Freie Universität Berlin, Institut für Biologie - Botanik, Altensteinstraße 6, D-14195 Berlin, Germany.
Mol Phylogenet Evol. 2013 Sep;68(3):604-18. doi: 10.1016/j.ympev.2013.04.009. Epub 2013 Apr 19.
Recent studies all indicated that both the affinities and subdivision of Boraginaceae s.str. are unsatisfactorily resolved. Major open issues are the placement and affinities of Boraginaceae s.str. in Boraginales and the major clades of the family, with especially the large tribes Cynoglosseae and Eritrichieae repeatedly retrieved as non-monophyletic groups, and the doubtful monophyly of several larger genera, especially Cynoglossum and Omphalodes. The present study addresses and solves these questions using two plastid markers (trnL-trnF, rps16) on the basis of a sampling including 16 outgroup taxa and 172 ingroup species from 65 genera. The phylogeny shows high statistical support for most nodes on the backbone and on the individual clades. Boraginaceae s.str. are sister to African Wellstediaceae, Wellstediaceae-Boraginaceae s.str. is sister to African Codonaceae. Echiochileae are retrieved as sister to the remainder of Boraginaceae s.str., which, in turn, fall into two major clades, the Boragineae-Lithospermeae (in a well-supported sister relationship) and the Cynoglosseae s.l. (including Eritrichieae). Cynoglosseae s.l. is highly resolved, with Trichodesmeae (incl. Microcaryum, Lasiocaryum) as sister to the remainder of the group. Eritrichieae s.str. (Eritrichium, Hackelia, Lappula) are resolved on a poorly supported polytomy together with the Omphalodes-clade (incl. Myosotidium, Cynoglossum p.p.), and the Mertensia-clade (incl. O. scorpioides, Asperugo). The Myosotideae (Myosotis, Trigonotis, Pseudomertensia) are retrieved in a well-supported sister-relationship to the core-Cynoglosseae, the latter comprising all other genera sampled. Cynoglossum is retrieved as highly para- and polyphyletic, with a large range of generic segregates embedded in Cynoglossum, but other species of Cynoglossum are sister to Microula or to the American "Eritrichieae" (Cryptantha and allied genera). Representatives of the genus Cynoglossum in its current definition are segregated onto six independent lineages, members of Omphalodes onto three independent lineages. At least 11 of the genera here sampled are deeply nested in other genera. The data show that individual details of nutlet morphology (e.g., winged margins, glochidia) are highly homoplasious. Conversely, a complex of nutlet characters (e.g., characters of the gynobase and cicatrix together with nutlet orientation and sculpturing) tends to circumscribe natural units. Geographical distribution of major clades suggests that the family originated in Africa and western Asia and radiated to eastern Eurasia, with several independent dispersal events into Australia and the New World.
最近的研究表明，滨藜科的亲缘关系和分支都没有得到令人满意的解决。主要的悬而未决的问题是滨藜科在滨藜目中的位置和亲缘关系，以及该科的主要分支，尤其是大的 Cynoglosseae 和 Eritrichieae 族被反复发现为非单系群，并且几个较大属的单系性值得怀疑，尤其是 Cynoglossum 和 Omphalodes。本研究利用两个质体标记（trnL-trnF，rps16），在包括 65 个属的 16 个外群和 172 个内群物种的基础上，解决了这些问题。系统发育显示，大多数骨干和个别分支的节点都具有很高的统计支持。滨藜科与非洲 Wellstediaceae 密切相关，Wellstediaceae-Boraginaceae 与非洲 Codonaceae 密切相关。Echiochileae 被认为是滨藜科的姊妹群，而滨藜科则分为两个主要分支，Boragineae-Lithospermeae（在一个支持良好的姊妹关系中）和 Cynoglosseae s.l.（包括 Eritrichieae）。Cynoglosseae s.l.得到了高度解析，其中 Trichodesmeae（包括 Microcaryum、Lasiocaryum）与该组的其余部分为姊妹群。Eritrichieae s.str.（Eritrichium、Hackelia、Lappula）在一个支持不佳的多系中与 Omphalodes 分支（包括 Myosotidium、Cynoglossum p.p.）和 Mertensia 分支（包括 O. scorpioides、Asperugo）一起得到解决。Myosotideae（Myosotis、Trigonotis、Pseudomertensia）在一个支持良好的姊妹关系中被认为是核心 Cynoglosseae 的姊妹群，后者包括所有其他取样的属。Cynoglossum 被认为是高度并系和多系的，其包含大量的属分群嵌入 Cynoglossum 中，但 Cynoglossum 的其他物种与 Microula 或美洲的“Eritrichieae”（Cryptantha 和相关属）为姊妹群。当前定义中的 Cynoglossum 属的代表被分离成六个独立的谱系，而 Omphalodes 的成员则分离成三个独立的谱系。在此取样的至少 11 个属都深深嵌套在其他属中。数据表明，坚果形态的个别细节（例如，有翅边缘、小钩）具有高度的同形性。相反，一组坚果特征（例如，雌蕊基部和疤痕以及坚果方向和雕刻的特征）往往会限定自然单位。主要分支的地理分布表明，该科起源于非洲和西亚，并辐射到东亚，有几个独立的扩散事件进入澳大利亚和新世界。
