QE905 .H3 1913 Halle, T Some remarks on the classification of fossil plants T. G. HALLE SOME REMARKS ON THE CLASSIFICATION OF FOSSIL PLANTS P^ 'RIKTED FROM GEOLOGISKA FORENINGENS 1 STOCKHOLM FuRilAXDLlNGAR NOV. 1913) -H3 GEOL. FUKEX. F»"»R1IANDL. 15(1 ;'..".. Iliift. •'.. Xov. 1 0 1 ii. '.Wu Somo remarks on the classification of fossil plants. By T. G. I^ALLE. (With Plates 0 and 10). The dithcultit'S attending the determination of fossil plants are due, to a Jarge extent, to the diiference in the principles adopted by different writers in the delimitation of species. (Jne of the most important points of dispute is whether, in doubtful cases, a broad or a narrow delimitation of species should be preferred. In a recent paper ^ I have advocated a narrow classification as being less apt to cause confusion in nomenclature and hasty conclusions in geological and palaeo- geographical matters. This course has later been criticized by Professor A. C. Seward ^ in his review of the paper men- tioned. In addition to his general reflections on the princip- les, Professor Seward has also made several interesting re- marks on the determinations of the Antarctic material. These remarks, which are closely connected with the general stand- point of Prof. Sewaud, call for an explanation from myself; and it would appear, too, that some points in my discussion of the principles would need emphasizing. It is naturally hopeless to attempt a reconciliation of the two opposite prin- <'iples, and a continued abstract discussion would serve no ' Hallk. T. G. The Mesozoic Flora of (Jraham Land. Wissens<'h. Ergebn. (). Srhwedisihen Siidpolar-Expcdition 1901—1903. Bd. 3, Lief. 14, Stockholm 1913. - Seward. A. C An extinct Antarctir Hora. The New Phytologist. Vol. 12. Nos. 4 Sc f). London 1913. 368 T. G. HALLE. [Nov. 1913. useful purpose, but a few of the points in question may deserve some further consideration. i After quoting my opinion 'that it is a lesser evil to keep j forms separated which are identical than to identify such as are distinct' Prof. Sewakd argues as an antithesis that it is a sound general principle 'to keep the mind open when there is no sufficient warrant for closing it'. It is a strange illu- stration of the difference in the conception of the problem that this very principle is the first and chief reason also for the exactl}^ opposite course advocated by the present writer. Indeed, I believe that a brief consideration of the process of classification of fossil plants will show the narrow delimita- tion of species to represent the less committing way. The individual specimens constituting the material of the palaeobotanist are grouped into species according to the degree of morphological similarity — very much in the same manner as in the classification of recent plants. There is the important difference, however, that the palaeobotanical specimens, in the overwhelming majority of cases, do not consist of individuals but only of portions of such. Therefore different organs of one individual may be referred to not only different species but different genera, this being the rule in many groups. And in cases of dimorphism, and even of great variation according to position on the plant, different specimens of one and the same kind of organ must often be described under different specific names though they really belong to the same species. These consequences, as every one knows, are unavoidable, being with necessity connected with the nature of the palaeobota- nical material. And they cannot be eliminated by means of even the very broadest classification: there will always be cases of dimorphism in which it is impossible to trace the connection of the different t^^pes, and it is only exceptionally that the material to be described is sufficiently large to give a satisfactory idea of the range of variation of the species. It follows, therefore, as a generally accepted fact, that the i 1 :;.";. II. i;.] CLAssiFirvTiox of fossil plants. 309 jalaeobotanical species, especially in critical frnjujjs, is not a taxononiical unit in llic same sense as species in the recent flora. The fact of two groups of forms being described under different names does not imply, therefore, that they are ne- cessarily distinct species but only that tliey cannot at pre- sent be ]) roved to be identical. And this, it must be re])eat('(l. holds true even in the case of a broad classification. The process of classification of fossil plant- remains, in other words, must naturally be, on the whole, a synthetic one. It consists in grouping together specimens into form-units (arti- ficial species) and piecing together such different parts and variations of a plant which have been described as artificial species, in order to get as accurate an idea as possible of the whole plant and its variations (the natural species). The writer who prefers in a dou])tful case to regard two similar forms as different species therefore only confesses his inability to establisl^ an identification at the moment. He cautiousl}* makes a pause in the synthetical process just because he believes it to be a sound principle 'to keep the mind o])en when there is no sufficient warrant for closing it'. The fact that the palaeobotanical species cannot be acce])ted as necessarily coinciding w4th the taxonomical may be used and has been used also as an argument for a broad classifica- tion. Since it is hopeless to strive at a natural delimitation of the species in all cases, it may be as well, it is argued, to use the specific name in a liberal sense, as a designation for a certain general ty])e of the organ in question. In referring a questionable form to a certain palaeobotanical species, it is said, the former has not been stated actually to belong to the same natural species but only to show a certain resem- blance of form. — This way of reasoning raises the (|uestion: 'what's in a name?' It cannot l)e helped that the palaeo- botanical species to a large extent is treated as a natural one. An observation made on one or some few specimens of a ])alaeobotanical species is generally and naturalU' regarded 37U T. G. HALLE. [Nov. 1913. as applicable to the whole species. Palaeobotany in general is, indeed, dependent on the possibility of so nsing, with all possible precaution, the palaeobotanical species, and the em- ])loyment of specific designations mainly serves just to mark utf such units. It is clear, therefore, that it must be an im- perative demand to make these units as pure and unmixed as possible, so that a statement made in regard to some spe- cimens of the species may with the greatest possible proba- bility apply to all others designated by the same name. An example may serve to illustrate the dangers w^hich, from this point of view, attend a broad classification. We may imagine J two sets of specimens of fern-leaves, which an adherent of h narrow classification would prefer to treat as two species but which, with a liberal delimitation of the species, are regarded as one. Specimens of the first of these sets are later found to have sporangia of, for instance, Osmundaceous type, and the result will be that the whole species will be stated to belong to that group. If then, after some time, specimens of; the second set are found to belong to stems showing the ana-^ tomieal features of the Marattiales, this character, by still regarding the species as a unit, will probably be applied to the species as a whole. The simple explanation may be that the name is used for two different natural species which may well belong to quite different groups. — It is clearly imposs- ible to guard satisfactorily against this risk, even with a narrow delimitation of species, and no doubt a large group of mistakes in palaeobotany is of this nature. But the use of specific names in a narrow sense will naturally tend to lessen this risk, whereas it hardly brings any other danger instead of it. In the case quoted, a mistake in this direction would only mean that the process of reconstruction of the. particular species is delayed. However one may look at the problem, it would appear that the use of specific names in a restricted sense would be less apt to cause confusion and mistakes. The most conspicuous objection, r.< I :•..'.. II. f.. CLAPSIl rCATlON (iK FOSSIL I'l^ANTS. 371 iind the (nic most often exj)r»'sse(l, is that this cniirsf imolves the creation of lari^^e niunhers of transient speeitic names, due onh' to the mistakes' of the respective authors. Havini; r^'gard to thr limited nature of the i)alaeobotanical material, it nf'»'(l hardly be feared that the mere nunilM-r of names will cause serious troubles. And there is this very important tylonhlatter. Hota- nisches Centraildatt. Vol. 2^). iMSfi. \t^ — i:i022U. G. F. /•'. /!'/.y. :\r2 T. (i. HALLE. [Nov. 11)13. iiu restricted use of new specific names. The aim must always be identification of those forms whose identity appears probal)le, and there is no doubt that with the increase of the material more and more 'species' will be found to be iden- tical. Professor Seward, indeed, has clearly demonstrated, how a large material makes it possible to recognize as mere varieties forms which in case of a smaller number of speci- mens no doubt would be described as different species — linffordia Gocpperti of the AVealden-flora being a very good instance. <)nly, it would appear that, when a less extensive material has to be described, a narrow delimitation would represent the more cautious course. The chief point is, of course, that the material should be sufficient for a description. It need not be large, since even a fairly small material may often be valuable, but it must be distinct and it must be well described and w^ell figured. If these demands are met the species may be safely used as a basis for future discus- sion and thus answers its purpose, even if it is ultimately found to be a mere synonym. After this abstract discussion the most important of Pro- fessor Seward's remarks on the determinations of the Antarctic plant-remains will be considered. These remarks, as already mentioned, for the most part are closely connected with the general standpoint of Professor Seward, and in such cases need no other discussion than the general exposition given above. The case of the only Eqnisetites-sjyecies of the Hope Bay Flora may be mentioned, however, as a good illustration. The close resemblance of some specimens of the Antarctic species E. approxiniatiis Natii. to E. rajinalialensis Schimp. is noted in my memoir, with the remark that both forms may be identical. The Indian material of the species is very poor, however, consisting only of diaphragms, short pieces of stems and broken leaf-sheaths with teeth indistinct or wanting.^ ^ The specimens in Feistmantei/s PI. 35, figs. 3, 4, are probably inverted. IM 3.',. 11. tj.] CLASSIFICATIOX n|.- FnS.SH, I'LANTS. 378 The charactors displayed l)y this material are so vague that they may well occur in many different species. The r^^sem- hlance between some of the more impei't'eet specimens of leat*- sheatiis from (-rrahani Land and tlie t'<|uall\- imperfect Indian ones, therefore, does not seem suttieient to warrant the use nf the name given to the latter: the identitication would he l»ased chietly on the lack of characters in (jne of the forms, in this case — which is worse — in the type-specimens. There is undoubtt'dJv a very ffreat risk in extendino: a name siv^'u to such imperfect specimens to a much bettei* characterized form, as thereby the name, on very insufficient grounds, is made to mean so much more than it has done originally and, therefore, perhaps something fpiite different. The danger is tliat forms from other parts of the world would probably be identitied hy other authors with the Indian E. rajmaludensis because of their resemblance not to the type-specimens but to the Antarctic ones. And the true Indian E. rajmahaJrusis may ultimately be found to l)e something different: this is, at any rate, just as ])ossible as that it is identical with the Antarctic form. The confusion arising then may be easily imagined, whereas, if the two forms are temporarily kept separate, an ultimate identification would l)ring no such con- sequence. Regarding the use of the generic name Schroptcris^ the reas- ons have been set forth in my discussion of P(icl/yj>trrfs (1. c, !». oD). To a reference of my S. f'nrcafa to the genus Dicho- jifriis I must most strongly object. As types of Didiojiti ris should be regarded the large Italian species for which Zigno founded the genus. The difference, in regard to both shape and venation of the pinnules, between Scbroptrfis f'nrcafa and Dichopttris risianica Zkjno is so obvious as to need no further explanation, and the coincidence in the forking of the rachis cannot be regarded as being of any importance. If Dirhoptrris is extended so as to embrace forms like Schro- })trris f'urcata it would much sooner come to include the genus 374 T. (J. HALLE. [Nov. 1913. Pachypti r'is. And as Fachi/pteris is the older name and has been commonly used — also by Professor Seavard in the 'York- shire Flora' — the name Dicliopteris should be rejected. The only ])lan to retain the genus Bichopicris is evidently to em- plo}' it in a very restricted sense for forms like those from the Italian Oolite, in which case my species of ScJerojyteris naturally cannot be included in it. Whether the venation and the peculiar habit of the Italian forms are sufficiently important characters to warrant this course, will not be dis- cussed here, but it is certain that if Dicliopteris is used in the sense of Prof. Seward's ^ it cannot be upheld but must be rejected in favour of Facliyptcris. Whichever course be followed, therefore. Dichopjfrris has only a secondary impor- tance for the delimitation of the genera Pachypteris, Tlmui- feldia and Sclcropteris. This is the reason why I did not consider it in the discussion of these genera, judging it wiser to leave this question to somebody who could make a thorough compara- tive study of the Italian forms. — From what has been said above regarding the relation of Pachyptteris and Dichopteris, it will already be clear why P. dalmatica, which was probably cor- rectly^ referred by v. Kerner to Pachypteris^ has not been in- cluded in Dichopfcris. The nomenclature of Zamitean fronds, on which Professor Seward also remarks, is a very difficult matter, and as yet there hardly exists any method of classification which is not beset by difficulties. In my Grraham Land paper I have dis- cussed the relations of the genera ZamiUs. Ptilopliyllum, and Otozamites: and Professor Seward remarks that I have used Zamites 'for several forms which agree much more closely with Ptilophyllnm fronds than with Zamites as usually em- ployed'. This remark is interesting as it shows that Profes-j sor Seward accepts the generic name Ptilophyllam. If both' Zamites and Ptilopliylluni are employed it is naturally neces- sary to establish some agreement as to how the genera are * Fossil Plants. Vol. 2, p. 552. 15(1 3o. II. •;.] CLASSIKrCATIOX nj' Ko-SIL I'LANTS. 375 to be delimited, in order that the nauit's may not he used difterently l)y difterent writers. I have tried to chdimit Pfi- lophylhtm by aceeptini^ Feistmantki/s definition which makes the downward trend of both pinna-edges at the base the ge- neric character. This is thus no new idea, the genus having been accepted in this sense also by other palaeobotanists af- ter the days of Feistmantel. This character, it must be ad- mitted, is not a very opportune one, since it is often only seen with difficult}-, and it does not mark off the genus very distinctly from all forms of Zamifrs, yet it seems to l^e the better one under the circumstances. As typical members of the genus Za mites may be regarded the species with large fronds of which the pinna-bases are distinctly rounded and characte- rized by a more or less marked callosity, Z. gigas being the best known representative of this group. The callosity can- not always be observed, however, and its occurrence is gener- ally not considered as a conditio sine qua non for a reference to Zcunites. To the same genus are often referred smaller forms without callosity and with the pinna-basis very slightly rounded. As well-known representatives of this group of forms may be regarded the species of Zamitcs described by Heer* from Greenland; and it was because of the resemblance of the Antarctic fronds to these Grreenland forms that I referred the former to Zamites. The difference between these two groups is in many cases obvious, and I have considered it in the paper quoted by distinguishing the two types as different sections of the ge- nus, resp. Kuzaiiiitcs and Snhzaniitis. It may be that the latter section would ])e better designated by a new generic name, but I did not judge such a course sufficiently war- ranted. If, on the other hand, the choice is between includ- ing these forms in Zdiuitts and referring them to PtilojtlniJ- turn, it would appear that the former genus is decidedly t(> be preferred. * Flora I'ossilis arrtica. Vol. 3, li>73, p. •;:', :ni.l lull . VU 11 ami 1.'.. 37(^ T. G. IIALLE. [Nov. 1913. It is true that VtUopliylhim as originally defined would em- brace also fronds of the type in question. But, as in the case of Zaniitcs and other old genera of C^^cadophytean fronds, it has become necessar}^ to substitute a more distinct definition for the original one: otherwise the genus would come into collision with other genera now commonl}^ accepted. This has been done by Feistmantel, whose definition of the genus seems to meet the most important demand to be made on a new definition of an old generic name, viz. to be based on the specimens for which the name was created.^ AVith exception of the numerous illustrations given by Feistmantel of the Indian species of FtUophylhwi, very few forms of that genus have been figured with due regard to the shape of the pinna- bases. In addition to a reproduction (PL 9, fig. 1) of one of Feistmaxtel's figures of Ftnophyllum aciitifoUum I have there- fore given, in PL 9, figs. 2 — 5, some photographs, in twice the natural size, of the pinna-bases of one of the English forms of the genus. This form, which is one of those commonly known among English palaeobotanists as Williamsonia pecten^ is identical with Cycaditcs pectinoides Phillips and should be named Ftilophyllum pectinoides (Phill.) Morr. It is seen from the figures to show the generic character of Ftilo- phyllani in Feistmaxtel's sense by having an asymmetric pinna-base with both edges decurrent on the rachis. And it must be admitted that this is a very characteristic type of pinna-base which well deserves to be distinguished from that of typical Zamites species, such as Z. gigas. In order to decide whether the Antarctic species of Zamites should be retained in that genus as a somewhat less typical group or included in PtiloplnjUnm, they should be compared with typical specimens of these two genera. The text-figures ^ Regarding the relation of Feistmantel's definition to Morris' type-speci- mens see Halle, T. Gr.: Some plant-bearing deposits in Patagonia and Tierra del Fuego and their floras. K. Sv. Vet. Akad. Handl., Bd 51, N:o 3, 1913, p. 36. l!'l l'"'. II.'"..' (I.ASSri irATIO.N OF FOSSIL I'LANTS. 377 \'2 and lo, pp. r)7 and ')!!, res])., in my ^lesu/oic Flora of Gra- liani Land, the illustrations given hy Hkkii (I. c; 1^1. 14; Pi. 1'), tigs. 1 — 10) and es])peially the photographs in I'l. 10. figs, l-fi. of thr present paper may he regarded as showing the typi- < sense. It would be difficult to find any sufficient generic character o78 T. G. HALLE. [Nov. 1913. of this wider genus Ftilophyllnni. From the typical Zamitcs it would diifer mainly by the less marked rounding of the basal corners and by the smaller size. The former distinction may be available in some cases but it would only be a dif- ference of degree, since even the very slightly rounded base of some of the Antarctic Zamites-forms belongs to the same type as that of Z. gigas and other typical species. If the speci- mens figured in PL lU, fig. 7, of this paper are compared with those in figures 1 — () of the same plate, which show typical pinna-bases of different species, from Greenland and (Iraham Land, referred by the present writer to Zamitcs^ it will be difficult to find any dift'erence sufficiently distinct to be expressed in a diagnosis. Indeed, it is not impossible that, w^ith the classification advocated by Professor Seward. the specimens in PI. 10, fig. 7, might be referred by some authors to Ptilopliijlliim. Yet this figure shows very typical specimens of Zamites gigas: only, these are reduced to the average size of the Ftiloj^hyUiim-fronds. It would appear from a consideration of this figure that the more stringent definition of Zamites is based to a great extent on the size. A difference of size may be a character of some importance; but it would not seem to be convenient to base on such a distinction a separation of two form-genera of a kind of fronds which show a rather great variation in size. The choice is thus between distinguishing Zamites and Ftilo- pliyllum in accordance with either the difference of shape of the pinna-base or the difference in the degree of rounding of the latter and in the size of the frond. As the latter course would involve both a mixing of two types with a diff'erent symmetry of the pinna-bases in one genus and a vaguer gene- ric distinction between the genera, it does not appear to pre- sent any advantages. It remains to discuss the question of the classification of sterile Coniferous shoots. I have created a new name, Elato- cladus, to be used for such sterile shoots of Conifers, whether IM ;;:>. 11. t"..' classification of fossii, im.ants. 37i> of (lorsiventral or of radial habit, which cannot be included in any of the c^encra instituted for more i)cciiliar forms. Pro- fcs.sor Skwaiid, who admits tliat the nomendaturt' for these forms needs revision, remarks on tlie ohjecti(jn against the name, alread\' discussed in my paper, that this generic term -includes forms with leav»\s (tf the Tcuitcs type as well as branches with leaves like those of Siilienolepidium, Elatidcs^ and other genera'. He proposes as an alternative plan 'to retain Taxitrs ill the wide sense in which it is used by most authors for twigs bearing linear and usually distichous leaves similar to those of Taxus, certain species of Podocarpus, Sequoia .senqx r- riirns, and other recent Conifers, and to adopt the name ]*'n/(02)Jn/Uu)H, for forms with radially disposed leaves like those of Elatides, Sphcnolepidium, and Chcirolcpis' . As to the use of Ta.iites for shoots of dorsiventral habit (with pseudo- distichous leaves) there is nothing to be said against this plan. Brongxiart instituted the genus for forms with only a general resemblance in habit to the recent spe- cies of Taxus. He employed it, however, especially for Ter- tiary forms which ])ossibly might be compared with the re- cent genus with greater probability than those from older forma- tions. U this (juestion is considered alone, it would seem con- venient to retain Taxitcs as advocated by Professor Seward; and 1 at first intended to employ the name in this sense. In regard to the other group, shoots with radial symmetr}^ the question is more difficult. To use Pagioplnillnm in a wide sense, for this group, would not seem to l)e an imjtrovement. Fa(/ioj)ht/llu))i is usually employed as designation for a fairly natural form-genus, which undoubtedly should be retained for forms with short and thick, often more "or less triangular and strongly keeled leaves, the species figured by Saporta (Plau- tes jurassiques. Vol. T), 1884, Pis. 4;")— o')), under the name of FarJu/jtl/i/lhoH being good representatives of the group, in ad- dition to the type-species, ra. Fig. 1. Ptilophyllum acutifolium MORR. Reproduction, in natural size, of the twice enlarged fig. 1 a, pi. 2. of Feistmantel: Ueber die indischen Cycadeengattungen Ptilophijllum und Dictyozaniites, 1876. Figs. 2 — 5. Ptilophyllum pectinoides (PlllLL.) MORR. Portions of fronds from the Lower Estuarine Series of the Inferior Oolite, at Whitby, Yorkshire, showing the pinna-bases, X 2. PI. 10. Figs. 1—3. Zamites antarcticus Halle. from the Jurassic of Hope Bay, Graham Land. Fig. 1, portion of the frond in text- fig. 13 c of Halle : The Mesozoic Flora of Graham Land; nat. size; tig. 2, two pinna-bases from the same specimen, showing indications of callosity, X 5; fig. 3, jnnna-bases of another specimen, x 5. Fig. 4. Zamites pusillus Halle. Pinna-bases of the specimen in PI. 7, fig. 12, and text-flg. 12 a of Halle : The Mesozoic Flora of Graham Land, X 5. Figs. 5, 6. Zamites hrevipennis Heer, from the Cretaceous of Western Greenland, X 2^ 2. Fig. 5, part of the specimen in PL 15, fig. 10; fig. 6. of the one in PI. 15, fig. 9, of Heer: Die Kreide-Flora der arctischen Zone. Flora fossilis arctica. Vol. 3. Fig. 7. Zamites gigas (Lindl. & HuTT.) AIORR. Large slab with several fronds from the Lover Estuarine series in the In- ferior Oolite of Whitby in Yorkshire, reduced to ^ '4 of the nat. size. All the specimens figured, with exception of the reproduction in PL 1. fig. 1. are in the Palaeobotanical Department of the State Mu- seum of Natural History at Stockholm. Geol. Foren:s FOrhandl. Bd 35. PI. 9. CcderqulsU Graf. A.-U., Sthlm. Gool. [-orciKS Fiirhandl. Bd 35 X5 /¥''''< X 2 • li 6 X' 4 CeOerquisis tir;ii. A.-H , Mhlm. Bolanicat Garden Libra QE905.H3 1913 Hall Thore Gustaf/Some remarks on the c gen llilllllll 3 5185 00092 2565