This is a species living in an extreme environment. There is unregulated international trade in this species. Levels of trade are not known, but are clearly large and growing. The fungus is also threatened in some places by road and urban development and by the effects of war.

AFRICA: Algeria, Egypt, Libya, Morocco, Tunisia. ASIA: Bahrain, Cyprus, Iran, Iraq, Israel, Kuwait, Saudi Arabia, Syria, Turkey, Yemen. ATLANTIC OCEAN: Spain (Islas Canarias). EUROPE: Italy (Sardinia, Sicily), Spain. Native throughout its recorded range. This fungus was one of several illustrated on postage stamps of Djibouti in 1987 (MOSS & DUNKLEY, 1988); it is possible that the fungus occurs there, but no verifiable record was found. Inconclusive evidence suggests this species may also extend into Mauritania and Western Sahara in the west (VOLPATO ET AL., 2013), and into Oman in the east [www.squ.edu.om/Portals/33/almasar/Horizonnew207.pdf, accessed 3 October 2013]. No information about altitudinal distribution has been found.

Over 30 records from scientific sources (specimens, databases and bibliographic sources combined, excluding duplicates) from at least April 1888 to April 2009, with observations in February, March, April and May. Most of the evidence used in the present study derives from collections or market purchases for scientific study. There is almost no information currently available from rural collector-suppliers, or from their commercial buyers about geographical distribution and abundance, and high quality information from such sources is necessary for evaluating threats and conservation status.

Not known. Said to be rare in Spain (MORENO ET AL., 2002), infrequent in Turkey (AKYÜZ ET AL., 2012), and abundant in Israel (KAGAN-ZUR & ROTH-BEJERANO, 2008). There is no information about population levels in north Africa and most of southwest Asia, the main areas of distribution of this species.

MINTER (2013) evaluated this species globally, using IUCN Categories & Criteria, as Vulnerable.

This fungus is associated with a wide variety of different organisms. ANIMALIA. Jaculus jaculus L., 1758, J. orientalis Erxlebel 1777; Suilla gigantea Meigen 1830. FUNGI. Alternaria alternata (Fr.) Keissl., A. tenuissima (Kunze) Wiltshire; Aspergillus flavus Link, A. fumigatus Fresen., A. niger Tiegh., A. ochraceus G. Wilh., A. terreus Thom; Chaetomium bostrychodes Zopf; Cladosporium herbarum (Pers.) Link; Cochliobolus australiensis (Tsuda & Ueyama) Alcorn [as Drechslera australiensis Bugnic. ex M.B. Ellis], C. hawaiiensis Alcorn [as Drechslera hawaiiensis Bugnic. ex M.B. Ellis], C. lunatus R.R. Nelson & F.A. Haasis [as Curvularia lunata (Wakker) Boedijn]; Fusarium oxysporum Schltdl.; Melanocarpus albomyces (Cooney & R. Emers.) Arx [as Thielavia albomyces (Cooney & R. Emers.) Malloch & Cain]; Mucor circinelloides Tiegh., M. hiemalis Wehmer; Neocosmospora vasinfecta var. africana (Arx) P.F. Cannon & D. Hawksw.; Penicillium digitatum (Pers.) Sacc.; Picoa lefebvrei (Pat.) Maire [as Phaeangium lefebvrei Pat.]; Rhizomucor pusillus (Lindt) Schipper [as Mucor pusillus Lindt]; Rhizopus microsporus Tiegh., R. stolonifer (Ehrenb.) Vuill.; Sordaria sp.; Terfezia claveryi Chatin, T. metataxasi Chatin; Tirmania nivea (Desf.) Trappe, T. pinoyi (Maire) Malençon; Ulocladium chartarum (Preuss) E.G. Simmons, U. tuberculatum E.G. Simmons. MONERA. Azotobacter chroococcum Beijerinck; Bacteria indet. PLANTAE. Artemisia monosperma Delile; Cistus albidus L., C. incanus L. (mycorrhizal), C. monspeliensis L., C. salviifolius L.; Fumana procumbens Gren. & Godr.; Halimum halimifolium (L.) Willk.; Helianthemum apenninum Mill., H. canariense Pers., H. eremophilum Pomel, H. hirtum var. deserti Cosson & Durieu, H. kahiricum Delile, H. ledifolium (L.) Mill., H. lippii (L.) Dum. Cours., H. monspeliense [not traced], H. salicifolium (L.) P. Mill., H. salviifolium [not traced], H. sessiliflorum (Desf.) Pers. (mycorrhizal), Helianthemum sp. (mycorrhizal); Kobresia bellardii (All.) Degel. (mycorrhizal); Plantago albicans L.; Schismus barbatus (L.) Thell.; Tuberaria guttata (L.) Fourr.

It forms mycorrhizas with a range of flowering plants. AL-WHAIBI (2009) provided a general review of desert plants and mycorrhizas. In Iran ascomata of T. boudieri occur about 5–10 cm below the soil surface, and field and laboratory studies have shown that it forms ectomycorrhizas with Kobresia bellardii (AMMARELLOU ET AL., 2007; AMMARELLOU & SAREMI, 2008). In Tunisia, experiments with H. sessiliflorum plants showed that individuals bearing mycorrhizas of T. boudieri grew higher, with greater leaf production and higher levels of N, P and K than those without (SLAMA ET AL., 2012). The same experiments also found enhanced growth on gypsum soils compared with sandy soils. Depending on conditions, T. boudieri may form endomycorrhizas or ectomycorrhizas with Cistus incanus (ZARETSKY ET AL., 2006a; ZARETSKY ET AL., 2006b). Studies in Saudi Arabia (BOKHARY & PARVEZ, 1992a; BOKHARY & PARVEZ, 1992b; BOKHARY ET AL., 1990) and Egypt (MOHAWED ET AL., 2001) have shown that ascomata of T. boudieri are associated with many other mostly ascomycetous fungi, and with bacteria (ROUGIEUX, 1963). ROUGIEUX (1963) noted a fungal-microbial-plant interaction in the field involving T. boudieri and Azotobacter chroococcum, a free-living nitrogen-fixing bacterium, with microbial populations in general much higher near ascomata than in soil 50 cm away, and with A. chroococcum present only near ascomata. In the laboratory, T. boudieri extracts stimulated growth of the Azotobacter, but inhibited Staphylococcus aureus Rosenbach 1884. CHATIN (1891) observed that fruitbodies of T. boudieri provide food and shelter for two species of the rodent genus Jaculus (desert jerboas). SLAMA (2010) reported that in Tunisia the fly Suilla gigantea feeds on ascomata. ROTH-BEJERANO ET AL. (2004) demonstrated that mycelial growth of this fungus is affected by levels of calcium. Physical and chemical qualities of soils associated with T. boudieri have been studied in Turkey (AKYÜZ ET AL., 2012). Species of Terfezia and Tirmania need a certain minimum amount of precipitation in a given year before they produce ascomata. In Kuwait, that minimum was reported to be 180 mm well distributed from October through to March (AWAMAH & ALSHEIKH, 1979). Terfezia boudieri has been been recorded from the following habitats: amenity & protected areas (national parks); coastal (maritime sands); desert (arid scrub, dunes, semi-desert); grassland.

In desert truffles wind is an important agent of distribution: ascomata of desert truffles have large thin-walled cells in the peridium and gleba. These take up water during rare moist conditions, permitting swelling of the underground ascomata and maturation of ascospores. This results in cracks in the soil surface. When the normal drier conditions resume, the covering sandy soil is blown away by the wind and the ascomata are exposed. The thin-walled cells of the drying fruitbodies collapse into powdery fragments exposing the spores, which are then released by abrasion from blowing sand, and become wind borne (TRAPPE ET AL., 2008).

Living in the difficult environment of dry deserts, this species is adapted to survive at levels of heat and water stress which would be very unfavourable for other fungi. As a result, it already lives in conditions near the limit for sustainable life. Climate change and global warming in particular are likely to be significant long-term threats. Deserts are ecosystems on which humans tend to place little monetary value. Habitat destruction through war, irrigation, development of recreational facilities such as golf courses, disturbance of soil (for example by tourist safaris using 4-wheel drive vehicles), construction of solar energy facilities, establishment of refugee camps, and similar developments are all likely to threaten the ecosystems where this fungus occurs. MOUBASHER (2010) reported that, in Egypt, of the two main areas known for desert truffles, one on the Mediterranean coast west of Alexandria was being destroyed by construction of factories, recreational areas, retirement homes, roads, tourist villages and similar developments, while the other, in the Sinai Peninsula was seriously disturbed by military activities. ALSHEIKH (1989) reviewed the serious and damaging impact of war on desert truffle populations in Kuwait in the 1990s, and in other parts of southwest Asia and north Africa during the 20th century. In many areas where this species occurs, social upheaval and war continue. Paradoxically, the civil unrest experienced in many countries where this fungus occurs may help to protect it: the possibility of encountering landmines is likely to deter many collectors (ALSHEIKH, 1989). Pollution is also sometimes a serious threat. The firing of Kuwait oil wells as an act of war in 1991 affected populations of desert truffles over a wide area (ALSHEIKH, 1989). Nothing is known about the effect of oil pollution in soils on these fungi, but it is likely to be long-term. Ascomata of T. boudieri are known to accumulate radionucleotides (GUILLÉN GERADA, 2002), but the impact of this on the species is not known. Formerly, harvesting of this species as a wild crop was carried out by rural populations for sustainable local consumption, but in the past twenty years there has been a significant increase in commercial harvesting in connexion with international trade,and the impact of this has never been evaluated, although there are reports that harvesting by refugees as their sole form of income is having a negative impact on populations (VOLPATO ET AL., 2013). There has been considerable interest in the possibility of cultivating desert truffles. If that happens, there will be the danger that a few genotypes favourable to cultivation will be used, and these may swamp the wild populations resulting in a loss of genetic diversity. At a local level, threats listed by GRAVITO HENRIQUES (2012a) for T. arenaria (overgrazing, trampling, and encroachment of scrub etc.), are also likely to apply to the present species.

Awareness of the importance of these desert truffles is very low even at governmental level: a survey of the most recent relevant national action plans and reports for the Rio Convention on Biological Diversity [www.cbd.int/nbsap/search/default.shtml, accessed 4 October 2013] by Algeria, Bahrain, Egypt, Georgia, Greece, Iran, Iraq, Israel, Italy, Jordan, Kuwait, Lebanon, Libya, Mauritania, Morocco, Oman, Portugal, Qatar, Romania, Saudi Arabia, Spain, Syria, Tunisia, Turkey, and the United Arab Emirates (all countries from which species of Terfezia have been recorded) showed that only Morocco and Saudi Arabia had any conservation plans. Both of these countries recognized that there might be a problem of over-exploitation; Saudi Arabia also described this species as of high conservation priority, and expressed concern about unregulated harvesting and damage by off-road use of vehicles. There is thus a strong need to make the appropriate conservation authorities in these countries more aware of the conservation needs of this species.

There is a significant and international on-line trade in these fungi. Ascomata of Terfezia and Tirmania not identified to species level are collectively marketed on the Internet as “desert truffles”, and commercial websites exist offering them for sale. On one site the price was €35-75 per kilogram with a minimum order of 100 kilograms and a claim by the vendor that 1000 kilograms per day could be supplied [www.alibaba.com/showroom/fresh-desert-truffles.html, accessed 29 October 2013]. The trade is secretive, with no easy access to addresses, and at most only very general information about the sources of the product. The English language websites are likely to be only a small part of the total market, and much of the trade and negotiations now seems to be conducted in Arabic through social networking sites like Facebook (G. SOLIMAN, pers.comm.). Much more information about this trade is needed.

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