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The Middle Eocene plant taphocoenosis from Eckfeld (Eifel, Germany)
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Review of Palaeobotany and Palynology 101 (1998) 7–28
The Middle Eocene plant taphocoenosis from Eckfeld (Eifel, Germany) Volker Wilde a,Ł , Herbert Frankenha¨user b b
a Forschungsinstitut Senckenberg, Pala ¨ obotanik, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz, Reichklarastr. 10, D-55116 Mainz, Germany
Received 10 February 1997; revised version received 14 August 1997; accepted 9 September 1997
Abstract More than 15,000 plant macrofossils have been collected during scientific excavations in an isolated occurrence of organic-rich clay=silt (‘oilshale’) with coarse turbiditic intercalations at Eckfeld near Manderscheid (Eifel, Germany). The sediments were deposited in a meromictic maar lake, and have been dated by mammals as late Middle Eocene (late Geiseltalian, MP13). Initial studies on leaves, fruits=seeds and flowers confirm the considerable diversity of the taphoflora indicated by about 200 ‘species’ of pollen and spores. The macrofossils suggest an interpretation as a zonal flora surrounding a more or less isolated inland lake with considerable inclination of the slopes. The assemblage has a number of taxa in common with the slightly older taphocoenosis of Messel, but with the exception of the ferns there are no species common to the Middle Eocene of the Geiseltal area. This may be explained by the regional palaeogeography with the Geiseltal area as a near-coastal peat-forming lowland, while Eckfeld and Messel were lakes at some distance from the coastal lowlands. 1998 Elsevier Science B.V. All rights reserved. Keywords: Eocene; Germany; lake; palaeobotany; palaeoecology; taphonomy; volcanism
1. Introduction The German Middle Eocene taphofloras of the Geiseltal area and of Messel have been known for a long time and studied in considerable detail. Other nearby plant assemblages dating from the same time interval like those of the Weißelster Basin and Helmstedt, have only been investigated in a preliminary fashion. Another locality with plant remains of Tertiary age was noted for the first time in the middle of the last century (Weber, 1853) in the vicinŁ Corresponding
author.
ity of Eckfeld near Manderscheid (Eifel, Germany; Fig. 1). The plant remains mentioned by Weber (1853) were never figured or described, and the material was apparently lost. Due to the excavations carried out by the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz since 1987 (Neuffer et al., 1996), the Eckfeld locality has become a well known fossillagersta¨tte, and was finally dated as Middle Eocene. Due to their palaeogeographic position, and the large number of macrofossils collected (>15,000), the plant assemblage from Eckfeld is of major importance in reconstructing the Middle Eocene vegetation of Germany. There was considerable taxonomic
c 1998 Elsevier Science B.V. All rights reserved. 0034-6667/98/$19.00 PII S 0 0 3 4 - 6 6 6 7 ( 9 7 ) 0 0 0 6 7 - 5
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Fig. 1. Palaeogeographic situation of the Middle Eocene of Western Europe (after Franzen, 1995), with localities mentioned in the text. Large dots D marine; fine stipple D land.
diversity as indicated by the presence of about 200 ‘species’ of spores and pollen (Nickel, 1996); knowledge on the macrofossils is, however, still limited. It is the aim of the present paper to give a more sophisticated overview of the present state of knowledge on the Middle Eocene macrofossils from Eckfeld, including some of its implications and comparisons with coeval plant assemblages. All the material is housed in the collections of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz.
2. Geological setting The Eifel is the German part of the outcropping Variscan Basement west of the river Rhine and north of the river Mosel. Most of the area is characterized by Devonian sediments with remnants of a Permian to Mesozoic cover (e.g. Meyer, 1994). Volcanic activity started in the Cretaceous and continued into Holocene times. As a result of the volcanic activ-
ity, there are numerous more or less circular volcanogenic depressions, today often occupied by a swamp or a lake. Traditionally, they have been called ‘maar’ by the local population. The term ‘maar’ is now internationally accepted for small monogenetic structures of phreatomagmatic origin with the base of the crater below the original surface (Lorenz, 1973; White, 1991). As reported for the first time by Weber (1853), an isolated occurrence of browncoal was discovered near Eckfeld in 1839, but mining activities had failed due to unfavourable rock conditions. The sediments were dated by Weber (1853) by comparison with some plant remains previously described from Oligocene localities near Bonn (Rott and Orsberg). The locality was for a long time overlooked. However, Pflug (1959) discovered a specimen of ‘coal’ from Eckfeld in the collections of the Geological Institute of the University of Cologne. Its sporomorphs were compared to the ‘Borkener Bild’ which at that time was thought to be of Late Eocene age. Based on new samples from the locality, the ‘Borkener
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Bild’ was later confirmed (Von der Brelie, 1969) at Eckfeld, but its range had by then been extended down into the Middle Eocene by mammal data from the type locality (Tobien, 1961). A Middle Eocene age (late Geiseltalian=MP 13) for the sediments exposed in Eckfeld by ongoing excavations was finally proven by a characteristic association of mammalian taxa (Franzen, 1994). The Tertiary sediments at Eckfeld were first explained by Weber (1853) as a filling of a primarily isolated maar-like depression. For some time, an interpretation as remnants of more extensive deposits (Pflug, 1959; Von der Brelie, 1969), possibly secondarily preserved in a volcanogenic depression (Von der Brelie, 1969), was favoured. Lo¨hnertz (1978) later recalled the maar hypothesis which was for the first time confirmed by drilling down into coarse pyroclastics (Negendank et al., 1982). Detailed geological and geophysical investigations recently gave final support to a maar origin for the Eckfeld structure (Pirrung, 1992b, 1993; Pirrung and Bu¨chel, 1994).
3. Sedimentary succession The maar structure at Eckfeld was cored to a depth of 66.5 m (Negendank et al., 1982). The sequence displayed by the core was roughly subdivided into basal pyroclastics followed by sideritic bituminous laminites and inorganic clay and silt. Within the bituminous laminites, a change from frequent diatomitic intercalations to non-diatomitic sediments was noted. The succession was interpreted as representing three successive stages of a maar lake. As a result of detailed field studies, the upper inorganic clay and silt is now interpreted as weathered bituminous laminites (Pirrung, 1992a). This is strongly supported by the mode of preservation of the plant remains with the organic material highly degraded or even missing. The Eckfeld structure is completely surrounded by a coarse marginal facies including varying proportions of volcanoclastic components (Pirrung, 1992a, 1993). Due to field conditions, scientific excavations in Eckfeld are still limited to a few meters of the nondiatomitic bituminous laminites which have been studied in detail. They are characterized by laminitic
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‘oilshales’ and siltstones (Mingram, 1994) with frequent turbiditic intercalations (Lutz, 1993a; Mingram, 1994). The organic fraction is mainly composed of fine-grained plant debris and the resistant remains of algae, for the most part chlorococcaleans like Botryococcus and Tetraedron (Wilde et al., 1993; Mingram, 1994). Apparently, plant macrofossils are more or less evenly distributed throughout the sequence under investigation. However, coarse and robust plant debris such as wood fragments and large fruits=seeds is concentrated in the turbiditic layers.
4. Physical setting of the flora The Palaeogene Eifel was traditionally thought of as a gently dipping peneplain devoid of any relief. Recently, a distinct magnetic anomaly with remnants of deeply eroded volcanic structures of Eocene to early Miocene age in the centre (‘Kelberger Hoch’) was interpreted as probably indicating major volcanogenic uplift north of Eckfeld (Bu¨chel and Pirrung, 1993; Pirrung and Bu¨chel, 1994) at that time. More evidence for some pre-Oligocene relief and contemporaneous drainage systems in the area was summarized by Lo¨hnertz (1994). Following Lo¨hnertz (1994), the Eckfeld maar today is located just between the completely eroded ‘Kelberger Hoch’ and an area with remnants of the Palaeogene surface. The Eckfeld maar and two adjacent volcanic structures are aligned along a NNE–SSW-trending major fault that can even be recognized in the Variscan basement (Meyer et al., 1994). After eruption(s) came to an end, the maar crater was occupied by a lake which soon became meromictic with algae flourishing in the mixolimnion (Wilde et al., 1993). Algal blooms were most probably seasonally controlled (Mingram, 1994). At least at times, the water may have been slightly alkaline, and some change in the water chemistry is possibly indicated by the disappearance of diatoms with time (Wilde et al., 1993). The occurrence of unionid bivalves may be taken as evidence that the lake at times was somehow connected to a fluvial system (Groh and Jungbluth, 1994). The dimensions of the Middle Eocene crater of Eckfeld have been estimated mainly from field data
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by Pirrung (1993). The diameter may have been approximately 850–950 m, with an inclination of the initial slopes of about 20º. Water depth never exceeded 170 m (Pirrung, 1993), and was probably not more than 50 m (Mingram, 1994). There is some evidence supporting some persistent instability of the slopes, and any shallows never reached significant extent (this paper).
5. The plant macrofossils Except for diatoms and chrysophycean cysts from the core (Zolitschka, 1993; Wilde et al., 1993; Mingram, 1994), all the plant material presently known from the Eckfeld site was collected during the excavations. As a consequence, all of the information on the plant macrofossils (leaves, fruits=seeds, flowers; Table 1 and Appendix A) presented below has been derived from a limited part of the succession. No attention has been paid as yet to the frequently occurring wood remains. After a first account of the plant remains had been published by Wilde (1989b), Wilde and Frankenha¨user (1993) presented a more extensive summary (for fruits=seeds see also Ja¨ckel and Frankenha¨user, 1994). It includes a list of taxa that was recently corrected by Wilde (1995). Detailed work on selected taxa started with algae (Wilde et al., 1993), bryophytes (Wilde, 1990), and ferns (Frankenha¨user and Wilde, 1993b). More recently, some angiosperm taxa have been studied, including pterocaryoid and engelhardioid juglandaceous fruits (Frankenha¨ user and Wilde, 1994; Wilde and Frankenha¨user, 1995), possible fruits of Burseraceae (cf. Canarium Linne´; Mai, Wilde and Frankenha¨user, in prep.), and a distinct type of scleromorphic dicotyledonous leaves (Pungiphyllum waltheri Frankenha¨user et Wilde, 1995). 5.1. Cryptogams Fossil bryophytes in general are rare. Surprisingly, there are now more than 40 specimens existing from Eckfeld. Most of them apparently represent a distinct type with tufts of leafy stems still adhering to a piece of wood or bark (Plate I, 2; Wilde, 1990) thus indicating an epiphytic habit. Except for
one specimen (Plate I, 1), there are no sporophytes preserved. Most remarkable is a single tiny fragment of a Frullania-like jungermannialean leafy liverwort that may have been epiphyllous (work in progress). There is a suite of ferns in Eckfeld that is common to other Middle Eocene localities like Messel and the Geiseltal area (Frankenha¨user and Wilde, 1993b; Wilde, 1995). Most common in Eckfeld are Schizaeaceae with Lygodium kaulfussii, including a considerable number of fertile pinnae fragments (Plate I, 6,7), and a single sterile frond fragment of Ruffordia, an extinct genus. In addition, there are some fragments of sterile pinnae of Osmunda lignitum (Osmundaceae; Plate I, 8), and a single fertile frond fragment of ‘Rumohra’ recentior (Polypodiaceae s.l.; Plate I, 3, 4). Most recently, a fragment of a fertile pinna was found that is apparently quite similar to extant Didymochlaena or Lindsaea (Polypodiaceae s.l.; Plate I, 5; work in progress). An interesting fact is the common occurrence of pinnae fragments of the ‘mangrove fern’ Acrostichum (Polypodiaceae). At present the genus is more or less restricted to distal mangrove-related habitats. Its occurrence in the sediments of Eckfeld is yet another example for a shift in the ecological amplitude since Palaeogene times. 5.2. Gymnosperms As in most European Tertiary localities, remains of gymnosperms are limited to conifers in Eckfeld. There are a single badly preserved twig fragment and a possible cone scale (Plate IV, 1) of taxodiaceous affinity. Cephalotaxus cf. messelensis is represented by an isolated leaf with diagnostic cuticular structures. Another unique specimen is the terminal part of a cupressaceous branch with fused facial and marginal leaves and Thuja-like cuticular structures. Because fusion of facial and marginal leaves is unknown from extant taxa as previously noted by Ferguson (1971), the specimen is assigned to Libocedrites salicornioides (Unger, 1841) Endlicher, 1847 (manuscript in preparation). 5.3. Monocotyledons There are a number of monocotyledonous leaf fragments from Eckfeld, most of them of unknown
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Table 1 The plant macrofossils of Eckfeld after the 1995 campaign with taxa that have been assigned at least to family level Bryophyta Dicranites sp. Muscites sp. Jungermanniales sp. (cf. Frullania sp.) Filices Osmundaceae (C) Schizaeaceae Polypodiaceae s.l. (C)
Coniferae Taxodiaceae ? Cupressaceae (C) Cephalotaxaceae Dicotyledoneae Anacardiaceae (?) Betulaceae (C) Burseraceae (?) Caprifoliaceae Hamamelidaceae (?) Juglandaceae (C)
Lauraceae Mastixiaceae (C) Menispermaceae (C) Moraceae Myricaceae (C) Papilionaceae (D Leguminosae) (C) Rosaceae (?) Rutaceae (?) Sargentodoxaceae Theaceae Ulmaceae (C) Vitaceae (C) fam. inc. sed. Monocotyledoneae Araceae Arecaceae (D Palmae) (C)
Cyperaceae (C) Liliaceae s.l. (?)
Osmunda lignitum (Giebel, 1857) Stur, 1870 Lygodium kaulfussii Heer, 1861 (C) cf. Ruffordia subcretacea Barthel, 1976 (C) Acrostichum sp. ‘Rumohra’ recentior (Unger, 1841–1847) Barthel, 1976 fragment of a fertile pinna similar to Didymochlaenaand Lindsaea Filicopsida inc. sed. [?Blechnum dentatum (Goeppert, 1836) A. Braun, 1852] fragment of a leafy twig and single cone scale Libocedrites salicornioides (Unger, 1838) Endlicher, 1847 (Cephalotaxus cf. messelensis Wilde, 1989a) cf. Pentoperculum Tiffney, 1994 (fruits) Carpinus-like bracts Canarium n.sp. (fruits) Sambucus sp. (seeds) flower with valvate anthers engelhardioid compound leaves and leaflets engelhardioid fruits (Palaeocarya sp.) Hooleya-like pterocaryoid fruits fragments of platycaryoid male inflorescences cf. Laurophyllum sp. (leaves) cf. Daphnogene sp. (leaves) fruits 3 distinct types of fruits=seeds at least ?Ficus sp. sensu Wilde (1989a), leaves Comptonia sp. (leaves) pods and leaflets compound leaves of Rosa vel Rubus type seeds, at least 1 type cf. Sargentodoxa sp. (fruits) cf. Polyspora saxonica Walther et Kvaˇcek, 1984 (leaves) cf. Ternstroemites dentatus Wilde, 1989a (leaves) ulmoid leaves Zelkova-like leaves vitioid leaves and seeds Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (leaves) a single fragment of a leaf comparable to ‘Araceae sp. 2’ (sensu Wilde, 1989a,b) 1 distinct type of flower fragments of palmate leaves fragments of pinnate leaves fragments of stout spinose rhachises cyperaceous fruits cf. Smilax (leaves)
Families definitely or possibly also represented by spores or pollen are marked by ‘(C)’ and ‘(?)’, respectively (after Nickel, 1994, 1996; pers. commun., 1995)
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affinity. The Araceae are represented by a few fragments of a leaf type commonly occurring in Messel. The most frequent type (>130 specimens) of all flowers in Eckfeld is a trimerous palm flower (Plate IV, 2,3) with monocolpate pollen preserved in situ. Fragments of palm leaves are not rare, including both pinnate (Plate II, 12) and palmate organization. Furthermore, there are fragments of stout petioles with hook-like spines referable to the Arecaceae. Some shoots with sheathing bases of linear leaves probably ought to be interpreted as submerged Hydrocharitaceae (Plate II, 8). A few leaves are similar to Smilax (Smilacaceae), while Cyperaceae are known from a number of well preserved fruits (Plate V, 5).
5.4. Dicotyledons Dicotyledons are represented in Eckfeld by numerous leaves, fruits=seeds, and a considerable number of flowers. Juglandaceae are by far the most important element in the assemblage (Plate III) with a great number of leaves, leaf fragments, and leaflets of different types. The juglandaceous material is characterized by peltate glands with a unicellular base that are typical for the extant members of the family (Plate III, 10). Number and affinities of genera and species included in the Eckfeld material will be the subject of a future study. Hooleya-like fruits are common and were assigned to
PLATE I Cryptogams. Scale always 1 cm. 1. Moss with sporophytes and sporangia; PB 1995=364 LS. 2. Epiphytic moss still adhering to a fragment of bark; PB 1995=365 LS. 3. Frond fragment of ‘Rumohra’ recentior sensu Barthel (1976); PB 1995=276 LS. 4. Detail from 3 with fertile pinnae (arrows at sporangia). 5. Fertile pinna fragment of a fern similar to extant Didymochlaena or Lindsaea; PB 1995=274 LS. 6. Lygodium kaulfussii, fertile frond fragment with a number of sporangiophores; PB 1995=278 LS. 7. Lygodium kaulfussii, fertile frond fragment with a number of sporangiophores; PB 1995=277 LS. 8. Sterile pinna fragment of Osmunda lignitum; PB 1995=275 LS. PLATE II (see p. 14) Common leaf types. Scale always 1 cm. 1. Ternstroemites cf. dentatus Wilde, 1989a (Theaceae); PB 1995=208 LS. 2. Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (fam. inc. sed.); PB 1995=82 LS. 3. Compound rosaceous leaf with basal stipules; PB 1995=168 LS. 4. Ulmaceous leaf; PB 1990=374 LS. 5. Leaf of unknown systematic affinity with entire margin and three veins emerging from the base; PB 1995=207 LS. 6. Leaf of unknown systematic affinity with coarse undulate margin; PB 1995=178 LS. 7. Leaf of unknown systematic affinity with asymmetric base and irregular margin; PB 1995=198 LS. 8. Leafy shoot of an aquatic monocotyledon (?Hydrocharitaceae); PB 1995=195 LS. 9. Leaf of Comptonia sp. (Myricaceae), composite figure including part and counterpart; PB 1995=167 LS. 10. cf. Ficus sp. sensu Wilde (1989a,b); PB 1995=199 LS. 11. Vitioid leaf (cf. Vitaceae, Menispermaceae, Aceraceae, etc.); PB 1995=205 LS. 12. Fragment of a pinnate palm leaf (Phoenicites sp.); PB 1995=206 LS. PLATE III (see p. 15) Juglandaceae. Scale 1 cm except for 10, 11. 1. Entire large compound leaf with leaflets attached; PB 1990=59 LS. 2, 3. Isolated leaflets, type 1, with chewed margins; PB 1995=171, 172 LS. 4. Leaflet, type 2; PB 1995=175 LS. 5. Fragment of a small compound leaf with leaflets attached; PB 1995=176 LS. 6, 7. Small entire leaf with leaflets attached; PB 1995=174, 170 LS. 8. Leaflet, type 3, with compound teeth; PB 1995=173 LS. 9. Engelhardioid winged fruit (Palaeocarya sp.); PB 1995=69 LS. 10. Typical juglandaceous peltate trichomes from pterocaryoid fruit. Scale 100 µm. 11. Hooleya-like pterocaryoid fruit. Scale 0.5 cm; PB 1994=267 LS.
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PLATE I
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PLATE II
For description see p. 12.
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PLATE III
For description see p. 12.
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PLATE IV
Flowers. Scale 0.1 cm, except for 12. 1. ?Taxodiaceous cone scale. PB 1990=22 LS. 2, 3. Male palm flowers; PB 1995=389, 390 LS. 4. Platycaryoid male inflorescence; PB 1990=78 LS. 5–9. Different flowers of unknown affinity; PB 1995=391 LS, PB 1991=14 LS, PB 1990=83 LS, PB 1990=20 LS, PB 1990=346 LS. 10, 11. Probable flowers of Rutaceae; PB 1990=343 LS, PB 1995=392 LS. 12. Pollen from flower in 11. Epifluorescence, scale 20 µm.
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the pterocaryoid alliance (Frankenha¨user and Wilde, 1994). In comparison, only two specimens of engelhardioid winged fruits (Palaeocarya sp.) have been found (Wilde and Frankenha¨user, 1995). Furthermore, there are pollen-bearing male inflorescences, some of them probably platycaryoid (Plate IV, 4), which is apparently supported by a distinct type of pollen that has been observed using epifluorescence. Apart from juglandaceous foliage, there are a number of more or less common leaf types. However, at the moment only a few of them can be assigned to a natural taxon. For example, there are a great number of ulmaceous leaves (Plate II, 4). Another common leaf type is represented by Ternstroemites cf. dentatus (Theaceae) which was previously described from Messel (Plate II, 1). Quite characteristic are those leaves of unknown systematic affinity that have recently been described as Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (Plate II, 2). A number of large leaves of a vitioid type (cf. Vitaceae, Menispermaceae, Aceraceae etc.) is represented mostly by fragments (Plate II, 11). There are a number of fern-like leaves of Comptonia (Myricaceae) in Eckfeld (Plate II, 9) which although common in Tertiary plant assemblages, only constitute a small percentage of the macrofossils. Another rarity is the two compound rosaceous leaves from Eckfeld, one of which even has stipules preserved (Plate II, 3). Legumes are represented in Eckfeld not only by fragments of leaves and isolated leaflets, but also by a few pods. As with the leaves, many of the fruits=seeds can not be assigned to a natural taxon. There is a taphonomic bias between the turbiditic layers and the regular finegrained sediments. The coarse-grained turbiditic material is characterized by an association of robust fruits=seeds, especially including large specimens that are otherwise rare. Among those, Mastixiaceae (Plate V, 4), Vitaceae, Rutaceae, Anacardiaceae (cf. Pentoperculum Manchester, 1994; Plate V, 13), Sargentodoxaceae (cf. Sargentodoxa; Plate V, 9; Tiffney, 1993), Menispermaceae, and possibly a new species of Canarium (Burseraceae) that was initially assigned to a three-loculed Nyssa have been identified preliminarily. Infructescences (Plate V, 10), tiny fruits and fruits with delicate parts, e.g. winged and fleshy fruits, are mainly known from the fine-grained ‘background’ sediments. Most common (>120 specimens) are Hooleya-like pterocaryoid winged fruits (Plate III,
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10, 11; Frankenha¨user and Wilde, 1994), and a distinct type of petiolate four-winged fruits of still unknown systematic affinity (Plate V, 7–8). There are also some nice specimens of hairy fruits (Plate V, 11, 12). Some of the taxa known from the turbiditic layers are also represented in the fine-grained sediments, e.g. at least three different species of Menispermaceae fruits=seeds, Rutaceae seeds, and few specimens of Sambucus (Caprifoliaceae) seeds. Due to their delicate nature, flowers and inflorescences are limited to fine-grained layers. At the moment, more than 30 different species may be distinguished with a total of more than 600 specimens. There are both anemophilous and entomophilous flowers, most of them with pollen preserved in situ (Plate IV). It is especially important, that most of the species are known not only from single specimens, but from a large number in some cases. However, due to the fact that studies on the flowers are still in an initial stage, only few of them can be assigned to extant families, like palms, Juglandaceae, and probably Rutaceae.
6. Interpretation A number of mechanisms are relevant for supplying macroscopic plant material to lakes (e.g. Fritz, 1986; Ferguson, 1993). Due to the physical character of the surrounding area, their contributions may be different as pointed out by Ferguson (1993) in dealing with the Middle Eocene lake of Messel. For the slightly younger maar lake of Eckfeld, a basically similar scenario may be envisaged. In both lakes, leaves, fruits, and flowers fell directly from overhanging twigs. Because of taphonomic filtering effects their fossilization potential may have been lower in Messel. The majority of the plant fragments were blown by wind onto the surface of the water. Surface runoff triggered by rainfall washed material from the surrounding slopes into the lakes, but the contribution of tributaries was probably low. It is difficult to estimate the importance of animal vectors in transporting macroscopic plant fragments, especially diaspores, but their contribution may have been considerable if fleshy fruits indicate endozoochory. As a typical maar structure, the Middle Eocene lake of Eckfeld was surrounded by slopes with a
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PLATE V
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high inclination which led to potential instability. Thus, some plant material slid down the slopes directly into the lake. Instability of the slopes in Eckfeld is indicated by frequent coarse-grained layers of turbiditic origin, and by the fact that there is only scarce evidence for anchored aquatic plants like charophytes, Nymphaeaceae, and Hydrocharitaceae. Accordingly, plants possibly related to a herbaceous fringe along the shoreline are missing or rare in the pollen and macrofossil record of Eckfeld. In Messel, such turbiditic layers have rarely been observed while remains of floating Nymphaeaceae and marginal helophytes (ferns, monocots) are common. Therefore, more gentle slopes and a transitional vegetation fringing the lake in Messel may have prevented material from directly sliding into the water. As a further consequence, taphonomic filtering effects were of minor importance in Eckfeld compared to Messel. This may be the reason for a conspicuous amount of woody material in Eckfeld, including twig and bark fragments with tufts of mosses still adhering. It is even possible that some of this material was directly derived from overhanging twigs. Coarse and resistant plant debris, such as wood=bark fragments and tough fruits=seeds was probably initially concentrated along the shoreline. From time to time these accumulations were affected by sliding due to instability of the slopes and subsequently incorporated into turbiditic layers. Therefore, this kind of material is especially abundant in the turbiditic intercalations. A considerable number of Lygodium pinnules in Eckfeld may also be explained by the absence of a taphonomic filter between the slopes of the crater and the lake itself. Possibly, these fast
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growing climbing ferns preferentially covered fresh surfaces that were repeatedly created by slumping. The overall diversity of the upper Middle Eocene vegetation of the Eckfeld area is indicated by about 200 ‘species’ of spores and pollen, and by a great number of leaf and fruit=seed types. However, at present it is difficult to reconstruct the vegetation of the Eckfeld area in detail because the systematic position of some of the more important fruit=seed and leaf types of the taphoflora is still unknown or uncertain. As pointed out above, anchored aquatics and marginal helophytes were underrepresented due to the physical character of the maar structure. It is obvious from leaves, fruits and pollen, that some Juglandaceae were common or even dominant in the surrounding forest. Palms were present, but their importance is difficult to estimate in spite of a number of flowers and some leaf fragments (extant palms usually produce a large number of flowers but do not shed their leaves). Conifers as well as Lauraceae were definitely scarce. Pinaceae and Fagaceae have not been proven by macrofossils, and their contribution to the pollen assemblage is only moderate. Therefore, they probably grew at some distance from the maar lake. A diversity of climbers (e.g. Lygodium, Menispermaceae, Vitaceae) could have covered the edge of the forest as well as fresh surfaces.
7. Interactions Some research has recently been focused on interactions, especially plant–fungal and plant–animal relationships (e.g. Scott and Taylor, 1983; Scott
PLATE V Fruits and seeds. Scale 0.5 cm except for 5 and 6 (scale 0.1 cm). 1–3. Different types of Menispermaceae seeds; PB 1995=370 LS, PB 1993=111 LS, PB 1993=116 LS. 4. Mastixioid fruit; PB 1995=379 LS. 5a–c. Isolated fruit of Cyperaceae; PB 1995=374 LS. 6. Seed of Sambucus sp. (Caprifoliaceae); PB 1995=366 LS. 7. Four-winged fruit of unknown affinity, wings flattened on bedding plane as in most specimens; PB 1993=728 LS. 8. Laterally compressed fruit as in 7, with petiole (arrow) and fruit proper attached; PB 1995=140 LS. 9. Fruit=Seed of cf. Sargentodoxa sp. (Sargentodoxaceae); PB 1995=381 LS. 10. Fragment of infructescence possibly related to Emmenopterys (Rubiaceae) (Manchester, 1994); PB 1995=380 LS. 11. Fruit of unknown affinity showing single seed with tuft of hairs (partly obscured by a leaf fragment below); PB 1993=746 LS. 12. Fruit=seed of unknown affinity with a hairy tail (‘Cypselites’ sp.); PB 1993=747 LS. 13a, b. Single specimen of cf. Pentoperculum Manchester, 1994 (Anacardiaceae), fruit; PB 1995=375 LS. a. From above. b. From below.
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PLATE VI
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and Paterson, 1984; Chaloner et al., 1991; Scott, 1991, 1992; Taylor and Osborn, 1996). As at Messel (Schaarschmidt, 1988), a lot of evidence for a wide spectrum of interactions has been collected at Eckfeld. Because detailed studies of the material are for the most part still outstanding, they will be mentioned only briefly. Epiphytic and even epiphyllous bryophytes are most noteworthy as representing rare evidence of plant–plant relationships. Examples of parasitic=saprophytic plant–fungal associations may be observed particularly on leaves. Most common is a diversity of fungal hyphae in cuticular preparations. In addition, there are some examples of cone-shaped ascomycete fructifications on leaves and fruits (Plate VI, 4, 5). In many of the leaves parts of the lamina have obviously been gnawed away. Different outlines of the resulting injury indicate a considerable diversity of animals feeding on leaves (Plate VI, 6–9), probably most of them arthropods. Another kind of leaf attack related to feeding is documented by traces of mining (Plate VI, 9; e.g. Lang et al., 1995). That some of the mammals fed on leaves is proven by bits of cuticles in the gut of a complete specimen of Propalaeotherium voigti (Plate VI, 11). Another obvious result of plant–animal interactions are galls (Scott et al., 1994). Many of them are caused by insects attaching their eggs to plant parts, thereby modifying growth of the respective tissues. There are a variety of galls preserved on the leaves from Eckfeld (Plate VI, 1–3). A spectacular example is the three-dimensionally preserved cone-shaped structure on some specimens of juglandaceous leaflets (Plate VI, 1).
21
In Eckfeld, insect pollination is already indicated by a number of rarely occurring types of large pollen grains (Nickel, 1996). More indirect evidence is derived from a typical architecture in some of the flowers, including highly advanced types like a dorsiventral flower with a tubular corolla (Frankenha¨user and Wilde, 1993a). Moreover, from the Eckfeld site there is even direct proof of a potential insect pollinator with the oldest honeybee presently known. It is still carrying its pollen load in place (Plate VI, 10; Lutz, 1993b).
8. Comparisons with other localities The Eckfeld plant assemblage should finally be compared with other nearby Middle Eocene plant localities in Germany (Fig. 1), like Messel, Helmstedt, the Geiseltal area and the Weißelster Basin. As shown in Table 2, most of the localities are not strictly contemporaneous, while the Geiseltal plant remains represent a number of localities spanning a considerable period of time. However, preliminary Table 2 Stratigraphic range of the localities mentioned in the text Eckfeld Messel Geiseltal >MP14 MP14 MP13 C MP12 MP11
Helmstedt Weißelsterbecken C
C
C C C C
? C C C
PLATE VI Interactions. 1a, b. Juglandaceous leaflet with 3-dimensionally preserved galls (arrows). PB 1995=385 LS. a. Overview. Scale 1 cm. b. Detail from 1a. Scale 0.5 cm. 2, 3. Leaves with different types of circular galls. PB 1995=386, 387 LS. Scale 1 cm. 4. Fruiting bodies of ascomycetes on a leaf surface. PB 1990=6 LS. Scale 0.1 cm. 5. Fruit affected by fungal growth; PB 1995=388 LS. Scale 0.5 cm. 6–9. Gnawed leaves. PB 1995=382-384 LS, PB 1990=68 LS. Scale 1 cm. 10a, b. Bee (Eckfeldapis electrapoides Lutz, 1993b) with pollen load in the metatarsal brush. PE 1992=614 LS. Scale 0.5 cm. b. Small tricolp(?or)ate pollen grains from metatarsal brush of a (unidentified). Negative print from slide, CLSM picture courtesy of LEICA GmbH, Bensheim. Scale 5 µm. 11a, b. Cuticles from gut of Propalaeotherium voigti. a. Scale 100 µm. b. Scale 30 µm.
22
V. Wilde, H. Frankenha¨user / Review of Palaeobotany and Palynology 101 (1998) 7–28
comparisons of Middle and Upper Eocene plant assemblages already suggest that differences in age within the Middle Eocene are not as important as different facies represented by their respective sediments (Wilde, 1989a). Therefore, a comparison of these assemblages may provide a simple approach to assess some small-scale differentiation of the Middle Eocene vegetation in Europe (Wilde, 1989b, 1995; Wilde and Frankenha¨user, 1993). Unfortunately, any such comparison can only be based on a qualitative approach and has to remain preliminary at present. Leaves have been described from all of the localities (see compilation in Wilde, 1995), but in none of the assemblages have they been monographed exhaustively. There is a large number of ‘species’ known from the Geiseltal area (Wilde, 1995), but still comparatively few from Helmstedt (Wilde, 1989a) and the Weißelster Basin (Fischer, 1991). With the exception of the Geiseltal area (Mai, 1976), knowledge of fruits and seeds is quite limited. In Messel (Collinson, 1988), and even more so in Eckfeld (this paper) only some of the numerous taxa have been identified or even studied in detail. There is hitherto no record of fruits and seeds from the other localities. Due to different modes of preservation, comparison of the respective taphofloras from the Geiseltal area (sieved material) and from Messel and Eckfeld (compressions) is difficult. Systematics of pollen and spores has been studied extensively in Messel and Eckfeld (Thiele-Pfeiffer, 1988; Nickel, 1994, 1996), but there is no comparably complete record for any horizon of the Geiseltal area or either of the other localities (Krutzsch, 1976, 1992). Similarity of the pollen and spore assemblages of Eckfeld and Messel is expressed by a great number of taxa common to both localities. However, published data from the Geiseltal area already suggest striking differences (Nickel, 1996). Accordingly, most of the ferns, and all of the seed-plant families hitherto recognized in the macrofossil assemblage of Eckfeld are also known from Messel (except for Betulaceae that are represented in Eckfeld by still debatable Carpinus-like bracts). There are many species definitely common to both localities, e.g. conifers (Cephalotaxus messelensis), Caprifoliaceae (more or less identical seeds of Sambucus), Juglandaceae (some common types of leaves and fruits), cf. Sargentodoxa seeds, Theaceae (Polyspora sax-
onica, Ternstroemites dentatus), Ulmaceae (common types of ulmoid and zelkovoid leaves), and Araceae (leaves: ‘Araceae sp. 2’ sensu Wilde, 1989a,b). Furthermore, identical or closely related species are highly probable but have still to be proven in Menispermaceae (seeds), Myricaceae (leaves of Comptonia), Rosaceae (leaves), Rutaceae (seeds), vitioid leaves, Cyperaceae (fruits), and Liliaceae s.l. (leaves of cf. Smilax). In contrast to Messel with one ‘species’ of coniferous foliage and 6 ‘species’ of angiosperm leaves that have also been described from the Geiseltal area, none of the phanerogamous leaves or fruits=seeds of Eckfeld has yet been proven to be identical to a species from the Geiseltal area. Surprisingly, most of the ferns from Eckfeld are common to both Messel and the Geiseltal area (Frankenha¨ user and Wilde, 1993b). Most interesting is the diversity of Juglandaceae represented in the taphocoenoses of Eckfeld and Messel with no macrorecord of the family in the Geiseltal area, Helmstedt and the Weißelster Basin. Conversely, leaves and fruits of Fagaceae are common in the Geiseltal area, but still completely missing in Eckfeld and Messel. The extreme diversity of lauraceous leaves in Messel and a number of species in the Geiseltal area (probably also in Helmstedt and the Weißelster Basin) is in strange contrast to the few lauraceous leaves in Eckfeld. Taxa like Doliostrobus (?Taxodiaceae), and Rhodomyrtophyllum (Myrtaceae) are especially common in the coal bearing areas (Geiseltal area, Helmstedt, Weißelster Basin), have been found in Messel, but are hitherto unknown from Eckfeld. Palaeogeographically, the peat-forming lowlands like the Geiseltal area, Helmstedt and the Weißelster Basin were situated fairly close to the Middle Eocene coastline (Fig. 1). Their respective taphofloras most probably represent the azonal floras of a coastal plain with some peatswamps. In contrast, Messel and Eckfeld were more or less isolated lakes at some distance from the coast. They were surrounded by a zonal flora flourishing in the central part of the Middle Eocene Western European Island. Different edaphic conditions are apparently not reflected in the ferns whose distribution was governed by the humid climate.
V. Wilde, H. Frankenha¨user / Review of Palaeobotany and Palynology 101 (1998) 7–28
Acknowledgements Excavations at the Eckfeld site were initiated and continued since 1987 by Dr. F.-O. Neuffer, head of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz. Important field observations were supplied by Dr. Herbert Lutz who supervised most of the fieldwork actually done by staff of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz with the help of volunteers. Some of the plant material was prepared by Uta Jaeckel (Mainz). Dr. Birgit Nickel (Frankfurt) generously provided us with her unpublished data on the pollen assemblage from Eckfeld. Among others, Drs. Margaret Collinson (London) and Kurt Goth (Freiberg) spent some time with us discussing the Eckfeld fruits and seeds. Excavations and research in Eckfeld have been subsidized by the government of Rheinland-Pfalz and by the Deutsche Forschungsgemeinschaft (Project Nr. II C Ne-440=1 and =2). Our fruitful cooperation has been steadily supported by both the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz and the Forschungsinstitut Senckenberg, Frankfurt am Main. Thanks are due to Professor David K. Ferguson, Vienna, for improving our English.
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Dissertation A, Berlin, 118 pp. (unpubl.). Frankenha¨user, H., Wilde, V., 1993a. Flowers from the Middle Eocene of Eckfeld (Eifel, Germany)—first results. In: Negendank, J.F.W., Zolitschka, B. (Eds.), Paleolimnology of European Maar Lakes. Lecture Notes Earth Sci. 49, 491–497. Frankenha¨user, H., Wilde, V., 1993b. Farne aus der mitteleoza¨nen Maarfu¨llung von Eckfeld bei Manderscheid in der Eifel. Mainzer Naturwiss. Arch. 31, 149–167. Frankenha¨user, H., Wilde, V., 1994. Zweiflu¨gelige JuglandaceenFru¨chte aus dem Mitteleoza¨n von Eckfeld bei Manderscheid (Eifel). Vorla¨ufige Mitteilung. Mainzer Naturwiss. Arch. Beih. 16, 143–150. Frankenha¨user, H., Wilde, V., 1995. Stachelspitzige Bla¨tter aus dem Mitteleoza¨n von Eckfeld (Eifel). Abh. Staatl. Mus. Mineral. Geol. Dresden 41, 97–115. Franzen, J.L., 1994. Neue Sa¨ugerfunde aus dem Eoza¨n des Eckfelder Maares bei Manderscheid (Eifel). Mainzer Naturwiss. Arch. Beih. 16, 189–211. Franzen, J.L., 1995. Die Equoidea des europa¨ischen Mitteleoza¨ns (Geiseltalium). Hallesches Jahrb. Geowiss. B 17, 31–45. Fritz, W.J., 1986. Plant taphonomy in areas of explosive volcanism. In: Gastaldo, R.A. (Ed.), Land Plants, Notes for a Short Course. Univ. Tennessee Dep. Geol. Sci. Stud. Geol. 15, 1–26. Groh, K., Jungbluth, J.H., 1994. Vorla¨ufige Mitteilung zur Najadenfauna (Mollusca: Bivalvia: Unionidae) des Eckfelder Maares (Mittel-Eoza¨n). Mainzer Naturwiss. Arch. Beih. 16, 151–165. Ja¨ckel, U., Frankenha¨user, H., 1994. Fru¨chte und Samen der mitteleoza¨nen Fundstelle Eckfeld=Eifel. Mitt. Rheinischen Naturforsch. Ges. 15, 85. Krutzsch, W., 1976. Die Mikroflora der Braunkohle des Geiseltales Teil IV: Die stratigraphische Stellung des Geiseltalprofils im Eoza¨n und die sporenstratigraphische Untergliederung des mittleren Eoza¨ns. Abh. Zentr. Geol. Inst. 26, 47–92. Krutzsch, W., 1992. Pala¨obotanische Klimagliederung des Alttertia¨rs (Mitteleoza¨n bis Oberoligoza¨n) in Mitteldeutschland und das Problem der Verknu¨pfung mariner und kontinentaler Gliederungen (klassische Biostratigraphien–pala¨obotanisch-o¨kologische Klimastratigraphie–Evolutions-Stratigraphie der Vertebraten). Neues Jahrb. Geol. Pala¨ontol. Abh. 186, 137–253. Lang, P.J., Scott, A.C., Stephenson, J., 1995. Evidence of plant– arthropod interactions from the Eocene Branksome Sand Formation, Bournemouth, England: introduction and description of leaf mines. Tertiary Res. 15, 145–174. Lo¨hnertz, W., 1978. Zur Altersstellung der tiefliegenden fluviatilen Tertia¨rablagerungen der SE-Eifel (Rheinisches Schiefergebirge). Neues Jahrb. Geol. Pala¨ontol. Abh. 156, 179–206. Lo¨hnertz, W., 1994. Grundzu¨ge der morphologischen Entwicklung der su¨dlichen Eifel im a¨ltesten Tertia¨r. Mainzer Naturwiss. Arch. Beih. 16, 17–38. Lorenz, V., 1973. On the formation of maars. Bull. Volcanol. 37, 183–204. Lutz, H., 1993a. Zur Sedimentologie der Leithorizonte des ‘Eckfelder Maares’ bei Manderscheid=Eifel (Mittel-Eoza¨n; Deutschland). Mainzer Naturwiss. Arch. 31, 65–83. Lutz, H., 1993b. Eckfeldapis electrapoides nov. gen. n. sp., eine
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‘Honigbiene’ aus dem Mittel-Eoza¨n des ‘Eckfelder Maares’ bei Manderscheid=Eifel Deutschland (Hymenoptera: Apidae, Apinae). Mainzer Naturwiss. Arch. 31, 177–199. Mai, D.H., 1976. Fossile Fru¨chte und Samen aus dem Mitteleoza¨n des Geiseltales. Abh. Zentr. Geol. Inst. 26, 93–149. Manchester, S.R., 1994. Fruits and seeds of the Middle Eocene Nut Beds Flora, Clarno Formation, Oregon. Palaeontogr. Am. 58, 1–205. Meyer, W., 1994. Geologie der Eifel, 3rd revised ed. Schweizerbart, Stuttgart, 618 pp. Meyer, W., Pirrung, B.M., Stets, J., 1994. Der variscische Sockel in der Umgebung des Eckfelder Maares. Mainzer Naturwiss. Arch. Beih. 16, 9–16. Mingram, J., 1994. Sedimentologie und Zyklizita¨t laminierter ¨ lschiefer von Eckfeld=Eifel. Mainzer Naturwiss. eoza¨ner O Arch. Beih. 16, 55–85. Negendank, J.F.W., Irion, G., Linden, J., 1982. Ein eoza¨nes Maar bei Eckfeld nordo¨stlich Manderscheid (SW-Eifel). Mainzer Geowiss. Mitt. 11, 157–172. Neuffer, F.O., Gruber, G., Lutz, H., Frankenha¨user, H., 1996. Das Eckfelder Maar—Zeuge tropischen Lebens in der Eifel. Landessammlung fu¨r Naturkunde Rheinland-Pfalz, Mainz, 102 pp. Nickel, B., 1994. Neue palynologische Untersuchungen am mit¨ lschiefer von Eckfeld bei Manderscheid=Eifel. teleoza¨nen O Erste Ergebnisse. Mainzer Naturwiss. Arch. 32, 7–25. Nickel, B., 1996. Die mitteleoza¨ne Mikroflora von Eckfeld bei Manderscheid=Eifel. Mainzer Naturwiss. Arch. Beih. 18, 1– 147. Pflug, H.D., 1959. Die Deformationsbilder im Tertia¨ r des rheinisch-saxonischen Feldes. Freib. Forschungsh. C 71, 1– 110. Pirrung, B.M., 1992a. Geologische und geophysikalische Untersuchungen am tertia¨ren ‘Eckfelder Maar’, Su¨dwesteifel. Mainzer Naturwiss. Arch. 30, 3–21. Pirrung, B.M., 1992b. Zur Frage der Entstehung eoza¨ner Sedimente im ‘Eckfelder Maar’ bei Manderscheid, Su¨dwesteifel. Mitt. Pollichia 79, 139–158. Pirrung, B.M., 1993. Weitere Sondierungen und geophysikalische Untersuchungen am eoza¨nen Eckfelder Maar bei Manderscheid, Su¨dwesteifel. Mainzer Naturwiss. Arch. 31, 37–63. Pirrung, B.M., Bu¨chel, G., 1994. Das Eckfelder Maar—ein tertia¨res Maar der Hocheifel. Mainzer Naturwiss. Arch. Beih. 16, 39–53. Schaarschmidt, F., 1988. Der Wald, fossile Pflanzen als Zeugen eines warmen Klimas. In: Schaal, S., Ziegler, W. (Eds.), Messel—ein Schaufenster in die Geschichte der Erde und des Lebens. Kramer, Frankfurt am Main, pp. 29–52. Scott, A.C., 1991. Evidence for plant–arthropod interactions in the fossil record. Geol. Today 7, 58–61. Scott, A.C., 1992. Trace fossils of plant–arthropod interactions. In: Maples, C.G., West, R.R. (Eds.), Trace Fossils. Short Courses Paleontol. 5, 197–223. Scott, A.C., Paterson, S., 1984. Techniques for the study of plant=arthropod interactions in the fossil record. Geobios Me´m. Spec. 8, 449–455.
Scott, A.C., Taylor, T.N., 1983. Plant=animal interactions during the Upper Carboniferous. Bot. Rev. 49, 259–307. Scott, A.C., Stephenson, J., Collinson, M.E., 1994. The fossil record of leaves with galls. In: Williams, M.A.J. (Ed.), Plant Galls. Syst. Assoc. Spec. Vol. 49, 447–470. Taylor, T.N., Osborn, J.M., 1996. The importance of fungi in shaping the paleoecosystem. Rev. Palaeobot. Palynol. 90, 249– 262. Thiele-Pfeiffer, H., 1988. Die Mikroflora aus dem mitteleoza¨nen ¨ lschiefer von Messel bei Darmstadt. Palaeontographica B O 211, 1–86. Tiffney, B.H., 1993. Fruits and seeds of the Tertiary Brandon Lignite. VII. Sargentodoxa (Sargentodoxaceae). Am. J. Bot. 80, 517–523. Tobien, H., 1961. Ein Lophiodon-Fund (Tapiroidea, Mamm.) aus den niederhessischen Braunkohlen. Notizbl. Hess. Landesamtes Bodenforsch. Wiesbaden 89, 7–16. Von der Brelie, G., 1969. Neue Untersuchungen im Alttertia¨r von Eckfeld bei Manderscheid (Eifel). Fortschr. Geol. Rheinland Westfalen 17, 27–40. Weber, C.O., 1853. Ueber das Braunkohlenlager von Eckfeld in der Eifel. Verh. Naturhist. Ver. Preuss. Rheinlande Westfalens 10, 409–415. White, J.D.L., 1991. The depositional record of small, monogenetic volcanoes within terrestrial basins. In: Fisher R.V., Smith, G.A. (Eds.), Sedimentation in Volcanic Settings. SEPM Spec. Publ. 45, 155–171. Wilde, V., 1989a. Untersuchungen zur Systematik der Blattreste aus dem Mitteleoza¨n der Grube Messel bei Darmstadt (Hessen Bundesrepublik Deutschland). Courier Forsch.-Inst. Senckenberg 115, 1–213. Wilde, V., 1989b. Vorla¨ufige Mitteilungen zur Flora aus dem Alttertia¨r von Eckfeld—Ergebnisse einer ersten Durchsicht des Fundmateriales aus den Grabungen von 1987 und 1988. Mainzer Naturwiss. Arch. 27, 23–31. Wilde, V., 1990. Moosreste aus dem Alttertia¨r von Eckfeld bei Manderscheid in der Eifel. Mainzer Naturwiss. Arch. 28, 1–6. Wilde, V., 1995. Die Makroflora aus dem Mitteleoza¨n des ¨ bersicht und Vergleiche. Hallesches Geiseltalgebietes, kurze U Jahrb. Geowiss. B 17, 121–138. Wilde, V., Frankenha¨user, H., 1993. Initial results on the importance of a flora from the Middle Eocene of Eckfeld (Eifel, W.-Germany). In: Negendank, J.F.W., Zolitschka, B. (Eds.), Paleolimnology of European Maar Lakes. Lecture Notes Earth Sci. 49, 499–503. Wilde, V., Frankenha¨user, H., 1995. Flu¨gelfru¨chte engelhardioider Juglandaceen aus dem Mitteleoza¨n von Eckfeld bei Manderscheid (Eifel). Mainzer Naturwiss. Arch. 33, 47–52. Wilde, V., Frankenha¨user, H., Lutz, H., 1993. Algenreste aus den mitteleoza¨nen Sedimenten des Eckfelder Maares bei Manderscheid in der Eifel. Mainzer Naturwiss. Arch. 31, 127–148. Zolitschka, B., 1993. Palaeoecological implications from the sedimentary record of a subtropical maar lake (Eocene Eckfelder Maar; Germany). In: Negendank, J.F.W., Zolitschka, B. (Eds.), Paleolimnology of European Maar Lakes. Lecture Notes Earth Sci. 49, 477–484.
Appendix A Summary of the probable life forms, dispersal mechanisms and physiognomic signatures of the Eckfeld taxa, based on the representation of the different plant parts in the assemblages (compiled by D.K. Ferguson) Family ALGAE Botryococcaceae
Chrysophyceae Zygnemataceae Bryophytes Dicranaceae s.l. Jungermanniaceae Incerta sedis
Other organs
Primary dispersal mechanism
Botryococcus Tetraedron Chara
coenobia unicells gyrogonites
gen. indet. Ovoidites sp.
cysts zygospores
exozoo-nautochory exozoo/nautochory endozoo-/exozoo/ nautochory exozoo-/nautochory exozoo-/nautochory
Dicranites† cf. Frullania Muscites†
FERNS AND FERN-ALLIES Lycopodiaceae Camarozonosporites eocaenicus Osmundaceae Osmunda lignitum/Baculatisporites primarius Polypodiaceae s.l. Acrostichum sp. cf. Didymochlaena/Lindsaea Laevigatosporites discordatus, L. haardtii Polypodium (Verrucatosphorites tenellis) Rumohra recentior Schizaeaceae Cicatricosisporites dorogensis, C. paradorogensis Lygodium kaulfussii/Leiotriletes maxoides, L. microadriennis cf. Ruffordia subcretacea Schizaea (Microfoveolatosporites granuloides)
Pollen= spores
Diaspores
x x x
capsules
wind wind wind
chamaephyte chamaephyte chamaephyte
wind wind
chamaephyte hemicryptophyte
x(2)
wind wind wind
geo-/hemicryptophyte hemicryptophyte geo-/hemicryptophyte
x
wind
geo-/hemicryptophyte
x(2)
wind wind
geo-/hemicryptophyte geo-/hemicryptophyte
x(2)
wind
geo-/hemicryptophyte
wind wind
geo-/hemicryptophyte geo-/hemicryptophyte
wind endozoochory wind endozoochory endozoochory wind wind wind
phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte
x x x x
sporangia
x
x
sporangia
x
sporangiophores x
x x x x
x
Life form
x x x x
x
Deciduous= Evergreen
Leaf margin entire
E E E E E E E D/E
25
GYMNOSPERMS Araucariaceae? Araucariacites europaeus Cephalotaxaceae Cephalotaxus cf. messelensis Cupressaceae s.s. Cupressacites insulipapillatus Libocedrites salicorniodes Ephedraceae Ephedra (Ephedripites eocenipites) Pinaceae Cathaya (Pityosporites microalatus) Pinus (Pityosporites labdacus) Taxodiaceae gen. indet.
Leaves
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Charophyceae
Taxon
26
Appendix A (continued) Family
Taxon
Aquifoliaceae Araliaceae
Pollen/ spores
?
x
Diaspores
Other organs
x
Primary dispersal mechanism
Life form
Deciduous/ Evergreen
Leaf margin entire
wind endozoochory
phanerophyte phanerophyte incl. liana phanerophyte phanerophyte—liana
D/(E) D/E
yes/no yes/no
D/E E
yes/no yes
D/E
yes/no
D D
no no
D
no
D/(E)
yes
E D
yes no
E
rarely
D/E
yes/no
chamaephyte/ phanerophyte phanerophyte
D/E
yes/no
D
no
chamae-/geo-/ hemicrypto-/ phanero-/therophyte phanerophyte phanerophyte phanerophyte phanerophyte incl. liana phanerophyte phanerophyte phanerophyte
D/E
yes/no
E D/E D/E E
yes/no yes/no yes/no mostly
D D/E D
no no no
llex (llexpollenites iliacus) Hedera (Araliaceoipollenites reticuloides) gen. indet. (Araliaceoipollenites euphorii, A. sp.)
x x
endozoochory endozoochory
x(2)
endozoochory
Alnus (Ainipollenites verus) Betula (Trivestibulopollenites betuloides) Carpinus type
x x
wind wind/endozoochory
x(3)
Burseraceae Caprifoliaceae
Bombacacidites kettigensis, B. tilioides, B. sp. Canarium Sambucus sp.
Chloranthaceae
Emmapollis pseudoemmaensis
x
endozoochroy
Clethraceae?/ Cyrillaceae? Ericaceae
Tricolporopollenites megaexactus
x
wind/endozoochory
Ericipites callidus, E. ericius, E. sp. Eucommia (Tricolporopollenites parmularius) Spinaepollis spinosus
x(3)
wind/endozoochory
x
wind
?
Archetype Quercus (Tricolpopollenites asper) gen.indet Compositoipollenites rhizophorus
x x ? x
auto-/endozoo-/ exozoo-/myrmeco-/ nautochory dyschory dyschory autochory/wind wind/endozoochory
Betulaceae s.l.
Bombacaceae
Eucommiaceae Euphorbiacaea
Fagaceae Hamamelidaceae Icacinaceae Juglandaceae
Carya (Caryapollenites circulus) Engelhardia type Platycarya type
x
wind/endozoochory wind/endozoochory
x x
x x
x x x
endozoochory endozoochory
flowers
x x
flowers
dyschory wind wind
(hemicrypto)-/ phanerophyte incl. liana phanerophyte chamaephyte/ phanerophyte chamaephyte/ phanerophyte phanerophyte phanerophyte (hemicryptophyte)/ phanerophyte hemicrypto-/ phanerophyte phanerophyte
V. Wilde, H. Frankenha¨user / Review of Palaeobotany and Palynology 101 (1998) 7–28
ANGIOSPERMS DICOTYLEDONS: Aceraceae Acer (Aceripollis striatus) Anacardiaceae cf. Pentoperculum†
Leaves
Apendix A (continued) Family
Taxon
Leaves
Loranthaceae s.l.
Gothanipollis gothanii
Mastixiaceae Menispermaceae
gen. indet. gen. indet.
?
Moraceae
Ficus
?
Myricaceae
Comptonia Triatriopollenites bitutius, T. rurensis Duplopollis myrtoides Monocolpopollenites crassiexinus Nyssa (Tricolporopollenites kruschii) Olax (Olaxipollis matthesii)
x
Lauraceae
Myrtaceae Nymphaeaceae Nyssaceae Olacaceae Oleaceae Onagraceae
Diaspores
Other organs
x(4)
x x x
x
x
x
x
x
Life form
Deciduous/ Evergreen
Leaf margin entire
?
phanerophyte
D/E
yes/no
wind endozoochory endozoochory autochory/wind/ nautochory endozoochory
D E E D/E
no yes yes yes
(D)/E
yes
E D/E
yes no
(D)/E
yes
D D/E
no yes/no
endozoochory endozoochory
x x
endozoo-/nautochory endozoo-/nautochory
phanerophyte hydrophyte
E H
yes H
x
endozoochory
phanerophyte
D
yes/no
x
endozoochory
E
yes
x
wind/endozoochory
phanerophyte incl. liana phanerophyte
D/E
yes/no
x(2)
wind/endozoochory
H
H
?
endozoochory
D/E
no
D D/E
yes yes/no
D
rarely
D/E
mostly
x x
x x(3C)
endozoochory endozoochory
endozoochory
Rosaceae
Rosa/Rubus
Rubiaceae Rutaceae
Emmenopterys gen. indet.
?
Salicaceae
Tricolpopollenites retiformis
x
wind
Sapotaceae
Tetracolporopollenites spp. div.
x(6-7)
endozoochory
? x(1C)
flowers
wind autochory/wind/ endozoochory
(chamae-/geo-)/ hemicrypto-/ (phanerophyte) (hemicryptophyte)/ phanerophyte incl. liana phanerophyte (hemicryptophyte-)/ phanerophyte incl. liana chamaephyte/ phanerophyte phanerophyte
27
x(2)
phanerophyte phanerophyte phanerophyte chamae-/hemicrypto-/ phanero-/therophyte chamaephyte/ phanerophyte phanerophyte (hemicryptophyte)/ phanerophyte, mostly liana phanerophyte incl. liana phanerophyte phanerophyte
x
Tricolporopollenites microreticulatus Corsinipollenites oculusnoctis, C. verrucatus
Primary dispersal mechanism
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Leguminosae
Plicatopollis hungaricus, P. lunatus, P. plicatus, P. potoniei Pterocarya type cf. Laurophyllum† cf. Daphnogene† gen. indet.
Pollen/ spores
28
Appendix A (continued) Taxon
Sargentodoxaceae Sterculiaceae
cf. Sargentodoxa sp. Reevesia (Reevesiapollis triangulus vel eocaenicus) Porocolpopollenites calauensis, P. rarobaculatus, P. vestibulum cf. Polyspora saxonica cf. Ternstroemites dentatus Intratiporopollenites ceciliensis, I. minimus, I. cf. maxoides
Symplocaceae Theaceae Tiliaceae
Ulmaceae s.l.
Leaves
Incerta sedis
Punigiphyllum waltheri
x
Smilacaceae
Milfordia hungarica, M. incerta, M. minima cf. Smilax
Sparganiaceae
Sparganiaceaepollenites spp.
Primary dispersal mechanism
Life form
Deciduous/ Evergreen
Leaf margin entire
x
endozoochory wind
phanerophyte-liana phanerophyte
D D/E
yes mostly
x(3)
endozoochory
phanerophyte
(D)/E
yes/no
phanerophyte phanerophyte (chamae-/ hemicrypto-)/ phanerophyte phanerophyte phanerophyte
E E D/E
yes/no no rarely
x x(2)
wind endozoochory wind/endozoo-/ exozoo-/ nautochory endozoochory wind
D/(E) D
yes/no no
x(4)
wind wind endozoochory
phanerophyte phanerophyte phanerophyte, mostly liana ?
D D D/(E)
no no no
?
no
x(3)
Vitaceae
gen. indet. Graminidites eifelensis, G. sp.
Other organs
x x
x x x
hydrocharitaceae Poaceae (Gramineae) Restionaceae
Diaspores x
Celtipollenites intrastructurus Polyporopollenites undulosus, P. verrucatus Ulmus type Zelkova type gen. indet.
MONOCOTYLEDONS: Araceae gen. indet. Arecaceae gen. indet. (Palmae) Cyperaceae gen. indet.
Pollen/ spores
x x(2CG)
?
x x
?
x
? x(2) x(3)
x
flowers, spines x
endozoochory? endozoo-/ nautochory endozoo-/exozoo-/ nautochory nautochory wind/endozoo-/ exozoochory ? endozoochory?
x(2)
endozoo-/ nautochory
hemicryptophyte? phanerophyte incl. liana geo-/hemicrypto-/ therophyte hydrophyte geo-/hemicrypto-/ phanero-/therophyte geo-/ hemicryptophyte phanerophyteliana? hydrophyte
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Family
Review of Palaeobotany and Palynology 101 (1998) 7–28
The Middle Eocene plant taphocoenosis from Eckfeld (Eifel, Germany) Volker Wilde a,Ł , Herbert Frankenha¨user b b
a Forschungsinstitut Senckenberg, Pala ¨ obotanik, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz, Reichklarastr. 10, D-55116 Mainz, Germany
Received 10 February 1997; revised version received 14 August 1997; accepted 9 September 1997
Abstract More than 15,000 plant macrofossils have been collected during scientific excavations in an isolated occurrence of organic-rich clay=silt (‘oilshale’) with coarse turbiditic intercalations at Eckfeld near Manderscheid (Eifel, Germany). The sediments were deposited in a meromictic maar lake, and have been dated by mammals as late Middle Eocene (late Geiseltalian, MP13). Initial studies on leaves, fruits=seeds and flowers confirm the considerable diversity of the taphoflora indicated by about 200 ‘species’ of pollen and spores. The macrofossils suggest an interpretation as a zonal flora surrounding a more or less isolated inland lake with considerable inclination of the slopes. The assemblage has a number of taxa in common with the slightly older taphocoenosis of Messel, but with the exception of the ferns there are no species common to the Middle Eocene of the Geiseltal area. This may be explained by the regional palaeogeography with the Geiseltal area as a near-coastal peat-forming lowland, while Eckfeld and Messel were lakes at some distance from the coastal lowlands. 1998 Elsevier Science B.V. All rights reserved. Keywords: Eocene; Germany; lake; palaeobotany; palaeoecology; taphonomy; volcanism
1. Introduction The German Middle Eocene taphofloras of the Geiseltal area and of Messel have been known for a long time and studied in considerable detail. Other nearby plant assemblages dating from the same time interval like those of the Weißelster Basin and Helmstedt, have only been investigated in a preliminary fashion. Another locality with plant remains of Tertiary age was noted for the first time in the middle of the last century (Weber, 1853) in the vicinŁ Corresponding
author.
ity of Eckfeld near Manderscheid (Eifel, Germany; Fig. 1). The plant remains mentioned by Weber (1853) were never figured or described, and the material was apparently lost. Due to the excavations carried out by the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz since 1987 (Neuffer et al., 1996), the Eckfeld locality has become a well known fossillagersta¨tte, and was finally dated as Middle Eocene. Due to their palaeogeographic position, and the large number of macrofossils collected (>15,000), the plant assemblage from Eckfeld is of major importance in reconstructing the Middle Eocene vegetation of Germany. There was considerable taxonomic
c 1998 Elsevier Science B.V. All rights reserved. 0034-6667/98/$19.00 PII S 0 0 3 4 - 6 6 6 7 ( 9 7 ) 0 0 0 6 7 - 5
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Fig. 1. Palaeogeographic situation of the Middle Eocene of Western Europe (after Franzen, 1995), with localities mentioned in the text. Large dots D marine; fine stipple D land.
diversity as indicated by the presence of about 200 ‘species’ of spores and pollen (Nickel, 1996); knowledge on the macrofossils is, however, still limited. It is the aim of the present paper to give a more sophisticated overview of the present state of knowledge on the Middle Eocene macrofossils from Eckfeld, including some of its implications and comparisons with coeval plant assemblages. All the material is housed in the collections of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz.
2. Geological setting The Eifel is the German part of the outcropping Variscan Basement west of the river Rhine and north of the river Mosel. Most of the area is characterized by Devonian sediments with remnants of a Permian to Mesozoic cover (e.g. Meyer, 1994). Volcanic activity started in the Cretaceous and continued into Holocene times. As a result of the volcanic activ-
ity, there are numerous more or less circular volcanogenic depressions, today often occupied by a swamp or a lake. Traditionally, they have been called ‘maar’ by the local population. The term ‘maar’ is now internationally accepted for small monogenetic structures of phreatomagmatic origin with the base of the crater below the original surface (Lorenz, 1973; White, 1991). As reported for the first time by Weber (1853), an isolated occurrence of browncoal was discovered near Eckfeld in 1839, but mining activities had failed due to unfavourable rock conditions. The sediments were dated by Weber (1853) by comparison with some plant remains previously described from Oligocene localities near Bonn (Rott and Orsberg). The locality was for a long time overlooked. However, Pflug (1959) discovered a specimen of ‘coal’ from Eckfeld in the collections of the Geological Institute of the University of Cologne. Its sporomorphs were compared to the ‘Borkener Bild’ which at that time was thought to be of Late Eocene age. Based on new samples from the locality, the ‘Borkener
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Bild’ was later confirmed (Von der Brelie, 1969) at Eckfeld, but its range had by then been extended down into the Middle Eocene by mammal data from the type locality (Tobien, 1961). A Middle Eocene age (late Geiseltalian=MP 13) for the sediments exposed in Eckfeld by ongoing excavations was finally proven by a characteristic association of mammalian taxa (Franzen, 1994). The Tertiary sediments at Eckfeld were first explained by Weber (1853) as a filling of a primarily isolated maar-like depression. For some time, an interpretation as remnants of more extensive deposits (Pflug, 1959; Von der Brelie, 1969), possibly secondarily preserved in a volcanogenic depression (Von der Brelie, 1969), was favoured. Lo¨hnertz (1978) later recalled the maar hypothesis which was for the first time confirmed by drilling down into coarse pyroclastics (Negendank et al., 1982). Detailed geological and geophysical investigations recently gave final support to a maar origin for the Eckfeld structure (Pirrung, 1992b, 1993; Pirrung and Bu¨chel, 1994).
3. Sedimentary succession The maar structure at Eckfeld was cored to a depth of 66.5 m (Negendank et al., 1982). The sequence displayed by the core was roughly subdivided into basal pyroclastics followed by sideritic bituminous laminites and inorganic clay and silt. Within the bituminous laminites, a change from frequent diatomitic intercalations to non-diatomitic sediments was noted. The succession was interpreted as representing three successive stages of a maar lake. As a result of detailed field studies, the upper inorganic clay and silt is now interpreted as weathered bituminous laminites (Pirrung, 1992a). This is strongly supported by the mode of preservation of the plant remains with the organic material highly degraded or even missing. The Eckfeld structure is completely surrounded by a coarse marginal facies including varying proportions of volcanoclastic components (Pirrung, 1992a, 1993). Due to field conditions, scientific excavations in Eckfeld are still limited to a few meters of the nondiatomitic bituminous laminites which have been studied in detail. They are characterized by laminitic
9
‘oilshales’ and siltstones (Mingram, 1994) with frequent turbiditic intercalations (Lutz, 1993a; Mingram, 1994). The organic fraction is mainly composed of fine-grained plant debris and the resistant remains of algae, for the most part chlorococcaleans like Botryococcus and Tetraedron (Wilde et al., 1993; Mingram, 1994). Apparently, plant macrofossils are more or less evenly distributed throughout the sequence under investigation. However, coarse and robust plant debris such as wood fragments and large fruits=seeds is concentrated in the turbiditic layers.
4. Physical setting of the flora The Palaeogene Eifel was traditionally thought of as a gently dipping peneplain devoid of any relief. Recently, a distinct magnetic anomaly with remnants of deeply eroded volcanic structures of Eocene to early Miocene age in the centre (‘Kelberger Hoch’) was interpreted as probably indicating major volcanogenic uplift north of Eckfeld (Bu¨chel and Pirrung, 1993; Pirrung and Bu¨chel, 1994) at that time. More evidence for some pre-Oligocene relief and contemporaneous drainage systems in the area was summarized by Lo¨hnertz (1994). Following Lo¨hnertz (1994), the Eckfeld maar today is located just between the completely eroded ‘Kelberger Hoch’ and an area with remnants of the Palaeogene surface. The Eckfeld maar and two adjacent volcanic structures are aligned along a NNE–SSW-trending major fault that can even be recognized in the Variscan basement (Meyer et al., 1994). After eruption(s) came to an end, the maar crater was occupied by a lake which soon became meromictic with algae flourishing in the mixolimnion (Wilde et al., 1993). Algal blooms were most probably seasonally controlled (Mingram, 1994). At least at times, the water may have been slightly alkaline, and some change in the water chemistry is possibly indicated by the disappearance of diatoms with time (Wilde et al., 1993). The occurrence of unionid bivalves may be taken as evidence that the lake at times was somehow connected to a fluvial system (Groh and Jungbluth, 1994). The dimensions of the Middle Eocene crater of Eckfeld have been estimated mainly from field data
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by Pirrung (1993). The diameter may have been approximately 850–950 m, with an inclination of the initial slopes of about 20º. Water depth never exceeded 170 m (Pirrung, 1993), and was probably not more than 50 m (Mingram, 1994). There is some evidence supporting some persistent instability of the slopes, and any shallows never reached significant extent (this paper).
5. The plant macrofossils Except for diatoms and chrysophycean cysts from the core (Zolitschka, 1993; Wilde et al., 1993; Mingram, 1994), all the plant material presently known from the Eckfeld site was collected during the excavations. As a consequence, all of the information on the plant macrofossils (leaves, fruits=seeds, flowers; Table 1 and Appendix A) presented below has been derived from a limited part of the succession. No attention has been paid as yet to the frequently occurring wood remains. After a first account of the plant remains had been published by Wilde (1989b), Wilde and Frankenha¨user (1993) presented a more extensive summary (for fruits=seeds see also Ja¨ckel and Frankenha¨user, 1994). It includes a list of taxa that was recently corrected by Wilde (1995). Detailed work on selected taxa started with algae (Wilde et al., 1993), bryophytes (Wilde, 1990), and ferns (Frankenha¨user and Wilde, 1993b). More recently, some angiosperm taxa have been studied, including pterocaryoid and engelhardioid juglandaceous fruits (Frankenha¨ user and Wilde, 1994; Wilde and Frankenha¨user, 1995), possible fruits of Burseraceae (cf. Canarium Linne´; Mai, Wilde and Frankenha¨user, in prep.), and a distinct type of scleromorphic dicotyledonous leaves (Pungiphyllum waltheri Frankenha¨user et Wilde, 1995). 5.1. Cryptogams Fossil bryophytes in general are rare. Surprisingly, there are now more than 40 specimens existing from Eckfeld. Most of them apparently represent a distinct type with tufts of leafy stems still adhering to a piece of wood or bark (Plate I, 2; Wilde, 1990) thus indicating an epiphytic habit. Except for
one specimen (Plate I, 1), there are no sporophytes preserved. Most remarkable is a single tiny fragment of a Frullania-like jungermannialean leafy liverwort that may have been epiphyllous (work in progress). There is a suite of ferns in Eckfeld that is common to other Middle Eocene localities like Messel and the Geiseltal area (Frankenha¨user and Wilde, 1993b; Wilde, 1995). Most common in Eckfeld are Schizaeaceae with Lygodium kaulfussii, including a considerable number of fertile pinnae fragments (Plate I, 6,7), and a single sterile frond fragment of Ruffordia, an extinct genus. In addition, there are some fragments of sterile pinnae of Osmunda lignitum (Osmundaceae; Plate I, 8), and a single fertile frond fragment of ‘Rumohra’ recentior (Polypodiaceae s.l.; Plate I, 3, 4). Most recently, a fragment of a fertile pinna was found that is apparently quite similar to extant Didymochlaena or Lindsaea (Polypodiaceae s.l.; Plate I, 5; work in progress). An interesting fact is the common occurrence of pinnae fragments of the ‘mangrove fern’ Acrostichum (Polypodiaceae). At present the genus is more or less restricted to distal mangrove-related habitats. Its occurrence in the sediments of Eckfeld is yet another example for a shift in the ecological amplitude since Palaeogene times. 5.2. Gymnosperms As in most European Tertiary localities, remains of gymnosperms are limited to conifers in Eckfeld. There are a single badly preserved twig fragment and a possible cone scale (Plate IV, 1) of taxodiaceous affinity. Cephalotaxus cf. messelensis is represented by an isolated leaf with diagnostic cuticular structures. Another unique specimen is the terminal part of a cupressaceous branch with fused facial and marginal leaves and Thuja-like cuticular structures. Because fusion of facial and marginal leaves is unknown from extant taxa as previously noted by Ferguson (1971), the specimen is assigned to Libocedrites salicornioides (Unger, 1841) Endlicher, 1847 (manuscript in preparation). 5.3. Monocotyledons There are a number of monocotyledonous leaf fragments from Eckfeld, most of them of unknown
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11
Table 1 The plant macrofossils of Eckfeld after the 1995 campaign with taxa that have been assigned at least to family level Bryophyta Dicranites sp. Muscites sp. Jungermanniales sp. (cf. Frullania sp.) Filices Osmundaceae (C) Schizaeaceae Polypodiaceae s.l. (C)
Coniferae Taxodiaceae ? Cupressaceae (C) Cephalotaxaceae Dicotyledoneae Anacardiaceae (?) Betulaceae (C) Burseraceae (?) Caprifoliaceae Hamamelidaceae (?) Juglandaceae (C)
Lauraceae Mastixiaceae (C) Menispermaceae (C) Moraceae Myricaceae (C) Papilionaceae (D Leguminosae) (C) Rosaceae (?) Rutaceae (?) Sargentodoxaceae Theaceae Ulmaceae (C) Vitaceae (C) fam. inc. sed. Monocotyledoneae Araceae Arecaceae (D Palmae) (C)
Cyperaceae (C) Liliaceae s.l. (?)
Osmunda lignitum (Giebel, 1857) Stur, 1870 Lygodium kaulfussii Heer, 1861 (C) cf. Ruffordia subcretacea Barthel, 1976 (C) Acrostichum sp. ‘Rumohra’ recentior (Unger, 1841–1847) Barthel, 1976 fragment of a fertile pinna similar to Didymochlaenaand Lindsaea Filicopsida inc. sed. [?Blechnum dentatum (Goeppert, 1836) A. Braun, 1852] fragment of a leafy twig and single cone scale Libocedrites salicornioides (Unger, 1838) Endlicher, 1847 (Cephalotaxus cf. messelensis Wilde, 1989a) cf. Pentoperculum Tiffney, 1994 (fruits) Carpinus-like bracts Canarium n.sp. (fruits) Sambucus sp. (seeds) flower with valvate anthers engelhardioid compound leaves and leaflets engelhardioid fruits (Palaeocarya sp.) Hooleya-like pterocaryoid fruits fragments of platycaryoid male inflorescences cf. Laurophyllum sp. (leaves) cf. Daphnogene sp. (leaves) fruits 3 distinct types of fruits=seeds at least ?Ficus sp. sensu Wilde (1989a), leaves Comptonia sp. (leaves) pods and leaflets compound leaves of Rosa vel Rubus type seeds, at least 1 type cf. Sargentodoxa sp. (fruits) cf. Polyspora saxonica Walther et Kvaˇcek, 1984 (leaves) cf. Ternstroemites dentatus Wilde, 1989a (leaves) ulmoid leaves Zelkova-like leaves vitioid leaves and seeds Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (leaves) a single fragment of a leaf comparable to ‘Araceae sp. 2’ (sensu Wilde, 1989a,b) 1 distinct type of flower fragments of palmate leaves fragments of pinnate leaves fragments of stout spinose rhachises cyperaceous fruits cf. Smilax (leaves)
Families definitely or possibly also represented by spores or pollen are marked by ‘(C)’ and ‘(?)’, respectively (after Nickel, 1994, 1996; pers. commun., 1995)
12
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affinity. The Araceae are represented by a few fragments of a leaf type commonly occurring in Messel. The most frequent type (>130 specimens) of all flowers in Eckfeld is a trimerous palm flower (Plate IV, 2,3) with monocolpate pollen preserved in situ. Fragments of palm leaves are not rare, including both pinnate (Plate II, 12) and palmate organization. Furthermore, there are fragments of stout petioles with hook-like spines referable to the Arecaceae. Some shoots with sheathing bases of linear leaves probably ought to be interpreted as submerged Hydrocharitaceae (Plate II, 8). A few leaves are similar to Smilax (Smilacaceae), while Cyperaceae are known from a number of well preserved fruits (Plate V, 5).
5.4. Dicotyledons Dicotyledons are represented in Eckfeld by numerous leaves, fruits=seeds, and a considerable number of flowers. Juglandaceae are by far the most important element in the assemblage (Plate III) with a great number of leaves, leaf fragments, and leaflets of different types. The juglandaceous material is characterized by peltate glands with a unicellular base that are typical for the extant members of the family (Plate III, 10). Number and affinities of genera and species included in the Eckfeld material will be the subject of a future study. Hooleya-like fruits are common and were assigned to
PLATE I Cryptogams. Scale always 1 cm. 1. Moss with sporophytes and sporangia; PB 1995=364 LS. 2. Epiphytic moss still adhering to a fragment of bark; PB 1995=365 LS. 3. Frond fragment of ‘Rumohra’ recentior sensu Barthel (1976); PB 1995=276 LS. 4. Detail from 3 with fertile pinnae (arrows at sporangia). 5. Fertile pinna fragment of a fern similar to extant Didymochlaena or Lindsaea; PB 1995=274 LS. 6. Lygodium kaulfussii, fertile frond fragment with a number of sporangiophores; PB 1995=278 LS. 7. Lygodium kaulfussii, fertile frond fragment with a number of sporangiophores; PB 1995=277 LS. 8. Sterile pinna fragment of Osmunda lignitum; PB 1995=275 LS. PLATE II (see p. 14) Common leaf types. Scale always 1 cm. 1. Ternstroemites cf. dentatus Wilde, 1989a (Theaceae); PB 1995=208 LS. 2. Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (fam. inc. sed.); PB 1995=82 LS. 3. Compound rosaceous leaf with basal stipules; PB 1995=168 LS. 4. Ulmaceous leaf; PB 1990=374 LS. 5. Leaf of unknown systematic affinity with entire margin and three veins emerging from the base; PB 1995=207 LS. 6. Leaf of unknown systematic affinity with coarse undulate margin; PB 1995=178 LS. 7. Leaf of unknown systematic affinity with asymmetric base and irregular margin; PB 1995=198 LS. 8. Leafy shoot of an aquatic monocotyledon (?Hydrocharitaceae); PB 1995=195 LS. 9. Leaf of Comptonia sp. (Myricaceae), composite figure including part and counterpart; PB 1995=167 LS. 10. cf. Ficus sp. sensu Wilde (1989a,b); PB 1995=199 LS. 11. Vitioid leaf (cf. Vitaceae, Menispermaceae, Aceraceae, etc.); PB 1995=205 LS. 12. Fragment of a pinnate palm leaf (Phoenicites sp.); PB 1995=206 LS. PLATE III (see p. 15) Juglandaceae. Scale 1 cm except for 10, 11. 1. Entire large compound leaf with leaflets attached; PB 1990=59 LS. 2, 3. Isolated leaflets, type 1, with chewed margins; PB 1995=171, 172 LS. 4. Leaflet, type 2; PB 1995=175 LS. 5. Fragment of a small compound leaf with leaflets attached; PB 1995=176 LS. 6, 7. Small entire leaf with leaflets attached; PB 1995=174, 170 LS. 8. Leaflet, type 3, with compound teeth; PB 1995=173 LS. 9. Engelhardioid winged fruit (Palaeocarya sp.); PB 1995=69 LS. 10. Typical juglandaceous peltate trichomes from pterocaryoid fruit. Scale 100 µm. 11. Hooleya-like pterocaryoid fruit. Scale 0.5 cm; PB 1994=267 LS.
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PLATE I
13
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PLATE II
For description see p. 12.
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PLATE III
For description see p. 12.
15
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PLATE IV
Flowers. Scale 0.1 cm, except for 12. 1. ?Taxodiaceous cone scale. PB 1990=22 LS. 2, 3. Male palm flowers; PB 1995=389, 390 LS. 4. Platycaryoid male inflorescence; PB 1990=78 LS. 5–9. Different flowers of unknown affinity; PB 1995=391 LS, PB 1991=14 LS, PB 1990=83 LS, PB 1990=20 LS, PB 1990=346 LS. 10, 11. Probable flowers of Rutaceae; PB 1990=343 LS, PB 1995=392 LS. 12. Pollen from flower in 11. Epifluorescence, scale 20 µm.
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the pterocaryoid alliance (Frankenha¨user and Wilde, 1994). In comparison, only two specimens of engelhardioid winged fruits (Palaeocarya sp.) have been found (Wilde and Frankenha¨user, 1995). Furthermore, there are pollen-bearing male inflorescences, some of them probably platycaryoid (Plate IV, 4), which is apparently supported by a distinct type of pollen that has been observed using epifluorescence. Apart from juglandaceous foliage, there are a number of more or less common leaf types. However, at the moment only a few of them can be assigned to a natural taxon. For example, there are a great number of ulmaceous leaves (Plate II, 4). Another common leaf type is represented by Ternstroemites cf. dentatus (Theaceae) which was previously described from Messel (Plate II, 1). Quite characteristic are those leaves of unknown systematic affinity that have recently been described as Pungiphyllum waltheri Frankenha¨user et Wilde, 1995 (Plate II, 2). A number of large leaves of a vitioid type (cf. Vitaceae, Menispermaceae, Aceraceae etc.) is represented mostly by fragments (Plate II, 11). There are a number of fern-like leaves of Comptonia (Myricaceae) in Eckfeld (Plate II, 9) which although common in Tertiary plant assemblages, only constitute a small percentage of the macrofossils. Another rarity is the two compound rosaceous leaves from Eckfeld, one of which even has stipules preserved (Plate II, 3). Legumes are represented in Eckfeld not only by fragments of leaves and isolated leaflets, but also by a few pods. As with the leaves, many of the fruits=seeds can not be assigned to a natural taxon. There is a taphonomic bias between the turbiditic layers and the regular finegrained sediments. The coarse-grained turbiditic material is characterized by an association of robust fruits=seeds, especially including large specimens that are otherwise rare. Among those, Mastixiaceae (Plate V, 4), Vitaceae, Rutaceae, Anacardiaceae (cf. Pentoperculum Manchester, 1994; Plate V, 13), Sargentodoxaceae (cf. Sargentodoxa; Plate V, 9; Tiffney, 1993), Menispermaceae, and possibly a new species of Canarium (Burseraceae) that was initially assigned to a three-loculed Nyssa have been identified preliminarily. Infructescences (Plate V, 10), tiny fruits and fruits with delicate parts, e.g. winged and fleshy fruits, are mainly known from the fine-grained ‘background’ sediments. Most common (>120 specimens) are Hooleya-like pterocaryoid winged fruits (Plate III,
17
10, 11; Frankenha¨user and Wilde, 1994), and a distinct type of petiolate four-winged fruits of still unknown systematic affinity (Plate V, 7–8). There are also some nice specimens of hairy fruits (Plate V, 11, 12). Some of the taxa known from the turbiditic layers are also represented in the fine-grained sediments, e.g. at least three different species of Menispermaceae fruits=seeds, Rutaceae seeds, and few specimens of Sambucus (Caprifoliaceae) seeds. Due to their delicate nature, flowers and inflorescences are limited to fine-grained layers. At the moment, more than 30 different species may be distinguished with a total of more than 600 specimens. There are both anemophilous and entomophilous flowers, most of them with pollen preserved in situ (Plate IV). It is especially important, that most of the species are known not only from single specimens, but from a large number in some cases. However, due to the fact that studies on the flowers are still in an initial stage, only few of them can be assigned to extant families, like palms, Juglandaceae, and probably Rutaceae.
6. Interpretation A number of mechanisms are relevant for supplying macroscopic plant material to lakes (e.g. Fritz, 1986; Ferguson, 1993). Due to the physical character of the surrounding area, their contributions may be different as pointed out by Ferguson (1993) in dealing with the Middle Eocene lake of Messel. For the slightly younger maar lake of Eckfeld, a basically similar scenario may be envisaged. In both lakes, leaves, fruits, and flowers fell directly from overhanging twigs. Because of taphonomic filtering effects their fossilization potential may have been lower in Messel. The majority of the plant fragments were blown by wind onto the surface of the water. Surface runoff triggered by rainfall washed material from the surrounding slopes into the lakes, but the contribution of tributaries was probably low. It is difficult to estimate the importance of animal vectors in transporting macroscopic plant fragments, especially diaspores, but their contribution may have been considerable if fleshy fruits indicate endozoochory. As a typical maar structure, the Middle Eocene lake of Eckfeld was surrounded by slopes with a
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PLATE V
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high inclination which led to potential instability. Thus, some plant material slid down the slopes directly into the lake. Instability of the slopes in Eckfeld is indicated by frequent coarse-grained layers of turbiditic origin, and by the fact that there is only scarce evidence for anchored aquatic plants like charophytes, Nymphaeaceae, and Hydrocharitaceae. Accordingly, plants possibly related to a herbaceous fringe along the shoreline are missing or rare in the pollen and macrofossil record of Eckfeld. In Messel, such turbiditic layers have rarely been observed while remains of floating Nymphaeaceae and marginal helophytes (ferns, monocots) are common. Therefore, more gentle slopes and a transitional vegetation fringing the lake in Messel may have prevented material from directly sliding into the water. As a further consequence, taphonomic filtering effects were of minor importance in Eckfeld compared to Messel. This may be the reason for a conspicuous amount of woody material in Eckfeld, including twig and bark fragments with tufts of mosses still adhering. It is even possible that some of this material was directly derived from overhanging twigs. Coarse and resistant plant debris, such as wood=bark fragments and tough fruits=seeds was probably initially concentrated along the shoreline. From time to time these accumulations were affected by sliding due to instability of the slopes and subsequently incorporated into turbiditic layers. Therefore, this kind of material is especially abundant in the turbiditic intercalations. A considerable number of Lygodium pinnules in Eckfeld may also be explained by the absence of a taphonomic filter between the slopes of the crater and the lake itself. Possibly, these fast
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growing climbing ferns preferentially covered fresh surfaces that were repeatedly created by slumping. The overall diversity of the upper Middle Eocene vegetation of the Eckfeld area is indicated by about 200 ‘species’ of spores and pollen, and by a great number of leaf and fruit=seed types. However, at present it is difficult to reconstruct the vegetation of the Eckfeld area in detail because the systematic position of some of the more important fruit=seed and leaf types of the taphoflora is still unknown or uncertain. As pointed out above, anchored aquatics and marginal helophytes were underrepresented due to the physical character of the maar structure. It is obvious from leaves, fruits and pollen, that some Juglandaceae were common or even dominant in the surrounding forest. Palms were present, but their importance is difficult to estimate in spite of a number of flowers and some leaf fragments (extant palms usually produce a large number of flowers but do not shed their leaves). Conifers as well as Lauraceae were definitely scarce. Pinaceae and Fagaceae have not been proven by macrofossils, and their contribution to the pollen assemblage is only moderate. Therefore, they probably grew at some distance from the maar lake. A diversity of climbers (e.g. Lygodium, Menispermaceae, Vitaceae) could have covered the edge of the forest as well as fresh surfaces.
7. Interactions Some research has recently been focused on interactions, especially plant–fungal and plant–animal relationships (e.g. Scott and Taylor, 1983; Scott
PLATE V Fruits and seeds. Scale 0.5 cm except for 5 and 6 (scale 0.1 cm). 1–3. Different types of Menispermaceae seeds; PB 1995=370 LS, PB 1993=111 LS, PB 1993=116 LS. 4. Mastixioid fruit; PB 1995=379 LS. 5a–c. Isolated fruit of Cyperaceae; PB 1995=374 LS. 6. Seed of Sambucus sp. (Caprifoliaceae); PB 1995=366 LS. 7. Four-winged fruit of unknown affinity, wings flattened on bedding plane as in most specimens; PB 1993=728 LS. 8. Laterally compressed fruit as in 7, with petiole (arrow) and fruit proper attached; PB 1995=140 LS. 9. Fruit=Seed of cf. Sargentodoxa sp. (Sargentodoxaceae); PB 1995=381 LS. 10. Fragment of infructescence possibly related to Emmenopterys (Rubiaceae) (Manchester, 1994); PB 1995=380 LS. 11. Fruit of unknown affinity showing single seed with tuft of hairs (partly obscured by a leaf fragment below); PB 1993=746 LS. 12. Fruit=seed of unknown affinity with a hairy tail (‘Cypselites’ sp.); PB 1993=747 LS. 13a, b. Single specimen of cf. Pentoperculum Manchester, 1994 (Anacardiaceae), fruit; PB 1995=375 LS. a. From above. b. From below.
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PLATE VI
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and Paterson, 1984; Chaloner et al., 1991; Scott, 1991, 1992; Taylor and Osborn, 1996). As at Messel (Schaarschmidt, 1988), a lot of evidence for a wide spectrum of interactions has been collected at Eckfeld. Because detailed studies of the material are for the most part still outstanding, they will be mentioned only briefly. Epiphytic and even epiphyllous bryophytes are most noteworthy as representing rare evidence of plant–plant relationships. Examples of parasitic=saprophytic plant–fungal associations may be observed particularly on leaves. Most common is a diversity of fungal hyphae in cuticular preparations. In addition, there are some examples of cone-shaped ascomycete fructifications on leaves and fruits (Plate VI, 4, 5). In many of the leaves parts of the lamina have obviously been gnawed away. Different outlines of the resulting injury indicate a considerable diversity of animals feeding on leaves (Plate VI, 6–9), probably most of them arthropods. Another kind of leaf attack related to feeding is documented by traces of mining (Plate VI, 9; e.g. Lang et al., 1995). That some of the mammals fed on leaves is proven by bits of cuticles in the gut of a complete specimen of Propalaeotherium voigti (Plate VI, 11). Another obvious result of plant–animal interactions are galls (Scott et al., 1994). Many of them are caused by insects attaching their eggs to plant parts, thereby modifying growth of the respective tissues. There are a variety of galls preserved on the leaves from Eckfeld (Plate VI, 1–3). A spectacular example is the three-dimensionally preserved cone-shaped structure on some specimens of juglandaceous leaflets (Plate VI, 1).
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In Eckfeld, insect pollination is already indicated by a number of rarely occurring types of large pollen grains (Nickel, 1996). More indirect evidence is derived from a typical architecture in some of the flowers, including highly advanced types like a dorsiventral flower with a tubular corolla (Frankenha¨user and Wilde, 1993a). Moreover, from the Eckfeld site there is even direct proof of a potential insect pollinator with the oldest honeybee presently known. It is still carrying its pollen load in place (Plate VI, 10; Lutz, 1993b).
8. Comparisons with other localities The Eckfeld plant assemblage should finally be compared with other nearby Middle Eocene plant localities in Germany (Fig. 1), like Messel, Helmstedt, the Geiseltal area and the Weißelster Basin. As shown in Table 2, most of the localities are not strictly contemporaneous, while the Geiseltal plant remains represent a number of localities spanning a considerable period of time. However, preliminary Table 2 Stratigraphic range of the localities mentioned in the text Eckfeld Messel Geiseltal >MP14 MP14 MP13 C MP12 MP11
Helmstedt Weißelsterbecken C
C
C C C C
? C C C
PLATE VI Interactions. 1a, b. Juglandaceous leaflet with 3-dimensionally preserved galls (arrows). PB 1995=385 LS. a. Overview. Scale 1 cm. b. Detail from 1a. Scale 0.5 cm. 2, 3. Leaves with different types of circular galls. PB 1995=386, 387 LS. Scale 1 cm. 4. Fruiting bodies of ascomycetes on a leaf surface. PB 1990=6 LS. Scale 0.1 cm. 5. Fruit affected by fungal growth; PB 1995=388 LS. Scale 0.5 cm. 6–9. Gnawed leaves. PB 1995=382-384 LS, PB 1990=68 LS. Scale 1 cm. 10a, b. Bee (Eckfeldapis electrapoides Lutz, 1993b) with pollen load in the metatarsal brush. PE 1992=614 LS. Scale 0.5 cm. b. Small tricolp(?or)ate pollen grains from metatarsal brush of a (unidentified). Negative print from slide, CLSM picture courtesy of LEICA GmbH, Bensheim. Scale 5 µm. 11a, b. Cuticles from gut of Propalaeotherium voigti. a. Scale 100 µm. b. Scale 30 µm.
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comparisons of Middle and Upper Eocene plant assemblages already suggest that differences in age within the Middle Eocene are not as important as different facies represented by their respective sediments (Wilde, 1989a). Therefore, a comparison of these assemblages may provide a simple approach to assess some small-scale differentiation of the Middle Eocene vegetation in Europe (Wilde, 1989b, 1995; Wilde and Frankenha¨user, 1993). Unfortunately, any such comparison can only be based on a qualitative approach and has to remain preliminary at present. Leaves have been described from all of the localities (see compilation in Wilde, 1995), but in none of the assemblages have they been monographed exhaustively. There is a large number of ‘species’ known from the Geiseltal area (Wilde, 1995), but still comparatively few from Helmstedt (Wilde, 1989a) and the Weißelster Basin (Fischer, 1991). With the exception of the Geiseltal area (Mai, 1976), knowledge of fruits and seeds is quite limited. In Messel (Collinson, 1988), and even more so in Eckfeld (this paper) only some of the numerous taxa have been identified or even studied in detail. There is hitherto no record of fruits and seeds from the other localities. Due to different modes of preservation, comparison of the respective taphofloras from the Geiseltal area (sieved material) and from Messel and Eckfeld (compressions) is difficult. Systematics of pollen and spores has been studied extensively in Messel and Eckfeld (Thiele-Pfeiffer, 1988; Nickel, 1994, 1996), but there is no comparably complete record for any horizon of the Geiseltal area or either of the other localities (Krutzsch, 1976, 1992). Similarity of the pollen and spore assemblages of Eckfeld and Messel is expressed by a great number of taxa common to both localities. However, published data from the Geiseltal area already suggest striking differences (Nickel, 1996). Accordingly, most of the ferns, and all of the seed-plant families hitherto recognized in the macrofossil assemblage of Eckfeld are also known from Messel (except for Betulaceae that are represented in Eckfeld by still debatable Carpinus-like bracts). There are many species definitely common to both localities, e.g. conifers (Cephalotaxus messelensis), Caprifoliaceae (more or less identical seeds of Sambucus), Juglandaceae (some common types of leaves and fruits), cf. Sargentodoxa seeds, Theaceae (Polyspora sax-
onica, Ternstroemites dentatus), Ulmaceae (common types of ulmoid and zelkovoid leaves), and Araceae (leaves: ‘Araceae sp. 2’ sensu Wilde, 1989a,b). Furthermore, identical or closely related species are highly probable but have still to be proven in Menispermaceae (seeds), Myricaceae (leaves of Comptonia), Rosaceae (leaves), Rutaceae (seeds), vitioid leaves, Cyperaceae (fruits), and Liliaceae s.l. (leaves of cf. Smilax). In contrast to Messel with one ‘species’ of coniferous foliage and 6 ‘species’ of angiosperm leaves that have also been described from the Geiseltal area, none of the phanerogamous leaves or fruits=seeds of Eckfeld has yet been proven to be identical to a species from the Geiseltal area. Surprisingly, most of the ferns from Eckfeld are common to both Messel and the Geiseltal area (Frankenha¨ user and Wilde, 1993b). Most interesting is the diversity of Juglandaceae represented in the taphocoenoses of Eckfeld and Messel with no macrorecord of the family in the Geiseltal area, Helmstedt and the Weißelster Basin. Conversely, leaves and fruits of Fagaceae are common in the Geiseltal area, but still completely missing in Eckfeld and Messel. The extreme diversity of lauraceous leaves in Messel and a number of species in the Geiseltal area (probably also in Helmstedt and the Weißelster Basin) is in strange contrast to the few lauraceous leaves in Eckfeld. Taxa like Doliostrobus (?Taxodiaceae), and Rhodomyrtophyllum (Myrtaceae) are especially common in the coal bearing areas (Geiseltal area, Helmstedt, Weißelster Basin), have been found in Messel, but are hitherto unknown from Eckfeld. Palaeogeographically, the peat-forming lowlands like the Geiseltal area, Helmstedt and the Weißelster Basin were situated fairly close to the Middle Eocene coastline (Fig. 1). Their respective taphofloras most probably represent the azonal floras of a coastal plain with some peatswamps. In contrast, Messel and Eckfeld were more or less isolated lakes at some distance from the coast. They were surrounded by a zonal flora flourishing in the central part of the Middle Eocene Western European Island. Different edaphic conditions are apparently not reflected in the ferns whose distribution was governed by the humid climate.
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Acknowledgements Excavations at the Eckfeld site were initiated and continued since 1987 by Dr. F.-O. Neuffer, head of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz. Important field observations were supplied by Dr. Herbert Lutz who supervised most of the fieldwork actually done by staff of the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz with the help of volunteers. Some of the plant material was prepared by Uta Jaeckel (Mainz). Dr. Birgit Nickel (Frankfurt) generously provided us with her unpublished data on the pollen assemblage from Eckfeld. Among others, Drs. Margaret Collinson (London) and Kurt Goth (Freiberg) spent some time with us discussing the Eckfeld fruits and seeds. Excavations and research in Eckfeld have been subsidized by the government of Rheinland-Pfalz and by the Deutsche Forschungsgemeinschaft (Project Nr. II C Ne-440=1 and =2). Our fruitful cooperation has been steadily supported by both the Naturhistorisches Museum Mainz=Landessammlung fu¨r Naturkunde Rheinland-Pfalz and the Forschungsinstitut Senckenberg, Frankfurt am Main. Thanks are due to Professor David K. Ferguson, Vienna, for improving our English.
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Scott, A.C., Taylor, T.N., 1983. Plant=animal interactions during the Upper Carboniferous. Bot. Rev. 49, 259–307. Scott, A.C., Stephenson, J., Collinson, M.E., 1994. The fossil record of leaves with galls. In: Williams, M.A.J. (Ed.), Plant Galls. Syst. Assoc. Spec. Vol. 49, 447–470. Taylor, T.N., Osborn, J.M., 1996. The importance of fungi in shaping the paleoecosystem. Rev. Palaeobot. Palynol. 90, 249– 262. Thiele-Pfeiffer, H., 1988. Die Mikroflora aus dem mitteleoza¨nen ¨ lschiefer von Messel bei Darmstadt. Palaeontographica B O 211, 1–86. Tiffney, B.H., 1993. Fruits and seeds of the Tertiary Brandon Lignite. VII. Sargentodoxa (Sargentodoxaceae). Am. J. Bot. 80, 517–523. Tobien, H., 1961. Ein Lophiodon-Fund (Tapiroidea, Mamm.) aus den niederhessischen Braunkohlen. Notizbl. Hess. Landesamtes Bodenforsch. Wiesbaden 89, 7–16. Von der Brelie, G., 1969. Neue Untersuchungen im Alttertia¨r von Eckfeld bei Manderscheid (Eifel). Fortschr. Geol. Rheinland Westfalen 17, 27–40. Weber, C.O., 1853. Ueber das Braunkohlenlager von Eckfeld in der Eifel. Verh. Naturhist. Ver. Preuss. Rheinlande Westfalens 10, 409–415. White, J.D.L., 1991. The depositional record of small, monogenetic volcanoes within terrestrial basins. In: Fisher R.V., Smith, G.A. (Eds.), Sedimentation in Volcanic Settings. SEPM Spec. Publ. 45, 155–171. Wilde, V., 1989a. Untersuchungen zur Systematik der Blattreste aus dem Mitteleoza¨n der Grube Messel bei Darmstadt (Hessen Bundesrepublik Deutschland). Courier Forsch.-Inst. Senckenberg 115, 1–213. Wilde, V., 1989b. Vorla¨ufige Mitteilungen zur Flora aus dem Alttertia¨r von Eckfeld—Ergebnisse einer ersten Durchsicht des Fundmateriales aus den Grabungen von 1987 und 1988. Mainzer Naturwiss. Arch. 27, 23–31. Wilde, V., 1990. Moosreste aus dem Alttertia¨r von Eckfeld bei Manderscheid in der Eifel. Mainzer Naturwiss. Arch. 28, 1–6. Wilde, V., 1995. Die Makroflora aus dem Mitteleoza¨n des ¨ bersicht und Vergleiche. Hallesches Geiseltalgebietes, kurze U Jahrb. Geowiss. B 17, 121–138. Wilde, V., Frankenha¨user, H., 1993. Initial results on the importance of a flora from the Middle Eocene of Eckfeld (Eifel, W.-Germany). In: Negendank, J.F.W., Zolitschka, B. (Eds.), Paleolimnology of European Maar Lakes. Lecture Notes Earth Sci. 49, 499–503. Wilde, V., Frankenha¨user, H., 1995. Flu¨gelfru¨chte engelhardioider Juglandaceen aus dem Mitteleoza¨n von Eckfeld bei Manderscheid (Eifel). Mainzer Naturwiss. Arch. 33, 47–52. Wilde, V., Frankenha¨user, H., Lutz, H., 1993. Algenreste aus den mitteleoza¨nen Sedimenten des Eckfelder Maares bei Manderscheid in der Eifel. Mainzer Naturwiss. Arch. 31, 127–148. Zolitschka, B., 1993. Palaeoecological implications from the sedimentary record of a subtropical maar lake (Eocene Eckfelder Maar; Germany). In: Negendank, J.F.W., Zolitschka, B. (Eds.), Paleolimnology of European Maar Lakes. Lecture Notes Earth Sci. 49, 477–484.
Appendix A Summary of the probable life forms, dispersal mechanisms and physiognomic signatures of the Eckfeld taxa, based on the representation of the different plant parts in the assemblages (compiled by D.K. Ferguson) Family ALGAE Botryococcaceae
Chrysophyceae Zygnemataceae Bryophytes Dicranaceae s.l. Jungermanniaceae Incerta sedis
Other organs
Primary dispersal mechanism
Botryococcus Tetraedron Chara
coenobia unicells gyrogonites
gen. indet. Ovoidites sp.
cysts zygospores
exozoo-nautochory exozoo/nautochory endozoo-/exozoo/ nautochory exozoo-/nautochory exozoo-/nautochory
Dicranites† cf. Frullania Muscites†
FERNS AND FERN-ALLIES Lycopodiaceae Camarozonosporites eocaenicus Osmundaceae Osmunda lignitum/Baculatisporites primarius Polypodiaceae s.l. Acrostichum sp. cf. Didymochlaena/Lindsaea Laevigatosporites discordatus, L. haardtii Polypodium (Verrucatosphorites tenellis) Rumohra recentior Schizaeaceae Cicatricosisporites dorogensis, C. paradorogensis Lygodium kaulfussii/Leiotriletes maxoides, L. microadriennis cf. Ruffordia subcretacea Schizaea (Microfoveolatosporites granuloides)
Pollen= spores
Diaspores
x x x
capsules
wind wind wind
chamaephyte chamaephyte chamaephyte
wind wind
chamaephyte hemicryptophyte
x(2)
wind wind wind
geo-/hemicryptophyte hemicryptophyte geo-/hemicryptophyte
x
wind
geo-/hemicryptophyte
x(2)
wind wind
geo-/hemicryptophyte geo-/hemicryptophyte
x(2)
wind
geo-/hemicryptophyte
wind wind
geo-/hemicryptophyte geo-/hemicryptophyte
wind endozoochory wind endozoochory endozoochory wind wind wind
phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte phanerophyte
x x x x
sporangia
x
x
sporangia
x
sporangiophores x
x x x x
x
Life form
x x x x
x
Deciduous= Evergreen
Leaf margin entire
E E E E E E E D/E
25
GYMNOSPERMS Araucariaceae? Araucariacites europaeus Cephalotaxaceae Cephalotaxus cf. messelensis Cupressaceae s.s. Cupressacites insulipapillatus Libocedrites salicorniodes Ephedraceae Ephedra (Ephedripites eocenipites) Pinaceae Cathaya (Pityosporites microalatus) Pinus (Pityosporites labdacus) Taxodiaceae gen. indet.
Leaves
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Charophyceae
Taxon
26
Appendix A (continued) Family
Taxon
Aquifoliaceae Araliaceae
Pollen/ spores
?
x
Diaspores
Other organs
x
Primary dispersal mechanism
Life form
Deciduous/ Evergreen
Leaf margin entire
wind endozoochory
phanerophyte phanerophyte incl. liana phanerophyte phanerophyte—liana
D/(E) D/E
yes/no yes/no
D/E E
yes/no yes
D/E
yes/no
D D
no no
D
no
D/(E)
yes
E D
yes no
E
rarely
D/E
yes/no
chamaephyte/ phanerophyte phanerophyte
D/E
yes/no
D
no
chamae-/geo-/ hemicrypto-/ phanero-/therophyte phanerophyte phanerophyte phanerophyte phanerophyte incl. liana phanerophyte phanerophyte phanerophyte
D/E
yes/no
E D/E D/E E
yes/no yes/no yes/no mostly
D D/E D
no no no
llex (llexpollenites iliacus) Hedera (Araliaceoipollenites reticuloides) gen. indet. (Araliaceoipollenites euphorii, A. sp.)
x x
endozoochory endozoochory
x(2)
endozoochory
Alnus (Ainipollenites verus) Betula (Trivestibulopollenites betuloides) Carpinus type
x x
wind wind/endozoochory
x(3)
Burseraceae Caprifoliaceae
Bombacacidites kettigensis, B. tilioides, B. sp. Canarium Sambucus sp.
Chloranthaceae
Emmapollis pseudoemmaensis
x
endozoochroy
Clethraceae?/ Cyrillaceae? Ericaceae
Tricolporopollenites megaexactus
x
wind/endozoochory
Ericipites callidus, E. ericius, E. sp. Eucommia (Tricolporopollenites parmularius) Spinaepollis spinosus
x(3)
wind/endozoochory
x
wind
?
Archetype Quercus (Tricolpopollenites asper) gen.indet Compositoipollenites rhizophorus
x x ? x
auto-/endozoo-/ exozoo-/myrmeco-/ nautochory dyschory dyschory autochory/wind wind/endozoochory
Betulaceae s.l.
Bombacaceae
Eucommiaceae Euphorbiacaea
Fagaceae Hamamelidaceae Icacinaceae Juglandaceae
Carya (Caryapollenites circulus) Engelhardia type Platycarya type
x
wind/endozoochory wind/endozoochory
x x
x x
x x x
endozoochory endozoochory
flowers
x x
flowers
dyschory wind wind
(hemicrypto)-/ phanerophyte incl. liana phanerophyte chamaephyte/ phanerophyte chamaephyte/ phanerophyte phanerophyte phanerophyte (hemicryptophyte)/ phanerophyte hemicrypto-/ phanerophyte phanerophyte
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ANGIOSPERMS DICOTYLEDONS: Aceraceae Acer (Aceripollis striatus) Anacardiaceae cf. Pentoperculum†
Leaves
Apendix A (continued) Family
Taxon
Leaves
Loranthaceae s.l.
Gothanipollis gothanii
Mastixiaceae Menispermaceae
gen. indet. gen. indet.
?
Moraceae
Ficus
?
Myricaceae
Comptonia Triatriopollenites bitutius, T. rurensis Duplopollis myrtoides Monocolpopollenites crassiexinus Nyssa (Tricolporopollenites kruschii) Olax (Olaxipollis matthesii)
x
Lauraceae
Myrtaceae Nymphaeaceae Nyssaceae Olacaceae Oleaceae Onagraceae
Diaspores
Other organs
x(4)
x x x
x
x
x
x
x
Life form
Deciduous/ Evergreen
Leaf margin entire
?
phanerophyte
D/E
yes/no
wind endozoochory endozoochory autochory/wind/ nautochory endozoochory
D E E D/E
no yes yes yes
(D)/E
yes
E D/E
yes no
(D)/E
yes
D D/E
no yes/no
endozoochory endozoochory
x x
endozoo-/nautochory endozoo-/nautochory
phanerophyte hydrophyte
E H
yes H
x
endozoochory
phanerophyte
D
yes/no
x
endozoochory
E
yes
x
wind/endozoochory
phanerophyte incl. liana phanerophyte
D/E
yes/no
x(2)
wind/endozoochory
H
H
?
endozoochory
D/E
no
D D/E
yes yes/no
D
rarely
D/E
mostly
x x
x x(3C)
endozoochory endozoochory
endozoochory
Rosaceae
Rosa/Rubus
Rubiaceae Rutaceae
Emmenopterys gen. indet.
?
Salicaceae
Tricolpopollenites retiformis
x
wind
Sapotaceae
Tetracolporopollenites spp. div.
x(6-7)
endozoochory
? x(1C)
flowers
wind autochory/wind/ endozoochory
(chamae-/geo-)/ hemicrypto-/ (phanerophyte) (hemicryptophyte)/ phanerophyte incl. liana phanerophyte (hemicryptophyte-)/ phanerophyte incl. liana chamaephyte/ phanerophyte phanerophyte
27
x(2)
phanerophyte phanerophyte phanerophyte chamae-/hemicrypto-/ phanero-/therophyte chamaephyte/ phanerophyte phanerophyte (hemicryptophyte)/ phanerophyte, mostly liana phanerophyte incl. liana phanerophyte phanerophyte
x
Tricolporopollenites microreticulatus Corsinipollenites oculusnoctis, C. verrucatus
Primary dispersal mechanism
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Leguminosae
Plicatopollis hungaricus, P. lunatus, P. plicatus, P. potoniei Pterocarya type cf. Laurophyllum† cf. Daphnogene† gen. indet.
Pollen/ spores
28
Appendix A (continued) Taxon
Sargentodoxaceae Sterculiaceae
cf. Sargentodoxa sp. Reevesia (Reevesiapollis triangulus vel eocaenicus) Porocolpopollenites calauensis, P. rarobaculatus, P. vestibulum cf. Polyspora saxonica cf. Ternstroemites dentatus Intratiporopollenites ceciliensis, I. minimus, I. cf. maxoides
Symplocaceae Theaceae Tiliaceae
Ulmaceae s.l.
Leaves
Incerta sedis
Punigiphyllum waltheri
x
Smilacaceae
Milfordia hungarica, M. incerta, M. minima cf. Smilax
Sparganiaceae
Sparganiaceaepollenites spp.
Primary dispersal mechanism
Life form
Deciduous/ Evergreen
Leaf margin entire
x
endozoochory wind
phanerophyte-liana phanerophyte
D D/E
yes mostly
x(3)
endozoochory
phanerophyte
(D)/E
yes/no
phanerophyte phanerophyte (chamae-/ hemicrypto-)/ phanerophyte phanerophyte phanerophyte
E E D/E
yes/no no rarely
x x(2)
wind endozoochory wind/endozoo-/ exozoo-/ nautochory endozoochory wind
D/(E) D
yes/no no
x(4)
wind wind endozoochory
phanerophyte phanerophyte phanerophyte, mostly liana ?
D D D/(E)
no no no
?
no
x(3)
Vitaceae
gen. indet. Graminidites eifelensis, G. sp.
Other organs
x x
x x x
hydrocharitaceae Poaceae (Gramineae) Restionaceae
Diaspores x
Celtipollenites intrastructurus Polyporopollenites undulosus, P. verrucatus Ulmus type Zelkova type gen. indet.
MONOCOTYLEDONS: Araceae gen. indet. Arecaceae gen. indet. (Palmae) Cyperaceae gen. indet.
Pollen/ spores
x x(2CG)
?
x x
?
x
? x(2) x(3)
x
flowers, spines x
endozoochory? endozoo-/ nautochory endozoo-/exozoo-/ nautochory nautochory wind/endozoo-/ exozoochory ? endozoochory?
x(2)
endozoo-/ nautochory
hemicryptophyte? phanerophyte incl. liana geo-/hemicrypto-/ therophyte hydrophyte geo-/hemicrypto-/ phanero-/therophyte geo-/ hemicryptophyte phanerophyteliana? hydrophyte
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Family