ARKIV FÖR KEMI, MINERALOGI OCH GEOLOGI. RAND 26 A. N:o 13.

An Outline of the Succession and Migration of

Non-Crinoid Pelmatozoan Faunas in the Lower

Paleozoic of Scandinavia.

By

GERHARD REGNELL.

With l table and 4 figures in the text.

Communicated April 28th 1 948 by ERIK STENSIÖ and PER GEIJER.

Con tents. Page

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l Previous work on the regional distribution of the non-crinoid pelmatozoans 4 Cambrian......................................................... 8 Ordovician........................................................ l l

Tremadocian.................................................. l l Skiddavian................................................... 1 2 Llandeilian. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l 7 Caradocian. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Ashgillian.... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3

Silurian. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 6 Llandoverian.................................................. 3 7 Wenlockian. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Ludlovian. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Silurian undefined. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

Concluding remarks................................................ 45 Literature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 7

lut.-oduction.

The Pelmatozoa cannot be said on the whole to play a very important role in the fossil record of the Cambro-Silurian deposits of Scandinavia, either with regard to number of genera and species, or to individuals. This is true first and foremost of the non-crinoid forms, whereas crinoids are a fairly conspicuous element as rock-builders especially in the Silurian of Gotland.

Arkiv /Cr kemi, mineralogi o. geologi. Bd 26 A. N:o 13. l

2 ARKIV FÖR KEMI, MINERALOGI O. GEOLOGI. BD 26 Å. N:O )3.

Apart from a few horizons in which Hydrophoridea appear in great masses , non-crinoid pelmatozoans are as a rule found more or less fortuitously in Sweden. This also applies to Norway. The Old-Paleozoic echinoderm faunas of that country, however, are as yet somewhat less thoroughly investigated from a taxonornie point of view. As to Denmark, the Cambro-Silurian rock floor is exposed in the Island of Bornholm only. No statements of cystoids from Bornholm are to be found in the literature. Nor do the collections of the Paleontological Museum in Copenhagen contain any spe­cimens, according to kind information (in titt. ) from Professor CHR. POULSEN.

The reason why non-crinoid Pelmatozoa are found so relatively seldom can hardly be - as in the case of certain other animal groups - that their remains have been destroyed to a great extent . On the contrary, the calcified external skeleton of most echino­derms 111ust have been especially resistant to the destructive power of environmental agencies . So it may be assumed, instead, that echinoderms are proportionally more frequent in the thanato ­coenoses than in the biocoenoses from which these were derived. The generally_ scarce occurrence of non-crinoid pelmatozoans, therefore , must be considered mainly a primary condition.

The migration of the sessile echinoderms now under discussion must have occurred during the stage of ontogeny in which the larvae were unattached, pelagic organisms. 1 In recent crinoids the free-living existence of the larvae is of short duration. Accord­ing to DELAGE & HEROUARD (1903 , p. 369) it lasts for two or three days only. In some cases it is considerably longer, however. MoRTENSEN (193 7 , p. 61 ) found that the embryos of the comatulid Tropiometra audouini are free-swimming 5-6 days, or even longer, before attaching themselves . Of course we do not know what were the conditions in this respect in the non-crinoid pelmatozoans living in the Paleozoic seas . Y et it is most likely that their larvae had much the same cycle of development as have recent crinoids. If that be so, the shortness of the free-swimming stage in the larval development was a factor acting more or less unfavourably upon the migration of the individual species , which could hardly spread over long distances of the open sea. At any event this can not have been possible except under especially favourable conditions , in that the larvae were transported by currents rapid enough to carry them from one littoral zone to another in the course of a few days. For i t must be kept in mind that all pelmatozoans , including the crinoids, so far known from Paleozoic deposits were living in fairly shallow water (cf . DEECKE 1915, p. l seq. , and RADDING 1933 , pp. 51-52) as indicated by the lithologic character of the

1 As regards the theory of the adult of Echinosphaerites as a planktonic form, cf. REGNELL ( 1 945, p. 146 ) .

GERHARD REGNELL, NON-CRJNOID PELMATOZOAN l<'AUNAS. 3

rock and by the associated fauna. With regard to environmental conditions they were consequently very different from recent stalked crinoids, which are confined exclusively to the deep sea . B ut this implies, too, that the restraining effect of the special larval development just mentioned was selective to a certain degree, in so far as the crinoids, and the biastoids as well , were not affected to the same extent as the other Pelmatozoa, as indicated by their rich differentiation and development.

As mentioned above, the remains of non7crinoid pelmatozoans are embedded in sediments deposited in a shallow sea. But they do seldom occur in sediments of the actual shore-line, i. e. con­glomerates and sandstones, as was already stated by SALTER (1866, p . 272: >>-- the limestorre has more or less of an aranaceous character, and i t is the rarest thing for Cystidae to occur in s u ch a rock») . Among forms origirrating from Scandinavia, one species only has been found in a sandstone, viz . the edrioasteroid Stromato­cystites balticus J AEKEL. 1 The y are extremely rare in pyrite­bearing sediments , and are almost absent in graptolite shales and other planktonic deposits2 ; nor are they common in other shales. In Sweden marly beds of the red Tretaspis shale of Dalarna, and the Staurocephalus shale and the so-called Cystoid shale of Scania have yielded some material. But species enclosed in limestorre series rather often occur in marly or argillaceous la y ers ( cf. RAD­DING 1933, p: 53, foot-note 1 ) . In the Oslo region, Norway, there are also shaly, lime-free sediments (especially in zones 4a�X-4b1X) hearing Hydrophoridea, which is also true of calciferous shales of zone 3c� (REGNELL 1948 a, p. 23 ) . Thus the non-crinoid Pelma­tozoa cannot be said , as a whole, to be typical facies fossils . To what extent the individual species are dependent on lithologic facies, however, is a question which will not be disenssed in this connexion.

The crinoids have survived up to the present time with a number of species, small in comparison with the vast amount of forms flourishing in ages past and especially during the Paleozoic. The non-crinoid pelmatozoans , on the other hand, became extinct in the Permian already (Blastoidea) , most of its groups still earlier . The limited stratigraphic range of the non-crinoid pelmatozoans,

1 From other regions we have examples of cystoid faunas occurring in psammitic rocks, e . g. the Starfish Bed of Girvan and the Schistes quartzeux of Sombre-et-Meuse (BATHER 1 9 1 3 , §§ 6, 559) .

2 In one instance only has the writer noticed hydrophorid remains (Cheiro­crinus sp . ) associated with graptolites, viz. in a slab of a hard, 1ime-free shale from Fossum, Norway (spec. No. Ec 5 1 65 in the Paleozool. Dept. of the State Museum of Natural History, Stockholm) , probably origirrating from the Upper Didymograptus shale (4a(X) . The Lower Devonian hydrophorids and carpoids described by DEHM ( 1 933 , 1 934) occur in the Hunsriick siates of Bundenbach, which contain, however, a preponderatingly benthonic fauna.

4 ARKIV FÖR KEMI, MINERALOG I O. GEOLOGI. BD 26 Å. N:O 13.

or, in other words, their short period of flourish, might be regarded as a function of the relatively feeble expansion power of most of the forms concerned.

Previous Work on the Regional Distr·ibution of the Non­

Crinoid Pelmatozoans.

The postulate of an attempt at analysing the migrations and history of development of a certain group of fossil animals within a definite area should be a thorough knowledge of the strati­graphic appearance all over the world of the several genera and species of the group which, in its turn, requires careful taxonornie investigations. It is evident that this demand is nothing but a vain wish : the fossil record is incomplete, and our research into the material available fragmentary or entirely wanting.

In the case of the non-crinoid Pelmatozoa, our present state of knowledge admits of but a rough outline of their history in the Oambro-Silurian seas covering part of the S c a n d i n a v i a n P e n­i n s u l a of to-day. The bulk of the Swedish material available has been re-examined recently by the present writer (REGNELL 1945) . Much of the old collections stored in the museums is not very fit for stratigraphical purposes, however, because of its inadequate labelling. The N orwegian material has not been studied in any detail as yet, but it has been thought worth while to consider in this connexion as far as it is known. ANGELIN (1878 ) , BRÖGGER (1882 ) (largely revised by the present author) , JAEKEL (1926 ) , and REGNELL (1948 a) have described a few forms. Some information on the matter can be gathered from some other papers as well, but these do not present any descriptions but mere statements on the occurrence of certain species .

The knowledge of the fossil record of the Ordovician-Silurian deposits of E s t o n i a and the L e n i n g r a d d i s t r i c t is notoriously of the utmost importance for the understanding of the Ordovician­Silurian faunas of Scandinavia. Good descriptions of non-crinoid pelmatozoans and observations on their stratigraphic range were given by several old er authors (P AND ER 1830, LEUCHTENBERG 1843 , VERNEUIL 1845, VOLBORTH 1846 i. a . , EICHWALD 1860 i. a . , F. ScHMIDT 1874 i . a . ) . In a paper published more than a hundred years ago EICHWALD (1845 or 1846) made some special comments of the cystoid fauna of Seandinavia as compared with that of the East Baltic Provinces , and on paleogeographical conditions in Sweden. It is true that these remarks are primitive and nu­acceptable to-day, being based on false identifications of species and on confused ideas in general, but they may be quoted here as a first , and hitherto almost unique, attempt at disenssing the

GERH.I.RD .REGNE LL, NON-CRINOID PELMATOZOAN FAUNAS. 5

stratigraphical occurrence and regional distribution of the non­crinoid pelmatozoans in the Scandinavian-Baltic region:

>>Mit dem Thonschiefer wechsellagert bei Christiania [Oslo] ein schwarzer Kalkstein, der ansser mehreren Arten von Asaphus, vorziiglich aus der Familie der Crinoideen die zu den ältesten Gattungen gehörigen Sphaeroniten enthält , die auch um Pawlowsk liberall in so grosser Menge vorkommen; dahin gehört vorziiglich Sphaeronites aurantium und S. pomum [foot-note : »Die Art von Pawlowsk ist durch die Zeichnung der Täfelchen von Sphaero­nites pomum völlig verschieden und gehört offenbar mit meinem Pro iocrinites aviform is in e i n e Gattung, da in beiden die 5 rinnenartigen Spalten bemerkt werden»] , die beide in denselben Schichten um Pawlowsk vorkommen . Auch ist die erstere Art schon von LINNE auf der Insel Oeland gefunden warden und da­durch diese Insel als Fortsetzung des Festlandes von Schweden anzusehen; sie kann also nicht, gleich Gottland, eine urweltliche Klippe oder ein Korallenriff in Meere der Vorwelt gebildet haben. Sphaer. pomum findet sich nicht in Norwegen, aber Sph. granatum nicht selten auf Gottland; diese ist bei uns nach nicht gefunden worden, während sich Sph. testudinarius Hrs . auch selten um Paw­lowsk, so wie auf Oeland findet. Ebenso wird Heliocrinus balticus in der Gegend von Christiania beobachtet , so wie um Pawlowsk und in Esthland, jedoch dort nirgends ein Hemicosmites, Cryptocri­nites , Gonocrinites [ = Echinoencrinites] o der ähnliche Arten, die unsere esthländischen Schichten so sehr auszeichnen. >> (EICHW ALD 1845? , pp . 103-104 . ) In the same work (p . 93) he states that in Sweden >>sphaeronites>> are especially characteristic of the clay­shales. This , as we have seen, is definitely not the case.

A good deal of the material dealt with in JAEKEL's fundamental work of 1899 originated from Estonia and Russia, and much light was thrown upon the morphology and appearance ·of the Edrioasteroidea and Hydrophoridea.

More recently, most important contributions to our knowledge of the Ordavieian non-crinoid pelmatozoan fauna of the East Baltic area have been published by HECKER in a series of papers (HECKER 1923, 1938, 1939, 1940 ) . 0RVIKU'S (1927 ) study on Echinosphaerites may also be mentioned, as well as a paper by PHLEGER (1935) and a note by LURA (1940) on a Silurian form.

Inconsiderable remains of Hydrophoridea were reported from P o l a nd by e. g. CzARNOCKI (1928 ) .

V.Asc.ÅUTANU (1931) gave same information on hydrophorids )

in the Upper Ordavieian of R o u m a n ia. vVhat is known about the non-crinoid pelmatozoan fauna of

B o h e m ia is due mainly to the work of BARRANDE (1887 ) (cf. also PERNER 1900 ) . A survey of the geographical and stratigraphical distribution of these forms in general is also included in BAR-

6 ARKIV FÖ R KE:&II, MINERALOGI O. GEOLOGI. BD 26 Å. N:O 13.

RANDE's great monograph. On p . 220 there is a table showing the number of cystoid genera (24 in all) then known within the dif­ferent stages of the Swedish series of s trata. It is based on in­formation from G. LINDSTRÖM. - The papers of POMPECKJ (1896 a) and RuzrcKA (1939) may also be referred to .

LORETZ (1884) and FREYBERG (1923) described Hydrophoridea from the Ordavieian of G e r m a ny, and RICHTER (1930) and DEHJVI (1933, 1934) have reported on Edrioasteroidea, Carpoidea , and Hydrophoridea from the Lower Devonian.

Apart from BATHER's (1913 ) admirable monograph on the Ashgillian Hydrophoridea and Carpoidea of Girvan, very little has been published on the taxonomy of non-crinoid pelmatozoans from the Lower Paleozoic of G r e a t Bri t a in after the memoir by FORBEs (1848) . This work, which thus appeared one hundred years ago, is quite naturally fairly out of date. SALTER (1866 ) described forms from N orth W ales. T here is a short article of special interest in this connexion, however, viz . the summary by SPENCER (1938) on the distribution and migration of the star­fishes and hydrophorids in the British Lower Paleozoic fauna . Part of this problem was touched upon slightly already by MARR (1882 , p. 325) . BEGG (1934, 1939) has dealt with Cyclocystoides. Of course there are a number of notes on cystaids in papers with mainly geological contents .

The Garadocian (sensu latu) cystoid material from B e l g i u m is still insufficiently known, bu t will be studied by the present writer. MAILLIEUX (1926 ) published a note on a Devonian Cyclocystoides. A study by Dr. HERTHA DoRECK on this and Rhine Devonian forms will appear in >>Senckenbergiana>> .

Our knowledge of the Hydrophoridea and Carpoidea of F r a n c e has increased considerably thanks to the important investigation carried out by CHAUVEL (1941 ) . Other contributions have been made by KOEKE� (1886 ) , THORAL (1935 a, 1935 b ) and by DREY­FUSS (1939 ) .

The inconsiderable material o f cystaids from the Lower Paleo­zoic of I t a l y has been reported on by BATHER (1910) , PILOTTI (1924 ) , and VINASSA DE REG�Y (1941 ) , of S p a i n by F AURA Y SANS (1913 ) , and of P o r t u g a l by DELGADO (1908 ) .

A description o f the cystoid fauna o f M o r o c c o i s being pre ­pared by G. [DELPEY-]TERMIER. A find of a Lower Devonian carpoid from S o u t h A f r i c a was announced by RENKIE (1936 ) .

BATHER (1906) and REED (1906, 1917 i . a . ) reported on the fairly multiform hydrophorid fauna of I n d i a. SuN (1936 ) de­scribed Ordavieian species from C h i n a and drew paleogeographical conclusions from the occurrence of certain European genera.

N o non-crinoid pelmatozoans have been recorded from the Lower Paleozoic of A u s t r a l i a, as far as the writer is aware. The

GERHARD REGNI<� LL, NON-CRINOID PELMATOZOAN FAUNAS. 7

so-called Cyamoidea (with the genus Peridionites) described by WHITEHOUSE (1941, p. 5 seq. ) from the lowest Middle Cambrian of Queensland and recognized as the ancestral group of the Pelmato­zoa cannot be accepted as echinoderms at all , according to GISLEK (194 7 ) .

The contents o f non-crinoid pelmatozoans i n the Old Paleozoic deposits of N o r t h A m e ri c a have been elucidated by a number of authors : HALL (1852 , 1867 i. a . ) , BILLINGS (1858 i. a . ) , SALTER & BILLINGS (1858 ) , S . A. MILLER (1889 i. a . ) , SCHUCHERT (1904, 1919 i. a . ) , RAYMOND (1913 i . a . ) , FOERSTE (1914, 1916 i . a . ) , RUEDE­}'[ANN (1925) , BASSLER (1935, 1936, 1943 ) , A. E . WILSON (1946 ) , and others . The importance o f various cystoid genera for the intercontinental correlation between Europe and America is con­sidered briefly in BASSLER 1943. Of considerable use to the student of Paleozoic pelmatozoans are the bibliographies published by the author just mentioned (BASSLER 1915, 1938) . This is of course also true of the bibliographic and faunal index of Paleozoic pelma­tozoans compiled by BASSLER & MOODEY (1943 ) . This work presents useful faunal lists of different areas , arranged according to stratigraphic units . For reasons easily understood the faunal lists and the references suffer from certain unavoidable errors and shortcomings. Where Sweden and N orway are concerned, this is largely due to the fact that the recent revisions of the non-crinoid pelmatozoan faunas of these countries had not yet appeared when the index was published.

We have very meagre information about the appearance of non­crinoid pelmatozoans in the Lower Paleozoic of S o u t h A m e r i c a. Insignificant remains were mentioned by HARRINGTON (1937 , p . 101 ; 1938 , p . 117 ) and KOBAYASHI (1937 , p . 417 ) .

The A r c t i c has not yielded much material either. I t has been recorded by POULSEN (1927 , 193 7 , 1946) 1 , TROEDSSON (1928), and ROY (1941 ) .

I n this short review of our present information o n the non­crinoid pelmatozoan faunas within various Lower Paleozoic areas such papers mainly have been considered as have more or less hearing on the subject to be dealt with below. Before concluding, it is appropriate to eaU attention to the magnificent work by JAEKEL (1899) , briefly mentioned above. In that work JAEKEL treated the Edrioasteroidea and Hydrophoridea not only taxo­nomically but also , and in the first place, from general points of view, thus also with regard to distribution in time and space.

1 The remains of cystaids mentioned from the Lower Ordavieian of Cape Clay, N. W. Greenland, by PouLSEN 1927 (p. 282) and from Cape Webster, Ella Island, by PouLSEN 1937 (pp. 27-28) , may be related to Macrocystella, accord­ing to kind information from Prof . PouLSEN (in litt. ) .

8 ARKIV FÖR KEMI, l!IINERALOGI O. GEOLOGI. BD 26 Å. N:O 13.

A survey of the evolution and distribution of the echinoderms in the Lower Paleozoic was presented recently by G. DELPEY [ -TERMIER] (1944 ) . (N. B. the Addendum to Literature, p. 54 below. )

Cambrian.

� Only a small number of pelmatozoans are known from the Cambrian deposits of the globe. From the Lower Gambrian there are two or three species (Stromatocystites walcotti SCHUCHERT, S. w. minor ScHUCHERT, Lepidocystis wanneri FOERSTE) ; from the Upper Gambrian no more than one species has been described as far ( Trachelocrinus resseri ULRICH; >>Obscure remains of cystoids>> were further reported from the Upper Cambrian of Andalnsia by R. & E. RICHTER, 1940, p . 52 , and of southern France, ibid. , p. 64 , after MIQUEL, mentioned by THORAL, 1935 a , pp. 90, 92, and 1935 b, p. 10 etc . , as well ) . In comparison with this very meagre representation in the lower and upper parts of the system, the Middle Gambrian pelmatozoan faunas are remarkably well devel­oped within a few areas , viz . Bohemia and southern France­Spain. Here the carpoids reached a vigorons flourish with a con­siderable differentiation of forms. N o traces of these carpoid faunas have been discovered in the Scandinavian series of strata . But the single pelmatozoan genus known so far from the Middle Gambrian of Scandinavia, the edrioasteroid Stromatocystites balti­cus J AEKEL\ has congeneric forms in the areas mentioned, i. e. the Bohemian S. pentangularis POMPECKJ and the Herault species S. cannati (MIQUEL) .

One of the first attempts at tracing the main features o f Gam­brian paleogeography is that by FRECH in 1899. Since that time a wealth of facts has been accumulated regarding the Gambrian sequences and faunas within different areas . And yet the data available are not sufficient (and will of course never be) for the constructing of reliable paleogeographical maps . This is clearly demonstrated by GRABAU's magnificent attempt at a synthesis of an immense material, aiming at a general cosmography for the several periods of the Paleozoic (GRABAU 1936-1938) . The work was never accomplished but remains a gigantic and interesting torso. In 1940, however, GRABAU published a comprehensive volume, in w hi ch the »pulse-beat>> of the earth was followed from the pre-Gambrian up to the coming of Man.

What is of special interest to us in this connexion is to consider

1 Two specimens only have been found of this species, and not in bed rock but in the drift, of north Germany. From the lithologic character of the rock it seems to be possible to conclude, however, that the specimens originate from the Middle Cambrian of Sweden, and presurnably from Öland (cf. REGNELL 1 945, pp. 1 98, 200) .

GERHARD REGNELL, NON-CRINOID PEJ�MA'l'OZOAN FAUNAS. 9

for a while the map showing the paleogeography of Europe in Middle Cambrian time (GRABAU 1936 a , PL II b ) and the cor­responding map of Pangaea (ibid. , PL II) which may be the most recent maps of their kind published. 1 One important correction was proposed by R. & E. RICHTER (1941 b, p. 34, textfig. 2) who showed that strong objections can be raised against the Tethys geosyncline as interpreted by GRABAU. Of still greater importance to us is the shape given to the northern section of the map . 2 The Caledonian geosyncline is taken in a sense different from that generally accepted. .As is well known, the Caledonian geosyncline in Europe skirts the N.W. coast of the continent, from Ireland via Norway to Spitsbergen, the S .E . foreland being represented in Seandinavia by the Central Fennoscandian continental trunk. The Paleozoic sediments of the Baltic Shield S. and S .E . of this con­tinental trunk are , quite naturally, typical shallow water deposits formed in an epicontinental sea (cf. e . g. HaLTEDAHL 1920, text­fig. 12; FREBaLD 1928; ÅSKLUND 1929, text-fig. l; ÅSKLUND 1938, textfig. 3 ; BAILEY & HaLTEDAHL 1938, text-figs . 1 , 16 ) . 3 The Baltic Basin was an integral part of the sea-province characterized by the .Acado-Baltic fauna, covering further .Acadia , England, Bohemia, Middle Germany, Southern France, Sardinia, parts of Spain and North .Africa (cf . R. & E. RICHTER 1940, pp . 71, 74 ; 1941 a , pp. 68, 69; 1941 b , p . 31) . In view of this, i t is not surpris­ing to find representatives of Stromatoeystites in the Middle Cam­brian of Southern Scandinavia, Bohemia, and Southern France.

For this .Acado-Baltic Basin GRABAU (1936 a, Pl. II b) used the term >>Caledonian geosyncline>> , which is of course illegitimate (cf. R. & E. RICHTER 1940, p . 72 ) . The Scandinavian section of the true Caledonian geosynclinc , on the other hand, coincides tolerably well with part of GRABAU's St. Lawrence geosyncline.

1 A synthetic map of the Cambrian in Europe is found on p. 99 in S . VON BuBNOFF's »Einfiihrung in die Erdgeschichte>> , I , Berlin 1941 . Somewhat modified reproductians of his maps of Pangaea were given by GRABAU 1 940.

2 We do not pay any attention to the fact that the map is drawn in polar pro­j ection with the North Pole in the region of modern Egypt, but Jet compass di­rections apply to modern maps. Nor is it necessary to discuss here GRABAU's supposition that, in Paleozoic times, the contirrents were in j uxtaposition (WE­GENER position) , forming one land-mass.

3 It should not be concealed, however, that in the text GRABAU ( 1 936 a, p. 634) declared that the »northern part of the Caledonian syncline [in the special sense of GRABAU] i . e. the part invalving southern Sweden and Bornholm, as weil as that of the Oslo region, represents marginal shelf deposits, with repeated floodings as outlined in the text. Whether the central part of the geosyncline was more rapidly subsiding and has thicker deposits is unknown.>> In their papers quoted repeatedly in this connexion, R. & E. RICHTER ( 1 940, p. 72; 1 9 4 1 b, pp. 36-37 ) object, rightly, against the wide -spread application of the term »geosyncline>> in paleogeographical discussions. The term should be restricted to designate the zones from which the folded sedimentary mountain chains developed.

10 ARKIV FÖR KEJIII, JIIINERALOGI O. GEOLOGI. BD 2() Å. N:O 13.

This is the northern extension of the Appalachian geosyncline , dismembered in Middle Cambrian time from the southern brancl1 by the upheavel of the so-called Albany axis . The Appalachian trdugh (including the St. Lawrence geosyncline) must be sup­posed to have erossed the Atlantic west of , and almost paraHel to, the Acado-Baltic province. We have to assume, further, that an exchange of faunal elements was possible between the Appalachian trough and the European part of the Acado-Baltic Basin, which communicated in an Arctic Sea. Known facts show (i. a. by the presence of a Middle Cambrian trilobite fauna in Bennett Island north of the New Siberian Group, described by HoLM & WEsTER­GÅRD 1930, closely related to that of southern Scandinavia) that the Baltic Basin was in connexion with the Siberian Sea , which, according to GRABAU (1936 a, p . 495) , may have been the centre of evolution and dispersal of the European Middle Cambrian faunas . 1

The presumed northern path-way from the Appalachian pro­vince to the Acado-Baltic would also explain the appearance of the American genus Stromatocystites in European series of strata. For Stromatocystites must be looked upon as an American element, as the oldest known representative of the g enus originates from the Lower Cambrian of Newfoundland (S. walcotti ScHUCHERT 1919 ) . The migration in a northern direction from Newfoundland was of course initiated in Lower Cambrian time already, because the area now occupied by western N ewfoundland seems to have been elevated over the sea in Middle Cambrian time (cf . maps I and II in GRABAU 1936 a ) . 2 It may be mentioned parenthetically, moreover , that the same migration route may have brought certain elements from the Appalachian province into the upper Lower Cambrian fauna of Aistjakk in Lappland, North Sweden, described by KAUTSKY (1945) , as indicated by the close affinity between Lingulella westergårdi KAUTSKY and the East Greenland species L. prisca POULSEN. And it was suggested by PouLSEN (1932 , p . 65) that )>the East Greenland geosyncline should b e regarded a s a continuation of the St. Lawrence geosyncline•> . Botsfordia may as well have invaded from the north , being known from the Lower Cambrian of Northwest Greenland (POULSEN 1927 , pp. 250, 339 ) , but may also have arrived in Lappland from another direction, for the genus occurs also within the Atlantic coast region of N orth America (POULSE::-< 1927 , p. 338 ) , belonging to the Acado-Baltic faunal province.

1 Cf. also the discussion of a possible Siberian Middle Cambrian strait con­necting the Acado -Baltic and 'Vestern Pacific provinces in R. & E. RICHTER 1941 b, p. 38 seq.

2 It can also be imagined in this and similar cases that forms common to the Appalachian and Acado-Baltic fauna! provinces have spread from a centre of evolution and dispersal in the Arctic (Boreal) Sea.

GERHAl{D REGNELI,, NON-CRINOID PELJ\IATOZOAN FAUNAS. ] l

0l'dovician. In Seandinavia the Hydrophoridea are by far the most im­

portant group of the non-crinoid pelmatozoans. In this region, as well as in other areas, they reached their fullest development during the Ordovician period. As will appear from the following survey, they are very poorly represented in the Lower Ordovician, however, which may be a consequence of the decrease in the pelmatozoan stock as a whole in Upper Gambrian time.

In order to facilitate the comparison with foreign areas, the author has thought it wisest to review the contents of non-crinoid pelmatozoans in the Ordovician (and the Silurian) sequence ac­cording to the British standard section, although the correlation with Scandinavian strata is not quite definite throughout.

Tremadocian.1

Cystoids of Tremadocian age are practically unknown from Scandinavia. The only statement of which the writer is aware is that of TULLBERG (1882 a, p. 230), who mentioned >>Cystoid frag­ments>> from the Ceratopyge limestorre of southern Öland. In their monograph on the Ceratopyge fauna, MOBERG & SEGERBERG (1906, p. 107) did not pay any attention to the so-called cystoids referred to by TULLBERG, on account of insufficient information supplied by that author. Obviously they had no opportunity of studying the material alluded to, nor had the present writer . So the true nature of these fossils must be left undecided . B ut we can hardly doubt that they were remains of pelmatozoans .

The lower section of the Tremadocian of Scandinavia , the Dictyonema shale , has so far not yielded any fossils of this kind.

Thus, for the present, we lack actual data for considering the Tremadocian pelmatozoan fauna of Seandinavia in relation to that of other areas. As emphasized above, the total of non-crinoid pelmatozoans (and pelmatozoans in general) is most insignificant in Lower Ordovician deposits. The richest fauna hitherto dis­covered origirrates from the Upper Tremadocian zone of Asaphel­lina barroisi of Languedoc (THORAL 1935 b, p. 330). Macrocystella, and allied forms (Mimocystites etc. ) 2 , seems to have had a wide, possibly cosmopolitic distribution in the Tremadocian, being recorded from France (THORAL L c . ) , England (STUBBLEFIELD & BULMAN 1927, pp. 113 , 118), Bohemia (BARRANDE 1887, p. 164 ) ,

1 Following LAPWORTH 1 879 ( O n the tripartite classification o f the Lower Palaeozoic rocks. - Geol. Mag. 6. Pp. l-15) , British stratigraphers usually classify the Tremadocian with the Upper Cambrian.

2 Referred to the Eocrinoidea by JAEKEL (1918, pp. 26-2 7 ) and, in accord­ance with him, by the present writer (REGNELL 1 945, p. 3 7 ) as weil.

12 ARKIV FÖR KEMI, MINERALOGI O. GEOLOGI. BD 26 Å. N:O 13.

Bavaria (POMPECKJ 1896 b, p . 2 ) , GREENLAND (cf . above, p . 7 , foot-note 1 ) , further from the South American Cordilieras (Ko­BAYASHI 193 7 , p . 417 ; HARRINGTON 193 7 , p . 101 ; 1938, p . 117 ) , and Korea (KOBAYASHI 1935, p . 59 ; the horizon, the Eoorthis zone, is taken by KoBAYASHI as the top-most zone of the Upper Cam­brian sequence) .

As mentioned above, the Tremadocian fauna o f Seandinavia does not call for any discussion of paleogeographical conditions with regard to the distribution of the pelmatozoans . Maps relating to that epoch have been compiled e. g. by GRABAU (1916, p . 621, text-fig. 10; 1936 b , Pl. III-a; 193 7 , Pl. III ) , HOLTEDAHL (1920, p. 6 , text-fig. 3) , and KOBAYASHI (1936, p . 491, text-fig. 1) . The several sea areas (and faunal provinces ) seem to have been in closer connexion with one another than during the Cambrian (N.B. that the term >>Caledonian geosyncline>> is used by GRABAU in the same improper sense as earlier) . The main paleogeographical features and migration routes were summarized by KoBAYASHI (1936, pp. 490-492 ) . As to Europe, a general survey of the dis­tribution and relationships of the faunal elements was presented by BRÖGGER (1896 ) in his well-known paper. THORAL (1935 b , p . 341 ) , b y his investigations of the Lower Ordovician of Southern France , also arrived at the conclusion that there existed an obvious affinity between the Tremadocian faunas of that region and of the part of the Acado-Baltic province ranging from eastern N. America to Seandinavia and Bohemia.

Skiddavian (Arenigian).

In the lowest Skiddavian strata of Sweden, made up of the Planilimbata stage (of the Asaphus series , or Orthoceratite lime­stone) , Hydrophoridea are extremely uncommon. In fact one unique specimen only has been found, belonging to Cheirocrinus holmi REGNELL, from the green Planilimbata limestone of the Island of Öland.

From the superimposing division, the Limhata stage, no remains of non-crinoid pelmatozoans have been recorded in the literature, nor have such fossils been noted by the present writer. It would not be preposterons to suspect that the total lack, or at least utmost rarity, of cystoids and similar forms in these sedi­ments is to some degree due to the special conditions under which the Limbata limestone was deposited. If we leave Scania and the Oslo region out of account, in which this zone (3c<X in Norway) i s not very well developed, the rock is usually, though not always, a red limestone. Its red colour is due to the contents of iron oxide and/or its compounds. MoBERG (1904 ) , who made the problem the subject of a special investigation, arrived at the conclusion

G E RHAR D R EGN E LL, N O N - C RINO ID P E Lnf.A.T O Z OAN FAUNåS. 13

(op . cit. , p. 142 ) that the red colouring of certain divisions of the Cambro-Silurian sequence was effected thus that the mud was transported into a shallow and shelving sea. Then it was exposed to oxidation and complete red weathering during periods of emer­gence when the tide was low. MOBERG (1904, p . 140) also pointed out that the red beds of the Orthoceratite limestone as a rule contain a smaller number of species than do the grey ones and are sometimes characterized by peculiar dwarf faunas. It cannot be altogether denied that the genesis of the red Limhata limestone thus outlined might have acted unfavourably upon the devel­opment of cystoids. It is true that Hydrophoridea and their allies are wanting in the upper, partially red-coloured portion of the Planilimbata limestone as well, but, on the other hand, there are a few forms w hi ch are confin ed to the red Platyurus limestone ( cf . below) . It should be remembered, too , that cystoids were repre­sented very sparsely not only in Scandinavian deposits but in general at the time of the formation of the Planilimbata and Limhata limestones. In Norwegian equivalents to these divisions of the Swedish Orthoceratite limestone no non-crinoid pelmato­zoans have been met with , as far as the author's knowledge goes.

The upper part of the Skiddavian is designated in Sweden (in shelly facies) as Asaphus limestone, and in Norway as 3c� (Ex­pansus shale and limestone) . Here the hydrophoridean fauna reached its first flourish in Sweden, with several remarkable forms, and mass occurrence where Sphaeronites is concerned. In the Oslo region the first representatives of the group made their appearance.

In the south and middle of Öland two faunistically and litho­logically characteristic divisions can be recognized within the Asaphus limestone, separated from each other by a bed crammed with thecae of Sphaeronites pomum (GYLLENHAAL) . The Sphaero­nites bed is further developed in Västergötland and, presumably, in Östergötland (REGNELL 1945, pp. 166-167 ) . Boulders of a grey limestone collected in 1947 at Lanna in Närke by Professor E. STENSIÖ , Stockholm, contain numerous poorly preserved spe­cimens of Sphaeronites. The rock very likely derives from the Asaphus limestone, the youngest member of the Paleozoic sequence in Närke. The find may indicate that the Sphaeronites bed has been present locally in Närke as weiL

As pointed out previously by the writer (REGNELL 1945, p . 166 ) , Sphaeronites pomum is undoubtedly a complex species com­prising several forms. In most cases, however, the material of Sphaeronites of the pomum-group is in such a poor state of pre­servation as not to permit the investigator to recognize characters reliable enough for the establishing of separate forms.

It may be worth while to make some comments on two spe­cimens of Sphaeronites pomum from Fågelsång in Scania (RM Ec

14 A RIUV FÖl� KEMI, :MINERALOGI O. GEOLOGI. BD 2(} Å. N:O ]3.

5025-5026).1 Provided that they were collected from the bed rock - the rock attached to the specimens agrees lithologically with the black Orthoceratite limestorre found at Fågelsång - they have some hearing on the stratigraphy of the Orthoceratite lime­storre at that locality (cf. REGNELL 1945, p . 168, footnote 9 ) . The Orthoceratite limestorre in S .E . Scania, and at Röstånga as well (the small exposure in Kvarnbäcken is accessible at very low water only) , is known to reach into the Asaphus limestone. But at Fågelsång, where the Orthoceratite limestorre is in direct contact with the Upper Didymograptus shale, the limestorre series does not include strata younger than the stage of Megalaspis limbata , ac­cording to the latest information available in the literature (FUNK­QUIST 1919, p. 44; EKsTRÖM 1937, p. 49 ) . 2 However, Sphaeronites pomum indicates clearly that some part of the Asaphus limestorre is present. Among the trilobites obtained from the Orthoceratite limestorre of Fågelsång (cf. the lists published by TULLBERG 1883 b , p . 244, and MoBERG 1907, p . 257) there are also several pointing in the same direction , e . g. Harpina scanica, Megalaspis extenuata , Pterygometopus sclerops3, Ampyx volborthi (the last-mentioned species associated with Sphaeronites pomum, a typical pygidium being present in the same hand-specimen as RM Ec 5026)4•

Consequently, the sub-zone of Phyllograptus nobilis (lowest sub-zone of the Upper Didymograptus shale) cannot be equivalent to at least the basal portion of the Asaphus limestone. According to EKSTRÖM (1937, p. 49, and the section on Pl . 10) there is, be­tween the Limhata limestorre and the Upper Didymograptus shale , a transition series (0 . 4-1 .1 m thick) of limestorre layers, inter­bedded with marl, shaly clay, and marl shale . This series has yielded a pygidium of Megalaspis limbata (not indicated whether from the lower or upper part of the transition series) , and on ac-

1 RM = Paleozoological Department of the State Museum of �atural History, Stockholm.

2 PouLSEN ( 1 936 , pp. 50-5 1 ) also contributed to the characterization of the stratigraphic position of the Fågelsång Orthoceratite limestorre but did not discuss its upper delimitation. It may be mentioned in this connexion that Cyclopyge umbonata {ANGELIN) has been found at Röstånga also {TULLBERG 1 883 a, p. 5 ) . In reproducing the stratigraphic scheme compiled by TuLLBERG ( 1 883 b, table facing p. 258) MoBERG ( 1 9 1 1 , p. 1 34) placed the Upper Didymo· graptus shale immediately above >)the black Fågelsång limestorre with M egalaspis /imbata S. & B . >) . In TULLBERG's table, however, the limestorre below the zone of >)Phyllograptus cf. typuS>) was referred to as >)Orthocerenkalk>), with no further attribute. The dividing of this camplex into faunistically characteristic zones was not done until 1 890 by MOBERG.

3 In Öland these three species are confined to the Asaphus limestone (cf . WIMAN 1 906, pp. 1 04, 105 ) .

4 Ampyx volborthi F . SCHMIDT was found by POULSEN {1936, pp . 48 , 51 ) in the Umbonata limestorre of Bornholm, but as a rule seems to appear at higher horizons. It was recorded from the Lower Asaphus limestone of Öland by the present writer {REGNELL 1 942, p. 4) .

GERHARD REGN ELL, NON·Cl�INOID PELMATOZOAN l<'AUNAS. li)

count of that it was referred by EKsTRÖM to the stage of Megal­aspis limbata. It is not unreasonable to suggest, however, that the upper portion of the so-called transition series represents part of, or the total of, the Lower Asaphus limestone. A view of the correlation of the Upper Didymograptus shale similar to that advanced here has been, moreover, expressed by TROEDBSON (1928, stratigraphic table on p. 179 ; 1931, stratigraphic table on p. 327) , who located the Upper Didymograptus shale above the Asaphus limestorre and put its lower part as an equivalent of the Gigas limestone.

As mentioned above , the first hydrophorid to appear in the Skiddavian of Seandinavia was a species of Cheirocrinus. In the Upper Skiddavian, Cheirocrinus and other genera of the Super­family Glyptocystitida became the most significant pelmatozoan element along with Sphaeronites pomum (in Sweden) . But con­trary to that species all Glyptocystitida are rare forms in the Scandinavian fossil record, most of them being represented as yet by some few specimens only. In Swedish strata the following species have been met with : Cheirocrinus leuchtenbergi (ANGELIN} , Ch. cf. nodosus (JAEKEL} , Echinoencrinites cf. reticulatus JAEKEL, E. senckenbergii MEYER, and Proctocystis monstruosa REGNELL. Zone 3c� of the Oslo region has yielded Cheirocrinus hyperboreus REGNELL , Echinoencrinites senckenbergii acutangulus REGNELL, and Erinocystis bröggeri REGNELL. Our list of species known so far from the Upper Skiddavian is complete after adding Hemicos­mites� sp . REGNELL (1948 a ) from the Oslo region\ and a species from the Asaphus limestorre of Röstånga, Scania, mentioned by TULLBERG (1883 a, p. 5) as >>cystoid>>2 • The material upon which TuLLBERG based his statement could unfortunately not be re­found.

Like the extra geosynclinal shelly faunas of the Scandinavian Paleozoic in general , the Skiddavian non-crinoid pelmatozoan assemblage is decidedly East Baltic in aspect. It is true that Sphaeronites pomum3, so characteristic of the Asaphus limestorre

1 The stratigraphic position of this species is not undisputed. According to BRÖGGER ( 1 882, p. 42 ) , who referred to it as Hemicosmites sp., the single specimen available, an isolated thecal plate, origirrates either from the Upper Skiddavian ( 3c�) or from the Lower Llandeilian ( 3cy).

2 In a review article, speaking of the Orthoceratite limestorre at Röstånga, TuLLBERG ( 1 883 b, p. 245) put it thus: >>hier aber sind auch Cystideen angetrof­fen>>.

3 No representative whatever of Sphaeronites has been described from the East Baltic area. Yet, according to kind information ( in litt. ) from Professor R. HECKER, Moscow, there are forms in the Leningrad province which have much in common with the Swedish Sphaeronites. It is due to a mistake, however, when BASSLER & MooDEY ( 1 943, pp. 36, 1 90) report Sph. pomum from the Leningrad province. They refer to VoLBORTH ( 1 870, Pl.-fig. l), who in the figure quoted reproduced a detail figure of Sph. pomum communicated by Lov:EN

J G A R KIV FÖR K E M I , l\II N E R A LOG I O. GEO LOGI . B D 26 Å . :N :O 13.

of Sweden, has not been recorded from areas outside this country, but the Glyptocystitida show a close affinity to East Baltic forms. In stage Bn (Megalaspis limestone) 1 in Estonia and the Leningrad district , corresponding to the stages of M egalaspis planilimbata and M. limbata in Sweden, we find species of the genera Cheiro­crinus and Echinoencrinites2 • These are represented by a multitude of forms in the subsequent stage, Bm (Vaginatum limestone, equivalent to the Asaphus limestone) , which also contains Erino­cystis and, probably, Proctocystis. Species common to the East Baltic area and Sweden are: Echinoencrinites cf. reticulatus JAEKEL and E. senckenbergii MEYER, and to the USSR and Norway: E. senckenbergii acutangulus REGNELL. 3 N o Skiddavian forms from Sweden are identical with such forms from Norway, nor is there any species common to Scandina,ia and any other region but the East Baltic Provinces.

The Norwegian Hemicosmites'? sp . is a little surprising, if it can be trusted that it origirrates from the Upper Skiddavian. Neither in Sweden, nor with any certainty in the East Baltic area has the genus any representatives until in the Llandeilian (the Upper Llandeilian as far Sweden is concerned) .4

( 1 867 ) . VoLBORTH ( 1 846, p. 1 86 ) stated that Sph. pomum had not been met with in Russia, hut had been identified erronously with other species (cf. the list of non-synonyms in REGNELL 1 945, p. 1 6 2 ) . It is wrong as well to report Sph. pomum from the Oslo region (BASSLER & MooDEY 1 943, pp. 37 , 1 90 ) . The statement must be due to the accidental misconception that the mountain Kinnekulle is situated in Norway. In a list of the Swedish Ordavieian cystoid fauna Sph. pomum is reported properly from »Kinnekulle, V•>[estrogothia].

1 The letter indices for the divisions in the Estonian geological column used in this paper are according to ÖPIK ( 1 930) .

2 As to the stratigraphic range of the East Baltic forms, see e. g. BASSLER & MoODEY ( 1 943, p. 37 seq . ) or ScuPrN ( 1 928, pp. 1 7 1� 1 7 2 ) . In some cases the statements do not agree. Information on East Baltic forms is also found in REGNELL 1 945.

3 The atfinity between the Norwegian and the East Baltic echinoderm faunas is further accentuated by the presence of the ophiocistid Volehovia in both regions (cf. REGNELL 1 948 a ) .

' The stratigraphic range of the different forms is as a rule indicated vaguely in the writings of the older authors and cannot always be made out with cer­tainty . According to JAEKEL ( 1 899, p . 308) , the oldest form of Hemicosmites known from the East Baltic area is H. malum (PANDER 1 830; not 1 840, as stated erronously by JAEKEL 1 899, p. 309, and REGNELL 1 945, p. 100) . BASSLER & MOODEY ( 1 943, p. 165 ) , however, report >>Hemicosmites elongatus PANDER•> from the >>Kunda or Walchow fm. >> which correspond, the former to the Vaginatum limestone, the latter to the Megalaspis limestone. In searching for information about H. elongatus in the original Jiterature, the preEent writer arrived at the conclusion that such a species does not exist. The only place where the name in question is used, as far as the writer is aware, is in ErCHWALD 1 856 (p. 124 ) , referred to by BASSLER & MooDEY (by a lapsus calami these authors give 1 867 as the year of publication of ErcHWALD's paper) . All that i s to be read here i s as follows: >>Hemicosmites elongatus PAND. Findet sich im untern Grauwackenkalke von Pulkowa und Popofka. •> Thus no description is given, and none was given

GER H ARD R E GNELL, NO N-CR INOID p �;Llli A 'l'OZOAN F AUNAS . 1 7

As just mentioned, no species of Skiddavian non-crinoid pelma­tozoans of Seandinavia are known from other areas except the East Baltic Provinces. This is also true of the genera, with the possible exception of Cheirocrinus. l THORAL (1935 a, p. 115) de­scribed, as a matter of fact, a species from the Lower Skiddavian of the Montagne Noir, which he designated as Ch. ? languedocianus. Judging from the figures published and the description, the deter­mination would seem to be correct .

It can hardly be doubted that the Skiddavian echinoderm fauna invaded Seandinavia via the East Baltic Provinces. Con­temporarily, certain elements migrated in a westerly direction along the northern border of the Caledonian-Hungarian barrier (or girdle of islands) thought to have separated intermittently the Bohemian Basin from the North-European Sea in Early and Middle Ordovician time (cf . e . g. REGNELL 1940, p. 11, with further references) .

Llandeilian.

As the Lower Skiddavian, so the lower part of the Llandeilian is characterized by a very poor echinoderm fauna. In Sweden this section of the series of strata in limestone facies is made up of the red Gigas and Platyurus limestones. With regard to lithologic development these stages are rather similar to the red limestones at the base of the Skiddavian barren of cystoid remains (cf . above, p . 13 ) . It is possible that the same factors as acted unfavourably upon the flourish of echinoderm faunas in early Skiddavian time were of some importance in this instance as well.

The Gigas limestone in Sweden has yielded no non-crinoid pelmatozoans but Sphaeronites? sp . from Öland (REGNELL 1945, p . 171 ) and Glyptosphaerites leuchtenbergi (VOLBORTH) from Öster­götland. As regards the last-mentioned species, the stratigraphic

by PANDER ( 1 830) either, who knew no species with that name. »Hemieosmites elongatus>> , therefore, is not even a nomen nudum but probably got inta the literature by accident. Very likely EICHWALD intended to refer to Remieosmiles oblongus (PANDER, sub Eehinosphaerites, 1 830, p. 146 ) . The stratigraphic position of the so-called H. elongatus does not either appear from EICHWALD 's statement j ust quoted. Exactly the same formulatian is used e . g. in the case of H. verrueosus, which, according to JAEKEL ( 1 899, p. 3 1 0) , origirrates from F p a camplex o n the Ordovician-Silurian boundary, known formerly a s Lyck. ho lm. H. oblongus, on the other hand, which has probably given rise to the who le discussion, is said by BASSLER & MooDEY ( 1 943, p. 1 66) to appear in stage C , , the lowest part o f which i s a t least not older than the Asaphus limestone. »H. oblongus>>, reported by FoRBEs ( 1 848, p. 5 1 1 ) , and others, with or without a mark of interrogation, from Wales might be, according to JAEKEL ( 1 899, p . 3 1 5) , a Caryoerinites or Caryoeystites ( = Helioerinites, as defined by REGNELL 1 945) .

1 In a previous paper (REGNELL 1 945, p . 98) Remieosmiles was listed, with a mark of interrogation, to appear in the Lower Ordavieian of France. The species upon which the statement was based may be of Upper Ordavieian age , however (cf. THORAL 1 935 a, p. 159 ) .

A r kiv för kemi, mineralogi o . geologi. B d 26 A. N : o 1 .1. 2

l !) ARK I V FÖR KEJ!II, MINERALOGI O. GEOLOGI. B D 26 Å. N:O 13. determination is not quite conclusive (REGNELL 1945, p . 156 ) . Glyptosphaerites leuchtenbergi has a wide range i n the East Baltic sequence (Vaginatum limestone-Uhaku) , and is one of the two Hydrophoridea known from the stage of Asaphus platyurus in Sweden . This stage is an equivalent of the Aseri (C1 r; ) in Estonia. The Swedish material has been derived mainly from Öland, further from Dalarna and, presurnably, the N orth Baltic area. The seeond species occurring in this stage is Echinosphaerites aurantium suecicus JAEKEL from Öland. As pointed out by REGNELL (1945, p . 149 ) , this form is very closely related to - if not identical with - the East Baltic Echinosphaerites aurantium infra HECKER from the corresponding horizon. Thus the intimate connexion between the East Baltic and Swedish Ordovician pelmatozoan faunas is demonstrated again . Cheirocrinus, recorded from equi­valent or somewhat younger strata of the Oslo region, has also East Baltic affinities, as we have seen. The genus is represented by Ch. nodosus (JAEKEL) from the Ogygia shale (4acx) of Heden­stad (JAEKEL 1899, p. 221 ) . Further material has been obtained partly from the Ogygia shale of Tortberg at Frogner, Oslo (GRORUD 1940, pp. 158, 159 ) , and partly from the Ampyx limestorre (4a� ) of Hovindsholm, Helgoya, Mjosen (ANGELIN 1878, Pl. 12 , figs. 16-17; HOLTEDAHL 1909, p . 9 ) . The species was referred by ANGELIN to Ch. leuchtenbergi (ANGELIN ) , by JAEKEL (1899, p. 221) to Ch. nodosus (JAEKEL) , by HaLTEDAHL to Ch. penniger EICHWALD, and by GRORUD to Ch. cf. leuchtenbergi. As stated by REGNELL (1945 , p. 76 )\ we are probably cancerned with a form which can be designated as Ch. cf. granulatus (JAEKEL) .

The Platyurus limestorre o f the Swedish Orthoceratite lime­storre is superiroposed by the stage of Illaenus schroeteri upon which follows, in Dalarna, the Crassicauda stage (JAANUSSON 1947) . This is equivalent with the Uhaku (Cg-C2.) of Estonia, and possibly with the Ancistroceras limestorre of Öland ("1\IoBERG 1911 , p. 104) 2 and in part with the Ampyx limestorre of Norway. In Seandinavia non-crinoid pelmatozoans are as a rule scarce in this sequence, which is made up largely of grey limestones. The Nor­wegian Cheirocrinus cf. granulatus was just mentioned. 3 According

1 GRORUD 's paper ( 1 940) was then not taken into account. 2 This question was left open by JAANUSSON ( 1 947, p. 50) . 3 After the present paper was already completed, the writer had occasion

to examine a few samples of limestone originating from 4a�-4b0( (Ampyx limestone-Lower Chasmops shale; locs . : Kullerud, Ringerike; Abbediengskollen, near Oslo ) hearing remains of a Cheirocrinus (the specimens belong to the Pale­ontologicaJ Museum in Oslo ) . Certain axial ridges of the thecal plates are pro­vided with protuberances similar to those of Ch. nodosus, but the test is finely granulated, as in Ch. granulatus, a character which - as far as known - is not found in Ch. nodosus. This form very Iikely represents a new species which may be referred to as Ch. aff. nodosus until more complete material is available. It is evident that this species - as weil as other Norwegian hydrophorid material ­requires further attention.

G E R H A R D R E G N EL L , N O N- C R I N O I D PEL JII A TO Z O A N F A U N A S . 1 9

to BRÖGGER (1887, p. 17) , Echinosphaerites aurantium (GYLLEN­HAAL) appears rather frequently at the top of the Ampyx limestorre (cf. also STORMER 1934, p. 25 ) . BRÖGGER (l. c . ) also mentioned mumerous columnals of cystoids>> (translated here ) , but of course >>cystoids>> must be taken as a synonym for >>pelmatozoans>> in this instance. From the upper Ancistroceras limestorre of Öland and equivalent strata of Västergötland Echinosphaerites (aurantium?) has been reported (cf . REGNELL 1945 , p. 147) . Otherwise only two species seem to have been met with in the Swedish Schroeteri limestone, viz . Cheirocrinus cf. nodosus (JAEKEL) from Väster­götland, mentioned previously from the Skiddavian (Asaphus limestone) of that district, and Cryptocrinites sp. from Gräsgård in Öland. 1

It was emphasized by JAANussoN (1947, p. 48) that there is a remarkable difference in the composition of the fauna in the Crassicauda stage of Dalarna and the Uhaku, its equivalent in the East Baltic district , inasmuch as the rich cystoid fauna of the Uhaku is absent , or nearly so , in the Crassicauda beds. This dif­ference is supposed to be due to differences in facies.

If we turn to the upper part of the Llandeilian in Scandinavia, such a difference in faunal character is no longer predominant , especially not in Sweden (though it should be remembered that the majority of the Paleozoic pelmatozoans of Norway have as yet not been the subject of taxonomical investigation ) . In fact, an op­timum of non-crinoid pelmatozoan development was initiated during the Upper Llandeilian. The abrupt changes in the fre­quency of certain faunal elements - in this special case the non­crinoid pelmatozoans - indicate that the series of strata does not represent a complete and uninterrupted sequence, but a complex formed during repeated transgressions separated by periods of upheaval. Very often the hiatuses are more or less masked dis­conformities (TROEDSSON 1925, pp. 192-193 ) . The stratigraphic gap between the middle and upper parts of the Llandeilian is demonstrated in certain districts of Sweden (Jämtland) by the presence of a conglamerate at the base of the Chasmops series (cf. THORBLUND 1940, pp. 13, 69 seq. , 121, 183 ) .

The top of the Llandeilian i s thus represented i n Sweden by the lower part of the Chasmops series, made up mainly of the Cystoid limestorre or Lower Chasmops limestone. Norwegian equivalents are the Lower Chasmops shale (4bcx) and the Lower Chasmops limestorre (4b� ) . 2

1 The Swedish Cryptocrinites (ono specimen only known) will be described in a forthcoming paper. The find of this genus is interesting, because the pre­seneo of Eocrinoidea in the Swedish series of strata has been indicated hitherto by Bockia? sp. only (REGNELL 1 945, p. 6 7 ) .

2 STORMER ( 1 934, p. 25) and BAILEY & HOLTEDAHL ( 1 938, P J . l) place the Llandeilian-Caradocian boundary below the Lower Chasmops shale. Follow-

:!0 A R K I V F Ö R K E M I , M I N ER A L O G I O . G E O LO G I . B D 26 Å. N:O )3 .

The earrelation between shelly and graptolite facies in the Upper Llandeilian of Sweden was discussed at some length by THORBLUND (1940, pp. 121-122 ) . On geological grounds he suggested >>that an upper part of the zone of N emagraptus gracilis be considered equivalent in age to the basal Chasmops series , instead of to the uppermost part of the Schroeteri limestone, i . e . the so-called Ancistroceras Iimestone•> (THORBLUND 1940, p . 121 ) . The paleontological evidence wanted b y THORBLUND seems t o be supplied by a graptolite associated with the shelly fauna of the Lo w er Chasmops Iimestone at B öda, N orth Öland. In examining specimens collected by him at the locality mentioned I, the present writer detected a fragment of a graptolite attached to the theca of a specimen of Echinosphaerites aurantium. The graptolite frag­ment was determined, with a high degree of probability, as a branch of a species belonging to the form-group of Nemagraptus gracilis (HALL) . If that be true, it seems to imply that the zone of N emagraptus gracilis is equivalent, not only with the basal con­glamerate and sandstone as indicated by THORSLUND (l. c . ; the sequences in Jämtland and Södermanland are considered) , but with part of the Lower Chasmops limestone as weil. 2 Som e fe"VI· fragments of graptolites , probably Glyptograptus sp . , were also found associated with Echinosphaerites aurantium in the flinty Lower Chasmops limestone of Västergötland.3 These may not shed any further light on the problem of correlation, however.

The non-crinoid Pelmatozoa known from the Lower Chasmops limestone in different districts of Sweden are as follows:

Scania: H el iocrinites granatum (W AHLENBERG) ( cf. REGNELL 1945, p. 126 ) , Echinosphaerites� sp . (REGNELL 1945, p. 153 ) , Haplo­sphaeronis oblonga (ANGELIN ) , >>cystoids>> (from the zone of >>Oaly­

mene dilatata>> , TULLBERG 1882 b, p . 19 ) . Öland: Oheirocrinus sp . (REGNELL 1945 , p . 77) , Hemicosmites

oelandicus REGNELL, Oaryocystites angelini (HAECKEL) , Heliocrini­tes granatum (WAHLENBERG) , H. ovalis (ANGELIN ) (boulders) , H . guttaeformis REGNELL� (boulder) , Echinosphaerites aurantium (GYLLENHAAL) , Sphaeronites globulus (ANGELIN ) .

Gotland (drilling core ) : Echinosphaerites sp . (THORSLUND 1938, p. 28) , Haplosphaeronis sp. (THORBLUND, l . c . ) .

Västergötland: Oheirocrinus sp. (REGNELL 1945, p . 77) , Sticho­cystis al�ttacea (ANGELIN )4, Heliocrinites ovalis (ANGELIN) , Echino-

ing THORSLUND ( 1 940, p. 1 83 } , the Lower Chasmops shale and the Lower Chasmops limestorre are here included in the Llandeilian.

l The material is in the PaleontologicaJ Institution of Lund University. 2 JAANUsSON & MARTNA ( 1 948, p . 1 90) suggest that the >>Crassicauda lime­

stone appears to be contemporaneous with the zone of Nemagrapt1ts gracilis» . 3 Specimen in the PaleozoologicaJ Dept. of the State Museum of Natural

History, Stockholm (No. Ec 3249 ) . 4 Harizon not established with certainty.

GERH ARD R EGNELL, NO N - C RINO I D PELJ\I A 'l'OZO A N F A UNAS. 2 1

sphaerites aurantium (GYLLENHAAL) , Echinosphaerites sp. (REGNELL 1945, p. 152 ) , Sphaeronites globulus (ANGELIN ) , Haplosphaeronis oblonga (ANGELIN ) , >>cystoid>> (W ALLERIUS 1894, p. 300; cf. REG­NELL 1945, p . 184 ) .

Östergötland: Heliocrinites angustiporus REGNELL, H. grana­tum (W AHLENBERG ) , H. oval is (ANGELIN ) , H. guttaeformis REGNELL, Echinosphaerites aurantium (GYLLENHAAL) , E. grandis JAEKEL? , Sphaeronites globulus (ANGELIN ) .

Södermanland: Echinosphaerites sp. (THORSLUND 1940, p . 114) . Dalarna: Bockia? s p . , Cheirocrinus leuchtenbergi (ANGELIN ) ?

(cf. REGNELL 1945, p . 75 ) , Caryocystites aff. angelini (HAECKEL) \ C. s p . 2 , Heliocrinites angustiporus REGNELL, H. granatum ( W AH­LENBERG) , H. cf. granatum (JAANUSSON 1947 , p. 4 7 ) , H. ovalis (ANGELIN ) ? , H. prominens (ANGELIN )3, Echinosphaerites aut·an­tium (GYLLENHAAL) , E. grandis JAEKEL? Sphaeronites globulus (ANGELIN ) , Haplosphaeronis oblonga (ANGELIN) , mndeterminable cystoid>> (JAANUSSON 1947 ) . 4

Jämtland: Caryocystites sp . (REGNELL 1945 , p . 118 ) , Helio­crinites granatum (WAHLENBERG) (boulders ) , Echinosphaerites aurantium (GYLLENHAAL) , E. sp. (THORSLUND 1940, pp . 68, 82 ) , Haplosphaeronis sp. (THORSLUND 1940, pp. 5 5 , 91 ) .

The North Baltic area (boulders ) : Heliocrinites prominens (ANGELIN ) , Echinosphaerites aurantium (GYLLENHAAL) .

The follöwing species of non-crinoid Pelmatozoa originate from the Lower Chasmops limestone of N orway: Echinosphaerites aurantium (GYLLENHAAL) (STORMER 1934, p. 25 ; etc. ) , Glypto ­sphaerites sp . (REGNELL 1945, p . 154 , foot-note 1 ) 5 , Sphaeronites globulus (ANGELIN) ? (cf. REGNELL 1945, pp. 161, 1 71 ) .

After this survey o f the stratigraphic range and regional dis ­tribution within Seandinavia of the Llandeilian non-crinoid pel­matozoan faunas, we shall try to trace their relations to con-

1 Specimen from Fj äcka (coll. J. MARTNA 1 945), not y et described (Paleonto­logical Institution, Uppsala) .

2 Specimen from Fj äcka (coli. G. HoLM 1 880) , not yet described (Paleonto­logical Dept . , State Mus. Nat. Rist . , Stockholm) .

3 The holotype o f H . prominens was found i n the collections o f the Paleo ­zoo!. Dept. of the State Mus. Nat. Rist . , Stockholm, after the author's mono ­graph of 1 945 was already printed (cf. REGNELL 1 945, p. 1 3 1 ) .

4 Some forms have been recorded from strata above the Lower Chasmops limestorre s. str. A specimen referred to Heliocrinites cf. ovalis (ANGELIN) has been collected by JAANUSSON & MARTNA in the >>beds of plastic clay (probably bentonite) >> . This section follows upon the Lower Chasmops limestorre s . str. and is superiroposed by beds of undetermined stratigraphic position but included provisionally in the Lower Chasmops limestone s . l . (JAANUSSON & MARTNA 1 948, p. 1 8 9 ) . According to the authors mentioned, these beds have yielded i . a. Echinosphaerites aurantium.

5 On the label, written by G. HOLM, who collected the specimen, the horizon is given as >>Chasmops series•>. Locality: Baadbugten.

22 A R KI V FÖR K E lll i , MINERALOGI O . GE OLOGI. B D 26 Å. N : O 13.

temporary extra-Scandinavian echinoderm faunas , which was briefly mentioned above.

starting with the Lower Llandeilian, we have had occasion already to emphasize its affinity to the East Baltic area, as de­monstrated by Glyptosphaerites leuchtenbergi, and by representa­tives of Cheirocrinus, Echinosphaerites , and Cryptocrinites. There is a statement (PILOTTI 1924, p. 32 ) that Glyptosphaerites , and possibly G. leuchtenbergi , appears in Sardinia, which seems to require further confirmation, however. The genus was not men­tioned by VrNASSA DE REGNY (1941 ) . If the statement be true , however, it would indicate an expansion towards the south of the Baltic echinoderm fauna. T hen i t seems most reasonable to assume that the migration was made possible by transgressions over the Caledonian-Hungarian barrier, in Middle Ordavieian time (cf . a bov e, p . 1 7 ) . Ordavieian strata of Baltic facies hearing Echino­sphaerites are present in the St. Cross Mountain between Warsaw and Cracow (CZARNOCKI 1928, p. 557 ; as regards the distribution of Lower Paleozoic rocks in Poland, see also SUJKOWSKI 1946, pp . 193-194, Pl. 14 ) . It may be that the sea in which these were deposited reached as far as the Bohemian Basin, which, in its turn , stood in connexion with the sea covering parts of Britain (SPENCER 1938) , South Europe, and North Africa. As a whole, the rich hydrophorid faunas of Bohemia have not much in common with the Baltic ones . But it should be noticed that the very genus Glyptosphaerites that gave rise to this discussion is represented in Bohemia by G. ferrigenus (BARRANDE) . According to PERNER (1900, p. 145 ) , the harizon of the single specimen known is Dd1 (Komarov beds, of Skiddavian age; cf. KETTNER and BoucEK 1936, Table 4 ) . This is admittedly an obstacle for the supposed Baltic origin of Glyptosphaerites in Bohemia. For in the East Baltic area the genus does not appear until in the Vaginatum limestone, as we have seen. This is equivalent to the lower part of the Sarka beds , which overlie the Komarov beds .

Glyptosphaerites has also been recorded from other areas , viz . from Thuringia, N.W. of the Bohemian Basin (LORETZ 1884; FREY­BERG 1923, p . 265 ) , and from Spain, S.W. of it (JAEKEL 1899, p. 425 ) . In the first-mentioned case, Glyptosphaerites is said to have been derived from strata equivalent to Dd5 in Bohemia, i. e. the very top of the Ordavieian sequence. If the specific determina­tions are correct , the genus is associated with hydrophorids appear­ing at considerably lower horizons, e . g. Protocrinites fragum (EICHWALD) from the Vaginatum limestone. On the other hand, there are species such as Orocystites helmhackeri thuringiae (JAE­KEL) and Heliocrinites confortatus (BARRANDE) , which are closely allied to, or identical with forms of BARRANDE's stage Dd4• This is Caradocian-Ashgillian in age. Evidently some forms of the

G E R H A R D R E G N E L L , N O N - C R I N O I D P E LJ\I A T O Z O A N F A U N A S . 23 Ordavieian cystoid assemblage of Thuringia have not been de­termined properly and offer no reliable basis for the assumption of a connexion between the Baltic and the Bohemian Basin via Thuringia in Lower Llandeilian time. Such conditions may rather have been prevailing during the Upper Ordovician. - As regards Glyptosphaerites sp. from Spain, JAEKEL (1899, p. 425 ) stated only that it origirrates from the Upper Ordavieian of Trasno.

It is well known that species belonging to the form-group of Echinosphaerites aurantium have an almost cosmopolitic distribu­tion. This is true of E. aurantium suecicus as well , if FIELD ' s statement (1919, p . 4 1 9 ) can be trusted that the species mentioned occurs in the Rodman formation of Pennsylvania. ! In Sweden it is confined to the Platyurus limestone; the most closely related E. aurantium infra appears in the equivalent Aseri of the East Baltic area. The North American form holds a considerably higher stratigraphic position, a fact which is in itself not quite unreasonable. 2 But hydrophorid genera of B altic aspect have recently been reported from Oklahoma strata equivalent to the Lower Llandeilian (BASSLER 1943 ) . These are Cheirocrinus (Cool Creek formation) and Caryocystites (Falls formation) , both Cha­zyan. From the Bromide formation (Black River, probably equivalent to the Swedish Schroeteri stage) BASSLER (1943 , p . 703) recorded Echinoencrinites� O f these, Cheirocrinus and Echi­noencrinites are represented, as we have seen, already in the Skid­davian of the Baltic area. As far as our present knowledge goes , this cannot be said of Caryocystites (as defined by REGNELL 1945, p . 112 ) , which does not appear until in the Upper Llandeilian of Sweden; the age of C. esthoniae (JAEKEL) is not known with any certainty, the unique specimen being obtained from an erratic boulder (JAEKEL 1899, p. 339-340) . Anyhow, we can hardly doubt that the genera under discussion are either immigrants from the Baltic region - transformed (at least some of them) into other species , it is true - or else are descendants from common ancestors in some centre of faunal evolution connected with the Baltic area, on the on e hand, and with N orth America, on the other.

1 Echinosphaerites aurantium suecicus was not mentioned from USA by BASSLER & MOODEY ( 1 943, p. 155 ) .

2 FIELD ( 1 9 1 9, p. 419 ) took the Rodman formation as •>the closing phase of Stones River time». According to WILMARTH ( 1 938, p . 1 83 7 ) , it is the upper formation of Black River, overlying the Lowville limestorre and underlying the Trenton. RuFFMAN ( 1 945, Middle Ordavieian earrelation table on p. 1 73 ) , finally, took i t a s the youngest formation o f the Nealmont, which forms the lower part of the Trenton. Thus the Rodman would be equivalent to part of, or the whole of the Crassicauda stage in Sweden. It may be mentioned that -according to CooPER ( 1 945, p. 275) - >>Stones River as a general time-rock term should no longer be used, nor should the name be revived if a post -Chazyan, pre­Black River group is recognized>>.

:2-1 AR K I V FÖR K E ni i , l\I I N E R A L O G I O . G E O L O G I . H D 26 Å. N : O 1 3 .

Then the question arises: What routes of migration did they use to arrive at their sites in the Western hemisphere, or, where was the common centre of evolution?

The general paleogeographical conditions of Middle Ordovician time were discussed i . a . by GRABAU (1923, p. 47 seq. ) , who pub­lished a map of Boreal paleogeography in Black River time (op. cit. , fig. 31 , p . 56) . The same subject, with reference to the Ameri­can and Eurasian circumpolar regions, was dealt with by HOLTE­DAHL (1918, p. 93; 1920, pp. 5-6; 1924, p. 109 seq. ) , and by TROEDsSON (1928, p. 176 seq. ) . It seems likely that there existed a narrow Scandinavian-Scottish land barrier (HoLTEDAHL 1920, p . 5 ) separating the Baltic and the Arctic Sea provinces from each other (cf . TROEDssoN 1928, p . 183, who also reviewed the divergent opinions on the faunal connexion between N orth America and the East Baltic area arrived at by BASSLER on the basis of his studies on bryozoans ) . The migration of faunal elements, therefore, was effected across the Arctic Sea. A >>true circumpolar distribution of the sea>> in Lower Ordovician time was assumed by HOLTEDAHL (1924, p . 121 ) , who further advocated the view that at a >>Somewhat younger [than Tetragraptus time] Ordavieian time the Khabarova fauna of Northern Russia and the Pterygometopus selerops fauna of the Tunguska etc. tell of a more or less open connection with the South Scandinavian Baltic region, while the Black River-Trenton faunas of B ear Island, N orth Greenland, the American Arctic Archipelago and riorthwestern North America indicate a very wide distribution of the North American-Arctic Sea at this period, the sea reaching further south (across the Baffin Land-Hudson Bay area) than in the preceding, Lower Ordavieian time>> (HoLTEDAHL 1924, p. 122 ) .

TROEDSSON (1928, p . 1 8 7 ) seems t o b e inclined t o accept largely the views set forth by HOLTEDAHL and went a little more into details: >>If - in this case - the geosyncline ex tending from N o­vaya Zemlj a to Alasca means a coast-line this interpretation is in accordance with the statements made here. Furthermore, there was a central (deep) basin, the Arctic sea, which formed an ob­stacle in the way of the litoral fauna but, in return, made it easier for the pelagic forms or forms which were able of pelagic spread­ing. >>

The present writer cannot see any possibility so far to decide conclusively whether the Hydrophoridea now under discussion are immigrants from the East Baltic area to North America via the Arctic sea, or whether the ancestral forms arose in the Arctic sea and from there invaded the B altic area as well as North America. As far as the writer is aware, no Lower and Middle Llandeilian Hydrophoridea have been recorded from the Arctic. This fact -which may be due to our still imperfect knowledge of the Lower

G E R H A R D R E GNE L L , N O N - C R I N OI D P E L DI A T O Z O A N F A U N .A S . 2 5

Paleozoic faunas o f these regions - cannot o f course be used as an argument for either o f these alternatives . The only species com­mon to the Baltic area and North America, as we have seen, is Echinosphaerites aurantium suecicus, provided that FIELD's de­termination is true (cf. above, p . 23 ) . If any importance can be attached to this matter, it is rather in favour of the theory sug­gesting a Baltic origin, all the more as the species in question ap­pears earlier in Sweden than in Pennsylvania. It should be noted, however, that according to FIELD , E. aurantium suecicus is as­sociated i . a . with E. grandis, which appears in Sweden in the Upper Llandeilian and the Lower Caradocian. One of the two determinations , therefore, seems to be wrong.

Now we shall consider briefly the non-crinoid pelmatozoans of the Upper Llandeilian in Scandinavia, and their contemporaries.

The Eocrinoidea are probably represented by a species of Bockia. This genus is known otherwise from the East Baltic area only, appearing first in stage Bu (Skiddavian) and ranging into stage 03 , which is equivalent (at least in part) to the Swedish Lower Chasmops limestone.

Several of the Hydrophoridea have East Baltic affinities, but as to species, a number of them seem to be endemic to Sweden. Reviewing the genera in systematic order, Cheirocrinus is richly differentiated in the East Baltic area, where it has a considerable vertical range. It may be remembered, too, that there are earlier representatives of this genus in Sweden and in Norway as well. Further species are found in North America. In Seandinavia no post-Llandeilian forms of Cheirocrinus have been met with so far , but the genus reached a wide distribution, having been recorded from Caradocian, or younger, strata of the East Baltic area, Bohemia (BARRANDE 1887, p . 160) , Great Britain (Girvan) (BATHER 1913, p. 442 ) , Portugal (DELGADO 1908, p . 83 ) , North America (cf . e. g. BASSLER 1915 ) , Baffin Land (RoY 1941, p. 80 ) , and Green­land (TROEDSSON 1928, p . 105 ) .

A s regards the early occurrence o f Hemicosmites i n Norway and the East Baltic area, cf. above, p. 16 . One species was described by SuN (1936 , p. 481 ) from Kweichou, China. The age of the Shihtzepu shale, from which Hemicosmites (associated with other non-crinoid pelmatozoans) was collected, was said to be Llandei­lian. l Since the fauna also contains a species designated as >>Sticho-

1 SFN ( 1 936, p . 48 1 ) suggested that the Shihtzepu sha1e is equivalent to the Shihtien beds in Yunnan hearing Sinocystis loczyi REED, which is also found in the Kweichou fauna. According to YIN & L u ( 1 936-3 7 ) , the term >>Shihtien beds>> should be restricted to the lower part of the series of strata, which is probably Llanvirnian. The superimposing section is called >>Hengshuitang beds•> , the lower part being characterized by Sinocystis and the upper by Gamarocrinus. The authors did not express any opinion as to the age of the Hengshuitang beds, but they may belong to a middle or upper part of the Llandeilian, which would

26 A R KIV FÖR K E M I , MINERAL OGI O . G E O L O G I . BD 26 Å. N : O 13. cystis sp . (cfr geometricus)», it will hardly be older than the Upper Llandeilian. In Sweden Stichocystis geometrica (ANGELIN } appears in a somewhat younger horizon, the Maeronrus beds (Caradocian) . St. alutacea (ANGELIN } , o n the other hand, seems t o have been derived from the Lower Ollasmops limestorre of Västergötland.

Caryocystites angelini (HAECKEL} is a significant element in the Lower Ollasmops limestorre fauna of northern Öland. A related form is known from corresponding strata of Dalarna . C. sp. from Jämtland shows more affinity to C. lagenalis REGNELL (cf. REG­NELL 1945, p . 118 ) . Some remarks on the genus have been given above (p. 23 ) .

Whereas Caryocystites had a very restricted regional distribu­tion during the Upper Llandeilian-Caradocian, Heliocrinites was widely spread, especially in Garadocian time . In Sweden there are at least five species in the Upper Llandeilian Ollasmops limestone, but from younger Ordovician strata one species only has been recorded. l If we have a general look at Heliocrinites, it is evident that it approached Seandinavia from the east, for the East Baltic series of strata contains several species that are older than the Swedish orres . In the far East, too, Heliocrinites has representa­tives, in part prior to those of Seandinavia ( species origirrating from the Naungkangyi beds of Burma, which are equivalent to the so-called Echinosphaerites limestone, Cl r;-Cul , of the East Baltic Provinces , according to REED 1906, p. 85 ) , and in part fairly equivalent to them (a species from the Shihtzepu shale of China, cf. above, p. 25, foot-note 1 ) . Later on, as we have seen, Heliocrinites reached far outside the Baltic Basin. 2 Thus Caradocian, or Ashgil­lian, forms have been reported from the following extra Baltic­Scandinavian countries : Thuringia (FREYBERG 1923, p. 263, H. confortatus , sub Echinosphaerites), Bollernia (BARRANDE 1887 , p . 153 , as the preceding) , Great Britain (BATHER 1913, p . 494 ) , B elgium (BATHER, L c. ) , France (KOENEN 1886, H. rouvillei , sub Caryocystites ; cf. also REGNELL 1945, p. 137 ) , Spain (FAURA Y SANs 1913, p . 112 ) , and Portugal (JAEKEL 1899, p . 330 ) . Of the Heliocrinites forms occurring within foreign areas none is identical specifically with any Swedish form, as far as known.

thus be the age of the Shihtzepu shale as weil. According to SuN & SzETU ( 1 947, p. 22) , the (Llanvirnian) Upper Shihtien formation of W. Yunnan is super­imposed by the Lower Pupiao formation, of Llandeilian age. This is correlated with the Upper Naungkangyi series of Burma, whereas the Lower Naungkangyi series is taken as an equivalent of the Lower and Upper Shihtien formations, which are said to range from the Tremadocian ? to the Llanvirnian.

1 It should be noted, however, that the stratigraphic range of the t wo species of Heliocrinites mentioned below as Silurian is not quite conclusive.

2 Heliocrinites granatum mentioned from Great Britain by FORBES ( 1 848, p. 5 1 2 ) and by numerous subsequent authors might be designated as H. balticus EICHWALD ? (cf. REGNELL 1 945, p. 1 2 2 ) .

G E R H A U D R E G N E L L , N O N - C R I N O I D P E L lll A T O Z O A N F A U N A S . 27

The writer bad occasion above (p. 23 ) to point out that Echino­sphaerites aurantium, and allied forms, bad an almost world-wide distribution. The relations between the forms of the various areas are not very well known, however. The Scandinavian material, too, is fairly heterogeneous , as indicated also by the rather wide strati­graphic range of this form-group. We have seen that it was re­presented in strata below the Upper Llandeilian, and it appears in Garadocian beds as well. In Sweden there are also a few forms of Echinosphaerites in the Upper Llandeilian, which are clearly distinguished from E. aurantium. One large species occurring in Östergötland and Dalarna (Upper Llandeilian-Lower Carado­cian) was referred tentatively to E. grandis JAEKEL by the present writer (REGNELL 1945 , p. 149 ) . It was suggested that E. pogrebowi HECKER may be identical with E. grandis. If that be so, E. grandis is an example of a species common to Sweden and the East Baltic Provinces . Within the area last mentioned, E. pogrebowi was recorded from the Kukruse, which is equivalent to the Lower Chasnwps limestone. A closely related form, designated by HECKER (1923 , p. 3 7 , Pl. 2 , fig. 3 ) as E. cf. pogrebowi, origirrates from the Island of Vaigach, from Lower Llandeilian strata , since the horizon was said to be equivalent to the stage of Asaphus platyurus. Further, E. grandis was mentioned from the Rodman formation of Pennsylvania by FIELD (1919, p. 419 ) , thus from a horizon some­what below the Lower Ollasmops limestorre in Sweden (cf. above , p . 23 , foot-note 2 ) . The present writer has had no opportunity to form a personal opinion of the correctness of FIELD 's determina­tion.

Another species of Echinosphaerites , with large pyriform theca , from Västergötland, was described and figured by REGNELL (1945 , p . 152 , Pl. 8, fig. 6 ) , but not designated by a specific name on ac­count of its unsatisfactory state of preservation . It is possible that this form is conspecific with a form from the Kukruse of Estonia, described by SCHMIDT [1858, p . 221 (223 of sep. print) ] as follows : >>Eine grosse, dieser Species [E. aurantium] nahestehende, birn­förmige Art von l bis 2 Zoll Längsdurchmesser, grundet sich bisher auf nur unvollkommen erhaltene Exemplare. >>

FUNKQUIST (1919, p . 38) mentioned E. aurantium from the so­called Cystoid shale of Scania, which probably represents the Lower Ollasmops beds. The present writer (REGNELL 1945, p. 153 ) is not inclined to confirm the determination, not even the genus, but preferred (l. c . ) to refer to the species as >>Echinosphaerites? sp . >> .

Glyptosphaerites? sp . from the Chasmops limestorre of the Oslo region is younger than the East Baltic representatives of the genus . Y et within other areas - in Sweden as well - there are species of Glyptosphaerites comparable in age with the N orwegian on e, or younger ( cf. above, p. 22 ) .

::! 8 A R K I V FÖI� K E llf l , l\J I N E ia L O G I o . G E O I. O G I . n D 2 6 A . N : o 1 3 . Sphaeronites globulus (ANGELIN ) is very common in the Lower

Chasmops limestone of northern Öland, but has been found also in a few other regions in Sweden (cf . above, p. 21 ) . Apart from BRÖGGER's (1882 , p. 42 , foot-note l ) statement of a variety of S. globulus from the Oslo region, the reliability of which the writer has had no opportunity to control , the species has not been re­corded from outside Sweden. The affinities of S. shitienensis (REED ) from West Yunnan (REED 191 7 , p . 12 ) and Karakoram (GORTANI 1934, p. 26 ) , appearing at a stage with a supposed equivalence to the Platyurus limestone, seem to be with S. globulus rather than with S. pomum (cf . REGNELL 1945, p. 170 ) .

According to data published (cf . REGNELL 1945, p . 171 ) , the genus Haplosphaeronis has been met with in Seandinavia and, possibly, in Yunnan ( >>Sphaeronis>> lobiferus REED) , but is also present in the Leningrad Province, according to kind information (in litt . ) from Professor R. HECKER of Moscow. The stratigraphic range of the East Baltic Haplosphaeronis is unknown to the writer. The contingent East Asiatic species is older than the Scandinavian ones. In Sweden, Haplosphaeronis appears first in the Lower Chasmops limestone, has its main distribution in the Maeronrus beds (Caradocian) , and still survived during the formation of the Lower Boda reef limestone of Dalarna (mainly Ashgillian ) . In Norway, Haplosphaeronis occurs in the Sphaeronid shale and lime­stone of Hadeland which constitute a northern facies of the Upper Chasmops limestone (4h&) . These deposits have been sug­gested to be correlated with the Kullsberg limestone of Dalarmt (STORMER 1945, p. 397 ) , which is a reef facies of the stratified »normal» Maeronrus beds . !

Summing up the principal conclusions to be drawn from this review of the non-crinoid fauna of the Upper Llandeilian of Scandi­navia, we find that the majority of its elements are of a distinct East Baltic type. It is true that certain forms stand fairly isolated and that the species common to Seandinavia and the East Baltic Provinces are easily counted, yet the assemblage of genera is largely the same. It may be mentioned that also the crinoids offer points of contact , as demonstrated by closely related forms of Hoplocrinus occurring in the Lower Chasmops limestorre on the

1 The writer takes advantage of the opportunity to correct a disturbing mi sprint on p. 1 7 4 in his memoir of 1 945. In Iine 12 from a bo ve i t is state d that the Norwegian Haplosphaeronis harizon forms the top of the >>Ordovician se ­quence•>, which should of course read >>Middle Ordavieian sequence», according to the designatians of the various portions of the Lower Paleozoic deposits adopted in that paper, REGNELL 1 945, pp. 63-64. This also agrees with the view advocated by ST0R!HER 1 945, p. 396, that the Tretaspis shale should be regarded as the base of the Upper Ordavieian of Scandinavia, sinc 3 in »the southern part of the Oslo Region, and probably also in Sweden, the typical black Tretaspis shale with Tretaspis seticornis forms a characteristic transgression zone following an interval of no sedimentation>>.

G E R H A R D R �; G N E LT,, N O N - C R I N O I D P E L M AT O Z O AN FA UN AS . 29

orre hand, and in the Kukruse, on the other (REGNELL 1948 b ) . W e note further that a faunal communication existed between the Baltic area and the far East, via Central Asia, as indicated by the occurrence of genera common to the two regions (Hemicosmites , Stichocystis , Heliocrinites , Echinosphaerites, Sphaeronites, ?Haplo­sphaeronis). On the basis of the Llandeilian cystoid fauna of Kweichou described by him, SuN (1936, p. 4 7 7 ) recognized >>a certain connection between Bohemia and China during Middle Ordavieian time; and makes it probable that the fauna originated in the Indo-Pacific Ocean and then migrated westward to Bohe­mia, bu t never extended in to the Baltic province [ concluded at from the fact that Aristocystites is present in the East Asiatic and Bohemian faunas but absent from that of the East Baltic region] . This supposition is greatly strengthened by the fact that the Aristocystis fauna of S. China occurs earlier (Llandeilian) than that of Bohemia (Oaradocian) . >>1 The particulars of the account quoted may give rise to some discussion. First, the evidence of Hemicosmites , Stichocystis , and Heliocrinites must not be over­laoked since they are characteristic of the Baltic cystoid fauna, but cannot be said to be characteristic of the Bohemian, the two first-mentioned genera not being represented at all . Secondly, as mentioned above {p. 22 ) , the East Baltic and the Bohemian Basin may not have been separated altogether but seem to have communicated on some occasions, in Lower and Middle Ordavieian time, presurnably via Poland. Several East B altic Llandeilian genera appearing in Garadocian and Ashgillian strata of western and south-western Europe may have migrated along this route. SuN may be right or not in assuming that the Aristocystites fauna originated in the Indo-Pacific Ocean.2 By bringing the East Baltic Basin, too, in connexion with the East Asiatic Sea, we must not take it for granted, however, that the fauna of the Baltic region is of Indo-Pacific origin. Both regions may have received immigrants from the Arctic Sea, from where faunal elements were brought to North America as well (cf. the map of KoBAYASHI 1936, p. 491; see also above, p. 24 ) . Anyhow, the exchange of faunal elements between Europe and Central Asia is a significant feature of various periods of the Lower Paleozoic. Thus it was suggested e. g. by KoBAYASHI (1936, p. 490) for Middle Gambrian -Lower Ordavieian faunas. The same was pointed out by TROEDssoN (1937 , p. 14 ) for the Oambro-Ordovician faunas of Eastern T 'ien-shan, and is evident from the Middle Ordavieian

1 According to CHAUVEL ( 1 94 1 , p . 64, etc . ) , all the Bohemian species of Aristocystites revised by him originate from dE1, which may fall within the Upper Llandeilian.

2 SuN's view was not corroborated by CHAUVEL ( 1 94 1 , p. 70) , who pointed out that it does not agree with the fact that Aristocystites appears in the Armo­rican Llandeilian.

30 Aim i V FÖit K E M I , M I N E R A L O G I O . G E O L O G I . BD 26 Å . N : O 13.

faunas of Karakoram described by GoRTANI (1934) . Dealing with the Siluro-Devonian faunas of Eastern T 'ien-shan the present writer arrived at a similar conclusion (REGNELL 1941, p. 5 7 ) .

Caradocian.

In Scandinavia, the Caradocian cystoid faunas are not very different from the Upper Llandeilian ones . A certain decrease is con­ceivable , however, less with regard to the quantity of individuals than with regard to the multitude of species. Most of the genera are the same, which is also true of several species. Y et a new ele­ment is brought in with the first representative of the Blastoidea.

All Caradocian Hydrophoridea described hitherto from Sweden originate from the Maeronrus beds of Dalarna (a cystoid fauna of Slandrom ls . age has not yet been studied) . They are as follows:

Hemicosmites extranous EICHWALD, Caryocystites lagenalis REGNELL, Stichocystis geometrica (ANGELIN ) , Echinosphaerites aurantium (GYLLENHAAL) , E. aurantium f . , E. grandis JAEKEL? , Glyptosphaerites suecicus (ANGELIN ) , Sphaeronitcs? sp. (REGNELL 1945, p. 171 ) , Haplosphaeronis oblonga (ANGELIN) ? . >>Megacystis ovalis>> ANGELIN (cf . REGNELL 1945, p. 184) appears in the >>Lep­taena limestone>> , according to LINDSTRÖM (1888, p . 23 ) ; we do not know, however, whether it was derived from the Ordovician or the Silurian portion of this complex.

The Upper Chasmops series (Macrourus beds) of Östergötland has yielded a unique specimen of Paracystis ostrogothicus SJÖBERG. Quite recently, through the courtesy of Mr. JAANussoN of Uppsala, the present writer got the opportunity of examining an echinoderm fragment from Macrourus beds in the southern quarry at Am tjärn, Dalarna. This small fragment seems to be referable to Paracystis.

The only cystoids known so far from the Caradocian strata of Norway are Echinosphaerites sp. , Haplosphaeronis kiaeri JAEKEL and H. kiaeri norvegicus JAEKEL. For a discussion of their rela­tions to the Swedish H. oblonga, see REGNELL 1945, pp. 173-17 4 .

There is hardly any need to comment further upon Echino­sphaerites aurantium, E. grandis , Haplosphaeronis oblonga, H. kiaeri (from the >>Sphaeronid shale>> of Hadeland) , H. kiaeri norve­gicus (from the superimposed >>Sphaeronid limestone>> ) , since all of them were treated above in disenssing the Upper Llandeilian forms.

H emicosmites extranous is common to Sweden and the East Baltic Provinces . As stressed by the present writer (REGNELL 1945, pp. 101, 102 ) , statements are not univocal concerning the stratigraphic distribution of this species in the East Baltic area. According to JAEKEL (1899 , p . 310) and BASSLER & MOODEY (1943 , p. 165) it occurs in stage D 1 (Johvi-Jewe) . This stage has to be

G E R H A R D R �: G N E LL , N O N - C R I N O I D P E L M A T O Z O A N l<' A U N A S . 3 1

correlated with a lower part of the Maeronrus beds, which seems to be the horizon of H. extraneus in Sweden as well. The species, which is the last representative of its genus in Scandinavia, is not known from any extra-Baltic area. In Estonia, Hemicosmites surviv ed until Silurian time (H. grandis J AEKEL, F 2 , Porkuni­Borkholm) . H.? loczyi JAEKEL from Yunnan, based on some detached plates , was said to have been derived from an Upper Ordavieian rock (JAEKEL 1899, p. 315; BATHER 1906, p. 29). Our information regarding this form is insufficient, however, H.? loczyi being in fact a nomen nudum. Some forms referred to Hemi­casmites were described by FORBEs (1848, pp. 510-511 ) , and have been mentioned by several subsequent writers. Their systematic position is uncertain and requires a thorough revision, which is also true of the rest of the British cystoid material , except that treated by BATHER (1913). According to JAEKEL (1899, p. 314), H.? squamosus FoRBEs does not fit in to the plan of organization of Hemicosmites and allied genera. As regards H. ? oblongus (PANDER) quoted by FoRBEs, cf. above, p. 16 , foot-note 4 . H. rugatus FORBEs ( syn. H. pyriformis FORBEs) was referred to Oocystites by DREYFUSS (1939, p. 129).

Two specimens of Stichocystis geometrica (ANGELIN ) 1 recently placed at the author's disposal by Mr. JAANussoN come from the upper part of the Lower Maeronrus beds . The material available previously was not determined so exactly with regard to horizon. Apart from some specimens obtained from erratics in north Ger­many recorded by JAEKEL (1899, p. 327), the only report on spe­cimens possibly belonging to this species is that of SuN (1936, p. 483), from the Llandeilian of Kweichou in China (cf. above) .

Garyocystites lagenalis REGNELL has been recorded from Da­larna only, but it is so characteristic of the Maeronrus beds (espe­cially the lower ones ) of that region that it should be regarded as an index fossil. The species seems to be endemic to Sweden.

Glyptosphaerites suecicus (ANGELIN ) is not very well known, either with regard to morphology or to stratigraphic range. It occurs exclusively in Dalarna and is the last species of Gly pto­sphaerites in the Scandinavian-East Baltic region.

Haplosphaeronis oblonga (ANGELIN) is present at least in the lower and middle Maeronrus beds .

>>Megacystis ovalis ANGELIN>> is undefinable, since no material referable to it could be recognized (cf . REGNELL 1945, p. 184). Since its stratigraphic appearance is not known either, it was thought better not to record this form in the table showing the stratigraphic distribution of Scandinavian non-crinoid Pelmatozoa.

1 In the author's recent description o f Stichocystis geometrica a n error un · fortunately crept in (REGNELL 1 945, p. 1 09, line 1 2 from the top) . In the line quoted, read h y d r o p o r e for g o n o p o r e .

32 A R K I V FÖR K E lll l , 1\I I N E R A I. O G I O . G E O L O G I . ll D 26 Å . N : O 13.

Finally, the coronate blastoid Paracystis has not been found outside Sweden and is extremely rare in this country, too, as men­tioned just above. It seems to be fairly isolated , but its affinities are with the Stephanoblastidae (cf. REGNELL 1945, pp. 189-190 ) .

As we have seen, the general character of the Caradocian non­crinoid pelmatozoan fauna of Seandinavia is not very divergent from that of the Upper Llandeilian. Thus it follows that the echinoderm fauna is first of all comparable with that of the East Baltic Provinces , although it must be said to be definitely im­poverished . A certain local development is also recognizable if we consider the predominance of Caryocystites and Haplosphaer·onis , which play a very unimportant role in the East Baltic area.

The Scandinavian-Baltic Caradocian faunas now under dis­cussion have little in common, however, with the contemporary echinoderm faunas of the rest of Europe and with those of Morocco . The Bohemian Basin and the Massif armoricain of Brittany are characterized by an Aristocystites and Craterina fauna which is very much the same in the two districts (cf . CHAUVEL 1941, p. 252 ) . Moreover, the latter fauna was essentially developed already in Upper Llandeilian time, being present in Portugal (DELGADO 1908 ) , Spain, and Morocco as well. According to SPENCER (1938, p . 415 ) , the Bala echinoderm faunas of Wales and Shropshire are related to Asiatic and Bohemian orres on the one hand and to Trenton faunas of Canada. and Kentucky on the other. From Ireland two species of Echinosphaerites were mentioned by J AEKEL (1899, p. 337 ) , viz . E. granulatus McCoY and E. globosus anglicus JAEKEL, the latter considered as closely related to E. globosus JAEKEL from Estonia. As to Wales, some species of Baltic aspect were listed by BASSLER & MOODEY (1943, p. 40 ) . These are Helio­crinites balticus (EICHW ALD) , H. granatum (W AHLENBERG) , Echino­sphaerites aurantium (GYLLENHAAL) , Eucystis� litchi (FORBEs) , and three species referred t o Hemicosmites ( a s regards the so-called Hemicosmites, cf. above, p. 31 ) .

The determination of Heliocrinites balticus may be correct. An impression of a part of the test in limestorre shale from Bwlch-y­Gaseg, North Wales (the TÖRNQUIST collection, Paleontol. Instit. , Lund) , is the only specimen that the present writer has had an opportunity to examine. This , however, seems to be referable to the species mentioned. l The forms of FoRBEs (1848) and sub­sequent authors denominated Caryocystites granatum and O. davi­sii McCoY probably also belong here (JAEKEL 1899, p. 330; BATHER 1906, p. 1 7 ) . As regards the British Echinosphaerites aurantium,

1 It is possible that this is the specimen alluded to by TöRNQviST ( 1 879, p . 7 1 ) , who was inclined to refer it to Echinosphaerites aurantium. On the label written by him accompanying the specimen it is designated as >>Echinosphaem baltica EICHW.>> , however.

G E R H A l� D l� E G N ELJ, , N O N - C R I N O I D P E I, ni A T O Z O A N l<' A U N A S . 3 3

the writer has had n o opportunity t o form a personal opinion. Eucystis� litchi, finally, is no typical Eucystis , bu t may be regarded - along with >>Caryocystites pyriformis>> and C. � munitus - as a species intermediate between that genus and Sphaeronites (JAEKEL 1899, p. 406 ) .

Though many points are still obscure concerning the British Garadocian echinoderm fauna, it does not seem unreasonable to assume that a communication existed between Great Britain and the Baltic region, principally on the evidence of H el iocrinites balticus . This species occurs in the Kukruse of the East Baltic Provinces , a stage which must be coi'related with the Upper (al­though not the topmost) Llandeilian. l It was observed by TÖRN­QUIST (1879, p . 71 ) already that faunal elements common to the Scandinavian-Baltic region and Great Britain (where the easterly facies is concerned) appear earlier within the former area. 2 Re­cognizing them as easterly migrants , he found this fact quite na­tural. The same problem was considered by MARR (1882 , p . 325) : vYho explained it by >>the remoteness of Britain from land in Arenig and Lower-Bala times ; for there are no shallow-water forms of },Tenig age, and very few of Lower-Bala age, in the southern parts of Britaim. And he had just stated that >>the migration of shallow­water forms seems to have taken place along the coast-lines , and to have proeeeded much more slowly than that of the deep-water forms>> .

Ashgillian.

In the Swedish Ordovician sequence, the Red Tretaspis shale and the Staurocephalus shale must be correlated with the Ashgil­lian, which is consequently true also of their equivalent in reef facies, the Lower Boda reef limestone of Dalarna. Whether the >>Middle Tretaspis limestone>> (JAANUSSON & MARTNA 1948) should be included in the Ashgillian or the Garadocian is not settled as yet . From this horizon there is some undescribed cystoid material.

In Norway, the Ashgillian is represented by the Tretaspis limestone and Upper Tretaspis shale, the Isotelns beds, and the Upp

_�r_most Ghasmops zone (stages 4c�-5a) . 3

1 FAURA Y SANS ( 1 9 1 3, p. 9 7 ) mentioned >>Echinosphaerites cf. balticus d 'EICHW. >> from a Caradocian greywacke at Montcada, province of Barcelona. The present writer cannot j udge to what extent this statement can be trusted, but it cannot be accepted unreservedly, since the same species (with a mark of interrogation, that is true) was Iisted (op. cit . , p. 1 1 2 ) from Tremadocian strata.

2 Since the Ashgillian series was not yet recognized (established by MARR, in 1 905) TöRNQUIST ( 1 879, p. 70) referred the Tretaspis shales to the Caradocian, which can be accepted only in part to -day. It should be noted that in the paper quoted TöRNQUIST (1. c . ) presented creditable ideas of the alte rnating East Bal­tic and British invasions in the Scandinavian fauna during the Lower Paleozoic.

3 The upper part of 5a (hearing i . a. Dalmanitina mucronata) probably belongs to the Silurian (cf. STORMER 1 934, pp. 24, 30) . As regards the lower boundary of the Ashgillian in Norway, cf. STORMER 1 945, p. 397 .

A rkiv för kemi, mineralogi o. geologi. B d 2 6 A . N : o 13. 3

34 A R K I V FÖ R J{ E: M I , lll i N E R A L O G I O . G E O L O G I . B D 2() Å . N : O 1 3 .

The impoverishment of the Scandinavian non-crinoid pelmato­zoan fauna perceivable in the Caradocian proceeded, and is de­monstrated clearly in the Ashgillian. The present writer is not inclined to ascribe this fact in the first hand to conditions of litho­logic facies in this case either, but to a general tendency affecting the non-crinoid pelmatozoan stock in Upper Ordovician time within several areas . l The Saunja and Vormsi stages, forming the top of the Ordovician sequence in Estonia (cf. JAANUSSON 1944 ) , have yielded the genus Glaphyrocystis only (two species ) , and the non-crinoid pelmatozoan fauna of the Bohemian Basin - honsing previously a multiform assemblage - is remarkably poor. In North American Cincinnatian faunas Hydrophoridea are entirely absent, the non-crinoid pelmatozoans being represented by some genera of Edrioasteroidea and Cyclocystoidea (BASSLER & MOODEY 1943, pp. 32-33 ) . Fairly rich non-crinoid pelmatozoan faunas are found, on the other hand, in Languedoc, in the Girvan district , and in Algiers .

The cystoids of the Ashgillian of the Montagne N oir were studied i. a. by KOENEN (1886 ) and DREYFUSS (1939 ) . This fauna contains genera such as ,Juglandocrinus, Corylocrinus, and Oocystis. Certain elements of the Languedoc fauna are known from other regions as well, namely Portugal, the Pyrenees, Sardinia, the earnie Alps, and India (DREYFUSS 1939, p. 131, with further references; to these we have to add the Urals, from where Corylo­m·inus was recorded by YAKOVLEV, 1940, and Algiers2) . This sug­gests that a communication between these regions and Languedoc existed in Ashgillian time. DREYFUSS (1943 , p. 132) found in­dications of a connexion with Great Britain as weiL

According to SPENCER (1938) , the starfish Bed fauna of Girvan is related to the Richmondian of the Ohio B asin together with a new fauna, where the starfish are concerned, being quite different from the Welsh-Bohemian faunas. >>Cystid fauna has same two constituents, i. e. American and a new fauna which is really an old fauna as developed in Languedoc. The number of peculiar genera common to both is very remarkable. An explanation might be an Eastern reservoir sending migrants by an Arctic route to Girvan. starfish are not known from the contemporary \Velsh, Irish or Baltic beds. Cystids in these three areas are as a whole different

1 The author hopes to get an opportunity of discussing in a forthcoming paper the interrelations of fluctuations in the intensity of non-crinoid pelmato ­zoan faunas and major events in the earth's history.

2 Mme TERMIER has been so kind as to place at the writer's disposal a manuscript entitled >>Decouverte de I 'ordovicien en Algerie Atlasienne>> by A. LAMBERT, H. TERMIER and G. TERMIER. In this paper the following Hydro­phoridea are announced: Heliocrinites rouvillei (KOENEN), Juglandocrinus cf. erassus KOENEN, Echinosphaerites sp. , and Caryocystites sp. They were as ­sociated with Nicolella actoniae (SowERBY).

G E R H A R D R E G N E L L , N O N - C R I N O I D P E L JH A T O Z O A N F A U N A S . 3 6

from those o f Girvan and have a common element, apparently Baltic in origin» (SPENCER, l. c . ) .

I t was mentioned above (p. 2 8 ) that Haplosphaeronis persisted during the Upper Ordovician, H. oblonga appearing in the Lower Boda reef limestone. From Dalarna there are another two forms, Caryocrinites septentrionalis REGNll�LL and O. ? sp. (REGNELL 1945, p . 106 ) , both of which may come from strata belonging to the horizon of the Red Tretaspis shale. Eucystis quadrangularis REGNELL is known to appear in the Lower Boda reef limestone (REGNELL 1945, p. 183 ) .

With our present knowledge, we must imagine that Caryo­crinites is a migrant from East .Asia, for the oldest known member of this genus is O. aurora (BATHER) from the Llandeilian Naung­kangyi beds of Burma (BATHER 1906, p. 2 9 ) . C. ? sp . (GORTANI 1934, p. 25 ) from Karakoram may be approximately contempora­neons with the species mentioned. Å form designated as >>Caryo­crinus ornatus SAY >> was reported by DELGADO (1908, pp. 62, 83) from the sections at Bugaco and in the Tage Basin (from here a >>Caryocrinus? sp. >> as well ) . The horizon was said to be Upper Ordovician, but judging from the associated echinoderm fauna it belongs to the Upper Llandeilian or Caradocian. In any case the Niagaran O. ornatus SAY of North America cannot be taken into account. l All other species of Caryocrinites known so far are Silurian. >>Caryocrinus sp . >> (KIJER 1897 , pp. 1 7 , 75 ; cf. also REG­NELL 1945, p . 105 ) from stage 5b in the Oslo region shows that the genus survived for some time in Scandinavia. According to a statement by F . SCHMIDT (1858, p . 221 ; 223 of sep. print ) , Caryo­crinites ornatus appears in stage 7 2 of Estonia. This statement seems to have been overlooked altogether by subsequent writers, and it is impossible to judge of its valne. SC:Hl\iiDT did not doubt the correctness of the generic determination, however, an in­complete theca being available to him >>der wenigstens die Gattung sicher erkennen lässt>> . Apart from the forms just mentioned, all Caryocrinites are from the Niagaran of N orth America, being represented by a considerable number of species.

TROEDssoN (1918, p . 21) recorded >>Caryocrinites>> sp . from the

1 BAsSLER & MooDEY ( 1 943, p. 138) list a further Ordavieian species, C. buchianus (HAECKEL, sub Enneacystis) from »Russia (vicinity St. Petersburg) >> . However, the erection of this species (as weil as of the genus Enneacystis) was a complete mistake, as pointed out by JAEKEL ( 1 897 , p. 392) : >>Die folgende neue Gattung Enneacystis stellt ihr Autor auf Formen auf, die angeblich von v. BucH aus dem Untersilur von Russland beschrieben sein sollen, in Wahrheit aber gar nicht existieren, denn die citierten Exemplare, die v. BucH beschrieb, gehören sämmtlich dem typischen Caryocrinus ornatus SAY aus dem Obersilur von Lockport im Staate New York an. >> C. buchianus, therefore, is a synonym of C. ornatus.

2 This earresponds (F. SCHMIDT 1 882, p. 527) to stage I, or the Lower Oesel group, of Wenlockian age.

3ti .A i m l V l<'ÖR K E M I , M I N im A L O G I O . G E O L O G I . B D 26 A . N : O 13 .

zone of Staurocephalus clavifrons at Tommarp in Scania. The present writer (REGNELL 1945, p . 137 ) referred the unique spe­cimen to a new species of Heliocrinites (not named) . A few more come from the Seanian Staurocephalus shale: >>Echinosphaerites sp . >> (TROEDSSON 1918, p . 23 ) , which is no Echinosphaerites (REGNELL 1945, p. 153 ) , and Eucystis? sp. (REGNELL 1945, p. 184 ) . It is not necessary here to comment further on these forms.

The only species reported from the Ashgillian (5aj l of Norway is Heliocrinites cf. balticus EICHWALD (KrAm 1901, p. 11 , sub Echinosphaerites), which was said to occur very commonly at Djupalstenen in Asker. In view of the divergent stratigraphic range of Heliocrinites balticus in the East Baltic area (cf . above, p. 33 ) , it is better not to say anything definite until the actual fossil material has been examined.

Summing up, the Ashgillian cystoid fauna of Seandinavia does not show any close relationship to the contemporary fauna of the East B altic Provinces , nor to that of any other region. But i t may be suggested, in view of the stratigraphic range and regional distribution of Caryocrinites in Eurasia, that the richly differen­tiated Middle Silurian Caryocrinites fauna of N orth America, devel­oped from immigrants from the sea covering part of Seandinavia in Upper Ordovician time. This migration may have taken place via the Arctic route, but it is more likely that it went across the Atlantic, for there are reasons to believe that the barrier separat­ing the South Scandinavian-Baltic region from the American Arctic one did not exist any longer (HoLTEDAHL 1920, p. 7; 1924, p . 122; TROEDssoN 1928, p . 186 ) . A recent support of this view is the observation by STORMER (1945, p . 398) that the Tretapsis faunas of Hadeland contain some North American trilobite genera. One form is identical even as to species.

S i l u ri an . The transition from the Ordovician to the Silurian system

eaused no great changes in the character of the non-crinoid pel­matozoan fauna of Scandinavia. Some elements are new but others are descendants from Ordovician ancestors within the South Scandinavian-Baltic Basin (cf . , however, the reservation below) .

A s regards Sweden, a certain rise of the fauna just under con­sideration is noticeable in the Lower Silurian, with regard to both number of species and individuals. In Norway, however, this part of the sequencc has yielded hitherto one species only, whereas no species have been recorded from younger strata. On account of the predominance of graptolite shales above the Dalmanitina series

1 N.B. above, p. 33, foot-note 3 .

G E R H A l{D It E G N E L L , N O N - C R I N O I D P E L M A T O Z O A N FA U N A S . 3 7

within most Silurian districts in Sweden, younger non-crinoid pelmatozoan faunas are known solely from Gotland. Contrary to expectation no material of non-crinoid pelmatozoans has been obtained from the so-called Pentamerus limestorre of Jämtland (mainly of Lower Handoverian age) ; nor is there any material of non-crinoid pelmatozoans from the Öved-Ramsåsa series of Scania [Upper Ludlow(-Downtonian? ) ] .

Llandoverian.

The only hydrophorid reported from the Silurian of Norway is Caryocrinites s p . (KiiER 1897 , pp. 1 7 , 7 5 ) w hi ch was f o und in the basal Silurian stage 5b (N.B. above, p . 33, foot-note 3 ) . This species has already been referred to above (p. 35 ) .

The lowest portion o f the Swedish Silurian sequence i s made up of the Dalmanitina series . In Dalarna, from where the main part of the non-crinoid pelmatozoan material comes, the Dalmanitina shales are replaced partially by the Upper Boda reef limestone. The stratigraphic position of the reef limestorre eomplex, formerly known as the >>Leptaena limestone>> , was not fixed in its main features until fairly recently. The Leptaena limestone turned out to have been formed during various periods of reef-building, alternating with periods of mormah sedimentation . Besides the Silurian part, as we have seen, it comprises two Ordavieian parts: an Upper Ordavieian part, the Lower Boda reef limestone, equi­valent mainly to the Staurocephalus and Red Tretaspis shales , and a Middle Ordavieian part, the Kullsberg reef limestone, equivalent to the principal part of the stratified Maeronrus beds . From the old museum labels , hearing the designation >>Leptaena limestone>> , it is often impossible to form a definite opinion on the age of the fossils conccrned, especially in those eases where no loeality is given. It must be admitted, therefore, that the forms from Dalarna referred to below may occur in the Upper Ordavieian portion of the Boda reef limestone as well, or perhaps exclusively within that portion. If that be so, there would naturally be a fairly pronouneed faunistical break between the Ordavieian and the Silurian systems where the non-erinoid pelmatozoan fauna is eoncerned.

The following forms have been found in Dalarna: Heliocrinites stellatus REGNELL, H. variabilis REGNELL, Eucystis raripunctata ANGELIN, E. angelini RECiNELL, E. acuminata REGNELL, E. quadr­angularis REGNELL (cf . p. 35 above) , E. sp . (REGNELL 1945, p . 183 ) , Sphaeronites? sp . (REGNELL 1945, p . 184) , >>Sphaeronis? dalecarlica>> ANGELIN, Tormoblastus bodae J AEKEL, Cyathotheca suecica JAEKEL , Cyathotheca? sp. l

1 This species from Boda (TöRNQUIST collection, Paleontological Institution of Lund University) has not yet been described.

38 A R K I V F Ö R K E lii i , lii i N E R A L O G I O . G E O L O G I . BD 2G A . N : O 1 3 .

From the Dalmanitina series o f Västergötland Holocystites? sp . (REGNELL 1945, p . 183, = >>Megacystis alternata HALL var. >> , ANGELIN 1878, p . 30) , and an undeterminable >>cystoid>> (TROEDS­SON 1921, p . 11 ; cf . REGNELL 1945, p . 184) have been reported.

Finally, in a boulder of so-called West Baltic Leptaena lime­storre from Hulterstad in Öland, there is an imperfect stem-frag­ment of Dendrocystites sp. (REGNELL 1945, p. 194) .

Of the non-crinoid pelmatozoans j ust mentioned, the two species of Heliocrinites are members of a genus well represented especially in the Middle Ordovician . H. stellatus and H. variabilis are somewhat different in type as compared with the Ordovician species, and are the youngest representatives of Heliocrinites known. The general distribution in time and space of the genus was reviewed above (p. 2 6 ) .

A s pointed out b y THORSLUND (1936 , p . 26 ) , the hydrophorid Eucystis is as characteristic of the Boda limestone as is Haplo­sphaeronis of the Kullsberg limestorre (we leave out of considera­tion the fact that Haplosphaeronis has been found also in the Lower Boda limestone) . There are several species of Eucystis , and some of them occur abundantly. Eucystis may have an easterly origin, for the oldest species possibly referable to this genus is a form from Yunnan described by REED (191 7 , p. 11 ) as Eucystis cf. raripunctata (Middle Llandeilian) . The specific determination is of course incorrect. Another doubtful form, coming next in age , is E. ? litchi (FORBEs, sub Caryocystites ; cf. above, p. 33 ) , from the Garadocian of Great Britain. E.? sp. from the Staurocephalus shale has been referred to above (p. 36 ) . In Dalarna the genus reached a rich development, but it has not been reported from the East Baltic area. From Seandinavia Eucystis spread over a good deal of central and western Europe, so that it is known (partially sub Proteocystites , Proteocystis , Carpocystites, and Carpocystis) from Lower Devonian deposits of Germany (JAEKEL 1899, p. 407 ) , Bohemia (JAEKEL l . c . ) , and France (FERRONI:ERE 1922, p . 7 2 ) . It occurs in the Devonian of Morocco as well (G. TERMIER, unpub­lished ) . Bulbocystis RuziCKA (1939, p . 292) from the Middle Devo­nian of Bohemia may be regarded as a descendant from Eucystis .

If the generic determination of Sphaeronites? sp. (REGNELL 1945, p. 184) can be trusted, it demonstrates that the stratigraphic range of Sphaeronites reaches considerably higher than the Middle Ordovician.

A very great number of species (about 50) of Holocystites have been described (mostly by S . A. MILLER) from the Niagaran of North America. H. alternata (HALL) , which ANGELIN (1878, p. 30) was inclined to regard as closely related to the Swedish species , occurs in the Racine and Cedarville dolomites (BASSLER & MoODEY 1943, p. 167 ) of the Upper Niagaran, corresponding to a middle or

G E R H A R D R E G N E L L , N O N - C R I N O I D P E L M A T O Z O A N F A U N A S . 3 9

upper part o f the Wenlockian (SWARTZ e t al. 1942 ) . If the con­siderably old er Swedish species is congeneric with the N orth American representatives of Holocystites , we must assume that these originate from migrants from north-western Europe.

The first blastoid entered the Swedish fauna in Garadocian time. In the Lower Silurian appeared the seeond representative of this subclass, which is one of the rarest of all fossil groups in the Swedish Paleozoic. Tormoblastus bodae J AEKEL, as well as the Garadocian Paracystis, may be derived from ancestral forms in the Bohemian Basin (the Middle and Upper Ordavieian genus Mespilo­cystites BARRANDE) . Here the Stephanoblastidae survived, devel­oping in to the r .. udlovian Stephanoblastus J AEKEL. At an earlier stage they had reached North America, as demonstrated by the multiform Niagaran Stephanocrinus CONRAD .

Also the Carpoidea and Edrioasteroidea are in Seandinavia confined chiefly to the Silurian part of the Paleozoic sequence. The only exception is the Middle Gambrian Stromatocystites.

Dendrocystites - one of the two carpoids known from Sweden - was distributed almost all over the world during the Ordavieian (see REGNELL 1945, pp. 194, 195-196 ) . The oldest species re­corded is D. vidali THORAL from the Upper Skiddavian of Herault . There are also some stem fragments from the Skiddavian of Korea which may belong here (KOBAYASHI 1934, p. 525 ) . Since the Swedish species seems to be fairly closely related to the Garadocian D. sedgwicki BARRANDE, it is possible that the genus invaded Sweden from the Bohemian Basin. But it may of course also have come either from the East Baltic Provinces , or from Scotland. The present species is one of the last members of Dendrocystites , being succeeded only by the Lower Devonian D. globulus DEHM of Germany (DEHM 1934, p . 20 ) .

In the case of Cyathotheca , its East Baltic origin cannot be doubted, the only Congeneric form being C. corallum (JAEKEL) from the Vaginatum limestone (Upper Skiddavian) of the Lenin­grad district .

We shall now have a look at the younger non-crinoid pelmato­zoans of the Silurian of Sweden, all of them found in Gotland. A few of these may fall within the Upper Llandoverian, viz . those probably derived from the Upper Visby marl . l The species can­cerned are Pyrgocystis cylindrica (AURIVILLIUS) and P. procera (AURIVILLrus) . The latter species occurs in younger strata as well where it is accompanied by a few more species belonging to the same genus. Pyrgocystis makes its appearance in the Skiddavian of the Leningrad Province (HECKER 1939) . 2 From the East Baltic

1 According to HEDE ( 1 942, p. 22 ) , the Upper Visby mar! is probably Upper Liandoverian in age.

2 A pyrgocystid, ))a very primitive form, as it seems•' (translated by the

40 A HI\ I V F Ö n K E lli i , ll! I N E R A L O G I O . G E O L O G I . BD 26 Å . N : O J 3 .

area Pyrgocystis spread to North America (P. sardesoni BATHER; Black River, 1\Iiddle Llandeilian; further P. batheri RUEDEMANK ; Cayugan, Bertie waterlime, Aymestry ) , Scotland (P. grayac BATHER; starfish bed , Ashgillian) , England (P. ansticei BATHER; Wenlockian) , Sweden, and Germany (P. octogona RICHTER; Lower Devonian) .

T o summarize what has been said above concerning the Liando­verian non-crinoid pelmatozoan fauna of Sweden, we note that the Hydrophoridea and Edrioasteroidea have their affinities with East Baltic, or, at any rate, easterly forms, just as during the Ordovician. Only Holocystites� and, to a certain extent, Eucystis are fairly isolated. A remarkable feature is that the Swedish cystoid faunas are much more developed than the contemporary East Baltic ones . The Blastoidea and Carpoidea - two groups really foreign to the Scandinavian Paleozoic fauna - quite likely invaded Sweden from the Bohemian Basin.

W enlockian.

The only hydrophorid hitherto recorded from Swedish 'Ven­lockian strata is the diploporit Gomphocystites gotlandicus (ANGE ­LIN } . This species appears in the Högklint group (Lower Wen­lockian) and the Slite group (W enlockian, not y o ung er than the zone of Cyrtograptus ellesi, both statements according to HEDE 1942 , p . 22 ) . Of the pcculiar Cyclocystoidea a few species have probably been obtained from the Högklint limestone: Cyclocystoides lindströmi REGNELL, C. insularis REGNELL, and a third species no t yet described. Finally, the Edrioasteroidea are rcpresented by three species, viz . Pyrgocystis procera (AURIVILLIUS) - a survivor from the Liandoverian - , P. sulcata (AURIVILLIUS} , and P. varia (AURIVILLIUS} . These are from the Mulde marl , which , according to HEDE (1942 , p. 22 ) probably earresponds in age to the Wenlockian zone of Cyrtograptus lundgreni or, alternatively, forms transitional beds to the Ludlovian .

• �part from the species of Pyrgocystis , which are descendants from ancestors of East Baltic origin (cf . yet below) , this Wen­lockian fauna offers an entirely new aspect as campared with earher non-crinoid pelmatozoan faunas. For both Gomphocystites and Cyclocystoides must be regarded as North American elements. In earher instances where we have found genera, or even species , to be common to Europe and North America, we have invariably had reason to believe that they appeared earller in Europe than they did in America. In the case just under discussion, on the other hand, the oldest North American representatives of the

present writer) was mentioned from Norway by JAEKEL ( 1 927 b, p . 4 ) , who dicl not, however, say anything about its stratigraphic horizon.

G E R H � l� D R E G N E L L , N O N- C R I N O I D P E L M A 'l' O Z O A. N F A O N A. S . 4 1 !l l l l

l N . A M E:. R I CA i E:.NG LAND J sCOTLAND G OTL Afi D B[LGIUM !

l -!-- t - � Lower Devonian l l

� l' l l il + -t- -r- -

L udlovian l l i l

Mnlocluan

Caradocwn

Llandedwn Cyc/ocys/oides Fig. l . Diagram showing the regional distribution and approximate strati­graphic range of the lmown species of Cyclocystoides (figures within a circlet represent species referred to Narawayella by some writers) . l, raymondi FoER· STE; 2, anteceptus HALL; 3 , salteri HALL; 4, halli BILLINGS; 5, cincinnatiensis MILLER & F ABER; 6, minor MILLER & DYER; 7, m undula MILLER & DYER; 8, nitida FABER; 9, parva MILLER & DYER; 1 0 , belZulus MILLER & DYER; l l , magnus MILLER & DYER; 1 2 , huronensis BrLLINGS; 1 3 , illinoisensis MILLER & GuRLEY; 14 , caractaci SALTER MS. ; 15 , marstoni SALTER MS. ; 1 6 , davisii SAV!' ER; 1 7 , decussatum BEGG; 1 8 , wrighti BEoo; 1 9 , insularis REGNELL; 20, lindströrn·i

REGNELL; 2 1 , sp. ; 22, sp. MAILLIEUX 1 926.

genera in question are prior to the European ones . As regards Gomphocystites, which is confined to North America and Gotland, this fact is not very prominent, however . l Y et the Osgood forma­tion of Indiana, from which G. indianensis S. A. MILLER originates , is partially older than the basal Wenlockian (SwARTZ et al. 1942, earrelation table) . G. tenax HALL, from the Lockport-Shelby dolomite of New York and the Louisville limestone of Kentucky (BASSLER & MooDEY 1943, p. 161 ) , seems to be fairly contempora­neons with the Gotland species . Three other Niagaran forms are younger. 2

'\Ve cannot ascertain what migration route Gomphocystites used to reach the waters covering Gotland of to-day. Perhaps we can form an opinion of this by considering the regional distribution of Cyclocystoides : �Iiddle Ordovician-Lower Silurian forms in North America, Middle-Upper Ordavieian forms, and a Lower

1 It should be emphasized again that a common centre· of evolution and dispersal could also be taken into consideration.

2 From the Middle Devonian of California the doubtful Gomphocystites ( 'l ) californicus 8TAUFFER has been recorded (cf. BASSLER & MOODEY 1 943, p . 1 6 1 ) .

i l l l

-+-1

i -,

42 A R K I V F Ö R K E U I , M I N E R A L O G I O . G E O L O G I . B D 26 Å . N : O 1 3 .

Silurian form i n England, Upper Ordovician forms i n Scotland, Middle Silurian forms in Sweden, and, finally, a Lower Devonian form in Belgium (BEGG 1939, p. 21 seq . ; BASSLER & MOODEY 1943, pp. 147-148; REGNELL 1945, p . 215 seq. ) . It seems justifiable to infer from these facts that the genus came to Gotland across the Atlantic. The systematic position of Cyclocystoides is disputable .

Also Pyrgocystis , in spite of its East Baltic origin , may fit into this pattern. As mentioned above, the genus spread over great areas , so as to be present in North America in Black River time. From here it may have been brought back to the Baltic Basin in Silurian time, via Scotland and England.

Ludlovian.

The gradual decrease of the non-crinoid pelmatozoan stock discernible in the Silurian of Gotland led to a complete extinction after the end of the Lower Ludlovian.

In Lower Ludlovian time the Hydrophoridea Rhombifera, which are absent in the lower strata of Gotland, became repre­sented by the genus Lovenicystis REGNELL, including the single species L. angelini (JAEKEL) . This species probably occurs in the Eke group (cf . REGNELL 1945, p. 92, foot-note 20 ) .

The explanation of this sudden appearance of a member of the Callocystitidae in the Upper Silurian of Gotland is undoubtedly that it is a migrant from the west, for the Apiocystinae are de­cidedly N orth American in origin. This subfamily commenced in the Upper Ordovician by Lepadocystis, followed in Lower Silurian time by Brockocystis . In Niagaran time, the Apiocystinae had split up into Apiocystites , Tetracystis , and Hallicystis , the first mentioned developing into Lepocrinites (Upper Silurian and Lower Devonian) and Lipsanocystis (Middle Devonian) . Jaekelocystis (Lower Devonian) seems to have developed from Tetracystis . All the genera mentioned are N orth American. Bu t in the course of time they extended their area of distribution. Thus Apiocystites and Lepocrinites are f o und in the W enlockian of England.

From this it might easily be assumed that the British Wen­lockian species had spread further eastwards , changing into the Lower Ludlovian Lovenicystis of Gotland. The morphological analysis shows, however, that Lovenicystis lies in the straight line of the Lower Silurian Brockocystis ( cf. the scheme published by REGNELL 1945, p. 91 ) . LUHA (1940, pp. 98-99) described an Upper Silurian callocystid from Oesel, which he called Lepocrinites kailuka. His report does not contain sufficient details to determine the exact taxonornie position of the species . It would not be surprising, however, if it turned out to be referable to Lovenicystis , demonstrating a migration of that genus eastwards . Lepocrinites

G E R H A R D J{EGN E L L , N O N - C R I N O I D P E L lli A. T O Z O A N l!, A UN A S . 4 3

N. A M ER I CA ENG LA N D GOTLAND

Upper Wen loclrian

11zdd/eWen1ocklan

Lower Wen

ljpperLlandov�rian Plsocri.nus

Fig. 2 . Diagram showing the regional distribution and approximate strati­graphic range of the known species of Pisocrinus [P. glabellus (RowLEY), N. America (Bainbridge ls. : Wenlockian) , and the doubtful P. ? yassensis ETHER­IDGE and P. ? yassensis lobatu.s ETHERIDGE, Y ass district, New South Wales (age of the Y ass Is. : Upper Wenlockian-Lower Ludlovian, according to HILL 1 943, p. 64) , have not been recorded in the diagram] . l, pyrijormis (RINGUE ­BERG); 2, bacula MILLER & GURLEY; 3, benedicti S. A. MILLER; 4, campana S. A. MILLER; 5, gemmiformis S. A. MILLER; 6, globosus (RINGUEBERG) ; 7 , g01·byi S . A. MILLER; 8, granulasus ROWLEY; 9, quinquelobus BATHER; 1 0, sphae ­J'icus ROWLEY; 1 1 , tennesseensis (ROEMER); 1 2 , pilula DE KONINCK; 1 3 , sp.

BATHER 1 893; 1 4, pocillum ANGELIN; 15, ollula ANGELIN.

kailuka was found in the Kaugatuma stage of the Upper Oesel group . .According to LUHA, this stage corresponds approximately to the Gotland strata bearing Lovenicystis.

It is hardly necessary to add anything to the remarks on Lovenicystis, since this genus is the only representative known of Ludlovian Hydrophoridea - and non-crinoid Pelmatozoa on the whole - in Sweden. It may only be pointed out once more that the Upper and Middle Silurian non-crinoid pelmatozoan faunas differ fundamentally from the Lower Silurian and Ordovician ones: Westerly elements have almost entirely replaced those of easterly origin. Taking the South Scandinavian-East Baltic province as a whole, the change in character may also be expressed thus ( somewhat generalized, that is true) : The province changed from an emigration area to an immigration one, as far as the group of animals now considered is concerned. ParaHel to this process there occurred a gradual decrease in the non-crinoid Pelmatozoa, leading to their total extinction in these waters in Ludlovian time. Their role was finished; it was taken over definitely by the Cri­noidea.

Let us in this connexion make a digression. It is quite in­structive to consider the Crinoidea as well with regard to migration

44 A R K I V F Ö it K E U I , l\I I N E R A L O G I O . G B O L O G I . BD 26 Å . N : O 1 3 .

N . A M E R I CA

Lower Lud!ovian

llpperWenlodian

/1iddle Wenlocklan

Fig. 3 . Diagram showing the regional distribution and approximate strati­graphic range of the known species of Pycnosaccus [P. tenuibrachiatus SPRINGER, Lower Devonian, and P. canadensis (WHITEAVES) , Upper Devonian, both N. America, have not been recorded in the diagram]. l, calyculus (HALL) ; 2 , laurelianus SPRINGER; 3 , dubius SPRINGER; 4, patei SPRINGER; 5 , americanus WELLER; 6, welleri SPRINGER; 7 , bucephalus (BATHER); 8, nodulasus ANGELIN;

9, scrobiculatus (HrsiNGER) ; 10, bucephalus bohemicus Bou.<KA.

directions , since they seem to conform very well with the ideas expressed above. The only crinoid group of the Ordovician of Sweden known tolerably well , the Hybocrinida, is decidedly East Baltic in origin (REGNELL 1948 b ; cf. also p . 28 above) , whereas the Silurian Gotland crinoids may have immigrated mainly from the west. Time is not yet ripe for a general ventilation of this question - nor is this paper the forum for it - but the statement is Supported by real facts in the few instances investigated. Some arbitrarily ehosen examples will prove this , here plotted in dia­grams for the sake of perspicuity. The diagrams are based mainly on records in BASSLER & JVIooDEY (1943 ) , completed by data con­tained in BousKA 1942 . The correlation between the North Ameri­can and British sequences is in accordance with SWARTZ et al. (1942 ) . Similar diagrams were constructed for Gissocrinus and Crotalocrinites (distribution in Bohemia according to BoUSKA 1943, 1946 ) . They point in the same direction as those of Pisocri ­nus and Pycnosaccus reproduced here.

Our thesis is corroborated by an example from the Asteroidea as well , viz . Palasterina antiqua (HISINGER) , which occurs in the upper parts of the Burgsvik sandstorre in Gotland (cf. MUNTHE 1921, p. 40 ) , of Upper Ludlovian age. The species, which , accord­ing to SPENCER (1922 , p. 220) , passes directly into the Devonian P. follmanni STURTZ, of Germany, is namely found in England and Scotland in Wenlockian and Lower Ludlovian strata (SPENCER 1922 , pp. 228-229) .

G ER H A R D l� E G N E L L , N O N - C R l N O l D P E L JII A T O Z O A N F A U N A S . 45

Silurian Undefined.

under this heading a few words may be said about Placo­cystites sp . from Gotland, since it is one of the two carpoids re­corded from Sweden. The author has not exaroined the specimen himself, and the stratigraphic position is not clear from the state­ments in the literature (cf. HEGNELL 1945, pp. 196-197 ) . Yet, it contributes a little to the characterization of the Silurian non­crinoid pelmatozoan fauna in Seandinavia and its migrations. The Gotland species is , as a matter of fact, closely related to Placo­cystites forbesianus DE KONINCK from the W enlockian of England. Hecently , CHAUVEL (1941 , p . 215 ) advocated that Anomalo­cystites bohemicus BARRANDE be referred to Placocystites . The species mentioned is from section d.-, (Ashgillian) of Bollernia and is thus the oldest representative of Placocystites . The migration history of Placocystites , therefore , is much the same as that of the Liandoverian Dendrocystites. In both cases , this foreign element was brought into the Swedish fauna by an episodic migration from the Bohemian Basin , conceivably via Great Britain .

Concl u d i ng Uemarl\s.

The author is well aware of the impropriety in drawing conclu­sions concerning migration routes and paleogeographical conditions on the basis of one group of animals only. But he hopes that this review of the succession of the significant non-crinoid pelmato­zoans within a restricted area and th e discussion on their relations to contemporary assemblages will contribute to some extent to the understanding of the faunas contained in the Old Paleozoic sequence of Scandinavia.

In conclusion, the writer wishes to point out a few facts which have bearing on discussions like that presented on the preceding pages .

First, we have to remember that all results derived from a study of fossil faunas are liable to envolve errors and roistakes due to the incompleteness of the geological record.

Secondly, we must realize that identical environmental con­ditions may produce similar morphological types independently in different areas. If such a phenomenon as that indicated here is not merely hypothetical, the very principle of drawing paleo­geographical conclusions on the basis of identical (morphologically but not phylogenetically) forms appearing in different regions would be disturbed seriously. This matter was considered by SIMPSON (1947, p. 622 , foot-note 1 ) , who was of opinion that it must be )>highly exceptional and can be ignored in an over-all

46 AR K I V FÖR K E M I , MIN ERA L O G I O . G E O L OGI. ll D 26 A . N : O 13 . g [/re qrouv � ' � Hemse gro up

-l I K/rn feberq grouv Nu /de m a r!

i l Hal/o Is. --�--� � � -:�0_ �rouo --�-0:: � ! lOlla !s ::l h'o q.41int gro up _J 1--+_.:._ _ __:c___---+-<rl " l Uvver lh sby m o r!

i l L o wer Vuby ma�0

' /'z _ _ EJ

Q !:. !

1 /T'a. .s irdes ser-z-es ] _ _

- Dalmandzno--;-;·;-:;�� !12 (,'IÖ��-0 c: 5fmA.TOCeDlu:du.; s:� -- (�/� : 1 Q 1------------·---c-

g B!aclr Tre lo J Dl .J -.� 1 { Slo. Ti drom Is 1 � l Ma( rouru-;zea1 J - -t Up�r Clta�-moJJsl.r t....'.1L_

ri ;,�;:l�:�:,�:,0s l c Z: Cystoz a' lr "- l � t L u -,e• C'�asmo .>.; l,

L Cras.ncaudu slo. qe 1 u o

; l � R f 0 _::3 latyu r u s s a:ge a: o

� A sanhu..r bed.: � Lt m. h a. ta y/aqe- _____l_ <A Pla ndt m 6cda stogl' ! ,.g Ce ro.fovyqe is: & sh.. 1 � Dzcfyo n e m a sit

.o

0 �/-_. Y2 ; ·,> L I'z

<>- - - - - 0 Locrin o{cieci

Hydrovh oridea

- ·8- - - - - -�- - Blasfoidea:

1 ____ J __ _L __ __.. __ __L___.L_ L .-J ___ �__..!...._..._. s • r • • m u u n � � M " M � N �

D Carpoideo: - -8- - - - - - -®- - [drioasferoidea

o Cyclocy..rloid e a .

Fig. 4 . Graph o f the stratigraphic distribution o f the non-crinoid Pelmatozoa in the Paleozoic sequence of Sweden, with reference to the number of species.

view>>. The present writer cannot see any reason to object to this opinion.

Thirdly, geography of the past is conceived on the modern base of the continental areas. This , however, >>is a greatly forshortened one in all the places where the roots of the ancient mountains and the peaks of the modern ones occur, and consequently the geo-

G E R H A R D R E G N E LI. , N O N - C R I N O I D P E I. M A 'f O Z O A N F A U N A S . 47

graphy of the present gives a more or less erroneous picture of the actual size and shape of the ancient seas in the areas older than the times of mountain making>> {SCHUCHERT 1929, pp. 118-119) .

Finally, it should be noted that peculiarities in the regional distribution of a taxonornie unit ascribed to the existence of a land barrier may very well in certain cases be due to great depths . For zoogeographically great depths of the sea are as great obstacles for the migration of many marine animal groups as are land barriers. This fact has been pointed out repeatedly, e . g. by TROEDBSON (1928 , p . 176 ) and by R. & E . RICHTER {1941 b , p . 31 ) .

The stratigraphic distribution of the non-crinoid Pelmatozoa so far known in the Paleozoic sequence of Sweden has been sum­marized in fig. 4. The graphs would probably have had a some­what different shape, had it been possible to represent the length of time required for the formation of the several stratigraphic divisions. Unfortunately, however, our present knowledge does not admit of the calculation of this factor.

The stratigraphic range of the individual species is in most cases restricted to one division only. So a graph showing the number of new species in each division is talerably in conformance with the graph representing the total number of species .

L i t e r a t u r e : Angelin, N. P. , 1878 . Iconographia crinoideorum in stratis Sueciae siluricis fossilium. Opus postumum edendum curavit Regia Academia Scientiarum Suecica. Holmiae . Pp . IV + 1-32. Pls. 1-29. (Pp. 28-32: Cystidea, ed. S . Lov:EN. ) - Asklund, B . , 1 929. Norrlands strandflate. Geol. Fören. Förh. 5 1 . Stockholm. Pp. 1 33-145. - --, 1 938. Hauptzi.ige der Tektonik und Stratigraphie der mittieren Kaledoniden in Schweden. Sver. Geol. Unders. Ser. C. 417. Stockholm. Pp. 1-99. Pl. l . Tabs. 1-2. -Bailey, E. B. & Holtedahl, 0 . , 1 938 . Northwestern Europe Caledonides. Reg. Geol. Erde. (Herausgegeb. von K. ANDREE, H. A. BROUWER u. W. H. BucHER . ) 2 : 2 . Leipzig. Pp. 1-76 + Index. Pls. 1-2 . - B arrande, J., 1887 . Classe des echinodermes. Ordre des cystidees. Ouvrage posthume de feu JoACHIM BARRANDE, publie par le Doct. W. WAAGEN. Syst. Sil. Centre Boh. 1ere partie. 7 : l . Prague. Pp. xvu + 1-233 . - Bassler, R. S . , 1 9 1 5 . Bibliographic index of American Ordavieian and Silurian fossils . 1-2 . Smiths. Inst. U. S . N at. Mus. Bull . 92. (2 pts. ) Washington. Pp. vm + 1-152 1 . Pls. 1-4. ---, 1 935. The classification of the Edrioasteroidea. Smiths. Mise . Coll. 93 : 8. Washington. Pp . 1-1 1 . Pl. l . - --, 1936. New species of American Edrio ­asteroidea. Ibid. 95 : 6. Pp. 1-33. Pls. 1-7. - --, 1 938. Pelmatozoa palaeozoica (generum et genotyporum index et bibliographia) . Foss. Cat. I: 83. ' s -Gravenhage. Pp. 1-194. - --, 1 943. New Ordavieian cystidean echinoderms from Oklahoma. Amer. J. Sci . 241 . New Haven. Pp. 694-703. Pl. l . - B assler, R. S. & Moodey, M. W., 1 943. Bibliographic and fauna! index of Paleozoic pelmatozoan echinoderms. Geol. Soc. Amer. Spec. Pap. 45. Baltimore. Pp. VI + 1-734. - Bather, F. A . , 1906. Ordavieian Cystidea from Burma. Pp. 6-40. Pls. 1-2. In REED , F. R. C . , The Lower Palaeozoic fossils of the Northern Shan States. Burma. Mem. Geol. Surv. India. Palae-

48 A R IU V I<'Öl� K E lii i , M I N E R A L O G I O . G E O L O G I . BD 2f:i Å . N : O 1 3 . ontol. Indica. N . S . 2 : 3 . Calcutta. - --, 1 9 10 . Ordavieian Cystidea from the Carnic Alps. Riv. Ital. Paleontol. Anno XVI. 1-2. Gatania (Perugia) . Pp . 38-54 ( 3-19 of sep . print ) . PI . 2 . - --, 1913 . Garadocian Cystidea from Girvan. Trans. Roy. Soc. Edinb. 49. 2: 6 . Edinburgh. Pp. 359-529 . Pls. 1-6. - --, 1 9 1 5 . Studies in Edrioasteroidea. 6 . Pyrgocystis. Geol. Mag. N. S. ( 6 ) . 2. London. Pp. 5-12, 49-60. Pls. 2-3. - Begg, J. L . , 1 934. On the genus Cyclocystoides. Ibid. 7 1 . Pp. 220-224. PI. 1 1 . - · -- , 1 939. On the genus Cyclocystoides , with the description of a new species from the Ashgillian of Girvan. Trans. Geol. Soc. Glasgow. 20 : l. Glasgow. Pp. 2 1-29. PI. l. - Billings, E . , 1 858 . On the Cystidae from the Lower Silurian rocks of Canada. Figures and descriptions of Ganadian arganie remains. Dec . 3 . ( Geol. Surv. Canada. ) Montreal. Pp . 9-74. Pls. l-7. - Bouska, J . , 1 942. Krinoidengattung Pycnosaccus ANGELIN im böhmischen Silur . Mitt. Tschech . Akad. Wiss. 1 942. J g. 52 : 2 1 . Prag. Pp . 1-4. PI. l . (Summary of the Czech text in Rozpravy Il. Tr. Cesk. Akad. Roi nik 52: 2 1 . Praha. Pp. 1-5. PI. l . ) - --, 1 943. Die Vertreter der Gattung Gissocrinus ANGELIN im böhmischen Silur. Ibid. 1 943 . Jg. 53: 44. Pp. l-10. Pls. l-2 . ( Summary of the Czech text in Rozpravy II. Ti·. Cesk. Akad. Roinik 53: 44. Pp. l-12. Pls. l-2. ) ­·--, 1 946. On Crotalocrinitidae (ANGELIN) from the Silurian and Devonian of Bohemia. Ibid. 1 946. Anmie 47 : 4. Pp. 1-17 . Pls. l-4. (Summary of the Czech text in Rozpravy II. Tf. Cesk. Akad. Ro: nik 56: 4 . Pp. l-24. Pls. 1-4. ) - Brögger, W. C., 1 882. Die silurisohen Etagen 2 und 3 im Kristiania­gebiet und auf Eker, ihre Gliederung, Fossilien, Schichtenstörungen und Con­tactmetamorphosen. Kristiania (A. W. BRÖGGER) . Pp. VIII + l-376 . Pls. 1-12. l map. - --, 1 887 . Geologisk kart over oerne ved Kristiania. Nyt Mag. Naturvidensk. 31 : 2. Kristiania. Pp . l-36. l map . - --, 1 896. Uber die Verbreitung der Euloma-Niobe-Fauna ( der Ceratopygenkalkfauna) in Europa. Ibid. 35. Pp. 1 64-240. - Chauvel, J., 1 9 4 1 . Recherches sur les cystaides et les carpoides armoricains. Theses Fac . Sci. Univ. Rennes. (C ) . 3. Rennes. Pp . l-286. Pls. 1-7. - Cooper, B . N., 1 945. Stones River equival­ents in the Appalachian region. J. Geol. 53. Chicago, Il!. Pp. 262-275. -Czarnocki, J. , 1 928 . Le profile de l'ordovicien inferieur et superieur a Zalesie pres Lag6w campare a celui des autres regions de la partie centrale du massif de Ste Croix. Bull. Serv. Geol. Pol. 4 : 3-4. Warszawa. Pp . 555-5 8 1 . (Polish with French translation; pp. 569-5 8 1 . ) - Deecke, W., 1 9 1 5 . Paläontologische Betrachtungen. 8. Uber Crinoideen. N. Jb. Mineral. etc. Jg. 1 9 1 5 : 2. Stutt ­gart. Pp . l-18. - Dehrn, R., 1 933 . Cystoideen aus dem rheinischen Unter ­devon. Ibid. Beil. Bd. 69. Abt. B. Stuttgart. Pp . 63-93. PI . 2 . - --, 1 934. Untersuchungen an Cystoideen des rheinischen Unterdevons. S. B. Akad. Wiss. Munchen. Math. -naturw. Abt. 1 934. Pp. 1 9-43. Pis. l-2. - Delage, Y. & Herouard, E., 1 903. Traite de zoologie concrete . 3 . Les echinodermes . Paris (ScHLEICHER Freres) . Pp. x + l-495. Pis. 1-53. - Delgado, J. F. N., 1 908. Systeme silurique du Portugal. Mem. Com. Serv. Geol. Port. Lisbonne. Pp. l-245. Pls. 1-8. - Delpey, GenevH�ve, 1 944. Histoire des echinodermes jusqu'au devonien inferieur. Bull. Soc. Geol. France. ( 5 ) . 14. Paris. Pp . 247-278 . - Dreyfuss, M . , 1939 . Les cystaides de l'ordovicien superieur du Languedoc . Ibid. ( 5 ) . 9. Pp. l l 7-l 34. Pls. 10-1 2 . - Eichwald, E . , 1 845 ( 1 846? ) . Die Urwelt Russlands, durch Abbildungen erlaeutert. 3 . Einige ver­gleichende Bernerkungen zur Geognosie Seandinaviens und der westlichen Pro­vinzen Russlands. St. Petersburg & Moscau. Pp . 1-156. Pls. l-2. - --, 1 856. Beitrag zur geographischen Verbreitung der fossilen Thiere Russlands. Alte Periode. Bull. Soc. Imp. Nat. Moscou. 29 : l. Moscou. Pp. 88-1 2 7 . ­--, 1 860. Lethaea Rossica ou paleontologie de la Russie . l . Premiere sect. de l'ancienne periode. Stuttgart. Pp. XIX + l-68 1 . Atlas with Pls. l-59 + pp. l-8. - Ekström, G., 1937 . Upper Didymograptus shale in Scania. Sver . Geol. Unders. Ser. C. 403. Stockholm. Pp. 1-53. Pls. 1-l l . - Faura y Sans, M., 1 9 1 3 . Sintesis estratigråfica de los terrenos primarias de los yaci ­mientos fossiliferos mas principales. Mem. R. Soc. Espaii . Hist. N at . 9. Madrid.

G E RH A RD R E G N E L L , N O N · C RIN OID P E L M A T O Z O A N J<' A U N A S . 49 Pp. 5-202. - Ferroniere, G . , 1 922 . L e calcaire d e l a Grange, pres Chalonnes (Maine et Loire ) . Bull. Soc. Sci. Nat. Nantes. (4) . 2. Nantes. Pp . l-37 . Pls. l-2. - Field, R. M . , 1 9 1 9 . The Middle Ordavieian of Central and South Central Pennsylvania. Amer. J. Sci. ( 4 ) . 48. New Haven. Pp. 403-428. ­Foerste, A. F., 1 9 14 . Notes on Agelacrinidae and Lepadocystinae, with de­scriptions of Thresherodiscus and Brockocystis. Bull. Sci. Lab. Denison Univ. 17. Granville. Pp. 399-487. Pls. l-6. - --, 1 9 1 6. Oomarocystites and Oaryocrinites. Cystids with pinnuliferous free arms. Ottawa Naturalist. 30. Ottawa. Pp. 69-79, 85-93, 10 1-1 1 3 . Pis. 2-5. - Forbes, E., 1 848. On the Cystidae of the Silurian rocks of the British Islands. Mem. Geol. Surv. Great Brit . a . Mus. Pract. Geol. 2 : 2. London. Pp. 483-534. Pis. l l-23. ­Frebold, H., 1 928 . Grundziige im Characterbilde der epirogenen Bewegungen Skandinaviens und des Baltikums im Kambro-Silur. Geol. Rundschau. 19. Berlin. Pp. 8 1-105. - Frech, F., 1 899 . Die geographische Verbreitung und Entwickelung des Cambrium. Congr. Geol. Int. 7me Sess. Mem. pres Congr. 6. St. Petersbourg. Pp . 1 2 7-1 5 1 . l table. - Freyberg, B. von, 1 923 . Die Fauna und Gliederung des Thiiringer Untersilurs. Z. Deutsch. Geol. Gesellsch . 74. 1 922 . Berlin. Pp. 237-276 . Pis. 4-5. - Funkquist, H. P. A. , 1 9 1 9 . Asaphusregionens omfattning i sydöstra Skåne och p å Bornholm. K. Fysiogr. Sällsk. Hand!. N. F. 31 . Lund. Pp. 1-56. Pis. 1-2. - Gisll!n, T., 1 94 7 . On the Haplozoa and the interpretation o f Peridionites. Zoo!. Bidr. Uppsala . 2 5 . ( = Festskr. tillägn. N. VON HoFSTEN. } Uppsala & Stockholm. Pp. 402-408 . - Gortani, M . , 1 934. Fossili ordoviciani del Caracorum. Sped. Ital. Filippi Himalaia etc. ( 1 9 1 3-14) . ( 2 ) . 5. Bologna. Pp. 1-93. Pis. l-19 . - Grabau, A. W., 1 9 1 6 . Comparison of American and European Lower Ordovician forma­tions. Bull. Geol. Soc. Amer. 27. New York. Pp . 555-622. - --, 1923 . Stratigraphy of China. l . Palaeozoic and Older. Geol. Surv. China. Peking 1 923-1924. Pp. XVIII + 1-528. Pis. 1-6. - --, 1 9 3 1 . Palaeozoic centres of faunal evolution and dispersaL Bull. Geol. Soc. China. 1 1 . Peking 193 1-32. Pp. 227-239. - --, 1 936-1938 . Palaeozoic formations in the light of the pulsation theory. Peiping (Univ. Press, Nat. Univ. Peking) . 1. Lower and Middle Cambrian pulsations. 1 9 3 6 a ( see. ed. ) . Pp . XXIV + 1-680. Pis. I, n, II a-c. 2. Cambrovician pulsation. l. Caledonian and St. Lawrence geo ­synclines. 1 936 b. Pp. xxn + l-75 1 . PI. III a. 3. Cambrovician pulsation. 2. Appalachian, Palaeocordilleran, pre-Andean, Himalayan and Cathaysian geosynclines. 1 9 3 7 . Pp. xxx + 1-850. Pis. III, III b-c. Correl. charts III: 1-6. 4. Ordovician pulsation. l. Ordovician formations of the Caledonian geosyncline, with a review and summary of the Skiddavian pulsation system. 1 938. Pp. XXXIII + 1-942. - --, 1 940. The rhythm of the ages. Earth history in the light of the pulsation and polar control theories. Peking (H. VETCH ) . Pp. XVII + 1-56 1 . Pis. l-25. - Grorud, H. H., 1 940. Et profil gjennem Ogy­giaskiefer og Ampyxkalk på Tortberg, Frogner ved Oslo . Norsk Geol. Tidsskr. 20. Oslo . Pp. 1 57-160. - Hadding, A., 1933 . The pre- Quaternary sediment­ar-y rocks of Sweden. 5. On the organic remains of the limestones. K. Fysiogr. Sällsk. Hand!. N. F. Avd. 2. 29 : 4. Lund. Pp. 1-93. - Hall, J., 1 852. Palaeontology of New-York. 2 . Natural History of New-York. Albany. Pp . VIn + l-362. Pis. l-85 + a number of lettered pls. ( = 104 pis. ) . - --, 1 86 7 . Account of some new or little known species of fossils from rocks of the age of the Niagaran group. 20th Ann. Rep. New York State Cab. Nat. Rist. Albany. Pp . 305-40 1 . ( Originally printed in advance, for the 1 8th Rep . , 1 864 . ) - Harrington, H. J. , 1 9 3 7 . On some Ordavieian fossils from Northern Argentina. Geol. Mag. 74. London. Pp. 97-124. Pis. 5-7. - --, 1938 . Sobre las faunas del Ordoviciano inferior del norte Argentino. Rev. Mus. La Plata. (N. S . ) Sec. Paleontol. 1 . Buenos Aires. Pp. 1 09-289. Pis. l-14. -Hecker, R. F., 1923 . Echinosferiti russkago silura [Echinosphaeritidae of the Silurian of Russia] . Trav. (Trudi) Mus. Geol. Mineral. Pierre le Grand Acad. Sci. Russie. 4 : l . 1 9 1 9-23. Petrograd. Pp. 1-68. Pis. 1-2. (Rus-

Arkiv för kemi, mineralogi o. geologi. Bd 26 A. N :o 13. 4

50 A R K I V l!' Ö R K E M I , �fi N E R A L O G I O . G E O L O G I . B D 26 Å . N : O 13. sian. ) - --, 1938 . New data on Rhipidocystis JKL (order Digitata n. o . , class Carpoidea JKL ) and o n a new genus Bockia ( subclass Eocrinoidea JKL, class Crinoidea MILL . ) from the Ordavieian of Leningrad province, USSR, and Estonia. C . R. (Dokladi) Acad. Sci. URSS. N. S . 1 938 . 19 : 5 . Moscou. Pp. 42 1-424. - --, 1 939 . Pyrgocystis from the Ordavieian of Leningrad province. Bull. (Izvestia) Acad. Sci. URSS. Ser. Bio!. 1 939. Moscou. Pp. 241-246. PI. l. ( Russian with English summary, pp. 245-246 . ) - --, 1 940. Carpoidea, Eocrinoidea and Ophiocistia des Ordoviziums des Leningrader Gebietes und Estlands. Acad. Sci. URSS . Trav. (Trudi) Inst . Paleontol. 9 : 4 . Moscou & Leningrad. Pp. 5-82. Pis . 1-10. (Russian with German summary, pp. 56-7 6 . ) - Hede, J. E . , 1 942. On the earrelation of the Silurian of Got ­land. Lunds Geol. Fältkl. 1 892-1942. Lund. ( = Medd. Lunds Geol. -Mineral . Inst. 1 0 1 . ) Pp. 205-229. - Hill, D., 1 943. A re-interpretation of the Austral­ian Palaeozoic record, based on a study of the rugose corals. Proc. Roy. Soc . Queensl. 54 : 6. Brisbane. Pp. 53-66. l table. - Holm, G. & Westergård, A. H., 1 930. A Middle Gambrian fauna from Bennett Island. Mem. Acad. Sci. URSS . ( 8 ) . C!. Phys. -Math. 21 : 8 . Leningrad. Pp. 1-25. Pis. 1-4. -Holtedahl, O . , 1 909 . Studien iiber die Etage 4 des norwegischen Silursystems beim Mjösen. Videnskabs-Selsk. Skr. l. Math. -Naturv. Kl. 1 909: 7. Christia­nia. Pp. 1-76. - --, 1 9 1 8 . Notes on the Ordavieian fossils from Bear Island collected during the Swedish expeditions of 1 8 9 8 and 1 899 . Norsk Geol. Tids­skr. 5. Kristiania. Pp. 79-94. Pis. 9-1 1 . - --, 1 920. Paleogeography and diastrophism in the Atlantic-Region during Paleozoic time. Amer. J. Sci. (4 ) . 49. New Haven. Pp. 1-25. - --, 1 924. On the rock formations of Novaya Zemlya, with notes on the Paleozoic stratigraphy of other Arctic lands. Videnskaps-Selsk. Kristiania. Rep. Sci. Res. Norw. Exp . Novaya Zemlya 1 92 1 . 22. Kristiania. Pp. 1-1 83. Pis. 1-44. - Huffman, G. G . , 1 945. Middle Ordavieian Iimestones from Lee County Virginia to Central Kentucky. J. Geol. 53. Chicago, III. Pp. 1 45-1 74. - Jaanusson, V., 1 944. Ubersicht der Stratigraphie der Lyckholm-Komplexstufe. C . R . Soc. Geol. Finlande. 16. Helsinki. Pp. 92-100. - --, 1947 . Zur Fauna und zur Korrelation der Kalksteine mit Illaenus crassicauda ( sog. Flagkalk) im Silj an­Gebiet Dalarnas. Geol. Fören. Förh. 69. Stockholm. Pp. 4 1-50. - Jaanu s ­son, V. & Martna, J. , 1 948. A section from the Upper Chasmops series to the Lower Tretaspis series at Fj äcka Rivulet in the Silj an area, Dalarne. Bull. Geol. Inst. Ups. 32. Uppsala. Pp. 1 83-1 93. Pis. 1 7-18 . - Jaekel, O., 1 897 . [Review article of] E . HAECKEL, Die Amphorideen und Cystoideen. Beiträge zur Morphologie und Phylogenie der Echinodermen. (Festschr. f. C. G EGENBAUR, 1896 . ) N. Jb. Mineral. etc. 1 897 : l. Stuttgart. Pp. 386-395. - --, 1 899. stammesgeschichte der Pelmatozoen. l . Thecoidea und Cystoidea. Berlin (SPRINGER) . Pp . x + 1-422 . Pis. 1-1 8. - --, 1 9 1 8 . Phylogenie und System der Pelmatozoen. Palaeontol. Z. 3. Berlin. Pp. 1-128. -- -, 1 926 . Uber zwei Cystoideen und ihre morphologische Bewertung. Norsk Geol. Tidsskr. 9. Oslo . Pp. 1 9-22 . PI. l . - --, 1 927 a. Uber Tormoblastus n. g . , eine coro ­nate Blastoidee, aus dem Ordovicium Schwedens. Ark. Zoo!. 19 A: 4. Stock­holm & Uppsala. Pp. 1-6 . PI. l. - --, 1 92 7 b . Gyathotheca suecica n. g. n. sp. , eine Thecoidee des schwedischen Ordoviciums. Ibid. 19 A: 5 . Stock­holm & Uppsala. Pp. 1-5. PI. l . - Kautsky, F., 1 945. Die unterkambrische Fauna von Aistjakk in Lappland. Geol. Fören. Förh. 67. Stockholm. Pp. 129-2 1 1 . Pis. 9-1 8 . - Kettner, R. & Boucek, B., 1 936 . Tableaux synop ­tiques des formations du Barrandien. Trav. Inst. Geol. Paleontol. Univ. Charles a Praha. 1 936 : 1 8 . Praha. - Kirer, J. , 1 897 . Faunistische Uebersicht der Etage 5 des norwegischen Silursystems. Videnskabs- Selsk. Skr. l . Math . ­Naturv. K l . 1 897 : 3 . Kristiania. Pp. 1-7 6 . - --, 1 9 0 1 . Etage 5 i Asker ved Kristiania. Studier over den norske Mellemsilur. Norges Geol. Unders. Aarb. 1 902: l. Christiania. Pp. 1-l l l . - Kobayashi, T . , 1 934. The Cambro­Ordovician formations and faunas of South Chosen. Palaeontology. 2. Lower Ordavieian faunas. J. Fac . Sci. Imp. Univ. Tokyo . ( 2 ) . 3 : 9. Tokyo. Pp .

G E R H A R D R EGNE L L , N O N - U R I N O l D P E L M A 1' 0 Z O A N P A U N A S . i) }

52 1-585. Pls. 1-8. - --, 1935 . The Cambro-Ordovician formations and faunas of South Chosen. 3. Gambrian faunas of South Chosen, with a special study of the Gambrian trilobits genera and families. Ibid. 4 : 2. P p. 49-3 4 4 . Pls. 1-24. - --, 1 936. The natural boundary between the Gambrian and Ordovician systems from the Asiatic standpoint. Intern. Geol. Congr. Rep. 1 6 . Sess. U. S . A. 1 933. 1. Washington. Pp. 4 85-493. - --, 1 9 3 7 . The Cambro -Ordovician shelly faunas of South America. J . Fac . Sci. Imp. Univ. Tokyo. (2 ) . 4 : 4 . Tokyo . Pp. 369-522. Pls. 1-8. - Koenen, A. von, 1 886 . Ueber neue Cystideen aus den Caradoc-Schichten der Gegend von Montpellier. N. Jb. Mineral. etc. 1 886: 2. Stuttgart. Pp. 246-254. Pls. 8-9. - Leuch ­tenberg, Maximilian, Herzog von, 1 843. Beschreibung einiger neuen Thisr­reste der Urwelt aus den si1urischen Kalkschichten von Zarskoj e-Selo. St. Petersburg. Pp. 1-26. Pis. 1-2. - Lindström, G., 1 888. List of the fossil faunas of Sweden. l. Gambrian and Lower Silurian. Stockholm. Pp. 1-24. ­Loretz, H., 1 884. Ueber Echinosphärites und einige andere organische Reste aus dem Untersilur Thiiringens. Jb. K. Preuss. Geol. Landesanst. 1 883 . Ber­lin. Pp. 1 36-158 . - Loven, S., 1 867 . Om Leskia mirabilis GRAY. Öfvers . K. Vet . -Akad. Förh. 1 867 : 5. Stockholm. Pp . 43 1-440. ( Summarized in English in Geol. Mag. 5. London 1 868 . Pp. 1 7 9-1 84. ) - Luha, A., 1 940. Paar uut kivistist Saaremaa lademeist. l . Tsiistiididae esindaj a Eesti gotlandiumist [Some new fossils from the series of strata of Saaremaa (Oesel) . l. A representa­tive of Cystoidea from the Silurian of Estonia] . Eesti Loodus. 2-3. 1939 . Tartu. Pp. 9 8-99. (Estonian. ) - Marr, J . E., 1 882 . On the Gambrian {SEDGW. ) and Silurian rocks of Scandinavia. Quart. J. Geol. Soc. 38. London. Pp. 3 1 3-32 7 . - Miller, S. A., 1 889. N orth American geology and paleontology for the use of amateurs, students, and scientists. Cincinnati, Ohio. Pp. 1-664. - Moberg, J. C . , 1 904. Om rödfärgade lager inom Sveriges kambro-silur. Geol . Fören. Förh. 26. Stockholm. Pp. 1 34-144. - --, 1 907 . Aeglina umbonata n. sp. Ibid. 29. Pp. 257-264. Pl. 3. - --, 1 9 1 1 . Historical­stratigraphical review of the Silurian of Sweden. Sver. Geol. Unders. Ser. C . 229. Stockholm. Pp. 1-2 10. l p i . - Moberg, J . C. & S egerberg, C. O . , 1 906. Bidrag till kännedomen o m ceratopygeregionen med särskild hänsyn till dess utveckling i Fogelsångstrakten. K. Fysiogr. Sällsk. Hand!. N. F. 17 Lund. Pp. 1-1 1 6 . Pis. 1-7. - Mortensen, Th. , 1937. Contributions to the study of the development and larva! forms of echinoderms. 3. D. Kg! . Danske Vidensk. -Selsk. Skr. Nat. Math. Afd. R. 9. 7 : l. K0benhavn. Pp. 1-65. Pl. 1-15. - Munthe, H., 1 9 2 1 . Beskrivning till kartbladet Burgsvik j ämte Ho­burgen och Ytterholmen. Sver. Geol. Unders. Ser. Aa. 152. Stockholm. Pp. 1-172 . Pl. l . - Öpik, A., 1 930. Brachiopoda Protremata der estländischen ordovizischen Kukruse-Stufe. Act. Comm. Univ. Tartu. A 17 : l . (Pub!. Geol. Inst. Univ. Tartu. 20. ) Tartu. Pp. 1-262. Pis. 1-22. - Orviku, K. Jaan ­soon - , 1927 . Die Rautenvariationen bei Echinosphaerites aurantium GYLL . und ihre stratigraphische V erbreitung im estnischen Ordovicium. Pub! . Geol. lust. Univ. Tartu. 8. Tartu. Pp. 1 - 1 6 . - Pander, C. H., 1 830. Beiträge zur Geognosie des russischen Reiches. St. Petersburg. Pp. xx + 1-165. Pls. 1-4, 4 B-C, 5-1 5, 1 6 A-B, 1 7-3 1 . l map. - Perner, J., 1 900. Nomenklatura a systematicke postaveni >>Barrandovych cystidei>> [Nomenclature and system­atic position of >>BARRANDE' s cystoids >>) . Vestn. Ceske Akad. Cis. Franti �ka Jo­sefa. 9. Praha. Pp. 1 42-148. (Czech. ) - Phleger, F. B., jun. , 1 935. Some Ordovician cystoids from Russia. Bull. Mus. Comp . Zoöl. Harvard College. 76 : 5. Cambridge, Mass. Pp . 1 9 1-20 1 . l pl. - Pilotti, C . , 1 924. Sul siluriano inferiore di regione S. Marco ( Iglesias) . Boll. Soc. Geol. Ital. 43. Roma. P. 32 . - Pompeckj , J. F., 1 896 a. Die Fauna des Cambrium von Tejrovic und Skrej in Böhmen. Jb. K. Geol. Reichsanst. 45. 1 895. Wien. Pp . 495-6 1 3 . Pis . 1 3-17 . - --, 1 896 b. Ein neuentdecktes Vorkommen von Tremadoc ­Fossilien bei Hof. Ber. Nordoberfränk. Ver. Natur- , Gesch. Landeskunde . 1 896 . Hof. Pp. 1-17 ( in sep. print) . PI. 2. - Poulsen, Chr. , 1927 . The Cambrian, Ozarkian and Ganadian faunas of Northwest Greenland. Medd.

52 A R KIV F Ö R K E lll l , liii N E R A L O G I O . G E O L O GI . B D 26 A . N : O 13. Greml. 70 : 2. Copenhagen. Pp. 233-348. Pis. 1 4-2 1 . - --, 1 932. The Lower Cambrian faunas of East Greenland. Ibid. 87. Pp. l-66. Pis. l-14. - --, 1 936 . Ubersicht uber das Ordovizium von Bornholm. Medd. Dansk Geol. Foren. 9. Kobenhavn. Pp . 43-66. - --, 1937 . On the Lower Ordo ­vician faunas of East Greenland. Medd. Gronl. 1 19 : 3. Copenhagen. Pp. l-72. Pls. l-8. - --, 1 946. Notes on Cambro-Ordovician fossils collected by

the Oxford University Ellesmere Land Expedition 1 934-5. Quart. J. Geol. Soc. 102. London. Pp. 299-337 . Pis. 1 9-24. - Raymond, P. E., 1 9 1 3 . Notes o n Gyclocystoides. Bull. Victoria Mem. Mus . l . Canada Geol. Surv. Pp. 23-32. Pl. 3: l-4. - Reed, F. R. C., 1 906. The Lower Palaeozoic fossils of the Northern Shan States, Burma. Mem. Geol. Surv . India. Palaeontol. Indica. N. S. 2 : 3 . Calcutta. Pp . 1-154. Pls. 1-8. - --, 1 9 1 7 . Ordavieian and Silurian fossils from Yunnan. Ibid. 4 : 3. Pp. III + 1-69. Pis. l-8. -Regnen, G., 1 940. Om faunan i planilimbatakalkstenen vid Köping på Öland. K. Fysiogr. Sällsk. Lund Förh. 10 : l . Lund. Pp. 1-15. Pl. l . - --, 1 9 4 1 . On the Siluro-Devonian fauna o f Chöl-tagh, Eastern T'ien-shan. I . Anthozoa. Rep . Sci. Exp. N. -W. Prov. China Leadership SvEN HEDIN . ( 5 ) . 3. (Also in Palaeontol. Sinica. ) Stockholm & Nanking. Pp. l-64. Pls. 1-12 . - --, 1 942. Stratigraphical and paleontological remarks on the Lower Ordavieian of Central and Northern Öland, with special reference to Köpings klint. Lunds Geol. Fältkl. 1 892-1942. Lund. ( = Medd. Lunds Geol . -Mineral. Inst . 99 . ) Pp. 1 68-1 8 4 ( 1-17 o f sep . print ) . - --, 1 945. Non-crinoid Pelmatozoa from the Paleozoic of Sweden. A taxonornie study. Medd . Lunds Geol . ­Mineral. Inst . 108 . Lund. Pp. vm + l-255. Pls. l-15. - --, 1 948 a. Echinoderms (Hydrophoridea, Ophiocistia) from the Ordavieian ( Upper Skiddavian, 3c� ) of the O slo region. Norsk Geol. Tidsskr. 27. O slo. Pp. 1 4-57. Pls. 1-2. - --, 1 948 b. Swedish Hybocrinida (Crinoidea Inadunata Disparata; Ordovician-Lower Silurian) . Ark. Zool. 40 A: 9. Stockholm. Pp. l-2 7 . Pls. 1-4. - Richter, R., 1 930. Schuppenröhren als Anzeiger von zwei im deutschen Devon neuen Echinodermen-Gruppen (Edrioasteroidea BILLINGS und Ophiocistia SoLLAS ? ) . Senckenbergiana. 12. Frankfurt a. M. Pp. 279-304. - Richter, R. & E., 1 940. Studien im Paläozoikum der Mit ­telmeerländer. 5. Die Saukianda-Stufe von Andalusien, eine fremde Fauna im europäischen Ober-Kambrium. Abh. Senckenb. Naturforsch. Gesellsch. 450 . Frankfurt a. M. Pp. l-88. Pls. 1-5. - --, 1 9 4 1 a. Studien im Paläo ­zoikum der Mittelmeer-Länder. 6. Die Fauna des Unter-Kambriums von Cala in Andalusien. Ibid. 455. Pp. 1-90. Pls. 1-4. - --, 1 9 4 1 b. Studien im Paläozoikum der Mittelmeer-Länder. 7. Das Kambrium am Toten Meer und die älteste Tethys. Ibid. 460. Pp. 1-50. Pls. 1-2. - Roy, S. K., 1 9 4 1 . The Upper Ordavieian fauna o f Frabisher Bay, Baffin Land. Field Mus. Nat. Rist . Geol. Mem. 2. Chicago. Pp. 1-2 1 2 . - Ruedemann, R., 1 925. Some Silurian ( Ontarian) faunas of New York. New York State Mus . Bull. 265. Albany. Pp. 5-1 34. Pls. l-24. - Ruzicka, R., 1 939 . Bulbocystis, a new cystoid from the Devonian of Bohemia. Pfiroda. 32 : 9 . Brno . Pp. 292-293. (Czech, with English summary, p . 293. ) - Salter, J. W., 1 86 6 . On the fossils of North Wales. ( Appendix to A. C. RAMSAY, The geology of North Wales. ) Mem. Geol. Surv. Great Brit . a. Mus. Pract . Geol. 3. London. Pp. 239-3 8 1 . Pls. l-26. - Salter, J . W . & Billings, E . , 1 858. O n Gyclocystoides, a new genus of Echinodermata from the Lower and Middle Silurian rocks. Figures and descriptions of Canadian organic remains. Dec. 3. ( Geol. Surv. Canada. ) Montreal. Pp. 86-90. P l . l O bis. - Schmidt, F., 1 858. Untersuchungen \iber die silurische Formation von Ehstland, Nord-Livland und Oesel. Arch. Naturkunde Liv - , Ehst- u. Kurlands. ( l ) . 2. Dorpat. Pp. l-248 ( l-250 of sep. print) . l map. - --, 1 874. Uber einige neue und wenig bekannte baltisch-silurische Petrefacten. (Miscellanea Silurica 2 . ) Mem. Acad. Sci. St. ­Petersb . ( 7 ) . 21 : I l . St. Petersbourg. Pp. 1-48. Pls. l-4. - --, 1 882 . On the Silurian (and Cambrian) strata o f the Baltic Provinces o f Russia, as campared with those of Seandinavia and the British Isles. Quart. J. Geol.

GERHARD R E G N E L L , N O N- C R I N O I D P E L J\1 A T O Z O A N F A U N A S . Ö3 Soc. 38. London. Pp . 5 1 4-536. PI. 2 3 . - Schuchert, Ch., 1 904. O n Siluric and Devonic Cystidea and Camarocrinus. Smiths. Mise . Coli. 47 : 2. Washing­ton. Pp. 20 1-272. Pis. 34-44. - --, 1 9 1 9 . A Lower Gambrian edrioasterid, Stromatocystites wctlcotti. Ibid. 70 : l. Pp. l-7. PI. l. -- --, 1 929. The making of paleogeographic maps. Leopoldina. 4. Leipzig. Pp. 1 1 6-125. -Scupin, H., 1 928. Ostbaltikum. l . Algonkium, Paläozoikum und Mesozoikum. Die Kriegsschauplätze 1 9 1 4-1 9 1 8 geologisch dargestellt. 9. Berlin. Pp. vu + l-270. Pis. l-3 . l map. - Simpson, G. G., 1 9 4 7 . Holarctic Mam­malian faunas and continantal relationships during the Cenozoic . Bull. Geol. Soc. Amer. 58. Baltimore. Pp. 6 1 3-688. - Spencer, W. K., 1 922. A mono­graph of the British Palaeozoic Asterozoa. 5. Palaeontogr. Soc. 1 920. London. Pp. 1 9 7-236. Pis. 1 4-1 7 . - --, 1938 . The distribution and migration of certain animal groups in the British Lower Palaeozoic fauna. 4. The Star· fishes and cystids. Rep. Brit . Assoc. Adv. Sci. London. 108. 1 9 3 8 . London . P. 4 1 5 . - Starmer, L., 1 934. Cambro-Silurian zones of the O slo region, with a brief earrelation between the British and Norwegian sections. Pp. 23-3 1 in HOLTEDAHL, 0 . , et al. , The geology of parts of southern Norway: The Oslo region and adj acent sparagmits district. Geol. Assoc. London. - --, 1945 . Remarks on the Tretaspis (Trinucleus) shales of Hadeland, with description of trilobits faunas. Norsk Geol. Tidsskr. 25. Oslo. Pp. 3 7 9-426. Pis. l-i. -Stubblefield, C. J. & Bulman, O. M. B . , 1927 . The Shineton shales of the \Vrekin district: with notes on their development in other parts of Shropshire and Herefordshire. Quart. J. Geol. Soc. 83. London. Pp. 96-145. Pis. 3-5 . ­Suj kowski, Z., 1 946. The geological structure of East Poland and West Russia: A summary of recent discoveries. Ibid. 1 02. Pp. 1 89-20 1 . Pl. 14. -Sun, Y. C . , 1936 . On the occurence of Aristocystis faunas in China. Bull . Geo!. Soc. China. 15. Nanking. Pp. 477-488. Pis. 1-2. - Sun, Y. C. & Szetu, S. S . , 1 947. Preliminary notes on the stratigraphy and structure of the Paoshan region, W. Yunnan. Contrib. Geol. Inst . Nat. Univ. Peking. 32. Peiping. Pp. l-26. Pis. l-2 . - Swartz, Ch. K., et al. , 1 942. Correlation of the Silurian formations of North America. Bull. Geol. Soc. Amer. 53. New York. Pp. 533-538. l pl. - Thoral, M., 1 9 3 5 a . Contributian a l'etude geologique des Monts de Lacaune et des terrains cambriens et ordoviciens de la Montagne Noire. Paris & Liege. Pp. 1-3 1 9 . Pis. l-5. - --, 1 9 3 5 b. Contributian a l'etude paleontologique de l 'ordovicien inferieur de la Montagne Noire et revision sommaire de la faune cambrienne de la Montagne Noire . ::\<Iontpellier. Pp. l-362. Pis. l-35. - Thors1und, P. , 1936 . Silj ansområdets brännkalkstenar och kalkindustri. Sver. Geol. Unders. Ser. C. 398. Stock­holm. Pp. l-64. Pls . 1-3. - --, 1 9 3 8 . Ordavieian and Silurian. Pp . 25-42 . PI. 4, in THORSLUND, P. & WEsTERGÅRD, A. H . , Deep boring through the Cambro-Silurian at File Haidar, Gotland. Ibid. 415 . - --, 1 940. On the Chasmops series of Jemtland and Södermanland (Tvären) . Ibid. 436 . Pp. 1-1 9 1 . Pls. 1-1 5. - Törnquist, S. L., 1 879 . Berättelse om en med understöd af allmänna medel utförd vetenskaplig resa i England, W ales och Skotland, afgifven till Kong!. Vetenskaps-Akademien. Öfvers. K. Vet. -Akad. Förh. 1 879 : 2. Stockholm. Pp. 63-79. - Troedsson, G. T., 1 9 1 8 . Om Skånes brachiopodskiffer. K. Fysiogr. Sällsk. Hand!. N. F. 30 : 3. Lund. Pp. 1-1 1 0 . Pls. l-2. - - -, 1 92 1 . Bidrag till kännedomen o m Västergötlands yngsta ordovicium j ämte ett försök till parallellisering av de ordovicisk-gotlandiska gränslagren i Sverige och N. Amerika. Ibid. Avd. 2. 1 7 : 3. Pp. 1-18. ---, 1 925. Grunddragen av skandinavisk-baltiska områdets paleografi i kambrosilurisk tid. Förh. 1 7de Skand. Naturforskaremötet ( Göteborg 1 9 2 3 ) . Göteborg. P p . 1 9 1-198. - - - , 1 928. On the Middle and Upper Ordavieian faunas of Northern Greenland. 2. Medd. Gran!. 72. Kobenhavn. Pp. 1-197 . Pis. 1-56. - --, 1931 . In RAMSAY, W., Geologiens grunder. 3 . upp!. Om­arb. av P. EsKOLA, B . AsKLUND, G. TROEDSSON, M . SAURAMO. 2. Stockholm ( HOLGER SCHILDT) . Pp. 1-48 1 . - --, 1 9 3 7 . On the Cambro-Ordovician faunas of Western Quruq tagh, Eastern T'ien-shan. Rep. Sci. Exp. N. -\V. Prov.

54 A R K I V }'Ö R K E lii i , l\I I N E R A L O G I o . G E O L O G I . B D 26 A . N : O 1 3 . China Leadership D r SvEN HEDIN. ( 5 ) . l . ( = Palaeontol. Sinica. N . S . B . 2 . ) Stockholm & Nanking. P p . 1-74. Pis. 1-10. - Tullberg, S . A . , 1 882 a . Förelöpande redogörelse för geologiska resor på Öland. Geol. Fören. Förh. 6 . Stockholm. Pp. 220-236. (Also Sver. Geol. Unders. Ser. C . 53. Pp. 1-1 9 . ) ­--, 1 882 b. Skånes graptoliter. l . Allmän öfversigt öfver de siluriska bild­ningarna i Skåne och j emförelse med öfriga kända samtida aflagringar. Sver. Geol. Unders. Ser. C. 50. Stockholm. Pp. 1-44. - --, 1 883 a . Skånes graptoliter. 2. Graptolitfaunorna i cardiolaskiffern och cyrtograptusskiff· rarne. Ibid. 55. Pp. 1-43. Pis. 1-4. - --, 1 883 b. Ueber die Schichten· folge des Silurs in Schonen, nebst einem V ergleiche mit anderen gleichalterigen Bildungen. Z. Deutsch. Geol. Gesellsch. 35. Berlin. Pp. 223-269. PI. 10 . -Väscäufanu, Th. , 1 9 3 1 . Les formations siluriennes de la rive roumaine du Dniester. Ann. Inst. Geol. Romånici. 15 . 1 930. Bucure�ti. Pp. 425-663. Pis. 1-12. (Roumanian with French summary, pp. 585-663 . ) - Verneuil, E. de, 1 845. In MURCHISON , R. I . , et al. , Geologie de la Russie d'Europe et des montagnes de l'Oural. 2. Troisieme partie. Paleontologie. Londres & Paris. Pp. xxxn + 1-5 1 1 . Pis. 1-43, E-G. - Vinassa de Regny, P., 1 9 4 1 . Cisti · dee itaiiane. Rendic . Real. Istit. Lombardo Sci. Lett . C!. Sci. Mat. Nat. ( 5 ) . 74 : 2 . Milano . Pp. 446-448. - Vo1borth, A., 1 846. Uber die russisohen Sphaeroniten, eingeleitet durch einige Betrachtungen iiber die Arme der Cystideen. Verh. Russ. -Kaiserl. Mineral. Gesellsch. St. Petersb. Jahre 1 845-46. St. Petersburg. Pp. 1 6 1-198. Pis. 9-1 0. - --, 1 870. Uber Achrado­cystites und Cystoblastus, zwei neue Crinoideen-Gattungen, eingeleitet durch kritische Betrachtungen iiber die Organe der Cystideen. Mem. Acad. Imp. Sci. St . -Petersb. ( 7 ) . 16 : 2. St . -Petersbourg. Pp . 1-14. l pi. - Wallerius, 1. , 1 894. Geologiska studier i Västergötland. Geol. Fören. Förh. 16. Stockholm. Pp. 298-306. - Whitehouse, F. W., 1 9 4 1 . Early Cambrian echinoderms similar to the larva! stages of recent forms. Mem. Queensl. Mus. 12 : l . Bris­bane. Pp. 1-28. Pis. 1-4. - Wilmarth, M. G., 1 938 . Lexicon of geologic names of the United States. U. S. Geol. Surv. Bull. 896. (2 pts. ) Washington. Pp. 1-2396. - Wilson, Alice E., 1 946. Echinodermata of the Ottawa forma· tion of the Ottawa-St. Lawrence lowiand. Canada Dept. Mines Res. Mines Geol. Branch. Geol. Surv. Bull. 4. Ottawa. - Wiman, C . , 1 906. In HEDSTRÖM, H. & WIMAN, c., Beskrifning till blad 5 . Sver. Geol. Unders. Ser. A l , a. stock· holm. Pp. 1-124. Pis. 1-9 . - Yakov1ev, N., 1 940. Le cystide Corylocrinus et la subdivision du silurien inferieur de l'Oural. C. R. (Dokladi) Acad. Sci. URSS. 28. Moscou. Pp. 764-765. - Yin, T. H. & Lu, C. H., 1 936-1 937 . On the Ordavieian and Silurian beds of Shihtien, Western Yunnan. Bull. Geol. Soc. China. 16. Nanking. Pp. 4 1-56. PI. l .

A d d e n d u m t o t h e l i s t o f L i t e r a t u r e : The writer wishes to call attention to certain papers which have some hearing on the subj ect dealt with above, but which had either escaped his notice when preparing the manuscript or had not appeared until the present paper was already in the press.

Melendez , B . M . , 1 944. Contribucion al estudio del Paleozoico Aragones. Trab. Inst. J. de Acosta. Ser. Geol. 3: l. Madrid. Pp. 1-1 49. Pis. 1-25. l table. - --, 1 946. Cistideos de Espafia. Las Ciencias. 1 1 : 2 . Madrid. Pp. 275-285. Pis. 1-3. - Melendez , B. M. & Meiendez, Isabel Hevia de , 1 947. La fauna Ashgilliense del silurico Aragones. Bol. Univ. Granada. 19 : 83. Granada. Pp. 247-259. PI. l . [These papers have hearing on the relation between the North European and the South European cystoid faunas.] -Sinclair, G . W . , 1 948. Three notes on Ordovician cystids. J. Paleontol. 22 . Menasha, Wis. Pp. 3 0 1-3 1 4. Pis. 42-44. [Important contribution to the knowiedge of cystoids of the Paleozoic of Canada.] - Sun , Y. C . , 1 948. The early occurrence of some Ordavieian and Silurian cystaids from Western Yunnan and its significance. Paleontol. Novitates. l . Nanking (print. in

G E R H A R D R E G N E L L , N O N - C R I N O I D P E L M A T O Z O A N �' A U N A S . f>i) Peiping). Pp . 1 - 9 . PI . l . [The importance of cystoids for the determination of the fatmal origin is emphasized. The cystoid faunas are proposed to haYe originated in the Indo -Pacific , from where they migrated to southern Europe and reached America in still later time. Some new species are described.] ­Ulrich , E. 0 . , 1 9 1 1 . Revision of the Paleozoic systems. Bull. Geol . Soc. Amer. 22. Washington. Pp. 28 1-680. Pis. 25-29. [On pp. 483-505 mat ­ters relating to the centres of origin and dispersal of fossil marine faunas, and connected problems are discussed.) - Weeks , L. G . , 1 948. Paleogeo­graphy of South America. Bull. Geol. Soc. Amer. 59 . Baltimore. Pp. 249 -282. Pis. 1-16.

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l l Bockia ? s p . REGNELI, 1 945 2 C1·yp tocrinites sp. 3 Cheit·ocrinus holmi R E G N E L L

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l 6 - sp. RE GNELL 1 9 4 5 7 EchinoenC?"inites c f. relicu lalus J A E K E L

l 8 - senckenber,gii MEYER 9 Froctocystis monstnwsa RE G N ELL

! l O Lovenicystis angelin i (.J A E K E L) l 1 1 Hemicosrnitcs oelandicus HEGNELJ • • i 1 2 - extraneus E J CHWALD l 1 3 Caryoc1·inites septentriona lis REG:KELL

1 4 - ? sp. REGNEJ, L 1 94 5 l 1 5 Stichocystis ,geornet1·ica ( A N G E L I N)

1 6 - a lutacea (A N G E L J N) 1 7 Ca ryocystites angelini (H A E C K E L) 1 8 - aff. angelini (H A E K E L) 1 9 - lagena/is REGNELL

20 - spp. 2 1 Heliocrinites anqttstiporu s REGN ELI.

2 2 - _gr-anafurn (WAHLENBERG) 2 3 - ova lis (A N G ELIN) 24 - cf. ovalis (A N G E LTN) 25 - guttaefonnis RE G N ELl.

2 6 - prominens (A NGELIN) 2 7 - stellaitts RE G NtLL

28 - variabilis REGNELL

29 - n . s p . REGNELT. 1 94 5 30 Echinosphaerites auranthtm (G Y LLE ::\ H A A L) .

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series

--- - · - ---, � � "' � aj ..ci � ..ci b.() "' "' ..ci "' b.() m - w w .e " w b.() b.() o$ w 'O "' w w b.() 'O m e6 o$ ""' . p. Q) P, � 00

·a w m .e 01 'O ""' ""' 00 00 o ..0 o ·a w p ..0 00 00 - s m ·a el m "' <il w s s m <il el ""' ..0 m

·� '"O m .2 ""' m "' <il w .... 'O p w o "' o$ ..el ..0 p 00 p "' p ·

� e6 .... <il "' ""' p, .., .... _., o ..el .... .... � <il ..el o$ e6 p ..el 'O .... E-i "' "' b.() p "' <:) -;; Q c Q E-i .... <:) ..0 � h o p "' E-i s i3 'fil h ._ .... .... o$ o ·a w _., .... Q "' <:) "' .!<: ;a .... <il ..el m w � � e6 � <il p. <:) 'O p el p:; <:) .... ::S P. el 'O "' <Il p:; w. Q o P:l � � _., B 8 w.

+ l + +

+ ? + ?

+ + + +

+ +

+ + +

+ (?) + + + + + + + + + +

+ ? + +

+ +

+ + +

Silurian

Liandoverian l (Val entian ) Wenlockian

l l Cyrtograptus series

l "' l "' ·o:: "'

l .� "' m .... -;:: -"' .... p. e6 o$ el 00 s s p p. l' m c

:;; h h .... � p � "' ..0 ..0 b.() o ·a ;!:! .... .... w w ""' el b.() .:s <il ;; ;; _., .9 ;::: s 00 "' o; <il o '"2 .... .... J;; ;!:! D:! � E-i "' "' b.() w q � p. 'o o p. � >-1 p l

i l

? '?

l Lodlovian

l l Ove� Colonus series Ram�åsa

senes

p. p - o p. .... .... p p. el b.() o p s .... o b.() b.() .... .... b.() "' "' "' 'O ..0 00 "' � "' s ... ::s _., "' � .9 � g

'? +

l l � l "' <':! .... "' :te 'O 'O .... ...

� o$ t, o$ � 'O 'O <':! "' <:) e6 'O � � � "' � � :.3 'O '§ l' d 'o 'o .!<: .... .:s el � el � "L .... el ""' .... bC ,.;s d <:) ,;s .... el s >:Q o "' z '"i:i w. C!) ""' "' o ,.;s m ""' ..el <Il 'el "' '""' ""' s l> 'o .... o ....

z "' 'O 'o w.

l ' + l

+ 2 + 3 '? + + + ? 4

+ 5 + + 6

+ 7 + 8 + 9

+ l O + 1 1

+ 1 2 + 1 3 + 1 4 + 1 5

+ 1 6 + 1 7

+ 1 8 + 1 9 + + 20

+ + 2 1 + + + + + 22

+ + + ? 23 + 24

o + 25 + + 26 + 2 7 + 28

+ 2 9 + + + + + + 30

�+- :n + 32

+ + 3 3 + 34 l

A R K I V FÖR K E M I , M I N E R A L O G I O . G E O L O G I . B D 26 Å. N : O 13.

3 5 3 6 3 7 3 8 3 9 4 0 4 1 4 2 4 3 4 4 4 5 4 6 4 7 48 49

l 50

1 5 1 5 2 5 3 54 55 5 6 5 7 58 5 9 60 6 1 6 2 6 3 6 4 65 66

l 67 68

Range and Geographical Distribution of the Non-Crinoid Pelmatozoa so far known

from Sweden

Echinosphaerites spp . . -? sp. REGNJ�LL 1 9 45 Glypto.qphaerites leuchtenber_qi (VOLBORTH) - suecicus (ANGELIN) Gornphocystifes _qotlandicus (A N GELIN) Sphaeronites pornurn ( GYI-LEN HAAL) - _qlobulus (AN GELIN) - spp . Hap losphaeronis oblon_qa (A NGELIN) - spp. Eucystis raripuncia ta AN GELIN - angelini REGNELL - acurninata REGNELL - quadrangularis REGNELL - sp. REGNELL 1 94 5 - ? s p . REGN ELI� 1 9 4 5 ( p . 1 8 4 ) Holocystites ? s p . REGNELL 1 9 45 »Sphaeronis ? daleca1'lica>> ANGELIN Paracystis ostrogoth icus SJÖBERG -? sp. Torrnoblastus bodae JAEKEL Dendrocystites sp. REGNELJ- 1 9 45 Placocystites sp. REGNELL 1 9 45 Strornatocystites baltic�ts JAEKEL Cyathotheca suecica J AEKEL . - ? sp. . . . . . Pyrgocystis sulca ta (AURIVIL LIUS) - procera (At;RIVILLIUS) . - vada ( AURIVJLLJUS) 1 - cylind1·ica (A URIVILLIUfl) Cyclocystoides lindsb·önti REGNELL 1 - insularis RE GNELL - sp. G e n n . & spp.

Gambrian

+

Tremado­cian

Ordavieian

A shgillian Skiddavian l l l (Arenigian)

Llandeilian Caradocian f ·---- ____ ___!_ ____ -;---__ _!___-;--------''-------1 C hasmops series l Ceratopyge

series Asaphus series

(Orthoceratite l s .) Tretaspis series

-- ----- --,------,--,------.---;---+--.,---�----o=+--.,------c--:.�----.----c--

+

+

+ + +

+

+

-� ... "' .., "' o ... .!:l "' w

+ +

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�

� !J

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j 2 � � + +

+ + +

-\-

+

+

+ +

+ +

? ') +

? ? ?

'! ?

+ + ?

'! +

+ +

+

Liandov e rian ( Valentian)

l l + f

Silurian

W en I ockian

+

Silurian undefined

+ •) '!

+ + +

l Ludlovian

+ +

+ + + +

+

+

+

+

+

+

+

-\-

-\­+ + + + + +

+ +

+ -\­+ + +

+

+

+

+

+ -\­+ + + + + + + -\-+ +

-\-+ +

+ +

+

+

+

+

+ 3 5 3 6 l 3 7 l 3 8 3 \J 40 4 l 4 2 4 3 44 45 46 4 7 4 8 4 9 50 5 1 5 2 5 3 5 4 55 56 5 7 58 l 5 () 60 6 1 6 2 6 3 6 4 65 66 6 7 68