Revision Of Cenomanian Flora From The Maletín Sandstone Greguš, Josef Kvaček, Jiří Acta Musei Nationalis Pragae Series B 2015 2015-12-31 71 3 - 4 315 364 Heer 1926 [266,585,672,695] Magnoliopsida Magnoliaceae Magnoliaephyllum GBIF Plantae Magnoliales 13 328 Tracheophyta species alternans  Pl. 11, Fig. 1–5; Pl. 12, Fig. 1–7; Pl. 13, Fig. 1–5; Pl. 14, Fig. 1–2 1866  Magnolia alternans HEERin CAPELLINIet HEER, p. 20, pl. 3, fig. 2–4. 1866  Magnolia capellinii HEERin CAPELLINIet HEER, p. 21, pl. 3, fig. 5–6. 1869  Ficus krausiana HEER, p. 15, pl. 5, fig. 3–6. 1869  Ficus mohliana HEER, p. 15, pl. 5, fig. 2. 1869  Daphnophyllum fraasii HEER, p. 17, pl. 6, fig. 1–2. 1869  Daphnophyllum crassinervium HEER, p. 18, pl. 7, fig. 2, pl. 11, fig. 5. 1869  Magnolia speciosa HEER, p. 20, pl. 7, fig. 1, pl. 9, fig. 2, pl. 10, fig. 1–2, pl. 11, fig. 1. 1869  Magnolia amplifolia HEER, p. 21, pl. 8, fig. 1–2, pl. 9, fig. 1. 1869  Juglans crassipes HEER, p. 23, pl. 6, fig. 3. 1874  Magnolia capellinii HEERin CAPELLINIet HEER; Heer, p. 115, pl. 33, fig. 1–4. 1874  Magnolia alternans HEERin CAPELLINIet HEER; Heer, p. 116, pl. 33, fig. 5–6, pl. 34, fig. 4. 1913  Magnolia alternans HEERin CAPELLINIet HEER; Fritel, p. 289, text-fig. 3–8. 1927  Magnoliaephyllum alternans( HEERin CAPELLINIet HEER) SEWARD, p. 120, text-fig. 25. 1999  Dicotylophyllumcf. alternans( HEERin CAPELLINIet HEER) KNOBLOCH, p. 41, pl. 13, fig. 8. L e c t o t y p e: Pl. 11, Fig. 1, designated here; Capellini and Heer 1866, p. 20, pl. 3, fig. 3. It will be necessary to choose a lectotypefrom Oswald Heer’s collection, which were not available to the present study.   Ty p e l o c a l i t y: Tekamah, Nebraska, USA. T y p e h o r i z o n: Dakota Formation, Cenomanian, Upper Cretaceous ( Knowlton 1898, Upchurch and Dilcher 1990). M a t e r i a l s t u d i e d: GBA_2, 3a, 6, 7, 8, 9, 11, 12, 17, 18, 19, 22, 23, 26, 27a–c, 28, 29, 33, 41, 42, 44, 45, 48, 49, 50, 51, 53, 54, 55, 56, 57, 60a, b, 61, 62, 63, 64, 65, 68, 69, 71, 72, 73, 75, 80, 81, 82, 83, 84, 85, 86, 88, 89, 90, 91, 92, 93, 94, 97, 99, 100, 101, 102, 103, 104, 113, 116, 121, 130, 131, 135, 137, 144, 153, 154, 155, 156; GPIT/PL_623, 626, 628, 635, 637, 647, 655, 663, 664, 665, 666, 668, 669, 670, 677, 679, 683, 688, 689, 693, 694, 695, 696, 697, 698, 699, 700, 701, 703, 704, 705, 706, 707, 708, 709, 710, 711, 712, 713, 714, 715, 719, 734, 735, 746, 747, 748, 752, 753, 754, 756; MMT (M)/G_323, 325, 326, 352, 353, 354, 357, 358, 365, 367, 375, 381, 382, 384, 385, 388, 390, 393, 394, 400, 404, 405, 406, 425, 434, 435, 436, 437, 442, 446, 449, 452, 463, 465, 475, 476, 477, 480, 482, 490; MMT (Z)/ G_317, 327, 355, 359, 374, 383, 386, 387, 389, 451; MZM/118_3, 7, 8, 9, 11, 13, 15, 17, 20, 21, 23, 24, 27, 29, 35, 36, 38, 39, 43, 45, 47, 54, 55, 56, 57, 59, 61, 64, 65; NHMW_8, 10, 13, 14, 15, 18, 19, 20, 21, 22, 24, 26; SMNS/650_1, 2, 5, 8, 9, 10, 11, 13, 14, 15; UGV_2, 3, 4, 7, 9, 10, 11, 12, 13, 14, 17, 18, 21; VMO_30.104, 6296, 8088, 8089, 8092, 8093. O c c u r r e n c e i n C z e c h R e p u b l i c: Maletín, Kunštát, Rudka, Březina, Vyšehořovice, Slivenec (Peruc-Korycany Formation). E m e n d e d d i a g n o s i s. Leaves entire-margined medium to large, ovate and longitudinally to widely elliptic. Apex acute to attenuate, base cuneate. Venation pinnate, brochidodromous to eucamptodromous. Occasional shorter and straight intersecondary veins present. Venation of higher order subtle, often not apparent. D e s c r i p t i o n. The suggested lectotype(Pl. 11, Fig. 1) described by Heer (Heer in Capellini and Heer 1866) from Dakota Cretaceous Formation (Tekamah locality, Nebraska). The lectotypeis an elliptical leaf with petiole, without preserved uppermost part of apex. Entire-margined leaf shows a cuneate base. Fragment of apical part tapers into an attenuate apex. Secondary veins brochidodromous, departing from relatively robust midvein at 40° angle. Short and subtle intersecondary veins occur between secondary veins, they arise approximately in the middle of the leaf lamina. The specimens described by Heer (1869)as  D. crassinervium(GPIT/PL_719, Pl. 11, Fig. 5; SMNS/650_9, Pl. 11, Fig. 4),  D. ellipticum(GPIT/PL_734, Pl. 11, Fig. 3) and  Daphnophyllum fraasii(SMNS/650_8, Pl. 11, Fig. 2) are entire-margined, elliptical to wide elliptical leaf impressions with cuneate base. They are widest at one-third or one-half of their length. Acute apex is preserved only in specimen no. GPIT/PL_734. All leaf impressions have fragmentarily preserved petioles. Relatively small leaf impression no. GPIT/PL_734 is only 118 mmlong and 58 mmwide (L/W=2). The second almost preserved leaf impression no. SMNS/650_8 is 182 mmlong and 65 mmwide (L/W=2.8). Secondary veins run from the relatively robust midvein at 35–40° angles. Leaf impression no. GPIT/PL_734 shows in the right side of the leaf lamina an apparent dichotomous branching of secondary veins (Pl. 11, Fig. 3). Similar branching is also apparent to the right upper side of specimen no. GPIT/PL_719 (Pl. 11, Fig. 5). This atypical forking is not apparent in the rest of the specimens. Specimens no. GPIT/PL_719 and GPIT/PL_734 show an apparent suprabasal vein, which runs almost parallel with the cuneate base. On the left half of leaf impression no. GPIT/PL_734 veins of third order run out from the first secondary vein, which are connected to the suprabasal vein. Occasional intersecondary veins are present. Leaf impression no. SMNS/650_5 (Pl. 12, Fig. 1), described by Heer (1869)as  Ficus mohlianais a 217 mmlong and 52 mmwide, entire-margined impression of longitudinally elliptic leaf, with narrow cuneate base and attenuate apex. This leaf impression is the widest in its middle part. Length to width ratio is 4.2. A well pronounced midvein is remarkably narrowed in the direction of the apical part. Poorly preserved secondary veins run out from the primary vein at an angle of 40°. Other specimens described by Heer (1869)as  Ficus krausianaare two impressions of entire-margined elliptical leaves. These impressions are widest in their lower half. The first impression (SMNS/ 650_2, Pl. 12, Fig. 2) has an acute apex. A small part of the missing base suggests a cuneate shape. Estimated length of the impression is 200 mm. Width is 53 mm; length to width ratio is 3.7. The primary midvein is relatively wide. Poorly preserved secondary veins depart from midvein at an angle of 35–40°. Near the margin of the leaf lamina, some veins show brochidodromous loops. Another, better preserved specimen (SMNS/650_1, Pl. 12, Fig. 3) is an approximately 175 mmlong and 46 mmwide leaf impression, with a cuneate base and oblong attenuate apex. Length to width ratio is 3.8. Small parts of the base and apex are not preserved. Brochidodromous secondary veins depart the midvein at a 40° angle. Leaf impression (GPIT/PL_746, Pl. 12, Fig. 4), described by Heer (1869)as  Juglans crassipesis entire-margined, elliptic with a cuneate base and not preserved apical part. It is widest in its middle part. Estimated length of the leaf impression is 165 mm, and width is 63 mm; length to width ratio is 2.6. The leaf shows a cuneate base with relatively long ( 50 mm) and wide petiole, and missing apex. Secondary venation is brochidodromous. Intersecondary veins are present.  The leaf impressions described by Heer (1869)as  Magnolia speciosais entire-margined, widely elliptical with cuneate bases and attenuate apices ( GPIT/ PL_735, Pl. 12, Fig. 7; GPIT/ PL_747, Pl. 12, Fig. 6; GPIT/ PL_752, Pl. 13, Fig. 1; GPIT/ PL_754, Pl. 13, Fig. 2; GPIT/ PL_756, Pl. 12, Fig. 5). The largest leaf impression no. GPIT/ PL_756 is 250 mmlong and 92 mmwide ( L/ W=2.7). Other leaf impressions are approximately 20 mmlong and 8 mmwide ( L/ W=2.5). Primary venation pinnate; secondary venation of brochidodromous typedeparting from relatively wide midvein at a 45–50° angle. All impressions have at least partially preserved petiole. Specimen no. GPIT/ PL_754 has a well pronounced long petiole ( 41 mm). Leaf impressions of magnolioid-type described by Heer (1869)as  Magnolia amplifolia(GPIT/PL_753, Pl. 13, Fig. 4; SMNS/650_13, Pl. 13, Fig. 3) are entire-margined, widely elliptic to obovate with cuneate base. The leaves are widest in the first third of their length. Apical parts are not preserved; judging by the convergent margins of leaf lamina, they were probably attenuate. Specimen (GPIT/PL_753) could be originally up to 220 mmlong and 106 mmwide (L/W=2.1). Secondary venation of brochidodromous to eucamptodromous typewith veins depart a relatively wide midvein at a 40° angle. Some veins do not reach the margin, and end at approximately 2/3 of length of other veins. Intersecondary veins are sometimes present. The newly discovered counterpart (GPIT/PL_666, Pl. 13, Fig. 5) to Heer's (1869)original (GPIT/PL_753, Pl. 13, Fig. 4) shows a clearly cuneate base and roughly sinuate margin. Other material from Starý Maletín consists of approximately 200 specimensof complete and nearly complete leaf impressions, and 25 fragments. All specimens are entire-margined, elliptic and widely elliptic to ovate. The best preserved specimens are 115–280 mmlong and 42–150 mmwide (average: L= 202 mm, W= 84 mm). Length to width ratio is 1.5–3.7 (average: 2.5). Venation is in most cases well preserved. Venation is pinnate, with secondary veins showing brochidodromous loops. Intersecondary veins are present. A few specimens have venation poorly preserved or missing. However, a number of specimens also show third order venation, which is percurrent (GBA_18, Pl. 14, Fig. 1a, b; GBA_29, Pl. 14, Fig. 2a, b). Most leaf impressions have preserved at least fragments of the petiole. Specimens no. GPIT/PL_746 (Pl. 12, Fig. 4) and no. GPIT/PL_754 (Pl. 13, Fig. 2) show very long petiole. D i s c u s s i o n. Heer ( Capellini and Heer 1866, Heer 1869) described eight new species of magnolioid-type leaf impressions, which he classified under the genera  Daphnophyllum,  Ficus,  Juglansand  Magnolia. Leaf impressions described by Heer are entire-margined, with similar elliptical shape, with cuneate base and attenuate apex. During his life, the leaf impressions were freshly collected, so the venation was more apparent. Despite this, he described only secondary venation. Today, higher order venation is seen on only a few specimens. Venation of these leaf impressions has very similar or identical architecture. Because Heer had only a few specimens, he classified the species according to their variable secondary characters, such as a moderately elongated apical part. In this case, we studied and compared almost 200 specimens, and concluded that among these leaf impressions, there is great variability, with smooth transitions in shape, size (including length to width ratio) and venation. This is confirmed by Fritel (1913), who described and illustrated the modern species  Magnolia grandiflora LINNAEUS, which has clear variability in elliptical leaf, with transition to ovate leaf, but also obovate leaf. With regards to great variability, Fritel (1913)synonymised the species  Ficus krausiana HEER,  Daphnophyllum crassinervium HEER,  D. ellipticum HEER,  D. fraasii HEER,  Juglans crassipes HEER,  Magnolia amplifolia HEER,  M. capellinii HEER,  M. speciosa HEERto the species  Magnolia alternans. Knobloch (1999)is also inclined to regard these as the same species. Knobloch classifies these impressions to the artificial genus  Dicotylophyllum SAPORTA. Dueto the character of these impressions from Maletín, it is not possible to determine systematic position in recent botanical classification. Leaf impressions from the NebraskaCretaceous described by Heer ( Capellini and Heer 1866) as  Magnolia alternansare, with their shape and simple typeof venation, morphologically identical to the Cenomanian species  Magnolia amplifolia, which Heer (1869)described from the Bohemian Cretaceous Basin.   Fossilleaves of the magnolioid-type, with preserved epidermal anatomy were described by Upchurch and Dilcher (1990)from the SE Nebraska Cretaceous( Rose Creek, USA). Forthis fossil material, a new genus  Pandemophyllum UPCHURCHet DILCHERwas established, with the typespecies  Pandemophyllum kvacekii UPCHURCHet DILCHER.  Upchurch and Dilcher (1990)remark in their publication that there is a possible relationship between the new genus  Pandemophyllumand Heer's (1869)genera. As Z. Kvaček (1983)mentions,  Magnolia amplifolia,  Daphnophyllum crassinerviumand  D. ellipticum(in this paper classified in genus  Magnoliaephyllum) are very similar in their anatomical features to  Myrtophyllum geinitzii HEER, which in this paper we describe as  Myrtoidea geinitzii( HEER) J. KVAČEKet GREGUŠ comb. nov.  Myrtoidea geinitziiis defined by an intramarginal vein, which is missing in leaves of  M. alternans. High-quality and complete leaf impressions of  M. alternansare very typical of the Maletín plant assemblage, and quite numerous compared to other species. This situation is, beside frequent occurrence of this leaf typein the Maletín sandstone, caused by human factor: during intensive quarrying of Maletín sandstone, it was easy for quarry workers to collect fossil impressions, and naturally, they only selected the best pieces. As Pek and Mikuláš (1997)mention, leaves of  M. alternansare curved, waved and have cross grooves and ridges, and even structures similar to recent insect burrows, although this observation is not new – Heer (1869)observed these same structures (“Insektengallen”) over a century earlier. [144,653,1812,1835] T United States of America Tekamah 13 328 1 Nebraska GPIT, PL, L, W 14 329 1 United States of America For 15 330 Fossil Rose Creek 14 329 1 Nebraska