Cearania heterophylla gen. nov et sp nov., a fossil gymnosperm with affinities to the Gnetales from the Early Cretaceous of northern Gondwana

Vegetative and fertile shoots of a shrub-like seed plant from the late Aptian Crato Formation of Brazil are described as Cearania heterophylla Kunzmann, Mohr and Bernardes-de-Oliveira, gen. nov. et sp. nov. Anatomical details of the axes, epidermal features and separate ovulate and pollen producing organs indicate the gymnospermous nature of this plant. The vascular tissue of the axes includes tracheids with bordered pits and fiber tracheids. Vegetative shoots comprising at least three branching orders bear opposite-decussately arranged ovate to lanceolate, dorsiventrally flattened, parallelodromous, rather thick leaves that vary tremendously in size. The amphistomatic leaves bear (brachy-)paracytic stomatal complexes arranged in simple longitudinal files. The ovulate structure is interpreted as a terminally attached single globular ovule/seed surrounded by at least five to six lanceolate bracts. A terminally attached pollen-cone like structure grows on a lateral leafy shoot. The unusual character combination may indicate that the fossils belong to a hitherto unknown group with affinities to ephedroid Gnetales. Sterile shoots formerly often described as Podozamites, Nageiopsis or Lilites that are at least partly congeneric with C. heterophylla Kunzmann, Mohr and Bernardes-de-Oliveira, gen. nov. et sp. nov. had a wide geographic distribution during the Early Cretaceous. (C) 2009 Elsevier B.V. All rights reserved.

Error, Bias, and Long-Branch Attraction in Data for Two Chloroplast Photosystem Genes in Seed Plants

Sequences of two chloroplast photosystem genes, psaA and psbB, together comprising about 3,500 bp, were obtained for all five major groups of extant seed plants and several outgroups among other vascular plants. Strongly supported, but significantly conflicting, phylogenetic signals were obtained in parsimony analyses from partitions of the data into first and second codon positions versus third positions. In the former, both genes agreed on a monophyletic gymnosperms, with Gnetales closely related to certain conifers. In the latter, Gnetales are inferred to be the sister group of all other seed plants, with gymnosperms paraphyletic. None of the data supported the modern ‘‘anthophyte hypothesis,’’ which places Gnetales as the sister group of flowering plants. A series of simulation studies were undertaken to examine the error rate for parsimony inference. Three kinds of errors were examined: random error, systematic bias (both properties of finite data sets), and statistical inconsistency owing to long-branch attraction (an asymptotic property). Parsimony reconstructions were extremely biased for third-position data for psbB. Regardless of the true underlying tree, a tree in which Gnetales are sister to all other seed plants was likely to be reconstructed for these data. None of the combinations of genes or partitions permits the anthophyte tree to be reconstructed with high probability. Simulations of progressively larger data sets indicate the existence of long-branch attraction (statistical inconsistency) for third-position psbB data if either the anthophyte tree or the gymnosperm tree is correct. This is also true for the anthophyte tree using either psaA third positions or psbB first and second positions. A factor contributing to bias and inconsistency is extremely short branches at the base of the seed plant radiation, coupled with extremely high rates in Gnetales and nonseed plant outgroups. M. J. Sanderson,* M. F. Wojciechowski,*† J.-M. Hu,* T. Sher Khan,* and S. G. Brady

买麻藤目的化石记录及其与 被子植物起源关系的探讨

二十世纪初，前人在中国的东北地区发现了大量保存完好的动物化石，其中以狼鳍鱼最具代表性，科学家将在该地区发现的化石生物群命名为“热河生物群”。“热河”这一名称得名于该化石群的经典产地，即当时的热河省东部地区。建国后，热河省被撤消，其西部划归河北省，东部划归辽宁省。原热河生物群的经典产地因此落在了今天辽宁省的西部，即辽西地区。但是，“热河生物群”这一在地质古生物学界具有深刻影响的名称仍然保留着，而且近几年来在该地区又有大量的鸟类以及恐龙的化石被发现。该地区现在已经成为世界级的古生物宝库。通过不同的方法对该地区进行时代测定的结果认为该地区的时代为白垩纪早期。 与发现众多、研究深入的动物群相比，在该地区开展的植物学研究起步较晚，但在最近几年取得了很大的进展。目前为止，已经发现的植物类群就有苔藓、蕨类、银杏、苏铁、松柏类和被子植物。其中，银杏、苏铁、松柏类尤为丰富。理论上，被子植物也正是从该生物群所代表的白垩纪早期开始出现并逐步走向繁荣的。近年来在热河生物群中就有不少关于被子植物早期类群的报道，如古果属A rchaefructus和里海果属Hyrcantha等。传统上认为，买麻藤目植物与被子植物起源的关系非常密切，但是最近的分子系统学研究却将该类群推离了被子植物，而作为裸子植物高等类群松柏类的姐妹群对待。但是，在热河生物群中的一些新买麻藤目植物的化石标本与被子植物早期类型化石标本的发现却提供了新的思路或证据。如近年来，我国学者已经在该生物群中报道的麻黄科下的2属4种。这些发现的类群都与买麻藤植物的基部类群麻黄属密切相关。这种新发现带来的证据或许可以为被子植物起源这一世界难题的解决提供新的思路。 在对前人关于买麻藤目植物化石标本的研究进行整理的时候，我们发现我们的部分化石与前人发表的一个种Ephedrites chenii在标本的形态学特征方面完全相同。但是在对该类群及其所在的属与麻黄的现代类群作对比研究后发现，该种植物的繁殖器官的特征完全符合麻黄属的特征，因此将该类群转移到麻黄属中作为新组合对待。另外，根据前人对该种在种加词的命名上的修改，我们将该新组合命名为Ephedra cheniae。 在调查该生物群中的买麻藤植物时，我们发现在部分化石类群中出现了前人没有记载过的新性状，比如在麻黄科类植物中发现了互生的分枝方式，并据此命名一个新的单种属Alloephedra xingxuei。为了探讨互生分枝这一性状对于麻黄科的分类意义，我们调查了国内外不少标本馆中的标本，并在野外做了取样统计。结果发现，在不少现代麻黄的枝条上都出现了不同程度的分枝发育不均衡，表现为在同一节上对生的两个分枝中一侧分枝能够正常发育而另一侧发育迟缓甚至不发育，这种不均衡的发育造成了在现代麻黄中出现了类似互生的分枝状况。在整理前人对麻黄分枝方式研究的基础上，结合我们对野外类群形态学性状的调查，我们认为由于在同一节上对生的两个分枝中一侧延迟或不发育而引起的类似互生的性状是较为广泛存在的，它不应该成为属一级的分类依据，而只能作为种一级的分类依据对待。因此我们将AHoephedra xingxuei转移到麻黄属中作为新组合Ephedra xingxuei处理。 除此之外，我们也发现了不少其他的买麻藤目化石标本，在对买麻藤目以往的化石记录了解以及对这些标本形态学性状把握的基础上，我们将这些新发现的化石标本放置在麻黄科麻黄属中作为新种对待，并根据其叶片以及苞片的特征分别将他们命名为披针叶麻黄（Ephedra lanceoleta）、裂叶麻黄（Ephedra divisa），卵叶麻黄（Ephedra ovata）以及双苞麻黄（Ephedra bibracta）。 与此同时，我们也发现了保存完好的被子植物的化石标本。该标本中具有5个离生的心皮、分枝方式兼具侧生和二叉分枝两种、并具有多裂的叶片。在辽西地区同时代发现的被子植物早期类型共有两个，分别是Archaefructus和Hyrcantha。其中前者具有许多个离生的螺旋状排列在可育枝的顶端1cm内的子房／心皮，而后者只有2-4个子房／心皮。在对我们新发现的化石标本与这两个被子植物的早期类型在叶片形态、分枝式样、果实大小、果实构成、果实排布、心 皮数目等形态学特征对比的基础上，我们认为新的化石标本描述了一个被子植物早期类群的新形态，并根据其具有五个心皮以及多裂的叶片的特征，命名为裂叶文采果Wentsaia divisa gen.＆sp. nov。 由于我们有幸能在辽西同时发现了买麻藤植物的化石标本和被子植物早期类型的标本，这就给了我们一个讨论二者在起源关系方面的机会。在对二者的营养器官特征、繁殖器官结构组成、繁殖器官性别构成以及对二者生活环境理解进行对比的基础上，我们认为，买麻藤目植物的早期类型与被子植物的早期类型之间存在相关性。但是就目前的证据而言，尚无法推测二者之间是否存在性状上的演化关系，而该问题的解决需要更多的化石证据的积累。

A putative gnetalean gymnosperm Cariria orbiculiconiformis gen. nov et spec. nov from the Early Cretaceous of northern Gondwana

Cariria orbiculiconiformis gen. nov. et spec. nov., a gymnosperm with gnetoid characters is described from the upper Aptian Crato Formation of the Araripe Basin in northeastern Brazil. Gross-morphology and anatomical details have been studied and characters have been discussed in respect to various seed plants. Several of these characters fit best with those of Gnetales and their putative fossil allies. However, the fossil plant cannot be assigned to any known extinct or extant group of seed plants in their current circumscription. Stem gross-morphology, xylotomical characters and epidermal features indicate a gnetophytic relationship, whereas characters of the reproductive organs are rather distinct from those found in extant taxa. The reproductive unit of the new taxon represents a triple organ consisting of two dichasial ovulate structures and one median pollen-producing structure containing smooth, monosulcate, boat-shaped pollen in-situ. Each ovulate structure consists of two distinct pairs of bracts, a sterile one at the base and a fertile one forming a terminal orbicular capsule. Stiff processes found in the apex of the ovulate structure may represent micropylar tubes of seeds, as seen in the Bennettitales-Erdtmanithecales-Gnetales group. C orbiculiconiformis gen. nov. et spec. nov. was ans herbaceous or semi-shrub-like plant that may have been adapted to the r-strategy in a stressful environment. (C) 2011 Elsevier B.V. All rights reserved.