True truffle diversity in Iran




Iran, true truffles, Tuber borchii, Tuber brumale, Tuber macrosporum, Tuber rufum, Tuber excavatum, Tuber fulgens


Inside commercial batches of T. aestivum imported from Iran to Italy the ascomata of six Tuber spp. (Tuber borchii, Tuber brumale, Tuber macrosporum, Tuber rufum f. lucidum, Tuber excavatum and Tuber fulgens) were found and described. Sixteen specimens were analyzed; most of them were completely immature probably because truffles in Iran are harvested by digging all the soil forest without using trained dogs. The morphology of the Iranian ascomata and their spore dimensions are similar to those of the European ascomata and phylogenetic analyses based on their ITS sequences placed them inside the European clades of Tuber. In Iran, truffles are found in the northern part of the country with the highest production in the Hyrcanian region (between Guilan and Golestan provinces). To date, Iran seems to be the most Eastern location where European species of Tuber are found.


Ammarellou A, Alvarado P (2018) First report of Tuber aestivum var. uncinatum from Iran based on morphological and molecular characteristics. Rostaniha 18 (2):227-228

Badalyan S, Hovsepyan R, Iotti M, Zambonelli A (2005) On the presence of truffles in Armenia. Flora Mediterranea 15:683–692.

Bagi I, Fekete O (2010) Identification of Tuber aestivum habitats in the South Caucasus, Azerbaidjan. Communication at 2nd congress of the Tuber aestivum/uncinatum. European Scientific Group, 20-22.8.2010, Juva, Finland.

Bahram M, Põlme S, Kõljalg U, Zarre,S ,Tedersoo L (2012) Regional and local patterns of ectomycorrhizal fungal diversity and community structure along an altitudinal gradient in the Hyrcanian forests of northern Iran. New Phytologist 193:465-473.

Benucci GMN, Csorbai AG, Falini LB, Marozzi G, Suriano E, Sitta N, Donnini D (2016) Taxonomy, Biology and Ecology of Tuber macrosporum Vittad. and Tuber mesentericum Vittad. In: True truffle (Tuber spp.) in the world (Zambonelli A, Iotti M, Murat C, eds) Soil Biology 47, Springer International Publishing, Cham, Switzerland, pp. 69-86.

Bonito G, Gryganskyi A, Vilgalys R, Trappe JM (2010) A global meta-analysis of Tuber ITS rDNA sequences: Species diversity, host specificity, and long-distance dispersal. Molecular Ecology 19(22):4994–5008.

Bonito G, Smith ME, Nowak M, Healy RA, Guevara G, Cázares E, Kinoshita A, Nouhra ER, Domínguez LS, Tedersoo L, Murat C, Wang Y, Moreno BA, Pfister DH, Nara K, Zambonelli A, Trappe JM, Vilgalys R. (2013) Historical biogeography and diversification of truffles in the Tuberaceae and their newly identified southern hemisphere sister lineage. PLoS One 8(1):e52765.

Bonuso E, Zambonelli A, Bergemann S E, Iotti M, Garbelotto M (2010) Multilocus phylogenetic and coalescent analyses identify two cryptic species in the Italian bianchetto truffle, Tuber borchii Vittad. Conservation Genetics 11(4):1453-1466.

Büntgen U, Egli S, Camarero JJ, Fischer E M, Stobbe U, Kauserud H, Tegel W, Sproll L, Stenseth NC (2012) Drought-induced decline in Mediterranean truffle harvest. Nature Climate Change 2:827–829.

Ceruti A, Fontana A, Nosenzo C (2003) Le specie europee del genere Tuber: una revisione storica, vol 37. Museo Regionale di Scienze Naturali, Torino.

Chen J, Li SC, Li JM, Yan B, Liu PG (2018) Distribution, biological diversity, and conservation of Chinese black truffle. Scientis Sinica Vitae 48(3):319-331.

Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9:772

Gardes M, Bruns TD (1993) ITS primers with enhanced specificity for basidiomycetes-application to the identification of mycorrhizae and rusts. Molecular Ecology 2:113–118.

Glime JM, Wagner DM (2017) Laboratory techniques: preservation and permanent mounts. In: Bryophyte ecology (Glime JM, ed) Vol. 3. Methods. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Available from:

Gógán Csorbai A, Bratek Z, Dimény J (2007) A new tool for rural development: truffle cultivation. Cereal Research Communications 35(2):413–417.

Gross G (1987) Zu den europäischen Sippen der Gattung Tuber. In: Atlas der Pilze des Saarlandes (Derbsch H, Schmitt JA, eds), Teil 2, Verlag der Derlattinia, Saarbrücken, Germany, pp 79–100.

Hall IR, Brown GT, Zambonelli A (2007) Taming the truffle. The history lore and science of the ultimate mushroom, Timber, Portland.

Hall TA (1999) BioEdit: a user-friendly biological sequence alignment program for Windows 95/98/NT. Nucleic Acids Symposium Series 41:95–98

Hamza B (1994) Investigation and identification of plant communities and constituent elements of Lesakuti forests (third series) in the south of Tonekabon. Forest and Rangeland Research Institute report, 129. Ministry of Agriculture, Tehran, Iran [in Persian].

Healy RA, Bonito GM, Smith ME (2016) A brief overview of systematics, taxonomy and ecology of the Tuber rufum clade. In: True truffle (Tuber spp.) in the world (Zambonelli A, Iotti M, Murat C, eds) Soil Biology 47, Springer International Publishing, Cham, Switzerland, pp. 125–136.

Iran Meteorological Organization (IRIMO) (2021) The average 10-years climatic parameters [in Farsi].

Jamali S (2017) First report of identification and molecular characterization of Tuber aestivum in Iran. Agroforest System 91:335–343.

Læssøe T, Hansen K (2007) Truffle trouble: what happened to the Tuberales? Mycological Research 111:1075–1099.

Le Tacon F, Marçais B, Courvoisier M,. Murat C, Montpied P, Becker M (2014) Climatic variations explain annual fluctuations in French Périgord black truffle wholesale markets but do not explain the decrease in black truffle production over the last 48 years. Mycorrhiza 24:115–125.

Leonardi M, Iotti M, Pacioni G, Hall I, Zambonelli A (2021) Truffles; biodiversity, ecological significances, and biotechnological applications. In: Industrially important fungi for sustainable development (Abdel-Azeem AM, Yadav AN, Yadav N, Usmani Z, eds) Biodiversity and Ecological Perspective, vol. 1, Fungal Biology. Springer International Publishing, Cham, Switzerland (in press).

Merényi Z, Varga T, Bratek Z (2016) Tuber brumale: a controversial Tuber species. In: True Truffles (Tuber spp.) in the world (Zambonelli A, Iotti M, Murat C, eds). Soil Biology 47, Springer International Publishing, Cham, Switzerland, pp. 49-68.

Merényi Z, Varga,T, Geml J, Orczán ÁK, Chevalier G, Bratek Z (2014) Phylogeny and phylogeography of the Tuber brumale aggr. Mycorrhiza 24:101–113.

Montecchi A, Sarasini M (2000) Funghi ipogei d'Europa. Associazione Micologica Bresadola. Centro studi micologici, Trento.

Pegler DN, Spooner BM, Young T.K (1993) British truffles. A revision of British hypogeous fungi. Royal Botanic Gardens, Kew.

Puliga F, Illice M, Iotti M, Baldo D, Zambonelli A (2020) Tuber iranicum, sp. nov., a truffle species belonging to the Excavatum clade. Mycologia 112(5):932-940.

Riousset L, Riousset G, Chevalier G, Bardet MC (2001) Truffes d’Europe et de Chine. INRA, Paris.

Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient bayesian phy- logenetic inference and model choice across a large model space. System Biology 61:539–542.

Silvestro D, Michalak I (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution 12:335–337.

Thompson JD, Higgins DG, Gibson TJ (1994). CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22:4673-4680.

Wedén C, Danell E, Tibell L (2005) Species recognition in the truffle genus Tuber - the synonyms Tuber aestivum and Tuber uncinatum. Environmental Microbiology 7(10):1535-1546.

White TJ, Bruns T, Lee S, Taylor JL (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: PCR Protocols: A Guide to Methods and Applications (Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds) Academic Press, San Diego, pp. 315–322.

Zambonelli A, Rivetti C, Percurdani R, Ottonello S (2000) TuberKey: a DELTA-based tool for the description and interactive identification of truffles. Mycotaxon 74(1):57-76.

Zambonelli A, Iotti M, Murat C (2016) True Truffle (Tuber spp.) in the World, Soil Biology 47. Springer International Publishing, Cham, Switzerland.




How to Cite

Puliga, F., Illice, M., Iotti, M., Leonardi, P., Baghdadi, A. ., Mozafari, A. A. ., & Zambonelli, A. (2021). True truffle diversity in Iran. Italian Journal of Mycology, 50, 52–62.