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New Species of Angiosperm Pollen from the Dakota Formation (Cenomanian, Upper Cretaceous) of Minnesota, U.S.

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Author(s): Shusheng Hu, David M. Jarzen, and David L. Dilcher Source: Palynology, 32(1):17-26. 2008. Published By: AASP: The Palynological Society URL: http://www.bioone.org/doi/full/10.2113/gspalynol.32.1.17

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NEW SPECIES OF ANGIOSPERM POLLEN FROM THE DAKOTA FORMATION (CENOMANIAN, UPPER CRETACEOUS) OF MINNESOTA, U.S.A.
SHUSHENG HU Paleobotany and Palynology Laboratory Florida Museum of Natural History University of Florida Gainesville, Florida 32611-7800, U.S.A. current address: Department of Biology Indiana University Southeast New Albany, Indiana 47150, U.S.A. e-mail: hus@ius.edu DAVID M. JARZEN DAVID L. DILCHER Paleobotany and Palynology Laboratory Florida Museum of Natural History University of Florida Gainesville, Florida 32611-7800, U.S.A. e-mail: dmj@flmnh.ufl.edu; dilcher@flmnh.ufl.edu

S. Hu, D.M. Jarzen, and D.L. Dilcher: New species of Cenomanian angiosperm pollen, Cenomanian of Minnesota, U.S.A.

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Abstract
Five new species of angiosperm pollen described from three localities in south-central Minnesota support a Cenomanian age for the Dakota Formation in this area. The new taxa, Cupliferoidaepollenites microscabratus, Dryadopollis minnesotensis, Nyssapollenites microfoveolata, Phimopollenites striolata, and Tricolpites labeonis, are described using both light and scanning electron microscopy, are easily recognizable, and are useful age indicator species. Because of the extent and transgressive nature of the Dakota Formation, age determination has been difficult. The new angiosperm pollen species described here may help to support age determination for the Dakota Formation over a broader geographic area. Key words: Late Cretaceous, Cenomanian, Dakota Formation, angiosperm pollen, taxonomy, biostratigraphy, paleoecology.

INTRODUCTION The Dakota Formation is a lithostratigraphic unit across a vast area of central and west-central North America (Ravn and Witzke, 1995). The name has often been used without consideration of the type Dakota Formation, either lithostratigraphically or chronostratigraphically (Witzke et al., 1983). Therefore the age and lithology of the Dakota Formation are probably not the same from the west margin to the east margin of the Western Interior Seaway. Currently, the age of the Dakota Formation in southwest Minnesota is thought to be Cenomanian (Setterholm, 1994). However, because of the absence of marine fossils, this suggestion of a Cenomanian age has mainly been based upon interpretations from the paleobotanical work of

Lesquereux (1895), and the palynological studies of Pierce (1961). The paleofloras of Lesquereux (1895) and the palynofloras of Pierce (1961) require reexamination and reinterpretation (Upchurch and Dilcher, 1990; Wang, 2002; Hu et al., 2004). Austin (1972) proposed that the nonmarine Cretaceous sediments in the Minnesota River Valley are approximately middle Cenomanian based upon clay mineralogy. Setterholm (1994) further proposed that the upper mudstone unit of the Dakota Formation in east-central Minnesota may be coeval with the Graneros Shale in western Minnesota, and that both units are late Cenomanian. Hu (2006) examined the palynofloras from the Dakota Formation of three clay pits in southern Minnesota. The Cretaceous sediments from these clay pits provide a rich

Palynology, 32 (2008): 1726 2008 by AASP Foundation

ISSN 0191-6122

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and diverse assemblage of palynomorphs, allowing refinement of the age (Hu, 2006). This paper is based on Hu (2006), and describes five new angiosperm pollen species identified from the Dakota Formation, and discusses their implications for age determination. GEOLOGIC SETTING During the Early Cretaceous, two epicontinental seas, the Boreal in the north and the Gulf in the south, were present on continental North America. The Boreal Sea advanced southward, meeting the northward-advancing Gulf Sea (Obradovich and Cobban, 1975) to form the Western Interior Seaway. This was a continuous sea extending from the Arctic to the Gulf of Mexico across North America during the late Albian (Witzke and Ludvigson, 1996). The Western Interior Seaway was transgressive during the late Albian to the Cenomanian; it was bordered to the west by the Cordilleran thrust belt, and on the east by the cratonic platform (Dyman et al., 1994). The type area of the Dakota Formation is located along the Missouri River in northeastern Nebraska and northwestern Iowa (Ravn and Witzke, 1995). The Dakota Formation is a sequence of nonmarine to marginal marine facies that are the oldest Cretaceous sediments in southwestern Minnesota (Witzke and Ludvigson, 1994). It includes two lithostratigraphic units, the lower sandstone unit and the upper mudstone unit; these are respectively similar to the Nishnabotna and Woodbury members of the Dakota Formation in age and lithology (Setterholm, 1994). The age of the Dakota Formation in southwest Minnesota is currently thought to be Cenomanian (Setterholm, 1994). Courtland Clay Pit (localities UF 19006, UF 19007) The sediments at Courtland Clay Pit (Nicollet County, MN, 4416'29" N, 9423'13" W) are dominated by laminated mudstone. Hajek et al. (2002) interpreted the sedimentary environment as a large lake based upon the millimeter to centimeter scale laminae, scattered, well-preserved leaves, and siderite (FeCO3) concretions. Lake Drummond (3636'12'' N, 07628'06'' W) in Virginia is a typical coastal lake, and may represent a similar environment to sediments of this Cretaceous lake exposed at Courtland Clay Pit. Highway 4 Clay Pit (locality UF 19000) The sediments at Highway 4 Clay Pit (Brown County, MN, 4426'05" N, 9443'37" W) consist predominantly of tabular cross-bedded sandstone and carbonaceous siltstone. Hajek et al. (2002) interpreted the sedimentary environ-

ment as a tidally influenced meandering river system based on inclined heterolithic stratification and tabular crossbedded fine-grained sandstone. Oxbow lakes, ridge and swale, and levee are important environments associated with meandering river systems. Ochs Clay Pit (locality UF 15750) The sediments at Ochs Clay Pit (Brown County, MN, 4413'26" N, 9500'42" W) are dominated by silty mudstone and siltstone. Sloan (1964) indicated that the sediments below a prominent lignite layer probably represent a lacustrine environment, based on the varved mudstone and abundant leaf fossils. By contrast, the sediments above the lignite layer probably represent an estuarine environment, based upon silty and sandy mudstone, and several shark vertebrae. The lignite may represent the distal side of the coastal swamp, which is not close to active beach barrier systems. The lowermost lacustrine sediments at Ochs Clay Pit appear to represent a low-energy lake paleoenvironment (Twenhofel, 1932; Visher, 1965). The sediments at these three clay pits in southwestern Minnesota are isolated from each other. It is difficult to make correlations between these three pits based upon lithology. Leaf megafossils are not satisfactory for comparative dating between these localities because a Dakota Formation leaf biostratigraphy has not been formulated. However, plant microfossils including pollen and spores are abundant in these sediments, and pollen has been successfully used for stratigraphic zonations of the Dakota Formation in other areas (Brenner et al., 2000). Based on the frequent occurrences of Fraxinoipollenites constrictus, Liliacidites reticulatus, Phimopollenites striolata, and Tricolpites cf. vulgaris, in sample UF 19007036710 at Courtland Clay Pit, sample UF 19000-046517 at Highway 4 Clay Pit, and sample UF 15750-046522 at Ochs Clay pit, these sediments are coeval. It is therefore possible to make stratigraphic comparisons between the three clay pits. MATERIALS AND METHODS Samples were collected from the Courtland Clay Pit, the Highway 4 Clay Pit, and the Ochs Clay Pit in southwestern Minnesota by Shusheng Hu in 2003 and 2004 (Text-Figure 1). Five stratigraphic sections were measured (Text-Figure 2). Pollen samples were collected at about 30 cm intervals from each of these sections. Standard processing techniques were used for all samples (Traverse, 2007). At the Courtland Clay Pit, 23 samples were collected; nine samples were processed of which eight samples contained abundant palynomorphs. At Highway 4 Clay Pit, 12 samples were

S. Hu, D.M. Jarzen, and D.L. Dilcher: New species of Cenomanian angiosperm pollen, Cenomanian of Minnesota, U.S.A.

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Text-Figure 1. Map of Minnesota illustrating the location of the three sites studied.

collected; eight samples were processed, with three samples providing abundant palynomorphs. At the Ochs Clay Pit, 27 samples were collected, from which eight samples were processed, with seven samples having abundant palynomorphs. All the samples were processed at the Paleobotany and Palynology Laboratory at the Florida Museum of Natural History (FLMNH), Gainesville, Florida, U.S.A. At least two slides of each sample were scanned in order to build a catalogue of pollen and spore morphotypes. Pollen counts were of at least 300 palynomorphs, and were made using a ZEISS Axiophot microscope. An AxioCam digital camera and imaging capturing software were used for the palynomorph identification and photography. Scanning electron microscopy was carried out using a Hitachi S4000 field emission scanning electron microscope (SEM) at the Department of Botany, University of Florida. Pollen and spore identifications were made by comparison with images and descriptions in the literature (e.g. Agasie, 1969; Brenner, 1963; Doyle and Robbins, 1977; May and Traverse, 1973; Pierce, 1961; Ravn, 1981; Ravn and Witzke, 1995; Romans, 1975), and the holotypes of Hedlund (1966), in the Sam Noble Oklahoma Museum of Natural History, Norman, Oklahoma, U.S.A. Slides, residues, SEM stubs, and unprocessed samples are deposited at the Paleobotany and Palynology Laboratory, Florida Museum of Natural History, Gainesville, Florida, U.S.A., as localities UF 15750, UF 19000, UF 19006, and UF 19007. The locations of the specimens illustrated in Plates 1 and 2 are given by reference to the

Text-Figure 2. Stratigraphic sections of the five measured sections from the three clay pits studied in Minnesota. The sections are arranged in relative stratigraphic position to each other. Note the vertical centimeter scale to the right. The horizontal distances between the sections are not to scale.

Paleobotany and Palynology Locality, the FLMNH (UF number), followed by a specimen number, with a slide number (PY01.PYn). The microscope stage coordinates are listed as the England Finder Slide (EFS) locations. SYSTEMATIC PALEONTOLOGY Anteturma POLLENITES Potoni 1931 Turma PLICATES Naumova 1939 emend. Potoni 1960 Subturma TRIPTYCHA Naumova 1939 emend. Potoni 1960 Genus Cupuliferoidaepollenites Potoni et al. 1950 ex Potoni 1960

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Type species Cupuliferoidaepollenites liblarensis (Thomson in Potoni et al., 1960) Potoni 1960 Cupuliferoidaepollenites microscabratus sp. nov. Plate 1, figs. 14 Diagnosis. Pollen grains free, isopolar; subprolate, prolate to perprolate (P/E=1.252.38), amb circular to subcircular; tricolpate, colpi nearly extending to the poles, somewhat thickened colpi margins, apocolpia small; exine thin, ca. 0.61.0 m, two-layered, nexine thin, sexine baculate, forming a microreticulate sculpture, lumina encircling 35 capita, sexine supratectally scabrate. Dimensions. Equatorial view 7(11)17 x 10(16)25 m (10 grains); polar view 12(16)22 m (5 grains). Holotype. UF 15750-046522-PY01A, EFS X45. Remarks. Psilatricolpites psilatus Pierce (1961) is larger, measuring 21 x 29 m. Occurrence. Courtland Clay Pit and Ochs Clay Pit. Name derivation. The specific epithet, microscabratus, is derived from the supratectal scabrate sexine. Genus Tricolpites Cookson 1947 ex Couper 1953 emend. Jarzen & Dettmann 1989 Type species Tricolpites reticulatus Cookson 1947 by subsequent designation of Couper 1953 Tricolpites labeonis sp. nov. Plate 1, figs. 59 Diagnosis. Pollen grains free, isopolar; subprolate, prolate (P/E=1.212.00), amb circular to subcircular; tricolpate, colpi nearly extending to the poles, apocolpia small; exine thin, ca. 0.5 m, ?two-layered, sexine columellate, pila short; microreticulate, lumina less than 0.5 m. SEM studies have shown that there is about a 0.7 m wide margin along the colpi on which the lumina are small (less than 0.1 m in diameter), and rare or absent. As this feature is not

observed under transmitted light microscopy, the surface is considered evenly reticulate over its entire surface, and therefore is included in Tricolpites. Dimensions. Equatorial view 6(10)14 x 9(14)19 m (14 grains); polar view 18 m (1 grain). Holotype. UF 15750-046526-PY03A, EFS Y37. Remarks. Jarzen and Dettmann (1989), following Ward (1986), emended the diagnosis of Tricolpites to include tricolpate grains with a regular (homobrochate) reticulum of uniform size measuring <1 m in diameter over the entire surface of the grain. Gunnerites and Retitricolpites are considered junior synonyms of Tricolpites. Tricolpites labeonis is similar to Tricolpites minutus (Brenner 1963) Dettmann 1973 (see Dettmann, 1973) in size and ornamentation, but it is differentiated from Tricolpites minutus by its narrow margin along the colpi on which the lumina are reduced or absent. Tricolpites labeonis sp. nov. is similar in shape, size, and ornamentation to in situ pollen of early to middle Albian flowers described as Aquia brookensis from the Bank near Brooke locality, Virginia (Crane et al., 1993), but differs in lacking the verrucose surface of colpi membranes in the pollen of Aquia brookensis. Occurrence. Courtland Clay Pit and Ochs Clay Pit. Name derivation. The specific epithet, labeonis, means one with large lips, based on the appearance of the margins of the colpi as seen using the SEM (Plate 1, fig. 8). Subturma PTYCHOTRIPORINES Naumova 1939 Genus Dryadopollis Srivastava 1975 Type species Dryadopollis argus Srivastava 1975 Dryadopollis minnesotensis sp. nov. Plate 1, figs. 1015 Diagnosis. Pollen grains free, isopolar; prolate spheroidal, subprolate to prolate (P/E=1.111.88), tricolporate,

PLATE 1
The scale bar represents 10 m unless otherwise noted. 14 Cupuliferoidaepollenites microscabratus sp. nov. 1: holotype, single grain, mid-focus. UF 15750-046522PY01A, X45. 2: UF 15750-046522-PY01A, Q18/4, pollen clump. 3: SEM, UF 19007-036708 stub 12, pollen clump. 4: as 3, detail of surface of single grain. Tricolpites labeonis sp. nov. 5, 6: holotype, single grain at high and mid-focus levels respectively, UF 15750-046526-PY03A, Y37. 7: SEM UF 1575046533, stub 3, scale bar = 2 m. 8: UF 15740-46533 stub 3, scale bar = 2 m. 9: UF 15750-46533-PY03A, R35/1 pollen clump. Dryadopollis minnesotensis sp. nov. 10: UF 19007036708 >10 , N36, mid-focus, pollen clump. 11: UF 19007-036708 >10 , M35/1, mid-focus, pollen clump. 12: holotype, UF 19006-036694-PY01A, S31/1, high focus. 13: as 12 mid-focus. 14: SEM, UF 19007036708, stub 12, pollen clump. Scale bar = 6 m. 15: as 14, close-up of the pore. Scale bar = 1 m.

1015

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S. Hu, D.M. Jarzen, and D.L. Dilcher: New species of Cenomanian angiosperm pollen, Cenomanian of Minnesota, U.S.A.

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apocolpia small; exine 1 m, ?two-layered, sexine columellate, pila short; microreticulate, lumina 0.11.0 m, muri ca. 0.5 m wide, lumina decreasing toward colpi and poles (heterobrochate of Ward, 1986, p. 25), ora small, about 0.8 m in diameter. Dimensions. Equatorial view 8(15)19 x 15(19)23 m (5 grains); polar view 16(17)19 m (13 grains). Holotype. UF19006-036694-PY01A, EFS S31/1. Remarks. Srivastava (1975) erected Dryadopollis to accommodate tricolporate pollen with a reticulum that is finer at the apocolpia and colpi margins than in the mesocolpal regions. This species is distinct from Dryadopollis argus, and Dryadopollis vestalis Ward 1986, in its smaller lumina and ora, and thinner exine. Occurrence. Courtland Clay Pit. Name derivation. The specific epithet, minnesotensis, is derived from Minnesota, where the fossil pollen was recovered. Genus Phimopollenites Dettmann 1973 Type species Phimopollenites pannosus (Dettmann & Playford 1968) Dettmann 1973 Phimopollenites striolata sp. nov. Plate 2, figs. 18 Diagnosis. Pollen grains free, isopolar; prolate spheroidal, subprolate to prolate (P/E=1.061.62), amb circular to subcircular; tricolporoidate, colpi slightly ragged and thickened, apocolpia small; exine 12 m, two-layered, nexine thinner than sexine, sexine columellate, pila dense; sexine microreticulate, lumina less than 0.5 m. SEM studies show the muri to be composed of closely spaced capita whose apical ends protrude supratectally, and give the appearance of a striate muri surface (Plate 2, fig. 8). Dimensions. Equatorial view 12(14)19 x 16(19)21 m (10 grains); polar view 15(18)24 m (7 grains). Holotype. UF 19006-046517-A1, >10 m, EFS N29/3.

Remarks. This species has a striate structure (muri with weak, transverse striations), that is different from other species of Phimopollenites when seen under the SEM. Occurrence. Courtland Clay Pit, Highway 4 Clay Pit, and Ochs Clay Pit. Name derivation. The specific epithet, striolata, is diminutive for furrow, as the muri appear to be furrowed (Plate 2, fig. 8). Genus Nyssapollenites Thiergart 1937 Type species Nyssapollenites pseudocruciatus (Potoni 1931) Thiergart 1937 Nyssapollenites microfoveolata sp. nov. Plate 2, figs. 915 Diagnosis. Pollen grains free, isopolar; prolate spheroidal, subprolate to prolate (P/E=1.001.56), amb rounded triangular; tricolporate, colpi nearly extending to the poles, colpi with thickened margins, pore ca. 1 m, apocolpia small; exine ca. 0.81.0 m, ?two-layered; sexine scabrate to microfoveolate, foveolae small, <1 m. Dimensions. Equatorial view 9(11)14 x 11(14)17 m (7 grains); polar view 11(14)15 m (3 grains). Holotype. UF 15750-046533-PY03A, EFS W30/3. Occurrence. Courtland Clay Pit and Ochs Clay Pit. Remarks. This species is distinct from Nyssapollenites albertensis Singh 1971, which has a larger and thickened pore (ca. 2.5 m in diameter), thickened colpi margins, and a scabrate (not microfoveolate) exine. Tricolporopollenites triangularis Groot et al. 1961 is psilate. Name derivation. The specific epithet, microfoveolata, is derived from the fine foveolate surface. DISCUSSION The five new pollen species described herein provide additional information on the age of the Dakota Formation

PLATE 2
The scale bar represents 10 m unless otherwise noted. 18 Phimopollenites striolata sp. nov. 13: holotype, single grain, near polar view, high, mid and low focus respectively, UF 19006-046517-A1 >10 , N29/3. 4, 5: single grain, equatorial view, high and low focus respectively, UF 19006-046517-A1, >10 , R22/1. 6: pollen clump, UF 19006-046517-A1, >10 , F14. 7: SEM, UF 19007046517 stub 1, polar view. Scale bar = 2 m. 8: as 7, detail of pollen surface, showing the closely spaced capita giving the striate appearance to the muri. Scale bar = 1 m. 915 Nyssapollenites microfoveolata sp. nov. 911: holotype, UF 15750-046533-PY03A, W30/3, high, mid and low focus respectively. 12, 13: UF 15750-046533PY03A, X42, equatorial view, mid and low focus respectively. 14: SEM UF 15750-046533 stub 3. Scale bar = 2 m. 15: as 14, detail of aperture region and surface ornamentation, showing the microreticulate to foveolate surface pattern. Scale bar = 1 m.

S. Hu, D.M. Jarzen, and D.L. Dilcher: New species of Cenomanian angiosperm pollen, Cenomanian of Minnesota, U.S.A.

Plate23 2

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in south central Minnesota. Several pollen taxa have traditionally been used to determine the age of non-marine sediments within the Dakota Formation (Nichols, 1994). Nyssapollenites microfoveolata sp. nov. is a key taxon for age dating. Nichols (1994) revised the palynostratigraphic zonation for the Upper Cretaceous non-marine sediments in the Rocky Mountain region of the United States, and proposed a scheme based upon a comparison with marine ammonite zones. Nichols (1994) noted that the first occurrence of psilate tricolporate pollen, such as Nyssapollenites microfoveolata sp. nov. and obligate tetrads (e.g. Artiopollis indivisus Agasie 1969), are indicative of a middle Cenomanian to Coniacian age (Nichols and Sweet, 1993; Nichols, 1994). Both Artiopollis indivisus and Nyssapollenites microfoveolata sp. nov. occur in Ochs Clay Pit, and Nyssapollenites microfoveolata sp. nov. occurs in Courtland Clay Pit (Hu, 2006). Based on these occurrences, the sediments in Courtland Clay Pit, Highway 4 Clay Pit and Ochs Clay Pit are considered no older than late Cenomanian in age. The megaspore Balmeisporites glenelgensis Cookson & Dettmann 1958 was found only in the lignite of Ochs Clay pit (Hu, 2006). This species also occurs in the Cenomanian sediments in the Peace River area of northwestern Alberta, Canada (Singh, 1971; 1983). Balmeisporites glenelgensis also occurs in the Sergeant Bluff lignite and the Stone Park lignite in northwestern Iowa and northeastern Nebraska. The age of these lignites was determined to be middle to late Cenomanian (Ravn and Witzke, 1995). Based upon common Balmeisporites glenelgensis, the lignite at the Ochs Clay Pit can be correlated with the Sergeant Bluff lignite and Stone Park lignite. This suggests that the age of the lignite, and the sediments above the lignite, at Ochs Clay Pit are probably middle Cenomanian. By comparison with Singh (1983), Nichols (1994), and Ravn and Witzke (1995), the age of the sediments in Courtland Clay Pit, Highway 4 Clay Pit, and Ochs Clay Pit is middle Cenomanian. The new species described in this paper provide new information on the diversity of angiosperms during the Cenomanian. Although angiosperm diversification was apparently rapid (Lidgard and Crane, 1988; Upchurch and Dilcher, 1990), triporate pollen had not evolved by the middle Cenomanian at the three clay pits studied herein. ACKNOWLEDGMENTS Appreciation is expressed to Karen Kelley and Lynda Schneider for SEM technical assistance. Scott Gooler and Terry Lott are recognized for their help during the 2003 and 2004 field seasons. Financial support was provided by:

Dilcher-Becker Funds, 2003; Evolving Earth Foundation, 2004 grant; Sigma Xi Grant In Aid of Research, 2004; Graduate Student Council of the University of Florida for Travel Grants, 2003 and 2004; Danker Fund, 2004, from the Department of Geological Sciences, University of Florida; and the Deep Time Project, NSF DEB-0090283. Hongshan Wang helped in finding locality data. We thank the Minnesota land owners for allowing collection of samples from their properties. Susan A. Jarzen assisted in manuscript preparation. We are very grateful for the comments and suggestions made by Douglas J. Nichols and Satish K. Srivastava as reviewers, and to James B. Riding for his editorial skills.

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