Revision of Late Devonian (Famennian) and some Early Carboniferous (Tournaisian) crinoids and...

R E V I S I O N O F L A T E D E V O N I A N ( F A M E N N I A N ) A N D

S O M E E A R L Y C A R B O N I F E R O U S ( T O U R N A I S I A N )

C R I N O I D S A N D B L A S T O I D S F R O M T H E T Y P E

D E V O N I A N A R E A O F N O R T H D E V O N

by N . G . L A N E , C . G . M A P L E S and J . A . W A T E R S

ABSTRACT. The crinoids and blastoids from the Pilton (Beds) Formation of the type Devonian of north Devonshire arerevised. These fossils were monographed by the Rev. G. F. Whidborne in 1898, but have not been studied since thattime. Recent studies on various groups of fossils from the Pilton and related rocks in North Devon con®rm that thegreat majority of these fossils are Famennian, although three specimens from Fremington are probably EarlyCarboniferous (Tournaisian). We identify four blastoid taxa from a fauna that is sparse and poorly preserved; twospiraculates, one ®ssiculate, and one taxon unidenti®able at the ordinal level. Mesoblastus cf. M. crenulatus from theGattendor®a Zone (Lower Carboniferous) near Fremington is the oldest known representative of this genus. Thecrinoid fauna is somewhat more diverse, but the preservation is equally poor. No changes are made in the ¯exiblecrinoids. Among camerate crinoids, one species is reassigned to Eumorphocrinus and one is retained in Actinocrinites.Specimens of some crinoids, such as Rhodocrinites and Megistocrinus, are so poorly preserved that certainidenti®cation was not possible. The hexacrinoid Adelocrinus, relegated to uncertainty for 150 years, is here shownto be a valid genus that is very similar to Arthroacantha, but not synonymous with it. Among the cladid crinoids, thedominant groups are those within the Superfamily Scytalocrinacea, which includes Bridgerocrinus, Sostronocrinus,and Scytalocrinus, all of which are placed in the new family Sostronocrinidae. One new species, Glossocrinuswhidbornei, is named. Non-pinnulate cladids, common in older Devonian rocks, do not occur. The fauna showsconsiderable similarity with faunas from eastern North America and Germany. It shows less resemblance to theextensive Famennian crinoid and blastoid fauna of north-western China, despite some remarkable congruencies,especially the occurrence of very similar species of Actinocrinites in these widely separated areas.

KEYWORDS: Devonian, Famennian, Carboniferous, Tournaisian, England, crinoid, blastoid.

T H I S paper is a revision of the Late Devonian crinoids and blastoids described and illustrated byWhidborne (1896, 1898) from the Pilton Formation of North Devon. We do not include otherechinoderms, star®shes and echinoids, nor do we include Middle Devonian crinoids from Devon,mainly Hexacrinites, that Whidborne described in earlier parts of his extensive monograph on thefaunas of the type Devonian.

The rocks were ®rst described by De la Beche (1839). For the general setting of the Devonian of NorthDevon see House and Selwood (1964), Edmonds et al. (1975), Goldring et al. (1978), Durrance andLaming (1982), House and Dineley (1985), Bluck et al. (1988), and Dineley (1992). All of the LateDevonian crinoids described by Whidborne were collected from a relatively small area near Barnstaple, onthe Taw estuary (Hambling and Rogers 1910). This area is on the north ¯ank of a major synclinorium inDevonian and younger rocks, with older rocks to the north, in the Ilfracombe area, and younger, EarlyCarboniferous, rocks to the south.

Both the age of the echinoderm fossils and their taxonomic assignments have changed, more or lessarbitrarily, despite the lack of taxonomic revision or careful inspection of the specimens reported heresince Whidborne's time. Although Whidborne (1896, 1898) considered the crinoids to be Devonian in age,he used a series of generic names that were almost exclusively characteristic of Early Carboniferouscrinoids at the time he worked. These include Actinocrinites, Platycrinites, Poteriocrinites, and

[Palaeontology, Vol. 44, Part 6, 2001, pp. 1043±1080, 6 pls] q The Palaeontological Association

Scytalocrinus. Whidborne did not discuss this Early Carboniferous aspect of the fauna, but it is apparentthat he must have believed that the crinoids had distinct Early Carboniferous af®nities. By 1898 numerousgeneric names had been applied to Devonian crinoids in North America, especially to those described fromNew York State (e.g. Hall 1859, 1862), yet Whidborne did not cite any of this work. Those NorthAmerican studies that he did cite all concerned Mississippian crinoids.

In 1930, Schmidt assigned species described by Whidborne to the Early Carboniferous and changedsome of the generic assignments. He placed several species in Hydriocrinus, a Late Carboniferous genus,and assigned Adelocrinus hystrix Phillips, 1841 to Arthroacantha. He compared the entire fauna from thePilton Formation with the crinoids of the Etroeungtian Stage of Germany, regarding both as EarlyCarboniferous in age.

In 1943, Bassler and Moodey followed Schmidt's lead and essentially repeated his taxonomic and ageassignments, placing the Pilton Formation in the Lower Carboniferous and changing generic assignments.No activity directly related to the Pilton crinoids and blastoids has been undertaken since 1943, althoughmuch modern, detailed work has been done on other groups of fossils, especially those useful inbiostratigraphy, that occur in these beds (see below).

A G E O F E C H I N O D E R M - B E A R I N G U P P E R D E V O N I A N S T R A T A

A great deal of detailed stratigraphic work has been accomplished on the type Devonian of south-westEngland during the past 25 years (House and Selwood 1964; Webby 1966; House 1977; Dineley 1986).This includes several studies of the conodonts (Dineley and Rhodes 1956; Austin et al. 1985; Castle 1985),goniatite cephalopods (Butcher and Hodson 1960; House 1963), trilobites (Goldring 1955; Kennedy1994), and ostracodes (Gooday 1975). A majority of these studies have been in areas other than those thathave yielded Late Devonian crinoids or blastoids, and hence provide only indirect evidence for the age ofthe Pilton (Beds) Formation, from which the echinoderms were collected (Ziegler and Sandberg 1990;Oliver and Chiupac 1991). All of these fossil groups provide some direct evidence for the age of the PiltonBeds. Kennedy (1994) provided a long list of trilobite localities in the Barnstaple area, many of which havealso yielded crinoids. All such localities in the Pilton Beds north of the Taw estuary are regarded here asLate Famennian or Famennian in age. Only exposures south of the Taw estuary, near Fremington, wereregarded as latest Famennian±Early Carboniferous in age by Goldring (1955) and Kennedy (1994). Basedon these age determinations, it seems that all of the localities cited in Whidborne, except for those atFremington, should be regarded as Famennian in age. The Fremington fossils are either earliest EarlyCarboniferous or Famennian. These latter include one specimen of Taxocrinus macrodactylus, onespecimen of Abactinocrinites? sp., and the spiraculate blastoid Mesoblastus cf. M. crenulatus.

For the purposes of this study, all crinoid and blastoid localities north of the Taw estuary are regarded asFamennian in age. Those localities south of the estuary are regarded as earliest Early Carboniferous. Atleast some of the northern localities are probably latest Famennian and time-equivalent to the Etroeungtbeds of Belgium and Germany.

Conodonts collected by Dineley and Rhodes (1956, p. 244, their localities 6 and 7) from Pilton Bedsnear Saunton are judged to be Late Devonian. Dineley later collected conodonts from Croyde Bay(reported in Goldring 1970, p. 811) that also are judged to be Late Devonian. Cephalopods collected fromthis area include Gattendor®a from Pilton B and Ammonellipsites from Pilton C (Prentice 1960). Trilobitesalso provide age information, especially Phacops in older rocks and Cyrtosymbole in younger beds.Palynomorphs from the Barnstaple area have been reported by O'Liathain (1992). He recognized that thelower Pilton Beds were latest Famennian, in the LN palynomorph zone of Higgs et al. (1988), which is thehighest zone in the Famennian. He also placed the Devonian/Carboniferous boundary at Fremington, thusessentially agreeing with other assessments that Pilton Beds north of the Taw estuary are Famennian andbeds south of the estuary are Tournaisian. Brachiopods are also useful for comparison of Pilton faunas withthose elsewhere in Great Britain and in western Europe (Paul 1937, 1939).

Conclusions based on these various fossils almost unanimously agree now that the Baggy Formation,immediately below the Pilton Beds, is in the Clymenia cephalopod zone and is slightly older than theEtroeungtian. Pilton Beds units A1±A3 are all in the Wocklumeria Zone and Etroeungtian in age. Pilton B,

1044 P A L A E O N T O L O G Y , V O L U M E 4 4

which belongs to the Gattendor®a Zone (Hangenberg Kalk), is Early Carboniferous, and has yielded a fewcrinoid and blastoid specimens from the Fremington area. Pilton C and D are clearly both EarlyCarboniferous in age and have yielded no crinoids or blastoids.

L O C A L I T I E S A N D L O C A L S T R A T I G R A P H Y

All but one (Brushford) of the localities cited by Whidborne from which crinoids or blastoids werecollected from the Pilton or Marwood beds occur in north Devon in the area around, or a few miles north-west of, Barnstaple (Text-®g. 1). The Brushford locality is in west Somerset, on the south-east border ofExmoor National Park. It is from this area that Phillips (1841) obtained his specimens of Taxocrinus andAdelocrinus. All of the localities cited by Whidborne from north Devon are shown in their relativegeographic position in Text-®gure 1. These sites form a north-west±south-east band of localities,re¯ecting the regional strike of these rocks, and thus are all within relatively close stratigraphic proximity.Goldring (1970) estimated the thickness of the Pilton Beds to be about 700 m. Serious structuralcomplexities, both folding and faulting, make detailed correlation and measurement dif®cult. Never-theless, the crinoid localities seem to indicate that comparable facies provide a coherent pattern. The PiltonBeds, or Pilton Formation, are underlain by the Baggy Formation, which also was considered Famennianin age by Kennedy (1994). An older name, no longer in use, is the Marwood (�Baggy) beds (Text-®g. 1).The only crinoids cited by Whidborne from these slightly older rocks are two species listed as occurringsouth-west of Sloly. Baggy Point and the rocks around Marwood have not yielded any crinoids orblastoids.

Goldring (1955) divided the Pilton Formation into three members: A±C (a higher unit, D, wasproposed by Prentice, 1960). Unit A, the oldest, includes three subunits: 1±3. The lower part of A(units A1 and A2) was studied in coastal sections at Croyde Bay. These beds have yielded few, if any,crinoids or blastoids. Unit A3, from which most of the crinoids and blastoids are judged to have beencollected, is based on exposures at Poleshill Quarry in Barnstaple. Other nearby crinoid and blastoidlocalities in and around Barnstaple are presumed to be of approximately the same age. Unit B is basedon rocks near Fremington that have yielded a few specimens of crinoids and a blastoid. These rocksare thought to be highest Famennian (upper Etroeungt) and earliest Early Carboniferous in age. Piltonunit C, named from exposures at the Ladywell Woods quarry, has yielded no identi®able crinoids orblastoids, although crinoid debris is common. This unit and unit D are both de®nitely EarlyCarboniferous in age.

Three Pilton localities that have yielded crinoids have been included in Joint Nature ConservationCommittee list of marine Devonian localities worthy of conservation. We quote here, with permission, theunpublished statements of the Joint Committee review series regarding these localities.

Baggy Point. This site is the type section of the Baggy Formation but also exposes good, representativesections of the Upcott Beds and Lower Pilton Beds. The Baggy Formation displays an extensive rangeof sedimentary structures, including numerous bioturbation traces, which are of great importance inestablishing a shallow marine, deltaic environment for these sediments, thus assisting the interpretation ofthe local and regional palaeoecology and palaeogeography. The site also includes Laticosta Cave, a well-known Pilton Beds fossil locality. The site as a whole demonstrates a wide range of contrasting facies(marine and non-marine) in the Famennian (Upper Devonian).

Downend. In the coastal section around Downend the lithological and sedimentological characters of theLower Pilton Beds (Upper Devonian) are well displayed. A rich fauna is indicative of a late Famennianage. Tuff bands are present in the section, which indicate the occurrence of penecontemporaneous volcanicactivity in the area, and these form useful local marker-horizons, particularly as this section has undergonetectonic thickening. In the late Devonian, a transgression northwards brought about deeper-waterconditions during the deposition of the Pilton Beds. The transition from shallow-water sandstones andsiltstones to deeper-water shales with cherts approximately coincided with the faunal changes marking theDevonian/Carboniferous boundary.

L A N E E T A L . : C R I N O I D S A N D B L A S T O I D S 1045

Fremington Quay. While the boundary between Devonian and Carboniferous has been recognized indisturbed thrust-sheets in north Cornwall, Fremington Quay is unique in displaying the boundary in aconformable succession, little affected by tectonism. The beds here are a uniform sequence of dark shalesthat are richly fossiliferous. Late Devonian (Wocklumeria Zone) and Early Carboniferous (Gattendor®aZone) faunas have been recognized here, and this section offers a unique opportunity for the detailedsampling of fossils to establish precisely the succession of faunas across the boundary.

C O L L E C T O R S A N D C O L L E C T I O N S

Echinoderms from the type Devonian area are scattered among several museums in Great Britain. Below,we list these museums and the collections they hold that either we have visited personally or from whichwe have been able to borrow specimens.

The North Devon Museum in Barnstaple holds a large collection of crinoids assembled by TownsendM. Hall (1845±1899) from the Pilton Beds. Whidborne utilized many specimens from the Hall collection,but he failed to credit Hall for any of them. All specimens cited by Whidborne as coming from the`Barnstaple Athaneum,' unless otherwise attributed, are from the Townsend Hall collection. For manyyears the museum was known as the North Devon Athaneum. A change in ownership and name occurredin 1986. In addition to his collection, Hall published several papers on the local geology and fossils (Hall1867, 1878). None of these papers deals with fossil echinoderms. The museum also has a few specimenscollected in the 19th Century by a Miss Partridge. We were unable to learn any more about this collector.One of her specimens was described and illustrated by Whidborne as Poteriocrinus tensus (1898, p. 226,pl. 38, ®g. 1). The museum also has a few specimens donated by Dr and Mrs Coomara Swamy. One ofthese specimens, Platycrinus? anguliferus, was described and illustrated by Whidborne (1891, p. 223,


TEXT-FIG. 1. Map of crinoid and blastoid localities cited by Whidborne (1898) in the Pilton and Marwood beds ofNorth Devon. Towns and villages are shown in larger type, outcrops or fossil sites in smaller type; modi®edfrom Ordinance Sheet 180, Barnstaple and Ilfracombe area, 1:50,000; inset shows position of detailed map in

south-west England.

pl. 37, ®g. 8). In addition to the crinoid specimens, two specimens of blastoids (Petaloblastus? ovalis) fromAshford and Bradiford are housed in the collections.

The Sedgwick Museum, University of Cambridge, holds two important collections of North Devoncrinoids. These are the William Porter collection and the Rev. G. F. Whidborne collection. All of thespecimens in the museum are illustrated Whidborne types, including his type of the blastoidPentremitidia phillipsii. The museum has no non-type specimens, nor any specimens collected afterWhidborne's publication in 1898. All of the specimens are mounted on blue boards with glue and areon display.

The British Geological Survey in Nottingham holds several important specimens, including the typespecimens of Phillip's (1841) Adelocrinus hystrix and Taxocrinus macrodactylus. In addition, it has twospecimens collected in the 20th Century by Wyatt Edgell.

The Natural History Museum, London, holds several specimens donated by J. G. Hamling in 1916. Mostof these specimens were collected in the 19th Century. The bulk of Hamling's collection is in the TorquayNatural History Museum. Whidborne credited Hamling with one specimen in his monograph. The NaturalHistory Museum, London, also holds one specimen donated or purchased by A. Champernowne in1909, and a specimen obtained from Townsend M. Hall in 1886. In addition, the museum has aphaenoschismatid blastoid collected by Roland Goldring in 1954.

The Department of Geology, University of Bristol, holds four specimens collected by Roland Goldringin 1953±54. These include one good specimen of a blastoid (Mesoblastus cf. M. crenulatus) fromFremington. The Bristol City Museum has no specimens pertinent to this study.

The Torquay Natural History Museum holds the Hamling collection, which is quite large and in quitegood curatorial condition. The collection was donated by Hamling in 1904. Unfortunately it containsrelatively few crinoids from the Pilton Beds. There are numerous large specimens of Hexacrinites from theMiddle Devonian and numerous non-echinoderm specimens from the Lower Carboniferous Culm.

The Royal Albert Memorial Museum in Exeter holds the Inkerman Rogers collection. This wasoriginally in the Bideford Museum, which has closed; the collection was transferred in 1981. It is in a verypoor condition, dif®cult to access and study, and unorganized. The museum holds original maps by Rogersof the Lower Culm on the south side of the Taw estuary. These are very detailed and every outcrop ismarked and described. Rogers made these maps in 1906. They show the rocks around Fremington as beingall in the Lower Carboniferous. The museum houses the P. W. Sladen collection of echinoderms, which ison display and quite well curated. However, the specimens are mainly North American or from Clitheroe,England. None was of use for this study. A small collection donated by N. Whitley contains a fewspecimens from Barnstaple and Marwood.

The University of Exeter holds no specimens of crinoids or blastoids from North Devon (E. B. Selwood,pers. comm. 1994).

T E C H N I Q U E S

All of the Pilton specimens seen by us are moulds that occur in highly cleaved, ®ne-grained mudstones tophyllites. Some specimens have been sheared and distorted. Nevertheless, many specimens reveal ®nedetails of ornament and plate boundaries. Silicone rubber casts were made for all specimens observed.Each specimen was sprayed ®rst with KrylonTM clear plastic spray, matte ®nish, to seal the surface. A thincoat of petroleum jelly also was applied to the specimens as a release coating. Because the jelly wasapplied with Q-tipsTM, small cotton ®bers caught in the jelly were encased in the rubber. Therefore, ourcasts were given a `shave' before they were photographed. We do not recommend using Q-tipsTM to applythe petroleum jelly. The silicone rubber used was RTV-2, GI-570, base and catalyst (produced by SiliconeInc., PO Box 363, High Point, NC 27261, USA). This rubber was chosen from many because it was softenough to release without damaging the specimens and it stiffened overnight so that casts could beremoved the day after they were poured.

Photography of the casts proved to be dif®cult. Ordinarily, the specimen would be coated with Kodakspotting opaque, either red or black, and then a coating of ammonium chloride sublimate applied. Thisprovides maximum contrast and helps bring out plate boundaries. Virtually nothing we tried would adhere


to the rubber. Eventually, however, we found that a coating of clear acrylic spray sealed the rubber andmade the coating of specimens easier.

S Y S T E M A T I C P A L A E O N T O L O G Y

Most genera and species are discussed here in the same order as they occur in Whidborne's (1898)monograph: ®rst blastoids, then ¯exibles, camerates, and cladids. Some re-allocations were necessary as aresult of different familial and generic assignments (Table 1).

Abbreviations. Measurements: W, width; H, height; R, radial; DC, dorsal cup; Cr, crown; BC basal circlet.Repositories: BGS, British Geological Survey, Nottingham; NHM, The Natural History Museum, London; UB,University of Bristol, Department of Geology; NDM, North Devon Museum, Barnstaple; TNHS, Torquay NaturalHistory Society; SM, Sedgwick Museum, University of Cambridge.


TABLE 1. Devonian (Famennian) crinoid and blastoid taxa reported in Whidborne (1896, 1898), their current taxonomicassignments, and veri®ed number of specimens for each taxon. Specimens not seen by Whidborne are indicated by ±.Taxa are listed in the same order in which they appear in the text.

Whidborne (1896, 1898) Lane et al., herein No. of specimens

BlastoideaPentremitidea phillipsii Petaloblastus? ovalis 3Codonaster conicus Order, family, and genus indeterminate 1

± Phaenoschismatid, genus and species indeterminate 1Crinoidea

FlexibiliaTaxocrinus macrodactylus Taxocrinus macrodactylus 9Taxocrinus stultus Taxocrinus stultus 3

CamerataRhodocrinus?, ?sp. Order, family, genus, and species indeterminate A 1Megistocrinus?, sp. Order, family, genus, and species indeterminate B 1Actinocrinus porteri Eumorphocrinus porteri 6Actinocrinus batheri Actinocrinites batheri 1

± Platycrinites sp. 1Platycrinus? anguliferus (part) Platycrinites? sp. 3Platycrinus? anguliferus (part) Order, family, genus, and species indeterminate C 3Adelocrinus hystrix Adelocrinus hystrix 8

CladidaScaphiocrinus transcisus Poteriocrinites transcisus 1

± Glossocrinus whidbornei sp. nov. 2Poteriocrinus sp. (part) Glossocrinus? sp. 2Poteriocrinus sp. (part) Glossocrinus? salebrosus 1Scaphiocrinus? salebrosus Glossocrinus? salebrosus 1Cyathocrinus barumensis Scytalocrinus barumensis 2Poteriocrinus stadiodactylus Bridgerocrinus stadiodactylus 1Poteriocrinus batheri Bridgerocrinus stadiodactylus 1Scytalocrinus stadiodactylus Bridgerocrinus stadiodactylus 3Scytalocrinus arachnoideus Bridgerocrinus? arachnoideus 4

± Bridgerocrinus? sp. 1Mariocrinus? [sic] mundus Sostronocrinus mundus 2Poteriocrinus tensus Aphelecrinus tensus 2Scaphiocrinus sp. Aphelecrinus? sp. 1Scaphiocrinus? plumifer Pachylocrinus? plumifer 5Coeliocrinus n. sp. Genus and species indeterminate 1Scaphiocrinus? inordinatus Family and genus uncertain 3

Class BLASTOIDEA Say, 1825Order GRANATOCRINIDA Bather, 1900 (emend. Waters and Horowitz, 1993)

Family GRANATOCRINIDAE Fay, 1961

Genus MESOBLASTUS Etheridge and Carpenter, 1886

Mesoblastus cf. M. crenulatus (Roemer, 1851)

Plate 1, ®gure 4

*1851 Pentatrematites crenulatus Roemer, pp. 366±367, pl. 4, ®g. 15a±d.1854 Pentremites crenulatus (Roemer); De Koninck and Le Hon, pp. 199±200, pl. 7, ®g. 4a±d.1862 Pentremites crenulatus (Roemer); Dujandin and Hupe, p. 96.1878 Pentremites crenulatus (Roemer); Bigsby, p. 233.1886 Mesoblastus crenulatus (Roemer); Etheridge and Carpenter, pp. 183±184, pl. 4, ®gs 1, 2; pl. 6,

®gs 8±10.1899 Mesoblastus crenulatus (Roemer); Bather, pp. 16±17.1903 Mesoblastus crenulatus (Roemer); Hambach, p. 65.1937 Mesoblastus crenulatus (Roemer); Fritz and Cline, p. 311.1943 Mesoblastus crenulatus (Roemer); Bassler and Moodey, pp. 222±223.1943 Mesoblastus crenulatus (Roemer); Chi, p. 112.1961 Mesoblastus crenulatus (Roemer); Fay, pp. 76±77, pl. 47, ®gs 1±13, text-®gs 151±157.1967 Mesoblastus crenulatus (Roemer); Macurda, pp. 477±480, pl. 60, ®gs 1±11, 13±15; pl. 61, ®gs 31±34,

47; text-®g. 6E.

Remarks. Roland Goldring collected a partial mould of a small blastoid in 1954 from Fremington that hetentatively identi®ed as Pentremites ovalis Goldfuss. The specimen is UB 18343. It consists of a thecalmould in lateral view, two sets of pinnules and partial stem. The oral surface is not preserved and thebasalia are only partially preserved. Although a mould, the surfaces of the plates are well preserved,showing prominent growth lines. The proximalmost 4 mm of stem is preserved and consists of thin discs(approximately 6 per mm). Morphology of the facets could not be observed. Two sets of brachioles arepreserved with an observed length of approximately 5 mm.

Although the oral surface, which is critical in generic identi®cation of blastoids, is notpreserved, enough morphology is retained to reject Goldring's identi®cation of this specimen asPentremites ovalis (now Petaloblastus ovalis) and questionably to assign the specimen toMesoblastus crenulatus (Roemer). P. ovalis has an elongate cone-shaped basalia (Fay 1961), butUB 18343 has slightly protuberant basals very similar to those of M. crenulatus (Macurda 1967,p. 477) specimens of equivalent size. The ambulacra of P. ovalis are broadly lanceolate withlancets fully exposed, and have a ¯at to slightly concave cross-sectional pro®le. In contrast, UB18343 has more narrowly lanceolate ambulacra with the suggestion that lancets were only partiallyexposed and a convex cross-sectional pro®le. These are ambulacral characteristics of M. crenulatus.Finally, UB 18343 shows two prominent hydrospire pores per side plate and a well-de®ned RDsector with distinct growth lines. Both of these characters are present in M. crenulatus, but absentin P. ovalis.

Macurda (1967) reviewed the occurrences of Mesoblastus as a part of his monograph on the Tournaisianblastoids of Belgium. Mesoblastus is a part of that fauna and is late Tournaisian; however, it is also foundin Lower Visean rocks of northern England (Macurda 1967). Species of Mesoblastus described from China(Chi, 1943) belong to other genera and await taxonomic reassignment.

Goldring assigned a stratigraphic position of Pilton Beds B, Gattendor®a Zone, to the Fremingtonlocality where Mesoblastus cf. M. crenulatus was collected. The Gattendor®a Zone is the lowestammonoid zone in the Hastarian Stage (Harland 1989) of the Tournaisian (Tn1b); therefore, theFremington occurrence of Mesoblastus is older than the occurrences in Belgium. As currently understood,Mesoblastus is con®ned biogeographically to the British Isles and north-western Europe, and ranges in agefrom Early Tournaisian to Visean.


Order PENTREMITIDA Matsumoto, 1929 (emend. Waters and Horowitz, 1993)Family HYPEROBLASTIDAE Fay, 1964

Genus PETALOBLASTUS Fay, 1961

Petaloblastus? ovalis (Goldfuss, 1829)

*1829 Pentremites ovalis Goldfuss, p. 161, pl. 50, ®g. 1a±c.v. 1898 Pentremitidea phillipsii Whidborne, vol. 3, p. 212, pl. 29, ®gs 5±6.v 1930 Pentremites ovalis Goldfuss; Schmidt, pp. 68±69, ®g. 13a±d.

1943 Pentremites ovalis Goldfuss; Bassler and Moodey, p. 242.v 1962 Petaloblastus ovalis (Goldfuss); Fay, p. 20, pl. 2, ®gs 1±3.

Remarks. Whidborne based his description on three specimens, none of which was complete enough in thecritical oral area for positive generic identi®cation. Schmidt (1930) placed Whidborne's species insynonymy with Pentremites ovalis as a part of his description of the Etroeungtian echinoderm faunas fromGermany. He also described a second species, Pentremites boletus, from the Etroeungt beds. Fay (1961)placed both of these species in a new genus, Petaloblastus, and did not comment on Schmidt's decision tosynonymize Whidborne's species from the type Devonian. Our observations of the specimens availableare consistent with Schmidt's placement of P. phillipsii; however, the preservation is too poor to enable usto place the specimen with absolute certainty in the taxon Petaloblastus ovalis. Therefore, in the absenceof specimens that are better preserved, we questionably accept Schmidt's assignment.

One specimen of this species, collected from Wrafton Lane in Barnstaple, is present in the Whidbornecollection (SM H219). Two additional specimens (NDM 338, 339) were collected from Ashford andBradiford.

Order FISSICULATA Jaekel, 1918Family PHAENOSCHISMATIDAE Etheridge and Carpenter, 1886

Genus and species uncertain

Plate 1, ®gures 1±2

Remarks. NHM E30486 is similar in shape to Whidborne's holotype of Codaster conicus, but clearly is nota codasterid. The specimen is a poorly preserved internal cast collected from Croyde Bay that does


E X P L A N A T I O N O F P L A T E 1

Figs 1±2. Internal cast of a phaenoschismatid, genus and species uncertain, collected from Croyde Bay (NHME30486); ´ 8. 1, lateral view. 2, oral view showing cast of hydrospire in lower left; slight lateral crushing ofspecimen is visible from this view.

Fig. 3. Bridgerocrinus? sp., NHM E21847, Hamling Collection, lateral view of crushed calyx, proximal part of stem,and partially preserved arms; ´ 3.

Fig. 4. Mesoblastus cf. M. crenulatus (Roemer, 1851), UB 18343 from Fremington (lowermost Carboniferous); lateralview of theca, two sets of pinnules, and partial stem; ´ 2.

Fig. 5. Bridgerocrinus stadiodactylus (Whidborne, 1896), SM H192; lateral view of partial cup (C-D interray) andproximal arms; ´ 2.

Fig. 6. Eumorphocrinus porteri (Whidborne, 1896), BGS 103584 from Braunton; apical view of partial tegmen withunattached, but articulated arms; ´ 1.

Fig. 7. Lateral view of isolated Glossocrinus? sp., anal sac, BGS 7155; ´ 0´9.Fig. 8. Pachylocrinus? plumifer (Whidborne, 1898), specimen SM H187; lateral view of crown and proximal

stem; ´ 1´9.

P L A T E 1

LANE et al., blastoids, crinoids

preserve some of the structure of the oral area. Although it cannot be assigned to a genus, we believe it isreferable to the Phaenoschismatidae. The phaenoschismatids have a long geologic record and are wellknown from Middle Devonian and Lower Carboniferous faunas.

Order, family, and genus uncertain

v*? 1898 Codonaster conicus Whidborne, p. 214, pl. 29, ®g. 4±4a.1899 Codaster conicus (Whidborne); Bather, p. 4.1943 Codaster conicus (Whidborne); Bassler and Moodey, p. 215.

v 1972 Codaster conicus (Whidborne); Breimer and Macurda, pp. 38, 48.

Remarks. Breimer and Macurda (1972) declared Codaster conicus (Whidborne) a nomen dubium becauseit was based on unrecognizable fragments. None of the currently recognized species of codasteridsoccurred before Visean (Early Carboniferous) time, although we (Waters et al. in review) have described acodasterid from the Famennian of north-western China.

Class CRINOIDEA Miller, 1821Subclass FLEXIBILIA Zittel, 1895

Order TAXOCRINIDA Springer, 1913Family TAXOCRINIDAE Angelin, 1878

Genus TAXOCRINUS Phillips, in Morris, 1843

Taxocrinus macrodactylus (Phillips), 1841

Plate 2, ®gure 1

* 1841 Cyathocrinus? macrodactylus Phillips, p. 29, pl. 15, ®g. 41.1841 Isocrinus monodactylus Phillips, p. 30.1843 Cladocrinites macrodactylus (Phillips); Austin and Austin, p. 198.1843 Taxocrinus macrodactylus (Phillips); Phillips, in Morris, p. 90.1848 Cladocrinites monodactylus (Phillips); Bronn, p. 304.1848 Cyathocrinus? monodactylus (Phillips); Bronn, p. 1217.1848 Cyathocrinus? macrodactylus (Phillips); Bronn, p. 1217.1848 Isocrinus macrodactylus (Phillips); Bronn, p. 1217.1848 Taxocrinus macrodactylus (Phillips); Bronn, p. 1217.1850 Cyathocrinus? monodactylus (Phillips); d'Orbigny, p. 103.1851 Cyathocrinus? macrodactylus Phillips; Roemer, p. 237.1875 Taxocrinus macrodactylus (Phillips); Baily, pl. 30, ®g. 3a±b.

v 1898 Taxocrinus macrodactylus (Phillips); Whidborne, p. 215, pl. 33, ®gs 2±4.



Fig. 1. Taxocrinus macrodactylus (Phillips, 1841) from the Lilycot quarry, Goodleigh; crown, proxistele, anddististele; lateral view; NHM E21867; ´ 1´5.

Figs 2±3. Oral views of Order, family, genus, and species indet. A from the Marwood beds. 2, counterpart showingpartial tegmen and oral view of branching arms; BGS 7143; ´ 2. 3, plate showing tegmen and aboral view of arms;BGS 7142; ´ 1´5.

Figs 4, 6. Eumorphocrinus porteri (Whidborne, 1896); ´ 1´5. 4, lectotype, from Barnstaple; calyx, partial arms, andproximal stem; ?posterior view; SM 180A. 6, BGS 7140 from Braunton; distal part of dorsal cup with armspreserved through tertibrachs; lateral view.

Fig. 5. Taxocrinus stultus Whidborne, 1896 from Pilton, William Porter collection; lateral view of crown throughproximal tertibrachs and proximal stem; SM H177B; ´ 3´1.

P L A T E 2

LANE et al., crinoids

1920 Taxocrinus macrodactylus (Phillips); Springer, p. 381, pl. 51, ®gs 1±4.1929 Taxocrinus macrodactylus (Phillips); H. Schmidt, p. 85, pl. 21, ®g. 3.

v 1930 Taxocrinus macrodactylus (Phillips); W. E. Schmidt, p. 62, pl. 1, ®gs 9±10.1938 Cyathocrinus? monodactylus (Phillips); Bassler, p. 181.1943 Taxocrinus macrodactylus (Phillips); Bassler and Moodey, p. 703.1943 Cyathocrinus? macrodactylus Phillips; Bassler and Moodey, p. 703.1978 Taxocrinus macrodactylus (Phillips); Moore et al., p. T768, ®gs 511-1, 512-5c.1993 Taxocrinus macrodactylus (Phillips); Webster, p. 300.

Remarks. This species, the type for the genus, has been described several times, most completely bySpringer (1920) and Schmidt (1930). We have little to add to these descriptions except to note thatPhilip's single surviving syntype shows three primibrachials in one ray and seemingly only twoprimibrachials in the other preserved ray. This variation seems to be indicated in Whidborne's (1898,pl. 33, ®g. 1) illustration of the syntype as well as in Moore et al. (1978, ®g. 512-5c). The principaldifference between Taxocrinus and Eutaxocrinus is the presence of two or three primibrachials,respectively. Whidborne (1898, p. 216) noted that he had two specimens of Taxocrinus macrodactylusfrom Fremington in the Porter Collection. We were unable to locate these. However, we did locate aspecimen (in the Hamling Collection) from Fremington that was unavailable to Whidborne at the timeof his monograph.

Material. In addition to Phillips' syntype, Whidborne had three other specimens available for study: BGS 7149(103603), 7149 (103602), and 7150. Phillips' syntype is BGS 7148, labelled Pilton for locality. The other specimensare labelled Marwood. In addition to these specimens that were available to Whidborne, two others were collected byJ. G. Hamling in the early part of the 20th Century. One is TNHS 69/12, from Fremington. This specimen is thusyounger than the other fossils and indicates that the species may have persisted from the Famennian into theTournaisian. The other Hamling specimen, NHM E21867, is from the Lily Cot quarry at Goodleigh, near Barnstaple. Itwas presented to the museum by Hamling in 1916.

Measurements. NHM E21867: preserved HCr, 35 mm.

Taxocrinus stultus Whidborne, 1896

Plate 2, ®gure 5

v* 1896 Taxocrinus stultus Whidborne, p. 377.v* 1898 Taxocrinus stultus Whidborne; Whidborne, p. 216, pl. 34, ®gs 1±3.

. 1920 Taxocrinus stultus Whidborne; Springer, p. 381, pl. 51, ®g. 5a±c.1943 Taxocrinus stultus Whidborne; Bassler and Moodey, p. 705.

Diagnosis. Crown small, without interradial plates. Stem does not widen just below calyx.

Remarks. This species differs from T. macrodactylus, with which it occurs, in being smaller, more slender,and with narrow arms. The proxistele remains the same width throughout, unlike T. macrodactylus, inwhich the proxistele expands in width proximally. The mesistele is distinctly heteromorphic.

Material. We here designate SM H177A, illustrated by Whidborne (1898, pl. 34, ®g. 1), as the lectotype. Thecounterpart of this mould is SM H177B (Whidborne, 1898, pl. 34, ®g. 2). These specimens and SM H178(Whidborne, 1898, pl. 34, ®g. 3) were all collected by William Porter from Pilton, who gave them toWhidborne. An additional specimen collected by Hall, NDM 403, is without locality information. This may ormay not be the same specimen noted by Whidborne as being in that collection and as having been collected atRoborough.

Measurements. SM H177a: WC, 3´9 mm; HC, 3´3 mm; HCr, 19 mm.


Subclass CAMERATA Wachsmuth and Springer, 1885

Order, family, genus, and species uncertain A


v. 1898 Rhodocrinus?, ?sp. Whidborne, pp. 217±218, pl. 31, ®gs 3±3d.1943 Rhodocrinus? sp. Whidborne; Schmidt, p. 87.

Remarks. This taxon was erected by Whidborne for a single specimen, part and counterpart, of a distortedcup and some attached arms. Whidborne (1898, p. 218) noted, `To what genus this fossil belongs is mostuncertain. There seem some slight grounds for supposing that it might belong to Rhodocrinus, andtherefore with much hesitation I have placed it tentatively there.' The cup has a rusty coating inside themould so that plates cannot be discerned. It is possible that the specimen may be an actinocrinoid, althoughdifferent from the other described actinocrinoids in the fauna. Based on the poor preservation of this singlespecimen, we have chosen to place it in open nomenclature as an indeterminate camerate crinoid.

Material. The part and counterpart are BGS 7142 and 7143. The label for the specimen states that it is from theMarwood beds of North Devon, but a speci®c locality is not known.

Measurements. BGS 7143: crushed WDC, 19´9 mm; crushed HDC, 11´1 mm.

Order, family, genus, and species uncertain B

Plate 3, ®gure 6

v. 1898 Megistocrinus?, sp. Whidborne, p. 218, pl. 37, ®g. 5.1930 Megistocrinus? sp. Whidborne; Schmidt, p. 87.

Remarks. The single specimen, SM H4047, from Barnstaple, is wrapped around and crushed against alarge segment of crinoid stem. As far as we can determine, Whidborne's sketch of the plate arrangement isaccurate, although there is insuf®cient evidence to assign the specimen to Megistocrinus as opposed tosome other small, multiplated camerate with a wide base and a bowl shape. Consequently, we have chosento place this single specimen in open nomenclature as an indeterminate camerate crinoid.

Order, family, genus, and species uncertain C

Plate 3, ®gure 3

v 1898 Platycrinus? anguliferus Whidborne, p. 223, pl. 37, ®gs 9±11.v. 1930 cf. Platycrinus anguliferus Whidborne; Schmidt, pp. 8±9, pl. 3, ®g. 16.

vp. 1943 Platycrinites anguliferous (Whidborne); Bassler and Moodey, p. 615.

Description. Three radial plates. SM H202 with three longitudinal diverging ridges trending from centre of distal edge,just below facet, to proximal edge; with ®ve or six horizontal, lateral, parallel ridges on each side of longitudinal ridgebundle; horizontal ridges do not touch longitudinal diverging ridges. SM H203 also with three longitudinal ridges, butonly two left-side ridges extend to edge of facet; two very short, vertical longitudinal ridges outside the central threeridges, each of which terminates in a lateral ridge; right-side longitudinal ridge truncated by third lateral ridge downfrom facet; 5±6 lateral ridges on each side of longitudinal ridge bundle. SM H204 cast is less complete than thedrawing shown by Whidborne, but with seven longitudinal ridges that are shorter towards bottom of plate.

Remarks. We have serious doubts that these radial plates with ridges, illustrated upside-down byWhidborne, belong to Platycrinites. We are not aware of any platycrinoid with raised ridges on theradial plates that form patterns like those shown by these specimens. This plate sculpture more closelyresembles that found on the radials of some dichocrinoids, hexacrinoids, and actinocrinoids. However, in


the absence of any complete calyces, we can do no better than to regard these specimens as indeterminatecamerate calyx plates.

Material. Three isolated radial plates from the Porter collection: SM H202 (Whidborne 1898, pl. 37, ®g. 9) fromRoborough; SM H203 (Whidborne 1898, pl. 37, ®g. 10) from Pilton; SM H204 (Whidborne 1898, pl. 37, ®g. 11) fromRoborough.

Measurements. SM H202: length 8´2 mm; width 7´8 mm. SM H203: length 9´6 mm, width 7´4 mm.

Order MONOBATHRIDA Moore and Laudon, 1943Family ACTINOCRINITIDAE Austin and Austin, 1842

Subfamily EUMORPHOCRININAE Ubaghs, 1978

Genus EUMORPHOCRINUS Wright, 1955

Eumorphocrinus porteri (Whidborne, 1896)

Plate 1, ®gure 6; Plate 2, ®gures 4, 6; Plate 4, ®gure 1

v* 1896 Actinocrinus porteri Whidborne, p. 377.v. 1898 Actinocrinus porteri Whidborne; Whidborne, pp. 220±221, pl. 5, ®g. 21; pl. 30, ®g. 8; pl. 31, ®gs 1±1a,

2, 5; pl. 32, ®g. 1.1943 Actinocrinites porteri (Whidborne); Bassler and Moodey, p. 273.

Diagnosis. Two ®xed secundibrachials in each half-ray; ®xed brachials grouped, but not extended intoprotuberant brachial lobes; plates nodose; anal tube may or may not be present.

Description. Cup with rounded, convex sides; basals low, only slightly wider than proximal columnal. Radials andprimibrachials large. First primibrachial hexagonal, second axillary, supporting two secundibrachials in each half-ray,resulting presumably in 20 arms, although no one specimen reveals all ®ve rays. Interray areas slightly depressed;interradials large, numerous, in broad contact with small tegmen plates. Tegmen conical, composed of numerous smallplates, with central anal opening, but presence of a tube is uncertain. Cup plates with ridges and nodes. Arms biserial,branching once high above the cup. Stem round, heteromorphic, with nudinodals and two orders of 5±7 internodals.

Remarks. Although the arms are grouped, the lack of well-developed protuberances on the calyx clearlyplaces these specimens in the Eumorphocrininae. Among other genera in the Eumorphocrininae, thisspecies differs from species of the genera Cytidocrinus and Manillocrinus in the structure of the arms. Itdiffers from Maligneocrinus in having two ®xed secundibrachs, rather than one, in each half-ray. It alsoresembles Dialutocrinus, but again differs in arm structure and number of ®xed secundibrachs.



Figs 1, 5. Platycrinites sp., E21854, Hamling collection, Downsend, showing several lengths of stem with loose radialplates and some attached arms. 1, magni®ed view of right end of 5 showing two radial plates, with two primibrachs;´ 2. 5, ´ 1.

Fig. 2. Glossocrinus? sp., SM H184 from Barnstaple, lateral view of radials, primibrachs, and secundibrachs; ´ 1´2.Fig. 3. Camerate order, family, genus, and species indet. C, SM H202, radial plate with ornamentation; ´ 1´7.Fig. 4. Glossocrinus? salebrosus (Whidborne, 1898), SM H200, lateral view of cup, proximal stem, and partially

disarticulated arms and anal sac; ´ 1´6.Fig. 6. Camerate order, family, genus, and species indet. B, SM H4047 from Croyde Bay, Barnstaple; specimen,

indicated by arrow, is wrapped around and crushed against a large segment of crinoid stem in centre of ®gure; ´ 2.Fig. 7. Abactinocrinites? sp., NDM 395, Townsend Hall Collection, Fremington; lateral view of upper part of cup (with

tegmen missing) and arms; ´ 1.

P L A T E 3


Eumorphocrinus is known from the Tournaisian of Scotland and Ireland, and questionably from theRedwall Limestone (Osagean) of Arizona.

Material. A lectotype is selected here, SM H180A, from Barnstaple, illustrated by Whidborne (1898, pl. 30, ®g. 8).Other specimens include BGS 7138 and 7139, part and counterpart from Braunton, BGS 7138 (Whidborne 1898, pl. 31,®g. 1), and BGS 7139 (Whidborne 1898, pl. 31, ®g. 2). BGS 7140 (Whidborne 1898, pl. 31, ®g. 5) and BGS 7141(Whidborne 1898, pl. 32, ®g. 1) are both from Braunton. In addition to these specimens available to Whidborne, theBritish Geological Survey holds two specimens acquired more recently: BGS 103591 from Barnstaple, in the WyattEdgell collection, and BGS 103584.

Measurements. SM H180a: WDC, 16´2 mm; HDC, 14´5 mm; BGS 7140: crushed WDC, 29´0 mm; BGS 7138: HDC,8´9 mm; WDC, 15´1 mm.

Subfamily ACTINOCRINITINAE Austin and Austin, 1842

Genus ACTINOCRINITES Miller, 1821

Actinocrinites batheri (Whidborne, 1896)


v* 1896 Actinocrinus batheri Whidborne, p. 377.v. 1898 Actinocrinus? batheri Whidborne; Whidborne, p. 222, pl. 32, ®gs 2±3b.

1930 Actinocrinus batheri Whidborne; Schmidt, p. 87.1993 Actinocrinites? batheri (Whidborne); Bassler and Moodey, p. 267.

Diagnosis. Small actinocrinoid with 20 arm openings, protuberant, grouped ®xed brachials, numeroussmall tegminal plates, and irregularly hexagonal ®rst primibrachials.

Description. Calyx small, cup low, cone-shaped. Basals small, low. Radials and other cup plates with radiating ridges.First primibrachial hexagonal in at least one ray, other rays too poorly preserved to be certain of plate shape. First orsecond ®xed secundibrachs axillary, resulting in four arm openings to a ray, or 20 free arms. Interradials few, but incontact with tegmen. Tegmen low, composed of numerous, small, nodose plates. Anal tube not preserved. Arms ofeach ray grouped and protuberant. Posterior side of cup damaged (shown in dark shading by Whidborne). Armsbiserial, unbranched. Proximal columnal round, with wide areola, narrow crenularium and epifacet, lumen?pentagonal.

Remarks. This is one of the more interesting specimens in the various collections. It bears closeresemblance to a Chinese Famennian species that we (Lane et al. 1997) named Uperocrinus zhaoe, butthat we now believe is an actinocrinoid. This new assignment is based on reconsideration of the nature ofthe ®rst primibrachial. In the Chinese specimens this plate is either ®ve- or six-sided, and if six-sided thetwo lateral sutures are quite short, commonly of unequal length, which gives a quasi-quadrangular aspectto the plate. It was for that reason, and because there are three rather than two secundanals, that anassignment to the batocrinoids was made. Now, we believe that, strictly speaking, the plates of the Chinesespecimen are hexagonal, and that it should be assigned to Actinocrinites. Both A. batheri and A. zhaoe areclosely similar to A. depressus Wright from the Tournaisian of Scotland, which also shows variation in thenumber of sutures and suture length in its primibrachials.

Among other genera that we considered for assignment of A. batheri, we excluded Blairocrinus becauseit has only a few large tegmen plates; Abactinocrinites because it has ten rather than 20 arm openings;Nunnacrinus because it has less strongly grouped arms; and Eumorphocrinus because it lacks protuberantarms.

Material. We here select BGS 7146 (Whidborne 1898, pl. 32, ®g. 3) as the lectotype. The counterpart is BGS 7145(Whidborne 1898, pl. 32, ®g. 2).

Measurements. BGS 7146: WDC, 13´4 mm; HDC, 5´5 mm; WBC, 3´4 mm.


Genus ABACTINOCRINITES Laudon and Severson, 1953

Abactinocrinites? sp.

Plate 3, ®gure 7

Description. Upper part of a large dorsal cup with attached biserial arms. Tegmen low and broad, with numerousplates, some of which bear small nodes. Radial plates not fully preserved, but radial facets occupy about one-half platewidth. First primibrachial axillary, bearing at least two large uniserial brachials on each side. At least two ®xedprimibrachials, second primibrachial axillary. Protuberant brachial lobes separated by ®xed interprimibrachialsbroadly connected with tegmen. Base of cup and stem not preserved.

Remarks. Numerous plates in the calyx, coupled with distinctive grouping of the arms and well-developedcalyx protuberances place this specimen within the subfamily Actinocrinitinae. The entire subfamily ispredominantly Early Carboniferous in age. Although the specimen resembles Actinocrinites, we chose toplace it in the genus Abactinocrinites with question because we cannot see the bottom of the ®xedbrachials, which precludes us from making a de®nitive generic assignment, and because the specimen hasten arm openings per ray, as opposed to 20 in Actinocrinites. This is probably an Early Carboniferousspecimen because it is from Fremington, which is south of the Taw estuary.

Material. A single specimen, NDM 395, in the Townsend Hall collection from Fremington.

Measurements: NDM 395: width of calyx 25 mm; width of protuberant brachial lobe 11 mm; preserved arm length53 mm; proximal arm width 4´8 mm; height of tegmen 6´3 mm.

Family PLATYCRINITIDAE Austin and Austin, 1842

Genus PLATYCRINITES Miller, 1821

Platycrinites sp.

Plate 3, ®gures 1, 5

Description. Four pleuricolumnals; columnal twisting continuous and gradual throughout stem length; all columnalsnodose for entire length. Two radials, both nodose, height and width subequal. Proximal arms preserved on bothradials; one primibrachial, axillary; two secundibrachials, the second axillary. Edge of nodose basal circlet possiblypresent below the two radials.

Remarks. One specimen collected by J. G. Hamling from Pilton Beds at Downsend, Saunton, and labelledPlatycrinites sp., cf. P. guttifer Schmidt, 1930, is an undoubted specimen of Platycrinites stem. Inaddition, two radial plates with arm attachments are present at one end of the stems. Although Whidbornereported Platycrinites from Barnstaple, we have been unable to verify the existence of this genus based onany of the specimens available to him at the time. However, this specimen from the Hamling Collection,which was not available to Whidborne, con®rms the presence of Platycrinites in the Famennian in thisarea.

Material. NHM E21854, stem cluster with two radial plates, proximal arms, and possible basal circlet preserved.

Measurements: Maximum stem length 100 mm, with three other shorter segments; stem width 4´3 mm; radial width7´2 mm, radial height 6´7 mm; facet width at arm base 3´2 mm.

Platycrinites? sp.

v*? 1898 Platycrinus? anguliferus Whidborne, p. 223, pl. 37, ®gs 8, 12.non 1898 Platycrinus? anguliferus Whidborne, p. 223, pl. 37, ®gs 9±11.


v. non 1930 cf. Platycrinus anguliferus Whidborne; Schmidt, pp. 8±9, pl. 3, ®g. 16.vp. 1943 Platycrinites anguliferous (Whidborne); Bassler and Moodey, p. 615.

Remarks. We have been unable to ascertain whether Whidborne's (1898, pl. 37, ®g. 8) principal specimenof this genus is a Platycrinites or not. Given the poorly preserved specimens illustrated by Whidborne, andthe likelihood that several of them do not belong to the genus Platycrinites, we consider his species,P. anguliferus, to be a nomen dubium.

We note that Schmidt (1930) described four species of Platycrinites from the Etroeungt beds ofGermany, two of them indeterminate and based on isolated plates.

Material. NDM 414, Barnstaple, Saunton Hotel, Coomara Swamy collection (Whidborne 1898, pl. 37, ®g. 8); NDM399, a set of biserial arms labelled Platycrinites, from Braunton, Townsend Hall collection; and NDM 394,indeterminate specimen labelled Platycrinites, Top Orchard, Townsend Hall collection.

Family HEXACRINITIDAE Wachsmuth and Springer, 1885

Genus ADELOCRINUS Phillips, 1841v* 1841 Adelocrinus Phillips, p. 30.

. 1898 Perischodomus Tornquist, p. 103.v 1898 Adelocrinus Phillips; Whidborne, 1898, pp. 223±224.. 1903 Adelocrinus Phillips; Drevermann, p. 518.

v 1930 Arthroacantha Williams; Schmidt, p. 19.p 1943 Arthroacantha Williams; Bassler and Moodey, p. 282.

Diagnosis. A hexacrinoid with tubercles on basals and radials that support small, slender spines, a singlelarge axillary primibrachial, and 20 arms.

Remarks. Numerous specimens of this crinoid are in the Pilton collections. Several of these have the smallspines preserved, laid over from the numerous small tubercles to which they were attached. AlthoughPhillips had only a single basal circlet when he described the genus, there is little doubt that the specimensreferred to Adelocrinus here are conspeci®c with his specimen. Subsequently, Phillips' distinctivebasal circlet came to be misidenti®ed as a columnal (see Jeffords 1978, p. T932). Whidborne (1898,pp. 223±224) clearly recognized that the genus was allied to Hexacrinites or to Arthroacantha, as didBather (1900, p. 201). Discovery of the spines still in association with the basals and radials proves thatArthroacantha and Adelocrinus are closely related. The two genera differ in that Arthroacantha has twoprimibrachials, rather than one, has arms that branch repeatedly, and has a different arrangement oftegminal plates (Williams 1883; Hinde 1885, 1886). It is likely that some of the species assigned toArthroacantha over the years should be placed in Adelocrinus. However, most of these specimens haveneither arms nor tegmen preserved; therefore, it is impossible to differentiate the two genera based only ondorsal cup characters. As currently known, Arthroacantha occurs in Devonian rocks below the Famennian,whereas Adelocrinus occurs in the Famennian of Britain and Germany.

Adelocrinus hystrix Phillips, 1841

Plate 4, ®gures 4±6, 9

v* 1841 Adelocrinus hystrix Phillips, p. 30, pl. 16, ®g. 42a±b.1857 Adelocrinus hystrix Phillips; Pictet, p. 333.

v. 1898 Adelocrinus hystrix Phillips; Whidborne, p. 224, pl. 30, ®gs 1±7a.1900 Adelocrinus hystrix Phillips; Bather, in Lancaster, p. 202.1900 Perischodomus magnus Tornquist; Bather, in Lancaster, p. 203.1903 Adelocrinus hystrix Phillips; Drevermann, p. 518, pl. 14, ®gs 23±23a.

v. 1930 Arthroacantha hystrix (Phillips); Schmidt, p. 191938 Adelocrinus hystrix Phillips; Bassler, p. 34.


1943 Adelocrinus hystrix Phillips; Bassler and Moodey, p. 282.v. 1978 Subclass and order uncertain, Moore and Teichert, p. T932.

Diagnosis. Basal circlet low, cone-shaped; cup spines short and slender.

Description. Basals three, equal, fused, sutures obscure, six-sided in outline, with small, raised tubercles; radials nearlyquadrangular, bearing 25±35 evenly scattered tubercles. Anal plate not observed. Radial facets narrow. One largetegminal plate bearing a single central tubercle was observed. First primibrachial axillary, bearing two secundibra-chials on each side, the second axillary. Lower brachials medially constricted. Arms long, slender, narrow and deep,gradually becoming biserial above secundaxils. Pinnules long, slender, directed obliquely distally; pinnules long andslender. Proxistele round, heteromorphic, with alternating higher and wider nodals and smaller priminternodals;remainder of stem round, heteromorphic with nudinodals, number of internodals uncertain; lumen round, articularfacet with wide crenularium.

Remarks. This species has also been reported from the Etroeungt beds of Germany by Schmidt (1930). Onespecimen (NDM 383) in the Townsend Hall Collection from Barnstaple is labelled Hexacrinius?, but morethan likely is a specimen of Adelocrinus hystrix arms.

Material. Phillips' holotype is BGS 7157, from Brushford, Somerset, which was illustrated by Whidborne (1898,pl. 30, ®g. 4). The specimen has been somewhat distorted laterally. BGS 7156 from Braughton is a calyx with stemattached and arm fragments. SM H197, William Porter collection, from Pilton (Whidborne 1898, pl. 30, ®g. 1); NDM340 from Top Orchard (Whidborne 1898, pl. 30, ®g. 5); NDM 341 (Whidborne 1898, pl. 30, ®g. 6) from Top Orchard;NDM 342 (Whidborne 1898, pl. 30, ®g. 3) from Bradiford; NHM E25495, in the A. Champernowne collectionobtained in 1909, is a nearly complete crown from a quarry near Dulverton Station-on-the-North, Somerset; UB 18341is a well-preserved basal circlet from Pilton Beds A, Wocklumeria Zone, north part of Croyde Bay, collected byR. Goldring in 1956.

Measurements. NHM E25495, HCr, 59 mm; HR, 8´5 mm; WR, 6´5 mm. BGS 7156: HDC, 14´1 mm; WDC, 11´6 mm.SM H197: WR, 10´9 mm; HR, 10´0 mm. UB 18341: HBC, 6´4 mm; WBC, 13´1 mm.

Subclass CLADIDA Moore and Laudon, 1943

Remarks. With respect to these crinoids, Whidborne (1898, p. 225) wrote, `In the maze of this orderit almost requires a necromancer to bring down a species to its rightful place amid the kaleidoscopicgenera that appear, change, and vanish.' How much more true that statement is today with the manygenera of the paraphyletic inadunates that have been named since Whidborne's time! The majority ofcladid specimens available for this study can be placed in the Superfamily Scytalocrinacea. Virtuallyall specimens have infrabasals visible in side view with a cone-shaped or high, bowl-shaped cup. Inmost cases the arms branch once or twice, ®rst on either the ®rst or second primibrachial. All butone species have radial facets that occupy the full width of the radials. All specimens seen arepinnulate.

We have chosen to follow the recommendation of Simms and Sevastopulo (1993) that the order Cladidabe elevated to a subclass. Ausich (1997) also followed this usage, agreeing that the group is paraphyletic.However, we have chosen not to follow the example of Kammer and Ausich (1996), in which no familieswere used.

Family POTERIOCRINITIDAE Austin and Austin, 1842

Remarks. Whidborne (1898) placed two species in this family, both in the genus Poteriocrinites. Webelieve neither belongs in this family because both have radial facets that are plenary rather thanangustary. In addition, both species have the ®rst primibrachial axillary rather than the numerousprimibrachials of Poteriocrinites. However, another Whidborne species, Scaphiocrinus transcisus,which does have angustary radial facets and strongly ridged cup plates, is here transferred toPoteriocrinites.


Genus POTERIOCRINITES Miller, 1821

Poteriocrinites transcisus (Whidborne, 1898)

Plate 4, ®gure 7

v* 1898 Scaphiocrinus transcisus Whidborne, p. 230, pl. 38, ®g. 3.1930 Hydriocrinus transcisus (Whidborne); Schmidt, pp. 57, 87.

Diagnosis. Cup high, with prominent radiating ridges on cup plates; anal sac stout, with large platesbearing radiating ridges.

Description. Dorsal cup large, high, conical. Cup plates bearing prominent straight ridges that radiate from the centresof plates. Infrabasals high and large, clearly making up about one-quarter of cup height. Basals large; radials short,with facets that are peneplenary to angustary. Anal plates in cup three, large, with radiating ridges, bearing large analplates above that also have ridges. Arm branching uncertain. Primibrachial 2 may or may not be axillary. If so,secundibrachial three is axillary; if not, primibrachial 5 is axillary. Arms not preserved very far above the one certainaxillary plate. Proximal columnals pentagonal, remainder of stem not known.

Remarks. This is a very distinctive species quite unlike others in the fauna. The high, conical cup withprominent infrabasals and radial facets distinctly narrower than the radials are important characters. Thespecies closely resembles species of Poteriocrinites and Springericrinus. It resembles the latter genus inthe strong ridges on cup and anal sac plates, and in the stout, prominent anal sac. It does not exhibit thesecondary ridges and troughs typical of that genus. Unlike this specimen, species of Hydriocrinus haveonly one primibrachial. Whether primibrachial 2 or 5 is axillary does not prohibit assignment toPoteriocrinites, in which primibrachial 1±14 may be axillary. The species can be compared withPoteriocrinites plicatus Austin and Austin, 1842 from the British Tournaisian.

Material. Although Whidborne cited two specimens of this species, we have only found the illustrated one (1898,pl. 38, ®g. 3), which is SM H201. We designate this specimen the lectotype.

Measurements. SM H201: HDC, 19´9 mm.

Family GLOSSOCRINIDAE Goldring, 1923

Remarks. Webster and Ha¯ey (in Webster et al. 1999) revised the family Rhenocrinidae Jaekel, 1918, andremoved the family Glossocrinidae Goldring, 1923 from synonymy with it. According to Webster and



Fig. 1. Eumorphocrinus porteri (Whidborne, 1896); BGS 7141 from Braunton; ¯attened crown; distal part of cup andproximal arms; ´ 1.

Figs 2±3. Actinocrinites batheri (Whidborne, 1896) from Braunton; ´ 2. 2, tegmen and proximal parts of arms; BGS7146. 3, aboral view of cup and proximal arms; lectotype, BGS 7145.

Figs 4±6, 9. Adelocrinus hystrix Phillips, 1841. 4, BGS 7156 from Braunton; lateral view of dorsal cup with axillaryprimibrach attached and proximal portion of stem; ´ 1´5. 5, UB 18341 from the north part of Croyde Bay; basalcirclet with bosses for movable spines; ´ 1´5. 6, SM H197 from Pilton; radial plate with movable spines attached;´ 1´8. 9, NHM E25495, Champernown collection, from quarry near Dulverton, Station-on-the-North, Somerset;compressed cup with spine bosses and arms; ´ 1´4.

Fig. 7. Poteriocrinites transcisus (Whidborne, 1898); lectotype, from Poleshill, William Porter collection; SM H201;anal sac and portion of one arm on right of photograph with some displaced cup plates; ´ 1´2.

Fig. 8. Aphelecrinus? sp., BGS 7152 from Braunton; lateral view of crown and stem; ´ 1´5.

P L A T E 4


Ha¯ey (in Webster et al. 1999, p. 468), the family Glossocrinidae contains the genera CatactocrinusGoldring, 1923, Charientocrinus Goldring, 1923, Corematocrinus Goldring, 1923, GlossocrinusGoldring, 1923, Liparocrinus Goldring, 1923, and Propoteriocrinus Schmidt, 1934, whereas the familyRhenocrinidae contains the genera Cydrocrinus Kirk, 1940, Hallocrinus Goldring, 1923, MaragnicrinusWhit®eld, 1905 and Rhenocrinus Jaekel, 1906. Finally, Webster and Ha¯ey (in Webster et al. 1999)reassigned the genus Araeocrinus Strimple and Watkins, 1969 from the family Rhenocrinidae to thefamily Corythocrinidae. Generally, the rhenocrinoids have higher cups and narrower interradial notchesthan do the glossocrinoids. In addition, the glossocrinoids have pentagonal stems (vs. circular stems inrhenocrinoids) and radial facets that are strongly convex on their outer sides (Webster and Ha¯ey, inWebster et al. 1999).

Genus GLOSSOCRINUS Goldring, 1923

Glossocrinus whidbornei sp. nov.


Derivation of name. Although somewhat belatedly, the species name is for the Rev. G. F. Whidborne, who ®rstmonographed the Pilton crinoid fauna.

Diagnosis. A glossocrinoid with smooth cup plates, cuneate brachials, and second primibrachial axillary.

Description. Crown tall and slender; dorsal cup high, narrow, cone-shaped, infrabasals making up about one-quarter ofcup height; plates smooth. Radials wider than high, with peneplenary facets and convex outer facet margins; interradialnotches very narrow; a gape is present between outer edge of facet and the facet for the ®rst primibrachial. Posteriorside of cup not observed. Anal sac tall, narrow, composed of numerous narrowly rectangular plates with slender ridgesconnecting all plates. Slits between plates present. Second primibrachial axillary, supporting cuneate, pinnulatebrachials. Higher branching not observed, but arms not well-preserved above secundibrachial 8. Proximal part of stempentagonally lobate.

Remarks. The distinctive radial facets, pentagonally lobate stem, and anal sac distinguish these twospecimens from all other crinoids in the fauna, and ally them with the Family Glossocrinidae. Prior to thisreport, Glossocrinus was known only from the Givetian±Frasnian of the central and eastern United States.Thus, the Pilton species is the youngest known for the genus and the only report of the genus outside NorthAmerica.

Material. NHM E21838, Top Orchard, Pilton, is here designated the holotype; NHM E21837, Top Orchard, Pilton, isthe paratype.

Measurements. NHM E21837: H anal sac, 32 mm; W anal sac, 5´4 mm; NHM E21838: HDC 6´8 mm; WDC, 6´1 mm.

Glossocrinus? salebrosus (Whidborne, 1898)

Plate 3, ®gure 4

v. 1898 Poteriocrinus sp. Whidborne, p. 228, pl. 36, ®gs 9±9a.? 1898 Poteriocrinus sp. Whidborne, p. 228, pl. 35, ®gs 4±5a.

v* 1898 Scaphiocrinus? salebrosus Whidborne, p. 232, pl. 37, ®g. 13.1930 Hydriocrinus salebrosus (Whidborne); Schmidt, p. 87.

Description. Cup about equal in height and width, plates smooth or slightly convex, infrabasals visible in side view,basals large; radials apparently relatively low; radial facets unclear, but apparently peneplenary. Proximal part of analsac composed of large, smooth plates. Brachials large, cuneate, branching ®rst on primibrachial 6. Stem pentagonal.


Remarks. We believe that Whidborne described this species under two different names. One specimen(BGS 7154) he called Poteriocrinus sp., a category in which Whidborne included a number of specimensthat do not appear to agree in their morphology. Whidborne's remarks under discussion of this group donot ®t this specimen, which has a relatively large, high cup, brachials that branch ®rst high above the cup,and a pentagonal stem. The other name used by Whidborne (1898) was Scaphiocrinus? salebrosus, whichwas based on three specimens: two in the Sedgwick Museum and one in the North Devon Museum. Onlyone specimen (SM H200) was illustrated. We describe here the Poteriocrinus sp. specimen (Whidborne1898, pl. 36, ®g. 9) and the sole illustrated specimen of his Scaphiocrinus? salebrosus.

We cannot assign these specimens to the Poteriocrinitidae because they do not have angustary facets andthe stem is pentagonal. They seem to ®t best among the Glossocrinidae (see Webster and Ha¯ey, inWebster et al. 1999). The arms suggest the glossocrinoid Liparocrinus, but the cup is quite different. Thecup resembles that of the rhenocrinoid Maragnicrinus, but the arms branch on primibrachial 3 in thatgenus and the stem is round. Among the possible genera of glossocrinoids, we judge Glossocrinus to be thebest ®t. It differs from Glossocrinus whidbornei sp. nov. primarily in that primibrachial 6 is axillary, asopposed to primibrachial 2 being axillary in G. whidbornei. This is clearly a distinctive species and ifbetter preserved specimens were available the species might belong to a new genus; therefore, we haveassigned the specimens to Glossocrinus, but have questioned the assignment.

A specimen labelled Scaphiocrinus salebrosus in the Townsend Hall collection from Bradiford (NDM343) is actually the basal circlet of a camerate crinoid. We were unable to locate the specimen from Upcottmentioned by Whidborne (1898).

Material. BGS 7154, from Braughton (Whidborne 1898, pl. 36, ®g. 9) and SM H200, from Poleshill, Pilton(Whidborne 1898, pl. 37, ®g. 13).

Measurements. BGS 7154: HDC, 15´3 mm; crushed WDC, 13´5 mm; Wstem, 5´0 mm; WBr, 4´1 mm. SM H200: WDC,6´7 mm; HDC, 6´1 mm; Wstem, 2´8 mm.

Glossocrinus? sp.

Plate 1, ®gure 7; Plate 3, ®gure 2

v. 1898 Poteriocrinus sp. Whidborne, p. 228, pl. 35, ®gs 4±5a.

Description (based on SM H184). Only the radials are preserved of the cup. Posterior interray is poorly preserved, withthree anal plates in primitive position. Arms contain three or four primibrachs; nine or ten secundibrachs; pinnulate;secundibrachs and tertibrachs weakly cuneate. Anal sac is cylindrical, consisting of several longitudinal rows of plates,each of which with an elevated central part. Anal sac plates much wider than high; elevated central parts aligned toform ridges; approximately six longitudinal rows in sac; ridges not as well preserved as other specimen; eachindividual plate with numerous slits; proximal anal sac plates with oblique ridges present only in lower 15 mm. Stemnot preserved.

Remarks. Plate 35, ®gure 5 in Whidborne (1898) is printed backwards. The sac is typical of theglossocrinoids, the arms ®t with Glossocrinus, but we do not have enough cup preserved to make ade®nite generic assignment.

Material. SM H184 from Barnstaple (Whidborne 1898, pl. 35, ®g. 5); BGS 7155 (Whidborne 1898, pl. 35 ®g. 4) fromMarwood beds at Braughton Downs.

Measurements. SM H184: ¯attened sac 11 mm wide; preserved length of sac is 40 mm; preserved length of arms39 mm; individual plates in sac 2 mm high, 5 mm wide; primibrach series 11 mm high, 4´8 mm wide; secundibrachseries 22´8 mm high, approximately 2´9 mm wide. BGS 7155: isolated anal sac preserved length 88 mm, preservedwidth ? mm; individual anal plates approximately 7 mm wide, 2 mm high.


Superfamily SCYTALOCRINACEA Moore and Laudon, 1943

Remarks. The great majority of Famennian cladid crinoids fall into this superfamily. They all haveinfrabasals visible in side view, some to a much greater extent than others. The cup may be straight-sidedand conical or have convex sides and a high bowl shape. The radial facets are, without exception, plenary.The arms branch on either primibrachial 1 or 2 in different species. The arms may remain unbranchedabove the primaxil, resulting in ten arms, or branch once more high above the cup, or branch more thanonce above the primaxil. Various combinations of these features characterize different families and generawithin the superfamily.

Those specimens that have the ®rst primibrachial axillary and that have ten arms are placed here inScytalocrinus. Those specimens that have a high, conical cup, primibrachial 2 axillary, and ten arms areplaced here in Bridgerocrinus. Those specimens that have primibrachial 2 axillary and have arms thatbranch once above the primaxil are placed in Sostronocrinus.

Family SCYTALOCRINIDAE Moore and Laudon, 1943

Genus SCYTALOCRINUS Wachsmuth and Springer, 1880

Scytalocrinus barumensis (Whidborne, 1896)

Plate 5, ®gures 4, 10

v* 1896 Cyathocrinus barumensis Whidborne, p. 377.v* 1898 Poteriocrinus barumensis Whidborne, p. 227, pl. 34, ®g. 6; pl. 35, ®g. 3.

1930 Hydriocrinus barumensis (Whidborne); Schmidt, p. 57.1930 Hydriocrinus? barumensis (Whidborne); Schmidt, p. 87.

Diagnosis. Cup and basals small; anal sac composed of stellate plates.

Description. Cup wider than high, straight-sided, infrabasals low, but visible in side view, basals and radials relativelysmall. Primibrachial 1 high, axillary, larger than radials, bearing cuneate, pinnulate secundibrachs. Anal sac long,cylindrical, composed of equidimensional stellate plates. Stem round, homeomorphic, remaining same width for20 mm below the crown.

Remarks. This species is placed in Scytalocrinus because it has ten arms and they branch on primibrachial1. In addition, the brachials are relatively low and the stem is round.



Figs 1±2. Glossocrinus whidbornei sp. nov. 1, holotype, NHM E21838, from Top Orchard, Pilton; lateral view of largeanal sac, proximal parts of two arms, cup, and proximal stem; ´ 1´5. 2, paratype, NHM 21837, from Top Orchard,Pilton; large anal sac, portions of arms, partially dislocated, and radial plates; ´ 2.

Figs 3, 9. Pachylocrinus? plumifer (Whidborne, 1898). 3, SM H186 from Barnstaple; lateral view of arms; ´ 1´2.9, lectotype, BGS 7153 from Braunton; lateral view of crown and proximal stem; ´ 1´3.

Figs 4, 10. Scytalocrinus barumensis (Whidborne, 1896). 4, lectotype, SM H182, from Barnstaple; lateral view ofdorsal cup and proximal arms; ´ 1´5; 10, SM H183 from Top Orchard, Pilton; lateral view of dorsal cup, arms, andproximal stem; partial anal sac also preserved between lower parts of arms; ´ 2.

Figs 5±6. Bridgerocrinus stadiodactylus (Whidborne, 1896). 5, lectotype, MND 365, from Roborough, TownsendHall collection; ¯attened dorsal cup, primibrachs, and proximal stem; ´ 2. 6, SM H191, from Pilton, William Portercollection; lateral view of dorsal cup, two primibrachs, lower part of anal sac, and disjunct part of stem; ´ 2.

Figs 7±8. Sostronocrinus mundus (Whidborne, 1898). 7, lectotype, NDM 365 from Braunton; lateral view of crusheddorsal cup, arms, and distal stem; ´ 2. 8, SM H179 from Croyde Rocks; lateral view of crushed dorsal cup, proximalparts of arms, and proximal stem; ´ 4.

P L A T E 5


Material. Whidborne (1898) ®gured two specimens of S. barumensis. We select SM H182 (Whidborne 1898, pl. 34,®g. 6) as the lectotype. The other specimen is SM H183 (Whidborne 1898, pl. 35, ®g. 3).

Measurements. SM H182: HDC, 5´0 mm; WDC, 6´3 mm.

Genus BRIDGEROCRINUS Laudon and Severson, 1953

Remarks. We note here an error in Moore et al. (1978, p. T640) with respect to this genus. The descriptionof this genus states that primibrachial 1, 2, or 3 may be axillary. The key preceding this description (Mooreet al. 1978, p. T639) states that primibrachial 2 is axillary. The type species of the genus is characterized byhaving two primibrachials per ray. Laudon and Severson (1953) did not mention any variation in thisnumber.

Lane et al. (1997) included in this genus two species from the Famennian of north-west China thatdiffer from the type species, one in having three, rather than two, primibrachials, and the other in havinga second arm branching high above the cup. We now believe these emendations were incorrect andreassign these two species. Bridgerocrinus minutus, with second arm branching, is reassigned here toSostronocrinus; Bridgerocrinus delicatulus is referred to Logocrinus, which may have 3 primibrachials toa ray.

Bridgerocrinus stadiodactylus (Whidborne, 1896)

Plate 1, ®gure 5; Plate 5, ®gures 5±6

v* 1896 Poteriocrinus stadiodactylus Whidborne, p. 377.v. 1896 Poteriocrinus batheri Whidborne, p. 377.

v* 1898 Scytalocrinus stadiodactylus (Whidborne); Whidborne, pp. 233±235, pl. 36, ®gs 2±6, 8; pl. 37, ®g. 14.1930 Scytatocrinus [sic] stadiodactylus (Whidborne); Schmidt, p. 87.1943 Scytalocrinus stadiodactylus (Whidborne); Bassler and Moodey, p. 678.

Diagnosis. A bridgerocrinoid with an anal sac that is in¯ated distally.

Description. Dorsal cup high, conical, straight-sided, plates smooth. Infrabasals high, occupying lower one-third ofcup height. Radial facets occupy full width of radials. Three anal plates in cup. Second primibrachials axillary,brachials narrow, high, pinnulate. Ten arms. Column slender, tapering in width just below the crown, round,homeomorphic.

Remarks. In every respect this species ®ts the diagnostic characters for Bridgerocrinus, and is closelysimilar to the type species, B. fairyensis Laudon and Severson, 1953, from Montana, differing mainlyin the structure of the anal sac. The distinctive anal sac is shown more clearly in a drawing byWhidborne (1898, pl. 37, ®g. 14) than in the actual specimen, but we believe that his drawing isessentially correct.

Material. We here select NDM 396, Townsend Hall collection, from Braunton, (Whidborne 1898, pl. 36, ®g. 2; pl. 37,®g. 14) as lectotype. This consists of a good dorsal cup and proximal parts of the arms. Other specimens include SMH193 from Barnstaple (Whidborne 1898, pl. 36, ®g. 3); NDM 401, Townsend Hall collection, from Roborough(Whidborne 1898, pl. 36, ®g. 5); SM H191, Porter collection, from Pilton (Whidborne 1898, pl. 36, ®g. 4); NDM 402,Townsend Hall collection, from Roborough (Whidborne 1898, pl. 36, ®g. 8); SM H192 (Whidborne 1898, pl. 36, ®g. 6)on the same small block of rock as Whidborne's sp. indet. (SM H194), an arm fragment (Whidborne 1898, pl. 36,®g. 7).

Measurements. NDM 402: HDC, 4´5 mm; WDC, 6´2 mm. SM H191: HDC, 6´3 mm; WDC, 6´2 mm.


Bridgerocrinus? arachnoideus (Whidborne, 1898)

Plate 6, ®gure 6

v* 1898 Scytalocrinus arachnoideus Whidborne, p. 235, pl. 38, ®gs 5±6.1930 Scytatocrinus [sic] arachnoides [sic] (Whidborne) [sic]; Schmidt, p. 87.1943 Scytalocrinus arachnoides [sic] Whidborne; Bassler and Moodey, p. 677.

Remarks. Neither specimen illustrated by Whidborne shows the dorsal cup well enough to verify hisdescription and illustration. Both specimens show the arms quite clearly, but the cup area is poorlypreserved. Whidborne (1898) stated that the cup is bowl-shaped, which we take to mean that it hasconvex sides. The infrabasals are clearly visible in side view in his plate 38, ®gure 5, and the cup isrelatively high. Three anals in the cup, elongate brachials and arms branching on primibrachial 2,without further subdivision, clearly place this species in Bridgerocrinus. Hence, we refer the speciesonly questionably to this genus because we cannot con®rm Whidborne's description. The stem is roundand heteromorphic.

Material. SM H195 is from Barnstaple (Whidborne 1898, pl. 36, ®g. 5). SM H196, is also from Barnstaple (Whidborne1898, pl. 36, ®g. 6), and is here selected as the lectotype. SM H191 (Whidborne 1898, pl. 36, ®g. 4) is from Pilton andin the Porter collection; MND 402 is from Roborough, and in the Townsend Hall collection.

Measurements. SM H196: HDC, 6´2 mm; WDC, 5´3 mm.

Bridgerocrinus? sp.

Plate 1, ®gure 3

Remarks. A single, small crushed specimen from the Hamling Collection (NHM E21847) is identi®ed asBridgerocrinus? sp. This specimen, which apparently was not seen by Whidborne, is a very small crushedcladid that resembles other specimens of Bridgerocrinus in the fauna.

Family SOSTRONOCRINIDAE fam. nov.

Diagnosis. Cup with infrabasals visible in side view, radial facets plenary; primibrachial 2 typicallyaxillary, rarely primibrachials 1, 3, or 4 axillary; with one additional higher bifurcation in each halfray; three anal plates in cup; column round; articulation between columnals homeomorphic orheteromorphic.

Type genus. Sostronocrinus Strimple and McGinnis, 1969.

Included genera. In addition to the type genus, Haertocrinus Moore and Plummer, 1940, and Tundracrinus Yakovlev,1928 are placed in this family.

Range. Late Devonian (Famennian) to Early Permian.

Remarks. These three genera are here removed from the Scytalocrinidae, where they were placed byMoore et al. (1978). They differ from typical scytalocrinoids in having 20, rather than ten, arms. Theydiffer from the Decadocrinidae in having a ¯at or convex base, rather than a concave base, in havingmore than ten arms, and in having infrabasals that make up a larger part of the cup side. They differfrom the Pachylocrinidae in having infrabasals visible in side view, and in lacking additional armbranching higher in the crown. The two younger genera, Haertocrinus and Tundracrinus, have armsthat branch on the ®rst primibrachial. The older genus, Sostronocrinus, has arms that branch on thesecond primibrachial in Lower Carboniferous species, and on the third or fourth primibrachial inDevonian species.


Genus SOSTRONOCRINUS Strimple and McGinnis, 1969

Sostronocrinus mundus (Whidborne, 1898)


v* 1898 Mariocrinus? [sic] mundus Whidborne, pp. 219±220, pl. 34, ®g. 5?; pl. 37, ®gs 6±7.v. 1930 Hydriocrinus mundus (Whidborne); Schmidt, p. 87.

1943 Hydriocrinus mundus (Whidborne); Bassler and Moodey, p. 517.

Diagnosis. Cup high, conical; cup plates smooth; brachials cuneate.

Description. Crown slender, ranging from small to medium-sized. Dorsal cup high, slender, straight-sided. Infrabasalsrelatively large and visible in side view. Arms pinnulate. Two primibrachials to a ray, with a second higher armbranching. Brachials cuneate. Stem round, heteromorphic proximally, becoming homeomorphic distally, nudinodalsseparated by 1±3 internodals.

Remarks. Whidborne mistook these two specimens for a camerate crinoid, mistaking primibrachials for®xed brachials. Mariocrinus (a mis-spelling of Mariacrinus Hall), to which he assigned the species withquestion, is now a junior synonym of Ctenocrinus Bronn, which belongs to the camerate familyMelocrinidae. The Devon specimens bear little resemblance to Ctenocrinus, so Schmidt (1930) rightlyconsidered them to be inadunate crinoids, although we believe his assignment to the Upper Carboniferousgenus Hydriocrinus is incorrect. It should be noted, however, that many of the genera now placed in theScytalocrinidae, like Sostronocrinus, had not been named in Schmidt's day. The specimens at hand differfrom Hydriocrinus in having a round, rather than a pentagonal, stem, and in having primibrachial 2 or 3axillary, rather than primibrachial 1. The specimens differ from most other scytalocrinoids in having 20,rather than ten, arms. Haude and Thomas (1989) recognized a species of Sostronocrinus in the Famennianof Germany. We believe their species is congeneric with the Pilton specimens, although they questionedthe generic identi®cation.

Material. The lectotype, here selected, is NDM 365 (Whidborne 1898, pl. 34, ®g. 5). The other specimen isSM H179.

Measurements. NDM 365: WDC, 4´6 mm; HDC, 5´1 mm.

Family APHELECRINIDAE Strimple, 1967

Remarks. The species described below is placed in this family because it has more than 20 arms and the®rst primibrachial is axillary. None of the genera in the families Scytalocrinidae or Sostronocrinidae has



Figs 1±2, 4, 8. Aphelecrinus tensus (Whidborne, 1896). 1±2, holotype, SM H181 from Barnstaple; lateral view ofcrushed dorsal cup, fragments of arms, and proximal stem. 1, ´ 1´7. 2, ´ 3´8. 4, 8, NHM E7197 from Top Orchard,Barnstaple, Townsend Hall collection. 4, oral view of arms with pinnules; ´ 1. 8, enlargement of portion of 4showing oral view of arms and pinnules; ´ 3.

Fig. 3. Genus and species indet., SM H216, from Barnstaple; lateral view of arms; ´ 1´5.Fig. 5. Pachylocrinus? plumifer (Whidborne, 1898), NDM 378 from Braunton; oblique view of cup and proximal

arms; ´ 1.Fig. 6. Bridgerocrinus? arachnoideus (Whidborne, 1898), lectotype, SM H196 from Barnstaple; lateral view of crown,

cup plates partially displaced, and several attached columnals; ´ 1´2.Fig. 7. Family and genus indet., SM H190, from Upcott Arch; lateral view of partial crown, dorsal cup, and arms; ´ 1´5.

P L A T E 6


this speci®c combination of features. Among other families in the superfamily, the Blothrocrinidae doescontain such genera as Ulrichicrinus, Woodocrinus, and Nebraskacrinus, but each of these genera hasunusual features that do not ®t our specimens.

Genus APHELECRINUS Kirk, 1944

Aphelecrinus tensus (Whidborne, 1896)

Plate 6, ®gures 1±2, 4, 8

v* 1896 Poteriocrinus tensus Whidborne, p. 377.v* 1898 Poteriocrinus tensus Whidborne, p. 226, pl. 35, ®gs 1±2; pl. 38, ®g. 1.

1930 Culmicrinus tensus (Whidborne); Schmidt, p. 87.1943 Culmicrinus tensus (Whidborne); Bassler and Moodey, p. 383.1945 Hydriocrinus tensus (Whidborne); Bassler and Moodey, p. 518.

Diagnosis. An aphelecrinoid with a large, high, dorsal cup composed of smooth plates; anal sac very long,cylindrical, composed of numerous rectangular plates.

Description. Dorsal cup conical, large, higher than wide, with large, upwardly ¯aring infrabasals; cup sides straight.Radial facets full width of radials; anal plates not observed, reported to be three by Whidborne. Primibrachial 1 large,axillary, supporting cuneate, pinnulate brachials that branch once well above the cup in at least some half rays. Stemround, consisting of alternating thick and thin columnals.

Remarks. This is the ®rst report of this genus and family from Devonian rocks. Numerous species ofAphelecrinus have been reported from throughout the Mississippian of North America, and a singlespecies was reported by Wright (1945) from the Visean of Scotland. A. tensus seems to be most closelyallied to Kinderhookian species, such as A. elegantulus (Wachsmuth and Springer), from the centralUnited States.

Material. Unlike most of his species, Whidborne explicitly stated that the Woodwardian (Sedgwick) Museumspecimen is the type of this species. This is SM H181, from Barnstaple. The other specimen is NDM 413 (Whidborne1898, pl. 38, ®g. 1).

Measurements. SM 181: WDC, 8´9 mm; HDC, 9´4 mm.

Aphelecrinus? sp.

Plate 4, ®gure 8

v. 1898 Scaphiocrinus sp. Whidborne, pp. 231±232, pl. 34, ®g. 8.1930 Hydriocrinus sp. (Whidborne); Schmidt, p. 87.

Remarks. The dorsal cup of this specimen, BGS 7152, from Braughton, is suf®ciently damaged that we areunable to make a positive generic identi®cation. However, the cup is signi®cantly higher than indicated byWhidborne, who noted that it was very shallow and bowl-shaped. It appears to us to be a high bowl shape,with infrabasals clearly visible in side view. The arms are as indicated by Whidborne, branching on the®rst primibrachial and then once higher on the arm, resulting in 20 arms. These features taken togetherindicate that the specimen can be questionably assigned to Aphelecrinus. The stem is round, slender, andindistinctly heteromorphic.

Measurements. BGS 7152: HDC, 4´8 mm; WDC, 3´0 mm.


Family PACHYLOCRINIDAE Kirk, 1942

Genus PACHYLOCRINUS Wachsmuth and Springer, 1880

Pachylocrinus? plumifer (Whidborne, 1898)

Plate 1, ®gure 8; Plate 5, ®gures 3, 9; Plate 6, ®gure 5

v*? 1898 Scaphiocrinus? plumifer Whidborne, pp. 228±229, pl. 31, ®gs 4±4c; pl. 33, ®g. 1; pl. 36, ®gs 1±1a;pl. 38, ®g. 2.

1930 Hydriocrinus plumifer (Whidborne); Schmidt, p. 57.1930 Scytactocrinus [sic] plumifer (Whidborne); Schmidt, p. 87.

Description. Dorsal cup low, base ¯at to slightly concave, tips of infrabasals visible in side view. Cup plates withridges, depressed or with pits at plate corners. Radial facets plenary. Primibrachial 2 axillary in several observed rays,but variation from one to several primibrachials occurs in different specimens. Arms branch at least twice aboveprimaxils. Stem pentagonal to pentalobate, consisting of nudinodals separated by two internodals proximallybecoming homeomorphic distally.

Remarks. This species closely resembles species of Pachylocrinus, differing mainly in having tips ofinfrabasals visible in side view. In the light of the age difference between this older species and younger,Early Carboniferous, species of the genus, this may be an evolutionary difference, with Early Carbonifer-ous pachylocrinoids evolving from Late Devonian forms like P.? plumifer by having the infrabasalscon®ned to the base of the cup in younger species. In some respects P.? plumifer also resemblesSostronocrinus, differing mainly in having a less conical, more bowl-shaped cup, and in having additionalarm branching. It has a less conical cup and less stout arms than Blothrocrinus and other blothrocrinoids.Given the differences noted above, and the lack of better preserved specimens, we have chosen to placeWhidborne's species only questionably in Pachylocrinus.

Material. We here designate BGS 7153 from Braunton as the lectotype. SM H187, from Barnstaple, is morecomplete, but not as well preserved. Other specimens include SM H185 (Whidborne 1898, pl. 31, ®g. 4) and SMH188a. Our cast fails to reveal any of the details shown in Whidborne (1898, pl. 31, ®g. 4a). His plate 33, ®gure 1 isSM H186. BGS 7153 also was illustrated by Whidborne (1898, pl. 36, ®g. 1). In addition, NDM 378 from Brauntonincludes a good, small partial crown. The TNHS holds a specimen consisting of a set of arms that is labelled?S. plumifer, which may or may not belong to this species. The fossil is in the Hamling collection, no. 73/12, fromnorth of Chelfham.

Measurements. BGS 7153: HDC, 4´2 mm; WDC, 4´2 mm. SM H187: WDC, 6´5 mm; HDC, 4´2 mm. NDM 376: HDC,5´5 mm; WDC 7´3 mm.

Family CERCIDOCRINIDAE Moore and Laudon, 1943

Genus and species indet.

Plate 6, ®gure 3

v. 1898 Coeliocrinus n. sp. Whidborne, p. 236, pl. 38, ®g. 7.1930 Coeliocrinus? sp. Whidborne; Schmidt, p. 87.

Remarks. Whidborne applied the name Coeliocrinus to a set of endotomously branching arms. ThreeLower Mississippian genera (Cercidocrinus Kirk, 1938; Ascetocrinus Kirk, 1940; Coeliocrinus White,1863), all of which have endotomously branching arms, are currently placed in the Family Cercidocri-nidae. Because a dorsal cup is not present on Whidborne's specimen, allocation below the family level isnot possible.

Material. SM H216 from Barnstaple (Whidborne, 1898, pl. 38, ®g. 7).


Family and genus indet.

Plate 6, ®gure 7

v. 1898 Scaphiocrinus? inordinatus Whidborne, pp. 230±231, pl. 34, ®g. 7?; pl. 35, ®gs 6±6a; pl. 38, ®g. 4.1930 Pachylocrinus inordinatus (Whidborne); Schmidt, p. 87.

Remarks. Although we have a total of ®ve casts labelled as this species, none is suf®ciently well preservedto allow us to either write a new description or assign it to a presently accepted genus. Therefore, we havechosen to place these specimens in open nomenclature and to consider the name Scaphiocrinus?inordinatus as a nomen dubium.

Material. According to Whidborne (1898, p. 230), his specimen from Upcott Arch was the best preserved and mostcomplete, and we here designate it the lectotype for the species. The specimen is given as part and counterpart, SMH190a and H190b (Whidborne 1898, pl. 38, ®g. 4). SM H189 is from Barnstaple (Whidborne 1898, pl. 35, ®gs 6±6a).BGS 7151 also was illustrated by Whidborne (1898, pl. 34, ®g. 7).

D I S C U S S I O N

Etheridge and Carpenter (1886, p. 129) stated that `the Devonian rocks of the British Isles have yielded butthe scantiest evidence of Blastoid Life and we are not in a position to af®rm with certainty the genericrelations of specimens hitherto found.' The situation has not improved markedly in the past 114 years. Inour observations of scant blastoid material from the Upper Devonian of Great Britain present in the variousmuseums, we have found only a single specimen that can be assigned with certainty at the generic level,and it is Early Carboniferous, not Devonian. The specimens are typically partials, preserved as moulds,and all lack clarity in the oral area, which is critical in the generic identi®cation of blastoids. The onecomplete blastoid specimen collected from this area since the publication of Etheridge and Carpenter(1886) of which we are aware (NHM E30486) is an internal mould of a pyriform ®ssiculate genus. Fromour analysis, as many as four blastoids are present in this fauna. One is the godoniform spiraculate that canquestionably be assigned to Petaloblastus, which is also known from reasonably well-preserved materialfrom the Etroeungt beds of Germany (Lane et al. in press). The other undoubted spiraculate, Mesoblastuscf. M. crenulatus, is from Fremington, which is Early Carboniferous. One specimen is a pyriform®ssiculate genus that questionably belongs to the Family Phaenoschismatidae, based on our study of theinternal mould mentioned above. The last specimen was assigned by Whidborne (1898) to the ®ssiculategenus Codonaster; however, the fragmentary nature of the material precludes assignment even to order.Where known, all of these blastoid families are represented in coeval Famennian faunas from north-western China described by Lane et al. (1997) and Waters et al. (in review). The Chinese faunas are muchmore diverse and abundant, and are much better preserved than the blastoid material from the typeDevonian.

Given the close geographical proximity of south-west Britain to the eastern part of North Americaduring Famennian time (McKerrow and Scotese 1990, ®g. 16), it is not surprising that crinoids from thesetwo areas show a number of similarities. Certainly they show greater similarity to each other than eitherone does with the very far removed Famennian echinoderms of north-western China (Lane et al. 1997).

The ¯exible crinoids of the Pilton Beds show relationships to those from North America, primarilythrough the occurrence of species of Taxocrinus in both areas. However, they fail to show any relation toChinese faunas that are dominated by Eutaxocrinus, a taxon that is unknown in the Famennian of Europe,but does occur in both older and younger European rocks.

With regard to camerate crinoids, Eumorphocrinus, identi®ed here, is the ®rst report of this genus belowthe Carboniferous. It does occur in the Tournaisian of Scotland, Ireland, and Australia, and possibly in thewestern United States. The presence of Actinocrinites here shows a relationship with the LowerCarboniferous of Britain, Ireland, and North America, but not with Asia. The occurrence of platycrinoidsin the Famennian of Britain, Germany, and China, but not in North America, shows a pattern differentfrom that for other camerates. The general lack, or rarity, of platycrinoids in North America both belowand above the Lower Carboniferous is different from that on other continents where platycrinoids are


common in the Upper Carboniferous or Permian. Finally, the occurrence of the hexacrinoid Adelocrinus inthe Famennian of Germany and Britain, and the occurrence of the closely related genus Arthroacantha inthe pre-Famennian Devonian of North America, again points to a biogeographical relationship amongthese areas. These crinoids are not known from Asia, although specimens of Hexacrinites have been foundin China.

With regard to cladid crinoids, the Pilton Beds, as well as Chinese and North American faunas, aredominated by pinnulate forms that are generally allied with the scytalocrinoids. These include severalfamilies and genera; however, the exact composition of each fauna is somewhat different. There is littleevidence for any of the older, more primitive, non-pinnulate cladids. One feature of the Pilton fauna is thepresence of true Poteriocrinites and of glossocrinoids, both of which show a relationship to the NorthAmerican faunas, where glossocrinoids largely are con®ned.

While the above discussion centres on geographic differences and similarities, we can also considerrelationships based on age (holdovers and precursors). Some of the crinoids represent last appearances oftheir genus or family. Adelocrinus is a good example, as are the glossocrinoids in the fauna. Theseholdovers are in a minority, however, when compared with the precursors that show kinship with LowerCarboniferous fossils. They include a variety of scytalocrinoids, the actinocrinoids, and Platycrinites. Thetypical earlier Devonian cladids that lack pinnules are not present, nor are the various large, many-plated,boxy camerates that typify many earlier Devonian crinoid faunas. In these respects, the Pilton fauna isclearly foreshadowing the dominant features of Lower Carboniferous crinoid faunas in both Europe andNorth America.

Acknowledgements. We gratefully acknowledge the helpful reviews of this manuscript by Steve Donovan and AndyKing. We especially thank the various curators of collections who helped make this study possible. These includeMichael Dorling, Sedgwick Museum, University of Cambridge; Allison Mills and Jerry Lee, North Devon Museum,Barnstaple; E. J. Loef¯er, University of Bristol; Steven Tunnicliff, British Geological Survey, Nottingham; DavidLewis and Andrew Smith, The Natural History Museum, London; and Michael Bishop, Torquay Natural HistorySociety Museum, Torquay. In addition, we received help and advice from George Sevastopulo, Kenneth Higgs,Michael House, and Roland Goldring. The photographs were taken by George Ringer. Thanks go to Ruth Droppo forproducing Text-®gure 1. Partial support for this study was provided by National Science Foundation grant EAR9406116.

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N. G. LANE

C. G. MAPLES

Department of Geological SciencesIndiana University

Bloomington, IN 47405, USAe-mail [email protected]

[email protected]

J. A. WATERS

Department of GeologyState University of West Georgia

Carrollton, GA 30118, USAe-mail [email protected]

Typescript received 26 August 1999Revised typescript received 17 January 2001


Revision of Late Devonian (Famennian) and some Early Carboniferous (Tournaisian) crinoids and...

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