Corresponding author: A. Martyn Ainsworth (
Academic editor: Scott Redhead
Rigorous diagnostics and documentation of fungal species are fundamental to their conservation. During the course of a species-level study of UK waxcap (Hygrophoraceae) diversity, two previously unrecognized species were discovered. We describe
In Europe, waxcap mushrooms (
Waxcaps are regarded as nitrogen-sensitive organisms because fruiting is inhibited by applications of nitrogenous fertilizers (
Taxonomic treatments of European waxcaps have recognized from one to seven genera (e.g.
Waxcap identification in Britain and Ireland currently adheres to
Recent developments in DNA-based methods of identification (“DNA barcoding”) are revolutionizing rapid diagnosis of diversity in mushrooms and other
This paper focuses on our treatment of two unusual waxcaps that, because of their viscid pilei and subregular hymenophoral tramal hyphae, are assigned to the segregate genus
A total of 20 collections corresponding to the
Collection and voucher information for the specimens used in this study.
GenBank<br/> Accession No. | Taxon | Fungarium<br/> Accession No. | Collection code/ seq. literature ref. | Source | Notes |
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K(M)181241 | E.J.M.Arnolds WX359 | UK, Wales, Merionethshire | |
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K(M)181245 | D.J.Harries DJH064A WX663 | UK, Wales, Pembrokeshire | |
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K(M)181246 | D.J.Harries DJH064B WX664 | UK, Wales, Pembrokeshire | Holotype |
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MICH10924 | A.H.Smith 21491 | USA, Michigan, Cheboygan Co. | |
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MICH45363 | T.E.Brooks 1098 | USA, Michigan, Cheboygan Co. | |
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MICH45364 | T.E.Brooks 1099 | USA, Michigan, Cheboygan Co. | |
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K(M)121495 | E.W.Brown | Channel Islands, Jersey | as |
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K(M)166625 | N.W.Legon | UK, England, South Somerset | as |
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MICH45365 | A.H.Smith 34029 | USA, Michigan, Chippewa Co. | |
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K(M)127070 |
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UK, N. Ireland, Derry | |
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K(M)127194 |
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UK, N. Ireland, Tyrone | |
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K(M)90029 |
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UK, England, Buckinghamshire | |
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K(M)90674 |
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UK, England, East Sussex | |
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Hungary, Kétvölgy | ||
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Hungary, Apátistvánfalva | ||
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K(M)156265 | R.Winnall | UK, England, Worcestershire | Holotype |
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K(M)181115 | R.D.Foster WCS15 WX115 | UK, England, Derbyshire | |
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K(M)181116 | R.D.Foster WCS26 WX126 | UK, England, Derbyshire | |
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K(M)181117 | R.Winnall WX459 | UK, England, Worcestershire | |
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K(M)181124 | J.E.Hodges DJH055 WX535 | UK, Wales, Pembrokeshire | |
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K(M)181126 | R.Winnall WX673 | UK, England, Worcestershire | |
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K(M)181127 | R.Winnall & A.M.Ainsworth WX694 | UK, England, Worcestershire | |
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K(M)181128 | R.Winnall & A.M.Ainsworth WX695 | UK, England, Worcestershire | |
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K(M)181129 | R.Winnall & A.M.Ainsworth WX696 | UK, England, Worcestershire | |
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K(M)181227 | R.Winnall WX461 | UK, England, Worcestershire | |
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K(M)41524 | C.Lovatt | UK, England, Staffordshire |
Spore measurements are rounded to the nearest half micron and preceded by associated data in square brackets. For example, [60, K(M)181128*, K(M)181129] would indicate that 60 spores in total were measured either in water from prints (
DNA was extracted using either an enzymatic digestion-glass fiber filtration protocol in 96-well plate format with a vacuum-manifold or the Whatman FTA® card method described in
Six additional sequences labelled as
The full ITS region was amplified and sequenced for 14 specimens. Only the ITS1 region was sequenced for all specimens from MICH, while only the ITS2 region was sequenced for K(M)181124. The ITS1 and ITS2 regions were amplified and sequenced separately for K(M)181245, and the two non-overlapping regions were concatenated and separated by 66 gaps corresponding to the 5.8S ribosomal subunit in the final alignment. Phylogenetic analysis resulted in a highly resolved tree with most nodes receiving strong bootstrap support (
Maximum likelihood phylogram using full and partial nuclear ribosomal internal transcribed spacers (ITS) sequences. Numbers above branches are nonparametric bootstrap values. Tree is arbitrarily rooted at the midpoint. Two well-supported terminal clades representing the new species
UNITED KINGDOM. England. Worcestershire (vice county 37): Bewdley, Willow Bank,
Pileus 15–55 mm diam., hemispherical to broadly conical or campanulate, initially with incurved margin, becoming applanate, often retaining broad umbo and irregular, lobed outline with indentations, folds and pleats, sometimes becoming radially furrowed, or split and flared, margin faintly to strongly translucently striate to half-way and becoming reflexed to highly revolute, viscid with gelatinous pellicle, sometimes with minutely rugose texture, at first usually dull violet purple (vinaceous grey to purple) with areas of pink, darker red or red-brown tones (rose, blood red to rusty tawny), sometimes more brownish (purplish date to dark brick), becoming paler and pinkish especially around margin which can also develop yellow (luteous) or yellow-brown (fulvous) tints, hygrophanous, dried pilei characteristically pale orange (saffron) flushed pink (rose). Lamellae ventricose, mostly narrowly adnate with some free, sinuate or broadly adnate elements, intervenose, concolorous with pileus near pileal attachment, becoming paler towards free edge, sometimes with yellow (luteous) or orange (saffron) tints. Stipe 15–70 × 5–15 mm, relatively stout, sometimes tapering upwards from the clavate base, hollow, often flexuose or tortuous, compressed or grooved, viscid but usually slightly less so than pileus, white often apically tinged with pileal colour and basally yellow (luteous) to pale orange (saffron) or becoming so, sometimes with purplish (vinaceous grey) blotches if frosted. Outer tissues of context concolorous with adjacent external surfaces, inner tissues paler. Dried lamellar trama (lens) often conspicuously dark pink (coral), contrasting with paler subhymenium and lamellar surfaces. Green pigments entirely absent. Without distinctive taste or smell. Spores [120, K(M)181126*, K(M)181127*, K(M)181128*, K(M)181129*] 6.0–8.5(-9.0) × 4.0–5.5(–6.0) µm, per-basidioma mean values 7.0–7.5 × 5.0 µm, Q=1.2–2.0, mean 1.5, short-ellipsoidal to ellipsoidal, not constricted. Basidia predominantly 4-spored, clavate, relatively long and slender with long attenuated base, (37–)40–63(–67) × 6–10 µm excluding sterigmatal length (4.0–8.0 µm). Clamp connections on basidia, within lamellar trama and pileipellis often with conspicuously looped hook cells (medallion clamps). Lamellar trama subregular with some interwoven elements, compartments 24–183 × 4–24 µm. Stipitipellis and pileipellis are ixotrichoderms.
Basidiomata of
Microscopic characters of
Known from a cemetery in West Wales (Pembrokeshire) and fields in central England (Worcestershire, Staffordshire and Derbyshire). The earliest known collection was made by C. Lovatt in Staffordshire in 1996 who noted that she had recorded similar specimens in 1994. It has fruited on private land at Willow Bank (Worcestershire) almost every year from 2000 onwards and recorded there in five discrete fruiting patches in a single field of ca. 0.8 ha.
In unimproved short (grazed or mown) acid-neutral rough pasture or other grassland. This species is often a relatively late fruiter and can continue producing basidiomata in January, long after other waxcaps have finished.
Latin
Collectors noted that although basidiomata of this species resembled
United Kingdom. England. Derbyshire (vice county 57): Edale, Lower Hollins Farm,
UNITED KINGDOM. Wales. Pembrokeshire (vice county 45): Hundleton, Somerton Farm,
Pileus 10–25 mm diam., hemispherical to conical or campanulate, sometimes with incurved margin, becoming plano-convex or remaining broadly conical, often umbonate, sometimes with irregular, lobed outline, margin faintly to strongly translucently striate to half-way, viscid or at least very lubricous, sometimes partially flared, at first usually pink-brown to orange-brown (brick, rusty tawny to fulvous), margin paler sometimes with orange (sienna to apricot) tints, hygrophanous, dried pilei dull orange (saffron to rust). Lamellae ventricose, mostly narrowly to broadly adnate with some slightly decurrent elements, intervenose, concolorous with pileus near pileal attachment, becoming paler towards free edge. Stipe 12–60 × 2–8 mm, cylindrical or compressed, sometimes with clavate base, hollow, sometimes flexuose, viscid but usually slightly less so than pileus, apically concolorous with pileus, paler below, sometimes basally tinted orange (apricot). Dried lamellar trama (lens) often darker than subhymenium and lamellar surfaces. Green pigments entirely absent. Without distinctive taste or smell, although one specimen [K(M)181241*] was noted to have a faint rubbery smell reminiscent of
Basidiomata of
Microscopic characters of
Distribution of
Identified from two sites in west Wales (Merionethshire and Pembrokeshire) supported by DNA sequence data. Fruiting was probably observed at the type locality by D.J. Harries on 6 August 2009 but no material was kept.
In unimproved short acid-neutral rough pasture in Merionethshire and found fruiting on bare soil near mosses on an almost vertical south-facing earth bank on farmland in Pembrokeshire.
Named to distinguish this European taxon from the morphologically similar
Initially, it seemed likely that historic British collections assigned to
United Kingdom. Wales. Merionethshire (vice county 48): Croesor, Cnicht,
The traditional
One of the novel taxa,
In Europe,
Our analysis showed that the ITS sequence derived from the holotype specimen of
We would like to thank Defra, Natural England and Scottish Natural Heritage for financial support and all those who collected and sent specimens of the two species described herein: E.J.M. Arnolds, R.D. Foster, D.J. Harries, J.E. Hodges and R. Winnall. Thanks also to R.D.F., D.J.H. and R.W. for allowing us to use their photographs.