塊菌屬

塊菌屬
科学分类
界：	真菌界 Fungi
门：	子囊菌门 Ascomycota
纲：	盤菌綱 Pezizomycetes
目：	盤菌目 Pezizales
科：	塊菌科 Tuberaceae
属：	塊菌屬 Tuber; P.Micheli（英语：P.Micheli） ex F.H.Wigg.（英语：F.H.Wigg.） (1780)
模式種
	Tuber aestivum; Vittad.（英语：Vittad.） (1831)
異名
	Tuber P.Micheli (1729); Tuberium Raf. (1815); Aschion Wallr.（英语：Wallr.） (1833); Ensaluta Zobel (1854); Lespiaultinia Zobel (1854); Oogaster Corda (1854); Tuber subgen. Vittadinion Zobel (1854); Vittadinion Zobel (1854); Terfeziopsis Harkn.（英语：Harkn.） (1899); Delastreopsis Mattir.（英语：Oreste Mattirolo） (1905); Mukagomyces S.Imai (1940)

塊菌屬（学名：Tuber），又稱西洋松露屬，為子囊菌門塊菌科的一屬，本屬目前已知222種，其中有幾種可以烹調，且價位極高^[2]。

描述

子囊果地下生（英语：Hypogeal），不具囊蓋（inoperculate），形態上成球狀、瘤塊狀，且略不規則。外表隨種類不同而有不同表現，可能佈滿金字塔型或圓錐形的突起鱗片，如黑松露（Tuber melanosporum）；也可能如義大利白松露（Tuber magnatum）呈平滑表面。顏色從黑色、紫黑色、紅棕色、黃褐色至灰白色不等^[3]。子囊果的巨觀構造可以大致分為最外層的包被（perido）和內部的產孢腔（gleba）^[4]。包被外層由外囊盤被（ectal excipulum）所構成，周邊皮質則由緻密的囊盤被髓質（medullary excipulum）構成。皮質常為假薄壁（pseudoparenchymatous）組織形態，有些則為長軸（prosenchymatous）組織形態^[3]。

產孢腔切面則可見因子囊盤的重疊壓縮所造成的迷宮樣緻密曲折紋路，這些紋路稱為脈（venae）。脈又分為外脈（venae externae）^{[註 1]}及顏色較白內脈（venae internae）^[3]^{[註 2]}。外脈為子實層，負責產生子實，組織內含有空氣；內脈不具生殖功能，在發育初期顏色呈白色，隨著個體逐漸成熟逐漸變深，其材質較為緻密且不含氣體^[3]。

子囊內含一至八顆孢子。本屬的孢子多有複雜紋飾，可分為兩類：一類有許多的棘狀突起，此類以黑孢松露為代表；另一類則具有蜂巢般的格紋，義大利白松露屬於此型^[4]。

生物學

一般盤菌的子囊前端有囊蓋（operculate），成熟時囊蓋會打開並釋出孢子。但塊菌生長於地下，又無法彈射孢子，因此會散發濃郁氣味，吸引哺乳動物挖掘食用，藉此散佈孢子^[5]。會食用塊菌的動物包括熊及豬，因此前人常使用豬的嗅覺來尋找塊菌，在母豬嗅聞到塊菌的位置後，再將松露掘出。但後來因為豬的力氣較大不易控制，因此改用狗來尋找塊菌^[5]。

種間關係

塊菌屬屬於菌根真菌，常會與其他植物產生菌根^[5]。歐洲最常見的共生喬木包含橡樹和榛樹。在亞洲則主要與主要與壳斗科、樺木科及松科等樹種的根系共生^[5]^[6]。

過往認為塊菌無法單獨生存，必須與其他植物形成共生性的外生菌根，以吸收孢外間質才能生存^[7]。但後來發現某些塊菌（如黑冬松露（英语：Tuber melanosporum））也會攝取環境的資源，甚至會使一些植物的根部退化，藉此吸收其殘留下來的營養和礦物質。因此塊菌與植物間的種間關係並不僅有互利共生，可能也會出現片利共生、寄生，及死體營養（necrotrophy）等交互關係^[7]。目前研究支持塊菌屬於輔助性腐生（auxiliary saprotrophic nutrition）生物，當植物的根系密集時，塊菌採取腐生模式，吸收孢外間質的養分；根系較稀疏時，塊菌會開始掠奪周邊的資源。然而，自由生長的塊菌由於缺乏營養供給，因此無法形成子囊果，且未必能與其他物種競爭^[7]。

除喬木的外生菌根之外，某些塊菌也被發現可以與蘭科植物形成蘭菌根，也能與杜鵑花類植物形成漿果鵑類菌根（arbutoid mycorrhizas）^[8]。

下属物种

本属包括以下物种：^[9]

夏块菌 Tuber aestivum (Wulfen) Spreng.
白脐块菌 Tuber alboumbilicum Y. Wang & Shu H. Li
Tuber anniae W. Colgan & Trappe
Tuber argenteum Gilkey
Tuber argentinum Speg.
Tuber arnoldianum Healy, Zurier & Bonito
阿萨块菌 Tuber asa-foetida
Tuber aztecorum G. Guevara, Bonito & M.E. Sm.
Tuber badium S.P. Wan
Tuber bellisporum Bonito & Trappe
Tuber bernardinii Gori
Tuber besseyi Gilkey
Tuber beyerlei Trappe, Bonito & G. Guevara
Tuber bisporum Gilkey
Tuber bituminatum Berk. & Broome
波密块菌 Tuber bomiense K.M. Su & W.P. Xiong
Tuber bonitoi G. Guevara & Trappe
Tuber bonnetii Roum.
勃氏块菌 Tuber borchii Vittad.，波氏块菌
勃氏块菌球孢变种 Tuber borchii var. sphaerosperma Malençon
Tuber brennemanii A. Grupe, Healy & M.E. Sm.
冬块菌 Tuber brumale Vittad.
Tuber brunneum G. Guevara, Bonito & Trappe
加州块菌 Tuber californicum
Tuber calosporum S.P. Wan
Tuber canaliculatum Gilkey
Tuber caoi L. Fan
Tuber castellanoi Bonito & Trappe
Tuber castilloi G. Guevara, Bonito & Trappe
Tuber cistophilum P. Alvarado, G. Moreno, Manjón, Gelpi & Jaime Muñoz
Tuber clarei Gilkey
Tuber crassitunicatum L. Fan & X.Y. Yan
Tuber cryptobrumale Merényi, T. Varga & Bratek
Tuber decipiens Bozac, Širić & Kos
Tuber depressum S.P. Wan
Tuber donnagotto Bozac, Širić & Kos
喜栎块菌 Tuber dryophilum Tul. & C. Tul.
深脈松露 Tuber elevatireticulatum K.F. Wong & H.T. Li
凹陷块菌 Tuber excavatum Vittad.
Tuber ferrugineum Vittad.
Tuber flavidosporum Hir. Sasaki, A. Kinosh. & Nara
Tuber floridanum A. Grupe, Sulzbacher & M.E. Sm.
臭块菌 Tuber foetidum Vittad.
台湾块菌 Tuber formosanum H.T. Hu & Y. Wang
粉状块菌 Tuber furfuraceum H.T. Hu & Y.I. Wang
Tuber gardneri Gilkey
Tuber gennadii (Chatin) Pat.
Tuber giganteum Gilkey
巨孢块菌 Tuber gigantosporum Y. Wang & Z.P. Li
光巨孢块菌 Tuber glabrum L. Fan & S. Feng
Tuber guevarae Bonito & Trappe
Tuber guzmanii Trappe & Cázares
Tuber harknessii Gilkey
喜马拉雅块菌 Tuber himalayense B.C. Zhang & Minter
Tuber hiromichii (S. Imai) Trappe
湖北块菌 Tuber hubeiense L. Fan
会东块菌 Tuber huidongense Y. Wang
会泽块菌 Tuber huizeanum L. Fan & C.L. Hou
Tuber incognitum Piña Páez, Bonito, Guevara & Castellano
印度块菌 Tuber indicum Cooke & Massee，异名：Tuber sinense K. Tao& B. Liu)
Tuber intermedium Bucholtz
Tuber irradians Gilkey
Tuber itzcuinzapotl de la Fuente & Rosales-Rosales, 2024 ^[10]
日本块菌 Tuber japonicum Hir. Sasaki, A. Kinosh. & Nara
金沙江块菌 Tuber jinshajiangense L. Fan
朝鲜块菌 Tuber koreanum H.Park & A.H.Eom, 2021^[11]
Tuber lacunosum Mattir.
Tuber lannaense Suwannar. & Lumyong
宽孢块菌 Tuber latisporum Juan Chen & P.G. Liu，阔孢块菌
Tuber lauryi Trappe, Bonito & G. Guevara
Tuber levissimum Gilkey
辽东块菌 Tuber liaotongense Y. Wang
Tuber lignarium (Harkn.) Gilkey
丽江块菌 Tuber lijiangense L. Fan & J.Z. Cao
Tuber linsdalei Gilkey
Tuber lishanense L. Fan & X.Y. Yan
小西氏石櫟松露 Tuber lithocarpii K.F. Wong & H.T. Li
刘氏块菌 Tuber liui A-S. Xu
李玉块菌 Tuber liyuanum L. Fan & J.Z. Cao
Tuber longisporum Gilkey
Tuber lucentum Bordallo
莱氏块菌 Tuber lyonii Butters
Tuber macrosporum Vittad.
斑点块菌 Tuber maculatum Vittad.
Tuber magentipunctatum Merényi, I. Nagy, Stielow & Bratek
白松露 Tuber magnatum Picco, 1788
Tuber malacodermum E. Fisch.
Tuber malenconii Donadini, Riousset, G. Riousset & G. Chev.
Tuber maresa Font Quer
黑松露 Tuber melanosporum Vittad.
Tuber melosporum (G. Moreno, J. Díez & Manjón) P. Alvarado, G. Moreno, Manjón & J. Díez
膜皱块菌 Tuber mesentericum Vittad.
Tuber mexiusanum G. Guevara, Bonito & Cázares
Tuber michailowskianum Bucholtz
小孢块菌 Tuber microspermum L. Fan & J.Z. Cao
小球孢块菌 Tuber microsphaerosporum L. Fan & Y. Li
密网孢块菌 Tuber microspiculatum L. Fan & J.Z. Cao
细疣块菌 Tuber microverrucosum L. Fan & C.L. Hou
Tuber miquihuanense G. Guevara, Bonito & Cázares
Tuber monosporum (Mattir.) Vizzini
山地块菌 Tuber montanum Chatin, 1892
Tuber moravicum Velen.
Tuber moretii Maire
Tuber multimaculatum Parladé, Trappe & I.F. Alvarez^[12]
网孢凹陷块菌 Tuber neoexcavatum L. Fan & Yu Li
光果块菌 Tuber nitidum Vittad.
少孢块菌 Tuber oligospermum (Tul. & C. Tul.) Trappe
Tuber oregonense Trappe, Bonito & P. Rawl.
Tuber pacificum Trappe, Castellano & Bushnell
攀枝花块菌 Tuber panzhihuanense X.J. Deng & Y. Wang
小块菌 Tuber parvomurphium L. Fan
Tuber petrophilum Milenković, P. Jovan., Grebenc, Ivančević & Marković
Tuber phlebodermum (Gilkey) Trappe
Tuber piceatum L. Fan, X.Y. Yan & M.S. Song
Tuber polymorphosporum S.P. Wan
多孢块菌 Tuber polyspermum L. Fan & J.Z. Cao
假柄块菌 Tuber pseudobrumale Y. Wang & Shu H. Li
假凹陷块菌 Tuber pseudoexcavatum Y. Wang, G. Moreno, Riousset, Manjón & G. Riousset
假喜马拉雅块菌 Tuber pseudohimalayense G. Moreno, Manjón, J. Díez & García-Mont.
拟白块菌 Tuber pseudomagnatum L. Fan
Tuber pseudoseparans G. Guevara, Bonito & Trappe
拟球孢块菌 Tuber pseudosphaerosporum L. Fan
短毛块菌 Tuber puberulum Berk. & Broome
Tuber pustulatum M. Leonardi, A. Paz, G. Guevara & Pacioni
Tuber queletianum Bataille
Tuber quercicola J.L. Frank, D. Southw. & Trappe
曲靖块菌 Tuber qujingense S.P. Wan
芜菁味块菌 Tuber rapaeodorum Tul. & C. Tul.
Tuber regianum Montecchi & Lazzari
Tuber regimontanum G. Guevara, Bonito & Julio Rodr.
棕红块菌 Tuber rufum Pollini
Tuber scleroneuron Berk. & Broome
Tuber separans Gilkey
歇尔氏块菌 Tuber shearii Harkn.
Tuber shidianense S.P. Wan & F.Q. Yu
Tuber shii L. Fan & Y.W. Wang
中国块菌 Tuber sinense K. Tao & B. Liu
中华夏块菌 Tuber sinoaestivum J.P. Zhang & P.G. Liu
中华白块菌 Tuber sinoalbidum L. Fan & J.Z. Cao
中华凹陷块菌 Tuber sinoexcavatum L. Fan & Yu Li
中华巨孢块菌 Tuber sinomonosporum J.Z. Cao & L. Fan
Tuber sinoniveum S.P. Wan
中华短毛块菌 Tuber sinopuberulum L. Fan & J.Z. Cao
中华球孢块菌 Tuber sinosphaerosporum L. Fan, J.Z. Cao & Yu Li
松露块菌 Tuber songlu S.P. Wan
球孢块菌 Tuber sphaerospermum (Malençon) P. Roux, Guy García & M.C. Roux
Tuber sphaerosporum Gilkey
Tuber spinoreticulatum Uecker & Burds.
Tuber suave Pacioni & M. Leonardi, 2021^[13]
Tuber suaveolens Ant. Rodr. & Morte 2021
亚球孢块菌 Tuber subglobosum L. Fan & C.L. Hou
太原块菌 Tuber taiyuanense B. Liu
Tuber tequilanum G. Guevara, Bonito & Trappe
德州块菌 Tuber texense Heimsch
泰国块菌 Tuber thailandicum Suwannar., Kumla & Lumyong
Tuber theleascum M. Leonardi, A. Paz, G. Guevara & Pacioni
姜黄块菌 Tuber turmericum L. Fan
脐状块菌 Tuber umbilicatum Juan Chen & P.G. Liu
钩块菌 Tuber uncinatum Chatin, 1888
单色块菌 Tuber unicolor Gilkey
Tuber vacini Velen.
Tuber verii Pacioni & Lalli
Tuber verrucosivolvum S.P. Wan
囊被块菌 Tuber vesicoperidium L. Fan
Tuber walkeri Healy, Bonito & G. Guevara
Tuber wanglangense L. Fan
汶川块菌 Tuber wenchuanense L. Fan & J.Z. Cao
Tuber whetstonense J.L. Frank, D. Southw. & Trappe
乌蒙块菌 Tuber wumengense L. Fan
黄孢块菌 Tuber xanthomonosporum Qing & Yun Wang bis
西藏块菌 Tuber xizangense A-S. Xu
宣化块菌 Tuber xuanhuaense L. Fan
Tuber yigongense L. Fan & W.P. Xiong
中甸块菌 Tuber zhongdianense X.Y. He, Hai M. Li & Y. Wang

參見

註釋

^ 其他別稱還有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。
^ 其他別稱還有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

參考文獻

^ Tuber P. Micheli ex F.H. Wigg. 1780. MycoBank. International Mycological Association. [2011-07-13].
^ Kirk PM, Cannon PF, Minter DW, Stalpers JA. Dictionary of the Fungi 10th. Wallingford, UK: CABI. 2008: 708. ISBN 978-0-85199-826-8.
^ ^3.0 ^3.1 ^3.2 ^3.3 Hanlin, Richard T. Illustrated genera of ascomycetes. TUBER P. Mich. ex A. Wigg.. St. Paul, Minn.: APS Press. 1990: 126. ISBN 0890541078. OCLC 21300121.
^ ^4.0 ^4.1 ROOT TIP: the terminal portion of a root of a tree.ASCUS: the spore bearing sac.. www.tuber.it. [2018-08-12]. （原始内容存档于2018-08-12）.
^ ^5.0 ^5.1 ^5.2 ^5.3 國立自然科學博物館. 菌類學習資源. [2018-08-13]. （原始内容存档于2016-03-24）（中文（臺灣））.
^ 歷時5年調查世界新種「深脈松露」發表揭開台灣真菌多樣性珍寶. 環境資訊中心、行政院農業委員會林務局. 2019-03-06 [2023-07-04]. （原始内容存档于2023-07-05）.
^ ^7.0 ^7.1 ^7.2 Gryndler, Milan. True Truffle Host Diversity. True Truffle (Tuber spp.) in the World. Cham: Springer International Publishing. 2016: 267–281 [2018-08-13]. ISBN 9783319314341. doi:10.1007/978-3-319-31436-5_16. （原始内容存档于2018-08-13）（英语）.
^ Taschen, Elisa; Sauve, Mathieu; Taudiere, Adrien; Parlade, Javier; Selosse, Marc-André; Richard, Franck. Whose truffle is this? Distribution patterns of ectomycorrhizal fungal diversity inTuber melanosporumbrûlés developed in multi-host Mediterranean plant communities. Environmental Microbiology. 2015-02-14, 17 (8): 2747–2761. ISSN 1462-2912. doi:10.1111/1462-2920.12741.
^ Tuber P.Micheli ex F.H.Wigg.. GBIF. [2023-01-25]. （原始内容存档于2023-01-25）.
^ Javier Isaac de la Fuente, Wendy Rosales-Rosales, César Ramiro Martínez-González, Magdalena Martínez-Reyes, Andrea Carolina Elizondo-Salas and Jesus Perez-Moreno. 2024. Tuber itzcuinzapotl sp. nov. (Tuberaceae, Pezizomycetes), the First Edible Truffle reported from Mexico with Traditional Biocultural Importance. Phytotaxa. 635(3); 206-216. DOI: 10.11646/phytotaxa.635.3.2
^ Hyeok Park, Ju-Hui Gwon, Jong-Chul Lee & Ahn-Heum Eom (2021) Report on a New Truffle Species, Tuber koreanum sp. nov., from Korea, Mycobiology, 49:6, 527-533, DOI: 10.1080/12298093.2021.1992089
^ Isabel F. Alvarez, Javier Parladé & James M. Trappe (1992) Loculotuber Gennadii Gen. Et Comb. Nov. and Tuber Multimaculatum Sp. Nov., Mycologia, 84:6, 926-929, DOI: 10.1080/00275514.1992.12026228
^ Leonardi, M.; Salvi, D.; Iotti, M.; Rana, G.L.; Paz-Conde, A.; Pacioni, G. Multilocus Phylogeography of the Tuber mesentericum Complex Unearths Three Highly Divergent Cryptic Species. J. Fungi 2021, 7, 1090. https://doi.org/10.3390/jof7121090

外部連結

塊菌屬在Index Fungorum中的資訊
The Lightning and the Truffle （页面存档备份，存于互联网档案馆）

[5] 其他別稱還有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

[6] 其他別稱還有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

[urlMycoBank:_Tuber-1] Tuber P. Micheli ex F.H. Wigg. 1780. MycoBank. International Mycological Association. [2011-07-13].

[Kirk2008-2] Kirk PM, Cannon PF, Minter DW, Stalpers JA. Dictionary of the Fungi 10th. Wallingford, UK: CABI. 2008: 708. ISBN 978-0-85199-826-8.

[:0-3] 3.0 ^3.1 ^3.2 ^3.3 Hanlin, Richard T. Illustrated genera of ascomycetes. TUBER P. Mich. ex A. Wigg.. St. Paul, Minn.: APS Press. 1990: 126. ISBN 0890541078. OCLC 21300121.

[:1-4] 4.0 ^4.1 ROOT TIP: the terminal portion of a root of a tree.ASCUS: the spore bearing sac.. www.tuber.it. [2018-08-12]. （原始内容存档于2018-08-12）.

[:2-7] 5.0 ^5.1 ^5.2 ^5.3 國立自然科學博物館. 菌類學習資源. [2018-08-13]. （原始内容存档于2016-03-24）（中文（臺灣））.

[:02-8] 歷時5年調查世界新種「深脈松露」發表揭開台灣真菌多樣性珍寶. 環境資訊中心、行政院農業委員會林務局. 2019-03-06 [2023-07-04]. （原始内容存档于2023-07-05）.

[:3-9] 7.0 ^7.1 ^7.2 Gryndler, Milan. True Truffle Host Diversity. True Truffle (Tuber spp.) in the World. Cham: Springer International Publishing. 2016: 267–281 [2018-08-13]. ISBN 9783319314341. doi:10.1007/978-3-319-31436-5_16. （原始内容存档于2018-08-13）（英语）.

[10] Taschen, Elisa; Sauve, Mathieu; Taudiere, Adrien; Parlade, Javier; Selosse, Marc-André; Richard, Franck. Whose truffle is this? Distribution patterns of ectomycorrhizal fungal diversity inTuber melanosporumbrûlés developed in multi-host Mediterranean plant communities. Environmental Microbiology. 2015-02-14, 17 (8): 2747–2761. ISSN 1462-2912. doi:10.1111/1462-2920.12741.

[11] Tuber P.Micheli ex F.H.Wigg.. GBIF. [2023-01-25]. （原始内容存档于2023-01-25）.

[12] Javier Isaac de la Fuente, Wendy Rosales-Rosales, César Ramiro Martínez-González, Magdalena Martínez-Reyes, Andrea Carolina Elizondo-Salas and Jesus Perez-Moreno. 2024. Tuber itzcuinzapotl sp. nov. (Tuberaceae, Pezizomycetes), the First Edible Truffle reported from Mexico with Traditional Biocultural Importance. Phytotaxa. 635(3); 206-216. DOI: 10.11646/phytotaxa.635.3.2

[13] Hyeok Park, Ju-Hui Gwon, Jong-Chul Lee & Ahn-Heum Eom (2021) Report on a New Truffle Species, Tuber koreanum sp. nov., from Korea, Mycobiology, 49:6, 527-533, DOI: 10.1080/12298093.2021.1992089

[14] Isabel F. Alvarez, Javier Parladé & James M. Trappe (1992) Loculotuber Gennadii Gen. Et Comb. Nov. and Tuber Multimaculatum Sp. Nov., Mycologia, 84:6, 926-929, DOI: 10.1080/00275514.1992.12026228

[15] Leonardi, M.; Salvi, D.; Iotti, M.; Rana, G.L.; Paz-Conde, A.; Pacioni, G. Multilocus Phylogeography of the Tuber mesentericum Complex Unearths Three Highly Divergent Cryptic Species. J. Fungi 2021, 7, 1090. https://doi.org/10.3390/jof7121090

[2]

[1]

[3]

[4]

[註 1]

[註 2]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]