Two LITTLE KNOWN SPECIES OF SPHENOPHYLLUM FROM THE STEPHANIAN OF SPAIN AND FRANCE

ROBERT H. WAGNER Geology Department, University of Sheffield, England, U.K.

ABSTRACT

The impression species Sphenophyllum incisum 'Wagner is fully described for the first time. with due reference to intra-specific variation as shown by abundant mate• rial from a single locality in the Stephanian B of North-west Spain. The associated stro• bilus is also recorded. Sphenophyllum nageli Grand'Eury, a middle to late Canta• brian species, is redescribed from the type region in the Cevennes, southern France. Similar material is illustrated from Cantabrian and Stephanian A localities in North-west Spain. Brief comments are made on several other species of Sphenophyllum from the Stephanian of western Europe, and a range chart is provided.

INTRODUCTION basis of a large collection of specimens from a single locality. Another little known species, Spheno• Doubingeris well overand twentyVetter (1954)years agopublishedsince phyllum nageli Grand'Eury, was actually ITa brief not~ on the various species of described in 1890, but hardly mentioned Sphenophyll1&m recorded from Stephanian afterwards. Doubinger and Vetter ignored strata. A monographic paper by Abbott it entirely in their synthesis of 1954, and (1958) filled in gaps with regard to the the species did not reappear in the literature American species; a brief note by Obrhel until 1970 when it Was mentioned again (1967) brought the description of Spheno• from the original type area, the Cevennes, phyllum nemejci Obrhel from the Stephanian by Bouroz, Gras and Wagner. The material of Bohemia; and recent work on the reported in that paper includes several Stephanian floras of Portugal and Spain well-preserved specimens which allow a has added several new species such as redescription of Sphenophyllum nageli Grand• Sphenophyllum guerreiroi Teixeira, Spheno• 'Eury, as provided in the present paper. phyllum incisum Wagner, Sphenophyllum Although, it would undoubtedly be of galeanoi Broutin, Sphenophyllwm casetas• interest to present a full synthesis of ense Knight, and Sphenophyllum crenulat1,em Stephanian species of Sphenophyllum, as Knight. Even the well studied Stephanian known at present from West European of France yielded a new species, Spheno• localities, this must be regarded as outside phyllum miravallis Vetter. The stratigraphic the scope of the present paper. Only a few distribution of the various Stephanian species comments may be made on some of the of Sphenophyllum has also become better more important species, stressing a few known. Vogellehner (1967) introduced a points of taxonomic interest and highlighting new species, Sphenophyllum intermedium, some of the more recently acquired informa• for material resembling S. emarginatum. in tion on stratigraphic ranges. Stephanian A-B strata. Most of these new species have been SYSTEMATIC DESCRIPTION adequately figured and described (Teixeira, 1951; Vetter, 1968; Broutin, 1973; Knight, Sphenophyllum incisum Wagner 1975), but Sphenophyllum incisum has only PIs. 1-4, figs. 1-12; PI. 6, fig. 20 been diagnosed briefly on the basis of a single holotype (Wagner, 1964), and cannot 1964 Sphenophyllum incisum n. sp., Wag• yet be regarded as adequately known, ner: C.R. 5e Congres Carbonijere, despite further illustration of this species by Paris 1963, II, p. 846, pI. III, fig. 24. Wagner (1965) and Stockmans and Williere 1964 Sphenophyllum incisum Wagner, Bou• (1965). This will be remedied in the present reau: Traiti de Paleobotanique, tome paper by providing a full description on the III. Sphenophyta, p. 66, fig. 41 5~i1 566 TIlE PALAEOBOTANIST

(same as Wagner, 1965, pI. 51, figs. 104, 105). TABLE46353·5Breadth47564·55·53Length45'53·53'522·553Central1 - CentralMEASUREMENTScleft ON 59 98 1965 7·5897'59 cleft LEAVESOFATSPHENOPHYLLUMLOC.FROM983DIFFERENTIN THE INCISUMLA SPECIMENSMAGDALENAWAGNER SphenophylZ,um11 incisum Wagner: "Vag•COALFIELD (PROVINCE LEON, NW SPAIN) ner, Mem. Inst. geol. Min. Espana, Length Breadth LXVI (pubI. March, 1966), pp. 52, 142, pI. 51, figs. 104, 105. 1965 5phenophyUum incisum Wagner, Stockmans & Williere: M em. Inst. R. Sci. nat. Belgique, 79 (pubI. April, 1966), pI. XXXII, figs. 6-9. 1970 SphenopJ~yllum incisum Wagner: Wag• ner & Artieda, La Cuenca Minera Cinera-Matallana, lam. XVII, fig. D (holotype. x 3). 1971 Sphenophyllum 1:ncisum Wagner: Wag• ner, Trab. Geol. Fac. Ci. Univ. Oviedo, 4, p. 42l. 1975 Sphenophyllum incisum Wagner: Al• varez-Ramis, Estudios Geol., XXXI, p. 786, lam. II, fig. 3 (after Wagner, 1964, pI. III, fig. 24 - holotype), non fig. 2 (probably Sphenophyllum angustifolium Germar). Description - Ribbed axes of variable width with closely spaced, overlapping verticils of six leaves showing straight sides and a deeply dissected distal border with elongate, pointed teeth, 4 or 6 in number. Length-breadth ratio of the leaves varying between 1'3 and 3 (generally about 2) and central cleft reaching most commonly a little over halfway down the leaf (compare Table 1 & Text-fig. 1). Leaf sizes between 4-11 mm length and 2-6 mm width (as measured on the distal border). Nervation corresponds closely to the segments of the of the La Magdalena Coalfield in the province deeply incised leaf. Leon, NW Spain. Material from this locality Branches occur as one per node at (Wagner loco 983) was figured previously by intervals of several internodes. Leaf verticils Wagner (1965) and Stockmans and Williere very densely arranged at the end of branches (1965). Although other plant species are where each verticil may overlap with several associated, 5phenophyllum incisum is the others. Lengthwise compression often gives dominant sphenophyte in the bed which was a false appearance of simple leaves. sampled extensively over a number of years. Fructification (associated, not in organic Associated sphenophytes are Annularia connection) consists of a strobilus with sphenophylloides (Zenker) von Gutbier and verticils of curved, rather spiky, apparently Asterophyllites equisetiformis (von Schlot• simple bracts carrying relatively large heim) Brongniart, a point worth remember• sporangia in the axils, probably in the ing when assigning some associated strobili proportion of one sporangium per bract. to Sphenophyllum incisum Wagner. Only Sporangiophores not observed. one of these strobili shows a ribbed axis Remarks on Figured M aterial- The above similar to those of Sphenophyllum incisum description is based on some eighty samples (not an infallible criterion anyway), the collected from a single locality in 10 to other strobilar axes being too thin for 20 em of rock at some distance above a ribbing to oCCur. A comparison with pub• thin coal, which is the first to occur above lished strobili of Annularia sphenophylloides the basal breccia on the northern edge (i.e. Calamostachys ealathifera Weiss) and WAGNItR- TWO titTLE KNOW SPECiES OF SPHENOPHYLLUM 567

Asterophyllites equisetiformis (i.e. Calamo• good preservation of these compressions and stachys germanica Weiss) show strongly impressions have allowed the fairly accurate divergent characters, particularly in view of measurement of length, breadth and relative the axillar position of the sporangia in the amount of incision at the central cleft of strobili associated with SphenophyUum in• each leaf. Altogether, 59 leaves were mea• cisum. These appear to conform to the sured for Table 1 (see also Text-fig. 1), Koinostachys type of strobilus in the Spheno• each leaf being taken from a different phyllales, a rather wide grouping of fructi• specimen and using completeness of preserva• fications (Remy, 1955) which is based mainly tion as the sole criterion for the selection of on the axillar position of sporangiophores specimens to be measured. The length• carrying either single or clustered sporangia breadth ratio falls within reasonably narrow (as a result of successive dichotomies of the limits and the majority of specimens occupy sporangiophore) and on the absence of a a virtually straight line on the graph shield in the peltate arrangement of sporan• plotting length against breadth. The depth gia. Both strobili and individual whorls of the central cleft is only a little more with axillar, branching sporangiophores (as variable (right hand graph of Text-fig. 1), in Sphenophyllum majus Brong & Spheno• in relation to the length of the leaf. phyllum k£dstoni Hemingway) have been The apparently rather minor variation in assigned to Koinostachys by Remy. length-breadth ratio and in the relative In the sterile foliage of Sphenophyllum depth of incision at the central cleft is incisum Wagner it is noticeable that the matched by the other characteristics such amount of variation is strictly limited. as the straight lateral sides, with only The large number of specimens collected occasional flaring near the distal border from a single locality, and the reasonably (probably as a result of the deep incisions

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o 2 3 456 7 o 2 3 456 7 breodth(mm) central cleft (mm)

TEXT-FIG. 1,- Graphs showin~.length of leaves of Sphenophyllum incisum plotted against breadth, and length against depth of mClSlOn at central cleft (actual number of specimens recorded with' j.'elative size of dot denoting position of larger or smaller number of specimens with certain ratios). 568 THE: PALAEOBoTA 1ST and at least partly attributable to preserva• number of teeth on the distal border. tion), and the invariable presence of deeply Sphenophyllum kidstoni is also characterized dissected leaves with long, pointed slips. by groups of 4- sporangia per sporangiophore There is none of the extreme polymorphism in the axils of apparently normal leaves, which characterizes some of the other species and does not possess strobili as such. This of Sphenophyll1~m, e.g. Sphenophyllum cunei• species is a middle Westphalian element, folium Sternberg, and more entire leaves are whereas Sphenophyllwn incisurn occurs in conspicuously absent in the assemblage. Stephanian B. An important characteristic of Spheno• There may be a vague resemblance to the phyllum incisum seems to be the close forma saxifragaefolium of the extremely poly• spacing of leaf verticils, a feature which morph Sphenophyllum cuneifolium (Stern• becomes even more pronounced near the tip berg) Zeiller, since this form also possesses of the ulhmate branches. a highly dissected leaf which may also have Compression is either lengthwise or as four sharp teeth on the distal border. How• spread-out verticils, with somewhat oblique ever, Sphenophyllum incisum shows highly compression being a not uucommon feature. dissected leaves throughout, whereas Spheno• Longitudinal compressions may not show phyllum c1~neifoliumshows a number of leaf the entire length of leaves and commonly shapes, the most common of which is an produce the somewhat misleading aspect of almost entire leaf with short, pointed teeth simple leaves, whereas these are not, in fact, on a distal border without central cleft. present. The narrowness of the leaves may The stratigraphic ranges of these two species have helped in producing this kind of are very different, since Sphenophyllum preservational aspect. Only two successive cuneifolium occurs in middle to upper dichotomies are present in the case of four Namurian and lower to middle Westphalian teeth being present on the distal border. strata, and Sphenophyllum incisum in An additional dichotomy of inner slips of Stephanian B. each pair in a foursome produced the six Remarhs on Synonymy - Sphenophyllum teeth that are very often present in the incisum was briefly diagnosed in Wagner, leaves of Sphenophyllum incisum. Also with 1964-, p. 84-6 (Compte rendu 5th Carboni• regard to the presence of either four or ferous Congress,Paris, 1963): " Leaf verticils six teeth and the concomitant rather narrow characterized by rather slender, deeply width of the distal border, the species shows incised leaves with almost straight lateral relatively constant characteristics. . sides and long, pointed teeth on the distal Comparison - Sphenophyllum angustt- side. The leaves are usually about 7 to folium Germar, a characteristic element ?f 10 mm long and about 3 to 4- mm wide Stephanian C floras and thus rather close m on the distal border." Comparison was stratigraphic occurrence to ~phenophylZ,~m made with Sphenophyllum cuneifolium forma incisum which has been found m Stephaman saxifragaefolium Germar. Only a single B, has narrower and longer leaves with a specimen (holotype) was figured at the time. length-breadth ratio of about 5. Its leaves It was refigured. X 3, in Wagner and are also not quite as deeply cleft as in Artieda (1970) as part of a documentation Sphenophyllum incisum (i.e. in relation to ~he of the Stephanian B flora of the Cinera• length of the leaves). The rat~er. maSSIve Matallana Coalfield, but a truly representative strobili of Sphenophylltmt angustifoltum pro• collection of several hundred specimens from vide another distinctive feature of this a single locality was obtained in later species, those of Sphenophyllum incisum years from the basal coal-measures of the being more slender and, above all, more neighbouring La Magdalena area, further laxly built. west in northern Leon (NW Spain). This The deeply incised distal border of Spheno• locality (983) yielded the specimen figured phyllum incisum oc.casionally app~ars to be in the 'Traiti de Paleobotanique' and in flaring, thus showmg a superfiCial resem• Wagner, 1965, as well as those illutsrated by blance to Sphenophyllum kidstoni Heming• Stockmans and Williere, 1965. All the way. However, the leaves of Sphenophyllt,m specimens recorded in the present paper are kidstoni have a stiffer appearance and also from this locality at La Magdalena. They show either a larger number of leaves show the full range of variation as described per verticil (i.e. more than 6) or a larger above. WAGNER - TWO LITTLE KNOWN SPEcIES OF . PHENOPHYLLUM 569

On the basis of the few specimens illus• trated in the literature, Alvarez-Ramis (1975) TABLE 2-DIMENSIONS OF LEAVES IN has suggested that Sphenophyllum incisum INDIVIDUAL SPECIMENS OF SPHENOPHYLLUM NAGELI GRAND'EURY might represent the basal portions of Spheno• FROM THREE LOCALITIES IN ZONE 2 phyllu.m angustijolium Germar. This view (PANISSIERE) IN THE CEVENNES finds no support from the extensive collec• COALFIELD OF SOUTHERN FRANCE tion of remains of Sphenophyllum incisum Length Breadth52·532 reported in the present paper, which does 87·5765 not show a sufficiently wide range of varia• 9 tion for the incorporation of this species in Sphenophyllum angustijoliuJn. The specimen from the Tineo Coalfield in western Asturias (North-west Spain) which Alvarez-Ramis (1975, lam. II, fig. 2) figures under the name Sphenophyllum incisum, probably does be• long to Sphenophyllum angustijolium which is a fairly common species at Tineo (compare even in the limited number of specimens Wagner, 1965, pp. 142-144, pI. 70). collected (Table 2); length ranging from Remarl.s on Preservation - The holotype 5 to 9 mm, width from 2 to 5 mm; length• of Sphenophyllum incisum Wagner is a breadth ratio ranging from 1·6 to 3; lateral black imprint and stain on a dark grey borders straight and distal borders convex, mudstone from the Pastora Formation of with short rounded teeth which may not the Ciiiera-Matallana Coalfield. The rank of always be apparent where preservation is coals in this coalfield (12-14% volatiles) slightly defective; nervation fairly close and seems to preclude the recovery of a cuticle. consisting of a single initial vein splitting The specimens from the La Magdalena rapidly and dichotomising into numerous Coalfield, recorded in the present paper, are ultimate veins each of which corresponds to preserved as dark carbonaceous stains on a rounded tooth on the distal border. rust-coloured imprints on slightly silty mud• Remar1Permian The latter (PI. 5, fig. 16; PI. 6, figs. 17-18) speCles. also possess six leaves per whorl and they In fact, the general shape of the leaves show the convex distal border with short, in Sphenophyll161n nageli Grand'Eury is quite rounded teeth that helps to characterize similar to that of Sphenophyl um thoni var. this species. However, their lea,-es are minor Sterzel, a late Stephanian B form marginally more truncate than those of with entire leaves (as against the fringed Sphenophyllum nageli from the Ceve!1I1es. leaves of Sphenophyllum thoni proper). In fact, they may be regarded as mter• There is, however, a fundamental difference mediate forms between this species and in that some of the veins end on the lateral Sphenophyllum emarginatum forJ?a tr1,f,ncatu:m borders in 5phenophylh611t thoni var. minor, Schimper. This is the form With an entll'e whereas they all terminate on the distal WAGNER- TWO LITTLE I(NOWN SPECIES OF SPHENOPHYLLUM 571

border in Sphenophyllum nageli. The leaves near Mercedes in the Barruelo Coalfield also tend to be a little larger in Spheno• (provirlce Palencia), Barruelo Formation, phyllum thoni var. minor, but it must be Calero Member, early Stephanian A; Loc. noted that the size variation in this species 2208 - tip of San Francisco Mine (Sanfesa overlaps with that of Sphenophyllum nageli. Company), south of San Salvador de Cant• The small crenulations that appear on the amuda (province Palencia), Salvador Forma• distal border of Sphenophyllum nageli are tion, middle Cantabrian. apparently absent in Sphenophyllum tlwni var. minor. REMARKS ON SOME OTHER SPECIES OF Comparison can be made also with 5pheno• SPHENOPHYLLUM FROM THE phyllwn crenulatum Knight which has entire STEPHANIAN leaves with rounded tops showing bluntly acute teeth becoming slender and awl-like An alphabetical order is followed for some in some cases. This is a species of the brief remarks on several other species group of Sphenophyllum thoni and thus of Sphenophyzz.um from the Stephanian of shows the presence of vein endings on both western Europe, and the same order is the lateral and distal borders which are fully followed for the accompanying range chart merged into one another. Also, Knight's (Table 3). species seems to show generally larger Sphenophyllum alatifolium Renault, a leaves than occur in Sphenophyllum nageli species with rather large and broad leaves Grand'Eury. with straight lateral borders and a truncate Sphenophyllum verticillatum (von Schlot• distal margin set with short, rounded teeth, heim) Zeiller is very clo~ely similar to has often been confused with Sphenophyllum Sphenophyllum nageli. It also shows all the costae Sterzel which may be distinguished veins ending on the distal margin which is by its somewhat convex distal border. convex and set with small, rounded teeth. These two species are possibly synonymous The size and length-breadth ratios of the (as was proposed by Doubinger & Vetter, leaves are also comparable and the two 1954), and in this case Sphenophyllum species could well be confused if Spheno• alatifolium has priority (see comments in phyllum verticillatmn had not shown the Wagner, 1965, pp. 140-142). extreme dimorphism recorded for this species There are no problems attached to the which possesses deeply incised leaves on the identification of Sphenophyllum angustifolium main axes. This dimorphism is unknown Germar which is easily recognized on its for Sphenophyllum nageli, even though it rather slender leaves with 2 to 4 elongate, should be noted that the number of speci• pointed teeth. This species is a reliable mens is still rather small and its full index for Stephanian C and Autunian. An range of variation possibly unknowll as American species, Sphenophyllum tenuifolium yet. Fontaine & White, which shows similar Distribution - Undoubted specimens of characteristics, has sometimes been recorded Sphenophyllum nageli Grand'Eury are known mistakenly from Europe after specimens only from the type region, i.e. the Cevennes belonging to Sphenophyllum angustifolium in southern France. Recent collection (Bou• Germar. roz, Gras & Wagner, 1970) has shown its One of the two species recently introduced presence in Zone 2 (Panissiere), of middle by Knight (1975), Sphenophyllum caseta• to late Cantabrian age: Loc. 2015 - old sense, has somewhat flaring, dissected leaves tips at Le Maze!, seams III, IV and V of with a sharply toothed distal border, a Zone 2 in its lower part, near the Bannelle length-breadth ratio of 1·25 to 2 (ranging Fault; Loc. 2016 - tip of Le Souterrain, in length from 4 to 10 mm) and only a Ruisseau de la Combe de Banne, Zone 2 certain amount of dimorphism. Comparison in its middle part. has been made primarily with Sphenophyl• Specimens comparable to Sphenophyllum lum kidstoni Hemingway, which has similar nageli and apparently transitional to Spheno• leaf types, but which shows tetrasporangiate phyllum e11larginatum forma truncatzlnz clusters amongst whorls of ordinary leaves, Schimper occur in middle Cantabrian and whereas Sphenophyllum casetasense bore small Stephanian A strata of north-west Spain: strobili on ultimate branches. The latter Loc. 1784 --- old tip on the path to Petrita, has been described from Stephanian B 572 THE PALAEOBOTAN1ST

TABLE 3-STRATIGRAPHIC RANGES OF THE DIFFERENT SPECIES OF SPHENO• PHYLLUM IN STEPHANIAN ROCKS OF FRANCE AND SPAIN (STEPHANIAN SERIES CONSISTING OF FOUR STAGES CANTABRIAN, STEPHANIAN A, STEPHANIAN B, STEPHANIAN C). ST. .,.., • emarginatumA truncatumST.AUTUN.-BST.formaC --. CANTAB. _? ,.• - - S. miravallis Vetter S. nageliverticillatumS. S.crenulatumGrand'EurycostaeS.S. thoni(vanSterzelKnightSchlotheim)van Mahr Schimper I - S. casetasense KnightS. angustifoliumS. longifolium GermarGermar S. incisum Wagner - S. thoni var. minor Sterzel - S. oblongifolium (Germar & Kaulfuss) S. galeanoiS. guerreiroi-Broutin-i Teixeira -- I S. alatifolium Renault

strata in the Sabero Coalfield of north-west The very common species Sphenophyllum Spain, whereas Sphenophyllum kidstoni emarginatum (Brongniart) Koenig is mainly occurs in Westphalian B. recorded from Westphalian C and D, but it Sphenophyllum crenulatum Knight is an• persists with large numbers into Cantabrian other Stephanian B element from the Sabero strata and still occurs sporadically in Coalfield, but it also occurs in the late Stephanian A (Knight, 1975). From late Stephanian B of Saarland (Germany). It Westphalian D onwards it is the forma was illustrated from Saarland by Germar, truncatum Schimper which is present, the Kneuper and Wagner (1968) under the typical form with a central cleft being name Sphenophyllum novo sp. aff. thoni von restricted apparently to Westphalian C and Mahr, and by Germar (1971) as Spheno• early to middle Westphalian D. phyllum miravallis Vetter. Sphenophyllum Sphenophyllum guerreiroi Teixeira is a crenulatum is a species in the group of form with small leaves (up to 4·5 mm long) Sphenophyllum thoni with which it agrees which shows a strongly developed median in showing leaves with the distal border cleft, somewhat convex lateral borders and merging gradually into the lateral borders, four pointed teeth on the distal border and the veins ending partly on the lateral (Teixeira, 1951, pp. 14-15). It was described margins. Sphenophyllum crenulatum is fur• originally from the late Westphalian D of ther characterized by its bluntly acuminate Portugal but has been found since in upper teeth, normally with only weak incisions Westphalian D and middle Cantabrian rocks in between but, particularly on the top• of Palencia, north-west Spain. lateral margin, the teeth may be slender, Sphenophyllum longifolium Germar is a awl-like and incised up to 2·5 mm. In this very characteristic species with fairly large, respect it differs markedly from Spheno• relatively long and dissected leaves. It phyllum miravallis which possesses leaves occurs in Stephanian Band C as well as with an entire margin. The size and length• in the lower Autunian. Storch (1966) has breadth ratios of the leaves are quite similar recertly figured Sphenophyllum longifolium in both species, but it is also to be noted from Westphalian D strata in Saxony, that Sphenophyllum miravallis Vetter shows thus showing its range to be appreciably anisophyllous whorls. longer. It is generally of rare occurrer~ce The two species described by Knight and its early appearance in the Zwickau• (1975) are in an unpublished Ph.D. thesis Lugau-Oelsnitz Coalfield of Saxony may which is currently in the press. well be linked to considerations of habitat. WAGN~R- TWO LITTLE KXOWN SPECIES OF SPHENOPHYLLUM 573

Sphenophyllum miravallis Vetter, a rare species representing the fi"st of a group of species of the Sphenophyllum thoni group, is mainly Permian Sphenophyllu:n with merging characterized by the veins ending Oil both lateral and distal borders. It first appears the lateral and distal borders which merge in upper Stephania'1 B with the variety very gradually. The size of the leaves is minor Sterzel which is characterized by fairly large (up to 20 mm long), with a somewhat smaller leaves with an entire length-breadth ratio of up to 2·5. They margin. In the Stephanian C it is joined possess rounded, entire margins, and the by the typical form of SphenophylluYn thoni general aspect of this species is not too which has generally larger leaves with a dissimilar to Sphenophyllum speciosum Royle fringed border. of the Permian of Gondwanaland. The only Sphenophyllum verticillatu:n (van Schlot• reliable records of this species are from the heim) Zeiller has smaller leaves with only an Massif Central of France, where Vetter (1968) apparent merging of lateral and distal records its occurrence from Stephanian B borders which possess very short, rounded to Autunian. teeth. This species shows a high degree of Sphenophyllum oblongijoliu1n (Germar & polymorphism ranging from the typical form Kaulfuss) Unger is undisputably the most with an entire margin to highly dissected common species ofSphenophyllum in Stephan• leaves. It occurs from Stephanian B on• ian strata. It is easily recognized by its wards and ranges into the Autunian. convex lateral borders, central cleft and pointed teeth on the distal border. Aniso• ACKNOWLEDGEMENTS phylly is often a characteristic feature of this species which ranges from middle-late The writer is grateful to Drs A. C. Higgins Cantabrian to Autunian. and J. A. Knight for constructive criticism Broutin (1973) has described a similar helpful to the final version of the manus• species, Sphenophyllum galeanoi, from the cript, and to Dr R. Coquel for the provision Autunian rocks of Guadalcanal (province of a xerox copy of Grand'Eury's (1890) of Sevilla, southern Spain). It differs from description and illustration of Sphenophyllum Sphenophyllum oblongijolium mainly on the nageli. He also acknowledges the efficient basis of cuticle characteristics. and skilful guidance of Drs A. Bouroz and Sphenophylluln thoni van Mahr is a strati• H. Gras during a collecting trip in the graphically important, well-characterized Cevennes Coalfield in 1969.

REFERENCES

ABBOTT, M. L. (1958). The American species of Surv. Czechoslovakia, 18, Pal.,: 297-307, Asterophyllites, Annularia, and Sphenophyllum. pIs. I-IV. Bull. Amer. Paleont., 38 (174): 285-372, pIs. DOUBINGER, J. & VETTER, P. (1954). Sur les 35-49. Sphenophyllum du Stephanien. Contribution Ii ALv'AREZ-RAMIS,C. (1975). Consideraciones sobre la revision du genre Sphmophyllum. Ann. semejanzas y sinonimias de las para-especies de Soc. geol. Nord, 74: 106-121- Sphenophyllum del Paleozoico espano!. Estu• GERMAR,R. (1971). Leitfossilien in der Schichten• dios Geol., 31: 785-793, lams I-III. folge des Saarkarbons. Beiheft Geol. Landes• BOUREAU, E. (1964). Traite de Paleobotanique. aufnahme des Saarlandes, 3: 1-124, Textabb. III. Sphenophyta, Noeggerathiophyta. Masson 1-89, Tafn I-XVIII. et Cie, Paris: 1-544, figs. 1-436. GERMAR, R., KNEUPER, G. & WAGNER, R. H. BOUROZ, A., GRAS, H. & WAGNER, R. H. (1970). (1968). Zur vVestfal/Stefan-Grenze und zur A propos de la limite Westphalien-Stephanien Frage der asturischen Faltungsphase im Saar• et du stephanien inferieur. In: Colloque sur brucker Hauptsatte!. Geologica et Palaeonto• la stratigraphie du Carbonifere, Congr. Co!l. logica, 2: 59-71, Tafn 1-2. Univ. Liege, 55: 205-225. GRAND'EuRY, F. C. (1890). Geologie et Paleonto• BROUTIN, J. (1973). Le Bassin Autuno-Stepha• logie du Bassin Houiller du Gard. Theolier et nien de Guadalcanal (Espagne du Sud). Don• Cie, St. Etienne: 1-354, pIs. I-XXII. nees paleobotanique et aspects stratigraphiques JONGMANS,W. J. (1911). Anleitung zur Bestim• et paleogeographiques. These 3e Cycle, Univ. mung der Karbonpflanzen West-Europas, mit Paris VI: 1-134, pIs. I-XVII. besonderer Berucksichtigung der in den Nieder• DAKov'A-PoKoRNA', S. (1951). Two new Spheno• landen und den benachbarten Ui.ndern gefun• phylla from the Lower Gray beds of the Coal denen oder noch zu erwartenden Arten. 1. basins of Central Bohemia. Sbor1'Zi'k Geol. Thallophyta, Equisetales, Sphenophyllales. 5'74 TilE PAtAEOBOTA1\IST

M eded. Rijksopsporing van Delfsloffen, 3: 1• Houill. Bassin d'Aquitaine, Albi, 1-194, pIs. 1• 482, figs. 1-390. XLIX. KNIGHT, J. A. (1975). The systematics and strati• VOGELLEHNERD. (1967). Die Flora de, Stefans graphic aspects of the Stephanian flora of the an der Hohengeroldseck bei Lahr (Mittlerer Sabero Coalfield. Ph.D. Thesis, Univ. Sheffield: Schwarzwald). 1. Sphcnophyta (Sphenophylla• 1-197 (pt. I), 1-676, pIs. 1-52 (pt. II). les und Equisetales). Ber. Naturf. Ges. Freibttrg OBRHEL, J. (1967). Sphenaphyllum n~mejci n. sp. i. Br., 57: 133-168, Abb 1-15. - neue Art aus dem b6hmischen Oberkarbon. \VAGNER,R. H. (1964). Stephanian Floras in ~W Spain, with special reference to the \Vestphalian V~stnik Ost~. Oslo Geol., 42 (2): 119-121, Tafn D - Stephanian A boundary. C.R. 5e Congres I-II. Carbonifere, Paris 1963, II: 835-851, pIs. I-III. REMY, W. (1955). Untersuchungen von kohlin WAGNER, R. H. (1965). Palaeobotanical dating erhaltenen fertilen und sterilen Sphenophylleg of Upper Carboniferous Folding Phases in NW und Formen unsicherer systematischer Stellung. Spain. Mem. Inst. Geol. Min. Espana, 66: Abh. dt. Akad. Wiss. Berlin, ](l. Chemie, Geologie 1-169, pIs. 1-77. ttnd Biologie, Jg. 1955, Nr. 1: 1-40, Tafn. 1-13. WAGNER, R. H. (1971). The stratigraphy and REMY, \V. & REMY, R. (1959). Pflanzenfossilien. structure of the Ciiiera-Matallana Coalfield (prov. Akademie-Verlag Berlin: 1-285, Abb. 1-209. Le6n, N.\V. Spain). In : The Carboniferous REMY, W. & REMY, R. (1966). Atlas wichtiger of North-west Spain" pt. II. Trabajos de stratigraphischer Leit- und Charakterarten im Geologia, Fac. Ci. Univ. Oviedo, 4: 385-429, eu ramer ischen Floren bereich. A rgumenta Palaeo• figs. 1-24. botanica, 1: 55-86, Tafn 1-18. WAGNER, R. H. & ARTIEDA, J. I. (l970). La STOCKMANS,F. & WILLrimE, Y. (1965). Documents Cuenca Minera Ciiiera-Matallana. S. A. Hullera paleobotaniques pour l'etude du Houiller dans Vasco-Leonesa, Leon, : 1-288, lams I-XIX. Ie Nord-Ouest de I'Espagne. Mem. Insl. R. WAGNER, R. H., PARK, R. K., \VINKLER PRINS, Sci. nat. Belgique, 79, pIs. I-XXXVIII. C. F. & Lvs, M. (1977). The Post-Leoni an STORCH, D. (1966). Die Arten der Gattung Basin in Palencia: A report on the Stratotype Sphenophyllum Brongniart im Zwickau-Lugau• of the Cantabrian Stage. In: Symposittm on Oelsnitzer Steinkohlenrevier. Ein Beitrag zur Carboniferous Stratigraphy. Spec. Publ. Geol. Revision der Gattung. Palaeont. Abh., (B), 2 Surv. Prague: 89-146. (2): 193-326, Tafn I-XXVII. WAGNER, R. H. & VARKER, \Y. J. (1971). The dis• TEIXEIRA, C. (1951). Plantas f6sseis do Permo• tribution and development of post-Leonian Carbonico portugues. II. Comun. Servo Geol. strata (upper Westphalian D, Cantabrian, Ste• Portugal, 32: 1-17, Est. I-XXII. phanian A) in northern Palencia, Spain. In: VETTER, P. (1968). Geologie et paleontologie des The Carboniferous of North-west Spain, pt. I L bassins houillers de Decazeville, de Figeac et Trabajos de Geologia, Fac. Ci. Univ. Oviedo, 4: du Detroit de Rodez. II. Etude paleontologique. 533-601, pIs. 1-2.

EX PLANA TION OF PLATES

(All figures natural size unless indicated othe'rwise) PLATE 1 PLATE 3

(Sphenophyllum incisum Wagner) (Sphenophyllum incisum \Vagner) 1, 2. Longitudinal compression of leaf whorls 5a, 8a. Spread-out, deeply incised leaves. x 3. showing the standard kind of deeply incised leaf 9. Spread-out verticil of very deeplv incised (figs. la & 2a 3). . . . leaves together with ribbed axis and lateral branch 3. Longitudinal compressIOn showmg partIal cross in longitudinal compression (fig. 9a. X 3). sections of leaves in closely spaced whorls on lateral lOa. Spread-out leaf verticils. X 3 (natural size branch (fig. 3a. X 3). on PI. 4).

PLATE 2 PLATE 4

(Sphenophyllum incisum Wagner) (Sphenophyllum incisum \Vagner) 4. Part of a strobilus with spiky bracts and axil• 10. Spread-out leaf verticils. iar sporangia (fig. 4a. X 3). 11. Strobilus together with axes showing ordinary 5, 6. Spread-out leaf verticils showing the long leaf verticils. (fig. 11a, strobilus three times en• pointed teeth on deeply incised leaves (figs. 5a. larged in order to show the alternate arrangement 6a. X 3). of bracts with axillar sporangia). 7. Longitudinal compression of closely spaced 12. Longitudinal compression of axes showing leaf verticils showing mainly cross sections of leaves cross sections of leaves (on the left a fragment of a which thus appear to be simple or only once bifurcate strobilus), (fig. 12a. X 3). (fig. 7a. X 3). 8. Spread-out leaves showing central cleft and N.B. All specimens on PIs. 1-4 from loco 983 in long, pointed teeth. the La Magdalena Coalfield, Stephanian B. PLATE 5 PLATE 6 (5phenophyllmn nageli Grand'Eury) (figs. 13-15) and Sphenophyllum d. nageli Grand'Eury (figs. 16-18). S. d. nageli (fig. 16) S. nageli (fig. 19), and Sphenophyllum incisum Wagner (fig. 20) 13. Spread-out leaf verticils showing the charac• teristic shape of lea\'es with rounded tops (fig. 13a, 16a. Semi-longitudinal compression of verticils three times enlarged in order to show the short, showing leaves with convex but almost truncate rounded teeth on the distal border). distal margins. X 3 (natural size figured on PI. 5). 14. Semi-longitudinal compression of axes with 17, 18. Spread-out leaf verticils showing rounded slender leaves showing the characteristic rounded tops set with short, blunt teeth (figs. 17a & tops (fig. 14a, three times enlarged to show venation). 18a. X 3). This specimen is similar to that figured by Grand' 19. Spread-out leaf verticils with rather narrow Eury (1890) as the type of Sphenophyllum nageli. leaves with characteristically rounded tops (fig. 15. Exceptionally small leaves in spread-out 19a. x 3). verticils (fig. 15a. X 3). 20. Longitudinal compression showing cross sec• 16. Semi-longitudinal compression of inter• tions of leaves. X 3. mediate form between S. nageli and S. emarginatum forma tntttcatum Schimper, showing somewhat less N.B. Fig. 16a from loco 2208 in northern Palen• rounded tops to the leaves. cia, middle Cantabrian; figs. 17-18 from loco 1784 in northern Palencia, lower Stephanian A; fig. 19 N.B. Figs. 13-14 from loco 2015 in Cevennes Coal• from loc. 2015 in Cevennes Coalfield, middle to field, middle to upper Cantabrian; fig. 15 from loco upper Cantabrian; fig. 20 from loc. 983 in La Magda• 2016 in Cevennes Coalfield, middle to upper Canta• lena Coalfield, Stephanian B. All figured specimens brian; fig. 16 from loco 2208 in northern Palencia. are in the author's collection at the University of middle Cantabrian. Sheffield, England. THE PALAEOBOTANlST

PLAT:E 1 ~AGl\ER-T\VO LITTLE ](XOWX SPECIES OF SPHElI'OPIlYLLUlII 511

~-'. PLATE 2 tHE PALAEOBOTANtSt

fLATE 3 WAGNER- TWO LITTLE K OWN SPECIES OF SPHENOPHYLLUM 51~ 580 THE PALAEOB01'ANTST

PLATE 5 WAGXER- TWO LITTLE K~OW~ SPECIES OF SPHENOPHYLLUM 581

fLATE Q