two new late carboniferous neuropteris species (medullosales) from saarland, germany and their...

Botanical Journal of the Linnean Society, 2002, 139, 193–205. With 24 figures

INTRODUCTION

During the Westphalian Epoch (300–310 Ma, Menninget al., 2000) swamp vegetation covered large areas oftropical land and its fossilized remains are now foundabundantly in North America, Europe and the FarEast. The vegetation is often referred to as the CoalForests because of the thick coals that it generated.This was an ‘ice-house’ world and provides the bestavailable pre-Quaternary analogue for conditionstoday (Gastaldo et al., 1996; Cleal & Thomas, 1999).Understanding the relationship between tropical vegetation change and climate in the Westphalianmay thus provide insights into what is influencingclimate change today.

During the Westphalian, Europe was undergoingsignificant geographical changes due to Variscan tec-tonic activity (the result of the collision between Gondwana and Laurussia). A broad belt of lowlandswamps persisted during most of this time across theVariscan Foreland. The extensive coal forests growingon these lowland swamps have resulted in the series

of coalfields now found between Britain and Poland.Further south, Variscan tectonics were pushing up a chain of mountains, in which several relatively small intramontane basins with upland coal forestsdeveloped, most notably in Saar-Lorraine, CentralBohemia, Zwickau (Saxony) and Svoge (western Bul-garia). Stanislav Oplustil (pers. comm.) has estimatedthat the Central Bohemian Basin was 1400–1800mabove sea level, but the elevation of the other basinssuch as Saar-Lorraine is unknown. Determining theelevations of these upland basins and their forests isobviously important to understand properly theVariscan evolution of Europe.

Differences in composition between the upland andlowland coal forests may help solve this problem.Gothan (1951, 1954) argued that the upland forestswere significantly different in the Westphalian butthis view has since been challenged (e.g. Storch, 1980).The discovery of two new and apparently endemicspecies of Neuropteris in the Bolsovian (middle West-phalian) of Saar-Lorraine (Cleal, 1985) therefore hassome importance for this problem. These species areformally described and named in this paper, and theirpalaeogeographical significance discussed.

© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 193–205 193

Two new late Carboniferous Neuropteris species(Medullosales) from Saarland, Germany and theirpalaeobiogeographical significance

CHRISTOPHER J. CLEAL

Department of Biodiversity & Systematic Biology, National Museums & Galleries of Wales, Cathays Park, Cardiff, CF10 3NP, UK

Received November 2001; accepted for publication February 2002

Two new species (Neuropteris terminiscus and N. kneuperi) are described from the Bolsovian (‘Westphalian C’)of Saarland. N. terminiscus had previously been misidentified as Paripteris pseudograndinioides, but the pinnaeare clearly imparipinnate and thus cannot belong to that morphogenus. N. kneuperi had previously been mis-identified as Neuropteris obliqua and N. heterophylla, both which appear to be absent from Saar-Lorraine. This isconsidered further evidence of the distinctive nature of the Westphalian vegetation of the Saar-Lorraine Basin, espe-cially among the seed-plants. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society,2002, 139, 193–205.

ADDITIONAL KEYWORDS: cuticles – coal forests – pteridosperm – Westphalian.

Corresponding author. E-mail: [email protected]

MATERIAL AND METHODS

All of the hand-specimens are in the collections of the Saarbrücken Mining School, Saarbrücken,Germany. They were photographed using a 35-mmSLR camera with extension tubes, mounted on a copy-ing stand; lighting was with four background flood-lights and two low-angle spot-lights. Terminology for describing the frond architecture is as used byCleal & Shute (1991). Cuticles were prepared usingstandard maceration techniques (Cleal & Zodrow,1989) and mounted on glass slides with CanadaBalsam. The slides are now stored in the Palaeon-tology Department, Natural History Museum,London. The cuticles were photographed using a LeitzOrtholux II microscope with brightfield illumina-tion or differential interference phase (Normarski)contrast.

SYSTEMATIC PALAEOBOTANY

MORPHOGENUS NEUROPTERIS (BRONGN.) STERNB.,1825, NOM. & ORTH. CONS.

Diagnosis: Following the emendation by Cleal et al.(1990) and Cleal & Shute (1995) the main diagnosticfeatures are as follows. Fronds bipartite, with tri- oroccasionally quadripinnate primary rachis branches.Orbicular cyclopterids absent from the lower part offrond. Pinnules basally constricted. Lateral veinsbroadly arched or flexuous. Pinnules hypostomatic.Stomata anomocytic, brachyparacytic, or occasionallycyclocytic, and subsidiary cell periclinal walls notthickly cutinized relative to the other cells. Trichomesusually abundant only on the abaxial pinnule surface.Adaxial epidermal cells differentiated between costaland intercostal fields. Abaxial cuticle with prominentintercellular flanges.

Basionym: Filicites (Sect. Nevropteris) Brongn., 1822.

Type species: Neuropteris heterophylla (Brongn.)Sternb. 1825. The type of this species (and thus of the genus) is now taken as that figured by Brongniart (1831: pl. 71). This has been conservedagainst the specimen figured by Brongniart (1822:pl. 2, fig. 6) with the generic protologue (see Laveine,1998).

NEUROPTERIS TERMINISCUS CLEAL SP. NOV.1971 Neuropteris pseudogigantea Potonié; Germer,

p. 29; text-fig. 32; pl. 6, fig. 4.1975 Paripteris pseudogigantea (Potonié) Gothan;

Doubinger & Germer, p. 19; text-fig. 10; pl. 9,figs 1–4 (non pl. 8, fig. 6 = P. pseudogigantea)

Etymology: The epithet refers to the small apical pinnules.

Diagnosis: Ultimate pinnae parallel-sided except near the apex, where they taper. Apical pinnulessmall, subrhomboidal, 10–15mm long. Lateral pin-nules 7–20mm (average ª 12mm) long. Larger pinnules robust (length/breadth ratio ª 2), usually linguaeform, rarely subfalcate; smaller pinnulesrounder and more isodiametric. Pinnule apex roundedor slightly flattened. Pinnule base cordate with promi-nent basiscopic and acroscopic auricles, becomingpartly fused to the rachis towards the pinna terminal.Prominent midvein extends for up to two-thirds ofpinnule length. Lateral veins broadly arched, fork fourto five times, and meet pinnule margin at 80–90°; nervation density 40–45 per cm on pinnule margin.

Holotype: Specimen C/1200, Saarbrücken MiningSchool (Figs 1,2 and 5). Seam 13 (middle SulzbachFormation, middle Bolsovian), Friedrichsthal Colliery,12 km NNE of Saarbrücken, Saarland, Germany.

Paratypes: In addition to the holotype, there are sixother specimens in the Saarbrücken Mining SchoolCollection that show large parts of ultimate or penultimate pinnae (C/5, C/6, C/510, C/1186, C/1198,C/1699). They all originated from the roof of Seam 13,from three localities near Friedrichsthal in the Saarland Coalfield (the Friedrichsthal, Helene andMaybach collieries).

Description: The fronds are at least tripinnate, withrachises that have irregular but marked longitudinalstriations. All orders of rachis are relatively robust:antepenultimate rachises are ª 5mm wide, penulti-mate rachises 1–5mm wide, and ultimate rachises1–2mm wide. The most complete specimen is of a trip-innate frond segment with five penultimate pinnae in a subopposite arrangement (Fig. 8). The ante-penultimate rachis is difficult to make out but the

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© 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 193–205

Figures 1–7. Neuropteris terminiscus sp. nov. From Seam 12 (Sulzbach Formation, Bolsovian), Saarland, Germany.Fig. 1. C/1200 (holotype). Scale bar = 6.7mm. Figs 2 and 5. Close-ups of pinnules in Fig. 1. Scale bar = 1.7mm. Fig. 3.C/1186. Scale bar = 5mm. Fig. 4. C/1699. Scale bar = 1.7mm. Fig. 6. C/6. Scale bar = 5mm. Fig. 7. C/510. Scale bar = 5mm.Figs 1,2 and 5 from Friedrichsthal Colliery, Friedrichsthal; Figs 3,6 and 7 from Helene Colliery, near Friedrichsthal; Fig. 4,locality not recorded.

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TWO NEW GERMAN SPECIES OF NEUROPTERIS 195


1 2 3

4 5

6 7

penultimate rachises appear to have been originallyattached at ª 70–80°. From the way the pinnules lobe,the fragment is clearly from high in the antepenulti-mate pinna. However, none of the attached penulti-mate pinnae are complete, and thus this specimenprovides no evidence of the shape of the antepenulti-mate segment. No intercalated pinnae or pinnules canbe seen attached to the antepenultimate rachis, butthis is probably due to poor preservation. It is notice-able that the ultimate pinnae are becoming shorterand less divided towards their proximal end, whichwould support the idea that there were intercalatedpinnae attached to the antepenultimate rachis(compare with middle left of the figure shown inLaveine et al. 1998: pl. 9, fig. 2). It is likely thereforethat the preserved fragment probably represents anear-terminal part of a primary rachis branch (sensuCleal & Shute, 1991).

The penultimate pinnae are parallel-sided for mostof their length, but taper slightly in their proximalpart, near their attachment with the antepenultimaterachis. No specimen has been found to show the formof the penultimate terminals. The penultimaterachises carry ultimate pinnae either alternately orsuboppositely arranged. The last order pinnae are

parallel-sided for most of their length (Figs 3 and 6).In their distal parts, these pinnae taper gradually toa small, subrhomboid apical pinnule, 10–15mm long(Figs 1 and 7). Lateral pinnules are attached alter-nately or suboppositely to the ultimate rachises,usually at 80–90°. (Figs 1–6), tending to as little as 50°towards the pinna apex (Fig. 7).

The lateral pinnules can be up to 23mm long (Doubinger & Germer, 1975: pl. 9, fig. 2), but12–15mm is the more normal length (Figs 2,4,5).Typical pinnules tend to be robust (length :breadthratio about 2) and parallel-sided for most of theirlength (Figs 4,5), but there are also some more elon-gate subfalcate forms (Fig. 2). The pinnule apex isusually rounded (e.g. Fig. 4) sometimes tending tobluntly pointed (Fig. 2). Along most of the pinnae, the pinnules have a constricted, cordate base, oftenwith well developed acroscopic and basiscopic lobes(Figs 4,5). Towards the pinna apex, however, the pin-nules become more broadly fused to the rachis (Fig. 7).

A strongly marked midvein extends for one half totwo-thirds of the length of the pinnule, before dividingto produce a fan of finer, dichotomous veins (Figs 2,4and 5); in the smaller pinnules it may be less than halfthe length of the pinnule (Fig. 7). Lateral veins are

196 C. J. CLEAL


Figure 8. Neuropteris terminiscus sp. nov., C/1198. From Seam 12 (Sulzbach Formation, Bolsovian), Helene Colliery,near Friedrichsthal, Saarland, Germany. Scale bar = 5mm.

alternately emitted at <10° from the midvein, andthen arch broadly to meet pinnule margin at 80–90°.Between the midvein and the pinnule margin, alateral vein may fork at <10°, between two and fourtimes, depending partly on the width of the pinnule.Nervation density averages 40–45 veins per cm onpinnule margin.

Adaxial cuticles have well-developed anticlinalflanges of more or less elongate cells, whose long axeslie approximately parallel to the veins (Fig. 9). Alongthe veins, the cells are mainly elongate and subrec-tangular, up to 200mm long and 10–20mm. Between

the veins, they are less elongate and more polygonal,up to 100mm long and 30mm wide. Very rare 50-mmwide multicellular trichomes occur, mainly along themidvein (Fig. 10).

Only a few fragments of abaxial cuticle wereobtained and these show a clear division between thecostal and intercostal fields. Costal epidermal cells areelongate subrectangular to subrhomboidal, up to110mm long typically 15–30mm wide (Fig. 12). Inter-costal cells are irregularly polygonal, usually some-what elongate, with their long axis aligned parallel to the venation, up to 70mm long and 20mm wide



9 10

11 12

Figures 9–12. Neuropteris terminiscus sp. nov. Cuticles from pinnules from Seam 13 (Sulzbach Formation, Bolsovian). Fig. 9. V.62985; adaxial cuticle. Fig. 10. V.69989; trichome on adaxial cuticle. Figs 11,12. V.62989; abaxial cuticles. Fig. 9 photographed under brightfield illumination, scale bar = 50 mm; Figs 10–12 photographed using NomarskiContrast, scale bars = 25 mm.

(Fig. 11). The faint impression of stomata can be seenin the intercostal fields. The stomata appear to beanomocytic, but otherwise their structure is difficultto ascertain.

Comparisons: The holotype of Neuropteris doubravicaPurkynová (1971), from the Langsettian Karviná For-mation of Upper Silesia, has very similar linguaeformpinnules, venation density, and small apical pinnules(Purkynová, 1971: pls 6–7). However, some of the otherspecimens figured by Purkynová (1971: pls 8–9) havemore elongate pinnules with a thick, Alethopteris-likemidvein and the species in fact probably belong to Neuralethopteris Cremer ex Laveine (1967) (see Cleal& Shute, 1995). None of the Saarland specimens showsuch neuralethopterid-like pinnules.

The larger pinnules of Neuropteris bourozii Laveine(1967) are similar to N. terminiscus (e.g. Laveine,1967: pl. 25, figs 1 and 5), but the more typical pin-nules of Laveine’s species are squatter and morerounded. N. bourozii can also be distinguished by itsdenser veins (ª 50 veins per cm along pinnule margin)and mostly larger apical pinnules.

Neuropteris resobae Cleal (1981) from the upperDuckmantian or lower Bolsovian of Palencia, some-times has similar linguaeform pinnules with basalswellings, and has about the same venation density(e.g. Cleal, 1981: pl. 2, Fig. 2). However, the veins ofN. resobae are more oblique to the pinnule margin(60–70°). Also, the larger pinnules of N. resobae tendto be subfalcate to subtriangular, quite different toanything seen in N. terminiscus. The apical pinnulesof N. resobae are also significantly larger (15–30mmlong).

Remarks: This species is known only from strata associated with Seam 13 in Saarland, although from several different localities. Germer (1971) andDoubinger & Germer (1975) described examples asParipteris pseudogigantea, and their figured speci-mens have lateral pinnules that indeed resemble thatspecies (cf. Laveine, 1967: pls 73–76). However, one oftheir illustrations (Doubinger & Germer, 1975; pl. 9,Fig. 2) shows pinnules from a large specimen that isclearly imparipinnate (shown in the present paper onFig. 8). Two other specimens with clearly identicallateral pinnules also show imparipinnate pinna ter-minations (Figs 1 and 7). This character clearly pre-cludes an assignment of this material to Paripteris(Laveine et al. 1993).

This species has a midvein that is no more than two-thirds of the pinnule length, a feature that favours anassignment to Neuropteris. Pinnules of LaveineopterisCleal, Shute & Zodrow, 1990, in contrast, usually havea stronger midvein (two-thirds to three-quarters of the pinnule length). Its fronds are at least tripinnate

(Fig. 8) and so is unlikely to belong to Macroneu-ropteris, which has essentially bipinnate fronds (Clealet al. 1996).

NEUROPTERIS KNEUPERI CLEAL SP. NOV.1953 Neuropteris obliqua (Brongn.) Zeiller; Guthörl,

p. 166; pl. 22, fig. 1.1957 Neuropteris obliqua (Brongn.) Zeiller; Guthörl,

p. 51; pl. 14, fig. 2A,B.1975 Neuropteris obliqua (Brongn.) Zeiller; Doubinger

& Germer, p. 8; pl. 2, Figs 4,5 and 6; pl. 3, fig. 1 (nonpl. 3, Fig. 2 = Neuropteris ovata Hoffmann – seeCleal, 1985).

1975 Neuropteris heterophylla Brongn.; Doubinger &Germer, p. 11; pl. 4, figs 2,3.

Etymology: The species is named in honour of the late Professor Gottfried Kneuper, formerly of the Saarbrücken Mining School, in recognition of hismajor contribution to geological research in the Saarland Coalfield, and for his enthusiastic support ofpalaeobotanical research there.

Diagnosis: Ultimate pinnae tapered in distal part.Apical pinnules large (up to 20mm long), lanceolate,with sinuous lateral margins. Lateral pinnules5–20mm long, 3–7mm wide. Larger pinnules elon-gate, subfalcate to subtriangular; smaller pinnulesmore isodiametric, biconvex with rounded apex. Pinnules basally constricted in proximal part ofpinnae, partly fused to rachis towards pinna terminal.Strongly marked midvein extends for up to three-quarters of pinnule length. Lateral veins often flexu-ous and widely forking, and meet pinnule margin atª 90°; average marginal nervation density 35 veins per cm. Adaxial costal epidermal cells elongate-subrectangular, up to 200mm long and 25mm wide;intercostal cells slightly less elongate and regular, upto 100mm long and 30mm wide. Abaxial costal epider-mal cells elongate, subrectangular to polygonal, up to250mm long and 30mm wide; intercostal cells are moreor less isodiametrically polygonal, 25–30mm in size.Stomata cyclocytic.

Holotype: Specimen C/2830, Saarbrücken MiningSchool (Fig. 13). Grey silty shale in the Geisheck Formation (upper Bolsovian), Füllengarten Colliery,near Saarbrücken, Saarland, Germany.

Paratypes: In addition to the holotype, there are five large specimens showing bipinnate or tripinnatefrond fragments (C/2873, C/3420, C/3421, C/3912,C/5509). They all originated from the middle GeisheckFormation (upper Bolsovian), from the following localities: Franziska Colliery, Friedrichsthal Colliery,

198 C. J. CLEAL


Füllengarten Colliery, Burbach and the Hangard 2Borehole.

Description: Little is known of the form of the ante-penultimate pinnae. One large but poorly preservedspecimen (Fig. 20) shows the medial part of a frondand this has intercalated pinnae attached to an antepenultimate rachis. In this part of the frond, the penultimate pinnae are alternately attached to the antepenultimate rachis at about right-angles. Asecond specimen shows a more distal part of an antepenultimate pinna, with the penultimate rachisesattached at about 30° and one short intercalated pinna(Fig. 14).

The penultimate pinnae taper only slightly in theirproximal part (Figs 14 and 20), but taper moremarkedly towards the apex (e.g. Figure 19). Ultimatepinnae are attached alternately or suboppositely tothe penultimate rachis at 70–80°.

The last order pinnae are parallel-sided for most oftheir length (Fig. 13) but taper towards the pinnaapex. They are terminated by a single apical pinnule,up to 20mm long and 10mm wide (Figs 17 and 20).The larger apical pinnules are sublanceolate and oftenhave sinuous margins (Fig. 17). Much smaller, moreslender apical pinnules may also occur (Doubinger &Germer, 1975: pl. 2, fig. 5).

The rachises have irregular, longitudinal markings.Antepenultimate rachises are ª 7mm wide, penulti-mate rachises 1–2mm wide and ultimate rachis 1mmwide.

The smallest pinnules are round, ª 3–4mm in size,and tending to be broadly attached to the rachis(Figs 13 and 20). The slightly larger pinnules(5–15mm long) become more elongate with curvedlateral margins and a round to bluntly acuminate apex(Fig. 17). The larger lateral pinnules (15–20mm long,5–8mm wide) have a more distinctly subtriangularaspect and are inserted perpendicularly on the rachis(Fig. 15). In the proximal part of such pinnules, thelateral margins are fairly straight and parallel, orsometimes slightly curved, but in the distal parts ofthe pinnule the margins taper or curve inwards. Thistaper or curvature may be more pronounced on thebasiscopic side, giving the pinnules an asymmetricalaspect (Fig. 15). The pinnules have a bluntly acumi-nate or narrowly rounded apex. The base of thepinnule is normally slightly cordate and is narrowlyattached to the rachis. A basiscopic and sometimes anacroscopic basal lobe may be present.

A strongly marked midvein extends for two-thirds tothree-quarters of the length of the pinnules, beforedividing to form a fan of dichotomous veins (Fig. 15).The midvein shows little or no evidence of decurrenceat the base. Lateral veins are emitted at <20° from themidvein. The laterals fork three to four times. Near

the pinnule margin, these dichotomies are at a verynarrow angle, but near the midvein the angle becomeswider; this wide angle forking in the middle of thepinnule gives the venation a somewhat flexuousappearance. The laterals meet the pinnule margin atabout right angles, with a nervation density of 25–45veins per cm (average 35 veins per cm).

The adaxial pinnule surface is strongly cutinizedand has well developed intercellular flanges (Fig. 21).These show the adaxial epidermis is weakly differen-tiated between costal and intercostal fields. The costal cells are elongate and subrectangular, 150–200mm long and 20–25mm wide; intercostal cellsslightly less elongate and regular, up to 100mm longand 30mm wide. Neither trichomes nor stomata wereobserved.

The abaxial pinnule surface is less stronglycutinized (Figs 22–24). Intercellular flanges can beseen, but less clearly than on the adaxial cuticle. Thecostal cells are elongate, subrectangular to polygonal,up to 250mm long and 30mm wide (Fig. 22). In contrast, the intercostal cells are more or less isodia-metrically polygonal, 25–30mm in size (Fig. 23). Nodefinite evidence of trichomes was found, but numer-ous subcircular holes occur along the veins, ª 30mm indiameter, which might be trichome bases or hydath-odes (Fig. 24).

Stomata occur on intercostal fields of abaxial cuticle.They are not well enough preserved to establish thedetailed structure of the stomata, but they appear to have a single ring of subsidiary cells, although the subsidiaries are not significantly more thicklycutinized than the other epidermal cells (Fig. 23). Thestomata are aligned approximately parallel to theveins.

Comparisons: Guthörl (1953, 1957) and Doubinger &Germer (1975) identified specimens of this species asNeuropteris obliqua (Brongn.) Zeiller. The confusionappears to have arisen because, like N. obliqua, it hassomewhat flexuous veins and a combination of largersubtriangular pinnules and smaller more rounded pin-nules. However, the smaller pinnules in the N. obliquafrond tend to be more slender and more broadlyattached to the rachis than in N. kneuperi (comparewith Laveine, 1967: pl. 50, figs 1 and 4; pls 51–52).Also, the larger subtriangular pinnules in N. obliquaare mostly from the basal part of the frond, whereasin N. kneuperi they occur in more distal positions,mainly in the distal ends of penultimate pinnae. Theabaxial cuticles of N. obliqua are also different, beingstrongly papillate and with anomocytic stomata (Cleal & Shute, 1992). No unequivocal examples ofN. obliqua have been described from Saar-Lorraine.

Laveine (1967: 232) identified the specimen figuredby Guthörl (1957: Fig. 2) as Neuropteris ovata



200 C. J. CLEAL


13 14

15 16 17

18 19



Figures 13–19. Neuropteris kneuperi sp. nov. From the Geisheck Formation (upper Bolsovian), Saarland, Germany.Fig. 13. C/2830 (holotype); Füllengarten Colliery. Figs 14 and 18. C/3912; Franziska Colliery. Figs 15 and 19. C/5509; 0.80m Seam, Friedrichsthal Colliery, Friedrichsthal. Fig. 16. C/2873; Füllengarten Colliery. Fig. 17. C/4081; locality notrecorded. Scale bar = 5mm in all except Figs 15 and 18, where it is 1.7mm.

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Figure 20. Neuropteris kneuperi sp. nov., C/3420. From below Tonstein 2, Geisheck Formation (upper Bolsovian),Friedrichsthal Colliery, Friedrichsthal, Saarland, Germany. Scale bar = 5mm.

Hoffmann. However, this species does not have apicalpinnules with a sinuous margin as seen in Guthörl’sspecimen. Also, the latter is late Bolsovian in age andthus rather older than is normal for N. ovata.

Doubinger & Germer (1975) identified two otherspecimens of N. kneuperi as Neuropteris heterophyllaBrongn. However, the latter species has oval to lin-guaeform pinnules with a rounder apex, a less flexu-ous venation and anomocytic stomata surrounded bypapillae (Cleal & Shute, 1991: Figs 1–8).

The nearest comparison is with Neuropteris britan-nica (Gutbier), the larger pinnules of both speciestending to be subtriangular, with a flexuous venation,and in which the stomata are cyclocytic. However, theveining in N. britannica is significantly less dense(ª 20 veins per cm along the pinnule margin – seeDaber, 1955: pl. 23, fig. 3; pl. 24, figs 2 and 3; pl. 25;fig. 1; Daber, 1957: pl. 5). The pinnules of N. britan-nica also tend to be more broadly fused to the rachis,such that von Gutbier (1835) originally assigned the

202 C. J. CLEAL


2221

23 24

Figures 21–24. Neuropteris kneuperi sp. nov., cuticles from pinnules. From 0,80m Seam, Geisheck Formation (upperBolsovian), Friedrichsthal Colliery Saarland, Germany. Fig. 21. V.62969. Adaxial cuticle under phase contrast. Scalebar = 40mm. Fig. 22. V.62969. Abaxial cuticle under phase contrast. Scale bar = 40mm. Fig. 23. V.62969. Abaxial cuticleunder Nomarski Contrast. Scale bar = 20 mm. Fig. 24. V.62960. Abaxial cuticle under brightfield illumination. Scalebar = 80mm.

species to Odontopteris. The apical pinnules of N. kne-uperi, with their sinuous margins, are very differentfrom anything described for N. britannica (compare forinstance the terminals figured by Remy & Remy, 1959:fig. 149c). The adaxial epidermis of N. britannica hasmore uniformly distributed cells over most of thepinnule except on the midvein and the ‘strongest’lateral vein (Barthel, 1962). The adaxial cells are alsoless elongate.

Remarks: This species is so far unknown outside ofSaarland. Previously, the best documentation was byDoubinger & Germer (1975), who assigned specimensvariously to N. obliqua and N. heterophylla. As arguedabove, however, these identifications are incorrect andthe material belongs to a new species.

The generic position of N. kneuperi presents a fewdifficulties. The gross morphology tends to point to itbeing a Neuropteris, in particular that the frond is atleast tripinnately divided, that there are intercalatedpinnae on the antepenultimate pinnae, and that thevenation is somewhat flexuous. The differentiatedadaxial epidermis is also characteristic of Neuropteris.However, Neuropteris as emended by Cleal et al.(1990) includes species with usually anomocyticstomata, whereas those of N. kneuperi have a clearlydeveloped ring of subsidiary cells. This may mean thatthe circumscription of Neuropteris needs to beexpanded to accommodate species with such stomatalstructures. Alternatively, there may exist a distinctsubset of neuropteroid species with such cyclocyticstomata (perhaps also including Neuropteris britan-nica) that would merit taxonomic distinction.

DISCUSSION

The two new species described in this paper add to thelist of endemic taxa from the upland coal forests ofSaar-Lorraine. Furthermore, N. kneuperi accounts formost of the earlier records of two of the common neuropterid species found in the lowland coal forestsof northern Europe: N. obliqua and N. heterophylla.Two other pinna fragments were figured as N. obliquaby Guthörl (1948: pl. 16, Fig. 4, with a ‘cf.’) and de Jong (1974: pl. 25, fig. 1), but are too small to iden-tify. De Jong (1974: pl. 25, fig. 2) compared anotherspecimen with N. obliqua, but this is clearly identicalto Neuropteris triangularis P. Bertrand (1930), whichLaveine (1967) has transferred to N. ovata. Theabsence of N. obliqua from Saar-Lorraine should notreally be surprising, as N. obliqua only rarely extendsabove the middle Duckmantian (Laveine, 1967, 1986;Cleal & Shute, 1995), which is stratigraphically lowerthan the main Saar-Lorraine Basin succession.

The only other illustrated record of N. heterophyllafrom Saar-Lorraine is the holotype (Brongniart, 1822:

pl. 2, fig. 6). However, N. heterophylla is now conservedwith a different type (Brongniart, 1831: pl. 71) whichis not conspecific with the holotype. Brongniart’s 1822specimen in fact belongs to Laveineopteris tenuifolia(Sternb.) Cleal et al. (1990) (Laveine & Blanc, 1996),a species that is abundant in Saar-Lorraine (Bertrand,1930). In contrast, Neuropteris heterophylla, as repre-sented by the conserved neotype, appears to be absentfrom Saar-Lorraine.

In Table 1, the neuropteroid components of the Saar-Lorraine upland forests and of the lowland paralicforests are compared for the middle WestphalianEpoch (Duckmantian and Bolsovian Ages). This isbased partly on data extracted from Cleal & Shute(1995) and integrated with the new data given in thepresent paper. Unlike the Cleal & Shute (1995) study,the evidence from all of the paralic basins have been combined together, as there is no significant dif-ference between their floras. The Parispermaceae neuropteroids have been excluded as they appear tobe biologically distinct from the other medullosaleans(Laveine et al., 1993). The Table clearly reveals themarked differences in the neuropteroids betweenthese two areas, with only two species being commonto both areas. There are 11 species found only in theparalic coalfields and three species found only in theSaar-Lorraine. This cannot be explained by differencesin the extent to which the two areas have been inves-tigated; Saar-Lorraine has virtually as long a historyof research as the paralic basins. It must reflect realdifferences in the original vegetation and wouldappear to reinforce the view of Gothan (1951, 1954)that the upland vegetation of the Saar-Lorraine washighly distinctive.

The Table also reveals that both the true neu-ropterids and laveineopterids were significantly more diverse in the lowland paralic forests than theupland forests. This suggests that most of the plantsrepresented by these morphogenera were essentiallyadapted to the lowlands forests. In the few caseswhere species had evolved that were adapted to themore upland conditions, this adaptation seems to have prevented them also occupying the lowlands. The only known exception to this was L. tenuifolia,which occurred in both habitats. However, recent work on L. tenuifolia (Cleal & Shute, in press) sug-gests that there may in fact be at least two taxa (?‘varieties’) within the traditional concept of L. tenuifo-lia and one of these (typically found in Langsettian to middle Duckmantian floras) was restricted to theparalic habitats. Only the later L. tenuifolia-bearingplants (typically late Bolsovian to early WestphalianD) were able to adapt to both upland and lowland habitats.

Storch (1979, 1980) argued that the phytogeographi-cal differences between the various intramontane



and paralic basins of Europe have been significantlyexaggerated. For instance, he reported that CentralBohemia, Zwickau, and Saar-Lorraine had nearly two-thirds of their Sphenophyllum species in common.However, these are all upland intramontane basins,presumably representing broadly comparable habi-tats. We do not know whether or not they were origi-nally connected but this may not have been critical forthe migration of spore-bearing plants such as Spheno-phyllum. It would be expected that seed-plants suchas the Medullosales would have found such migrationmore difficult and the species would thus tend to haveless wide geographical distribution. This seems to be supported by the present study, although moredetailed taxonomic studies on the medullosaleansfrom the other intramontane basins will be neededbefore the full picture is revealed.

ACKNOWLEDGEMENTS

This work was funded by the Deutsche Forschungs-gemeinschaft, as part of a study for a PhD degree atthe University of Sheffield. The work was supervisedby Professor Bob Wagner (now of the Jardin Botánicode Córdoba, Spain), to whom I am extremely grateful.I would like to thank the authorities of the Saar-brücken Mining School for allowing access to the speci-mens described in this paper. During visits to thegeological collections of the mining school, then storedat von der Heydt, I was assisted by my redoubtable

friend, Herr Berndt-Arwed Richter, to whom I will beeternally grateful. Thanks go to Drs Deborah Spillardsand Imogen Poole for help with preparing the illus-trations. Finally, I must express my deep gratitude tothe late Professor Gottfried Kneuper, who initiated theproject and arranged the funding.

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204 C. J. CLEAL


Table 1. Distribution of Neuropteris, Laveineopteris and Macroneuropteris species in the paralic basin and Saar-Lorraineduring the Duckmantian and Bolsovian

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two new late carboniferous neuropteris species (medullosales) from saarland, germany and their...

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