__ THE MATURAL TORY MUSEUM | 5% OCT 2001 | PRE St a] TE H Geology Series wy THE NATURAL HISTORY MUSEUM VOLUME 57 NUMBER2 29 NOVEMBER 2001 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Geology Series is edited in the Museum’s Department of Palaeontology Keeper of Palaeontology: | Dr N. MacLeod (acting) Editor of Bulletin: Dr M.K. Howarth Assistant Editor: Mr C. Jones Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. 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(Geol.) © The Natural History Museum, 2001 Geology Series ISSN 0968-0462 Vol. 57, No. 2, pp. 83-162 The Natural History Museum Cromwell Road London SW7 5BD Issued 29 November 2001 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Geol.) 57(2): 83-93 Issued 29 November 2001 The Cenozoic Brachiopod Terebratula: its type species, neotype, and other included species -— a THE NATURAL DAPHNE E. LEE | oz or ; ee eh at : ee 3° OCT 2001 Department of Geology, University of Otago, P.O. Box 56, Dunedin, New Zealand C.H.C. BRUNTON PRESENTED Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD | | GENE RAL LIBRAR\ Y EMMA TADDEI RUGGIERO a eae Dipartimento di Scienze della Terra della Universita’ di Napoli ‘Federico II’, Largo San Marcellino 10, 80138 Napoli, Italy MASSIMO CALDARA & ORONZO SIMONE Dipartimento di Geologia e Geofisica, Campus Universitario, Via Orabona 4, 70125 Bari, Italy Synopsis. Terebratula terebratula (Linnaeus, 1758) has a long and complex history. The specimen now recognised as the type was first illustrated by Colonna in 1616, and the first use of “Terebratula’ is attributed to Lhwyd, 1699. Colonna’s specimen was refigured by Klein, 1753, and the species Anomia terebratula was described by Linnaeus, 1758, with reference to the Colonna and Klein illustrations. The genus Terebratula was proposed by Miiller in 1776, and Anomia terebratula Linnaeus designated as the type species by Lamarck in 1799, although it was not an originally included species. In spite of this history, the type of the genus was never formally ratified, the whereabouts of the type specimen was unknown, and the age and exact position of the type locality was uncertain. This paper summarises the history of Terebratula terebratula (Linnaeus) from 1616. Anomia terebratula Linnaeus is now accepted as the type species of Terebratula (ICZN ruling, 2000). We have collected new material from a locality near Andria, Italy, from which Colonna collected specimens of Terebratula, and selected a neotype from the Calcarenite di Gravina Formation which is Pliocene in age. Two existing species, Terebratula sinuosa (Brocchi) and T: calabra Seguenza, are placed in synonymy with 7: terebratula. Three species are currently recognised in Terebratula, ranging in age from Miocene to Early Pleistocene when the genus became extinct. probably because of ocean cooling in the Mediterranean region. INTRODUCTION The brachiopod genus Terebratula has a long and complex geologi- cal and nomenclatural history. The nominal genus Terebratula was proposed by Miiller in 1776, and as pointed out by Muir-Wood (1955), it ‘is the first valid post-Linnean brachiopod genus’. Terebratula terebratula (Linnaeus) is the name-bearer for the Order Terebratulida which encompasses loop-bearing brachiopods of Devonian — Recent age and includes most brachiopods living today. The name Terebratula has been widely used for over 200 years: more than 850 specific names were applied to the genus between 1800-1850 alone (Sherborn, 1932). In spite of the ubiquity of the name, the genus and species on which it was originally based have, until recently, been little studied and many basic questions about Terebratula terebratula remained unanswered. As pointed out nearly acentury ago (Buckman, 1907), not only was the type species of the genus unconfirmed, but its type locality and age were uncertain. The present study had several aims. The first objective was to ratify the type of the genus Terebratula for inclusion in the revised Brachiopod Treatise, following the recommendation made by Muir- Wood in the 1965 Treatise volume on the Order Terebratulida. Secondly, we wished to summarise the complex nomenclatural history of the genus and species, Terebratula terebratula, since it was first illustrated and described in 1616. The third aim was to locate the type specimen and/or type locality of T. terebratula, or, if this proved impossible, select a neotype to act as namebearer for the order. Finally, we wished to describe the age and relationships of species currently included in Terebratula. The first objective was achieved with an application to the Inter- national Commission on Zoological Nomenclature to validate the selection of Anomia terebratula Linnaeus as type-species of the © The Natural History Museum, 2001 genus Terebratula as designated by Lamarck in 1799 (Lee & Brunton, 1998; Ruling of the Commission, September 2000). The remaining objectives are achieved in this paper. DERIVATION OF THE NAME The name Terebratula was first used in print by Lhwyd (1699) (Little etal., 1973), and is the oldest generic name in the Phylum Brachiopoda in current use. Terebratula is ‘so-called from the perforated beak of the ventral valve’ (Little et al., 1973: 2265), and is a quasi-diminutive of Latin terebratus, the past participle of terebrare ‘to bore’. The brachiopods listed as Terebratula in Lhwyd’s catalogue of shells in the collections of the Ashmolean Museum, Oxford, were ‘poorly figured specimens from Witney’ (Muir-Wood, 1955: 2). The name came into fairly common use in the 18th century, and some of the numerous brachiopods referred to as Terebratula by other pre- Linnean authors were mentioned by Muir-Wood (1955). The species name terebratula was first used in a valid binomial by Linnaeus in his description of Anomia terebratula in Systema Natu- rae (1758: 703). Linnaeus gave no illustration, but referred to figures in Colonna (1616c), Lister (1678) and Klein (1753). The Colonna, and Colonna/Klein, illustrations are reproduced in Figs 1, 2 THE COLONNA ILLUSTRATION OF TEREBRATULA (FIG. 1) Fabio Colonna (1567—1650) (Fabius Columna) was born in Naples and was one of the first natural historians to use copper plates for engraving botanical and zoological figures. He wrote extensively on 84 D.E. LEE, C.H.C. BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE 22 SOAS a Fig. 1 Reproduction of page 22 in Purpura (Colonna, 1616). fossils, and “was one of the first to place them . . . in a primarily biological context... Colonna also applied the same precise nomen- clature to his fossils as to his living animals, distinguishing different kinds of related fossils with more accuracy then ever before’ (Rudwick, 1985: 42). Colonna was a member of the Accademia dei Lincei (Academy of Lynxes), and in 1606, he published a work on natural history: Minus cognitarum stirpium aliquot, ac etiam rariorum nostro coelo orientium [ecphrasis] . . De aquatilibus, aliisque animalibus quibusdam paucis libellus [plants pp.3-340; animals I-LXXIII]. A new edition was published in 1616 (Colonna, 1616a—c), including part II, Purpura (of which De purpura, aliisque testaceis rarioribus, ARN CGN oes Concha tnamia wertice rofrate Faby Columnez CED ED pp. 1-29, and De glossopetris dissertatio, pp. 31-39, are two chap- ters), in which he described and figured a number of shells, some fossil and some living. His illustration on page 22 (Fig. 1 herein) was a woodcut of five shells. The upper three specimens are double- valved brachiopods, while the lower specimens are internal molds of bivalves. The plate is not numbered, and the five specimens are distinguished by brief captions placed above each specimen. Linnaeus (1758: 703) made three separate references to the illus- trations of brachiopods on this plate in his discussion of species of Anomia. Under Caputserpentis. 200., he gave a brief description, and referred to Column. purp. 22. f.2, 1.e. the smooth brachiopod on the upper right. Brunton & Cocks (1967) discussed in detail the THE CENOZOIC BRACHIOPOD TEREBRATULA Fig. 2 Figures of Concha anomia, the holotype of Anomia terebratula. 2a, woodcut from Colonna (1616); 2b, woodcut from Major (1675); 2¢, engraving from Klein (1753). ambiguities which arose when Linnaeus (1767) changed his descrip- tion of A. caputserpentis from a smooth, fossil brachiopod to a capillate living species (now Terebratulina retusa), although he retained the reference to the Colonna figure. An application to the International Commission on Zoological Nomenclature to change the type species of Terebratulina d’Orbigny, 1847 from Anomia caputserpentis Linnaeus, 1758, to Anomia retusa Linnaeus, 1758, disposed of the ambiguity caused by Linnaeus himself when he altered his definition of A. caputserpentis between 1758 and 1767 (Brunton & Cocks (1967: 295); Ruling of the Commission, 1968). Under Terebratula. 201., Linnaeus listed three separate illustra- tions. The first, ‘Column. purp. 22. f.1, refers to the upper left figure of a smooth, strongly folded brachiopod. The second refers to “List. angl. 240.t. 8. f. 46’, which is a non-plicate Jurassic shell from England (Lister, 1678). The third reference, which confirms that Linnaeus did indeed intend the upper left brachiopod to be the type of terebratula, is unmistakably the same (redrawn) Colonna illustra- tion reproduced in a figure by Klein (1753) (Fig. 2 herein). Buckman (1907: 528) pointed out that Lister’s figure did not match the description given by Linnaeus, and that the Colonna-Klein figure “must be taken as the holotype, which, in fact, has been the usual practice’. Linnaeus included the central figure in Colonna’s plate in his ‘Hysterita. 203. Mus. Tess. 90. t.5.f.1,2,2. Column. purp. 22. f. 3? Trilobos.’ This large, strongly ribbed and folded fossil rhynchonellid does not correspond to the Mus. Tess. illustration listed by Linnaeus, which is an internal mold of Schizophoria (Brunton et al., 1967). The two upper brachiopods in the Colonna figure are both smooth terebratulid brachiopods with large open foramens and strongly delineated growth bands. Across both drawings is the caption: ‘“Con- cha anomia vertice rostrato’, and beneath the left figure, although probably relating to the central brachiopod on the plate, is the word Altera (another). On the adjacent page 23, in Purpura, the chapter is headed Concha rarior Anomia vertice rostrato. I. Cap. XII (or ‘rare Anomia shell with apical beak’). Although there is no scale on the illustration, the brachiopod on the upper right is at natural size (icon magnitudinem aequat on page 23 in Purpura). CONCHA ANOMIA VERTICE ROSTRATO Some of the problems which have made it difficult for earlier and present authors to define Terebratula arise from the lack of corre- spondence between the figures on page 22 of Colonna, and the text on the following pages. In particular, several specimens mentioned on pages 23—24 are not illustrated at all, although they were num- bered sequentially by the author. Thus, Lee & Brunton (1998) assumed that the strongly folded figure on the upper left on page 22 was that described in the text on the facing page (Cap. XII) under the heading ‘Concha rarior Anomia vertice rostrato’, and accordingly they concluded that this specimen was that collected by Colonna from Andria. More recently, we have found that this might not be correct. In five original copies of Colonna’s Purpura (1616c) held by the University of Naples ‘Federico IV (two in the Library, and three in the Department of Biologia Vegetale and the Botanical Garden) and in another original copy held in the Botany Library of the Natural History Museum, London, the two figures at the top of page 22 are marked with ‘4’ (upper left), and ‘TV (upper right). These small numbers are not shown in the two published reproductions of Colonna’s plate (Dollfus & Dautzenberg, 1932; Muir-Wood, 1955), nor in the 1675 edition of Colonna’s Purpura edited by Major. The following section attempts to clarify the problems we have encountered. 1. Colonna described four Conchae in the text on pages 23—24 and marked them with a Roman or Arabic notation. Of these, only the first (1) and the fourth (4) are figured on page 22. The four Conchae are: (1) Concha rarior Anomia vertice rostrato. |. Cap. XII (‘TV on page 22 and ‘I’ on page 23). The description of ‘I’ corre- sponds to figure ‘I’ (page 22, top right). (11) minor 2 (‘2’ at the edge of the margin of line 29 on page 23). This specimen is not figured by Colonna. (111) Altera Neptunia maior II. imbricata. Cap. X10 (‘UT on page 24). This specimen is not figured by Colonna. (iv) Concha Anomia IV. margine undosa. Cap. XIV. (‘4 on page 22 and ‘IV’ of page 24). The description of ‘IV’ corresponds to figure “4° (page 22, top left). . The first shell, (that is the smooth, unfolded specimen on the upper right on page 22) comes from Andria. The third comes from Nettuno (50 km south of Rome). The second and the fourth come from the Museum of Ferrante Imperato in Naples and their provenance is not given. 3. Colonna thought the four Conchae (ie., the figured and unfigured specimens described on pp. 23-24) were similar to each other. Indeed, in the index on p. 41 three of these are included under a single name “Concha anomia quae, rarior vertice rostrato Plin. i) 86 D.E. LEE, C.H.C. BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE descript. 23. icon. 22. Altera imbricata. 24. altera margine undosa, ibid. & 25°. The two figures on the top of page 22 are joined under the same caption ‘Concha anomia vertice rostrato . The figure in the centre of page 22, described on pages 24—25 as ‘Concha altera Anomia striata [trilobos] rarior. 1. Cap. XV.’, is considered as another shell and its numeration starts again from ‘T’. Much of the confusion over the identity of the two figures on the top of page 22 stems from the fact that Colonna described the shell placed on the upper right first, and on a following page discussed the shell portrayed on the upper left, using a different practice from that which became well established in the following centuries. The confusion increased further when later authors assigned to these figures two numbers that Colonna had never employed. Thus Linnaeus (1758), following modern convention, designated the strongly folded specimen on the upper left as f. 1 (ie., figure 1) and that on the upper right as f. 2 (ie., figure 2). Brocchi (1814), and the present authors, until lately, have done the same. The problem of deducing which shell description accompanied which illustration was compounded in a second edition of Colonna’s work, with the associated text from the earlier edition subdivided into numbered paragraphs, which was reprinted posthumously in 1675 by J.D. Major. In this edition, the redrawn figure of the strongly folded specimen from the upper left of page 22 in the 1616 edition (1.e. Anomia terebratula of Linnaeus) appeared on page 32 within the text referring to the Andria locality. Similarly, in the Dictionarium this figure was noted by Major as that described in Cap. 12, of Colonna (1616c: 23). A comparison of the three figures (Fig. 2 herein) shows that the Klein figure listed by Linnaeus (Klein, 1753; Tab. Nostra XI. n. 74), was redrawn from this later edition (Major, 1675) (Fab. Columnae de Purp. Cap. XII $.3. pag. 32 [Klein 1753, p. 171-2]). Dollfus & Dautzenberg (1932) correctly interpreted the refer- ences to Major’s figures, but added to the confusion, by giving the same citation for both the upper right and upper left figures: te. ‘p. 22, fig.1 (2e)’. These authors also assumed that the caption ‘Altera’ pertained to the upper left figure, whereas it undoubtedly refers to the figure in the center (“Altera [trilobos]’), because the captions in all seven figures in Colonna’s Purpura (pp. 13, 16, 20, 22, 27, 30, 33) are placed over, and never under, the relevant figures. DISCUSSION AND COMMENTS This discovery, as our manuscript was almost ready for submission and after the ICZN had approved the selection of a neotype for Terebratula terebratula (Linnaeus), raises some issues which need further discussion. If indeed the provenance of the specimen selected by Linnaeus as the type of Anomia terebratula is unknown and the specimen is lost, then the basis for the species, and consequently the genus, family and superfamily would remain uncertain. However, since Lee & Brunton (1998) have already nominated a neotype from Colonna’s locality near Andria, the neotype locality now becomes the type locality for the species, regardless of the locality of the original Colonna brachiopod (International Code of Zoological Nomenclature Article 75f). Is it possible then to determine the provenance of the original specimen of Colonna selected by Linnaeus as the type of Anomia terebratula? Firstly, it is obvious that Colonna’s caption: ‘Concha anomia vertice rostrato’ applies to both of the brachiopods figured on the upper right and upper left of page 22, the first of which (that on the upper right) certainly came from Pliocene strata near Andria, and that Colonna himself regarded these specimens as similar to one another. Secondly, six original copies of Colonna (1616c) that we have consulted have the numbers:’I’, “4’ and ‘Il’ written beside the three brachiopods figured on page 22. These numbers are not included in Major’s edition. Linnaeus (1758, 1767) and Brocchi (1814), if aware of these numbers, used different, modern numbering (i.e. f.1 for Colonna’s specimen 4 on the upper left), and Major (1675) and Lee & Brunton (1998) considered that Colonna’s description on page 23 referred to the figure on the upper left. Thirdly, even if the specimen of Colonna’s (1616c: 22) upper left figure is from an unknown locality, it was filled with white, loose sediment (Colonna, 1616c: 24), and it is possible that it was collected from the same Pliocene calcarenites at Andria. From a close examination of the Colonna woodcut, it seems likely that his specimen was somewhat deformed. It has a large, open fora- men, and two strongly developed plicae/folds on the dorsal valve that begin at an early stage of growth, and would have resulted in a strong sulciplication (margine undosa) of the anterior commissure. The artist (?Colonna himself) may have exaggerated the depth of the folds, although the depiction of the other brachiopods on the plate seem to be faithful to reality. No undeformed specimens collected by the authors have folds as strongly developed as those depicted in the woodcut. From the many specimens of 7erebratula collected by the authors from Colonna’s Andria locality and from elsewhere in Italy, it is apparent that the brachiopods in any fossil assemblage/population vary considerably in the degree of folding and may be rectimarginate to biplicate or sulciplicate. Thus, both specimens labelled by Colonna as Concha anomia vertice rostrato are species of Terebratula (sensu lato), and given the wide variation in populations of Neogene Terebratula, might be conspecific. Certainly, the specimens of Terebratula terebratula figured by us in this paper (Figs 6—9) fall somewhere in the middle of the two short-looped brachiopods illustrated in Colonna’s woodcut. The specimen of Pliocene Terebratula terebratula from Monte Mario selected and figured by Buckman (1907) and illustrated in the 1965 Treatise, is crushed and deformed in a similar manner. In the Natural History Museum, London, there are several collections of Terebratula from this same horizon at Monte Mario, near Rome. These specimens, which are almost certainly conspecific, vary from small rectimarginate (juve- nile) specimens (labelled 7: depressa) to large rectimarginate individuals (named T. grandis), to examples with moderate sulciplication (labelled 7: ampulla or T. terebratula). Thus, our selection of a neotype from a locality described and collected by Colonna closely follows the recommendations of the Code. It should be noted that Muir-Wood’s (1955: fig. 2) caption for her reproduction of the original Colonna figure is misleading. The caption reads ‘Reproduction of early drawings of Brachiopods figured as “Concha anomia’, and taken from Fabio Colonna’s first edition of de Purpura, 1616,p.22.The first figure is of aspecimen from Mte. Mario, near Rome, and is probably of Tertiary (?Pliocene) age; the other two figures may represent Jurassic forms’. However, as shown here, the first figure is from an unknown locality, while the second specimen is of Pliocene age and came from near Andria, not Rome. TYPE LOCALITY OF TEREBRATULA TEREBRATULA Colonna (1616c: 23) described the locality from which he collected his specimen of Terebratula shown on the top right of his figure THE CENOZOIC BRACHIOPOD TEREBRATULA al es if Sg? della om ad a SL eo a2 : C. deSiliigf > .- A Vefconuaw BPal. Diucale LS Benedetto DSP Ee ae B drea OPotta dS Andrea P. Porta della Batra QPeorta del. oS Mo BS. hiars' IS Angelo TLa Corte V.Molmra X Annungicts cant 2S AL de. erg Beas tele PooG: urls pa 87 M. iy Wit os oS . Re q eee a iin SSS See oes ~— = — —— 5 Lge zratells HS Cir eae : Z ichafiano Z,ZLoccolantrt, Capytte: Yous 55. Vite 6S. Ang. JIL: ; Pe Fig. 3 Reproduction of Pacichelli’s late 17" century map of Andria. North is at the bottom of the figure, and the church of Santa Maria dei Miracoli (2 in the key) is west of the city near the right hand edge of the map. (Colonna, 1616: 22) as follows: ‘We found this shell full of the white sediment | tophacea concretione’ | on which that whole sloping area or hillis made. This is constituted not so much of loose sediments, as of fragments of various shells and unbroken shells too. We collected this one and others in the small valley or ditch a little below the Church of D. Maria de Andria, which is situated one mile outside the city’ (see the Appendix for a translation of Colonna’s pp. 23 and 24). In the modern town of Andria the names of five churches include the word ‘Maria’, and to find which of them was the one referred to by Colonna it was necessary to consult Pacichelli’s late seventeenth century map of Andria (Fig. 3). The churches of Santa Maria Nova (4 on Fig. 3) and Santa Maria dei Miracoli (2 on Fig. 3) are both situated outside the town to the west (west is on the right hand side of Fig. 3), but only the latter church is built directly on the tophacea concretione’ (= Calcarenite di Gravina Formation). On either side of a small natural valley (now dry) adjacent to the church of Santa Maria dei Miracoli 1-3m high cliffs of white, well-cemented calcarenite out- crop sporadically (Fig. 4), and specimens of Terebratula are scattered throughout the calcarenite showing that this locality is undoubtedly the one visited by Colonna. The brachiopods are not uncommon, but are often fragile and/or broken. The basement rocks of the region around Andria are Lower Cretaceous in age. Overlying these with angular unconformity are 25 to 30m of coarse-grained highly fossiliferous marine biocalcarenites and calcirudites of the Calcarenite di Gravina For- mation which is widespread in this area. In the vicinity of the Madonna d’Andria church, the sequence consists of 2m of fine, bioturbated, massive calcarenites lacking macrofossils. Above this is a coarse, bioturbated calcarenite up to 4m thick which includes oysters, scattered pectinids, echinoids and brachiopods (Fig. 5). The upper 2m thick bed is a well-cemented, very fossiliferous calcarenite with a discontinuous oyster bed near its base. Fossils include Ostrea, Chlamys, internal moulds of bivalves such as Veneridae, echinoids, calcareous algae and brachiopods. On macrofossil (Caldara, 1987; Caldara & Gissi, 1993) and microfossil (Taddei Ruggiero, 1996) evidence, the lower part of the Calcarenite di Gravina Formation is locally Middle to Upper Pliocene, or possibly Upper Pliocene to Lower Pleistocene in age. No trace of a cool-temperature Pleistocene macrofossil fauna was found during our visit. Thus, the type locality for the neotype of Terebratula terebratula (Linnaeus) is adjacent to the church of Santa Maria dei Miracoli, about 2 km west of Andria, Puglia, Italy in the Calcarenite di Gravina Formation, of Upper Pliocene age. The specimen selected as neotype comes from beneath a small overhang about 200 m north of the church. The outcrops are difficult to access, and involve crossing private property. SYSTEMATIC DESCRIPTIONS Order TEREBRATULIDA Moore, 1952 Superfamily TEREBRATULOIDEA Schuchert, 1913 Family TEREBRATULIDAE Gray, 1840 DIAGNOSIS. Medium to large, ventribiconvex, rectimarginate to uniplicate or biplicate, rarely unisulcate or sulciplicate, smooth or 88 D.E. LEE, C.H.C. BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE Fig. 4 Outcrops of the Calcarenite di Gravina Formation, from which the neotype was collected, in the small valley north of the church. THE CENOZOIC BRACHIOPOD TEREBRATULA 89 Tyrrhenian Sea Fig.5 Location map, geological map and stratigraphic column for the Andria region, Puglia, Italy. A, alluvial deposits (Holocene); B, terraced alluvial deposits (Pleistocene); C, terraced marine deposits (Pleistocene); D, Calcarenite di Gravina Formation (Upper Pliocene); E, Calcare di Bari Formation (Lower Cretaceous); F, Colonna’s locality; G, outcrop from which the neotype of Terebratula was collected (see stratigraphic column). 1, Pectinidae; 2, Veneridae; 3, gastropods; 4, brachiopods; 5, echinoids; 6, calcareous algae; 7, oysters; 8, bioturbations; 9, mud pebbles; 10, fine calcarenite; 11, coarse calcarenite. with fine radial capillae; loop short, triangular; outer hinge plates usually concave or flat, commonly attached to dorsal edge of crural base, inner hinge plates rarely developed. Subfamily TEREBRATULINAE Gray, 1840 Genus TEREBRATULA Miller, 1776: 249 DIAGNOsIS. Medium to large, subpentagonal to broadly oval, smooth; anterior commissure rectimarginate to uniplicate or sulciplicate; beak short, erect; foramen large, symphytium partly visible. Pedicle collar short; cardinal process flat and semielliptical to a thickened boss; outer hinge plates narrow or lacking; crural processes may be long; loop short, broadly triangular; transverse band narrow, forming a low arch. TYPE SPECIES. Anomia terebratula Linnaeus, 1758, by the subse- quent designation of Lamarck (1799: 89). GEOGRAPHIC RANGE. _ Italy, Sicily, Malta, Spain, Algeria. STRATIGRAPHIC RANGE. Miocene — Early Pleistocene. REMARKS. The great majority of the thousand or more specific names attributed to Terebratula have long been accommodated in other genera. However, although T. terebratula is the oldest available name for the medium to large, smooth, short-looped Miocene — Pleistocene terebratulids from Italy and the circum-Mediterranean region, it has rarely been used in the literature or in identification of specimens for the reasons outlined above. This is due mainly to the uncertainties surrounding the identity, age and type locality of the Colonna specimen. The Colonna work is rare, and no translation has hitherto been available. Secondly, there was doubt as to the correct type species for the genus. Gaetani & Sacca (1985), in a paper dealing with systematics and shell structure of brachiopods of Miocene — Pleistocene age from southern Italy, commented on the problem ofrecognising T: terebratula. They concluded that there were two valid species: Terebratula sinuosa (Brocchi) and T: calabra Seguenza which were restricted to the Upper Miocene and Pliocene respectively. Cooper (1983) identified large sulciplicate specimens of Pliocene age from Monte Mario, Rome, as T. ampulla. Other authors (Taddei Ruggiero, 1986, 1990, 1994, 1996) have identified large, shellbed-forming Pleistocene specimens as T. scillae Seguenza. Until variation within large populations of the species placed in Terebratulahas been studied, we are able to recognise only three valid species of Terebratula — T. terebratula (Linnaeus, 1758), T: ampulla (Brocchi, 1814) and T: scillae Seguenza, 1871. Terebratula terebratula (Linnaeus, 1758) Figs 6-9 1758 Anomia terebratula Linnaeus: 703. SYNONYMS 1. Anomia sinuosa Brocchi, 1814: 468, is an objective synonym because Brocchi gave no figure, but referred to ‘Column. 22, f.1°, which is the holotype of Anomia terebratula. 2. Terebratula calabra Seguenza, 1871: 64 3. Terebratula costae Seguenza, 1871: 67; Taddei Ruggiero, 1994: 206. 90 D.E. LEE, C.H.C. BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE 9b 8b Figs 6-9 Neotype and 3 topotypes of Terebratula terebratula (Linnaeus) from a Colonna locality at Madonna dei Miracoli, Andria, Italy. 6a—c, dorsal, lateral and anterior views of neotype, NHM BG152 (length 55.4mm); 7a—c, dorsal, ventral and anterior views of topotype, NHM BG194 (length 50.8mm); 8a, b, ventral and anterior views of topotype, NHM BG195, showing M-shaped anterior commissure; 9a, b, ventral views of loop of topotype, NHM BG196. All figures natural size, except Fig. 9b. THE CENOZOIC BRACHIOPOD TEREBRATULA TYPE SPECIMENS. The holotype, the specimen figured by Colonna (1616c), is lost. We here nominate as neotype (Fig. 6a—c), an entire specimen from the Calcarenite di Gravina Formation near a locality collected by Colonna, in the collection of the Natural History Mu- seum, London (BMNH BG152), collected 22 March 1998 by M. Caldara & O. Simone. Dimensions of the neotype are: length 55.4 mm, width 43 mm, thickness 30mm. A well-preserved complete topotype with the dimensions: length 50.8 mm, width 42 mm, thickness 28 mm, is also figured (Figs 7a—c). Two further topotypes, one with a complete loop (Figs 9a, b), and a dorsal valve with a strongly-M-shaped anterior commissure (Figs 8a, b) are also illustrated. MATERIAL. The brachiopods from the Calcarenite di Gravina For- mation are frequently broken across mid-valve, or are separated valves. Most specimens are infilled with coarse, white, hard, moder- ately cemented calcarenite, and few have a complete anterior commissure. A number of topotypes (BG153—161, BG194—196) are held in The Natural History Museum, London. TYPELOCALITY. The type locality is an outcrop of Calcarenite di Gravina Formation, of Pliocene age, on the east side of a small dry valley, about 200m north of the church of Madonna dei Miracoli (41°13'52"N; 16°16'00"E), about 2km west of Andria, Puglia, Italy. AGE. Late Miocene (Tortonian), Pliocene. DISTRIBUTION. Puglia, Calabria, Tuscany, Emilia (Piacentino), Abruzzi, Spain (Alicante). DESCRIPTION. Shell of medium to large size, biconvex, anterior commissure rectimarginate, uniplicate to sulciplicate, two broad plicae may be developed; beak short, massive, suberect; foramen large, mesothyrid to permesothyrid; symphytium narrow, partly concealed. Pedicle collar short; hinge teeth with moderately swollen bases. Cardinal process semielliptical, moderately protuberant; outer hinge plates narrow; no inner hinge plates; crural bases fused to socket ridges to form a deep V-shaped trough, crural processes long; loop broadly triangular, about 0.3 valve length, transverse band narrow, forming medially flattened low arch. Terebratula ampulla (Brocchi, 1814) 1814. Anomia ampulla Brocchi: 466 AGE. Pliocene. DISTRIBUTION. Italy (Emilia (Piacentino), Tuscany, from Brocchi’s list), but not Calabria (see Seguenza, 1871). COMMENTS. A medium-sized, strongly bisulcate species. Terebratula scillae Seguenza, 1871 1871 Terebratula scillae Seguenza: 39 AGE. Early Pleistocene. DISTRIBUTION. Calabria, Puglia, Sicily. COMMENTS. The largest species of Terebratula (up to 95mm in length), which formed extensive shellbeds in the Early Pleistocene. ECOLOGY AND EXTINCTION OF TEREBRATULA Terebratula grandis Blumenbach, 1803, now included in the genus Pliothyrina (see Cooper, 1983), may be the ancestor of Terebratula 9] sensu stricto. This large species from the Oligocene of Germany needs further study. Pliothyrina appears to have been widespread in northern Europe and England (see collections in the Natural History Museum, London), whereas Terebratula was abundant in the Med- iterranean region. Both groups became extinct in the Plio-Pleistocene. Terebratula was widely distributed in the Mediterranean region from the Miocene until the early Pleistocene. It lived in circalittoral environments on muddy, biodetrital seafloors, attached to substrates which included bivalves and other brachiopods. During the Messinian, when the Mediterranean basin became too saline to support normal marine life, the brachiopod fauna disappeared from the region. With flooding of the Atlantic sea into the Mediterranean basin, the Mediterranean was recolonised by a brachiopod biota with Atlantic affinities (Logan, 1979). This does not account for the reappearance of Terebratula, which may have survived in refugia. In the Pliocene, Terebratula was an important component of a widespread brachiopod fauna which included Aphelesia and Megerlia (Gaetani, 1986; Taddei Ruggiero, 1994, 1995). The youngest records are of very large Terebratula scillae which formed vast shellbeds in the Early Pleistocene (Taddei Ruggiero, 1986, 1994). The extinction of Terebratula by the Middle Pleistocene appears to be related to the reduction in sea temperatures as Pleistocene cooling proceeded. ACKNOWLEDGEMENTS. Professor Maurizio Gaetani gave initial introduc- tions to geologists in Italy. We thank Eileen Brunton and Sarah Long at the The Natural History Museum, London, for help with bibliography and for photography of the brachiopods. We also thank Professor John Barsby, University of Otago, for translations of Colonna’s work, Bill Lee and Doug Campbell for improvements to the manuscript, Andrew Grebneff for speci- men preparation and Ewan Fordyce for photography. The Royal Society of New Zealand contributed towards travel to the British Museum (Natural History) in 1991, and funding from the University of Otago Division of Sciences, and the Department of Scienze della Terra of the University of Napoli, is gratefully acknowledged. Professor Giovanni Aliotta, I] University of Napoli, gave us new information on Colonna, Professor Roberto Taddei, University of Napoli, translated some passages of Purpura, and the Library of the University and the Department of Biologia Vegetale of the University of Napoli ‘Federico IV allowed photographic reproduction of Colonna’s Purpura. REFERENCES Brocchi, G. 1814. Conchiologia Fossile Subappennina, 2 vols., 712 pp, 32pls. Milan, Stamperia Reale. Brunton, C.H.C. & Cocks, L.R.M. 1967. Terebratulina d’ Orbigny, 1847 (Brachiopoda): Proposed designation of a type-species under the plenary powers. Z.N. (S.) 1809. Bulletin of Zoological Nomenclature, 24: 294-296. . & Dance, S.P. 1967. Brachiopods in the Linnaean Collection. Proceedings of the Linnean Society of London, 178: 161-183, pls 14. Buckman, S.S. 1907. Brachiopod Nomenclature: the Genotype of Terebratula. Annals & Magazine of Natural History, series 7, 19: 525-531, pl. XI. Caldara, M. 1987. Segnalazione di pliocene medio e superioire nelle Murge Settentrionali (Puglia). Bollettino della Societa Geologica Italiana 106: 153-162. & Gissi, F. 1993. Le ‘Biocalcareniti di Gravina’ del margine ofantino delle Murge: considerazioni stratigrafiche, paleoambientali e tettoniche. Bonifica, 8, 3: 153-171. Colonna, F. 1606. Minus cognitarum stirpium aliquot, ac etiam rariorum nostro coelo orientium ecphrasis, etc. (1st edition) Rome, G. Facciotto. 1616a. Minus cognitarum rariorumque nostro coelo orientium stirpium ecphrasis, etc. 2nd edition. Rome, J. Mascardo. — 1616b. Minus cognitarum stirpium pars altera. \st edition. Rome, J. Mascardo. — 16l6c. Purpura: Hoc est de Purpura ab Animali testaceo fusa, de hoc ipso Animali, aliisquibus rarioribus Testaceis quibusdam. iv + 42 pp, 2pls. Rome, J. Mascardo. Cooper, G.A. 1983. The Terebratulacea (Brachiopoda), Triassic to Recent: A Study of the Brachidia (Loops). Smithsonian Contributions to Paleobiology, 50: 290pp, 77 pls. 92 1D).18), ILJ5} 3, C.lal{C- Dollfus, G. & Dautzenberg, P. 1932. Les mollusques de Fabius Columna. Journal de Conchyliologie, 76: 283-333. 15 pls. Gaetani, M. 1986. Brachiopod communities from the Plio/Pleistocene of Calabria and Sicilia (Italy). Jn, Racheboeuf, P.R. & Emig, C. (editors), Les Brachiopodes fossiles et actuels. Biostratigraphie du Paleozoique, 4: 281-288, | tab., 2 figs. & Sacca, D. 1985. Brachiopodi Neogenici e Pleistocenici della Provincia di Messina e della Calabria Meridionale. Geologica Romana, 22: 1-43. Klein, J.T. 1753. J.T. Klein Tentamen methodi Ostracologicae, sive dispositio naturalis Cochlidum et concharum. \77pp, 12pls. Lugduni Batavorum. Lamarck, J.B.P.A. de M. 1799. Prodrome d’ une Nouvelle Classification des Coquilles. Mémoires Sociétié d'Histoire Naturelle de Paris, 1: 63-91. Lee, D.E. & Brunton, C.H.C. 1998. Terebratula Miiller, 1776 (Brachiopoda): pro- posed designation of Anomia terebratula Linnaeus, 1758 as the type species. Bulletin of Zoological Nomenclature, 55 (4): 220-223. Lhwyd, E. 1699. Lithophylacii Briiannici Ichnographia, sive Lapidum aliorumge; Fossilium Britannicorum singulari figura insignium. 145pp, 17pls. Londini & Lipsiae. Linnaeus, C. 1758. Systema naturae, sive Regna tria Naturae systematice proposita per Classes, Ordines, Genera et Species, 10th edition, Tom 1: Regnum Animale, pp. (II) 1-824. Stockholm. 1767. Systema naturae, sive Regna tria Naturae systematice proposita per Classes, Ordines, Genera et Species, 12th edition, Tom 1, part 2: 533-1327. Stock- holm. Lister, M. 1678. Historia Animalium Angliae . . . (et) de lapidibus ejusdem insulae ad cochlearum quandum imaginem figuratis. vi + 250 pp., 9 pls. London. Little, W., Fowler, H.W., Coulson, J., Onions, C.T. & Friedrichsen, G.W.S. 1973. The Shorter Oxford English Dictionary on Historical Principles. 2 volumes, 3rd edition. Oxford, Clarendon Press. Logan, A. 1979. The recent Brachiopoda of the Mediterranean Sea. Bulletin de I’ Institut Océanographique de Monaco, 72 (1434): 1-112, 10pls., 22 figs. Major, J.D. 1675. F Columnae...Opusculum de Purpura... iterum luci datum opera ac studio J.D. Majoris ... cujus ... accesserunt annotationes quaedam: (Doctrinae BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE de Testaceis, inordinem congruum redactae specimen, tabulis aliquot comprehensum . cum brevi Dictionario Ostracologico, de partibus Testaceorum auctore J.D.M.Med.D. xii + 72 pp, text illustrations. Kili. Muir-Wood, H.M. 1955. A history of the classification of the Phylum Brachiopoda. British Museum (Natural History), London. 124pp. — 1965. Mesozoic and Cenozoic Terebratulidina. In, Moore, R.C. (editor), Treatise on Invertebrate Paleontology, Part H, Brachiopoda, vol. 2: 762-816. Geological Society of America and University of Kansas Press, Lawrence. Miiller, O.F. 1776. Zoologiae Danicae Prodromus seu Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. xxxii + 282pp. Havniae (Copenhagen). Rudwick, M.J.S. 1985. The Meaning of Fossils: Episodes in the History of Palaeontol- ogy. 2nd edition. 287pp. University of Chicago Press, Chicago and London. Seguenza, G. 1871. Studii paleontologici sui Brachiopodi terziarii dell Italia meridionale. Bollettino Malacologica Italiano, 4: 1-79, 6 plates. Sherborn, C.D. 1932. Index Animalium. Part Il: Index to Generic Names, showing the trivial names associated with each, from 1801 to 1850. Cambridge University Press, London. Taddei Ruggiero, E. 1986. Croissance allométrique de Terebratula scillae Seguenza. In, P-R. Racheboeuf & C. Emig (editors), Les Brachiopodes fossiles et actuels. Biostratigraphie du Paleozoique, 4: 381-389, \pl., 1 tab., 4 figs. 1990. Analisi paleoecologica di un affioramento di Calcareniti di Gravina. Atti 1V Simposio di ecologiae Paleoecologica delle Comunita Bentoniche, pp. 443-454. 5 figs. 1994. Neogene Salento brachiopod palaeocommunities. Bollettino Societa Paleontologica Italiana, 33 (2): 197-213, 3 pls., 2 tabs., 16 figs. 1995. Le paleocomunita’ circalitorali a brachiopodi plio-pleistoceniche della Puglia. Atti museo Geologico Paleontologico Monfalcone, quaderno speciale 3: 89— 5). —— 1996. Biostratigrafia e Paleoecologia delle Calcareniti di Gravina nei dintorni di Cerignola (Brachiopodi e Foraminiferi). Memorie Societa Geologica Italiana, 51: 197-207, 6 tabs., 5 figs. APPENDIX Translation of Colonna, 1616c, page 23 and part of page 24, provided by Professor John Barsby, Department of Classics, University of Otago, and Professor Roberto Taddei, Dipartimento di Biologia Vegetale, Universita’ di Napoli ‘Federico II’. Page 23, Concha rarior Anomia vertice rostrato. I. Cap.XII. “Now we will discuss the one found in the city of Andria. If anyone of greater curiosity would seek them in that place, he will find many of the more rare ones, still unrecognised and unseen. And he will notice that nature has had a lot of fun in forming them. The appearance of this [shell] is smooth, depressed, a little elongated (longer than broad), differing from other shells especially in the fact that one of the two valves is longer and that it extends its umbo and the whole apex which is longer and rounder and sharper and sticks out above the apex of the other valve, so that the last apex is connected beneath the umbo of the first one. The shell is small, white, thin, and a little bit wrinkled transversely by additions [=with the surface marked by growth lines], but not for that reason rough, but smooth. We found this shell full of the white sediment on which that whole sloping area or hill is made. This is constituted not so much of loose sediments, as of fragments of various shells and unbroken shells too. We collected this one and others in the small valley or ditch a little below the Church of D. Maria de Andria, which is situated one mile outside the city. We were there to pay our thanks for favours received from most holy Mother of God, among the others who assemble there in great crowds every day to pay their vows. The church is adorned with large gifts and signs of miracles: the church itself has a sumptuous structure, as does the monastery. We observed one shell like this at the Museum of our very learned Imperato, a rich treasure of all natural things. This shell has a little sinuous margin and the longer valve has a slight groove in the back, another in the middle, protruding in opposite way (?). All the shells have the same particular feature, i.e. an orifice in the rostrate, prominent, apex, from which they can, as a turbine, suck and eject water, in the manner of a ‘sylvester Lepas’ or an ‘Auris marinea’. The figure is natural size. A stony shell like this, but much greater, is figured in the first part, under the name Concha gibbosa.’ Page 24, Altera Neptunia maior III. imbricata. Cap. XIII. ‘Another twice larger, with sinuous margin too, we found in tufaceous or sandy materials near Albano, in which is the ditch of Castello (Castle) or Arce. And there are many different shells never complete, but all piled-up and tangled . . . The shell is 3 inches long, 2 wide, and in the middle has like another shell built on. Concha anomia IV. margine undosa. Cap. XIV “This differs from the previous similar one in the colour which is on the pale side, but was full of white, loose sediment too, for the wavy and curly margin so that it is like an “M’ letter, for the back is inflated and not hollow, because in the other valve a triple groove is recognisable, but all have a pierced apex. I had these, among the others, from the Museum of our Imperato.’ THE CENOZOIC BRACHIOPOD TEREBRATULA Purpura. 23 Goncha rarior Anomiavertice rofirato. I. Cap. XII. A Nomias Conchas illas effe dicimus , quarumaltera pars co- herensaliquo modo ab altera effigie , aut maguitadine aut vtroque modo differat wrouas , quidem contrarium eft vetbi ¢Aer#t, quod eft , fimilis, par, equalis, (cilicet , diffimilis,impar, inequalis : Lib.g.c.3§ ideo inter ceteras notas 4 Plinio memoratas, quibus Concharum» varietates diftinguit plurimas , cum inzqualitatis notam non inue- netimus, Anomias Conchas appellare libuit, vel Plinij more illasy vertice roftrato,dicere; quarum etiam differentiz func plures : Nunc de hac in Ciuitate Andriz reperta verba faciemus, in quo loco ft quis curiofior perquirere vellet, illas etiamrariores non paucas ine ueniet adhucincognitas, & inuifas. Naturamd; in his efforman- dis multum lufiffe animaduertet. Huius igitur effigies leuis,depref= fa,parum oblonga,ab alijs Conchis in hoc precipué differens ,quod altera Conchz pars oblongior eft, collum, ceruicemd; totam, quz oblongior eft, & rorundior,atque acutior, prominetd; fupra cerui- cem alterius diffundit, ve infra illius collum altera ceruix conne- @atur.- Concha parua‘eft, candida, tenuis , obliqu¢ parumaddita- mentis rugofa , fednon obid afpera , fed leuis. Kepletam inueni- mus candida tophacea concretione, qua totus ille clinofus locus, fiue collis eft editus , qui non magis terrena concretione.tophacea, guam variarum Concharum fragmentis, & integris etiam innume- ris eft compa@us: hanc & alias in vallecula illa, fiue fotla quadam parum fubtus Ecclefiam D. Mariz de Andria,extra vrbem miliario fita,excepimus :illic enim ob gratiasa San@ifsima Dei Genitrices acceptas referendas fuimus, ficuti & alij magna cum frequentia vo- ta foluentes concurrunt quotidie : Ecclefia quidem illa magnis do- nis, & miraculorum fignis ornatur,nec non fumpruola ftrucura ip- fa Ecclefia, & Monafterium . Huius fimilem apud do@ifsimum Im- minors; peratum noftrum in fuo Mufzo,rerum omnium nature fatis copio- fo chefauro, obferuauimus , que margine erat parum vndola , & longiore conche parte canalem vix cofpicuum in dorfo,altero verd in medio,c6trario modo extuberante,omnefque peculiar! nota sut reditz,;quod ceruice prominence roftrata,pertufa oriuntur,qua tur sinatorum more aquam haurire & expuere, fyluefiris Lepadis , aut Auris marinz modo poffunt: icon magnitudinem zquat.Huic fimi- Jem maiorem multo lapideam depinximus in prima parte,nomine Conchz gibbofe. ae Altera Fig. 10 Reproduction of Colonna, 1616c, pp. 23, 24. 24 Fabij Columnz Altera Neptania maior III, imbricata. Cap. XIII. D Vplo maiorem alteram Neptuni reperimus Albanen. dio- cefis etiam vndofa margine in tophaccis , fiue fabulofis illis concretionibus, in quibus Caftellifiue Arcis toflact, & maxima variara tcftaricongeries con(picitur ,nec vna reperies integram » fed omnes inter fe congeltas, & implicatas, ve non Natura,fed mae sis impulfu fra@as, & eieG&as cum (abulone, & molem illam to- phacea conftruxiffe teftari fragmentis repleta fit cenfendum:quare mare aliquando varijs in regionibus excreuifle & zftuaffe fatis co- ftat,locaq;immutata. Mirum quidem eft huiufinodi teltas recentes, & viuas hodie non reperiri, qaamobrem ¢longé marts alluuiones profe@as, & adueG@as cenfemus potius , quam Naturam defijife fi- miles parere.tres pollices longa,duos lata e(t celta, habetq; in me- dio veluti aleeram concham fuper clatam imbricis modo,vt in alijs obferuatur, prefertim pectunculis . Concha Anomia IV.margine undofa. Cap. XIV. D Iffert hzc 4 fuperiore congenere colore ad pallidum inclinan- te, que etiam repleta erat concretione candida terrea, 5 quod margine fit vndofa in {p'ras contorta , M, litteram reprafen- tance, dorfumg; habeat elatum, noncauum; altera parte triplici canali diftinguatur,fed omnes ceruicem habent perforatam,¢x ime perati noftri mufzo habuimus inter alias . Concha altera Anomia firiata vs thoBss rarior. I. Cap. XV. E X eodem genere & hze videtur,maior Concha,& crafsior;cu- ius obefa valua minor eft,atque tribus fimul iunctis teftis,me dia magis excuberante conftru-ta videtur, & fenis ftrijs, cotidemd; ftrigibus in Gingulis lobis , quibus margines denticulatz fiunt , infe- cta;preterquam parsinteriecta interlobos,quz recta linea margine definit. Alcera parte, qua value in caput prominet, medium habet lobum deprefsiorem, & oblongiorem,reliquis 4 latere brenioribus, & elatis, eodemq; modo ftriatis ; qua parte tota concha expantis alis auiculam incuruam reprafencace videtur Concha cortex ca- ftanea - —- e 9 inom the ay v » , & ; 7 es is, , _ = eo" ; ® : a c =n =7@ ¥ . A it a = a —— ee a - A Sr ., --o-,, —— if ae J he = 3 ea ey a ~~ « - a es a ~ j . : not Denial 4 . — at¢ vv = : : ei = : On ee i i) = oe i= 7 ‘ 1s a ; i ey oF RM \ a 1 a a 2 “1 t ee b-—P % s fey 1 Gok 7 Aye BO Galo | Cue iabege oi Fe Clo 9 624i be Caape Bull. nat. Hist. Mus. Lond. (Geol.) 57(2): 95-107 Issued 29 November 2001 Gough’s Cave 1 (Somerset, England): a study of the pectoral girdle and upper limbs STEVEN E. CHURCHILL Department of Biological Anthropology and Anatomy, Duke University, Durham NC 27708, USA SYNOPSIS. The pectoral girdles, arms and forearms of Cheddar Man (Gough’s Cave 1) are well preserved and, with the exception of a missing left scapula, are completely represented. These remains, which derive from early Holocene deposits in Gough’s Cave, are described here. Comparative evaluation of the Gough’s Cave | remains reveals that his upper limb skeleton was somewhat gracile, but certainly within the range of variation in strength measures of his contemporaries. INTRODUCTION The upper appendicular skeleton of Cheddar Man is represented by both claviculae, the right-side scapula, the humeri, ulnae and radii of both sides, and elements of both hands. The preservation of the upper limb elements is generally good. The elements of the pectoral girdles, arms and forearms are described below (the manual remains are described elsewhere), followed by a discussion of the overall morphology of the upper limbs. MATERIAL The description of the Gough’s Cave 1 upper limb remains includes both traditional osteometric (Tables 1, 2, 4-7, 9, 10, 12 and 13) and diaphyseal cross-sectional data (Tables 3, 8, and 11). Mean osteo- metric and cross-sectional data are provided for other European Mesolithic and later Upper Paleolithic specimens to provide a com- parative framework for evaluating the morphology of Gough’s Cave 1. These additional specimens range in geological age from the terminal Pleistocene to the mid-Holocene, and include Arene Candide 2 to 5, 10, and 12 to 15, Bruniquel 24, Cap-Blanc 1, Chancelade 1, Farincourt 1, Gramat 1, Hoédic 1, 2, and 4 to 10, Neuessing 2, Oberkassel 1 and 2, Rochereil 1, Romito 3 and 4, Romanelli 1, St. Germain-la-Riviére 4, Téviec 1, 7 to 9, 11 and 16, and Veyrier 1 (Stasi & Regalia, 1904; Verworn et al., 1919; Grazzini, 1921; Péquart & Péquart, 1934; Bonin, 1935; Boule & Vallois, 1937, 1946; Vallois, 1941-1946, 1972; Lacam et al., 1944; Pittard & Sauter, 1945; Sauter, 1957; Graziosi, 1962; Genet-Varcin & Miquel, 1967; Billy, 1969; Gieseler, 1977; Paoli et al., 1980). All comparative data were collected by the author on original specimens. Diaphyseal cross-sections of the claviculae, humeri and ulnae were reconstructed from radiographs and external contour molds for the midshaft (claviculae and humeri) or mid-proximal (ulnae) dia- physes. Subperiosteal contour molds were taken perpendicular to the diaphyseal axis, using dental putty molds (Cuttersil Putty Plus: Heraeus Kulzer Inc.), at 50% (midshaft) or 65% (mid-proximal) of biomechanical length (measured from the distal end). The molds were photostatically reproduced on paper to provide the subperio- steal (outside) contour of the cross-section. In the case of claviculae, ventral, dorsal, superior and inferior cortical dimensions were meas- ured from superoinferior and dorsoventral radiographs. For humeri and ulnae, anterior, posterior, medial and lateral cortical thicknesses were measured from mediolateral and anteroposterior radiographs. © The Natural History Museum, 2001 Subperiosteal dimensions from the original specimens were com- pared with those from the radiographs to determine the degree of parallax distortion and thus allow for algebraic correction of cortical thickness measurements. The cortical dimensions were used along with the subperiosteal contour to interpolate the endosteal contour. The resultant cross-sections were manually digitized and geometric properties were computed using a PC-DOS version (Eschman, 1990) of SLICE (Nagurka & Hayes, 1970). SLICE calculates the total subperiosteal (TA) and cortical (CA) areas, 2nd moments of area about the superoinferior (clavicle) or anteroposterior (humerus and ulna) (I,) and dorsoventral (clavicle) or mediolateral (humerus and ulna) (I) axes, and the maximum (I). and minimum (I) 2nd moments of area. Geometric analysis of cross-sections provides measures of the contribution of bone geom- etry to the resistance of biomechanical loads: in the case of cortical area, to axial compressive and tensile loads; for 2nd moments of area, to bending loads. Medullary area (MA) can be determined from total and cortical areas (MA = TA — CA). The polar moment of area (J, or L.) is a measure of torsional rigidity and overall strength, and can be determined as the sum of any two perpendicular 2nd moments of area (J =[I, + I] = Ha. + Tnind): Since cross-sectional measures of strength co-vary positively with body size, humeral and ulnar cross- sectional values were standardized by powers of bone length to produce measures of relative strength, or robusticity. Following the rationale described in Churchill (1994), measures of cross-sectional area (TA, CA and MA) were standardized as: PUREE seared = (AREA, andardized! SL’) * 10° where AL = humeral or ulnar articular length. 2nd (and polar) moments of area (I, I, Tax? Amin? 1) Were similarly standardized as: 2nd Moment, ,. jardizea = (20d Moment, sardizeg/ AL”) * 10°. In addition to the measures of bone rigidity outlined above, three cross-sectional shape indices were computed to better illustrate the morphology of the Gough’s Cave 1 upper limb material. Percent cortical area (%CA = 100*CA/TA) serves as a simple measure of the degree of cortical occlusion of the medullary space. Ratios of 2nd moments of area provide information about diaphyseal shape (at the location of the cross section) with respect to anatomical axes (1/1) or with respect to the axis of maximum bending rigidity (I, , /I,, ). max min/* CLAVICULAR REMAINS Both claviculae are preserved and are virtually complete (Figs 1, 2). Both lack the sternal epiphysis but have well preserved metaphyseal plates, making it clear that the sternal secondary centres of ossifica- tion were unfused. The right clavicle is damaged on the inferior and 96 2a S.E. CHURCHILL Figs 1,2 Gough’s Cave | claviculae, in superior (a), ventral (b), inferior (c) and dorsal (d) views; 1, right-side clavicle; 2, left-side clavicle; x 0.66. dorsal aspects of the shaft around the costoclavicular ligament attachment area, and is patched with plaster in this location. The right-side bone also suffers from erosion and damage over the superior, dorsal and inferior surfaces of the acromial process. The left-side clavicle is undamaged. Both claviculae are fairly short and stout and not markedly curved. The acromial ends are short and more curved than the proximal shafts, which have only a slight curvature. The sternal end of the right clavicle is round in medial view (with a maximum dorsoventral diameter of 20.9mm, and a maximum superoinferior diameter of 23.6mm), while the left is a dorsoventrally narrow oval (maximum DV diameter = 17.9mm, maximum SI diameter = 27.3mm). The right clavicle exhibits a greater degree of torsion in the shaft than the left clavicle. With the acromial processes held horizontally, in me- dial view the long axis of the articular surface (metaphyseal plate) of the right-side sternal end is inclined at a 45° angle (superoventral to inferodorsal) to the coronal plane. In the left clavicle, the long axis of the sternal articulation is also oriented superoventrally to inferodorsally, but it forms only a 15° angle with the coronal plane. The overall size and shaft curvatures is largely symmetrical between the sides (Table 1). The medial shafts are elongated superodorsal to inferoventral (7.e., the long axes of the shaft cross-sections run in that direction). The costoclavicular ligament attachment area (on the left clavicle, this area is obscured by plaster on the right clavicle) is a dorso- ventrally narrow but mediolaterally long scar. The left-side costoclavicular ligament insertion area is dorsally placed, so much so that it is almost positioned on the dorsal surface of the shaft. On both sides the anterior medial shafts (in the vicinity of the M. pectoralis major attachment area) are flattened and are mostly smooth (there is some mild rugosity along the inferomedial edge of Table 1 Dimensions (mm) of the Gough’s Cave | claviculae. Right Left Maximum length (M-1)* (135) 135.9 Articular length*” 128.3 132.2 Conoid length** 102.9 104.4 Midshaft maximum diameter* 13.9 13.9 Midshaft minimum diameter® 9.6 9.8 Midshaft circumference (M-6) 40 39 Mid-proximal superoinferior diameter® 14.6 11.4 Mid-proximal anteroposterior diameter® 12.3 12.6 Mid-proximal circumference* 42 40 Proximal epiphyseal superoinferior diameter’ 23.3 26.4 Proximal epiphyseal anteroposterior diameter' D2 18.2 Costal impression mediolateral diameter® = il8).5) Costal impression dorsoventral diameter® - 2)S) Conoid superoinferior diameter® PF 13.5 Conoid anteroposterior diameter” Neg 17.8 Acromial superoinferior diameter’ 11.5 11.7 Acromial anteroposterior diameter’ - 2ACS Martin numbers (M-#: Martin, 1928) for measurements are provided where appropriate. * Jength lacking the unfused sternal epiphysis. » direct distance between the mid-points of the proximal and distal epiphyses. © direct distance from the mid-point of the proximal epiphysis to the middle of the conoid tubercle. “ midshaft determined relative to articular length. © taken at mid-conoid length. "maximum (SI-superoinferior) and minimum (AP-anteroposterior) diameters of the proximal epiphysis. ® mediolateral and dorsoventral diameters of the costoclavicular ligament attachment area. " taken at the conoid tubercle perpendicular (SI) and parallel (AP) to the superior surface of the bone. ‘acromial diameters taken perpendicular (SI) and parallel (AP) to the superior surface of the bone. GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS Table 2 Comparative clavicular osteometrics (mean, SD, n). 97 Gough’s Cave | LUP/Meso 3 LUP/Meso 2 Right claviculae Maximum length (135) Articular length 128.3 Conoid length 102.9 Mid-proximal SI diameter 14.6 Mid-proximal AP diameter 12.3 Conoid SI diameter 27) Conoid AP diameter INTe7/ Left claviculae Maximum length 135.9 Articular length 132.2 Conoid length 104.4 Mid-proximal SI diameter 11.4 Mid-proximal AP diameter 12.6 Conoid SI diameter es) Conoid AP diameter 17.8 All measurements are in mm and are defined in Table 1. Table 3 Mid-shaft clavicular cross-sectional properties (mean, SD). Gough’s Cave | Right claviculae Total area (TA) (mm?) 124.9 Cortical area (CA) (mm?) 87.3 Medullary area (MA) (mm?) 37.6 SI 2nd moment of area (I,) (mm‘*) 1319.8 DV 2nd moment of area (I) (mm‘*) 1154.2 Maximum 2nd moment of area (I...) (mm”) S21 Minimum 2nd moment of area (I) (mm*) 721.9 Polar moment of area (J) (mm‘*) 2474.0 Percent cortical area (%oCA) 69.9 1/1, 1.14 max min 2.43 Left claviculae Total area (TA) (mm?) 118.9 Cortical area (CA) (mm?) 73.9 Medullary area (MA) (mm?) 45.0 SI 2nd moment of area (I,) (mm’) 999.6 DV 2nd moment of area (I_) (mm*) 1084.4 Maximum 2nd moment of area (I,,,.) (mm*) 1505.1 Minimum 2nd moment of area (I,,) (mm*) 578.9 Polar moment of area (J) (mm*) 2084.0 Percent cortical area (%CA) 62.2 1, 0.92 1 Me 2.60 143.4, 6.4, 8 129 ANDES 138.2, 6.7, 8 126.3, 3.6, 5 MOM 210 100.4, 7.6, 7 11.8, 1.7, 10 9.8, 0.5, 7 11.7, 0.8, 10 10.3, 0.9, 7 12.6, 3.7, 10 9.8, 1.4, 10 GLI, LS), 0) 15:2 eS10 145.4, 9.0, 10 128.0, —, 2 140.6, 9.1, 10 125.6, —, 2 109.6, 9.8, 13 98.8, 5.4, 4 11.8, 2.1, 16 10.0, 0.8, 6 11.7, 0.8, 16 9.8, 1.3, 6 iO), 7, 1S) 10.0, 1.1, 8 Na SO}, 15) VA eek 7 LUP/Meso 3 LUP/Meso 2 (n=4) (n=2) 107.4, 8.0 68.8 84.0, 7.8 57.0 WZ WA 11.8 869.5, 198.1 450.9 884.1, 96.2 305.4 1001.2, 131.3 473.0 752.5, 139.5 283.3 1753.7, 264.2 756.3 78.3, 6.1 82.9 0.981, 0.185 1.487 1.343, 0.113 1.690 (n=4) (n=3) 122.0, 24.5 Sis, 179 91.2, 23.9 64.1, 13.1 30.8, 3.7 73.5). 1316.6, 671.8 5359:3, 215.9 1016.9, 325.7 498.9, 192.0 1398.1, 626.5 591.6, 177.9 935.5, 364.1 466.7, 232.5 2333.5, 990.4 1058.3, 405.5 VE DET 79.0, 2.2 1.240, 0.307 1.127, 0.081 1.468, 0.099 1.433, 0.517 See text for definition of measurements. the M. pectoralis major origin area on the right-side bone; on the left-side there is some mild rugosity along the entire superior margin of the attachment area for this muscle). On the left clavicle there is a clearly defined insertion area for M. sternocleidomastoideus, beginning with a small but well defined projecting crest medially. This crest continues laterally as a nar- row, mildly rugose strip extending about 25mm along the superior edge of the shaft. The right-side clavicle lacks a clearly marked insertion for M. sternocleidomastoideus, but instead has some poorly defined, mild rugosity in this area. On both sides, the M. subclavius attachment falls on a narrow ridge medially (between the M. pectoralis major origin area ventrally and the dorsal sur- face of the shaft dorsally) but broadens out as it nears the conoid tubercle on the proximal part of the acromial process. Here there is some mild rugosity (on the right, the left-side is largely non- rugose) and the attachment area is defined dorsally by a ridge running to and joining the conoid tubercle. There is also a slight ridge, more prominent in the right clavicle, separating the M. subclavius origin area ventrally from the M. deltoideus attachment area. The area of origin of M. deltoideus is weathered and dam- aged on the right-side, but on the left it can be seen as a well defined, rugose and projecting crest. The conoid tubercles are large and project both inferiorly and dorsally. Both have long ridges extending medially from the tuber- cles (these ridges form the dorsal margins of the M. subclavius attachment areas). The trapezoid line on the left clavicle (this area is damaged on the right clavicle) is not overly rugose but is clearly visible. The acromial facets are poorly defined. The M. trapezius attachment areas are rugose on both acromial processes. The Gough’s Cave | claviculae are compared to those of late Upper Paleolithic and Mesolithic (LUP/Meso) males and females in Tables 2 and 3. Fig. 3 Gough's Cave | right scapula, in ventral (a), lateral (b), dorsal (c) and superior (d) views; x 0.8. GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS SCAPULAR REMAINS Only the right-side scapula is preserved (Fig. 3). The bone is largely complete, lacking only the superior angle (including some of the body on the M. supraspinatus surface) and vertebral border (and the adjacent 1-1.5cm of the body) from just distal of the root of the spine to the inferior angle. Scapular metrics are provided in Tables 4 and 5. All of the observable secondary centres of ossification are fully fused and the epiphyseal lines are obliterated. These include the subcoracoid centre, the infraglenoid centre, the acromial centre, and the vertebral border centre (at least at the root of the spine — the only place this centre can be evaluated). It is possible that the vertebral border — inferior angle centre of ossification was not fully fused along its entire length, and that the preserved portion of the inferior angle represents an epiphyseal surface. Reconstructive materials obscure observation of the inferior angle, making evaluation of the state of fusion of the growth centre difficult. The glenoid fossa is piriform (pear shaped) and has a very strong dorsal orientation. The dorsal margin of the articular surface rounds off onto the base along the dorsal edge, especially on the dorsoinferior margin. The articular surface of the glenoid fossa is irregular but there does not appear to be a central pit (McCown & Keith, 1939) (again, reconstructive materials obscure the articular morphology). The attachment for the glenoid labrum can be made out along the ventral margin of the articular surface, but disappears along the dorsal edge where the articular surface rounds off onto the base. The infraglenoid tubercle begins as a very large attachment area on the dorsoinferior edge and dorsal part of the inferior edge of the glenoid. The tubercle is very large and elongated superoinferiorly (extending 32.6mm below the inferior margin of the glenoid fossa). The surface of the infraglenoid tubercle itself is finely rugose. The axillary crest (the attachment site for the fascia separating M. sub- scapularis and M. teres minor) is blunt and rounded, and runs along the dorsal portion of the axillary border for most of its length (Fig. 3b). At the distal end of the inferior M. teres minor attachment area the axillary crest veers dorsally and forms a rather strong crest on the dorsolateral edge of the axillary border. This morphology Table 4 Dimensions (mm) of the Gough’s Cave | right scapula. Morphological length (M-2) 100.0 Infraspinatus breadth (M-5a) 124.0 Basal spinous length (M-8) 79.4 Spino-acromial length* 130.0 Spinal thickness? 1-5) Acromio-glenoid distance® 45.4 Axillary border length (M-3) 126.6 Functional axillary border length* 139.0 Mid-axillary border thickness* 12.5 Spino-glenoid angle (M-22) 86° Axillo-glenoid angle (M-17) Bhs) Axillo-spinal angle (M-16) 60° Glenoid maximum length (M-12) 38.0 Glenoid maximum breadth (M-13) 26.1 Glenoid articular length‘ 34.7 Glenoid articular breadth’ 24.8 Martin numbers (M=#: Martin, 1928) for measurements are provided where appropriate. “vertebral border at the spine to the tip of the acromion. > maximum superoinferior thickness along the middle of the spine. “center of glenoid fossa to the tip of the acromion. ‘center of glenoid fossa to the most caudal point on the inferior angle. © dorsoventral diameter of the mid-axillary border, including dorsal and ventral pillars as present. ‘ glenoid fossa length and breadth taken across the internal margins of the glenoid labrum attachment. 99 suggests that the M. subscapularis extended dorsally around the axillary border and had some fibres attached to the dorsal side of the lateral edge of the scapula at this level. The dorsal pillar (see Churchill, 1994) is laterally placed (running along the lateral-most edge of the axillary border) and well developed, and forms a deep sulcus on the body just medial of the pillar. The ventral pillar is medially positioned and is very strong, leaving a long, wide and shallow sulcus on the ventral aspect of the axillary border. Gough’s Cave 1|’s scapular axillary border thus presents a ventral sulcus morphology similar to that seen in high frequencies in recent sam- ples of European (and European-descent) populations (Trinkaus, 1977; Frayer, 1992; Churchill, 1996). On the dorsal aspect of the axillary border, there is what appears to be a vascular groove separat- ing the infraglenoid tubercle and the superior portion of the attachment area for M. teres minor. This groove may indicate an accessory artery in the scapular anastomotic complex, one arising either from the root of the subscapular artery or perhaps stemming directly from the axillary artery. The groove for the circumflex scapular artery can be seen Clearly dividing the M. teres minor origin further distally on the border. The M. teres minor attachment areas are poorly defined, the superior attachment area is not flattened and neither area is rugose. There is a small lateral flange for M. teres major and some flattening of the distal end of the axillary border. There is little-to-no curvature of the body of the scapula. With the scapula held in anatomical position, the lateral edge of the acromion is broad dorsoventrally (measuring 49mm). The lateral edge of the acromion bears a rugose strip for the attachment of M. deltoideus along its entire length. The superior/dorsal surface of the spine is sinuous. The M. trapezius insertion area is not rugose. On the coracoid process, the lateral edge bears two small depressions with small crests for the M. coracobrachialis and M. biceps brachii short head attachments. The scapular notch is relatively broad. HUMERAL REMAINS Both humeri are virtually complete (Figs 4, 5). The right-side has some damage to the dorsal surface of the head and this area is filled in with plaster. The left-side has some erosion to the medial surface of the head and what appears to be an erosional defect on the superoventral surface of the trochlea (see below). Otherwise, the two bones are very well preserved. There is a very slight trace of an epiphyseal line on the anterior and medial surfaces of the proximal metaphysis just below the lesser tubercle and the articular surface of the head. The line is more apparent on the right humerus than on the left. On both humeri the line is largely obliterated on the dorsal and lateral surfaces. Even though the line is visible, the head 1s fully fused and the lines are near obliteration. Distally, the distal epiphyses and the medial epicondyle secondary centres of ossification are fully fused and the lines oblit- erated on both humeri. The two sides are largely symmetrical with respect to size, robusticity and most details of morphology (Table 6). Although both humeri are similar in size and rugosity of muscle markings, the left- side seems to be the generally more muscle marked of the two. The humeri are not markedly robust. There is no notable curvature to the diaphyses in any plane. The deltoid tuberosities are neither large nor projecting, such that the diaphyses are of uniform dimension (save for a slight swelling at the deltoid tuberosity) along the shaft from every perspective. The head of the right humerus is dorsomedially directed (producing a torsion angle of 141°), while the left has a more medially oriented head (torsion angle = 166°). 100 4a 5a 4b 5b S.E. CHURCHILL 4c 5¢ 4d Sd Figs 4,5 Gough’s cave | humeri in ventral (a), medial (b), dorsal (c) and lateral (d) views; 4, right-side humerus; 5, left-side humerus; x 0.37. Table 5 Comparative scapular dimensions (mean, SD, n) Gough’s Cave | LUP/Meso 2 (right + left) LUP/Meso 3 (right) (right + left) Morphological length 100.0 139.4, 34.6, 4 - Axillary border length 126.6 IPAS)-2), 3).6), ©) 123.0,—, 2 Glenoid articular length 34.7 35) 1/ 2d) €83 Bye, MoT Glenoid articular breadth 24.8 26.6, 1.7, 6 23.1, 1.4, 7 Glenoid index* 71.5 Wek 2) 72.1, 4.1, 7 All measurements are in millimeters and are defined in Table 4. *Glenoid index = (articular breadth/articular length) x 100. The articular surfaces of the heads are fairly round, the left-side seems to be somewhat more oval shaped (with the long axis running superoinferiorly) than the right. The articular surfaces are smooth (save for erosional damage) and free of any signs of degeneration. The right humerus has some erosion over the lateral surface of the greater tubercle, and the only muscle attachment area not eroded is the ‘facet’ for the insertion of M. teres minor. On the left-side the greater tubercle is well preserved, revealing a distinct ‘facet’ for M. teres minor and amore diffuse attachment area for M. supraspinatus and M. infraspinatus. The attachment area for these muscles is smooth and there is no clear separation between the facets for the two muscles. Both sides have pronounced intertubercular sulci formed medially by superoinferior running pillars extending from the lesser tuberosities along the insertion areas for M. latissimus dorsi and M. teres major, and laterally by pronounced crests for M. pectoralis major. The pillars are non-rugose (except at the muscle attachment areas) and the M. pectoralis major scars are only mildly rugose. The deltoid tuberosities are not pronounced, but appear instead as slightly projecting, rugose crests. Indentations can be seen on both sides at the intersection of the proximal deltoid tuberosity with the inferior portion of the attachment of the pectoralis major muscle (the indentation is sightly more pronounced on the right-side). The indentation is best seen in medial or lateral view (Figs 4b, 4d, Sb, 5d). On the right-side there is a very slight musculoskeletal stress marker GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS Table 6 Dimensions (mm) of the Gough’s Cave | humeri. Right Left Maximum length (M-1) 3 322 Articular length (M-2) 316 318 Midshaft maximum diameter (M-5) 22.2 21.9 Midshaft minimum diameter (M-6) 17.0 15.2 Midshaft circumference (M-7a) 65 63 Distal minimum circumference (M-7) 62 61 Head anteroposterior diameter (M-10) (40.2) 40.1 Epicondylar breadth (M-4) 58.0 S75 Distal articular breadth (M-12a) 43.0 43.0 Capitular breadth (M-12) 21.0 19.0 Trochlear breadth (M-11) 20.2 21.1 Trochlear max. anteroposterior diam.(M-13) 24.9 23.9 Medial epicondyle projection* 18.3 16.7 Lateral epicondyle projection? 19.7 20.1 Olecranon fossa breadth (M-14) 23.8 23.9 Olecranon fossa depth (M-15) 12.2 12.2 M. pectoralis major sulcus breadth* 39 5.6 Deltoid tuberosity width? 11.5 13.3 Deltoid tuberosity circumference* 68 64 Cubital angle (M-16) inige 101° Torsion angle (M-18) 141° 166° Martin numbers (M-#: Martin, 1928) for measurements are provided where appropriate. * medial trochlear margin to the most projecting portion of the epicondyle. » Jateral trochlear margin to the most projecting tip of the epicondyle. © maximum breadth (mediolateral) of the middle of the muscle insertion area. 4 distance between the apices of the delimiting crests of the tuberosity taken at 5/12’s of humeral maximum length, following Endo (1971). © shaft circumference taken at 5/12’s of humeral maximum length, following Endo (1971). (enthesopathy’: Dutour, 1986; Hawkey & Merbs, 1995) in the anterior crest of the deltoid just inferior of this depression. A similar but smaller projection can be seen at the same location on the left- side. The deltoid tuberosities are narrow in anterolateral view and are two-crested on both humeri. The posterior crest on the left humerus Table 7 Comparative humeral osteometrics (mean, SD, n). Gough’s Cave 1 Right humeri Maximum length* 322 Midshaft maximum diameter 22.2 Midshaft minimum diameter 17.0 Distal minimum circumference 62 Head anteroposterior diameter (40.2) Epicondylar breadth 58.0 Distal articular breadth 43.0 Deltoid tuberosity width ils) Deltoid tuberosity circumference 68 Left humeri Maximum length* 322 Midshaft maximum diameter 21.9 Midshaft minimum diameter 2) Distal minimum circumference 61 Head anteroposterior diameter 40.1 Epicondylar breadth 57.5 Distal articular breadth 43.0 Deltoid tuberosity width 13.3 Deltoid tuberosity circumference 64 101 is more rugose than the posterior crest on the right humerus (the right-side crest is very mildly rugose). The other muscle attachment areas on the shaft are non-rugose (including M. brachialis, and the medial and lateral heads of M. triceps brachii). There is some slight rugosity along the M. coraco- brachialis insertions on both humeri. Distally, the medial epicondyles are very large in superoinferior dimension but do not project very far mediolaterally, giving them a squat appearance. Both medial epicondyles present smooth but irregular surfaces where the common flexor tendon attaches. On the left-side there is a distinct crest on the anterosuperior edge of the attachment area, between the medial epicondyle proper and the shaft of the humerus. No such crest is apparent on the right-side. The lateral supracondylar ridges also seem well developed. The common extensor tendon origins are represented by small, smooth areas, and the M. anconeus origins are represented on both sides by smooth, thin semi-circular ridges that do not join the common extensor attachment areas. The distal articular surfaces are relatively small, and have deep medial and lateral grooves. The lateral grooves (separating the lateral trochlear margin from the capitulum) are very wide and distinct. The capitullae are very small superoinferiorly and are slightly indented on their inferolateral surfaces, giving them a slightly laterally pointed appearance in anterior view. The left trochlea has an oval eroded patch (measuring 13mm mediolaterally by 9.5mm superoinferiorly) on its anterosuperior surface. This appears to be post-mortem in nature, and there is no corresponding defect on the left ulna, ruling out osteoarthritic eburnation. No other signs of degeneration are present on the distal articulations. From the dorsal perspective, the distal medial pillars are relatively broad. The lateral pillars are also broad, and the surfaces behind the capitullae are flat and at angles of ca. 145° to the dorsal surfaces of the lateral pillars. The olecranon fossae are relatively small but deep. Comparative data on other late Pleistocene/early Holocene human humeri are provided in Tables 7 and 8. LUP/Meso 3 LUP/Meso 2 SOS WAL I7/ 287.0, 13.7, 9 23.9, 1.8, 17 207 ess 10, 77, U40), 19/ 15.4 1.6, 10 64.1, 4.1, 18 Sp ysh nS) 43.6, 2.8, 13 Mit okey I 61.9, 2.2, 16 56.4, 2.9, 4 42.8, 2.4, 16 39.2 4.3,7 GP Pets iF) Shh, As il 71.5, 3.9, 16 62.5, 4.4, 11 S842 12) 287.4, 7.5, 10 23.4, 2.5, 13 20.2, 1.2,9 17.0, 2.0, 14 15.8, 1.0, 9 625 Seine 55.9, 2.6, 8 43.1, 1.6, 14 SR A)5 Was)3 3) Colles Sisile iLI 54.7, 1.8; 7 43.2,2.1, 10 Shh) Poh 7 16.9, 3.4, 13 13.6, 2.3, 10 69.5, 6.7, 13 59.7, 3.8, 10 All measurements are in millimeters and are defined in Table 6. “includes estimated lengths for some comparative specimens 102 Table 8 Mid-shaft humeral cross-sectional properties, standardized* (mean, SD). Right humeri Total area (TA) Cortical area (CA) Medullary area (MA) ML 2nd moment of area (1,) AP 2nd moment of area (I) Maximum 2nd moment of area (i) Minimum 2nd moment of area (Lain) Polar moment of area (J) Percent cortical area (%oCA) I/I max min Left humeri Total area (TA) Cortical area (CA) Medullary area (MA) ML 2nd moment of area (I,) DV 2nd moment of area (I,) Maximum 2nd moment of area (I...) Minimum 2nd moment of area (I...) Polar moment of area (J) Percent cortical area (%CA) l/l ty at max min See text for definition of measurements. S.E. CHURCHILL Gough’s Cave | LUP/Meso ¢ LUP/Meso 2 (n=17) (n=10) 307.3 363.2, 59.4 314.7, 36.0 DIN 260.8, 52.1 224.4, 43.2 86.2 102.4, 32.5 90.2, 26.2 682.6 955.0, 296.5 USDA WOT 783.9 1104.0, 404.6 750.1, 207.5 952.4 1241.1, 487.1 1048.9, 292.4 514.1 718.5, 232.8 SSA IS3i7 1466.5 2059.1, 684.7 1539.8, 384.5 71.9 71.7, 8.0 70.2, 8.3 0.871 0.893, 0.132 1.079, 0.168 1.853 1.712, 0.313 1.901, 0.426 (n=12) (n=9) Piel 350.3, 59.9 2977.1, 32.0 180.2 253.4, 53.6 207.0, 32.6 90.9 96.8, 29.3 9077733:9 452.3 832.2, 251.9 659.2, 145.2 649.0 1106.7, 423.3 662.0, 152.4 723.9 1313.3, 412.9 811.6, 187.8 377.4 730.0, 246.7 473.5, 91.8 1101.3 1938.8, 659.3 1321.2, 275.9 66.5 122.13 71.2, 10.7 0.697 0.803, 0.201 1.006, 0.184 1.918 1.837, 0.318 1.695, 0.136 * Cross-sectional strength measures standardized to humeral articular length (HAL). Areal measures (TA, CA and MA) were divided by HAL’? and multiplied by 10°. 2nd moments of area (I,1,1. ,I x? Ty? “min? “max Table 9 Dimensions (mm) of the Gough’s Cave 1 ulnae. Measurement , J) were divided by HAL* and multiplied by 10°. Maximum length (M-1) Articular length (M-2) Olecranon length (M-8) Olecranon height (M-7) Olecranon breadth (M-6) Trochlear notch chord (M-7(1)) Coronoid height* Radial facet maximum diameter? Radial facet minimum diameter? Coronal trochlear angle (M-15) Diaphyseal sagittal trochlear angle (M-15a) Proximal anteroposterior diameter‘ Proximal transverse diameter® Proximal circumference‘ Crest anteroposterior diameter (M-11) Crest mediolateral diameter (M-12) Midshaft anteroposterior diameter Midshaft mediolateral diameter Midshaft circumference Distal minimum circumference (M-3) Pronator quadratus crest maximum diameter® Pronator quadratus crest minimum diameter* Pronator quadratus circumference? Head breadth* Right Left = (270) z 250 14.8 = FG = 24.0 = 23.5 = 36.3 34.9 19.9 18.3 10.9 10.6 14° es Ble = 18.3 18.4 18.8 18.7 51 49 15.3 13.7 14.3 15.0 12.2 ye) = G9) - 42 37 2 12.8 = 10.6 2 38 = 16.9 17.0 Martin numbers (M-#: Martin, 1928) for measurements are provided where appropriate. “maximum anteroposterior diameter from the dorsal surface of the bone to the anterior tip of the coronoid process (McHenry et al., 1976). ® maximum and minimum diameters of the articular facet for the radial head. “ taken at the level of the distal border of the ulnar tuberosity (McHenry er al., 1976). ® shaft maximum and minimum diameters and circumference at the most projecting portion of the pronator quadratus crest. “mediolateral diameter of the ulnar head. ULNAR REMAINS Both ulnae are largely complete (Figs 6, 7, Table 9). The right-side is somewhat better preserved than the left, but is missing a 4cm portion of the diaphysis from about midshaft distally. This area has been reconstructed in plaster. With the exception of the missing diaphy- seal fragment, the bone is complete. The left-side is missing the anterior half of the olecranon process (from the anterior edge of the M. triceps brachii insertion), including some of the olecranon articu- lar surface in the trochlear notch. There is also some damage to the surface of the left-side distal shaft in the region of the M. pronator quadratus crest. Otherwise, the bone is very well preserved. The proximal and distal epiphyses are fully fused. The left-side ulna has a closed but still (barely) visible epiphyseal line between the head and shaft. All of the other epiphyseal lines are fully obliterated. Both ulnae are of moderate robusticity (Tables 10 and 11). The right-side shaft has a lateral curve to the distal end (as seen in volar view), beginning in the area around the M. pronator quadratus crest. The left-side diaphysis is straight, but has had some damage to and refitting of the distal end, perhaps affecting the appearance of distal shaft. The shafts are also straight in lateral view. The trochlear notch on the right-side (the left is damaged) opens anteroproximally (Fig. 8). Neither side has a very marked separation (as either a crest or sulcus) between the coronoid and olecranon articular surfaces in the notch, nor does either side express a sagit- tally oriented crest in the notch. There is no evidence of degenerative changes to the trochlear or radial notches of either side. The radial notches are relatively large, and are oriented superolaterally. The M. triceps brachii insertion on the olecranon process is mildly rugose (on the right ulna, this region can’t be fully evaluated in the left), with vertical striations on the posterior edge of the olecranon but no spurring or indications of musculoskeletal stress lesions. The GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS 6b 7b 103 Figs 6,7 Gough’s Cave | ulnae, in ventral (a), medial (b), dorsal (c) and lateral (d) views; 6, right-side ulna; 7, left-side ulna; x 0.5. Fig.8 Gough’s Cave | right ulna in lateral view; x 1. M. anconeus insertion area is mildly rugose, occupying a narrow ridge (between the medial and lateral surfaces of the proximal shaft) on the dorsal surface of the shaft. This mildly rugose line continues distally until it joins with the line marking the aponeurotic attach- ment of M. flexor digitorum profundus, M. flexor carpi ulnaris and M. extensor carpi ulnaris. Supinator crests of both sides are well developed, beginning as pronounced crests arising from the inferodistal margins of the radial notches (Fig. 8). The crests con- tinue distally to a half centimeter or so below the M. brachialis scar, where they stop and then resume again another 10—20mm distally. The distal portions of the M. supinator ridges follow the interosseous crests (lying just posterior to and on the lateral surface of the interosseous crests), about one-third of the way down the shaft. The posterior edge of the proximal portion of the M. supinator attach- ment area is also marked by a crest on the lateral surface below the trochlea. The M. brachialis scars are moderately rugose, and present themselves as small raised patches (roughly 13mm proximodistally x 9mm mediolaterally on both sides). The area between the scars and the coronoid processes is mildly rugose. No markings are evident on either side for M. pronator teres. There is a large tubercle present on the anteromedial corner of the coronoid process in the area of the origin of the ulnar head of M. flexor digitorum superficialis on the right-side. This area is somewhat damaged in the left-side ulna, but it is certain that the tubercle, if indeed it existed, was not nearly as large as that of the right. 104 S.E. CHURCHILL Table 10 Comparative ulnar osteometrics (mean, SD, n). Gough’s Cave | LUP/Meso 3 LUP/Meso 2 Right ulnae Olecranon length 14.8 W/siL, 20, 115) 16.7, 2.0, 4 Olecranon height 27.7 Pel, We), 13} 23.8, 1.4, 4 Olecranon breadth 24.0 Pte Ms), 23.6, 1.1, 4 Trochlear notch chord 23-5) 27.0, 3.6, 10 DPA Pla 3) Coronoid height 36.3 37.4, 1.6, 12 SYN PM Radial facet maximum diam 19.9 16.8, 1.8, 9 16.4, —, 2 Radial facet minimum diam 10.9 11.6, 0.6, 9 11.2,-,2 Proximal AP diameter 18.3 IS\.7/, Woy, 13) 22-2085 Proximal transverse diam 18.8 16.8, 2.0, 13 EIR IEE S Proximal circumference Sil 50.5, 3.8, 12 45.0, 2.6, 5 Midshaft AP diameter 12.2 14.6, 1.9, 5 12.8, -, 1 Pron. quadratus max diam 12.8 13.6, 1.4, 11 AE LED Pron. quadratus min diam 10.6 10.3, 0.9, 11 9.6, 0.9, 4 Pron. quadratus circum 38 Bie O), Sh)! tl 35.3, 2.9, 4 Left ulnae UP/Meso 3 UP/Meso 2 Maximum length* (270) 263.9, 10.6, 14 245.8, 8.3, 4 Articular length* 250 241.6, 10.6, 15 224.7, 7.4,7 Coronoid height 34.9 S67/ le 33.3, 1.0, 4 Radial facet maximum diam 18.3 16.8, 1.3, 9 16.4, 1.6, 3 Radial facet minimum diam 10.6 Util, ee) 10.1, 0.4, 4 Proximal AP diameter 18.4 WS), 22, 15.4, 1.6, 8 Proximal transverse diam 18.7 sesh, 770335 10 13.4, 1.7, 8 Proximal circumference 49 48.0, 6.3, 11 44.2, 3.5,7 Midshaft AP diameter WP Se M723) 13.5, 0.6, 4 Midshaft ML diameter 13.2 14.5, 1.5, 3 11.7, 0.5, 4 All measurements are in millimeters and are defined in Table 9. “includes estimated lengths for some comparative specimens. Table 11 Mid-proximal ulnar cross-sectional properties, standardized* (mean, SD). Gough’s Cave 1 LUP/Meso 3 LUP/Meso 2 (n=4) (n=2) Right ulnae Total area (TA) 228.8 266.8, 36.7 192.0 Cortical area (CA) 210.1 PPV) B\3\3) 9a Medullary area (MA) 18.7 44.1, 14.7 14.9 ML 2nd moment of area (1,) 417.9 534.8, 235.8 BO283 AP 2nd moment of area (I) 491.5 655.3, 143.7 278.3 Maximum 2nd moment of area (ee) 536.8 714.5, 184.3 360.9 Minimum 2nd moment of area (I, ,.) By 2a 475.6, 150.0 269.8 Polar moment of area (J) 909.5 1190.1, 325.7 630.7 Percent cortical area (%CA) 91.8 83.5, 5.5 92.5 I/, 0.850 0.814, 0.306 1.269 iy ines 1.440 1.528, 0.203 1.338 Left ulnae (n=3) (n=3) Total area (TA) 221.8 Te VAP 244.0, 19.3 Cortical area (CA) 195.7 242.9, 27.4 221.6, 23.5 Medullary area (MA) 26.1 34.8, 24.5 22.9, 12:6 ML 2nd moment of area (I, ) 3315)57/ 632.9, 187.8 562.9, 86.1 DV 2nd moment of area (I) 487.3 76.1, 141.1 443.4, 120.3 Maximum 2nd moment of area ce 488.9 784.4, 129.1 610.4, 110.3 Minimum 2nd moment of area (1 334.2 524.6, 79.4 395.9, 52.6 Polar moment of area (J) 823.1 1309.1, 179.3 1006.3, 160.4 Percent cortical area (%CA) 88.2 87.5, 8.7 90.8, 5.0 1/1, 0.689 0.975, 0.376 1.319, 0.331 ley 1.463 1.507, 0.267 1.537, 0.124 max min See text for definition of measurements. * Cross-sectional strength measures standardized to ulnar articular length (UAL). Areal measures (TA, CA and MA) were divided by UAL? and multiplied by 10°. 2nd moments of area (I,, 1), 1,,,,. 1...» J) were divided by UAL* and multiplied by 10°. min? ~ mai On both sides, the distal end of the interosseous crest branches into left ulna) is relatively small but is clearly discernable as a narrow, two crests—more strongly so on the right-side but also to some extent on projecting crest. the left. This may reflect a division of the membranes for M. flexor The styloid processes are relatively long. There are no signs of digitorum profundus anteriorly and M. extensor indicus posteriorly. any degenerative changes to the distal articular surfaces on either The right-side M. pronator quadratus crest (this area is damaged in the side. GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS Table 12 Dimensions (mm) of the Gough’s Cave | radii. Measurement Right Left Maximum length (M-1) - 248 Articular length (M-2) - 235 Proximal anteroposterior diameter* 113)-5) 13.1 Proximal mediolateral diameter* 2 12.8 Proximal circumference* 4] 4] Crest anteroposterior diameter (M-5) 12.4 11.7 Crest mediolateral diameter (M-4) lel 15.6 Midshaft anteroposterior diameter (M-5a)° 12.0 11.9 Midshaft mediolateral diameter (M-4a)? 16.9 15.1 Midshaft circumference (M-5(5))? 46 42 Distal minimum circumference (M-3) 4] (41) Head-neck length (M-1a) 33.0 3183 Neck-shaft angle (M-7) 6° 6° Head anteroposterior diameter (M-5(1)) 21.1 22.4 Head mediolateral diameter (M-4(1)) 21.2 21.3 Head circumference (M-5(3)) 69 71 Neck anteroposterior diameter (M-5(2)) 14.7 15.1 Neck mediolateral diameter (M-4(2)) 12.4 14.7 Neck circumference (M-5(4)) 44 46 Bicipital tuberosity lengths 22.4 21.5 Bicipital tuberosity breadth‘ 14.1 14.2 Distal breadth (M-5(6)) - Pf Distal depth* - 15.4 Carpal articular breadth‘ - y-3} Martin numbers (M-#: Martin, 1928) for measurements are provided where appropriate. “taken midway between the bicipital tuberosity and the proximal end of the interosseous crest. » right-side midshaft position was estimated using the more complete left-side radius. © maximum proximodistal diameter of the tuberosity. “ maximum dorsovolar diameter of the tuberosity. “maximum dorsovolar diameter of the distal epiphysis, not including the dorsal (Lister’s) tubercle. ‘ maximum mediolateral diameter of the articular surface. Table 13 Comparative radial osteometrics (mean, SD, n). Gough’s Cave | Right radii Midshaft AP diam 12.0 Midshaft ML diam 16.9 Head-neck length 33.0 Neck-shaft angle 6° Head AP diam 21.1 Head ML diam 21.2 Neck AP diam 14.7 Neck ML diam 12.4 Bicip. Tuberosity length 22.4 Bicip. Tuberosity breadth 14.1 Left radii Maximum length* 248 Articular length* 235 Midshaft AP diam 11.9 Midshaft ML diam 15.1 Head-neck length 31.3 Neck-shaft angle 6° Head AP diam 22.4 Head ML diam PWN3} Neck AP diam 15.1 Neck ML diam 14.7 Bicip. tubersoity length PNNIES) Bicip. tuberosity breadth 14.2 105 RADIAL REMAINS Both radii are preserved (Figs 9, 10). The right radius is missing the distal end from the region of the distal interosseous crest. The diaphysis has also been broken and refitted near midshaft. The left- side is more complete but has some damage to its distal end. The anterior surface of the left distal shaft, from slightly below midshaft to the epiphyseal end, is eroded and damaged, and the anterior surface of the distal metaphyseal area is missing. The proximal halves of both radii are in a good state of preservation. The proximal (right and left) and distal (left) epiphyses are fully fused and the epiphyseal lines are obliterated. No degenerative changes are evident in any of the preserved articular surfaces. The heads of both radii are slightly (but only slightly) oblong, with the long axis running anteromedial to posterolateral. The neck of the right radius is narrower in both anterior and lateral views compared to the left (Table 12). The radial tuberosity and anterior oblique crest also have a slightly greater development on the left-side. The radial tuberosities are proximodistally short (Table 12), miderately wide, but projecting medially (the left more so than the right). The anterior oblique crests extend about 3 cm below the radial tuberosities, and are well developed. The M. supinator insertion areas are smooth. The proximal end of the left radius is generally larger and more robust than the right (in the head, neck and radial tuberosity dimen- sions: Table 12), but the right clearly has a greater development of the interosseous crest and shaft. The shafts are relatively straight (in both anterior and lateral views) and are of moderate robusticity (Table 13). The M. pronator teres insertions are clearly defined and of moderate rugosity. The attachment sites for M. pronator quadratus and M. brachioradialis are too damaged on the left-side (and absent on the right) to say anything about. On the dorsal aspect of the distal left shaft, the dorsal (‘Lister’s’) tubercle is very well developed. LUP/Meso 3 LUP/Meso 2 12.3, 0.4, 4 10.1, 0.9, 3 16.0, 1.4, 4 3}-3}5 Lhe}, 3) 31.8, 2.6, 12 SHO) ils7/55) 8.8°, 3.4,12 Mee S a6 22.7, 1.4, 8 20.8, 1.7, 3 21.8, 1.4, 10 20.6, 1.8, 3 PBL Ses IO) 11.9, 1.4,6 Sesh 1,3}, @) TES ORG PPIs, 2A}, 2) 22S 21657. 42S oll 2212 aah sh, 9 UP/Meso 3 UP/Meso 2 243.9, 7.4, 10 226.8, 8.8, 4 223 nO Ses PIN SP, SES) 11.9, 0.7, 4 10.0, 0.7, 3 14.9, 1.5, 4 12 lkORS Sify, WEA, 1 31.0, 3.3, 4 9.0°, 4.1, 10 WPT F/A0) 5) 22.9, 1.0, 12 TEI / 200), 8) 21.7, 0.6, 8 18.9, 2.3,4 14.0, 1.0, 12 12.4, 0.9, 6 12.3, 3.9, 14 11.4, 1.0, 6 PIS DY SUEY. ANS! 19.4, 1.8,6 14.0, 1.7, 14 Ibe aa OF) i TT All measurements are in millimeters and are defined in Table 12. “includes estimated lengths for some comparative specimens. 9a S.E. CHURCHILL 10c 9d 10d Figs 9,10 Gough’s Cave | radii, in ventral (a), medial (b), dorsal (c) and lateral (d) views; 9, right-side radius (note that the distal end is reconstructed), 10, left-side radius; x 0.5. COMPARATIVE MORPHOLOGY OF THE UPPER LIMB REMAINS In most measures, the Gough’s Cave | upper limb skeleton is comparable to that of other European males alive during the final Pleistocene and early Holocene. In both gross external and cross- sectional dimensions of the humerus, radius and ulna, Gough’s Cave 1 is unremarkable compared to his contemporaries. These bones tend to be slightly longer (on average) than those of his male contemporaries, yet to have epiphyseal dimensions and diaphyseal diameters roughly equivalent to the mean values observed in the comparative sample males (Tables 7, 10 and 13). Thus relative to length, the upper limb long bones of Gough’s Cave 1 are somewhat gracile when seen in the context of other similarly aged fossils. This characterization holds for size-standardized measures of the me- chanical strength of the humerus and ulna as well (Tables 8 and 11), suggesting that the habitual biomechanical loads incurred in the upper limbs of Gough’s Cave 1 were somewhat meager relative to other Upper Paleolithic and Mesolithic foragers. Perhaps the most remarkable feature in the upper limbs (used here in the broad sense to include the pectoral girdles) is the robusticity of Gough’s Cave 1’s right clavicle. In terms of length, the right clavicle falls between the mean values for males and females in the compara- tive sample (Table 2). However, in external dimensions (Table 2) and unstandardized cross-sectional properties (Table 3), Gough’s Cave 1’s right clavicle tends to fall on or above the male sample means. The superoinferior, dorsoventral and maximum 2nd moments of area, as well as the polar moment of area, are two or more standard deviations above the (admittedly small) male sample means (Table 3). Given that Gough’s Cave 1’s claviculae were relatively short, the clavicular strength values are all the more striking. Consideration of the cross-sectional geometric properties presented in Table 3 shows that the large values for bending rigidity (2nd and polar moments of area) in Gough’s Cave 1’s right clavicle are due in part to the overall size of the cross-sections (TA). The right clavicle has a midshaft cortical area (CA) close to the comparative male mean, but this cortical bone is distributed further from the geometric centre of the section. This results ina relatively large medullary cavity, arelatively low percent cortical area, and relatively high 2nd moments of area. GOUGH’S CAVE PECTORAL GIRDLE AND UPPER LIMBS A similar consideration of Gough’s Cave 1’s left clavicle shows it to be relatively gracile, at least in cross-sectional measures of strength (Table 3) if not in external dimensions (Table 2). Bilateral asymme- try in upper limb strength measures is certainly to be expected (see Churchill, 1994), and the degree of asymmetry seen here is not likely to be outside the range observed in fossil and recent, moderately to highly active foragers. What is curious about Gough’s Cave | is the robusticity of the right clavicle in the context of an otherwise relatively gracile upper limb skeleton. Gough’s Cave 1’s right clavicle has an I/I, ratio that is above but within one standard deviation of the male comparative sample mean (yet below the mean of females: Table 3), yet has an I, /I,,. ratio considerably higher than the mean of the comparative samples. The angle formed between the plane of greatest bending rigidity and the dorsoventral plane passing through the centre of the section is 130° in the Gough’s Cave | right clavicle, indicating an adaptation to bending in the superodorsal to inferoventral direction (i.e., about an inferodorsal to superoventral axis). This might suggest an habitual generation of dorsoventrally oriented forces operating on the clavi- cle during full abduction of the humerus (in which scapular rotation on the thorax necessitates conjunct posterior axial rotation of the clavicle, bringing the superodorsal-inferoventral axis of the bone into the transverse plane of the body). While it is the case that the muscle scars for M. deltoideus, the major abductor of the humerus, are rugose on the clavicle (at least on the left side, this region is damaged in the right), scapulae and humeri, it is also the case that the humeral deltoid tuberosities are weakly developed. This, combined with the overall gracility of the other upper limb bones, makes it difficult to imagine a behavior pattern that could differentially load the clavicle more than the other bones of the limb. The asymmetry evident in the claviculae is also not apparent in the other upper limb remains (and in fact the reverse pattern is seen in the radii, with the left being slightly more robust than the right). Actions involving forceful right-side scapular abduction (as in pushing on a heavy object with the arms abducted and flexed, that is, out in front of the body) or adduction (as in forcefully pulling something using motion at the shoulder) would be expected to generate bending moments in the transverse plane of the clavicle. However, such movements would also be expected to create bending stresses in the arm and forearm. Thus if Gough’s Cave | were routinely engaged in an activity requiring forceful pulling or pushing with one limb (perhaps involving woodworking in the manufacture of tools, or perhaps trap setting), we would expect a marked degree of right-left asymmetry in bone strength and muscle scarring in the upper limbs overall. Occupationally induced habitual heavy loading of the bones and joints of the upper limb is known to promote clavicular robusticity. For example, Lane (1887) reported marked robusticity in the lateral half of the clavicle in milkmen that routinely carried heavy milk pails with their arms by their sides. While observations like this support the possibility that the robusticity of the Gouch’s Cave 1 right clavicle reflects habitual labor induced (‘occupational’) loads, the nature of the robusticty (both in terms of bone shape and the observed asymmetry) and the lack of associated degenerative changes to the clavicular joints does not fit with known patterns of occupational changes to upper limb morphology (re- viewed in Kennedy, 1989). Thus the behavior patterns that produced this morphology in Cheddar Man remain, for the time being, un- known. 107 REFERENCES Billy, G. 1969. Le squelette post-cranien de |1’'Homme de Chancelade. L’Anthropologie,73: 207-246. Bonin, G. von 1935. 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A’ Gre T (Aiwasewr Sr — 7 ee < > ~~ - i Poa 4) le eo % - = = - - — ® = =. - = a - o = ai a, 7 — Hie a ie y tah y yi _ im @gue a 4 i 4 i] - aaa hd y Ly m=] 7 : r) a ae — i = 7 = : - a te > 5 : ; . iy -_ 7 a - ; _— 7 ill - — i es mae = - 7 5 7 : £ Paseo ya re: = ie b-ccast M - i ~ 25 ae Ka \i 2 @he ve \ a “ * ¢ . ee ; aan Ee yin oy . a “ee Wi< Das we owe rr in ae, 0 Pre. pe Las » 7 is tiem 1 eat ae ic +9 Lm <9 . _ qrhpe r ‘a ‘ hae i Bull. nat. Hist. Mus. Lond. (Geol.) 57(2): 109-114 Issued 29 November 2001 Systematic affinity of Acroporella assurbanipali Elliott (Dasycladaceae), with notes on the genus Neomeris FILIPPO BARATTOLO & ROBERTA ROMANO Dipartimento di Scienze della Terra, Universita degli Studi di Napoli Federico II, Largo San Marcellino 10, 80138 Napoli, Italy Synopsis. The holotype of the extinct Dasycladacean alga Acroporella assurbanipali Elliott is re-described and a reconstruc- tion is presented. The status of the species is reviewed, and it is assigned to the genus Neomeris, subgenus Larvaria. INTRODUCTION This paper deals with the systematic affinity of the Lower Creta- ceous alga Acroporella assurbanipali, which is known only from the type-locality at Erbil Liwa, Iran. We present a new description based on a reinterpretation of the holotype, which is deposited in the Natural History Musem, London (Elliott collection). A. assurbanipali is reassigned to the genus Neomeris, subgenus Larvaria. The taxo- nomic status of Acroporella and Neomeris 1s also discussed. ACROPORELLA ASSURBANIPALI AND ITS RELATIONSHIP WITH THE GENUS ACROPORELLA (PRATURLON) EMEND. PRATURLON & RADOICIC The genus Acroporella was established by Praturlon in 1964 and based on algal specimens from Upper Jurassic — Lower Cretaceous shelf limestones of the Central Apennines (Marsica, Central Italy). In that paper, Praturlon characterized the genus as simple thallus, with euspondyle and acrophorous primary laterals (= branches), and probably endosporous. The type species Acroporella radoicici has the following characters: cylindrical, not segmented dasycladaceae; branches acrophorous, simple, oblique to axis (15—45° on the hori- zontal), arranged in euspondyle alternating whorls; calcification reaching the axial cell; dimensions in mm (between brackets the extreme value): D=0.55 (0.36—-0.70); d=0.25 (0.17—0.33); p=0.06 (0.05—0.07); w=8-12; h=0.1 (0.085—0.11); maximum measured length of the fragments=5.4; reproductive organs unknown. Praturlon & Radoicic (1974), following the rules of the Inter- national Code of Botanical Nomenclature, changed the epithet from A. raidoicici to A. radoicicae. Moreover, on the basis of better preserved Barremian-Aptian material from Dinarides, they showed that the species bears four short secondary laterals per primary lateral set distally. Accordingly they proposed an emendation of the genus and gave the following diagnosis for Acroporella: ‘Cylindri- cal, not segmented dasycladaceae, having whorls of primary branches distally ramified in clusters of button-like secondary twigs’. However six years before the emendation by Praturlon & Radoicic (1974), Elliott had established the species A. assurbanipali from the Lower Cretaceus of Erbil Liwa, Iran. According to Elliott (1968) the species is characterised by: ‘Cylindrical tubular calcified dasyclad, external diameter 1.36 mm, internal diameter 0.55 mm (40% of external); successive near-horizontal verticils, probably 3 or 4 per © The Natural History Museum, 2001 mm of tube-length, of perhaps twelve radial branches each. The single branches communicate with the stem cavity by a pore of about 0.052 mm diameter: they then swell out to a fig- or flask-shaped cavity of 0.182 mm maximum diameter, narrow to a slightly curved tube of 0.078 mm diameter, and then at the outer surface flare out to a shallow terminal diameter of 0.156 mm’. Since its proposal, A. assurbanipali has been overlooked by most authors. Granier & Deloffre (1993) re-assessed its taxonomic posi- tion and confirmed the affinity with Acroporella, but with reservations. One of the authors of the present paper (F. Barattolo) recently examined the Elliott collection at the Natural History Museum, London. Re-examination of the single specimen in thin section (holotype) and Elliott’s description, allows us to propose a new interpretation of the anatomy of the alga. Accordingly, we assign the species to the genus Neomeris. NEOMERIS (LAMOUROUX, 1816), ITS SUBGENERA AND ALLIED GENERA The genus was defined on an extant species from Jamaica (Neomeris dumetosa Lamouroux). The alga is characterised by a cylindrical thallus bearing whorls of primary laterals. Each primary lateral bears at its outer end a single ovoid ampulla together with two secondary laterals. The laterals, usually those of secondary order, may produce a cortical layer (Valet, 1968; Berger & Kaever, 1992). Seven species are known to live in tropical and subtropical seas where they populate protected microenvironments of the reef plat- form (Valet, 1979), in moderately exposed environments (e.g. Neomeris mucosa) and in tidal pools (e.g. Neomeris cokeri) (Taylor, 1972). According to Konishi & Epis (1962) a depth of 0-10 m and a temperature of 15—20°C are their optimal environmental conditions. The main diagnostic characters utilised in botanical description of specific rank in extant Neomeris are: shape of primary laterals; shape of secondary laterals and their distal ends making a cortex or not; shape of fertile ampulla and cyst; shape of the plug set at the connection between primary laterals and ampulla; calcification (Génot, 1980). In spite of its modest role in the modern marine environment, the genus Neomeris is well represented in the fossil record, but incom- plete calcification does not allow observation of some characters that are useful for classification. The earliest occurrence of the genus (Neomeris cretacea) is in the Albian of Orizaba (Mexico; Barattolo, 1987). C. Neomeris subgenus Neomeris Figs 1A—C__ Reconstruction of fossil species belonging to three subgenera of Neomeris. A, Neomeris (Larvaria) assurbanipali Elliott noy. comb.; B, Neomeris (Drimella) drimi Radoicic 1984; C, Neomeris (Neomeris) cretacea Steinmann 1899. On the right, the relationships (not to scale) of the primary lateral (R,; broken lined circles), ampulla (A; grey circles) and two secondary laterals (R,; white circles) are shown. SYSTEMATIC AFFINITY OF ACROPORELLA ASSURBANIPALI 111 PLATE 1 Fig. 1 Neomeris (Larvaria) assurbanipali Elliott noy. comb. Oblique section. Lower Cretaceous (Valanginian — Hauterivian), Garagu formation, Erbil Liwa, Iran; V.52032, x 34. Fig.2 Neomeris (Larvaria) assurbanipali Elliott nov .comb. Detail of the left side of Fig.1, showing the ampulla set on distal end of a primary lateral. Note the secondary lateral set beneath the fertile ampulla. Lower Cretaceous (Valanginian — Hauterivian). Garagu formation, Erbil Liwa, Iran. V.52032, x 54. Fig.3 Neomeris (Larvaria) assurbanipal Elliott nov. comb. Detail of the lower part of Fig.1, showing the tufts of two phloiophorous laterals making a sort of cortex at the periphery. Lower Cretaceous (Valanginian — Hauterivian), Garagu formation, Erbil Liwa, Iran; V.52032, x 48. 112 In 1913, Leon & Jean Morellet added greatly to our knowledge of Tertiary fossil species of Neomeris and related forms. Their work was based on a rich microflora collected by Munier-Chalmas from the Paris Basin at the end of the 19" century. In addition to document- ing new fossil material, they established several neomerid genera (e.g. Larvaria, Vaginopora, Meminella) that were later considered as junior synonymous of Neomeris (Génot 1980). Deloffre & Génot (1982) gave the following description of Neomeris: “Two orders of branches. Each primary branch bears one fertile ampulla and two (sometimes three in living Neomeris) secondary branches. These are set on both sides of the fertile ampulla (Neomeris s.s.) or setin a plane located beneath the plane containing fertile ampullae (sub-genus Larvaria) . Later, Radoicic (1984) established the new subgenus Drimella, based on a Late Cretaceous species from Serbia, and gave the following diagnosis: “Neomeris with secondary branches laterally located on one side of the fertile ampulla, on vertical or more or less oblique plane. In adjacent verticils, secondary branches can be arranged: in one on the same, and in another on the opposite side of the fertile ampulla, or in each verticil on the same side’. The subdivision into three subgenera (Neomeris, Larvaria, Drimella) was confirmed by Génot (1987). Their structural organi- sation is illustrated in Fig. 1. Phylum CHLOROPHYTA Pascher, 1914 Class CHLOROPHYCEAE Kiitzing, 1843 Order DASYCLADALES Pascher, 1931 Family DASYCLADACEAE Kiitzing, 1843 Tribe NEOMEREAE Pia, 1920; emend. Bassoullet et al., 1979 Genus NEOMERIS Lamouroux, 1816; emend. Deloffre, 1970 Neomeris assurbanipali Elliott, noy. comb. 1968 Acroporella assurbanipali Elliott: 18, pl. 1, fig. 5. HOLOTYPE. V.52032 (1), slide. HORIZON AND LOCALITY. Lower Cretaceous, Valanginian/ Hauterivian. Garagu Formation. Well no.116, Kirkuk, Iraq. REFERRED MATERIAL. G. F. Elliott Colln., presented 1966. DESCRIPTION. The character of the species can be deduced from the single specimen in oblique section (pl. | fig. 2). Assessment of Table 1 Main biometrical parameters of Neomeris assurbanipali Elliott nov. comb. All size parameters are given in mm. d: inner diameter of the calcareous skeleton; D: outer diameter of the calcareous skeleton; e: thickness of the calcareous wall; pv: vertical width of primary laterals; pvd: vertical width of primary laterals at their distal end; /: length of primary laterals; w: number of primary laterals per whorl; /: distance between two subsequent whorls; p’: width of secondary laterals; /’: length of secondary pores; da: diameter of the ampulla. This paper From Eliott, 1968 d 0.55 0.55 D 1.4 1.36 e 0.43 pv 0.16 — 0.20 0.182 pvd 0.156 I 0.25 0.78 WwW 11-12 12 h 0.23 — 0.25 p 0.010 —0.012 | 0.20 da 0.11 F. BARATTOLO AND R. ROMANO Fig. 2 Neomeris (Larvaria) assurbanipali Eliott noy. comb. Longitudinal reconstruction of the thallus. Left side: central siphon, primary laterals, secondary laterals, ampullae and calcification (in black) in axial section. Right side: perspective view of the primary laterals, secondary laterals, ampullae and central siphon deprived of calcification. On the left frontal part only the junction of the laterals and the ampullae are drawn. At the base is a perspective view of the cortex built from the secondary laterals drawn in hexagonal meshes ( x 45). SYSTEMATIC AFFINITY OF ACROPORELLA ASSURBANIPALI the variability of the species and some qualitative characters such as the shape of thallus, laterals and reproductive organs would have required more abundant material. The thallus is apparently cylindrical and simple. Primary laterals are arranged in moderately close whorls. Their position between whorls, alternated or in continuity, is not evident. However they are phloiophorous and rather strong, almost perpendicular to the stem axis. The transverse section of the primary pores is rectanglar in the middle part and square in the distal part. The number of primary laterals per whorl can be estimated as 1 1-12. This value corresponds well with that supplied by Elliott. Two strong secondary laterals originate from the distal end of each primary lateral. They show a subcylindrical inner part of modest length, 0.11—0.12 mm, and 0.10—0.12 mm in diameter, followed by a part that flares out quickly, so making a cortex at their distal end (pl.1, fig. 3). The two secondary laterals are nearly placed on an horizontal plane, i. e. parallel to the verticillar plane. Elliott did not identify the twofold laterals, but he interpreted the shape of primary and secondary laterals in longitudinal section as parts of a single lateral. He correctly described the first inner part (primary) as ‘a fig- or flask-shaped cavity’, and the second outer part (secondary lateral) as ‘a slightly curved tube’. The holotype occasionally shows structures referable to reproduc- tive organs (ampullae: pl.1, fig. 2). The ampulla is set at the distal end of each primary lateral, over the plane where the secondary laterals are placed. It is of spherical shape, 0.11 mm in diameter, joined to the primary lateral probably by means of a short peduncle. The difficulty in observing the reproductive organs can be explained by the fact that most of them were empty and were destroyed after the re-crystallisa- tion of the aragonitic calcareous skeleton. Only those filled by matrix have a chance of being preserved and recognised in thin section. The calcification consists of a strong calcareous skeleton. It envelopes the primary laterals, the ampullae and the secondary laterals up to their swollen distal end corresponding to the inner edge of the cortex. The smooth and regular inner cavity indicates that calcification probably reached the wall of the central stem. More- over, in this area the primary laterals open with a reduced pore. The main biometrical values of the alga are given in Table1. PALAEONTOLOGICAL RECONSTRUCTION The reconstruction in Figs. 2 and 3 takes into account the biometrical values and the morphological characters previously given. The number of pores ina whorl has been chosen to be twelve, and a cortex of hexagonal meshes set horizontally has been supposed. The latter seems to be the most likely arrangement for that number of pores in a whorl and the distance between whorls. Considering the apex of the thallus, no elements can be deduced from the single specimen at our disposal, and therefore it has been drawn by inference from extant species of Neomeris (e.g.. N. annulata). TAXONOMIC ATTRIBUTION The species is attributed to the genus Neomeris from the evidence of primary laterals in verticils that bear distally two secondary laterals and an ampulla. The fact that the species exhibits two secondary laterals arranged in an horizontal plane beneath an ampulla, allows us to refer N. assurbanipali to the subgenus Larvaria. The species, if compared with others of the same genus, shows such a combination Fig. 3 Neomeris (Larvaria) assurbanipali Elliott noy. comb. Transverse reconstruction of the thallus. Upper half: part of a verticil showing the central siphon, primary laterals, secondary laterals and ampullae without calcification. Lower half, right: transversal section through the primary and secondary laterals, calcification in black. Lower half, left: transversal section through the primary laterals and ampullae, calcification in black ( x 45). of characters as to maintain the taxonomical validity. In addition Neomeris (Larvaria) assurbanipali differs strikingly from all the other counterparts by having a calcareous skeleton that envelops the primary laterals entirely, and seemingly reaches the central stem. REFERENCES Barattolo, F. 1987. Remark on Neomeris cretacea Steinmann (Chlorophyta, Dasycladales) from the Cretaceous of Orizaba (type- locality), Mexico. Bollettino della Societa Paleontologica Italiana, 29: 207-218. Berger, S. & Kaever, M.J. 1992. Dasycladales — An illustrated monograph of a fascinating algal order. Stuttgart-New York (Georg Thieme Verlag). 247 pp. Deloffre, R. 1970. Un niveau a Algues dans le Sparnacien de la region de Plagne (Petites-Pyrénées-Haute-Caronne) et observations sur le genre Neomeris LAMOUROUX 1816. Bulletin du Centres de Recherches de Pau—SNPA., 4 (2): 353— 379. — & Génot, P. 1982. Les algues Dasycladales du Cenozoique. Bulletin du Centres de Recherches Exploration-Production Elf-Aquitaine, memoire 4: 447 pp. Elliott, G. F. 1968. Permian to Palaeocene calcareous algae (Dasycladaceae) of the Middle East. Bulletin of the Natural History Museum, London, Geology, supplement 4 (1): 11 1pp. Génot, P. 1980. Les Dasycladacées du Paleocéne supérieur et de 1’ Eocéne du Bassin de Paris. Mémoires de la Société géologique de France, 138 (2): 25. — 1987. Les Dasycladacées du Paleogene d’Europe nord-occidentale (Bassin de Paris, Bretagne et Cotentin). These de doctoral d'etat, Université de Nantes, 1: 499 Pp: Granier, B. & Deloffre. 1993. Inventaire critique des Algues dasycladales fossiles. II’ partie: Les algues dasycladales du Jurassic et du Cretace. Revue de Paléobiologie, 12 (1): 19-65. Konishi, K., & Epis, R. C. 1962. Some early Cretaceous calcareous algae from Cochise Country, Arizona. Micropaleontology, 8 (1): 67-76. Morellet, L. & J. 1913. Les Dasycladacees du Tertiaire parisien. Mémoires de la Société géologique de France, Paleontologie 47: 43 pp.. 3pls. Praturlon, A. 1964. Calcareous Algae from Jurassic-Cretaceous limestone of Central Apennines (Southern Latium-Abruzzi). Geologica Romana, 3: 171-201. — & Radoicic, R. 1974. Emendation of Acroporella (Dasycladaceae). Geologica Romana, 13: 17-20. 114 Radoicic, R. 1984. New species and new subgenus of Neomeris (Dasycladaceae, Green Algae) from the Upper Cretaceous of Metohija. Bulletin de l’Academie Serbe des Sciences; classe des sciences mathematiques et naturelles, 25: 17-33. Taylor, W. R. B. 1972. Marine algae of the eastern tropical and subtropical coast of the Americas, pp.97-107. Ann Arbor University, Michigan Press. F. BARATTOLO AND R. ROMANO Valet, G. 1968. Contribution a I’ étude des Dasycladales. Nova Hedwigia, 16: 21-82 —— 1979, Paleontological approach to dasycladales from the ecology of recent forms. Bulletin du Centres de Recherches Exploration-Production Elf-Aquitaine, 3 (2): 852 pp. Bull. nat. Hist. Mus. Lond. (Geol.) 57(2): 115-162 Ge a cS Issued 29 November 2001 Palynological zonation of Mid-Palaeozoic sequences from the Cantabrian Mountains, NW Spain: implications for inter-regional and interfacies correlation of the Ludford/Pridoli and Silurian/Devonian boundaries, and plant dispersal patterns JOHN B. RICHARDSON Department of Palaeontology, Natural History Museum, Cromwell Road, London SW7 5BD ROSA M. RODRIGUEZ Departamento de Ingenieria Minera, Universidad de Leon, C/Jesus Rubio 2, 24071 Leon, Spain STUART J. E. SUTHERLAND Department of Geology, University of British Columbia, 6339 Stores Road, Vancouver, British Columbia, V6T 1Z4, Canada CONTENTS ETRE UU UO OTN es ote sees ose cao toc vna vices ouvan vo eeeVuse da Sn eeos er neat at ive sed edae sat enena cout eet cvsswsedvacuecoasesaebdscecusdsvsegedssdsivestesaseacesas ast vonesceqsensareeaess Geographical & geological setting ...........:c::ccceseeeeereeeeees SAM SATO MOMMA OM reece recesses tecenes-cereeetneersenereees Age and biozones of the San Pedro Formation IS OLE Sener terse cent cee neat n ee MeO cnet ncesevvagenresncurerecssn cer evstsecevasuors sarscnnssescecase st dunedhneutnatanarage svassusreaas ioatnescsceonaseneaanansecares Age of the biozones and inter-regional palynological Correlation ..............:::cccecssceseseesesesesescscecesesevscscsscsesesecsessescssseacaseeeeesees 123 GJPOIS DIOZOVGTSS cossecorgccoce:coorcoaebososes:dedanes60087ce50s800 2805004 2cOF CSREES OO ESOH AOC CN = 225000 SEE OEE AEH aN GOCE COO CORSSDEC LED BECUC ES pec SCEECoROSBCEAESRONeU dor 123 Laurussia-Gondwana spore distribution patterns 126 (CURT OTA CRA craccencaceenbaceg acta coc aSC COLE O-RSECURCGIEEOS RIG BOERS Eb SCG CCO ORAL ROC EREE CEO EOO ECSC Oo SODSGEca SCH COBEOE SO See co EOF pu soon ag aoe sPaceceecEeee 129 (GSTS ceosedgencooncgnecogdecben7 ias0463 5 Sood 50 Hor 2s Seco G eae a cceCEE GEASS CE EBEG EEL CORAL ECOL eSECSHORO PEACE DEE SEES EO SEE ot bad coor aacoceLuocoocbanoectcete 129 ENTIBEONET (Oo -necocceaqeecnencentsceeec o256382260020240C0C 200949 NCOTEE a0 07UTCESCC OOS ROGDOR ESL conccnegzo cen aon oe coe eaSaecoscocbeccoonccrecoceecocccoscncenos.cececoeneceooot 130 TE AN 1 Cl seen et Nea. ene Pe Se Nae ee cee eA nue civ uc ecu stius et ettetataruse ctcaceneseses stnsa esturtans:Sitazstates stinacwssrenetunsttesepetsteas 131 [Lg JPRS -cnancsoneneanonnon’ soedasse.300 share coboe SoagogeoCecacn eGo sa6c6 se cbeCac2 Se BE ERACaD EO EDO ose gen .Baa0L0dBsecspeboace ;poroocoengandoaasbodeccectoscaascarcacneercocacen6 132 Comparison of the chitinozoans from the Cantabrian Mountains with coeval sections and the global biozonation scheme .. 133 Silunanpeloball@hitinoZOAanlbiOZONa tomer erscscece-ceecesces-essneses-senenenasactecransetere -tetenersnenvenened etter er erererenctate renter eenr i ressten ters tee 135 Biostratieraphical CONCIISLOMS sceccccc-tenec--ecesssscesvereseseceesevececnsnseurnesenersssevtsnscnsesasteasesctenereterserreverrsrericucesosestecsssecensusrerersersncesrez= 135 136 Materials and methods Maxonomiy Of Selected) Spore CAKG ..2....c.cscecc-cecessaseessececarensesoasvecnassccsadeneusncesceescncteanesnerscuevanecuesessisincetsnessrieeseseeesssesansn Antetunma Sporites H. Potonié 1893 or-....-c....cecccececsssecasernscrvoscusccrsnsnscessnscrencccceceesenesetersceescorneceraccestnsnseseanernsasess Subturma Azonotriletes Luber 1935 Genus Retusotriletes (Naumova) Richardson 1965 ...........cc:::cccececeeeeees Genus Apiculiretusispora Stree] 1964 .......cccccccccteeseteeeteeseseeeeeseeenenees Genus Breconisporites Richardson et al. 1982 ..........-. Infraturma Murornati Potonié & Kremp 1954 ............e Genus Emphanisporites McGregor 1961 ..........:.:11e Subturma Zonotriletes Waltz 1935 in Luber & Waltz 1938 ..... Infraturma Crassiti Bharadwaj & Venkatachala 1961 .......... Genus Ambitisporites Hoffmeister 1959 ........-.:.-00 Genus Scylaspora Burgess & Richardson 1995........... Genus Concentricosisporites Rodriguez 1983 .........-.-- Genus Iberoespora Cramer & Diez 1975 ..... Genus Leonispora Cramer & DieZ 1975 .....cccecceecceeeteecteeseeeseesssesscneeerssessecsenessouensenenscnsneaneneneerececuececesereuatectts Genus Streelispora (Richardson & Lister) RAiGhardSOmenailOS Dnamcscccececcesevsesteerenieveas: Genus Aneurospora Stree] 1964 .....c.sseccccecsesccssessesssseeseesesecsecseticercrecsscassnsssssccnesrscsscessressesseisanenscnsancaneetenecnennatnstes Genus Coronaspora Rodriguez 1979 emend, ........cssesseessesecesessecseeseensesesscesssesseesccscseenseneaneaenesneanceucensuecnecnscisees 149 Infraturma Patinati Butterworth & Williams 1958 ...........c.ccccescsceseseseeeesesenerscsessesnscnscsesasavssssusesessnscsciseecsenassssenensenens 155 Genus Chelinospora Allen 1965. ......s:ssessssessecsecseetessesseneesenneesensssessesusenssnesscascassnerscsessecnecasansnuenecnsaneanecsacuanecustiatys 155 © The Natural History Museum, 2001 116 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND GenusiGyrbosporites Allen Sieeesereseerenansceecsess cater nter rns snes ct tee ee ee meena ee rie tee ane cere 159 Taxonomiy/of @hitinoZOalerecccessscecnesnessesseee-e= Order Prosomatifera Eisenack 1972 ............. Family Lagenochitinidae Eisenack 1931 .. Subfamily Angochitininae Paris 1981............ GenuseRaMOGHILiNGlS OME TCZAVATINE OC KE ILO GAs mereeeneenteee a etee es cectac ce ate sene ane senetereeaeerenetante serene rere ener 160 Acknowledgements... 1RXSY RSIS NT EIS i see eSona cb ehonabtosdonoreanéeendaecosoccaccbo-posococaeaceedebacac coseance bons oSeesca0b00}00;G0 Joss soseenguedaeec 2c CsoeGaanbsESOuecdb Teen coe ebuoacer sBseenoscccoaacLS: 160 SYNOPSIS. Mid-Palaeozoic strata from the Cantabrian Mountains (north-west Spain) contain rich assemblages of spores, acritarchs and Chitinozoa. Fossil maturation is variable but generally high. The stratigraphical distribution of miospores and cryptospores from four sections through the San Pedro and lower La Vid Formations is sufficiently consistent for the establishment of a sequence of six biozones (including four new biozones). In ascending order the biozones are: 1, Scylaspora vetusta —S. sp. B (V) Spore Biozone; 2, Artemopyra brevicosta — Hispanaediscus verrucatus (BV) Spore Biozone; 3, Coronaspora reticulata — Chelinospora sanpetrensis (RS) Spore Biozone; 4, Chelinospora hemiesferica (H) Spore Biozone; 5, Scylaspora elegans — Iberoespora cantabrica (EC) Spore Biozone; 6, Streelispora newportensis — Emphanisporites micrornatus (MN) Spore Biozone. The elegans — cantabrica (EC) Spore Biozone is divided into two sub-biozones based on the first appearance of the genus Aneurospora and the lower part of the MN Biozone is distinguished as a separate subzone, namely the Streelispora newportensis — Leonispora argovejae (NA) Spore Assemblage Sub-Biozone (also present in England). The Ludford/Pridoli boundary is probably within the upper part of the reticulata — sanpetrensis Spore Biozone, and the Aneurospora Sub-Biozone and succeeding MN Biozone, allow inter-regional correlation with basal and Lower Lochkovian strata. Comparisons between England, Spain, and North Africa, show that regional differences in spore floras are most marked in the Pridoli. In the Lower Devonian the differences are less pronounced, and the appearance of variants of the genus Aneurospora marks a significant event in both the Laurussian and Gondwanan regions, reflecting wide dispersal of their parent plants. Chitinozoans occur throughout most of the sequence and several species have potential for inter-regional correlation e.g. Ramochitina villosa (RS Biozone) upper part of the urna (lower EC Biozone) and Eisenackitina bohemica (lower MN Biozone). INTRODUCTION Because of their abundance and species diversity land plant spores have a much greater potential for the determination of ancient floral regimes than their parent plants. Further, where their fertile parts are known, all Silurian to Lower Devonian (Lochkovian) plants are homosporous with small spores (often c. 20 um, and below 100 um). Factors which should ensure that their parent plants are widely dispersed. In fact, mid-Palaeozoic spores and marine palynomorphs are found on each of the major drifting continental blocks studied, namely Laurussia and Gondwana. Comparisons of sequences of spores from the Anglo-Welsh area with those from the Cantabrian Mountains and North Africa should aid in the resolution of the nature of these Palaeozoic floras. However, a prerequisite is the study of spore sequences in the three areas. In this study miospores from Southern Britain and the Cantabrian Mountains are studied in detail along with some preliminary work on the chitinozoans of northern Spain. Work on North Africa has been partially published (Richardson & Ioannides, 1973) and additional North African areas are currently being studied. Seward (1931) stated that it is inconceivable that the climate of the Earth was ever uniform, and influences on vegetation might be expected to be profound where crustal pieces drifted through different latitudinal belts. By providing an accurate biozonal framework this work aims to facilitate more accurate comparisons of spore floras in the three areas. The resultant floral dispersal patterns vary through the time interval studied and further work is needed to answer some of the many questions raised. One of the important features of this study is the completeness of the Pridoli sections in the Cantabrian Mountains in a marine but continentally influenced succession. This probably represents the most complete miospore zonation through the Pridoli Stage. In the continental sections in England there are some of the longest, well- preserved, miospore successions, with a sequence of miospore assemblages from Upper Aeronian to Lower Pridoli and uppermost Pridolf to Lower Lochkovian (Silurian to Lower Devonian). How- ever, there is a gap, equivalent to the upper Lower Pridoli, Middle Pridoli, and lower Upper Pridoli, where no miospore assemblages are known. Between these productive beds of the Lower Devonian (Aneurospora Subzone and MN Zone) and those from the Lower Downtonian (Lower Pridolt) below, there is, therefore, in England, a considerable gap in the Pridoli spore occurrence. Although there are spores in graptolite-bearing marine sediments in Podolia these spores have been inadequately described and illustrated. The spores illustrated by Shepeleva (1963) from the Lower Devonian of Podolia possibly belong to the MN Zone and her species Lophozonotriletes decoratus closely resembles Streelispora newportensis, the nominal species for the base of the MN Zone. GEOGRAPHICAL & GEOLOGICAL SETTING (FIG. 1) The Cantabrian Mountains, in the north and northwest of Spain, extend through the Provinces of Asturias, Le6én, Cantabria and Palencia. Geologically, most of the Cantabrian Mountains belong to the ‘Cantabrian Zone’ of the Iberian Hercynian Massif and occupies the outermost part of it. The “Cantabrian Zone’ is composed of unmetamorphosed Palaeozoic sedimentary rocks deformed during the Variscan Orogeny. The “Cantabrian Zone’ has been divided into five tectono-sedimentary units (Julivert, 1967), arranged in concen- tric bands parallel to the ‘Narcea Anticlinorium’, an arch of Precambrian rocks on the western boundary of the ‘Cantabrian Zone’ (see Fig. 1). Following to the east of the ‘ Narcea Anticlinorium’ is the ‘Folds and Nappes Unit’ where the most complete Palaeozoic sequences are found. This Unit occupies the western part of the Asturias and the northern part of the Leon Provinces. Within this structural unit four sections in a Silurian to Lower Devonian clastic succession have been selected for study. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN 117 La Peral 14 a Faults and Thrusts — Outcrop of the San Pedro Foramtion po Mesozoic / Tertairy PreCambrian — Zo 20km SS Fig.1 Simplified geology of the Cantabrian Region, NW Spain (based on Rodriguez, 1983). San Pedro Formation This formation was described in two main areas: a, the ‘Ferruginous Sandstones of Furada’ (Barrois, 1882) on the northern flank of the Cantabrian Mountains, and b, the San Pedro Formation (Comte, 1937) in the south. These two names refer to the Upper Silurian Formations of the ‘Folds and Nappes Unit’ of the Cantabrian Zone. Three of our sections were originally designated as the San Pedro Formation (Geras de Gordon, La Vid de Gordon and Argovejo). The fourth section was included in the Furada Formation (La Peral). As both formations constitute a single lithostratigraphic unit they are all referred to in the text as San Pedro Formation. The San Pedro Formation is predominantly sandy and overlies the Formigoso Formation; the latter is Silurian in age. The Formigoso Formation is composed of black shales, with alternating shales and quartzites in the upper part. The San Pedro Formation appears to be in stratigraphical continuity with the underlying Formigoso Forma- tion in the sections near the Precambrian of the ‘Narcea Anticlinorium’, but the contact is sharp and erosive in the sections situated farther east. Informally three members can be distinguished in this formation. The Lower Member consists mainly of red ferruginous, coarse- grained sandstones, sometimes with oolites, having cross-trough stratification. At some localities there are conglomeratic beds, with phosphatic, sideritic and silty clasts and intraclasts; some of these clasts have a volcanic origin (Van den Bosch 1969; Suarez de Centi, 1988). The Middle Member is composed of medium to fine grained sandstones with current and wave ripples, and grey to green intensively bioturbated mudstones and siltstones. The Upper Member has fine-grained quartzitic sandstones, with cross stratification, interbedded with dark to black shales. The San Pedro Formation was deposited in a shallow epeiric sea in intertidal to inner platform conditions (Suarez de Centi, 1988) in a transgressive-regressive cycle (Rodriguez, 1983). In this latter inter- pretation the middle member represents the deepest conditions. Age and biozones of the San Pedro Formation The age of this formation has been established on macrofauna, microplankton and spores. A macrofauna of brachiopods, mol- luscs, connularia, trilobites and graptolites has been found, but they are not abundant and are restricted to certain beds (Comte, 1934, 1959; Llopis Llad6, 1958, 1960, 1964; Poll, 1962, 1963, 1970). The lower half of the formation is of Ludlow age, based on brachiopods and connularia (Comte, 1934). In some sections, situ- ated to the west of the ‘Folds and Nappes Unit’ (see Fig. 1), near one of our localities (La Peral), Poll (1963, 1970) found graptolites 20 to 50 m. below the top of the formation: Saetograptus fristichi fristichi, S. fr. linearis, S. chimaera salweyi belonging to the nilssoni, scanicus, and probably leintwardinensis Biozones and indicating a Gorstian to Lower Ludfordian age. The highest part was dated as Lochkovian on its macrofauna (Comte, 1934, 1937, 1959; Poll, 1970). In contrast to the macrofauna, microplankton and miospores are extremely abundant and well preserved. Microplankton (acritarchs, 118 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND chitinozoans) and miospores were extensively studied by Cramer 1964a, b, 1966b, c, 1967, and 1970; by Cramer & Diez 1968, 1975 and 1978a, b; and by Rodriguez 1978a, b, c, 1979, and 1983. Several biozones based on chitinozoans (Cramer & Diez, 1978b) SPORES Biozonation (Figs 2-6) and on acritarchs and miospores (Rodriguez, 1983) have been The San Pedro Formation has been sampled in four sections: at described. These biozones were established on composite sections Argovejo, Geras de Gordon and La Vid de Gordon (referred to in the and thus they do not constitute formal biostratigraphical units text as Geras and La Vid respectively) and La Peral. Productive according to the rules of the International Code of Stratigraphic samples from the La Peral section occur at a lower horizon than in the Nomenclature. Instead they constitute respectively, chitinozoans, other sections (Fig. 5). Strata from the underlying Formigoso Forma- acritarchs and spore associations, each one arranged in chrono- tion were sampled in the Argovejo and La Vid sections only. The logical order, ranging from the Upper Wenlock? to Lower samples from this formation are practically barren of spores but may Lochkovian. Based on the miospore associations found in the contain chitinozoans. The absence of smaller palynomorphs, such as Geras Section, and on correlation with other palynological assem- spores, may indicate turbulent to winnowing environments, and blages found in several different localities, Rodriguez (1983, fig. possible reworking of at least some chitinozoans. Alternatively, there 13) placed the Silurian/Devonian transition about 5 m below the may be a disconformity either within, or at the top of, the Formigoso stratigraphic level adopted in the present work. This author also Formation (Cramer & Diez, 1978b), and possibly within the lower showed that the boundaries of this formation are highly San Pedro Formation. The La Vid Formation, overlying the San diachronous (eg. the upper San Pedro Formation is younger in Pedro strata, has been sampled only at the La Vid and Argovejo sections to the south and west of the “Folds and Nappes Unit’ than sections, but all samples taken from shale intercalations in the sandy in those closer to the Precambrian Asturian Arch). Argovejo, Geras dolomites were barren. and La Vid are in the south part of this unit and all these sections Four new spore zones, and one subzone, are proposed. These have Lochkovian strata at the top. zones, with the exception of the hemiesferica Biozone, contain two Chitinozoans of Pridolf and Lochkovian age from the Carazo nominal taxa, at least one of which starts at the base of the zone. The Formation, of the ‘Pisuerga-Carrion Unit’, east of the “Cantabrian zones are described in ascending order and reference sections are Zone’, were described by Schweineberg (1987). The Carazo Forma- given for the base of each zone, or its lowest known occurrence. In tion differs in lithology from the San Pedro Formation but, although addition, in two sections, possible equivalents of the Synorisporites the precise lateral continuity between these two formations has not libycus — Chelinospora poecilomorpha (LP) Assemblage Spore been established, the San Pedro Formation also includes sediments Biozone (Richardson & McGregor, 1986) occur, and in the La Peral of Pridoli and Lochovian age. Section the Artemopyra brevicosta — Hispanaediscus verrucatus (3) a s 4 3 = : aoa Ae -Angochitina elongata Mid Gorstian - Early Pndoli {Sandstone 1S They &8 5 & "SG G «| At- Angochitina tsegelnjucki_ Lochkovian Bishate =» Phosphatic nodules S Si S = Sas Be te ws $ = 5 | Eb - Eisenackitina bohemica Lochkovian EQ) sandstone & Shale fx) Dolomites Sc) iS Ls S e iS RS 3) g = y 8 SS || Rv Ramochitina villosa Late Ludfordian HB Sandstone / Oolites FA Limestones 8 N fe S oS 8 x aS = a = 68 > S Ss Me - Margachitina elegans Mid Pndoli mesh, ~ EJ Quarwites ES ga SS 528 8S 72 JS 8 FS | Ss - Sphaerochitina sphaerocephala Mid Gorstian - Early Pridoli ree SHAS ESS S v= 8 2 Rn Oe & & «| Pe - Psuedoclathrochitina carmenchui Mid Pridoli Ss iS $ 8 2 Sa 8 acs 8 $ 3 Sis s Uu - Urnochitina urna Pridoli - Earliest Lochkovian SESALSS PREVSE SS LSPS SA RSASREESSSS RR ERE RS SS SO PS aS aS Bae ©: S48 ‘ : WES Chitinozoan SSSSSSSSESSSRSESS ESS Spore 2 DRasSv_eVaBaFS SV vxwgoeV@s V4; ti Zonation xe Oe FS ot FOSS FSET E VES A onation VUOUHRVVOVKHOHVITCOITNAD SESS) | | ts | [ Ger 18 mMn|Loc Ger17 MN Loc. OM |S! Ger 16 Lochkovian (na) bo |B} Ger 15B (na) exeiloae el Ger 15A (a) Isu @ Ger14 _ _ _-—Eb/At (el | | Ger 13 Pri. g e Ger 12 (a) —= a EG cle| | & | Ger 11 ro | §| [Ger10 EC [7 ° @ Gero _ a Po/M = Kea ig Soo eia|) aie Pridoli fe | Ger7 F DB | rEerGL =p — Pri.| H Ger5 IL Ger4 Ger3 es I | | Ger 2B BS }— Ss/Ae RS ?Pri [ | | Ger 2A (LP t— Rv —-— =) = —= = 2 | Ger1 a —! | —— 7 ba . ines Geta a Ludfordian Fig. 2 Selected spore distribution at Geras. Chitinozoan zonation after Verniers et al. (1995): bo: bohemica, su: superba, ele: elegans, ko: kosovensis, ba: barrandei, ph: philipi. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN (BV) Spore Biozone may be present. The latter zone is based on material from the Anglo-Welsh region (Burgess & Richardson, 1991) where the base of the zone, in the type Wenlock area, is recorded in the late /undgreni Graptolite Biozone (Lower Homerian. Wenlock). Scylaspora vetusta — Scylaspora cf. kozlica (Dufka) (V) Spore Assemblage Biozone DEFINITION. Based on the co-occurrence of the two nominal taxa with Scylaspora scripta (Burgess & Richardson, 1995). NOMINAL SPECIES Scylaspora vetusta (Rodriguez) comb. nov. Rodriguez, 1978b: pl. 1, fig 8. Scylaspora cf. kozlica (Dufka) comb. nov. Dufka 1995: pl. 3, figs 10a—10b. CHARACTERISTIC SPECIES. The species Ambitisporites abundo (Rodriguez) comb. nov. and Concentricosisporites (2sp.) appear in the upper part of the zone. Within this zone the morphology of Scylaspora vetusta is highly variable. ACCESSORY SPECIES. Long ranging laevigate species such as Ambitisporites avitus Hoffmeister 1959, A. dilutus (Hoffmeister) Richardson & Lister 1969, A. warringtonii (Richardson & Lister) comb. nov., and Archaeozonotriletes chulus (Cramer) Richardson & Lister 1969 are present, along with murornate Synorisporites sp. REMARKS. The base of the Scylaspora vetusta Biozone is not seen as both the nomimal species occur in the lowest sample of the La 119 Peral section. The upper boundary is defined by the incoming of the nominal species of the succeeding Artemopyra brevicosta- Hispanaediscus verrucatus (BV) Biozone. OCCURENCE. Strata of the S. vetusta—S. cf. kozlica Spore Biozone are found only in the La Peral section but the nominal species persist into higher zones in other sections. Higher records are discontinuous and may be, at least in part, due to reworking (La Vid Coronaspora reticulata — sanpetrensis and lower Chelinospora hemiesferica (H) Biozones). AGE. Possibly lower Upper Homerian. Chitinozoa indicate a doubt- ful Llandovery age and may be reworked. Artemopyra brevicosta — Hispanaediscus verrucatus (BV) Spore Biozone REMARKS. Defined in the Anglo-Welsh area (Richardson & McGregor, 1986, Burgess & Richardson, 1995: 26, text-fig. 8) this zone ranges through the middle and upper parts of the Homerian but is unknown in the lower part of the /undgreni Graptolite Biozone at the base of this stage in the type area. The spore zone in the type area is divided into three subzones, namely: Artemopyra brevicosta, Hispanaediscus lamontii, and Emphanisporites protophanus (Bur- gess & Richardson, 1995). In the La Peral sequence the simultaneous appearance of A. brevicosta and H. verrucatus (sample LP4) is followed by the occurrence of Hispanaediscus lamontii in the next higher sample (LP5, 12 m above) and the nominal species persist into the overlying RS Spore Biozone. All of these species are typical of the Homerian, but two (H. lamonti and H. verrucatus) continue Spore Chitinozoan & BW Dolomites Zonation Zonation | 3% : E Limestones MN & RS £3) WiSandstone /Oolites [J Sandstone L L =) iS, S 84 2 In Oc. Oc. = na s & Se) ee 8 : Wi Shale bo a $3 2 3 & Qe « m & & . s BB Ferrous Oolites e iS oy kel 3 es 2 u= Se | es S Ss 8 8 © 3. aS) Sy S vy 5 e+ Phosphatic nodules FjSandstone & Shale ee ee = SS S @ 8 = EC | ie S S| S is ars) iS YS Ss a 2 3 S LL ES Al -Ancyrochitina libyensis ?Late Gorstian / Pidoli laps Sas ER OH ES BSS) S SS sas 2S Ac - Angochitina chlupaciLochkovian le! Ss Nes SSN Ry Ss oS Ge ax e SI Se S £ Soy oS 8 $ = ~ % Sys Ws AL Eb - Eisenackitina bohemica Lochkovian a Ss SS) ks a Se iS y oe Q g O58 Cb - Cyathochitina sp. B Late Llandovery Saas y i] NJ S 28 g SJ S es ws NJ 8 Me - Margachitina elegans Mid Pndoli H 3 5 8 3 iS) 5 2 8 § d x 8 S$ SB eS) 5 Ss - Sphaerochitina sphaerocephala Mid Gorstian - early Pridoli AA A RAO & & 7 8 5 Q g S & Q | Pc-Psuedoclathrochitina carmenchui Mid Pridoli S$ $ Ny 3 3 3 Sy 8 8 & = Q S a 3 is) é Uu - Urnochitina urnaPridoli - Earliest Lochkovian = 2 SS SC) = && Pri) | Pri Seesessseeorg sess 7 ‘ > = ee va K 5 N Se os CESSES SOEREAS & =| BCUOADVOVOOHLOOVKRHACNOOUR Key Cantabrian po Chitinozoan taxa 5 _ Arg 14 iS = . 5 1 -Agis MN|_ Ac Lochkovian E | Arg 12 (na 22kol elie 5 we cre p AN) iE (a) |— 7Eb ee ee ee 3 Arg 11A oO aa - RS es 2) EC 2? s | Arg 10 —! = 5 J ica 2 _Ag 9 ir ea Ag 8 | Uu/ Pe | Ag7 /Me Pridoli L2 aaa | Arg 5G/6 lh a2 Ludf. |p| p.. _ Arg 5F a Pri. Pages — — : | @ _ Arg 5B Ludf, | AgsA RS | Ss ee ee Tal | ml mm es sep . ~—T AS aay, =~ . pAg4 LP LAl Ludfordian g 5 _Ag3 oy |e HE | Arg 2 Cb fe r LAg1 l— Fig.3 Selected spore distribution at Argovejo. Chitinozoan zonation after Verniers ef al. (1995): bo: bohemica, su: superba, ele: elegans, ko: kosovensis. 120 into lower part of the Synorisporites libycus —Chelinospora poecilomorpha (LP) Spore Biozone in the Anglo-Welsh area. Regard- ing the age of the La Peral samples, there are two main possibilities: the spores represent either an impoverished Gorstian (LP) assem- blage with reworked fossils from the Homerian, or b, in situ Homerian spores. Since older spores may be reworked and deposited inside clasts in younger rocks, they may be preferentially retained, whereas the lower density of contemporary palynomorphs may cause them to be selectively removed by currents. OCCURENCE. Found only in the La Peral section. AGE. Homerian. In the type Wenlock the brevicosta and lamontii Sub-Biozones are more or less equivalent to the upper /undgreni and nassa to lower ludensis Graptolite Biozones respectively. There is no chitinozoan evidence for sample LP4 but in the underlying sample (LP3) the chitinozoan species Conochitina rudda also indicates a Upper Wenlock to Lower Ludfordian age. Therefore, the chitinozoan evidence fits with an assignment to either the upper brevicosta — verrucatus (BV) Biozone (Middle to Upper Homerian), or to the succeeding spore zone in the Ludlow. Coronaspora reticulata — Chelinospora sanpetrensis (RS) Spore Biozone (Figs 3-5) NOMINAL SPECIES Coronaspora reticulata sp. nov. Knoxisporites? riondae Cramer & Diez 1975 (pars): pl. 1, fig. 17. Chelinospora sanpetrensis (Rodriguez) comb. nov. Rodriguez, 1978c: pl. 1, fig. 13 (figs i, j, 1). DEFINITION. Based on the incoming of either of the two nominal species. In some sections the two species appear at the same horizon (La Peral), in others C. sanpetrensis appears earlier. The zone represents the interval between the first appearance of the nominal species and the first occurrence of Chelinospora hemiesferica (Cramer & Diez) comb. nov. ACCESSORY SPECIES. Chelinospora poecilomorpha, Coronaspora cromatica, Emphanisporites splendens, Chelinospora cantabrica. REMARKS. This biozone is present in all four sections. In the La Peral section the base lies above a thick sandstone in sample 6, c.71 m above the base of the San Pedro Formation. However, in the Argovejo section several species that occur together in the lowest RS Spore Biozone in the other samples follow one another in sequence. This may mean that the oldest part of the zone is seen in the Argovejo sequence. In all sections, however, Chelinospora cantabrica appears a little above the first appearance of C. sanpetrensis, which suggests that all sections begin at a similar stratigraphical level. The concur- rence of the two species may form a useful sub-biozone. Current data (see also Richardson & McGregor, 1986) indicate that trilete spores, with a regular distal reticulum of even lacunae and well-developed muri, first appear in the Pridoli. However, precise corroboration is lacking, and spore/chitinozoan assemblages from the lower parts of the Cantabrian Mountains sections studied are often poor. Consequently, the exact relation of this sculptural event to the Ludfordian/Pridoli boundary in the Cantabrian Mountains is not clear. Distally reticulate spores first appear in the RS Spore Biozone and the spore evidence indicates an Upper Gorstian age for the base of the biozone. Other rare taxa include /nsolisporites sp. (Pl. 3, fig. 3) and similar taxa are found in Britain in the Ludlow. Chitinozoans in Ger 2A indicate an Upper Ludfordian age and consequentely the upper part of the RS Spore Biozone is probably of Pridoli age. REFERENCE SECTION. La Peral, Cordillera Cantabrica, Province of J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND Asturias, NW Spain. The ‘base’ of the zone is between 49 and 71 m above the base of the San Pedro Formation. The strata in this 22 m interval are dominated by red sandstones and were not sampled. The zone is at least 22 m thick in the La Peral section. DISTRIBUTION. La Peral, Geras, La Vid and Argovejo sections. In the Argovejo sequence the thickness of the zone is uncertain due to a 40 m interval where samples are barren. In the other sections the thickness of the zone is around 15—25 m. ARGOVEJO SECTION. The lowest eleven samples are barren or do not contain diagnostic palynomorphs. In sample ARG/4 poecilomorpha occurs with cf. sanpetrensis and may indicate the presence of the upper part of the LP Biozone ( Upper Ludfordian in England and Wales). Chitinozoa from the same sample are long ranging from Upper Gorstian to Pridoli and therefore do not conflict with the presence of the upper LP Spore Biozone but are insuffi- ciently precise. GERAS SECTION. Chelinospora sanpetrensis, one of the nominal species of the RS Spore Biozone, occurs in the basal sample (Ger 1) of the Geras section. Chelinospora poecilomorpha and Stellatispora inframurinatus var. cambrensis appear in the overlying samples (Ger 2A and Ger 3 respectively) and are also found in the upper Gorstian and Ludfordian of England and Wales but are absent from the succeeding Downtonian there. The presence of the chitinozoan Ramochitina villosa indicates a Late Ludfordian age for sample Ger 2A within the RS Biozone. Emphanisporites splendens appears in Ger2A to Ger6. This species 1s found in North Africa in association with typical lower Downton (Lower Pridolt) species in the upper part of its known range there. Thus, at least part of the RS Spore biozone is of Upper Ludfordian age and, on present evidence, the Ludfordian/ Pridoli boundary probably occurs within the higher parts of the RS Biozone. The Geras section is faulted at the base and no further samples were collected below Ger 1. Coronaspora primordiale (PI. 6, fig. 3) occurs rarely in samples Ger 2B and Ger 3, but in the Argovejo and La Peral sections is found sporadically only in the overlying H Biozone. Emphanisporites spp. (eg. Pl. 2, fig. 9) occur rarely in sample Ger 2B and also extend into the overlying H Biozone in the La Peral section. Other species typical of the Homerian and Gorstian occur erratically in this section and are most probably reworked, eg. Hispanaediscus lamonti. LA VID SECTION. The lowest productive sample in the San Pedro Formation is LV 6, which belongs to the RS Biozone. However, spores similar to S. libycus and indistinguishable from C. poecilomorpha (nominal species of the LP Biozone) occur. In England and South Wales these species die out near the top of the Ludfordian. Part of the RS Biozone is probably, therefore, Ludfordian and may be equivalent to the /ibycus — poecilomorpha Biozone of the Anglo-Welsh area. However, in the Cantabrian sections examined the nominal species of the LP Biozone are erratically distributed. Reworking is also evident in this part of the sequence and taxa typical of the Homerian and Lower Gorstian occur there. These finds are associated with an incursion of thick sandstones. AGE. Ludfordian to lowermost Pridoli. Chitinozoa in samples from the RS Biozone in the Argovejo, Geras and La Peral sections mainly have a long range, from Mid-Gorstian to Lower Pridolt. Chelinospora hemiesferica (H) Spore Interval Biozone (Figs 2-5) NOMINAL SPECIES Chelinospora hemiesferica (Cramer and Diez) comb. nov.; Cramer and Diez 1975: pl. 2, figs 34-36, (PI. 9, figs 3, 5). PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN LV 1 Ath - Angochitina thadeui Early Gorstian - Late Ludfordian Cb - Cyathochitina sp .B Late Llandovery i Agglomerate Me - Margachitina elegans Mid Pridoli Uu - Urnochitina urna Pridoli - Earliest Lochkovian ff Sandstone s 2 s W shale x = 3 ~ wees S = g C2) Sandstone & Shale oS = 3 = : Spore | Chitinozoan wan R 2 BS 28 3 KO BR i e [X Dolomites Zonation | Zonation LV 17 a8 ahs Seyi Sas gas 2 3 J Limestones = LV 16 UR eg RI SEY Gish ey TS al eg, aS ; MN pense Evins CUS TG Me EN he OS soa on cee mR Cm NGOS ceiver s [-} Quartzites eet LV 14 BRE SESS, REESBSELGESSGME YS _—————— TER Se PS POR SSes s 7 SRS eB BB Sandstone 2 BYAIS SHAS WSS HS SESE Mee ee OA Ss ?su Rete SIMS GEOIERY Qi ust ps coll Clare iis Mali Sol IG Hi Oolites | ele a2 See geo Gog Sig Goa cig SENS a Pri. | xo(8| ae NIQMCRERSICC HS SOE Se Se SS SG Ne eee . = Rg Rs > ow iy Sgskro~ss Sunes — aie) & > £98 SS 5 as > ak a9 3 Qa SNES aa 3 q & = .*Phosphatic nodules / clasts z SSSERSHE AR SASSER PS ORS Sa sac a = Ria tS ROUZ ety SR ISS eee? RUS: SWS SRS Ris Ca oe Ss 2 See ass AS Ole Tinga ana eye) RBBoeoSsHoegst sous os (@)-—Uu |e Pte > Be iva ; Pegeemmaean|* LV 7 aah idoli Ludf. Lv = oe H Pridoli RS tLV 8a # 5 | aa VIG 2 eal = -LV 7b ; E +LV 7a RS 6 ELV 6 o ? So | eT ay || aes ae ee E aacecasay LV saab LV 4 - LV 4 Ludfordian Lian. ly peered LV 3 r IF a i ee eal | IL pLV 3 ; ne oa eae i LV 2 L_cb we EV i- | | fLV 4 Fig. 4 Selected spore distribution at La Vid. Chitinozoan zonation after Verniers ef al. (1995): su: superba, ele: elegans, ko: kosovensis. Coronaspora subornata (Cramer and Diez) comb. nov.; Cramer & Diez 1975: pl. 1, fig. 7, (PI. 8, fig. 7). REFERENCE SECTION. San Pedro Formation, Geras, Province of Le6én, Cordillera Cantabrica, NW Spain. The base of the zone is between samples Ger 5 and 6, and the base of sample 6 is c.16.2 m above the base of the section. The zone is about 22 m thick in the Geras section. DEFINITION. The base of the zone is based on the first appearance of C. hemiesferica (Cramer & Diez). In the lower part of the zone Chelinospora sanpetrensis (Rodriguez), Coronaspora reticulata sp. noy. and Emphanisporites splendens (Richardson & Ioannides, 1973) persist. This biozone represents the interval between the appearance of the nominal species and the first occurrence of Scylospora elegans and Iberoespora cantabrica (EC) Biozone. ACCESSORY SPECIES. Coronaspora subornata (occurs erratically within the zone), Chelinospora canistrata sp. nov. Concentrico- sisporites agradabilis (Rodriguez) Rodriguez, 1983. All the accessory spores have been found in three of the four sections. REMARKS. Chelinospora hemiesferica (Cramer & Diez) shows considerable variation and a morphological series can be traced in these sections. C. hemiesferica (Silurian, Piidolf) has narrow genicu- late muri and higher in the zone is accompanied by C. cf. hemiesferica with broader muri, and then C. lavidensis and C. media, which both have wide lumina and occur along with typical Lower Devonian (Lochkovian) species Chelinospora cassicula Richardson & Lister. So far the variant C. cf. hemiesferica has been found in three of the four sections: Geras (EC Biozone), La Peral (H, and EC Biozones), and La Vid (upper EC (Aneurospora Sub-Biozone), lower MN Sub- Biozone). AGE. Lower and Middle Pridolf. The chitinozoan Urnachitina urna occurs in the basal sample (LP10) of the hemiesferica Biozone in the La Peral section, U. urna has arange from Pridoli to lowermost Lochkovian. Scylaspora elegans — Iberoespora cantabrica (EC) Spore Assemblage Biozone (Figs 2-5) NOMINAL SPECIES Scylaspora elegans sp. nov. (PI. 5, figs 24, PI. 6, fig. 8). Iberoespora cantabrica Cramer & Diez 1975: pl. 2, figs 24, 26-28 (Pl. 6, fig. 4; Pl. 7, fig. 4). REFERENCE SECTION FOR BASE OF BIOZONE. La Vid section, Cantabrian Mountains, Province of Leén, NW Spain. The lowest occurrence of Scylaspora elegans was found in sample LV 10, c. 102 m above the base of the section. DEFINITION. The base of the zone is defined by the incoming of one, or both, of the nominal species. Usually Jberoespora cantabrica occurs stratigraphically above S. elegans. The first appearance of Aneurospora spp., lacking proximal papillae, defines a sub-biozone in the upper part of the zone. ACCESSORY SPECIES. Retusotriletes bipellis Rodriguez, 1978c, Cymbosporites sp. B, and several Chelinospora morphs (variants) 129) with narrow and coarse muri and an irregular reticulum are common accessory spores. Retusotriletes coronadus (Rodriguez, 1983: pl. 8, fig. 1) appears in the lower part of the zone. REMARKS. The upper part of the EC Biozone (Aneurospora Sub- Biozone) is recognisable in England (Hereford and Worcester) in the Raglan Marl Formation below the first appearance of the MN Biozone in the St. Maughans Formation (Richardson, Rodriguez & Sutherland, 2000). In the La Vid section (Fig. 4) the earliest definite specimens of Scylaspora elegans and Aneurospora spp. occur together, consequently the lower part of the EC biozone has not been found there. OCCURENCE. The zone is found in all four sections. AGE. Upper Pridoli to lowermost Devonian. The appearance of the genus Aneurospora is probably close to the base of the Devonian (Richardson, Ford & Parker 1984; Richardson, Rodriguez & Suther- land, 2000). Chitinozoans in the lower EC Biozone are of Middle Pridolfage (basal EC, La Peral, lower EC, Geras and La Vid sections). The same chitinozoans indicating a mid-Pridoli age (Margachitina elegans) and Pridolfto earliest Lochkovian age (Urnochitina urna) are both present in the upper part of the EC Biozone (Aneurospora Sub- Biozone) and in the top of the underlying H Biozone. Aneurospora spp. (A) Sub-Biozone (Figs 2-5) REFERENCE SECTION FOR BASE OF SUB-BIOZONE. Upper San Pedro J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND Formation, La Vid section, Cantabrian Mountains, NW Spain. The lowest occurrence of Aneurospora spp. was found in sample 10, c. 106 m above the base of the section. DEFINITION. Appearance of the genus Aneurospora without proxi- mal papillae. Represents the interval between the appearance of Aneurospora and the incoming of Streelispora newportensis of the succeeding zone. ACCESSORY SPECIES. Spores of the Chelinospora hemiesferica (Cramer & Diez) complex, C. cf. hemiesferica, C. cassicula Richardson & Lister 1969, C. lavidensis sp. nov., C. media sp. nov., Cymbosporites sp. B, and Retusotriletes coronadus s.s. OCCURENCE. Foundinallfoursections inthe Cantabrian Mountains. AGE. Earliest Devonian (Lower Lochkovian), assessment based on comparison with sections in the Lower Old Red Sandstone of England. Streelispora newportensis — Emphanisporites micrornatus (MN) Spore Biozone Richardson & McGregor, 1986 (Figs 2-4); Streelispora newportensis — Leonispora argovejae Spore Assem- blage (NA) Sub-Biozone NOMINAL SPECIES Streelispora newportensis (Chaloner & Streel) Richardson & Lister 1969: pl. 41, figs 3-6 (PI. 7, fig. 8). Spore Chitinozoan Zonation | Zonation Ae -Angochitina echinata Mid Gorstian - Early Pridloi —- ——-—| Cr - Conochitina rudda Late Wenlock - Early Ludfordian Loc.|(na)| a Me - Margachitina elegans Mid Pridoli (a) 3 Ss - Sphaerochitin sphaerocephalaMid Gorstian - Early Pridoli > ; OD 3 S - Spinachitina sp. ?Llandovery HY EC ele ae xs = Lay Pe - Psuedoclathrochitina carmenchui Mid Pidoli = = 3 3 aS} ~ 5 SOR Uu - Urnochitina urna Pridoli - Earliest Lochkovian SSKSegERSON SB SL 7 H 2 Pridoli 3 H ive — ae mm? A . Ger2A&8 o As | = ae = : 2 | Ludf. / E No spores | GOrst. 3 Ea aa | ?Hom. E wvéass | ; ic 17 SN y [2:cb] Fig.6 Correlation of Cantabrian sections. Spore zonation new, except for LP Biozones after Richardson & McGregor (1986), MN Biozone after Holland & Richardson (1977) and Richardson & McGregor (1986), and BV Biozone after Burgess & Richardson (1995); MN: micrornatus — newportensis, (na): newportensis — argovejae Subzone, (a): Aneurospora Subzone, EC: elegans — cantabrica, H: hemisphaerica, RS: reticulata — sanpetrensis, LP: libycus — polymorpha, BV: brevicosta — verrucatus, V: vetusta. Key Cantabrian chitinozoan taxa: Ac: Angochitina chlupaci Lochkoyian, Ae: Angochitina echinata Mid Gorstian — Lower Pridoli, Al: Ancyrochitina libyensis Late Gorstian? — Lower Pyridoli, Ath: Angochitina thadeui Lower Gorstian-Late Ludfordian, At: Angochitina tsegelnuki Lochkovian, Cb: Cyathochitina sp. B Late Llandovery, Cr: Conochitina rudda Late Homerian-Lower Ludfordian, Eb: Eisenackitina bohemica Lochkovian, Me: Margachitina elegans Mid Piidoli. Pe: Pseudoclathrochitina carmenchui Mid Pridoli, Rv: Ramochitina villosa Late Ludfordian, Ss: Sphaerochitina sphaerocephala Mid Gorstian to Lower Piidolt, S: Spinachitina sp. Llandovery?, Uu: Urnachitina urna Pridoli- earliest Lochkovian. Artemopyra brevicosta — Hispanaediscus verrucatus (BV) Spore Biozone (Burgess & Richardson 1995). Both the nominal species are found in samples LP3 and LPS. The fact that Hispanaediscus lamontii also occurs in the latter sample may indicate that the junction between the A. brevicosta and H. lamontii Sub-Biozones lies between these two samples. In south Wales this sub-zonal boundary occurs in the Middle Homerian between the /undgreni and nassa Graptolite Biozones. Chelinospora reticulata — C. sanpetrensis (RS) Spore Biozone. This biozone is represented in all four sections. In the La Peral section both the nominal species appear together in sample LP6. Neither species has been found in the type area of England and Wales. However, the RS assemblages include Concentricosisporites sagittarius and Stellatisporites inframurinatus var. cambrensis first found in the Lower and lower Upper Gorstian respectively, and spores similar to Chelinospora obscura in the Middle Gorstian. All of these species can occur together in the lower part of the cambrensis Sub-Biozone. On current data, therefore, we tentatively, regard the lower boundary of the RS Biozone as lower Upper Gorstian, approxi- mately lower incipiens Graptolite Biozone. Chelinospora hemiesferica (H) Spore Biozone. The base of the hemiesferica Biozone is found in sample LP 10 in the La Peral section. Associated spores are mainly found in the Cantabrian Moun- tains and North Africa. Several species, however, also occur in the Anglo-Welsh area. One of these, Chelinospora poecilomorpha occurs throughout the Ludlow Series in south Wales but has not been found in the Lower Pridoli there. In North Africa this species has a similar range and occurs with Emphanisporites splendens, a co-occurrence also found in LP9. In sample LP12 Amicosporites splendidus occurs and this species is also found in the uppermost Ludfordian to Downtonian (Lower Pridoli) in England but is rare there. The base of the biozone is placed tentatively, therefore, in the Lower Ptidoli but could be also of latest Ludfordian age. Scylaspora elegans — Iberoespora cantabrica (EC) Spore Biozone. The first appearance of Scylaspora elegans occurs in the La Peral section in sample LP 16 coeval with chitinozoans typical of the Middle Pridoli. This is consistent with chitinozoan finds in the sections of Argovejo (Arg 8), Geras (Ger 10), and La Vid (LV 10, LV 11). The base of the zone is therefore placed in the Middle Pridoli with some confidence. Aneurospora (A) Spore Sub-Biozone. The first occurrence of this genus is found in three of the four sections studied: La Peral (sample PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN LP19), Geras (sample Ger 12) and La Vid (sample LV 10). This spore event occurs in England in sections from Hereford and Worcester in strata below the micrornatus — newportensis (MN) Zone. Although the Aneurospora Sub-Biozone in the Cantabrian Mountains occurs a little below (c. 5 m) the occurrence of the typical Lochkovian chitinozoans Eisenackitina bohemica and Angochitina tsegelnjucki (Geras), we regard the base of this Sub-Biozone as close to the Silurian/Devonian Boundary (Richardson, Rodriguez & Sutherland, 2000). This correlation is based partly on comparisons with material from Podolia described by Arkhangel’skaya (1980). In Podolia, below the base of the equivalents of the MN Biozone (Chortkov Group), there is distinct spore assemblage in the Mitkov Formation (Borshchoy Group). This lower assemblage probably contains non-papillate spores of the genus Aneurospora and is above the first occurrence of the graptolite Monograptus uniformis uniformis providing support for the ideathat the Aneurospora (A) Sub-Biozone is Devonian. Based on the distribution of key chitinozoan species, Paris & Grahn (1996) used Pacific Ocean 0 1000km —@ — ee Chukotka ay © A) Sy 125 the base of Monograptus uniformis uniformis, the higher of the two critical graptolite subspecies, to mark the base of the Devonian. In particular Eisenackitina bohemica, also found by us in the Cantabrian Mountains, and otherrestricted chitinozoan species occur in the upper Tajna Formation along with M. uniformis uniformis. Although it is difficult to be certain, because of the nature of the spore illustrations (Arkhangel’skaya, 1980), this Lower Podolian spore assemblage appears to include non-papillate species/varieties of Aneurospora and may, therefore, be equivalent to the Aneurospora Sub-Zone. Assem- blages with tripapillate spores (Aneurospora/Streelispora?) occur in the Chortkoy Group c. 70 m above the Mitkov assemblage. The Silurian/Devonian Boundary is currently placed at the first appear- ance of M. uniformis angustidens at base of the Borshchov Group (Tajna Formation, Nikiforova, 1977) but no spore assemblages have been described from the lower part of the Borshchov Group. Even so, the position of the Silurian/Devonian boundary as determined by Paris & Grahn seems reasonable from the spore evidence currently available. Lomonosov Fig.7 Piidolf palaeotectonic-palaeogeographic map. Simplified from Ziegler (1988). 126 LAURUSSIA - GONDWANA SPORE DISTRIBUTION PATTERNS (FIGS 7-9) Upper Silurian and Lower Devonian spore floras are diverse and homosporous but little is known of their geographical dispersal. Silurian palaeogeographical maps show Europe and North America at low latitudes, between the Tropic of Capricorn and the equator (Witzke & Heckel, 1988), separated by an ocean from Gondwana (with Iberia in close proximity on its northern flanks) in higher southern latitudes (McKerrow & Scotese, 1990). The map (Fig. 7) based on Ziegler (1988) shows the Cantabrian Terrane separated from Gondwana by Proto-Tethys and on its northern flank the Rheic Ocean lies between the Aquitaine Cantabrian Terrane and northern France and Britain. McKerrow & Scotese show a wedge-shaped ocean narrowing westward to the point where the Americas were in contact. Using this reconstruction, and unless there was a major climatic barrier, there would have been a possible land plant migra- tion route between Laurussia and Gondwana. With favourable winds, airborne transport would have been feasible for many of the spores Proto Arctic Ocean Proto Arctic \ Ocean < & = a4 8 é E cS o 2 12) 4, z seocl. athon - Ouachitas°® J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND because of their small size (Mogensen, 1981) and it is worth noting that many moss species have achieved wide dispersal in the northern hemisphere since the last ice age, some 10,000 years ago (Schofield, 1985). Such a time span is negligible in terms of geological time and is essentially instantaneous. Theoretically therefore, since there was no major physical barrier to dispersal apart from the availability of suitable habitats, and possibly climate, large parts of the tropical flora should have been uniform. As size affects dispersal (Mogensen, 1981), smaller spores, corresponding to homosporous plants with spores less than 25 um, and appearing in the Pridoli and lowermost Lochkovian, should have had the widest distribution. Before the Pridloli, multiple unit cryptospores from the Llandovery and Wenlock, and to a lesser extent in the Upper Ordovician are apparently closely similar and widespread. Such assemblages occur in North America, Europe, North Africa, the Middle East and China. Was this wide dispersal due to efficient spore dispersal mechanisms, closer prox- imity of the main continents when their parent plants evolved, climate, or some other factors? The Ordovician data may not be representative, as few assemblages have been described in detail, but the uniformity of Lower Silurian cryptospore assemblages form a Ural Ocean Fennosarmatian High Pe ae Cantabrian Positive areas 3 Convergence direction Fig. 8 Emsian palaeotectonic-palaeogeographic map. Simplified from Ziegler (1988). PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN 30° Ludlow - Pridoli B05 0° 30° Early Devonian 127 ==> Cold current =P Warm curent He Lana Barrier Fig.9 Oceanic circulation through the Ludlow-Lower Devonian. Based on Le Hérissé et al. (1997). contrast to some of the Upper Silurian trilete associations showing regional variations in species distribution. Well illustrated accounts of spore data from Upper Silurian- Lower Devonian sequences are limited to comparatively few areas, viz. Britain (Richardson & Lister, 1969), the Cantabrian Mountains (Rodriguez, 1983), and North Africa (Richardson & Ioannides 1973, and work in progress). More limited stratigraphical data are avail- able from Belgium, France, Poland, and China. Upper Silurian and Lower Devonian spore assemblages have been described from graptolite-bearing sequences only in England and Wales (Richardson & Lister, 1969, Burgess & Richardson, 1995) and Podolia (Arkhangel’skaya, 1980). However, many of the published illustra- tions of the Podolian material are insufficient for detailed comparisons and key taxa cannot be identified with certainty at the species level. Nevertheless the assemblages enable general correlation of these important graptolite-bearing sequences with Britain. The following discussions are therefore based mainly upon the authors’ previous and current work, and consequently the present discussion is depend- ent on data from a limited geographic area representing three separate blocks, namely Laurussia, Iberia-Aquitaine and northern Gondwana. Several spore taxa (eg. Streelispora newportensis), although rare in parts of Gondwana, appear to be geographically widespread. Some cryptospore taxa, eg. Tetraletes variabilis Cramer 1966a, may have a restricted distribution and so far have been recorded only from Gondwana, Iberia and Brittany, whereas other cryptospores (PI. 1, figs 1, 7, 9-11) occur in Laurussia as well. The planar tetrads of Tetraletes occur in Silurian-Lower Devonian sections from north- west Spain and North Africa, with a single specimen recorded from Brittany (d’Erceville, 1979). It is interesting to note that similar planar tetrads are found in the modern thalloid liverwort Riccia perssonii from South Africa (Perold, 1989). Both the fossil Tetraletes and the modern Riccia spores are permanently adherent and are joined by more or less smooth bands of attachment separating areas of spinose sculpture. Many cryptospore taxa are found in the Silurian of North America, Britain, Western Europe, the Baltic States, Podolia, Spain, and North Africa, whereas others are apparently regionally confined. As more records are forthcoming it should be possible to 128 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN determine how many of these ubiquitous spores had evolved before the break-up of the southern continent. On the other hand, trilete species, which evolved after many of the regionally dispersed cryptospores (e.g. Tetrahedraletes medinensis (Strother & Traverse), Pl. 1, fig. 10), apparently show a dichotomy with several distinctive trilete species currently known only from the Cantabrian Mountains and North Africa. As more spore data accumulates, therefore, the dispersal patterns of early land floras may provide constraints to palinspastic reconstructions. In addition, new insights into the corre- lation of marine with non-marine plant-bearing strata, will help to date more accurately events in land plant evolution, recorded in greatest detail in continental rocks. Further, the joint study of chitinozoans and spores in the same sequences will eventually permit an understanding of inter-facies correlation, not only in marine shelf environments, but also with stratotype sequences mainly established in sediments deposited in distal marine environments where spores are often rare and badly preserved. There is some support from the chitinozoan data for the changing regional differences in the spore data outlined above. Though the nature of sampling for chitinozoans (see below) makes any direct comparisons difficult. However, there does appear to be an increas- ing similarity between chitinozoan assemblages of northwest Spain and North America/Europe in the upper/later parts of all four sec- tions studied. Due to a lack of data it is not possible to determine if there is a corresponding ‘dissimilarity’ with Gondwanan assem- blages. Paris et al. (1995) noted similarities between chitinozoan assemblages in northern Spain and Gondwana and dissimilarity with coeval assemblages in Baltica during the Llandovery. This evidence was used to support the existence of a wide mid-European Rheic Ocean during the Llandovery. Such temperal changes in our assemblages, i.e. a greater similarity with Gondwanan spore floras in the Lower Silurian, ard increasing similarity with Baltic/Avalonian assemblages in the Upper Silurian/ Lower Devonian, would be consistent with a drift of northwest Spain away from Gondwana and towards Laurentia/Avalonia through this period. CHITINOZOA General distribution In sections where the Formigoso Formation is represented (Argovejo and La Vid) chitinozoans belonging to the genera Cyathochitina and PLATE 1 129 Conochitina are common. The lower parts of the San Pedro Forma- tion often contain Angochitina and Sphaerochitina with species belonging to Plectochitina (e.g. Plectochitina caminae Cramer & Diez 1978a, figs 8q, 8r and Plectochitina rosendae Cramer & Diez 1978a, fig. 8p) and heavily ornamented species such as Ancyrochitina Jjavieri Schweineberg 1987 (figs 8f, g) becoming more common in the central parts of the formation. Certain species such as Calpichitina velata Wrona (1980: figs 6i, j) and Margachitina catineria Obut (1973: fig. 6r) occur in ‘blooms’ at certain horizons, for example C. velata at La Vid (LV6) and M. catenaria (PI. 11, fig. 13) at Geras (Gerl6, 17 and 18). In addition, the chitinozoans Vinnalochitina horrentis (Wrona, 1980), Cingulochitina ervensis (Paris, 1979) and Cingulochitina serrata (Taugourdeau & de Jekhowsky, 1960) form common components of assemblages in the San Pedro Formation (see PI. 11, figs 8, 11a, 11b and PI. 12, fig. 9 respectively). Geras (Fig. 10) Unlike the other three sections, no Llandovery chitinozoans were recovered. Sphaerochitina sphaerocephala (Eisenack 1932) and Angochitina echinata Eisenack 1932 (Pl. 12, figs 6 and 10 respec- tively) were used as accompanying species in the Global Chitinozoan Biozonation of Verniers et al. (1995). Their presence in Ger 2b, 1.5 metres above the base of the section, suggests a mid-Gorstian to Lower Pridoli age for the lower part of the San Pedro Formation at Geras. Greater resolution is provided by Ramochitina villosa (Laufeld, 1974) (recovered from sample Ger 2a, 2.4 metres above the base of the section and Ger 2b; PI. 13, fig. 7) which has been reported from the top of the Lower Leintwardine Formation and the Upper Whitcliffe Formation of the type Ludlow, UK (Sutherland 1994: 65) and from strata of similar Upper Ludfordian age on Gotland (Hamra and Sundre Beds, Laufeld, 1974: 96) and Estonia (Kuressaare Re- gional Stage, Nestor, 1990: 85). Ancyrochitina valladolitana Schweineberg 1987 (figs 8d, e) was recovered from Ger 4, 10 m above the base of the section within the San Pedro Formation. This species was described from the Palencia region of Northern Spain by Schweineberg (1987) and assigned to graptolite biozones 34/35 (Gorstian) of Elles & Wood (1901-18). However, it was noted that the chitinozoan species probably ranges into the Pridolif (Schweineberg 1987: 77). Ancyrochitina javieri Schweineberg (1987: figs 8f, g) has also been recovered from Ger 4. This species was described as occurring between graptolite biozones 32 to 35 (late Homerian-Lower Gorstian) of Elles & Wood (1901-18) in the Palencia Region of Northern Spain Figs 1,2 Artemopyra? sp. A. 1, BM 137698, oblique compression showing radial folds on the proximal and distal surfaces adjacent to the curvatural ridge, x 2000, stub Ger92/9/1; 2, BM 130612, specimen with more delicate muri than that in Fig. 1, oblique compression, x 2000, sample Ger92/8. Fig. 3 Fig. 4 Fig. 5 Fig. 6 Fig. 7 Fig. 8 Fig. 9 geniculate disjointed muri, x 2000. Chelinohilates sp. BM 137155, dyad showing distal and subequatorial muri, sample Geras 92/9/1. Cymbohilates cf. allenii var. magnus Richardson 1996. BM 115329, proximal view showing remnants of hilum, sample Arg92/14. Hispanaediscus cf. lamontii Wellman 1993. BM 135296, proximal view, x 2000, sample Ger92/2B/2. Cymbohilates cf. horridus Richardson 1996. BM 130858, sample LV92/13; 6a, proximal view; 6b, detail of spines; x 2500. Hispanaediscus lamontii Wellman 1993. BM 137471, dyad, sample Arg13/D1. Tetraletes variabilis Cramer 1966a. BM 137848, tetragonal (cruciform) tetrad, sample Ger92/6. Velatitetras rugulata? Burgess 1991. BM 465, LV92/6; 9a, tetrad, lines of attachment probably covered by closely adherent envelope; 9b, detail of Fig.10 Tetraletes medinensis (Strother & Traverse) emend. Wellman & Richardson 1993. BM 130623, showing lines of attachment and microgranulate distal surfaces, sample Ger92/8/1. Fig. 11 Pachytetras sp. BM 138111, LV92/8/DD; 11a, unfused tetrad showing sculpture sloughing off revealing lines of attachment; 11b, BM 138113, detail of microverrucate-murornate sculpture, x S000. Fig.12 Tetraletes variabilis? Cramer 1966a. BM 132691, LV92/10; 12a, BM 132691; 12b, detail of microverrucate sculpture, x 2000. All are Scanning Electron Photomicrographs at x 1000 unless otherwise stated; BM numbers refer to photographic negatives in the archives of the Natural History Museum, London. 130 Sphaerochitina sphaerochepala cyrochitina valladolitana Angochitina echinata Angochitina ambrosi ambrosi Ancyrochitina fragilis brevis Cingulochitina serrata Palenchitina pisuergensis Angochitina elongata ©! Cingulochitina ervensis Margachitina elegans Be Ramochitina villosa J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND Sandstone Mishale (Sandstone & Shale =. Phosphatic nodules Kj Dolomites B Limestones BB Sandstone / Oolites Quartzites B Ferrous Oolites Pseudoclathrochitina carmenchui Urnochitina urna Vinnalochitina horrentis Plectochitina carminae Eisenackitina bohemica Calpichitina velata Margachitina catenaria Spore Zonation Chitinozoan Zonation =) a a) @ pa co aie An | Lochkov. ] Bono) Ancyrochitina javieri c £ 5 ee 2) 5) o 5 n Ludford. LP c= = = Fault Fig. 10 Selected chitinozoan distribution at Geras. Chitinozoan zonation after Verniers er al. (1995): bo: bohemica, su: superba, ele: elegans, ko: kosovensis, ba: barrandei, ph: philipi. (Schweineberg, 1987: 70). A similar species, Ancyrochitina sp. A, was described by Paris from the Formation du Val in the section at Heuzé, Brittany (Paris, 1981) and is associated with Pterochitina perivelata Eisenack (1937), a chitinozoan with a well established range from the Upper Ludfordian to the Upper Pridoli (barrandei Biozone to early part of the superba biozone of Verniers etal. (1995). The presence of Angochitina elongata Eisenack (1931, figs 8h, 1) in sample Ger 9 may be the result of reworking. Urnochitina urna (Eisenack, 1934) is a very useful Pridoli marker in the stratotype area (Barandian region, Central Bohemia). U. urna occurs very close to (within 50 cm) or at the base of the Pridoli (although some atypical specimens do occur a few centimetres below the first occurrence of Monograptus parultimus (Kriz et al., 1986). U. urna is recovered from the Geras section at Ger 10, 55 metres above the base of the section in the San Pedro Formation. In addition to U. urna, Pseudoclathrochitina carmenchui Cramer (1964b) and Margachitina elegans (Taugourdeau & de Jekhowsky, 1960: pl. 7, figs 92, 93) also occur in Ger 10. Both species are important markers for the mid-Pridoli elegans Biozone of Verniers et al. 1995. In Bohemia, Eisenackitina bohemica (Eisenack, 1934) occurs 50cm above the base of the Devonian at Klonk, the Silurian/Devonian boundary stratotype, and Karlstejn (Paris et al., 1981). At Geras, E. bohemica 1s recovered from Ger 14, 70 metres above the base of the section in the San Pedro Formation (Pl. 11, fig. 14). It is found associated with Angochitina tsegelnjucki Paris & Grahn (1996, fig. 7r) (PI. 12, fig. 8). In Podolia this species is reported from the upper part of the Mitkoy Formation which, on the basis of its association with E. bohemica and Margachitina catenaria Obut (1973), was assigned to the Lochkovian by Paris & Grahn (1996). In the highest part of the section, between 74 and 76 m above the base (samples Ger 16, 17 & 18), significant numbers of a species resembling M. catenaria of Lochkovian age occur consistently (see p. 134). However, only a tentative assignment is made here to the Cantabrian forms as no individuals were recovered that demon- strated a convincing peduncle. Stage and Series Boundaries On the basis of chitinozoans the Silurian/Devonian boundary would be placed below the first occurrence of E. bohemica at Ger 14. The Ludlow/Pridoli boundary is less easy to define but will occur some- where between the first occurrence of typical Pridoli chitinozoans (U. urna, P. carmenchui and M. elegans) in Ger 10 but above the last occurrence of R. villosa in Ger 2b. Argovejo (Fig. 11) An Upper Llandovery age for the Formigoso Formation at Argovejo (Argovejo 1—3; 21—-10.5 metres below the base of the San Pedro Formation) 1s suggested by the presence of Cyathochitina sp. B Paris (1981: pl. 11, fig. 11) (PI. 11, fig. 12). This species is reported from Upper Llandovery strata in the Massif Armorican by Paris (1981: 299). The discovery of the Llandovery chitinozoan Cyathochitina elenitae Cramer (1964b) in sample Arg 3 (PI. 11, fig. 6) supports this age assignment. The latter species (PI. 11, fig. 5) was also recovered with another Llandovery chitinozoan, Conochitina alagarda Cramer 1967 in Arg 4 (Pl. 11, fig. 2). However, as both of these species are found associated with spores from the RS Biozone (late Ludfordian- Lower Pridoli) they may represent reworking. Ancyrochitina libyensis Jaglin (1986) was recovered from Arg 4, PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN La Vid Formation IN Dolomites ro Limestones WB Sandstone / Oolites Mi Ferrous Oolites s 3 = 3 x 5 <= > Az) S > = = F 3 seh 2s anes “24 5 i 3 Se ,s 428e = 8 = S2eS8u 8, Ss (J Red Sandstones f.} Dolomites VG Sos Bay ed os OS 2a gu 8S8 ey LV 12 Re) aS g Bs ese 5 = 2 8, fs -° Phosphatic nodules /clasts [Gj/Limestones a ¢ Ha SF Ss os iS) SPF : eo V8) > 5s 8 Sas Ege g8 3 f] Agglomerate [7] Quartzites g PSTD iv io aos SS GLRSS SRP SIR 3 Peer SESPesSSeeta lasses SSS CHESSER ES SSE SES e SS 3 = mm & S RS a 5 82 % y %& 8 ° Chitinozoan Spore NN 3 Q F F 2 A = & See nee oS aes as 3 Sa eh me s SSOa< Zonation Zonation = LV 18 - Linon nye alc a LV17 - il i Lop ay LV8A LV16 - a | | | | LV 16 2) Die [| LN Ts) Lochkov. ae - || ry ie Lochkovy. : e ares 9 }-——- LW) | LV 12 Middle i =e rvs) LV 1 = . Oval Pridoli toes EC NY NY = LV 10 = a eee ko = ps EVES LV9 aoe a SVT TY BB _ LV 8B - LV6 ater LV 8A LV 8A Pridoli Ni aa es : tae LNT ?Ludlow Le a r= LV 6 oe’) ce) won|? — Z LV 5b | + LV5b ciate : 2 : | ed | LV 5b LN Oa LV 5a Gorst. Ludford. & = | | Lv 5a aes e LV 4 [Lv4 ar ay LV 3 = =) rvs LV2-. LV2 "a a ?Llan Z\ [ivi LV! -|file LV1 Fig. 12 Selected chitinozoan distribution at La Vid. Chitinozoan zonation after Verniers ef al. (1995): su: superba, ele: elegans, ko: kosovensis. LV1 and LV2 (42 m and 38.8 m below the base of the San Pedro Formation) suggests an Upper Llandovery age for the Formigoso Formation. Angochitina echinata E1senack (1931) and Angochitina thadeui Paris (1981) (Pl. 12, fig. 11) were recovered from LV 5a. The latter has been reported from strata assigned to Graptolite Biozones 32 and 33 (Homerian to Lower Gorstian) of Elles & Wood (1901-18) by Schweineberg (1987: 59). A Llandovery age is also supported by the presence of C. elenitae in LV1. The recovery of this species (39 metres above the base of the San Pedro Formation at the level of sample LV 7b) is probably the result of reworking. A. echinata is a common species in the Middle Gorstian to Lower Pridoli (Verniers er al. 1995). Typical Piidoli chitinozoans (e.g. Urnochitina urna—PI. 12, fig. 4), associated with Plectochitina (PI. 13, fig. 8), arenotrecovered until LV 10, 102.2 metres above the base of the San Pedro Formation. Margachitina elegans (PI. 12, fig. 5), used as an index species for the mid-Pridoli by Verniers et al. (1995), is first encountered in LV11, 107.8 m above the base of the San Pedro Formation. Ancyrochitina fragilis brevis Taugourdeau & de Jekhowsky (1960) was recovered from LV 10(PI. 13, fig. 5). This species was regarded as a ‘late Ludlow’ species by Cramer & Diez (1978a). Samples LV 10, 11, and 12 are regarded as lowermost Devonian based onthe spores butno chitinozoans were recovered that could be positively identified as Devonian. The spores in sample LV 13, c. 1 m above LV 12, are typical of the lower MN Biozone (NA Subzone) of Lower Lockhovian age. Stage and Series boundaries No Devonian chitinozoans were recovered from La Vid and typical Pridoli forms such as Pseudoclathrochitina carmenchui and M. elegans were still recovered from the top of the section. However, the first appearance of Aneurospora is used tentatively to place the base of the Devonian below LV10 (see also Richardson, Rodriguez & Sutherland, 2000). The base of the Pridoli is placed somewhere below LV 10 with the first occurrence of U. urna. As with Argovejo, the presence of Upper Llandovery chitinozoans in the Formigoso Formation may suggest the presence of a disconformity or the reworking of Lower Silurian faunas. The spacing of some samples, because of unsuitable lithology and poor recovery from others, makes a more precise biostratigraphy difficult at this time. La Peral (Fig. 13) Because Spinachitina has not been reported above the Llandovery, the presence of Spinachitina sp. in LP2 (Pl. 11, fig. 4) 18 meters above the base of the San Pedro Formation may suggest either a pre-Wenlock age for this part of the section or possibly evidence for reworking. Conochitina rudda Sutherland (1994), present in LP3, 24.6 metres above the base of section, has been described from the Upper Wenlock/Lower Ludlow of the type area in the Welsh Borderlands (Sutherland, 1994: 48). Conochitina pachycephala Eisenack 1964, present in LP3 (PI. 11, fig. 1), 1s used as an index species for the mid-Upper Wenlock but also ranges well into the Gorstian (Verniers ef al. 1995). The presence of Cyathochitina elenitae (P1. 11, fig. 3) in the same sample may be the result of reworking. The recovery of Angochitina echinata (Pl. 12, fig. 10) suggests an age of at least Middle Gorstian to Lower Pridoli (Verniers et al., 1995) for LP 8, 77.5 m above the base of section. A. elongata, recovered from the same sample, was used as a mid-Ludlow zonal indicator in the biozonation of PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN Red Sandstones [i Sandstone fj Dolomites BB Shale 3 = Fj Limestones [J Sandstone & Shale = g 2 5 fj Quartzites BB Sandstone / Oolites = 8 5S § 5 ater 2 S aces » . = -& S|s-*Phosphatic nodules / clasts Ml Ferrous Oolites = $ ~~ =— S re vw PI = SS Bs RES eats 5S fay WS 2 WS sss aes SSPE ES sess 1-1.5um high and wide. Proximal equatorial areas, outside the contact areas, and distal surface laevigate. DIMENSIONS. 28-43 um. COMPARISONS. This species is distinguished by curvaturae PLATE 3 Fig. 1 Retusotriletes? saturnus sp. nov. BM 129970, proximal view, sample Arg92/10. Fig.2 Telisporites sp. A. LV92/13; 2a, BM 130849, proximal view, x 500; 2b BM 130850, distal spines and curvatural ridge; 2c BM 130851, distinct sculpture on contact area, x 2500. Fig. 3 Insolisporites sp. Ger92/2B; 3a, BM 134643, proximal view showing distal sculpture and proximal apical cones; 3b, BM 134644, detail of proximal apical sculpture, x 2000. Fig.4 Raistrickia sp. Ger15B; 4a, BM 130714, proximal view, x 2000; 4b, BM 130713, detail of baculate sculpture and labrate laesurae, x 2500. Fig.5 Scylaspora vetusta (Rodriguez) comb. nov. Ger2B/1; 5a, BM 135149, obliquely compressed specimen with proximal crenulate muri near the equator; 5b BM 135150, detail of proximal sculpture x 2000. Fig.6 Emphanisporites cf. decoratus Allen 1965. BM 132902, slightly oblique proximal view showing spaced distal cones, sample LV 13/1. Fig.7 Emphanisporites rotatus McGregor 1961. BM 134062, proximal view, Arg92/5A. Fig. 8 Emphanisporites ct. splendens Richardson & Ioannides 1979. BM 134633, proximal view?, showing annulus, apical thickening and radial muri, Ger92/2B. Fig.9 Emphanisporites splendens Richardson & Ioannides 1979. 9a, BM 140339, proximal view showing well-developed paired lips, faint radial muri and three prominent inter radial thickenings, x 2000, LP10/DD; 9b, BM136882, proximal view showing a kyrtome-like structure, Ger2B/1, x 1000. All figs x 1000, unless stated otherwise. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND 140 PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN perfectae formed from sculptural elements and absence of sculpture outside the contact areas towards the amb and over the distal surface. OCCURRENCE. Rare in the Cantabrian Mountains, Argovejo, La Vid, hemiesferica (H) to micrornatus — newportensis (MN) Biozones. Pridoli to Lochkovian. In England (Shropshire & Her- efordshire) where this species occurs in the Lower Pifdoli and spores of this type have been found also in sporangia (Fanning et al., 1991: 3.2 b). Similar spores are found higher in the sequence in the Lower Lochkovian but the higher spores have a variably greater cover of apiculate to spinose sculpture; in extreme forms the spore is com- pletely covered with cones/spines (Richardson, unpublished). REMARKS. In the case of Form 2 the curvatural bands are defined by sculpture, sometimes barely discernible, sometimes forming broad bands of variable grana and coni paralleling the trilete mark and merging radially into curvatural sculpture. This species, Apiculiretusispora arcidecus sp. nov. occurs in the Lower Pridoli of England and has been found in sporangia (Fanning et al., 1991). Stratigraphically higher spores are variable but include specimens with more pronounced sculpture. There is considerable variation; variants in the Lower Devonian have dense distal sculpture, whereas the proximal surface is completely covered with cones, or short spines, with only three small inter-radial areas remaining smooth (interradial papillae). A third species, Breconisporites? cf. coronadus shows that zonate spores may also have a distinct proximal pattern of sculpture similar to that of A. arcidecus sp. nov. Genus BRECONISPORITES Richardson et al. 1982 Breconisporites? spp. (cf. Retusotriletes coronadus Rodriguez, 1983, pars) Pl. 2, figs 5—7; Pl. 4, figs 2-4 This form has equatorial wedge-shaped curvaturae extending equatorially (into a zona?). On the proximal surface there is a band of spines and cones in each of the inter-radial areas. These sculptured bands parallel the curvaturae and join with the sculpture which parallels the lips. The ‘curvatural’ sculpture is composed of fimbriae, spines and cones. Sculpture size c. 1-6 um. The proximal central area, within the arcuate sculpture, may be depressed. There is a +/— circular thickening over the distal polar area. REMARKS. Specimens of Breconisporites from South Wales (lo- cus typicus) (Richardson et al., 1982) have a bizonate cingulum and some species have a proximal area of coni; the Cantabrian spores 14] have some similarities but are not distinctly bizonate. Rare laevigate or proximally rugulate zonate spores that are more clearly bizonate occur near the top (sample LV 13) of the sampled part of the La Vid Section (PI. 4, fig. 5). In specimens with prominent proximal spines (PI. 4, fig. 3) the distribution of the spines, in arcuate, ‘curvaturate’ zones, linked to areas of spines bordering the lips, is similar to the pattern of sculpture seen in Apiculiretusispora arcidecus sp. nov. where the sculpture is much finer. The curvatural thickening is extended to form a zona and there are three arcuate bands of spines and cones on the proximal face. The latter spores, and the heavily sculptured spores of type 2, have been found only in the Devonian so far, and form part of a complex morphon, which is being further, investigated. Infraturma MURORNATI Potonié & Kremp 1954 Genus EMPHANISPORITES McGregor 1961 TYPE SPECIES. fig. 7). Emphanisporites rotatus McGregor 1961 (PI. 3, Emphanisporites splendens Richardson & Ioannides 1979 Pl. 2, fig. 8, Pl. 3, fig. 9 1973, Emphaenisporites? pseudoerraticus Richardson & Ioannides: 275-76, pl. 3, figs 12-15; pl. 4, figs 1-4, 7. Emphanisporites? pseudoerraticus Rodriguez: 417, pl. 2, fig. 17. 1979 Emphanisporites splendens Richardson & Ioannides: 111. 1983 ?Emphanisporites pseudoerraticus Rodriguez: 40, 41, pl. 3, fig. 20. 1978c DIMENSIONS. 26—63um (17 specimens); 50—78um (24 specimens) (Richardson & Ioannides 1973). REMARKS. The Cantabrian and North African specimens are highly variable, but the specimens in this study include much smaller forms than any found in North Africa. Also the ‘annulus’ is sometimes formed of three inter-radial tangential thickenings which together form a triangle resembling a proximal kyrtome. However, in some of the Cantabrian and the previously described North African speci- mens, there is no annulus but only irregular inter-radial thickenings. In other specimens (PI. 3, fig. 8) with an annulus, radial ribs, and polar, or near polar, thickenings that may be proximal, there is no clear trilete mark. These are referred to herein as Emphanisporites cf. splendens. PLATE 4 Fig. 1 Aneurospora richardsonii (Rodriguez) comb. nov. Tilt 45°, Arg97/13/1P; la, BM 2118, oblique proximal compression; 1b, BM 2119, showing distal coni, x 2000; 1c, BM130290, specimen in oblique compression showing rigid equatorial crassitude, Arg/92/13. Fig. 2 Breconisporites sp. A. Sample La Vid 92/13/1; 2a, BM 132895, proximal view; 2b, BM 132896, distal proximal sculpture, x 2000. Fig. 3 Breconisporites sp. C. LV92/13/1; 3a, BM 132899, proximal view; 3b, BM 132900, spinose proximal sculpture forming zones parallel with the equator and paralleling the trilete sutures, x 2000. Fig.4 Breconisporites sp. B. BM 130860, proximal view showing rugulate sculpture, x 500, LV92/13. Fig.5 Zonate spore. Proximal view, specimen with laevigate contact face, BM 130859, LV92/13. Figs 6,10 Chelinospora cf. hemiesferica (Cramer & Diez) comb. nov. 6, BM 130308, Arg92/10; 6a, BM 130307, oblique compression; 6b, detail of radial folds around the hilum, x 2000; 10, BM 132469, oblique view with spaced uncluttered distal muri, LP92/19. Fig.7 Chelinospora cantabrica sp. nov. BM 135304, partial tetrad, upper specimen distal view showing large broad lacunae, lower specimen giving a partial proximal view of the equator surrounding hilate area and subequatorial sculpture, Ger92/2B/2. Fig.8 Chelinospora sanpetrensis (Rodriguez) comb. noy. 8a, BM 136891, distal view showing low murornate sculpture and narrow, occasionally branching lumena, Ger92/2B/3; 8b, BM 135042, proximal/oblique view showing granulate — micro rugulate curvatural sculpture, thin broken hilum, and subequatorial radial lacunae, Ger92/2A; 8c, BM 134063, spore tetrad showing sub-equatorial radial lacunae, sample Arg92/5A. Fig.9 Chelinospora hemiesferica (Cramer & Diez) comb. nov. BM 132687, oblique view, LV92/10. All figs x 1000, unless stated otherwise. 142 OCCURRENCE. Lowerand Middle San Pedro Formation, RS and H Biozones; Upper Ludfordian and Lower Piidoli, possibly some specimens are reworked. Subturma ZONOTRILETES Waltz 1935 in Luber & Waltz 1938 Infraturma CRASSITI Bharadwaj & Venkatachala 1961 Genus AMBITISPORITES Hoffmeister 1959 Ambitisporites warringtonii (Richardson & Lister, 1969) comb. nov. BASIONYM. Retusotriletes warringtonii Richardson & Lister 1969. 1969 =Retusotriletes warringtonii Richardson & Lister 1969: pl. 37, figs 7, 8. REMARKS. The curvyaturae are thickened and form a crassitude that is more or less coincident with the equator. These forms are closely similar to A. avitus except for the size and thickness of the crassitude. Ambitisporites? eslae (Cramer & Diez, 1975) comb. nov. JL, 5 tora, BASIONYM. Retusotriletes eslae Cramer & Diez 1975 (pars), 343, pl. 1, fig. 11 only. 1973 Ambitisporites sp. B, Richardson & Ioannides: 277, pl. 6, fig. 8 only. 1975 Retusotriletes eslae Cramer & Diez (pars): 343, pl. 1, fig. 11 only (see discussion under Scylaspora elegans sp. nov.). 21976 Retusotriletes maculatus McGregor & Camfield: 26, pl. 1, fig. 6. 21995 Ambitisporites tripapillatus Moreau-Benoit; Burgess & Richardson: 16, pl. 6, fig. 16. HOLOTYPE. Cramer & Diez 1975: pl. 1, fig. 11. COMPARISON AND REMARKS. A. tripapillatus Moreau-Benoit (1976: 37, pl. 7, figs 2-4) has a darkened area along the Y-rays at the spore apex with straight to concave sides. Scylaspora elegans has a large darkened apical area and proximal rugulate sculpture. Retusotriletes maculatus McGregor & Camfield (1976) and, in part, Ambitisporites sp. B (McGregor & Camfield, 1976: pl. 6, fig. 8 only), appear to have a narrow equatorial crassitude, but are otherwise similar to A. eslae (Cramer & Diez). They all show an equatorial crassitude except at the radial apices where the curvaturae invagi- nate. A similar feature is seen in some specimens of Ambitisporites, eg. A. avitus. Several Lower Devonian laevigate spores are proxi- mally retusoid (McGregor & Camfield, 1976) and from the limited data available these spores appear to be pandemic, although many of J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND the records are from Spain, North Africa and South America. The age of similar spores to this species is varied. A. eslae, A. tripapillatus and R.. maculatus are all from the Lower Devonian but the Burgess & Richardson record is of Lower Pridoli age. OCCURENCE. Lochkovian). Argovejo, EC (a) subzone and MN zone (Lower Genus SCYLASPORA Burgess & Richardson 1995 TYPE SPECIES. Scylaspora scripta Burgess & Richardson 1995. REMARKS. Some of the spores placed in the genus Rugosisporites by Dufka (1995) resemble those of Scylaspora; however, the type species of Rugosisporites is Retusotriletes chartulatus McGregor (1978) which is retusoid with distinct invaginations at the radial apices. McGregor’s species is similar, but not identical, to some Middle and Upper Devonian species also placed in the genus Retusotriletes, namely R. rugulatus and R. phillipsii. Consequently, since the type species of Rugosisporites is regarded here as belong- ing to Retusotriletes, as McGregor determined, then removal of the type species necessitates the invalidation of the genus Rugosisporites by placing it in synonymy with the genus Retusotriletes. Scylaspora elegans sp. nov. Pl. 5, figs 2-4, Pl. 6, fig. 8 1975 Retusotriletes eslae Cramer & Diez (pars): 343, pl. 1, fig. 12 only. 1983 = Retusotriletes eslae Cramer & Diez; Rodriguez, 1983: 47, pl. 5, figs 14, 19, 20. HOLotTyPe. FM 1490(PI.5, fig. 2), diameter 43 um; sample La Vid 13, slide D3/2, co-ord. 090 1036; E.F. no. J34/3; upper San Pedro Formation, La Vid section, Cantabrian Mountains, Province of Leon, northwest Spain. DERIVATION OF NAME. Latin a. elegans, refined, elegant, referring to the delicate proximal sculpture. DIAGNOsIs. A Scylaspora with a narrow +/— equatorial curvatural crassitude, three distinct proximal inter-radial papillae, Y-sutures bordered by darkened (?thickened) narrow areas tapering to the equator, proximal surface with contorted microrugulae, accompan- ied by short radial microrugulae near the curvaturae, microrugulae become less distinct polewards; distal exine laevigate. DESCRIPTION. Amb rounded to subtriangular. Proximal surface with straight to sinuous microrugulae adjacent to the curvaturae, <0.5 um wide and high, and decreasing in height towards the proximal pole; straight microrugulae confined to the curvatural area. Under the SEM the microrugulae are close-packed and contorted PLATE 5 Fig. 1 Ambitisporites? eslae (Cramer & Diez) comb. noy. FM 1505, Arg92/12 (510) 3, 206 1169. Figs 2-4 Scylaspora elegans sp. nov. 2, FM 1490 (holotype), proximal view showing dark apical triangle, narrow crassitude, curvaturae perfectae and inter-radial papillae, LV92/13 (D3) 2, 0901036; 3a, 3b, FM 1506, 3b, x 1500 showing minute radial ridges outside the curvatural crassitude, Ger92/12 (371) 3, 100 1131; 4, FM 1507, LV92/17 (D3) 3, co-ord. 207 0908, E.E.V21/3. Figs 5-8 Scylaspora vetusta (Rodriguez) comb. nov. showing variations. 5, FM 1508, oblique compression showing +/— radial muri around the curvatural crassitude, LV92/8 (D3) 2, 106 0991; 6, FM 1509, proximal view, thin-walled specimen showing equatorial radial thickenings becoming indistinct towards the proximal pole, LP92/8 (D2) 1, 025 1014: 7a—e, FM 1510, proximal view showing irregular and anastomosing proximal muri; 7e, x 1500; LV92/8 (D3) 2, 058 1127; 8, FM 1511, specimen showing paired lips, Arg92/5a (473) 3, 029 0910. Fig.9 Scylaspora cf. scripta Burgess & Richardson 1995. FM 1512, x 1500, LV97/8a (D16) 2, 096 1123. Fig. 10 Scylaspora ct. kozlica (Dufka) comb. nov. FM 1513, LV92/8 (D3) 1, 163 0983; 10a, proximal polar compression, showing proximal microrugulae and grana; 10b, x 1500, showing detail of proximal sculpture. All figs x 1000, unless stated otherwise. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN ie 144 becoming sinuous, and less distinct, poleward. Trilete sutures +/— equal the spore radius or slightly less, accompanied by raised, often darkened areas, tapering towards the amb. A wider thinner proximal triangular area reaches c. 4/Sths of the spore radius and inter-radially often reaches the papillae; papillae distinct, rounded to ovoid in plan view, with a slightly raised profile, maximum diameter 4—9 um. Trilete-folds, or smooth lips, occur in some specimens and are narrow over the apical area and expand towards the equator. DIMENSIONS. 27-58 um (59 specimens measured). COMPARISONS. Scylaspora elegans sp. nov. differs from proxi- mally tripapillate Ambitisporites species by its proximal microrugulae and apical darkened area (lips) bordering the trilete mark. The photograph of the holotype of Ambitisporites ? eslae Cramer & Diez (1975: pl. 1, fig. 11) lacks this feature and has a wide crassitude. Retusotriletes maculatus has proximal curvaturae invaginated at their radial apices. REMARKS. The current investigation revealed that Retusotriletes eslae Cramer & Diez includes two distinct forms. The original holotype has a broad equatorial crassitude, trilete folds and appears to lack the apical, gradually tapering and darkened areas that flank the trilete mark. Specimens with a broad crassitude are rare in the samples studied (eg. Pl. 5, fig. 1) and it is not known whether such forms have a proximal microverrucate sculpture. It has not been possible to study the holotype for this purpose and, although Cramer & Diez described microrugulae, their diagnosis states that the proximal surface is ‘smooth to microrugulate’. Pending re-examination of the type material the forms with a broad crassitude are retained but referred to as Ambitisporites ? eslae (Cramer & Diez). The more commonly occur- ring spores, withanarrow, more or less equatorial, curvatural crassitude, distinct trilete area and proximal microrugulae, are transferred to the new species Scylaspora elegans. The microrugulae can be seen under the light microscope in some specimens but under the SEM all spores of this species show microrugulae that decrease in size towards the proximal pole. Both species occur in all sections but data from the present study shows that S. elegans sp. nov. occurs at the base of the EC Biozone whereas A? eslae (Cramer & Diez) comb. nov. occurs first in the Aneurospora Subzone at the top of the EC Biozone. In the Cantabrian Mountains Scylaspora elegans sp. nov. occurs in strata equivalent to the Upper/uppermost Pridoli and extends into the Lower Lochkovian (Devonian) and may, therefore, help in establishing the Silurian/Devonian boundary. PLATE 6 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND OCCURRENCE. Upper San Pedro Formation, Argovejo, Geras, La Peral and La Vid sections, EC Biozone to Lower MN Sub-Biozone, Upper Pridoli to Lower Lochkovian. Scylaspora cf. scripta Burgess & Richardson 1995 Il, Dp WS) DIMENSIONS. — 23—29um (3 specimens measured). OCCURRENCE. La Vid, samples 6, 7B, 8A. Scylaspora vetusta (Rodriguez) comb. nov. Pl. 3, fig. 5; Pl. 5, figs 5—8; Pl. 6, fig. 1 BASIONYM. Archaeozonotriletes vetustus Rodriguez, 1978b: 219, pl. 1, fig. 8. 1973. Emphanisporites? sp. D, Richardson & Ioannides (pars): 276, pl. 3, fig. 9 only. 1978b Archaeozonotriletes vetustus Rodriguez: 219, pl. 1, fig. 8. 1978c Archaeozonotriletes chulus Rodriguez: pl. 1, fig. 14. 1978c Archaeozonotriletes vetustus Rodriguez: 414, pl. 2, figs 7, 8. 1983 Archaeozonotriletes vetustus Rodriguez: 32, pl. 1, figs 2, 18, 20. 1995 Rugosisporites cf. chartulatus (McGregor); Dufka (pars): 71, pl. 2, figs 9-14. HOLOTYPE ANDTYPELOCALITY. Rodriguez, R.M. 1978b: pl. 1, fig. 8; Corniero village, Province of Len, Cantabrian Mountains, north- west Spain, lower San Pedro Formation, ?Ludfordian. EMENDED DIAGNOSIS. A Scylaspora with closely spaced, more or less radial, anastomosing, irregularly sinuous, muri/rugulae on the proximal surface; distally laevigate apart from a narrow subequato- rial area with +/— radial muri. DIMENSIONS. 28-72 tum (60 specimens measured). REMARKS. Spores of this species are highly variable. In some cases there are irregular muri/rugulae that reach the pole, in others cuneiform elements are prominent over the equatorial crassitude but sculpture is faint to indistinguishable over the rest of the proximal surface. Intermediates occur where distinct equatorial sculpture becomes increasingly faint leaving a laevigate apical zone. The equatorial crassitude is variable in width. Fig. 1 Scylaspora vetusta (Rodriguez) comb. nov. la, BM 140019, proximal view, LP92/10/DP; 1b, BM 134179, proximal view showing subequatorial radial muri and scabrate apical area, sample Geras 92/7; 1c, BM 134195, proximal view, showing irregular proximal sculpture, Arg92/4A/1. Fig. 2 Scylaspora sp. BM 135151, strong broad equatorial crassitude and coarse sinuous proximal muri, Ger2B/2. Fig. 3 9 Coronaspora primordiale (Rodriguez) Rodriguez 1983. BM 137485, proximal view showing inter-radial thickenings, grooved crassitude and lips, Ger2B/2. Fig.4 Iberoespora cantabrica Cramer & Diez 1975. 4a, BM 116051, proximal view showing chevron-shaped elements forming the labra, Arg92/7; 4b, BM 137480, proximal view showing inter-radial thickenings, Arg13/Dii. Fig.5 Iberoespora glabella? Cramer & Diez 1975. BM 129724, distal view showing concentric furrow inside equatorial crassitude and foveolate- murornate sculpture over central area, Arg92/11B. Fig. 6 Coronaspora cromatica (Rodriguez) Jansonius & Hills 1979. 6a, BM 134510, proximal view showing variably developed verrucate lips, kyrtome and inter-radial muri, Ger92/2B; 6b, BM 134199, proximal view, showing kyrtome, inter-radial thickenings and verrucate labra, Arg5A/1; 6c, BM 137269, distal view showing annulus and irregular muri in polar area, Ger92/2B/2; 6d, BM 135144, oblique compression showing equatorial crassitude and distal annulus, Ger92/2B/1; 6e, BM 134634, proximal view showing well-developed verrucate labra, Ger92/2B; 6f, BM 134634, detail of proximal apical area in Fig. 6e, x 2000. Fig.7 Coronaspora reticulata sp. nov. 7a, BM 138958, oblique proximal view showing kyrtome, LV92/7b/DD1; 7b, BM 138395, proximal view showing kyrtome and thin apical area. Fig. 8 Scylaspora elegans sp. nov. 8a, BM 2548, proximal view, Arg 92/11B/2; 8b, BM2551, detail of proximal microrugulae, tilt 45°, x 4000. All figs x 1000, unless stated otherwise. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN “yet 146 Pl. 5, fig. 10 Cf. 1995 Rugosisporites kozlicus Dufka (pars): 72, pl. 2, figs 15—17. Scylaspora cf. kozlica (Dutka) comb. nov. DESCRIPTION. Amb circular to subcircular. Equatorial crassitude 1—2 um wide. Trilete mark labrate, c. 4/5 spore radius, lips less than lum wide, merge with curvaturae perfectae which form a narrow equatorial crassitude with a thicker inner portion and a narrow (<0.5 um), apparently membranous, extension. Proximal surface covered with narrow irregular rugulae c. 0.5 um wide and possible micrograna. Sculpture less distinct near the proximal pole. Distal surface laevigate except for sub-equatorial rugulae with scattered micrograna. DIMENSIONS. 31—44 um (30 specimens measured). COMPARISONS. Similar to S. scripta in having radially aligned muri near the equator and distal sculpture, but has more radial geniculate and anastomosing robust muri. S. kozlicus (Dufka) is similar but is distally laevigate and lacks the paired lines of thicken- ing on either side of the Y-mark. S. sp. A (this study) has geniculate muri on the proximal and distal surfaces and is rare. REMARKS. Scylaspora cf. scripta is similar to forms described from the Homerian and may be reworked. The spores have a mem- branous extension, seen at the equator on some specimens, and may be double-layered with the thin outer part of the exoexine forming the rugulae over the proximal surface. S. vetusta has coarser rugulae but is otherwise closely similar. OCCURRENCE. La Vid section, upper RS and lower H Biozones (Upper Ludfordian — Pridolt). Genus CONCENTRICOSISPORITES Rodriguez 1983 TYPE SPECIES. REMARKS. Jansonius & Hills (1990: card 4630) considered that the central dark area in the type species may represent an inner body. Concentricosisporites sagittarius (Rodriguez) 1983. Concentricosisporites agradabilis (Rodriguez) Rodriguez 1983 AL, We, 2 1978c Stenozonotriletes agradabilis Rodriguez: 421, pl. 3, figs 7, 5 Hil. 1983 Concentricosisporites agradabilis Rodriguez: 35, pl. 4, figs Wy Wil, 113). HOLOTYPE AND TYPE LOCALITY. Rodriguez, 1978b: pl. 3, fig. 8; Torrestio, Province of Leon, Cantabrian Mountains, northwest Spain, lower San Pedro Formation. DIAGNOSIS. See Rodriguez, 1978c: 421. PLATE7 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND DIMENSIONS. 26-39 ym (18 specimens measured); Rodriguez, 1978b, 24-35 um. COMPARISONS. This species shows some resemblance to spores of the genus Emphanisporites but the proximal radial ribs are not always distinct and the equatorial crassitude is relatively wide (3-6 uum) in relation to the spore radius. OCCURRENCE. Lower San Pedro Formation; upper RS and lower H Biozones; sporadically occurs in EC and lower MN Biozones where they may represent reworking. In the higher specimens the distal annulus is lobed and the radial ‘muri’ more distinct. Upper Ludfordian and Lower Pridoli. Concentricosisporites sagittarius (Rodriguez) Rodriguez, 1983 Pie 7.-tig. 1 1978b Stenozonotriletes sagittarius Rodriguez: 219, pl. 1, fig. 7. 1978c Stenozonotriletes sagittarius Rodriguez: 421, pl. 2, figs 13, 14. 1983 Concentricosisporites sagittarius (Rodriguez); Rodriguez: 36, pl. 3, fig. 21. 1995 Concentricosisporites sagittarius Rodriguez; Burgess & Richardson: 17, pl. 6, figs 14, 15. HOLOTYPE. Rodriguez, 1978b: pl. 1, fig. 7. DIAGNOSIS. See Rodriguez, 1978b: 219. DIMENSIONS. 29-44 um (10 specimens measured); 25-40 pm (Rodriguez, 1978). OCCURRENCE. San Pedro Formation, Upper RS & H Biozones La Vid, upper RS & lower H Biozones La Peral, lower MN Biozone Geras (? reworked). Upper Ludfordian and Lower Pridoli. Genus IBEROESPORA Cramer & Diez 1975 TYPE SPECIES. Iberoespora cantabrica Cramer & Diez 1975. Iberoespora cantabrica Cramer & Diez 1975 Pl. 6, fig. 4; Pl. 7, fig. 4 HOLOTYPE. Cramer & Diez 1975: pl. 2, fig. 24. DIAGNOsIS. See Cramer & Diez 1975: 339, pl. 2, figs, 24, 26-28, 310), Sil DIMENSIONS. 30 to 45 um (Cramer & Diez, 1975); 27 to 42 um (40 specimens measured). Fig. 1 Concentricosisporites sagittarius (Rodriguez) Rodriguez 1983. FM 1514, Ger92/15a (475) 1, 044 1032. Fig. 2 Concentricosisporites agradabilis (Rodriguez) Rodriguez 1983. FM 1515, LV92/13 (495) 2, 098 0982; 2a, proximal focus; 2b, distal focus, showing annulus. Fig.3 /beroespora glabella? Cramer & Diez 1975. FM 1516, Argovejo 92/12 (510) 3, 139 0874. Fig.4 /beroespora cantabrica Cramer & Diez 1975. FM 1517, Arg92/12 (510) 3, 212 0962; 4a, 4b, showing verrucate lips, distal geniculate muri, and inter-radial thickenings; 4a, x 1500; 4c, median focus showing margin of the central body invaginated. Fig.5 Leonispora argovejae Cramer & Diez 1975. FM 1519, LV92/13 (D3) 1, 051 0913. Fig.6 Iberoespora mariae Rodriguez 1983. FM 1518, LP92/20 (D8) 2, Geras 92/11 (D8) 1, 030 0969 (see Fig.2, sample GER 11). Fig.7 Aneurospora sp. Tripapillate form. FM 1520, Ger92/14 (468) 7, 132 0975; 7a, proximal focus; 7b, proximal focus, showing curvaturae perfectae. Fig. 8 Streelispora newportensis (Chaloner & Streel) Richardson & Lister 1969. FM 1521, LV13 (D3) 1, 082 0986; 8a, 8b, x 1500. Fig.9 Aneurospora richardsonii (Rodriguez) comb. noy., FM 1522, Gerl4 92/14 (468) 7, 135 0838; 9b, shows sculpture in detail, x 1500. Fig. 10 Chelinospora sanpetrensis (Rodriguez) comb. nov. FM 1523, Arg92/5a (473) 3, 123 1092; 10a, distal focus showing foveolate exine; 10b, with narrow arcuate fold around proximal hilum, x 1500. All figs x 1000, unless stated otherwise. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN 147 148 J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN COMPARISON. Distally foveolate specimens with a laevigate crassitude are referred to as Iberoespora glabella? (P1. 6, fig. 5; Pl. 7, fig. 3). The original illustrations (Cramer & Diez, 1975: pl. 2, figs 22, 29) do not show the distal surface clearly. REMARKS. Cramer & Diez do not explain how they differentiate /. guzmani from I. cantabrica. Both species have the same structure, radial muri on the crassitude, crenulate lips, a distal sculpture of reticulate to convolute muri, and thickenings in the inter-radial areas. However, the size range given for J. guzmani is only 10-20 um whereas that for /. cantabrica is 30-45 um. The specimens in this study are all larger than 26 um. OCCURRENCE. Upper San Pedro Formation in Argovejo, Geras and La Vid sections in the Province of Leon and La Peral section in the Province of Asturias, upper EC — lower MN Biozones. Genus LEONISPORA Cramer & Diez 1975 TYPE SPECIES. Leonispora argovejae Cramer & Diez 1975. Leonispora argovejae Cramer & Diez 1975 Cramer & Diez 1975, pl. 1, fig. 3. DESCRIPTION. Cramer & Diez (1975: 342). DIMENSIONS. 25-35 um (ibid. 1975); 30-38 um (14 specimens measured, present study). PI. 7, fig. 5 HOLOTYPE. COMPARISONS. The genus is closely similar in structure to Streelispora but is laevigate. OCCURRENCE. Upper San Pedro Formation, Geras, La Vid sec- tions; MN Miospore Biozone, NA Subzone. Genus STREELISPORA (Richardson & Lister) Richardson, Streel, Hassan & Steemans 1982 TYPE SPECIES. Streelispora newportensis (Chaloner & Streel); Richardson & Lister 1969. Streelispora newportensis (Chaloner & Streel); Richardson & Lister 1969 Pl. 7, fig. 8 1968 Granulatisporites newportensis Chaloner & Streel: 92, pl. 19, figs 7, 8, 11. 1969 — Streelispora newportensis (Chaloner & Streel); Richardson & Lister: 230-31, pl. 41, figs 3-6. HOLOTYPE. Chaloner & Streel 1968: 92, pl. 19, figs 7, 8, 11. 149 DIMENSIONS. 28-39 um (13 specimens measured), 17-48 um (80 specimens measured) Richardson & Lister 1969. OCCURRENCE. Upper San Pedro Formation, Argovejo, Geras, La Vid, MN Biozone, NA Sub-Biozone. Genus ANEUROSPORA Stree! 1964 TYPE SPECIES. Aneurospora goensis Streel 1964 Aneurospora richardsonii (Rodriguez) comb. nov. VA Ah, wis, 2 TAL, If, taney, © BASIONYM. Streelispora richardsonii Rodriguez, 1983: 50, pl. 8, figs 6, 7, pl. 9, fig. 3. 1978c Geminospora spinosa Allen; Rodriguez: 416, pl. 5, figs 7, 8. 1983 Streelispora richardsonii Rodriguez: 50, pl. 8, figs 6, 7, pl. 9, fig. 3. HOLOTYPE AND LOCUS TYPICUS. See Rodriguez, 1983: 50, pl. 8, fig. 3. EMENDED DIAGNOSIS. An Aneurospora with no proximal papillae, distal sculpture consists of spaced slender spines (fimbriae), ele- ments dominantly parallel-sided with blunt, rounded apices. COMPARISON. The slender fimbriae distinguish this species. Anapiculatisporites raistrickiaeformis Schultz (1968) has coarser, more irregular elements (see also Steemans 1989). DIMENSIONS. 38 to 50 um (Rodriguez, 1983); 32 to 47 um (28 specimens measured). OCCURRENCE. Lower part of lower MN Biozone, NA Subzone, Argovejo, Geras and La Vid sections. REMARKS. The species Aneurospora richardsonii lacks the proxi- mal papillae and associated tangential folds and is herein transferred to the genus Aneurospora. Tangential folds around the proximal papillae distinguish Streelispora and Leonispora from tripapillate species of Aneurospora (eg. Pl. 7, fig.7). Genus CORONASPORA Rodriguez 1979 emend. 1978b Coronaspora mariae Rodriguez: 218, pl. 1, figs 1-3. 1979 Coronaspora mariae Rodriguez: 232. TYPE SPECIES. Coronaspora mariae Rodriguez, 1978. HOLOTYPE. 9 2. Coronaspora mariae Rodriguez, 1978b: pl. 1, figs 1, PLATE 8 Figs 1-4 Coronaspora cromatica (Rodriguez) emend. 1, FM 1524; 1a, showing proximal kyrtome and broad verrucate lips, 1b, intermediate focus, Ic, distal view showing annulus and sculpture of sinuous muri and rounded verrucae; Ger92/5 (542) 2, 098 1006; 2, FM 1525; 2a—c, proximal to distal levels of focus; 2a with narrow lips and distinct kyrtome; 2b relatively narrow crassitude; 2c, distal sculpture of small interconnected narrow verrucae Arg92/Sa, (473) 3, 023 0983; 3, FM 1526, proximal view, Arg92/Sa (473) 3, 139 1133; 4, FM 1527, proximal view, specimen with inter-radial thickenings on the kyrtome, Arg92/Sa (473) 3, 099 0938. Figs 5,6 Coronaspora reticulata sp. nov. 5, FM 1528, proximal focus showing kyrtome with wide inter-radial thickenings, LV92/7b (D3) 3, 156 0917; 6a, b, FM 1491 (holotype), showing distal reticulum (the radial extremities are visible in Fig. 6a), LV92/7b (D3) 3, 043 0965. Fig. 7 Coronaspora subornata (Cramer & Diez) comb. nov. FM 1529, specimen showing narrow Y-folds, equatorial crassitude, distal annulus and irregular distal thickening; Ger92/9 (456) 1, 035 0472. Fig.8 Chelinospora canistrata sp. nov. FM 1492 (holotype), Ger92/5 (542) 2, 083 1024. Fig.9 Coronaspora infraornata (Rodriguez) comb. nov. FM 1530, showing curvatural crassitude, distal annulus and distal polar thickening at two focal levels, Ger92/9 (456) 1, 066 0942. All figs x 1000. 150 EMENDED DIAGNOSIS. Equatorial crassitate miospores with rounded or convexly triangular amb, a proximal kyrtome of thick- ened arcuate ridges extending to the radial apices where it is flat-topped and joins with the trilete lips; distal surface with, or without, a distal annulus; distal exine laevigate, verrucate, irregu- larly murornate, or reticulate. COMPARISON. The rounded amb and distinct kyrtome distinguish this genus. There is no kyrtome on the type species of Amicosporites (A. splendidus) but two other species, A. infraornatus Rodriguez 1978b and A. subornator Cramer and Diez 1975 have this structure and are herein transferred to Coronaspora. The kyrtome in Ahrensisporites Potonié & Kremp, 1954 and Concavisporites Pflug in Thomson and Pflug, 1953 are distinct because the radial parts are prominent and inter-radially the amb is straight to concave. The kyrtome tapers evenly towards the equator in Kyrtomisporis Madler 1964 (see also Achilles, 1981) whereas in Coronaspora the kyrtome is a radially broad and flattened (truncated) thickening where the inter-radial branches of the kyrtome meet, and may invaginate to become confluent with the lips. In cross-section the Silurian kyrtomes have a semicircular-shape. REMARKS. Rodriguez validated the genus in 1979 when she desig- nated Coronaspora mariae as type species. Coronaspora cromatica (Rodriguez) emend Pl. 6, fig. 6; Pl. 8, figs 14 1978b Coronaspora cromatica Rodriguez: 218, pl. 1, fig. 4. 1979 | Coronaspora cromatica (Rodriguez) Jansonius & Hills: card number 3543. 1983 Coronaspora cromatica Rodriguez: pl. 3, fig. 4. HOLOTYPE. Rodriguez, 1978b: 218, pl. 1, fig. 4. EMENDED DIAGNOSIS. A Coronaspora with distinct labra, a proxi- mal kyrtome with inter-radial thickenings, distal annulus, and variable muri and verrucae in the distal polar region. DESCRIPTION. Amb subcircular to subtriangular with convex sides. Spores preserved in polar compression suggesting an original of more or less lenticular shape. Exine proximally with a distinct complete kyrtome with inter-radial thickenings, radial ‘bars’, con- necting these to a narrow (c. 3 ym), +/— smooth to irregularly thickened, equatorial crassitude. Kyrtome distinct, formed by raised ridges (+/— semi-circular in profile) paralleling the lips in the inter- radial areas, flattened at radial extremities where it may show distinct invaginations; lips variably developed, may be broad and verrucate often taper sharply to the apices, or may be laevigate and sinuous; in forms with verrucate lips sutures distinct. Distally an annulus has smooth to irregular margins c. 9 um wide, outer border J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND of annulus reaches to 2/3 of the spore radius. Distally polar area sculptured by broad muri 3—7um wide, 9-18 um long, muri highly variable in shape and orientation, forming linear or sinuous bars which may be smooth-sided, or show a series of constrictions giving a beaded appearance, due to the mixture of verrucae and short muri, or coalescent verrucae. DIMENSIONS. 28-47 uum (58 specimens measured); Rodriguez, 1978b, 20-30 um; width of the cingulum c. 2 um. COMPARISONS. Coronaspora mariae also has a distal annulus and is closely similar but has large rounded verrucae over the distal pole. REMARKS. Rare specimens with sinuous laevigate lips, like the type material (Rodriguez, 1978b), occur in the Upper part of the San Pedro Formation. So far the forms with smooth lips have only been found in the upper H to EC Zones. Specimens with verrucate lips occur in the upper RS Zone and may prove to be a useful marker for the lower boundary of the Pridoli. OCCURRENCE. Forms with verrucate lips (PI. 6, fig. 6, Pl. 8, figs 1— 4) occur in the Lower San Pedro Formation; lower upper RS Biozone Geras, Argovejo, lower H La Vid and La Peral sections; spores with laevigate lips, C. cf. cromatica, Upper San Pedro Formation, upper H & EC Biozone La Peral, EC & lower MN Biozones La Vid. Coronaspora reticulata sp. nov.P1. 6, fig. 7; Pl. 8, figs 5, 6 1975 Knoxisporites? riondae Cramer & Diez (pars): 341, pl. 1, fig. 17 only. Knoxisporites ? riondae Cramer & Diez; Rodriguez : 42, 43, pl. 2, figs 18, 19. 1983 Knoxisporites? riondae Cramer & Diez; Rodriguez (pars): 42, 43, pl. 4, figs 2, 3. 1978c HOLOTYPE. FM 1491 (PI. 8, fig. 6), diameter 36 um; sample La Vid 7B, slide D3/3, co-ord. 043 0965; E.F. no. E27/1; lower San Pedro Formation, La Vid section, Province of Leon, Cantabrian Mountains, northwest Spain. DIAGNOSIS. A Coronaspora with broad distal muri forming a reticulum; kyrtome thickened especially in the inter-radial areas; kyrtome surrounds a clover-leaf shaped area of thin exine; laesurae labrate, labra narrow, smooth and sinuous. DESCRIPTION. Amb circular to subcircular. Spores usually pre- served in polar compression, suggesting an original, more-or-less lenticular cross-section. Equatorial crassitude smooth +/— 2 um. Kyrtome smooth, invaginated and increasing in width towards the spore apex in the inter-radial areas, 24 pm, joined at the radial apices where they may be slightly invaginated at the junction with PLATE 9 Fig. 1 Chelinospora cantabrica sp. nov. FM 1493, (holotype); 1a, distal focus showing a reticulum of broad muri, several with characteristic narrow restrictions; 1b, shows equatorial margin and muri in plan, Arg92/5a (473) 3, 144 1082. Fig. 2 Chelinospora lavidensis sp. nov. FM 1494 (holotype), showing narrow muri forming an irregular reticulum with subequatorial muri normal to the equator, Arg92/12 (510) 3, 093 1019. Figs 3,5 Chelinospora hemiesferica (Cramer & Diez) comb. noy. 3, FM 1531; 3a, proximal view showing thick crassitude and narrow concentric fold at the equatorial edge of the hilum; 3b, distal surface showing close-packed narrow geniculate muri, Ger92/9 (450) 1, 147 1050; 5, FM 1532 tetrad, showing parallel subequatorial muri normal to the equator and geniculate muri over the distal patina, Arg92/14 (468) 11, 080 1069. Fig.4 Chelinospora cf. hemiesferica (Cramer & Diez) comb. noy. 4, FM 1533; 4a, distal view, muri geniculate, but broader and more spaced than in the species; 4b, proximal focus showing margins of the hilum, Arg92/11a (441) 4, 074 0915. Fig.6 Chelinospora media sp. nov. FM 1495, holotype, Arg92/12 (510) 3, 093 1019. Fig.7 Cymbosporites cf. dittonensis Richardson & Lister 1969. FM 1534, Arg92/13 (455) 1,049 1014. All figs x 1000. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN 151 Nn i) J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN the laesurae, lips smooth, narrow, decreasing in width over the thinner polar area, <1 tm wide. Distal surface with a reticulum of broad muri c.1—4 ym wide and c. 1 wm high, surrounding +/— polygonal lumine 3—6 um maximum width. DIMENSIONS. 26-44 tum (52 specimens measured). COMPARISONS. The presence of an equatorial crassitude, proximal kyrtome and distal reticulum, distinguish this species from Knoxisporites ? riondae Cramer & Diez 1975, the latter has a ‘marked concentrical ring’ around the distal pole and distal warts (see Cramer & Diez 1975: 341, reconstruction fig. 2.3, and distal focus of holotype pl. 1, fig. 16). REMARKS. The proximal surface is seen best under the SEM (PI. 6, fig. 7b) but the kyrtomal thickenings and thin trilobed proximal apical area are also seen under the light microscope. OCCURRENCE. La Peral RS and lowermost H Biozone, La Vid RS Biozone, and doubtful, poorly preserved specimens in upper EC and lower MN Biozones; Upper Ludfordian and Pridolf; possibly re- worked in Upper Pridoli and Lower Lochkovian, Argovejo. Coronaspora subornata (Cramer & Diez) comb. nov. Pe Salton BASIONYM. Amicosporites subornator Cramer & Diez 1975: 338, pl. 1, fig. 7. 1973 Spore type D; Richardson & Ioannides: 281, pl. 9, fig. 2. 1975 Amicosporites subornator Cramer & Diez: 338, pl. 1, fig. 7. 1978c Amicosporites subornator Cramer & Diez; Rodriguez: 412— 13, pl. 2, fig. 2. 1983 Amicosporites subornator Crame & Diez; Rodriguez: 30, pl. 3, fig. 13. HOLOTYPE. Cramer & Diez 1975; pl. 1, fig. 7; San Pedro Forma- tion, Argovejo section, Province of Leén, Cantabrian Mountains, northwest Spain. DIAGNOsISs. A Coronaspora with a distal annulus and smooth exine. Description. Amb subcircular to subtriangular with convex sides. Spores preserved in polar compression, suggesting an original of more or less lenticular shape. Exine with a distinct proximal kyrtome, equatorial crassitude +/— smooth to irregularly thickened, and a narrow distal annulus (c. 3 um wide); annulus smooth, and reaches to c. 4/5 of the spore radius. Distal? pole surrounded by a dark ring (?annulus). Trilete sutures reach the crassitude, which is invaginated at the radial apices; lips smooth, distinct, and parallel-sided. PLATE 10 15s DIMENSIONS. 30-40 pm (Cramer & Diez, 1975); 31-44 um (14 specimens measured, present study). COMPARISONS. Differs from other members of the genus, with the exception of C. infraornata, in having a laevigate exine, C. infraornata has a dark distal polar area. REMARKS. Cramer & Diez (1975: 338) refer to the kyrtome as ‘three segments of a circular inspissation’ and a dark polar ring as a proximal inspissation. The spores differ from typical members of the genus Amicosporites because they have a distinct proximal kyrtome (Cramer & Diez, 1975; fig. 2, 2) similar, if not identical, to those of C. mariae and C. cromatica. We therefore transfer this species to Coronaspora. OCCURRENCE. La Peral EC Biozone, La Vid, uppermost H, or lowermost EC Biozone, Argovejo, H Biozone; Middle and Upper Pridoli. Coronaspora infraornata (Rodriguez) comb. nov. Pl. 8, fig. 9 BASIONYM. Amocosporites infraornatus Rodriguez 1978b: 217, pl. 1, figs 5, 6. 1978b Amocosporites infraornatus Rodriguez: 217, pl. 1, figs 5, 6. 1978c Amocosporites infraornatus Rodriguez: 412, pl. 2, fig. 3. 1983. Amocosporites infraornatus Rodriguez: 29, pl. 2, figs 14, 16. HOLOTYPE. Rodriguez, 1978b: 217-18, pl. 1, figs 5, 6; San Pedro Formation, Corniero section, Province of Le6n, Cantabrian Moun- tains, northwest Spain. EMENDED DIAGNOSIS. A Coronaspora witha distal annulus, smooth exine, and a dark circular area over the distal pole. DESCRIPTION. Amb subcircular to subtriangular with convex sides. Spores preserved in polar compression, suggesting an original of more or less lenticular shape. Exine with a distinct proximal kyrtome, equatorial crassitude +/— smooth to irregularly thickened, and a narrow distal annulus; annulus smooth and 2—3 um wide, reaches to c. 4/5 of the spore radius. Distal pole surrounded by a circular dark area. Trilete sutures reach the crassitude, which is invaginated at the radial apices; lips smooth, distinct, and parallel-sided. DIMENSIONS. 15-35 um Rodriguez (1978b); 25-47 um (25 speci- mens measured, present study). COMPARISONS. Differs from Coronaspora subornata in having the circular dark area at the distal pole. In the present study, forms with a dark ring can be distinguished from those with a circular dark area Fig. 1 Chelinospora hemiesferica? (Cramer & Diez) comb. nov. BM 137697, proximal view, x 2000, Ger92/9/D1. Fig.2 ?Cymbosporites sp. A. BM 134646, proximal view, Ger92/2A. Fig.3. Cymbosporites sp. B. Arg92/10; 3a, BM 130309, proximal/oblique view, x 2000; 3b, BM 130310, detail of sculpture showing some grana fused into groups, x 2500. Figs 4,5 Cymbosporites sp. 4a, BM 133533, Ger92/14, equatorial view; 4b, same specimen showing detail of multi-layered wall structure and acinoform sculpture of muri and microconi, x 2000; 5, sample Geras 92/8; 5a, BM 130610, equatorial view; 5b, BM 130611, detail of irregular distal coni, x 2000. Fig.6 Chelinospora (Lophozonotriletes?) poecilomorpha (Richardson & Ioannides) comb. noy. 6a, BM 135046, oblique compression showing hilum and large distal verrucae, Ger92/2A; 6b, BM 141007, proximal view, LP10/DD2:; 6c, BM 132242, distal view, Arg92/13, BM 116097; 6d, distal view showing discrete, rounded to flat-topped verrucae variable in size, LP92/16. Fig.7 Chelinospora (Lophozonotriletes?) poecilomorpha (Richardson & Ioannides) comb. nov. BM 141007, proximal view, LP92/10/DD2. Fig.8 Archaicusporites torrestionensis? Rodriguez 1983. BM 130522, proximal/oblique view, Ger92/9. Fig.9 Cf. Pachytetras sp. Possible permanent cryptospore tetrad, Ger92/2B/2; 9a, BM 137274, tetrad showing rugulate distal sculpture; 9b, BM137275, triple contact. All figs x 1000, unless stated otherwise. J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN but in the original description Rodriguez separated Amicosporites infraornator from A. subornata on the distal position of the dark area. Cramer & Diez (1975) on the other hand thought that the dark ring in A. subornator was proximal. If further specimens show that in A. subornator the dark ring is distal then the two species may have to be combined. REMARKS. The spores differ from typical members of the genus Amicosporites in having a proximal kyrtome (see under C. cromatica). Rodriguez (1978b) included in her species both spores with a distal annulus, not well defined at the inner margin, and others where the annulus was dissected into a series of verrucate-like structures (Rodriguez, 1978b: pl. 1, fig. 12). OCCURRENCE. Geras, upper RS, H and lower EC Biozones, La Vid, upper RS to top H Biozones, Argovejo, RS, H, EC and MN Biozones, La Peral upper H and lower EC Biozones; Piidolf and lowermost Devonian. Infraturma PATINATI Butterworth & Williams 1958 emend. Smith & Butterworth 1967 Genus CHELINOSPORA Allen 1965 TYPE SPECIES. Chelinospora concinna Allen 1965. Chelinospora canistrata sp. nov. Pl. 8, fig. 8. DERIVATION OFNAME. Latin canistrumn. wicker basket, canistrata resembling a basket. HOLOTYPE. FM 1492, sample Geras 92/5, slide 2 (542), co-ord. 0831024, E.F. no. J33/1; Lower San Pedro Formation, Geras section, Province of Leon, Cantabrian Mountains, northwest Spain. DIAGNOsIs. A Chelinospora with low large verrucae-muri and closely packed convolute microrugulae. DESCRIPTION. Proximally hilate, distally patinate miospores; hilum thin, sometimes diaphanous, frequently torn, or may be collapsed or lost, with a narrow concentric fold just inside the crassitude; narrow equatorial curvatural crassitude with short proximal radial muri extending to the hilum margin seen on a few specimens; distal sculpture of low verrucae and muri, often indistinct, and closely packed sinuous microrugulae < 1 um wide. Y-mark with narrow ‘folds’ extend c. 2/3 spore radius and merge into crassitude at the hilum margin. DIMENSIONS. 25-43 um (based on 18 specimens). COMPARISONS. The sculpture of low verrucae-muri and microrugulae distinguish Chelinospora canistrata sp. noy. from PLATE 11 155 other Chelinospora species. OCCURRENCE. Found in all sections apart from La Vid, uppermost RS, H, and lower EC Biozones. Chelinospora cantabrica sp. nov. PI. 4, fig. 7; Pl. 9, fig. | DERIVATION OF NAME. Spain. Named after the Cantabrian region, NW HOLOTYPE. FM 1493 (Pl. 9, fig. 1), diameter 39 um, sample Argovejo 92/5a, slide 3 (473), co-ord. 1441082, E.F. no. P39/1; Lower San Pedro Formation, Argovejo section, Province of Leén, Cantabrian Mountains, northwest Spain. DIAGNOsIs. A Chelinospora with a reticulum composed of broad muri with occasional constrictions and narrow sutures, lumina wide, polygonal to irregular. DESCRIPTION. Proximally hilate miospores, equatorially 2-4 um thick, distal exine thick (not measured); amb subcircular to subtriangular. Proximal exine thin, frequently broken, or absent, trilete mark not seen. Equatorially and distally sculptured with a broad reticulum, muri 1-10 um wide, 1-3 um high, lumina wide and polygonal 2.5-10 um wide; muri show constrictions broken by narrow sutures at the junction with adjacent polygonal fields. DIMENSIONS. 5-51 um (based on 20 specimens). COMPARISONS. The broad muri with distinct constrictions and sutures, and wide +/— polygonal lumina, distinguish this species from other species of Chelinospora. Coronaspora reticulata sp. nov. has similar reticulate sculpture but has an equatorial crassitude and a proximal kyrtome. OCCURRENCE. Argovejo, Geras, and La Vid sections, Lower San Pedro Formation, RS and lowermost H Biozones, occasional finds in higher zones in the La Vid section. Chelinospora cassicula Richardson & Lister 1969 1969 Chelinospora cassicula Richardson & Lister: 242-43, pl . 42, figs 10-12. 1978a Chelinospora mariae Rodriguez (pars); Rodriguez: 10, pl. 1, fig. 3 only. REMARKS. Rodriguez, 1978a (pl. 1, figs 3-5) placed several spore morphologies in this taxon but from the illustration the holotype belongs in C. cassicula. In the Spanish material examined for this paper several Chelinospora species with an equatorial band of radial muri form an evolving ‘lineage’ (morphon) beginning with C. hemiesferica, Fig. 1 Conochitina pachycephala Eisenack 1964. FC 901, lateral view, x 170, LP3/SB4/45. Fig. 2 Conochitina alargada Cramer 1967. FC 91, lateral view, x 220, Arg4/SB7/3. Figs 3,5,6 Cyathochitina elenitae Cramer 1964. 3, FC 92, lateral view, x 200, LP3/SB4/48; 5, FC 93, lateral view x 200, Arg4/SB7/3; 6, FC 94, Arg3/ SB7/1; 6a, lateral view x 150; 6b, detail of carina, x 750. Fig. 4 Spinachitina sp. FC 95, LP2/SB4/43; 4a, lateral view, x 300; 4b, basal margin detail, x 1750. Fig. 7 Calpichitina velata (Wrona 1980). FC 96, Ger15b/SBGeras-1; 7a, apertural view, x 500; 7b, detail of operculum x 1000. Fig. 8 Vinnalochitina horrentis (Jaglin 1986). FC 97, Ger13/SBGeras-2; 8a, apertural view, x 500; 8b, detail of operculum x 1000. Fig. 9 Cingulochitina wronai Paris & Kriz 1984. FC 98, lateral view, x 500, Arg7/541/2, (R/34). Fig. 10} Urnochitina urna Eisenack 1934. FC 99, lateral view, x 300, LP20/SB5/55. Fig. 11 Cingulochitina ervensis (Paris 1979). FC 100, Ger11/458/7, (N31/2); 11a, lateral view, x 250; 11b, detail of carina, x 1000. Fig.12 Cyathochitina sp. B, Paris 1981.FC 101, Arg3/SB7/1; 12a, lateral view, x 300; 12b, detail of carina, x 800. Fig. 13 ?Margachitina catenaria Obut 1973. FC 102, apertural view, x 400, Ger16/471/3. Fig.14 Eisenackitina bohemica (Eisenack 1934). FC 103, lateral view, Ger14/468/14, (Q26/4). 156 followed by C. cf. hemiesferica in the H Biozone, accompanied by C. lavidensis in the upper part of the H zone. Chelinospora media and C. cassicula are confined to the higher parts of the section studied Lower Lochkovian (upper EC & lower MN Biozones). OCCURRENCE. Argovejo, Geras, and La Vid sections Upper San Pedro Formation, lower MN (NA) Sub-Biozone. Chelinospora hemiesferica (Cramer & Diez) comb. nov. a sis, Se ell, sae 3 3) BASIONYM. Iberoespora hemiesferica Cramer & Diez, 1975: 341, pl. 2, figs 34-36. 1978c Cymbosporites dittonensis Richardson & Lister; Rodriguez: pl. 3, fig. 18. HOLOTYPE AND LOCUS TYPICUS. Cramer & Diez 1975: 334, 336; pl. 2, figs 34, 35: “base of a thick green shale 10 m below the exposed top of the San Pedro Formation at the westernmost outcrop in the village of Argovejo in the province of Leon, Spain’. EMENDED DIAGNOSIS. A Chelinospora with closely spaced nar- row muri, geniculate in plan, over the distal hemisphere and an equatorial to subequatorial area, where the muri become radial. Proximal exine thin, trilete mark labrate, labra narrow. DIMENSIONS. 30—40um (Cramer & Diez, 1975); 29-44 um (based on 75 specimens, present study), muri <1 um wide and high. REMARKS. The spores may be double-layered and on some speci- mens a membranous curvatural zone is seen bearing radial extensions of the distal muri. COMPARISON. The tightly packed geniculate narrow distal muri distinguish this species. Cramer & Diez (1975) refer to a ‘pro- nounced sculpture of brain-like muri’ distally. OCCURENCE. Distributed through the H to lower MN Biozone (NA Sub-Biozone). Chelinospora cf. hemiesferica (Cramer & Diez) comb. nov. Pl. 4, figs 6, 10; Pl. 9; fis: 4; Pl 10; fig: 1 DESCRIPTION AND COMPARISON. As for C. hemiesferica, but with broader, more widely spaced distal muri; muri more than 1—3 um wide and 1—2 um apart. OCCURRENCE. Found in all sections, Argovejo, Geras, La Peral J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND and La Vid sections, Upper San Pedro Formation; range hemiesferica, EC Biozones to lower MN (NA) Sub-Biozone; found in the hemiesferica Biozone only in the La Peral Section. Chelinospora lavidensis sp. nov. 1978c PI. 9, fig. 2 Iberoespora hemiesferica Cramer & Diez; Rodriguez: 418, pl. 3, fig. 15. 1983 Dictyotriletes geriense Rodriguez: pl. 6, fig. 6 only. DERIVATION OF NAME. HOLOTYPE. FM 1494 (PI. 9, fig. 2), diameter 33 um; sample Ger 9, slide D1/1, co-ord. 195 1056; E.F. no. U/36, upper San Pedro Formation, Geras section, Province of Leon, Cantabrian Mountains, northwest Spain. Named for the La Vid locality. DIAGNOsIS. A Chelinospora with narrow low muri, | um or less wide and >lum high, forming an irregular reticulum with broad lumina. DESCRIPTION. Proximally hilate miospores, equatorially 2-4 um thick, distal exine thick (not measured), amb subcircular to sub- triangular. Proximal hilum thin, frequently broken or absent, trilete mark not seen. Equatorially and distally sculptured with an irregular reticulum, muri | um or less wide, <1 um high, lumina highly variable in shape and size on a single spore, at the distal pole large and polygonal 8—10 um wide, subequatorially c. 1-2 um wide; muri become radial at the equator. Y-mark with narrow ‘folds’ extends c. 2/3 spore radius and merge into crassitude at the hilum margin, frequently gape over much of the hilum forming a distinct triangular area. DIMENSIONS. 29-43 um (based on 17 specimens). COMPARISON. Chelinospora hemiesferica (Cramer & Diez) comb. nov. has a distal sculpture of closely packed geniculate muri but resembles C. lavidensis sp. nov. because both species have an equatorial band of radial muri. REMARKS. Several species of Chelinospora (C. hemiesferica, C. sanpetrensis, C. media and C. lavidensis) have a subequatorial band of muri normal to the equator, which may indicate affinity or possibly represents a feature related to the development of the tetrad in murornate spores. OCCURRENCE. Upper part of H and EC Biozones, rare in lower MN (NA) Sub-Biozone, Upper Pridoli and lowermost Devonian, Argovejo, Geras and La Vid sections. PLATE 12 Figs 1,2 Pseudoclathrochitina carmenchui (Cramer 1964). 1, FC 104, lateral view, x 350, Arg11A/508/12, (Q27/4); 2, FC 105, LP20/SB5/54; 2a, lateral view x 400; 2b, basal margin detail, x 1100. Fig. 3 Margachitina elegans var. corta (Cramer 1964). FC 106, lateral view, x 300, Arg12/510/8, (K30/4). Fig.4 Urnochitina urna, Eisenack 1934. FC 107, chain of 3 individuals, lateral view, x 220, LV10/492/9, (N34). Fig.5 Margachitina elegans Taugourdeau & de Jekhowski 1960. FC 108, lateral view, x 300, LV12/494/8, (N25). Fig.6 Sphaerochitina sphaerocephala (Eisenack 1932). FC 109, lateral view, x 700, Arg5A/SB7/4. Fig.7 Fungochitina kosovensis Paris & Kriz 1984. FC 110, LP19/SB5/57; 7a, lateral view, x 200; 7b, basal margin detail, x 1250. Fig.8 Angochitina tsegelnjucki Paris & Grahn 1996. FC 111, lateral view, x 200, Ger14/468/8, (P31/4). Fig.9 Cingulochitina serrata (Taugourdeau & Jekhowski 1960). FC 112, LV12/494/8, (N24), 9a, chain of 11 individuals, lateral view, x 130; 9b, detail of connection, x 500. Fig. 10 Angochitina echinata Eisenack 1931. FC 113, LP8/CSB1/1; 10a, lateral view, x 350; 10b, neck detail, x 750. Fig. 11 Angochitina thadeui Paris 1981. FC 115, LV5a/CSB1/8; 11a, lateral view, x 200; 11b, apertural detail, x 650. Fig. 12 Angochitina ambrosi ambrosi Schweineberg 1987. FC 116, Ger8/461/8, (N38/3); 12a, lateral view, x 200; 12b, apertural detail, x 750. Fig. 13 Angochitina chlupaci Paris, Laufeld & Chlupac 1981. FC 117, lateral view, x 250, Arg13/455/2, (J28/2). Fig. 14 Angochitina echinata Eisenack 1931. FC 114, Ger9/456/7, (K31/1); 14a, lateral view, x 300; 14b, spiny ornament detail, x 850. PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND PALYNOLOGICAL ZONATION OF MID-PALAEOZOIC, NW SPAIN Pl. 9, fig. 6 Chelinospora mariae Rodriguez (pars): 10, pl. 1, fig. 4. Chelinospora media sp. nov. 1978a DERIVATION OF NAME. Latin medius a., intermediate. HOLOTYPE. FM 1495, sample Argovejo 92/12 (510) 3, 093 1019, E.F. no. K32/2 J32/4; Lower San Pedro Formation, Argovyejo section, Province of Leon, Cantabrian Mountains, northwest Spain. DIAGNosis. A Chelinospora with low muri, <1 um wide and < lum high, forming a regular reticulum with broad lumina of more or less even size. DESCRIPTION. Proximally hilate miospores, equatorially 2-4 um thick, distal exine thick (not measured); amb subtriangular. Proximal hilum thin, frequently broken, or absent, trilete mark not usually seen. Exine equatorially c. 5 um, distally thick equatorially, but not measured. Equatorially and distally sculptured with a reticulum, muri | um or less wide, 1-2 um high, lumina relatively large, polygonal and 6-14 um wide; muri become radial at the equator. A few specimens show a Y-mark, with sutures to the inner margin of the patina. DIMENSIONS. 31-60 um (based on 12 specimens); England, 37-67 um (based on 37 specimens). COMPARISON. Differs from Chelinospora lavidensis by the more regular reticulum with broad, more or less even, lumina. The holotype of Chelinospora mariae Rodriguez (1978a: pl. 1, fig. 3) appears to be conspecific with C. cassicula Richardson & Lister 1969, but other specimens figured by Rodriguez (1978a: pl. 1, fig. 4) appear identi- cal to C. media sp. nov. REMARKS. There appears to be a morphological gradation between Chelinospora lavidensis, C. media, and C. cassicu:la with some overlap in the ranges of the species. C. lavidensis occurs in the uppermost Pridoli and Lower Devonian (upper H, EC Biozones and lower MN Sub-Biozone) of the Cantabrian Mountains. C. media occurs near the Silurian/Devonian Boundary and in the lower MN Sub-Biozone, and is rare in the middle MN Sub-Biozone; C. cassicula occurs both in Spain and England in the lower MN Sub-Biozone, and especially the middle MN Sub-Biozone in England. Two tryads/dyads (Richardson, 1996a: pl. 3, figs 1, 2) show such close similarities in sculpture to C. media that they probably were derived from the same, or closely similar, plants. All these Chelinospora species represent part of an evolving spore morphon and were probably derived from a group of closely related plants. The dyads of the species Chelinohilates erraticus Richardson 1996b: pl. 7, figs 5—7, have identical sculptural patterns. OCCURRENCE. Upper San Pedro Formation, La Vid EC Biozone, Argoyejo and Geras sections, lower MN Biozone, uppermost Silurian and Lower Lochkovian. Occurs in the lower part of the lower MN (NA) Sub-Biozone, Shropshire and Herefordshire in England. 159 Chelinospora (Lophozonotriletes?) poecilomorpha (Richardson & Ioannides) comb. nov. Pl. 10, figs 6, 7 BASIONYM. Lophozonotriletes? poecilomorpha Richardson & Ioannides, 1973: pl. 7, figs 9-15; pl. 8, figs, 1, 4-6. HOLOTYPE. Richardson & Ioannides, 1973: pl. 7, fig. 10, BP 10/22- 831060, C1-34 borehole, Libya. REMARKS. This taxon is transferred to the genus Chelinospora because specimens studied under SEM and light microscopy show a distinct distal pattern with a thin polar proximal membrane. Chelinospora sanpetrensis (Rodriguez) comb. nov. Pl. 4, fig. 8; Pl. 7, fig. 10 BASIONYM. Brochotriletes sanpetrense Rodriguez, 1978c: 414, pl. 1, fig. 13. 1983 Brochotriletes sanpetrensis Rodriguez: 34, pls 1, 10, 19. HOLOTYPE AND LOCUS TYPICUS. Rodriguez, 1978c: pl. 1, fig. 13; Torrestio, Province of Le6n Cantabrian Mountains, northwest Spain, lower part of San Pedro Formation. EMENDED DIAGNOSIS. A Chelinospora with a densely murornate, foveolate, and irregular foveo-reticulate sculpture with broad, sinu- ous, low muri, and narrow lumina. DESCRIPTION. As in Rodriguez, 1978c: 414, but forms with a reticulum of wide muri and large rounded, polygonal and irregular lumina are classified as a separate species (C. cantabrica sp. nov.). DIMENSIONS. 24, 29-49 um (based on 60 specimens); 30-45 um. (Rodriguez, 1978c). REMARKS. The structure is patinate and consists of a thick distal exine and a thin proximal hilum, frequently ruptured, with a narrow fold on the inner margin of the curvaturae. Under the SEM some specimens show a sculpture of microrugulae and micrograna in the curvaturate zone. Specimens with a dominantly foveolate sculpture are referred to as Chelinospora cf. sanpetrensis but probably grade into the reticulate forms. OCCURRENCE. Argovejo, Geras, La Peral and La Vid sections, lower and middle San Pedro Formation; range RS and lower to middle H Biozones; isolated specimens found in lower EC Sub- Biozone (Geras) and lower MN (NA) Sub-Biozone (La Vid). Genus CYMBOSPORITES Allen 1965 TYPE SPECIES. Cymbosporites cyathus Allen 1965. Cymbosporites cf. dittonensis Richardson & Lister 1969 Pl. 9; fig. 7 DIMENSIONS. 22-35 um (based on 6 specimens). PLATE 13 Figs 1,3c Palenchitina pisuergensis Schweineberg 1987. FC 118, Ger7/SB7/6; 1a, lateral view, x 300; 1b, ornament detail, x 1000; 3c, enlargement of part of Fig. la, x 1000. Fig. 2 Ancyrochitina valladolitana Schweineberg 1987. FC 119, Ger4/CSB3/S; 2a, lateral view, x 355; 2b, appendix detail, x 750. Fig.3 Ancyrochitina javieri Schweineberg 1987. FC 120, Ger7/SB7/6; 3a, lateral view, x 300; 3b, basal margin detail, x 650. Fig. 4 Angochitina elongata Eisenack 1931. FC 121, Ger9/456/7, (M32/1); 4a, lateral view, x 300; 4b, ornament detail, x 850. Fig.5 Ancyrochitina fragilis brevis Taugourdeau & Jekhowski 1960. FC 122, LV10/492/9, (L36/4); 5a, lateral view, x 300; 5b, appendix detail, x 1000. Fig. 6 Ancyrochitina libyensis Jaglin 1986. FC 123, Arg4/SB7/3; 6a, lateral view, x 500; 6b, appendix detail, x 850. Fig. 7 Ramochitina villosa (Laufeld 1974). FC 124, Ger2B/CSB3/3; 7a, lateral view, x 400; 7b, apertural detail, x 500. Fig.8 Plectochitina rosendae Cramer & Diez 1978. FC 125, lateral view, x 400, LV 10/492/9, (H35/1). Fig.9 Plectochitina carminae Cramer 1964. FC 126, LV12/494/7, (P22/4); 9a, lateral view, x 300; 9b, appendix detail, x 600. 160 REMARKS. The Cantabrian material has close-packed wide based coni to spinose elements, some biform. The elements are not mixed with minute cone as in typical forms of the species. OCCURRENCE. Rare, Upper San Pedro Formation, Argovejo, Lower MN (NA) Sub-Biozone. Cymbosporites sp. B Pl. 10, fig. 3 DESCRIPTION. Amb subtriangular to subcircular, usually preserved in polar compression. Proximal surface laevigate, contact areas covered by a thin diaphanous exine (hilum), near the periphery of the contact areas are distinct arcuate folds, which form a constant character, usually destroyed except near the margin. Outside the contact areas the exine is much thicker, equatorially and distally, 1.5—2 um (measured equatorially). Sculpture confined to the equato- rial margin and distal hemisphere; consists of broad based coni rounded in plan and evenly tapered in profile, c. 1.5 um wide and >1 um high; sculptural elements spaced, but with insufficient distance between them for elements of a similar size. Triradiate mark not seen apart from curvatural invaginations at the radial apices. DIMENSIONS. 26-31 um (based on 11 specimens). COMPARISONS. Similar to Cymbosporites cf. catillus Allen 1965 in Richardson & Lister 1969 except that the coni are broad-based and larger. REMARKS. Other specimens which may belong to the genus have been found in the Argovejo and Geras sections, namely ?Cymbosporites sp. A (Pl. 10, fig. 2) and Cymbosporites sp. (Pl. 10, figs 4, 5). OCCURRENCE. Upper San Pedro Formation, Argovejo, Geras, and La Vid, EC Biozone and rare lower MN (NA) Sub-Biozone. TAXONOMY OF CHITINOZOA As the main purpose of this paper is to define a spore biozonation for the Cantabrian region, only Ramochitina villosa (formerly Gotlandochitina Laufeld 1974) is described briefly here. A more detailed investigation of the chitinozoans from the Cantabrian Area is to be presented in a forthcoming publication. All material is deposited in the collections of the Natural History Museum, London. Order PROSOMATIFERA Eisenack 1972 Family LAGENOCHITINIDAE Eisenack 1931, emend Paris 1981 Subfamily ANGOCHITININAE Paris 1981 Genus RAMOCHITINA Sommer & van Boekel 1964, emend. Paris et al. 1999 TYPE SPECIES. 426, pl. 1, fig. 3. Ramochitina ramosi Sommer & van Boekel, 1964: Ramochitina villosa (Laufeld 1974) AL, WS), tie, 7) 1974 Gotlandochitina villosa Laufeld: 95, figs 7a, b. 1994 Gotlandochitina villosa Laufeld; Sutherland: 65, pl. 17, figs 7-14. HOLOTYPE. LO 4587 T, figured Laufeld, 1974: fig 56a. DIAGNOsIS. (After Laufeld, 1974: 95). Body spheroidal, neck cylindrical to subcylindrical; broadly rounded flexure. Ornament of spines which may exhibit simple branching; spines on apertural J.B. RICHARDSON, R.M. RODRIGUEZ AND S.J.E. SUTHERLAND portion of vesicle curved in aboral direction and in opposite direction on the apertural portion of vesicle; spines on neck decrease in size towards oral pole. DISCUSSION. In the generic revision of Paris, Winchester-Seeto, & Grahn (2000) Gotlandochitina Laufeld 1974 is considered a junior synonym of Ramochitina Sommer & van Boekel 1964. G. villosa is here transferred to Ramochitina. ACKNOWLEDGEMENTS. We are indebted to the Natural Environment Re- search Council for research grant GR3/09624: to Katie Bagshaw and Denise Darwin for technical assistance and for preparing material and picking spores for light and SEM microscopy: Peter York is thanked for the light photomicrographs, Chris Jones and Alex Ball for help with SEM studies; and Derek Adams, Natural History Museum Photographic Unit, who composed the plates electronically. REFERENCES Achab, A. & Asselin, E. 1993. Upper Silurian and Lower Devonian chitinozoan microfaunas in the Chaleurs Group, Eastern Canada. Special Papers in Palaeontol- ogy, 48: 7-15, pl. 1. Achilles, H. 1981. Die ratische und liassiche Microflora Frankens Palaeontographica, B 179: 1-86, pls 1-17. Allen, K. C. 1965. Lower and Middle Devonian spores of North and Central Vestspitsbergen. Palaeontology, 8: 687-748, pls 94-108. Arkhangel’skaya, A. D. 1980. Plant spores from some Lower Devonian sections of the western regions of the Russian Platform (in Russian). Jn, Urvanova, V. N. & Nemtinova, G. I., Palynological research in the Proterozoic and Phanerozoic of oil- and gas-bearing regions of the USSR. Trudy Vnigni, Moscow, 217: 26-46, 140-44, pls 5-11. Barrois, Ch. 1882. Récherches sur les terrains des Asturies et la Galice. Memoires de la Société Géologique du Nord, Lille, 2: 630 pp., 20 pls. Burgess, N.D. 1991. Cryptospores and miospores from the Type Llandovery Area, south-west Wales. Palaeontology, 34: 565-99, pls 1, 2. Burgess, N. D. & Richardson, J. B. 1991. Silurian cryptospores and miospores from the type Wenlock area, Shropshire, England. Palaeontology, 34: 601-28, pls 1, 2. & 1995. Late Wenlock to Early Pridoli cryptospores and miospores from south and south-west Wales, Great Britain. Palaeontographica, B 236: 1-44, pls 1— 10. Chaloner, W. G. & Streel, M. 1968. Lower Devonian spores from South Wales. Argumenta Paleobotanica, 1: 87-101, pls 19, 20. Comte, P. 1934. Sur les couches intermédiares entre le Silurien de Dévonien dans les Asturies. Comptes Rendu de |’Academie des Sciences, 198: 1-166. 1937. La série Cambrienne et Silurienne de Le6n. Compte Rendu de |’'Academie de Sciences, 204: 604 pp. 1959 Recherches sur les terrains anciens de la Cordillére Cantabrique. Memorias Instituto Geolégico y Minero de Espana, Madrid, 60: 13-425. Cramer, F.H. 1964a. Some acritarchs from the San Pedro Formation (Gedinnian of the Cantabric Mountains in Spain). Bulletin de la Société Belge de Géologie, de Paléontologie et d’Hydrologie, 73: 33-38, pls 1, 2. — 1964b. Microplankton from three Palaeozoic formations in the province of Leon (NW Spain). Leidse geologische mededelingen, 30: 255-361, pls 1-24. —— 1966a. Palynomorphs from the Siluro-Devonian boundary in NW Spain. Notas y Comunicaciones del Instituto Geologico y Minero de Espana, 85: 71-82, pls 1-3. 1966b. Chitinozoans of a composite section of Upper Llandovery to basal Lower Gedinnian sediments in northern Leon, Spain. A preliminary report. Bulletin de la Société Belge de Géologie, 75 (1): 69-129, pls 1-5. 1966c. Hoegispheres and other microfossils incertae sedis of the San Pedro Formation (Siluro-Devonian Boundary) near Val porquero, Leon, NW Spain. Notas y Comunicaciones, del Instituto Geologico y Minero de Espana, 86: 69-129, pls 1, 2. 1967. Palynology of Silurian and Devonian rocks in northwest Spain. Boletin del Instituto Geologico y Minero de Espana, 77: 223-286, pls 1+. — 1970. Distribution of selected Silurian acritarchs. Revista Espanola de Micropaleontologia Numero extraordinario, 1: 203 pp., 23 pls. —— & Diez, M. d. C. R. 1968. Consideraciones taxonémicas sobre los acritarcos del Silurico medio y superior del Norte de Espana. Los acritarcos acantomorfiticos. Boletin del Instituto Geoldgico y Minero de Espana, 79: 541-574, 21 pls. — & 1975. 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Pp. 1-83. £43.40 No. 2 Terebratula californiana Kiister, 1844, and reappraisal of west coast north American brachiopod species referred to the genus Laqueus Dall, 1870—Late Campanian-Maastrichtian corals from the United Arab Emirates-Oman border region— Rhombocladia dichotoma (M‘Coy, 1844) [Fenestrata, Bryozoa]: designation of a lectotype—The Gough’s Cave human fossils: an introduction—The Creswellian (Pleistocene) human axial skeletal remains from Gough’s Cave (Somerset, England)—The Creswellian (Pleistocene) human lower limb remains from Gough’s Cave (Somerset, England). 2000. Pp. 85-161. £43.40 CONTENTS 82 The Cenozoic Brachiopod Terebratula: its type species, neotype, and other included species D.E. Lee, C.H.C. Brunton, Emma Taddei Ruggiero, Massimo Caldara & Oronzo Simone 95 Gough’s Cave 1 (Somerset, England): a study of the pectoral girdle and upper limbs S.E. Churchill 109 Systematic affinity of Acroporella assurbanipali Elliott (Dasycladaceae), with notes on the genus Neomeris Filippo Barattolo & Roberta Romano 115 Palynological zonation of Mid-Palaeozoic sequences from the Cantabrian Mountains, NW Spain: implications for inter-regional and interfacies correlation of the Ludford/Ptidoli and Silurian/Devonian boundaries, and plant dispersal patterns John B. Richardson, Rosa M. Rodriguez, Stuart J. E. Sutherland Bulletin of The Natural History Museum GEOLOGY SERIES Vol. 57, No. 2, November 2001