XIII EJIP Conference Proceedings

Current Trends in Paleontology and Evolution Libro de resúmenes / Conference proceedings XIII Encuentro en Jóvenes Investigadores en Paleontología (XIII EJIP) XIII Meeting of Early-Stage Researchers in Paleontology (XIII EJIP) Cercedilla, 15 - 18 de Abril de 2015 Laura Domingo, M. Soledad Domingo, Omid Fesharaki, Blanca García Yelo, Ana Rosa Gómez Cano, Verónica Hernández-Ballarín, Daniel Hontecillas, Juan L. Cantalapiedra, Paloma López Guerrero, Adriana Oliver, Jonathan Pelegrín, Miriam Pérez de los Ríos, María Ríos, Óscar Sanisidro & Alberto Valenciano (Editors) Designed by Juan L. Cantalapiedra and Óscar Sanisidro “LEGO, el logotipo de LEGO y la Minifigura son marcas comerciales del Grupo LEGO. ©2015 The LEGO Group“ ISBN 978-84-606-7282-1 XIII EJIP Conference Proceedings

A new window to an old world: reconstructing the Middle Eocene of Messel for Hessisches Landesmuseum Darmstadt

Una nueva ventana a un viejo mundo: reconstruyendo el Eoceno medio de Messel para el Hessisches Landesmuseum Darmstadt Oscar Sanisidro1*, Norbert Micklich2 y Oliver Sandrock2

Las reconstrucciones paleontológicas son una potente herramienta para transmitir el conocimiento y/o hipótesis científicas al público general. El presente trabajo presentamos las ilustraciones científicas incluidas en la nueva exposición del Hessisches Landesmuseum Darmstadt (Alemania). El ecosistema de Messel se ha dividido en 6 hábitats diferentes con sus propias comunidades de fauna y flora. Estos son: la orilla del mar, una vista diurna del dosel arbóreo, una vista nocturna del mismo emplazamiento, el suelo del bosque, un claro en el bosque y las zonas pantanosas adyacentes.

Palabras clave: Eoceno, Messel, paleoilustración, reconstrucción anatómica, museografía.

Paleontological reconstructions are a powerful tool to transmit current scientific knowledge and/or hypotheses to the general public. Here we present the scientific illustration of the Messel fossil site included in the new exhibit of the Hessisches Landesmuseum Darmstadt (Germany). The Messel ecosystem has been divided into six different habitats with their own floral and faunal communities. These are: the maar’s shoreline and open water area, a diurnal view of the canopy, a nocturnal view of the same spot, the ground of the rainforest, a forest clearing, and the swampy neighboring areas. An additional overview of the entire Messel area was also created.

Keywords: Eocene, Messel, paleoillustration, anatomical reconstruction, museology.

Introduction review of the geological settings and biota found in the pit are described in Schaal and Ziegler (1992), Habersetzer and Schaal ocated about 8 km North-East of Darmstadt, the fossil (2004), Harms and Schaal (2005), Schaal (2005), or Gruber and site of Messel (Fossillagerstätte Grube Messel) is Micklich (2007). The long faunal and floral list of the Messel known worldwide for its exceptionally well preserved fossil site has been most recently tabulated by Morlo (2004), earliest Middle Eocene biota (Ypresian/Lutetian L Wilde (2004), and references included therein. boundary, 48.27 - 48.11 Ma). Messel Pit was a maar, a deep, tropical-subtropical lake partially embedded within a volcanic Material and Methods caldera surrounded of lush vegetation and a diverse fauna. The Institutional abbreviations: HLMD-Be = Behnke collection of high episodic mortality was probably produced by poisonous Hessisches Landesmuseum Darmstadt; FIS = Forschungsinstitut gases emitted periodically from the lake (Richter and Storch, Senckenberg, Frankfurt a.M.). 1980; Franzen and Köster, 1994). The anoxic conditions of Paleoenvironmental reconstruction: To create an overall the bottom favored the preservation of soft tissue structures aerial view of the Messel area, we first translated the topographic (e.g., Schaal et al. 1988; Koenigswald and Storch, 1998) and profiles published by Schaal and Ziegler (1992) into a 16-bit prevented the decay of the carcasses at the lake floor. Anaerobic grayscale readable digital elevation model (DEM). Subsequently, bacteria were mainly responsible for the soft tissue preservation. the 2D DEM file was converted into a 3D topographic relief with Over 100 years of excavations of the bituminous claystone the software e-on Vue 7 XStream. Finally, procedural-generated deposits have yielded an overwhelming body of knowledge trees (with a mean height of 30 m), clouds and water were added about the Eocene ecosystems from Central Europe. A general to the scene. The scene was not included in the final exhibit and view of the taphonomic background together with an exhaustive

1Museo Nacional de Ciencias Naturales – CSIC. C/Pinar, 25. 28006, Madrid (Spain). *Email: [email protected] 2 Hessisches Landesmuseum Darmstadt, Department of Natural History. Friedensplatz 1, 64283 Darmstadt (Germany).

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Figure 1. Paleoenvironmental reconstruction of the canopy during the daytime of the Messel Pit Fossil site (Middle Eocene) exhibited in the Hessisches Landesmuseum Darmstadt (Germany). Drawing by OS.

is planned for future educational purposes. It was rendered in illustrations and species represented, the taxa depicted in each 8-bitcolor, a size of 60 (w) x 33 (h) cm, a resolution of 28,346 scenario are easily seen and densely packed, ranging from 15 to pixels/cm, and saved in JPEG format. 31 species (including both fauna and flora). They were selected The remaining six illustrations, which represent particular according to three criteria: its relative abundance in the fossil habitats of the Messel area and pertain to the new museum site, its scientific relevance, and the presence of corresponding exhibit, were made using the software Adobe Photoshop CS specimens on display in the new exhibit. 5 (which allows multi-layered rendering - indispensable for In paleontology, scientific illustration is the only means preparing complex drawings) together with the digital pen tablet, available to visualize extinct landscapes and organisms. Wacom Intuos3. Photographs taken by OS in Kalimantan Timur The usually vast amount of scientific knowledge on which a and Sumatra (Indonesia) during 2011 and 2013 expeditions paleontological reconstruction relies must be synthetized to were used in order to obtain graphic references of extant maars simplify communication to the general public. That implies and tropical rainforests. that, unlike scientific drawing of living biological organisms, paleontological reconstruction of a fossil species is a direct Results and discussion reflection of the scientific conceptions at that moment andis Six different habitats were selected in order to show the broad subject to later updates. The methodological approach used diversity of the Messel maar and its neighboring surroundings. to achieve a reliable reconstruction of a fossil species differs These are (ordered according to their proximity to the maar): according to the group of study. the maar’s shoreline and open water area, a diurnal view of Because plants play a key role in tropical and equatorial the canopy (see figure 1), a nocturnal view of the same spot, forests, it is important to understand the vegetation structure the ground of the rainforest, a forest clearing, and the swampy of extant tropical forests as well as stress the main differences neighboring areas. The sizes of the drawings range from 85 (w) with those from the Eocene times in order to represent a realistic x 80 (h) cm to 210 (w) x 110 (h) cm, and have a resolution of paleoenvironmental reconstruction. Wilde (2005) linked the 28,346 pixels/cm. They were prepared in color. floral assemblage found in Messel with extant paratropical forests Despite their diversity, vertebrates living in tropical (according to the category defined by Wolfe, 1979). These forests forests are often elusive and represent but a fraction of the are dominated by macrophyllous trees. However, the late Eocene total animal biomass. Nevertheless, in order to enhance the floras from Central Europe (i.e. Messel, Kučlín, and Lábatlan) didactic component of the work and optimize the number of clearly differ in leaf categories, as they are dominated by noto-

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to microphyllous vegetation (Kvaček and Wilde, 2010). This and insertion of muscles is determined by rough areas, crests, particularity led Kvaček (2010) to erect a new vegetation unit for and ridges on the bone. However, muscle scars in the bones the extinct European Paleogene-Eocene vegetation, the “Mid- are not always evident and their determination often requires Latitude Broad-Leaved Notophyllous Evergreen Forests”. These the recognition of homologous relations with extant species. forests were dominated by Lauraceae, Juglandaceae, Ulmaceae, More external muscles are subsequently added layer by layer Leguminosae, and some coniferous genera (e.g. Doliostrobus). configuring the general outline of the body. The last step requires A total of 96 flowering plant families have been found in Central adding additional soft tissues (such as cartilage, fat, and, finally, Europe so far, a small part of the original diversity present at that skin) to the muscular reconstruction. Direct evidence of these time. Pollen, flowers, fruits, seeds, leaves, and a comparatively tissues are unknown in the vast majority of fossil species and small amount of branches have been unearthed. Despite sharing their reconstruction (along with color selection) is the most some common taxa, the species composition of each drawing speculative step of the whole process. Fortunately, hair and reflects the particular habitat conditions. For example, the edge feather cover is preserved in the Messel fossil site as a thin and of a forest clearing is expected to be full of climbing vines such dark layer of siderite and/or pyrite deposited over the bacteria that as grapes, moonseeds, and rattan palms trying to profit from the coated it (so-called Hautschatten or ‘skin shadow’). This dark sunny spot. On the other hand, the ground of a fully-grown forest outline not only gives information regarding length and density area would have been clear of vegetation but full of dead leaves of the hair or feather cover but, in some cases, contrasting dark of the trees above (such as walnuts, legumes, or laurels). Their and light patterns (e.g. HLMD-Be 178b), provide a transformed foliage would have housed a great diversity of both vertebrates reflection of the original color. and invertebrates (represented in the figure 1). Besides the exceptionally preserved faunal and floral Invertebrates are one of the most abundant groups in Messel. remains, the high informative resolution of the fossils provides Insects are usually not preserved in the fossil record. The direct evidence about the ecologic interactions. A selection of exceptions are the Konservat-Lagerstätte, like the Messel fossil some of these interactions has been included in the presented site, where the fossil insects are abundant and even show highly landscapes, divided into plant-insect associations and vertebrate colorful patterns. When reconstructing fossil insects, the overall gut-dentition contents (following Labandeira et al. 2011). Plant- morphology is determined by the wings and/or the exoskeleton insect associations include heliozelid damage on Laurophyllum, of the species. The level of detail preserved in the compressions nepticulid mine on dicot, armoured scale insects on dicots, usually depends on the amount of heavy-chitinous components circular holes on Lauraceae, or chew marks on Araciphyllites of the species: while stag beetles (Lucanidae) or jewel beetles tertiarius. Vertebrate gut-dentition contents are represented (Buprestidae) have hard elytra and are often represented by by seed feeding on the artiodactyl Messelobunodon schaeferi, finely-preserved fossils (e.g. HLMD-Be 104 or HLMD-Be 117) Lauraceae in gut contents of the primitive horse Eurohippus others like reduviid hemipterans (e.g. FIS coll. –nr. Mel 7203) messelensis, plant ingestion by the rodent Ailuravus macrurus, sometimes present more blurred outlines. Unfortunately, despite or moth predation by the bat Palaeochiropteryx. Additionally, the color patterns observed in Messel’s insects, which truly some reproductive behaviors, such as pregnant females of resemble distinct pigmentations or hue variations, the original Eurohippus messelensis or an ovoposition by an alticine beetle colors in life were altered during fossilization (McNamara have also been included. et al. 2011). Therefore, depicted colorations should be taken with caution as studies dealing with color reconstruction are Conclusions still scarce and limited to taxa with preserved color-producing Messel Pit Fossil site is of paramount importance for the nanostructures (e.g. ditrysian moth; McNamara et al. 2011). comprehension of the fauna and flora of the Middle Eocene, Vertebrate fossil reconstruction emerged with the work as demonstrated by its inclusion in the World Heritage Site of of Charles R. Knight (Antón and Sánchez, 2004). Since fossil biotas. The collections extracted from the fossil site have then, fossil vertebrate reconstruction has proliferated and generated the bulk of our scientific knowledge regarding various attained a highdegreescientific accuracy and methodological aspects such as systematics, taxonomy, ecology, biodiversity, or standardization. The work can be divided into a series of stages geology. As most fossils were produced from extinct organisms (summarized in Antón and Sánchez, 2004): firstly, missing that no longer exist, illustrations are an essential expositive bones and/or bony parts are completed using the closest taxon resource in paleontology. The illustrations of the Messel biota available with the needed area preserved. Consequently, more for the new Darmstadt exhibit can assist as a last link in the distantly related reference species can give rise to larger errors. process of translating the scientific discourse for society in a This step can be largely omitted in the case of fossil Lagerstätten direct and visual way. like Messel, where vertebrate skeletons are habitually nearly complete and found in anatomical position. The second step involves the reconstruction of muscular masses. The origin

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Acknowledgments 322 pp. Schaal S (2005). Messel Pit Fossil Site – Snapshots from the We thank Francisco Gascó, Mary Parrish, and the editorial Eocene, Vernissage. UNESCO World Heritage Sites Series, board for their valuable suggestions and comments. Heidelberg, Vernissageverlag, 21(5), 68 pp. References Wilde V (2004). Aktuelle Übersicht zur Flora aus dem mitteleozänen “Ölschiefer” der Grube Messel bei Darmstadt Antón M, Sánchez I (2004). Art and science: the methodology (Hessen, Deutschland). Courier Forschungsinstitut and relevance of the reconstruction of fossil vertebrates. Senckenberg, 252: 109–114. Miscelánea en Homenaje a Emiliano Aguirre. Paleontología, Wilde V (2005). The green Eocene. The diverse flora of a 2(4): 2–22. paratropical climate. Vernissage UNESCO World Heritage Franzen JL, Köster A (1994). Die eozänen Tiere von Messel - Sites Series 21(151): 14–19. ertrunken, erstickt oder vergiftet? Natur und Museum, 124: Wolfe, JA (1979) Temperature parameters of humid to mesic 91–97. forests of eastern Asia and relation to forests of other Gruber G, Micklich N (eds) (2007). Messel, treasures of the regions of the northern hemisphere and Australasia. United Eocene. The book to the exhibition »Messel on Tour« of States Geological Survey, Professional Paper, United States the Hessisches Landesmuseum Darmstadt. Hessisches Government Printing Office, Washington, 1106 pp. Landesmuseum Darmstadt, 159 pp. Habersetzer J, Schaal S (eds) (2004). Current geological and palaeontological research in the Messel Formation. Courier Forschungsinstitut Senckenberg, 252: 1–245 Harms F-J, Schaal S (eds) (2005). Current geological and palaeontological research in the Messel Formation. Courier Forschungsinstitut Senckenberg, 255: 1–236. Koenigswald WV, Storch G (eds) (1998). Messel, ein Pompeji der Paläontologie, Thorbecke, Sigmaringen, 152 pp. Kvaček Z (2010) Forest flora and vegetation of the European early Palaeogene – a review. Bulletin of Geosciences, 85(1): 3–16. Kvaček Z, Wilde V (2010) Foliage and seeds of malvacean plants from the Eocene of Europe. Bulletin of Geosciences, 85(1): 163–182. Labandeira CC, Dunne JA, Williams RJ (2011). The Messel food web. The World at the Time of Messel, 22nd International Senckenberg Conference, Frankfurt, 15-19 November, 95- 97. McNamara ME, Briggs DEG, Orr PJ, Wedmann S, Noh H, Cao H. (2011) Fossilized Biophotonic Nanostructures Reveal the Original Colors of 47-Million-Year-Old Moths. PLoS Biology, 9(11): e1001200. doi:10.1371/journal. pbio.1001200 Morlo M, Schaal S, Mayr G, Seiffert C (2005) An annotated taxonomic list of the Middle Eocene (MP 11) Vertebrata of Messel. Courier Forschungsinstitut Senckenberg, 252: 95- 108. Richter G, Storch G (1980). Beiträge zur Ernährungsbiologie eozäner Fledermause aus der Grube Messel. Natur und Museum, 110: 353–367. Schaal S, Ziegler W (eds) (1988). Messel – Ein Schaufenster in die Geschichte der Erde und des Lebens, Kramer, Frankfurt am Main. 315 pp. Schaal S, Ziegler W (eds) (1992). Messel – An Insight into the History of Life and of the Earth, Clarendon Press, Oxford,

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