微生物学报

Treatment	ANOSIM	Treatment	ANOSIM
JY vs. JJ	0.198	0.001	JJ vs. XT	0.947	0.001
JY vs. BG	0.195	0.003	JJ vs. BD	0.897	0.001
JY vs. XT	0.436	0.001	BG vs. XT	0.133	0.025
JY vs. BD	0.439	0.001	BG vs. BD	0.171	0.001
JJ vs. BG	0.723	0.001	XT vs. BD	0.095	0.046

Treatment	ANOSIM	Treatment	ANOSIM
JY vs. JJ	0.198	0.001	JJ vs. XT	0.947	0.001
JY vs. BG	0.195	0.003	JJ vs. BD	0.897	0.001
JY vs. XT	0.436	0.001	BG vs. XT	0.133	0.025
JY vs. BD	0.439	0.001	BG vs. BD	0.171	0.001
JJ vs. BG	0.723	0.001	XT vs. BD	0.095	0.046

Group	Taxonomy	Indicator value	P	Relative abundance (%)	Group	Taxonomy	Indicator value	P	Relative abundance (%)
JY	Candida	0.402	0.011	1.744	XT	Cladosporium	0.379	0.003	11.620
	Cephaliophora	0.596	0.004	1.582		Cystobasidium	0.569	0.002	0.611
	Trichomerium	0.912	0.001	0.918		Preussia	0.410	0.002	0.224
	Wallemia	0.567	0.001	0.504		Neosetophoma	0.495	0.009	0.118
	Moesziomyces	0.464	0.007	0.388		Papiliotrema	0.305	0.036	0.112
	Strelitziana	0.495	0.001	0.233		Coniothyrium	0.449	0.011	0.067
	Neoroussoella	0.307	0.029	0.095		Naganishia	0.314	0.021	0.038
	Keratinophyton	0.270	0.009	0.057		Inopinatum	0.222	0.012	0.030
	Ustilaginoidea	0.358	0.048	0.056		Arxiella	0.336	0.003	0.021
	Cutaneotrichosporon	0.372	0.001	0.030	BD	Curvularia	0.605	0.001	2.658
	Tilletia	0.595	0.001	0.028		Amphobotrys	0.400	0.002	0.950
	Gaertneriomyces	0.288	0.001	0.025		Albifimbria	0.510	0.006	0.645
	Paraconiothyrium	0.350	0.003	0.014		Purpureocillium	0.458	0.019	0.551
	Trichosporon	0.310	0.007	0.014		Rhizophlyctis	0.352	0.009	0.163
	Epichloe	0.284	0.009	0.011		Acremonium	0.345	0.018	0.141
JJ	Kazachstania	0.604	0.001	27.100		Talaromyces	0.441	0.002	0.134
	Diutina	0.584	0.001	2.855		Pseudopithomyces	0.320	0.019	0.133
	Debaryomyces	0.712	0.001	0.669		Neopestalotiopsis	0.293	0.006	0.102
	Botryotrichum	0.578	0.001	0.026		Cryptococcus	0.342	0.014	0.100
BG	Alternaria	0.339	0.026	18.230		Sarocladium	0.361	0.001	0.050
	Geotrichum	0.342	0.024	0.460		Moesziomyces	0.150	0.038	0.043
	Aureobasidium	0.484	0.043	0.455		Paramyrothecium	0.307	0.047	0.039
	Selenophoma	0.392	0.045	0.348		Betamyces	0.246	0.032	0.030
	Meyerozyma	0.337	0.008	0.147		Ustilago	0.332	0.010	0.020
	Mucor	0.349	0.017	0.118		Robillarda	0.155	0.047	0.016
	Buckleyzyma	0.348	0.031	0.057		Emericellopsis	0.230	0.009	0.013
	Coprinopsis	0.200	0.015	0.044

Group	Taxonomy	Indicator value	P	Relative abundance (%)	Group	Taxonomy	Indicator value	P	Relative abundance (%)
JY	Candida	0.402	0.011	1.744	XT	Cladosporium	0.379	0.003	11.620
	Cephaliophora	0.596	0.004	1.582		Cystobasidium	0.569	0.002	0.611
	Trichomerium	0.912	0.001	0.918		Preussia	0.410	0.002	0.224
	Wallemia	0.567	0.001	0.504		Neosetophoma	0.495	0.009	0.118
	Moesziomyces	0.464	0.007	0.388		Papiliotrema	0.305	0.036	0.112
	Strelitziana	0.495	0.001	0.233		Coniothyrium	0.449	0.011	0.067
	Neoroussoella	0.307	0.029	0.095		Naganishia	0.314	0.021	0.038
	Keratinophyton	0.270	0.009	0.057		Inopinatum	0.222	0.012	0.030
	Ustilaginoidea	0.358	0.048	0.056		Arxiella	0.336	0.003	0.021
	Cutaneotrichosporon	0.372	0.001	0.030	BD	Curvularia	0.605	0.001	2.658
	Tilletia	0.595	0.001	0.028		Amphobotrys	0.400	0.002	0.950
	Gaertneriomyces	0.288	0.001	0.025		Albifimbria	0.510	0.006	0.645
	Paraconiothyrium	0.350	0.003	0.014		Purpureocillium	0.458	0.019	0.551
	Trichosporon	0.310	0.007	0.014		Rhizophlyctis	0.352	0.009	0.163
	Epichloe	0.284	0.009	0.011		Acremonium	0.345	0.018	0.141
JJ	Kazachstania	0.604	0.001	27.100		Talaromyces	0.441	0.002	0.134
	Diutina	0.584	0.001	2.855		Pseudopithomyces	0.320	0.019	0.133
	Debaryomyces	0.712	0.001	0.669		Neopestalotiopsis	0.293	0.006	0.102
	Botryotrichum	0.578	0.001	0.026		Cryptococcus	0.342	0.014	0.100
BG	Alternaria	0.339	0.026	18.230		Sarocladium	0.361	0.001	0.050
	Geotrichum	0.342	0.024	0.460		Moesziomyces	0.150	0.038	0.043
	Aureobasidium	0.484	0.043	0.455		Paramyrothecium	0.307	0.047	0.039
	Selenophoma	0.392	0.045	0.348		Betamyces	0.246	0.032	0.030
	Meyerozyma	0.337	0.008	0.147		Ustilago	0.332	0.010	0.020
	Mucor	0.349	0.017	0.118		Robillarda	0.155	0.047	0.016
	Buckleyzyma	0.348	0.031	0.057		Emericellopsis	0.230	0.009	0.013
	Coprinopsis	0.200	0.015	0.044

Group	Taxonomy	Contribution (%)	Group	Taxonomy	Contribution (%)
JY vs. JJ	Kazachstania	35.30	JJ vs. XT	Kazachstania	36.90
	Alternaria	13.70		Alternaria	12.00
	Diutina	5.90		Cladosporium	10.50
	Cephaliophora	3.70		Thelebolus	7.00
	Thelebolus	3.40		Diutina	3.80
JY vs. BG	Kazachstania	22.10	JJ vs. BD	Kazachstania	37.40
	Alternaria	17.00		Cladosporium	7.10
	Cladosporium	7.00		Alternaria	6.70
	Cephaliophora	2.80		Thelebolus	4.90
	Diutina	2.70		Diutina	3.80
JY vs. XT	Kazachstania	21.30	BG vs. XT	Alternaria	17.40
	Alternaria	13.80		Cladosporium	11.60
	Cladosporium	10.90		Thelebolus	7.90
	Thelebolus	7.70		Kazachstania	5.90
	Iodophanus	4.00		Iodophanus	4.50
JY vs. BD	Kazachstania	21.00	BG vs. BD	Alternaria	15.40
	Alternaria	10.70		Cladosporium	7.70
	Cladosporium	6.90		Kazachstania	5.70
	Thelebolus	5.20		Thelebolus	4.90
	Curvularia	3.10		Curvularia	3.50
JJ vs. BG	Kazachstania	36.40	XT vs. BD	Alternaria	13.50
	Alternaria	16.20		Cladosporium	11.00
	Cladosporium	6.90		Thelebolus	10.80
	Diutina	4.50		Iodophanus	5.40
	Aspergillus	1.80		Curvularia	3.70

Group	Taxonomy	Contribution (%)	Group	Taxonomy	Contribution (%)
JY vs. JJ	Kazachstania	35.30	JJ vs. XT	Kazachstania	36.90
	Alternaria	13.70		Alternaria	12.00
	Diutina	5.90		Cladosporium	10.50
	Cephaliophora	3.70		Thelebolus	7.00
	Thelebolus	3.40		Diutina	3.80
JY vs. BG	Kazachstania	22.10	JJ vs. BD	Kazachstania	37.40
	Alternaria	17.00		Cladosporium	7.10
	Cladosporium	7.00		Alternaria	6.70
	Cephaliophora	2.80		Thelebolus	4.90
	Diutina	2.70		Diutina	3.80
JY vs. XT	Kazachstania	21.30	BG vs. XT	Alternaria	17.40
	Alternaria	13.80		Cladosporium	11.60
	Cladosporium	10.90		Thelebolus	7.90
	Thelebolus	7.70		Kazachstania	5.90
	Iodophanus	4.00		Iodophanus	4.50
JY vs. BD	Kazachstania	21.00	BG vs. BD	Alternaria	15.40
	Alternaria	10.70		Cladosporium	7.70
	Cladosporium	6.90		Kazachstania	5.70
	Thelebolus	5.20		Thelebolus	4.90
	Curvularia	3.10		Curvularia	3.50
JJ vs. BG	Kazachstania	36.40	XT vs. BD	Alternaria	13.50
	Alternaria	16.20		Cladosporium	11.00
	Cladosporium	6.90		Thelebolus	10.80
	Diutina	4.50		Iodophanus	5.40
	Aspergillus	1.80		Curvularia	3.70

Group	JY	JJ	BG	XT	BD
Nodes	313	261	303	208	307
Edges	5 368	3 856	4 536	3 710	4 944
Density	0.055	0.057	0.050	0.086	0.053
Modularity	0.821	0.807	0.864	0.728	0.847
Degree (average)	17.150	14.774	14.970	17.837	16.104
Network diameter	14	13	14	9	11
Clustering coefficient	0.814	0.877	0.842	0.923	0.808
Path length (average)	4.262	5.252	4.946	3.656	4.513
Betweenness centrality (average)	513.170	568.050	525.840	116.465	548.570
Eigenvector centrality (average)	1.000	1.000	0.714	0.016	1.000

Group	JY	JJ	BG	XT	BD
Nodes	313	261	303	208	307
Edges	5 368	3 856	4 536	3 710	4 944
Density	0.055	0.057	0.050	0.086	0.053
Modularity	0.821	0.807	0.864	0.728	0.847
Degree (average)	17.150	14.774	14.970	17.837	16.104
Network diameter	14	13	14	9	11
Clustering coefficient	0.814	0.877	0.842	0.923	0.808
Path length (average)	4.262	5.252	4.946	3.656	4.513
Betweenness centrality (average)	513.170	568.050	525.840	116.465	548.570
Eigenvector centrality (average)	1.000	1.000	0.714	0.016	1.000

Group	Plant saprotroph			Avian pathogen
JY	0.70±0.05a	0.04±0.04b	3.70±1.69a	0.40±0.30a	2.10±0.72a	0.21±0.20b
JJ	0.95±0.62a	0.06±0.07b	3.53±1.17a	0.39±0.22a	2.16±0.83a	0.30±0.18a
BG	0.65±0.74a	0.07±0.08b	2.61±1.54b	0.32±0.28a	1.50±0.76b	0.11±0.09c
XT	0.79±0.80a	0.15±0.18a	0.79±0.70c	0.03±0.06b	0.50±0.19c	0.01±0.01d
BD	0.75±0.72a	0.06±0.06b	1.95±1.36b	0.10±0.10b	1.35±0.93b	0.07±0.06cd

Group	Plant saprotroph			Avian pathogen
JY	0.70±0.05a	0.04±0.04b	3.70±1.69a	0.40±0.30a	2.10±0.72a	0.21±0.20b
JJ	0.95±0.62a	0.06±0.07b	3.53±1.17a	0.39±0.22a	2.16±0.83a	0.30±0.18a
BG	0.65±0.74a	0.07±0.08b	2.61±1.54b	0.32±0.28a	1.50±0.76b	0.11±0.09c
XT	0.79±0.80a	0.15±0.18a	0.79±0.70c	0.03±0.06b	0.50±0.19c	0.01±0.01d
BD	0.75±0.72a	0.06±0.06b	1.95±1.36b	0.10±0.10b	1.35±0.93b	0.07±0.06cd

Pathogenic species	Symptoms	Infected target
Aspergillus terreus	Respiratory aspergillosis	Human, bird, poultry, etc.
Aspergillus flavus	Invasive aspergillosis	Human, bird, poultry, etc.
Malassezia restricta	Malassezia infection	Human, bird, cattle, etc.
Malassezia globosa	Malassezia infection	Human, bird, dog, horse, etc.
Cladosporium halotolerans	Respiratory tract infection	Human, bird, frog, etc.
Exophiala nishimurae	Phaeohyphomycosis	Human, fish, amphibian, etc.
Exophiala castellanii	Phaeohyphomycosis	Human, fish, etc.
Trichosporon asahii	Trichosporonosis	Human, reptile, etc.
Cyphellophora fusarioides	Onychomycosis	Human
Cyphellophora suttonii	Skin or nail infection	Human

Pathogenic species	Symptoms	Infected target
Aspergillus terreus	Respiratory aspergillosis	Human, bird, poultry, etc.
Aspergillus flavus	Invasive aspergillosis	Human, bird, poultry, etc.
Malassezia restricta	Malassezia infection	Human, bird, cattle, etc.
Malassezia globosa	Malassezia infection	Human, bird, dog, horse, etc.
Cladosporium halotolerans	Respiratory tract infection	Human, bird, frog, etc.
Exophiala nishimurae	Phaeohyphomycosis	Human, fish, amphibian, etc.
Exophiala castellanii	Phaeohyphomycosis	Human, fish, etc.
Trichosporon asahii	Trichosporonosis	Human, reptile, etc.
Cyphellophora fusarioides	Onychomycosis	Human
Cyphellophora suttonii	Skin or nail infection	Human

巢湖野鸟与同域禽类肠道真菌群落结构特征

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王凌怡 ¹^,² , 黄浩圆 ¹ , 水酷 ¹ , 项兴佳 ¹^,²^,^*

微生物学报 | 研究报告 2026,66(1): 428-442

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微生物学报 | 研究报告 2026, 66(1): 428-442

巢湖野鸟与同域禽类肠道真菌群落结构特征

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王凌怡¹^,², 黄浩圆¹, 水酷¹, 项兴佳¹^,²^,^*

作者信息

^1.安徽大学资源与环境工程学院，安徽合肥

^2.湿地生态系统保护与修复安徽省重点实验室，安徽合肥

Characteristics of fungal community structures in guts of wild birds and sympatric poultry in Chaohu Lake

Lingyi WANG¹^,², Haoyuan HUANG¹, Ku SHUI¹, Xingjia XIANG¹^,²^,^*

Affiliations

^1.School of Resources and Environmental Engineering, Anhui University, Hefei, Anhui, China

^2.Anhui Province Key Laboratory of Wetland Ecosystem Protection and Restoration, Hefei, Anhui, China

出版时间: 2026-01-04 doi: 10.13343/j.cnki.wsxb.20250597

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摘要

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【目的】 鸟类具有独特的生活史特征，是研究肠道微生物的理想模型。野鸟与家禽生态位重叠，增加了病原菌交互传播的风险。本研究重点关注巢湖野鸟(八哥、小天鹅、白骨顶鸡)与同域家禽(家鸭和家鸡)肠道真菌及病原菌的群落特征。 【方法】 采用高通量测序技术(Illumina MiSeq)对中国巢湖流域野鸟及同域家禽肠道真菌群落进行分析，并着重关注各物种肠道病原菌的特征。 【结果】 家鸭与白骨顶鸡肠道真菌多样性显著高于家鸡、八哥和小天鹅。不同物种肠道真菌群落组成存在显著差异。家禽因投喂谷物饲料，其肠道中显著富集子囊菌门(Ascomycota)、被孢霉门(Mortierellomycota)及哈萨克斯坦酵母属(Kazachstania)等与谷物降解相关的真菌类群。小天鹅作为植食性水鸟，其肠道中高效降解植物的真菌枝孢霉属(Cladosporium)在群落中占据主导地位，维持了很高的腐生真菌相对丰度。与家禽相比，野鸟肠道真菌群落构建以确定性过程为主导，表明野鸟具有更强的肠道过滤能力。此外，野鸟携带的病原菌多样性与相对丰度更低。 【结论】 野鸟和家禽肠道真菌群落特征呈现出显著的宿主特异性。家禽因投喂谷物饲料，其肠道中显著富集与谷物降解相关的真菌类群。野鸟肠道呈现更强的过滤能力，降低了病原真菌的多样性与相对丰度。

关键词

肠道真菌 / 病原菌 / 巢湖野鸟 / 禽类

Abstract

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[Objective] Birds, with unique life history characteristics, are ideal models for studying gut microorganisms. The niche overlap between wild birds and poultry increased the risk of interactive transmission of pathogens. This study focused on the community characteristics of gut fungi and pathogens in wild birds (crested myna, tundra swan, and common coot) and sympatric poultry (domestic duck and domestic chicken) in Chaohu Lake. [Methods] High-throughput sequencing (Illumina MiSeq) was employed to analyze the fungal communities in guts of wild birds and sympatric poultry in Chaohu Lake of China, and the characteristics of gut pathogens of each species were particularly studied. [Results] The gut fungal diversity of domestic duck and common coot was significantly higher than that of domestic chicken, crested myna, and tundra swan. There were significant differences in gut fungal community composition among different species. Due to grain-based diets, the guts of poultry were significantly enriched with the fungal taxa related to grain degradation, such as Ascomycota, Mortierellomycota, and Kazachstania. Tundra swan is herbivorous waterfowl. The genus Cladosporium, efficient plant-degrading fungi, dominated in the gut of tundra swan. The gut of tundra swan maintained higher relative abundance of plant saprotroph. The fungal community assembly in guts of wild birds was dominated by deterministic processes, which indicated that wild birds had a stronger gut filtering capacity. In addition, wild birds had lower diversity and relative abundance of pathogens. [Conclusion] The characteristics of gut fungal communities in wild birds and domestic poultry showed significant host specificity. Due to grain-based diets, the guts of poultry were significantly enriched with fungal groups related to grain degradation. The guts of wild birds had a stronger filtering capacity, which reduced the diversity and relative abundance of pathogens.

Key words

gut fungi / pathogen / wild birds in Chaohu Lake / poultry

引用本文

王凌怡, 黄浩圆, 水酷, 项兴佳. 巢湖野鸟与同域禽类肠道真菌群落结构特征. 微生物学报, 2026 , 66 (1) : 428 -442 . DOI: 10.13343/j.cnki.wsxb.20250597

Lingyi WANG, Haoyuan HUANG, Ku SHUI, Xingjia XIANG. Characteristics of fungal community structures in guts of wild birds and sympatric poultry in Chaohu Lake[J]. Acta Microbiologica Sinica, 2026 , 66 (1) : 428 -442 . DOI: 10.13343/j.cnki.wsxb.20250597

正文

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动物肠道微生物群落是宿主的重要组成部分，可影响宿主的营养吸收、免疫状态和行为认知等^[1-2]。肠道微生物群落与宿主之间长期进行双向互作，形成了协同进化且结构复杂的共生关系^[3]。因此，对动物肠道微生物群落结构和功能的研究至关重要^[4]。肠道微生物群落在不同物种间、同一物种不同个体间以及同一个体的不同生长时期均表现出显著差异^[5-6]。研究表明宿主的遗传背景、饮食结构及环境因子共同驱动着肠道微生物群落的动态变化^[7]，其中食性对宿主肠道微生物群落的影响最为显著^[8]。有研究表明从食肉性动物到杂食性、草食性动物，其肠道微生物多样性呈梯度递增趋势^[9]。肠道微生物多样性特征不仅是维持肠道生态系统稳定的基础，还能通过代谢产物互作网络为宿主提供多重生理益处^[10]。

大量研究结果证明肠道细菌群落结构与鸟类食性高度关联。例如，水禽肠道中富集的梭菌属(Clostridium)和拟杆菌属(Bacteroides)细菌可高效降解湿地植物中的纤维素，而陆禽肠道中的优势细菌类群为乳杆菌属(Lactobacillus)和肠球菌属(Enterococcus)，能够消化陆生植物种子和昆虫^[11-12]。白鹤从湿地转向农田觅食时乳杆菌科(Lactobacillaceae)的丰度增加2倍以适应高碳水化合物饮食^[13]。近年来，鸟类肠道真菌群落的研究逐渐受到关注，其群落结构呈现出显著的宿主特异性。水禽肠道中水生真菌，如壶菌门(Chytridiomycota)的定殖与湿地生境密切相关，而陆禽则富含土壤来源的真菌类群(如子囊菌门)^[14]。候鸟群体因迁徙行为面临低温环境和病原体感染等生态挑战^[15]。对欧洲柳莺(Phylloscopus trochilus)的研究发现，越冬期其肠道中耐低温酵母菌(Rhodotorula)的丰度显著升高，该酵母菌可通过合成不饱和脂肪酸维持肠道黏膜的低温适应性^[16]。Wu等^[17]研究发现，白头鹤可调节其肠道真菌群落的构建过程，增强肠道过滤能力，通过定向选择或排除特定微生物类群降低病原菌丰度。

野鸟偏好湿地环境作为栖息地，但湿地周围存在居民区，且饲养着很多家禽。野鸟与家禽的生态位重叠为病原菌的跨物种传播创造了条件^[18-19]。宏基因组学研究显示，越冬水鸟肠道中常携带致病性真菌，如新型隐球菌(Cryptococcus neoformans)、烟曲霉(Aspergillus fumigatus)及念珠菌(Candida albicans)等，其孢子可通过粪便在湿地环境中长期存活^[20]。家禽与野鸟的密切接触可能导致病原真菌双向传播。在鄱阳湖湿地的案例中，从家鸭分离的曲霉菌株与白鹤(Grus leucogeranus)粪便中的菌株具有高度遗传同源性^[21]，这提示野鸟与家禽之间存在持续的病原菌交换与感染。这类人畜共患真菌不仅威胁家禽，甚至会导致禽类从业人员患病。

巢湖作为长江中下游典型的通江湖泊湿地，是中国五大淡水湖之一^[22]。其广阔的浅水滩涂与水生植被为野鸟提供了良好的栖息地，每年可支撑超过5万只水鸟越冬^[23]。巢湖周边家禽养殖密集，年存栏量超200万只，导致野鸟与家禽在湖滨带形成显著的生态位重叠，增加了野鸟和家禽之间病原体的交叉传播风险^[24]。鉴于此，本研究采用高通量测序方法探究巢湖野鸟与同域家禽肠道真菌和病原菌群落结构特征，研究结果在野鸟种群保护和家禽流行性疾病防疫等方面具有重要的科学意义。

1 材料与方法

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1.1 研究区域概况

巢湖(31°16′-32°00′N，117°25′-117°58′E)位于安徽省合肥市境内，是我国长江中下游迁徙水鸟重要的聚集区域。每年，大量迁徙水鸟于10月下旬迁入长江中下游越冬地，次年4月初迁离，在越冬地停留时间约5个月。环巢湖湿地共有3条候鸟迁徙通道，其中1条为水鸟迁徙路线，2条为林鸟迁徙路线。途经巢湖的水鸟主要来自黄河三角洲和江苏盐城等沿海地区。近年来，迁徙至巢湖的候鸟数量和种类逐年增加，环巢湖十大湿地已成为珍稀候鸟重要的补给地和越冬地。此外，巢湖湿地存在大量留鸟种类，因此选择该湖泊作为研究地点。

1.2 样品采集

本研究于2023年1月2日在巢湖的三沙河段进行采样。采集越冬期2种候鸟小天鹅(Cygnus columbianus)及白骨顶鸡(Fulica atra)、1种留鸟八哥(Acridotheres cristatellus)和2种家禽鸡及鸭的粪便样本，用于研究野鸟与同域家禽肠道真菌及病原菌群落特征。本研究严格界定同域采样标准，即家禽采样位点与野鸟主要活动位点的直线距离≤1 km，且家禽和野鸟生境类型高度一致，以排除空间距离过远或生境异质性对宿主菌群组成的影响。在野鸟集群觅食地，采用体外粪便采样法(非损伤取样)采集野鸟新鲜粪便样本，每种野鸟粪便样本采集不少于10份。采集样本前通过双筒望远镜观察觅食野鸟至少1 h，确认觅食野鸟的分布地点。采样时仔细辨别并采集新鲜的野鸟粪便样本，每个样本不少于2 g，放入事先准备好的无菌管中，做好样本标签后置于放有冰袋的保温箱中进行野外保存和运输，回实验室后立刻冻存于-20 ℃冰箱中备用。与野鸟粪便样本采集时间同步，采集同域家禽(鸡及鸭)新鲜粪便样本，选取无临床症状的成年个体，每种家禽粪便样本不少于10份。

利用粪便DNA提取试剂盒(Qiagen QIAamp公司)提取粪便样本中的DNA，通过线粒体分子标记技术确认粪便样本的宿主类型。本研究最终采集样本数93个，包括家鸭样本20个、家鸡样本19个、八哥样本20个、小天鹅样本14个以及白骨顶鸡样本20个。采用真菌专用引物ITS1F (5′-CTTGGTCATTTAGAGGAAGTAA-3′)和ITS2R (5′-GCTGCGTTCTTCATCGATGC-3′)对野鸟及同域家禽粪便样本DNA进行扩增。PCR反应体系(50 µL)：DNA聚合酶(2 U，TaKaRa公司) 25 µL；DNA模板(200 μmol/L) 1 µL；正、反向引物(0.4 μmol/L)各0.5 µL；ddH₂O 23 µL。PCR扩增程序：95 ℃ 5 min；94 ℃ 45 s，55 ℃ 45 s，72 ℃ 45 s，共35个循环；72 ℃ 10 min。扩增产物经2%琼脂糖凝胶电泳检测，将扩增合格的PCR产物送往上海美吉生物公司测序。测序原始数据采用QIIME (v1.9)软件进行生物信息学分析。首先删除低质量(Q<30)和较短(<250 bp)的序列，高质量的序列通过97%相似性聚类得到操作分类单元(operational taxonomic unit, OTU)。进一步去除嵌合体后采用核糖体数据库项目(ribosomal database project, RDP)方法将序列与ITS真菌数据库(UNITE Database)进行比对注释，获得真菌分类信息。由于测序得到的每个样本序列数不同，本研究对每个样本统一抽取17 563条序列数以研究肠道真菌群落组成和多样性特征。

1.3 数据分析

本研究使用单因素方差分析(analysis of variance, ANOVA)方法评估野鸟和同域家禽肠道真菌α多样性及优势菌门类相对丰度的差异。通过非度量多维度分析(non-metric multidimensional scaling, NMDS)^[25]和相似性分析(analysis of similarities, ANOSIM)^[26]比较野鸟和同域家禽肠道真菌群落组成的差异。通过β零模型评估肠道真菌群落构建模式，以估计随机性过程和确定性过程对群落构建影响的相对重要性。通过共现网络推断不同宿主肠道真菌之间的相互作用，计算肠道真菌的拓扑特征。运用FUNGuild功能预测软件对所有真菌物种进行分析，预测肠道真菌的潜在功能。通过FUNGuild预测的动物致病菌物种进一步作为关键词在Web of Science进行检索，确定其感染宿主类型。

2 结果与分析

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2.1 肠道真菌α多样性分析

本研究采用肠道真菌的Chao1指数和ASV richness来表征肠道真菌的多样性。如图1所示，家鸭与白骨顶鸡肠道真菌的Chao1指数及ASV richness显著高于其他物种；家鸡、八哥及小天鹅肠道真菌的Chao1指数及ASV richness较低，且差异不显著。

2.2 肠道真菌群落结构

基于非度量多维度分析(NMDS)研究野生鸟类与同域饲养家禽肠道真菌群落组成，结果显示不同物种间肠道真菌群落组成存在显著差异(P<0.05；图2A和表1)。本研究采用基于相对丰度的β零模型方法，通过计算β零偏差值定量评估确定性过程和随机性过程对野鸟和家禽肠道真菌群落构建的相对贡献度。其中，β零偏差值趋近于0表征随机性过程起主导作用，趋近于1则表明确定性过程占据优势地位。研究结果显示，相较于同域分布的家禽，野鸟肠道真菌群落构建表现出更强的确定性过程(图2B)。

野鸟与同域家禽肠道真菌优势门类为子囊菌门(Ascomycota)、担子菌门(Basidiomycota)、壶菌门(Chytridiomycota)和被孢霉门(Mortierellomycota)。研究表明，家禽肠道中子囊菌门相对丰度高于野鸟，其中家鸡样本呈现出最高值。家鸡肠道中担子菌门相对丰度最低，白骨顶鸡肠道中壶菌门相对丰度最高。小天鹅和白骨顶鸡肠道中被孢霉门相对丰度低于其他物种(图3)。本研究共鉴定出228个真菌属，其中真菌优势属主要为哈萨克斯坦酵母属(Kazachstania)、链格孢属(Alternaria)、枝孢霉属(Cladosporium)和寡囊盘菌属(Thelebolus)。与野鸟相比，家禽以哈萨克斯坦酵母属为绝对优势属，其相对丰度显著高于野鸟。野鸟肠道中枝孢霉属相对丰度显著高于家禽。八哥和小天鹅肠道中链格孢属相对丰度高于其他物种。小天鹅肠道中寡囊盘菌属的相对丰度最高(图4)。

指示菌是能特异性反映特定宿主肠道中显著富集的微生物类群，可作为区分不同宿主肠道真菌群落差异的关键标志。基于属水平的指示物种分析表明，5类宿主肠道真菌群落呈现显著的特异性富集特征(表2)。家鸭肠道内显著富集的真菌属主要有念珠菌属(Candida)、头梗霉属(Cephaliophora)、多臂菌属(Trichomerium)、节菌属(Wallemia)等15种；家鸡肠道内显著富集的真菌属主要有哈萨克斯坦酵母属(Kazachstania)、链状念珠菌属(Diutina)、得巴利酵母菌属(Debaryomyces)和球孢毛葡孢霉属(Botryotrichum) 4种；八哥肠道内显著富集的真菌属主要有交链孢霉属(Alternaria)、地霉属(Geotrichum)、短梗霉属(Aureobasidium)、月痣菌属(Selenophoma)等8种；小天鹅肠道内显著富集的真菌属主要有枝孢霉属(Cladosporium)、囊担菌属(Cystobasidium)、光黑壳属(Preussia)、新刚毛座菌属(Neosetophoma)等9种；白骨顶鸡肠道内显著富集的真菌属主要有弯孢属(Curvularia)、葡萄孢属(Amphobotrys)、白毛菌属(Albifimbria)、淡紫紫孢霉属(Purpureocillium)等17种。

SIMPER分析结果显示，家禽类群(家鸭、家鸡)与野生鸟类(八哥、小天鹅、白骨顶鸡)的肠道真菌群落差异主要由3类核心属驱动，其中哈萨克斯坦酵母属(Kazachstania)在家禽与野鸟肠道真菌群落组成的差异贡献率最高；其次是链格孢属(Alternaria)和枝孢霉属(Cladosporium)。在家禽之间，哈萨克斯坦酵母属(Kazachstania)和链格孢属(Alternaria)是导致家鸭与家鸡肠道真菌群落组成差异的主要菌属。野鸟之间的分析表明，Alternaria和Cladosporium是引起八哥与小天鹅肠道真菌群落组成差异的主要菌属；Alternaria是引起八哥与白骨顶鸡肠道真菌群落组成差异的主要菌属；小天鹅与白骨顶鸡肠道真菌群落组成差异主要由Alternaria、Cladosporium及Thelebolus导致(表3)。

2.3 野鸟与同域家禽肠道真菌网络分析

为研究野鸟与同域家禽肠道真菌种间关系，本研究绘制了种间相关性Co-occurrence共现网络图(图5)。野鸟与家禽肠道真菌网络节点主要属于子囊菌门和担子菌门。家鸭肠道真菌网络复杂度最高，包含313个节点和5 368条边。八哥和白骨顶鸡肠道真菌网络复杂度较高，分别包含303个和307个节点，4 536条和4 944条边。家鸡肠道真菌网络复杂度较低，包含261个节点和3 856条边。小天鹅肠道真菌网络节点与边数最少，仅有208个节点和3 710条边，且模块化程度最低(表4)。

2.4 肠道真菌功能预测

基于肠道微生物功能预测分析，本研究对野鸟肠道真菌中植物腐生菌、动物病原菌及鸟类病原菌的特征进行了深入分析(表5)。植物腐生菌多样性在野鸟与同域家禽之间未观察到显著差异，但小天鹅肠道中植物腐生菌的相对丰度显著高于家禽及其他鸟类(P<0.05)。本研究进一步鉴定出10种具有潜在致病特征的动物病原菌(表6)。野鸟肠道动物病原菌的多样性显著低于家禽(P<0.05)。进一步对动物病原菌深入分析得到5种鸟类病原菌，包括土曲霉菌(Aspergillus terreus)、黄曲霉菌(Aspergillus flavus)、限制马拉色菌(Malassezia restricta)、球形马拉色菌(Malassezia globosa)及耐盐枝孢霉菌(Cladosporium halotolerans) (表6)。野鸟肠道中鸟类病原菌多样性和相对丰度显著低于同域家禽(P<0.05)。小天鹅肠道中鸟类病原菌多样性和相对丰度最低。

3 讨论

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本研究系统分析了巢湖流域野生鸟类(小天鹅、白骨顶鸡、八哥)与同域家禽(家鸡、家鸭)肠道真菌群落及病原菌的组成特征。研究表明小天鹅肠道中真菌多样性最低，这可能是由于小天鹅的食性造成的。小天鹅是植食性水鸟，其他野鸟和家禽是杂食性禽类。食物单一可能是导致小天鹅肠道真菌多样性下降的重要原因^[27]。与家鸡和八哥相比，家鸭和白骨顶鸡肠道真菌多样性更高，这可能是由栖息地环境造成的。家鸡与八哥是陆生禽类，而家鸭与白骨顶鸡偏爱水生环境。作为长江中下游典型的淡水湿地，巢湖流域的浅水区及沼泽生境中富集了大量腐生真菌与寄生真菌^[28]。水生栖息地中多样真菌孢子的摄入是导致家鸭和白骨顶鸡肠道真菌多样性增加的重要原因^[29]。

不同宿主肠道真菌群落组成呈现出显著差异，说明肠道真菌具有显著的宿主特异性^[30]。进一步研究发现，不同野鸟间肠道真菌群落组成的差异小于野鸟与家禽间群落组成的差异。家禽由于涉及食物投喂以及抗生素使用等，导致其与野鸟肠道真菌群落组成出现更大的差异^[31]。进一步基于β零偏差模型的群落构建机制分析显示，野鸟肠道真菌群落构建过程以确定性过程为主导。这一结果可能与野鸟在自然环境中面临的病原菌压力而进化出的特异免疫过滤机制相关。野鸟频繁接触土壤、水体等复杂生境，长期高强度的病原菌暴露驱使野鸟定向筛选肠道真菌类群^[32]。因此，长期自然选择压力造成宿主对不适应菌群产生定向淘汰作用，促使真菌群落结构与宿主生态需求高度匹配^[14]。因此，野鸟宿主免疫筛选和肠道环境过滤等生态过程在其肠道真菌群落构建过程中起关键作用^[33]。相比之下，家禽肠道真菌群落构建过程中随机性过程的贡献更高，这可能源于养殖场均质化环境削弱了生态位限制，或者谷物投喂降低了菌群间资源竞争强度^[31]。此类人为干预使得家禽菌群定殖更多依赖随机扩散与生态漂变，从而形成以随机性过程为主导的群落构建模式^[34]。

Zhang等^[35]研究发现，物种基因型显著影响宿主肠道微生物群落结构，表明2个物种亲缘关系越近其肠道微生物群落组成差异越小。然而，本研究结果不支持以上结论。家鸭和小天鹅同属于雁形目，两者亲缘度高，与其他物种亲缘关系较远，但两者肠道真菌多样性及群落组成差异较其他物种更大。其他物种间亲缘关系与肠道微生物群落相似度也未呈现显著关系。这可能是由于栖息地环境及食性等原因造成的，说明外部环境或者食性在塑造宿主肠道真菌群落结构中的贡献可能高于宿主基因型的影响。

野鸟与家禽肠道真菌群落组成在门和属的水平上呈现显著分异。家禽肠道真菌群落中子囊菌门和被孢霉门显著富集，这一现象与家禽投喂谷物密切相关。子囊菌门包含大量能发酵碳水化合物的酵母菌，可高效降解谷物多糖^[36]；被孢霉门偏好富氮环境(如家禽高蛋白饲料)，可为宿主累积养分^[37]。SIMPER分析揭示Kazachstania是驱动家禽与野鸟肠道真菌群落分异的核心菌属，该结果进一步佐证了宿主驯化过程通过改变饮食结构实现真菌群落的重塑与转型^[38]。作为典型的发酵型酵母菌，Kazachstania在家禽高谷物饲料的肠道环境中占据优势地位^[36]。八哥与小天鹅和白骨顶鸡肠道真菌群落的差异主要源于Alternaria。该菌广泛定殖于陆生植物体表，其孢子通过野鸟摄食果实进入肠道，而八哥作为典型陆生鸟类进而具有更高的相对丰度^[39]。

基于共现网络的微生物互作分析进一步佐证了宿主类型对肠道菌群内部关系的调控作用。研究表明小天鹅肠道真菌网络复杂度较其他野鸟或家禽最低，这与其特化的植食性摄食策略密切相关。植食性导致Cladosporium在群落中占据主导地位，该菌能够高效降解植物细胞壁，这种功能专一化的群落结构显著降低了种间互作需求，体现出特化食性对菌群互作的简化效应^[13]。小天鹅食性简单，其通过食性特化使菌群功能定向优化，从而提升环境适应性^[40]。基于肠道真菌功能预测结果进一步证实以上结论。小天鹅肠道中植物腐生菌的相对丰度显著高于其他鸟类。研究表明小天鹅通过摄食菹草(Potamogeton crispus)等沉水植物，选择性富集具有纤维素降解能力的真菌类群，协助宿主降解植物细胞壁^[41-42]。基于上述结果揭示植食性小天鹅肠道的高纤维环境通过正向选择维持了高丰度的腐生真菌^[43]。

动物病原菌的群落结构在野鸟与家禽间呈现显著分异。野鸟肠道中动物病原菌和鸟类病原菌的多样性和相对丰度均显著低于家禽，该现象可通过双重机制解释：一方面，野鸟承受极强自然选择作用，通过频繁接触病原体增强自身免疫耐受性，如野鸟肠道通过分泌黏多糖和抗菌肽等物质阻断病原菌定殖^[44]，或靶向破坏病原菌结构抑制病原菌生长^[45]。对抗病原菌能力较弱的野鸟会被自然淘汰，长期过程中野生鸟进化出高强度的肠道过滤作用，筛选特定肠道微生物类群，抑制病原菌定殖^[14]；另一方面养殖场高密度养殖环境导致病原菌暴露强度和交互感染程度显著高于自然生境^[46]。巢湖流域作为我国重要的水产养殖基地与候鸟迁徙通道，家禽粪便中的病原菌可通过水体或气溶胶扩散，威胁濒危水鸟。Wang等^[47]研究发现，IUCN濒危物种鸿雁在养殖密集区因频繁接触养殖废水，其肠道潜在病原菌如黄曲霉等的丰度显著高于自然保护区中的鸟类。此外，野鸟也可以把病原菌传播给家禽甚至人类，造成养殖场家禽大规模死亡和人类流行性疾病暴发^[48]。鉴于此，亟需关注野鸟与家禽病原菌交互传播研究，通过建立病原过滤屏障(如粪污厌氧发酵处理)和候鸟栖息地隔离带切断家禽-野鸟间的病原传播链，维护流域生态安全与公共卫生安全。

4 结论

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本研究系统分析了巢湖流域野鸟与同域家禽肠道真菌及潜在病原菌的群落特征，发现食性和栖息地环境在塑造宿主肠道真菌群落结构中的作用显著高于宿主基因型的影响。家禽由于投喂谷物饲料，肠道中显著富集与谷物降解相关的真菌类群。野鸟肠道具有更强的过滤作用，可降低病原菌定殖，增强环境适应性。野鸟和同域家禽肠道中检测出多种潜在病原菌，暗示亟需关注野鸟与家禽病原菌交互传播研究，进而为评估巢湖流域疾病传播风险及公共卫生安全提供理论依据。

基金

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国家自然科学基金(31801989)
安徽省优秀青年科研项目(2022AH030015)

参考文献

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doi: 10.13343/j.cnki.wsxb.20250597

接收时间：2025-07-31
首发时间：2026-01-12
出版时间：2026-01-04

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收稿日期：2025-07-31
录用日期：2025-10-17

基金

National Natural Science Foundation of China(31801989)

国家自然科学基金(31801989)

Anhui Province Outstanding Youth Research Project(2022AH030015)

安徽省优秀青年科研项目(2022AH030015)

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^1.安徽大学资源与环境工程学院，安徽合肥

^2.湿地生态系统保护与修复安徽省重点实验室，安徽合肥

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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TxT

Treatment	ANOSIM	Treatment	ANOSIM
JY vs. JJ	0.198	0.001	JJ vs. XT	0.947	0.001
JY vs. BG	0.195	0.003	JJ vs. BD	0.897	0.001
JY vs. XT	0.436	0.001	BG vs. XT	0.133	0.025
JY vs. BD	0.439	0.001	BG vs. BD	0.171	0.001
JJ vs. BG	0.723	0.001	XT vs. BD	0.095	0.046

Treatment

ANOSIM

Treatment

ANOSIM

JY vs. JJ

0.198

0.001

JJ vs. XT

0.947

0.001

JY vs. BG

0.195

0.003

JJ vs. BD

0.897

0.001

JY vs. XT

0.436

0.001

BG vs. XT

0.133

0.025

JY vs. BD

0.439

0.001

BG vs. BD

0.171

0.001

JJ vs. BG

0.723

0.001

XT vs. BD

0.095

0.046

Group	Taxonomy	Indicator value	P	Relative abundance (%)	Group	Taxonomy	Indicator value	P	Relative abundance (%)
JY	Candida	0.402	0.011	1.744	XT	Cladosporium	0.379	0.003	11.620
	Cephaliophora	0.596	0.004	1.582		Cystobasidium	0.569	0.002	0.611
	Trichomerium	0.912	0.001	0.918		Preussia	0.410	0.002	0.224
	Wallemia	0.567	0.001	0.504		Neosetophoma	0.495	0.009	0.118
	Moesziomyces	0.464	0.007	0.388		Papiliotrema	0.305	0.036	0.112
	Strelitziana	0.495	0.001	0.233		Coniothyrium	0.449	0.011	0.067
	Neoroussoella	0.307	0.029	0.095		Naganishia	0.314	0.021	0.038
	Keratinophyton	0.270	0.009	0.057		Inopinatum	0.222	0.012	0.030
	Ustilaginoidea	0.358	0.048	0.056		Arxiella	0.336	0.003	0.021
	Cutaneotrichosporon	0.372	0.001	0.030	BD	Curvularia	0.605	0.001	2.658
	Tilletia	0.595	0.001	0.028		Amphobotrys	0.400	0.002	0.950
	Gaertneriomyces	0.288	0.001	0.025		Albifimbria	0.510	0.006	0.645
	Paraconiothyrium	0.350	0.003	0.014		Purpureocillium	0.458	0.019	0.551
	Trichosporon	0.310	0.007	0.014		Rhizophlyctis	0.352	0.009	0.163
	Epichloe	0.284	0.009	0.011		Acremonium	0.345	0.018	0.141
JJ	Kazachstania	0.604	0.001	27.100		Talaromyces	0.441	0.002	0.134
	Diutina	0.584	0.001	2.855		Pseudopithomyces	0.320	0.019	0.133
	Debaryomyces	0.712	0.001	0.669		Neopestalotiopsis	0.293	0.006	0.102
	Botryotrichum	0.578	0.001	0.026		Cryptococcus	0.342	0.014	0.100
BG	Alternaria	0.339	0.026	18.230		Sarocladium	0.361	0.001	0.050
	Geotrichum	0.342	0.024	0.460		Moesziomyces	0.150	0.038	0.043
	Aureobasidium	0.484	0.043	0.455		Paramyrothecium	0.307	0.047	0.039
	Selenophoma	0.392	0.045	0.348		Betamyces	0.246	0.032	0.030
	Meyerozyma	0.337	0.008	0.147		Ustilago	0.332	0.010	0.020
	Mucor	0.349	0.017	0.118		Robillarda	0.155	0.047	0.016
	Buckleyzyma	0.348	0.031	0.057		Emericellopsis	0.230	0.009	0.013
	Coprinopsis	0.200	0.015	0.044

Group

Taxonomy

Indicator value

Relative abundance

(%)

Group

Taxonomy

Indicator value

Relative abundance

(%)

Candida

0.402

0.011

1.744

Cladosporium

0.379

0.003

11.620

Cephaliophora

0.596

0.004

1.582

Cystobasidium

0.569

0.002

0.611

Trichomerium

0.912

0.001

0.918

Preussia

0.410

0.002

0.224

Wallemia

0.567

0.001

0.504

Neosetophoma

0.495

0.009

0.118

Moesziomyces

0.464

0.007

0.388

Papiliotrema

0.305

0.036

0.112

Strelitziana

0.495

0.001

0.233

Coniothyrium

0.449

0.011

0.067

Neoroussoella

0.307

0.029

0.095

Naganishia

0.314

0.021

0.038

Keratinophyton

0.270

0.009

0.057

Inopinatum

0.222

0.012

0.030

Ustilaginoidea

0.358

0.048

0.056

Arxiella

0.336

0.003

0.021

Cutaneotrichosporon

0.372

0.001

0.030

Curvularia

0.605

0.001

2.658

Tilletia

0.595

0.001

0.028

Amphobotrys

0.400

0.002

0.950

Gaertneriomyces

0.288

0.001

0.025

Albifimbria

0.510

0.006

0.645

Paraconiothyrium

0.350

0.003

0.014

Purpureocillium

0.458

0.019

0.551

Trichosporon

0.310

0.007

0.014

Rhizophlyctis

0.352

0.009

0.163

Epichloe

0.284

0.009

0.011

Acremonium

0.345

0.018

0.141

Kazachstania

0.604

0.001

27.100

Talaromyces

0.441

0.002

0.134

Diutina

0.584

0.001

2.855

Pseudopithomyces

0.320

0.019

0.133

Debaryomyces

0.712

0.001

0.669

Neopestalotiopsis

0.293

0.006

0.102

Botryotrichum

0.578

0.001

0.026

Cryptococcus

0.342

0.014

0.100

Alternaria

0.339

0.026

18.230

Sarocladium

0.361

0.001

0.050

Geotrichum

0.342

0.024

0.460

Moesziomyces

0.150

0.038

0.043

Aureobasidium

0.484

0.043

0.455

Paramyrothecium

0.307

0.047

0.039

Selenophoma

0.392

0.045

0.348

Betamyces

0.246

0.032

0.030

Meyerozyma

0.337

0.008

0.147

Ustilago

0.332

0.010

0.020

Mucor

0.349

0.017

0.118

Robillarda

0.155

0.047

0.016

Buckleyzyma

0.348

0.031

0.057

Emericellopsis

0.230

0.009

0.013

Coprinopsis

0.200

0.015

0.044

Group	Taxonomy	Contribution (%)	Group	Taxonomy	Contribution (%)
JY vs. JJ	Kazachstania	35.30	JJ vs. XT	Kazachstania	36.90
	Alternaria	13.70		Alternaria	12.00
	Diutina	5.90		Cladosporium	10.50
	Cephaliophora	3.70		Thelebolus	7.00
	Thelebolus	3.40		Diutina	3.80
JY vs. BG	Kazachstania	22.10	JJ vs. BD	Kazachstania	37.40
	Alternaria	17.00		Cladosporium	7.10
	Cladosporium	7.00		Alternaria	6.70
	Cephaliophora	2.80		Thelebolus	4.90
	Diutina	2.70		Diutina	3.80
JY vs. XT	Kazachstania	21.30	BG vs. XT	Alternaria	17.40
	Alternaria	13.80		Cladosporium	11.60
	Cladosporium	10.90		Thelebolus	7.90
	Thelebolus	7.70		Kazachstania	5.90
	Iodophanus	4.00		Iodophanus	4.50
JY vs. BD	Kazachstania	21.00	BG vs. BD	Alternaria	15.40
	Alternaria	10.70		Cladosporium	7.70
	Cladosporium	6.90		Kazachstania	5.70
	Thelebolus	5.20		Thelebolus	4.90
	Curvularia	3.10		Curvularia	3.50
JJ vs. BG	Kazachstania	36.40	XT vs. BD	Alternaria	13.50
	Alternaria	16.20		Cladosporium	11.00
	Cladosporium	6.90		Thelebolus	10.80
	Diutina	4.50		Iodophanus	5.40
	Aspergillus	1.80		Curvularia	3.70

Group

Taxonomy

Contribution (%)

Group

Taxonomy

Contribution (%)

JY vs. JJ

Kazachstania

35.30

JJ vs. XT

Kazachstania

36.90

Alternaria

13.70

Alternaria

12.00

Diutina

5.90

Cladosporium

10.50

Cephaliophora

3.70

Thelebolus

7.00

Thelebolus

3.40

Diutina

3.80

JY vs. BG

Kazachstania

22.10

JJ vs. BD

Kazachstania

37.40

Alternaria

17.00

Cladosporium

7.10

Cladosporium

7.00

Alternaria

6.70

Cephaliophora

2.80

Thelebolus

4.90

Diutina

2.70

Diutina

3.80

JY vs. XT

Kazachstania

21.30

BG vs. XT

Alternaria

17.40

Alternaria

13.80

Cladosporium

11.60

Cladosporium

10.90

Thelebolus

7.90

Thelebolus

7.70

Kazachstania

5.90

Iodophanus

4.00

Iodophanus

4.50

JY vs. BD

Kazachstania

21.00

BG vs. BD

Alternaria

15.40

Alternaria

10.70

Cladosporium

7.70

Cladosporium

6.90

Kazachstania

5.70

Thelebolus

5.20

Thelebolus

4.90

Curvularia

3.10

Curvularia

3.50

JJ vs. BG

Kazachstania

36.40

XT vs. BD

Alternaria

13.50

Alternaria

16.20

Cladosporium

11.00

Cladosporium

6.90

Thelebolus

10.80

Diutina

4.50

Iodophanus

5.40

Aspergillus

1.80

Curvularia

3.70

Group	JY	JJ	BG	XT	BD
Nodes	313	261	303	208	307
Edges	5 368	3 856	4 536	3 710	4 944
Density	0.055	0.057	0.050	0.086	0.053
Modularity	0.821	0.807	0.864	0.728	0.847
Degree (average)	17.150	14.774	14.970	17.837	16.104
Network diameter	14	13	14	9	11
Clustering coefficient	0.814	0.877	0.842	0.923	0.808
Path length (average)	4.262	5.252	4.946	3.656	4.513
Betweenness centrality (average)	513.170	568.050	525.840	116.465	548.570
Eigenvector centrality (average)	1.000	1.000	0.714	0.016	1.000

Group

Nodes

313

261

303

208

307

Edges

5 368

3 856

4 536

3 710

4 944

Density

0.055

0.057

0.050

0.086

0.053

Modularity

0.821

0.807

0.864

0.728

0.847

Degree (average)

17.150

14.774

14.970

17.837

16.104

Network diameter

Clustering coefficient

0.814

0.877

0.842

0.923

0.808

Path length (average)

4.262

5.252

4.946

3.656

4.513

Betweenness centrality (average)

513.170

568.050

525.840

116.465

548.570

Eigenvector centrality (average)

1.000

0.714

0.016

1.000

Group	Plant saprotroph			Avian pathogen
JY	0.70±0.05a	0.04±0.04b	3.70±1.69a	0.40±0.30a	2.10±0.72a	0.21±0.20b
JJ	0.95±0.62a	0.06±0.07b	3.53±1.17a	0.39±0.22a	2.16±0.83a	0.30±0.18a
BG	0.65±0.74a	0.07±0.08b	2.61±1.54b	0.32±0.28a	1.50±0.76b	0.11±0.09c
XT	0.79±0.80a	0.15±0.18a	0.79±0.70c	0.03±0.06b	0.50±0.19c	0.01±0.01d
BD	0.75±0.72a	0.06±0.06b	1.95±1.36b	0.10±0.10b	1.35±0.93b	0.07±0.06cd

Group

Plant saprotroph

Animal pathogen

Avian pathogen

Diversity

Relative abundance (%)

Diversity

Relative abundance (%)

Diversity

Relative abundance (%)

0.70±0.05a

0.04±0.04b

3.70±1.69a

0.40±0.30a

2.10±0.72a

0.21±0.20b

0.95±0.62a

0.06±0.07b

3.53±1.17a

0.39±0.22a

2.16±0.83a

0.30±0.18a

0.65±0.74a

0.07±0.08b

2.61±1.54b

0.32±0.28a

1.50±0.76b

0.11±0.09c

0.79±0.80a

0.15±0.18a

0.79±0.70c

0.03±0.06b

0.50±0.19c

0.01±0.01d

0.75±0.72a

0.06±0.06b

1.95±1.36b

0.10±0.10b

1.35±0.93b

0.07±0.06cd

Pathogenic species	Symptoms	Infected target
Aspergillus terreus	Respiratory aspergillosis	Human, bird, poultry, etc.
Aspergillus flavus	Invasive aspergillosis	Human, bird, poultry, etc.
Malassezia restricta	Malassezia infection	Human, bird, cattle, etc.
Malassezia globosa	Malassezia infection	Human, bird, dog, horse, etc.
Cladosporium halotolerans	Respiratory tract infection	Human, bird, frog, etc.
Exophiala nishimurae	Phaeohyphomycosis	Human, fish, amphibian, etc.
Exophiala castellanii	Phaeohyphomycosis	Human, fish, etc.
Trichosporon asahii	Trichosporonosis	Human, reptile, etc.
Cyphellophora fusarioides	Onychomycosis	Human
Cyphellophora suttonii	Skin or nail infection	Human

Pathogenic species

Symptoms

Infected target

Aspergillus terreus

Respiratory aspergillosis

Human, bird, poultry, etc.

Aspergillus flavus

Invasive aspergillosis

Human, bird, poultry, etc.

Malassezia restricta

Malassezia infection

Human, bird, cattle, etc.

Malassezia globosa

Malassezia infection

Human, bird, dog, horse, etc.

Cladosporium halotolerans

Respiratory tract infection

Human, bird, frog, etc.

Exophiala nishimurae

Phaeohyphomycosis

Human, fish, amphibian, etc.

Exophiala castellanii

Phaeohyphomycosis

Human, fish, etc.

Trichosporon asahii

Trichosporonosis

Human, reptile, etc.

Cyphellophora fusarioides

Onychomycosis

Human

Cyphellophora suttonii

Skin or nail infection

Human