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Culcita Remberi Sp. Nov., an Understory Fern of Cyatheales from the Miocene of Northern Idaho

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Culcita Remberi Sp. Nov., an Understory Fern of Cyatheales from the Miocene of Northern Idaho Int. J. Plant Sci. 179(8):635–639. 2018. q 2018 by The University of Chicago. All rights reserved. 1058-5893/2018/17908-0004$15.00 DOI: 10.1086/698938 CULCITA REMBERI SP. NOV., AN UNDERSTORY FERN OF CYATHEALES FROM THE MIOCENE OF NORTHERN IDAHO Jerald B. Pinson,1,* Steven R. Manchester,† and Emily B. Sessa* *Department of Biology, University of Florida, Box 118525, Gainesville, Florida 32611, USA; and †Florida Museum of Natural History, University of Florida, Box 118525, Gainesville, Florida 32611, USA Editor: Michael T. Dunn Premise of research. Excavations at the Clarkia fossil beds in northern Idaho have recovered numerous ex- ceptionally well-preserved plant fossils over the past 45 years. We report on a compression fossil of a fertile frond from a tree fern found at the Emerald Creek locality. Methodology. Intact leaf material was liberated from the substrate by wetting with hydrofluoric acid fol- lowed by submersion in a water bath. The lamina was preserved between sheets of acetate and photographed, and fertile pinnules with sporangia were mounted on aluminum stubs and imaged with scanning electron micros- copy. Spores were released from sporangia after incubation in Schulz’s solution and treatment with KOH. Pivotal results. The specimen is similar to members of several extant fern lineages, including Hymenophyl- laceae, Dennstaedtiaceae, and the tree fern order Cyatheales. It shares with these groups the possession of oblong, marginal sori, although the sori of the specimen are most similar to the cup-shaped sori of Dennstaedtiaceae and the bivalvate sori of Culcitaceae. The fossil also has sporangia, each with a complete, uninterrupted, oblique annulus, which is strongly associated with Culcitaceae. Conclusions. On the basis of its overall lamina morphology and details of the sori and especially the sporan- gia, we conclude that this specimen belongs to an extinct species of the monogeneric tree fern family Culcitaceae. The occurrence of a tree fern in the temperate plant community of the Miocene Clarkia locality is unexpected but not incompatible with the habitat preferences of this genus, as the two extant species of Culcita are found in mon- tane altitudes and temperate forests elsewhere in the world. Keywords: fossil, Miocene, Polypodiopsida, sporangia, tree fern. Introduction suggests that temperatures in Idaho were warmer during the mid- Miocene than they are today (Smiley and Rember 1981). The Clarkia fossil beds in northern Idaho have been the source This study reports on the discovery of a compression fossil of of exceptionally well-preserved impression and compression fos- a fertile frond from a tree fern found at the Emerald Creek local- sils since their discovery in 1972 (Smiley and Rember 1981). De- ity near Clarkia, Idaho. While ferns are nowhere near as ubiq- position occurred during the middle Miocene, when a volcanic uitous as spermatophytes at the site, fossilized remains of Equi- eruption dammed the adjacent St. Maries River at approximately setum, Osmunda, and Polypodium have been reported (Smiley 16–15.4 Ma (Ladderud et al. 2015). Many of the leaf fossils re- et al. 1975). Still, the recovery of only one fossilized tree fern tain their original pigmentation until exposed to air, after which fragment, compared to the thousands of conifer and angio- they rapidly oxidize, and their pristine condition has even al- sperm specimens at Clarkia (Smiley and Rember 1981), under- lowed for isolation of flavonoids (Giannasi and Niklas 1977) scores the rarity of the specimen. Here we present a description and DNA in a few instances (Golenberg 1991; Soltis et al. 1992; of the morphology of the leaf, sporangia, and spores, paying Kim et al. 2004). Studies reconstructing the plant communities particular attention to characters that distinguish it from species present at the time of deposition lead to the conclusion that the with similar morphology and allow us to hypothesize its closest local flora was a temperate deciduous forest consisting mainly living relatives. of conifers (Kvaček and Rember 2000) and angiosperms (Smiley and Rember 1985), taxonomically similar to forest communities Material and Methods present today in both eastern Asia and the southeastern United States (Smiley et al. 1975; Smiley and Rember 1981, 1985). This Material was obtained from the Emerald Creek locality known as University of Idaho P-37 (UF locality 18630, 47701.9720N 1 Author for correspondence; email: jbp4166@ufl.edu. 116720.3070W) in an excavation adjacent to the Rember resi- Manuscript received January 2018; revised manuscript received May 2018; dence, Benewah County, Idaho. The intact compression fossil electronically published August 3, 2018. was lifted from the sediment by wetting the surface of the lam- 635 This content downloaded from 128.227.126.194 on November 05, 2018 07:56:04 AM All use subject to University of Chicago Press Terms and Conditions (http://www.journals.uchicago.edu/t-and-c). 636 INTERNATIONAL JOURNAL OF PLANT SCIENCES ina with concentrated hydrofluoric acid for a few minutes, then clude members with similar marginal, oblong sori, taxa in the immersing the specimen in a water bath and manipulating the Hymenophyllaceae have extremely thin laminar tissue that is loosened lamina onto a piece of plastic window screening, only one or a few cell layers thick and indusia that are generally which was carefully removed by hand from the water bath and more circular than elongated (Tryon and Tryon 1982), neither inverted onto a piece of acetate previously trimmed to an appro- of which applies to the fossil. Of the two remaining groups, priate size. Drops of glycerine were then added and another sheet Dennstaedtiaceae and the tree ferns, the fossil’s sori are most of acetate applied to archive the lamina as a glycerine fossil sand- similar to both Dennstaedtiaceae and Culcitaceae. Members wich on acetate. Portions of pinnules with sori were detached of Dennstaedtiaceae have cup-shaped sori that are similar in ap- and mounted on aluminum stubs for scanning electron micros- pearance to the bivalvate sori of Culcitaceae (Tryon and Tryon copy (SEM). These were coated with palladium with a Denton 1982). However, because of the oxidation of the sori in the fos- DeskV sputter-coater and imaged with a Hitachi SU5000 Schottky sil specimen, we were unable to determine which of these two Field-Emission SEM. Transmitted-light microscopy of sporan- the fossil possesses. gia and spores was performed with a Nikon Eclipse microscope. Sporangia in Dennstaedtiaceae have a vertical to slightly oblique To free the spores from the surrounding indusia and sporan- annulus interrupted at the stalk (Tryon and Tryon 1982). The gia, maceration of tissue was first attempted with KOH, to no fossil, however, displays sporangia with a complete, uninterrupted, effect. We then incubated the tissue with Schultz’s solution for oblique annulus (fig. 2), characters strongly associated with the 5 min, which, when washed in water and followed by brief treat- family Culcitaceae (Sen 1968; Tryon and Tryon 1982). The ment with 5% KOH, cleared the sporangia and released the unique leaf morphology of the fossil fragment separates it from spores. other potentially closely related (and otherwise morphologically similar) genera of tree ferns; for example, the lamina is at least Results bipinnate and is highly dissected, the latter feature distinguishing it from genera such as Cibotium. On the basis of these characters Taxonomic Treatment considered jointly, we conclude that the fossil is an extinct mem- ber of the monogeneric tree fern family Culcitaceae. Other char- Class—Polypodiopsida acters, such as the grooved rachis and unornamented trilete spores, while not unique to Culcitaceae, further support the consignment Order—Cyatheales of the fossil to this family. Family—Culcitaceae Discussion fl Species—Culcita remberi Pinson, Manchester & Sessa This extinct species belonged to a ora that included many sp. nov. (Figs. 1, 2D, 2F) temperate elements, for example, Metasequoia, Cunninghamia, Calocedrus, Taxus, Platanus, Liquidambar, Cercidiphyllum, Holotype. UF 18630-61360, designated here, deposited in Betula, Ostrya, Paliurus, Prunus, Pterocarya, and Fagus (based the Paleobotanical Collection of the Florida Museum of Nat- on examination of specimens in the University of Florida Paleo- ural History, University of Florida. botanical Collection at the Florida Museum of Natural History). Diagnosis. The fossil is a fragment of a fertile leaf approx- The presence of a tree fern in a such an environment might ini- imately 3 cm in width and 7 cm in length. The fragment is at tially seem suspicious. The two extant Culcita species, however, least twice pinnate and the margins entire. Pinnules are deltoid do not grow exclusively, or even primarily, in warm tropical en- and taper slightly toward the apex of the fragment. The adaxial vironments, nor do they have the prominent vertical trunk asso- surface of the rachis is grooved. Sori are oblong and marginal, ciated with most other extant tree ferns. The fronds are large, attached to a single prominent vein and with an indusium com- reaching up to 3 m in length, but the trunk consists only of a posed partially of laminar tissue. The sori are located exclusively small, prostrate rhizome, relegating these plants to the under- one per pinnule lobe. Sporangia have an oblique, complete an- story (Large and Braggins 2004). These rhizomes have only “ ” nulus composed of approximately 34 cells. Spores are trilete the semblance of trunks
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