treatments included AIP on green buds (AIP G) and the use of exclusion bags for the OSP and AIP methods. Inter-specific hybrids (CT x CCV, CT x CH and CT x CCC) were successfully created using all three pollination methods. The AIP Y treatment provided the highest seed yields and achieved time savings of >41% over the conventional and OSP methods, resulting in up to five-fold increases in operator productivity. However, the AIP Y treatment also had the highest C. torelliana contamination levels (9.3–13.2%). The use of exclusion bags with the AIP method had minimal effect on contamination rates, indicating a high proportion of selfpollen contamination. Contamination rates varied between maternal parents, suggesting variation in selfcompatibility for C. torelliana individuals. AIP using semi-ripe green buds was not effective at reducing selfing and had low operator productivity. The AIP method is suitable for use in a large-scale hybrid breeding program for C. torelliana. When self-pollination effects are managed, it could greatly reduce the costs associated with the production of seed of elite family crosses for commercial forestry deployment.