Danny Haelewaters’s research while affiliated with University of Colorado Boulder and other places

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Publications (195)


Fungal diversity and function in metagenomes sequenced from extreme environments
  • Article
  • Full-text available

December 2024

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52 Reads

Fungal Ecology

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Dongying Wu

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Fungi are increasingly recognized as key players in various extreme environments. Here we present an analysis of publicly-sourced metagenomes from global extreme environments, focusing on fungal taxonomy and function. The majority of 855 selected metagenomes contained scaffolds assigned to fungi. Relative abundance of fungi was as high as 10% of protein-coding genes with taxonomic annotation, with up to 289 fungal genera per sample. Despite taxonomic clustering by environment, fungal communities were more dissimilar than archaeal and bacterial communities, both for within-and between-environment comparisons. Relatively abundant fungal classes in extreme environments included Dothideomycetes, Eurotiomycetes, Leotiomycetes, Pezizomycetes, Saccharomycetes, and Sordariomycetes. Broad generalists and prolific aerial spore formers were the most relatively abundant fungal genera detected in most of the extreme environments, bringing up the question of whether they are actively growing in those environments or just surviving as spores. More specialized fungi were common in some environments, such as zoosporic taxa in cryosphere water and hot springs. Relative abundances of genes involved in adaptation to general, thermal, oxidative, and osmotic stress were greatest in soda lake, acid mine drainage, and cryosphere water samples.

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Fig. 1 MycoBank.org data on tropical and non-tropical fungal name depositions. Cumulative counts of fungal species name depositions in Mycobank.org from tropical and non-tropical locations, and annual percentage of tropical fungal names deposited in Mycobank from 1823 to November 2020
Fig. 2 New, unusual, and endemic tropical taxa from the UPRS. Clavulina craterelloides, unique morphology for this genus a, Meredithblackwellia eburnea, new genus b, Craterellus pleurotoides, unique morphology for this genus c, Pseudotulostoma volvata, new genus d, Guyanagaster necrorhizus, new genus e1-2, Fusarium xyrophilum, new life history strategy for this genus f, Guyanagarika pakaraimensis, new genus g, Hygrocybe sp., unique morphology for this genus h. Scale bars a, d-h 2 cm; b 10 µm; c 1 cm
Fig. 4 IUCN Red List data on tropical fungi, plants, and animals. Cumulative percent of IUCN evaluations for species of fungi, plants, and animals from tropical habitats a and percent fungi, plants, and animals considered threatened or data deficient from tropical and non-tropical locations b through June 2023
The contribution of tropical long-term studies to mycology

November 2024

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232 Reads

IMA Fungus

Fungi are arguably the most diverse eukaryotic kingdom of organisms in terms of number of estimated species, trophic and life history strategies, and their functions in ecosystems. However, our knowledge of fungi is limited due to a distributional bias; the vast majority of available data on fungi have been compiled from non-tropical regions. Far less is known about fungi from tropical regions, with the bulk of these data being temporally limited surveys for fungal species diversity. Long-term studies (LTS), or repeated sampling from the same region over extended periods, are necessary to fully capture the extent of species diversity in a region, but LTS of fungi from tropical regions are almost non-existent. In this paper, we discuss the contributions of LTS of fungi in tropical regions to alpha diversity, ecological and functional diversity, biogeography, hypothesis testing, and conservation—with an emphasis on an ongoing tropical LTS in the Pakaraima Mountains of Guyana. We show how these contributions refine our understanding of Fungi. We also show that public data repositories such as NCBI, IUCN, and iNaturalist contain less information on tropical fungi compared to non-tropical fungi, and that these discrepancies are more pronounced in fungi than in plants and animals.


‘Invisible and uncharismatic’ fungi need taxonomy champions, too

November 2024

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19 Reads


Classes and phyla of the kingdom Fungi

October 2024

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1,759 Reads

Fungal Diversity

Fungi are one of the most diverse groups of organisms with an estimated number of species in the range of 2–3 million. The higher-level ranking of fungi has been discussed in the framework of molecular phylogenetics since Hibbett et al., and the definition and the higher ranks (e.g., phyla) of the ‘true fungi’ have been revised in several subsequent publications. Rapid accumulation of novel genomic data and the advancements in phylogenetics now facilitate a robust and precise foundation for the higher-level classification within the kingdom. This study provides an updated classification of the kingdom Fungi , drawing upon a comprehensive phylogenomic analysis of Holomycota , with which we outline well-supported nodes of the fungal tree and explore more contentious groupings. We accept 19 phyla of Fungi, viz . Aphelidiomycota , Ascomycota , Basidiobolomycota , Basidiomycota , Blastocladiomycota , Calcarisporiellomycota , Chytridiomycota , Entomophthoromycota , Entorrhizomycota , Glomeromycota , Kickxellomycota , Monoblepharomycota , Mortierellomycota , Mucoromycota , Neocallimastigomycota , Olpidiomycota , Rozellomycota , Sanchytriomycota, and Zoopagomycota . In the phylogenies, Caulochytriomycota resides in Chytridiomycota ; thus, the former is regarded as a synonym of the latter, while Caulochytriomycetes is viewed as a class in Chytridiomycota . We provide a description of each phylum followed by its classes. A new subphylum, Sanchytriomycotina Karpov is introduced as the only subphylum in Sanchytriomycota . The subclass Pneumocystomycetidae Kirk et al. in Pneumocystomycetes , Ascomycota is invalid and thus validated. Placements of fossil fungi in phyla and classes are also discussed, providing examples.


Recent fieldwork and fungarium studies double known diversity of Chlorosplenium and improve understanding of species distributions

August 2024

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78 Reads

Chlorosplenium is a small genus comprising five species of inoperculate discomycetes in the order Helotiales (Leotiomycetes) often recognizable by their bright yellowish-green colors and gregarious growth on wood. In this study, we describe five new species-C. aotearoa, C. australiense, C. cusucoense, C. epimorsicum, and C. hawaiiense-based on a combination of recent fieldwork and examination of previously collected fungarium specimens. We use an integrative taxonomic approach to support the distinction of new species, incorporating morphology and DNA sequence data with biogeography. Macro- and micromorphological features of apothecia for all species and culture characteristics for four of the five new species are documented. A multilocus phylogeny based on nuc rDNA internal transcribed spacer region ITS1-5.8S-ITS2, partial large subunit nuc ribosomal DNA (28S nuc rDNA), and A-B regions of the largest subunit of RNA polymerase II (RPB1) gene is presented. Additionally, we report Chlorosplenium chlora from Europe for the first time and expand our knowledge of the diversity and distributions of species in this genus in America, Australia, and New Zealand.




Phylogenomics, divergence times and notes of orders in Basidiomycota

July 2024

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2,147 Reads

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10 Citations

Fungal Diversity

Basidiomycota is one of the major phyla in the fungal tree of life. The outline of Basidiomycota provides essential taxonomic information for researchers and workers in mycology. In this study, we present a time-framed phylogenomic tree with 487 species of Basidiomycota from 127 families, 47 orders, 14 classes and four subphyla; we update the outline of Basidiomycota based on the phylogenomic relationships and the taxonomic studies since 2019; and we provide notes for each order and discuss the history, defining characteristics, evolution, justification of orders, problems, significance, and plates. Our phylogenomic analysis suggests that the subphyla diverged in a time range of 443–490 Myr (million years), classes in a time range of 312–412 Myr, and orders in a time range of 102–361 Myr. Families diverged in a time range of 50–289 Myr, 76–224 Myr, and 62–156 Myr in Agaricomycotina, Pucciniomycotina, and Ustilaginomycotina, respectively. Based on the phylogenomic relationships and divergence times, we propose a new suborder Mycenineae in Agaricales to accommodate Mycenaceae. In the current outline of Basidiomycota, there are four subphyla, 20 classes, 77 orders, 297 families, and 2134 genera accepted. When building a robust taxonomy of Basidiomycota in the genomic era, the generation of molecular phylogenetic data has become relatively easier. Finding phenotypical characters, especially those that can be applied for identification and classification, however, has become increasingly challenging.


New insights into the DNA extraction and PCR amplification of minute ascomycetes in the genus Laboulbenia (Pezizomycotina, Laboulbeniales)

June 2024

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61 Reads

IMA Fungus

Molecular studies of fungi within the order Laboulbeniales (Ascomycota, Pezizomycotina) have been hampered for years because of their minute size, inability to grow in axenic culture, and lack of reliable and cost-efficient DNA extraction protocols. In particular, the genus Laboulbenia is notorious for low success with DNA extraction and polymerase chain reaction (PCR) amplification. This is attributed to the presence of melanin, a molecule known to inhibit PCR, in the cells. We evaluated the efficacy of a standard single cell-based DNA extraction protocol by halving the recommended amount of reagents to reduce the cost per extraction and adding bovine serum albumin (BSA) during the multiple displacement amplification step to reverse the effect of melanin. A total of 196 extractions were made, 111 of which were successful. We found that halving the reagents used in the single cell-based extraction kit did not significantly affect the probability of successful DNA extraction. Using the halved protocol reduces cost and resource consumption. Moreover, there was no significant difference in the probability of successfully extracting DNA based on whether BSA was added or not, suggesting that the amount of melanin present in cells of the thallus has no major inhibitory effect on PCR. We generated 277 sequences from five loci, but amplification of the internal transcribed spacer region, the mitochondrial small subunit rDNA, and protein-coding genes remains challenging. The probability of successfully extracting DNA from Laboulbeniales was also impacted by specimen storage methods, with material preserved in > 95% ethanol yielding higher success rates compared to material stored in 70% ethanol and dried material. We emphasize the importance of proper preservation of material and propose the design of Laboulbeniales-specific primers to overcome the problems of primer mismatches and contaminants. Our new insights apply not only to the genus Laboulbenia; Laboulbeniales generally are understudied, and the vast majority of species remain unsequenced. New and approachable molecular developments will benefit the study of Laboulbeniales, helping to elucidate the true diversity and evolutionary relationships of these peculiar microfungi.


Annual data on observations, observers, and species in nine organismal groups on iNaturalist.org from 2008 to 2022: (a) total observers; (b) total observations; (c) total unique species observed; (d) average number of observations by observer; (e) percentage of total described species observed; (f) percentage of total estimated species observed. Total described and estimated species in (e) and (f) derived from https://www.dcceew.gov.au/science‐research/abrs/publications/other/numbers‐living‐species/executive‐summary.
Number of species assessed on the IUCN Red List of Threatened Species (IUCN, 2023): (a) assessments completed for three eukaryotic kingdoms; (b) assessments completed for the kingdom Fungi. Data taken from https://www.iucnredlist.org/resources/summary‐statistics.
The power of citizen science to advance fungal conservation

March 2024

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611 Reads

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13 Citations

Fungal conservation is gaining momentum globally, but many challenges remain. To advance further, more data are needed on fungal diversity across space and time. Fundamental information regarding population sizes, trends, and geographic ranges is also critical to accurately assess the extinction risk of individual species. However, obtaining these data is particularly difficult for fungi due to their immense diversity, complex and problematic taxonomy, and cryptic nature. This paper explores how citizen science (CS) projects can be leveraged to advance fungal conservation efforts. We present several examples of past and ongoing CS‐based projects to record and monitor fungal diversity. These include projects that are part of broad collecting schemes, those that provide participants with targeted sampling methods, and those whereby participants collect environmental samples from which fungi can be obtained. We also examine challenges and solutions for how such projects can capture fungal diversity, estimate species absences, broaden participation, improve data curation, and translate resulting data into actionable conservation measures. Finally, we close the paper with a call for professional mycologists to engage with amateurs and local communities, presenting a framework to determine whether a given project would likely benefit from participation by citizen scientists.


Citations (70)


... The genus Russula Pers., a type genus of the family Russulaceae, is recognized as the largest genus within the phylum Basidiomycota [1,2]. It comprises approximately 1100 described species, with estimates suggesting the existence of at least 3000 species [3,4]. ...

Reference:

Morphological Studies and Phylogenetic Analyses Unveil Two Notable New Species of Russula Subg. Heterophyllinae from China
Phylogenomics, divergence times and notes of orders in Basidiomycota

Fungal Diversity

... Where more taxonomic approaches to research are needed or used, such as in nationwide conservation strategies, shadow diversity can be employed as a reflexive tool to check for adequate coverage of various groups of life ( Figure 5), echoing calls made on behalf of a historically underfunded taxon, such as fungi (Oyanedel et al. 2022) and a more conscious employment of the factors perpetuating ignorance of shadow diversity (Figure 4). By grouping 13 terms (Figure 1b) under the umbrella of shadow diversity, complementary concepts are more visible and we can more readily share advances in one area of shadow biodiversity as a potential model for elevating greater inclusion for other lifeforms still deeper in shadows (such as success in momentum for improved inclusion of fungi, outlined in Haelewaters et al. (2024)). Rather than discussing "unknown unknowns", shadow diversity further provides shared linguistic means ( Figure 3 and Table 1) to distinguish more precisely the extent to which a particular area of biodiversity is unknown and for whom, compared to the overlapping and unclear set of terms highlighted by our review (Lack of clarity section). ...

The power of citizen science to advance fungal conservation

... Neither Pseudophialophora species have been previously described on a non-grass host as also any other DSE have been reported associated with the palm Acrocomia aculeata. Therefore, we emphasize that, as suggested by Cazabone et al. [50], the description of new taxa, even if singleton-based, is an emerging need within mycology, especially when it comes to rare and still little-explored groups such as DSE and its host. We hope that this work will encourage mycologists and taxonomists to increasingly study and describe DSE fungi present in different environments on the planet. ...

Singleton-based species names and fungal rarity: Does the number really matter?

IMA Fungus

... As of 1 June 2024, iNaturalist had recorded 11,817,129 observations of 22,491 fungal species from 789,727 contributors. The platform's scientific contributions include reassessing fungal taxa (e.g., Mitchell et al. 2022), updating species distribution (e.g., de Groot et al. 2024), studying host specialization of fungi (Gange et al. 2011;Heilmann-Clausen et al. 2016), providing data for local checklists (e.g., Shumskaya et al. 2023), and assessing fungal species' conservation status (Mueller et al. 2022). ...

Beetlehangers.org: harmonizing host–parasite records of Harmonia axyridis and Hesperomyces harmoniae

Arthropod-Plant Interactions

... axyridis), highly invasive in many parts of the world [21], was recorded on Tenerife in 2003 and 2004 [10,31], and its next records known to us are those from 2021 and 2022 reported here. Interestingly, while on Tenerife, H. axyridis appears to be well established (a reproducing population recorded) and associated with its specific natural enemies [32,33], no other island of the Canarian archipelago has been reported to host this ladybird. It can be assumed that the species will appear on other islands of the archipelago in the near future. ...

Parasitism of ladybirds by Hesperomyces in the Canary Islands

Arthropod-Plant Interactions

... (1971), a common jelly fungus in the reserve. Interestingly, members of our lab have described a mycoparasite inhabiting this species, Slooffia micra (Bourdot & Galzin) (Schoutteten 2023) [74], which, despite targeted efforts, has not been found in the sampled area. S. micra was incorporated in the latest version of UNITE [53]) as SH0867261.10FU ...

Diversity of colacosome-interacting mycoparasites expands the understanding of the evolution and ecology of Microbotryomycetes

Studies in Mycology

... Over an extended period, DNA-based techniques have played a pivotal role in uncovering both inter-and intra-species phylogenetic variations, essential for describing new species (Kidd et al. 2023). While the ITS region stands as a universal barcode marker for fungal identification, its applicability to entomophthoroid fungi is hindered by high intragenomic variation (Schoch et al. 2012;Hyde et al. 2023). Fortunately, the development of the full ribosomal operon and additional gene loci encoding proteins as fungal barcodes has addressed some of these challenges (James et al. 2006;Wurzbacher et al. 2019;Voigt et al. 2021;Zhao et al. 2023). ...

Global consortium for the classification of fungi and fungus-like taxa

mycosphere

... & C.P. Robin were supplemented with sequences downloaded from NCBI Gen-Bank (https://www.ncbi.nlm.nih.gov/genbank/) following the dataset in Van Caenegem et al. (2023b). All sequences included in our analyses are listed in Table 2. ...

Botryandromyces, a morphology-based genus concept scrutinized by molecular data

Mycological Progress

... These facts, combined with the decline in the number of taxonomists worldwide, deepening the Linnean shortfall, which refers to the unknown diversity of organisms. This shortfall hampers conservation efforts and monitoring of environmental impacts (Haelewaters et al. 2024;Hortal et al. 2015). In the current scenario of climate and biodiversity crisis, it is possible that microbial species are going extinct without even being known, and key microbial functions are being lost, leading to the loss or alteration of ecosystem services (Hutchins et al. 2019). ...

Biological knowledge shortfalls impede conservation efforts in poorly studied taxa—A case study of Laboulbeniomycetes
  • Citing Article
  • September 2023

Journal of Biogeography

... Due to the WGA, there is a higher risk to amplify contaminants. Yet, it has been successfully used in many other studies by Haelewaters and colleagues (Haelewaters et al. 2018b(Haelewaters et al. , 2019a(Haelewaters et al. , 2019bWalker et al. 2018;Haelewaters and Pfister 2019;Haelewaters and De Kesel 2020;Liu et al. 2020; Van Caenegem et al. 2023a, 2023b. Haelewaters et al. (2019b) used a modified protocol, in which they halved the use of every component, to save products and reduce costs per extraction. ...

Five new species of Gloeandromyces (Fungi, Laboulbeniales) from tropical American bat flies (Diptera, Streblidae), revealed by morphology and phylogenetic reconstruction