Studies in Hyaloscyphaceae associated with major vegetation types in the Canary Islands II: a revision of Hyaloscypha

Abstract: Four species of the genus Hyaloscypha are presented for the Canary Islands. The study is based on recent collections and 12 previous records. The earlier reports of the genus (Hyaloscypha fuckelii, H. hyalina and H. leuconica) are corrected. All of the reported species are new to the Canarian archipelago (H. aureliella, H. intacta, H. spiralis and H. strobicola), and only one has been reported before from the Macaronesian region (H. aureliella). A key, descriptions, illustrations and notes about ecology are provided. Citation: Quijada L., Huhtinen S., Negrín R. & Beltrán-Tejera E. 2017: Studies in Hyaloscyphaceae associated with major vegetation types in the Canary Islands II: a revision of Hyaloscypha. — Willdenowia 47: 31–42. doi: https://doi.org/10.3372/wi.47.47104 Version of record first published online on 13 February 2017 ahead of inclusion in April 2017 issue.


Introduction
The genus Hyaloscypha Boud. contains c. 38 species worldwide (Kirk & al. 2008). This generic name appeared for the first time in a floristic work of Gillet (1879), but it was really established six years later by Boudier (1885), who characterized the genus by its fleshy and broadly sessile apothecia with few hairs; cylindric-clavate, branched and septate paraphyses not protruding above the asci; and aseptate ascospores with drops. Boudier (1885) gave two species as examples, i.e. Helotium vitreolum (P. Karst.) P. Karst. and Peziza dentata Pers., and later (Boudier 1907) included 34 species, but it was Velenovský (1934Velenovský ( , 1939Velenovský ( , 1947 who totally blurred the generic limits of the genus, including 70 taxa (Svrček 1985). Also Dennis (1949Dennis ( , 1956) had apparent problems in delimiting the genus. Huhtinen (1989) included 20 species in his monograph of Hyaloscypha, five of them with two varieties each and one with three varieties. Also, a historical overview of the taxonomic problems in the genus after Boudier was given by Huhtinen (1989). Hyaloscypha vitreola (P. Karst.) Boud. had earlier been proposed as the conserved type of the generic name (Huhtinen & Cannon 1987) and was later accepted.
Although some authors have pointed out the possibility of substrate specificity (Velenovský 1934;Dennis 1949), nowadays it has been observed that the host seldom influences species differentiation. Huhtinen (1989) recognized six major ecological groups: (1) growing on softwoods (wood of gymnosperm trees such as conifers), (2) growing on hardwoods (wood of angiosperm trees, deciduous or evergreen), (3) growing on both softwoods and hardwoods, (4) restricted to oak wood, (5) inhabiting all types of litter, and (6) confined to herbaceous litter. The genus is considered widespread, mainly distributed in the temperate N hemisphere; there are also some reports from the S hemisphere in Australia, Argentina, Chile, New Zealand, Philippines, South Georgia, and Tristan da Cunha (GBIF; Huhtinen 1989). The phenology of the different species of Hyaloscypha was presented in Huhtinen (1989). He observed that fruiting appears during the whole frost-free season and tends to diminish from September onward.
Three species of the genus have been reported in the Canary Islands: Hyaloscypha fuckelii Nannf., H. hyalina (Pers.) Boud. and H. leuconica (Cooke) Nannf. in Beltrán- Tejera & al. (2004Tejera & al. ( , 2008 and Ribes (2009). The aim of this investigation is to contribute to the knowledge of the genus Hyaloscypha in the Canary Islands, providing revision of previously collected specimens, detailed descriptions, keys and ecological data.

Material and methods
Methods for collection, types of vegetation explored, and macro-and microscopic techniques for examination of apothecia follow Del Arco & al. (2010) and Quijada & al. (2015). All previously reported specimens were revised to confirm or correct their identity. Specimens are deposited at the mycological section of the herbarium of the University of La Laguna (TFC; herbarium code follows Thiers 2016+). Colour coding refers to ISCC-NBS (Anonymous 1976). Municipalities and names for localities follow IDE-Canarias visor 3.0 (http://visor.grafcan.es/visorweb/). The {number of studied specimens} is indicated in curly brackets, except if only one collection was found.
All specimens were collected in pine forests on Pinus wood except one collection (TFC Mic. 24101), and all have blunt hairs with resinous exudate and without dextrinoid reaction, and non-guttulate cylindric to suballantoid ascospores, which fit perfectly in Hyaloscypha aureliella. The short description in Beltrán- Tejera & al. (2004Tejera & al. ( , 2008 did not remark on the characteristics of hairs, but her personal notes and drawings showed the resinous exudate over the hairs, microscopically confirmed after the revision of the samples (Fig. 5). Hyaloscypha aureliella may be confused with H. fuckelii if no attention is paid to the resinous granules in the hairs, which can happen when the sample is mounted in a medium that disolves the resinous matter, but another feature helping the distinction is the presence of amyloid nodules in the excipulum of H. aureliella (but only in c. 30 % of the populations). In Ribes (2009), the resinous exudate was observed, but the sample was erroneously identified as H. fuckelii (Fig. 5 Svrček in Česká Mykol. 40: 209. 1986. -Fig. 2.
Distribution and ecology -The species has been reported in the N hemisphere in Europe (Czech Republic, Denmark, Finland, France, Germany, Luxembourg, Spain, Sweden, Russia, Ukraine, United Kingdom) and in North America (Canada). Growing on hardwoods (Betula L., Carpinus L., Castanea Mill., Populus L., Prunus L., Salix L. and Sorbus L.), apparently also one stray collection on Juniperus. Occurring in all seasons, being more abundant from autumn to spring (Svrček 1986;Huhtinen 1989;Baral 1992 Remarks -The present description fits in very well with the consulted literature (Svrček 1986;Huhtinen 1989;Hansen & Knudsen 2000;Raitviir 2004;Morozova 2014).
The main difference could be the slightly more fusiformclavate morphology in the ascospores studied by us, which can be explained as an over-mature morphology before germination. Hyaloscypha spiralis (Velen.) J. G. Han & al. in Fungal Biol. 118: 161. 2014. -Fig. 3.
Distribution and ecology -The species has been found in the N hemisphere in Europe (Italy) and North America (United States). Growing on dead cone scales of Pinus.
Remarks -Hyaloscypha strobilicola is easy to identify due to the ecology, i.e. growth on cone scales, and its morphology, i.e. minute apothecia with resinous matter in hairs, short amyloid asci with croziers, and small ascospores without guttules. The closest species is H. aureliella, but it has larger apothecia (0.2 -0.5 mm vs 0.1 -0.2 mm), hairs (to 44 µm vs *to 34.5 µm) and ascospores (*6 -9 µm vs *4 -5 µm). Hyaloscypha strobilicola had been found in the Canary Islands before this study, but the specimens were erroneously reported under H. leuconica. All the characters of our sample fit in well with the description of the holotype (Huhtinen 1989;Raitviir 2004).    Tejera & al. 2004).

Discussion
Our revision of the genus Hyaloscypha in the Canary Islands showed that all species previously reported (H. fuckelii, H. hyalina and H. leuconica) have been erroneously identified (Fig 5). None of the taxa treated in this paper had been reported before in the archipelago. The ecology of the Canarian specimens agrees with the general knowledge reported before for each species (Huhtinen 1989). Hyaloscypha spiralis has been reported mainly on hardwoods, occurring more abundantly from spring to autumn. In the Canarian archipelago it follows the same pattern, but its development seems to be mainly restricted to one type of vegetation (laurel forest), and here it is reported for the first time on the following hardwoods: Arbutus L., Erica L., Laurus L., Morella Lour. and Picconia DC. Its phenology is shorter than previously reported, restricted mainly to the spring. As Huhtinen (1989) indicated in his monograph, H. aureliella is a very common species restricted mainly to softwoods, which is the case also in the Canarian archipelago. Although the phenology of the species is different in the temperate and boreal zones, the peak of fruiting is reached between summer and autumn. In the Canary Islands this peak is reached in the period of maximum rainfall and mild temperatures, between autumn and spring. In summer, in the dry period with high temperatures, only one specimen was found. For H. intacta and H. strobilicola, we have too few collections to permit remarks on its phenology. Hyaloscypha intacta grows on hardwoods, and here we widen its distribution to lower latitudes.
All the specimens collected in the Canary Islands have been found in laurel or pine forests in middle elevations, between 750 m and 2060 m. In some species, substrate plays a major role independently from climatic conditions, such as in Hyaloscypha aureliella (Pinus wood) or H. strobilicola (cone scales). On the other hand, H. spiralis grows independently from the substrate, and is linked to the most humid areas influenced by the trade winds.