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Research Article
Two new species and a new record of Hypoxylaceae (Xylariales, Ascomycota) from Mexico
expand article infoPamela E. Reyes‡, Juan Pablo Pinzón‡, Ricardo Valenzuela§, Tania Raymundo§, Juan Tun-Garrido‡, Ricardo García-Sandoval|
‡ Universidad Autónoma de Yucatán, Mérida, Mexico
§ Instituto Politécnico Nacional, Ciudad de México, Mexico
| Universidad Nacional Autónoma de México, Ciudad de México, Mexico

Corresponding author: Ricardo Valenzuela ( rvalenzg@ipn.mx )

Academic editor: Jian-Kui Liu
© 2024 Pamela E. Reyes, Juan Pablo Pinzón, Ricardo Valenzuela, Tania Raymundo, Juan Tun-Garrido, Ricardo García-Sandoval.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Reyes PE, Pinzón JP, Valenzuela R, Raymundo T, Tun-Garrido J, García-Sandoval R (2024) Two new species and a new record of Hypoxylaceae (Xylariales, Ascomycota) from Mexico. MycoKeys 111: 111-127. https://doi.org/10.3897/mycokeys.111.133046
Open Access

Abstract

The family Hypoxylaceae has a cosmopolitan distribution with greater diversity in tropical regions, its growth habit is saprotrophic, endophytic and potentially phytopathogenic. From the revision of herbarium specimens and field collections from the Yucatan Peninsula, two new species were described: Annulohypoxylon fusisporum, characterized by having fusiform spores and Hypoxylon xmatkuilense which is distinguished by having stromata vinaceous and dark brown KOH-extractable pigments. The species are described based on morphological characters and Bayesian Inference analyses of four molecular markers (ITS, LSU, RPB2 and TUB2). In addition, one new record from Mexico is presented: Hypoxylon bellicolor. The presence of Daldinia eschscholtzii, H. lenormandii, H. lividipigmentum and Entonaema liquescens is confirmed with molecular data.

Key words

Diversity, Neotropics, phylogeny, Yucatan Peninsula

Introduction

The family Hypoxylaceae (Xylariales, Ascomycota) contains 22 genera and 495 species (Wijayawardene et al. 2022; Bánki et al. 2024) 75 species have been cited from Mexico: Annulohypoxylon (6), Daldinia (13), Durotheca (1), Entonaema (4), Hypomontagnella (2), Hypoxylon (36), Jackrogersella (2), Parahypoxylon (1), Phylacia (8), Pyrenopolyporus (2) (CONABIO 2024).

They are characterized by having erect, glomerate, pulvinate, discoid, effused-pulvinate, hemispherical, spherical or peltate stromata; solitary or confluent, brightly colored, dark or black, pruinose or smooth, with or without extractable pigments that are evident with 10% KOH; spherical, obovoid or tubular perithecia with spherical, umbilicate or papillate ostioles, with or without discs formed by dehiscence of the surrounding tissue. They share a nodulisporium-like asexual state (which is one of the features that sets them apart from Xylariaceae) and are distributed in tropical and temperate areas of the world (Ju and Rogers 1996). Their lifestyle is mainly endophytic, saprotrophic and even associated with insect vectors (Wendt et al. 2018).

Among the most important taxonomic works that have been published about the genera that at this moment are considered as part of Hypoxylaceae, we can find the monograph of Hypoxylon by Ju and Rogers (1996), the revision of Child (1932) who published the first monograph of Daldinia with 11 species, furthermore Ju et al. (1997) described 19 species of the genus. Stadler et al. (2014a) recognized 47 taxa of Daldinia based on morphological, chemotaxonomic and phylogenetic evidence; Sir et al. (2016) described D. korfii from the Yungas region in Argentina; Chlebicki (2022) presented a review of the genus from Poland.

The genus Entonaema, was erected by Alfred Möller (1901) who described to E. liquescens and E. mesentericum, currently as Xylaria mesenterica (Stadler et al. 2008). Pošta et al. (2023) accepted six species: E. liquescens, E. cinnabarinum, E. dengii, E. moluccanum, E. globosum, and E. siamensis.

In the study by Wendt et al. (2018), based on analysis with four molecular markers (ITS, LSU, RPB2 and TUB2) Hypoxylaceae was formally recognized, clearly segregated from Xylariaceae with the following genera: Acrostaphylus, Annulohypoxylon, Anthocanalis, Ascoporia, Chlorostroma, Daldinia, Entonaema, Henningsina, Hypoxylina, Hypoxylon, Jackrogersella, Phylacia, Pyrenomyxa, Pyrenopolyporus, Rhopalostroma, Rostrohypoxylon, Ruwenzoria, Sphaeria, Thamnomyces and Thuemenella.

Lambert et al. (2019) erected Hypomontagnella based on a study of Hypoxylon monticulosum and its allies. The features that characterize them are the stromata woody to carbonaceous lacking colored granules, papillate ostioles usually with a black annulate disc, without apparent KOH-extractable pigments in mature stromata and perispores smooth or with transversally striate ornamentations.

Recently, Cedeño-Sanchez et al. (2023) proposed Parahypoxylon as a new genus including P. papillatum and P. ruwenzoriense; Lambert et al. (2023) studied Phylacia species in Argentina and found a close relationship with Rhopalostroma, Thamnomyces and Daldinia species, which all have similar secondary metabolites.

In Mexico, 75 species of the family are known on all types of vegetation, standing out the study of San Martín et al. (1999b) in which they reviewed the genus Hypoxylon including some members of Section Annulata (now genus Annulohypoxylon); San Martín and Lavin (1997) described three species of Entonaema; Medel et al. (2006) made a review of Phylacia, recording eight species; Barbosa-Reséndiz et al. (2020) reported an updated list of Daldinia in Mexico; Raymundo et al. (2014, 2017, 2021) and Reyes et al. (2020) have reported species of Hypoxylaceae from Protected Natural Areas: Chamela-Cuixmala, Cozumel Island, Lagunas de Chacahua National Park, Sierra de Álamos-Río Cuchujaqui and El Cielo. For this work, several specimens from herbaria and field collections in the Yucatan Peninsula have been studied taxonomically. Morphological and molecular analyses have revealed some taxonomic novelties that we report in this document.

Materials and methods

Collections sites and sampling

In the current paper, 240 specimens were studied, of which 148 are deposited in the “Alfredo Barrera Marín” Herbarium of the Universidad Autónoma de Yucatán (UADY) and the “Gastón Guzmán Huerta” Fungi Collection of the Herbarium of the Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional (ENCB), while 77 were collected in the Campeche, Quintana Roo and Yucatan States between 2021–2023. In this study, nine specimens were analyzed morphologically and sequenced.

Morphological characterization

The description of macromorphological features was carried out with water and 10% KOH to examine perisporium dehiscense and to view the stromatal pigments, Melzer’s reagent to show the amyloid reaction of the ascal apical apparatus, specialized literature in each genus was reviewed (San Martín et al. 1999b; Stadler et al. 2014a; Kuhnert et al. 2016; Lambert et al. 2019; Reyes et al. 2020), colors were described according to Rayner (1970).

DNA extraction, PCR and sequencing

The stromata were macerated in liquid nitrogen, then the QIAGEN DNeasy kit (Hilden, Germany) was used according to the manufacturer’s specifications. The primers used were reported by White et al. (1990) (ITS1, ITS4), Gardes and Brunes (1993) (ITS1-F) for ITS (internal transcriber spacer gene); Vilgalys and Hester (1990) (LR07, LR0R) for LSU (large subunit ribosomal gene); Liu et al. (1999) (RPB2-5F, -7cR) for RPB2 (partial second largest subunit of the DNA-directed RNA polymerase II gene); O’Donnell and Cigelnik (1997) (T1, T22) for TUB2 (beta-tubulin gene). PCR reactions were made using 25 µl reaction:12.5 µl GoTaq® (Promega, Madison, USA) “Master Mix”, 2-3 µl DNA, 0.5 µl of each primer (100 µM), BSA 2.5 µl and 7 µl of sterilized water. The PCR conditions are detailed in Table 1. Sequencing was performed by the Sanger technique at Macrogen Inc.

Table 1.
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PCR conditions used in this study.

DNA locus Initial denaturation Cycles Denaturation Annealing Elongation Final denaturation
ITS 94 °C-5 mins 35 94 °C-1 min 55 °C-1 min 72 °C-1 min 72 °C-10 mins
LSU 94 °C-5 mins 34 94 °C- 1 min 52 °C-1 min 72 °C -2 mins 72 °C-10 mins
RPB2 94 °C- 5 mins 38 94 °C-30 s 53 °C-1 min 72 °C-1.30 mins 72 °C-10 mins
TUB2 94 °C-5 mins 38 94 °C -30 s 47 °C-30 s 72 °C-2.30 mins 72 °C-10 mins

Molecular phylogenetic analyses

A concatenated matrix of the four molecular markers was constructed using 274 sequences from 71 species as a reference from the GenBank (Appendix 1) and 36 obtained in the present study. Sequences were aligned using MAFFT software (Katoh and Kuma 2002) and edited in BioEdit (Hall 1999). The evolution model that best fit each group of sequences was obtained using the j Model Test 2.1.10 software (Posada and Crandall 1998) supported by Bayesian Information Criterion (BIC). A Maximum Likelihood analysis was performed in RaxML 8.2.12 (Stamatakis 2006) with 1000 bootstrap replicates. Additionally, a Bayesian analysis was carried out with the same matrix in Mr.Bayes 3.2.6. (Ronquist and Huelsenbeck 2003) using four MCMC chains, 10000000 generations, taking samples every 1000 generations, applying a burnin of 25%, Xylaria polymorpha was used as an outgroup.

Results

Phylogenetic analysis

The molecular matrix of the four concatenated loci (ITS, LSU, RPB2 and TUB2) was 7283 bp in length, having 1640 for the first, 2417 for the second, 1200 for the third and 2026 for the fourth, applying the following substitution models: SYM+I+G for ITS, HKY+I+G for TUB2 and GTR+I+ G for LSU and RPB2.

The family Hypoxylaceae was represented in the phylogenetic tree by nine genera among which we can find: Phylacia (Phy) with two specimens, Thamnomyces (Tha) one specimen, Rhopalostroma (Rho) one specimen, Daldinia (D1 and D2) 14 specimens, Ruwenzoria (Ru) one specimen, Pyrenopolyporus (Py) three specimens, Annulohypoxylon (A) nine specimens, Jackrogersella three specimens, Rostrohypoxylon (Ro) one specimen, Hypomontagnella (Hyp) two specimens, Durotheca (Du) three specimens, Parahypoxylon (Pa) two specimens, Entonaema two specimens, Hypoxylon (H1, H2, H3, H4) 36 specimens (Fig. 1), which were selected by having molecular information supported by several studies (Wendt et al. 2018; Cedeño-Sanchez et al. 2023; Lambert et al. 2023; Pošta et al. 2023).

Figure 1. 

Inferred molecular phylogenetic tree obtained by Bayesian Inference using a multigene alignment (ITS, LSU, RPB2 and TUB2). The tree was rooted with Xylaria polymorpha. The sequences generated in the present work and the new combination are highlighted. Bayesian posterior probability values ≥ 0.98 and Bootstrap support values ≥ 70 from the Maximum Likelihood analysis are indicated on the branches.

In the phylogram resulted from the Bayesian Inference analysis (Fig. 1), Hypomontagnella, Jackrogersella, Parahypoxylon, Phylacia, Pyrenopolyporus, Rhopalostroma and Thamnomyces appear as monophyletic; Daldinia as paraphyletic with supports in each group, D1 with 1/95 and D2 with 1/100; Annulohypoxylon also paraphyletic with values of 1/96; Hypoxylon as polyphyletic in four groups, H1 with values of 1/97, H2 with support of 1/100, H3 with 1/92, H4 + E with 1/95.

The identity of taxa previously reported for the country such as Daldinia eschscholtzii, Hypoxylon lenormandii, H. lividipigmentum and Entonaema liquescens is confirmed, of which there was limited molecular information for Mexican specimens.

According to the morphological revision of the specimens and the topology of the trees, two new species arise: Annulohypoxylon fusisporum and Hypoxylon xmatkuilense; the presence of H. bellicolor is confirmed in the country.

Taxonomy

New species from Mexico

Annulohypoxylon fusisporum P. Reyes, Pinzón, R. Valenz. & Raymundo, sp. nov.

MycoBank No: 851003
Fig. 2

Gen Bank

ITS (OR807998), LSU (OR807987), RPB2 (OR825472), TUB2 (OR825468).

Diagnosis

It is characterized by having fusiform spores 10–12 × 4–5 µm, grayish green to dull green KOH-extractable pigments, ¼ perithecial mounds exposed, with straight germ slit in spore-length on the convex side.

Etymology

in reference to the fusiform spores.

Holotype

Mexico • Quintana Roo, Bacalar Experimental Station, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias; 10 March 2023; P. Reyes leg.; UADY 83.

Description

Stromata effused-pulvinate 20–70 mm long × 20–30 mm wide × 1 mm thick, surface dark brown to black (Fig. 2a) inconspicuous perithecial mounds up to 1/4 exposed (Fig. 2b) dark brown to black granules beneath surface, KOH-extractable pigments Grayish Green (50) changing to Dull Green (70) after 1 minute (Fig. 2c) perithecia 0.7–0.8 mm diam, spherical, conical- papillate ostioles surrounded with a truncatum type disc 0.2–0.25 mm diam; asci 8-spored cylindrical, the spore bearing parts 60–75 µm long × 5–7 µm wide, stipes 20–45 µm long, with amyloid, discoid apical apparatus, 1–2 µm high × 2 µm wide (Fig. 2d); ascospores 10–12 × 4–5 µm, unicellular, fusiform with narrowly rounded ends, with faint, straight germ slit along the spore on the convex side (Fig. 2e), light brown, perispore dehiscent 10% KOH (Fig. 2f).

Figure 2. 

Annulohypoxylon fusisporum sp. nov. a general view of stromata b perithecia spherical with truncatum type ostiolar disc c KOH-extractable pigments d asci e straight germ slit f perispore dehiscent 10% KOH.

Host

Growing on dead wood of Brosimum alicastrum (Moraceae) in subevergreen forest.

Notes

It is similar to A. subeffusum (Hladki and Romero 2009) due to the color, shape of the stromata and the grayish green pigments, however the ascospores of this one are smaller 7–10 × 3–3.8 µm and have inconspicuous perithecial mounds. With A. leptascum it shares fusiform shape spores, unlike the length of the germ slit which is short, and the pigments are olive-colored, not changing after 1 minute in 10% KOH; Kuhnert et al. (2016) found BNT, truncatone A and truncatone C in aforementioned taxa.

Hypoxylon xmatkuilense P. Reyes, Pinzón, R. Valenz. & Raymundo, sp. nov.

MycoBank No: 855643
Fig. 3

Gen Bank

ITS (OR807999), LSU (OR807990), RPB2 (OR825476), TUB2 (OR825467).

Diagnosis

It characterized by having stromata effused-flattened, surface vinaceous, perithecia obovoid and dark brown KOH-extractable pigments.

Etymology

from the Mayan Xmatkuil “place where God is asked” in reference to the place where it was collected for the first time.

Holotype

Mexico • Yucatán, east from cemetery, comisaría Xmatkuil; 14 Oct 2021; P. Reyes leg.; UADY 3.

Paratypes

Mexico • ibid.; 11 July 2023; P. Reyes leg.; UADY 118 • ibid; P. Reyes leg.; UADY 119.

Description

Stromata effused-flattened, 25–100 mm long × 5–20 mm wide × 1 mm thick, surface pruinose, young stromata have a thin layer Dark Vinaceous (82), when mature this layer is lost, leaving remains that appear Brown Vinaceous (84) (Fig. 3a), subsurface blackish, composed of weakly carbonaceous tissue and inconspicuous brownish black granules; perithecia 1.2–1.5 mm diam, obovoid inconspicuous with umbilicate ostioles (Fig. 3b); KOH-extractable pigments dark brown (Fig. 3c); asci not seen; ascospores 10–12 × 4–5 µm, unicellular, ellipsoid, inequilateral, narrowly rounded ends, with straight 2/3 spore-length germ slit on the convex side (Fig. 3d), dark brown, perispore dehiscent in 10% KOH (Fig. 3e).

Figure 3. 

Hypoxylon xmatkuilense sp. nov a general view of stromata b perithecia obovoid c KOH-extractable pigments d straight germ slit on the convex side e perispore dehiscent 10% KOH.

Host

Growing on dead wood of Gymnopodium floribundum Rolfe (Polygonaceae) in deciduous forest

Notes

It shares the effused-pulvinate to flattened stromata and vinaceous surface with H. lateripigmentum, H. pulicicidum, H. investiens, H. hinnuleum, H. olivaceopigmentum, it differs by having perithecia obovoid, dark brown KOH-extractable pigments and perispore dehiscent (Ju and Rogers 1996; Bills et al. 2012; Kuhnert et al. 2014a; Sir et al. 2019); Fournier et al. (2024) described H. aeneipigmentatum as a new taxon from Saül, French Guiana, based on ITS sequences, and noted that it belongs H. investiens complex; moreover reported about a basal perithecial nucleus as a new differential character in this group.

New record from Mexico

Hypoxylon bellicolor Cedeño-Sanchez, L. Wendt & L.C. Mejía 2020 Mycosphere 11(1): 1464 (2020).

Fig. 4

Description

Stromata effused-pulvinate 20–90 mm long × 10–30 mm wide, Rust (39) (Fig. 4a) pruinose surface with rust granules beneath the surface (Fig. 4b), KOH-extractable pigments Luteous (12) (Fig. 4c); perithecia ovoid, umbilicate ostioles (Fig. 4d); asci 8-spored cylindrical, the spore bearing part 40–80 µm long × 5–6 µm wide, stipes 30–35 µm long, with amyloid, discoid apical apparatus 2–3 µm high × 2 µm wide (Fig. 4e); ascospores 10–12 µm long × 4–6 µm wide, with germ slit straight less than spore-length on the convex side, perispore dehiscent in 10% KOH with coil-like ornamentation (Fig. 4f).

Figure 4. 

Hypoxylon bellicolor a general view of stromata b pruinose surface c KOH-extractable pigments d perithecia ovoid e asci f perispore dehiscent 10% KOH with coil-like ornamentation.

Specimens examined

Mexico • Campeche, Champotón-Campeche highway; 19 Jan 2018; P. Reyes leg.; UADY 145 • Quintana Roo, Centro de Conservación y Educación Ambiental, Cozumel; 20 Jan 2018; R. Valenzuela leg.; ENCB 17942 •Yucatán, east from the cemetery, comisaría Xmatkuil; 15 Oct 2021; P. Reyes leg.; UADY 7 • ibid.; P. Reyes leg.; UADY 9 •Southern Ecological Park, Mérida; 21 Sept 2022; P. Reyes leg.; UADY 45 • Ría Celestún Biosphere Reserve; 29 Oct 2022; P. Reyes leg.; UADY 55.

Host

Growing on dead wood of Lysiloma latisiliquum (L.) Benth. (Fabaceae), Lonchocarpus sp. (Fabaceae) in deciduous forest; on dead wood of Conocarpus erectus L. (Combretaceae) in coastal dune vegetation.

Known distribution

Panama (Cedeño-Sanchez et al. 2020).

Notes

It shares a stromata color and yellow pigments with H. perforatum, however this one is characterized by having a white disc surrounding the ostioles; in addition it has a cosmopolitan distribution (Ju and Rogers 1996).

Discussion

Since the segregation of Hypoxylaceae from Xylariaceae (Wendt et al. 2018), many questions have arisen, which have been answered over the years based on the review of groups as Hypoxylon that appears as polyphyletic, and from which the genera Hypomontagnella (Lambert et al. 2019) and Parahypoxylon (Cedeño-Sanchez et al. 2023) have emerged; likewise, Daldinia appears grouped in two different clades that are supported in the present study, so it requires a review from integral taxonomy approach to elucidate several questions that will continue to arise within the family.

In the phylogenetic tree of the present study it can be seen that most of the clades of the known genera have good statistical support, Parahypoxylon, Durotheca, Jackrogersella, Pyrenopolyporus, Hypomontagnella, Rhopalostroma, Thamnomyces, Phylacia and Ruwenzoria noted as monophyletic.

As for Hypoxylon, it appears as polyphyletic forming four groups (labeled here as H1, H2, H3, and H4, Fig. 1). Group H1 is formed by the following species: 1) H. investiens which has a wide distribution, mainly in a tropical climate; it is distinguished by having tubular perithecia. 2) H. pulicicidum segregated from the previous one by Fournier et al. (2015b) and Bills et al. (2012) who consider it a rare taxon; it is distinguished by having slightly papillated ostioles and lanceolate perithecia. 3) H. lateripigmentum has olive-yellow granules beneath the surface, yellowish brown pigments in 10% KOH and perispore dehiscent. 4) H. olivaceopigmentum is distinguished by having the largest spores in the complex 9–15.7 × 4.5–7.2 µm and has been recorded from monocotyledonous hosts. 5) H. hinnuleum, proposed by Sir et al. (2019) stands out by having ostioles conical black papillate. Finally, the new species H. xmatkuilense is characterized by having perithecia obovoid, dark brown KOH-extractable pigments and perispore dehiscent. This species is sister to H. lateripigmentum, which is only known from Martinique.

The members of this group have tropical distribution, H. xmatkuilense and H. lateripigmentum are Neotropical, so far reported from Caribbean area; H. olivaceopigmentum and H. hinnuleum from subtropical area, while H. investiens and H. pulicicidum are Pantropical.

H2 is made up only by H. lividipigmentum with two Mexican specimens; this one is characterized by having a sigmoid germ slit in the ascospores, a taxonomic character that is less frequent in the genus.

H3 where the type H. fragiforme stands out, so this clade stands as Hypoxylon sensu stricto, share distribution with H. howeanum, H. ticinense of temperate climate, meanwhile H. crocopeplum, H. fendleri, H. lenormandii, H. haematostroma and H. rickii have been recorded from tropical areas (Wendt et al. 2018; Cedeño-Sanchez et al. 2023).

H4 consists of species of varied distribution and specific plant associations as H. fuscum recorded on hosts of the family Betulaceae, in addition to Acer and Salix; H. vogesiacum associated with Acer, H. porphyreum with Quercus; regarding H. bellicolor, a new record from Mexico, we registered it growing on Fabaceae and Combretaceae hosts; furthermore, the sequences of the four markers were obtained from two Mexican specimens enriching the knowledge of this taxon, since they were only available ITS and TUB2 sequences (Ju and Rogers 1996; Fournier et al. 2015b; Cedeño-Sanchez et al. 2020).

The polyphyletic origin of Hypoxylon encourages more morphological, chemical and phylogenetic studies, including a better sampling of the species to resolve Hypoxylon evolutionary history and probably segregate new genera.

On the other hand, our sequences of Entonaema liquescens (ENCB:RV_19274) matched with those of the specimen CNF 2/11263 (Pošta et al. 2023). However, they had no similarity with those from the strain ATCC 46302 (Wendt et al. 2018). This pattern is explained by Stadler et al. (2020) and Cedeño-Sanchez et al. (2024) who point out heterogeneity and polymorphisms among multiple copies of different ITS and LSU loci.

The genus Daldinia resulted to be paraphyletic, in agreement with Stadler et al. (2014a), forming two groups; in D1 group, most of the species are exclusively tropical such as D. bambusicola, D. eschscholtzii, D. placentiformis and D. theissenii; the D2 group it would be the genus sensu stricto, this clade has a widely distribution and there is a recurrence in the hosts of Betulaceae and Lauraceae (Stadler et al. 2014a).

Annulohypoxylon also appears as paraphyletic, having Rostrohypoxylon terebratum nested inside the clade, which agrees with Tang et al. (2009) and Wendt et al. (2018); it is closely related to the new species A. fusisporum, proposed in the current study; Stadler (2011) mentioned that R. terebratum represents a lineage that has evolved from Annulohypoxylon, while differing in some morphological features and the production of unique secondary metabolites. It is worth mentioning that this taxon could be a turning point for segregating the genus, but more taxonomic and genomic samples are needed.

Conclusion

The present study is important because it provides information about the species from a tropical area, particularly from the Yucatan Peninsula, which has a valuable biogeographic history that makes the family have a great diversity, and therefore two new species are described and reports a new record from this region in Mexico; the confirmation of the identity of four species through molecular phylogenetic data is also relevant, since cryptic or semicryptic species may appear, especially in tropical areas.

The proposal of new species is fundamental for the understanding of a recently established family, in which there are still questions to be clarified. These questions will have to be supported by a polyphasic approach that provides comprehensive tools for the interpretation of the phylogenetic relationships in the group.

Acknowledgements

The authors are grateful for the facilities granted to Fernando Arellano Martín, researcher at the Chetumal Experimental Field of the Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP). To Orlando Poot, mayor of Tres Garantías, Quintana Roo. PhD, Juan José Ancona for the identification of hosts in the municipality of Tekax, Yucatán. To the Mesoamerican Floristic Resources Laboratory and the Tropical Ecology Laboratory of the Universidad Autónoma de Yucatán for the use of the facilities and equipment and Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional for the loan of herbaria material. Reyes acknowledges the Consejo Nacional de Humanidades, Ciencia y Tecnología (CONAHCYT) for the grant awarded.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was financed by Instituto Politécnico Nacional (SIP20230017; SIP20240029; SIP20240367).

Author contributions

Conceptualization: JPP. Project administration: RV. Supervision: JTG, TR. Visualization: RGS. Writing - original draft: PER.

Author ORCIDs

Pamela E. Reyes https://orcid.org/0000-0003-0882-5245

Juan Pablo Pinzón https://orcid.org/0000-0002-0964-2996

Ricardo Valenzuela https://orcid.org/0000-0001-6596-522

Tania Raymundo https://orcid.org/0000-0002-7525-0034

Juan Tun-Garrido https://orcid.org/0000-0001-8796-3405

Ricardo García-Sandoval https://orcid.org/0000-0001-8641-3305

Data availability

All of the data that support the findings of this study are available in the main text.

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Appendix 1

Table A1.
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List of sequences used in phylogenetic analyses obtaneid from Gen Bank. Sequences derived from type material are marked as type (T), holotype (HT), epitype (ET) isotype (IT) and paratype (PT), the sequences obtained in this study are highlighted.

Species Strain/voucher number Origin and status ITS LSU RPB2 TUB2 Reference
Annulohypoxylon annulatum CBS 140775 USA (ET) KY610418 MK287546 KY624263 KX376353 (Kuhnert et al. 2016; Wendt et al. 2018)
Annulohypoxylon atroroseum ATCC 76081 Thailand AJ390397 KY610422 KY624233 DQ840083 (Kuhnert et al. 2014a; Wendt et al. 2018)
Annulohypoxylon fusisporum sp. nov. UADY:PR_83 Mexico (HT) OR807998 OR807987 OR825472 OR825468 This study
Annulohypoxylon leptascum MFLUCC 13-0587 Thailand KU604576 KU604580 Sir et al. (2016a)
Annulohypoxylon michelianum CBS 119993 Spain KX376320 KY610423 KY624234 KX271239 Wendt et al. (2018); Kuhnert et al. (2014a)
Annulohypoxylon moriforme CBS 123579 Martinique KX376321 KY610425 KY624289 KX271261 Wendt et al. (2018); Kuhnert et al. (2016)
Annulohypoxylon nitens AXL030 Thailand KJ934991 KJ934992 KJ934994 KJ934993 Daranagama et al. (2015)
Annulohypoxylon stygium MUCL 54601 French Guiana KY610409 KY610475 KY624292 KX271263 Wendt et al. (2018)
Annulohypoxylon truncatum CBS 140778 USA (ET) KY610419 KY610419 KY624277 KY624277 Wendt et al. (2018); Kuhnert et al. (2016)
Daldinia andina CBS 114736 Ecuador (HT) AM749918 KY610430 KY624239 KC977259 Bitzer et al. (2008)
Daldinia bambusicola CBS 122872 Thailand (HT) KY610385 KY610431 KY624241 KY624241 Wendt et al. (2018); Hsieh et al. (2005)
Daldinia caldariorum MUCL 49211 France AM749934 KY610433 KY624242 KC977282 Wendt et al. (2018); Kuhnert et al. (2014a); Bitzer et al. (2008)
Daldinia concentrica CBS 113277 Germany AY616683 KY610434 KY624243 KC977274 Wendt et al. (2018); Kuhnert et al. (2014a); Triebel et al. (2005)
Daldinia dennisii CBS 114741 Australia (HT) JX658477 KY610435 KY624244 KC977262 Stadler et al. (2014a); Kuhnert et al. (2014a); Wendt et al. (2018)
Daldinia eschscholtzii MUCL 45434 Benin JX658484 KY610437 KY624246 KC977266 Stadler et al. (2014a); Kuhnert et al. (2014a); Wendt et al. (2018)
Daldinia eschscholtzii UADY:PG_136 Mexico OR808001 OR807988 OR825473 OR825469 This study
Daldinia loculatoides CBS 113279 UK (ET) AF176982 KY610438 KY610438 KX271246 Johannesson et al. (2000); Wendt et al. (2018)
Daldinia macaronesica CBS 113040 Spain (PT) KY610398 KY610477 KY624294 KX271266 Wendt et al. (2018)
Daldinia petriniae MUCL 49214 Austria (ET) AM749937 KY610439 KY624248 KC977261 Bitzer et al. (2008); Kuhnert et al. (2014a); Wendt et al. (2018)
Daldinia placentiformis MUCL 47603 Mexico AM749921 KY610440 KY624249 KC977278 Bitzer et al. (2008); Kuhnert et al. (2014a); Wendt et al. (2018)
Daldinia steglichii MUCL 43512 Papua New Guinea (PT) KY610399 KY610479 KY624250 KX271269 Wendt et al. (2018)
Daldinia theissenii CBS 113044 Argentina (PT) KY610388 KY610441 KY624251 KX271247 Wendt et al. (2018)
Daldinia vernicosa CBS 119316 Germany
(PT)		 KY610395 KY610442 KY624252 KC977260 Kuhnert et al. (2014a); Wendt et al. (2018)
Durotheca crateriformis GMBC0205 China (T) MH645426 MH645425 MH645427 MH049441 De Long et al. (2019)
Durotheca guizhouensis GMBC0065 China (T) MH645423 MH645421 MH645422 MH049439 De Long et al. (2019)
Durotheca rogersii GMBC0204 China MH645433 MH645434 MH645435 MH049449 De Long et al. (2019)
Entonaema liquescens CNF 2/11263 USA OQ869784.1 OQ865124.1 OQ877106.1 OQ877117.1 Pošta et al. (2023)
Entonaema liquescens ENCB:RV_19274 Mexico OR807997 OR807993 OR825474 OR825466 This study
Hypoxylon bellicolor UCH9543 Panama MN056425.1 MK908139 Cedeño-Sanchez et al. (2020)
Hypoxylon bellicolor UADY:PR_9 Mexico OR808002 OR807989 OR825470 This study
Hypoxylon bellicolor ENCB:PR_145 Mexico OR808004 OR807994 OR825475 OR825471 This study
Hypoxylon carneum MUCL 54177 France KY610400 KY610480 KY624297 KX271270 Wendt et al. (2018)
Hypoxylon cercidicola CBS 119009 France KC968908 KY610444 KY610444 KC977263 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon crocopeplum CBS 119004 France KC968907 KY610445 KY624255 KC977268 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon fendleri MUCL 54792 French Guiana KF234421 KY610481 KY624298 KF300547 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon fragiforme MUCL 51264 Germany (ET) KC477229 KM186295 KM186296 KM186296 Stadler et al. (2013); Daranagama et al. (2015); Wendt et al. (2018)
Hypoxylon fuscum CBS 113049 Germany (ET) KY610401 KY610482 KY624299 KX271271 Wendt et al. (2018)
Hypoxylon griseobrunneum CBS 331.73 India (HT) KY610402 KY610483 KY624300 KC977303 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon haematostroma MUCL 53301 Martinique (ET) KC968911 KY610484 KY624301 KC977291 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon hinnuleum ATCC 36255 USA (T) MK287537 MK287549 MK287562 MK287575 Sir et al. (2019)
Hypoxylon howeanum MUCL 47599 Germany AM749928 KY610448 KY624258 KC977277 Bitzer et al. (2008); Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon hypomiltum MUCL 51845 Guadeloupe KY610403 KY610449 KY624302 KX271249 Wendt et al. (2018)
Hypoxylon investiens CBS 118183 Malaysia KC968925 KY610450 KY624259 KC977270 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon investiens TBRC 16251 Thailand OQ108848.1 OQ144968.1 Suetrong et al. (2023)
Hypoxylon lateripigmentum MUCL 53304 Martinique (HT) KC968933 KY610486 KY624304 KC977290 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon lenormandii CBS 119003 Ecuador KC968943 KY610452 KY624261 KC977273 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon lenormandii UADY:PR_82 Mexico OR808003 OR807991 OR825477 This study
Hypoxylon lividipigmentum BCRC 34077 Mexico (IT) JN979433 AY951735 Hsieh et al. (2005)
Hypoxylon lividipigmentum UADY:PR_1 Mexico OR808000 OR807992 OR825478 This study
Hypomontagnella monticulosa MUCL 54604 French Guiana (ET) KY610404 KY610487 KY624305 KX271273 Wendt et al. (2018)
Hypoxylon musceum MUCL 53765 Guadeloupe KC968926 KY610488 KY624306 KC977280 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon olivaceopigmentum DSM 107924 USA (T) MK287530 MK287542 MK287555 MK287568 Sir et al. (2019)
Hypoxylon perforatum CBS 115281 France KY610391 KY610455 KY624224 KY624224 Wendt et al. (2018)
Hypoxylon petriniae CBS 114746 France (HT) KY610405 KY610491 KY624279 KX271274 Kuhnert et al. (2016); Wendt et al. (2018)
Hypoxylon pilgerianum STMA 13455 Martinique KY610412 KY610412 KY624308 KY624315 Wendt et al. (2018)
Hypoxylon porphyreum CBS 119022 France KC968921 KY610456 KY624225 KC977264 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon pulicicidum CBS 122622 Martinique (HT) JX183075 KY610492 KY624280 JX183072 Bills et al. (2012); Wendt et al. (2018)
Hypoxylon rickii MUCL 53309 Martinique (ET) KC968932 KY610416 KY624281 KC977288 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon rubiginosum MUCL 52887 Germany (ET) KC477232 KY610469 KY624266 KY624311 Stadler et al. (2013); Wendt et al. (2018)
Hypoxylon samuelsii MUCL 51843 Guadeloupe (ET) KC968916 KY610466 KY624269 KC977286 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypomontagnella submonticulosa CBS 115280 France KC968923 KY610457 KY624226 KC977267 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon ticinense CBS 115271 France JQ009317 KY610471 KY624272 AY951757 Hsieh et al. (2005); Wendt et al. (2018)
Hypoxylon trugodes MUCL 54794 Sri Lanka (ET) KF234422 KY610493 KY624282 KF300548 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon vogesiacum CBS 115273 France KC968920 KY610417 KY624283 KX271275 Kuhnert et al. (2014a); Wendt et al. (2018)
Hypoxylon xmatkuilense sp.nov. UADY:PR_3 Mexico (HT) OR807999 OR807990 OR825476 OR825467 This study
Hypoxylon xmatkuilense sp. nov. UADY:PR_118 Mexico (PT) OR807996 OR807995 PP239283 PP239284 This study
Jackrogersella cohaerens CBS 119126 Germany KY610396 KY610497 KY624270 KY624314 Wendt et al. (2018)
Jackrogersella minutella CBS 119015 Portugal KY610381 KY610424 KY610424 KX271240 Kuhnert et al. (2016); Wendt et al. (2018)
Jackrogersella multiformis CBS 119016 Germany (ET) KC477234 KY610473 KY624290 KX271262 Kuhnert et al. (2014a); Wendt et al. (2018)
Parahypoxylon papillatum ATCC 58729 USA (T) KC968919 KY610454 KY624223 KC977258 Kuhnert et al. (2014a), Wendt et al. (2018)
Parahypoxylon ruwenzoriense MUCL51392 Congo (T) ON792786 ON954156 OP251039 ON813078 Cedeño-Sanchez et al. (2023)
Phylacia globosa STMA 18042 Argentina OQ437889.1 OQ437885 OQ453168 OQ453172 Lambert et al. (2023)
Phylacia lobulata STMA 18040 Argentina OQ437893.1 OQ437883 OQ453164 OQ453170 Lambert et al. (2023)
Pyrenopolyporus hunteri MUCL 52673 Ivory Coast (ET) KY610421 KY610472 KY624309 KU159530 Kuhnert et al. (2016); Wendt et al. (2018)
Pyrenopolyporus laminosus MUCL 53305 Martinique (HT) KC968934 KY610485 KY624303 KC977292 Kuhnert et al. (2014a); Wendt et al. (2018)
Pyrenopolyporus nicaraguensis CBS 117739 Burkina Faso AM749922 KY610489 KY624307 KC977272 Bitzer et al. (2008); Wendt et al. (2018)
Rhopalostroma angolense CBS 126414 Ivory Coast KY610420 KY610459 KY624228 KX271277 Wendt et al. (2018)
Rostrohypoxylon terebratum CBS 119137 Thailand (HT) DQ631943 DQ840069 DQ631954 DQ840097 Tang et al. (2007); Fournier et al. (2010)
Ruwenzoria pseudoannulata MUCL 51394 D.R. Congo (HT) KY610406 KY610494 KY624286 KX271278 Wendt et al. (2018)
Thamnomyces dendroidea CBS 123578 French Guiana (HT) FN428831 KY610467 KY624232 KY624313 Stadler et al. (2010b); Wendt et al. (2018)
Xylaria polymorpha MUCL 49884 France KY610408 KY610464 KY624288 KX271280 Wendt et al. (2018)
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