Chase Mark W, Williams Norris H, de Faria Aparacida Donisete, Neubig Kurt M, Amaral Maria do Carmo E, Whitten W Mark
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK.
Ann Bot. 2009 Aug;104(3):387-402. doi: 10.1093/aob/mcp067. Epub 2009 Apr 4.
Floral morphology, particularly the angle of lip attachment to the column, has historically been the fundamental character used in establishing generic limits in subtribe Oncidiinae (Orchidaceae), but it has also been long recognized that reliance on this character alone has produced a highly artificial set of genera. In essence, lip/column relationships reflect syndromes associated with pollinator preferences; most genera of Oncidiinae as previously defined have consisted of a single floral type. Here, the degree to which this has influenced generic delimitation in Brazilian members of the largest genus of Oncidiinae, Oncidium, which previous molecular (DNA) studies have demonstrated to be polyphyletic, is evaluated.
Phylogenetic analyses of the following multiple DNA regions were used: the plastid psbA-trnH intergenic spacer, matK exon and two regions of ycf1 exon and nuclear ribosomal DNA, comprised of the two internal transcribed spacers, ITS1 and ITS2, and the 5.8S gene. Results from all regions analysed separately indicated highly similar relationships, so a combined matrix was analysed.
Nearly all species groups of Brazilian Oncidium are only distantly related to the type species of the genus, O. altissimum, from the Caribbean. There are two exceptions to this geographical rule: O. baueri is related to the type group and O. orthostates, an isolated species that lacks the defining tabula infrastigmata of Oncidium, is not exclusively related to any previously described genus in the subtribe. Several well-supported subclades can be observed in these results, but they do not correspond well to sections of Oncidium as previously circumscribed or to segregate genera as defined by several recent authors. In spite of their floral differences, these groups of Oncidium, formerly treated as O. sections Barbata, Concoloria pro parte, Crispa, Ranifera, Rhinocerotes, Rostrata (only O. venustum), Synsepala, Verrucituberculata pro parte and Waluewa, form a well-supported clade with Gomesa (including Rodrigueziella and Rodrigueziopsis) embedded in it. Two often recognized segregate genera, Baptistonia and Ornithophora, and the recently described Carriella are also embedded within the Brazilian clade. The level of variation within major subclades of the Gomesa clade is low and similar to that observed within other genera of Oncidiinae.
Convergence on a stereotypical syndrome of floral traits associated with pollination by oil-collecting bees has resulted in these characters not being reliable for producing monophyletic taxa, and the genus Oncidium, defined by these characters, is grossly polyphyletic. Vegetative and a few floral/inflorescence characters link these taxa with a mainly Brazilian distribution, and they were all transferred to Gomesa on this basis rather than separated from Gomesa based on their floral differences, which we hypothesize to be simple shifts in pollination strategies. Other authors have described a large number of new genera for these former members of Oncidium, but most of these are not supported by the results presented here (i.e. they are not monophyletic). A new genus, Nohawilliamsia, is described for O. orthostates because it does not fit in any currently recognized genus and is only distantly related to any other member of Oncidiinae.
花的形态,尤其是唇瓣与蕊柱的附着角度,一直是历史上用于界定文心兰亚族(兰科)属级界限的基本特征,但长期以来人们也认识到仅依赖这一特征会产生一套高度人为划分的属。本质上,唇瓣/蕊柱关系反映了与传粉者偏好相关的综合征;如之前所定义,大多数文心兰亚族的属都由单一花型组成。在此,评估这种情况在文心兰亚族最大属——文心兰属的巴西成员中对属级界定的影响程度,之前的分子(DNA)研究已证明该属是多系的。
对以下多个DNA区域进行系统发育分析:质体psbA - trnH基因间隔区、matK外显子以及ycf1外显子的两个区域和核糖体DNA,后者由两个内部转录间隔区ITS1和ITS2以及5.8S基因组成。分别分析所有区域得到的结果显示出高度相似的关系,因此对合并矩阵进行了分析。
巴西文心兰的几乎所有物种组与该属的模式种、来自加勒比地区的高文心兰亲缘关系都很疏远。这一地理规律有两个例外:鲍氏文心兰与模式组相关,而直立柱文心兰是一个孤立物种,缺乏文心兰属典型的柱头下板,它并不完全与该亚族中任何先前描述的属相关。在这些结果中可以观察到几个得到有力支持的亚分支,但它们与之前界定的文心兰属的组或几位近期作者定义的分立属并不完全对应。尽管它们的花存在差异,但这些文心兰组,以前被视为文心兰属的髯毛组、部分纯色组、皱边组、瘤瓣组、犀角组、具喙组(仅细纹文心兰)、合萼组、疣瘤组部分以及瓦卢埃瓦组,与嵌入其中的戈梅萨属(包括罗氏兰属和拟罗氏兰属)形成了一个得到有力支持的分支。两个经常被认可的分立属,巴氏兰属和鸟舌兰属,以及最近描述的卡里兰属也嵌入在巴西分支内。戈梅萨分支主要亚分支内的变异水平较低,与在文心兰亚族其他属中观察到的情况相似。
趋同于与采油蜂传粉相关的花部特征的刻板综合征,导致这些特征对于产生单系类群不可靠,并且由这些特征定义的文心兰属是高度多系的。营养体特征以及一些花部/花序特征将这些主要分布在巴西的类群联系起来,基于此它们都被转移到戈梅萨属,而不是基于它们的花部差异与戈梅萨属分开,我们推测这些花部差异只是传粉策略的简单转变。其他作者为这些以前的文心兰属成员描述了大量新属,但这里呈现的结果并不支持其中大多数(即它们不是单系的)。为直立柱文心兰描述了一个新属——诺哈威廉姆斯兰属,因为它不符合任何目前认可的属,并且与文心兰亚族的任何其他成员亲缘关系都很疏远。