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2018-OK-Trochodendraceae.Pdf Int. J. Plant Sci. 179(8):663–676. 2018. q 2018 by The University of Chicago. All rights reserved. 1058-5893/2018/17908-0006$15.00 DOI: 10.1086/699282 NEWLY RECOGNIZED DIVERSITY IN TROCHODENDRACEAE FROM THE EOCENE OF WESTERN NORTH AMERICA Steven R. Manchester,1,* Kathleen B. Pigg,† Zlatko Kvaček,‡ Melanie L. DeVore,§ and Richard M. Dillhoff∥ *Florida Museum of Natural History, Dickinson Hall, 1659 Museum Road, Gainesville, Florida 32611, USA; †School of Life Sciences and Biodiversity Knowledge Integration Center, PO Box 874501, Arizona State University, Tempe, Arizona 85287, USA; ‡Faculty of Science, Institute of Geology and Palaeontology, Charles University, Albertov 6, 128 43 Prague 2, Czech Republic; §Department of Biological and Environmental Sciences, Georgia College and State University, Milledgeville, Georgia 31061, USA; and ∥Evolving Earth Foundation, PO Box 2090, Issaquah, Washington 98027, USA Editor: Patricia G. Gensel The Eocene flora of the Okanogan Highlands in the Pacific Northwest of North America has been recognized previously to include extinct species of both extant genera of the Trochodendraceae. Here, using microcomputed tomography (mCT) scanning to augment traditional methods, we recognize additional diversity, including two new fruit types. Concavistylon wehrii sp. nov. is documented by a fertile twig with attached leaves and an infruc- tescence, allowing for an unusually complete reconstruction of this extinct genus. Concavistyon wehrii infruc- tescences are racemes bearing fruits on short pedicels. Fruits are apically dehiscent capsules with four to six styles. The leaves resemble those of modern Trochodendron in pinnate venation, glandular teeth, and epidermal anatomy but have short petioles. The second new type of infructescence, Pentacentron sternhartae gen. et sp. nov., resembles extant Tetracentron in having small, sessile, apically dehiscent capsules but consistently has five, rather than four, styles. The mCT X-ray imaging demonstrates that fruits of both Concavistylon and Pentacentron differ from those of extant Trochodendraceae in having styles that are concave with stigmas directed inward rather than outward. These fossils, together with previously recognized fossil fruits and leaves of Trochodendron and leaves of Tetra- centron from the same beds, indicate that the Trochodendraceae family was more diverse 50–52 Ma than it is today. Keywords: British Columbia, Concavistylon, fruits, infructescences, leaves, McAbee, Republic, Washington, Tertiary, Tetracentron, Trochodendron. Introduction known also from the Eocene One Mile Creek site in British Co- lumbia. In addition to the extant genera, it has become clear that Eocene shales from the Okanogan Highlands of the Pacific North- some extinct representatives of the family resided in western west in North America document early diversity of the eudicot North America during the Tertiary. For example, Concavistylon family Trochodendraceae. Both of the extant genera, Trochoden- Manchester, Pigg & DeVore was recognized recently on the basis dron Sieb. & Zucc. and Tetracentron Oliver, have been docu- of infructescences from the middle Miocene of Oregon (Man- mented previously in Eocene shales of Washington and British chester et al. 2018). Columbia (Pigg et al. 2001, 2007). This is interesting because Here we recognize two more representatives of Trochoden- these genera are now monotypic and endemic to east Asia. Paleo- draceae from the Okanogan Highlands Eocene flora: Concavis- botanical evidence indicates that both of them were distributed tylon wehrii sp. nov. and Pentacentron sternhartae gen. nov. in western North America through at least the Middle Miocene New microcomputed tomography (mCT) from X-ray scan data, (e.g., Manchester et al. 2018). Reproductive axes found at the showing details hidden within the shale, has provided important Republic, Washington, locality in association with leaves of Tro- characters for the identification of these new species. A fertile chodendron nastae Pigg, Wehr & Ickert-Bond are quite similar to twig bearing both fruits and foliage provides a firm basis for those of fossil Miocene and extant forms, except for their small describing vegetative as well as fertile morphology of Concavis- size (Pigg et al. 2001). A more robust infructescence, Trocho- tylon. In addition, specimens with cuticle preserved allow for the dendron drachukii Pigg, Dillhoff, DeVore, and Wehr was de- description of epidermal characters previously undocumented for scribed from McAbee, British Columbia (Pigg et al. 2007). Tetra- fossil leaves of Trochodendraceae. centron hopkinsii Pigg, Dillhoff, DeVore, and Wehr leaves are In the present study, we describe these additions to the fossil record of Trochodendraceae and summarize all of the taxa now recognized from the Eocene Okanogan Highlands flora for com- 1 Author for correspondence; email: steven@flmnh.ufl.edu. parison with fossil occurrences of this family known elsewhere in Manuscript received April 2018; revised manuscript received May 2018; elec- the Northern Hemisphere. These records indicate greater morpho- tronically published September 7, 2018. logical diversity in this family during the Eocene than today. 663 This content downloaded from 128.227.196.016 on September 27, 2018 00:34:22 AM All use subject to University of Chicago Press Terms and Conditions (http://www.journals.uchicago.edu/t-and-c). 664 INTERNATIONAL JOURNAL OF PLANT SCIENCES Material and Methods uniformly toward infructescence apex. Styles persistent, 4, 5, or 6 (variable); 1.9–2.9 mm long, arising subequatorially, curv- We focused on fossils from two main sites within the early- ing toward the fruit apex; elliptical bulge (interpreted as nec- middle Eocene Okanogan Highlands localities, ca. 50–52 Ma tary) beneath each style. Capsules smooth, with a basal thickening (DeVore and Pigg 2010; Greenwood et al. 2016): the Republic and whorl of perianth scars at junction with pedicel. Dehiscence flora from the town of Republic (Ferry County), northeastern apical, with separation initiating immediately above each style Washington (Wolfe and Wehr 1987; Wehr and Hopkins 1994), and joining apically to form a 4-, 5-, or 6-rayed starlike opening. and the McAbee flora near Cache Creek, British Columbia (Dill- Twig bearing simple leaves in alternate phyllotaxy. Terminal hoff et al. 2005; Gushulak et al. 2016). Specimens are archived bud 1 cm long, 5 mm wide, elongate, pointed apically; axillary at the Burke Museum of Natural History and Culture, University bud 8 mm long, 1.8 mm wide; terminal bud scale scars 5–8, prom- of Washington, Seattle (UWBM); the Stonerose Interpretive Cen- inent, encircling stem. Petiole 8–12 mm long, nonpulvinate, ter, Republic, Washington (SR); Thompson Rivers University, short in relation to lamina length (9%–11%). Lamina obovate, Kamloops, British Columbia (TRU); and Florida Museum of Nat- 10.2–12.5 cm long # 3.1–3.5 cm wide; margin serrate with reg- ural History, Gainesville, Florida (UF). The holotype specimen ularly spaced rounded, glandular, teeth. Venation pinnate with of Concavistylon wehrii did not initially show the physical con- a straight midvein and 8–15 pairs of brochidodromous second- nection of fruits and foliage (fig. 1C), but the connection was ary veins. Intersecondary veins absent. Tertiary and higher-order subsequently exposed by chipping away the overlying sediment veins forming an irregular mesh. Adaxial cuticle smooth, cells po- with fine needles while observing it with a dissecting microscope lygonal, anticlinal walls straight to slightly curved, almost equally (fig. 1B,1D). sized, 30–45 mm in diameter. Abaxial cuticle showing unspe- Cuticles from leaves in the shales of the Boot Hill locality at cialized cells, straight-walled, stomatal complexes rounded-oval, Republic (UWBM locality B4131) and McAbee (UWBM local- 35–40 # 40–50 mm in diameter, stomatal ledges thick, border- ity B5751) were recovered by mechanical removal from the rock ing the broadly oval aperture and not reaching to the poles. Tri- surface, cleaned in hydrofluoric acid, and macerated slightly with chome bases lacking on both abaxial and adaxial cuticles. 5% potassium hydroxide. Fragments were washed in distilled Holotype. UWBM PB 101336. Specimen bearing both infruc- water and placed in glycerine, covered with a coverslip, and pho- tescence and leaves (figs. 1A–1E,2A,4A). From Boot Hill, Re- tographed with Nomarski interference contrast optics. In addi- public, Washington. Burke Museum locality B4131, collected tion, cuticles were examined by epifluorescence with a Zeiss pho- and donated by Dwyane Day in August 1999. tomacroscope using an H546 green excitation filter set (15) with Paratypes. Isolated infructescence from Republic. USNM a wavelength of 546 nm. 537360 from USGS locality 2871, collected by Henry Landis in Leaves and infructescences were photographed by reflected 1901. Previously figured as Trochodendron sp. (fig. 2; fig. 58 light with a Canon Rebel XSI digital single-lens reflex camera in Manchester et al. 2009). with an EF-S60-mm macro lens. Additional images were obtained Type locality. Boot Hill, Republic, Ferry County, Washing- from data sets compiled by mCT. The scanning of impression ton. UWBM locality B4131, Tom Thumb Ash Member of the specimens in shale was performed at the University of Florida Klondike Mountain Formation, latest early Eocene. College of Engineering Nanoscale Research Facility with a GE Additional locality. McAbee, British Columbia, latest early Phoenix VFtomeFxm240 CT scanner, using a Tungsten
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