红菇科Russulaceae真菌是一类重要的外生菌根菌,多数物种子实体艳丽,外形醒目,易于采集,是从极地冻原到热带森林生态系统中大型真菌的重要成员（Knudsen & Borgen 1982;Verbeken & Walleyn 2012）。在传统的伞菌分类系统中,红菇科被分为两个属：具乳汁、菌褶前端菌髓由菌丝构成的归入乳菇属Lactarius,不具乳汁、菌褶菌髓具球状胞的归入红菇属Russula（Moser 1983;Singer 1986）。尽管历史上曾经有研究者试图将乳菇类真菌分为多个属（Redeuilh et al. 2001）,在分子系统发育研究方法引入该科研究之前,普遍接受的仍然是划分为两属的分类系统。
Buyck et al.（2008）选取红菇科的代表物种,采用3个基因（片段）构建系统发育树后发现,红菇属和乳菇属均不为单系。如果将Russula subsect. Ochricompactae和乳菇属的L. furcatus构成的分支另立新属,再将乳菇属拆分为两个属,则属的并系问题就能得到解决。由此,红菇科真菌被分为4个属：红菇属、乳菇属（狭义Lactarius,下简写为L.）、多汁乳菇属（Lactifluus,下简写为Lf.）以及叉褶菇属Multifurca（图1）。这是目前普遍采纳的红菇科分类系统,也是很多乳菇类真菌和少数红菇属物种的名称在近10年来发生变化的原因所在。现在,人们普遍认识到一些具乳汁的腹菌如Zelleromyces和Arcangeliella实为乳菇属的成员,不具乳汁的Cystangium、Macowanites和Martellia应归入红菇属（Nuytinck et al. 2004;Elliott & Trappe 2018）。
图1上图有个别错误：乳菇状组的拉丁名为sect. Lactarioides。请使用更新过的图1。乳菇属下小红乳菇亚属的中的L. subzongrius应改为L. subzonarius。在红菇属粘盖亚属下增加一个中国的种R. glutinosoides。请使用更新过的图1。 基于DNA序列构建的红菇科系统发育树和属下分类系统以及代表食毒物种 树的拓扑结构由下列研究整合而来：Buyck et al.（2008）关于红菇科的系统发育关系,de Crop et al.（2017）关于多汁乳菇属的属下分类系统,Buyck et al.（2018）和Wang et al.（2019）关于红菇属的属下分类系统,Wang et al.（2018）关于叉褶菇属的属下分类系统. 加粗的分支在以上研究中获得显著支持率. 枝长仅为示意图,并不必然与上述研究中显示的成比例
图1
图1
基于DNA序列构建的红菇科系统发育树和属下分类系统以及代表食毒物种
树的拓扑结构由下列研究整合而来：Buyck et al. (2008) 关于红菇科的系统发育关系，de Crop et al. (2017) 关于多汁乳菇属的属下分类系统，Buyck et al. (2018)和Wang et al. (2019)关于红菇属的属下分类系统，Wang et al. (2018) 关于叉褶菇属的属下分类系统. 加粗的分支在以上研究中获得显著支持率. 枝长仅为示意图，并不必然与上述研究中显示的成比例
Fig. 1
Molecular phylogeny of Russulaceae and representative edible and poisonous species.
The tree is constructed by integrating the following studies: Buyck et al. (2008) on Russulaceae, de Crop et al. (2017) on Lactifluus, Buyck et al. (2018) and Wang et al. (2019) on Russula and Wang et al. (2018) on Multifurca. Thick branches received significant supports in these studies. Branch lengths are of schematic diagram, not necessarily corresponding to the exact values.
红菇科真菌是全球范围内最为广泛采食的野生真菌类群之一。据不完全统计,全球有39个国家将94种乳菇作为食用菌,有28个国家将128种红菇作为食用菌（Boa 2004）。Wu et al.（2019）记录了中国的51种可食乳菇和78种可食红菇。李国杰等（2010,2015）列出了中国的82种可食红菇并提供了部分可食与有毒物种的照片。在云南野生食用菌市场,王向华和刘培贵（2002）报道了14种红菇和15种乳菇（图2A）。在马达加斯加,红菇和鸡油菌Cantharellus spp.是当地最常见的野生食用菌（Buyck 2008）。近年来,在北美和亚洲又发现和描述了多种可食乳菇和红菇的新种,增加了该科可食用物种的数目。在红菇科中,除叉褶菇属的物种由于较为少见、缺少关于可食性的报道外,其他3属中均包含大量的野生食用菌。其中,松乳菇（图2B）与美味牛肝菌Boletus edulis、鸡油菌C. cibarius、松茸Tricholoma matsutake和块菌Tuber spp.齐名,为全球最为畅销、最受赞誉的野生食用菌（Hall et al. 2003）。
图2
图2
红菇科中的代表食用菌
A：云南市场上多种红菇科真菌混合出售的场景,内有红汁乳菇、稀褶茸多汁乳菇、假稀褶多汁乳菇和变绿红菇;B：松乳菇;C：云杉乳菇;D：红汁乳菇;E：黄褐乳菇;F：香亚环乳菇;G：假稀褶多汁乳菇;H：薄囊体多汁乳菇;I：多变多汁乳菇;J：蓝黄红菇;K：致密红菇近似种;L：密褶红菇近似种;M：灰肉红菇;N：灰肉红菇近似种;O：变绿红菇
Fig. 2
Representative edible species of Russulaceae.
A: Several species of Russulaceae sold together in Yunnan market, including L. hatsudake, Lf. gerardii, Lf. pseudohygrophoroides and R. virescens; B: L. deliciosus; C: L. deterrimus; D: L. hatsudake; E: L. cinnamomeus; F: L. subzonarius; G: Lf. pseudohygrophoroides; H: Lf. tenuicystidiatus; I: Lf. versiformis; J: R. cyanoxantha; K: R. cf. compacta; L: R. cf. densifolia; M: R. griseocarnosa; N: R. aff. griseocarnosa; O: R. virescens.
整个红菇科中目前所知的剧毒物种仅有特产于东亚的亚稀褶红菇R. subnigricans。该种误食后致死率达51%（陈作红等 2016）。误食诱吐红菇R. emetica和日本红菇R. japonica引起胃肠炎型中毒（Moser 1983;Li et al. 2020b）,误食毛头乳菇L. torminosus（图3A）引起肢体炎症（Kawamura 1911）。据中国疾病预防控制中心的数据（Li et al. 2020b）,仅2019年中国发生的与日本红菇及其近似种有关的中毒案例就达41起,仅次于由大青褶伞Chlorophyllum molybdites引起的中毒案例数（54起）。表1列出了中国常见的可食和有毒（含慎食）红菇科真菌。
红菇科真菌子实体较大,醒目,出菇量大,易于采集,是最易获得的野生菌类群之一。有毒物种和可食物种（如亚稀褶红菇与稀褶红菇R. nigricans、日本红菇与短柄红菇R. brevipes、诱吐红菇与鳞盖红菇R. lepida）外形相似,有些物种即使是专业人员如果不借助专业的手段也无法作出完全准确的判断。所以,了解红菇科真菌的分类特点和名称更新,预知识别中可能存在的风险,对于保证与野生食用菌有关的食品安全有重要意义。
红菇科的传统分类强烈依赖于子实体的外观、孢子纹饰、盖表皮式样和囊状体的有无及形态（Hesler & Smith 1979;Buyck 1990;Verbeken & Walleyn 2012）。由于子实体的大小和颜色容易受自身发育和环境因子影响而改变,人们对物种的划分存在诸多争议（Adamčík et al. 2019）。DNA数据的应用较为有效地解决了一部分物种的分类问题,这些研究结果带来了一系列名称的变化,但同时也对该科物种的分类提出了新的问题和挑战。本文就该科三属代表可食类群中存在的分类问题进行评述,以期帮助读者理解名称的变化和由来,应对名称的变化带来的不便,以及正确看待形态和DNA数据各自在红菇科食用菌物种识别中的优势和局限性。
1 乳菇属Lactarius
乳菇属的模式种为毛头乳菇,这与该属原初的模式种L. piperatus不同。导致这一变化的原因是,乳菇类真菌在Buyck et al.（2008）的系统发育树中位于两个独立的分支（图1）,如果继续沿用原来的命名学处理,即把Lf. piperatus作为乳菇属的模式,则大部分乳菇物种由于与Lf. piperatus不在一个分支内将不得不给予新的属名,这显然不利于名称的稳定。Buyck et al.（2010）建议将L. torminosus作为乳菇属的保留模式（conserved type）,重新启用Roussel（1806）发表的多汁乳菇属（Lactifluus,以Agaricus lactifluus为模式）去容纳那些与Lf. piperatus位于同一个分支内的物种。这种处理能够最大程度地稳定物种名称,随后被国际植物命名法规（墨尔本法规）所认可。
几内亚（Verbeken et al. 2002）,北至中国河南（笔者研究结果）,需引起注意。全球可食的物种大部分居于乳菇亚属,个别位于小红乳菇亚属,两亚属中均有有毒的物种。
表1 中国常见可食与有毒红菇科真菌
Table 1
| 物种 Species | 食性 Edibility | 在中国的分布 Distribution in China | 共生树种或生境 Host trees or habitats |
|---|---|---|---|
| 冷杉乳菇 Lactarius abieticola | 可食 Edible | 中部-西南亚高山 Subalpine Central and Southwest | 冷杉 Abies |
| 橙红乳菇 Lactarius akahatsu | 可食 Edible | 广布 Widely distributed | 松 Pinus |
| 高山毛脚乳菇 Lactarius alpinihirtipes | 有毒 Poisonous | 西南亚高山 Subalpine Southwest | 高山栎 Quercus sect. Heterobalanus |
| 水环乳菇 Lactarius aquizonatus | 慎食 Not recommended | 东北、西南亚高山 Northeast and subalpine Southwest | 云杉、高山松 Picea and Pinus densata |
| 暗褐乳菇 Lactarius atrobrunneus | 有毒 Poisonous | 热带 Tropical | 壳斗科 Fagaceae |
| 暗缘乳菇 Lactarius atromarginatus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 南方喙状乳菇 Lactarius austrorostratus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 鸡足山乳菇 Lactarius chichuensis | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 黄褐乳菇 Lactarius cinnamomeus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科、松 Fagaceae and Pinus |
| 胶联乳菇 Lactarius conglutinatus | 慎食 Not recommended | 亚热带 Subtropical | 混交林 Mixed forest |
| 松乳菇 Lactarius deliciosus | 可食 Edible | 北温带、亚带 North temperate and subtropical | 松林 Pinus |
| 云杉乳菇 Lactarius deterrimus | 可食 Edible | 北温带和西南亚高山 North temperate and subalpine Southwest | 云杉、五针松 Picea and five-needle pines |
| 北欧乳菇 Lactarius fennoscandicus | 可食 Edible | 北温带和西南亚高山 North temperate and subalpine Southwest | 云杉、五针松 Picea and five-needle pines |
| 糙鳞乳菇 Lactarius furfuraceus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 细弱乳菇 Lactarius gracilis | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 红汁乳菇 Lactarius hatsudake | 可食 Edible | 广布 Widely distributed | 松林 Pinus |
| 横断山乳菇 Lactarius hengduanensis | 可食 Edible | 西南亚高山 Subalpine Southwest | 云杉林 Picea |
| 毛脚乳菇 Lactarius hirtipes | 有毒 Poisonous | 亚热带 Subtropical Southwest | 壳斗科 Fagaceae |
| 印度黄汁乳菇 Lactarius indochrysorrheus | 慎食 Not recommended | 北温带和亚热带 North temperate and subtropical | 混交林 Mixed forest |
| 蜡蘑状乳菇 Lactarius laccarioides | 慎食 Not recommended | 热带 Tropical | 壳斗科林 Fagaceae |
| 黑色乳菇 Lactarius nigricans | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 橄榄褐乳菇 Lactarius olivaceoumbrinus | 慎食 Not recommended | 西南亚高山 Subalpine Southwest | 冷杉 Abies |
| 橄榄乳菇 Lactarius olivinus | 慎食 Not recommended | 西南亚高山 Subalpine southwest | 云杉 Picea |
| 欧姆斯乳菇 Lactarius oomsisiensis | 有毒 Poisonous | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 波宁乳菇 Lactarius porninsis | 可食 Edible | 东北 Northeast | 落叶松 Larix |
| 假红汁乳菇 Lactarius pseudohatsudake | 可食 Edible | 西南亚高山 Subalpine Southwest | 云杉 Picea |
| 绒边乳菇 Lactarius pubescens | 有毒 Poisonous | 东北、西南亚高山 Northeast and subalpine Southwest | 桦木 Betula |
| 红褐乳菇 Lactarius rubrobrunneus | 有毒 Poisonous | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 似白乳菇 Lactarius scoticus | 有毒 Poisonous | 东北、西南亚高山 Northeast and subalpine Southwest | 桦木 Betula |
| 窝柄黄乳菇 Lactarius scrobiculatus | 慎食 Not recommended | 东北、西南亚高山 Northeast and subalpine Southwest | 云杉、泠杉 Picea、Abies |
| 中华环纹乳菇 Lactarius sinozonarius | 慎食 Not recommended | 西南亚热带 Subtropical Southwest | 混交林 Mixed forest |
| 多鳞乳菇 Lactarius squamulosus | 慎食 Not recommended | 热带 Tropical | 阔叶林 Broad-leaved forest |
| 近大西洋乳菇 Lactarius subatlanticus | 有毒 Poisonous | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 近短柄乳菇 Lactarius subbrevipes | 慎食 Not recommended | 西南亚热带 Subtropical Southwest | 松 Pinus |
| 近毛脚乳菇 Lactarius subhirtipes | 有毒 Poisonous | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 亚靛蓝乳菇 Lactarius subindigo | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 香亚环乳菇 Lactarius subzonarius | 可食 Edible | 亚热带 Subtropical | 壳斗科 Fagaceae |
| 毛头乳菇 Lactarius torminosus | 有毒 Poisonous | 北温带和西南亚高山 North temperate and subalpine Southwest | 桦木 Betula |
| 堇紫乳菇 Lactarius uvidus | 慎食 Not recommended | 东北 Northeast | 桦木 Betula |
| 靓丽乳菇 Lactarius vividus | 可食 Edible | 亚热带 Subtropical | 松 Pinus |
| 稀褶茸多汁乳菇 Lactifluus gerardii | 可食 Edible | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 长毛多汁乳菇 Lactifluus longipilus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 淡黄多汁乳菇 Lactifluus luteolus | 可食 Edible | 亚热带 Subtropical | 混交林 Mixed forest |
| 灰绿多汁乳菇 Lactifluus glaucescens | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 狮黄多汁乳菇近似种 Lactifluus aff. leoninus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 长绒乳菇 Lactifluus pilosus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 宽囊体多汁乳菇 Lactifluus pinguis | 可食 Edible | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 辣味多汁乳菇 Lactifluus piperatus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 假稀褶多汁乳菇 Lactifluus pseudohygrophoroides | 可食 Edible | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 亚辣味多汁乳 Lactifluus subpiperatus | 慎食 Not recommended | 亚热带-热带 Subtropical-tropical | 混交林 Mixed forest |
| 近粉绒多汁乳菇 Lactifluus subpruinosus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 薄囊体多汁乳菇 Lactifluus tenuicystidiatus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 中华热带多汁乳菇 Lactifluus tropicosinicus | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 多变多汁乳菇 Lactifluus versiformis | 可食 Edible | 西南亚高山 Subalpine Southwest | 高山栎 Quercus sect. Heterobalanus |
| 多汁乳菇 Lactifluus volemus | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 冷杉红菇 Russula abietina | 可食 Edible | 东北 Northeast | 针叶林、混交林 Coniferous and mixed forests |
| 辣褶红菇 Russula acrifolia | 可食 Edible | 东北、西南亚高山 Northeast and subalpine Southwest | 混交林 Mixed forest |
| 烟色红菇 Russula adusta | 可食 Edible | 广布 Widely distributed | 针叶林 Coniferous forest |
| 白红菇 Russula albida | 可食 Edible | 广布 Widely distributed | 阔叶林 Broad-leaved forest |
| 白黑红菇 Russula albonigra | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 铜绿红菇 Russula aeruginea | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 革质红菇 Russula alutacea | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 暗绿红菇 Russula atroaeruginea | 可食 Edible | 西南亚高山 Subalpine Southwest | 云杉 Picea |
| 金红菇 Russula aurea | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 天蓝红菇 Russula azurea | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 短柄红菇 Russula brevipes | 可食 Edible | 北温带、西南亚高山 North temperate and subalpine Southwest | 混交林 Mixed forest |
| 裘氏红菇 Russula chiui | 慎食 Not recommended | 西南亚高山 Subalpine Southwest | 高山栎 Quercus sect. Heterobalanus |
| 致密红菇 Russula compacta | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 蓝黄红菇 Russula cyanoxantha | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 类致密红菇 Russula compactoides | 可食 Edible | 西南亚高山 Subalpine Southwest | 冷杉 Abies |
| 美味红菇 Russula delica | 可食 Edible | 广布 Widely distributed | 针叶林、阔叶林、混交林 Coniferous, broad-leaved and mixed forests |
| 密褶红菇 Russula densifolia | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 诱吐红菇 Russula emetica | 有毒 Poisonous | 广布 Widely distributed | 混交林 Mixed forest |
| 非白红菇 Russula exalbicans | 可食 Edible | 北温带、西南亚高山 North temperate and subalpine Southwest | 针叶林、混交林 Coniferous and mixed forests |
| 臭红菇 Russula foetens | 慎食 Not recommended | 广布 Widely distributed | 针叶林、混交林 Coniferous and mixed forests |
| 绵粒红菇 Russula granulata | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 可爱红菇 Russula grata | 可食 Not recommended | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 灰肉红菇 Russula griseocarnosa | 可食 Edible | 亚热带-热带 Subtropical-tropical | 壳斗科 Fagaceae |
| 日本红菇 Russula japonica | 有毒 Poisonous | 亚热带 Subtropical | 混交林 Mixed forest |
| 蜜味红菇 Russula melliolens | 可食 Edible | 西南亚热带 Subtropical Southwest | 阔叶林 Broad-leaved forest |
| 厚皮红菇 Russula mustelina | 可食 Edible | 广布 Widely distributed | 针叶林、混交林 Coniferous and mixed forests |
| 稀褶红菇 Russula nigricans | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 黑绿红菇 Russula nigrovirens | 可食 Edible | 西南亚高山带 Subalpine Southwest | 冷杉 Abies |
| 黄白红菇 Russula ochroleuca | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 青黄红菇 Russula olivacea | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 沼泽红菇 Russula paludosa | 可食 Edible | 广布 Widely distributed | 针叶林、阔叶林 Coniferous and broad-leaved forests |
| 拟蓖形红菇 Russula pectinatoides | 可食 Edible | 广布 Widely distributed | 针叶林、阔叶林 Coniferous and broad-leaved forests |
| 致密红菇 Russula pseudocompacta | 可食 Edible | 西南亚热带 Subtropical Southwest | 混交林 Mixed forest |
| 致密红菇 Russula pseudodelica | 可食 Edible | 亚热带 Subtropical | 混交林 Mixed forest |
| 假拟蓖形红菇 Russula pseudopectinatoides | 慎食 Not recommended | 西南亚高山带 Subalpine southwest | 冷杉、落叶松、云杉 Abies, Larix and Picea |
| 点柄红菇 Russula punctipes | 慎食 Not recommended | 亚热带 Subtropical | 混交林 Mixed forest |
| 凯莱红菇 Russula queletii | 有毒 Poisonous | 广布 Widely distributed | 针叶林、混交林 Coniferous and mixed forests |
| 玫瑰红菇 Russula rosea | 可食 Edible | 广布 Widely distributed | 混交林 Mixed forest |
| 黄茶红菇 Russula sororia | 可食 Edible | 广布 Widely distributed | 针叶林、阔叶林、混交林 Coniferous, broad-leaved and mixed forests |
| 血红菇 Russula sanguinea | 可食 Edible | 广布 Widely distributed | 针叶林、混交林 Coniferous and mixed forests |
| 亚稀褶红菇 Russula subnigricans | 剧毒 Deadly poisonous | 亚热带 Subtropical | 壳斗科 Fagaceae |
| 菱红菇 Russula vesca | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 堇紫红菇 Russula violacea | 可食 Edible | 广布 Widely distributed | 阔叶林 Broad-leaved forest |
| 变绿红菇 Russula virescens | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
| 红边绿菇 Russula viridirubrolimbata | 可食 Edible | 亚热带 Subtropical | 阔交林 Mixed forest |
| 黄孢红菇 Russula xerampelina | 可食 Edible | 广布 Widely distributed | 阔叶林、混交林 Broad-leaved and mixed forests |
图3
图3
红菇科中的代表毒菌及慎食物种
A：毛头乳菇;B：窝柄黄乳菇;C：毛脚乳菇;D：蜡蘑状乳菇;E：红褐乳菇;F：近毛脚乳菇;G：橄榄褐乳菇;H：欧姆斯乳菇;I：堇紫乳菇;J：日本红菇（李海蛟拍摄）;K：近似诱吐红菇;L：亚稀褶红菇（陈作红拍摄）
Fig. 3
Representative poisonous and inedible species of Russulaceae.
A: L. torminosus; B: L. scrobiculatus; C: L. hirtipes; D: L. laccarioides; E: L. rubrobrunneus; F: L. subhirtipes; G: L. olivaceoumbrinus; H: L. oomsisiensis; I: L. uvidus; J: R. japonica (photo by H.J. Li); K: R. aff. emetica; L: R. subnigricans (photo by Z.H. Chen).
1.1 松乳菇组Lactarius sect. Deliciosi
乳菇亚属的松乳菇组是以全球著名食用菌L. deliciosus为模式的、广布于北温带的类群,也是红菇科中最易识别的食用菌类群。其关键识别特征为：子实体特别是菌褶橙色、胡萝卜色、灰红色、靛蓝色,伤后发蓝绿色,盖表常具环纹,盖缘无毛,菌褶伤后有少量橙色、暗血红色或靛蓝色乳汁。有别于上述关键特征,本组中有3个物种的乳汁为白色,分别为亚洲的L. aurantiozonatus、欧亚的波宁乳菇和北美的L. splendens（Nuytinck et al. 2007,2017;Lee et al. 2019）。其中,波宁乳菇可食,其他两种食性未知。该组中,除靛蓝乳菇L. indigo和亚靛蓝乳菇生于壳斗科林下外,其余种类均与松科植物如松属、冷杉属、云杉属、黄杉属和落叶松属植物共生（Nuytinck et al. 2007）。这种形态和生态上的同质性为识别这一大类食用菌带来了极大的便利,使其成为最为安全的野生食用菌类群之一。
松乳菇组全球共知46种,物种表现出极强的地区特有性,即各大洲有各自的种类,大洲之间共享的物种很少（Nuytinck et al. 2007;Nuytinck & Ammirati 2014;Wang et al. 2015b;Wang 2016;Han et al. 2019）。目前所知仅松乳菇、云杉乳菇、北欧乳菇和波宁乳菇广布于欧亚大陆（笔者所采标本鉴定结果）。在中国,原被定为L. sanguifluus的实为红汁乳菇,被定为L. semisanguifluus的极有可能是橙红乳菇,被定为L. thyinos的为冷杉乳菇,被定为L. indigo的为亚靛蓝乳菇。
由于全组种类在形态上的高度同质性,在松乳菇组内进行种一级的划分和识别十分困难。很多物种,如云杉乳菇、L. quieticolor、L. vinosus都曾被作为另一个物种如松乳菇的变种。即使在欧洲,不同的研究者对同一名称的理解和诠释也不同（Nuytinck 2004）。外表差别较大的个体,如云杉乳菇的菌盖可以从明亮的橙黄色到蓝绿色,被DNA数据证实为同一物种,而一些物种如中国的冷杉乳菇和日本的悦色乳菇L. laeticolor在外形上高度相似,均为明亮的橙色且与冷杉属植物共生,却被DNA数据证明为不同的物种,二者甚至不为姊妹种（Wang 2016）。即使是长期从事乳菇分类学研究的人员,如果不了解样品的来源地和共生树种,不借助DNA数据,也很难在相近物种间作出完全准确的判断。
DNA数据应用于该类食用菌并非能解决所有的分类问题。Nuytinck et al.（2007）采用核糖体RNA基因内转录间隔区（internal transcribed spacer,ITS）和甘油醛-3-磷酸脱氢酶基因（glyceraldehyde-3-phosphate dehydrogenase）的序列对全球松乳菇组的物种进行研究后发现,该组物种间不仅在形态上高度相似,在DNA序列上也极为相近,这表现为系统发育树中极短的枝长、一部分分支较低的支持率和并非所有的样品都能聚于末端分支内。不仅如此,该类群中存在着一定程度的“种间杂交”现象。Nuytinck & Verbeken（2005）发现云杉乳菇和北欧乳菇之间的区分并不清晰。笔者在中国采集了大量属于这两个种的标本,发现了同样的现象,即在ITS片段上,用以区分云杉乳菇和北欧乳菇的信息位点处,有较多的个体存在碱基的简并现象。即使采用多基因数据,也依然不能在两者之间做出清晰的划分。这表明,之前依据少量样品的形态特征和DNA数据进行的物种划分可能并不合适。如同形态特征会存在过渡性、连续性一样,由于不完全谱系分选、杂交和网状进化,DNA数据也不会在所有的情况下都能清晰分组。由于该组物种与共生树种间具有高度专一性,如靓丽乳菇专性与松属松亚属的植物共生,横断山乳菇专性与云杉共生,在识别这一大类食用菌时,了解标本的产地和共生树种,依赖形态上的同质性和相似性,比依赖DNA数据具有更强的可操作性。
与松乳菇组的物种相比,乳菇亚属中有些物种并不表现出强烈的地区特有性,而是广布于北温带或欧亚大陆,如毛头乳菇（图3A）、水环乳菇、茶绿乳菇L. necator、橄榄乳菇、绒边乳菇、橄榄褐乳菇（图3G）、复生乳菇L. repraesentaneus、窝柄黄乳菇（图3B）、堇紫乳菇（图3I）等。这些或盖表具长毛,或乳汁苦辣、白色变黄色或紫色的物种应避免食用（Arora 1986）。
1.2 小红乳菇亚属Lactarius sug. Russularia
小红乳菇亚属包含了一大类子实体纤细、红褐色、褐色、盖表较为光滑的物种。与松乳菇组的物种不同,这个类群的物种尽管在外观上较为相似,在显微特征上却总能找到较为稳定的区别。与乳菇亚属不同,该亚属仅有少数物种在欧亚地区广布,如桔色乳菇L. aurantiacus、易烂乳菇L. tabidus、棕红乳菇L. badiosanguineus（Wisitrassameewong et al. 2016;笔者所采标本鉴定结果）。在中国曾记载将L. camphoratus作为食用菌（Chiu 1945;应建浙和臧穆 1994）。笔者对L. camphoratus的引证标本进行研究后发现,它们都不是L. camphoratus,而是香亚环乳菇（图2F）、毛脚乳菇（图3C）、高山毛脚乳菇和暗乳菇L. obscuratus。迄今,在中国没有发现L. camphoratus,只发现了它的近缘物种疣孢乳菇L. tuberculatus,分布于云南和贵州（Wang 2018）,但并不作为食用菌采集。同样在中国作为食用菌记载的油味乳菇L. quietus（应建浙和臧穆1994）,到目前为止尚未被证实,而是发现了与其形态和亲缘关系相近的东亚特有种东方油味乳菇L. orientaliquietus和东亚乳菇L. asiae-orientalis,广布于东北到西南等地（Wang 2018;Wang et al. 2018）。中国西南高山栎林下有北美L. rubidus（candy cap）的近缘种假脆乳菇L. pseudofragilis（= L. serilactifer）,但子实体少有香味（Wang 2017）,未见采食。在中国,具有特别香味被采食的为香亚环乳菇（图2F）。该种子实体较小,菌盖红褐色具明显环纹,乳汁白色或水样液,全子实体具类似当归的香气,仅具褶缘囊状体。
中国西南民间采食的黄褐乳菇也属于小红乳菇亚属,昆明及贵阳市场常有出售（图2E）。该种发表于1945年（Chiu 1945）,后Wang（2007）研究了其模式标本并对形态进行了重新描述,但未提供DNA序列。由于GenBank上无匹配序列,Wisitrassameewong et al.（2015）将其作为新种L. kesiyae发表。令人遗憾的是,依据泰国研究者的研究结果,陈言柳等（2019）又将其作为新记录种“思茅乳菇”在中国进行了报道。与其在东南亚生于思茅松下不同,该种在中国西南常生于纯的壳斗科林下或具壳斗科树种的林下,在ITS序列和形态特征方面,中国的材料与东南亚的材料无异。关于其可食性由于缺乏足够的证据暂无定论,但从民间广泛采食并在市场上出售这一事实看,应未发生过中毒事件。Wang et al.（2018）描述的胶联乳菇与黄褐乳菇在形态上相似,在中国中东部,民间采食的类似于黄褐乳菇的有可能是食性未知的胶联乳菇。
本亚属中的毛脚乳菇、近毛脚乳菇（图3F）、暗褐乳菇、红褐乳菇（图3E）食后有引起胃肠炎型中毒的案例（Li et al. 2020b;中国疾病预防控制中心2020年样品鉴定结果）。与这些种外形相近较为近缘的还有高山毛脚乳菇、鸡足山乳菇、橙褐乳菇L. aurantiobrunneus、红皱乳菇L. rubrocorrugatus、南方喙状乳菇和近大西洋乳菇。虽然这些物种有时具有一定的香味,如鸡足山乳菇和暗褐乳菇,考虑以往曾经发生过的中毒事件,它们的食性值得怀疑。这些物种普遍子实体小而细弱,柄的长度常明显长于菌盖的直径,盖表光滑具棘状突起,子实体为红褐色或黄褐色。在采食乳菇属真菌时,应避免采食具有如上特征的物种。原初描述自泰国的蜡蘑状乳菇在云南南部有分布（图3D）,外形极似食用菌红蜡蘑Laccaria laccata,它与毛脚乳菇的近缘关系使得我们有理由怀疑它的可食性,应引起注意。
2 多汁乳菇属Lactifluus
在红菇科的系统发育树上,多汁乳菇属与乳菇属分属于两个独立的系统发育分支（图1）。尽管两者的区分更多地基于DNA数据,但在特征上也表现出一定的偏好性。以下特征仅见于多汁乳菇属（但并非所有的种类）：1）盖表或柄表天鹅绒状;2）盖表、柄表或子实层上具厚壁菌丝;3）菌褶菌髓具球状胞（Verbeken & Nuytinck 2013）。而下列特征则仅见于乳菇属（并非所有种类）：1）盖表具环纹,粘,全盖或边缘具刺毛;2）菌柄具窝斑;3）盖表皮粘菌丝平伏型。在地理分布上,多汁乳菇属具有明显的热带性质,多样性集中于亚热带-热带亚洲、热带非洲、美洲中南部,而乳菇属的多样性中心则在北温带（Buyck et al. 2008）。
多汁乳菇属由4个亚属构成：裸果亚属Lf. subg. Gymnocarpi、拟乳菇亚属Lf. subg. Lactariopsis、多汁乳菇亚属Lf. subg. Lactifluus和假裸果亚属Lf. subg. Pseudogymnocarpi（de Crop et al. 2017）（图1）。该属真菌在全球广泛采食,各亚属均有代表性可食物种。除辣味多汁乳菇复合群和绒白多汁乳菇Lf. vellereus复合群的部分物种乳汁具辣味食后可能会引起不适外（Li et al. 2020b）,还未发现真正有毒的物种。如前所述,由于属名的变化,很多可食物种的名称发生了变化。原来长期使用Lactarius这一属名的物种,现被移入Lactifluus中（Stubbe et al. 2012;Verbeken et al. 2012a,2012b）。与乳菇属不同,多汁乳菇属的分类表现为如下特点：在物种的层面上,形态相近甚至相同的物种DNA数据的相似性较低（Verbeken & Nuytinck 2013）,而在属下分类群的层面上,形态特征差别较大的类群却表现出较近的亲缘关系（Wang et al. 2015a;de Crop et al. 2017）。地理分布上,多汁乳菇属中到目前为止仅发现伯尔弟多汁乳菇Lf. bertillonii分布于欧亚大陆（笔者所采标本鉴定结果）,换言之,多汁乳菇属的物种具有极强的地区特有性。甚至在亚属或组的地理分布上,也表现出较强的地理偏好性,如L. subg. Lactifluus的物种仅分布于北半球,而Lf. subg. Lactariopsis的多数组则仅见于热带非洲（Verbeken & Walleyn 1999;de Crop et al. 2017）。
2.1 薄囊体多汁乳菇复合群
尽管薄囊体多汁乳菇（图2H）及其相近种在云南已采食、销售多年,且在中国广东、泰国和越南均有分布,该种直到2006年才以一份采自云南腾冲市场的标本作为模式发表（Wang & Verbeken 2006）。该种在云南被称为“奶浆菌”,由于形态上的相似性,曾被误定为Lf. rugatus（王向华等 2004）。它与Lf. rugatus的区别在于子实层上具细弱的大囊体。此后,基于更多的近似于薄囊体多汁乳菇的标本（这些标本中的相当数量采自云南野生菌市场）, Wang et al.（2015a）采用3个基因片段发现该种实为一个种复合群,其下包含5个系统发育种,构成了多汁乳菇属的一个新组薄囊体组Lf. sect. Tenuicystidiati。其中,Lf. tropicosinicus和Lf. subpruinosus由于在形态上与薄囊体多汁乳菇有别而作为新种发表,剩余的两个种处理为Lf. tropicosinicus的隐形种。目前发现的该组所有的物种都可食。该组的物种尽管与L. rugatus在形态特征上高度相似（除了后者子实层上不具有大囊体外）,两者并不居于同一亚属,而那些与薄囊体多汁乳菇居于同一亚属的类群,如多汁乳菇组Lf. sect. Lactifluus、稀褶茸多汁乳菇组Lf. sect. Gerardii、辣味多汁乳菇组L. sect. Piperati在形态上与其相去甚远。对该类群进行的研究表明,在多汁乳菇属的分类中,对于物种的鉴定,应注重根据形态特征确定其所属的复合群,不必过于追求复合群下的物种细分,而对于某一复合群系统位置的确定,则需依靠多基因片段确定,形态特征很可能会给出错误的暗示。
2.2 其他种复合群
在多汁乳菇属中,由于DNA数据的应用,近年来的分类学研究使得该属物种数目急剧膨胀。这些发现为野生食用菌增加了新的成员。例如,历史上,在著名的食用菌多汁乳菇复合群内,欧洲被广泛接受的仅有Lf. volemus,北美有Lf. volemus和Lf. corrugis,亚洲尽管有本土的名称Lf. princeps和王氏乳菇L. wangii,但也基本沿用欧美的名称。van de Putte et al.（2010,2012）根据DNA序列上的差异和微妙的形态特征差异,在亚洲发表了9个新种,其中常见的有宽囊体多汁乳菇和多变多汁乳菇（图2I）,这两个种在我国广布,而欧洲的Lf. volemus也被证实由3个物种构成（van de Putte et al. 2016）。该复合群内发现的系统发育物种超过20个。相似地,在包含可食物种的稀褶茸多汁乳菇组中,也发现了37个系统发育物种,其中部分形态上可以识别的物种如Lf. hora、Lf. leae和Lf. limbatus得以发表（Stubbe et al. 2010;de Crop et al. 2018）。在包含可食物种的稀褶多汁乳菇复合群中,一直被定为北美的Lf. hygrophoroides的亚洲材料,新近被描述为一个亚洲特有种假稀褶多汁乳菇（图2G）,同时发现的还有新种L. luteolamellatus和多汁乳菇复合群的新种Lf. holophyllus（Tibpromma et al. 2017）。
由于大量隐形种的存在,对于多汁乳菇属下新物种的描述和确定,最大的难度在于研究者无法清晰地知道在采用DNA数据所构建的系统发育树上,哪些分支对应于历史上已经发表的物种如Lf. gerardii、Lf. hygrophoroides、L. princeps、白绒多汁乳菇Lf. puberulus、王氏乳菇等。研究发现,在这些物种的原初描述地,如北美、欧洲和亚洲,存在着不止一个系统发育种。解决这一难题需要在这些物种的模式产地,采集与原始描述或模式标本特征高度吻合的样品,将其作为附加模式（epitype）,提供条形码片段,用以诠释物种的概念,稳定种名的使用。另外一个不得不面对的现实是,随着一些形态上稍有区别的系统发育分支被作为新种发表,在系统发育树上遗留了大量无法命名的分支,这些分支并不形成单系,但在形态上高度相似。当使用DNA数据去与已发表的研究结果进行对比、拟合时,最终不得不回到“大种”的名称上,而这些“大种”（如多汁乳菇）采用形态特征即可以确定。多汁乳菇属中大量的分类学实践和民间知识证明,属于同一个种复合群的物种在可食性方面表现一致,所以,对于可食物种的鉴定,根据形态特征确定其所属的复合群,比过分追求复合群下的物种细分具有更强的实际意义和可操作性。
3 红菇属Russula
尽管DNA数据的应用更新了红菇科的属下分类系统,除R. subsect. Ochricompactae被从红菇属中移出外,该属的范围并没有发生太大的变化。红菇属的分类是大型真菌分类学的难点,表现为：种类多,特征易变,特征重叠,很多物种间在DNA序列上的区分度较低（Adamčík et al. 2019）。整体上,红菇属的分类学研究滞后于乳菇属和多汁乳菇属。这一状况不仅表现在亚洲、非洲和美洲,也存在于真菌分类学的发源地欧洲（与J. Nuytinck交流）,很多原初描述自欧洲的物种的概念尚未被充分理解（Caboň et al. 2018）。
按照红菇属的最新分类系统,该属分为8个亚属（Buyck et al. 2018,2020）（图1）。除包含物种极少的厚壳亚属R. subg. Crassotunicata和粘盖亚属R. subg. Glutinosae外,其他6个亚属下均有可食的种类。关于哪些物种可食,并没有统一判断标准,而只能依据民间已有的知识和经验积累。一些文献认为,菌肉味柔和为判断可食红菇的特征（Arora 1986;Härkönen et al. 2003;Hall et al. 2003）,但日本红菇具有柔和的菌肉（Hongo 1954）却引起多起胃肠炎型中毒事件（Li et al. 2020b）。整体上看,异褶亚属R. subg. Heterophyllidia和红菇亚属R. subg. Russula是拥有食用菌最多的两个亚属,但在红菇亚属中存在一定数目的毒菌。致密亚属R. subg. Compactae和短柄亚属R. subg. Brevipes既包括食菌也包括毒菌。与乳菇类真菌相比,红菇属真菌用于分类的特征较少,特征更为多变,食菌与毒菌间的区分更为困难。
3.1 火炭菌
“火炭菌”是中国中南部的一类著名食用菌的统称,指菌盖灰白色至黑褐色、子实体伤后变黑的一类红菇,属致密亚属。该亚属物种子实体灰黑色、黑褐色,坚硬致密,有别于多数红菇较为鲜艳松软的子实体,菌褶不等长,与多数红菇等长的菌褶不同。致密亚属由两个组构成（Buyck et al. 2018）（图1）：黑色组R. sect. Nigricantinae包括子实体伤后变红、变黑的种类,如在北温带广泛采食的白黑红菇、烟色红菇、密褶红菇（图2L）、稀褶红菇和著名的毒菌亚稀褶红菇（图3L）;多褶组R. sect. Polyphyllae包含的物种菌褶初白色后变为粉红色,不变红或黑,如北美的R. cortinarioides、R. eccentrica和R. polyphylla（Liu et al. 2015）,文献中缺乏对该组物种是否可食的记载。被Shimono et al.（2014）定为亚稀褶红菇的部分样品（clades B-2、B-3、B-4）实际是近似R. eccentrica的物种（Park et al. 2014;笔者所采标本鉴定结果）。尽管这些种与稀褶红菇并不近缘且在形态上表现出差异,如菌盖色稍浅,盖表易龟裂为颗粒状,菌褶成熟后粉红,受伤不变色,但它们的菌肉与亚稀褶红菇一样不会变黑。这一特征可以作为将可食的火炭菌与（潜在）有毒的物种区分开来的关键特征。Shimono et al.（2014）和笔者对更多材料的研究发现,剧毒的亚稀褶红菇和可食的稀褶红菇确实近缘。这种遗传分化上并不显著的类群在食毒性方面表现出如此大的差异的现象提醒我们,关于食用菌和毒菌的确定,要结合大量民间实践和实验数据,对于不了解、不熟悉的类群应避免食用。
包括大量可食物种的黑色组的分类较为困难。在较早期的分类中,由于已描述的物种较少,依据少数几个特征如菌肉和菌褶的变色类型、菌褶的疏密、菌盖的颜色以及是否胶粘和孢子的大小和纹饰类型可以相对清楚地划分物种。随着描述物种的增多,一些物种间的界限开始变得重叠、模糊,如由Shaffer（1962）描述的R. atrata与R. albonigra的区别仅在于盖表皮较厚,而这一特征可能易受到环境因素的影响。R. dissimulans与R. nigricans相比菌褶较密,孢子较大。在GenBank中,被定为R. dissimulans的ITS序列有3条,来自美国西部的EU303008与来自东部的KY848515和KT933979之间的相似度仅91.4%,而一条被定为R. nigricans的序列GU289649与KY848515的相似度却达99.8%。这很可能是由于R. dissimulans与R. nigricans之间的界限并不如原初发现的那么清晰造成的。什么是真正的R. dissimulans值得探究。即使在欧洲也有一些中间类型,如R. adusta f. rubens兼具烟色红菇、密褶红菇和稀褶红菇的特征（Shaffer 1962）。Shimono et al.（2014）发现,形态上被定为密褶红菇的样品形成了6个末端分支。当把GenBank中大量属于黑色组的序列和笔者从中国样品上获得的序列进行分析后发现,具有密褶红菇和稀褶红菇特征的序列/样品均不聚为单系。黑色组中物种的概念和真实的物种数目尚属未知。
如果黑色组的物种像多汁乳菇属那样具有高度的地区特有性,亚洲的问题会变得比较简单。然而数据显示的情况并非如此。在同一形态型下,欧洲、北美、亚洲的样品并不按照地理区域表现出明显的聚类关系。亚洲、欧洲、北美甚至中美洲的部分ITS序列间差异很小。对于那些系统发育树上的具有较长枝长的分支,固然可以根据微妙的形态特征将它们作为“新种”进行描述,但产生的一个后果是,基因树上有大量的样品处于不能被分类的境地。在这种情况下,除了挑选式发表新种,是不是可以既考虑样品间的共性,又考虑它们在序列上表现出的差异?在充分取样的基础上,在尊重同一个物种的前提下,描述若干种下分类群,如Shaffer（1962）所做的那样,是不是比发表若干形态上几乎无法区别的新种,更便于人们使用呢?
3.2 蓝黄红菇复合群
蓝黄红菇复合群是一类极为常见的食用红菇（图2J）,在北温带广泛采食,分类学上这一复合群相当于异褶亚属的蓝黄亚组R. subsect. Cyanoxanthinae。对于蓝黄红菇复合群,一个较为关键的野外识别特征是,菌褶白色,用手指来回触摸菌褶,如有油脂感（greasy）,则属本类。其他关键识别特征为：孢子印白色,孢子表面纹饰为离散的疣突,少有细线相连,盖表皮用结晶蓝染色呈异染色性（metachromatic）,盖表皮具丰富的囊状体。
由于蓝黄红菇形态上的多变性,其下描述有多个变种（型）,有些变种如var. peltereaui和var. variata有时又会被处理为独立的种（www.index fungorum.org/names）。最近亚洲地区描述了数个蓝黄红菇的近缘种,如鼎湖山红菇R. dinghuensis、莲红菇R. lotus、黑绿红菇R. nigrovirens、近白粉红菇R. subpallidirosea（Zhao et al. 2015;Zhang et al. 2017;Li & Deng 2018）。在这一复合群内,一些形态上相似的物种ITS序列的相似度较低,如莲红菇与近白粉红菇（89.65%）,而一些形态特征相差较大的物种ITS序列反而更为相似,如黑绿红菇因其黑绿色具龟裂斑状的菌盖明显有别于蓝黄红菇,甚至与亲缘关系较远的变绿红菇相似（Zhao et al. 2015）,但在ITS序列上两者却有97%的相似度。随着蓝黄红菇复合群下发表的新种越来越多,试图依据DNA序列上的归类去寻找形态特征上的差异会变得越来越困难,最终这一复合群的分类可能呈现由DNA数据支持的形态学物种和隐形种并存的状态。从确定食性的角度来看,如注重掌握其关键的形态特征,即可实现正确鉴定,在实际操作中,没有必要识别更多的“小种”。
与蓝黄红菇居于同一亚属盖表为绿色的著名食用菌“青头菌”的相关分类学问题见本专刊“中国南方变绿红菇近似种类青头菌的物种多样性”一文。
3.3 大红菌
“大红菌”是中国南部对一类子实体较大、菌盖红色至黑红色、盖缘无颗粒状条纹、菌肉柔和常呈灰色的一类红菇的俗称,属红菇属的核心类群红菇亚属。与以上已讨论的类群不同,大红菌的界限较不明确。红菇亚属中子实体较大、菌盖红色的种类极多。在欧美作为食用菌的R. xerampelina、R. decolorans、R. rosea、R. integra的菌盖均为红色,且R. decolorans的菌肉伤后或与空气接触后也会变为灰色至黑色。在中国,大红菌一直被定为R. vinosa,直到Wang et al.（2009）依据采自云南西双版纳的标本将其作为新种灰肉红菇发表并提供了ITS和LSU序列后,该种才有了自己的名称。典型的“大红菌”应具备R. griseocarnosa的基本特征：子实体大型,菌盖无放射状条纹,红色、黑红色,湿时粘,菌肉灰色,柔和,囊状体大,盖表皮栅状,无盖表囊状体,孢子具刺状疣突,刺疣间偶具连线。
Li et al.（2010）基于DNA序列发现中国南方俗称的“大红菌”包含3个互为单系的系统发育分支,灰肉红菇只是其中的一个分支。这3个系统发育分支形成了一个单系,与欧洲的R. vinosa形成姊妹分支。如在分析中加入GenBank中新近上传的一些来自中国样品的序列,KJ957323-KJ957340和KU552086,广义R. griseocarnosa下的分支会变得更为多样和复杂。笔者在云南哀牢山和马关县采集的个别标本（图2N）虽然总体上与灰肉红菇相似,但菌盖为黑黄褐色,与Das et al.（2011）定为灰肉红菇的印度样品极为接近。这些样品应作为不同于灰肉红菇的新物种还是它的种下分类群有待深入研究。根据笔者研究的标本,与灰肉红菇近缘的分支内的一些材料菌肉并不总是呈典型的灰色,盖表皮菌丝也表现出一定差异。所以,中国民间所称的“大红菌”真正的范围和物种数目还需进一步结合形态证据进行确认。Jiang et al.（2017）描述自中国的褐酒红菇R. brunneovinacea似与灰肉红菇近缘,但这种近缘关系很可能是由于取样不足引起的,新近上传至GenBank中的R. seperina的两条来自欧洲的序列（MG687341和MN130108）与褐酒红菇更为近缘。
3.4 臭红菇复合群
除以上主要的食用种类外,一些形态近似于臭红菇、桂樱红菇R. laurocerasi、点柄臭红菇R. senecis的种也频见于云南中南部、贵州等地市场（应建浙和臧穆 1994;王向华和刘培贵 2002）。在贵州,当地人将此类红菇用水煮沸后贮于罐中,使其发酵,作为酸蘑菇食用（应建浙和臧穆 1994）。在印度的西孟加拉地区,点柄臭红菇被当地居民作为美味食用菌采食（Khatua et al. 2015）。陈作红等（2016）认为这些物种食后会引起胃肠炎型中毒反应,故应慎食。这些物种菌盖湿时较粘、盖缘具有明显的颗粒状条纹,子实体常具不悦气味,属异褶亚属的劣味组R. sect. Ingratae（图1）。尽管该组是红菇属中最易识别的类群之一,组内物种间的界限却十分微妙。
Song et al.（2018）显示日本的R. senecis与中国的点柄红菇R. punctipes形成各自具有高支持率的姊妹分支。分析Song et al.（2018）使用的数据和GenBank中的数据可以看出,这是由于取样不足和数据的缺陷引起的。Song et al.（2018）所使用的日本的点柄臭红菇的ITS序列（AB531459和AB509717）仅有ITS2区,未收入同样来自日本的与AB531459和AB509717高度相似的环境样品序列AB259126、AB259127和AB605664以及来自韩国和印度的序列。如果将Lee et al.（2017）和Khatua et al.（2015）定为点柄臭红菇的DNA数据加入,会发现将Song et al.（2018）中的点柄臭红菇和点柄红菇作为一个种下的两个（地理）种群更为合适。如果这一分类学处理能够得到接受,R. punctipes这一种名由于早于R. senecis就成为正确的应被使用的名称。由于该组内多数ITS序列间的相似性较高,如不合理界定物种区分所使用的阈值,对ITS数据的质量不加严格控制,那些由于人为因素造成的数据错误或由于偶然的变异造成的碱基差异则会被解读为种间的遗传差异,从而产生大量的“小种”。在欧美地区,历史发表的大量物种与现代的DNA数据间的关联并没有解决。如在欧洲,定为臭红菇R. foetens的序列也并不聚集在一个分支上,在R. foetens这一名称的应用未被固定之前,无法确定那些序列上不同于真正的R. foetens的身份。在这样的类群中,过度依赖DNA序列尤其是ITS序列上的少量变异,过度依赖形态特征上的细微变化,最终会使得物种被划分得过细、过小。到了一定程度,一些物种间的界限会变得模糊、重叠,反而为人们认识物种带了诸多不便。
3.5 日本红菇
近年来,在中国中南部引起的一系列因误食毒菌引起胃肠炎型反应的事件中,日本红菇是最为常见的物种之一（Li et al. 2020b）。之所以特别提及该种,是因为它可能是最难与可食的近缘物种进行区分的有毒蘑菇。日本红菇所在的短柄亚属是红菇属中较小的亚属,包含了一类子实体中至大型、白色至淡黄褐色、肉厚致密、菌褶不等长、盖表皮不易剥离的物种,北温带和热带均有分布（Buyck et al. 2018）。著名的食用菌美味红菇、短柄红菇、灰绿红菇R. chloroides也属于这一亚属。由于形态上的相似性,历史上该亚属的很多物种都曾被作为另一物种的属下分类群,如灰绿红菇曾被作为R. delica的变种或变型,Kauffman（1918）将短柄红菇作为美味红菇的变种。尽管Shaffer（1964）从北美地区描述了多个新种,且Buyck & Adamčik（2013）对一些模式标本进行了研究,但GenBank中并没有鉴定为这些物种的序列。大量的样品还是被置于以上常见的名称下。Li et al.（2020a）基于中国的标本描述了与美味红菇近缘的3个新种,它们与相近种的区别更多表现在微妙的显微特征上,与欧美被定为美味红菇或短柄红菇的部分序列的相似度在98%以上。
与美味红菇等可食红菇不同,日本红菇并不属于乳菇状组R. sect. Lactarioidae,而属于淡孢亚组R. subsect. Pallidosporinae（图1）,它的孢子印色较深（ochraceous）,脐上区非淀粉质,盖表皮菌丝厚壁（Hongo 1954;Buyck & Adamčik 2013;Li et al. 2020a）。由于这些显微特征在野外识别中无法使用,加之日本红菇的菌肉柔和,采集者很容易将日本红菇误作美味红菇复合群（“石灰菌”、“背土菌”）的物种,甚至由于外形上的相似性将其误作辣味多汁乳菇采食。在GenBank中共有5条ITS序列被鉴定为R. japonica。分析这些序列可以发现,它们聚为两个分支,其中一支包含来自日本的序列（AB509603和AB597671）。日本红菇的真实身份尚需确定。
近年来,针对红菇科的分类学研究发现了一些完全不同于已知分支的新的系统发育分支。Wang et al.（2015a）在多汁乳菇属中发现了新组薄囊体组和异囊体组L. sect. Ambicystidiati,其中的异囊体乳菇是乳菇类真菌首次发现的具有退化乳汁管的物种;de Crop et al.（2017）在多汁乳菇属中发现了多个特有分布于热带非洲和南美的分支;Wang et al.（2019）基于采自云南东南部的样品,发现了红菇属的一个新的亚组近条纹亚组R. subsect. Substriatinae,这一亚组与热带非洲的类群有着较近的亲缘关系;Buyck et al.（2020）发现北美东部的R. glutinosa和云南中部的类胶粘红菇R. glutinosoides形成的分支代表了乳菇属的一个新亚属胶粘亚属R. subg. Glutinosae,且位于红菇属系统发育树的基部（图1）,有望为研究红菇属真菌的起源提供关键的证据。Wang et al.（2015a）在多汁乳菇属中发现的薄囊体组的物种在民间一直被作为食用菌采食并销售。这些发现一方面为鉴定红菇科可食物种带来了便利,同时也表明,红菇科真菌的多样性较我们所知的复杂。考虑到如亚稀褶红菇这类的剧毒物种与可食物种近缘,对于这些新描述的完全不同于已知的分类群,如果没有足够的来自民间或科学上的经验和证据,对其可食性应保持足够的警惕。
4 展望
近年来,DNA与形态学证据相结合发现了红菇科中大量的属下分类群,如亚属、组、亚组,它们均来自亚热带-热带地区,今后更多有趣的新发现将集中于亚热带、热带地区的新分类群,以及基于这些类群所建立的全球亚热带-热带地区之间的关联。全球亚热带-热带地区如东南亚、中美洲、非洲等地广泛采食的一些红菇科物种可能为新物种,有待描述发表。从已有的研究结果看,乳菇属和红菇属具有较强的温带性质（Looney et al. 2016）,这两属中存在一定数目的洲际广布种,这种现象在北方地区和高海拔地区表现得尤为突出（Adamčík et al. 2019）。所以,对北方和高海拔地区物种的识别,应充分考虑那些原初描述自欧洲、北美和亚洲的物种,必要时使用附加模式固定名称的使用。目前红菇科分类中的一个流行做法是,用ITS序列在GenBank中用BLASTn进行搜索对比,如果找不到高相似度的序列,即假定为新种,再从标本上寻找特征与已知的近缘物种进行区分。而事实上,红菇科中大量已发表的物种由于未被充分认识,在相关的DNA数据库中并无对应的序列。这种“BLAST不到相近序列即为新种”的观点和做法会割裂DNA数据与传统分类学研究成果间的关联,甚至产生多余的名称,需予以重视。
尽管Li et al.（2019）发现ITS-rpb2作为Barcode可以分辨出红菇的物种,但这很可能是因为其分析中仅包含了少量的物种（27种）且这些物种分散在不同的亚属中。在红菇科的研究中,无论是哪个类群,当来自全球各地的样品加入已有的DNA数据集时,均会出现以下现象,原有物种（分支）之间的界限变得模糊,分支的枝长变短,物种在形态上的网状交织关系变得愈发强烈。而一个令人担忧的趋势是,在取样不足或仅收录少量样品或物种时,依据形态和序列上相对明显的差异发表新种的做法变得越来普遍。最终的结果是,一些在形态和序列上具有过渡或混合式样的样品被遗留下来,无法归类。在今后的红菇科分类学研究中,随着发表的新种越来越多,一个需要重新考虑的问题是：那些新种是否应该被作为“种”对待,如果将它们作为种下分类群,如变种、变型甚至地理种群,是否更能体现它们之间的共性和形态及遗传上的交叉关系?多基因数据的应用被认为可以更好地界定分类群,但分类群等级的确定却需要更为全面的考量。需要根据每一类群的数据特点灵活设定物种的大小范围。在红菇科食用菌的鉴别中,如果考虑操作的可行性,应着眼于划分大种或种复合群,而不是用微妙的特征和DNA数据上的少量变化去识别更多的“小种”。
致谢：在开展与本论文有关的研究工作及论文撰写过程中,得到以下同行的帮助：湖南师范大学的陈作红教授提供亚稀褶红菇的照片并就毒菌问题进行过有益的讨论,中国疾病预防控制中心的李海蛟博士提供日本红菇的照片和相关DNA数据以及红菇科相关的中毒案例,河北农业大学的李国杰博士提供文献并核对相关DNA数据,云南省农业科学院的李树红研究员和苏开美研究员以及四川省农业科学院的何晓兰博士提供研究材料,贵州科学院的王晶和内蒙古大学的韩瀛参与野外调查、实验室工作和数据分析,法国自然历史博物馆的Bart Buyck博士就部分标本的野外鉴定提供指导并提供文献,中国科学院昆明植物研究所的杨祝良研究员、冯邦博士和葛再伟博士对论文的初稿进行了修改提高,曹书琴对文稿进行了校对,在此一并表示感谢。
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Although Sikkim belongs to one of the Global Biodiversity Hotspots, little is known about its ectomycorrhizal fungi, and even less about the main genera of Russulales, i.e. Lactarius, Lactifluus, Multifurca and Russula. Combining a multilocus genealogical and morphological study, we aimed to document the diversity within Lactifluus volemus sensu lato of Sikkim Himalaya. We compared nuclear ITS and LSU rDNA, nuclear rpb1 and rpb2 protein-coding, and mitochondrial atp6 protein-coding genealogies to determine species boundaries. Interspecific relationships were inferred from the combined dataset. Bayesian and maximum likelihood single-locus genealogies are concordant and support recognition of six species. Three of these could be identified by unique morphological characteristics and are described as new species: L. dissitus, L. leptomerus and L. versiformis.
Lactarius volemus sensu lato (Russulales) from northern Thailand: morphological and phylogenetic species concepts explored
DOI:10.1007/s13225-010-0070-0
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[本文引用: 1]
Lactarius volemus (Fr.: Fr.) Fr. is a well known and morphologically easily recognizable milkcap of the Northern hemisphere, forming ectomycorrhiza with both deciduous and coniferous trees. It was originally described from Europe, but is also reported in other continents. Although it is characterized by several unique macro- and micromorphological features, substantial variation in colour, lamellae spacing and changing and staining of the latex has been recorded and it is therefore considered as a putatively unresolved species complex. This study explores the concordance between morphological and phylogenetic species concepts within L. volemus sensu lato of northern Thailand, combining a critical morphological scrutiny with a multiple gene genealogy based on LSU, ITS and rpb2 nuclear sequences. Twelve strongly supported monophyletic clades and six terminal branches are discernable in all phylogenetic trees and represent 18 phylogenetic species. Six of the monophyletic clades can be morphologically distinguished and are described as new species: L. acicularis, L. crocatus, L. distantifolius, L. longipilus, L. pinguis and L. vitellinus. Five other clades also show some morphological differences, but these are too subtle and do not allow for a clear-cut species delimitation without the corroboration of molecular data. Lactarius volemus sensu lato of northern Thailand is therefore still considered as a partially cryptic species complex. Pleurolamprocystidia, pileipellis hairs and to a lesser degree also pileus colour are important diagnostic characteristics. Spore morphology, latex discoloration and pileus surface texture are less useful as diagnostic features. Whether this rich diversity is the result of in situ Pleistocene survival or post-glacial expansion and subsequent radiation, has yet to be revealed.
Lactifluus volemus in Europe: three species in one - revealed by a multilocus genealogical approach, Bayesian species delimitation and morphology
DOI:10.1016/j.funbio.2015.08.015
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PMID:26693681
[本文引用: 1]
This study provides morphological and molecular evidence (from nuclear ITS, LSU, and rpb2 DNA sequences) for three previously unrecognized species within the morphospecies Lactifluus volemus from Europe. Phylogenetic species are supported by both a multi-locus tree-based method and Bayesian species delimitation. Lactifluus volemus and Lactifluus oedematopus are provided with a new description, and a third species, Lactifluus subvolemus, is described as new to science. Lactifluus oedematopus can be easily recognized by its short pileipellis hairs. Both L. volemus and L. subvolemus have longer pileipellis hairs and can only be distinguished from each other based on cap colour. Intermediary colour forms, however, occur as well, and cannot be identified as either L. volemus or L. subvolemus without molecular data. Revealing that L. volemus--already considered extinct in the Netherlands and the Belgian Flemish region, and declining in other European countries--is actually a complex of three species that are even more vulnerable to extinction, this study emphasizes the fundamental role of taxonomy in species conservation.
Not every milkcap is a Lactarius
Studies in tropical African Lactarius species 7. A synopsis of the section Edules and a review on the edible species
Monograph of Lactarius in tropical Africa. Fungus flora of tropical Africa. Vol. 2
New combinations in Lactifluus. 1. L. subgenera Edules, Lactariopsis, and Russulopsis
DOI:10.5248/118.447 URL [本文引用: 1]
Lactarius (Basidiomycota, Russulaceae) in Papua New Guinea. 3. Two new Lactarius species in subgenus Plinthogali
DOI:10.1071/SB01036 URL [本文引用: 2]
Visiting Russula (Russulaceae, Russulales) with samples from southwestern China finds one new subsection of R. subg. Heterophyllidia with two new species
DOI:10.1007/s11557-019-01487-1 URL [本文引用: 4]
Type studies on the Lactarius species published from China
DOI:10.3852/mycologia.99.2.253
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PMID:17682778
Thirteen species and one variety of the genus Lactarius (Russulaceae, Russulales) have been published from China. Since the original descriptions and illustrations are often poor, it is difficult to understand the concepts of these taxa. Therefore, the type specimens of all these taxa were traced, reviewed and (or) re-examined by employing modern taxonomic methods. The type collections of L. atrosquamulosus and L. omeiensis could not be found. The type collection of L. minimus var. macrosporus was found to consist of mixed material belonging to two different species and a lectotype, which belongs to Lactarius subgen. Russularia, sect. Olentes was designated. Lactarius squamulosus turned out to be a synonym of L. gracilis and L. wangii was synonymized here with L. volemus. Except for L. wangii, the others were known only from China or Asia. In total, ten taxa were provided with modern or new macromorphological descriptions based on the dried basidiocarps and with illustrations of basidiospores, pleuromacrocystidia, pileipellis and (or) stipitipellis. As a result of this re-examination, the differences between these taxa and their closely related taxa were discussed and their taxonomic positions in four contemporary infrageneric classification systems of Lactarius were proposed. Based on recent conspecific collections, information on habitats and geographical distribution of some species was provided.
Three new species of Lactarius sect. Deliciosi from subalpine-alpine regions of central and southwestern China
DOI:10.7872/crym/v37.iss4.2016.493 URL [本文引用: 2]
Two new species of Lactarius subg. Russularia from subalpine regions of southwestern China
Fungal biodiversity profiles 71-80
DOI:10.7872/crym/v39.iss4.2018.419 URL [本文引用: 2]
Revisiting the morphology and phylogeny of Lactifluus with three new lineages from southern China
DOI:10.3852/13-393
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PMID:26240310
[本文引用: 2]
As a recent group mainly defined by molecular data the genus Lactifluus is in need of further study to provide insight into the morphological and molecular variation within the genus, species limits and relationships. Phylogenetic analyses of nuc rDNA ITS1-5.8S-ITS2 (ITS), D1 and D2 domains of nuc 28S rDNA (28S), and part of the second largest subunit of the RNA polymerase II (rpb2) (6-7 region) sequences of 28 samples from southern China revealed three new lineages of Lactifluus. Two of them are nested in a major clade that includes the type of Lactifluus and here is treated as two new sections: L. sect. Ambicystidiati and L. sect. Tenuicystidiati. Lactifluus ambicystidiatus, described here as a new species (= sect. Ambicystidiati), has both lamprocystidia and macrocystidia in the hymenium, a unique combination of features within Russulaceae. Furthermore, only remnants of lactiferous hyphae are present in L. ambicystidiatus and our results suggest that the ability to form a lactiferous system has been lost in this lineage. Lactifluus sect. Tenuicystidiati forms a strongly supported monophyletic group as a sister lineage to L. sect. Lactifluus. We recognize it based on the thin-walled macrocystidia and smaller ellipsoid spores with an incomplete reticulum compared with L. sect. Lactifluus. The former placement of L. tenuicystidiatus in the African L. sect. Pseudogymnocarpi is not supported. Using genealogical concordance we recognize five phylogenetic species within L. sect. Tenuicystidiati and describe two of these as new, L. subpruinosus and L. tropicosinicus. The third lineage, represented by L. leoninus, forms a sister group to L. subg. Lactariopsis sensu stricto. The three lineages provide further evidence for morphological features in Lactifluus being homoplasious. Some sections and species complexes are likely to be composed of more species and merit further investigations. Subtropical-tropical Asia is likely a key region for additional sampling.
Phylogeny, biogeography and taxonomic reassessment of Multifurca (Russulaceae, Russulales) using three-locus data
DOI:10.1371/journal.pone.0205840 URL [本文引用: 5]
Resources investigation and studies on the wild commercial fungi in Yunnan
Yunnan is an area rich in wild commercial fungi resources, namely, fungi sold in markets. Much attention has been paid to fungal resources in the past. However, comprehensive investigation and studies on the resources were wanting. Through investigation, collection and identification of species, the geographical distribution patterns, seasonal changes and ranking based on the trade volume and the economic value of the resources were documented continuously from 1997 to 2001. In total, 207 species (including varieties and forms) belonging to 64 genera are recognized, showing high diversity and high endemicity. Among the species, those of the genera Boletus, Tricholoma, Termitomyces, Thelephora, Russula, Lactarius, and Ramaria are main commercial fungi. Despite artificial selection, commercial fungi show obvious geographical distribution patterns: southern Yunnan is rich in tropical species and northwestern Yunnan is rich in temperate alpine and subalpine species. Central and southwestern Yunnan fungal communities display subtropical and temperate characters. Each area possesses its own typical species and they are related by some transitional ones. We ranked the commercial fungi species in the order of trade volume. Those of grade I and II are the dominant species. Almost 90% of the commercial fungi are edible and about 5% are medicinal, while 7% are poisonous. The economic value of each species is also provided.
Lactarius vividus sp. nov. (Russulaceae, Russulales), a widely distributed edible mushroom in central and southern China
Three new species of Lactarius subgen. Lactiflui
DOI:10.1127/0029-5035/2006/0083-0167 URL [本文引用: 1]
Russula griseocarnosa sp. nov. (Russulaceae, Russulales), a commercially important edible mushroom in tropical China: mycorrhiza, phylogenetic position, and taxonomy
DOI:10.1127/0029-5035/2009/0088-0269 URL [本文引用: 1]
Lactarius subgenus Russularia (Basidiomycota, Russulales): novel Asian species, worldwide phylogeny and evolutionary relationships
DOI:10.1016/j.funbio.2016.08.004 URL [本文引用: 1]
Lactarius subgenus Russularia (Russulaceae) in South-East Asia: 3. new diversity in Thailand and Vietnam
DOI:10.11646/phytotaxa.207.3 URL [本文引用: 1]
Resource diversity of Chinese macrofungi: edible, medicinal and poisonous species
DOI:10.1007/s13225-019-00432-7 URL [本文引用: 1]
Russula dinghuensis sp. nov. and R. subpallidirosea sp. nov., two new species from southern China supported by morphological and molecular evidence
DOI:10.7872/crym/v38.iss2.2017.191 URL [本文引用: 1]
Russula nigrovirens sp. nov. (Russulaceae) from southwestern China
DOI:10.11646/phytotaxa.236.3 URL [本文引用: 2]
云南野生贸易真菌资源调查及研究
野生贸易真菌即市场上出售的野生真菌，云南的野生贸易真菌资源十分丰富且长期以来受到关注。但以往对于这一资源的全面调查和研究较为缺乏。本项目通过市场调查、标本采集和鉴定，自1997～2001年连续4年对云南野生贸易真菌从种类、地理分布、季节变化、基于贸易量的优势度评价和经济价值5方面进行研究。云南野生贸易真菌已知共64属207种（含变种、变型），以担子菌中的牛肝菌属（Boletus）、口蘑属（Tricholoma）、鸡土从菌属（Termitomyces）、革菌属（Thelephora）、红菇属（Russula）、乳菇属（Lactarius）、丛枝瑚属（Ramaria）为主，表现出较高的多样性和较强的地区特有性。尽管具有人为选择的因素，但云南野生贸易真菌仍表现出较强的地理分布特异性：滇南地区包含较多的热带种类，具有较强的热带性质，而滇西北则具有较强的温带高山、亚高山性质，滇中、滇西南等地表现出亚热带和温带的过渡特征。以上各地区既有各自的代表种类，又通过某些过渡类群存在一定的联系。根据各个种贸易量的相对大小即优势度将贸易真菌分为4个等级，其中的优势Ⅰ级和优势Ⅱ级为主导种类。云南贸易真菌中约近90%的种类为食用菌，5%为药用菌，另有约7%为有毒种类。对每一种贸易真菌的经济价值给予了评价。
