Herrera Et Al_2011_Paleocene Menispermaceae Endocarps From Colombia & Wyoming

Botany

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December 2011 ◆ Volume 98 ◆ Number 12

AMERICAN JOURNAL OF

Offi cial Publication of the Botanical Society of America, Inc.www.amjbot.org

American Journal of Botany 98(12): 2004–2017. 2011.

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American Journal of Botany 98(12): 2004–2017, 2011; http://www.amjbot.org/ © 2011 Botanical Society of America

Menispermaceae are a pantropical angiosperm family within Ranunculales with approximately 71 genera and 450 – 500 spe-cies ( Thanikaimoni, 1984 ; Kessler, 1993 ). Species of the family are most common in tropical lowland rainforests although sev-eral genera have species adapted to cooler or drier biomes. Some of the morphological synapomorphies recognized for the

family include liana or vine growth habit, fl owers unisexual, plants usually dioecious, petioles often swollen at the base, ovules two (but one aborting), fruits in aggregates of drupes, endocarps with a condyle and seeds with a large, usually curved embryo ( Diels, 1910 ; Kessler, 1993 ; Ortiz et al., 2007 ; Hoot et al., 2009 ; Jacques, 2009a ).

Historically, Menispermaceae were divided into fi ve or eight tribes based largely on endocarp and seed characters ( Miers, 1851 ; Diels, 1910 ; Kessler, 1993 ). However, the recent boom in molecular analyses has resulted in revised hypotheses of tribal relationships, morphological evolution, and classifi cation for this family ( Jacques et al., 2007 , 2011 ; Ortiz et al., 2007 ; Wang et al., 2007 ; Jacques and Bertolino, 2008 ; Hoot et al., 2009 ). The endocarp types (i.e., boat-shaped, horseshoe-shaped, hairpin-shaped) that were used for the traditional classifi cation within the family appear to be highly homoplasious, and several tradi-tional tribes and genera based on these characters are poly-phyletic or paraphyletic when superimposed on a molecular phylogeny ( Jacques et al., 2007 ; Ortiz et al., 2007 ; Hoot et al., 2009 ). These new hypotheses of intrafamilial relationships in Menispermaceae indicate the need for caution in identifying affi nities for fossil taxa. Nonetheless, fossil endocarps are in-valuable for understanding the evolution and paleobiogeography of the family. Furthermore, their identifi cation to the tribe or ge-nus level will require a combination of fruit characters instead of relying on the historical endocarp types ( Jacques et al., 2011 ).

As recently listed in Doria et al. (2008) and Jacques (2009b) , dozens of leaf, wood, fl ower, pollen, and endocarp fossils

1 Manuscript received 15 November 2010; revision accepted 12 September 2011.

This research was made possible through support from the Evolving Earth Foundation, the Geological Society of America Foundation, the Asociaci ó n Colombiana de Ge ó logos y Geof í sicos del Petroleo-ARES, The Smithsonian Institution, the Gary S. Morgan Student Research Award, and the Lewis & Clark Foundation-American Philosophical Society to F.H., National Science Foundation (NSF) grants EF-0431266 and BSR-0743474 to S.R.M., NSF DEB-0919071 to M.R.C., NSF DEB-0733725 to C.J., and NSF DEB-0542679 to S.B.H. The authors thank L. Teicher, F. Chavez, and the geology team at Minas Cerrej ó n. J. Moreno, E. Cadena, A. Rincon, S. Moron, G. Bayona, S. L. Wing, J. Bloch, M. I. Barreto, G. Doria, and A. Rojas for their assistance with the Bogot á and Cerrej ó n fi eld trips; C. A. Mu ñ oz for his hospitality during fi eldwork in Bogot á . D. Bell (US) granted access to modern Menispermaceae collections, R. C. Ortiz and F. Jacques provided helpful discussions about the systematics of the fossils, and G. Bedoya provided nomenclatural suggestions. Two anonymous reviewers and T. Lott provided helpful reviews of the manuscript. F.H. thanks B. Himschoot for support.

6 Author for correspondence (e-mail: fherrera@fl mnh.ufl .edu)

doi:10.3732/ajb.1000461

PHYTOGEOGRAPHIC IMPLICATIONS OF FOSSIL ENDOCARPS OF MENISPERMACEAE FROM THE PALEOCENE OF COLOMBIA 1

Fabiany Herrera 2,3,6 , Steven R. Manchester 2 , Sara B. Hoot 4 , Keir M. Wefferling 4 , M ó nica R. Carvalho 3,5 , and Carlos Jaramillo 3

2 Department of Biology – Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611 USA; 3 Smithsonian Tropical Research Institute, Apartado Postal 0843-03092, Balboa, Anc ó n, Rep ú blica de Panam á ; 4 Department of Biological Sciences, P. O. Box 413, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin 53201 USA; and 5 Department

of Geosciences, Pennsylvania State University, University Park, Pennsylvania 16802 USA

• Premise of the study: Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifi cally within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene.

• Methods: Fossil endocarps of Menispermaceae were collected from the Cerrej ó n Formation, the recently discovered Bogot á fl ora, and Wyoming (~60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera.

• Key results: Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylo-genetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identifi cation and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania , which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania , Cissampelos , and Cyclea ; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades.

• Conclusions: The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleo-forests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous – Paleogene boundary.

Key words: Cissampelos ; Colombia; endocarps; fossils; paleobiogeography; Paleocene; rainforest; Stephania .

2005December 2011] Herrera et al. — Fossil endocarps of Menispermaceae from South America

Formation had accumulated in the lowlands ( Bayona et al., 2008 ). Pollen as-semblages from the Bogot á Formation belong to the zone T-03b- Foveotricol-pites perforatus of Jaramillo et al. (2011) , indicating a middle to late Paleocene age. The formation varies from extensive and thick siltstones, claystones, and paleosols to interbedded sandstones and sporadic conglomerates and breccias. Contrary to the Cerrej ó n Formation, the Bogot á sequence lacks coal deposits. Initial sedimentological analyses from the Bogot á Formation indicate deposi-tion in fl uvial environments. The specimen BF18-ING-0017 was found in a massive yellowish lacustrine claystone bed, about 40 cm thick underneath a thick red paleosol. The specimens BF19-ING-0012-0016 and BF18-ING-0023 are from grayish, laminated, and fi ne-grained sandstones that suggest deposi-tion in overbank deposits. The specimens BF20-ING-0019-0020, BF20-ING-0024, and BF20-ING-0026 were found along with abundant crocodile teeth and turtle shells in an ~20 cm reddish breccia with very angular grains and iron oxide cementing material. This bed may have been deposited during an aqueous debris fl ow. Some of the plant families and genera already identifi ed from the Bogot á fl ora include Annonaceae, Arecaceae, Fabaceae, Lauraceae, Malvaceae, Salicaceae, Theaceae, and the fl oating-aquatic fern Salvinia .

The Colombian fossils are stored at the paleontological collections of the Colombian Geological Institute (INGEOMINAS) in Bogot á , Colombia.

The specimen from Linch, Wyoming (UF18257-21872; Fig. 17 ) was recov-ered from a Paleocene (Tiffanian) locality of the Fort Union Formation (Pow-der River Basin), exposed at the west side of the highway 192 at the southern end of the road cut in Pine Ridge, WY quadrangle 7.5 ′ series, GPS 43 ° 37.800 ′ N 106 ° 12.371 ′ W. Pollen samples from the Fort Union Formation have corrobo-rated a middle-late Paleocene age for the stratigraphic sequence where the fossil was found ( Nichols, 1994 ). The menispermaceous endocarp is found together with predominantly deciduous taxa including Metasequoia , Aesculus , Cranea , Corylites , Fagopsiphyllum , and Juglandicarya simplicarpa .

We studied modern fruits representing 68 of the 71 extant genera recognized for Menispermaceae with materials from the U. S. National Herbarium (US) at the Smithsonian Institution in Washington, D.C., the Smithsonian Tropical Re-search Institute (STRI) in Panama, the University of Florida Herbarium (FLAS) in Gainesville, Florida, the U. S. National Seed Herbarium housed at the Na-tional Arboretum, Washington, D. C. (BARC), the Herbarium of the Arnold Arboretum (A) at Harvard University in Boston, Massachusetts, the Field Mu-seum of Natural History Herbarium (F) in Chicago, Illinois, and the University of Wisconsin-Milwaukee (UWM) Herbarium in Milwaukee, Wisconsin ( Figs. 18 – 48 ). To fully examine the endocarp morphology of the extant taxa (species listed in Appendix 1) and prepare them for photography, the mesocarp tissue was removed after hydration with boiling water or by soaking the fruits in tap water for 8 – 24 h, depending on their size and mesocarp thickness. The endo-carps were cleaned with a nylon toothbrush and rinsed until free of soft tissues. Photographs were taken using a Retiga 2000R digital camera with QCapturePro 5.1 software, attached to an Olympus SZX16 microscope in the UWM micros-copy laboratory. For the grayscale photos, exposures were adjusted and back-grounds removed in Photoshop CS2 (Adobe, San Jose, California, USA). Fully mature endocarps and drupes were measured with calipers; the average size was recorded when multiple specimens were examined. These studies were supplemented by consideration of published images and descriptions (e.g., Thanikaimoni, 1986 ; Jacques, 2009a ). Terminology for the description of fruits partly follows Jacques (2009a) . See Appendix 2 for a complete defi nition of endocarp characters.

To assess affi nities of the fossil endocarps with living genera within Menispermaceae, we searched for extant morphological synapomorphies based on the characters that are preserved in the fossil remains. Scoring criteria are available in Appendix 3, also at the MorphoBank web site [http://www.morphobank.org; Project (P407): Herrera et al_Endocarp Morphological Matrix Menispermaceae]. Character optimization was conducted with the pro-gram Mesquite 2.74 ( Maddison and Maddison, 2009 ). The morphological char-acters were superimposed on a simplifi ed DNA tree topology for Menispermaceae (only those extant clades with high support of PP ≥ 95 and BS ≥ 80 were main-tained; Hoot et al., 2009 ). This simplifi ed DNA tree was then used as the “ back-bone constraint ” for a molecular scaffold analysis ( Springer et al., 2001 ; Manos et al., 2007 ). Sagittaria sagittifolia (Alismataceae) and Fumaria indica (Papav-eraceae) were used to root the tree. The morphological matrix included 25 char-acters, 17 of them referring to the endocarp, the remaining eight characters were taken from Hoot et al. (2009) and correspond to habit, leaf and pollen features (Appendix 3). A heuristic search was carried out with 1000 replicates of random taxon addition and tree-bisection-reconnection (TBR) branch swap-ping in the program PAUP* 4.0b10 ( Swofford, 2003 ). Bootstrap analyses were performed on the morphological data without enforcing the backbone con-straint, using 100 replicates and full heuristic searches.

have been attributed to Menispermaceae ranging from the lower Cretaceous to the Pliocene. In South America, Paleo-gene fossil leaves have been attributed to Menispermaceae from the Paleocene of Colombia and Argentina ( Iglesias et al., 2007 ; Doria et al., 2008 ) and the Eocene of Brazil ( Dolianiti, 1949 ; Mello et al., 2000 ). Of these fossil records, just those reported from the Paleocene of Colombia ( Doria et al., 2008 ) have been fully described and compared with extant leaves of the family; the remaining reports of Menispermaceae in South America still require a detailed examination to confi rm their affi nity. Because of the homoplasy found in leaf characters in Menispermaceae (see Hoot et al., 2009 ), the four Paleocene fossil species from the Cerrej ó n Formation of Colombia could not be placed more specifi cally within recognized clades of the family ( Doria et al., 2008 ). Future comparative studies on cuticle and leaf architectural patterns among living Menisper-maceae might provide stronger evidence for placing such fossils.

Jacques et al. (2011) recently used a molecular scaffold analysis for the placement of 26 taxa of fossil menisperma-ceous endocarps. In general, their consensus tree agreed with the previous suggested affi nities of the fossil taxa, except in the case of some fossil species placed within the genera Coc-culus and Parabaena . Despite the drawbacks of the molecular scaffold approach (relatively low number of fossilizable char-acters compared to DNA data, the lack of interaction among data partitions, homoplasy, and the limited topology of the resulting trees), this method seems feasible for hypothesizing the placement of extinct species within a given topology of extant taxa ( Manos et al., 2007 ).

Here, we describe three Paleocene fossil taxa based on en-docarps collected from north and central Colombia (Cerrej ó n and Bogot á paleofl oras) and Wyoming (USA). A molecular scaffold analysis was implemented for the placement of the fossil taxa. The menispermaceous fossil endocarps comple-ment the earlier report based on foliage from the Cerrej ó n fl ora ( Doria et al., 2008 ) and provide greater resolution of in-trafamilial relationships. In particular, these new fossil spe-cies suggest a wider biogeographic distribution for the ancestors of extant African and Australasian lineages and sug-gest a Paleocene connection with extinct menispermaceous taxa from North America.

MATERIALS AND METHODS

The newly recovered South American endocarps are from two widely sepa-rated Paleocene fl oras from northern (Cerrej ó n fl ora) and central (Bogot á fl ora) Colombia ( Figs. 1 – 16 ). Three specimens were collected from the upper levels of the Cerrej ó n Formation, an ca. 700-m-thick stratigraphic sequence, composed of abundant and thick coals, fl uvial sandstones, and lacustrine siltstones that were deposited in a succession of tidal fl ats to coastal fl oodplains crossed by channels ( Bayona et al., 2004 , 2008 ). The Cerrej ó n Formation has been dated as middle to late Paleocene (~60 Ma) based on pollen zonation, correlations with stable carbon isotopic data, and marine microfossils ( Jaramillo et al., 2007 ; 2011 ). The Cerrej ó n endocarp fossils (CJ85-ING-1412 to CJ85-ING-1414) were found along with angiosperm leaf remains in an extensive gray siltstone bed around 60 – 90 cm thick that was deposited in a swampy-lacustrine environment. The local fl ora was dominated by Araceae, Arecaceae, Fabaceae, Lauraceae, Malva-ceae, Zingiberales, and several unidentifi ed ferns ( Herrera et al., 2008 ; G ó mez-Navarro et al., 2009 ; Wing et al., 2009 ; Carvalho et al., 2011 ).

Eleven endocarps were collected from a recently discovered fossil fl ora from the eastern Colombian Andes. The plant localities are from the Bogot á Formation, an outcropping at the Sabana de Bogot á between 2700 – 3000 m a.s.l. During the Paleocene, the Andes had not been uplifted, and the Bogot á

2006 American Journal of Botany [Vol. 98

Description — Impressions and casts of fi ve horseshoe-shaped endocarps; outline of endocarp obovate in lateral view with a smooth and broad-rounded dorsal crest and an asym-metrical and acute base; length 4.5 – 6.5 mm, mean 5.65 mm (SD = 0.89, N = 4), width 3.5 – 4.3 mm, mean 3.9 mm (SD = 0.43, N = 4); endocarp wall thickness ~0.6 – 0.7 mm (measured from the locule cast to the apex of the endocarp); the dorsal crest with one side broader (toward putative stylar end) than the other; lateral crest decorated with spine-like structures (from 10 to 13); condyle outline obovate; a straight ventral vascular tube ascends (from base of condyle area) and approaches the locule forming a strong curvature at the connection point (preserved only in one specimen CJ85-ING-1413; Figs. 3, 4 ); locule cast apically positioned within the outline of the endocarp, with un-equal limbs and rimmed by the lateral crest, beset with radially aligned ribs (from 10 to 13); locule cast limbs widely spaced, ~0.7 mm (measured from tip of the short limb to tip of the lon-ger limb); longer limb of locule cast appears to be at seed radi-cal end (also stylar end of fruit); ventral notch straight; a tiny, narrow (length ~1 mm, wide ~0.2 mm), and obovate perfora-tion piercing the endocarp near the base.

Derivation of specifi c epithet — From the prefi x “ palaeo ” for ancient and “ sudamericana ” from the continent where the fos-sils originate.

Age, source, and stratum — Middle to late Paleocene. Holo-type and paratypes CJ85-ING-1413 and CJ85-ING-1414 from Guajira Peninsula, Rancher í a Basin, Cerrej ó n coal mine, Cer-rej ó n Formation, Tabaco Extensi ó n localities 0705 and 0712, 11 ° 07 ′ 49.8N, 72 ° 34 ′ 60.5W, localities placed below coal bed 175. Paratypes BF18-ING-0017 and BF18-ING-0023 from Cundinamarca state, Cogua locality FH0903, Bogot á Forma-tion; 5 ° 04 ′ 36.1N, 73 ° 57 ′ 18.9W.

Systematic affi nity within Menispermaceae — Among mod-ern genera of the S-C-C clade ( Fig. 50 ), the fossil shows closest morphological similarity to Stephania . Some of the characters that support this affi nity include the horseshoe-shaped endo-carp, the length of the endocarp considerably less than 10 mm, a thin endocarp wall ( < 1 mm), only one lateral crest on each side of the endocarp, a straight ventral vascular tube, a long radical limb, conspicuous locule ribs, a straight ventral notch, and the lack of protrusions in the locule chamber ( Figs. 18 – 28 ). Although none of these traits by itself is synapomorphic for Stephania ( Fig. 49 ), the combination of all the characters men-tioned makes it possible to recognize endocarps of this genus and facilitates identifi cation of the Paleocene endocarps. Steph-ania exhibits a signifi cant amount of intrageneric variation in ornamentation (ribs and spines), perforation (presence/absence and size), endocarp shape (from obovate to rounded), number of dorsal crests (one or two), and spiny or nonspiny dorsal and lateral crests ( Figs. 18 – 28 ) and the Cerrej ó n and Bogot á endo-carps fi t well within this natural morphological variation. Cis-sampelos and Cyclea are closely related to Stephania ( Hoot et al., 2009 ; Jacques et al., 2011 ), but their endocarps are easily distinguished ( Figs. 29 – 33, 35 – 39 ). In these two genera, there are always two lateral crests per side, usually two dorsal crests (rather than one as in this fossil), and their endocarps are often imperforate.

Stephania palaeosudamericana also resembles members of the Sinomenium - Menispermum clade, but in these two genera the endocarps do not have a complete perforation (but rather

SYSTEMATICS

Fruits of Menispermaceae are single-seeded drupes with an ornamented to smooth endocarp; the shape of the endocarp and its seed varies strongly from completely straight, boat-shaped, to horseshoe-shaped as found in many extant genera ( Figs. 18 – 48 ). The curved-seeded endocarps result from an outgrowth of the ovary wall into the placental region; this outgrowth is known as the condyle ( Hoot et al., 2009 ). Curved seeds are not unique for Menispermaceae; for example, some genera in Schisan-draceae, Cochlospermaceae, and Alismataceae may resemble the typical horseshoe shape of some Menispermaceae. How-ever, the combination of characters mentioned already (orna-mented to smooth endocarp and condyle) makes the drupes of Menispermaceae unique within angiosperms. The three new species from the Paleocene of Colombia and Wyoming reported in this study present a unique combination of characters diag-nostic for the family.

The morphological characters mapped over the simplifi ed topology of Menispermaceae corroborate other recent studies indicating homoplasy among endocarp morphology features within this family ( Fig. 49 ; Appendix 3). Although we did not fi nd any strong synapomorphies for the simplifi ed clades in Menispermaceae, the character mapping provides hypoth-eses for the placement of the fossils. The three fossil taxa studied are very similar to the endocarps found in the follow-ing clades and genera ( Fig. 49 ): (a) Sinomenium - Menisper-mum , (b) Pericampylus (c) Stephania - Cissampelos - Cyclea ( S - C - C ). Morphological characters shared between these clades and the fossils treated here include horseshoe-shaped endocarps, the endocarps being less than 10 mm in length, the presence or absence of a basal sculpturing, the presence of lateral crests, and a spiny ornamentation on both dorsal and lateral crests.

The molecular scaffold analysis yielded 152 trees of 76 steps (CI 0.447, RI 0.548, RC 0.245). The resulting strict consensus tree was similar to the backbone constraint generated from the DNA data ( Fig. 49 ), but placed the three fossil species within the extant S - C - C clade ( Fig. 50 ). A more detailed comparison of the fossil endocarps with modern genera is discussed follow-ing the taxonomic descriptions.

Order — Ranunculales Dumortier

Family — Menispermaceae A. L. de Jussieu

Genus — Stephania Loureiro

Type species — Stephania palaeosudamericana Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo sp. nov.

Diagnosis — Endocarp thin-walled; dorsal crest broad-rounded, base asymmetrical and acute; with one lateral crest on each side of the plane of symmetry; lateral crests spiny; locule cast apically positioned and with unequal and closely spaced limbs and 10 to 13 ribs; straight ventral vascular tube; endocarp with a tiny perforation in the condylar area, near the base.

Holotype hic designatus — CJ85-ING-1412 ( Figs. 1, 2 ).

Paratypes — CJ85-ING-1413 ( Figs. 3, 4 ); CJ85-ING-1414 ( Fig. 5 ); BF18-ING-0017 ( Fig. 6 ); BF18-ING-0023 ( Fig. 7 ).


Figs. 1 – 17. Fossil taxa from Paleocene of Colombia and Wyoming, USA. Figs. 1 – 7. Stephania palaeosudamericana Herrera, Manchester, Hoot, Wef-ferling, Carvalho et Jaramillo sp. nov. Figs. 1, 2. Holotype (CJ85-ING-1412), part and counterpart; from upper to lower arrows: 1. Locule ribs, seed radical end, straight ventral vascular tube, stylar end; 2. lateral ridges. Figs. 3, 4. Paratype (CJ85-ING-1413), part and counterpart; from upper to lower arrows: 3. Dorsal crest, entry of vascular tube into seed locule; 4. lateral ridges, perforation. 5. Paratype (CJ85-ING-1414), from upper to lower arrows: broader dorsal crest, straight ventral notch. 6. Paratype (BF18-ING-0017), arrow: lateral spines. 7. Paratype (BF18-ING-0023), locule cast showing a longer limb. Figs. 8 – 11. Menispina evidens Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo gen. nov. sp. nov. 8. Holotype (BF19-ING-0014), upper arrow: conspicu-ous dorsal spine, lower arrow: basal sculpturing. 9. Paratype (BF19-ING-0015), arrow: basal sculpturing. 10. Paratype (BF19-ING-0012), arrow: broader dorsal crest. 11. Paratype (BF19-ING-0016), left arrow: pedicel, middle arrow: basal sculpturing, right arrow: broader dorsal crest. Figs. 12 – 17. Palaeoluna bogotensis Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo gen. nov. sp. nov. 12. Holotype (BF20-ING-0020), upper arrow: conical pits of spines, lower arrow: locule ribs. 13. Paratype (BF20-ING-0019), upper arrow: locule inner protrusions, lower arrow: thick endocarp wall. 14. Paratype (BF20-ING-0024), arrow: locule inner protrusions. 15. Paratype (BF20-ING-0026), showing thick endocarp wall and locule inner protrusions. 16. Paratype (BF2-ING-0018), arrow: putative stylar end. 17. Palaeoluna sp. (UF18257-21872), Paleocene of Wyoming, endocarp impression. Scale bars = 2 mm.


Systematic affi nity within Menispermaceae — No living ge-nus coincides fully with the morphology of Menispina evidens . Nevertheless, this fossil shares characters with three unrelated clades within the Menispermaceae ( Fig. 49 ; Appendix 3): S - C - C clade, Sinomenium - Menispermum , and Pericampylus .

Menispina evidens is similar to the Sinomenium - Menisper-mum clade and shares some characters (i.e., horseshoe-shaped endocarps, length signifi cantly less than 10 mm, thin endocarp wall ( < 1 mm), and the lack of locule inner protrusions; Fig. 49 ); however, the extant genera differ from the Paleocene species in the presence of a vasculature channel on the lateral crests, the absence or poor development of dorsal spines, and inconspicu-ous locule ribs ( Figs. 42 – 49 ).

On the other hand, Menispina evidens resembles the extant genus Pericampylus in having endocarps with limbs of equal length, basal sculpturing, a horseshoe-shaped endocarp, length signifi cantly less than 10 mm, a thin endocarp wall ( < 1 mm), conspicuous locule ribs, a straight ventral notch, and lack of locule inner protrusions ( Figs. 40, 41, 49 ). Pericampylus dif-fers, however, from Menispina in having two dorsal and lateral crests on each side of the endocarp, and in the presence of very thick spines ( Fig. 40 ).

The molecular scaffold analysis places Menispina within the S - C - C clade together with the two other fossil genera from the Paleocene of Colombia ( Fig. 50 ). Within this clade, Menispina is placed as sister to Palaeoluna , but without boot-strap support. Characters shared with the S - C - C clade include: the horseshoe-shaped endocarp, length signifi cantly less than 10 mm, thin endocarp wall ( < 1 mm), conspicuous locule ribs, a straight-margined ventral notch, and the lack of locule inner protrusions ( Fig. 49 ). Notable differences between M . evidens and other members of the S - C - C clade, namely the absence of endocarps with limbs of equal length and basal sculpturing, as well as the lack of synapomorphic endocarp characters to re-late this fossil genus to any other extant clade, lead us to con-sider M . evidens as of uncertain placement within the extant Menispermaceae topology.

Morphogenus — Palaeoluna Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo gen. nov.

Generic diagnosis — Endocarp thick-walled; dorsal crest or-namented with inconspicuous narrow spines; one lateral crest; locule cast with unequal and widely spaced limbs, and 15 to 16 prominent ribs; locule chamber with inner protrusions.

Derivation of generic name — From the prefi x “ palaeo ” for ancient and the Latin “ luna ” for moon, because of the resem-blance to a crescent moon-like shape.

Type species — Palaeoluna bogotensis Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo sp. nov.

Holotype hic designatus — BF20-ING-0020 ( Fig. 12 ).

Paratypes — BF20-ING-0019 ( Fig. 13 ); BF20-ING-0018 ( Fig. 16 ); BF20-ING-0024 ( Fig. 14 ); BF20-ING-G0026 ( Fig. 15 ).

Species description — Molds and casts of fi ve horseshoe-shaped endocarps; endocarp outline obovate in lateral view with broad-rounded dorsal side; length ~6.2 – 9.0 mm, mean 7.2 mm (SD = 1.2, N = 4), width 5.0 – 7.0 mm, mean 5.9 mm (SD = 0.9,

lateral apertures that do not traverse the endocarp) and are usu-ally rounded in outline, the dorsal crest is more or less spiny (more in Sinomenium ), and the locule ribs are inconspicuous ( Figs. 42 – 48 ).

Fossil endocarps of Stephania have been reported from the early Miocene of Kenya ( Chesters, 1957 ) and the late Pleisto-cene of Nepal ( Bhandari et al., 2009 ). Both fossil species have prominent ornamentation on the dorsal crest, which is quite dif-ferent from S. palaeosudamericana ; the species from Nepal also has a larger perforation.

Morphogenus — Menispina Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo gen. nov.

Generic diagnosis — Endocarp thin-walled; endocarp outline elliptic, circular and obovate in lateral view with single rows of conspicuous acute spines on the dorsal and lateral crests; locule cast extending near to the base, more or less smooth with equal and closely spaced limbs and marked with 11 to 14 radial ribs that are more or less aligned with the dorsal and lateral spines; apparently lacking a perforation, but with a basal sculpturing near the shorter limb; pedicel short.

Derivation of generic name — From the Greek word “ Meni-skos ” for crescent-shaped and the Latin word “ spina ” for spine.

Type species — Menispina evidens Herrera, Manchester, Hoot, Wefferling, Carvalho et Jaramillo sp. nov.

Holotype hic designatus — BF19-ING-0014 ( Fig. 8 ).

Paratypes — BF19-ING-0012 ( Fig. 10 ), BF19-ING-0015 ( Fig. 9 ), BF19-ING-0016 ( Fig. 11 ).

Description — Impressions and casts of four crescent and horseshoe-shaped endocarps; length 3.0 – 4.3 mm, mean 3.7 mm (SD = 0.53, N = 4), width 2.5 – 4.3 mm, mean 3.4 mm (SD = 0.8, N = 4); endocarp wall thickness ~0.6 – 1.0 mm (measured from the locule cast to the apex of the endocarp); outline of endo-carp elliptic, circular and obovate in lateral view with 13 to 16 conspicuous, stiff, acute spines on the dorsal and lateral crests; the longest dorsal spines are at least 1.5 mm in length; the thickest dorsal spine bases are mostly located near the stylar and pedicel ends (best seen in the holotype and paratype BF19-ING-0015, Figs. 8, 9 ); dorsal crest with one side broader (toward stylar end) than the other; condyle outline obovate and elliptic; central locule cast extending to the base and with equal limbs, marked with 11 to 14 radially aligned ribs and with rounded pits or grooves (probably spine bases); locule cast limbs closely spaced; apparently lacking a perforation, but with a basal sculpturing (manifested as a small concavity) near shorter limb; ventral notch straight-margined; pedicel short (~0.5 mm).

Derivation of specifi c epithet — From the Latin “ evidens ” : evident, clear.

Age, source, and stratum — Middle to late Paleocene. Bo-got á Formation, Cundinamarca state, Nemoc ó n locality, Checua mine. Holotype and paratypes BF19-ING-0014-16, locality FH0801, 5 ° 0.8 ′ 24.9N, 73 ° 50 ′ 80.4W. Paratype BF19-ING-0012, locality FH0806, 5 ° 0.8 ′ 14.5N, 73 ° 50 ′ 80.2W.


5 ° 0.8 ′ 41.5N, 73 ° 50 ′ 58.3W. Paratype BF20-ING-0018 comes from GPS location 5 ° 0.8 ′ 23.44N, 73 ° 50 ′ 27.54W.

Systematic affi nity within Menispermaceae — In the molec-ular scaffold analysis, Palaeoluna bogotensis is placed within the S - C - C clade ( Fig. 50 ). One of the main morphological char-acters supporting this placement is the presence of strongly un-equal locule limbs ( Fig. 49 ). Although this character is not synapomorphic for the S - C - C clade, it appears to be diagnostic in comparison with the remaining clades in the family that ex-hibit horseshoe-shaped endocarps with conspicuous locule ribs. The strongly unequal limbs combined with other characters (e.g., conspicuous locule ribs, endocarp length less than 10 mm, etc.) make this group recognizable. Within the S - C - C clade, Cissampelos has the closest morphological similarity with the fossil taxon ( Figs. 29 – 33 ); the almost J-shaped locules in this extant genus are very similar to those in P . bogotensis ( Figs. 30, 33 ). On the basis of these morphological similarities, we sup-port Palaeoluna as an extinct genus within the S - C - C clade.

N = 4); endocarp wall thickness ~1.0 – 2.1 mm (measured from the locule cast to the apex of the endocarp); impression speci-mens of the interior side of the dorsal crest preserve conical pits indicating the presence of narrow, spine-like structures; one lat-eral crest on each side; condyle outline ovate in shape; central locule cast with unequal limbs (appearing as a J-shaped locule), ornamented by 15 to 16 radially aligned ribs; opposite to the locule ribs are spine-like structures that correspond to inner protrusions of the locule chamber; locule cast limbs widely spaced, ~1.1 mm (measured from tip of the short limb to tip of the longer limb).

Derivation of specifi c epithet — From the Bogot á Formation, where the fossils were found.

Age, source, and stratum — Middle to late Paleocene. Bo-got á Formation, Cundinamarca State, Nemoc ó n locality, Checua mine. Holotype and paratypes BF20-ING-0019, BF20-ING-0024, and BF20-ING-G0026 are from locality FH0918,

Figs. 18 – 48. Extant Menispermaceae endocarps. Figs. 18 – 20. Stephania brevipes Craib (A96-895). 18. Lateral view (LV), endocarp shape obovate with a prominent perforation, arrow: ornamentation. 19. Sagittal view (SV), arrow: apical locule/seed. 20. Sagittal view empty (SVE), arrow: straight ventral vascular tube. Figs. 21 – 23. S . delavayi Diels (US599329). 21. LV, arrow: broad dorsal crest. 22. SV, endocarp with an apical locule/seed (arrow). 23. SVE, arrow: tiny perforation. Figs. 24 – 27. S . japonica (Thunb) Miers (US3461000, A3485). 24. LV, arrow: ornamentation. 25. SV, arrow: longer locule limb of the stylar end. 26. SVE, arrow: straight ventral vascular tube. 27. LV of the seed, arrow: conspicuous ribs. 28. S . hernandiifolia (Willd.) Walp. (US3345848), arrow: straight ventral notch. Figs. 29, 30. Cissampelos andromorpha DC (US1997934, US2575572). 29. LV, endocarp exhibiting typical morphology for this genus, arrow: outermost lateral crest. 30. SV, arrow: longer seed/locule limb. Figs. 31 – 33. C . pareira L. (F2742). 31. LV, arrow: lateral crest spines. 32. SVE, arrow: inner locule ribs. 33. LV of the seed, notice longer limb and arrow: conspicuous ribs. 34. Diploclisia glaucescens (Blume) Diels (A1501), SV, hairpin-shaped endocarp, arrow: inner locule protrusions derived from intense exterior ribbing. 35. Cyclea atjehensis Forman (A90-33), LV, notice two lateral and two dorsal crests, arrow: conspicuous spines. 36. Cy . hypoglauca (Schauer) Diels (US1754790), SV, left arrow: longer locule/seed limb, right arrow: straight ventral vascular tube. 37. Cy . peltata Diels (US2616394), LV, arrow: ornamentation of outermost lateral crest. 38. Cy . polypetala Dunn (US458824), LV, endocarp showing two dorsal and two lateral crests, arrow: one of the lateral crests. 39. Cy. racemosa Oliv. (A663), SVE, arrow: straight ventral vascular tube. Figs. 40, 41. Pericampylus glaucus (Lam.) Merr. (A96-662). 40. LV, arrow: broad dorsal crest, also notice conspicuous spines. 41. LV of the seed ex-hibits conspicuous ribs (arrow) and two limbs of equal length. Figs. 42 – 45. Menispermum canadense L. (UWM15599, US371274). 42. LV, arrow: vasculature channel. 43. SV, notice central seed. 44. SVE, notice smooth locule chamber wall; arrow: entry of lateral vascular tube. 45. LV of the seed exhibiting two limbs of equal length and without ribs. Figs. 46 – 48. Sinomenium acutum (Thunb.) Rehder & Wilson (A24889, US1990264). 46. LV, arrow: spines on the dorsal crest. 47. SVE, notice smooth locule chamber wall; arrow: entry of vascular tube. 48. LV of the seed shows inconspicuous ribs. Scale bars = 0.5 mm.


Fig. 49. Morphological characters superimposed on a simplifi ed tree topology of Menispermaceae (only those extant clades with high support PP ≥ 95 and BS ≥ 80 were maintained, modifi ed from Hoot et al., 2009 ). See Appendix 2 and 3 for a complete list of the genera and characters states. * Cocculus appears paraphyletic. This simplifi ed DNA tree was then used as the “ backbone constraint ” for a molecular scaffold analysis. Photos of endocarps correspond to Stepha-nia brevipes Craib (A96-895); lowercase letters indicate some of the morphological characters mapped. Gray boxes highlight the extant genera that show the closest similarity to the new Paleocene endocarps from Colombia and Wyoming. Fossil taxa ( † ) included at the bottom of the fi gure for comparison.


Palaeoluna bogotensis also resembles species of the fossil genus Palaeosinomenium Chandler, originally described from the early Eocene of England ( Chandler, 1961 ). Palaeosinome-nium is typifi ed by a horseshoe-shaped endocarp with one loc-ule limb much longer than the other, radial ridges that continue over the lateral crest toward the perforation, and the presence of abundant and conspicuous ribs. The new fossil morphogenus Palaeoluna can be differentiated from Palaeosinomenium by the thick endocarp wall, the lack of radial ridges that continue over the lateral crest, and the presence of inner protrusions in the locule chamber. This last character is only found in the un-related hairpin-shaped endocarps of modern Diploclisia ( Fig. 34 ). Affi nity with extant Diploclisia can be ruled out, however, based on differences in endocarp size, the shape of the locule chamber, and the ornamentation.

Chandler (1961) suggested Sinomenium and Menispermum as the extant genera most closely related to Palaeosinomenium . How-ever, after examining these two extant genera we question this af-fi nity. In the Sinomenium - Menispermum clade, the locule ribs are fairly inconspicuous ( Figs. 42 – 48 ). Also, in Palaeosinomenium the radial ridges may continue over the lateral crest toward the perfora-tion, not characteristic of Sinomenium or Menispermum . In addi-tion, the locule area is completely smooth and unribbed as observed in living and fossil taxa ( Figs. 42 – 48 ; Liu and Jacques, 2010 ).

We also placed an endocarp from the middle-late Paleocene (Tiffanian) Fort Union Formation near Linch, Wyoming, within the new genus Palaeoluna ( Fig. 17 ). The endocarp shows a strong similarity to the Colombian taxon, having an outline obovate in lateral view, strongly unequal (J-shaped) and widely spaced limbs, and the locule chamber has abundant inner protrusions.

Figs. 50, 51. 50. Strict consensus tree of morphological data analyzed under the backbone constraint shown in Fig. 49 , bootstrap values given for nodes > 50%; fossil taxa ( † ) are highlighted with a gray shaded box, solid circles correspond to the backbone constraint. See Appendix 3 for a complete list of the genera. * Cocculus appears paraphyletic. 51. Fossil locations and geographic distribution of three extant genera in Menispermaceae that are related to the Paleocene endocarps. Modern distribution based on literature (e.g., Thanikaimoni, 1986 ) and biodiversity occurrence data accessed through GBIF Data Portal, data.gbif.org, 2010-10-15. Base map courtesy of National Aeronautics and Space Administration Jet Propulsion Laboratory, California Institute of Technology.


DISCUSSION AND IMPLICATIONS FOR MENISPERMACEAE

The new menispermaceous endocarp taxa described in this study augment the list of fossils recently identifi ed from the Pa-leocene forests of northern South America. Other components of the Cerrej ó n fl ora identifi ed to extant taxa below the family level include the aroid Montrichardia ( Herrera et al., 2008 ), members of the palm subtribes Euterpeinae and Attaleinae ( G ó mez-Navarro et al., 2009 ), and leaves of Eumalvoideae, Malvaceae ( Carvalho et al., 2011 ). The new fossils reconfi rm the presence and diversity of Menispermaceae in the middle-late Paleocene of Colombia. Lianas are the predominant habit for extant Menisper-maceae and are typical of rainforests. The Cerrej ó n and Bogot á paleofl oras have all the characteristics representative of this biome ( Herrera et al., 2008 ; Head et al., 2009 ; Wing et al., 2009 ); hence, it is not surprising to recover this family in the fl ora.

The fossil leaves and endocarps from the Paleocene of Co-lombia constitute the earliest known record of this family in northern South America. The distinctive endocarps of this fam-ily seem to be lacking from the fl ora of the late Cretaceous (Maastrichtian) Guaduas Formation, which is stratigraphically beneath the Bogot á Formation (currently under study, C. Jaramillo and F. Herrera, unpublished data), so it is possible that the fam-ily was introduced to this region later toward the end of the Cretaceous or in the Paleocene during the interval immediately preceding the time in which the Cerrej ó n and Bogota fossils were deposited.

Modern neotropical rainforests are distinctive in family compo-sition ( Gentry, 1988 ; Terborgh and Andresen, 1998 ) and Menisper-maceae is one of the common elements in this biome. How Menispermaceae originated or reached South America is still un-known. There is little doubt that vicariance may have played an important role in the history of the family; however, based on the available fossil record, dispersal events and/or local extinctions

seem to explain better the current distribution for some clades of Menispermaceae. Here, we propose four hypothetical scenarios that explain the occurrence of these fossil taxa:

(1) Dispersal via Laurasia — The former presence of the genus Stephania , which grows today in equatorial Africa and Australasia ( Fig. 51 ), in Paleocene neotropical rainforests, indicates a broader distribution of this genus in the past. It is possible that this genus reached the neotropics by dispersal from Africa and/or Asia via Laurasia ( Fig. 52 ). The absence of Stephania endocarps of Paleo-cene age or older ages from Old World fl oras could also support a neotropical origin with subsequent local extinction of this genus. Paleobotanical localities of Paleocene age are too poorly repre-sented in Africa and Southeast Asia to refute the Laurasian hypoth-esis. However, the presence of Stephania endocarps in the Messel fl ora (~47 Ma) from the middle Eocene of Germany (Collinson et al., in press) supports this route of dispersion.

(2) Transpacifi c dispersal of Stephania between South America and Australasia ( Fig. 52 ) — There is increasing evi-dence supporting transpacifi c dispersal events in several extant plant groups ( Michalak et al., 2010 ; Renner et al., 2010 ), which makes this route seem plausible. Another fossil from the Cer-rej ó n Formation assigned to the modern palm genus Nypa shows a connection with southeastern Asia ( G ó mez-Navarro et al., 2009 ). Also, new fossil fruits from a late Eocene locality of Panama (Tonos í , Azuero Peninsula) show affi nities with ex-tant Australasian genera (F. Herrera and S. Manchester, per-sonal observation).

(3) Exchange of Stephania via Antarctica ( Fig. 52 ) — A Pa-leocene dispersal event via Antarctica is also probable. This route of exchange lasted until the opening of the Drake Passage and the resulting glaciation in the late Eocene ( Scher and Martin, 2006 ). This is also supported by several plant lineages of

Fig. 52. Hypothesized routes of dispersal for the Paleocene endocarps from Colombia and North America. Black star indicates the Bogota and Cer-rej ó n fl oras from Colombia; white star indicates the Fort Union Formation, Linch, Wyoming, USA. Paleocene map modifi ed from Scotese (2001) .


extant Australasian affi nity (e.g., Eucalyptus , Gymnostoma , and Papuacedrus ) together with menispermaceous leaves and endo-carps found in Eocene fl oras of Argentina ( Zamaloa et al., 2006 ; Iglesias et al., 2007 ; Wilf et al., 2009 ; Gandolfo et al., 2011 ).

(4) Caribbean exchange — Menispermaceae was also pres-ent in mid-latitude North America at about the same time as the deposition of the Colombian endocarps. Examples include cf. Canticocculus Chandler from the upper Paleocene of North Dakota, United States ( Crane et al., 1990 ), and the aforementioned specimen of Palaeoluna from the middle-late Paleocene of the Fort Union Formation, Wyoming ( Fig. 17 ). The morphological similarity between the latter specimen and those of Palaeoluna bogotensis is striking and suggests that phytogeographic ex-change occurred between these two regions during or perhaps immediately prior to the middle-late Paleocene. This phytogeo-graphic exchange could have been favored when the Caribbean volcanic arc briefl y connected South and North America during the Paleocene ( Fig. 52 ; Bayona et al., 2010 ; Cardona et al., 2010 ; Buchs et al., 2011 ). Palaeoluna is placed within the ex-tant S - C - C clade ( Fig. 50 ), and it shows close similarity to ex-tant Cissampelos ; this modern genus has a very widespread distribution ranging from North America, the neotropics, Af-rica, and Australasia ( Fig. 51 ). If Palaeoluna represents the an-cestor of Cissampelos , South and North America rise as candidates for the ancestral area of this lineage.

As yet, the place of origin of the Menispermaceae remains uncertain, although Thanikaimoni (1984 , 1986 ) suggested the Cretaceous lowland forests of Africa. Interestingly, Cretaceous fossil records for Menispermaceae remain unsubstantiated. The fossil genus Prototinomiscium Knobloch and Mai ( Knobloch and Mai, 1986 ) from the upper Cretaceous of Europe has been uncritically accepted as a valid record of Menispermaceae for molecular dating analyses (e.g., Anderson et al., 2005 ), although the endocarp fossil shows a surface cell pattern unlike any known for the family. In our opinion, the assignment of this fossil to Menispermaceae is dubious. Also, many fossil leaves called Menispermites from Cretaceous fl oras of North America and Europe remain uncertain as to their affi nity ( Jacques, 2009b ).

The menispermaceous diversity now recognized in the Pa-leocene of Colombia indicates that South America may have played an important role early in the radiation and dispersal of the family after the Cretaceous-Paleogene boundary, where the derived S - C - C clade was already present, and it was relatively abundant in the earliest neotropical rainforests. This pattern of radiation is also observed in the Paleocene of Colombia in Fa-baceae, Malvaceae, Araceae, Arecaceae, and Annonaceae ( Herrera et al., 2008 ; G ó mez-Navarro et al., 2009 ; Wing et al., 2009 ; Carvalho et al., 2011 ). Recent molecular divergence time estimates for Menispermaceae ( Jacques et al., 2011 ) suggest an early middle Eocene age of origination for the S - C - C clade ( < 53.4 Ma); however, the new fossils from Colombia suggest an older Paleocene age (~60 Ma).

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Appendix 1. Extant species of Menispermaceae photographed and analyzed for endocarp morphology.

Species name Herbarium no. Region Altitude (m a.s.l.)

Antizoma calcarifera Miers US1991350 South Africa — Cissampelos andromorpha DC. Coll. Fernandez, A. US1997934 Colombia — Cissampelos andromorpha DC. Coll. Mexia, Y. BARC13596 Ecuador — Cissampelos andromorpha DC. Det. Barneny, R. US2575572 Suriname 350 Cissampelos fasciculata Bentham. Coll. Maxon, W. US1181670 Costa Rica 1600 Cissampelos grandifolia Tr & Pl. Coll. Mexia, Y. US1663781 Ecuador 650 Cissampelos grandifolia Tr & Pl. Coll. Skutch A US1642714 Costa Rica 1040 Cissampelos pareira L. Coll. Cook, F. & Griggs, R. US407831 Guatemala — Cissampelos pareira L. Coll. George, P. BARC G011 Guatemala — Cissampelos pareira L. Coll. Morales, J. F2742 Costa Rica — Cissampelos tropaeolifolia DC. Coll. Mexia, Y. & Yaveo, T. BARC130993 Mexico — Cissampelos tropaeolifolia DC. Coll. Encarnaci ó n, F. US3011876 Peru — Cocculus carolinus L. (DC). BARC19889 US — Cocculus hirsutus Diels L. Coll. Barakrishnan, A. US2686633 Sri Lanka — Cyclea atjehensis Forman. Coll. Maxwell, J. A(90-33) Thailand — Cyclea hypoglauca (Schauer) Diels. Coll. Tsui, T.M. US1754790 China — Cyclea hypoglauca Diels. Coll. Tsui, T.M. BARC652 China — Cyclea peltata Diels. Coll. Saldanha C.J. US2616394 India — Cyclea polypetala Dunn. Coll. Henry, A. US458824 China — Cyclea racemosa Oliv. Chow, H. A(663) China — Dioscoreophyllum cumminsii (Stapf) Diels. BARC1970 Nigeria — Diploclisia affi nis Diels. Coll. Wilson, E.H. US599510 China — Diploclisia glaucescens (Blume) Diels. Coll. Chan, K. A(1501) Hong Kong — Jateorhiza macrantha Hooker. Coll. Fernandez, C. US3264726 Equatorial Guinea 240 Limaciopsis loangensis Engl, Coll. Louis, J. BARC7750 Belgian Congo — Menispermum canadense L. Coll. Harrisonburg. BARC94 DR US — Menispermum canadense L. Coll. Rhodes, D. US371274 US — Menispermum canadense L. Coll. Sylvester, C. UWM15599 US — Menispermum dauricum DC. BARC061791 — — Menispermum dauricum DC. Coll. Dorsett, M. BARC89750 China — Pericampylus glaucus (Lam.) Merr. Coll. Maxwell, J. A(96-662) Thailand — Pericampylus glaucus (Lam.) Merr. Coll. Wang, C. US1670049 China — Pericampylus incanus Lamk (Merr). Coll. Merril, E. & McC., F. BARC20102 — — Sinomenium acutum (Thunb.) Reh. & Wi. Coll. Bo. D. & Bart, B. A(24889) China — Sinomenium acutum (Thunb.) Rehder & Wilson. Coll. Lee, C. US1990264 China — Sinomenium acutum R. & W. Coll. Yat-Sen, C. & Szechwan. BARC114802 China — Stephania brevipes Craib. Coll. Maxwell, J. A(96-895) Thailand — Stephania capitata (Blume) Spreng. Coll. King. BARC5239 Malay Peninsula — Stephania delavayi Diels. Coll. Wilson, E.H. US599329 China 200 Stephania epigaea H.S Lo. Coll. Bartholomew, B. US3043598 China — Stephania hernandiifolia (Willd.) Walp. Coll. Huq. A. US3345848 Bangladesh — Stephania japonica (Thunb) Miers. Coll. Hartley, T. US3461000 New Guinea 1220 Stephania japonica (Thunb) Miers. Coll. Schodde, R. A(3485) Australia — Stephania rotunda Lour. Coll. SPI. BARC47804 India — Stephania rotunda Lour. Cur. Bot. Gard. Darjeeling. BARC94085 India — Tiliacora acuminata (Lam.) Miers. Coll. Jayasuriya, M. US2721394 Sri Lanka —

Although we studied 68 of the 71 extant genera, we only include here those for which the mesocarp tissue was removed to reveal endocarp characters. Herbarium acronyms follow Index Herbariorium ( http://sweetgum.nybg.org/ih/ ). Specimens from A include collector number in parentheses.


Appendix 2. List of morphological characters and states.

1. Endocarp type: horseshoe-shaped (0); straight (1); hairpin-shaped (2); reniform (3). 2. Endocarp Length (mm): small ( < 10 mm) (0); large ( > 10 mm) (1).3. Endocarp wall thick: yes ( > 1 mm) (0); no ( < 1 mm) (1).4. Outline endocarp shape: obovate (0); ovate (1); elliptic (2); rounded (3); other (4).5. Dorsal crest no.: 0 (0); 1 (1); 2 (2); ≥ 3 (3).6. Lateral crest no.: 0 (0); 1 (1); 2 (2); ≥ 3 (3).7. One dorsal crest broader: yes (0); no (1). 8. Spiny dorsal crest a : yes (0); no (1). 9. Spiny lateral crest a : yes (0); no (1).

10. Vascular tube course b : enters ventrally (0); enters laterally (1).11. Locule chamber position: apical (0); central (1); basal (2); along endocarp exterior (3). 12. One limb noticeably longer: yes (0); no (1). 13. Locule ribs conspicuous: yes (0); no (1).14. Ventral notch straight c : yes (0); no (1).

15. Perforation present d : yes (0); no (1). 16. Basal sculpturing e : yes (0); no (1).17. Locule inner protrusions: yes (0); no (1).18. Arborescent habit: liana or herbaceous vine (0); shrub or tree (1).19. Leaf venation: pinnate (0); actinodromous (1); acrodromous (2).20. Leaf peltate: yes (0); no (1).21. Leaf lobed: yes (0); no (1).22. Endosperm present: yes (0); no (1).23. Endosperm ruminate: yes (0); no (1).24. Stamen fusion: all stamens free (0); fi laments fused-anthers free (1); some stamens fused-others free (2); fusion of all stamens, anthers at least partially fused

and forming a ring-like structure (3).25. Pollen type: tricolpate (0); tricolporate (1); triporate (2); inaperturate (3).

a Characters 8, 9. Spines are defi ned here as a type of ornamentation that exceed or extend beyond the dorsal or lateral crests. b Character 10. Vascular tube course: direction of entry of the vascular tube into the locule; for example, in Stephania , the vascular tube enters ventrally

( Fig. 20 ), appearing as a straight tube. On the other hand, a lateral vascular tube enters the seed chamber/locule with some curvature either laterally or sublaterally; see Menispermum ( Fig. 44 ).

c Character 14. Ventral notch straight: lower ventral line of the endocarp that does not present any indentation or curvature ( Fig. 28 ). d Character 15. Defi ned as a complete perforation of the endocarp. This character is different from lateral apertures which are openings that do not

completely traverse the endocarp. e Character 16. Basal sculpturing. This character refers to basal concavities (observed on inner faces of the endocarp) and/or convex sculpturing (observed

on outer faces of the endocarp). Because of the uncertainty of the presence of either one of these characters in the fossil genus Menispina Herrera gen. nov. we have scored and combined them as basal sculpturing.

Characters in boldface are also presented in Fig. 49 .


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Documents

Herrera Et Al_2011_Paleocene Menispermaceae Endocarps From Colombia & Wyoming