A checklist of fungi isolated from honey (2000–2022)

Ivo Roxo; António Luís Amaral; António Portugal; João Trovão; Ivo Roxo; António Luís Amaral; António Portugal; João Trovão

doi:10.48130/SIF-2023-0014

2023 Volume 8

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ARTICLE Open Access

A checklist of fungi isolated from honey (2000–2022)

1.
FitoLab—Laboratory for Phytopathology, Instituto Pedro Nunes, Rua Pedro Nunes, Quinta da Nora 3030-199, Coimbra, Portugal
2.
Polytechnic Institute of Coimbra, Coimbra Institute of Engineering, Rua Pedro Nunes, Quinta da Nora 3030-199, Coimbra, Portugal
3.
CEB - Centre of Biological Engineering, Universidade do Minho, Campus de Gualtar 4710-057, Braga, Portugal
4.
LABBELS –Associate Laboratory, Centre of Biological Engineering, Universidade do Minho, Campus de Gualtar 4710-057, Braga/Guimarães, Portugal
5.
Instituto de Investigação Aplicada, Laboratório SiSus, Rua Pedro Nunes, Quinta da Nora 3030-199, Coimbra, Portugal
6.
University of Coimbra, Centre for Functional Ecology - Science for People & the Planet, Associate Laboratory TERRA, Department of Life Sciences, Calçada Martim de Freitas 3000-456, Coimbra, Portugal

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Corresponding author: ivoroxo@sapo.pt

Received: 13 April 2023
Accepted: 29 May 2023
Published online: 02 August 2023
Studies in Fungi 8, Article number: 14 (2023) | Cite this article

Abstract

Mycological studies focusing on fungal species thriving on honey related products have a series of critical applications ranging from the expansion of basic scientific knowledge, the exploration of their industrial utilization, understanding their contributions to food spoilage and even environmental pathogen monitoring. During the last two decades, several works dealing with the isolation and characterization of fungal species thriving on honey have been published. Nonetheless, a thorough summarization of these results has not yet been compiled. This work analyses and compiles a checklist of fungi isolated and identified/described from honey nectar, honey blossom and honeydew between the years 2000 and 2022. Based on this assessment, we detected that over 130 entries have been reported from honey samples worldwide. Consequently, this work provides a checklist of such fungi, that will be of interest to mycologists, microbiologists, food scientists working on the topic, and also beekeepers.
- Apiculture,
- Microfungi,
- Species list,
- Taxonomy,
- Yeasts
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Copyright: © 2023 by the author(s). Published by Maximum Academic Press, Fayetteville, GA. This article is an open access article distributed under Creative Commons Attribution License (CC BY 4.0), visit https://creativecommons.org/licenses/by/4.0/.

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Roxo I, Amaral AL, Portugal A, Trovão J. 2023. A checklist of fungi isolated from honey (2000–2022). Studies in Fungi 8:14 doi: 10.48130/SIF-2023-0014

Roxo I, Amaral AL, Portugal A, Trovão J. 2023. A checklist of fungi isolated from honey (2000–2022). Studies in Fungi 8:14 doi: 10.48130/SIF-2023-0014

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Introduction

Beekeeping (or apiculture) is a zootechnical activity that aims to sustain and exploit, economically and rationally, the domestic bee Apis mellifera, in an effort to obtain their direct products and by-products. Goods such as honey, propolis, pollens, royal jelly, bee bread, waxes and apitoxins (bee venoms) have a wide and diverse range of applications, including in the cosmetic, food, pharmacological and therapeutical industries^[1]. This is noted, for instance, in the expansion of apitherapy (an alternative therapy that uses products coming directly from honeybees) in Western cultures in the last few years^[2]. Moreover, beekeeping is also considered a valuable example of an environmentally sustainable production system, with notorious positive impacts on global biodiversity and agriculture. Overall, this activity is mainly characterized by three advantageous outcomes, namely: (1) a confluence of economic interests (by the production of honey and by-products of the hive, which can provide financial gains), (2) social impact (since it contributes to the fixation of the rural population in territories where other economic activities are hard to be develop or maintain), and (3) a contribution to environmental conservation, sustainability and health (through pollination of cultivated and wild fields)^[3].

According to the European Union and Portuguese legislation (Decree-Law 179/2004) and the Codex Alimentarius (CODEX STAN 12-1981), honey is a natural sweetener produced by honey bees from: (1) flower nectars (blossom honey or nectar honey), or (2) carbohydrate-rich secretions of plants or even excretions of plant-sucking phytophagous aphids (honeydew), after combination with the animal specific molecules, placement, dehydration, and storage in the honey comb (to ripen and mature)^[4−6]. Honey is composed by sugars turned into a super saturated solution containing mainly the monosaccharides (fructose and glucose, in a concentration not lower than 60%) and by a much lesser amount of oligosaccharides, organic acids, enzymes (amylases and α-glucosidase) and colloidal particles^[4].

The quality of honey is mainly determined by its sensorial, chemical, physical and microbiological characteristics. From a microbiological perspective, honey can have two sources of contamination by microorganisms: (1) primary sources: including pollen, the digestive tracts of honey bees, dust, air, soil and nectar (more difficult to control), and (2) secondary sources: arising from human honey manipulation, while also including risks related to air quality, food handlers, cross-contamination and the sanitary state of the equipment and buildings used in the process (more easily controlled by good manufacturing practices)^[7]. And though the European Commission sets maximum levels of mycotoxins for various types of food products, they are often incomplete when considering bee products^[8].

Honey spoilage is not an often-reported phenomenon, mainly due to their associated antimicrobial properties, which result from several different factors. These include contributions from the formation of hydrogen peroxide (H₂O₂), floral source, low pH, low moisture content, low redox potential, low protein content, high osmolarity, high viscosity and limitations to oxygen penetration^[9−12]. On the other hand, an important physicochemical property that can affect the development of microorganisms in honey is the substrate low water activity (a_w), which inhibits the growth of almost all organisms^[12]. Nonetheless, if the moisture content is high enough (above 21%), xerotolerant and xerophile microorganisms can develop causing honey fermentation and spoilage^[4,12]. The microbes of interest in honey are primarily yeasts, fungi and spore-forming bacteria, since their presence can influence the products stability and sanitary quality. Since bees collect pollens and nectars, yeast and fungi presence in honey is unavoidable^[5,7] and examples of filamentous fungi usually found in honey encompass Aspergillus, Penicillium, Mucor and Monascus, along with some osmophile yeasts such as Saccharomyces^[7,13]. Moreover, additional common fungal contaminants of honey are the obligate xerophiles Ascosphaera apis and Bettsia alvei, several xerotolerant species^[4], various species of plant pathogenic fungi^[14], mycotoxin-producing species^[15], and fungi commonly found in pollens and the digestive tract of bees^[16].

During the last two decades, several studies focusing on the isolation and characterization of fungal species thriving in honey have been published. However, a thorough summary of these results has not been compiled so far. With this in mind, the aim of this work is to elicit a checklist of fungi isolated from honey, honey blossom and honeydew, during the time period of 2000 to 2022. As such, this work provides critical information that can be helpful to mycologists, beekeepers and the industrial sector to improve honey and quality and production levels.

Materials and methods

The present checklist is based on a survey of scientific papers using morphological and/or molecular methods to identify fungal taxa isolated from honey, honey blossom and honeydew, during the time period of 2000 to 2022^{[4−7,13,15−29]}. Moreover, the checklist was annotated to contain currently accepted fungal names according to the Index Fungorum (www.indexfungorum.org) to provide an up-to-date analysis and facilitate future knowledge sharing.

Results

Checklist

Table 1. Fungi isolated and identified from honey, honey blossom and honeydew between 2000 and 2022.

Class	Order	Family	Genus		Synonym	Studied substrate:
*Phylum Ascomycota* Caval.-Sm.**
Dothideomycetes O.E. Erikss. & Winka	Capnodiales Woron.	Cladosporiaceae Chalm. & R.G. Archibald	Cladosporium Link	Cladosporium cladosporioides (Fresen.) G.A. de Vries.		Nectar Honey; Honeydew^{[5, 16, 27]}.
				Cladosporium sp. Link.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18, 23, 25−26].}
	Dothideales Lindau	Saccotheciaceae Bonord.	Aureobasidium Viala & G. Boyer	Aureobasidium pullulans (de Bary & Löwenthal) G. Arnaud.		Nectar Honey; Honeydew^[5].
				Aureobasidium sp. Viala & G. Boyer.		Honey Blossom; Honeydew; Nectar Honey^{[23, 26]}.
	Incertae sedis	Seuratiaceae Vuill. ex M.E. Barr	Atichia Flot.	Atichia sp. Flot..		Honey Blossom; Honeydew^[18].
			Seuratia Pat.	Seuratia sp. Pat..		Honey Blossom; Honeydew^[18].
	Mytilinidiales E. Boehm, C.L. Schoch & Spatafora	Mytilinidiaceae Kirschst.	Peyronelia Cif. & Gonz. Frag.	Peyronelia sp. Cif. & Gonz. Frag..		Honey Blossom; Honeydew^[18].
	leosporales Luttr. ex M.E. Barr	Pleosporaceae Nitschke	Alternaria Nees	Alternaria alternata (Fr.) Keissl..		Nectar Honey; Honeydew^{[5, 16]}.
				Alternaria multiformis (E.G. Simmons) Woudenb. & Crous.		Honey Blossom; Honeydew^[4].
				Alternaria sp. Nees.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18, 23, 26]}.
			Stemphylium Wallr.	Stemphylium sp. Wallr..		Honey Blossom^[26].
		Didymellaceae Gruyter, Aveskamp & Verkley	Epicoccum Link	Epicoccum sp. Link.		Honey Blossom; Honeydew^{[15, 18]}.
			Phoma Sacc.	Phoma sp. Sacc..		Honey Blossom; Honeydew^{[15, 18]}.
		Torulaceae Corda	Torula Pers.	Torula mellis Fabian & Quinet.		Honey Blossom; Honeydew^[18].
				Torula sp. Pers..		Honey Blossom; Honeydew^[18].
Eurotiomycetes O.E. Erikss. & Winka	Ascosphaerales Gäum. ex Benny & Kimbr.	Ascosphaeraceae L.S. Olive & Spiltoir	Ascosphaera L.S. Olive & Spiltoir	Ascosphaera atra Skou & K. Hackett.		Honey Blossom; Honeydew^[4].
				Ascosphaera sp. L.S. Olive & Spiltoir.		Honey Blossom; Honeydew^[18].
			Bettsia Skou	Bettsia alvei (Betts) Skou ex Pitt, Lantz, Pettersson & Leong.		Honey Blossom; Honeydew^{[4; 18]}.
	Chaetothyriales M.E. Barr	Cyphellophoraceae Réblová & Unter.	Cyphellophora G.A. de Vries	Cyphellophora jingdongensis H. Yang & K.D. Hyde.		Nectar Honey^[24].
	Eurotiales G.W. Martin ex Benny & Kimbr.	Aspergillaceae Link	Aspergillus P. Micheli ex Haller.	Aspergillus discophorus Samson, Zalar & Frisvad.	Emericella discophora.	Nectar Honey; Honeydew^[5].
				Aspergillus nidulans (Eidam) G. Winter.	Emericella nidulans.	Nectar Honey^[27].
				Aspergillus qinqixianii Y. Horie, Abliz & R.Y. Li.	Emericella qinqixianii.	Nectar Honey; Honeydew^[5].
				Aspergillus asperescens Stolk.		Honey Blossom; Honeydew^[4].
				Aspergillus candidus Link.		Nectar Honey; Honeydew^{[13, 16, 27]}.
				Aspergillus clavatus Desm..		Honey Blossom; Honeydew^[15].
				Aspergillus flavus Link.		Honey Blossom; Honeydew; Nectar Honey^{[13, 15−16, 27]}.
				Aspergillus fumigatus Fresen..		Honey Blossom; Honeydew; Nectar Honey^{[7, 13, 15−16, 27]}.
				Aspergillus montevidensis Talice & J.A. Mackinnon.		Honey Blossom; Honeydew^[4].
				Aspergillus niger Tiegh..		Honey Blossom; Honeydew; Nectar Honey^{[5, 7, 13, 15−16, 27]}.
				Aspergillus ochraceus G. Wilh..		Honey Blossom; Honeydew; Nectar Honey^[15−16].
				Aspergillus proliferans G. Sm..		Nectar Honey; Honeydew^[5].
				Aspergillus pseudoglaucus Blochwitz.		Honey Blossom; Honeydew^[4].
				Aspergillus sp. P. Micheli ex Haller.	Emericella sp. and Eurotium sp..	Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18, 25−26]}.
				Aspergillus spelunceus Raper & Fennell.		Nectar Honey; Honeydew^[5].
				Aspergillus terreus Thom.		Nectar Honey^[16].
				Aspergillus versicolor (Vuill.) Tirab..		Honey Blossom; Honeydew; Nectar Honey^{[15−16, 27]}.
			Paecilomyces Bainier	Paecilomyces sp. Bainier.		Nectar Honey; Honey Blossom^[23].
			Penicillium Link	Penicillium italicum Wehmer.		Nectar Honey; Honeydew^[5].
				Penicillium solitum Westling.	Penicillium crustosum.	Honey Blossom; Honeydew^[18].
				Penicillium apimei R.N. Barbosa, Souza-Motta, N.T. Oliveira & Houbraken.		Nectar Honey^[21].
				Penicillium aurantiogriseum Dierckx.		Honey Blossom; Honeydew^[15].
				Penicillium brevicompactum Dierckx.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18]}.
				Penicillium brocae S.W. Peterson, Jeann. Pérez, F.E. Vega & Infante.		Nectar Honey^[21].
				Penicillium camemberti Thom.		Honey Blossom; Honeydew^[4].
				Penicillium chrysogenum Thom.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18]}.
				Penicillium citrinum Thom.		Honey Blossom; Honeydew; Nectar Honey^{[4, 15, 21]}.
				Penicillium commune Thom.		Nectar Honey^[7].
				Penicillium corylophilum Dierckx.		Honey Blossom; Honeydew; Nectar Honey^{[4,5, 7, 15, 27]}.
				Penicillium cravenianum Visagie & K. Jacobs.		Honey Blossom; Honeydew^[4].
				Penicillium decumbens Thom.		Nectar Honey; Honeydew^[5].
				Penicillium echinulatum Biourge.		Nectar Honey; Honeydew^[5].
				Penicillium expansum Link.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15]}.
				Penicillium griseofulvum Dierckx.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15, 18]}.
				Penicillium sp. Link.		Honey Blossom; Honeydew; Nectar Honey^{[5, 13, 15,16, 18, 21, 23, 25,26]}.
				Penicillium meliponae R.N. Barbosa, Souza-Motta, N.T. Oliveira & Houbraken.		Nectar Honey^[21].
				Penicillium mellis R.N. Barbosa, Souza-Motta, N.T. Oliveira & Houbraken.		Nectar Honey^[21].
				Penicillium polonicum K.W. Zaleski.		Nectar Honey; Honeydew^{[5, 7]}.
				Penicillium raistrickii G. Sm..		Honey Blossom; Honeydew^[15].
				Penicillium sclerotiorum J.F.H. Beyma.		Nectar Honey^[21].
				Penicillium wotroi Houbraken, López-Quint., Frisvad & Samson.		Nectar Honey^[21].
			Talaromyces C.R. Benj.	Talaromyces affinitatimellis Rodr.-Andr., Stchigel & Cano.		Honey Blossom; Honeydew^[4].
				Talaromyces basipetosporus Stchigel, Cano & Rodr.-Andr..		Honey Blossom; Honeydew^[4].
				Talaromyces brunneosporus Rodr.-Andr., Cano & Stchigel.		Honey Blossom; Honeydew^[4].
				Talaromyces funiculosus (Thom) Samson, N. Yilmaz, Frisvad & Seifert.	Penicillium funiculosum.	Nectar Honey^[27].
				Talaromyces brasiliensis R.N. Barbosa, Souza-Motta, N.T. Oliveira & Houbraken.		Nectar Honey^[21].
				Talaromyces scorteus (Nakaz., Y. Takeda & Suematsu) S.W. Peterson & Jurjević.		Nectar Honey^[21].
			Xerochrysium Pitt	Xerochrysium xerophilum (Pitt) Pitt.		Honey Blossom; Honeydew^[4].
		Monascaceae J. Schröt.	Monascus Tiegh.	Monascus mellicola R.N. Barbosa, Souza-Motta, N.T. Oliveira & Houbraken.		Nectar Honey^[21].
				Monascus pilosus K. Satô ex D. Hawksw. & Pitt.		Honey Blossom; Honeydew^[4].
				Monascus purpureus Went.		Honey Blossom; Honeydew^[4].
				Monascus ruber Tiegh..		Honey Blossom; Honeydew^[4].
	Incertae sedis	Eremascaceae Engl. & E. Gilg	Eremascus Eidam	Eremascus albus Eidam.		Honey Blossom; Honeydew^[4].
	Onygenales Cif. ex Benny & Kimbr.	Helicoarthrosporaceae Stchigel, Rodr.-Andrade & Cano	Helicoarthrosporum Stchigel, Cano & Rodríguez-Andrade	Helicoarthrosporum mellicola Stchigel, Cano & Rodríguez-Andrade.		Honey Blossom; Honeydew^[4].
		Incertae sedis	Strongyloarthrosporum Rodr.-Andr., Cano & Stchigel	Strongyloarthrosporum catenulatum Rodr.-Andr., Cano & Stchigel.		Honey Blossom; Honeydew^[4].
Leotiomycetes O.E. Erikss. & Winka	Helotiales Nannf.	Incertae sedis	Coniothecium Corda	Coniothecium sp. Corda.		Honey Blossom; Honeydew^[18].
			Triposporium Corda	Triposporium sp. Corda.		Honey Blossom; Honeydew^[18].
		Myxotrichaceae Locq. ex Currah	Oidiodendron Robak	Oidiodendron mellicola Rodr.-Andr., Cano & Stchigel.		Honey Blossom; Honeydew^[4].
			Skoua A.A. Wynns	Skoua asexualis Rodr.-Andr., Cano & Stchigel.		Honey Blossom; Honeydew^[4].
				Skoua fertilis (Stoppel) A.A. Wynns.		Honey Blossom; Honeydew^[4].
		Sclerotiniaceae Whetzel	Botrytis Micheli ex Pers.	Botrytis sp. P. Micheli ex Pers..		Honey Blossom; Honeydew^{[15, 18]}.
Saccharomycetes G. Winter	Incertae sedis	Incertae sedis	Oosporidium Stautz	Oosporidium sp. Stautz.		Honey Blossom; Honeydew^[18].
	Saccharomycetales Luerss.	Incertae sedis	Candida Berkhout	Candida sp. Berkhout.		Honey Blossom; Honeydew^[18].
				Candida lundiana Saks., M. Suzuki, Lumyong, Ohkuma & Chantaw.		Nectar Honey^[29].
				Candida parapsilosis (Ashford) Langeron & Talice.		Nectar Honey^[6].
				Candida suthepensis Saks., M. Suzuki, Lumyong, Ohkuma & Chantaw..		Nectar Honey^[29].
			Starmerella C.A. Rosa & Lachance	Starmerella magnoliae (Lodder & Kreger-van Rij) C.A. Rosa & Lachance.	Candida magnoliae.	Honey Blossom; Honeydew; Nectar Honey^{[4, 6]}.
				Starmerella sorbosivorans (S.A. James, C.J. Bond & I.N. Roberts) C.A. Rosa & Lachance.	Candida sorbosivorans.	Honey Blossom; Honeydew; Nectar Honey^{[4, 6]}.
		Lipomycetaceae E.K. Novák & Zsolt	Lipomyces Lodder & Kreger-van Rij	Lipomyces sp. Lodder & Kreger-van Rij.		Honey Blossom; Honeydew^[18].
		Saccharomycetaceae G.Winter	Debaryomyces Klöcker	Debaryomyces hansenii (Zopf) Lodder & Kreger-van Rij.		Nectar Honey; Honeydew^[5].
			Pichia E.C. Hansen	Pichia sp. E.C. Hansen.		Honey Blossom; Honeydew^[18].
				Pichia membranifaciens (E.C. Hansen) E.C. Hansen.		Nectar Honey^[6].
			Saccharomyces Meyen	Saccharomyces sp. Meyen.		Honey Blossom; Honeydew; Nectar Honey^{[13, 18]}.
				Saccharomyces cerevisiae (Desm.) Meyen.		Honey Blossom; Honeydew; Nectar Honey^{[6, 18]}.
			Schwanniomyces Klöcker	Schwanniomyces sp. Klöcker.		Honey Blossom; Honeydew^[18].
			Zygosaccharomyces B.T.P. Barker	Zygosaccharomyces favi G. Péter, Čadež & Dlauchy.		Nectar Honey^[17].
				Zygosaccharomyces gambellarensis Torriani, M. Lorenzini, Salvetti & Felis.		Honey Blossom; Honeydew^[4].
				Zygosaccharomyces mellis Fabian & Quinet.		Honey Blossom; Honeydew; Nectar Honey^{[4−6, 18]}.
				Zygosaccharomyces priorianus Klöcker.		Honey Blossom; Honeydew^[18].
				Zygosaccharomyces rouxii (Boutroux) Yarrow.		Honey Blossom; Honeydew; Nectar Honey^{[6, 18]}.
				Zygosaccharomyces siamensis Saks., M. Suzuki, Chantaw., Ohkuma & Lumyong.		Honey Blossom; Honeydew; Nectar Honey^{[4, 29]}.
				Zygosaccharomyces sp. B.T.P. Barker.		Honey Blossom; Honeydew^[18].
		Trichomonascaceae Kurtzman & Robnett	Blastobotrys Klopotek	Blastobotrys meliponae R.N. Barbosa, Boekhout, G.A. Silva, Souza-Motta & N. Oliveira.		Nectar Honey; Honeydew^[19].
		Wickerhamomycetaceae Kurtzman, Robnett & Bas.-Powers	Wickerhamomyces Kurtzman, Robnett & Bas.-Powers	Wickerhamomyces sp. Kurtzman, Robnett & Bas.-Powers.	Hansenula sp..	Honey Blossom; Honeydew^[18].
Schizosaccharomycetes O.E. Erikss. & Winka	Schizosaccharomycetales O.E. Erikss.	Schizosaccharomycetaceae Beij. ex Klöcker	Schizosaccharomyces Lindner	Schizosaccharomyces sp. Lindner.		Honey Blossom; Honeydew^[18].
				Schizosaccharomyces octosporus Beij.		Honey Blossom; Honeydew^[18].
Sordariomycetes O.E. Erikss. & Winka	Hypocreales Lindau	Cordycipitaceae Kreisel ex G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora	Engyodontium de Hoog	Engyodontium sp. de Hoog.		Nectar Honey^[25].
		Hypocreaceae De Not.	Trichoderma Pers.	Trichoderma hamatum (Bonord.) Bainier.		Nectar Honey^[27].
				Trichoderma sp. Pers..		Honey Blossom; Honeydew^{[15, 18]}.
		Incertae sedis	Acremonium Link	Acremonium sp. Link.		Honey Blossom; Honeydew; Nectar Honey^{[16, 18]}.
			Sarocladium W. Gams & D. Hawksw.	Sarocladium strictum (W. Gams) Summerb..	Acremonium strictum.	Nectar Honey^[27].
				Sarocladium sp. W. Gams & D. Hawksw..	Cephalosporium sp..	Honey Blossom; Honeydew^[18].
		Nectriaceae Tul. & C. Tul.	Fusarium Link	Fusarium sp. Link.		Honey Blossom; Honeydew; Nectar Honey^{[7, 15−16, 18]}.
				Fusarium oxysporum Schltdl.		Nectar Honey^[27].
		Stachybotryaceae L. Lombard & Crous	Stachybotrys Corda	Stachybotrys sp. Corda.		Nectar Honey; Honey Blossom^[23].
	Incertae sedis	Apiosporaceae K.D. Hyde, J. Fröhl., Joanne E. Taylor & M.E. Barr	Arthrinium Kunze	Arthrinium sp. Kunze.		Honey Blossom; Honeydew; Nectar Honey^{[5, 26]}.
	Sordariales Chadef. ex D. Hawksw. & O.E. Erikss.	Chaetomiaceae G. Winter	Botryotrichum Sacc. & Marchal	Botryotrichum atrogriseum J.F.H. Beyma.		Nectar Honey^[27].
			Chaetomium Kunze	Chaetomium globosum Kunze.		Nectar Honey; Honeydew^[5].
				Chaetomium sp. Kunze.		Honey Blossom; Honeydew^[18]
			Trichocladium Harz	Trichocladium griseum (Traaen) X. Wei Wang & Houbraken.
	Xylariales Nannf.	Hypoxylaceae DC.	Daldinia Ces. & De Not.	Daldinia concentrica (Bolton) Ces. & De Not..		Nectar Honey; Honeydew^[5].
Phylum Basidiomycota R.T. Moore
Microbotryomycetes R. Bauer, Begerow, J.P. Samp., M. Weiss & Oberw.	Sporidiobolales Doweld	Sporidiobolaceae R.T. Moore	Rhodotorula F.C. Harrison	Rhodotorula sp. F.C. Harrison.		Honey Blossom; Honeydew^[18].
				Rhodotorula mucilaginosa (A. Jörg.) F.C. Harrison		Nectar Honey^[6].
Tremellomycetes Doweld	Filobasidiales Jülich	Filobasidiaceae L.S. Olive	Naganishia Goto	Naganishia uzbekistanensis (Á. Fonseca, Scorzetti; Fell) Xin Zhan Liu, F.Y. Bai, M. Groenew. & Boekhout.	Cryptococcus uzbekistanensis.	Nectar Honey; Honeydew^[5].
	Tremellales Fr.	Cryptococcaceae Kütz. ex Castell. & Chalm.	Cryptococcus Vuill.	Cryptococcus neoformans (San Felice) Vuill..	Cryptococcus neoformans var. grubii.	Honey Blossom; Honeydew^[18].
	Trichosporonales Boekhout & Fell	Trichosporonaceae Nann.	Cutaneotrichosporon Xin Zhan Liu, F.Y. Bai, M. Groenew. & Boekhout	Cutaneotrichosporon mucoides (E. Guého; M.T. Sm.) Xin Zhan Liu, F.Y. Bai, M. Groenew. & Boekhout.		Nectar Honey^[6].
			Vanrija R.T. Moore	Vanrija humicola (Dasz.) R.T. Moore.		Nectar Honey^[6].
Ustilaginomycetes Warm.	Ustilaginales Bek.	Ustilaginaceae Tul. & C. Tul.	Not determined.	Ustilaginaceae sp. Tul. & C. Tul..		Honey Blossom; Honeydew^[18]
Wallemiomycetes Zalar, de Hoog & Schroers	Wallemiales Zalar, de Hoog & Schroers	Wallemiaceae R.T. Moore	Wallemia Johan-Olsen	Wallemia hederae S. Jančič, Zalar & Gunde Cimerman..		Nectar Honey^[19].
				Wallemia mellicola Jancic, Nguyen, Seifert & Gunde-Cimerman..		Nectar Honey^[19].
Phylum Mucoromycota Doweld
Mucoromycetes Doweld	Mucorales Dumort.	Cunninghamellaceae Naumov ex R.K. Benj.	Cunninghamella Matr.	Cunninghamella elegans Lendn..	Cunninghamella bertholletiae.	Honey Blossom; Honeydew^[4].
		Mucoraceae Fr.	Mucor P. Micheli ex L.	Mucor hiemalis Wehmer..		Nectar Honey^[16].
				Mucor sp. P. Micheli ex L..		Honey Blossom; Honeydew; Nectar Honey^{[13, 15, 18, 23, 26]}.
				Mucor plumbeus Bonord..		Honey Blossom; Honeydew^[4].
				Mucor racemosus Bull..		Nectar Honey^[16].
			Rhizopus Ehrenb.	Rhizopus arrhizus A. Fisch..	Rhizopus oryzae.	Honey Blossom; Honeydew^[4]
				Rhizopus sp. Ehrenb..		Nectar Honey^[7].
				Rhizopus stolonifer (Ehrenb.) Vuill..		Nectar Honey^[16].
Peronosporaceae sp. de Bary and Pythium sp. Pringsh., have also been isolated, identified and noted as fungi, from Honey Blossom and Honeydew samples^[18].

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A checklist of fungi isolated from honey (2000–2022)

Abstract