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红火蚁(Solenopsis invicta Buren)原产南美洲[1],是世界上百种危险入侵生物之一[2-3]。由于其主动攻击性、高繁殖力、强竞争力、广食性、高破坏性的特点,对人体健康和公共设施设备及生态环境造成危害[4–7]。随着全球运输网络的建立和贸易往来的加速,地理屏障对红火蚁的阻隔作用越来越小。在人类活动参与下,阻止红火蚁扩散变得十分困难[8-9]。目前红火蚁已广泛入侵全球多个国家地区[4, 9-10]。大量研究表明,红火蚁入侵会对本地生物多样性产生威胁,导致本地物种多样性丧失[6, 11–14],甚至会破坏生态系统过程[14-15]。红火蚁入侵不同生境后,通过干扰、资源竞争等显著降低区域内蚂蚁群落多样性[16–24]。
温度是红火蚁觅食活动的重要影响因子[25],温度和降水决定红火蚁分布格局和能否成功入侵[26]。相比本地蚂蚁,红火蚁具有极强的温度适应能力[27-28],对恶劣环境胁迫具有极高的耐受性[27, 29-30],并且它们在受人类改造或干扰的生境中能够快速定殖和发展,可以实现较大的种群规模[25]。随着红火蚁进一步扩散和适应,其入侵并适应的生境类型更加多样化[31]。然而,在红火蚁发生次适宜区,相关研究还较少,其生物学特性、入侵机制、与本地蚂蚁群落的关系等是否会发生改变等应加强研究和探索。
云南省于2013年报道有红火蚁入侵,随后在10余个州(市)发现红火蚁疫情,并且呈现加重趋势,对当地生物多样性、人民生产生活以及公众健康造成了严重影响[32-33]。目前昆明市盘龙区、官渡区、五华区、西山区和呈贡区均有红火蚁发生,呈贡区的发生程度较重,多发生于城市绿地、人工林和撂荒地等生境,通过定期开展药剂防治,红火蚁发生发展处于可控状态。昆明地处滇中高原,经模型预测属于红火蚁中低度适生区,可能不适宜红火蚁长期生存繁殖[34-35]。然而,自发现红火蚁近十年来,红火蚁在本地区发生较为频繁,红火蚁显然对昆明地区生态环境产生了一定的适应性。例如,昆明地区红火蚁的活动盛期为6至11月,与广东吴川地区(盛期为5—6月和10—11月)有一定差异[33]。与起源地相比,昆明地区气温相对偏低,红火蚁在温度适宜的季节能够有一定程度发展,但种群发展总体处于较低水平,种群存在较大波动[32]。在此背景下,昆明地区作为红火蚁的次适宜区,红火蚁入侵及其适应过程对本地蚂蚁造成的影响需要进一步关注。本研究选取昆明两地5种生境类型调查蚂蚁群落,比较不同生境中蚂蚁群落多样性、群落结构差异,分析红火蚁发生密度与本地蚂蚁群落关系,探讨2个科学问题:1)次适宜区红火蚁入侵是否对本地蚂蚁群落产生影响,如何影响?2)次适宜区红火蚁入侵对当地蚂蚁群落产生影响的机制是通过蚁巢密度实现还是种群数量实现?为昆明地区红火蚁防控提供理论参考。
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研究地位于云南省昆明市虹桥路(25°02′13.66″ N,102°47'28.08" E,海拔1 995 m)和沙朗乡(25°09′59.29″ N,102°40′45.76″ E,海拔2 035 m)两地。该地区2019年的年平均气温为15.6 ℃,年均降雨量850 mm,最冷月平均气温8.6 ℃,极端低温−6.7 ℃[36]。在昆明市虹桥路和沙朗乡发现红火蚁发生,虹桥路红火蚁发生地为废弃工厂荒地及育苗地,场地为废弃状态,裸地覆盖有草本层;沙朗乡红火蚁发生地为废弃建筑用地,种植有绿化苗木,草本层不发达。两地相对封闭,人为活动相对较少,可排除人为干扰对蚂蚁群落的影响。为了避免人工药剂防治造成影响,所选择的调查样地调查前未开展过药剂防治,经与管理部门协商在调查期间未开展红火蚁防治。
在上述两个地点选取红火蚁发生和未发生区域,选择红火蚁发生的常见生境类型调查蚂蚁群落,每种类型选择2个重复样地,所选取的调查样地间距200 m以上,调查样地信息见表1。
表 1 昆明市虹桥路和沙朗乡蚂蚁调查样地信息
Table 1. Information of ant sample sites in Hongqiao and Shalang of Kunming
项目
Item代码
Code红火蚁蚁巢密度
Nest density of red fire ant/hm2生境类型
Habitat主要特点
Characteristics红火蚁发生地 HW1 20.2 高红火蚁巢密度荒地 红火蚁入侵超过5年,废弃场地,四周有围栏,基本无人为活动干扰,无乔木,阳光能直射地面。 HW2 17.1 EP1 7.0 桉树林 红火蚁入侵超过4年,与HW相邻,基本无人为活动干扰,种植桉树,树龄7年以上,郁闭度约30%,大部分区域透射阳光。 EP2 6.1 LW1 8.5 低红火蚁巢密度荒地 红火蚁入侵约3年左右,有车辆碾压及堆废弃建筑材料痕迹,人为活动干扰轻微,无乔木,阳光直射地面。 LW2 11.8 EC1 11.1 LW和SF交错区 有挖土、踩踏等人为活动痕迹,人为活动干扰轻微,部分区域有乔木荫蔽,地面在部分时段能阳光直射或透射。 EC2 9.5 红火蚁未发生 SF1 NA 云南松次生林 云南松林为主的杂木林,有枯落物,有捡菌、踩踏痕迹,人为活动干扰中等,郁闭度约70%,能透射阳光。 SF2 NA 注:红火蚁蚁巢密度计算未包括废弃蚁巢,统计蚁巢直径超过5 cm的独立蚁丘并且有红火蚁存活,仅在样地选择首次调查时统计
Notes: Abandoned ant nests were not included in the nest density calculation of red fire ant. Independent anthills with nests over 5 cm in diameter and surviving red fire ants were counted, only counted in the first survey -
5种类型样地的地表蚂蚁群落基于个体数的物种稀疏和预测曲线如图1。5条曲线的实线部分代表的是实际上抽样的个体总数以及物种总数,而虚线部分则代表的是对个体数和物种数的预测值,当虚线部分趋于平缓时说明抽样比较充分,阴影部分表示置信区间。由图可知,5种类型生境抽样曲线较为平缓,尾部虚线趋于平缓,表明5种类型样地的蚂蚁群落抽样较为充分。
图 1 不同栖境基于蚂蚁个体数的物种稀疏和预测曲线
Figure 1. Rarefaction and extrapolation curves of different habitats based on ant individuals
在昆明市虹桥路和沙朗乡两地先后2次共采集蚂蚁2 748头,隶属于4亚科19属34种(形态种)。共采集红火蚁1 920头,本地蚂蚁828头。在桉树林中采集到的红火蚁个体数最多,占比为99.2%,仅采到1种本地蚂蚁,其次为荒地(HW中红火蚁个体数占比94.2%,3种本地蚂蚁,LW中红火蚁个体数占比90.7%,7种本地蚂蚁),在云南松次生林与荒地的交错区(以下简称交错区)中采集到的红火蚁个体数最少,占比为44.7%,15种本地蚂蚁。在云南松次生林中采集到本地蚂蚁18种,未采集到红火蚁,本地蚂蚁物种丰富度及多度最多(表2)。
表 2 昆明市虹桥路和沙朗乡不同生境蚂蚁物种名录及多度
Table 2. Ant species and abundance of different habitats in Hongqiao and Shalang of Kunming
物种名称 Species EP HW LW EC SF 猛蚁亚科 Ponerinae 环纹大齿猛蚁 Odontomachus circulus 1 黄足短猛蚁 Brachyponera luteipes 1 1 扁头猛蚁属待定种1 Ectomomyrmex sp.1 4 扁头猛蚁属待定种2 Ectomomyrmex sp.2 1 扁头猛蚁属待定种3 Ectomomyrmex sp.3 1 切叶蚁亚科 Myrmicinae 棒刺大头蚁 Pheidole spathifera 13 2 大头蚁属待定种1 Pheidole sp.1 10 大头蚁属待定种2 Pheidole sp.2 1 8 厚结大头蚁 Pheidole nodifera 3 乌木举腹蚁 Crematogaster ebenina 1 1 盘腹蚁属待定种1 Aphaenogaster sp.1 3 阿普特铺道蚁 Tetramorium aptum 18 草地铺道蚁 Tetramorium caespitum 12 铺道蚁属待定种1 Tetramorium sp.1 7 铺道蚁属待定种2 Tetramorium sp.2 1 疏毛无刺蚁 Kartidris sparsipila 12 361 裸心结蚁 Cardiocondyla nuda 5 25 1 102 宽结摇蚁 Erromyrmalati nodis 2 红火蚁 Solenopsis invicta 624 571 556 169 臭蚁亚科 Dolichoderinae 无毛凹臭蚁 Ochetellus glaber 2 黑头酸臭蚁 Tapinoma melanocephalum 1 2 酸臭蚁属待定种1 Tapinoma sp.1 1 蚁亚科 Formicinae 开普刺结蚁 Lepisiota capensis 4 角弓背蚁 Camponotus cornis 50 25 弓背蚁属待定种1 Camponotus sp.1 1 弓背蚁属待定种2 Camponotus sp.2 11 弓背蚁属待定种3 Camponotus sp.3 1 长角立毛蚁 Paratrechina longicornis 1 林间毛蚁 Lasius hayashi 6 玉米毛蚁 Lasius alienus 34 74 毛蚁属待定种1 Lasius sp.1 6 缅甸尼氏蚁 Nylanderia birmana 9 黄足尼氏蚁 Nylanderia flavipes 2 斜结蚁待定种1 Plagiolepis sp.1 1 物种丰富度 Species richness 2 4 8 16 18 多度 Abundance 629 606 613 378 522 -
5种类型生境蚂蚁群落物种丰富度和Chao-1估计值有显著差异(物种丰富度:Kruskal-Wallis χ2 = 15.63,P<0.01;Chao-1估计值:Kruskal-Wallis χ2 = 15.62,P<0.01),其中,云南松次生林和交错区的蚂蚁物种丰富度和Chao-1估计值均显著高于桉树林(图2a,b)。5种类型样地蚂蚁群落多度无显著差异(图2c)。5种类型样地蚂蚁群落的优势度有显著差异(Kruskal-Wallis χ2 = 16.79,P<0.01),其中,桉树林、高红火蚁蚁巢密度荒地和低红火蚁蚁巢密度荒地的蚂蚁优势度显著高于交错区(图2d)。
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5种类型生境的蚂蚁群落结构相似性有显著差异(Global Test Sample statistic R =0.81,P<0.01),云南松次生林的蚂蚁群落结构明显不同于红火蚁发生生境。在红火蚁发生生境中,高密度红火蚁发生样地的蚂蚁群落结构与桉树林相似;交错区的蚂蚁群落结构与高密度红火蚁发生样地和桉树林不相似;低密度红火蚁发生样地的蚂蚁群落结构介于上述二者之间(图3)。
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红火蚁蚁巢密度与红火蚁多度、本地蚂蚁多度和本地蚂蚁物种丰富度均无显著关联。随着红火蚁蚁巢密度的增加,红火蚁多度无显著增加(图4a);随着红火蚁蚁巢密度的增加,本地蚂蚁多度及物种丰富度未发生明显变化(图4b,c)。
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随着红火蚁多度的增加,本地蚂蚁物种丰富度呈显著线性下降(t = −3.36,P<0.01),二者为中等程度负相关(Pearson相关系数=−0.67)(图5a);本地蚂蚁多度随着红火蚁多度的增加呈显著线性下降(t = −2.66,P<0.05),二者也为中等程度负相关(Pearson相关系数=−0.58)(图5b)。
红火蚁次适宜区本地蚂蚁群落研究——以昆明为例
Influence of Solenopsis invicta on Local Ant Communities in Sub-suitable Areas——An Example in Kunming
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摘要:
目的 红火蚁入侵导致了严重的生态问题。温度是制约红火蚁入侵并发展的关键因素。本研究在云南省昆明市不同生境调查蚂蚁群落,旨在揭示红火蚁在次适宜区对本地蚂蚁群落的影响。 方法 于2021年6月和8月,选择云南省昆明市虹桥路和沙朗乡区域5种生境作为研究样地,采用陷阱法调查了不同样地蚂蚁群落,分析不同生境蚂蚁群落多样性、群落结构相似性以及本地蚂蚁群落与红火蚁发生情况关系。 结果 5种生境基于蚂蚁个体数的物种稀疏和预测曲线较平缓,抽样充分;5种生境中共采集蚂蚁2 748头,隶属于4亚科19属34种,其中红火蚁1 920头;桉树林中采集到红火蚁个体数最多(99.2%),其次为荒地(高红火蚁蚁巢密度HW 94.2%,低红火蚁蚁巢密度LW 90.7%),云南松次生林与荒地交错区最少(44.7%);云南松次生林中未采集到红火蚁;桉树林中仅采集到1种本地蚂蚁,荒地中分别采集到3种(HW)和7种(LW),交错区中采集到15种,云南松次生林中采集到18种;云南松次生林和交错区蚂蚁物种丰富度和Chao-1估计值显著高于桉树林,而二者优势度指数则显著低于桉树林;5种生境的蚂蚁群落结构相似性有显著差异,有红火蚁发生的生境的蚂蚁群落结构与云南松次生林不相似;红火蚁蚁巢密度与红火蚁工蚁多度、本地蚂蚁多度、物种丰富度无显著关联;随着红火蚁工蚁多度的增加,本地蚂蚁物种丰富度和多度均呈显著线性下降。 结论 在昆明红火蚁次适宜区,红火蚁多发生于受干扰的开阔生境,特别是稀疏的桉树林,而很少入侵郁闭度高的生境;红火蚁入侵后导致本地蚂蚁群落多样性水平显著降低,改变蚂蚁群落结构;红火蚁入侵后,需要种群发展至较大规模才表现出种群数量优势,进而对本地蚂蚁产生显著影响;减少红火蚁适宜生境,对适宜生境加强巡查,及时对中大型蚁巢开展药剂治理将有利于降低红火蚁发生发展。 Abstract:Objective The invasion of Solenopsis invicta has caused serious ecological problems, and temperature is the key factor limiting the invasion and development of S. invicta. This study investigated the ant communities in different habitats in Kunming, Yunnan Province, to reveal the influence of S. invicta on local ant communities in sub-suitable areas. Methods In June and August 2021, based on five habitats in Hongqiao Road and Shalang, Kunming City, Yunnan Province, the ant communities in the different habitats were surveyed by pitfall traps, and the relationship between the local ant communities and the occurrence of S. invicta was analyzed. Results The thinning and prediction curves based on the ant individuals in five habitats were smooth and the sampling was sufficient; A total of 2 748 ants were collected from 5 habitats, belonging to 4 subfamilies, 19 genera and 34 species, of which 1 920 were S. invicta; the individual number of S. invicta collected from eucalyptus forest was the highest (99.2%), followed by wasteland (HW 94.2%, LW 90.7%), and the ecotone between Pinus yunnanensis secondary forest and wasteland had the lowest number of S. invicta collected (44.7%); S. invicta was not collected in P. yunnanensis secondary forest. Only 1 species of native ants was collected in the eucalypt forest, 3 species and 7 species were collected in the wasteland, and 15 species were collected in the ecotone. The species richness and Chao-1 estimate of ants in P. yunnanensis secondary forest and ecotone were significantly higher than those in eucalyptus forest, while the dominance index was significantly lower than that in eucalyptus forest. The similarity of ant community structure in five habitats was significantly different, and the ant community structure in habitats where S. invicta occurred was not similar with that in P. yunnanensis secondary forest; There was no significant correlation between nest density and abundance, local ant abundance and species richness of S. invicta. As the abundance of S. invicta increased, the species richness and abundance of local ants decreased linearly. Conclusion In the sub-suitable area of S. invicta in Kunming, S. invicta mostly occurs in disturbed open habitats, especially in eucalypt forests, but rarely invades habitats with high canopy density; After the invasion of S. invicta, the diversity level of the local ant community decreases significantly and the ant community structure changes; After the invasion of S. invicta, it is necessary for the population to develop to a large scale to show the population quantity advantage, and a small number of local ants can coexist with it during the population development; Reducing the suitable habitats of S. invicta, strengthening the inspection of suitable habitats, and carrying out chemical treatment on medium and large ant nests in time will help reduce the occurrence and development of S. invicta. -
Key words:
- Solenopsis invicta
- / sub-suitable region
- / local ant communities
- / adaptation strategy
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表 1 昆明市虹桥路和沙朗乡蚂蚁调查样地信息
Table 1. Information of ant sample sites in Hongqiao and Shalang of Kunming
项目
Item代码
Code红火蚁蚁巢密度
Nest density of red fire ant/hm2生境类型
Habitat主要特点
Characteristics红火蚁发生地 HW1 20.2 高红火蚁巢密度荒地 红火蚁入侵超过5年,废弃场地,四周有围栏,基本无人为活动干扰,无乔木,阳光能直射地面。 HW2 17.1 EP1 7.0 桉树林 红火蚁入侵超过4年,与HW相邻,基本无人为活动干扰,种植桉树,树龄7年以上,郁闭度约30%,大部分区域透射阳光。 EP2 6.1 LW1 8.5 低红火蚁巢密度荒地 红火蚁入侵约3年左右,有车辆碾压及堆废弃建筑材料痕迹,人为活动干扰轻微,无乔木,阳光直射地面。 LW2 11.8 EC1 11.1 LW和SF交错区 有挖土、踩踏等人为活动痕迹,人为活动干扰轻微,部分区域有乔木荫蔽,地面在部分时段能阳光直射或透射。 EC2 9.5 红火蚁未发生 SF1 NA 云南松次生林 云南松林为主的杂木林,有枯落物,有捡菌、踩踏痕迹,人为活动干扰中等,郁闭度约70%,能透射阳光。 SF2 NA 注:红火蚁蚁巢密度计算未包括废弃蚁巢,统计蚁巢直径超过5 cm的独立蚁丘并且有红火蚁存活,仅在样地选择首次调查时统计
Notes: Abandoned ant nests were not included in the nest density calculation of red fire ant. Independent anthills with nests over 5 cm in diameter and surviving red fire ants were counted, only counted in the first survey表 2 昆明市虹桥路和沙朗乡不同生境蚂蚁物种名录及多度
Table 2. Ant species and abundance of different habitats in Hongqiao and Shalang of Kunming
物种名称 Species EP HW LW EC SF 猛蚁亚科 Ponerinae 环纹大齿猛蚁 Odontomachus circulus 1 黄足短猛蚁 Brachyponera luteipes 1 1 扁头猛蚁属待定种1 Ectomomyrmex sp.1 4 扁头猛蚁属待定种2 Ectomomyrmex sp.2 1 扁头猛蚁属待定种3 Ectomomyrmex sp.3 1 切叶蚁亚科 Myrmicinae 棒刺大头蚁 Pheidole spathifera 13 2 大头蚁属待定种1 Pheidole sp.1 10 大头蚁属待定种2 Pheidole sp.2 1 8 厚结大头蚁 Pheidole nodifera 3 乌木举腹蚁 Crematogaster ebenina 1 1 盘腹蚁属待定种1 Aphaenogaster sp.1 3 阿普特铺道蚁 Tetramorium aptum 18 草地铺道蚁 Tetramorium caespitum 12 铺道蚁属待定种1 Tetramorium sp.1 7 铺道蚁属待定种2 Tetramorium sp.2 1 疏毛无刺蚁 Kartidris sparsipila 12 361 裸心结蚁 Cardiocondyla nuda 5 25 1 102 宽结摇蚁 Erromyrmalati nodis 2 红火蚁 Solenopsis invicta 624 571 556 169 臭蚁亚科 Dolichoderinae 无毛凹臭蚁 Ochetellus glaber 2 黑头酸臭蚁 Tapinoma melanocephalum 1 2 酸臭蚁属待定种1 Tapinoma sp.1 1 蚁亚科 Formicinae 开普刺结蚁 Lepisiota capensis 4 角弓背蚁 Camponotus cornis 50 25 弓背蚁属待定种1 Camponotus sp.1 1 弓背蚁属待定种2 Camponotus sp.2 11 弓背蚁属待定种3 Camponotus sp.3 1 长角立毛蚁 Paratrechina longicornis 1 林间毛蚁 Lasius hayashi 6 玉米毛蚁 Lasius alienus 34 74 毛蚁属待定种1 Lasius sp.1 6 缅甸尼氏蚁 Nylanderia birmana 9 黄足尼氏蚁 Nylanderia flavipes 2 斜结蚁待定种1 Plagiolepis sp.1 1 物种丰富度 Species richness 2 4 8 16 18 多度 Abundance 629 606 613 378 522 -
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