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Summary: Mycotaxon 135 (1)—Mycobiota new to www.mycotaxon.com
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Uploaded — February 2020
Macrofungi from the Hebron and
Jerusalem Hills of Palestine
MAXIMUS THERESA THALER1,2*, AYSHA AL-WAHSH2,
ALEA MEUSER2, ALYSSA ROOKS2, MAZIN QUMSIYEH2
1Department of Biological Sciences, Binghamton University,
4400 Vestal Parkway East, Binghamton, NY 13902-6000, USA
2Palestine Museum of Natural History, Palestine Institute for Biodiversity and Sustainability,
Mar Andreas campus, Bethlehem University, PS-90907 Bethlehem,
Occupied Palestinian Territories
*CORRESPONDENCE TO: maximusthaler@gmail.com
ABSTRACT—This study is based on specimens of macrofungi collected from biodiversity
hotspots in the southern West Bank of occupied Palestine during a four-week survey
period in December and January 2018–19. We identified 27 macrofungi species
representing 39 genera from six field sites. Samples were collected, photographed, and
archived in the Palestine Museum of Natural History herbarium. Species descriptions,
field site details, and a cladogram of the observed fungi are provided. The importance of
citizen science and accessible taxonomic inquiry is also discussed.
KEYWORDS—diversity, taxonomy
Introduction
Macrofungi are a polyphyletic fungal group which have spore-bearing
structures visible to the naked eye (mushrooms, brackets, puffballs, cup fungi)
and there are estimates of 53,000–110,000 species worldwide (Mueller et al et
al. 2007). Macrofungi typically fall under one of three clades: Basidiomycota,
Ascomycota, and (occasionally) Zygomycota. Finer cladistic resolution,
especially at the species level, is notoriously difficult to obtain with certainty
but advances in DNA sequencing technology have expanded knowledge
dramatically (Hibbet et al. 2016). But as our phylogenetic knowledge of
macrofungi is increasing, climate change, habitat loss, and other
anthropogenic forces are driving many species to extinction before they are
even described (Lees and Pimm 2015).
Macrofungi diversity in most western Asian regions remains understudied,
but in recent years a handful of studies have been published in the region
including Iran (Amoopour et al. 2016), Turkey, (Kaya 2009, Acar et al. 2015),
Iraqi Kurdistan (Suliaman et al. 2017, Toma et al. 2013), Saudi Arabia, (Abou-
Zeid & Altalhi 2006), Jordan, (Al Momany 2018), Egypt (Abdel-Azeem 2010),
and Syria (Abdel-Hafez 1983). Many of these studies comment on the paucity
of mycological data and need for additional studies.
Indeed, some key regions remain completely unstudied. For example, in a
recent survey of Morchella and Helvella in Israel, Barseghyan & Wasser
(2008) shows collections mostly in the Galilee and coastal areas of Historic
Palestine but largely excluded collections from the occupied West Bank; and
the few samples included from the region were all collected prior to 1971. The
ecology of the occupied West Bank has deteriorated significantly in the
decades since, making a thorough diversity survey of the area more urgent than
ever. This current study focuses on macrofungi diversity in selected vulnerable
areas of the Southern part of the occupied Palestinian territories.
Since 2014, the Palestine Museum of Natural history has been working to
fill this gap in our ecological knowledge of the region. Teams of citizen
scientists, made up of native Palestinians and international researchers, have
conducted surveys throughout those pockets of biodiversity remaining in the
West Bank. The importance of this is twofold: Firstly, as the region has
experienced unprecedented human population growth, the ecosystem services
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provided by undeveloped areas have become increasingly valuable. The most
common landscape of our surveying was semi-feral olive terraces, where olive
trees from a previous era were interspersed with wild scrub plants. These
terraces provide a home to so far uncounted numbers of fungal species, whose
impacts on soil fertility (and therefore olive harvests) are likely substantial and
varied. Secondly, in a region so starkly maligned by issues of land access and
ownership, few Palestinians are easily able to experience the landscape outside
of developed urban centers. This research provides opportunities for
Palestinians to develop a relationship with their environment which is rooted
in an understanding of the living world.
Materials and Methods
Mushrooms were photographed and collected from six areas in the southern part of
West Bank (Fig. 1) over a period of 24 days, from December 13, 2018 to January 6
2019. Our main sampling areas were the steep valleys west of Bethlehem, which are
noted biodiversity hotspots (Garstecki et al. 2010) but are also endangered by political
turmoil (Abdallah et al. 2011, Isaac & Hilal 2011).
Brief notations on each study site with longitudes and latitudes
1) WADI QUFF – 31.579896, 35.038929
Wadi Quff protected area of 2500 dunums is located in the western part of the
Hebron Governorate, and consists of two confluent valleys between Beit Kahil and
Tarqumiya. The habitat is dominated by oak (Quercus caliprinos) and introduced pine
trees (Pinus halepensis). The area has been studied for plants and animals, (see
Qumsiyeh et al. 2016) but never for fungi.
2) AL QARN – 31.619556, 35.127159
Al-Qarn is a hill of 50 dunums overlooking Beit Ummar town and Al Aroub
Refugee camp in the Hebron District. Its habitat is similar to Wadi Quff but there are
more Arbutus andrachne (strawberry tree) and less introduced pine trees.
3) WADI HUSAN – 31.717958, 35.130733
This valley is not considered a protected area like the previous two localities.
However, it is part of the biodiversity rich valley system that connects areas 3, 4, & 5.
4) AL MAKHROUR – 31.718557, 35.156388
Wadi Al-Makhrour is a valley located about 7 km south of the old city of Jerusalem
and about 6 km northeast of the old city of Bethlehem. It is connected to a larger valley
system which stretches southwest of Jerusalem. Water flows from the Walaja and
Cremisan valleys and then converges with waters from Al Makhrour itself (between
Beit Jala, Al-Khader, and Al-Walaja), to then drain into Battir and then Husan (locality
3 above) and Nahhalin valleys. Al Makhrour is the central part of the system that refills
the water aquifer of Bethlehem District area. Its many fresh water springs irrigate old
growth olive groves. Al Makhrour is also recognized as a UNESCO world heritage site
because of its rich agricultural heritage and beautiful ancient landscape.
5) CREMISAN VALLEY – 31.728257, 35.175298
Cremisan Monastery is the reason this valley was named. The valley belongs to the
Beit Jala and Al-Walaja areas but much of it was taken for colonial Israeli settlements.
The lower parts of the valley, and those closest to Beit Jala are cultivated (almonds,
apricots, olives, figs). Highland regions of the valley are protected from cultivation by
inaccessibility and contain natural habitats rich in live oak and native flowering annuals.
6) PMNH MAR ANDREA – 31.717711, 35.205431
This site of about 12 dunums was included in the study because it is essentially an
oasis in the urban areas of Bethlehem (see Qumsiyeh et al. 2017).
Macrofungi from the Hebron & Jerusalem Hills
…
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FIG 1: Photographs of typical habitats for each sample area. 1) Wadi Quff 2) Al Qarn 3) Wadi
Husan 4) Al Makhrour 5) Cremisan Valley 6) PMNH Mar Andrea Campus
FIG 2: A map of our study sites. Numbers as in FIG 1
Survey Methods
Survey methods consisted of traveling to our selected field sites, walking across
promising areas (habitats with high moisture) and sampling for macrofungi.
Mushrooms were collected and taken back to the lab at the Palestine Museum of
Natural history for spore prints and further microscopy. Some more delicate species
were photographed and then left in the field.
Collected mushrooms were dried slowly over a period of days using solar
radiation in a well aerated hallway (window drying). Larger samples were dried in
a conventional toaster oven set to 100 degrees Fahrenheit in one-hour long
increments until dry. Once fully dried, samples were numbered, labeled, and stored
in plastic for archival purposes. Mushrooms were identified using guidebooks and
consulting websites such as www.mushroomexpert.com. We received many useful
identifications through discussions and photo uploads on online
mycological communities like www.mushroomobserver.org. Photographs of
mushrooms were uploaded to mushroomobserver.org under the username
“maximusthaler”, and tagged with PMNH herbarium numbers, for future reference.
(https://mushroomobserver.org/observer/observations_by_user/6055).
Where appropriate, samples were examined using a Labomed optical light
microscope. Slides of spores were prepared using distilled water, and photographed
through a 100x oil immersion lens. We were unable to gather genetic data from these
samples, and so all of our species level identifications remain somewhat tentative, and
contingent upon higher quality molecular analysis. While we are confident in the
species level identifications we provide, in some cases the information available to us
was only sufficient to identify the sample in question to the genus level (see results).
Specimens are kept at the Palestine Museum of Natural History fungi collection
(PMNH-F).
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Results
We found 39 genera of mushrooms and were able to identify the following
27 macrofungi species:
Agaricus campestris L. (1753)
MYCOBANK 356498
COLLECTION: Al Qarn PMNH-F-1019
ECOLOGY: Saprotrophic, grows in grass (Kuo 2018a). Our samples were found on
a mixed woodland hillside.
DESCRIPTION: Cap 3–6 cm wide, white, with fine hairs sometimes visible. Gills free
from stem, pale pink aging to brown, covered with a partial veil in button stage. Stem
3–5 cm tall, 1–3 cm wide, with white remnants of a partial veil in a ring. Spores globose
to subglobose.
COMMENTS: This species is a dominant mushroom in grazing areas (Roberts and
Evans, 2011, Abate 1999), hence its common name of Horse Mushroom. This species
is characterized by a white cap. Its lamella has initially a pinkish color and eventually
becomes dark brown. It can be further identified by a negative KOH reaction (Mitchell
and Walter 1999, Kerrigan et al. 2005).
FIG 3. Agaricus campestris L. (1753)
Amanita ovoidea (Bull.) Link (1833)
MYCOBANK 156329
COLLECTION: Al Makhrour PMNH-F-1011, Cremisan (photographed)
DESCRIPTION: Cap 15–25 cm, convex, flattening with age, creamy white, with veil
fragments hanging from the cap margin when young. Gills free from the stem, crowded,
white. Stem 7–15 cm tall, 3–6 cm wide, white, with powdery ring of veil remnants,
emerging from a white volva underground.
ECOLOGY: Ectomycorrhizal; found under deciduous trees, notably oaks, sometimes
olive, on lime or alkaline soil (O’Reilly 2016). Our samples were found emerging deep
from the ground in a recently plowed olive grove.
Macrofungi from the Hebron & Jerusalem Hills
…
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COMMENTS: This species is part of a larger, more ambiguous group: Amanita sect.
Lepidella. Our samples may actually be Amanita gilbertii or one of several other A.
sect. Lepidella members. While we are confident in this identification of Amanita
ovoidea, A. sect. Lepidella remains underspecified and difficult to differentiate. Given
the understudied nature of the occupied Palestinian bioregion, there is a distinct
possibility that this sample belongs to some other A. sect. Lepidella species. Molecular
analysis (outside the scope of the current study) is required for completely confident
identification.
FIG 4. Amanita ovoidea (Bull.) Link (1833)
Arrhenia rickenii (Hora) Watling (1989)
MYCOBANK 125095
COLLECTION: Al Makhrour PMNH-F-1052
DESCRIPTION: Cap 5–25 mm wide, pale brown, convex funnel shaped, radially lined.
Gills decurrent, widely spaced, sometimes branching near cap margins. Stem 5–30 mm
long, 1–3 mm wide.
ECOLOGY: Saprophytic, nearly always on mosses on alkaline soil (O’Reilly 2016).
Our samples were found on moss-covered limestone gravel.
FIG 5. Arrhenia rickenii (Hora) Watling (1989)
Coprinellus micaceus (Bull.:Fr.) Vilgalys, Hopple & Jacq. Johnson (2001)
MYCOBANK 474361
COLLECTION: PMNH Mar Andrea PMNH-F-1037
DESCRIPTION: Cap 2–4 cm, radially lined, amber, covered in fine mica-like flakes.
Gills attached to the stem, pale at first, then darkening and dissolving with age into
black ink. Stem 2–5 cm long, 2–4 mm thick, pale amber, ridged.
ECOLOGY: Saprotrophic, grows in clusters on decaying wood. Its substrate is often
buried, causing the mushrooms to appear terrestrial (Kuo 2008a). Our samples were
found in grass at the base of an almond tree.
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FIG 6. Coprinellus micaceus (Bull.:Fr.) Vilgalys, Hopple & Jacq. Johnson (2001)
Coprinopsis friesii (Quél.) P. Karst. (1872)
MYCOBANK 318302
COLLECTION: Al Makhrour PMNH-F-1053
DESCRIPTION: Cap 5–15 mm, radially lined, grey, dissolving with age into black
ink. Gills free, black. Stem 1–3 cm long, 1–2 mm thick.
ECOLOGY: Saprotrophic, can be found in grass (Wood and Stevens 2015). This
sample was found in a grassy olive grove.
COMMENTS: This species is part of a larger, more ambiguous group Coprinopsis
sect. Picacei, 'stirps Friesii', and the specific identification as Coprinopsis friesii is
tentative. Species in this group can only be accurately identified using molecular
methods beyond the scope of this study. The name Coprinopsis friesii should be viewed
mostly as a placeholder until higher fidelity analysis can occur.
FIG 7. Coprinopsis friesii (Quél.) P. Karst. (1872)
Coprinus comatus (O.F.Müll.) Pers. (1797)
MYCOBANK 148667
COLLECTION: Al Makhrour PMNH-F-1042
DESCRIPTION: Cap 3–7 cm wide, cylindrical when young, then expanding and
dissolving into black ink with age. Cap surface white, tending to dark tan in the center,
with radially flaking scales. Gills free from the stem, initially white, aging to black ink.
Stem 5–12 cm long, 1–2 cm wide, separates easily from the cap.
ECOLOGY: Saprotrophic, grows in grass, on wood chips, or hard-packed ground
(Kuo 2008b). Our samples were found at the base of pines
Macrofungi from the Hebron & Jerusalem Hills
…
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FIG 8. Coprinus comatus (O.F.Müll.) Pers. (1797)
Cortinarius infractus (Pers.) Fr. (1838)
MYCOBANK 218869
COLLECTION: Al Qarn PMNH-F-1067
DESCRIPTION: Cap 4–10 cm wide, olive brown, convex, flattening with age, sticky
when young. Gills attached to the stem, densely packed, aging to deep orange brown
as spores mature, with weblike remnants of orange cortina hanging. Stem 4–8 cm long,
2–3 cm wide. Spores subglobose.
ECOLOGY: Mycorrhizal, found in association with hardwoods (Kuo 2012). Our
samples were found near Quercus and Arbutus.
COMMENTS: Cortinarius is one of the largest fungal genera, containing over 1000
species, many of which can only be identified molecularly (Kuo 2012). Cortinarius
distribution and diversity in the eastern Mediterranean is understudied, and molecular
phylogenetic research has shown this genus is significantly more complex than
previously thought (Garnica et al. 2003). Mycorrhizal host interactions with
Cortinarius can be extremely species specific, and there are likely Cortinarius species
with ecologies specific to the region which remain undescribed. While we are
reasonably confident in our determination of the species of this sample to be C.
infractus, it should be emphasized that our use of the name more accurately refers to a
group of closely related fungi rather than a unique species. Genetic analysis, and closer
analysis of our sample’s ecological interactions with its host are both required for a
more confident, regionally specific identification to be established. Brondz and Høiland
(Brondz and Høiland 2009) showed that Cortinarius infractus contains many alkaloid
compounds in the 5-Hydroxytriptophane synthesis pathway (5-HTP is an over the
counter antidepressant). Habitats containing Cortinarius infractus are thus of notable
biomedical importance.
FIG 9. Cortinarius infractus (Pers.) Fr. (1838)
Cryptomarasmius corbariensis (Roum.) T.S. Jenkinson & Desjardin (2014)
MYCOBANK 561778
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COLLECTION: Al Makhrour PMNH-F-1059
DESCRIPTION: Cap 2–3 mm, amber, radially lined. Gills white, free from stem,
widely spaced. Stem 2–4 cm tall, <1 mm wide, shiny black.
ECOLOGY: Saprotrophic, grows on rotting leaves of olive and other trees (Bozok et
al. 2018). Our samples were found growing on damp olive leaves.
FIG 10. Cryptomarasmius corbariensis (Roum.) T.S. Jenkinson & Desjardin (2014)
Cyathus olla (Batsch) Pers. (1800)
MYCOBANK 215509
COLLECTION: Al Qarn PMNH-F-1012
DESCRIPTION: Peridium 1 cm high, 1 cm wide. Cone shaped, with margins trumpet
shaped, cracking with age. Outer surface grey and rough. Inner surface silvery and
smooth. Peridioles flat, circular, lentil shaped, grey, with mycelium emerging centrally
from a lateral side and connecting to the base of the peridium.
ECOLOGY: Saprotrophic; can be found on the ground, in grass, woody debris, or
dead plant stems (Kuo, 2014). Our sample was found in grass in a mixed wood hillside.
FIG 11. Cyathus olla (Batsch) Pers. (1800)
Geopora arenosa (Fuckel) S. Ahmad (1978)
MYCOBANK 314400
COLLECTION: Cremisan PMNH-F-1014, Wadi Quff (photographed)
DESCRIPTION: Cup 1–2 cm across. Inner surface white, smooth. Outer surface
brown, rough. Margin irregularly cracked. Stem absent.
ECOLOGY: Thought to be mycorrhizal (O’Reilly 2016). Our samples were found
closely associated with moss, on limestone soils.
Macrofungi from the Hebron & Jerusalem Hills
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FIG 12. Geopora arenosa (Fuckel) S. Ahmad (1978)
Helvella lacunosa Afzel (1783)
MYCOBANK 147441
COLLECTION: Husan valley (PMNH-F-1065), Cremisan valley, Wadi Quff (photographed)
DESCRIPTION: Cap 2–3 cm, irregularly shaped, sometimes resembling a saddle,
black. Stem 3–4 cm long, 1–2 cm wide, grey, deeply ribbed with multiple holes.
ECOLOGY: Ectomycorrhizal, associated with oaks and pines (Nguyen et al. 2013).
Our samples were found near live oak.
COMMENTS: This species is common, and has global distribution. However,
Nguyen et al. (2013) note that what has historically been considered a single species
can actually be divided into several distinct clades. Nguyen has described new Helvella
species unique to western North America, and identifiable by ectomycorrhizal host. H.
vespertina is associated with pine, while H. dryophila is associated with oak. While
molecular data continues to corroborate the global distribution of Helvella lacunosa
(Nguyen 2013 describes some Helvella samples from Minnesota and Japan as 99%
genetically similar), it is highly likely that there are unidentified species being
misidentified as Helvella lacunosa in less studied regions, such as occupied Palestine.
Given the extreme partner specificity common in ectomycorrhizal fungi, it seems
possible that what we have identified as H. lacunosa could actually be a new Helvella
species, adapted to the live oak and pine species native to Palestine. Genetic analysis is
required to determine the relationship between our samples and H. lacunosa collections
from elsewhere in the world.
FIG 13. Helvella lacunosa Afzel (1783)
Hypomyces cervinigenus Rogerson & Simms (1971)
MYCOBANK 315654
COLLECTION: Wadi Quff PMNH-F-1071
DESCRIPTION: Powdery, pink and white aging to brown.
ECOLOGY: Parasitic on other fungi, this ascomycete is exclusively found growing
on Helvella species (Kuo 2006).
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COMMENTS: This species has global distribution and is found wherever its host,
Helvella lacunosa, is found. However, recent studies have noted that H. lacunosa is
actually several distinct species, and these distinctions have ramified into isomorphic
speciations among its parasites (Nguyen et al. 2013). Given the extreme host specificity
of Hypomyces, combined with the ectymycorizial partner specificity of its Helellva host,
it seems probable that what we have identified as Hypomyces cervinigenus could
actually be a new Hypomyces species, adapted to the (potentially also unique) local
Helvella population native to Palestine. Genetic analysis is required to determine the
relationship between our samples and Hypomyces collections from elsewhere in the
world.
FIG 14. Hypomyces cervinigenus Rogerson & Simms (1971)
Irpex lacteus (Fr.) Fr. (1828)
MYCOBANK 177211
COLLECTION: Al Qarn PMNH-F-1000
DESCRIPTION: Fruiting body white, mixed with shades of brown. Underside spore
surface developing small toothlike structures.
ECOLOGY: Saprotrophic, grows in clumps upon the undersides of fallen hardwood
(Kuo, 2007). Our samples were found on hardwood in a mixed woodland hillside.
FIG 15. Irpex lacteus (Fr.) Fr. (1828)
Lentinus arcularius (Batsch) Zmitr. (2010)
MYCOBANK 543135
COLLECTION: Al Makhrour PMNH-F-1006
DESCRIPTION: Cap 1–3 cm, usually convex, light brown covered in darker brown
concentric scales. Pores 0.5 – 2mm, smaller on cap margins and on the stem apex. Stem
2–4 cm long, 2–4mm wide. Dark brown at the top, whiter towards the base.
ECOLOGY: Saprobic, grows on decaying deciduous wood, often oak. Sometimes
these mushrooms grow from buried wood and appear terrestrial. Our samples appeared
terrestrial and were found near live oak.
Macrofungi from the Hebron & Jerusalem Hills
…
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COMMENTS: Detailed anatomy and morphological development were studied by
Hibbet et al. (1993). Some samples found in occupied Palestine lack the conspicuous
hairs on the cap fringe which North American records attest to (Kuo 2008c), and in this
way the samples more closely resemble L. brumalis. But pore structure and ecology
(samples were found amidst Quercus) make L. arcularius more likely. There is some
possibility that this is a new species or sub-species (or at least a notable phenotypic
pattern) unique to the region. Genetic analysis is required to test this hypothesis.
FIG 16. Lentinus arcularius (Batsch) Zmitr. (2010)
Lepista sordida (Schumach.) Singer (1951)
MYCOBANK 299524
COLLECTION: Al Makhrour PMNH-F-1020, Cremisan (photographed)
DESCRIPTION: Cap 3–8 cm, weakly convex, flattening with quickly with age. Lilac
purple when young, aging to pale tan. Gills free from the stem, crowded, also lilac
purple aging to pale tan. Stem 2–4 cm tall, 1–2 cm wide, lilac-tan, appressed-fibrillose.
Spores spheroid, with thin walls.
ECOLOGY: Saprotrophic, in open woody areas, usually where leaf litter collects and
rots (O’Reilly 2016). Our samples were found at the base of olive and oak.
FIG 17. Lepista sordida (Schumach.) Singer (1951)
Lycoperdon perlatum Pers. (1797)
MYCOBANK 220647
COLLECTION: Al Makhrour PMNH-F-1009
DESCRIPTION: Fruiting body 2–3 cm wide, inverted pear shaped, tan, covered in
small spines. Interior white, aging to olive. Spores emerge from a spilt at the apex of
the fruiting body with age. Spores globose, smooth walled.
ECOLOGY: Saprotrophic; can be found on the ground or on deadwood, or brush.
These samples were found on bare soil surrounded by loose twigs and leaf mulch.
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FIG 18. Lycoperdon perlatum Pers. (1797)
Omphalotus olearius (DC.) Sing. (1948)
MYCOBANK 288943
COLLECTION: PMNH Mar Andrea PMNH-F-1061
DESCRIPTION: Cap 8–12 cm, orange, appressed-fibrillose, Bioluminescent when
very young. Gills orange, decurrent. Stem 4–6 cm long, 1–2cm wide.
ECOLOGY: Saprotrophic; grows on stumps, buried roots, or on the base of
hardwoods, especially oaks and olive (Kuo, 2015b). This sample was found growing at
the base of olive.
FIG 19. Omphalotus olearius (DC.) Sing. (1948)
Peziza badia Pers. (1800)
MYCOBANK 141966
COLLECTION: Al Qarn PMNH-F-1013, Wadi Quff (photographed)
DESCRIPTION: Cup 1–2 cm across, pale brown, darker and smoother on the inner
surface. Margin inturned. Stem absent.
ECOLOGY: Saprotrophic, can be found on compacted heavy soils, particularly forest
footpaths (O’Reilly 2016). Our sample was found in a mixed wood hillside.
Macrofungi from the Hebron & Jerusalem Hills
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FIG 20. Peziza badia Pers (1800)
Psathyrella bipellis (Quél.) A.H.Sm. (1946)
MYCOBANK 438921
COLLECTION: Al Makhrour PMNH-F-1024, Cremisan PMNH-F-1064
DESCRIPTION: Cap 2–4 cm, radially lined, hygraphanous deep brown aging to dull
tan. Gills attached to the stem, deep brown. Stem 4–7 cm long, 2–5 mm thick.
ECOLOGY: Saprotrophic; grows in groups on lawns or in decaying plant matter (Kuo
2011). Our samples were found in damp decaying leaves, mostly olive and oak.
FIG 21. Psathyrella bipellis (Quél.) A.H.Sm. (1946)
Pseudoplectania nigrella (Pers.) Fuckel (1870)
MYCOBANK 188201
COLLECTION: Wadi Quff PMNH-F-1073
DESCRIPTION: Cup <1 cm across, black. Inner surface smoother than outer surface.
Stem absent.
ECOLOGY: Saprotrophic, grows on decaying pine debris (Dennis 1969). Our
samples were found in mossy limestone soils amid pine, cedar, and oak.
COMMENTS: This species has global distribution, and has been described in regions
between the Caribbean, South America, and Japan (Dennis 1954, Mardones-Hidalgo &
Iturriaga 2011, Paden 1983). It has also previously been described in Israel (Barseghyan
and Wasser 2008). Barseghyan and Wasser (2008) note that Israel has high biodiversity
of Pezizomycetes; its varied terrain and proximity to both northern Africa and western
Asia supports Pezizomycetes from both continents, 115 species in total. We too noted
a high diversity of Pezizomycetes in the field site of this collection, Wadi Quff. We
found five Pezizomycetes species in this small protected area, of which we were able to
identify three. This species is notable for its blackish cup or disc shaped apothecia (Korf
1973, Le Gal 1953, Medel et al. 2006). The mushroom forms cup shaped and as it ages
the disc flattens and sometimes can even become planar (Seaver 1913). Seaver (1913)
demonstrates that the cup margin has very small hairs, 5 mm to 1.5 cm in diameter.
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FIG 22. Pseudoplectania nigrella (Pers.) Fuckel (1870)
Sarcomyxa serotina (Pers.) P. Karst. (1891)
MYCOBANK 304280
COLLECTION: Cremisan PMNH-F-1080
DESCRIPTION: Cap 3–10 cm, kidney shaped, light brown. Gills white, adnate. Stem
reduced pseudostem.
ECOLOGY: Saprotrophic, grows on hardwoods and occasionally conifers (Kuo
2017). Our samples were found at the base of live oak.
FIG 23. Sarcomyxa serotina (Pers.) P. Karst. (1891)
Suillus collinitus (Fr.) Kuntze (1898)
MYCOBANK 306571
COLLECTION: Cremisan PMNH-F-1007, Mar Andrea & Al Makhrour (photographed).
DESCRIPTION: Cap 4–11 cm, covered in an easily pealed brown cuticle which is
viscid when damp. Stem 3–7 cm tall, 1–2 cm wide, often covered in brown markings.
Spore surface pale yellow, aging to brown. Spores irregularly ellipsoidal, some with
pointed ends.
ECOLOGY: Mycorrhizal, associated with pines, and prefers limestone soils (Assyov
2018). Our samples were all found within 10 meters of a pine species.
Macrofungi from the Hebron & Jerusalem Hills
…
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FIG 24. Suillus collinitus (Fr.) Kuntze (1898)
Tapinella panuoides (Batsch) E.-J. Gilbert (1931)
MYCOBANK 253796
COLLECTION: Cremisan Valley PMNH-F-1018
DESCRIPTION: Cap 3–6 cm, light brown, fan shaped, with fine hairs along the fringe.
Stem absent. Gills often crimped, white to beige. Spores ellipsoid, smooth walled.
ECOLOGY: Saprotrophic, grows on conifers (Kuo 2015a). Our samples were found
at the base of fallen pine.
COMMENTS: This species is fairly common, and an occasional subject of biomedical
research. Schneider et al (2008) have isolated atromentin compounds, and their
associated genes from this species. These compounds have been shown to have
antibiotic and anti-cancer properties (Zheng et al. 2006, Kim & Lee 2009). Thus
ecosystems containing Tapinella panuoides are of notable biomedical importance.
FIG 25. Tapinella panuoides (Batsch) E.-J. Gilbert (1931)
Trametes hirsuta (Wulfen) Pilát (1939)
MYCOBANK 267192
COLLECTION: Cremisan PMNH-F-1001
DESCRIPTION: Fruiting body forming bracket shelves 3–6 cm wide. Shelf surface
covered in concentric rings of white to brown hairs. Pore surface forms maze-like
structures close to the origin of growth, which condense into deep pores towards cap
margins.
ECOLOGY: Grows on stumps and fallen hardwood (Kuo 2010). Our samples were
found on fallen hardwood (likely almond) in an olive grove.
COMMENTS: Puri et al. (2006) found that it could be utilized as a novel source of
aryl tetralin lignans, which are important compounds used for the synthesis of
topoisomerase inhibitors. Habitats containing Trametes hirsuta are thus of notable
biomedical importance.
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Thaler & al.
FIG 26. Trametes hirsuta (Wulfen) Pilát (1939)
Tubaria furfuracea (Pers.) Gillet (1876)
MYCOBANK 197499
COLLECTION: Wadi Quff PMNH-F-1029
DESCRIPTION: Cap 1–2 cm, orange. Convex at first and broadening with age.
Radially lined at the edge where cap flesh is thinner. Gills adnate, orange. Stem hollow,
2–5 cm long, 2–4 mm wide.
ECOLOGY: On twigs and other woody debris (O’Reilly 2016). Our samples were
found amidst pine.
COMMENTS: Common name is scurfy twiglet.
FIG 27. Tubaria furfuracea (Pers.) Gillet (1876)
Volvopluteus gloiocephalus (DC.) Vizzini, Contu & Justo (2011)
MYCOBANK 518592
COLLECTION: Al Makhrour PMNH-F-1003
DESCRIPTION: Cap 5–10 cm, brown, darkening to black at the center. Sometimes
sticky when young. Appressed-fibrillose and radially lined towards the margins where
cap flesh is thinner. Gills free from the stem, crowded fairly densely. Pale at first but
quickly darkening to a deep brown. Stem 6–13 cm long, 2–3 cm wide, enlarging at the
base. Grey, covered with a fine white powder. Stem emerges from a white volva. Spores
ellipsoidal.
ECOLOGY: Saprotrophic, terrestrial, in grassy areas or composting organic matter
(Justo et al. 2011a). Our samples were found in thick grass.
COMMENTS: Kuo (2018b) notes that samples from California have streaked greyish
caps, while samples on the eastern side of North America are whiter. All of the samples
we found in occupied Palestine have streaked brown caps and deep brown gills. Justo
et al. (2011b) notes that the species has global distribution and it is unsurprising that
superficial morphology would vary with geography. There are likely underlying
genetic and environmental causes (and sub-species level taxonomic patterns) which are
responsible for this morphological variation. Genetic analysis of our samples is
Macrofungi from the Hebron & Jerusalem Hills
…
17
required to determine how phylogenetically distinct they are from V. gloiocephalus
elsewhere in the world.
FIG 28. Volvopluteus gloiocephalus (DC.) Vizzini, Contu & Justo (2011)
Xerocomellus redeuilhii Simonini, Gelardi & Vizzini (2016)
MYCOBANK 818381
COLLECTION: Al Makhrour PMNH-F-1005
DESCRIPTION: Cap 2–5 cm, red to yellow and cracking with age. Stem 3–4 cm tall,
1–2 cm wide, yellow with red streaks, becoming more concentrated at the base. Spore
surface yellow.
ECOLOGY: Mycorrhizal, associated with hardwoods, often oaks. Found on
calcareous soil (Simonini et al. 2016). These samples were found at the base of live oak.
COMMENTS: The taxonomy of the red capped, yellow spore surfaced boletes is in
an unusually large state of flux. Several genera, including Xerocomellus and
Hortiboletus, have been segregated from Boletus sensu lato, and distinguishing
between these genera can often only be accomplished with genetic data. While we are
reasonably confident in our identification of this sample as Xerocomellus redeuilhii,
other species in the genus, or perhaps even Hortiboletus remain possible alternatives.
Differentiating between these closely related taxa is not within the scope of this
diversity study. Genetic analysis is required for full confidence in this species
identification.
FIG 29. Xerocomellus redeuilhii Simonini, Gelardi & Vizzini (2016)
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Thaler & al.
Other taxa
In addition to the species detailed above, we made many observations of
fungi that could only be identified accurately to the genus level. Table 1 reports
all the genera observed, whether we were able to identify to species level or
not. In total, we recorded 39 genera, from 29 families (7 orders, 3 classes, 2
phyla) with at least 59 species (27 identified to the species level), from six field
sites over of 24 days, from December 13 2018 to January 6 2019.
TABLE 1. Observations /collections of mushrooms identified to the genus level.
GENUS
NAME
NUMBER OF
SPECIES
OBSERVATION IDENTIFICATION NUMBERS
Agaricus
2
PMNH-F-1019, PMNH-F-1004
Amanita
1
PMNH-F-1011
Arrhenia
1
PMNH-F-1052
Auricularia
1
PMNH-F-1016
Clavaria
1
PMNH-F-1017
Clitocybe
>5
PMNH-F-1015, PMNH-F-1025, PMNH-F-1027, PMNH-F-1036,
PMNH-F-1044, PMNH-F-1046, PMNH-F-1049, PMNH-F-
1050, PMNH-F-1051, PMNH-F-1062, PMNH-F-1078, PMNH-
F-1083, PMNH-F-1084, PMNH-F-1086
Coprinellus
4
PMNH-F-1022, PMNH-F-1037, PMNH-F-1039, PMNH-F-1060
Coprinopsis
1
PMNH-F-1053
Coprinus
1
PMNH-F-1042
Coriolopsis
1
PMNH-F-1023
Cortinarius
2
PMNH-F-1067, PMNH-F-1074
Cryptomarasmius
1
PMNH-F-1059
Cyathus
1
PMNH-F-1012
Geopora
1
PMNH-F-1014
Gomphidius
1
PMNH-F-1079
Helvella
1
PMNH-F-1065
Hypomyces
1
PMNH-F-1071
Inocybe
>5
PMNH-F-1031, PMNH-F-1032, PMNH-F-1033, PMNH-F-1034,
PMNH-F-1035, PMNH-F-1038, PMNH-F-1040, PMNH-F-
1043, PMNH-F-1047, PMNH-F-1056, PMNH-F-1068, PMNH-
F-1081, PMNH-F-1082
Irpex
1
PMNH-F-1000
Lactarius
1
PMNH-F-1070
Lentinus
1
PMNH-F-1006
Lepista
1
PMNH-F-1020
Lycoperdon
1
PMNH-F-1009
Melanoleuca
>1
PMNH-F-1028, PMNH-F-1030, PMNH-F-1075
Mycena
1
PMNH-F-1026
Nolanea
1
PMNH-F-1085
Omphalotus
1
PMNH-F-1061
Parisola
1
PMNH-F-1010
Peziza
1
PMNH-F-1013
Psathyrella
>5
PMNH-F-1021, PMNH-F-1024, PMNH-F-1041,PMNH-F-1045,
PMNH-F-1048
PMNH-F-1054, PMNH-F-1057, PMNH-F-1058, PMNH-F-1063,
PMNH-F-1064, PMNH-F-1069
Pseudoplectania
1
PMNH-F-1073
Sarcomyxa
1
PMNH-F-1080
Suillus
2
PMNH-F-1007, PMNH-F-1076
Stereum
1
PMNH-F-1077
Tapinella
1
PMNH-F-1018
Trametes
1
PMNH-F-1001
Tubaria
>2
PMNH-F-1008, PMNH-F-1029, PMNH-F-1066, PMNH-F-1072
Volvopluteus
1
PMNH-F-1003
Xerocomellus
1
PMNH-F-1005
TOTAL
>59
PNMH-F-1000 – PNMH-F-1086
DISCUSSION
In four weeks of surveying in December and January 2018–19, we
identified 27 macrofungi species from six field sites in the southern West Bank.
We made further observations of fungi which could be identified to the genus
level, for a total record of 39 genera containing at least 59 species. This survey
represents the first attempt since the British mandate period (nearly a century
ago) to document the diversity of macrofungi in this region of historic
Palestine (Reichert 1940). While this survey is by no means exhaustive, it
nonetheless makes a significant contribution to our understanding of the
Macrofungi from the Hebron & Jerusalem Hills
…
19
biodiversity of this understudied region. Climate change, combined with eight
decades of geopolitical unrest, have synergistically degraded Palestinian
ecosystems at an unprecedented rate (Qumsiyeh 2018). Thus, it is now more
important than ever to document the fungal species present in the region. As
climate change and the occupation continue to interact, the macrofungal
species diversity will very likely be reduced in future decades. This in turn will
almost certainly impact agricultural yields, particularly in areas which have
been traditionally used as village commons that provide polycultures of
minimally managed food sources.
The Palestine Museum of Natural History is building its capacity in
mycological work starting from the interest of a small staff and volunteers with
limited resources. This study represents an example of grassroots Citizen
Science, the value of which has become more widely appreciated in recent
years (Silverton 2009). Silverton notes that “projects that fit uneasily into the
standard model of hypothesis-testing research” still often have great value.
Thoroughly documenting the diversity of the region is critical for any
conservation efforts, as it is widely acknowledged that diversity is the primary
indicator of the wellbeing of ecological systems (Magurran 1988).
As this research continues, and more sophisticated methods are
implemented, we expect the number of unique species identified to grow. Our
study mostly used ecological markers and macroscopic morphological features.
Microscopy was used sparingly for the analysis of spore prints, and no genetic
data were gathered. As we implement molecular methods in future studies,
we expect to find that some fungi initially identified as a common species with
Eurasian or global distribution will actually prove to be locally adapted
variants. Most keys available were developed for Europe or North America,
and can only be applied superficially to Palestinian fungi. Indeed, because
global fungal diversity remains poorly understood and many species are yet to
be described (Mueller et al. 2007), it would be surprising if there weren’t
unique Palestinian species to be found.
For example, our observations point to two morphological oddities
(already described in the results section) which merit further inquiry: 1) North
American descriptions of Volvopluteus gloiocephalus describe it as having
grey or white caps, and light-colored gills, while the Palestinian samples we
found have brown cap and dark brown gills. This discrepancy likely has
phylogenetic significance. 2) North American descriptions of Lentinus
arcularus describe the mushroom as exhibiting a hairy fringe along the cap,
but the Palestinian samples we found were mixed (some with fringes and some
without). Future studies should compare North American and European
samples to the Palestinian specimens we collected to clarify these issues.
Figure 30 shows the phylogeny of the genera of macrofungi we identified
in Palestine. It is remarkable that they represent 29 families in a preliminary
study like this. With the advent of molecular methodologies, we will proceed
now to do DNA sequence analysis on expanded surveys of mushrooms from
throughout the region (and not just from the described areas of the southern
West Bank). For example, we did not sample in areas like the Jordan Valley
and the arid regions of Palestine which include Irano-Turanian, Saharo
Arabian, and Ethiopian-Sudanese phytogeographic elements (see Soto-
Berelov et al. 2015).
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Thaler & al.
FIG 30: A phylogeny of all macrofungi observed during the study
Future research should continue our efforts to map macrofungal diversity
in occupied Palestine, with the ultimate aim of generating a thorough
distribution map of all genera and species present in the region. Furthermore,
it should be emphasized that for these types of diversity studies, the
collaborative methods used are just as important as the results. While our
methods were simple, they allowed for a passionate community of citizen
scientists to re-open the field of Palestinian mycology after nearly a century of
neglect.
Macrofungi from the Hebron & Jerusalem Hills
…
21
Acknowledgments
Many thanks to all of the Palestine Museum of Natural History staff and volunteers
who contributed in ways big and small to this study. Jesse Qumsiyeh, Elias Handal,
Reena Saeed, Mohammed Najajrah, Mohammed Abusarhan, Ilona Baniusevic, Gabija
Kirkilaite, and Victor Chapinal Agudo. Identification assistance was provided by
Dimitar Bojantchev, Allan Rockefeller, and Davide Puddu. This research was made
possible in part by a generous grant from the National Geographic Society and the
Darwin Initiative.
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