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: Armenia, Azerbaijan, Bahrain, Cyprus, Georgia, Iran, Iraq, Israel, Jordan, Kuwait, Lebanon, Qatar, Saudi Arabia, Syria, Turkey, Turkmenistan. ATLANTIC OCEAN: Spain (Islas Canarias). EUROPE: Italy (mainland and Sardinia), Portugal, Spain. Native throughout its recorded range. 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.

Not known. Over 130 records from scientific sources (specimens, databases and bibliographic sources combined, excluding duplicates) from at least 1892 to March 2004, with observations in January, February, March, April, May, June, November and December. 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. Rare in Morocco (MALENÇON, 1973). Widespread and abundant from north Africa to southwest Asia, except Egypt (ALSHEIKH, 1995). Common in the Qatar desert (AL-THANI, 2010).

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

This species is associated with many other organisms. ANIMALIA. Oryx leucoryx Pallas 1777. FUNGI. Alternaria alternata (Fr.) Keissl.; Aspergillus flavus Link, A. fumigatus Fresen., A. niger Tiegh., A. terreus Thom; 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.; Mucor circinelloides Tiegh.; Penicillium digitatum (Pers.) Sacc., P. dupontii Griffon & Maubl. [as Talaromyces thermophilus Stolk], P. notatum Westling; Picoa lefebvrei (Pat.) Maire [as Phaeangium lefebvrei Pat.]; Rhizopus stolonifer (Ehrenb.) Vuill., Rhizopus sp.; Scopulariopsis brevicaulis (Sacc.) Bainier; Setosphaeria rostrata K.J. Leonard; Sordaria sp.; Terfezia boudieri Chatin; Tirmania nivea (Desf.) Trappe, T. pinoyi (Maire) Malençon; Ulocladium tuberculatum E.G. Simmons. MONERA. Bacteria indet. PLANTAE. Acacia hebeclada DC; Artemisia herba-alba Asso, A. monosperma Delile; Atractylis serratuloides (Cass.) DC.; Carex stenophyllum Wahlenb. (mycorrhizal); Cistaceae indet.; Cistus albidus L. (mycorrhizal), Cistus sp.; Fumana procumbens Gren. & Godr. (mycorrhizal); Helianthemum almeriense Pau (mycorrhizal), H. apenninum Mill. (mycorrhizal), H. eremophilum Pomel, H. kahiricum Delile (mycorrhizal), H. lavandulifolium Mill., H. ledifolium (L.) Mill. (mycorrhizal), H. lippii (L.) Dum. Cours. (mycorrhizal), H. salicifolium (L.) P. Mill. (mycorrhizal), H. viscarium Boiss. & Reut. (mycorrhizal); Pinus halepensis Mill.; Plantago albicans L.; Quercus coccifera L.; Thymelaea hirsuta (L.) Endl.; Tuberaria guttata (L.) Fourr. [as Helianthemum guttatum (L.) P. Mill.] (mycorrhizal).

This species forms mycorrhizas with a range of flowering plants. AL-WHAIBI (2009) provided a general review of desert plants and mycorrhizas. General aspects of the mycorrhiza formed by T. claveryi were described by DEXHEMIER ET AL. (1985). FORTAS & CHEVALLIER (1992) showed that in Algeria this fungus forms two different types of mycorrhiza with Tuberaria guttata [as Helianthemum guttatum], depending on soil fertility: ectomycorrhizas in phosphorus rich substrata, and ectendomycorrhizas in phosphorus deficient substrata. MORTE ET AL. (1994) developed techniques for in vitro production of mycorrhizas between this fungus and Helianthemum almeriense, later refined by GUTIÉRREZ ET AL. (1996), and GUTIÉRREZ ET AL. (2003) provided a further account of some other aspects of these mycorrhizas. In vitro studies of mycelium of Terfezia claveryi show that it is well adapted to tolerate water stress (NAVARRO-RÓDENAS ET AL., 2011). The function of aquaporins in roots of H. almeriense is affected by drought conditions which simultaneously trigger changes in the type of mycorrhiza formed with T. claveryi, and it has been postulated that this is a factor in drought response regulation in the plant (NAVARRO-RÓDENAS ET AL., 2012b; NAVARRO-RÓDENAS ET AL., 2013). The type of mycorrhiza formed between this fungus and that plant is also influenced by levels of organic phosphorus (NAVARRO-RÓDENAS ET AL., 2012a). JAMALI & BANIHASHEMI (2013) have shown that it is possible to use nested PCR techniques to identify the presence of T. claveryi in roots and therefore to demonstrate mycorrhizal associations with Helianthemum species in Iran. This species prefers calcareous soils, where T. arenaria (Moris) Trappe prefers acid soils (HONRUBIA ET AL., 1992).

Studies in Saudi Arabia (BOKHARY & PARVEZ, 1992a; BOKHARY & PARVEZ, 1992b; BOKHARY ET AL., 1990), Egypt (MOHAWED ET AL., 2001) and Iran (JAMALI & BANIHASHEMI, 2012b) have shown that ascomata of T. claveryi are associated with many other mostly ascomycetous fungi, and with bacteria. These include glucophilic and halophilic species, fungi which can produce mycotoxins, and some potential plant pathogens (JAMALI & BANIHASHEMI, 2012b). Nothing seems to be known about the relationships between these species, and their ecological role. The fungus has been associated with the endangered Arabian oryx in that occurs within areas set aside to protect that species (OSTROWSKI ET AL., 1998); this has been viewed as a problem for oryx conservation because of intrusion into the reserve by local people to collect ascomata; there seems to have been no consideration of the fungus as an important component of the natural habitat of the animal which may itself need protection. 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). This species has been recorded from the following habitats: amenity & protected areas (national parks); coastal (maritime sands); desert (arid scrub, dunes, semi-desert); grassland.

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. 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. Appropriate conservation authorities in these countries need to be made more aware of the need to protect this species.

Much more information is needed about the current unregulated international trade in this species.

This species is prized for its culinary qualities: a popular account with information about some ways in which it can be prepared was provided by LOIZIDES ET AL. (2012). CHATIN (1891) and CHATIN (1892) reported that, in a good season, up to 20 camel loads per day could arrive at the market in Damascus. Factors affecting possible commercial production of this species were explored in Morocco (BOUZIANI, 2009). 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.).

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