U ni versi ty of Queensland PAPERS
DEPARTMENT OF GEOLOGY
Volume III. 1951 Number 14
Upper Devonian and Middle Carboni fe rous Brachiopods of Queensland
By
W. G. H. MAXWELL, B.Sc . Department of Geology, University of Queensland.
UNIVERSITY OF QUEENSLAND PRESS BRISBANE
DATE O.'f PUBLICATION 30th SEPTEMBER, 1951 DEPARTMENT OF GEOLOGY
VoL. IlL 1951 NUMBER 14
UPPER DEVONIAN AND MIDDLE CA RBONI FEROUS BRACHIO PODS OF QUEENSLAND
By
VV G. H. MAXWELL, B.Sc.
University of Queensland
UNIVERSITY OF QUEENSLAND PRESS BRISBANE
DATE OF PUBLICATIO� 30th September, 1 !).) I UPPER DEVONIAN AND MIDDLE CARBONI FEROUS BRACHIOPODS OF QUEENSLAND By W. G. H. MAXWELL, B.Sc. Department of Geology, University of Queensland. (With 4 Text Figures, 2 Maps, and 4 Plates) . Abstract.-Brachiopods from the Upper Devonian and Middle Carboniferous beds of Queensland are described and classified. They have been compared with species of a similar nature from other parts of the world, and the age of the beds in which they occur has thus been confirmed. One new genus-Levipustula, of the sub-family Productinae, new species of Spirijerellina Fredericks, Neospir�fer Fredericks, Cyrtospir�fer Nalivkin, and Sinospirijer Grabau, are described. I. INTRODUCTION. The material used for these investigations was borrowed from the Geology Department of the University of Queensland, and from the Queensland Geological Survey. In addition, specimens were collected by the author and companions from the Neerkol and Mt. Barney districts. Several of the species found in the Queensland beds also occur in New South Wales. In the Lambie Series (N.S.W.), two spiriferoid species are present. One of these is found in Queensland and the other closely resembles a Queensland species. I have described this second species from the Lambie Series and, although the title of this paper includes Queensland Brachiopods only, I believe that its close relation to the Queensland forms warrants its description. The specimens have been preserved as external casts and internal moulds in indurated mudstones. Complete absence of shelly material has made impossible the examination of the structure and growth of the shell layers. The terminology adopted in this paper is the same as that used by Dr. H. M. Muir-Wood (1928) for the Protremata, and that used by P. E. Cloud (1942) for the Telotremata. However, there are several terms which seem to have caused controversy among recent workers on brachiopods, and so the following interpretation of these is used. I . Deltidium.---,--This term has been used for any plate or plates which close or partly close the delthyrium of species of the Telotremata. Since shelly material is lacking in the Queensland specimens, it is almost impossible to decide the direction a�d manner of growth of the deltidium, and whether it consists of one or two plates. 2. Delthyrial cavity.-This is the space directly ventral to the delthyrium. The Umbonal cavities are the two regions ventral to the area, and separated from the delthyrial cavity by the dental lamellae.
3. Dental lamellae or dental plates. -These are two structures extending from the under surface of the area to the floor of valve. They border the delthyrial cavity, and at their junction with the area two teeth are usually produced. The dental lamellae may be straight or curved, parallel or divergent. 4. Apical apparatus.-This term includes all features associated with the posterior part of the ventral valve, such as dental lamellae, deltidium and median septum. II. DEVONIAN BRACHIOPODS. Introduction The Devonian brachiopod species described in the following pages, were collected from the Upper Devonian beds in the Star River Basin and the Dotswood area near Townsville, and in the Mt. 1\lorgan District. One species from the Lambie beds at Solitary Creek, New South ·wales, is also described. The three species 4 UPPER DE VONIA N AND
described belong to the genera Cyrtospirifer Nalivkin and Sinospirijer Grabau, both of which are closely related. In addition to these, other brachiopod species are present in the fauna, but poorly preserved material has not allowed a description of these to be made. Stratigraphical Background : The Devonian deposits of Queensland are restricted to the eastern portion of the State. The sediments are mainly geosynclinal, some being marginal, while further west, fresh water deposits occur (Bryan and Jones 1945, p. 27). Lower Devonian beds may be represented in the far north by the upper part of the Chillagoe Limestone, and the Broken River Series. The Silverwood Series near Warwick has been placed in the Lower Devonian. Lower Devonian fossils have also been found in the Mt. Etna Series near Rockhampton. In the Clermont District, fossiliferous limestones of Middle Devonian age occur in the Douglas and Drummond Creeks. The Burdekin Series, near Townsville, consisting gf conglomerates, sandstones, calcareous shales and limestones, is of late Middle Devonian (Givetian) age. The limestones contain a rich coral fauna (Hill 1942) , and also the brachiopods Stringocephalus (Brown 1944) Atrypa and Gypidula, and the nautiloids Gyroceras and Phragmoceras. Devonian beds found at Ukalunda have not been accurately determined, but part of the series, at least, is as old as the Couvinian (Whitehouse 1929, p. 158). With the passage from Middle Devonian . to Upper Devonian in Queensland, there was a change, in many areas, from marine to lacustrine conditions. This change seems to have been gradual, as '' paralic " sediments occur in many localities. In the Dotswood Series which overlies the Burdekin Series, both plants and marine fossils are present. These conditions are also found at Mt. Wyatt. The Star Series, north-west of Townsville, contains Upper Devonian brachiopods, but part of the series may extend into the Lower Carboniferous. Systematic Descriptions : Order Telotremata Beecher 1891. Super-family Spiriferacea Waagen 1883. Family Spiriferidae King 1846. Sub-family Cyrtiinae Fredericks 1926.
Diagnosis. -Forms characterised by dental lamellae, a deltidium and an apical apparatus similar to that in Spirifer ostiolatus Hall and Clarke ; the dental lamellae extend to the floor of the valve and are thickened on their inner sides and across their dorsal ends to form the deltidium. Remarks.-The apical structure characteristic of this sub-family is illustrated by two sections through the rostral part of the shell.
Fig. 1. JIIDDL E CARBONIFEROUS BRA CHIOPODS OF QUEENSLA ND 5
Fredericks (1926, p. 410) divided the sub-family Cyrtiinae into four genera, according to the external ornament. These genera are : Cyrtella Fredericks, 1919, with strong radial plications. Cyrtia Dalman, 1828, which has the fine filamentous ornament characteristic of the Silurian spirifers. Cyrtospirife r Nalivkin, 1918, which is finely and evely plicate over the whole external surface. Spinocyrtia Fredericks, 1916, with radial plications and finer concentric ornament. To these may be added the genus Sinospirifer Grabau 1931 which has radial costae similar to those of Cyrtospir�fer but differs slightly from this genus internally. Genus Sinospirife r Grabau 1931. Grabau 1931, pp. 231-249, pl. XXVIII, figs. 4-6 ; pl. XXIX, figs. 1-3. Tien 1938, pp. 110-116, pl. XVI, figs. 1-3, text fig. 20 A-B. Genotype (by original designation) .-Sinospirifer sinensis Grabau 1931, p. 241, pl. XXVIII, figs. 4-6, pl. XXIX, figs. 1-3. Lungkouchung beds at Hsianghsiang, of Upper Devonian (Frasnian) age. Diagnos£s.--Shell subequally convex. Cardinal extremities acute or extended. Cardinal area arcuate. Sinal plications include two primary plications. Two diverging dental lamellae rest on the floor and curve outward, bounding the muscular impression. Umbonal part of the shell usually thickened. Remarks.-This genus was proposed by Grabau (1931) to include all those Upper Devonian spirifers in which the lateral plications are simple, while those in the sinus and on the fold increase by intercalation and bifurcation. Tien (1938, p. 110), after a re-study of this group, both on the basis of internal as well as external characters, emended the sub-genus and separated out two more sub-genera, Tenticospirifer and Hunanospirifer, from Grabau's original group. When Grabau (1931) proposed the sub-genus, he drew attention to the similarity between the Chinese species of Sinospirifer and the European forms belonging to the genus Cyrtospirifer. His distinctions between the two genera were mainly phylogenetic rather than morphological. The only morphological distinctions which he noted were the smaller size of the Chinese forms and the lirate character of the shell. The latter feature is seen in some species of Cyrtospirifer. Internally, the dental plates are less strongly developed and tend to be more curved in Sinospirifer than in Cyrtospirifer. These differences hardly seem to be of generic significance, but as I have been unable to compare specimens of the genotype of each genus, I have accepted the two groups as distinct. Range.-Many of the specimens of Sinospirifer s£nensis Grabau, figured by Grabau (1931) came from unknown localities and horizons. Tien (1938) collected this species from the lower part of the Lungkouchung beds at the type locality of Hsianghsiang, where it occurs �n a shaly limestone. It was also found in a weathered shale, capped by the Yohlu sandstone near Shehyenper, in Changsha. The significance of these localities and horizons can be understood after examining the stratigraphical table of Tien (1931, p. 6). The following is a summary of part of this table :- Upper Devonian. a. Hsikuangshan series (=Famennian), replaced by the Yohlu sandstone in Changsha and Hsiangtin districts. At the base is the Changlungchieh shale, then the Tutzutang limestone and the Makunao limestone. 6 UPPER DEVONIA N AND
b. Shetienchiao Series-characterised by the Sinospirifer fauna (=Frasnian). l. Shetienchiao beds-limestones, sandstones and shales, found in the Shaoyang, Hsianghsiang, Changsha, Ninghsiang and Anhua districts. 11. Lungkouchung beds-sandstones and shales, often with intercalations of limestones. Middle Devonian. a. Chitzechiao Series ( =Givetian)-characterised by the Stringocephalus fauna. b. Tiaomachien Series ( =Eifelian). Sinospirifer sinensis var. australis var. nov. (Plate I, Figs. 1-16, Plate II, Figs. 1-3). ·Holotype.-F ll927 University of Queensland. Type locality.-4! miles south-east of Dotswood Homestead along the Fanning River Road, Parish of St. Giles, County of Nares. Age.-The age of the beds, in which this species occurs, had not, up t<;> the present, been accurately determined, but part of the beds at least were believed to b� of UI?per Devonian Age. The occurrence of this species helps to confirm this previOus v1ew. Diagnosis.-Variety narrower and smaller than genotype. Otherwise diagnosis as for genus. Description.-External features. The width of the ventral valve is slightly greater than the length, the maximum width occurring at the hinge line. The dorsal valve is much wider than it is long. The cardinal extremities of both valves vary from rounded corners to pointed extensions. The hinge line is straight. Dimensions Ventral Valve Dorsal Valve Width 17 18 25 30 mm 16 24 26 mm Length 16 14 19 25 mm 13 20 19 mm The ventral valve is strongly convex, with a median sinus that varies from rounded to subangular. The cardinal area extends for the entire length of the hinge line and is triangular to subrectangular in outline. It may be fiat at the extremities, but it is always curved in the delthryial region. The delthyrium is a triangular slit, extending from the hinge margin almost to the apex of the valve. In all the specimens examined it was not possible to determine whether the deltidium consisted of one or two plates. Two sharp teeth occur at the anterior corners of the delthyrium. The dorsal valve is also convex and carries a median fold, which broadens anteriorly. The fold is bounded by two deep furrows, which are more prominent than the furrows between the plications on the rest of the valve. The area of the dorsal valve is very narrow, and often indistinct . . The ornament is the same on both valves, consisting of small radial plications, which are simple, on the lateral slopes. The plications in the sinus and on the fold are characteristic of this species. There are two primary plications in the sinus, and other minor plications which arise by intercalation or by bifurcation. The sinal plications, as defined by Grabau (1931, No. 2) are defined by the three following formulae, viz. :- a. 2 + I + 1 + ly + 1 + I + 2. b. 2 + I + 2 + I + 2. MIDDLE CARBONIFEROUS BRACHIOPODS OF QUEENSLA ND 7
c. 1 + I + 1 + ly + 1 + I + l. I =primary plication. y=an intercalated plication. The first sinalformula is more common in the larger, more mature individuals.
Fig. 2.
Figures illustrating the three types of sinal plications found in Sinospirifer sinensis var. a-ustralis.
The number of plications per 5 mm., measured across the lateral slopes at a distance of 5 mm. from the apex, varies from 9 to ll. In addition to these plications very fine radial lirae are present. These can be seen clearly only with the aid of a microscope. They average from 9 to 10 per plication, or 80 to 100 per 5 mm. This lirate ornament is restricted to the exterior of the valves. Concentric growth lines occur, but these are neither prominent nor abundant, except in the antero lateral regions.
Internal feat-ures. In the ventral valve, two dental plates diverge from the posterior part of the shell, extending downwards from the lateral margins of the delthyrium, sloping towards each other for half their height, and then curving away laterally. They merge into the floor of the valve, extending anteriorly for some distance, as small ridges bounding the muscle scar. They are thickened posteriorly, and are joined together by a transverse thickened fold, which lies across the posterior part of the delthyrial cavity. This apical structure is characteristic of the sub-family Cyrtiinae Fredericks. In most specimens a small vertical ridge projects downwards from this transverse thickening, and merges into the median part of the muscle scar. The muscle scar is an oval, striated area, occupying that part of the valve floor between the two dental plates. It extends well into the posterior part of the valve and, except for the small vertical ridge which merges into its posterior margin, there is no median septum dividing it. Anterior to the muscle scar, there is a low rounded fold, corresponding to the sinus on the exterior of the valve. The umbonal cavities, in the postero-lateral regions, are sharp, conical sockets. The sharp apices of the umbonal cavities are characteristic of all the specimens examined. Except for the umbonal cavities, and muscle scar, the interior of the valve carries an ornament of small radial plications and fine growth lines.
Specimens of the dorsal valve have been poorly preserved, and the internal features, in particular, are not distinct. A small striated cardinal process occurs at the centre of the posterior margin. Rising from its base is a '-minute median ridge, which extends for several mm. along the floor of the valve. A smooth area for muscle attachment occurs on either side of this ridge. The crural bases are small, indistinct structures, lateral to the cardinal process. UPPER DE!.ONhJ.N AND
Range.--Sinospirifer sinensis has been found only in beds of Upper Devonian age in China. and so I believe that the age of part of the beds, at least, in which the Queensland variety is found, is Upper Devonian. This variety occurs at Corner Creek in the Star Basin, in the Dotswood beds near Townsville, and in the Lambie Series at Solitary Creek, New South Wales. A spiriferid similar to this Queensland form has been found in beds of Frasnian age in North:Western Australia (Teichert, 1943). Dr. C. Teichert has allowed the author to examine some of the material which he collected from these beds, and although the species is different from the Queensland form, they both appear to belong to the genus Sinospirifer Grabau. Remarks.-Examination of specimens collected from the Lambie Series has shown that two species are present in this series. One resembles the true Spirifer disjunctus Sowerby, and the other is Sinospirijer sinensis var. australis. Specimens of this latter variety, which occurs in the Lambie Series, are of more constant size than the Dotswood and Star specimens. In the Queensland specimens, the dimensions show a fairly wide range, but mainly the larger individuals are present in the Lambie Series. Sinospirifer sinensis var. australis differs from Cyrtospirifer reidi sp. nov. in the curved nature of its dental plates and also in the contour of the dorsal valve. Other features of the two species are similar. Genus.-Cyrtospirifer Nalivkin. Genotype.-Spirifer verneuili Murchison 1840, p. 252, pl. II, figs. 3a. b, d. Ferques, Upper Devonian (Frasnian) ; description of genus reproduced by Grabau (1931 , p. 227). Diagnosis.-Medium sized shells, with costate fold and sinus. Intercalation and bifurcation of the costae in the sinus and fold, but never on the lateral slopes. Dental plates strong, divergent, and vertical or slightly curved, extending along the valve floor for half its length. Range.-The range of the genus has been discussed by the author in the paper "The Occurrence of an Upper Devonian Brachiopod (Cyrtospirifer reidi sp. nov.) m the :\It. Morgan District." Cyrtospirifer reidi Maxwell 1950.
Remarks.-This species has been described and figured m a prevwus paper (�Iaxwell, 1950) . Cyrtospirife r subdisjunctus sp. nov. (Plate I, fig. 17 a, b, c). Holotype.-F ll925, University of Queensland. Type locality.-Solitary Creek, near Bathurst, N.S.W. Synonomy.-Spirifera disjuncta (Sowerby), Brown I.A., and Joplin, G.A., 1938, p. 220.
Age. -This species occurs in the Lambie Series, which has been determined as Upper Devonian in age.
Diagnosis. -Approximately the same dimensions as the genotype ; width greater than length ; cardinal extremities pointed ; primary sinal plications not prominent. Thickening of the dental plates only in the delthyrial cavity. The dental plates are usually vertical, rarely slightly curved. Umbonal part of shell greatly thickened.
Description External features.-The ventral valve is almost twice as wide as it is long, the ears being very alate, and pointed. This valve is convex, carrying a median sinus which widens and becomes shallow anteriorly. Primary plications in the sinus are not prominent. The cardinal area is well developed, triangular and JJIDDLE CA RBONIFERO US BR.-l CHIOPODS OF QUEENSLAND 9
curved. A large triangular delthyrium intersects the area. This is closed by a deltidium. The umbo is only slightly incurved over the delthyrium. The area on the dorsal valve is not well developed. The outline of the dorsal valve is similar to that of the ventral valve, but the contour differs, in that a fold, bounded by two prominent furrows, occupies the median part of the valve. The ornament of both valves is the same, consisting of radial plications and fine growth lines. The plications are simple on the lateral slopes, but they increase by inter calation and bifurcation in the sinus and on the fold. The sinal formula is-
2 + I + 1 + ly + 1 + I + 2. Dimensions of the Ventral Valve : Length on Curvature Height Width Width of Area 31 26 62 5 33 30 60 7 18 15 35 12 10 24 2
Internal features. -Two strong, vertical, divergent dental lamellae extend from the posterior wall of the valve, for one-third the length of the valve floor. These dental lamellae are wedge-shaped and show no thickening except in the delthyrial cavity, in the older individuals. The umbonal cavities on the lateral sides of the dental plates are sharply contoured. The dental plates merge into the valve floor, extending anteriorly as small ridges which border part of the muscle scar. Under the deltidium, a small vertical ridge projects downward between the dental plates, merging into the posterior part of the muscle scar. The area of muscle attachment is not prominent. The dorsal valve carries a small median ridge, extending from the apex of the valve for one-third of the length of the floor. The crural bases have not been well preserved in any of the specimens examined.
Range. -This species has been found only in the Lambie Series of New South Wales, and appears to be restricted to the Upper Devonian. Remarks.-Cyrtospirifer subdisjunctus occurs in a 15' band of quartzite, strati graphically low in the marine sequence of the Mt. Lambie district. It was stated by Brown and Joplin (1938, p. 220) that this quartzite band contained internal moulds and external casts of Spirifera disjuncta and Rhynchonella pleurodon. Examination of this material has shown that both Cyrtospirifer and Sinospirifer are· represented in this band. The species Cyrtospirifer subdisjunctus is similar to Cyrtospirifer reidi found at Mt. Morgan, but it grows to a much greater size. Also, there is less distortion of the internal features, due to old age thickening, in the former species. It differs from Spirifer disjunctus Sowerby in that it lacks the well marked muscle scar in the ventral valve, and the stronger concentric ornament, seen in Sowerby's figures. The sharp, conical umbonal cavities of Cyrtospirifer subdisjunctus are not found in Spirifer disjunctus Sowerby.
III. CARBONIFEROUS BRACHIOPODS. Introduction : The species described were collected from the Middle Carboniferous beds of Neerkol, Yarrol and Mt. Barney, and include a species of Levipustula gen. nov., one species of N eospirifer, and a species of Spiriferellina. Three species, one belonging to the Orthoidea, and two belonging to the Spiriferidae, also occur in these beds, but poorness of preservation have not permitted their · description. lO UPPER DE VONIA N AND
Stratigrap hical Background : Marine strata of Carboniferous age outcrop at several localities in a belt extending from St. Helens, just south of Townsville, to the New South Wales border. The most complete sequence has been found in the Rockhampton district. The Rock hampton series begins with the Michelinia Limestone, which is regarded as the base of the Tournaisian (Bryan and Jones 1945, p. 33) . Succeeding this limestone is the Protocanites bed, which has been correlated with the Protocanites lyoni zone of Europe. At the top of the Rockhampton series is the Lion Creek lime�tone, which contains a coral fauna suggesting its position as near the top of the VIsean. Succeeding this limestone are the Neerkol beds, which contain the brachiopod fauna, later described in this paper. Out-crops of beds equivalent to the Neerkol beds occur at Yarrol and Mt . Barney. In the Rockhampton district, J. H. Reid believes that the heavy conglomerate at the base of the series indicates a distinct · tectonic break. The conglomerate is succeeded by the indurated mudstones which constitute the greater part of the beds. Carey and Browne (1938) suggest that the Late Visean orogeny which disturbed the sediments along eastern Australia, produced two zones of folding in Queensland, an easterly one of considerable intensity, with an axis running from Rockhampton, through Brisbane and into New South Wales, and a less intense, westerly one in the Drummond Range. In the intervening trough, there was little or no folding, thus accounting for the angular concordance of the Neerkol beds at Stanwell. Lacustrine sedimentation occurred in widely scattered areas to the west of the marine belt. The Drummond Beds, Silver Valley beds, and Pascoe River beds are fresh water sediments of Lower and Middle Carboniferous age. Systematic Descriptions : Order Protremata Beecher 1891. Super-family Strophomenacea Schuchert 1896. Family Productidae Gray 1840. Sub-family Productinae Waagen 1884. Genus Levipustula gen. nov. Genotype.-Levipustula levis sp. nov. .0Jo. F 11900 a, b, c, d. University of Queensland. Middle Carboniferous, �eerkol Series, Por. 5V, Parish of Neerkol, County of Livingstone, Queensland. Diagnosis .-Oval configuration, pustulose surface, irregular growth lines. Dorsal valve plane to geniculate, ventral valve convex, non-sulcate. Adductor muscle scars non-dendritic. Cardinal ridge present in dorsal valve. Brachial impressions indistinct or absent. Area, teeth, and dental plates absent . Remarks.-So far, the only species assigned to this genus is the type-species. The creation of new genera, particularly in the family Productidae, may not be desirable, but an examination of this new form will show that all its characters are common to no known genus. It resembles Pustula in ornament and shape, but the presence of non-dendritic muscle scars is sufficient to exclude it from this genus. The genus Leioproductus Stainbrook 1947, has many characters common to this new form, but differs in being sulcate and possessing, in some cases, an area. Levipustula has been compared with the genus Krotovia Fredericks, specimens of Krotovia spinulosus Sowerby, from the type-locality (Carboniferous Limestone, Linlithgowshire, Scotland), having been examined. The internal structure of this genus could not be studied, but the absence of growth lines on the exterior and its very proportions were sufficient to distinguish the two forms. MIDDLE CA RBONIFEROUS BRA CHIOPODS OF QUEENSLA ND 11
The position of this genus in the family Productidae is still problemati�al. Waagen's original diagnosis of the sub-family Productinae stated that forms belongmg to this group were characterised by dendritic muscle scars. This new ge�us has non-dendritic scars, thus suggesting that it may belong to the Strophalosnn.ae. or Chonetinae. However, the other characters of the genus (absence of areas, restnctwn of spines to the ventral valve, and hinge) distinguish it from these sub-families. Levipustula levis sp. nov. (Pl. II, Figs. 4-13). Holotype.-No. F 11900 a, b, c, d. University of Queensland. Typ e locality.-Portion 5V, Parish of Neerkol, County of Livingstone, Queens land. Synonomy.-Productus brachythaerus (G. Sowerby) Jack and Etheridge 1892, p. 248. Pl. 13, Fig. 5. Productus pustulosus (Phillips) Voisey 1940. pp. 196, 198. Age.-The age of the type locality, up to the present, had not been determined exactly. The association of this species with Neospirifer pristinus sp. nov., and Spiriferellina neerkolensis sp. nov. implies a Moscovian age. Diagnosis.-As for genus. Description, External features.-The outline of the shell is subcircular, the hinge being slightly less than the greatest width. The shell varies from plano convex to concavo-convex and sometimes it is geniculate. The v�nter is often fiat, but never sinuate. The flanks of the ventral valve are fairly steep, but the visceral cavity is never deep. The umbo is strongly incurved, stout and pointed, the umbonal angle varying from 85° to 100°. Area, delthyrium and pseudo-deltidium are absent. Ears are well developed, and are separated from the umbonal slopes by a shallow angle. The ornament consists of spine bases and irregular growth lines. The spine bases are distributed over the whole of the valve, and are arranged in quincunx. They are more abundant on the venter and ears than on the flanks. The spines arose from the anterior end of these ridge-like spine bases, which are more elongate on the venter and flanks than on the ears. The spines were directed obliquely towards the anterior. Spines also occurred along the cardinal margin and those which haw been preserved are very slender and curved. They vary from 4 mm. to 13 mm. in length, the longest ones rising laterally from the ears. The number of spine-bases per centimetre across the venter, measured at a distance of 10 mm. from the apex, varies from 5 to 7. The number of spine bases per centimetre, measured down the venter, varies from 2 to 3. The dorsal valve may be fiat, slightly concave or sharply geniculate. The anterior part of the visceral disc is more depressed than the posterior. The ornament consists of growth lines which are more prominent than those on the ventral valve. S�all pits, corresponding to the spine bases of the other valve, are arranged in qumcunx. Internal features.-In the ventral valve, the features are not very striking. Two very narrow, elongate-triangular adductor scars, separated by a poorly developed median ridge, occur in the posterior of the valve. Sometimes a lateral constriction produces the appearance of two pairs of adductors-posterior and anterior. The scars are slightly raised above the surrounding surface, and extend for one-fifth the length of the valve floor. The diductor muscle scars are two large, striated 1:! UPPER DE VONIEI. N EI.ND
scars, surrounding the adductors, and occupying a quadrate area extending from the umbo for a third of the length of the visceral disc. The area of attachment is raised but is not quite as high as the adductor scars. Usually the muscle scars of the ventral valve are not very distinct, this being due probably, to the extreme thinness of the shell. The rest of the valve contains small depressions corresponding to the spine bases on the exterior. The dorsal valve has a straight hinge line, rising centrally from which is the cardinal process. This is trilobed when viewed dorsally, bilobed when viewed ventrally, and it is never strongly developed. A notch occurs at the base of the cardinal process, this notch being characteristic of most Productids. Rising anteriorly from this notch is a median septum which extends for two-thirds the length of the visceral disc. In many specimens it bifurcates about 3 mm. below the cardinal process, each branch being attached to the base of this process. In some individuals, these two branches give rise to two more ridges, which extend from the cardinal process, for some distance along the lateral margins of the adductor scars. The two latter features have not been developed to the same degree in all the specimens examined. Cardinal ridges are given off laterally from the cardinal process and extend for the two-thirds of the width of the cardinal margin. Brachial impressions have been preserved in one individual only, and in this one they were not prominent. Their lack of preservation may be accounted for by the thinness of the shell. Growth lines and small pustules are distributed over those parts of the valve not occupied by the muscle scars. The adductor scars consist of two non-dendritic, triangular areas, which have been raised above the surrounding surface. They lie on either side of the median septum, the longest side of their triangular area being parallel to this septum. · The scars are thickened anteriorly, gradually disapp�aring posteriorly. The surface of the scars is not entirely smooth, a few small transverse ridges and depressions being present. The length of the adductors varies from one-third to one-quarter the length of the visceral disc. They are approximately three times as long as their maximum width. Dimensions of some individuals : I. Dorsal valve- Length from apex to geniculation 18 20 20 20 23 mm. Width 24 25 25 28 29 mm. Median septum 12 15 13 11 11 mm. Adductor scar 5 X 1.5 6x2 7x2 6 X 1 6x2 mm. II. Ventral Valve- Length 18 21 22 25 27 35 mm. Width 21 27 28 34 35 42 mm. Width/Length 1.16 1.28 1.3 1.35 1.26 1.2 Remarks.--This species shows closest relationship to the Carboniferous forms of Krotovia, Pustula and Leioproductus. Up till the present, it has not been found in beds of known Permo-Carboniferous nor pre-Carboniferous age. In addition to the Queensland specimens, individuals from the New South Wales Carboniferous Burindi Series of Gloucester, Bramble Bay and Drake have been examined. These were collected by A. H. Voisey, M.Sc., and were listed by him (1940, pp. 192-210) as Productus pustulosus Phillips. The New South Wales species is identical with Levipustula levis sp. nov. It does not belong to the genus Pustula, as the muscle scars in this species are non-dendritic, and it lacks the strong, concentric banding, so characteristic of Pustula. MIDDLE CARBONIFEROUS BRA CHIOPODS OF QUEENSLAND 13
Jack and Etheridge (1892, p. 248) described and figured a specimen from Portion 5V, Stanwell, and determined it as Productus brachythaerus G. Sowerby. The author has examined their specimen, and has found that it belongs to the species Levipustula levis. The specimen is neither Productus brachythaerus G. Sowerby, which is a Strophalosiid, nor is it Productus brachythaerus Morris, which is a Terrakea. Fredericks (1931, p. 213) described and figured the species Krotovia tolli from the Kharaulakh Mountains in Russia. His description and figure closely resemble those of Levipustula levis, except that Krotovia tolli has lateral :ridges, slightly curved hinge-line and a quadrifid cardinal process. Fredericks and other authors have used the genus Krotovia to cover a wide variety of forms and it is very doubtful whether all the species which have been assigned to the genus do really belong to it. Order Telotremata Beecher 1891. Super-family Spiriferacea Waagen 1883. Family Spiriferidae King 1864. . Sub-family Elythinae Fredericks 1926.
Diagnosis .-Spirifers characterised by dental lamellae and a small median ridge in the ventral valve. Remarks.-This sub-family was proposed by Fredericks (1926, p. 406) to include the following seven genera :- Eospirifer Schuchert 1908, which is finely striate, Spirifer Sowerby 1815-small costae, C horistites Fischer de Waldheim 1925, Schuchertia Fredericks 1919-with radial plications and superimposed lirae. Neospir�fer Fredericks 1919-fimbriate, lamellose, Elytha Fredericks 1918-plicate, lamellose, and spinose.
Fig. 3 The apical structure of this sub-family was illustrated by Fredericks (1926, p. 422) by two sections through the rostral part of the ventral valve. To this sub-family I have added the genus Trigonotreta Konig (1825, p. 3, fig. 70). A plaster cast of its genotype has been examined, and topotype material has been used to investigate the internal structure. As a result of this investigation, the author has placed this genus in the sub-family Elythinae. Further discussion of its relation to the other genera of this sub-family will be given under the remarks on the genus Neospiri fer Fredericks. Genus Neospirifer Fredericks 1924, emend. Ivanov and Ivanova 1937. Non Neospirifer Nikitin, 1900, p. 385. Neospirifer Fredericks 1924 (1), p. 311. l4 UPPER DE VONIA X AXD
Neospirifer Fredericks 1924 (2), p. 42. Neospirifer Fredericks 1926, p. 406. Neospirifer Paeckelmann 1931, p. 38. Neospirifer Ozaki 1931, p. 24. Neospiri fer Dunbar and Condra 1932, p. 326. Neospirifer emended, Ivanov and Ivanova 1937, pp. 17, 187. Genotype.-Spirifer fa sciger Keyserling. Diagnosis.-Shells subtriangular in outline ; costate and usually fasciculate, with concentric growth lines. Wide, linear, ventral area. Dental lamellae hang pendent as flanges under the palintrope, touching the floor in the apical region. Much thickened in the umbonal region. Heavy callosity between the dental lamellae. Remarks.-The genus Neospirifer was proposed by Fredericks (1924) and he cited (1926) as the type-species, Spirifer Jasciger Keyserling. The use of this species was unfortunate, and has led to the confusion which now exists regarding the genus Seospirifer. \Vhen Keyserling (1846, p. 231, pl. \'III, figs. 3, 3a, 3b) proposed the species Spirifer fasciger, he figured three syntypes, all of which were poorly preserved impressions. Examination of these figures shows that they may all be different forms. Although Fredericks proposed the genus, he did not select a lectotype, and in spite of all my investigations I have not been able to find whether one has since been chosen. �or did he give any real diagnosis of the genus. Dunbar and Condra (1932, p. 326) were the first to do this, but they based their interpretation on the American form, Spirifer cameratus :\lorton. Xo one has described the internal structure of the genotype, Spirifer Jasciger Keyserling, and as topotype material is not available, I have based my interpretation of the genus on Dunbar's and Condra's diagnosis. Ivanov and Ivanova (1937) emended the genus, indicating the difficult problem which had arisen due to Keyserling's choice of poorly preserved type material, and Fredericks' failure to select a lectotype. To overcome these difficulties, they suggested the substitution of the species Spirifer tegulattts Trautschold as the type species. However, this is contrary to the International Rules of Zoological �omen clature, and so their substitution of this species is not valid. The Status of Trigonotreta Konig 1825.-King {1850, p. 126) selected the syntype Trigonotreta stokesii Konig as the type species of this genus, the genotype consisting of an incomplete specimen shov.-ing the external characters of the dorsal valve, the ventral area and the ventral umbonal region. This specimen was collected from beds of Permo-Carboniferous age in Tasmania, and is characterised b\· the strong, linear, ventral area, and the fasciculation. Comparison with Keyserling's fig. 3a, pl. 8, suggests that the two forms may be the same externally. The internal structure of Trigonotreta stokesii has been determined from topotype material, and it has been found that this structure is very similar to that of Spirifer tegulatus as figured by Ivanov and Ivanova (1937), and many of the �orth American species of .Veospi rifer figured by Dunbar and Condra ( 1932). Since the validity of Trigonotreta Konig, as a genus must be accepted, then many of the species formerly assigned to S eospirifer Fredericks will have to be revised, as I beliew that many forms belonging to the former genus ha\·e been erroneously placed in the latter. It is hoped that further research on the genus Trigonotreta will be published in later work on the Queensland Permian brachiopods. Range.-Species of Seospirifer have not been found in beds older than the base of the Moscovian in Russia. The earliest f(Jrms in �orth America occur in the MIDDLE CARBONIFEROUS BRACHIOPODS OF QUEENSLAND 15 lower part of the Pennsylvanian period. Thus the range of the genus may be definedas from the very base of the Middle Carboniferous to the top of the Penman._ "King (1930) described forms from the late Permian beds of the Glass Mountains, Texas. N eospirifer pristinus sp. nov. (Pl. III, Figs. l-8, Pl. IV, Figs. l-4). Holotype-F 2005 a, b, c, d. Queensland Geological Survey. Type locality.-Portion 5V, Parish of Neerkol, County of Livingstone, Queens land. Synonomy.-Choristites aff. mosquensis (Fischer) \Vhitehouse, 1928, p. 441. Age.-The age of the type locality has been determined in this paper as Moscovian. Diagnosis.-Biconvex shell, with radiating costae. Fasciculation is scarcely developed. Two strong teeth are present. Heavy callosity obscures the base of the dental lamellae. Description, External features.-Almost equidimensional in outline, the length being slightly less than the width. The dimensions of t_ypical specimens are :-
Length : 51 40 44 50 50 mm. Width : 54 47 53 52 57 mm. In most cases, the hingeline is slightly less than the greatest width. The ventral valve is strongly convex, with a moderately incurved umbo, and two small ears, separated by a gentle curve from the umbo. The ears are never strongly developed, nor pointed. An elongate-triangular area extends for almost the whole length of the hinge-line. Its continuity is interrupted under the umbo, by a strong triangular delthyrium. The ventral valve carries a fairly prominent sinus, rising from the umbo, and gradually widening anteriorly. This sinus tends to be broad and gentle, rather than deep and angular. The dorsal valve is also strongly convex, but with a smaller umbo than that of the ventral valve, and also smaller ears. A narrow, triangular area is present, this carrying a broad low delthyrium. A median fold rises from the umbo, and broadens in the younger stages. Both valves are ornamented with small radiating costae, the fold and sinus carrying the same ornament as the lateral slopes. The costae are low and rounded, the intervening grooves being sharp and narrow. The number of costae per centimetre, measured at a distance of one centimetre from the umbo, varies from 6 to 8. The number of concentric lamellae per centimetre on the ears averages 8 to 10, although they are less abundant on other parts of the shell. Internal features.-In the ventral valve, two strong teeth rise from the comers formed on the hinge-line by the delfhyrium. They project dorsally and anteriorly, and merge into the dental lamellae, which slope posteriorly towards the floor of the valve. The junction of the dental lamellae with the valve floor is obscured by callosity. A gently curved surface is formed at the junction of these lamellae with the underside of the area, giving the impression that they grew downward from the hinge, and anteriorly from the posterior wall . ..-\ well marked groove, between the dorsal end of the lamellae and the cardinal area, was probably produced by the deposition of adventitious material about the dental lamellae. This groove is more 16 L.PP ER DE I.ONIAN AND prominent in the older forms. Muscle scars occur on a raised portion of the valve, extending along the floor on the valve for half its length. This area is bounded by two sharp ridges which merge anteriorly into the fold, corresponding to the' sinus on the exterior. The adductor scars are two, verv faint, narrow, elongate markings, occurring on either side of a minute median ridge, and surrounded by two large, striated diductor scars. The walls of the valve in the umbonal cavities are pitted. This feature is found in many of the Permo-Carboniferous Spirifers. The rest of the valve is ornamented with small, radial costae and growth lamellae . .-\n apical callosity is always present, and extends from the anterior ends of the dental lamellae into the umbo. It is usually fairly large, obscuring part of the muscle scar and the base of the dental lamellae. The great increase in the size of this callosity, the thickening of the dental lamellae, and the elevation of the area of muscle attachment seem to be old age tendencies. In the dorsal valve, the characteristic features are the wide delthyrium, the narrow triangular area, and the well developed crural bases, from which the spiralium diverges. On the inner sides of the crural bases are the two dental sockets. Between these is a raised, striated, cardinal process to which the diductor muscles were attached. Anterior to the cardinal process there is a hollow, out of which rises a fine median ridge, which separates the adductor scars. These scars are small, rounded, and smooth, and occupy a slightly depressed area. Radial markings and concentric growth lines cover the rest of the interior. Remarks.-The main characteristic of the genus Trigonotreta is its strong fasciculation. This may or may not be sufficient to separate the genus from .\'eospi rifer, but further work on the Australian Permo-Carboniferous spiriferids is necessary before any decision can be made. Since the genus N eospirifer has been used to include other non-fasciculate forms, and forms in which fasciculation is poorly developed, I have placed the species described above in this genus rather than in Trigonotreta. Internally this species shows close resemblance to the species N eospirifer tegulatus Trautschold, which Ivanov and I vanova (19il7) described as the type species of N eospirifer. Externally, N eospirifer pristinus lacks the fascicula tion common to most members of the genus, although in some specimens the tendency towards fasciculation may be discerned. Examination of species of .Veospiri fer figured by Dun bar and Condra ( 1932), Ivanov and I van ova ( 1937), and Muir-Wood and Oakley (1941) shows that there is less variation in the internal than in the external characters. The early forms of N eospirifer from the Pennsylvanian, figured by Dunbar and Condra, were non-fasciculate, but the tendency towards fasciculation became more pronounced in later species. Another trend in the genus is the development of stronger dental lamellae. Changes seen in the dental lamellae of different species are of degree and not kind. In N eospirijer pristinus, both these trends are indistinct, and it seems reasonable to conclude that this is one of the early species of the Middle Carboniferous. Range.-On comparison with other species, N eospirifer pristinus shows closest resemblance to Neospirifer cam ratus Morton (DlJnbar & _ � Condra 1932, p. 327). It resembles the latter species, m ornament, contour and dimensions, but in this American species, fasciculation is developed. N eospirifer cameratus Morton occurs in the Vanport Limestone of the lower Allegheny formation at Putnam Hill, Ohio. The Allegheny formation is equivalent to part of the Des Moines Series which Dunbar (1940) correlated with the Moscovian of Russia. Sub-family Delthyrinae Fredericks 1926, p. 406. Diagnosis.-Spirifers characterised by a median septum, and dental lamellae which slope laterally from the delthyrium to the valve floor. MIDDLE CA RBONIFEROUS BRA CHIOPODS OF QUEENSLA ND 17
Remarks.-The apical structure is illustrated by a section through the posterior part of the valve.
Fig. 4
This sub-family includes nine genera, both punctate and impunctate, each being characterised by distinctive ornament. These genera are :- Munia Fredericks, 1918-smooth. Spiriferellina Fredericks, 1919-radial folds. Spirijerinaella Fredericks, 1926-radial folds and concentric lines. Lytha Fredericks, 1919-costate. Reticularia M'Coy, 1844-spinose and lamellose. Delthyris Dalman, 1828-lamellose, finely striate and fimbriate. Reticulariina Fredericks, 1916-plicate, spinose and fimbriate. Maia Fredericks, 1919-lamellose and strongly plicate. Spiriferella Tschernyschew 1902-striated costae, arranged in bundles and a fimbriate border. To this sub-family may be added the genus Punctospirife r North 1920. It differs from Spirijerellina Fredericks in its stronger lamellosity and its more numerous plications.
Genus Spirijerellina Fredericks 1924. Spiriferina d'Orbiginy (partim), 1847, p. 268. Spiriferina d'Orbiginy (partim), 1850, p. 334. Spirijerellina Fredericks, 1924, {I), p. 299. Spirijerellina Fredericks, 1924, (2), p. 33. Spiriferellina Fredericks, 1926, p. 407. Spirijerellina Fredericks, 1927, pl. I, fig. 8. Spirijerellina Paeckelmann 1931, p. 50. Genotype (by original designation) .-Terebratulites cristatus Schlotheim. Diagnosis.-Shells small, punctate, with prominent median sinus and fold, with three to six lateral plications on either side of the median fold and sinus. Two small dental lamellae and a median ridge formed by a tight folding of the shell floor, occur in the ventral valve.
Remarks.-This genus was proposed by Fredericks in 1924 (pp. 289-299) and, as stated by Muir-Wood and Oakley (1941, p. 8), it is equal in part to Spiriferina of Tschernyschew (1902), Kozlowski (1914, p. 71, fig. 17), and to Trigonotreta Fredericks (1916). Fredericks (1927) figured a transverse section of Spirijerellina I l:l UPP ER DE VONIA N AND cristata to indicate the difference in internal structure from that of Spiriferina. This latter genus has dental lamellae, a delthyrial plate and a pseudo-septum formed of two distinct fused plates-outgrowths of the delthyrial plate. Spiriferellina has no delthyrial plate, and has a median septum in addition to the dental lamellae. Spiriferellina differs from Punctospirifer North, in that the latter genus �as more numerous and less angular plications, and a more flattened, broader median fold and shallower sinus. The lamellose ornament of Spiriferellina is less pronounced. Range.-The genus Spiriferellina has been recorded from beds of Mi�dle and Upper Carboniferous and Permian age. Branson (1948, p. 511),. has h�ted the localities of Permian age from which various authors have descnbed th1s genus.
Spiriferellina neerkolensis sp. nov.
(Plate IV. figs. 5-9). Holotype.-F 11926 a, b. University of Queensland.
Type locality. - Portion 5V, Parish of Neerkol, County of Livingstone, Queens land.
Age.-The age of the type locality has been determined in this paper as Moscovian.
Diagnosis. -Biconvex shell with radiating plications, median fold and sinus. Cardinal extremities slightly rounded. Concentric growth lamellae present. Median septum and dental plates in the ventral valve ; small crural plates and faint median ridge in the dorsal valve.
Description. External features.-The shell is wider than long, the greatest width occurring just anteriorly to the hinge-line. The shell is biconvex, and the ventral umbo is slightly incurved. A strong median sinus, with three to fivelateral plications on each side, occurs on the ventral valve, a corresponding fold and plications being present on the dorsal valve. These widen anteriorly, producing a sen;ated anterior margin . The plications become progressively weaker laterally, the two sinal plications being, by far, the most prominent. The concentric growth lamellae are not strongly developed, but can be seen on both valves. Both valves are finely punctate. Dimensions Ventral Valve Dorsal Valve
Length 11.5 1 3 . 8 11.0 12.8 mm. Width 14.0 21.2 16.6 18.3 mm.
Internal features.-Arising from the floor of the ventral valve, in the umbonal region, are two small, slightly divergent dental lamellae, and a slightly stronger median septum, which extends for a third of the length of the valve floor. Small indistinct muscle scars occur on either side of this septum. No apical callosity is developed, the delthyrial cavity being quite sharply contoured.
In the dorsal valve, the divergent crural processes are supported by two small vertical crural plates which diverge from the base of a minute, striated cardinal process. Between these crural plates a very low median ridge extends along the floor of the valve for a distance of several mms. MIDDLE CA RBONIFEROUS BRA CHIOPODS OF QUEENSLA ND 19
Range.-This species, so far, has been found only in the Neerkol beds in Queensland, and these are of Moscovian age. The species resembles the American, short-hinged form of Punctospirifer kentuckyensis Shumard (Dunbar and Condra 1932), which occurs in the Missouri Series of the Pennsylvanian. The Missouri Series has been correlated, by Dunbar (1940) , with the Triticites Zone in the Russian sequence, and therefore it is equivalent to the Russian Upper Carboniferous, over lying the Moscovian beds. This short-hinged form which Dunbar and Condra have placed in the species Punctospirifer kentuckyensis Sl)umard has many of the external features·characteristic of Spirijerellina, and may possibly belong to this latter genus. Remarks.-An undetermined species of Spirijerellina has been collected from beds of Lower Carboniferous age in the Rockhampton district (locality Fairy Bower) . This form differs from the younger Spiriferellina neerkolensis, in that it possesses more numerous, smaller plications. However, it has an outline and contour similar to that of the Neerkol form, and may, possibly, be ancestral to it. Species of Spirijerellina are abundant in the Permo-Carboniferous of Queensland, one particular species from the Ingelara stage having similar proportions to the Neerkol form, but differing in the possession of one less lateral plication. De Koninck (1895) and Dun (1902) have both described specimens which they have determined as Spirijerellina cristata Schlotheim, from the Clarencetown district of New South Wales. These were collected from beds of Tournaisian age, and although I have not examined de Koninck's nor Dun's original material, specimens collected from their original localities have been investigated. These New South Wales forms are considerably different from the Neerkol species, differing in outline, contour, angularity of plications and degree of lamellosity.
Spirijerellina neerkolensis differs markedly from -the Western Australian forms of Spirijerellina cristata, both internally and externally. The Western Australian form from the Wandagee beds is strongly lamellose, and the plications are stronger and more angular. In addition, the internal plates are more heavily developed.
IV. CONCLUSION AND ACKNOWLEDGMENTS. A Research grant, held wit�in the University of Queensland, and financed by Commonwealth funds, has enabled this work to be carried out. I am indebted to the Queensland Geological Survey, and the Department of Geology, University of Queensland, for the opportunity of studying the material on which this investiga tion has been based. Valuable information has been supplied by Dr. S. M. K. Henderson, Director of Museums, Glasgow ; Mr. W. Hood, Glasgow ; Mr. W. N. Edwards and Dr. H. M. Muir-Wood, British Museum (Nat. Hist.) ; Professor T. N. George, University of Glasgow ; Dr. A. B. Walkom, Australian Museum, Sydney ; Mr. E. D. Gill, National Museum, Melbourne ; Dr. C. Teichert, University of Melbourne ; and Miss E. Archer, C.S.I.R.O., Melbourne ; all of whom I wish to thank.
I am also grateful to Dr. F. W Whitehouse and Mr. F S. Colliver for their assistance and advice during the course of this work, and to :\hss M. McCarthy and Miss D. Watson for clerical assistance. Photographs were taken by Mr. Thompson of the University of Queensland. I wish to thank Mr. and Mrs. Hardgrave for the kind hospitality which they extended to me during my collecting trips at Mt. Barney. Invaluable assistance and guidance was afforded by Dr. D. Hill throughout the course of the investigations. 20 UPPER DE VONIA N AND
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BRANSON, C. C., 1948. "Bibliographic Index of Permian Invertebrates." Mem. geol. Soc. Amer. , XXVI, pp. 1- 1049. BROWN, I. A., 1944. "Stringocephalid brachiopoda in Eastern Australia." ]. roy. Soc. N.S. W., LXXVII. pp. 119-129, pis. IV-V BROWN, I. A., AND joPLIN, G.A., 1938. "Upper Devonian sediments at Mt. Lambie, N.S.W.'' Proc. Linn. Snc. N.S. W . . LXIII, Pts. 3-4, pp. 219-223. H., BRYAN, W. AND JoNES. 0. A . . 1 944. "A revised glossary of Queensland stratigraphy." Publ. Univ. Qd. Dept. of Ceo!. (N.S.) II, No. 11, pp. 1 -77. ------, 1945. "The geological history of Queensland : a stratigraphic outline." Publ. Univ. Qd. , Dept. of Geol. (N.S.) II, No. 12, pp. 1 - 103. CAREY, S. W., AND BROWNE. W. R., 1938. " Review of the Carboniferous stratigraphy, tectonics and palaeogeography of New South Wales and Queensland." ]. roy. Soc. N.S. W., LXXI, pp. ;)91-614. CLOUD, P. E., 1942. "Terebratuloid brachiopoda of the Silurian and Devonian." Spec. Pap. geol. Soc. Amer. , XXXVIII, pp. 1-182, pis. I-XXVI. CooPER, G. A., 1942. "Correlation of the Devonian sedimentary formations of North America." Bull. geol. Soc. Amer. , LIII, pp. 1729-1 793, 1 pl ., 1 text fig. DAVIDSON, T., 1860. "The Carboniferous system in Scotland, characterised by its brachiopoda." The Geologist, IV, p. 182. pl. III, figs. 23-24. ------, 1863. " Monograph of British fossil brachiopoda." Palaeontogr. Soc. II, Pt. 5, No. 4, pp. 1-280, pis. I-LV. ------, 1865. "Monograph of British fossil brachiopoda : the Devonian brachiopoda.'' Palaeontogr Soc. III, Pt. 6, pp. 1 - 128, pl. 1-20. DEMANET, F., 1938. "La faune des couches de passage du Dinantien au Namurien dans le synclinorium de Dinant." Mmz . Mus. Hist. nat. Belg. LXXXIV, pp. 1-201, pl. I-XIV. DuN, W. S., 1900. "Carboniferous brachiopods from Clarence Town." Rec. geol. Surv. N.S.W., VII, Pt. I, pp. 72-88. pl. XXI-XXIII. , 1 904. "Notes on some new species of Palaeozoic brachiopoda from New South Wales." Rec. geol. Surv. N.S. W., VII, Pt. 4, pp. 318-325, pl. LX-LXI. DUNBAR, C. 0., 1940. "Type Permian : Its Correlation." Bull. Amer. Ass. Petrol. Geoi. XXIV, Pt. 1, pp. 237-281 DuNBAR, C. 0. , AND CoNDRA, G. E., 1932. " Brachiopoda of the Pennsylvanian system in Nebraska." Bull. geol. Surv. Neb. V . . (Second series) , pp. 1-377, pls. I-XLIV. FENTON, C. L.. 1931. "Studies of evolution in .the genus Spirifer." Pub. Wag. Inst. Sci. II, pp. 1-435, pis. 1-L. FENTON, C. L., AND FENTON :\1. .\ . 1924. " Hackberry stage of the Upper Devonian." Pub. Unw. Mtch. Mus. geol. I. pp. 1-204, pis. I-XLV. FREDERICKS, G., 1916. "Palaeontological notes. 2. On some Upper Palaeozoic brachiopoda of Eurasia." Mem. Com. geol., St. Petersb. CLVI, pp. l-87, pls. I-XXIV. -----, 1924. ( 1) . " Etudes palaeonto1ogiques. 2. Sur. les Spiriferides du Carbonifere superieur de I'Oural." Bull. Com. geol. St. Petersb. XXXVIII, No. 3, pp. 295-324. (not seen). ------. 1 924. (2) . "Upper Palaeozoic of Ussuriland. I. Brachiopoda." Rec. geol. Com. Russ. Far East No. 28. pp. l-52, 1 pl. (not seen).
. , 1926. "Synopsis of the genera of the family Spiriferidae King.'' Bull. Acad. Set. URSS XX, 5-6, pp. 393-423. ------. 1928. "Contribution to the classification of the genus Productus." Bull. Com. geol. St. Petersb. XLVI, No. 7, pp. 773-792. MIDDLE CARBONIFEROUS BRACHIOPODS OF QUEENSLAND 21
------, 1931. "The Upper Palaeozoic fauna of the Kharaulakh Mountains." Bull. Acad. Sci. U.R.S.S. (series 7) , pp. 119-228, pis. I-IV. FoWLER, A., 1944. "A deep bore in the Cleveland Hills." Geol. Mag., Lond. , 1944, pp. 254-266 193-206. GossELET, J., 1894. ".Etude sur les variations du Spirifer verneuili." Mem. Soc. geol. Nord. , IV, Pt. 1, pp. 1-51, pis. I-VII. GRABAU, A. W., 1931. (1). "Devonian brachiopoda of China. I. Devonian brachiopoda from Yunnan and other districts in South China." Palaecont. sinica, B, III, Fasc. 3, pp. 1-545, pis. I-LIV.
------, 1931. (2) . "The significance of the sinal formula in Devonian and post-Devonian Spirifers." Geol. Soc. China, XI, No. 1, pp. 93-96, pis. I-II. HALL, j., 1867. "Natural History of New York. Palaeontology.'' IV, Pt. 1, pp. 1-422, pis. I-LXIII.
----- , 1894. "Natural History of New York. Palaeontology." VIII, Pt. 2, pp. 1-394, pis. I-LXXXIV. HILL, D., 1938-41. "A Monograph on the Carboniferous rugose corals of Scotland. Faunal and stratigraphical section." Palaeontogr. Soc. (Monogr ), pp. 1-213, pis. I-XI.
------, 1942. "The Middle Devonian rugosecorals of Queensland. III. Burdekin Downs, Fanning River and Reid Gap, North Queensland.'' Proc. roy. Soc. Qd. LUI, No. 14, pp. 229-268, pis. V-XI. -----, 1943. "A re-interpretation of the Australian Palaeozoic record, based on a study of the rugose corals.'' Proc. roy .. Soc. Qd. I-IV, No. 6, pp. 53-66. HUDSON, R. G. S., 1945. "The goniatite zones of the Namurian." Geol. Mag. Lond. 1945, pp. 1-10. HuDSON, R. G. S., AND CoTTON, G., 1946. "The Carboniferous rocks of the Edale anticline, Derbyshire." Quart. ]. geol. Soc. Lond. CI, pp. 1-36, pl. I. IVANOV, A. P., AND IVANOVA, E. A., 1937. "Brachiopoda of the middle and upper Carboniferous of the Moscow basin (Neospirifer and Choristites)." Trav. Inst. paleozool. Acad. Sci. U.S.S.R. VI , livr. 2, pp. 1-213, pis. I-XXIII. jACK, R. L., 1879. "The geology and mineral resources of the district between Charters Towers and the coast." Publ. geol. Surv. Qd. I, pp. 1-28. jACK, R. L., AND ETHERIDGE, R., 1892. "The geology and palaeontology of Queensland and New Guinea." Publ. geol. Surv. Qd. XCII, pp. 1-756, pis. I-LXVIII. KEYSERLING, A. voN, 1896. "Wissenschaftliche Beobachtungden auf einer Reise in das Petschora-Land." St. Petersburg, pp. l-465, pis. I-XXII (not seen). KING, R. E., 1930. "The geology of the Glass Mountains, Texas." Bull. Univ. Tex . Sci. Ser. 3042, pp. l-245, pis. I-XLIV. KING, W. , 1850. "A monograph of the Permian fossils of England." Palaeontogr. Soc. (Monogr.) , pp. 1-258, pis. I-XXVIII.
KozLoWSKI, R., 1914. "Les brachiopodes du Carbonifere superieur de Bolivie." Ann. Paleont. IX, pp. 1- 100, pis. I-XI.
LEGOINTRE, G., AND DELEPINE, G. "Etudes geologiques dans Ia Region Paleozoique comprise entre Rabat et Tiflet." Notes Serv. Min. Maroc., XXVIII, p. 69, pl. XIV.
MACGREGOR, M., 1931. "Scottish Carboniferous stratigraphy : An introduction to the study of the Carboniferous rocks of Scotland.'' Trans. geol. Soc. Glasg. XVIII, pp. 442-558.
MAXWELL, W. G. H. , 1950. "An Upper Devonian brachiopod (Cyrtospirifer reidi sp. nov.); from the Mt. Morgan District." Publ. Univ. Qd. Dept. of Geol., III, 12, pp. 1-8, pl. I.
MILORADOVITCH, B., 1937. "Morphogenesis of the ventral valve in Spiriferidae." Prob. Palaeontology , II-III, pp. 501-535.
MooRE, R. C., 1944. "Pennsylvanian formations." Bull. geol. Soc. Amer. LV, Pt. 1, pp. 657-706.
MuiR-Wooo, H. M. , 1927. Ann. Mag. Nat. Hist. XIX, p. 289 (not seen).
------, 1928. "The British Carboniferous Producti II." Mem. geol. Surv. U.K. Palaeontology . III. Pt. l, pp. 1-217 , 12 plates.
------, 1930. "The classification of the British Carboniferous brachiopod sub-family Productinae." Ann. Mag. Nat. Hist. (10), V, pp. 100- 108. 22 UPPER DE VONIA N AND
------, 1948. "Malayan lower Carboniferous fossils and their bearing on the Visean palaeogeography of Asia." British Museum (Nat. Hist.), June, 1948.
MUIR-WooD, H. :.\1., AND OAKLEY, K. P., 1941. "Upper Palaeozoic faunas of North Sikkim." Palaeont. indica. (N.S.), XXXI, 1, pp. l-91, pis. I-IV. MuRCHISON, 1840. Bull Soc. geol. Fr. XI, p. 252 (not seen) . NIKITIN, 1900. Bull. Com. geol. St. Petersb. XVIII, p. 385 (not seen). OzAKI, K., 1931. "Upper Carboniferous brachiopods from North China." Bull Shnnf!.hm Sci. Inst. I, No. 6, pp. 1-205, 15 plates (not seen) . OBRUTSCHEW, W. A., 1926. "Geologie von Siberien." Berlin.
PAECKELMANN, W., 1930. "Die brachiopoden des deutschen Unterkarbons I." Abh. preuss. geol. Landesanst. CXXII, pp. 1-326, 24 plates (not seen) .
------. 1931. " \'ersuch einer zusammenfassenden systemati:-: d.::i Spiriferidae." N . .Jb. Min. Geol. Palaont., B.B., LXVII, B, pp. 1-64 (not seen). PAUL, H., 1937. "Die transgression der Visestufe am Nordrande des Rheinischen Schieferge birges." Abh. preuss. geol. Landesanst. Neu folge CLXXIX, pp. 1-116, 3 plates. PRENDERGAST, K. L., 1943. "Permian Productinae and Strophalosiinae of Western Australia." ]. roy. Soc. W. Aust. , 1941-2, XXVIII, pp. 1-62, 6 plates. REED, F. R. C., 1944. "Brachiopoda and Mollusca from the Productus Limestones of the Salt range." Palaeont. indica (N.S.) XXIII, 2, pp. 1-678, pls. I-LXV. REID, J. H., 1930. "The Queensland Upper Palaeozoic succession." Pub. geol. Surv. Qd. 278, 96 pages, 9 plates. REID, J. H., AND MORTON, C. C., 1928. "Central Queensland Geological Section." Qd. Govt. Min. ]. No. 29, pp. 384-389.
SEMICHATOV, S. V., 1934. "Carboniferous fauna from the Don and Medveditza Rivers. I. Productinae." Trans. geol. prosp. serv . U.S.S.R. Fasc. 260, pp. 1-52, 3 plates.
ScHUCHERT, C., AND LE VENE, C., 1929. "Fossilium Catalogus. I. Animalia." Pt . ..J.2, Brachiopoda, pp. 1-140. SHIMER, H. W., AND SHROCK, R. R., 1944. "Index fossils of North America." Chapman and Hall, Ltd. , London. SoKOLSKAIA, A. N., 1941. "Lower Carboniferous and Devonian-Carboniferous brachiopods of the Moscow basin. (Tschernyschino, Upa and Malevka-Murajevnia Beds) , Pt. I. Spiriferidae." Trav. Inst. Paleont., Moscow, XII, 2, pp. 1 - 1 39, 12 plates (not seen) .
SoWERBY, J . , 1914. "Mineral Conchology " I. STAINBROOK, M. A., 1945. "Brachiopoda of the Independence Shale, Iowa." Mem. geol. Soc. Amer. XIV, pp. 1-74, 6 plates. TEICHERT, C., 1941. "Upper Devonian goniatite succession of Western Australia." Amer. ]. Sci. CCXXXIX, pp. 148-153.
------, 1943. "The Devonian of Western Australia. A preliminary review. Parts I and II." Amer. ]. Sci. CCXLI, pp. 69-94 and 167-184. THOMAS, I., 1 914. "The British Carboniferous Producti. I." Mem. geol. Surv. U.K . Palaeontology . Pt. 4, pp. 199-364, 4 plates. THOMAS, H. D., 193/'i. " The brachiopod Punctospirifer pulchra (Meek) ." Amer. Midl. Nat. XVI, pp. 203-207, pl. VII (not seen). TIEN, C. C., 1938. "Devonian brachiopoda of Hunnan." Palaeont. sinica (N.S.) B, 4 (whole series) 113, pp. 1-147, pis. I-XXII. TscHERNYSCHEW, T., 1902. "Die Obercarbonischen Brachiopoden des Urals und des Timan." Mem. Com. feol. Russe XVI, 2, pp. 1-749 (not seen) , 63 plates. VoiSEY, A. H., 1936. "The upper Palaeozoic rocks in the neighbourhood of Boorook and Drake, N.S.W." Proc. Linn. Soc. N.S. W LXI, Pt. 3-4, pp. 155-168, pl. VIII. ------, 1940. "The upper Palaeozoic rocks in the country between the Manning and Karuah Rivers, New South Wales." Proc. Linn. Soc. N.S.W. LXV, pp. 1 92-2 10, 1 pl. ------. 1945. "Correlation of some Carboniferous sections in New South Wales." Proc. Linn. Soc. N.S. W. LXX, pp. 34-40, pl. IV-V. WAAGEN, W., 1882-4. "Salt Range Fossils." Palaeont. indica. Ser. LXIII, IV, Fasc. I, 2, 4 (not seen) . MIDDLE CARBONIFEROUS BRACHIOPODS OF QUEENSLAND 23
WHIDBORNE, G. F. , 1896-1907. " A Monograph of the Devonian Fauna of the South of England. Vol. III." Palaeontogr. Soc. pp . 1-247, 38 plates. WHITEHOUSE, F. W., 1927. "Notes on Palaeozoic fossils from the Cannindah District." Qd. Govt. Min. ]. XXVIII, p. 189. 441-442. ----- , 1928. "Central Queensland Geology." Qd. Govt. Min. ]. XXIX, pp. Wyatt ------, 1929. "Preliminary report on Devonian fossils from the districts of Mt. and Ukalunda. " Qd. Govt. Min. ]. XXX, pp. 158-159.
VI. EXPLANATION OF PLATES.
Except where otherwise noted , the specimens are all in the University of Queensland collection. All figures approximately natural size.
PLATE I.
Sinospirifer sinensis Grabau var. australis var. nov.
Fig. l. F 11965. 4t miles S.E. of Dotswood Homestead, Parish of St. Giles. Upper Devonian. Plasticine model of ventral valve.
Fig. 2. F 11966. ibid.
Fig. 3. F 11964. ibid.
Fig. 4. F 2009 a. Queensland Geological Survey Collection. Corner Creek, Star River Basin. Upper Devonian. Plasticine model of the ventral valve.
Fig. 5. F 20096. Queensland Geological Survey Collection. Corner Creek, Star River Basin . Upper Devonian. Plasticine model of the ventral valve.
Fig. 6. F 2007 a. Queensland Geological Survey Collection. Corner Creek, Star River Basin. Upper Devonian. Plasticine model, sho.wing the internal features of the ventral valve.
Fig. 7. F 11928. 4! miles S.E. of Dotswood Homestead, Parish of St. Giles. Upper Devonian. Plasticine model showing the internal features of the ventral valve.
Fig. 8. F 2007 b. Queensland Geological Survey Collection. Corner Creek, Star River Basin. Upper Devonian. Plasticine model of interior of ventral valve.
Fig. 9. F 11949. Solitary Creek, Bathurst , N.S.W. Upper Devonian. a and b. Internal views of the ventral valve.
Fig. 10. F 2008. Queensland Geological Survey Collection. Corner Creek, Star River Basin . Upper Devonian. a. Plasticine model. b. Internal cast of ventral valve.
Fig. ll. F 11927, F 11928. 4! miles S.E. of Dotswood Homestead . Parish of St. Giles. Upper Devonian. F 11928 is the Holotype. = "b" in figure 11a. 11b=internal plasticine model of ventral valve of F 11927.
Fig. 12. F 11929-30. 4! Iniles S.E. of Dotswood Homestead. Internal moulds of the ventral valve.
Fig. 13. F 11963-11966. 4! miles S.E. of Dotswood Homestead. Upper Devonian. Rock specimen containing internal moulds and external casts. 24 UPPER DE VONIA N AND
Fig. 14. F 2010. Queensland Geological Survey Collection. Corner Creek, Star River Basin. Upper Devonian. Internal mould of ventral valve .
Fig. 15. � 2009. ibid.
Fig. 16. F 2007 a, b. ibid.
Cyrtospirife r subdisiunctus sp. nov.
Fig. 17. F ll950. Holotype. Solitary Creek, near Bathurst, N.S.W. Upper Devonian. a. Internal cast of ventral valve. b, c. Plasticine models of ventral interior.
PLATE II.
Sinospirifer sinensis Grabau var. australis var. nov.
Fig. I. F ll944. Solitary Creek, near Bathurst, N . S.W. Upper Devonian. Internal mould of dorsal valve.
Fig. 2. F ll934-4l. 4! miles S.E. of Dotswood Homestead , Parish of St. Giles, County of Nares. Upper Devonian. Rock specimen containing external casts of the dorsal and ventral valves.
Fig. 3. F ll943. Solitary Creek, near Bathurst, N.S.W. Upper Devonian. Internal mould of dorsal valve.
Levispustula levis sp. nov.
Fig. 4. F ll900 Holotype. Portion 5V, Parish of Neerkol, County of Livingstone, Queens- land. Moscovian. a. Plasticine model of the ventral valve (external) . b. Plasticine model of the dorsal valve (external) . c. Internal mould of the ventral valve. d. Plasticine model of the ventral valve (internal) . e. Plasticine model of the dorsal valve (internal).
Fig. 5. F ll909. Paratype. Portion 5V, Parish of Neerkol.. Moscovian. Plasticine model of external of ventral valve.
Fig. 6. F ll90l. Paratype. Portion 5V, Parish of Neerkol. Moscovian. External cast and plasticine model of ventral valve , showing the elongated nature of the spine bases.
Fig. 7. F ll967. Paratype. Portion 5V, Parish of Neerkol. Moscovian. External cast of the ventral valve.
Fig. 8. F ll913 b. Portion 200, Parish of Palen , County of Ward . Moscovian. Plasticine model of the internal surface of the ventral valve, showing the muscle scars.
Fig. 9. F ll908. Portion II, Parish of Baywulla , County of Yarrol. Moscovian. External cast of the ventral valve.
Fig. 10. F 11915. Portion 198, Parish of Palen , County of Ward. Moscovian. Plasticine model of the internal surface of the dorsal valve, showing Cardinal ridge, bifurcating median septum, and smooth adductor scars.
Fig. 11. F 11906. Portion 2162, Parish of Neerkol. County of Livingstone. Moscovian. a. Internal mould of dorsal valve. b. Plasticine model.
Fig. 12. F ll914. Portion 200, Parish of Palen, County of Ward. Moscovian. Internal mould of dorsal valve.
Fig. 13. F 13120. Portion 5V, Parish of Neerkol. Moscovian, External mould of ventral valve. JIIDDLE CA RBONIFEROUS BRA CHIOPODS OF QUEEJ.VSLA ND 25
PLATE III. Neospirzfer pnstinus sp. nov. Fig. l. F 1Hil6. PGrtion 5V, Parish of Neerkol. County of Livingstone. Moscovian. Internal mould of the ventral valve, showing the strong teeth, triangular area and delthyrium, poorly developed dental plates. and strong area of muscle attachment. (Plc:..,ticine model.)
Fig. 2. F 11920. Portion 5V, Parish of Neerko) , County of Livingstone. Moscovian. Umbonal region of the interior of the ventral valve (plasticine model). Fig. 3. F 11921. Portion 5V, Parish of Neerkol. Moscovian. Plasticine model, showing the external ornament of the ventral valve. Fig. 4. F 2005. Holotype. Queensland Geological Survey Collection. Portion 5V. Parish of Neerkol. Plasticine model, showing the internal structure of the ventral valve.
Fig. <� . F 2006. Queensland Geological Survey Collection. Portion 5V, Parish of Neerkol Plasticine model, showing a flattened dorsal valve, and the posterior region of the ventral valve. Fig. 6. F 11916. Portion 5V, Parish of Neerkol. Moscovian. Internal mould of the ventral valve.
Fig. i. F 11921. Portion 5V, Parish of Neerkol. Moscovian. External cast of ventral valve. Fi�. 8. F 11919. Portion 5V, Parish of Neerkol. Moscovian. External cast of the ventral valve.
PLATE n·. Neospir�fer pristinus sp. nov. Fig. l. F 11920. Portion 5V, Parish of Neerkol. Moscovian. Internal mould of the rostral part of the ventral valve. Fig. 2. F 11917. Portion 5V, Parish of Neerkol. Moscovian. Plasticine model of the dorsal valve. Fig. 3. F 11917. Portion 5V, Parish of Neerkol. Moscovian. Internal mould of the dorsal valve.
Fig. 4. F 11918. Portion 5V, Parish of Neerkol. Moscovian. Internal mould of the rostral part of the ventral valve. Spiriferellina neerkolensis sp. nov. Fig. 5. F 11926. Holotype. Portion 5V, Parish of Neerkol. Moscovian. a. External mould of ventral valve (magnified by lt). b. Internal mould of ventral valve. c. Plasticine model of interior of ventral valve (magnified by li)- d. Plasticine model of external of ventral valve (magnified by It). Fig. 6. F 13121. Portion 5V, Parish of Neerkol. Moscovian. a. Plasticine model of interior of dorsal valve. b. Internal mould of dorsal valve. Fig. 8. F 13121. Portion 5V, Parish of Keerkol. :\Ioscovian. Internal mould of dorsal valve. Fig. 9. F 13123. Portion 5V, Parish of Neerkol. !\Ioscovian. External mould of dorsal valve. Spir1jerellina sp.
Fig. i. F 13122. Portion 602, Parish of Neerkol, County of Livingstone. Lower Carboniferous. a. Plasticine model of external of ventral valve (magnified by It). b. External mould of ventral valve. This specimen is figured in order to show the slight difference between it and the Moscovian species. Sf>iriferellina nel'rkolensis sp nov .. �6 UPPER DE VONIA N .-J ND
DEVONIAN � Lac ustr1ne � Geos':JnclJn aiiiTIJ]] M«rtne ro ssds Q L&Jcuslr1ne Fo sst!s D
Both C
0
l ---
I I
,_- ..... - - .J
::YlAPOF QUEENSLAND SHOWING LOCALITIES WHERE DEVONIAN BEDS HAVE BEE:: Localities.-!. Chillagoe. 2. Clarke River. 3. Star River. 4. Burdekin Downs. .) . Fanning River and Dotswood. 6. Reid Gap. 7. Ukalunda and Mt. Wyatt. 8. Grasstree. 9. Douglas and Drummond Creeks. 10. Hunter and Marble Is. 11. Mt. Etna. 12. Morinish. 13. Cawarral. 14. l'lam and Marmor. 15. :vit. Morgan. 16. Raglan. 17. Mt. Larcom. 18. Kroombit. 19. \'arrol. 20. Silverwood. 21. Gilberton. 22. Drummond Range. .1IIDDLE CA RBONIFEROUS BRA CHIOPODS OF QL'EENSLA ND 27 CARBO NIFEROUS � Lacustr1ne � Geosyndrnal Mdrlnt!' Fo �s1/.J[I] 0 L11cusfrtn� f"Ds.s!I�O Both [] - - - T I I MAP OF QUEENSLAND SHOWING LOCALITIES WHERE CARBONIFEROUS BEDS HAVE BEEN FOUND (Reconstructed from Bryan and Jones 1945). Localities.-!. Pascoe River. 2. Newellton. 3. St. Helens. 4. Drummond Range. .'i . Lion «nd Neerkol Creeks. 6. Rockhampton. 7. Station Creek. :\H. Morgan. 8. Mt. Grim. 9. Diglum. 10. Cania. ll. �lany Peaks. 1:!. Cannindah. 13. Riverleigh and Mundubbera. 14. Texas. 15. \1t. Barney. 16. Bullock Creek, :\atal Downs. 17. ---- Plate I 2 3 4 5 7 8 9a 9b lOa Plale Ill Plate IV 2 Sc Sb 6a Sa Sd 6b 7a 7b 8 9