taxonID	type	description	language	source
4D2487A3EF5C8260FC4EFA856A4BF7DD.taxon	description	Text-fig. 2 d, e M a t e r i a l. USNM PAL 622101, 717666. L o c a l i t y. Dakin and Disbrow Creek. D e s c r i p t i o n. Winged seed round to rectangular with wings forming retuse connection at poles; 0.7 – 1.0 mm tall, 0.6 mm wide; seed body elliptical, in the center of the wings; polar ends of seed body in line with polar ends of wings; seed body 0.3 – 0.7 mm long, 0.2 mm wide. R e m a r k s. This seed bears a superficial resemblance to those of Metasequoia which has also been identified based on foliage from the Kishenehn Formation (Text-fig. 2 f; USNM PAL 776571, 2 g USNM PAL 776572). However, the extant Metasequoia glyptostroboides HU et W. C. CHENG has seeds that range between 4.2 – 5 mm long and 4.8 – 6 mm wide, about 4 – 5 times larger than the Kishenehn specimen (eFloras 2008 – 2022, Liu and Basinger 2009). Fossil Metasequoia seeds from other sites are also larger (e. g., 5 – 6 mm long and 6 – 7 mm wide; Falder et al. 1999). Metasequoia is a major component of the Early Eocene Falkland flora of British Columbia (Smith et al. 2012).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF5C8260FE86F8CA6B2BFCC1.taxon	description	Text-fig. 2 b M a t e r i a l. USNM PAL 622297. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Conifer leaf 19.3 mm long, 3.8 mm wide; petiole stout and rectangular, 0.8 mm long and 0.9 mm wide; leaf base asymmetrical, obtuse and rounded; apex not preserved or in matrix; midvein 0.6 mm; a pair of stomatal bands runs on both sides of the midvein, running parallel to it. Width of the stomatal bands varying between 0.2 – 0.3 mm. R e m a r k s. The affinities of this leaf remain is uncertain. Prominent stomatal bands can be found in leaves of Taxaceae (Amentotaxus PILG., Cephalotaxus SIEBOLD et ZUCC. ex ENDL., Taxus L.), Cupressaceae (Sequoia ENDL.) and Pinaceae (Abies L., Pseudotsuga CARRIÈRE and Tsuga (ENDL.) CARRIÈRE). However, these taxa have stomatal bands as wide or wider than the midvein, whereas our specimen has stomatal bands half of the width of the midvein. The leaf has narrower stomatal bands than those that have been recorded for Amentotaxus PILG. (eFloras 2008 – 2022). Modern Amentotaxus has leaves that are 20 – 115 mm long, 5 – 15 mm wide and with stomatal bands 1 – 2 mm wide (eFloras 2008 – 2022).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF5A8266FEC0F82D6B4EFDAC.taxon	description	M a t e r i a l. USNM PAL 620087, 620274. L o c a l i t y. Park. D e s c r i p t i o n. Scale leaves on axis measuring 8.0 mm long and 1.2 mm wide; branchlets sprawling; distance from facial leaf apices 1.7 mm; leaf height 1.4 mm, leaf width 0.9 – 1.0 mm; apex of facial leaf does not cover juncture of subsequent lateral leaves; leaf apex pointed to slightly rounded 99 – 110 °; leaves are keeled. R e m a r k s. Cupressoideae 2 differs from Cupressoideae 1 and Cupressoideae 4 by having apices that do not cover the subsequent lateral leaf junctions, having obtuse leaf apices and by having keeled leaves. Cupressoideae 2 differs from Cupressoideae 3 by having shorter leaves (1.4 mm vs. 2.1 mm, respectively). This fossil cannot be placed confidently in a modern genus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF5A8266FB9BFDDC6BDBFAC5.taxon	description	M a t e r i a l. USNM PAL 620184. L o c a l i t y. Park. D e s c r i p t i o n. Scaly leaf axis measuring 6.4 mm long, 1.8 mm wide at lateral leaf widest point; leaves 2.1 mm long, 1.1 mm wide; distance between facial leaf apices 2.6 mm; facial leaf apices do not cover subsequent lateral leaf junction; leaf apices rounded and acute 68 °. R e m a r k s. Cupressoideae 3 differs from Cupressoideae 2 by having longer leaves (2.1 mm vs. 1.4 mm), that are nonkeeled and acute apices. Cupressoideae 1 might be equivalent to Cupressoideae 3, with their different appearance due to abscission of leaves in the former. Abscission can occur in genera like Chamaecyparis such that the facial leaves can either cover or not cover the junction of the subsequent lateral leaves (Kotyk et al. 2003). Cupressoideae 3 differs from Cupressoideae 4 by having appressed leaf junctions instead of leaf junctions that flare outward. This fossil cannot be placed confidently in a modern genus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF5A8265FB9BFAE46D9FFD0B.taxon	description	M a t e r i a l. USNM PAL 621863. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Scaly leaf axis measuring 17.3 mm long and 1.0 mm wide; branchlets opposite; leaves decussate, leaves triangular measuring 1.3 mm long with a pointed acute apex measuring between 59.0 and 88.5 °; distance between apices on facial leaves 2.6 – 2.9 mm; apex of facial leaf covers junction of subsequent lateral leaves or may be below junction; lateral leaves separate on facial surface before angling outward. R e m a r k s. Cupressoideae 4 differs from Cupressoideae 1 and 2 by having longer leaves with pointed, rather than rounded apices and having longer internodes. Cupressaceae 4 differs from Cupressaceae 3 by having leaf junctions that flare outward instead of being appressed to the stem. The lateral leaves are not proportionally long enough to match those of Tetraclinis MAST. (Kvaček et al. 2000) and the facial leaves are more prominent than those of Fokienia A. HENRY et H. H. THOMAS (McIver 1992). Mesocyparis MCIVER et BASINGER has opposite instead of alternate branching (McIver et Basinger 1987, Kodrul et al. 2006). Alternate branching cupressaceous conifers including Thuja, Chamaecyparis and Platycladus SPACH have lateral leaf apices that can be adpressed to the side and are indistinguishable from one another (McIver and Basinger 1987, Wu et al. 2014). Thus, this fossil cannot be confidently placed into a genus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF598265FEDBFB006AA2FDB6.taxon	description	M a t e r i a l. USNM PAL 621424. L o c a l i t y. Park. D e s c r i p t i o n. Winged seed linear, 11.7 mm long, 5.3 mm wide at widest; seed body elliptical 3.8 mm long, 2.3 mm wide oriented with long axis parallel to medial edge; seed body on proximal side of seed; seed body contains numerous, round resin vesicles; abmedial margin begins approximately one quarter of the way up seed body extending so widest area of wing is distal; distal edge straight; medial edge convex in distal quarter. R e m a r k s. Winged seeds with resin vesicles and with the wings widest in the distal half belong to Abies (Wolfe and Schorn 1990). Abies pollen is known from the early Eocene McAbee and Quilchena floras of British Columbia (Dillhoff et al. 2005, Lowe et al. 2018, Mathewes et al. 2016) and Yellowstone, Wyoming (Wing 1987). Macrofossils are known from the early Eocene McAbee and Falkland floras of British Columbia (Dillhoff et al. 2005, Lowe et al. 2018, Smith et al. 2012), Republic flora of Washington (Pigg et al. 2011), Green River Formation (Brown 1929) and Oligocene Beaverhead flora of Montana (Wing 1987). Abies is also recognized based on seeds and foliage from the mid-Eocene Thunder Mountain flora of Idaho (Axelrod 1998). Although the identity of some of the specimens that were attributed to Abies deweyensis AXELROD, by Axelrod (1998) has been questioned due to disarticulated parts and a lack of diagnostic features (Erwin and Schorn 2005).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFE08FD446E76FB53.taxon	description	M a t e r i a l. USNM PAL 619981, 776558. L o c a l i t y. Park. D e s c r i p t i o n. Linear winged seed 7.9 – 9.9 mm long, 3.6 – 4.6 mm wide at widest point; seed body elliptical to obovate, on proximal side of seed, 2.7 – 4.0 mm long, 2.1 – 2.4 mm wide, concave notch on proximal medial side; wing begins halfway down seed body along abmedial side and less than one quarter of the way to the seed body’s terminus on the medial side; widest part of wing is just past the center more towards the distal end; medial and abmedial edges join in a smooth curve.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFE08FB6E6AFAFD39.taxon	description	M a t e r i a l. USNM PAL 776565, 776566. L o c a l i t y. Tunnel Creek. D e s c r i p t i o n. Winged seed linear 13.5 – 14.7 mm long, 6.2 – 6.9 mm wide; widest point of wing just distal of the halfway mark; wing encompasses most of seed body; seed body with concave notch on the proximal medial side; seed body obovate to elliptical 3.2 – 4.6 mm long, 1.6 – 2.0 mm wide.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFE08FB6E6AFAFD39.taxon	discussion	Comments on fossil Picea Picea seeds have a distinctive concave notch on the proximal medial side (Wolfe and Schorn 1990). Picea sp. 2 differs from Picea sp. 1 in being larger, with its widest point only slightly past the midpoint of the wing instead of more distal. The wing also encompasses more of the seed body. Pollen of Picea is recognized from the early Eocene Green River Formation (Wodehouse 1933, Nichols 2010), the Chuckanut Formation of Washington (Griggs 1970) and the Quilchena flora of British Columbia (Mathewes et al. 2016). Macrofossils are known from the early Eocene Republic flora (Klondike Mountain Formation of Republic, Washington) (Pigg et al. 2011), McAbee and Falkland floras of British Columbia (Dillhoff et al. 2005, Smith et al. 2012), late Eocene Beaverhead Basin of Montana (DeVore and Pigg 2010) and Florissant Formation of Colorado (MacGinitie 1953), the Oligocene Creede Flora of Colorado (Wolfe and Schorn 1990) and Ruby River Basin of Montana (Becker 1961). Axelrod (1998) initially recognized Picea coloradensis AXELROD, Picea deweyensis AXELROD, and Picea magna AXELROD, from the mid-Eocene Thunder Mountain flora of Idaho. The specimens assigned to these species were subsequently identified as another genus, indeterminate, cf. Picea or Picea sp. by Erwin and Schorn (2005). The presence of Picea suggests microthermal conditions; these seeds may have come from the surrounding mountains around the basin (Dawson and Constenius 2018).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFBA1FD466CE0FB45.taxon	description	M a t e r i a l. USNM PAL 619964. L o c a l i t y. Dishbrow Creek. D e s c r i p t i o n. Structure cylindrical, 4.9 cm long, 1.6 cm wide, composed of helically arranged rectangular outlines; five rectangles wide and 9 rectangles tall, 0.4 cm tall, 0.1 – 0.3 cm wide, lateral extensions 0.2 – 0.3 cm. R e m a r k s. The rectangular outlines in the structure may be obscured leaf scars or cone scale scars. The lateral extensions may be remnants of bracts or scales or taphonomic degradation.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFBA1FB656BDDFA50.taxon	description	M a t e r i a l. USNM PAL 776557. L o c a l i t y. Dishbrow Creek. D e s c r i p t i o n. Ovuliferous cone cylindrical, 4.5 cm long, 2.1 cm wide, scales 1.4 mm long.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFBA1FA506C0AF7F4.taxon	description	M a t e r i a l. USNM PAL 620025. L o c a l i t y. Park. D e s c r i p t i o n. Ovuliferous cone cylindrical, 5.5 cm long, 2.8 mm wide, at least 15 rows of scales, scales 1.3 cm long.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF57826BFBA1FA506C0AF7F4.taxon	discussion	Comments on the three cones of uncertain affinity All three cones appear to lack apophyses, so they are not Pinus. It is unclear whether they represent one or multiple species. bar 1 cm. q: USNM PAL 622423 Pinus subg. Pinus, Pinus sp. S 4 seed. Scale bar 2 mm. r: USNM PAL 625925 Pinus subg. Pinus, Pinus sp. S 5 winged seed. Scale bar 2 mm. s: USNM PAL 625839 Pinus subg. Pinus, Pinus sp. S 5 winged seed. Scale bar 3 mm. t: USNM PAL 622339 Pinus subg. Pinus, Pinus sp. S 6 winged seed. Scale bar 5 mm.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF56826AFB86FA216BECF8CC.taxon	description	Text-fig. 3 p M a t e r i a l. USNM PAL 626064. L o c a l i t y. Deep Ford. D e s c r i p t i o n. Cone scale 23.7 mm long; apophysis large, rhomboid, 7.1 mm tall and, 11.3 mm wide; umbo dorsal. R e m a r k s. Pinus subg. Pinus (also known as Diploxylon, the hard pines or yellow pines) have large, rhomboid apophyses and dorsal umbos.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF56826AFE91FB946E1BFA10.taxon	description	Text-fig. 3 k M a t e r i a l. USNM PAL 776570. L o c a l i t y. Coal Creek. D e s c r i p t i o n. Ovuliferous cone cylindrical; 7.7 cm tall, 4.6 cm wide; scale 1.7 cm long; at least 14 rows of scales, 3 – 4 scales wide, helically arranged; apophysis rhomboidal, 0.8 – 1.5 cm wide, 0.3 – 0.4 cm tall; umbo terminal; spine 0.1 cm long. R e m a r k s. The apophysis places this cone in the genus Pinus and the terminal umbo places this cone in the subgenus Strobus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF56826AFEEEFF066EE3FDC5.taxon	description	M a t e r i a l. USNM PAL 776556. L o c a l i t y. Tunnel Creek. D e s c r i p t i o n. Ovuliferous cone ovate; 5.9 cm tall, 1.2 cm wide; scales helical; apophysis narrow rhomboids 1.2 – 1.4 cm wide and 0.3 cm tall. R e m a r k s. Because the umbo is not clearly preserved, the fossil cannot be identified to subg. Pinus (dorsal umbo) or subg. Strobus (terminal umbo).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF53826FFEEEFB2B6BFBFED8.taxon	description	M a t e r i a l. USNM PAL 620300. L o c a l i t y. Park. D e s c r i p t i o n. Leaves 0.9 mm long, 0.5 – 0.6 mm wide; elliptical; aerenchyma present; pseudoroot 0.4 mm long, 0.2 mm wide with slanted terminus. R e m a r k s. The small size, elliptical leaves, presence of a pseudoroot and lack of venation are characters of the aquatic araceous subfamily, Lemnoideae. The lack of brown pigment cells, which we might expect to preserve in these cellularly preserved compressions (and which do preserve in the Spirodela fossils), rules out its placement in Spirodela or Landoltia and the broad elliptical shape of the fronds contrasts with the more narrow ellipses of Wolffia HORKEL ex SCHLEID and the narrow fronds of Wolfiella (Armstrong 2021). Note the name of the distinctive surficial cells is brown pigment cells. While they are brown in modern plants we would not necessarily be able to determine the color in the fossils. Thus, the leaf shape and lack of brown pigment cells support the placement of this specimen in Lemna. In North America, Lemna fossils are known from the Miocene of Mississippi (McNair et al. 2019). The Kishenehn fossils are the oldest record of Lemna, predating Oligocene occurrences in Eurasia (Dorofeev 1963). Another notable occurrence of Cenozoic Lemnoideae is that of the Limnobiophyllum leaf and root fossils from the Late Cretaceous through Oligocene of western North America (Hoffman 1995, Kvaček 1995, McIver and Basinger 1993, Stockey et al. 1997). Limnobiophyllum differs from our fossils by having veined leaves, branching roots and size (leaves are 1 – 2 cm wide) (Kvaček 1995, Stockey et al. 1997).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF53826FFB86FB0B6BBFF7DD.taxon	description	L o c a l i t i e s. Deep Ford, Disbrow Creek, Park and Spring. M a t e r i a l. USNM PAL 622670, 622675, 623125, 626043, 625757, DMNH EPI. 48225, 48188. D e s c r i p t i o n. Stem 0.2 – 0.7 mm wide; leaves opposite; nodes thick; leaves at least 24.9 mm long, 1.2 – 1.4 mm wide; 2 ° 1 ° 2 ° (where left to right the veins are thin, thick then thin, see fig. 1 a, b in Robil et al. (2021) for comparison) parallel venation type with veins joining to the center vein near leaf apex; apex rounded; margin entire; pigment or glandular cells present; fruits flat, circular drupes, 2.24 – 3.10 mm tall, 1.8 – 2.6 mm wide, 0.6 mm thick; style pointed with persistent stigmatic region; pedicel 0.1 mm long and 0.1 mm wide; four fruits on a raceme with peduncle 9 mm long and 0.2 mm wide. R e m a r k s. The aquatic genus, Potamogeton, is identifiable from leafy stems bearing infructescences. The genus is distinguished by its opposite narrow leaves, and circular, flat fruits with a pointed apex and persistent styles that are borne in racemes (Manchester 2001). Fruits and leaves of Potamogeton have also been recognized from the late Eocene Florissant Formation of Colorado (Manchester 2001) and leaves from the mid-Eocene Thunder Mountain flora of Idaho (Axelrod 1998).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF53826FFEBDFE286E6DFB9B.taxon	description	M a t e r i a l. USNM PAL 624518, 625611, 768304. L o c a l i t i e s. Dakin and Spring. D e s c r i p t i o n. Plants with elliptical fronds 1.1 – 3.9 mm long, 1.5 – 2.5 mm wide; prophylla present, 0.8 – 1.1 mm long, 1.1 – 1.4 mm wide; brown pigment cells present. The presence of large prophylla and brown pigment cells places these plants within Spirodela (Armstrong 2021). Prophylla are absent in Lemna L. and reduced in Landoltia LES et D. J. CRAWFORD (Armstrong 2021). We interpret the brown, circular dots that are concentrated near the prophylla and more dispersed in the distal portion of the leaves as brown pigment cells. Brown pigment cells are absent in Lemna but present in Spirodela (Armstrong 2021). Spirodela was described from the Paleocene of Alberta and Saskatchewan (Dawson 1875, Berry 1935, McIver and Basinger 1993) and the middle Eocene of Wyoming (MacGinitie 1974), but these fossils were subsequently transferred to the extinct araceous genus Limnobiophyllum KRASSILOV (Kvaček 1995, Stockey et al. 1997).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF53826FFC76FEE86A34FB9B.taxon	description	M a t e r i a l. USNM PAL 622956, 626140. L o c a l i t i e s. Deep Ford and Disbrow Creek. D e s c r i p t i o n. Frond triangular, 4.4 – 4.7 mm long, 0.3 – 0.7 mm wide at base, base flat to curved, budding pouch triangular, 0.9 – 1.2 mm tall, 0.3 – 0.7 mm wide, aerenchyma present. R e m a r k s. The slender triangular fronds with aerenchyma and budding pouches are diagnostic of Wolffiella (Armstrong 2021). This specimen is not the dispersal hair of a Typha fruit because it is a multicellular frond. Wolffiella plants can exhibit a variety of morphologies including ovate, “ tongue ” and sickle shaped (Acosta et al. 2021) (Text-fig. 4 g). The size range of these specimens also fits the size range of modern North American Wolffiella fronds (4.4 – 4.7 mm for the fossils, 1 – 9 mm for the modern) (Landolt 2020). Ours would fit in the “ tongue ” category, which does not occur in the other genera of Lemnoideae. These are the first reported Wolffiella fossils. Today there are 10 species of this floating aquatic plant across the Americas and Africa (Landolt 2020). Species in the US are concentrated in the Southwest and Southeast (Landolt 2020).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF52826DFEDCF8276E95F9F8.taxon	description	M a t e r i a l. USNM PAL 729860, 621989, 622997, 623087, 623129, 623144, 717449, 625390, 625721. L o c a l i t i e s. Dakin, Disbrow Creek, Park and Spring. D e s c r i p t i o n. Carpodium 2.5 – 6.8 mm long, 0.6 – 0.9 mm wide; carpodium body obovate with rounded apex, 0.6 – 3.5 mm long, 0.6 – 0.9 mm wide tapering to gynophore; gynophore 1.9 – 3.6 mm long, vein sometimes visible going into carpodium from gynophore; follicle cylindrical fusiform, 1.9 – 2.1 mm long, 0.7 – 0.8 mm wide; coat thin; edges straight,; one pole with small tuft (remnant style) 0.1 mm long, or rounded poles; lateral edges flat; sulcus running between poles; fruit body elongated ellipse, 1.1 – 2.1 mm long, 0.3 – 0.7 mm wide; cylindrical projections at pole (s) gently tapering distally 0.1 – 0.3 mm tall, 0.1 – 0.2 mm wide proximally, 0.1 mm distally; single longitudinal groove; psilate sculpture. R e m a r k s. Although the fossils are similar in shape and size to Wolffiella there is a stipe and vein in the developing Typha fruit that is not present in Wolffiella. The thin wall of the follicle can be observed in modern cattails (Iowa Plants 2012). This fossil bears some resemblance in shape and size to fruits of modern Elodea MICHX. (Hydrocharitaceae), a genus found throughout the Americas today (Kirkbride et al. 2000, Haynes 2020). Hydrilla RICH. is similar in morphology but is smaller (less than 1 mm) and is more sculptured with longitudinal grooves (Kirkbride et al. 2000). Similarly shaped seeds can also be found in Plantago L. (Plantaginaceae), which is globally distributed (except in Antarctica) (Canadian Food Inspection Agency 2017) and in Carex sp. (Bhandari et al. 2010, 2011). Despite similarities with Elodea, Hydrilla and Plantago, the fossil fruits have polar extensions that the aforementioned plants do not have. The lateral edges of Elodea (Kirkbride et al. 2000, Haynes 2020) and Carex L. fruits are more rounded rather than flat, as observed (Bhandari et al. 2010, 2011). Carex scales can be triangular but lack the polar extension and possess foliar lateral extensions near the base, which do not appear in the fossil (Minnesota Wildflowers 2023 a).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF51826DFE9CF9886AF2FEDB.taxon	description	Text-fig. 6 a M a t e r i a l. USNM PAL 625188. L o c a l i t y. Spring. D e s c r i p t i o n. Fruit elliptical, 2.9 mm long (body 2.6 mm long), 1.5 mm wide; basal spine 0.3 mm long, 0.2 mm wide; three short visible lateral spines on the antipodal side 0.1 mm long; small indentations on the surface. R e m a r k s. We interpret this as an eroded specimen retaining only remnants of the original spines. By inference from symmetry, a total of at least six short lateral spines can be inferred. Ceratophyllum muricatum CHAM. subsp. incertum (BERRY) HERENDEEN, LES et DILCHER is known from the early and mid-Eocene of the Green River Formation of Wyoming and the Claiborne Formation of Tennessee (Herendeen et al. 1990). However, C. muricatum subsp. inertum is larger (2.8 – 4.0 mm long and 2.0 – 2.3 mm wide) and has more lateral spines (8 – 11) (Herendeen et al. 1990).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF51826DFBAEFBB06B63F976.taxon	description	M a t e r i a l. USNM PAL 622299, 768140. L o c a l i t i e s. Dakin and Disbrow Creek. D e s c r i p t i o n. Achene obovate 8.8 – 8.9 mm long, 1.7 mm wide with flat base rounding to acute apex and elongate style 3.7 – 4.1 mm long and 0.2 mm wide, single seed obovate to fusiform preserved inside 3.3 – 5.1 mm long and 0.4 – 0.9 mm wide with thickened micropylar end, many dispersal hairs 0.5 – 1.8 mm long arise from base of achene directed apically. R e m a r k s. The presence of dispersal hairs distinguishes Platanus fruits from the fruit of Macginicarpa (the fossil genus used for fruits associated with Macginitiea J. A. WOLFE et W. WEHR in Manchester leaves) (Manchester 1986, Huegele and Manchester 2022). Platanus fruits are known from the mid-Eocene Clarno Nut Beds of (Manchester 1994). Platanus is an indicator for riparian environments (Nixon and Poole 2003).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF51826DFC66FEEB6AFAFC5E.taxon	description	Text-fig. 6 b M a t e r i a l. USNM PAL 722388. L o c a l i t y. Dakin. D e s c r i p t i o n. Follicle 2.6 mm long, 1.4 mm wide at the widest; bulbous with one end tapering to a point and the antipodal side ending with a small cylinder measuring 0.4 mm tall and 0.5 mm wide; seven parallel striations 0.1 mm wide extend longitudinally. R e m a r k s. Longitudinal ribbing along a chimney lamp-shaped follicle is characteristic of Thalictrum. Fossil Thalictrum fruits are known from the Oligocene and Miocene of Europe (Szafer 1961, Dorofeev 1963, Mai 1995) and the Pleistocene of Nepal (Bhandari et al. 2009, 2010, 2011). If confirmed, this would be the oldest Thalictrium ca. 20 Ma older than estimates for crown of that genus (Soza et al. 2013).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF518273FC7DF8B96FC6FEDB.taxon	description	M a t e r i a l. USNM PAL 621411, 622713, 622294, 623312, 624006, 624179, 624447, 624499, 624509, 624556, 624648, 624684, 624710, 624778, 717434, 717553, 712990, 625251, 625257, 625269, 625286, 625358, 625406, 625485, 625500, 625541, 625560, 625572, 625609, 625618, 625647, 625651, 625661, 625662, 625676, 625752, 625764, 625776, 625783. L o c a l i t i e s. Dakin, Disbrow Creek, Park and Spring. D e s c r i p t i o n. Winged seeds with seed body on one side; full seeds ranging from 2.5 – 5.5 mm long and 0.5 – 4.00 mm wide; seed bodies obovate to semi-fusiform, 1.1 – 3.3 mm long and 0.3 – 1.5 mm wide; seed body oblique with proximal end tilting to the ventral side of the wing and distal side tilting to the dorsal side of the wing; seed wings straight or curved with distal end straight or curved proximally; wing finely striate; raphe wrapping around the margin of the wing or turning dorsally near the middle to continue along the margin to the chalazal end of the seed body. R e m a r k s. Seeds with this morphology match the morphospecies Carpolithes lunatus HICKEY. Such seeds co-occur with or are a constituent of the fruits known as Jenkinsella Rᴇɪᴅ et CHANDLER, Joffrea CRANE et STOCKEY, Nyssidium HEER and are commonly associated with the leaves known as Trochodendroides BERRY (Crane and Stockey 1985, 1986, Golovneva and Alekseev 2017). Further investigations are needed to differentiate these plants based on seeds (Golovneva and Alekseev 2017). Brown (1939) documented cercidiphyllaceous elements from 30 localities in the western US and Dakotas from Cretaceous through Oligocene.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4F8273FC67FC7A6A81F86D.taxon	description	M a t e r i a l. USNM PAL 717328. L o c a l i t y. Disbrow Creek. Description. Samara 2.1 cm long, 0.7 cm wide at widest point of wing; nutlet elliptical 0.5 cm long and 0.3 cm wide; peduncle persistent, small projection pointing distally along the dorsal side of the wing near the apex of the nutlet; dorsal side of wing slightly convex; ventral side of wing becomes wider just past the nutlet apex and reaches full width ⅔ of the way to the distal end; veins originate near the nutlet apex and curve down to the ventral side of the wing, generally dichotomizing, occasionally anastomosing and cross veins with proximal veins more likely to anastomose. R e m a r k s. Fruits of Deviacer wolfei are known from the similarly aged Nut Beds of the Clarno Formation of central Oregon; they are samaras with elliptical nutlets, persistent styles, dorsal projections and fall within the same size range and ratios (Manchester 1994, Chen and Manchester 2015). Deviacer fruits that appear similar but are unassigned to a species are also known from the early Eocene Republic flora of eastern Washington (Pigg et al. 2008) and early Eocene Falkland flora of British Columbia (Smith et al. 2012). Late Paleocene fossil fruits with this morphology from Almont, North Dakota with anatomical preservation were named Paleosecuridaca PIGG, DEVORE et M. F. WOJC. (Pigg et al. 2008). According to those authors, similarities suggest that at least some of these fruits are assignable to Polygalaceae.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4F8273FEA8FC506EE4F989.taxon	description	Text-fig. 7 a M a t e r i a l. USNM PAL 722490. L o c a l i t y. Dakin. D e s c r i p t i o n. Endocarp elliptical 2.1 mm long, 1.1 mm wide; four to five lateral projections on each side measuring 0.2 – 0.4 mm long and 0.1 – 0.3 mm wide at the base. R e m a r k s. Myriophyllum fruits are rectangular to elliptical and have short, lateral projections. Pleistocene leaf fossils are known from Kashmir (Puri 1951) and endocarps from Nepal (Bhandari et al. 2009, 2010, 2011). Another fossil occurrence attributed to Haloragaceae is Tarahumara sophiae HERN. - CASTILLO et CEV. - FERRIZ from the Campanian- Maastrichtian of northern Mexico (Hernandez-Castillo and Cevallos-Ferriz 1999). The Kishenehn specimen may be the oldest macrofossil of the genus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4F8273FF5AFEEA6D6DFC51.taxon	description	M a t e r i a l. USNM PAL 620034. L o c a l i t y. Park. D e s c r i p t i o n. Raceme 23.4 mm long with three widely spaced follicles; peduncle 0.8 mm wide; follicles elliptical with truncated base and acute apex, 10.2 – 11.0 mm long, 3.2 – 4.0 mm wide. R e m a r k s. Infructescences of Jenkinsella knowltonii are generally 35 – 50 mm long, have 5 – 7 widely spaced follicles that are 10 – 16 mm long and 4 – 7 mm wide (Golovneva and Alekseev 2017). The Kishenehn specimen is slightly shorter, is lacking the preservation of reproductive shoots and has fewer follicles though this could be a preservational issue. J. knowltonii is known from the Paleocene Denver Formation of Colorado (Golovneva and Alexeev 2017).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4F8272FBB4F87D6A71F7DB.taxon	description	M a t e r i a l. USNM PAL 621987. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Legumes containing two circular seeds; fruit 3.5 mm long, 1.9 mm wide; fruit constricted between adjacent seeds such that width decreases to 1.2 mm between seeds; seeds 1.4 – 1.5 mm wide; style persistent and triangular 0.4 mm long and 0.2 mm wide at base.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4D8271FEE1FF266E8EFD95.taxon	description	M a t e r i a l. USNM PAL 625803. L o c a l i t y. Spring. D e s c r i p t i o n. Legume containing at least two circular seeds; fruit at least 15.9 mm long, 5.0 mm wide, constricted to 2.9 mm wide between seeds; seeds 5.7 – 5.8 mm wide; legume apex triangular 1.7 mm wide at base; 1.1 mm long, sutural wing 0.3 mm wide.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4D8271FEE1FF266E8EFD95.taxon	discussion	Comments on Fabaceae The fruit of Fabaceae sp. 2 can be differentiated from Fabaceae sp. 1 by its larger size and somewhat stronger constriction around the seeds.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4D8271FEFAF9CE6AF1FE9A.taxon	description	M a t e r i a l. USNM PAL 621976. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Fruit a samara, length 4.3 mm, width 1.9 mm, fruit body excluding stipe and styles 2.8 mm long, wing narrowing (0.3 mm) surrounding central seed body, seed body elliptical 2.1 mm long and 1.3 mm wide, two rectangular, prominent styles 1.2 mm long and 0.7 mm wide emerging apically in a V-shaped configuration. R e m a r k s. This fruit bears a resemblance to those of Ulmus okanaganensis DENK et DILLHOFF from the early Eocene of British Columbia and Washington (Denk and Dillhoff 2005). However, U. okanaganensis lacks a visible wing and has styles that are more tapered and curved to the center (Denk and Dillhoff 2005). There is also a similarity to fruits named as Ulmus sp. from the same time and area as U. okanaganensis (Denk and Dillhoff 2005), but the styles differ from this being more tapered and proximally curved.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4D8277FBB8FA966E79FB4F.taxon	description	M a t e r i a l. UF 19652 - 86483, 19652 - 86484, 19652 - 86485, 19652 - 86486, 19652 - 86487, 19652 - 86488. L o c a l i t y. M 3. D e s c r i p t i o n. Endocarp calcareous, ovoid, 4.7 mm long, 3.5 mm wide, with reticulate surface ridging, areoles of the reticulum square; endocarp 7 squares wide and about 8 squares tall. R e m a r k s. A calcified endocarp with reticulate sculpture enables us to place this fossil in Celtis. Such endocarps are commonly known from Eocene deposits (Manchester 1989, 1994). Family Juglandaceae DC. ex PERLEB Genus Palaeocarya SAPORTA Text-fig. 8 i M a t e r i a l. USNM PAL 622690. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Winged fruit incomplete, 2.1 cm long and 1.7 cm wide; main wing trilobed; lobes lanceolate diverging at 46.4 °, middle lobe 7.0 mm wide and lateral lobes 5.4 – 5.8 mm wide; wing lobes possessing three main veins, each with the midvein being the thickest; lateral ascending veins dissipate into loops distally, epimedial veins mostly perpendicular, exmedial veins branching apically, veins visible up to third order. Prophyllum ca. 0.8 cm long and 0.6 cm wide with straight, unbranched trichomes below the wing measuring between and 1.7 to 3.7 mm long. R e m a r k s. Palaeocarya is a fossil genus that accommodates fruits with trilobed wings, similar to those of the extant genera, Engelhardia LESCH. ex BLUME of Asia and Oreomunnea of Central America. Distinctions between species of the extant genera and in Palaeocarya can be made in part by wing venation patterns (Manchester 1987). Only the basal portions of the wings are seen in this specimen. Palaeocarya clarnensis MANCHESTER and P. uintaensis (MACGINITIE) MANCHESTER both appear to have three main veins in the basal portion of their wings, however, the lateral veins of P. uintaensis become secondary loops more apically (Manchester 1987) whereas P. clarnensis has a pubescent nutlet base (Manchester 1987) like this fossil. Because this fossil shares features of both species and P. uintaensis is known from only one specimen, this fossil is not being assigned to a species. More specimens may help resolve its taxonomic affinity. P. clarnensis has been observed in collections from the Green River Formation (Little Mountain, Wyoming) as well as from the Clarno Formation (West Branch Creek and Gosner Road, Oregon), with the Clarno localities being a similar age to the Kishenehn Formation (Manchester 1987). P. uintaensis is known only from the Green River Formation (Rainbow, Utah) (MacGinitie 1969, Manchester 1987).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4B8277FBC8FE896A00FD46.taxon	description	M a t e r i a l. USNM PAL 622732, 626069. L o c a l i t y. Deep Ford and Disbrow Creek. D e s c r i p t i o n. Seed body obovate 2.7 – 2.8 mm tall, 2.4 – 2.6 mm wide; base concave; apex retuse; two styles 0.3 – 0.4 mm tall, 0.1 – 0.2 mm wide with flat stigmas. R e m a r k s. These two specimens have wings that are not well differentiated from the seed body and have stout styles which resemble some species of Alnus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4B8277FBC8FD696B42FBA6.taxon	description	M a t e r i a l. USNM PAL 623147. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Seed body obovate 1.8 mm tall, 1.3 mm wide at widest; apex retuse; stylar base (s) present, 0.2 mm tall, 0.2 mm wide. R e m a r k s. The seed is smaller than Alnus sp. F 1 and has a thinner wing but still not as defined from the seed body as in most species of Alnus.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4B8277FEF1F94C6B35FF19.taxon	description	M a t e r i a l. USNM PAL 622756. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Infructescence cylindrical, 14.7 mm long, 7.1 mm wide; borne on a twig 1.2 – 3.8 mm wide. R e m a r k s. Although the attachment is not completely preserved, it appears that the peduncle on this specimen was short. Due to the thick, coalified bracts, this specimen likely belongs to Alnus rather than Betula, which has thinner, deciduous bracts.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4B8277FEE6FAAF6E28F95C.taxon	description	M a t e r i a l. USNM PAL 623160. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Fruit a samara, circular in outline, 2.5 mm tall, 2.4 – 2.6 mm wide; seed body elliptical, in center of fruit, 2.1 mm tall, 1.6 mm wide; styles two, persistent, 0.4 mm tall and 0.1 mm wide with flat stigmas. R e m a r k s. The specimen has a thin, differentiated wing as in most Betula species but has stout styles as in most Alnus species.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4A8276FEA9FB806E1FF89A.taxon	description	M a t e r i a l. USNM PAL 625742. L o c a l i t y. Spring. D e s c r i p t i o n. Samara 15.0 mm tall, 10.5 mm wide; pedicel 4.4 mm long, 0.4 mm wide; disc 0.5 mm tall, 1.7 mm wide; abortive fruit 1.1 mm in diameter; laminated vascular bundle emerging out of disc and into flat, circular seed body 1.0 mm in diameter, surrounded by circular wing 5.2 mm in diameter; veins radiate from center, dichotomizing occasionally between margin or halfway between margin and seed body; fimbrial vein surrounds wing. R e m a r k s. The presence of a hypogynous perianth and disc scar at the junction of the pedicel and fruit and shape of the samara conform to Dipteronia. Fimbrial veins may be absent in Dipteronia fruit such as Dipteronia sinensis OLIV. or present such as in Dipteronia dyeriana HENRY and Dipteronia brownii MCCLAIN et MANCHESTER (McClain and Manchester 2001). Dipteronia fruits are known from the Paleocene of Wyoming, the middle and late Eocene of British Columbia, Washington, Oregon and Colorado and the early Oligocene of Oregon (McClain and Manchester 2001), as well as from the early Oligocene of western China (Ding et al. 2018).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4A8276FB95FD2C6A4BF8DB.taxon	description	M a t e r i a l. USNM PAL 625792. L o c a l i t y. Spring. D e s c r i p t i o n. Fruit a bladder-like capsule, elliptical, 17.2 mm tall with three wings showing, each 6.8 mm wide, divided by a line of dehiscence; seeds circular, 1.0 mm long to 0.7 mm wide; one per locule attached near the midway point between base and apex of the fruit. This faintly preserved specimen shows quite clearly the longitudinal septum and darkened central seed attachments, but the outline of the surrounding bladder is seen only near the base and apex. The septa in Koelreuteria (Sapindaceae), terminate, distally about a third of the way up the fruit, where the seeds attach (Manchester et al. 2009), so that the locular cavities join into a single locule above the seed attachment. Our specimen does not show termination in the septa. Neither Craigia W. W. SM. et W. E. EVANS (Malvaceae), Urvillea KUNTH (Sapindaceae) nor Arfeuillea PIERRE ex RADLK. (Sapindaceae) possess this feature of our fossil. Species of Craigia vary in length from 9 – 35 mm and 7 – 33 mm in width, are septicidal and possess wings with radiating veins (Wang et al. 2021). Urvillea is 20 – 40 mm tall 15 – 18 mm wide (in U. venezuelensis FERRUCCI), is septifragal, one-seeded, has subparallel veins and its veins radiate outward (Ferrucci 2006, Wang et al. 2013, 2021). Arfeuillea is 50 mm long, 45 mm wide, is loculicidal and has net-like veins and a flat to slightly rounded apex. cf. Boniodendron is a potential candidate as the modern fruits are subglobose, 18 – 23 mm tall and have one seed per locule at the same level as in this fossil (Wang et al. 2013). Boniodendron has loculicidal dehiscence (Wang et al. 2013). Without a clear understanding of how the valves open or the venation, we are unable to assign this fossil to a genus. Based on the commonality of winged-bladder fruits in Sapindaceae, the fossil likely belongs to this family.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF4A8275FC69F82D6B1FF8BF.taxon	description	M a t e r i a l. USNM PAL 622108, 622276, 622650. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Winged fruit elongate-elliptical, with a central seed body; wing with pointed apex and base; fruit 25.1 – 32.12 mm long, 7.3 – 8.6 mm wide; seed body 4.9 – 6.4 mm long, 3.7 – 5.3 mm wide; ventral (intramarginal) vein of the wing prominent, marginal or submarginal; stylar vein present, minor vein small, mostly parallel, occasionally dichotomizing or anastomosing (more on dorsal side); seed bodies sub-circular. R e m a r k s. Ailanthus fruits in North America have traditionally been placed into either Ailanthus americana COCKERELL or Ailanthus lesquereuxi COCKERELL but are morphologically identical to those described earlier as Ailanthus confucii UNGER (Corbett and Manchester 2004). The fossil matches the description of A. confucci by having a marginal to submarginal ventral vein as opposed to having an inset vein as in Ailanthus tardensis HABLY (Corbett and Manchester 2004). A. confucii is recognized from the early Eocene through late Miocene of western North America (Corbett and Manchester 2004). Among extant species, it resembles the temperate eastern Asian native species, Ailanthus altissima, which is invasive today in North America and Europe.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF498274FB8BF8CD6D4BFCAF.taxon	description	Text-fig. 9 d M a t e r i a l. USNM PAL 623735. L o c a l i t y. Dakin. D e s c r i p t i o n. Winged seed at least 7.5 – 10.8 mm long, 3.4 – 4.9 mm wide at widest; seed body elliptical, on proximal side of wing, long axis of seed body almost parallel to wing, pointing slightly ventrally, 2.5 – 3.4 mm long, 1.3 – 2.1 mm wide; wing extending around whole seed body becomes wider halfway down seed body and becomes widest past the distal pole where it maintains width; wing contains small, straight striations that run parallel to the course of the wing. R e m a r k s. We were unable to observe the cellular pattern of the wing so are hesitant to place it firmly in the genus Cedrela because similar veinless single-winged seeds occur in many other taxa (for listing of examples, see Kvaček and Wilde 2010). Cedrela is reported from the early Eocene Kisinger Lakes and (rarely) in the Green River Formation, late Eocene Florissant Formation of Colorado (MacGinitie 1953) and early Oligocene Bridge Creek flora of the John Day Formation, Oregon (Meyer and Manchester 1997). Cedrela - like seeds are known from the late Eocene Whitecap Knoll flora of the John Day Formation, Oregon but differ in wing cell pattern (Manchester 2000).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF488274FB84FA546B82F8E4.taxon	description	M a t e r i a l. USNM PAL 624090. L o c a l i t y. Dakin. D e s c r i p t i o n. Seed circular, curved embryo, 1.3 mm long, 1.1 mm wide; eight rows of circular, concentric reticulae; hilar region 0.3 mm long, 0.1 mm deep; persistent funicle 0.1 mm wide, 0.1 mm wide.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF488274FF74FC3E6E81FA60.taxon	description	M a t e r i a l. USNM PAL 624113. L o c a l i t y. Dakin. D e s c r i p t i o n. Seed ovate 2.8 mm long, 2.1 mm wide at widest point with elongate lateral hilum 0.7 mm long; reticulate to punctate sculpture with round fossae. R e m a r k s. The fossil genus Rutaspermum is used for seeds with rutaceous affinity that cannot be placed into a modern genus. Whereas our seed has a similar shape to other Rutaspermum, the sculpture is more pronounced in our fossil (Gregor 1989). Modern rutaceous genera such as Zanthoxylum can extend from subtropical into temperate latitudes (eFloras 2008 – 2022).	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF488274FEC4FA606BD6FA95.taxon	description	M a t e r i a l. USNM PAL 622306, 623124, 623138 and DMNH EPI. 41847, 48190, 48216, 48222, 48239. L o c a l i t y. Disbrow Creek. D e s c r i p t i o n. Winged capsular fruit 14.3 mm long, 9.2 mm wide; pedicel 2.9 mm long; pedicel truncated; wing elliptical to ovate, with a rounded base and emarginate apex; approximately 2.5 mm between edge of central body and wing margin; fimbrial vein present; veins arise between 38 – 46 °. Arching apically about half-way, occasional bifurcations with the apical vein retaining the same thickness; veins lose thickness towards margin; central locular area thicker / darker than surrounding wing, fusiform, 7.2 mm long, 3.1 mm wide with longitudinal groove, slit or suture running the entire length; persistent styles 1.3 mm long; wing surrounds two styles. R e m a r k s. We considered various candidate families for the identification of these winged fruits before settling on the assignment to Thlaspi in the Brassicaceae. The fossils are distinctive in the presence of a central body, persistent styles and arching veins of the wings. The dispersed valves of Koelreuteria LAXM. (Sapindaceae) and Craigia W. W. SM. et W. E. EVANS (Malvaceae) capsular fruits also have bilateral symmetry and are known from Eocene deposits of North America (Kvaček et al. 2005, Wang et al. 2013). Craigia can be distinguished from Koelreuteria by the presence of a fusiform central locule in the former (Wang et al. 2013). Two species of Craigia are currently recognized, Craigia bronnii (UNGER) KVAČEK, BŮŽEK et MANCHESTER from Europe and western Asia and Craigia oregonensis (ARNOLD) KVAČEK, BŮŽEK et MANCHESTER from North America and Asia (Kvaček et al. 2005). These fruits have straight veins that do not lose gauge towards the wing margin unlike the fossil (Kvaček et al. 2005). We also considered the families Begoniaceae C. AGARDH and Polygonaceae JUSS. The sides of the proximal portion of the central body in Begonia L. is detached from the wing but the overall shape is obovate and there is persistent stamen and style (Manchester and O’Leary 2010). The wings on polygonaceous fruit are derived from the perianth and possess thin veins (Manchester and O’Leary 2010). The venation pattern, locule shape, persistent pedicel and style of these specimens are consistent with the previously described fossil Thlaspi primaevum H. F. BECKER from the early Oligocene Ruby flora from western Montana (Becker 1961, Beilstein et al. 2010). We also compared the specimen to Noccaea MOENCH but the veins of that genus are less defined than those in Thlaspi. The wings of Thlaspi primaevum do not fully surround the stylar area and the perianth scars are more pronounced (Becker 1961, Manchester and O’Leary 2010). This fossil from the Kishenehn is the oldest occurrence of Thlaspi. It would imply still an earlier radiation of the clade that includes Arabidopsis (Beilstein et al. 2010). The type species, T. arvense L., has a similar wing morphology, with the wings encompassing the stylar area, though the central body appears to be proportionally wider.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF488279FB84F8876DADFCFA.taxon	description	M a t e r i a l. USNM PAL 624372. L o c a l i t y. Dakin. D e s c r i p t i o n. Seed circular, 1.4 – 1.2 mm in diameter, funicle 0.3 mm long, 0.3 mm wide, sculpture psilate.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF488279FB84F8876DADFCFA.taxon	discussion	Comments on Chenopodium Chenopodium seeds are round with a funicle and range in sculpture from reticulate to psilate (Benet-Pierce and Simpson 2014). A reticulate sculptured Chenopodium is known from the Miocene-Pleistocene of Tibet, however, Chenopodium sp. 1 has larger fossa in the reticulae and Chenopodium sp. 2 is psilate (Huang et al. 2021). Modern Chenopodium seeds can be reticulate or psilate (Benet-Pierce and Simpson 2014). Here we figure Chenopodium pratericola RYDB. (Text-fig. 11 c, Minnesota Wildflowers 2023 b) and Chenopodium polyspermum L. (Text-fig. 11 d, Agricultura Romaneasca 2011) as comparisons. We also compared these two seeds to those of Corydalis DC. (Papaveraceae) which are of similar size, shape and may be reticulate or psilate. It would appear though that Corydalis seeds are slightly less circular and have a deeper indentation near the funicle (pers. obs.). Rutaspermum was also identified from the Kishenehn but we distinguish these Chenopodium from the Rutaspermum by their circular rather than reniform shape. Eurya THUNB. (Theaceae) is also reniform (Bhandari et al. 2009). We also looked at other genera in the Chenopodioideae. Stutzia E. H. ZACHARIAS has fruiting bracts that are arrowhead shaped (Zacharias 2012). Cycloloma fruits generally have attached, papery perianth, and although that could have worn off to leave the seed, the hilar scar is less prominent (image by Carole Ritchie, United States Department of Agriculture Natural Resource Conservation Service PLANTS Database in Vescovo 2023). Both Chenopodiastrum S. FUENTES and Cycloloma appear less coiled than Chenopodium (image by Stefan Lefnaer in Areces-Berazain 2022, image by Carole Ritchie, United States Department of Agriculture Natural Resource Conservation Service PLANTS Database in Vescovo 2023). This is not an exhaustive list of genera in Chenopodioideae but should serve to illustrate that these seeds are distinguishable from other dissiminules in the subfamily. These are the oldest recorded Chenopodium fossils.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF458279FBC1FA7C6A6EF889.taxon	description	M a t e r i a l. USNM PAL 722179. L o c a l i t y: Disbrow Creek. D e s c r i p t i o n. Schizocarpic achene; flat proximal edge and rounded distal edge, 4.3 mm tall, 3.6 mm wide (excluding spines); six lateral spines on distal apical end measuring between 0.6 and 1.5 mm long. R e m a r k s. Spiny achenes close in size and shape are found in Hydrocotyle L. (Araliaceae JUSS.) (e. g., H. scutellifera BENTH.) (Perkins 2019) and Apiaceae.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF458279FBB2FCFC6A46FA69.taxon	description	M a t e r i a l. USNM PAL 624316, 625396, 624316. L o c a l i t i e s. Dakin and Spring. D e s c r i p t i o n. Bi-winged seed, 5.9 – 10.2 mm long, 1.1 – 1.4 mm wide; distal wing margin hairy; wings never wider than central body; central body 3.0 mm long; seed elliptical, 1.3 – 2.6 mm long 0.6 – 1.4 mm wide. R e m a r k s. Bi-winged seeds are characteristic of many Bignoniaceae. The tribe Catalpeae DC. ex MEISN. includes the extant genera Catalpa SCOP. and Chilopsis D. DON, which both have distally shredded seed wings (Lersten et al. 2002). We choose to point out this similarity instead of placing our specimens in Catalpeae, out of caution, because we have not conducted a full survey of the family; other bignoniaceous tribes might also have distally shredded seed wings.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF458279FE91FCCA6D1AF87C.taxon	description	M a t e r i a l. USNM PAL 717259, 622651, 623020, 623513, 624027, 624249, 624336, 624374, 624717, 624808, 717508, 625160, 625523, 625756, 768185. L o c a l i t i e s. Dakin, Disbrow Creek, Park and Spring. D e s c r i p t i o n. Samaras 1.6 – 8.9 mm long to 0.7 – 4.5 mm wide; endocarp 1.0 – 6.5 mm long to 0.6 – 3.2 mm wide; fruits and seed bodies elliptical with seed bodies in the center of the samara; reticulate venation over the endocarp and a strong intramarginal or marginal vein along the wing margins; stigmatic cleft, positioned laterally near the apex; basal stipe present. R e m a r k s. Size and length width ratios are the key distinguishing factors for fossil Eucommia species endemic to east Asia (Call and Dilcher 1997, Manchester et al. 2009). These fossils fall within the size ranges of Eucommia montana which displays a wide range (Becker 1969, Call and Dilcher 1997). Eucommia montana is known from the early Eocene to Oligocene as far north as Quilchena, British Columbia and as far south and east as Florissant, Colorado (Call and Dilcher 1997). This species had smaller fruits than those known from the Eocene of Mississippi (Call and Dilcher 1997) and Oligocene of southern Mexico (Magallón-Puebla and Cevallos-Ferriz 1994). Eucommia is now endemic to China, but its fossil record includes North American and European as well as Asian occurrences (Manchester et al. 2009). Family Oleaceae HOFFMANNS. et LINK Genus Fraxinus L.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF45827FFC20F82C6D0DFE47.taxon	description	P l a n t F o s s i l N a m e s R e g i s t r y. PFN 003030 (for new genus). G e n e r i c d i a g n o s i s. Winged structure circular with dark, thick striations radiating around obovate seed body, seed body with rugulate sculpture. E t y m o l o g y. The genus is Greek for “ variegated winged seed ” with “ ptero- ” for “ wing ”, “ - heterochros- ” for “ variegated ” and “ - sperma ” for “ seed. R e m a r k s o n g e n u s. One species in newly erected genus, preserved on tan-gray paper-shale.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF43827FFF4AFE676D02F8E0.taxon	description	P l a n t F o s s i l N a m e s R e g i s t r y. PFN 003031 (for new species). E t y m o l o g y. The specific epithet is for the Horsefly, British Columbia where the first fossils of this species were described. H o l o t y p e. USNM PAL 722530, National Museum of Natural History, District of Columbia, USA, Text-fig. 12 a. Ty p e h o r i z o n a n d t y p e l o c a l i t y. DakinSite in the Kishenehn Formation, ca. 44 Ma. S p e c i f i c d i a g n o s i s. Winged structure circular; wing has thick striations (up to 0.1 mm) radiating around seed body starting from the seed body and mostly terminating at the margin of the wing, some striations bifurcating; seed body obovate with rugulate sculpture. S i z e. Structure 2.1 mm tall, 2.7 mm wide; seed body 1.28 mm tall, 0.77 mm wide. D e s c r i p t i o n. Roughly 27 striations radiating from the seed body, some bifurcating or trifurcating. R e m a r k s. Similar disseminules have been found from the early Eocene Okanogan sites of Republic, WA (Pigg and Wehr 2002), McAbee, BC and Horsefly, BC (UWBMB 4131 - 96008, DSCN 6543). Penhallow (1908) assigned specimens from Horsefly of this disseminule as Ulmus minuta GOEPP. (94 – 95), which was a leaf species, but he recognized that the fruit was much smaller than other known fossil and modern elms. U. minuta is a synonym for Ulmus pyramidalis (GOEPPERT) emend. ILJINSK. (Takhtajan 1982, Traiser et al. 2019). U. pyramidalis DIPPEL was already in use for a different extant elm and was recognized as a synonym of Ulmus glabra HUDS. U. glabra ’ s modern geographic range includes Europe and western Asia but its fruits lack the thick, darkened veins that appear in the fossil and possess two styles which the fossil lacks (Thomas et al. 2018, Sherman-Broyles 2021). These tiny disseminules lack the wing venation and stylar cleft and style diagnostic for Ulmus, and thus require a new binomial.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF43827FFC17FD7B6A7BF7F0.taxon	description	P l a n t F o s s i l N a m e s R e g i s t r y. PFN 003033 (for new species). E t y m o l o g y. The specific epithet is for the for the Kishenehn Formation. M a t e r i a l. USNM PAL 624649, 625447, 622695, 622402, 623987, 624154. H o l o t y p e. USNM PAL 625447, National Museum of Natural History, District of Columbia, USA, Text-fig. 12 c. Ty p e h o r i z o n a n d t y p e l o c a l i t y. Spring Site in the Kishenehn Formation, ca. 44 Ma. O t h e r l o c a l i t i e s. Dakin, Dishbow Creek and Spring Sites. S p e c i f i c d i a g n o s i s. Flower square in cross section, elliptical in longitudinal section; pedicel slightly curved; awl-shaped and apices point outward with concave calyx lamina curving up to connect corners; hair-like projections protruding from near the axis. S i z e. Flower 1.5 – 2.2 mm diagonal cross section, each side 1.0 – 1.5 mm long, 2.9 – 5.1 mm tall; pedicel 0.2 – 0.3 tall, basal corners of flower protrude outward 0.3 – 0.4 mm wide; hair like projections 0.5 – 0.8 mm long. D e s c r i p t i o n. Hair-like projections are in the same plane as the corners of the flower. R e m a r k s. Possible affinities for this structure are flowers or fruits in tribe Saniculeae and tribe Scandiceae subtribe Scandicinae in Apiaceae because of their shape, size and projections. PAL 722210 “ KF 1 ”. Scale bar 2 mm. g: USNM PAL 624877 KF 2. Scale bar 1 mm. h: USNM PAL 624832 KF 3. Scale bar 2 mm. i: USNM PAL 768303 KF 4. Scale bar 1 mm. j: USNM PAL 625629 KF 4. Scale bar 3 mm. k: USNM PAL 624455 KF 5. Scale bar 3 mm. l: USNM PAL 609598 KF 6 seed. Scale bar 1 mm. m: USNM PAL 722298 KF 7. Scale bar 1 mm. n: USNM PAL 625631 KF 7. Scale bar 1 mm. o: USNM PAL 625736 KF 8. Scale bar 2 mm. p: USNM PAL 625551 KF 9 Scale bar 1 mm. q: USNM PAL 621685 KF 9. Scale bar 1 mm. r: USNM PAL 620311 KF 9. Scale bar 1 mm. s: USNM PAL 626216 KF 9. Scale bar 1 mm. t: USNM PAL 625608 KF 9. Scale bar 1 mm.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
4D2487A3EF43827FFC48FF266A38FD4B.taxon	description	P l a n t F o s s i l N a m e s R e g i s t r y. PFN 003032 (for new genus). G e n e r i c d i a g n o s i s. Epigynous flower square in cross section with awl-shaped projections in each corner beginning at the base above the pedicel, connected by a lamina; base flat to slightly rounded; center with hair-like projections. E t y m o l o g y. The genus is Latin for “ four awl flower ” (“ quadra- ” for “ four ”, “ - subula- ” for “ awl ” and “ - flora ” for “ flower ”). R e m a r k s o n g e n u s. One species in newly erected genus, preserved on tan-gray paper-shale.	en	Smith, MacKenzie A., Greenwalt, Dale E., Manchester, Steven R. (2023): Diverse Fruits And Seeds Of The Mid-Eocene Kishenehn Formation, Northwestern Montana, Usa, And Their Implications For Biogeography. Fossil Imprint 79 (1): 37-88, DOI: 10.37520/fi.2023.004, URL: http://dx.doi.org/10.37520/fi.2023.004
