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The Importance of Fossils in Elucidating the Phylogeny And Macroevolution of Foraminifera Susan L. Richardson, Ph.D Introduction Molecular systematics has revolutionized our understanding of where Foraminifera fit on the Tree of Life; however, molecular phylogenies depict crown clades only, and thus cannot evaluate the evolutionary relationships of extinct taxa to modern species. This study summarizes the results of a cladistic analysis of 121 fossil and Recent foraminiferal taxa, including representative species from Paleozoic Lagenina, Paleozoic Fusulinacea, and Mesozoic Involutinina. Methods A cladistic analysis of a data matrix comprised of 121 foraminferal taxa and 170 characters (both binary and multistate) discrete morphological characters was analyzed in TNT (Tree analysis using New Technology) (Goloboff et al. 2003) using the New Technology search option. The search returned retained two trees with a best score of 688 after 15,014,536 rearrangements. Results The resulting evolutionary relationships of the fossil foraminiferal groups mentioned above as revealed by this analysis are as follows: 1. Fusulinacea (yellow) is a monophyletic clade that branches within the more inclusive clade Globothalamea (Node #2) as the sister group to Textularida (orange). 2. Involutinina (pink) is a monophyletic clade that branches as the sister group to the clade Tubothalamea (purple & Node 1). 3. “Lagenina” is a polyphyletic group comprised of: a. A monophyletic clade of two-chambered undivided tubular Paleozoic lagenids (red) that is more closely related to the clade Tubothalamea (purple). b. A monophyletic clade of multi-chambered septate Paleozoic “lagenids” (green) that branches as the sister group to the modern multi-chambered lagenids (aqua). 4. The clade Globothalamea (Node #2) is the sister group to a more inclusive clade comprised of the multi-chambered Paleozoic “lagenids” (green) + Recent “lagenids” (aqua). Results Phylogenetic Nomenclature Phylogenetic nomenclature defines taxon names in reference to published phylogenies. See the PhyloCode Cantino and de Queiroz (2010) for additional information on phylogenetic nomenclature. Examples of definitions for definiton of foraminiferal clade names are given below. Foraminifera is the clade originating with the most recent common ancestor of Marginopora vertebralis Quoy and Gaimard in de Blainville 1830, and all extant organisms that share a more recent common ancestor with Marginopora vertebralis Quoy and Gaimard in de Blainville 1830, than with Gromia oviformis Dujardin 1835 or Filoreta marina Bass and Cavalier-Smith 2009 or Haplosporidium nelsoni (Haskin, Stauber & Mackin 1966) or Paradinium poucheti Chatton 1910 or Plasmodiophora brassicae Woronin 1878 or Eucyrtidium hexastichum (Haeckel 1887) (Richardson and Lipps In review). Globothalamea can be defined as: the clade originating with the most recent common ancestor of Neogloboquadrina dutertrei (d’Orbigny, 1839) and Reophax sp. (GenBank accession: HE998675) as originally depicted on the reference phylogeny of Foraminifera in Fig. 1 of Pawlowski et al. (2013). Tubothalamea can be defined as: the clade originating with the most recent common ancestor of Marginopora vertebralis Quoy and Gaimard, 1830, and Ammodiscus sp. (GenBank accession: HE998691), as originally depicted on the reference phylogeny of Foraminifera in Fig. 1 of Pawlowski et al. (2013). References Cited Cantino, P. D., and K. de Queiroz. 2010. International code of phylogenetic nomenclature. Version 4c. http://www.ohio.edu/phylocode/index.html . Goloboff, P., J. S. Farris, and K. C. Nixon. 2003. TNT (Tree analysis using New Technology). Published by the authors, Tucamán, Argentina. Pawlowski, J., M. Holzmann, and J. Tyszka. 2013. New supraordinal classification of Foraminifera: Molecules meet morphology. Marine Micropaleontology 100(1-10). Richardson, S. L., and J. H. Lipps. In review. Foraminifera d’Orbigny 1826 [S. L. Richardson & J. H. Lipps], converted clade name. . In K. de Queiroz, and J. Gauthier, eds. PhyloCode Companion Volume. University of California Press, Berkeley, CA. tnt 2 Gromia oviformis Allogromia laticollaris Saccammina sphaerica Hyperammina sp Earlandia elegans Spirillina vivipara Ammodiscus sp Tournayella sp Brunsiina sp Cornuspira planorbis Spiroloculina sp Silicoloculina sp Quinqueloculina seminula Triloculina trigonula Planispirinella exigua Dendritina sp Peneroplis planatus Archaias angulatus Parasorites orbitolitoides Sorite orbiculus Angulodiscus communis Papillaconus sp Coscinoconus sp Semiinvoluta sp Trocholina sp Frentzenella perforata Trocholinopsis porosa Planispirillina papillosa Kristantollmanella sp Licispirella sp Wallowaconus sp Lamelliconus sp Coronoserra sp Coronipora sp Piriniella sp Tezaquina clivuli Amphoratheca iniqua Amphoratheca sp Syzrania amazonica Syzrania groves Syzrania pulchra Reophax scorpiurus Hormosina globularis Ammobaculites agglutinans Haplophragmoides canariensis Patellina corrugata Palaeobigenerina geyeri Textularia candeiana Spirotextularia floridana Sahulia kerimbaensis Siphonifereroides balearicus Nanicella gallowayi Millerella marblensis Endothyra bowmani Endothyra baileyi Plectogyra plectogyra Ozawainella huecoensis Pseudostaffella needhami Schubertella transitoria Boultonia guadalupensis Paraboultonia splendens Staffella Fusulina mysticensis Fusulina girtyi Parafusulina wordensis Parafusulina kingorum Skinnerina typicalis Polydiexodina capitensis Eopolydiexodina afghanensis Triticites moorei Schwagerina diversiforma Schwagerina nelsoni Turkmenella radiata Assilina ammonoides Heterostegina depressa Heterocyclina tuberculata Pararotalia spinigera Neorotalia calcar Calcarina spengleri Nonionella atlantica Nonion depressulum Elphidiella arctica Haynesina germanica Elphidium albiumbilicatum Elphidium williamsoni Elphidium macellum Elphidium margaritaceum Elphidium crispum Elphidium selseyense Elphidium excavatum Elphidium incertum Nodosinelloides pinardae Protonodosaria rauserae Vervilleina bradyi Colaniella lepida Colaniella parva Colaniella minuta Howchinella woodwardi Marginulina albatrossil Lenticulina iota Laevidentalina frobisherensis Mesodentalina matutina Prodentalina sp Pseudonodosaria subannulata Dentalina flintii Dentalina obliqua Dentalina antennula Nodosaria haliensis Nodosaria subsoluta Glandulina yunnanesis Barnardina thanetana Euglandulina inusitata Glandulina laevigata Tappanella arctica Geinitzina postcarbonica Pachyphloia ovata Multiseptida corallina Aratikhinella cannula Nodosinella digitata Eonodosaria evlanensis Spandelinoides nodosariformes TNT Tree #2 TNT Tree #1
