转Bt水稻对捕食性天敌黑肩绿盲蝽种群动态的影响

【目的】为研究转Bt水稻对其非靶标害虫稻飞虱的主要捕食性天敌黑肩绿盲蝽Cyrtorhinus lividipennis种群发生动态的影响。【方法】本研究以cry1Ac/cry1Ab融合基因型转Bt抗虫水稻"华恢1号"及其对照亲本"明恢63"为供试水稻,于2011、2012和2013连续3年在广西兴安县"转基因水稻试验基地"开展大田试验。【结果】与对照亲本稻田相比,转Bt水稻稻田黑肩绿盲蝽种群发生量有增加趋势,且在2012年出现极显著差异。此外,就被捕食者褐飞虱Nilaparvata lugens和白背飞虱Sogatella furcifera而言,无论在水稻品种,年份还是两者交互作用之间都存在极显著差异;而就两种稻飞虱总量而言,只在不同年份间存在极显著差异,在水稻品种及两者交互作用间不存在显著性差异。进一步通过相关分析表明,2011—2013连续3年黑肩绿盲蝽与褐飞虱、白背飞虱和两者总虫量之间均存在极显著正相关关系。【结论】转Bt水稻稻田捕食性天敌黑肩绿盲蝽的数量较大,这与非靶标害虫褐飞虱和白背飞虱种群发生量较大有关。     

黑肩绿盲蝽捕食褐飞虱卵的功能与数值反应

褐飞虱作为水稻的重要迁飞性害虫,至今未见有效的生物防治技术报道。黑肩绿盲蝽对褐飞虱卵及低龄若虫有较强的捕食能力,为评价黑肩绿盲蝽对褐飞虱的控害能力,研究了黑肩绿盲蝽若虫各虫期和雌雄成虫对褐飞虱卵的捕食功能反应与数值反应。功能反应研究结果表明,黑肩绿盲蝽对褐飞虱卵的捕食功能反应符合HollingⅡ型方程,捕食量在一定范围内随着猎物密度的增加而增加,若虫的捕食能力大于成虫;四龄若虫理论捕食量最大,为104.2粒卵/d。数值反应研究结果表明,不同褐飞虱卵密度对黑肩绿盲蝽的生长、发育、繁殖有着显著影响。随着猎物密度增大,黑肩绿盲蝽若虫发育速率、雌虫产卵量、孵化率及雌性比增大,成虫寿命减小,最后趋于稳定。猎物密度1、3、5卵/d分别是黑肩绿盲蝽若虫存活、成虫羽化及维持种群最低繁殖力的临界值。在天敌控害潜能的研究过程中,数值反应的研究目的在于探讨捕食者对猎物密度的依赖程度,探明能引起捕食者种群衰退的猎物密度临界值,为大田释放捕食者时增补替代寄主,增强捕食者持续控害能力提供理论依据。     

基于MaxEnt模型的两种捕食性盲蝽潜在分布区及其适生性分析

【目的】为了预测和分析稻飞虱的两种重要捕食性天敌中华淡翅盲蝽Tytthus chinensis和黑肩绿盲蝽Cyrtorhinus lividipennis的潜在分布区,研究气候环境因素对其潜在分布的影响。【方法】Max Ent是以最大熵理论为基础的预测物种地理分布的模型,本文运用Max Ent模型,根据这两种盲蝽的已知分布区及其气候环境影响因素,推测这两种盲蝽的适生性分布及气候环境因素的影响。【结果】中华淡翅盲蝽主要适宜分布区域为中国秦岭-淮河以南地区、东北亚的朝鲜半岛和日本、东南亚以及菲律宾和印度尼西亚群岛部分地区,即东洋区和大洋区北部群岛;黑肩绿盲蝽适宜分布区比中华淡翅盲蝽更广,可达山东、河北以南,陕西、云南、贵州以东的中国大部地区,海外除中华淡翅盲蝽分布区外,也可分布于南美亚马逊雨林地区。研究还发现,最热季度降雨量、平均日温差对中华淡翅盲蝽、黑肩绿盲蝽适宜性分布有重要影响;夏季降雨量大于500 mm、平均日温差低于10℃,有利于这两种盲蝽的分布。【结论】两种捕食性盲蝽适宜生境主要受最热季度降雨量、平均日温差影响。本研究为发挥这两种捕食性盲蝽在害虫生物防治中的作用提供了重要依据。     

Bt水稻对褐飞虱及其二种天敌种群动态的影响

【目的】评价转cry1Ab/Ac基因抗虫水稻(华恢1号,HH1)对非靶标害虫褐飞虱Nilaparvata lugens(St?l)及其2种天敌(蜘蛛和黑肩绿盲蝽Cyrtorhinus lividipennis)种群发生动态的影响。【方法】试验分HH1处理组、亲本明恢63(MH63)组2个处理,每个处理设置3块试验田,定期调查不同处理田块内的褐飞虱及其2种天敌蜘蛛和黑肩绿盲蝽的数量。【结果】转基因水稻HH1对褐飞虱、蜘蛛和黑肩绿盲蝽的种群动态有一定程度的影响,但整个水稻生育期的试验结果显示,与亲本对照水稻MH63相比,转基因抗虫水稻HH1对上述3种非靶标昆虫均无显著差异。【结论】转cry1Ab/Ac基因水稻HH1的种植不会对褐飞虱及蜘蛛、黑肩绿盲蝽产生负面影响。     

黑肩绿盲蝽的国内外研究概况

<正> 黑肩绿盲蝽Cyrtorhinus lividipennis(Reuter)亦称苍翅盲蝽,属半翅目、盲蝽科、嗜卵盲蝽属Cyrtorhinus(=Cyrtorrhinus)它分布广泛,是水稻产区稻飞虱等害虫的重要捕食性天敌。早在1920年就发现对飞虱有捕食作用(Usinger,R.J.1939),1952年日本首先研究了黑肩绿盲蝽的捕食行为(Suenaga,H.et al.,1952)。浙江农业大学植保系于70年代初已在东阳县病虫测报站开始观察其生活习性、田间消长情况和施药影响等。现将有关该天敌的研究概况综述于下。     

水稻挥发物在黑肩绿盲蝽捕食行为中的作用(英文)

在室内研究了水稻品种和氮肥水平对黑肩绿盲蝽对褐飞虱卵嗜好性的影响 ,及其与水稻挥发物的关系。双向选择试验表明 ,黑肩绿盲蝽对不同水稻品种上的褐飞虱卵有着不同的嗜好性 ,但对不同氮肥水平稻株上褐飞虱卵的嗜好性无明显差异。嗅觉试验表明 ,黑肩绿盲蝽对水稻健康苗挥发物的趋性明显地强于空白对照 ,对褐飞虱若虫为害苗挥发物的趋性强于健康苗 ,对褐飞虱雌成虫为害苗的趋性又强于若虫为害苗。不同品种的水稻挥发物对黑肩绿盲蝽的引诱作用存在明显差异 ,但不同氮肥水平的水稻挥发物间差异不显著。上述结果表明 ,水稻挥发物在黑肩绿盲蝽的捕食行为中起着重要作用。文中就如何改进水稻特性和栽培措施以增强天敌作用进行了讨论。     

稻田氮肥施用量对黑肩绿盲蝽捕食功能的影响

在实验室条件下研究了黑肩绿盲蝽Cyrtorhinuslividipennis Reuter在不同含氮量稻株上对褐飞虱Nilaparvata lugens Stal卵和低龄若虫的捕食能力、对褐飞虱卵的捕食功能反应以及褐飞虱蜜露和水稻伤流液对其捕食 能力的影响。结果表明,黑肩绿盲蝽对褐飞虱卵和若虫的捕食量均与寄主植物的含氮量呈显著 负相关。黑肩绿盲蝽在相同氮肥施用量的稻株上连续饲养2代后对褐飞虱卵的捕食能力没有改变 。黑肩绿盲蝽对褐飞虱卵的功能反应呈Holling Ⅱ型方程,其参数瞬时发现率(a)和处置时间(T_h)只与寄主含氮量有关,而与黑肩绿盲蝽种群和褐飞虱卵的来源无关。 在高氮量稻株上黑肩绿盲蝽种群对褐飞虱卵的瞬时发现率(a)下降导致了功能反应的减弱, 而在相同含氮量稻株上黑肩绿盲蝽种群之间的捕食功能没有明显差异。黑肩绿盲蝽成虫取食水 稻伤流液和褐飞虱蜜露时寿命明显延长,取食高氮稻株的褐飞虱分泌的蜜露对延长黑肩绿盲蝽 雌成虫寿命的作用最大。但是,在高氮稻株上褐飞虱蜜露显著降低黑肩绿盲蝽的捕食能力。这 些结果表明黑肩绿盲蝽对褐飞虱自然控制作用的下降是稻田过量施用氮肥后褐飞虱种群增加的 主要原因之一。     

“秕谷草-伪褐飞虱-中华淡翅盲蝽”载体植物系统的可行性

为明确用于防治稻飞虱的“秕谷草-伪褐飞虱-中华淡翅盲蝽”载体植物系统的可行性,本文开展了靶标作物水稻、载体植物秕谷草和靶标害虫褐飞虱、替代猎物伪褐飞虱组合对中华淡翅盲蝽成虫形态、种群增长参数、产卵选择性和取食选择性的影响研究.结果表明: 中华淡翅盲蝽水稻种群(取食褐飞虱卵)的雌雄成虫个体均显著大于秕谷草种群(取食伪褐飞虱卵).取食褐飞虱卵的中华淡翅盲蝽种群增长参数单雌产卵量、净增长率、内禀增长率、周限增长率明显高于取食伪褐飞虱卵的种群,但取食伪褐飞虱卵也有较高的种群增长能力,足以扩繁种群.对于靶标作物水稻和载体植物秕谷草,无论是水稻种群还是秕谷草种群均偏好在水稻上产卵.同时,2个种群的雌虫、雄虫和若虫对褐飞虱卵和伪褐飞虱卵捕食量没有显著性差异.在上述研究基础上,讨论了在田间建立“秕谷草-伪褐飞虱-中华淡翅盲蝽”载体植物系统的可行性.     

不同农事操作技术对稻田害虫和天敌种群动态的影响

【目的】通过分析常规稻作田、稻虾共作田和香根草Vetiveria zizanioides诱集田3种农事操作技术对水稻害虫及其天敌种群动态的影响,为湖北地区新型绿色农事操作技术防控田间害虫、天敌保育和稻田有害生物可持续治理策略提供重要依据。【方法】在水稻抽穗期,采用目测法、扫网法、盆拍法、剥查法对3种农事操作稻田(常规稻作田、稻虾共作田和香根草)内害虫(稻飞虱、稻叶蝉、稻螟虫)和天敌进行搜集和统计。【结果】香根草诱集田内稻飞虱种群数量显著低于稻虾共作田(P0.05),常规稻作田间稻叶蝉、稻螟虫种群数量显著高于稻虾共作田和香根草诱集田数量(P0.05);3种农事操作技术下稻田间天敌种类主要以捕食性天敌蜘蛛类、草蛉类、黑肩绿盲蝽Crytorrhinuslividipennis和青翅隐翅虫Paederus fuscipes为主。其中,稻虾共作田和香根草诱集田拟环纹豹蛛Pardosa pseudoannulata、拟水狼蛛Piratasubpiraticus和黑肩绿盲蝽Cyrtorhinuslividipennis种群数量显著高于常规稻作田(P0.05)。【结论】香根草稻作田间稻叶蝉、稻螟虫种群数量显著低于常规稻作田,稻虾共作田间稻叶蝉、稻飞虱种群数量显著低于常规稻作田;3种农事操作技术下稻田间天敌种类主要以捕食性天敌蜘蛛类、草蛉类、黑肩绿盲蝽为主。     

黑肩绿盲蝽耐饥饿能力的研究

【目的】为早期释放黑肩绿盲蝽Cyrtorhinus lividipennis提供理论依据,研究了黑肩绿盲蝽不同虫态耐受饥饿及低温的能力。【方法】室内条件下,比较了稻苗+水、无苗有水和无苗无水三种食物条件下黑肩绿盲蝽耐饥力大小。【结果】适温(26℃)下,黑肩绿盲蝽3龄若虫、5龄若虫和成虫在稻苗+水的存活时间显著高于无苗有水,无苗有水时的存活时间又显著高于无苗无水,表明稻苗对于黑肩绿盲蝽有营养作用,取食稻苗有助于提高黑肩绿盲蝽的耐饥力。黑肩绿盲蝽成虫耐饥力最强,3龄若虫和5龄若虫接近,有苗有水时,3龄若虫、5龄若虫和成虫的平均存活时间分别为2.10、2.22和4.25 d。在低温(15℃)下,水稻苗对黑肩绿盲蝽成虫的存活不利,有苗有水时雌雄成虫的存活时间均低于无苗有水处理,其中雌成虫两处理间的差异显著(P=0.001)。【结论】黑肩绿盲蝽对饥饿和低温有较强的耐受性,以成虫最强,水稻生长前期田间释放时以成虫为宜。     

Intra-guild predation relaxes natural enemy impacts on herbivore populations

Abstract. 1. To investigate the role of intra-guild predation in mediating the impact of the natural enemy complex on herbivore populations, a manipulative field experiment was conducted using uncaged plots (islets of Spartina cordgrass) on a North American salt marsh. The densities (moderate or low) of two invertebrate predators, the generalist wolf spider Pardosa littoralis and the specialist mirid bug Tytthus vagus , were manipulated in a 2 × 2 factorial design, and the resulting treatment effects on the population growth of their herbivorous prey, Prokelisia planthoppers, were assessed.
2. The abundance of wolf spiders on experimental islets was unaffected by the presence of mirid bugs, however the density of mirid bugs was influenced very negatively by the presence of the wolf spider.
3. The negative effect of the wolf spider on mirid bugs most probably resulted from the intra-guild predation of mirids by spiders because planthopper limitation by the wolf spider alone was significantly greater than when both predators were present.
4. As a result of intra-guild predation, planthopper population growth was positive in the presence of both predators, despite the fact that each predator alone promoted a decrease in planthopper population growth.
5. Notably, the occurrence of intra-guild predation diminished top-down impacts on planthopper populations in a relatively simple food web where strong top-down effects were expected. This result, however, was limited to habitats on the marsh with simply structured vegetation lacking leaf litter.     

Spider predation on a mirid pest in Japanese rice fields

Spiders are common generalist predators, and understanding their potential in biological control is important for the development of integrated pest management programs. In this study, predation by three groups of spiders on the mirid bug Stenotus rubrovittatus (Hemiptera: Miridae) in rice paddies was investigated using DNA-based gut-content analysis. A laboratory feeding study revealed that the detection half-lives of bug DNA in the spider gut at 25 °C was 3.4 days for Lycosidae and 1.5 days for Tetragnathidae. Individual spider predation on the mirid bug was investigated by detecting DNA of prey in field-collected spiders. In total, 1199 spiders were assayed from three spider groups: Pirata subpiraticus (Lycosidae), Tetragnatha spp. (Tetra-gnathidae), and Pachygnatha clercki (Tetra-gnathidae), which each differ in their preferred microhabitat as well as their predatory habits. Detection rates of prey DNA in spiders increased significantly with the density of prey across all spider groups. P. subpiraticus and Tetragnatha spp. predation showed a better fit to a saturated response curve to increasing prey density, while P. clercki showed a simple linear relationship with prey density. Densities of alternative prey species did not affect the detection rates of mirids. These results suggest that predation on pests by generalist predators in an agroecosystem is affected not only by prey abundance but also by predator preference for specific prey. Predator preference is therefore an important factor to consider when estimating the role of natural enemies as biological control agents.     

Niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three pest management strategies in paddy fields

The duck-rice and fish-rice were two important strategies of pest management in the process of organic rice. The niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three strategies of pest management in paddy fields were studied in order to understand the influences of different pest control methods on ecological relationships between the two main types of predators. The results of our study suggested that ducks might prey upon wolf spiders resting on the base of rice plants, thus the populations of wolf spiders decreased in duck-rice treatment. The impact of ducks on mirid bug was not significant because it preferred to congregate on the mid-part of rice plants. The temporal selectivity of wolf spiders was smaller than that of mirid bug, which meant occurrence frequency of wolf spiders was higher compared with mirid bug. In addition, competition between wolf spiders and mirid bug was significantly more intense in duck-rice or fish-rice treatment than in natural control treatment at different stages. Probably, ducks and fishes drove wolf spiders away from their original spatial niche, the base to the mid-part in rice plants, enhancing competition between wolf spiders and mirid bug in duck-rice or fish-rice treatment. Based on the results of this study, duck-rice or fish-rice, as a control pest method in the process of organic rice, should be prudentially treated.     

Niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three pest management strategies in paddy fields

The duck-rice and fish-rice were two important strategies of pest management in the process of organic rice. The niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three strategies of pest management in paddy fields were studied in order to understand the influences of different pest control methods on ecological relationships between the two main types of predators. The results of our study suggested that ducks might prey upon wolf spiders resting on the base of rice plants, thus the populations of wolf spiders decreased in duck-rice treatment. The impact of ducks on mirid bug was not significant because it preferred to congregate on the mid-part of rice plants. The temporal selectivity of wolf spiders was smaller than that of mirid bug, which meant occurrence frequency of wolf spiders was higher compared with mirid bug. In addition, competition between wolf spiders and mirid bug was significantly more intense in duck-rice or fish-rice treatment than in natural control treatment at different stages. Probably, ducks and fishes drove wolf spiders away from their original spatial niche, the base to the mid-part in rice plants, enhancing competition between wolf spiders and mirid bug in duck-rice or fish-rice treatment. Based on the results of this study, duck-rice or fish-rice, as a control pest method in the process of organic rice, should be prudentially treated.     

Stable isotope analyses document intraguild predation in wolf spiders (Araneae: Lycosidae) and underline beneficial effects of alternative prey and microhabitat structure on intraguild prey survival

Intraguild predation (IGP) is common among generalist predators and an important issue in food web theory, because IGP may destabilise communities by increasing extinction of species. Also, IGP may interfere with the effectiveness of generalist predators as biological control agents. In general, occurrence of IGP in laboratory or field studies is inferred from abundance data or direct observations only. We investigated if tracing stable isotopes allows distinction between different types of predation and confirmation of IGP. Wolf spiders were chosen as model organisms for generalist predators; IGP of third instar A. cuneata on second instar P. palustris was investigated in a laboratory experiment. The availability of alternative prey and the complexity of the microhabitat were manipulated, since both factors are thought to facilitate coexistence of predators.
Stable isotope analysis documented predation of A. cuneata on P. palustris and predation on alternative prey by both juveniles. Both the presence of alternative prey and the availability of shelter reduced mortality of juvenile P. palustris during the first week. During the second week mortality increased in complex structure without alternative prey presumably due to enhanced activity and cannibalism among starving P. palustris . Thus, microhabitat complexity and prey abundance may foster coexistence of wolf spiders in the field.
In conclusion, stable isotope analysis was proven a powerful tool to investigate animal behaviour without direct observation. The method allowed disentangling predator feeding behaviour when more than one type of prey was present.     

Development and preliminary application of a triplex real‐time polymerase chain reaction assay for evaluating predation on three planthoppers in a rice ecosystem

A multiplex real‐time quantitative polymerase chain reaction (PCR) assay was developed to simultaneously detect the DNA of three rice planthoppers, that is, Sogatella furcifera (Horváth) (white‐backed planthopper), Nilaparvata lugens (Stål) (brown planthopper) and Laodelphax striatellus (Fallén) (small brown planthopper), in the gut of their predators. The sets of primers and ALLGlo probes were targeted to the regions of internal transcribed spacer 2 (ITS2) genes in nuclear ribosomal DNA (rDNA). The sensitivity, specificity and interference test for the multiplex real‐time quantitative PCR assay were analysed. The assay's detection limits were 100, 1000 and 100 copies for the white‐backed planthopper, the brown planthopper and the small brown planthopper, respectively. The specificity tests showed no cross‐reactivity with genomic DNA from 30 other dominant herbivores, saprophagous insects and predators from rice ecosystem for each planthopper species. The assay was used in a preliminary study of predation events on the three planthoppers by three major spiders viz., Pardosa pseudoannulata (Bösenberg et Strand), Ummeliata insecticeps (Bösenberg et Strand) and Tetragnatha maxillosa Thorell which each differ in their preferred microhabitat as well as their predatory habits in rice field, and the results showed their predation on each planthopper species could be well evaluated using this method. Therefore, the multiplex real‐time quantitative PCR assay provides a new tool to study the mechanisms of prey shifting and natural regulation of the three rice planthoppers by generalist predators in rice ecosystem.     

Effects of intraguild predation and cannibalism in two generalist phytoseiid species on prey density of the pink citrus rust mite in the presence of high-quality food

BioControl - Intraguild predation (IGP) and cannibalism influence the effectiveness of biological control with generalist predators. In Japanese citrus orchards, two generalist phytoseiid species,...     

Resource quality,resource availability,and intraguild predation among omnivorous mirids

Intraguild predation (IGP) among predatory species can influence many plant-arthropod associations. However, the relevance of IGP is poorly understood for truly omnivorous species such as those that can complete development on both animal and plant diets. Here we test the hypothesis that IGP among two omnivorous mirids is more common when extraguild food is either absent or not suitable. Laboratory experiments were performed in experimental cages in order to determine the effect of intraguild prey densities and diet availability on direction and intensity of IGP between Dicyphus tamaninii and Macrolophus caliginosus (Heteroptera: Miridae). Intraguild predation was symmetrical between the two mirid species in the absence of alternative food. Increasing densities of intraguild prey enhanced drastically the incidence of IGP. Intraguild predation was reduced when mirids were in the presence of green or red tomato fruits, but the presence of any other extraguild resources had no impact on IGP level. However, when given before the experiments, all resources with the exception of tomato leaves significantly reduced IGP. A second experiment was performed on live plants to compare the results of the previous trials with that obtained in a more natural setting. No IGP was observed when both mirid species were present on a plant. However, development of the intraguild prey (the more vulnerable stage) was hindered by the presence of the intraguild predator. The potential of such results is discussed from community ecology and biological control perspective.     

用ELISA方法定量评价食虫沟瘤蛛对稻飞虱的捕食作用(英文)

对捕食性节肢动物的捕食作用进行评价是害虫生物防治研究的一个重要内容。本文利用酶联免疫吸附试验（ＥＬＩＳＡ）定量评价了食虫沟瘤蛛对稻飞虱的捕食作用。结果表明,方法特异性完全符合试验要求;食虫沟瘤蛛在捕食３头３－５龄白背飞虱或褐飞虱若虫后的检出期分别为９６小时和１２０小时;在早稻田中,捕食性天敌对白背飞虱和褐飞虱的阳性反应率分别为１９．０５％－４７．３４％和１９．０５％－６６．６７％,在飞虱密度较低时,捕食性天敌仍表现出较高的阳性率;捕食量随飞虱密度的增加而增加,但捕食率下降。捕食性节肢动物是调节褐飞虱种群动态的重要因子之一。     

Influence of intraguild predation among generalist insect predators on the suppression of an herbivore population

We evaluated the influence of intraguild predation among generalist insect predators on the suppression of an herbivore, the aphid Aphis gossypii, to test the appropriateness of the simple three trophic level model proposed by Hairston, Smith, and Slobodkin (1960). We manipulated components of the predator community, including three hemipteran predators and larvae of the predatory green lacewing Chrysoperla carnea, in field enclosure/exclosure experiments to address four questions: (1) Do generalist hemipteran predators feed on C. carnea? (2) Does intraguild predation (IGP) represent a substantial source of mortality for C. carnea? (3) Do predator species act in an independent, additive manner, or do significant interactions occur? (4) Can the experimental addition of some predators result in increased densities of aphids through a trophic cascade effect? Direct observations of predation in the field demonstrated that several generalist predators consume C. carnea and other carnivorous arthropods. Severely reduced survivorship of lacewing larvae in the presence of other predators showed that IGP was a major source of mortality. Decreased survival of lacewing larvae was primarily a result of predation rather than competition. IGP created significant interactions between the influences of lacewings and either Zelus renardii or Nabis predators on aphid population suppression. Despite the fact that the trophic web was too complex to delineate distinct trophic levels within the predatory arthropod community, some trophic links were sufficiently strong to produce cascades from higher-order carnivores to the level of herbivore population dynamics: experimental addition of either Z. renardii or Nabis predators generated sufficient lacewing larval mortality in one experiment to release aphid populations from regulation by lacewing predators. We conclude that intraguild predation in this system is wide-spread and has potentially important influences on the population dynamics of a key herbivore.