- Email: [email protected]
A ‘boaring’ tale: an interesting collection of Elaphomyces papillatus, a new British record
J!#K#LMN
A ‘boaring’ tale: an interesting collection of Elaphomyces papillatus , a new British record E !"#@! F
R
ecently (November 2017) I took the opportunity to attend the Mycena workshop that Derek Schafer organised in the Forest of Dean. I must confess that I had an ulterior motif for this visit and hope that no one else with a serious interest in Mycena was prevented from attending by my booking a place. A number of years ago I had collected immature material in the vicinity of an undescribed Pachyphlodes. This later became an isotype of P. nemoralis Hobart, Bona & Conde (see Healy et al. 2015) so I was hoping I might get better material. On the afternoon of arrival I visited the site and was shocked by the absolute devastation caused by the wild boar. The numbers of these
animals must have expanded hugely since I first visited the area as it was churned up to within inches of the tree trunks. The forestry commission report (https://www.forestry.gov.uk/forestry/infd9fyfc5 ) states that these feral boar (Fig. 1) have five to ten piglets per breed; breed three times a year and become fertile at one year old. The official estimate for 2017 is approximately 1600 animals in the Forest of Dean, a relatively small area. There was little hope of finding the P. nemoralis in what looked like a ploughed field. I therefore used the opportunity to look a little deeper than normal, taking advantage of the well rutted soil.
## "#! # "#O#! # ""O## "#! " #! #<"# ! ! ##E !"#@!
FD6#H ! O#:!O#H " "O#H! #.! C "#H#H
doi:10.1016/j.fldmyc.2018.07.011
/-/
FM Vol19(3) new version.qxp_Field mycology 07-16-2018 12:02 Page 102
J!#K#LMN I was pretty soon rewarded by some very small collections which I almost discarded, assuming them to be immature young Elaphomyces sp., either E. muricatus or E. granulatus. Luckily, they were put into the box to check which species I had collected. Later, after looking at them carefully under the microscope, I discovered I had three species, one mature collection and two other different immature collections, all growing within 10–15 cm of each other alongside an oak tree in an oak glade in mixed deciduous woodland. The mature collection proved very interesting. My collection of Elaphomyces papillatus Vitt. This tiny species (see Fig. 2) was no bigger than a pea ( largest 8 mm across ) with a smooth peridium. It was clearly something I had not seen before, nor was it known in Britain. The cortex (rind) when I examined it closely, was finely papillate, formed by tufts of darker hyphae (see Fig. 4). On cutting I could see the material was nicely mature as the dark spores filled the cavity (see Fig. 3). The inner peridial layer had an undefined mottling of dark brown and lighter brown soft tissue (again evident in Fig. 3). The spores of this collection were amazing, ornamented with striate ridges and forming an incomplete reticulum (Fig. 5). 51 spores measured in water averaged 15–18.5 µm. I had used Paz et al. (2017) to identify the material and these spores were bigger than the description of their Spanish material but within the range of a Norwegian variety (are spores larger further north where we have more rain?). These very distinctive diagnostic features of spore ornamentation, size of spores and structure of the peridium pointed to Elaphomyces papillatus var. striatosporus which they illustrate. Luckily I was able to pass the material by Thomas Læssøe, one of the co-authors of this paper, who agreed with my determination. Paz et al (2017) is the first major revision of European Elaphomyces and is the first overview of the genus in Europe supported by a ribosomal DNA-based phylogeny. It is a long overdue and welcome addition to the literature on Elaphomyces. As well as E. papillatus they have further included two varieties: E. papillatus var. striatosporus and E. papillatus var. sulphureopallidus in their subsection Papillati. I did not
question their interpretations, nor did I fully understand the implications of the sequencing, but my collection clearly had characters from both the type variety and the variety striatosporus. My collection was sent to the Royal Botanic Gardens Kew to be deposited. It was sequenced on arrival and results matched the sequence of the epitype of E. papillatus var. papillatus and not that of the var striatosporus which I had plumped for, and embarrassingly announced publically at the BMS Autumn meeting at Kew in Nov. 2017. I was as you can imagine puzzled. This collection, made on 1 Nov. 2017, is deposited at Kew as K(M) 237621. Thoughts on the status of E. papillatus var. striatosporus This variety was first described at species rank by Kers (1984), who collected it near Oslo, Norway in 1980. His description is most helpful and supplements the summary given in Paz et al. (2017). Kers describes the ecology and the finding of small clusters in close proximity to fine roots which he believed were hazel, as well as describing the morphological features of his collection. Most of his material was found in Sweden where Corylus avellana and Quercus ruber dominated. He speculates that his species grows in undisturbed primary vegetation (something we in England are short of}. Paz and Lavoise in Paz et al. (2017) reduced Kers’ species to a variety on the basis of DNA analysis, recognising that the spore ornamentation and peridial surface differed from their designated lectotype material; a further single collection was also described as a further variety. From the Kers description I am happy that my material falls comfortably into his concept as he describes changes to the cortex as the material ages, as being “black to sepia, smooth or somewhat scurfy (not warted), in young ascocarps, covered by a thin mantle of brown hyphae, sometimes minutely and indistinctly reticulated”. Exospore ornamentation is variable “but most spores show their characteristic pattern of more or less parallel running ridges”. The peridum he describes as “greyish brown and somewhat juicy”. The Paz et al. (2017) epitype of the type variety, illustrated in their paper on page 217, clearly has a similar cortex to my collection but a
J!#K#LMN
y , m y l s e . s y n n ##" ""! # ! !##E#@!
# M# " ""! # " !# ! # "# !O# ! !##E#@!
#6#" ""! " O # !
# "# "# ! !##E#@!
#=#" ""! ! " #;## " # ! !##E#@!
#4#" # ! !##E#@!
##" ! ! # ! !##E#@!
s
.
k , l . g s . e . n
d A r e . y e l r g n y e c . d
e , a
/-'
J!#K#LMN
References
very different spore ornamentation, yet it matches my collection’s ITS sequence. The variety from Norway, also on page 217, has identical spores to mine that feature the parallel ridges but a cortex that has little grey papillae on the surface, unlike mine and whilst the ITS sequence is very close it is not a match. This species is clearly variable and there needs to be more material collected to understand it fully. I suspect that had Paz et al. seen my material prior to publishing they might well have reconsidered their decision to recognise a variety based on the morphology and only a very small difference in ITS; or they would perhaps have utilised a more extensive analysis of different genetic regions of the DNA to separate the varieties in the group more conclusively. How far apart molecularly does it need to be for a variety or species to be erected? There is presently no defined diffrence, it is open to interpretation. But usually, I’m told, one needs a good number of collections from a range of sites to get a holistic view before recognising a variety with sequences this close.
Forestry Commission (2017). http://www.forestry.gov.uk/forestry/infd9fyfc5 Healy, R., Hobart, C., Tocci, G., Bona, L., Merényi, Z., Paz Conde A. & Smith, M. (2015). Fun with the discomycetes: revisiting collections of Korf’s anamorphic Pezizales and Thaxter’s New England truffles leads to a connection between forms and the description of two new truffle species: Pachyphlodes pfisteri and P. nemoralis. Ascomycete.org 7 (6): 363. Kers, L.E. (1984). Elaphomyces striatosporus found in Sweden. Nordic Journal of Botany 4(1): 71–74. Morris, S. (2017). Forest of Dean pixies fighting against the cull of wild pigs. Guardian 27:11 available at: https://www.theguardian.com/environment /2017/oct/12/boar-war-the- forest-of-deanpixies-fighting-against-the-cull-of-wild-pigs Paz, A., Bellanger, J. M., Lavoise, C., Molia, A., Lawrynowicz, M., Larsson,E., Ibarguren, I.O., Jeppson, M., Læssøe, T., Sauve, M., Richard, F. & Moreau, P. A. (2017). The genus Elaphomyces (Ascomycota, Eurotiales): a ribosomal DNA-based phylogeny and revised systematics of European ‘deer truffles’ in: Persoonia: Molecular Phylogeny and Evolution of Fungi 38: 197–239.
And what else? So what of the other two species collected at the site that were immature? These are illustrated above (Figs 6 & 7). Figure 6 I believe to show the second collection in the UK of E. decipiens and Fig. 7 possibly E. muricatus, but the peridium in this collection was not typical of the collections I normally see. Sadly extraction of DNA was not successful from either collection. A return visit is needed to collect mature specimens. And what of the wild boar? There is a ‘Boar war’ going on (Morris, 2017); the Forestry Commission have set a “target population” of at most 400 animals. They managed a cull of 492 in 2016 including road kill figures. The ‘Forest of Dean wild boar cull saboteurs’ are, as you can imagine, actively engaging in preventing the cull, suggesting the pigs promote much needed tourism in the area. Certainly my view is they are an asset in the forest for fungal dispersal and soil aeration but perhaps not in the numbers that presently exist. Acknowledgements. Thanks go to Kare Liimatainen for sequencing my three Elaphomyces collections at RBG Kew and to Martyn Ainsworth for supporting the work.
/-!
A ‘boaring’ tale: an interesting collection of Elaphomyces papillatus , a new British record E !"#@! F
R
ecently (November 2017) I took the opportunity to attend the Mycena workshop that Derek Schafer organised in the Forest of Dean. I must confess that I had an ulterior motif for this visit and hope that no one else with a serious interest in Mycena was prevented from attending by my booking a place. A number of years ago I had collected immature material in the vicinity of an undescribed Pachyphlodes. This later became an isotype of P. nemoralis Hobart, Bona & Conde (see Healy et al. 2015) so I was hoping I might get better material. On the afternoon of arrival I visited the site and was shocked by the absolute devastation caused by the wild boar. The numbers of these
animals must have expanded hugely since I first visited the area as it was churned up to within inches of the tree trunks. The forestry commission report (https://www.forestry.gov.uk/forestry/infd9fyfc5 ) states that these feral boar (Fig. 1) have five to ten piglets per breed; breed three times a year and become fertile at one year old. The official estimate for 2017 is approximately 1600 animals in the Forest of Dean, a relatively small area. There was little hope of finding the P. nemoralis in what looked like a ploughed field. I therefore used the opportunity to look a little deeper than normal, taking advantage of the well rutted soil.
## "#! # "#O#! # ""O## "#! " #! #<"# ! ! ##E !"#@!
FD6#H ! O#:!O#H " "O#H! #.! C "#H#H
doi:10.1016/j.fldmyc.2018.07.011
/-/
FM Vol19(3) new version.qxp_Field mycology 07-16-2018 12:02 Page 102
J!#K#LMN I was pretty soon rewarded by some very small collections which I almost discarded, assuming them to be immature young Elaphomyces sp., either E. muricatus or E. granulatus. Luckily, they were put into the box to check which species I had collected. Later, after looking at them carefully under the microscope, I discovered I had three species, one mature collection and two other different immature collections, all growing within 10–15 cm of each other alongside an oak tree in an oak glade in mixed deciduous woodland. The mature collection proved very interesting. My collection of Elaphomyces papillatus Vitt. This tiny species (see Fig. 2) was no bigger than a pea ( largest 8 mm across ) with a smooth peridium. It was clearly something I had not seen before, nor was it known in Britain. The cortex (rind) when I examined it closely, was finely papillate, formed by tufts of darker hyphae (see Fig. 4). On cutting I could see the material was nicely mature as the dark spores filled the cavity (see Fig. 3). The inner peridial layer had an undefined mottling of dark brown and lighter brown soft tissue (again evident in Fig. 3). The spores of this collection were amazing, ornamented with striate ridges and forming an incomplete reticulum (Fig. 5). 51 spores measured in water averaged 15–18.5 µm. I had used Paz et al. (2017) to identify the material and these spores were bigger than the description of their Spanish material but within the range of a Norwegian variety (are spores larger further north where we have more rain?). These very distinctive diagnostic features of spore ornamentation, size of spores and structure of the peridium pointed to Elaphomyces papillatus var. striatosporus which they illustrate. Luckily I was able to pass the material by Thomas Læssøe, one of the co-authors of this paper, who agreed with my determination. Paz et al (2017) is the first major revision of European Elaphomyces and is the first overview of the genus in Europe supported by a ribosomal DNA-based phylogeny. It is a long overdue and welcome addition to the literature on Elaphomyces. As well as E. papillatus they have further included two varieties: E. papillatus var. striatosporus and E. papillatus var. sulphureopallidus in their subsection Papillati. I did not
question their interpretations, nor did I fully understand the implications of the sequencing, but my collection clearly had characters from both the type variety and the variety striatosporus. My collection was sent to the Royal Botanic Gardens Kew to be deposited. It was sequenced on arrival and results matched the sequence of the epitype of E. papillatus var. papillatus and not that of the var striatosporus which I had plumped for, and embarrassingly announced publically at the BMS Autumn meeting at Kew in Nov. 2017. I was as you can imagine puzzled. This collection, made on 1 Nov. 2017, is deposited at Kew as K(M) 237621. Thoughts on the status of E. papillatus var. striatosporus This variety was first described at species rank by Kers (1984), who collected it near Oslo, Norway in 1980. His description is most helpful and supplements the summary given in Paz et al. (2017). Kers describes the ecology and the finding of small clusters in close proximity to fine roots which he believed were hazel, as well as describing the morphological features of his collection. Most of his material was found in Sweden where Corylus avellana and Quercus ruber dominated. He speculates that his species grows in undisturbed primary vegetation (something we in England are short of}. Paz and Lavoise in Paz et al. (2017) reduced Kers’ species to a variety on the basis of DNA analysis, recognising that the spore ornamentation and peridial surface differed from their designated lectotype material; a further single collection was also described as a further variety. From the Kers description I am happy that my material falls comfortably into his concept as he describes changes to the cortex as the material ages, as being “black to sepia, smooth or somewhat scurfy (not warted), in young ascocarps, covered by a thin mantle of brown hyphae, sometimes minutely and indistinctly reticulated”. Exospore ornamentation is variable “but most spores show their characteristic pattern of more or less parallel running ridges”. The peridum he describes as “greyish brown and somewhat juicy”. The Paz et al. (2017) epitype of the type variety, illustrated in their paper on page 217, clearly has a similar cortex to my collection but a
J!#K#LMN
y , m y l s e . s y n n ##" ""! # ! !##E#@!
# M# " ""! # " !# ! # "# !O# ! !##E#@!
#6#" ""! " O # !
# "# "# ! !##E#@!
#=#" ""! ! " #;## " # ! !##E#@!
#4#" # ! !##E#@!
##" ! ! # ! !##E#@!
s
.
k , l . g s . e . n
d A r e . y e l r g n y e c . d
e , a
/-'
J!#K#LMN
References
very different spore ornamentation, yet it matches my collection’s ITS sequence. The variety from Norway, also on page 217, has identical spores to mine that feature the parallel ridges but a cortex that has little grey papillae on the surface, unlike mine and whilst the ITS sequence is very close it is not a match. This species is clearly variable and there needs to be more material collected to understand it fully. I suspect that had Paz et al. seen my material prior to publishing they might well have reconsidered their decision to recognise a variety based on the morphology and only a very small difference in ITS; or they would perhaps have utilised a more extensive analysis of different genetic regions of the DNA to separate the varieties in the group more conclusively. How far apart molecularly does it need to be for a variety or species to be erected? There is presently no defined diffrence, it is open to interpretation. But usually, I’m told, one needs a good number of collections from a range of sites to get a holistic view before recognising a variety with sequences this close.
Forestry Commission (2017). http://www.forestry.gov.uk/forestry/infd9fyfc5 Healy, R., Hobart, C., Tocci, G., Bona, L., Merényi, Z., Paz Conde A. & Smith, M. (2015). Fun with the discomycetes: revisiting collections of Korf’s anamorphic Pezizales and Thaxter’s New England truffles leads to a connection between forms and the description of two new truffle species: Pachyphlodes pfisteri and P. nemoralis. Ascomycete.org 7 (6): 363. Kers, L.E. (1984). Elaphomyces striatosporus found in Sweden. Nordic Journal of Botany 4(1): 71–74. Morris, S. (2017). Forest of Dean pixies fighting against the cull of wild pigs. Guardian 27:11 available at: https://www.theguardian.com/environment /2017/oct/12/boar-war-the- forest-of-deanpixies-fighting-against-the-cull-of-wild-pigs Paz, A., Bellanger, J. M., Lavoise, C., Molia, A., Lawrynowicz, M., Larsson,E., Ibarguren, I.O., Jeppson, M., Læssøe, T., Sauve, M., Richard, F. & Moreau, P. A. (2017). The genus Elaphomyces (Ascomycota, Eurotiales): a ribosomal DNA-based phylogeny and revised systematics of European ‘deer truffles’ in: Persoonia: Molecular Phylogeny and Evolution of Fungi 38: 197–239.
And what else? So what of the other two species collected at the site that were immature? These are illustrated above (Figs 6 & 7). Figure 6 I believe to show the second collection in the UK of E. decipiens and Fig. 7 possibly E. muricatus, but the peridium in this collection was not typical of the collections I normally see. Sadly extraction of DNA was not successful from either collection. A return visit is needed to collect mature specimens. And what of the wild boar? There is a ‘Boar war’ going on (Morris, 2017); the Forestry Commission have set a “target population” of at most 400 animals. They managed a cull of 492 in 2016 including road kill figures. The ‘Forest of Dean wild boar cull saboteurs’ are, as you can imagine, actively engaging in preventing the cull, suggesting the pigs promote much needed tourism in the area. Certainly my view is they are an asset in the forest for fungal dispersal and soil aeration but perhaps not in the numbers that presently exist. Acknowledgements. Thanks go to Kare Liimatainen for sequencing my three Elaphomyces collections at RBG Kew and to Martyn Ainsworth for supporting the work.
/-!