Testudinaceae Arx, Persoonia 6(3): 366 (1971).

MycoBank number: MB 81456; Index Fungorum number: IF 81456; Facesoffungi number: FoF 08372, 20 species.

Saprobic on dead or decaying wood, parasitic on fungi, pathogenic on humans, isolated from soil, decaying plant materials and woody substrata, in terrestrial, freshwater and marine habitats. Sexual morph: Ascomata cleistothecial or perithecial, globose to subglobose, carbonaceous, dark- brown to black, immersed, clypeate, papillate, ostiolate or lacking ostioles, periphysate. Peridium multi-layered, thick-walled cells of textura angularis. Hamathecium comprising branched, septate, cellular or trabeculate pseudoparaphyses, evanescent to persistent. Asci 8-spored, thick-walled, bitunicate, fissitunicate, clavate to cylindrical, short pedicellate, lacking an apical structure or with an ocular chamber, evanescent to persistent. Ascospores 1-seriate, relatively small, ellipsoidal, brown or hyaline, 1-septate or multi-septate, or muriform, with or without ornamentation on spore surface, with or without furrows, verrucose to verruculose. Asexual morph: Undetermined.

TypeTestudina Bizz.

NotesTestudinaceae was established by von Arx (1971) to accommodate Testudina, Neotestudina Lepidosphaeria, Argynna (transferred to Argynnaceae) and Pseudophaeotrichum (synonymized as Neotestudina). This family is characterized by astomatous ascomata with a dark peridium, bitunicate asci and dark 2-celled ascospores (about 10μm long) and placed in Pseudosphaeriales (= Pleosporales). Hamathecium of Testudinaceae can be cellular (e.g. Lepidosphaeria and Muritestudina) and trabeculate pseudoparaphyses (e.g. Halotestudina and Lojkania). Hawksworth & Booth (1974) considered Testudinaceae as a synonym of Zopfiaceae, but based on SEM studies of the ascospores Hawksworth (1979) regarded Zopfiaceae as a synonym of Testudinaceae. Subsequently, Eriksson (1981) accepted both families and later Zopfiaceae was validated by Eriksson & Hawksworth (1992). The phylogenetic relationships of selected coprophilous Pleosporales were investigated by Kruys et al. (2006). The molecular studies revealed that Lepidosphaeria nicotiae, Neotestudina rosatii, Ulospora bilgramii and Verruculina enalia formed a strongly supported clade in Pleosporales (Suetrong et al. 2009). These genera are known to share similar morphological features, such as dark, 1-septate ascospores with or without germ- pores, and with or without ornamentation.

Verruculina enalia was earlier treated under Didymosphaeriaceae but multi-gene phylogenetic analyses by Schoch et al. (2006) and Suetrong et al. (2009) suggested it as a member of the Testudinaceae. Molecular studies by Suetrong et al. (2009) showed that Massarina ricifera, an obligate marine species, shares a sister group relationship with U. bilgramii, N. rosatii and Quintaria lignatilis formed a sister group to Testudinaceae with weak support. Hyde et al. (2013) also accepted Testudinaceae as a family in Dothideomycetes including five genera. A freshwater genus Angustospora was introduced by Li et al. (2016a) in Testudinaceae based on its morphology and molecular phylogeny. Phookamsak & Hyde (2015) transferred Lojkania from Fenestellaceae to Testudinaceae based on its morphological similarities with Verruculina. Muritestudina was established by Wanasinghe et al. (2017c) based on its distinct hyaline, ellipsoidal, muriform ascospores, in contrast to other genera and supported by multi-gene analyses. Currently, Testudinaceae comprises Angustospora, Halotestudina, Lepidosphaeria, Lojkania, Muritestudina, Neotestudina, Testudina, Ulospora and Verruculina. The identification of taxa from Testudinaceae was mainly based on a few uncertain morphological characters and limited molecular data. Hence this family requires fresh collections in order to provide molecular data and better taxonomic assignment.