954 American Journal of Botany 95(8): 954?973. 2008. The plant diversity in the tropics of South America is one of the greatest in the world; nevertheless, the evolutionary patterns of the neotropical fl oras across geologic time, the climatic con- ditions in which they appeared, and the mechanisms that ex- plain such variation are still poorly known ( Burnham and Johnson, 2004 ). The record of pollen and spore diversity from the Paleogene to early Neogene in the neotropics shows that plant diversity in the tropics has been variable through time and seems to correlate with long-term global climatic changes ( Jaramillo et al., 2006 ). The Paleocene Cerrej ? n fl ora of northern Colombia has been investigated in the last couple of years with more than 1500 specimens (mostly leaves) collected and about 57 morphotypes identifi ed ( Wing et al., 2004 ; Herrera et al., 2005 ). The Cerrej ? n fl ora, according to leaf margin and area analysis, corresponded to a rain forest that grew in coastal areas ( Herrera, 2004 ; Herrera et al., 2005 ). Here, we studied one of the most common morphotypes of the Cerrej ? n fl ora, that is related to Menispermaceae. The study of this morphotype can give us clues about the evolution of neotropical forests because lianas, which are common in Menispermaceae, are a key com- ponent of multistratifi ed forests ( Gentry, 1991 ). Menispermaceae is a diverse family in the order Ranuncula- les ( Cronquist, 1981 ; APG II, 2003 ) whose species are distrib- uted mainly in tropical lowlands and grow in low altitudes ( Kessler, 1993 ). The family is primarily composed of herba- ceous or woody climbers with dioecious and small inconspicu- ous fl owers. The fruits are drupes, probably dispersed by birds ( Kessler, 1993 ), and the common name of the family, the moon- seed family, is derived from their curved endocarp and embryo. The basic type of pollen is tricolpate, lacking operculum and costae, with perreticulate tectum, a collumellate infratectal layer, and a granular endexine ( Thanikaimoni, 1984 ). Extant Menispermaceae include ~500 species grouped into 70 genera ( Ortiz et al., 2007 ). The classifi cation is principally based upon fruit and seed characters, and the most important differences between the genera concern the endocarp ( Diels, 1910 ; Kessler, 1993 ). Several molecular phylogenetic analyses have confi rmed the monophyly of the family ( Hoot et al., 1999 ; APG II, 2003 ; Ortiz et al., 2007 ), but the intrafamiliar classifi - cation has been controversial ( Barneby, 1970 ; Forman, 1986 ; Kessler, 1993 ; Ortiz et al., 2007 ). Different intrafamiliar clas- sifi cations have been proposed ( Miers, 1851 ; Hooker and Thomson, 1855; Bentham and Hooker, 1862 ; Miers, 1864 ; Prantl, 1888 ; Diels, 1910 ; Kessler, 1993 ); however, the tradi- tional and most accepted one ( Diels, 1910 ) divides the family into eight tribes based on a combination of characters from the seeds, fl owers, and endocarps. The only phylogenetic hypothe- sis for Menispermaceae to date, based on molecular data ( Ortiz et al., 2007 ), provides evidence for the monophyly of four of the eight tribes circumscribed by Diels and shows that the four remaining tribes are polyphyletic. Modern biogeographic distribution of Menispermaceae shows predominance in West Gondwanaland ( Raven and Axelrod, 1974 ; Gentry, 1982 ). However, the cradle of the family is unknown, and different hypotheses have been proposed. Most authors think that the group has a Laurasian origin, as for many other Ranunculales, because the most basal species of the fam- ily occurs in the West Pacifi c and the fossil records to date come mainly from the northern hemisphere ( Diels, 1910 ; Takhtajan, 1 Manuscript received 16 July 2007; revision accepted 2 June 2008. The authors thank two anonymous reviewers, the Biostratigraphic Team at the Colombian Petroleum Institute, and the Department of Geology EAFIT University. They deeply thank F. Chavez, C. Montes, S Moodley, and the geology team at Carbones del Cerrej ? n LLC, S. Wing for help in collecting and morphotyping fossil leaves, J. Betancur and F. Gonzalez at Herbario Nacional Colombiano, R. Fonnegra, J. Rold ? n, and R. Callejas at Herbario de la Universidad de Antioquia. Thanks to N. Atkins for improving the grammar. C.J. thanks M. I. Barreto for continuous support and source of creative ideas. This work was supported by Carbones del Cerrej ? n, the Smithsonian Paleobiology Endowment Fund, the Unrestricted Endowments SI Grants, NSF (grant DEB-0733725) and the Colombian Petroleum Institute. 4 Author for correspondence (e-mail: jaramilloc@si.edu) doi:10.3732/ajb.2007216 MENISPERMACEAE FROM THE CERREJ ? N FORMATION, MIDDLE TO LATE PALEOCENE, COLOMBIA 1 Gabriela Doria, 2 Carlos A. Jaramillo, 2,4 and Fabiany Herrera 2,3 2 Center for Tropical Paleoecology and Archeology, Smithsonian Tropical Research Institution, Balboa, Ancon, Panama; and 3 Florida Museum of Natural History, Gainesville, Florida, 32611-2710 USA The origin and processes creating the high diversity of plant species in neotropical rain forests and their fl oristic composition and multistratitifi ed forest structure are still uncertain. Here, we studied one of the most common leaf morphotypes of the Cerrej ? n fl ora (middle-late Paleocene, ca. 60-58 Ma), Guajira, Colombia, that contains one of the oldest records of neotropical rain forest fl oras. Fifty-seven leaf specimens were carefully examined with a focus on general morphology, venation patterns, and cuticular characteristics. The analysis allowed us to recognize four new species that were assigned to the fossil-leaf genus Menispermites on the basis of an ovate leaf shape with cordate to truncate bases, actinodromous primary venation, brochidodromous secondary venation, percurrent tertiary venation, regular polygonal reticulate fourth and fi fth venation, well-developed poly- gonal areoles, entire margin, and the presence of a fi mbrial vein. This set of characters suggests a possible affi nity with the pantropical angiosperm family Menispermaceae. The predominantly climbing habit of this family suggests that the Cerrej ? n Paleocene tropical rain forest was already multistratifi ed. These fi ndings represent the earliest record for the family in northern South America. Key words: Colombia; leaf morphology; lower eudicots; Menispermaceae; paleobotany; Ranunculales. 955August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia 1969 ; Raven and Axelrod, 1974 ; Wolfe, 1977 ; Kessler, 1993 ). In contrast, Thanikaimoni (1984) hypothesized that because the primitive characters of Menispermaceae are found in African species, the family may have originated in the lowlands of Af- rica during the Cretaceous, when South America, Africa, Mad- agascar, and India were close to one another. The recently published phylogeny ( Ortiz et al., 2007 ) recovers two major clades within Menispermaceae, with Tinomiscium petiolare , an Indo-Malaysian centered taxon, as sister to the remainder of the family; however, no biogeographic interpretations have been drawn from this basis. Although Menispermaceae has an extensive fossil record that includes both extant and extinct genera ( Tables 1 ? 3 ), the present study reports and describes for the fi rst time fossil leaves from northern South America. The oldest record identifi ed with some confi dence is an endocarp from the Turonian (Upper Cre- taceous, 91 Ma) of central Europe ( Knobloch and Mai, 1984 , 1986 ), but fossil leaves found in North America ( Fontaine, 1889 ; Berry, 1916 ; Hollick, 1927 ; Seward, 1927 ; Hollick and Martin, 1930 ; Bell, 1956 ; Doyle and Hickey, 1976 ) and Asia ( Takhtajan, 1974 ; Spicer et al., 2002 ) suggest that the family can be traced to the Early Cretaceous. Fossil pollen has been widely reported since the Early Cretaceous in many different geographic areas ( Table 3 ). The tropical fossil record of Menispermaceae is very sparse and consists of endocarps ( Chesters, 1957 ) and leaves ( Jacobs and Kabuye, 1987 ) from the Miocene of Kenya, leaves from the Eocene of Borneo ( Andrews, 1970 ), and pollen from the Mio- cene of Tunisia ( M ? on and Tayech, 1986 ) and Taiwan ( Huang, 1980 ; Song et al., 2004 ) and Pleistocene of Ethiopia ( Bonnefi lle et al., 1987 ). In South America the fossil record of the family is reduced to fossil leaves from the Paleocene of Argentina ( Iglesias et al., 2007 ) and the Eocene ( De Lima and Salard-Cheboldaeff, 1981 ) Table 1. Seeds and endocarps of Menispermaceae in the fossil record. Age Area Locality References Paleocene Europe Czech Republic Knobloch, 1971 Eocene Europe Belgium England Fairon-Demaret and Smith, 2002 Chandler, 1925, 1962, 1963, 1964, 1978; Reid and Chandler, 1933; Scott, 1954; Eyde, 1970 Germany Collinson, 1988 France Jacques and De Franceschi, 2005 North America Oregon Scott, 1954; Manchester, 1994 Virginia Tiffney, 1999 Washington Pigg and Wehr, 2002 Oligocene Europe Czech Republic Kvacek and Walther, 1998 Germany Kirchheimer, 1957 Russia Szafer, 1938; Dorofeev, 1963; Tralau, 1963; Takhtajan, 1974 Miocene Africa Kenya Chesters, 1957 Asia Russia Dorofeev, 1969; Takhtajan, 1974 Pliocene Europe Netherlands Reid and Reid, 1910; Zagwijn, 1963 Neogene Australia New South Wales Mueller, 1876; Andrews, 1970 Table 2. Leaves of Menispermaceae in the fossil record. Age Area Locality References Early Cretaceous Asia Russia Takhtajan, 1974 ; Spicer et al., 2002 North America Alaska Hollick and Martin, 1930 British Columbia Hollick, 1927 ; Bell, 1956 Louisiana Berry, 1916 Virginia Fontaine, 1889 ; Doyle and Hickey, 1976 Greenland Seward, 1927 Late Cretaceous Asia Russia Kryshtofovich and Baikovskaia, 1960 ; Budantsev, 1968 ; Takhtajan, 1974 Europe Czech Republic Velenovsky, 1889 ; Hughes, 1976 North America Alaska Hollick and Martin, 1930 Alberta Bell, 1962 British Columbia Bell, 1957 New Mexico Robison et al., 1982 Wyoming Dorf, 1942 ; Van Boskirk, 1998 Paleocene Asia Russia Ablaev, 1971 ; Takhtajan, 1974 North America Alaska Wolfe, 1966 , 1972; Hickey, 1977 British Columbia Wolfe, 1966 , 1972; Hickey, 1977 Colorado Barclay et al., 2003 Greenland Wolfe, 1966 , 1972; Hickey, 1977 High Arctic McIver and Basinger, 1999 Louisiana Berry, 1922a North Dakota Hickey, 1977 Rocky Mountains and Great Plains Brown, 1962 ; R ? ffl e, 1968 South America Argentina Iglesias, et al., 2007 Eocene Asia Borneo Andrews, 1970 North America Diels, 1910 Alaska Wolfe, 1972, 1977 California Potbury, 1935 ; MacGinitie, 1941 Kentucky Berry, 1922b Sierra Nevada Central Wolfe, 1968 Utah MacGinitie, 1969 Washington Wolfe, 1968, 1977 ; Wolfe and Tanai, 1978 Wyoming Hickey, 1977 ; Wing et al., 1995 ; Wilf, 2000 South America Brazil De Lima and Salard-Cheboldaeff, 1981 Oligocene Asia Russia Iljinskaja, 1972 Miocene Africa Kenya Jacobs and Kabuye, 1987 Asia Japan Ozaki, 1991 Russia Takhtajan, 1974 North America Alaska Wolfe, 1966 Nebraska MacGinitie, 1962 Pliocene Europe France Hollick, 1927 South America Brazil Dolianiti, 1949 ; Mello, Sant ? Anna, and Bergqvist, 2000 956 American Journal of Botany [Vol. 95 (11 ? 1 ? N, 72 ? 45 ? W; Fig. 1 ). The Cerrej ? n coal mine is bounded to the north by the Oca Fault, to the west by the eastern foothills of the Sierra Nevada de Santa Marta, and to the east by northwest-verging thrust faults of the Perij ? Range ( Jaramillo et al., 2007 ). Lithofacies, sedimentary structures, and palyno- logical associations in the ~700-m-thick Paleocene succession in the Cerrej ? n coal mine suggest that a mixed platform was covered and buried by coal- bearing siliciclastic strata, which accumulated on coastal plains ( Jaramillo et al., 2007 ). Sedimentation was characterized by deltas toward the base of the formation and continental environments dominated by channel systems, fl oodplains, swamps, and lagoons upward ( Jaramillo et al., 2007 ). The Cerrej ? n Formation has been dated using mollusks ( Etayo-Serna, 1979 ) and pollen ( Van der Kaars, 1983 ; Bayona et al., 2004 ; Jaramillo et al., 2007 ) as middle to late Paleocene (~60 ? 58 Ma), following the time scale of Gradstein et al. (2004). The fossil remains consist of whole or fragmented leaf impressions and compressions. Sixty-eight specimens were collected from the morphotype in- formally named CJ6. Fifty-seven specimens were examined: four specimens from locality 0315, 24 from 0317, 21 from 0318, four from 0319, three from 0322, and one from 0323 ( Fig. 2 ). The CJ6 morphotype was described following and Pliocene of Brazil ( Dolianiti, 1949 ; Mello et al., 2000 ), one liana fossil wood probably related to Ranunculales from the Miocene of Argentina ( Lutz and Mart ? nez, 2007 ), and fossil pollen from the Oligocene to recent of British Guiana ( Van der Hammen and Wijmstra, 1964 ) and the Miocene of Suriname ( Amstelveen, 1971 ). Here, we studied one of the most common morphotypes of the Cerrej ? n fl ora. We describe it in detail and compare it with modern fl oras and fossil taxa to assess its phylogenetic affi nity. MATERIALS AND METHODS Fossil leaves were collected from six localities (0315, 0317, 0318, 0319, 0322 and 0323) at the Cerrej ? n Formation, an outcrop at El Cerrej ? n coal mine on the northern side of the Cesar-Rancheria basin, Guajira peninsula, Colombia Table 3. Pollen of Menispermaceae in the fossil record Age Area Locality References Lower Cretceous Asia Russia Papulov and Ediger, 1973 ; Srivastava, 1978 Upper Cretaceous Asia Russia Zaklinskaya, 1953 ; Boytsova and Pokrovskaya, 1954 ; Agranovskaya et al., 1960 ; Sedova, 1960 ; Samsonov, 1964 ; Andreeva et al., 1966 ; Grigoreva, 1966 ; Fradkina, 1967 ; Markova et al., 1967 ; Golbert and Markova, 1968 ; Chlonova, 1971 ; Pogodayeva, 1973 ; Laukhin and Kulkova, 1974 ; Blyakhova, 1976 ; Fradkina and Kiseleva, 1976 ; Aleksandrova et al., 1977 ; Boytsova et al., 1980 ; Samoilovich, 1980 ; Mikhelis, 1982 ; Levina et al., 1983 ; Markova and Skuratenko, 1983 ; Kalmeneva et al., 1986 ; Kruchinina et al., 1990 North America Delaware, Maryland Graham, 1964 Paleocene Asia Russia Pokrovskaya, 1960b ; Blyakhova, 1966 ; Pelipenko, 1966 ; Brattseva, 1969 ; Grinenko and Kiseleva, 1971 ; Kulkova and Laukina, 1975 ; Fradkina and Kiseleva, 1976 ; Tomskaya, 1981 ; Mikhelis, 1982 ; Zharikova et al., 1982 ; Teslenko, 1990 ; Fradkina, 1996 Eocene Asia Russia Zaklinskaya, 1953 ; Agranovskaya et al., 1960 ; Pokrovskaya, 1960a ; Andreeva et al., 1966 ; Barbashinova, 1966 ; Shakhmundes, 1966 ; Krayenva et al., 1967 ; Barbashinova, 1968 ; Rzhanikova, 1968 ; Brattseva, 1969 ; Barbashinova, 1971 ; Pulatova, 1973 ; Bykadorov et al., 1974 ; Oleynik and Oleynik, 1979 ; Zosimovich and Mikhelis, 1979 ; Korallova, 1981 ; Akhmeteev et al., 1996 ; Davidzon et al., 1996 ; Turdukulov and Fortuna, 1996 Age Area Locality References Europe France Gruas-Cavagnetto et al., 1980 North America Wyoming Leopold and MacGinitie, 1972 Oligocene Asia Russia Boytsova and Pokrovskaya, 1954, 1956 ; Blom, 1960 ; Andreeva et al., 1966 South America Guyana Van der Hammen and Wijmstra, 1964 Miocene Africa Tunisia M ? on and Tayech, 1986 Asia Taiwan Huang, 1980 ; Song et al., 2004 Asia Russia Shchekina, 1954 ; Pokrovskaya, 1956 ; Sedova, 1956 ; Dzhabarova, 1959 ; Manykin, 1960 ; Mchedlishvili, 1963 ; Andreeva et al., 1966 ; Ramishvili, 1969 ; Leye, 1980 Europe Hungary Istvan, 1964 France Bessedik, 1982 ; Bessedik et al., 1984 ; M ? on et al., 1989 Spain Durand-Delga et al., 1989 South America Surinam Amstelveen, 1971 Pliocene Asia Russia Mchedlishvili, 1963 Europe France Suc, 1978, 1980, 1981 ; Michaux and Suc, 1981 Portugal Diniz, 1984 Romania Drivaliari et al., 1999 Netherlands Zagwijn, 1963 Pleistocene Africa Ethiopia Bonnefi lle et al., 1987 Europe France Suc, 1978, 1980, 1981 ; Michaux and Suc, 1981 Polland Stachurska et al., 1973 Quaternary Asia China Li and Wu, 1978 957August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia actinodromous basal; lateral primaries ascending toward apex; secondaries brochidodromous, one to several ascending sec- ondaries merging from the midvein; tertiary veins alternate percurrent, opposite percurrent or mixed alternate-opposite percurrent; higher-order veins regular polygonal reticulate; po- lygonal areoles well developed; and ultimate marginal venation fi mbrial. Systematic description ? Family ? Menispermaceae Genus ? Menispermites Lesquereux. Species ? Menispermites cerrejonensis Doria, Jaramillo & Herrera sp. nov. Diagnosis ? Simple notophyllous to mesophyl- lous leaves. Ovate symmetrical lamina, 11.0 (8.5 ? 14) cm (4 specimens measured), 8 (5.4 ? 11.3) cm wide (8 specimens mea- sured). Base truncate to slightly cordate, base angle obtuse; apex acuminate, apex angle acute. Margin entire and unlobed. First vein category actinodromous basal, 5 ? 9 veins departing from the petiole insertion; central primary slightly wider than lateral primaries; fi rst pair of lateral primaries extending about two-thirds the distance to apex into apical half of lamina. Sec- ondary venation brochidodromous; one or few pairs of ascend- ing secondaries from the midrib, with the lowermost pair merging above the middle of the blade; central secondaries al- ternate or subopposite. Simple agrophic veins. Weak intersec- ondaries. Third vein category alternate percurrent. Fourth vein category regular polygonal reticulate, sometimes alternate per- current; fi fth vein category regular polygonal reticulate. Are- oles well developed, 4 ? 5 sided; free-ending veinlets absent, unbranched, or 1-branched. Fimbrial vein conspicuous. Holotype ? ING-CJ6 ? 0145 ? 0318, Fig. 3A, B . Paratypes ? ING-CJ6 ? 0907 ? 0318, Fig. 3C, D ; ING- CJ6 ? 0137 ? 0318. Type locality ? 0318, Pit Tabaco High Deep, Cerrej ? n coal mines, northern Cesar-Rancheria basin, Guajira peninsula, northern Colombia, located at 11 ? 13 ? N, 72 ? 55 ? W. Horizon ? Upper part of the Cerrej ? n Formation. Between coal seams 160 and 170. the parameters of the Leaf Architecture Working Group (LAWG, 1999), which focus on general morphology and venation patterns. Terminology of fossil taxa previously described was modifi ed following LAWG terminology when neces- sary, to make the comparison easier. Specimens were analyzed using a stereo- microscope and photographed using a Nikon (Washington, D.C., USA) D70s digital camera with low-angle lighting to reveal venation details. The drawings of veins were traced from the photographs. Cuticles were prepared using a modifi ed Schulze (1855) method. The better-preserved cuticles were isolated from the matrix with fi ne needles and treated with a saturated solution of potas- sium chlorate in 65% nitric acid for 1 ? 24 h. The cuticles were rinsed three times with distilled water when they turned clear, upon which they were submerged in 10% NaOH for 15 min. Finally, cuticles were rinsed again, and the two sur- faces were separated, one from the other, with needles when possible. The cu- ticles were mounted in glycerol on glass slides. Anatomical features were photographed using a Nikon digital camera DXM1200 in a Nikon Eclipse 80i microscope. All specimens are housed at the paleobotanical collections of IN- GEOMINAS (Instituto Colombiano de Geolog ? a y Miner ? a) in Bogota, Colom- bia. The leaves were compared with living and fossil taxa through consultation of the literature ( Troupin, 1962 ; Barneby, 1970 ; Krukoff and Barneby, 1970a , b , 1974 ; Barneby and Krukoff, 1971 ; Ott, 1997 ), observation of specimens from herbarium collections at Herbario Nacional Colombiano (COL) and Her- bario de la Universidad de Antioquia (HUA), and direct observation of extant species in the fi eld. Several angiosperm families with similar gross morphology (i.e., Aristolochiaceae, Dioscoreaceae, Euphorbiaceae, Malvaceae, Passifl o- raceae), were examined to establish the natural affi nities of the morphotype. Exemplars of 12 extant neotropical genera ( Abuta Aubl., Anomospermum Miers., Borismene Barneby, Chondrodendron Spreng., Cissampelos L., Cura- rea Barneby & Krukoff, Disciphania Eichl., Hyperbaena Miers ex Benth., Odontocarya Miers, Orthomene Barneby & Krukoff, Sciadotenia Miers, Telitoxicum Moldenke) were examined in the herbaria. Descriptions and illus- trations by Ott (1997) were used as the main source for the neotropical taxa not found in the visited herbaria (e.g., Elephantomene Barneby & Krukoff). Com- parisons including non-neotropical taxa were made with photographs from the C. V. Starr Virtual Herbarium, Neotropical Herbarium Specimens (http:// sciweb.nybg.org/VirtualHerbarium.asp). RESULTS Four different kinds of leaves constitute the CJ6 morpho- type. They are assigned here to four new species of Menisper- mites Lesquereux. Common characters for the leaf forms are as follows: blades simple; lamina ovate symmetrical; base cordate to truncate; apex acute; margin entire, unlobed; fi rst venation Fig. 1. Map of location of the Cerrej ? n coal mine, northern Colombia. 958 American Journal of Botany [Vol. 95 Fig. 2. Stratigraphical section of the Cerrej ? n Formation. Numbers indicate fossil plant localities. (Adapted from Bayona et al., 2004 ). 959August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia lated to a single taxonomic group on the basis of these charac- ters. In many species of Menispermaceae, different types of indumenta are present; in the fossil cuticles, however, indu- ment was not detected. Species ? Menispermites cordatus Doria, Jaramillo & Her- rera sp. nov. Diagnosis ? Simple mesophyllous leaves. Ovate symmetri- cal lamina, 14.1 (8.5 ? 24) cm long (11 specimens measured), 12.63 (6.7 ? 22) cm wide (10 specimens measured). Base cor- date to deeply cordate, base angle obtuse; apex acuminate when preserved, apex angle acute. Margin entire and unlobed. First vein category actinodromous basal; 5 ? 7 veins from the base; central and lateral primaries homogeneous. Secondary venation brochidodromous; one or few ascending secondaries from the midrib, with the lowermost pair merging above the middle of the blade; central secondaries alternate to subopposite. Main fi rst and second veins concave. Compound agrophic veins pres- ent. Third vein category mostly opposite percurrent, but alter- nate percurrent and mixed opposite-alternate types also present. Fourth vein category regular polygonal reticulate. Fifth vein category regular polygonal reticulate. Areolation well devel- oped, areoles 4 ? 5 sided. Free-ending veinlets unbranched to 1 ? 2 branched. Fimbrial vein conspicuous. Leaf texture chartaceous. Holotype ? ING-CJ6 ? 0128 ? 0317, Fig. 6A, B Paratypes ? ING-CJ6 ? 0154 ? 0317, Fig. 6C, D Type locality ? 0317, Pit Tabaco, Cerrej ? n coal mines, north- ern side of the Cesar-Rancheria basin, Guajira peninsula, north- ern Colombia, located at 11 ? 14 ? N. 72 ? 57 ? W. Horizon ? Upper part of the Cerrej ? n Formation. Between coal seams 100 and 102. Derivatio nominis ? The specifi c epithet refers to the charac- teristic cordate base shape. Studied material ? ING-CJ6 ? 0120 ? 0317, ING-CJ6 ? 0121 ? 0317, ING-CJ6 ? 0125 to ING-CJ6 ? 0134 ? 0317, ING-CJ6 ? 0155 ? 0317, ING-CJ6 ? 0156 ? 0317, ING-CJ6 ? 0158 ? 0317 to ING-CJ6 ? 0160 ? 0317, ING-CJ6 ? 0162 ? 0317 to ING-CJ6 ? 0164 ? 0317, ING-CJ6 ? 0167 ? 0317, ING-CJ6 ? 0176 ? 0315, ING-CJ6 ? 0177 ? 0317, ING-CJ6 ? 0179 ? 0315. Description ? Specimens assigned to this morphotaxon were found at localities 0315 and 0317. It is the most common form in the localities, with 25 specimens. Leaf shape is ovate with the base wide and cordate. Marginal petiole has been detected in four specimens (ING-CJ6 ? 0121 ? 0317, ING-CJ6 ? 0156 ? 0317, ING-CJ6 ? 0159 ? 0317, ING-CJ6 ? 0132 ? 0317), which showed it as either stout and straight or thin and curved ( Fig. 6C, D ). Central primary is slightly curved toward the apex, and central secondaries are subopposite or alternate. First pair of lateral primaries merge at an acute to perpendicu- lar (40 ? 90 ? ) angle from the central, reaching 2/3 ? 1/4 the dis- tance to the apex, looping with the central secondaries. The second pair of lateral primaries and subsequent pairs emerge at a wide acute to obtuse (70 ? 120 ? ) angle; all primaries with concave ascending course toward the apex. First and second pairs of lateral primaries branching distally at 1/2 the length. Several tertiaries merging distally from the exmedial pairs of primaries (simple or compound agrophics) at an obtuse to per- pendicular angle, joining superadjacent vein at an acute angle. In many of the specimens, conspicuous opposite percurrent third venation occurs, with the angles of the tertiaries becom- ing more obtuse with respect to the midrib, away from the axis Derivatio nominis ? The specifi c epithet refers to the type lo- cality at the Cerrej ? n Formation. Studied material ? ING - CJ6 ? 0135 ? 0318 to ING - CJ6 ? 0154 ? 0318, ING - CJ6 ? 0606 ? 0318, ING-CJ6 ? 0908 ? 0318. Description ? Specimens assigned to this morphotaxon were found in locality 0318. Leaf shape is ovate with the base rounded, truncate, or slightly cordate. Length to width ratio varies from 1 : 1 ? 2 : 1. The apex is rarely preserved. Central primary with straight course; central secondaries alternate, sometimes subopposite. The fi rst pair of lateral primaries merge at a narrow acute angle (20 ? 50 ? ) from the midrib, with a moderately curved course upward reaching to the distance to the apex, branching exmedially at the length of the vein and forming loops with the central secondaries; second pair of lat- eral primaries merging at a wide acute angle (40 ? 90 ? ), slightly curved reaching one-third the distance to apex, looping with lateral secondaries, with several secondaries merging distally (simple agrophic veins), each one joining superadjacent at an acute angle ( Fig. 3 ). Third venation mostly alternate percurrent ( Fig. 4 ). Shape of epidermal cells is irregular, with sinuous anticlinal walls at the areoles, and rectangular, with straight anticlinal walls at the veins ( Fig. 5A ); anomocytic stomata, with four cells adjacent to the guard cells not differentiated in any way from the normal epidermal cells ( Fig. 5B, C ), and a diffuse pattern of distribution, probably only present at the abaxial surface. Leaf texture membranaceous, cuticle well preserved. Comparisons ? Menispermites cerrejonensis differs from M. guajiraensis sp. nov. and M. horizontalis sp. nov. in having ab- sent or few central secondary veins from the midrib, with the lowermost vein pair originating above the middle of the blade, instead of having several pairs of central secondaries with the lowermost pair originating below the middle of the blade. Menispermites cordatus sp. nov. also presents similar venation patterns in the central secondaries; however, M. cordatus can be distinguished from M. cerrejonensis by having deeply cor- date bases vs. rounded, truncate, or slightly cordate, concave primaries vs. straight or slightly curved primaries, and tertiary venation opposite percurrent vs. alternate percurrent. Chondrodendron brasiliense Dolianiti from the Pliocene of Brazil ( Dolianiti, 1949 ) is similar in base shape and in the pat- terns of primaries and central secondaries. However, in C. brasiliense the uppermost secondaries loop near the margin, fusing with the fi mbrial vein, whereas in M. cerrejonensis they loop far from the margin. Moreover, third venation is opposite percurrent in Chondrodendron brasiliense , not alternate per- current as in M. cerrejonensis . Menispermites cerrejonensis presents several characteristics present in the neotropical Menispermaceae genera Curarea Barneby & Krukoff (clade II, Tiliacorae, Ortiz et al., 2007) and Disciphania Eichler (clade I, expanded Tinosporeae, Ortiz et al., 2007). This resemblance is particularly strong in M. cerre- jonensis having (1) few central secondaries, alternate or subop- posite, with the lowermost pair arising above the middle of the blade; (2) fi rst pair of lateral primaries branching once or few times exmedially along the length of the vein; and (3) mem- branaceous leaf texture. Epidermal characters provide useful systematic informa- tion in the Menispermaceae ( Ferguson, 1974 ; Cutler, 1975 ; Wilkinson, 1978 ; Hong et al., 2001 ). However, irregular epi- dermal cells with sinuous anticlinal walls and anomocytic stomata are widely represented within Menispermaceae ( Hong et al., 2001 ). Thus, Menispermites cerrejonensis cannot be re- 960 American Journal of Botany [Vol. 95 961August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia Fig. 3. Leaf architecture of Menispermites cerrejonensis Doria, Jaramillo & Herrera sp. nov. (A, B) ING-CJ6 ? 0145 ? 0318 (holotype). (C, D) ING- CJ6 ? 0907 ? 0318 (paratype). Arrows in A, B and C: fi mbrial veins. Circle in D shows third and fourth orders of venation. Scale bars = 10 mm. ? Fig. 4. Venation patterns of Menispermites cerrejonensis Doria, Jaramillo & Herrera sp. nov. (A) Secondary brochidodromous venation. Arrow: fi m- brial vein, ING-CJ6 ? 0137 ? 0317. (B ? D) ING-CJ6 ? 0908 ? 0318. (B) Tertiary, fourth and fi fth orders of venation. Arrow: sign of herbivory. (C) Close up of (B). (D) Areolation. Arrow: branched free ending veinlets. Scale bars: B = 5 mm, C = 3 mm, D = 1 mm. of symmetry, forming almost continuous concentric semicir- cles of tertiary veins; this pattern is not evident where the third venation is alternate percurrent or alternate-opposite percur- rent ( Fig. 6 ). Highest venation orders and fi mbrial vein are well preserved in many specimens, but it was not possible to prepare cuticles. Comparisons ? Menispermites cordatus , like M. cerrejonen- sis , presents one or few pairs of secondaries merging from the midrib with the lowermost pair arising above the middle of the blade; however, M. cordatus presents deeply cordate bases, concave ascending principal veins, and tertiary venation mostly opposite percurrent, whereas M. cerrejonensis presents rounded to truncate bases, straight or slightly curved ascending second- aries and mostly alternate percurrent tertiary venation. The combination of characters found in M. cordatus is also found in the neotropical genus Abuta Aublet (clade II-A, 962 American Journal of Botany [Vol. 95 ing 3/4 of the total length, looping with central secondaries, branching exmedially several times, each secondary merg- ing at an acute angle joining with the superadjacent vein at an obtuse angle; second pair of primaries merging at a wide acute to obtuse (70 ? 106 ? ) angle, with slightly curved course toward the margin reaching 1/4 of the total length of the blade, subparallel to the secondaries from the fi rst lateral prima- ries, looping with the lowermost secondary; several tertiaries merging distally at an acute angle, each one joining with the superadjacent at an acute angle ( Fig. 7A, B ). Leaf texture is chartaceous to coriaceous. Highest orders of venation are not easily recognizable. Comparisons ? Menispermites guajiraensis differs from M . cerrejonensis and M . cordatus in having several pairs of sec- ondaries merging from the midrib, with the lowermost pair aris- ing below the middle of the blade, not one to few central secondaries merging above the middle of the blade. Menisper- mites horizontalis presents a pattern of central secondaries dis- tribution similar to that of M. guajiraensis , but the secondaries in this taxon are horizontal and not with straight to smoothly curved course toward the margin. Thus, the characters that al- low us to differentiate M. guajiraensis from the other taxa here described are the number of secondary veins and their angle and course. Among extant Menispermaceae, the fossils present some similarities with Elephantomene Barneby & Krukoff (clade II-A, Anomospermae, Ortiz et al., 2007). This resemblance is particularly strong in the following characters: (1) several reg- ularly spaced medial secondary veins, departing from the mid- rib at an acute angle with straight course toward the margin; (2) fi rst pair of lateral primaries merging at an acute angle, branching exmedially in a series of straight parallel secondar- ies regularly spaced; and (3) exmedial pair of primaries merging at an obtuse angle reaching 1/4 or less the distance to the apex. In Elephantomene eburnea Barneby & Krukoff, the only ex- tant species of the genus, the leaves are broadly ovate to sub- orbicular, and the bases are rounded to truncate ( Ott, 1997 ), not cordate as in M. guajiraensis . Both E. eburnea and M. gua- jiraensis have major secondaries that form a narrow acute an- gle with the central primary, thus forming a fan ( Ott, 1997 ). Establishing the affi nities of this taxon is rather diffi cult be- cause the highest orders of venation ? reliable characters for confi rming affi nities within Menispermaceae ? were not well preserved. Anomospermae, Ortiz et al., 2007). Conspicuous opposite percurrent tertiaries, forming concentric semicircles, are also present in many Abuta species [e.g., A. rufescens, A. mycetandra Krukoff & Barneby, and A. pahni (Mart.) Krukoff & Barneby]. Menispermites cordatus differs from Abuta species with broadly ovate leaves in having secondaries and tertiaries looping far from the margin in rounded arches, whereas extant species of Abuta have veins looping conterminous to the fi mbrial vein oc- cur, at least at the basal part of the blade. Species ? Menispermites guajiraensis Doria, Jaramillo & Herrera sp. nov. Diagnosis ? Simple mesophyllous to megaphyllous leaves. Ovate symmetrical lamina, 18 (13 ? 30) cm long and 18.5 (11 ? 28) cm wide (3 specimens measured); base cordate, base angle obtuse; apex not preserved. Margin entire and unlobed. First vein category actinodromous basal, 5 ? 7 veins from the base; central primary wider than the lateral primaries. Secondary ve- nation brochidodromous; several pairs of ascending secondar- ies from the midrib, with the lowermost pair merging below the middle of the blade; central secondaries opposite to suboppo- site. Third vein category opposite percurrent. Fourth and fi fth venation orders regular polygonal reticulate. Areolation well developed. Fimbrial vein not observed, but probably present. Holotype ? ING-CJ6 ? 0169 ? 0319 ( Fig. 7A, B ) Paratype ? ING-CJ6 ? 0868 ? 0319 Type locality ? 0319, Pit Tabaco High Deep, Cerrej ? n coal mines, northern side of the Cesar-Rancheria basin, Guajira pen- insula, northern Colombia, 11 ? 66 ? N, 73 ? 31 ? W. Horizon ? Middle part of the Cerrej ? n Formation. Between coal seams 105 and 106. Derivatio nominis ? The specifi c epithet refers to the collec- tion area at Guajira Department. Studied material ? ING-CJ6 ? 0122 ? 0317, ING-CJ6 ? 0154 ? 0317, ING-CJ6 ? 0170 ? 0319, ING-CJ6 ? 0171 ? 0319. Description ? Specimens were collected from localities 0317 and 0319. The largest specimens of the morphotype CJ6, reaching the megaphyll size category, are included in this mor- photaxon. Petiole, where present, is marginal and stout. Cen- tral primary markedly wider than the lateral ones, with straight course. Central secondaries emerging at an acute angle, with straight to smoothly curved course toward the margin. The in- nermost pair of lateral primaries emerging at an acute (30 ? 60 ? ) angle from the central primary, with straight course, ascend- Fig. 5. Cuticular features of Menispermites cerrejonensis Doria, Jaramillo & Herrera sp. nov. (A) Epidermal cells. (B, C) Stomata in the areole area. Arrows in (B): guard cells. g = guard cell, s = subsidiary cell. Scale bars= 25 ? m. 963August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia Fig. 6. Leaf architecture of Menispermites cordatus Doria, Jaramillo & Herrera. (A, B) ING-CJ6 ? 0128 ? 0317 (holotype). (C, D) ING-CJ6 ? 0154 ? 0317. Scale bars = 10 mm. Species ? Menispermites horizontalis Doria, Jaramillo and Herrera sp. nov. Diagnosis ? Simple megaphyllous leaves. Ovate symmetrical lamina; base not preserved; apex acute, apex angle acute. Margin entire and unlobed. First vein category probably actinodromous basal, 3 (probably 5) veins from the base; central primary wider than the lateral primaries. Secondary venation brochidodromous; several pairs of horizontal secondaries from the midrib, with the lowermost pair merging below the middle of the blade; central secondaries opposite. Third vein category opposite percurrent. Fourth and fi fth venation orders regular polygonal reticulate. Areolation well developed. Fimbrial vein present. Holotype ? ING-CJ6 ? 0166 ? 0317 ( Fig. 8 ) Type locality ? 0317, Pit Tabaco, Cerrej ? n coal mines, north- ern side of the Cesar-Rancheria basin, Guajira peninsula, north- ern Colombia, located at 11 ? 14 ? N, 72 ? 57 ? ? W. 964 American Journal of Botany [Vol. 95 below the middle of the blade, but differs in having numerous horizontal instead of ascending medial secondaries. Moreover, the number of medial secondaries is higher in M. horizontalis . As in M. cordatus , M. guajiraensis has third venation opposite percurrent, but the two differ in the position and number of medial secondaries. Among extant Menispermaceae, several characters are similar to those of the neotropical Sciadotenia (clade II, Tiliacoreae, Ortiz et al., 2007), including (1) the presence of numerous horizontal secondaries, (2) tertiary venation opposite percurrent, and (3) ex- medial secondaries joining with the uppermost in an acute angle. In Sciadotenia usually the innermost pair of primary veins are pli- nerved (i.e., suprabasal actinodromous venation), diverging 0.3 ? 1 cm above the base ( Ott, 1997 ; G. Doria, personal observation); however; this character is not preserved in this specimen. DISCUSSION The general morphology observed in the four species de- scribed suggests a relationship to the angiosperm family Horizon ? Upper part of the Cerrej ? n Formation. Between coal seams 100 and 102. Derivatio nominis ? The specifi c epithet refers to the charac- teristic horizontal secondary venation. Studied material ? ING-CJ6 ? 0166 ? 0317 Description ? Specimen was collected in locality 0317. It is a fragmentary leaf that has a marginal vein, apparently palmate venation (the base is missing), two pairs of conspicuous pre- sumed lateral primaries with slightly curved course toward the apex ( Fig. 8A ). Numerous pairs of secondaries merging from the midrib at a perpendicular angle, the lowermost arising below the middle of the blade, medial secondaries opposite with horizontal course ( Fig. 8A, B ). Exmedial secondaries departing from lateral primaries at a wide acute to perpendicular angle, also with hori- zontal course, looping with the superadjacent at an acute angle ( Fig. 8C ). All secondaries (medial and exmedial) regularly and closely spaced. Tertiary veins opposite percurrent, perpendicular to secondaries ( Fig. 8C ). Leaf texture chartaceous. Comparisons ? Menispermites horizontalis , like M. guajiraensis , presents several medial secondaries with the lowermost merging Fig. 7. Leaf architecture of Menispermites guajiraensis Doria, Jaramillo & Herrera sp. nov. ING-CJ6 ? 0169 ? 0319 (holotype). Scale bars = 10 mm. 965August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia by the fusion of primary and secondary veins are present in Aristolochiaceae and Malvaceae as well; however, in Aristolo- chiaceae these arcs correspond to a brochidodromous festooned type of venation, whereas in Menispermaceae the venation is brochidodromous simple. In Malvaceae the arcs are mainly an- gular and sharp, not well rounded as in Menispermaceae. The generic name here adopted for the four new taxa, Menispermites Lesquereux ( Lesquereux, 1874 ), was originally proposed to describe broadly deltoid, peltate, or subpeltate, trilobed fossil leaves with actinodromous fi rst venation, from the Upper Cretaceous of North America, presumably related to Menispermaceae (e.g., Hickey and Wolfe, 1975 ). The genus is used now to encompass a wider morphological range including taxa with wide ovate to orbicular, peltate, subpeltate, or bas- ifi xed, unlobed to 3- or 5-lobed leaves with actinodromous or acrodromous fi rst venation ( Table 4 ). Several species have been Menispermaceae (basal eudicots, Ranunculales, APG II, 2003). Similar characteristics are pointed out by Wolfe (1968 , 1977 ) as diagnostic for fossil leaves of Menispermaceae. The princi- pal features of the new morphotaxa indicating a relationship with this family are (1) primary venation actinodromous; (2) secondary veins brochidodromous with conspicuous rounded loops; (3) tertiary veins opposite percurrent, alternate percur- rent, or mixed opposite-alternate percurrent; (4) fourth and fi fth venations polygonal reticulate; (5) areolation well developed; (6) margin entire; and (7) fi mbrial vein present. Other unrelated families, Aristolochiaceae and Malvaceae, also present the fi rst four characters, but they do not have a fi mbrial vein. The fi mbrial vein results from the fusion of higher vein orders into a vein running just inside the margin, and it is almost a constant character in the fossil specimens, except in the specimens of M. guajiraensis . Conspicuous loops formed Fig. 8. Leaf architecture of Menispermites horizontalis Doria, Jaramillo & Herrera sp. nov. (A ? D) ING-CJ6 ? 0166 ? 0319 (holotype). (C) Medial sec- ondaries. (D) Exmedial secondaries and perpendicular tertiaries. Scale bars = 10 mm. 966 American Journal of Botany [Vol. 95 T ab le 4 . Co m pa ra tiv e ta bl e of m or ph ol og ic al fe at ur es o f f os sil le af ta xa , p re su m ab ly re la te d to M en isp er m ac ea e. Ta x o n Le af sh ap e Lo ba tio n Pe tio la r at ta ch m en t M ar gi n Ba se sh ap e A pe x 1s t Ve n at io n N o. o f pr im ar ie s2 nd V en at io n N o. p ai rs o f c en tra l se co n da rie s 3r d Ve n at io n M ar gi na l u lti m at e v en at io n Ti m e Li te ra tu re M en is pe rm ite s a ce ri fol ia Le sq ue re ux D el to id 3 ? 5 Lo ba te N ot p el ta te (m arg in al ?) En tir e N ea rly tr un ca te to cu n ea te Lo be d ac ut e ap ex A ct in o 3 ? 5 ? 1 ? 2 ? ? Cr et ac eo us Le sq ue re ux , 1 87 4 M . a cu til ob u s Le sq ue re ux D el to id Lo ba te ? D en ta te B ro ad ly ro u n de d- n ea rly tr un ca te Lo be d ac ut e ap ex A ct in o 5 Cr as pe ? O pe n in os cu la tin g n et w o rk La te Cr et ac eo us B el l, 19 57 M . a m er ic an us B er ry Sl ig ht ly co rd at e (O va te ) U nl ob ed M ar gi na l En tir e B ro ad ly tr un ca te o r sli gh tly co rd at e B lu nt ly po in te d Pi nn at e 1 Ca m pt o (B roc hi. ) 4 ? 5 ? ? Eo ce ne B er ry , 19 30 M . b el li B er ry O va te U nl ob ed ? En tir e Co rd at e R ou nd ed A ct in o 5 Ca m pt o ? ? ? La te Cr et ac eo us D or f, 19 42 M . c oc ke re lli (K no w lto n) Do rf ? U nl ob ed Su bp el ta te En tir e ? u n du la te ? ? ? Ca m pt o ? O pp os ite pe rc ur re nt ? La te Cr et ac eo us D or f, 19 42 M . h ar de m an en sis B er ry O va te U nl ob ed M ar gi na l En tir e B ro ad ly tr un ca te d to s lig ht ly co rd at e ? A ct in o 3 (5? ) Ca m pt o ? br oc hi ? 2 ? 3 ? ? Eo ce ne B er ry , 19 30 M . k ni gh tii K no w lto n O rb ic ul ar Lo be d ? Lo be d- u n du la te Co rd at e R ou nd ed A ct in o 7 Ca m pt o ? ? ? La te Cr et ac eo us D or f, 19 42 M . l im ac io de s M ac G in iti e O va te U nl ob ed M ar gi na l En tir e Cu ne at e A cu m in at e A cr o Lo op in g ne ar th e m ar gi n ? O pp os ite pe rc ur re nt Fi m br ia l Eo ce ne M ac G in iti e, 1 96 9 M . o bt us ilo ba Le sq ue re ux R en ifo rm - de lto id O bs cu re ly 3- lo ba te Pe lta te D ee pl y u n du la te O bt us e, pe lta te , en la rg ed a nd tr un ca te ? A ct in o 5 Su bc ra sp e N on e ? fe w ? ? Cr et ac eo us Le sq ue re ux , 1 87 4 M . p ar va re o la tu s H ic ke y Ve ry w id e o v at e U nl ob ed M ar gi na l Sh al lo w ly lo be d O bt us e (C on ca v e) R ou nd ed - em ar gi na te A cr o 3 Lo op in g ne ar th e m ar gi n ? H ig h ac ut e an gl es Fi m br ia l Pa le oc en e H ic ke y, 19 77 M . p ot om ac en si s B er ry O rb ic ul ar to o v al , o v at e U nl ob ed M ar gi na l Cr en at e- u n du la te Cu ne at e R ou nd ed A ct in o 7 ? 13 ? ? ? ? La te Cr et ac eo us B el l, 19 56 , D oy le an d H ic ke y, 19 76 M . r en ifo rm is D aw so n Se m io rb ic ul ar to Se m ie lli pt ic al U nl ob ed M ar gi na l En tir e- sli gh tly u n du la te Co rd at e R ou nd ed A ct in o 7 Ca m pt o ? ? ? La te Cr et ac eo us B el l, 19 56 M . s al in en sis Le sq ue re ux B ro ad ly tr ia ng ul ar O bs cu re ly 5- lo be d o r D ee pl y u n du la te ly lo be d N ot p el ta te (m arg in al ?) D ee pl y u n du la te ly lo be d N ea rly tr un ca te o r at te nu at ed cu n ei fo rm ? A ct in o 5 B ro ch i ? ? ? Cr et ac eo us Le sq ue re ux , 1 87 4 M en is pe rm ite s s p . B el l ? Lo be d? ? ? A sy m m e- tr ic al -c or da te ? A ct in o 5 ? ? ? ? La te Cr et ac eo us B el l, 19 62 M en is pe rm ite s s p. (K no w lto n) n. co m b. B el l O va te U nl ob ed ? En tir e Co rd at e ? A ct in o ? ? ? O pp os ite pe rc ur re nt ? La te Cr et ac eo us B el l, 19 62 , (pl ate 20 , fi g. 3 ) M . t en ui ne vi s Fo n ta in e O va te ? Tr u n ca te to o bt us e A ct in o ? ? ? ? ? La te Cr et ac eo us D oy le a n d H ic ke y, 19 76 M . t or o su s B el l O va te -o rb ic ul ar U nl ob ed ? En tir e? Tr u n ca te - sli gh tly co rd at e ? A ct in o 5( ? 7) B ro ch i ? ca m pt o Fe w , re m o te fro m th e ba se Pe rc ur re nt , fa irl y str on g ? La te Cr et ac eo us B el l, 19 57 M . v irg in ie ns is Fo n ta in e O rb ic ul ar U nl ob ed ? Cr en ul at e Co rd at e ? A ct in o 10 ? ? ? ? La te Cr et ac eo us D oy le a n d H ic ke y, 19 76 967August 2008] Doria et al. ? Paleocene Menispermaceae from Colombia T ab le 4 . Co nt in ue d. Ta x o n Le af sh ap e Lo ba tio n Pe tio la r at ta ch m en t M ar gi n Ba se sh ap e A pe x 1s t Ve n at io n N o. o f pr im ar ie s2 nd V en at io n N o. p ai rs o f c en tra l se co n da rie s 3r d Ve n at io n M ar gi na l u lti m at e v en at io n Ti m e Li te ra tu re An am irt a m ill er i W o lfe O va te U nl ob ed M ar gi na l En tir e Co rd at e A cu m in at e A ct in o 7 B ro ch i ? ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 97 7 Ca lk in sia fra n kl in ie ns is W o lfe O bo v at e U nl ob ed M ar gi na l En tir e Cu ne at e A br up tly ac u te A ct in o 3 Ca m pt o or cr as pe 2 ? 4 ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 96 8 Ca lk in sia p la fke ri i W o lfe Q2 El lip tic al U nl ob ed M ar gi na l En tir e A cu te A cu m in at e A ct in o 5 Cr as pe 3 ? 4 ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 97 7 Ch on dr o de nd ro n br a si lie ns e D ol ia ni ti O va te U nl ob ed Su bp el ta te En tir e- u n du la te Sl ig ht ly co rd at e R et us e A ct in o 7 Cr as pe do - dr om ou s Fe w , ab ov e th e m id dl e o f th e bl ad e ? Fi m br ia l v ei n Pl io ce ne D ol ia ni ti, 1 94 9 Ci ss am pe lo s ro tu nd ifo lia Po tb u ry W id el y ov at e U nl ob ed M ar gi na l En tir e D ee pl y co rd at e- pe lta te R ou nd ed - sli gh tly re tu se A ct in o 3 B ro ch i 4? ? ? La te Eo ce ne Po tb u ry , 19 35 ? Co cc ul us ? fl a be lla (N ew be rry ) W o lfe El lip tic al to o bo v at e U nl ob ed M ar gi na l En tir e, cr en at e or sh al lo w ly lo ba te A cu te to o bt us e R ou nd ed - m u cr o n at e A cr o 3 Ca m pt o 0 ? 4 R eg ul ar po ly go na l re tic ul at e ? Pa le oc en e W o lfe , 1 96 6 , 19 72 ; H ic ke y, 19 77 Co cc ul us ro tu nd a M ac G in iti e Ci rc ul ar U nl ob ed M ar gi na l En tir e B ro ad ly ro u n de d- sli gh tly co rd at e R ou nd ed A ct in o 5 B ro ch i ? cr as pe 3? ? ? M io ce ne M ac G in iti e, 1 96 2 Co cc ul us sp . ? U nl ob ed ? En tir e R ou nd ed ? A ct in o 3 Ca m pt o ? ? ? Eo ce ne W o lfe , 1 97 7 ( pl at e 26 , fi g. 5 ) Co cc ul us sp . O za ki O va l t o el lip tic al Sh al lo w ly 3- lo be d M ar gi na l En tir e Co rd at e ? A cr o 3 Ca m pt o ? W ea kl y pe rc ur re nt ? M io ce ne - Pl io ce ne O za ki , 1 99 1 H yp er ba en a di for ma Po tb u ry O va te to o bo v at e U nl ob ed M ar gi na l En tir e Cu ne at e to co rd at e A cu te o r ac u m in at e A ct in o 5 B ro ch i 1 ? 2 ab ov e th e m id dl e o f th e bl ad e O pp os ite pe rc ur re nt ? La te E oc en e Po tb u ry , 19 35 H yp se rp a ca sh m an en sis W o lfe El lip tic al U nl ob ed M ar gi na l En tir e N ar ro w ly ro u n de d R ou nd ed A ct in o 5 B ro ch i Se v er al ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 96 8 H . fr a n kl in ie ns is W o lfe El lip tic al U nl ob ed M ar gi na l En tir e N ar ro w ly ro u n de d A cu te A ct in o 5 B ro ch i Se v er al ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 96 8 Li m ac ia st en op hy lla W o lfe O va te U nl ob ed M ar gi na l En tir e N ar ro w ly ro u n de d A cu te A ct in o 3 Ca m pt o ? A lte rn at e pe rc ur re nt Fi m br ia l v ei n Eo ce ne W o lfe , 1 97 7 M en is pe rm um da ur ic um oi de s B el l Su bp en ta go na l to s ub he x a- go na l U nl ob ed - lo ba te Pe lta te En tir e- de nt at e ? ? A ct in o 6 ? 8 Cr as pe 1 ? 2 Lo os e n et w o rk ? La te Cr et ac eo us B el l, 19 57 Pa ra tin om is ci um co n di tio na lis W o lfe O va te U nl ob ed M ar gi na l En tir e B ro ad ly ro u n de d A cu te A ct in o 5 Ca m pt o (fo rm ing lo op s) ? ? Fi m br ia l v ei n Eo ce ne W o lfe , 1 97 7 Py cn ar rh en a sp . El lip tic U nl ob ed ? En tir e R ou nd ed ? R ou nd ed ? A ct in o 5 B ro ch i ? ? ? Eo ce ne W o lfe , 1 97 7 No te s: A cr o, a cr od ro m ou s; A ct in o, a ct in od ro m ou s; Br oc hi , b ro ch id od ro m ou s, Ca m pt o, C am pt od od ro m ou s; Cr as pe , c ra sp ed od ro m ou s 968 American Journal of Botany [Vol. 95 diversifi cation seen in extant families during the Early Ceno- zoic are explained by canopy closure coupled with an increase in forest height and canopy stratifi cation. Our study suggests that Menispermaceae was present very early in the Cenozoic, being the earliest record for this family of lianas in the South American tropics. There are very few records of fossil fl oras from the Late Cretaceous of tropical South America. However, preliminary analysis of a new Maastrichtian fl ora (ca. 70 Ma) from central Colombia, the Guaduas fl ora, indicates the ab- sence of Menispermaceae ( Gutierrez and Jaramillo, 2007 ). The Paleocene Cerrej ? n paleofl ora could then be the oldest record of a multistratifi ed tropical rain forest in the neotropics, sug- gesting that multistratifi cation is an ancient characteristic of South American rain forests, one that evolved following the Cretaceous ? Paleocene transition. LITERATURE CITED Ablaev , A. G. 1971 . Remnants of fossil leaves of Menispermum from Primorye Territory. Botanical Journal of USSR 56 : 1356 ? 1358 . Agranovskaya , I. A. , Y. A. Alyushinskogo , E. F. Asatkina , E. P. Boytsova , A. D. Bocharnikova , Z. M. Martynovoi , I. M. Pokrovskaya , G. M. Pomanovskoi , M. A. Sedova , N. K. Stelmak , and N. A. Shchekina . 1960 . 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A palynostratigraphical correlation of seven test wells in the Coastal Plain of Surinam. Geologie en Mijnbouw 21 : 177 ? 182 . Andreeva , E. M. , E. P. Boytsova , T. T. Koltsova , N. I. Komarova , N. V. Kruchinina , A. A. Lyuber , M. V. Oshurkova , L. A. Panova , I. M. Pokrovskaya , G. M. Romanovskaya , I. A. Sivertseva , N. K. Stelmak , I. P. Tabachnikova , A. A. Yalysheva , and O. N. Zhezhel . 1966 . Opisanie iskopaemykh spor predstavitelei Bryophyta, Lycopsida, Sphenopsida, Filicinae i rastitelnykh mikrofos- silii neyasnogo sistematicheskogo polzheniya ? Rastitelnye mikrofossilii neyssnogo sistematicheskogo polozheniya. In I. M. Pokrovskaya [ed.]. Paleopalinologiya. Trudy VSEGEI, Novaya Seriya 141: 114 ? 135. Andrews , H. N. 1970 . Index of generic names of fossil plants 1820 ? 1965. U. S. Geological Survey Bulletin . Apg II [Angiosperm Phylogeny Group] . 2003 . An update of the Angiosperm Phylogeny Group classifi cation for the orders and fami- lies of fl owering plants: APG II. Botanical Journal of the Linnean Society 141 : 399 ? 436 . Barbashinova , V. N. 1966 . Spore-pollen complexes of Eocene deposits of Primorsk. Doklady Akademii Nauk SSSR 170 : 149 ? 151 . Barbashinova , V. N. 1968 . Pollen from old angiosperms in Tertiary de- posits in some far eastern districts. Doklady Akademii Nauk SSSR 179 : 1167 ? 1170 . Barbashinova , V. N. 1971 . Spore and pollen assemblages of the Uglovski Suite of the Artemovsk Coal Layers. Proceedings of the Institution of Higher Learning; Geology and Intellce Service 11: 8 ? 14. Barclay , R. S. , K. R. Johnson , W. J. Betterton , and D. L. Dilcher . 2003 . Stratigraphy and megafl ora of a K-T boundary section in the eastern Denver Basin, Colorado. Rocky Mountain Geology 38 : 45 ? 71 . proposed under this generic name from the Upper Cretaceous to Eocene of North America ( Fontaine, 1889 ; Berry, 1916 , 1922a , b ; Seward, 1927 ; Hollick and Martin, 1930 ; Dorf, 1942 ; Bell, 1956 , 1957 , 1962 ; MacGinitie, 1969 ; Doyle and Hickey, 1976 ; Hickey, 1977 ; Wing et al., 1995 ), Asia ( Kryshtofovich and Baikovskaia, 1960 ; Budantsev, 1968 ; Takhtajan, 1974 ; Spicer et al., 2002 ), and Europe ( Hughes, 1976 ). The four species described here fit into the broad diagnosis for the genus. Menispermites species cannot be assigned to a tribe within Menispermaceae, but they have many characteristics of leaf morphology and venation patterns that suggest an affi nity with Menispermaceae. Leaf characters are thought to be highly homoplasious within Menispermaceae ( Ortiz et al., 2007 ). Examination of the mor- phological transitions in the molecular phylogeny of the family shows that the typically simple leaf with actinodromous vena- tion is the plesiomorphic condition in the family ( Ortiz et al., 2007 ). Leaf characters in Menispermaceae have been reported as useful for identifi cation ( Ott, 1997 ), but the full variation of leaf morphology, including cuticles, among all 70 genera of Menispermaceae remains to be studied. Although similar to the leaves of several modern menisper- maceous taxa, each species of Menispermites described here has a unique combination of features not found in living Menispermaceae or other fossil leaves. However, some charac- ters could be used to relate the fossils to extant groups. In Menispermites cerrejonensis the membranaceous leaf texture is notable, suggesting a herbaceous habit, and the venation pattern of secondaries ( Figs. 3, 4 ) resembles that of several species of Disciphania , tropical American climbers currently centered in Amazonia ( Kessler, 1993 ). Leaf texture is hardly recognizable in the other three fossil taxa because of the type of fossiliza- tion (impressions vs. compressions in M. cerrejonensis ). In Menispermites cordatus , the most conspicuous characters are the strongly curved primaries and medial secondaries, and the opposite percurrent third venation ( Fig. 6A ? D ). This species resembles some species of the extant Abuta , the most speciose genus of Menispermaceae in the neotropics with ~32 species ( Kessler, 1993 ). Menispermites guajiraensis resembles Ele- phantomene , a monospecifi c genus from the South American tropics, in the pattern of first and second venation, where the major secondaries form a narrow acute angle with the cen- tral primary, forming a fan ( Fig. 7A, B ). The presence of hori- zontal secondaries in M. horizontalis ( Fig. 8 ) is characteristic of the extant Sciadotenia , a genus of ~20 species of climbers from tropical Central and South America ( Kessler, 1993 ). How- ever, a megaphyllous leaf size is not common in the extant Sciadotenia . Most of the extant species of Menispermaceae are lianas or vines. Woody habit probably is the ancestral condition in the group ( Kim et al., 2004 ; Ortiz et al., 2007 ). Lianas are an im- portant component in neotropical fl oras ( Gentry, 1991 ) and infl uence the dynamics of many forests ( Schnitzer et al., 2000 ; Schnitzer and Bongers, 2002 ; Londr ? and Schnitzer, 2006 ). 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