豆科英文名为Fabaceae，或者Leguminosae，“Fabaceae”这个名字来自已经不复存在的Faba属，Faba属现在被包含在Vicia中。术语“faba”来自拉丁语，似乎只是指“豆”。Leguminosae则是一个比较久远，但仍是有效的名称。这些植物的果实，称为豆类。

豆科是物种数量第三大的陆地植物家族，仅次于兰科和菊科，约有770属和约19 500种已知物种，约占所有开花植物种类的7% (Magallón and Sanderson, 2001; Judd et al., 2010; LPWG, 2013a; Christenhusz and Byng, 2016)。五个最大的属分别是Astragalus (超过3 000种)，Acacia (超过1 000种)，Indigofera (约700种)，Crotalaria (约700种)和Mimosa (约400种)，约占所有豆科植物种类的四分之一。

从经济上讲，豆科植物的重要性仅次于禾本科植物。例如，据估计，从1990年到2012年世界豆类出口增加了一倍多，从660万吨增加到1 340万吨，2012年豆类出口价值估计为95亿美元(粮农组织[FAO]: http://www.fao.org/pulses-2016/en/)。联合国大会将2016年定为国际豆类年，以提高人们对豆类的营养价值、对粮食安全、可持续农业的重要性以及对减少生物多样性丧失和气候变化的认识。豆科植物是重要的粮食作物，是高营养蛋白质和微量营养素的来源，可以极大地有益于健康和生计，尤其是在发展中国家(Yahara et al., 2013)。

自农业开始以来，豆科植物已经在世界不同地区与禾本科植物一起驯化，并在其早期农业发展中发挥了关键作用(Gepts et al., 2005; Hancock, 2012)。许多豆科植物几千年来一直是人类的主食，它们的使用与人类的进化密切相关(Dimitri, 1987)，在当今世界，豆科植物在制造木材、油和树脂、清漆、油漆、染料、药品等领域具有重要的作用，同样也是园林绿化的不可或缺的一部分。在温带和热带地区，豆科植物作为饲料和绿肥也具有独特的重要性。

豆科植物分布广泛，在全球几乎所有的生物群落中都是重要的生态组成成分，甚至在最极端的生境中也有分布(Schrire et al., 2005)。在非洲、南美洲和亚洲的热带森林中，它们是构成物种多样性和丰度的重要元素(Banda et al., 2016)，在整个热带地区，它们主宰干燥森林和热带稀树大草原，同样的情况也发生在地中海地区，沙漠和温带地区，高纬度和高海拔地区。豆科植物可以是大型的乔木、多年生或一年生草本植物、一年生或多年生的有卷须攀缘植物、灌木、木本藤本植物，以及较少见的水生植物。豆科植物花的对称性涵盖了从辐射对称到两侧对称和非对称的全部范围，这些花适应广泛的传粉媒介，如昆虫、鸟类和蝙蝠。

大多数豆科植物与土壤根瘤菌共生固定大气氮的能力可能是该家族最著名的生态特征，然而，并非所有的豆类都与固氮细菌有关。总而言之，该家族在形态、生理和生态上都异常多样化，是植物进化多样化的最显著例子之一。所有这些特征使得豆科生物学家对豆科的生物、家族多样性和进化、功能特征的进化以及家族的生态和生物地理学一直保持着浓厚的兴趣(Stirton and Zarucchi, 1989; Lavin et al., 2004; Champagne et al., 2007; Simon et al., 2009; Doyle, 2011; Simon and Pennington, 2012; Koenen et al., 2013; Moncrieff et al., 2014; Dugas et al., 2015)。

1豆科进化

Fabales包含约7.3%的双子叶植物物种，而这种多样性的大部分仅包含在该目所包含的四个科之一：Fabaceae中，该分支还包括Polygalaceae，Surianaceae和Quillajaceae。Fabales起源可追溯到9 400万至8 900万年前，而开始多样化则是在约7 900万至7 400万年前(Angiosperm Phylogeny Website. Version 7, May 2006. http://www.mobot.org/MOBOT/Research/APweb/orders/fabalesweb.htm#Fabaceae managed by Stevens, P. F)。事实上，豆科植物在第三纪早期已经多样化，与其他属于开花植物的科成为现代地球生物群中普遍存在的一部分(Herendeen et al., 1992; Lewis et al., 2005)。

豆科植物有丰富多样的化石记录，特别是在第三纪，这一时期的花、果、叶、木和花粉化石在许多地方都有发现(Crepet and Taylor, 1985; Crepet and Taylor, 1986; Herendeen, 2001)。而最早确定属于豆科的化石出现在古新世晚期(约5 600万年前) (Herendeen and Wing, 2001; Wing et al., 2004)。Cesalpinioideae，Papilionoideae和Mimosoideae是传统上被认为是代表豆科植物成员的3个亚科，以及这些亚科中的数量庞大的演化支成员——例如genistoides，发现在随后的一段时期内出现，从5 500万至5 000万年前开始(Herendeen et al., 1992)。事实上，从始新世中期到晚期的化石记录中，已经发现了代表豆科主要谱系的各种类群，表明现代豆科的大多数类群已经存在，并在这一时期出现了广泛的多样性(Herendeen et al., 1992)。

因此，豆科大约在6 000万年前开始多样化，而最重要的分支大约在5 000万年前分离，其中，主要的Cesalpinioideae分支为5600至3400万年前，而Mimosoideae基础族群则为(4 400±260)万年前(Lavin et al., 2005; Bruneau et al., 2008)。Mimosoideae和Faboideae之间的划分可以追溯到5 900万至3 400万年前，其中Faboideae基础族群为(5 860±20)万年前(Wikström et al., 2001)。尽管每个属内的物种多样化相对较晚，但仍有可能确定Faboideae内某些类群的分化年代。例如，Astragalus在大约1 600万到1 200万年前从Oxytropis中分离出来，此外，Neoastragalus的非整倍体物种的分离始于400万年前，Inga是Papilionoideae的另一个属，约有350种，在过去200万年中似乎发生了分化(Wojciechowski et al., 1993; Wojciechowski, 2003; Wojciechowski, 2005; Wojciechowski et al., 2006)。

根据化石和系统发育的证据，已有证据表明豆科植物最初是在古近纪时期沿Tethys seaway的干旱或半干旱地区进化而来的(Schrire et al., 2005)。然而，另一些人认为，还不能排除非洲(甚至美洲)是该族群的起源(Doyle and Luckow, 2003; Pan et al., 2010)。

目前关于根瘤所需基因进化的假设是豆科植物在多倍体事件后从其他途径获得(Yokota and Hayashi, 2011)。已经有几种不同的途径被认为是供给根瘤所需复制基因的途径，这些途径的主要供体为丛枝菌根共生基因、花粉管形成基因和血红蛋白基因，而参与植物-细菌识别的SYMRK基因则是丛枝菌根途径与根瘤化途径的主要共享基因之一(Markmann et al., 2008)。花粉管的生长与侵染线的生长相似，侵染线呈极性生长，与花粉管向胚珠的极性生长相似，这两种途径均需要相同类型的果胶降解细胞壁酶(Rodríguez-Llorente et al., 2004)。根瘤中的酶在固氮过程中需要大量的ATP，但同时对自由氧又非常敏感，因此，为了解决这一矛盾的情况，这些植物表达一种叫leghaemoglobin的血红蛋白，这种基因被认为是在多倍体事件后获得(Downie, 2005)。

2豆科分类系统构建

豆科植物的系统发育一直是世界各地研究小组的研究对象。这些研究小组利用形态学、DNA数据(叶绿体内含子trnL, 叶绿体基因rbcL和matK, 或核糖体间隔子ITS)和支链分析来研究该家族不同谱系之间的关系(Lavin et al., 1990; Käss and Wink, 1996; Sanderson and Wojciechowski, 1996; Käss and Wink, 1997; Bruneau et al., 2008; Cardoso et al., 2013; Félicien et al., 2018)。

早期学者主要依据豆科的形态学对其分类，该传统分类方法延续使用了很长的一段时间，典型的就是3亚科分类系统。但是随着科技的不断发展，分子系统学崛起，传统分类系统的弊端逐渐显现出来，此时学者意识到3亚科分类系统是不合理的，迫切需要一个新的分类系统来取代原有的系统。在2010年的时候，成立的豆科系统发育工作组(Legume Phylogeny Working Group (LPWG))就是针对研究新型分类系统，并在2017年正式提出了6亚科分类系统，一直沿用至今。

3豆科的3亚科分类系统

与一些其他大型被子植物家族相反，如禾本科(Grass Phylogeny Working Group, 2001; Grass Phylogeny Working Group, 2012)、菊科(Panero and Funk, 2002)，豆科植物亚科级一直是一个广泛使用的中心级别。在传统上习惯根据花形态的不同分为含羞草亚科(Mimosoideae)、云实亚科(Caesalpinioideae)和蝶形花亚科(Papilionoideae)3个亚科(Hutchinson, 1964; Cronquist, 1981)。这3个亚科的首次命名是在1825年，Candolle将豆科细分为四个亚目(=亚科)，除了第四个“亚目”Swartzieae外，其余3个“亚目”则为今天的亚科(Candolle, 1825)。Bentham (1865)对这3亚科分类系统进行了详细的阐述，其分类成为了随后140年豆科植物家族分类的基础(Taubert, 1891; Lewis et al., 2005)。

Hutchinson将3亚科提升至科级(Hutchinson, 1926; Hutchinson, 1964)，而在豆科系统学进展的第一卷中，这三组被认为是亚科(Polhill and Raven, 1981)。不管等级如何，自19世纪以来，这3个类群一直是被用作是鉴定和分类豆科植物的标准，在植物学、植物区系学和分类学课程中广泛教授，并且一直被世界各地的农学家，园艺学家和生态学家使用。

4探讨3亚科分类系统存在的问题

3亚科分类系统的分类基础基本上是基于一组显著的花特征，尤其是花瓣排列模式(Caesalpinioideae呈上升覆瓦状; Mimosoideae呈镊合状; Papilionoideae呈下降覆瓦状)和花的对称性(Caesalpinioideae为不确定性对称, 图1; 图2; 图3; Mimosoideae为辐射对称,图4; Papilionoideae为两侧对称, 图5; 图6; 图7)。虽然这些花的一些特征可能对定义Papilionoideae和Mimosoideae很有用，但它们在定义传统的Caesalpinioideae存在很大局限性(Tucker, 2003; Bruneau et al., 2014)。此外，即使对于Papilionoideae和Mimosoideae，这些花的大部分特征现在被认为是同源的(Pennington et al., 2000)。例如，以辐射对称花为特征的单个种或演化枝在Papilionoideae的基础上独立演化多次(Pennington et al., 2000; Cardoso et al., 2012b; Cardoso et al., 2013a; Ramos et al., 2016) (图5; 图6; 图7)。类似地，虽然辐射对称是Mimosoideae最显著的特征，但分布在MCC (Mimosoideae-Caesalpinieae-Cassieae)演化枝上密切相关的谱系也具有辐射对称的花(图3)。

无论分类单元或基因取样如何，豆科植物作为单系群在所有的分子系统发育分析中都得到了强有力的支持(LPWG, 2013a)。尽管豆科植物家族的近源物种存在不确定性(Dickison, 1981; APG III, 2009; Bello et al., 2009)，但自从家族成立以来，其单一性和独特性从未在形态学上受到质疑(Lewis et al., 2005; Bello et al., 2012)。除了少数例外，这个科最显著的特征是雌蕊通常由单心皮所组成，子房上位，1室，基部常有柄或无，沿腹缝线具侧膜胎座，胚珠2至多颗，悬垂或上升，排成互生的2列(Lewis et al., 2005)。然而，豆科系统学家早就意识到目前的亚科分类与新出现的系统发育结果之间的差异(Irwin, 1981; Käss and Wink, 1996; Doyle et al., 1997)，最出名的是并系群Caesalpinioideae，其许多族与亚族都非单系群，这意味着这些分支的系统发育结构并未直接反映在当前的分类中(Lewis et al., 2005)。

因此，豆科分类学界需要豆科新分类系统来取代传统3亚科分类系统也变得势在必行。

5豆科的6亚科分类系统

由于豆科分类学界对于支序分类体系比较热衷，2010年，豆科系统发育工作组(Legume Phylogeny Working Group (LPWG))成立，该国际团体致力于推动豆科系统发育研究，其主要任务是提出豆科系统发育研究概要和计划安排以及拟要解决的核心问题。LPWG于2013年，在南非约翰内斯堡召开第6届国际豆科植物会议，其主题为“豆科新分类系统”，于是豆科分类系统的重新修订被正式提上议事日程(LPWG, 2013)。

直至2017年，LPWG基于叶绿体matK序列数据，构建了一棵迄今为止取样最全(约91%的属和20%的种)的豆科系统发育树，并结合形态证据，提出了六个亚科的新分类系统，该分类系统中，仅传统的Papilionoideae (图5; 图6; 图7)被保留；传统的Mimosoideae不复存在，而成为新Caesalpinioideae (图3; 图4)一个分支mimosoid clade；除保留在新Caesapinioideae中的类群，原Caesapinioideae其余四个分支新拟为四个亚科：即Cercidoideae (图1A-F)、Detarioideae Burmeist (图2)、Dialioideae (图1H-L)和Duparquetioideae (图1G) (LPWG, 2017)。

毋庸置疑新的分类系统对豆科分类学、基因组学、发育生物学和进化生物学研究将会产生重要影响，而且随着研究不断深入，豆科的分类系统将会愈发完善。

6展望

不论是从经济价值，还是学术上的研究价值，豆科植物的重要性不言而喻。尤其是豆科植物几乎包罗了绝大部分的植物的形态、生长、发育等多个维度，如其花的形态、分布范围等，是植物多样性研究最好的物种之一。关于豆科植物分类系统的研究也一直都在不断探寻和摸索中，分类系统从3亚科过渡到6亚科，越来越合理，但是其中依然存在较多的争议和不能圆满解决的问题。不过科学是在不断进步的，豆科分类学的研究会进一步深化，会更加合理化，这不仅仅对豆科本身有重要的意义，同时对研究植物进化多样性的研究也起到重大的参考价值。

作者贡献

黎建辉负责文献收集、论文写作；张洁是项目负责人，负责论文修改和定稿；俞诗莹负责论文翻译；王浙茹、刘楚楚负责修改和校对。全体作者都阅读并同意最终的文本。

致谢

本研究由诸暨市翠溪生物技术研究院《翠溪创新研发项目基金》资助。

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