关中旱地小麦生育期土壤速效养分时空变异特征研究

为了揭示旱塬地区土壤主要养分的动态变化规律,探明其时空有效性,在较为精确的时空尺度条件下研究了关中地区小麦生育期间土壤剖面0～100 cm范围内硝态氮、碱解氮、速效磷和速效钾动态变化特征.结果表明:土壤硝态氮在0～100 cm范围内空间分布特征较为明显,在30～40 cm处为硝态氮累积峰值,第90～150天范围为剖面上硝态氮素快速消耗期;土壤碱解氮在0～100 cm范围内存在着二元变异结构,0～40 cm空间变异特征较明显,而40 cm以下则时间变异特征更为突出;土壤速效磷空间梯度变异明显,表现为从上至下速效磷含量递减,小麦生育期间土壤剖面速效磷几乎是同步递减;土壤速效钾则是时间变异特征比空间变异特征更显著,小麦生育期间剖面上各层土壤速效钾含量降低幅度均在50 mg/kg左右.     

旱地不同栽培模式下土壤水分和矿质氮含量的时空变化

以田间试验为研究手段连续两年研究了旱地不同栽培模式和施用氮肥对冬小麦生长期间土壤水分、硝态氮和铵态氮含量时空变化的影响.结果表明:与常规相比,覆膜和覆草均显著增加了0～5、5～10和10～20cm土层水分含量,而施用氮肥却是0～20 cm各土层水分含量均有所降低.不同栽培模式对土壤贮水量的影响园作物各生育时期的不同而不同,返青期覆膜和覆草模式土壤贮水量显著高于常规对照,拔节期覆草模武土壤贮水量显著高于覆膜模式,灌浆期各模式之同土壤贮水量的差异未达显著水平.冬小麦生长期间不同处理的土壤铵态氮含量均较低,时空变化较小;而不同处理的土壤硝态氮含量的时空变化较大,其含量的时空变化与施氮量、栽培模武和不同时期有密切关系.增加氮肥用量明显提高0～20 cm土层硝态氮含量和矿质氮的累积量,随着生长时期的推进,土壤硝态氮含量降低明显.0～100 cm土层中硝态氮的累积量为覆草＞常规＞覆膜.     

狭叶锦鸡儿灌丛沙堆土壤水分和肥力的时空分布

通过对内蒙古高原草原区、荒漠草原区和荒漠区狭叶锦鸡儿(Caragana stenophylla)不同发育阶段灌丛沙堆土壤有机质、全氮、全磷、速效磷和水分的空间分布研究。结果发现:3种生境中,灌丛土壤水肥含量表现为:草原区荒漠草原区荒漠区。在草原区和荒漠草原区,狭叶锦鸡儿灌丛沙堆土壤全氮和全磷随土层加深(0~10 cm、10~20 cm、20~40 cm)呈逐渐增加趋势,而速效磷和水分相反;草原区灌丛沙堆土壤有机质富集于0~10 cm土层,荒漠草原区富集于20~40 cm土层。在荒漠区,灌丛对水分和大部分养分(除速效磷)的层间分布影响不大,而20~40 cm土层的速效磷含量逐渐减少。3种生境中狭叶锦鸡儿灌丛内的养分含量均高于灌丛外相同土层,且随着灌丛增大,这种趋势越来越明显;在草原区,随着狭叶锦鸡儿冠幅增大,灌丛土壤水分含量呈显著下降趋势,在荒漠草原区和荒漠区,灌丛土壤水分含量随着灌幅增大而增加。这说明狭叶锦鸡儿灌丛改变了土壤水分和肥力的空间分布,这种效应是在灌丛沙堆的发育过程中逐渐形成,并体现出来的。     

毛乌素沙地天然臭柏灌丛地的土壤特性

采用样方调查法对毛乌素沙地天然臭柏灌丛地土壤特性调查结果表明 :臭柏灌丛地土壤水分、有机质、全氮、速效钾和碱解氮等养分资源分布具有空间异质性。在臭柏灌丛地(覆盖度 70 %～ 85 % ) 0～ 5 cm的土层 ,聚集了较多的中细粒子 ( 0 .2 5～ 0 .0 2 cm) ,平均含量为68.9% ,比油蒿覆盖区 (覆盖度 40 %～ 5 5 % )高 9% ,并随土层深度的加深 ,其含量降低。水分、有机质、全氮、速效钾和碱解氮等养分含量的分布 ,存在明显的“沃岛”现象。在 0～ 1 0 0 cm的土层范围内 ,臭柏灌丛地 60 %以上的土样含水率为 1 %～ 2 % ,变异系数高达 1 1 3%。土层深度 <2 0 cm时 ,臭柏灌丛土壤含水率由灌丛内向灌丛外逐渐减少 ,当土层 >2 0 cm时 ,则正好相反。70 %的土样速效磷含量 <2 mg/kg,除速效磷之外 ,土壤中的有机质、全氮、速效钾、碱解氮等在臭柏灌丛下 0～ 2 0 cm的土层内相对富集。在 0～ 1 0 0 cm的土层内 ,有机质的变异系数为 96.3% ,全氮为 1 0 1 .4% ,碱解氮为 80 .6% ,速效钾为 88.4%。臭柏灌丛地土壤特性的空间异质性 ,是生物和物理过程共同作用的结果。     

黄土高原沟壑区果园还耕对土壤水分的影响

在黄土高原沟壑区王东沟小流域,在塬面、梁地和坡地三种地貌类型上,分别选取了盛果期果园、老果园和退耕还田等3种土地利用与管理方式,测定土壤0～400 cm(塬面0～600 c m)水分含量分布,研究果园的利用管理方式变化对土壤剖面中水分含量变化的影响.结果表明,梁地和坡地上,盛果期果园、老果园土壤水分含量均接近作物凋萎湿度(10%);塬面上,盛果期果园、老果园土壤水分含量为15%.退果4年的耕地、坡地0～400 cm土层土壤储水量显著增加,梁地和塬面虽有增加但与老果园相比,剖面储水量并没有达到显著水平;退果4年耕地、坡地和梁地剖面中的60～140 cm和220～400 cm含水量提高最显著,塬面上22 0～600 cm土层中水分提高显著.     

滴灌时长对伊犁河流域土壤理化性质及作物产量的影响

为阐明新疆伊犁河南岸灌区不同滴灌应用年限下新开垦耕地土壤水分分布和土壤全氮及有机质分布,进一步了解干旱区荒地引水灌溉开垦过程中土壤养分变化,本文以伊犁河流域连续新开垦地区——伊犁河南岸灌区管理处为研究区,以冬小麦(Triticum aestivum L.)为研究对象,通过监测不同滴灌应用年限(1 a,2 a,3 a,4 a)以及未开垦荒地(CK) 0～60 cm剖面土壤水分、全氮含量和土壤有机质含量,揭示滴灌应用年限对滴灌条件下土壤理化性质以及冬小麦产量的影响。结果表明:新垦土地土壤含水率会随滴灌应用年限延长而增加,种植和灌溉会改善新开荒地的土壤水入渗条件,有利于耕作层水分调蓄;新垦土地各层土壤全氮会随滴灌应用年限增加而增加,且随深度增加分布趋势由开垦前先减少后增加变化为先增加后减少,在0～30 cm土层出现表聚现象;不同滴灌应用年限0～60 cm土壤有机质含量随年限增加而增加,开垦种植增加了0～40 cm土壤有机质含量,并在0～20 cm土层明显富集;根据全国第二次土壤普查养分分级相关标准,研究区荒地开垦后土壤养分全氮及有机质均提高了一级;新开垦耕地灌溉1～4 a,冬小麦产量随滴灌应用年限的增加而增加。养分及产量增加的主要原因在于当地现行的灌溉制度及作物秸秆残留。     

蔬菜日光温室栽培条件下土壤养分累积特性研究

测定了陕西杨凌示范区不同日光温室栽培土壤剖面的养分含量及变化。结果表明,日光温室土壤有机质、硝态氮、有效磷和速效钾含量呈现显著累积趋势,其中以0~20 cm土层有机质、0~40 cm土层有效磷和速效钾以及0~200 cm土层硝态氮累积量尤为突出。蔬菜生长过程中温室土壤剖面中铵态氮含量的变化相对较小,而硝态氮含量呈逐渐降低趋势。蔬菜收获后土壤剖面(0~200 cm)硝态氮残留量在707~1 161 kg/hm2之间,平均达954kg/hm2,明显高于农田土壤。温室栽培条件下土壤电导率明显高于农田土壤,以土壤表层尤为突出。土壤硝态氮、有效磷和速效钾与土壤电导率之间的关系均达显著水平,其中硝态氮含量与土壤电导率间的关系最为密切,说明过量施用肥料特别是氮肥,是引起土壤盐分累积的主要原因。     

保护性耕作对黄土高原春玉米田土壤理化特性的影响

为了探索黄土高原春玉米区保护性耕作农田土壤理化特性变化,测定分析了保护性耕作处理第二年春玉米生长不同时期农田土壤容重、水分和养分变化.结果表明,保护性耕作处理在玉米生长前期0～10 cm土壤容重呈下降趋势且小于传统耕作,但后期增加幅度较大;与传统耕作相比保护性耕作能显著增加玉米生长前期表层0～60 cm和后期100～200 cm土壤含水量 ,有较好保水、蓄水作用;保护性耕作下表层0～20 cm土壤养分指标除全磷外,均表现为稳定升高趋势,且能有效提高土壤全钾和速效钾含量;土壤有机质、全氮、速效氮、速效磷含量低于传统耕作,但变异系数较小.     

施用氮磷钾对黄土丘陵区山地红枣林土壤酶与土壤肥力的影响

采用野外试验与室内分析,研究了连续4 a施氮磷钾肥对黄土丘陵区山地枣园土壤酶活性及土壤养分含量的影响。结果表明:连续4 aNPK1施肥处理能够明显增强土壤脲酶、磷酸酶、过氧化氢酶和蔗糖酶的活性,提高土壤有机质、全氮、速效氮、速效磷和速效钾含量;在土壤剖面中总的趋势是土壤酶活性和土壤速效氮、速效磷和速效钾含量随土层深度加深而降低。土壤酶活性与土壤养分因子的相关分析表明,在0~20 cm土层,磷酸酶与有机质、速效磷,过氧化氢酶与有机质、全氮呈显著或极显著性正相关。在20~40 cm,脲酶与有机质、全氮、速效氮,磷酸酶与速效钾,过氧化氢酶与速效磷,蔗糖酶与有机质、速效钾呈显著或极显著性正相关。在40~60 cm,脲酶与全氮、速效磷,磷酸酶与有机质、速效氮,过氧化氢酶与速效磷,蔗糖酶与有机质、全氮、速效磷呈显著或极显著正相关。     

人工林土壤养分含量变化的对比研究

通过现场采样研究了内蒙古多伦县一家河牧场华北落叶松(Larix principis-rupprechtii)人工林和兴安杨(Populus hsinganica C.Wanget Skv.)人工林0-60cm深度土壤有机质、活性有机质、土壤全量养分和速效养分含量的变化。结果表明:在相同立地条件下,不同树种相同土壤层间土壤有机质、活性有机质(0-5cm土层除外)含量存在明显差异。两树种5cm以下土层土壤活性有机质含量的差异显著,落叶松人工林5-20cm土层、20-40cm土层和40-60cm土层土壤活性有机质含量分别比兴安杨人工林高35.3%、38.8%和45.9%。两树种土壤速效氮、磷、钾含量的表聚性明显,并且随着土层深度增加速效氮、磷、钾含量降低趋势显著。落叶松人工林0-5cm、5-20cm和20-40cm土层全钾含量分别比兴安杨人工林相应的土层高30.9%、35.4%和31.6%。兴安杨人工林0-5cm土层全氮含量比落叶松人工林高55.1%。     

Dermestid beetles in ‘evolution Canyon’, lower Nahal Oren, Mt. Carmel, including new records for Israel

Fifteen species of dermestid beetles were recorded at ‘Evolution Canyon’ (EC), Lower Nahal Oren, Mt. Carmel, Israel. They represent ~35% of known Israeli dermestid species. The following three species were recorded for the first time in Israel:Trogoderma svriaca Dalla Torre, 1911;Ctesias svriaca Ganglbauer, 1904; andAnthrenus (s.str.) jordaniens Pic, 1934. Adults of 13 species were collected on the more solar radiated, warmer and climatically more fluctuating south-facing slope (SFS); ten species were collected on the opposite, north-facing slope (NFS), which was cooler and climatically more stable. The abundance of adult dermestid beetles was 1.9 times higher on the SFS than on the NFS (86 and 47, respectively). Species richness and abundance distribution at EC (three collecting stations on each slope and one at the valley bottom) were significantly negatively correlated with the plant cover that consisted of trees and bushes (Spearmanr _s ,P=0.007 and 0.039, respectively) and perennials (Spearmanr _s ,P=0.039 and 0.077, respectively), indicating that non-woody plants were preferred by adult dermestid beetles.     

Hymenopteran Parasitoids on Fruit-infesting Tephritidae (Diptera) in Latin America and the Southern United States: Diversity, Distribution, Taxonomic Status and their use in Fruit Fly Biological Control

We first discuss the diversity of fruit fly (Diptera: Tephritidae) parasitoids (Hymenoptera) of the Neotropics. Even though the emphasis is on Anastrepha parasitoids, we also review all the information available on parasitoids attacking flies in the genera Ceratitis, Rhagoletis, Rhagoletotrypeta, Toxotrypana and Zonosemata. We center our analysis in parasitoid guilds, parasitoid assemblage size and fly host profiles. We also discuss distribution patterns and the taxonomic status of all known Anastrepha parasitoids. We follow by providing a historical overview of biological control of pestiferous tephritids in Latin American and Florida (U.S.A.) and by analyzing the success or failure of classical and augmentative biological control programs implemented to date in these regions. We also discuss the lack of success of introductions of exotic fruit fly parasitoids in various Latin American countries. We finish by discussing the most pressing needs related to fruit fly biological control (classical, augmentative, and conservation modalities) in areas of the Neotropics where fruit fly populations severely restrict the development of commercial fruit growing. We also address the need for much more intensive research on the bioecology of native fruit fly parasitoids.     

Diagnostics,genetic diversity and pathogenic variation of ascochyta blight of cool season food and feed legumes

Molecular diagnostic techniques have been developed to differentiate the Ascochyta pathogens that infect cool season food and feed legumes, as well as to improve the sensitivity of detecting latent infection in plant tissues. A seed sampling technique was developed to detect a 1% level of infection by Ascochyta rabiei in commercial chickpea seed. The Ascochyta pathogens were shown to be genetically diverse in countries where the pathogen and host have coexisted for a long time. However, where the pathogen was recently introduced, such as A. rabiei to Australia, the level of diversity remained relatively low, even as the pathogen spread to all chickpea-growing areas. Pathogenic variability of A. rabiei and Ascochyta pinodes pathogens in chickpea and field pea respectively, appears to be quantitative, where measures of disease severity were based on aggressiveness (quantitative level of infection) rather than on true qualitative virulence. In contrast, qualitative differences in pathogenicity in lentil and faba bean genotypes indicated the existence of pathotypes of Ascochyta lentis and Ascochyta fabae. Therefore, reports of pathotype discrimination based on quantitative differences in pathogenicity in a set of specific genotypes is questionable for several of the ascochyta-legume pathosystems such as A. rabiei and A. pinodes. This is not surprising since host resistance to these pathogens has been reported to be mainly quantitative, making it difficult for the pathogen to overcome specific resistance genes and form pathotypes. For robust pathogenicity assessment, there needs to be consistency in selection of differential host genotypes, screening conditions and disease evaluation techniques for each of the Ascochyta sp. in legume-growing countries throughout the world. Nevertheless, knowledge of pathotype diversity and aggressiveness within populations is important in the selection of resistant genotypes.     

Brief Guide for Authors

正Journal of Arid Land(JAL)is an international journal(ISSN 1674-6767;CN 65-1278/K)for the natural sciences,sponsored by the Xinjiang Institute of Ecology and Geography,Chinese Academy of Sciences and Science Press.It is published by Science Press and Springer-Verlag Berlin Heidelberg bimonthly.JAL publishes original,innovative,and integrative research from arid and semiarid regions,ad     

Epidemiology of Toxigenic Fungi and their Associated Mycotoxins for Some Mediterranean Crops

Recent data on the epidemiology of the common mycotoxigenic species of Fusarium, Alternaria, Aspergillus and Penicillium in infected or colonized plants, and in stored or processed plant products from the Mediterranean area are reviewed. Emphasis is placed on the toxigenicity of the causal fungal species and the natural occurrence of well known mycotoxins (aflatoxins, ochratoxins, fumonisins, trichothecenes, zearalenone, patulin, Alternaria-toxins and moniliformin), as well as some more recently described compounds (fusaproliferin, beauvericin) whose toxigenic potential is not yet well understood. Several Fusarium species reported from throughout the Mediterranean area are responsible of the formation of mycotoxins in infected plants and in plant products, including: Fusarium graminearum, F. culmorum, F. cerealis, F. avenaceum, F. sporotrichioides and F. poae, which produce deoxynivalenol, nivalenol, fusarenone, zearalenone, moniliformin, and T-2 toxin derivatives in wheat and other small grains affected by head blight or scab, and in maize affected by red ear rot. Moreover, strains of F. verticillioides, F. proliferatum, and F. subglutinans, that form fumonisins, beauvericin, fusaproliferin, and moniliformin, are commonly associated with maize affected by ear rot. Fumonisins, were also associated with Fusarium crown and root rot of asparagus and Fusarium endosepsis of figs, caused primarily by F. proliferatum. Toxigenic A. alternata strains and associated tenuazonic acid and alternariols were commonly found in black mould of tomato, black rot of olive and citrus, black point of small cereals, and black mould of several vegetables. Toxigenic strains of A. carbonarius and ochratoxin A were often found associated with black rot of grapes, whereas toxigenic strains of A. flavus and/or P. verrucosum, forming aflatoxins and ochratoxin A, respectively, were found in moulded plant products from small cereals, peanuts, figs, pea, oilseed rape, sunflower seeds, sesame seeds, pistachios, and almonds. Finally, toxigenic strains of P. expansum and patulin were frequently found in apple, pear and other fresh fruits affected by blue mould rot, as well as in derived juices and jams.     

A survey of the genera of<Emphasis Type="Italic">Microgaster</Emphasis> latreille and<Emphasis Type="Italic">Hygroplitis</Emphasis> thomson (Hymenoptera: Braconidae: Microgastrinae) from China

The genera ofMicrogaster Latreille 1804 andHygroplitis Thomson 1895 from China are presented systematically in this paper. Thirty-two species ofMicrogaster and three species ofHygroplitis are known in China. Diagnosis, character variation, distribution and host of each species among the two genera are presented, including its host and distribution. Keys to the species ofMicrogaster andHygroplitis are given. http://www.phytoparasitica.org posting Dec. 19, 2006.     

Plant Viruses Transmitted by Whiteflies

One-hundred and fourteen virus species are transmitted by whiteflies (family Aleyrodidae). Bemisia tabaci transmits 111 of these species while Trialeurodes vaporariorum and T. abutilonia transmit three species each. B. tabaci and T. vaporariorum are present in the European–Mediterranean region, though the former is restricted in its distribution. Of the whitefly-transmitted virus species, 90% belong to the Begomovirus genus, 6% to the Crinivirus genus and the remaining 4% are in the Closterovirus, Ipomovirus or Carlavirus genera. Other named, whitefly-transmitted viruses that have not yet been ranked as species are also documented. The names, abbreviations and synonyms of the whitefly-transmitted viruses are presented in tabulated form together with details of their whitefly vectors, natural hosts and distribution. Entries are also annotated with references. Whitefly-transmitted viruses affecting plants in the European–Mediterranean region have been highlighted in the text.     

The parasitoid complex ofLiriomyza cicerina on chickpea (Cicer arietinum)

Liriomyza cicerina (Diptera: Agromyzidae) is an important pest on chickpea in Turkey. The objective of this study was to determine the parasitoids and rates of parasitism ofL. cicerina on chickpea (Cicer arietinum L.) during the 2005 and 2006 seasons in ?anl?urfa province, Turkey. Leaves with mines were sampled weekly and kept in the laboratory to observe and count emerging leafminer and parasitoid adults. Eight parasitoid species were collected: the braconidsOpius monilicornis Fischer andOpius tersus Foerster and the eulophidsDiaulinopsis arenaria (Erdös) andNeochrysocharis formosa (Westwood), which occurred in both the winter and summer seasons;Diglyphus crassinervis Erdös,Neochrysocharis ambitiosa Hansson,Neochrysocharis sericea (Erdös) andPediobius metallicus (Nees), which occurred only in the summer growing areas.Diaulinopsis arenaria was the predominant parasitoid with 4–7.7% parasitism rate whileN. ambitiosa andO. monilicornis were the second and third most predominant species. The results of these trials show that sinceDia. arenaria occurred throughout every season, it could potentially be used for control of the leafminerL. cicerina.     

Plant–herbivore asynchrony necessitates augmentative releases of the exotic beetle,Zygogramma bicolorata,to enhance the biological control of P arthenium hysterophorus

Systematic information on the quantitative impact of Z ygogramma bicolorata on the biology of P arthenium hysterophorus is crucial as the seeds of this weed continue to germinate from the accumulated soil seed bank throughout the year in the form of different germinating flushes, while the activity of the beetle ceases during winter as it enters diapause. Therefore, plant–herbivore interactions need to be explored to develop predictions of the overall impact of the introduced beetle on the weed. The findings revealed that defoliation by Z . bicolorata had a significant impact on the plant height, density and flower production in flushes F ₃, F ₄ and F ₅, but not in F ₁ and F ₂ that exhibited longer periodicity, profuse branching, a longer flowering period and maximum flower production and contributed mostly to the existing seed soil bank. Therefore, total depletion of the existing soil seed bank was not possible. Consequently, the effect of augmentative field releases of laboratory‐reared beetles was explored on F ₁ and F ₂ in February for three consecutive years (2011–2013). Before initiating the trial, random soil samples were taken from the plots that were assigned to the paired treatments (i.e. with the beetle and without the beetle [insecticide‐treated]) and it was found that the seed bank in those samples did not differ. The single release of Z . bicolorata adults at five per plant at the six‐leaf stage significantly reduced the soil seed bank, compared to without the biocontrol agent, irrespective of the flushes at the end of the season.     

The use of Solacol® (validamycin) as a growth retardant in the isolation of soil fungi

Solacol^®, a formulation of the antibiotic validamycin, at 0.33% in 2% malt extract agar, reduced the spread of fungi on dilution plates drastically and allowed twice as much incubation time before subculturing; this resulted in an elevated number of species isolated. Using pure cultures of 62 common soil fungi, it was shown that all fast-growing species (exceptPythium ultimum) were efficiently inhibited but not completely suppressed. Inhibition was comparable to that by 0.5% oxgall, though, while this substance completely suppressed several species, Solacol very strongly inhibited onlyGaeumannomyces graminis, Gerlachia nivalis, Harzia acremonioides, Verticillium biguttatum andRhizoctonia solani. In a further experiment each separate constituent of Solacol was tested against 22 fungi at equivalent concentrations. Validamycin strongly inhibitedChaetomium globosum and two Basidiomycetes, though hardly more than the non-ionic detergent which mainly inhibited the other fungi. A few species were, however, more inhibited by Solacol than by the detergent alone. Solacol at 0.33% is a suitable aid in dilution plating of soil fungi, by increasing the number of colonies and species observed.Samenvatting Solacol^®, een formulering van het antibioticum validamycine, remde de groei van schimmels in verdunningsplaten met een concentratie van 0.33% in 2% moutagar en maakte het mogelijk de periode tot afenten met een factor 2 te verlengen; daardoor was het aantal geïsoleerde soorten duidelijk toegenomen. Met reincultures van 62 algemene grondschimmelsoorten werd aangetoond, dat alle snelgroeiende soorten (met uitzondering vanPythium ultimum) voldoende geremd, maar niet volkomen onderdrukt werden. Het remmingspercentage was vergelijkbaar met dat van 0.5% ossegal, hoewel dit laatste sommige soorten volkomen onderdrukte; Solacol remde alleenGaeumannomyces graminis, Gerlachia nivalis, Harzia acremonioides, Verticillium biguttatum enRhizoctonia solani zeer sterk. In een volgend experiment werden de componenten van Solacol t.o.v. 22 fungi apart getoetst in concentraties equivalent aan 0.33% Solacol. Validamycine remde alleenChaetomium globosum en twee basidiomyceten behoorlijk, maar nauwelijks meer dan de niet-ionische uitvloeier, die in hoofdzaak de overige remeffecten veroorzaakte. Enkele soorten werden echter door het complete Solacol veel sterker geremd dan door de uitvloeier alleen. Solacol in een verdunning van 0,33% wordt aanbevolen bij verdunningsplaten voor het isoleren van grondschimmels ten einde het aantal kolonies en soorten te verhogen.