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植物研究雑誌 J. J. Jpn. Bo t. 73: 73: 26-34 (1 998) 1 Permineralized Permineralized Matoniaceous Fossils from the Cretaceous of Japan ) Harufumi NISHIDA a，Akira YOSHIDA b and Makoto NISHIDA b aLaboratory aLaboratory of Geosciences and Biology ，Faculty of Science and Engineering ， Chuo University ， Kasuga Kasuga 1-13-27 ，Bunkyo-ku ，Tokyo ，112-0003 JAPAN; bpaleontological bpaleontological Laboratory ，Research Institute of Evolutionary Biology ， 2-4-28 2-4-28 Kamiyoga ，Setagaya-ku ，Tokyo ，158-0098 JAPAN (Received (Received on September 1 ，1997) A fossil matoniaceous rhizome ，Tricyclopteris japonocretacea gen. et sp. nov. is described described from the Upper Cretaceous of Hokkaido ，Japan. The rhizome has a tricyclic solenotele solenotele with endarch primary xylem. A matoniaceous petiole or rachis ，Matoniostipes mesozoica mesozoica Nishida (1973) from the Lower Cretaceous of Choshi ， Chiba Prefecture ，is emended and reexamined for comparison. It has loose parenchyma (Seward 1899) near protoxylem protoxylem like the petiole of living Matonia pectinata. Matonia pectinata is also characterized characterized by a small cavity parenchyma (Holder ， 1925: Ogura ， 1972) near protoxylem in in the rhizome ， while it does not occur in Tricyclopteris. (Continued (Continued from Y oshida et al.， Res. Ins t. Evolu t. Bio l. Sci. Rep. 8: 85-94 ， 1996) Introduction on a badl y preserved specimen from the Jurassic The Matoniaceae is a rather small family of of lndia (Sharma and Bohra 1978). leptosporangiate leptosporangiate fern. The family consists of Tidwell and Skog (1992) described two two living genera Matonia (2 spp.) and genera of different ages from Tasmania. Mid- Phanerosorus (1 sp.). Thespeciesarerestricted Mesozoic Tasmanopteris has a polycyclic to to the paleotropical regions and prefer habitat siphonostele and Tertiary Heweria has similar in in forest floor or margin ，though it is well structure. However ，Tidwell and Skog (1992) known from the fossil records that the family state possible Mesozoic origin of Heweria. was cosmopolitan during the Mesozoic. Matoniostipes mesozoica (Nishida 1973) is Fossils that are comparable to the the first record of permineralized matoniaceous Matoniaceae are being found ever since the fossil in the world from the Lower Cretaceous Triassic. Triassic. lmpression or compression fossils of Chiba Prefecture ，J apan. It is emended by such as Phlebopteris Brongniart (1 836) (= reexaminations of type specimen. The fossil Laccopteris Laccopteris Presll838) (the Upper Triassic to which will be described here is a fern rhizome the the Cretaceous) and Matonidium Schenk collected by the first and third authors from the (1871) (1871) (J urassic and Cretaceous) have shown Upper Cretaceous of Yubari City ，Hokkaido. worldwide distribution. Delo ωsorusSk ωog(1988) It has a tricyclic stele characteristic of a i臼s an Ame 口r-lC 咽 matoniaceous rhizome. The rhizome is com- Cretaceous. Cretaceous. These are all impression or com- pared to the early described fossils and living pression foliage. The first record of Matonia ，and is designated here as a new permineralized permineralized matoniaceous rhizome is based genus of the Matoniaceae. 一一 26 一一 February February 1998 Journal of Japanese Botany Vo l. 73 No. 1 27 Materials Materials and Methods consisting of scalariform tracheids and xylem Specimen No. 82301 is a permineralized parenchyma ， lacking cavity parenchyma fern fern rhizome 6 mm long and 4 mm in diameter ， near protoxylem. Each stele surrounded by with with fairly well preserved histology. It is cy- sclerenchyma tissue consisting of thick - walled lindrical lindrical in original shape ， but one-third of it is cells. degenerated degenerated or lost during the diagenesis. The Etymology. The genus name originates from specimen specimen is embedded in a calcareous nodule the tricyclic stele of rhizome. found found from the riverbed of the Yubari River ， Tricyclopteris japonocretacea H. Nishida ， along along downstream from Hakobuchi dam ， A. Y oshida & Nishida ， sp. nov. Ohyubari ， Yubari City ，Hokkaido ，J apan. The Species diagnosis. As that of the genus. nodule nodule containing the fossil also include ma- Etymology. The specific epithet means the rine rine fauna and shows lithological features of fossil occurrence (J apan) and age (Cretaceous). the the nodules derived from the Upper Creta- Deposition. Micropreparations of the holotype ceous ceous Middle or Upper Yezo Group (Turonian- specimen (No. 82301) is housed in the Labo- Santonian) ，which are widely distributed along ratory ofPhylogenetic Botany ， Faculty of Sci- the the Yubari Rive r. A cyatheaceous tree fern ence ，Chiba University ， Japan. petiole ，Thyrsopterorachis (H. Nishida and Description. Rhizome cylindrical ， without Nishida Nishida 1979) has been described from the prominent dorsiventrality ， 4.0 mm in diameter same area. GeologicaI information should be and with tricyclic stele. The outer and middle referred referred to the previous paper (H. Nishida and steles amphiphloic solenostele ，and the inner a Nishida Nishida 1979). Micropreparations were made protostele (Fig. 2). Epidermis eroded and los t. by ordinary peel method (Joy et al. 1956) using Hypodermallayer consists of sclerenchyma of 1.8% hydrochloric acid as an etching reagen t. 2- 3 cells thick; cells rectangular in cross sec- We used micropreparations of Matonia tion ，12.5-25μm in diamete r. Outer cortex 3- pectinata pectinata made by the late Pro f. P. Maheshwari ， 5 cells wide ，composed ofthick-walled paren- India ， for comparison. chyma ， while inner cortex 7-10 cells wide ， Micropreparations Micropreparations of the type specimen of composed of thin- walled ce Il s; ce Il s 65-75μm Matoniostipes Matoniostipes being deposited at Chiba Uni- in diameter (Fig. 2). Each stelar ring sur- versity versity were compared to the fossil rhizome. rounded by a sclerenchyma sheath of 5-7 cells thick. thick. The sclerenchyma cells 25-37.5μmm Results Results diameter(Figs. 3，4 and 6). Endodermis mostly Taxonomic treatment degraded. Diameter of stelar rings 2.0 mm in Class Class Filicopsida the outer ， 0.9-1.0 mm in the middle and 0.2 Order Filices Leptosporangiatae mm in the central protostele. In the external Family Matoniaceae C. Presl (1 847) stelar ring ， the xylem plate slightly undulates Genus Tricyclopteris H. Nishida ，A. in the inner contour (Fig. 2，between arrows). Yoshida & Nishida ， gen. nov. The xylem chiefly endarch or rarely mesarch Type species. T.japonocretacea H. Nishida ， with ca. 15 protoxylem points.. Metaxylem A. A. Yoshida & Nishida sp. nov. (Figs. 2-6). tracheids have scalariform wall thickening. Diagnosis Diagnosis of the genus. Permineralized fern The most part of phloem degenerated. Xylem rhizome with tricyclic stele; outer and middle mass of the central protostele centrarch and stelar stelar rings amphiphloic and with chiefly consists of less than 10 ce Il s (Fig. 4). Cavity endarch endarch or a few mesarch xylem; innermost parenchyma absent near protoxylem (Fig. 5). stele stele ectophloic centrarch protostele; xylem Affinity. the slender elongated rhizome with 28 28 植物研究雑誌第73 巻第1号 平成10 年2月 Fig. Fig. 1. Cross section of rhizome of living Matonia pectinata showing tricyclic solenostele. Each stelar stelar ring is surrounded by sclerenchyma sheath. Inner contour of the outer ring undulate. U pper side side of rhizome to the righ t. Scale bar: 1 mm. Fig. 2. Cross section of rhizome of Tricyclopteris japonocretacea japonocretacea gen. et sp. nov. ，showing tricyclic stele ， sclerenchyma sheath ，and xylem plate folding folding (between two arrowheads). Scale bar: 0.5 mm. Abbreviations. Abbreviations. Ab: abaxial side ，Ad: adaxial side ， ed: endodermis ， oc: outer cortex ， p: parenchyma ， ph: phloem ， px: protoxylem ，sc: sclerenchyma sheath ， vb: vascular bundle ， x: xylem. xylem. February February 1998 Journa1 of Japanese Botany Vo l. 73 No. 1 29 Figs.3-6. Figs.3-6. Tricyclopterisjaponomesozoica gen. et sp. nov. Fig. 3. Part of outer stelarring ， showing amphiphloic amphiphloic concentric bundle with endarch xylem and surrounding sclerenchyma sheath. Scale Scale bar: 100μm. Fig. 4. Stelar rings enlarged. Scale bar: 100μm. Fig. 5. Outer stelar ring enlarged ， showing absence of cavity parenchyma near protoxylem. Scale bar: 50μm. Fig. 6. Part of of middle stelar ring enlarged. Scale bar: 50μm. 30 植物研究雑誌第73 巻第1号 平成10 年2月 tricyclic tricyclic stele structurally characterizes some parts (Fig. 2，between arrowheads). Tricyclopteris. Tricyclopteris. The amphiphloic concentric The slender rhizome of Tricyclopteris re- bundles bundles and xy lem maturation ，which is chiefl y sembles that of Phanerosorus which is 3 mm endarch endarch or rarely mesarch ， are also character- in diameter ， while Matonia rhizomes range istic istic features. The stelar configuration and from 9 to 10 mm in diamete r. Howeve r. the presence of endodermis attribute the stele of Phanerosorus is dicyclic. Tricyclopteris Tricyclopteris to leptosporangiate ferns. In the Matonia rhizome is characteristic in having living living leptosporangiate ferns ， a polycyclic stele cavity parenchyma near protoxylem (Holder occurs occurs inAcrostichum ，Dennstaedtia ，Matonia ， 1925) (Figs. 7 and 8). Such cavity parenchyma Pteris ，Saccoloma ，and Thyrsopteris (Bower is absent in Tricyclopteris (Fig. 5). 1918 ，1926 ， 1928; Ogura 1972). Acrostichum ， Sharma and Bohra (1 978) reported a Saccoloma and Thyrsopteris differ from Matonia-like silicified rhizome with tricyclic Tricyclopteris Tricyclopteris in having nests of medullary stele
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    Bomfleur et al. BMC Evolutionary Biology (2015) 15:126 DOI 10.1186/s12862-015-0400-7 RESEARCH ARTICLE Open Access Osmunda pulchella sp. nov. from the Jurassic of Sweden—reconciling molecular and fossil evidence in the phylogeny of modern royal ferns (Osmundaceae) Benjamin Bomfleur1*, Guido W. Grimm1,2 and Stephen McLoughlin1 Abstract Background: The classification of royal ferns (Osmundaceae) has long remained controversial. Recent molecular phylogenies indicate that Osmunda is paraphyletic and needs to be separated into Osmundastrum and Osmunda s.str. Here, however, we describe an exquisitely preserved Jurassic Osmunda rhizome (O. pulchella sp. nov.) that combines diagnostic features of both Osmundastrum and Osmunda, calling molecular evidence for paraphyly into question. We assembled a new morphological matrix based on rhizome anatomy, and used network analyses to establish phylogenetic relationships between fossil and extant members of modern Osmundaceae. We re-analysed the original molecular data to evaluate root-placement support. Finally, we integrated morphological and molecular data-sets using the evolutionary placement algorithm. Results: Osmunda pulchella and five additional Jurassic rhizome species show anatomical character suites intermediate between Osmundastrum and Osmunda. Molecular evidence for paraphyly is ambiguous: a previously unrecognized signal from spacer sequences favours an alternative root placement that would resolve Osmunda s.l. as monophyletic. Our evolutionary placement analysis identifies fossil species as probable ancestral members of modern genera and subgenera, which accords with recent evidence from Bayesian dating. Conclusions: Osmunda pulchella is likely a precursor of the Osmundastrum lineage. The recently proposed root placement in Osmundaceae—based solely on molecular data—stems from possibly misinformative outgroup signals in rbcL and atpA genes.
  • Systematics and Biogeography of the Clusioid Clade (Malpighiales) Brad R

    Systematics and Biogeography of the Clusioid Clade (Malpighiales) Brad R

    Eastern Kentucky University Encompass Biological Sciences Faculty and Staff Research Biological Sciences January 2011 Systematics and Biogeography of the Clusioid Clade (Malpighiales) Brad R. Ruhfel Eastern Kentucky University, [email protected] Follow this and additional works at: http://encompass.eku.edu/bio_fsresearch Part of the Plant Biology Commons Recommended Citation Ruhfel, Brad R., "Systematics and Biogeography of the Clusioid Clade (Malpighiales)" (2011). Biological Sciences Faculty and Staff Research. Paper 3. http://encompass.eku.edu/bio_fsresearch/3 This is brought to you for free and open access by the Biological Sciences at Encompass. It has been accepted for inclusion in Biological Sciences Faculty and Staff Research by an authorized administrator of Encompass. For more information, please contact [email protected]. HARVARD UNIVERSITY Graduate School of Arts and Sciences DISSERTATION ACCEPTANCE CERTIFICATE The undersigned, appointed by the Department of Organismic and Evolutionary Biology have examined a dissertation entitled Systematics and biogeography of the clusioid clade (Malpighiales) presented by Brad R. Ruhfel candidate for the degree of Doctor of Philosophy and hereby certify that it is worthy of acceptance. Signature Typed name: Prof. Charles C. Davis Signature ( ^^^M^ *-^£<& Typed name: Profy^ndrew I^4*ooll Signature / / l^'^ i •*" Typed name: Signature Typed name Signature ^ft/V ^VC^L • Typed name: Prof. Peter Sfe^cnS* Date: 29 April 2011 Systematics and biogeography of the clusioid clade (Malpighiales) A dissertation presented by Brad R. Ruhfel to The Department of Organismic and Evolutionary Biology in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the subject of Biology Harvard University Cambridge, Massachusetts May 2011 UMI Number: 3462126 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted.