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VI.—The Classification of the Phacopidæ

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VI.—The Classification of the Phacopidæ 172 F. E. Cowper Reed—On the P/wcopidw. EXPLANATION OF PLATES. PLATE VII. FIG. 1 shows the original face A at the top of the stone. In front is the cleavage plane E, which, however, is not clearly visible owing to the light reflected from the somewhat irregular surfaces at the back of the crystal. On the extreme right is the cleavage plane H. FIG. 2 shows the cleavage plane F, which, on account of its favourable position relative to the camera, appears as a brilliantly illuminated surface; the irregular original faces C and D also appear in this view. PLATE VIII. FIG. 3 shows the triangular indentations on the irregular face D ; also portions of A and B. FIG. 4 shows the crystal resting on the cleavage plane E with the faces B and D exposed to view. The sharp bounding edges are formed as follows: at the bottom by E, on the right by A, and on the left by G. VI.—THE CLASSIFICATION OF THE PHACOPID.S:. By F. K. COWPER REED, M.A., F.G.S., of the Sedgwick Museum, Cambridge. f PHE family of Trilobites termed Phacopidae has been defined by I Beecher1 as follows:—" Glabella tumid, widest in front. Free cheeks continuous, united anteriorly. Suture extending from iu front of the genal angles inward to the eyes, and thence forward around the glabella. Eyes generally large, always with distinct facets, Bchizochroal. Thorax of eleven segments with grooved pleura. Pygidium usually lai'ge and of many segments; limb ribbed; margin entire or dentate." Nomenclature in use. While the general limits and characteristics of the family as thus given are universally recognised, much confusion and diversity of opinion still exist as to the generic groups which must be therein included. There is no precise agreement in the usage of many of the common generic names; and many subgenera have been from time to time established without general acceptance. Some palaeontologists (e.g. Salter) have been of the opinion that the family only contained one genus, Phacops, which might be split up into several subgenera. Barrande, however, recognised the presence of two genera, Phacops (in a more restricted sense than Salter) and Dalmania or Dalmanites ; and Hall & Clarke in 1888 2 adopted this view. Beecher in 1900 (op. cit.) mentioned six divisions or groups within the limits of the family, and put them all as of equal generic value. Other examples of differences in the classification of the family need not here be given ; but in order to show the multitude of generic or subgeneric groups which have been established, but of which only a few have been generally adopted, the following list 1 Beecher, Amer. Journ. Sci., vol. iii (1897), p. 202 ; and in Zittel's " Textbook of Palaeontology " (English translation, 1900), p. 636. 2 Hall & Clarke: Palseont. New York, vol. vii, pp. xxvii-xxxii. Downloaded from https://www.cambridge.org/core. INSEAD, on 18 Jul 2018 at 18:49:05, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756800131991 o c 2 O z D Downloaded from https://www.cambridge.org/core. INSEAD, on 18 Jul 2018 at 18:49:05, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756800131991 o o m X o c Downloaded from https://www.cambridge.org/core. INSEAD, on 18 Jul 2018 at 18:49:05, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756800131991 F. R. Cowper Reed—On the Phacopidce. 173 may be given, irrespective of questions of synonymy or pre- occupation, which will be dealt with subsequently :— Aeaste, Goldfuss, 1843. Metacanthus, Corda, 1847. Asteropyge, Corda, 1847. Monorakos, Schmidt, 1886. Chasmops, McCoy, 1849. Odontoeephalus, Conrad, 1840. Coronura, Hall & Clarke, 1888. Odontochile, Corda, 1847. Corycephalus, Hall & Clarke, 1888. Phaeops, Emmrich, 1839. Cryphaus, Green, 1837. Plenraeanthns, Milne-Edwards, 1840. Cryphina, Oehlert, 1889. Portlockia, McCoy, 1846. Ualtnania, Emmrich, 1844. Probolium, Oehlert, 1889. Dalmanites, Barrande, 1852. Pterygmnetopus, Schmidt, 1881. Hausmannia, Hall & Clarke, 1888. Somatrikelon, McMurtrie, 1819. Homalops, Remele, 1884. Synphoria, Clarke, 1894. Malladaia, Oehlert, 1896. Tnmerocephalus, McCoy, 1849. Of the above names, Pleuracanthus and Cryphaus are both pre- occupied, the former both by a genus of Coleoptera (Gray, 1832} and by a genus of fishes (Agassiz, 1837), the latter by a genus of Coleoptera (Klug, 1833). Cryphtsus was founded on the American species Cr. Boothi, Green, of which Cr. collitelus, Green, which possesses a median terminal spine to the pygidium as well as the live lateral pairs, was considered merely a variety by Hall.1 Corda, in dealing with the similar European species C. arachnoides (Hoeninghaus) and G. stellifer, Burmeister, considered the presence or absence of the terminal spine a feature of generic importance; and accordingly put them in separate genera, which he called Asteropyge and Metacanthus respectively. Salter2 was the first to deuounce the artificiality and weakness of this division; so that, as Cryphcem is not available as a generic or group name, Asteropyge must be used as its designation. Kayser3 has endeavoured to subdivide Cryphmus again into Corda's two sections. Though the type-species of Dalmania is D. caudatus (Briinn.) and that of Odontochile 0. Hausmanni (Brong.), (or as Barrande- declares D. Reussi, Barr.), yet it was early recognised that these species were congeneric. Both Dalmania and Odontochile were unfortunately preoccupied, the former by a genus of Diptera (Robineau Desvoidy, 1830) and the latter by a genus of Coleoptera (Odontocheila, Laporte, 1834), so that Barrande proposed the name Dalmanites, which though often used in a very wide sense, yet must hold good as the group - name of which the type - species is D. caudatus. Hall & Clarke4 employed the name Dalmanites in such a wide sense that they proposed the name Hausmannia for the eub-group containing the type-species ; but Oehlert5 considers it an unnecessary duplicate name. Acaste must be considered as preoccupied, the name Acasta having been given by Leach in 1811 to a genus of Cirripedes. There is no strict synonym for this group, so that the name must be 1 Hall & Clarke : op. cit., p. 45. 2 Salter: Mem. Geol. Surv., dec. ii, 1849, art. 1, p. 8. 3 Kayser: "Die Fauna alt. Dev. Ablag. Harzes," p. 33 (Abhandl. geol. spec. Kart. Preuss., Bd. ii, Heft 4, 1878). 4 Hall & Clarke: op. cit., p. xxxi. 5 Oehlert: Bull. Soc. Geol. France, ser. in, vol. xvii (1889), p. 758. Downloaded from https://www.cambridge.org/core. INSEAD, on 18 Jul 2018 at 18:49:05, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756800131991 174 F. JR. Coicper Reed—On the P/iacopidce. replaced by a new one if the morphological or phylogenetic claims of the group which it marks are suflicient to warrant a distinctive title. The divisional names, therefore, which remain after this pre- liminary weeding-out are the following :— ' Acaste.' Monorakos. Asteropyge. Odontocephahis. Chasmops. Phacops. Coronura. Portlockia. Corycephalus. Probolinm. Cryphina. Pterygometopus. Dalmanites. Sonuitrikelon. Homalops. Synphoi'ia. Malladaia. Trimerocephalus. Whether all the above divisions have sufficient morphological or ^)hylogenetic value to be retained in a natural system of classification is the question now to be discussed. Principles of Classification. Dismissing from our minds all ideas and prejudices derived from customary usage and past authorities, it is worth while considering in what manner we should group the members and species of the Phacopidaa supposing we were to start completely de novo. The principles for a natural arrangement are acknowledged to be found in the modifications of the head-shield, as we have learned especially from Professor Beecher's valuable researches. In the case of the members of a compact family such as the Phacopidae the changes in the glabella offer the best guide, and these changes are shown by the fusion or reduction or even disappearance of certain lobes. The modifications in the thorax, which always retains the fixed number of eleven segments, are of secondary morphological value and of minor developmental importance; and while there is an immense amount of variation in the characters of the pygidium in shape, number of segments, and ornamentation, yet the evidence of the trilobites as a whole and of individual families (e.g. the Cheiruridse) indicates that structural changes in this part of the body occur in less regular succession and pursue often an independent or eccentric development, so that for the purposes of a classification which is intended to be based on phylogeny they are of less significance than those in the head-shield. Starting, then, from the well-established view of the primitive pentamerous composition of the head-shield of trilobites, we may expect to find this pentameric segmentation best exhibited in the earliest and least modified members of the family. The phylogeny may be anticipated to follow the ontogeny which has been traced in more than one species of Phacops. The group termed Dalmanites by Barrande, using the name in its widest and most comprehensive sense, most clearly preserves the fivefold division of the head-shield. In the early post-embryonic stages of its species, as well as at maturity, the five segments are •well displayed. The retention of this primitive character throughout Downloaded from https://www.cambridge.org/core. INSEAD, on 18 Jul 2018 at 18:49:05, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756800131991 F.
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