Redmond N E, Morrow C C, Thacker R W, Diaz M C, Boury-Esnault N, Cárdenas P, Hajdu E, Lôbo-Hajdu G, Picton B E, Pomponi S A, Kayal E, Collins A G
*Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA; School of Biological Sciences, MBC, 97 Lisburn Road, Queen's University, Belfast BT9 7BL, UK; Department of Biology, University of Alabama at Birmingham, 1300 University Boulevard, Birmingham, AL 35294-1170, USA; Museo Marino de Margarita, Boulevard de Boca Del Rio, Boca del Rio, Nueva Esparta, Venezuela; IMBE-UMR7263 CNRS, Université d'Aix-Marseille, Station marine d'Endoume, rue de la Batterie des Lions, 13007-Marseille, France; Department of Systematic Biology, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, 752 36 Uppsala, Sweden; Departamento de Invertebrados, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, Brazil; **Departamento de Genética, IBRAG, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, RJ, Brazil; Department of Natural Sciences, National Museums of Northern Ireland, 153 Bangor Road, Holywood BT18 0EU, Northern Ireland, UK; Harbor Branch Oceanographic Institute-Florida Atlantic University, 5600 U.S. 1 North, Ft Pierce, FL 34946, USA; Laboratory of Analytical Biology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA; National Systematics Laboratory of NOAA Fisheries Service and Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA.
Integr Comp Biol. 2013 Sep;53(3):388-415. doi: 10.1093/icb/ict078. Epub 2013 Jun 22.
The most diverse and species-rich class of the phylum Porifera is Demospongiae. In recent years, the systematics of this clade, which contains more than 7000 species, has developed rapidly in light of new studies combining molecular and morphological observations. We add more than 500 new, nearly complete 18S sequences (an increase of more than 200%) in an attempt to further enhance understanding of the phylogeny of Demospongiae. Our study specifically targets representation of type species and genera that have never been sampled for any molecular data in an effort to accelerate progress in classifying this diverse lineage. Our analyses recover four highly supported subclasses of Demospongiae: Keratosa, Myxospongiae, Haploscleromorpha, and Heteroscleromorpha. Within Keratosa, neither Dendroceratida, nor its two families, Darwinellidae and Dictyodendrillidae, are monophyletic and Dictyoceratida is divided into two lineages, one predominantly composed of Dysideidae and the second containing the remaining families (Irciniidae, Spongiidae, Thorectidae, and Verticillitidae). Within Myxospongiae, we find Chondrosida to be paraphyletic with respect to the Verongida. We amend the latter to include species of the genus Chondrosia and erect a new order Chondrillida to contain remaining taxa from Chondrosida, which we now discard. Even with increased taxon sampling of Haploscleromorpha, our analyses are consistent with previous studies; however, Haliclona species are interspersed in even more clades. Haploscleromorpha contains five highly supported clades, each more diverse than previously recognized, and current families are mostly polyphyletic. In addition, we reassign Janulum spinispiculum to Haploscleromorpha and resurrect Reniera filholi as Janulum filholi comb. nov. Within the large clade Heteroscleromorpha, we confirmed 12 recently identified clades based on alternative data, as well as a sister-group relationship between the freshwater Spongillida and the family Vetulinidae. We transfer Stylissa flabelliformis to the genus Scopalina within the family Scopalinidae, which is of uncertain position. Our analyses uncover a large, strongly supported clade containing all heteroscleromorphs other than Spongillida, Vetulinidae, and Scopalinidae. Within this clade, there is a major division separating Axinellidae, Biemnida, Tetractinellida, Bubaridae, Stelligeridae, Raspailiidae, and some species of Petromica, Topsentia, and Axinyssa from Agelasida, Polymastiidae, Placospongiidae, Clionaidae, Spirastrellidae, Tethyidae, Poecilosclerida, Halichondriidae, Suberitidae, and Trachycladus. Among numerous results: (1) Spirophorina and its family Tetillidae are paraphyletic with respect to a strongly supported Astrophorina within Tetractinellida; (2) Agelasida is the earliest diverging lineage within the second clade listed above; and (3) Merlia and Desmacella appear to be the earliest diverging lineages of Poecilosclerida.
多孔动物门中种类最多、物种最丰富的纲是寻常海绵纲。近年来，鉴于结合分子和形态学观察的新研究，这个包含7000多种物种的进化枝的系统分类学发展迅速。我们增加了500多条新的、近乎完整的18S序列（增加了200%以上），试图进一步加深对寻常海绵纲系统发育的理解。我们的研究特别针对那些从未有过任何分子数据样本的模式种和属的代表性物种，以加速对这个多样谱系进行分类的进展。我们的分析确定了寻常海绵纲的四个得到高度支持的亚纲：角质海绵亚纲、粘海绵亚纲、单轴海绵亚纲和异轴海绵亚纲。在角质海绵亚纲中，树角海绵目及其两个科，达尔文海绵科和网枝海绵科，都不是单系的，而网角海绵目被分为两个谱系，一个主要由皮海绵科组成，另一个包含其余的科（艾氏海绵科、海绵科、厚皮海绵科和旋枝海绵科）。在粘海绵亚纲中，我们发现软骨海绵目相对于软海绵目是并系的。我们对后者进行修正，将软骨海绵属的物种包括在内，并建立一个新的软骨海绵目来包含软骨海绵目中其余的分类单元，我们现在摒弃软骨海绵目。即使增加了单轴海绵亚纲的分类单元样本，我们的分析结果仍与之前的研究一致；然而，哈氏海绵属的物种散布在更多的进化枝中。单轴海绵亚纲包含五个得到高度支持的进化枝，每个进化枝都比之前认为的更加多样，并且目前的科大多是多系的。此外，我们将刺尖简骨海绵重新归类到单轴海绵亚纲，并将菲尔霍氏芮氏海绵复活为简氏菲尔霍氏海绵，新组合。在庞大的异轴海绵亚纲进化枝中，我们基于其他数据确认了最近确定的12个进化枝，以及淡水海绵目和维图林科之间的姐妹群关系。我们将扇形斯氏海绵转移到斯氏海绵科内的斯氏海绵属，斯氏海绵科的分类地位不确定。我们的分析发现了一个大型的、得到强烈支持的进化枝，它包含除海绵目、维图林科和斯氏海绵科之外的所有异轴海绵亚纲物种。在这个进化枝中，有一个主要的分类，将斧海绵科、双针海绵目、四放海绵目、布氏海绵科、星骨海绵科、拉斯派利海绵科以及一些彼得罗米卡属、托普桑海绵属和轴海绵属的物种与凝胶海绵目、多鞭海绵科、板海绵科、穿贝海绵科、螺旋星海绵科、泰斯海绵科、巧板海绵目、海松海绵科、软海绵科和粗枝海绵科分开。在众多结果中：（1）螺旋海绵亚目及其泰氏海绵科相对于四放海绵目中一个得到强烈支持的星状海绵亚目是并系的；（2）凝胶海绵目是上述第二个进化枝中最早分化的谱系；（3）梅氏海绵属和盘皮海绵属似乎是巧板海绵目中最早分化的谱系。
