The fungal kingdom is vast, spanning ∼1.5 to as many as 5 million species diverse as unicellular yeasts, filamentous fungi, mushrooms, lichens, and both plant and animal pathogens. The fungi are closely aligned with animals in one of the six to eight supergroups of eukaryotes, the opisthokonts. The animal and fungal kingdoms last shared a common ancestor ∼1 billion years ago, more recently than other groups of eukaryotes. As a consequence of their close evolutionary history and shared cellular machinery with metazoans, fungi are exceptional models for mammalian biology, but prove more difficult to treat in infected animals. The last common ancestor to the fungal/metazoan lineages is thought to have been unicellular, aquatic, and motile with a posterior flagellum, and certain extant species closely resemble this hypothesized ancestor. Species within the fungal kingdom were traditionally assigned to four phyla, including the basal fungi (Chytridiomycota, Zygomycota) and the more recently derived monophyletic lineage, the dikarya (Ascomycota, Basidiomycota). The fungal tree of life project has revealed that the basal lineages are polyphyletic, and thus there are as many as eight to ten fungal phyla. Fungi that infect vertebrates are found in all of the major lineages, and virulence arose multiple times independently. A sobering recent development involves the species Batrachochytrium dendrobatidis from the basal fungal phylum, the Chytridiomycota, which has emerged to cause global amphibian declines and extinctions. Genomics is revolutionizing our view of the fungal kingdom, and genome sequences for zygomycete pathogens (Rhizopus, Mucor), skin-associated fungi (dermatophytes, Malassezia), and the Candida pathogenic species clade promise to provide insights into the origins of virulence. Here we survey the diversity of fungal pathogens and illustrate key principles revealed by genomics involving sexual reproduction and sex determination, loss of conserved pathways in derived fungal lineages that are retained in basal fungi, and shared and divergent virulence strategies of successful human pathogens, including dimorphic and trimorphic transitions in form. The overarching conclusion is that fungal pathogens of animals have arisen repeatedly and independently throughout the fungal tree of life, and while they share general properties, there are also unique features to the virulence strategies of each successful microbial pathogen.

真菌王国非常庞大，涵盖约 1.5 至 500 万种物种，包括单细胞酵母、丝状真菌、蘑菇、地衣以及植物和动物病原体。这些真菌与六到八个真核生物超类群之一的动物（后康生物）密切相关。动物和真菌界最后一次拥有共同的祖先是在十亿年前，比其他真核生物群体的时间要晚。由于其密切的进化历史以及与后生动物共享的细胞机制，真菌是哺乳动物生物学的特殊模型，但事实证明在受感染的动物中治疗起来更加困难。真菌/后生动物谱系的最后一个共同祖先被认为是单细胞的、水生的、具有后鞭毛的能动的，并且某些现存物种与这个假设的祖先非常相似。真菌界内的物种传统上被划分为四个门，包括基础真菌（壶菌门、接合菌门）和最近衍生的单系谱系，双核真菌（子囊菌门、担子菌门）。真菌生命树项目揭示了基础谱系是多系的，因此真菌门有八到十个之多。感染脊椎动物的真菌存在于所有主要谱系中，并且毒力独立出现多次。最近一项发人深省的发展涉及来自基底真菌门壶菌门的Batrachochytrium dendrobatidis物种，它的出现导致了全球两栖动物的减少和灭绝。 基因组学正在彻底改变我们对真菌界的看法，接合菌病原体（根霉、毛霉）、皮肤相关真菌（皮肤癣菌、马拉色菌）和念珠菌致病物种进化枝的基因组序列有望提供对毒力起源的见解。在这里，我们调查了真菌病原体的多样性，并说明了基因组学揭示的关键原理，涉及有性生殖和性别决定、基础真菌中保留的衍生真菌谱系中保守途径的丧失，以及成功的人类病原体的共享和不同的毒力策略，包括二态性以及形式上的三态转变。总体结论是，动物的真菌病原体在整个真菌生命树中反复且独立地出现，虽然它们具有共同的特性，但每种成功的微生物病原体的毒力策略也有独特的特征。