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 Botany. 
 
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 No. I, — Nematophyton Ortoni, n.sp. 
 
 BV 
 
 D. P. Penhallow. 
 
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 , [Reprinted from the Annals of Botany, Vol. X, No. 37, Mnrcli, 1896.] 
 
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 Nematophyton Ortoni, n.sp. 
 
 BY 
 
 D. P. PENHALLOW, 
 
 McGill University, Montreal. 
 With Plate V. 
 
 SINCE my last general summary of the genus Nemato- 
 phyton *, two new species from Europe have been re- 
 corded, one by Mr. C. A. Barber from the Tymawr quarry, 
 near Cardiff, Wales, and named by him N. Storriei'^ ; the 
 other from the shales in the neighbourhood of Grafrath on 
 the lower Rhine, and designated by Count Solms-Laubach 
 Nematophyton dechenianum ^ 
 
 I have now to describe a third, which differs in very im- 
 portant respects from all previously recorded species, from 
 the middle or upper division of the great Ohio shales. This 
 specimen was obtained by Professor Edward Orton, of the 
 Geological Survey of Ohio, from whom the facts relating to 
 its origin have been obtained, and after whom I would name 
 it Nematophyton Ortoni. 
 
 ' Notes on Devonian Plants. Tmns. R. See. Can. VII. iv. 19. 
 
 ' Nematophycus Slorriei, Ann. Hot. VI. 329. 
 
 ' Ueber devonische Fflonzenreste aus den Lenneschiefem der Gegend von 
 Grafrath am Niederrhein. Jahrbuch der konigl. preuss. gcologischeii Landes- 
 anstalt. 1894, 67. 
 
 lAnnats of Botany, Vol. X. No. XXXVII. March. 1896.] 
 
 B 
 
 'I 
 
42 
 
 Penhallow. — Ncmatophyton Orioni, n. sp. 
 
 The hand-specimen transmitted to us for examination is 
 15 cm. high and 14 cm. broad at the base, from which it 
 gradually tapers upward in such a way as to suggest that 
 it must be the base of a stem. This idea is strengthened by 
 the presence of a number of basal and lateral processes with 
 their outward terminations presenting fractured surfaces, show- 
 ing that they represent the bases of more or less prolonged 
 organs such as roots. Between these processes are rounded 
 indentations, such as may be supposed to have been made by 
 large pebbles. In fact, to those who are familiar with the 
 branching base of a Laminarian stipe, the resemblance between 
 the two is very striking. But Prof. Orton informs me that 
 this specimen was originally part of a very much larger one 
 which was broken in getting it out of the shale. The largest 
 fragment thus obtained measured about ao inches long and 
 8 to 10 inches wide. A photograph of it shows the surface 
 to be irregularly indented with several processes, the whole 
 presenting the general aspect of the smaller specimen taken 
 from it. It is clear then that the original plant must have 
 been one of rather large dimensions, on the scale of a tree, 
 and the general external characters of the specimens justify 
 the view that they represent the base of a stem or stipe at the 
 point whence the roots issue. 
 
 Internally the specimen is highly silicified throughout, 
 and shows no evidence whatever of concentric structure. 
 Externally there are limited areas covered with a friable, 
 carbonaceous film which, in places, attains a thickness of 
 about 1-5 mm., but for the most part it is very thin. It in 
 all probability represents the carbonised residue of the cortical 
 tissue, although microscopical examination fails to disclose 
 any definite structure. We can therefore only assume that it 
 represents cortex, a view, however, which finds its justification 
 in our general knowledge of the alterations in such structures 
 under similar conditions. 
 
 The first sections examined were received from Prof. Orton, 
 but they got badly broken in transit, in consequence of which, 
 and also for the purpose of ascertaining the relation of s»^^ruc- 
 
Penhallow. — Nematophyion Ortoni, n.sp. 43 
 
 ture to special locations, a new series of sections was prepared 
 at the Peter Redpath Museum. It was found upon examina- 
 tion that there were no material differences of structure 
 exhibited by the two sets, so that no special reference to 
 them is required. The structure was found to be most 
 beautifully preserved, and in this respect it is comparable 
 with the Gaspd specimens of N. Logani. 
 
 In transverse section the structure is seen to consist of 
 numerous round, thick-walled, and rather \videly separated 
 cells of rather uniform size, the larger having a diameter of 
 about 67 ju. Between these cells others are to be seen 
 running more or less transversely at various angles, and there- 
 fore interlacing with the first or longitudinal series. The 
 spaces between the cells are occupied by small crystals of 
 silica and a limited amount of carbonaceous matter, which 
 shows no disposition indicative of structure. So far very 
 searching examinations have failed to disclose anything of 
 the nature of smaller hyphae such as distinguish the structure 
 of N. Logani, N. crassum, and N. laxum, and it is impossible 
 to say if such were at any time present, though the presence 
 of carbonaceous matter among the siliceous crystals would 
 seem to suggest the possibility of hyphae, or at least of some 
 structure, having been present, since it has already been shown 
 in the case of N. Logani and N. crassum, that the carbona- 
 ceous matter of the original structure often becomes distributed 
 in such ways through the influence of crystallization ^ 
 
 There is no evidence whatever of radial spaces such as 
 were found in N. Logani. But there are rather numerous 
 isodiam i i': areas, about 0-30 mm. broad, occupied entirely 
 (Fig- 3) ^J]/ masses of very narrow and densely interlacing 
 hyphae, having a diameter of 4-8 fx. In the sections so far 
 submitted to examination, the structure of these hyphae has, 
 in most cases, been destroyed by crystallization, and in the 
 few instances where the structure was intact it was not possible 
 to ascertain if they were septate, though the occurrence of 
 
 ' M. Micr. Jill. X. 69, 70. Trans. R. Soc. Can. VII. iv. 23 ; Proc. U. S. Nat. 
 Mus. XVI. lis. 
 
 B a 
 
 I 
 
44 Penhallow, — Nematophylon Ortoni, n. sp. 
 
 septa in the similar hyphae of N. Logani, as determined by 
 Mr. Barber ^ would seem to indicate the possibility of their 
 presence in this case. In no instance do these hyphae appear 
 to cling to the walls of the large tubes of the medulla as in 
 N. S torrid, but they fill isolated cavities which, for want of 
 a better term, I may designate as the medullary spots. These 
 spots exhibit the same general form and dimensions in all 
 planes of section, and on smoothly cut surfaces of the hand- 
 specimen they often appear as small cavities about i-o mm. 
 broad, frequently containing minute crystals. We thus have 
 evidence that these spots are not always occupied by small 
 y hyphae, a fact which suggests either (i) that the spaces are 
 > normal, and the hyphae intrusive growths, or (2) that the 
 small hyphae are normal, and in some cases have been 
 removed by decay or other causes. Which of these views 
 is correct our material does not permit me to say; but the 
 view advanced by Mr. Barber, and strengthened by our know- 
 ledge of similar openings in the various known species of 
 Nematophyton, that these spaces have some connexion with 
 the aeration of the plant ^ would seem to offer a reasonable 
 explanation of their occurrence. It may also be pointed out 
 that the most marked alterations of structure, through decay (?) 
 "ind crystallization, to be met with in the present species are 
 found in the hyphae of these areas. With respect to the 
 occurrence of these medullary spots, the present species ap- 
 proaches somewhat closely to N. crasstim. 
 
 In longitudinal section it is wholly unnecessary to dis- 
 tinguish between the radial and tangential planes, since the 
 structure presents the same aspects in each case. The struc- 
 ture consists of broad, tubular cells running in a direction 
 generally parallel to the axis of growth, together with others 
 less numerous, but yet in large numbers, traversing the stem 
 in all directions, so that unlike the species hitherto described, 
 there is a want of definiteness in direction. Occasionally 
 these cells are exposed to a great length, but more commonly 
 the plane of section cuts them off at frequent intervals so 
 
 » Ann. Bot. VI. 333. » Ann. Bot. VI. 337. 
 
 I 
 
] 
 
 Penhallow. — Nematophyton Or tout, n.sp. 45 
 
 that only short fragments appear (Figs. 2 and 3). A marked 
 pecuh'arity of the cells in this plant is the frequency with 
 which local expansions of the lumen occur. These we can 
 only regard as representing the trumpet-hyphae and situations 
 of sieve-plates so common in the Laminarieae. Although in 
 the majority of cases no sieve-plate could be observed, in 
 a few instances the fact of such structures having been 
 present was quite obvious (Figs, a and 3). One of the 
 trumpet-hyphae is shown on a much larger scale in Fig. 5. 
 I have had no very good opportunity of instituting a com- 
 parison between these structures and the forms occurring in 
 the larger species of the Laminarieae. My chief comparison, 
 therefore, has been with the forms common to the North 
 Atlantic coast ; but through the kindness of Dr. W. G. Farlow, 
 it has been possible to take into consideration Macrocystis 
 pyrifera^. Although differing in detail, the general character 
 of these structures in Nematophyton and Macrocystis is so 
 similar as to suggest the belief that our fossil is related to 
 those modern types of seaweeds of which Macrocystis is an 
 example. As in the transverse section, no small hyphae are 
 to be found between the large cells, but the latter are seen 
 to branch somewhat frequently and always, so far as de- 
 termined from the present material, in the immediate vicinity 
 of a medullary spot (Fig. 4). 
 
 From the details thus outlined it is clear that the plant is 
 an Alga, and of an alliance with the Laminarias. Having 
 regard to the general character of the stem-structure, it is 
 evident from our specimens that the cortical layer was rela- 
 tively thin, the medulla strongly predominating, and in these 
 respects the stem presents features which are well represented 
 by Laminaria digitata. 
 
 This species differs from all others so far known, in the 
 very loose character of the medulla, a feature which may be 
 characteristic of the species as a whole, but which may belong 
 more particularly to certain regions of the plant, and this 
 
 * In this connexion compariion may be made with the varions forms of trumpet- 
 hyphae in Macrocystis, as figured by Prof. F. W. Oliyer in Ann. of Bot. I. 95. 
 
 1-. 
 I 
 

 V 
 
 46 Penhalloiv. — Ncviatophyton Or tout, n.sp. 
 
 view gains strength from the fact that our sections were 
 apparently taken from one of the hapteres near its junction 
 with the principal axis. 
 
 In his article on N, Storriei, Mr. Barber finds it difficult to 
 agree with my views respecting the organic connexion between 
 the large cells of the medulla and the small hyphae, basing 
 his objections upon examinations of N. Logani and N. Stor- 
 riei; but in view of the evidence at hand, it is impossible 
 for me to accept the modifications he suggests'. Evidently 
 when his article was written he had not seen my second 
 paper on Nematophyton ^ in which five species are described. 
 In speaking of N. crassmn ^ I then made use of the following 
 description : — 
 
 ' The most significant fact so far observed, consists in the discovery 
 of a distinctly branching system, similar in its general character to that 
 of N. Logani, though differing from it in some important respects. 
 In one case I found a branch projecting from the side of a large cell, 
 with a diameter of 5-8 /x and a length to the point where cut off of about 
 35 /i. Two other branches near together were each 4-6^; two more 
 were 2-3 fi and 4-6 /x; another 6-9 /* in diameter ... It was therefore 
 clear that the larger cells of this plant branch into a secondary 
 plexus as in N. Logani, and as all of the instances in which the 
 branches were seen to emanate from the larger cells occurred in the 
 open tracts above described, it would appear that these latter serve 
 as the special regions in which branching is effected.' 
 
 We have here, then, the fact that in A^. crassmn the large 
 cells do branch into small filaments of the same general 
 diameter as the small hyphae of the spaces, and that such 
 branching takes place where these hyphae are most abundant. 
 If this be taken in connexion with Mr. Barber's admission 
 that the !iranching is most frequent in the region of the 
 spaces, it will be seen that there are good grounds for 
 denying his contention with respect to N. Logani, N. crassmn^ 
 and N. laxiim, while the presumption would be in favour of 
 
 ' Ann. of Bot. VI. 335. 
 
 ' Trans. R. Soc. Can. VII. iv. 19. 
 ' L. c, p. 32. 
 
■TiR- 
 
 PenhallouK — Nematophyton Ortoni, n.sp. 47 
 
 regarding similar structural conditions to exist in the other 
 species where the evidence is not so well defined. Never- 
 theless, it must be remembered that the genus Nematophyton, 
 as we now know it, is made up of several apparently distinct 
 species of whose entire structure we know but little : and it is 
 quite within the range of possibility — as is even now sug- 
 gested by the striking structural differences presented by 
 N. Ortoni— that when we are able to reconstruct the entire 
 organism in each case, it may be found that more than one 
 genus is represented, or that some of those which now appear 
 distinct, may in reality be different parts of the same species. 
 Our present knowledge of the genus Nematophyton shows 
 that it embraces what appear to be eight distinct species as 
 follows: — 
 
 1. N. Logani, Dn. Lower Erian of Gaspd ; Silurian (Upper 
 Ludlow) of England and Silurian (Cap Bon Ami) of New 
 Brunswick. {Dazvson.) 
 
 2. N. Hicksii (Etb ;, Dn. Denbighshire grit (Silurian) of 
 Wales. (Bteks.) 
 
 3. N. orassum (Dn.), Penh. Middle Erian of Gasp^ (Be//) ; 
 Hamilton Group (Middle Erian) of New York. [C/arke and 
 Prosser^ 
 
 4. N. laxum (Dn.), Penh. Lower Erian of Gaspe. {Be//) 
 
 5. M". tenue (Dn.), Penh. Lower Erian of Gaspe. [Be//) 
 
 6. N. Storriei, Barb. Silurian (Wenlock Age) of Cardiff, 
 Wales. {Storrie.) 
 
 7. N. dechenianum, Solms-Laub. Upper Devonian of 
 Grafrath, Germany. {So/ms-Laubac/t.) 
 
 8. N. Ortoni, n. sp. Upper Erian of Ohio. (Orton.) 
 
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 48 Penhallmv. — Nematophyton Or torn, n. sp. 
 
 EXPLANATION OF FIGURES IN PLATE V. 
 
 Illustrating Prof. Penhallow's paper on Nematophyton Ortoni, n. sp. 
 Description of Figures. 
 
 Fig. 1. Transverse section showing the general character of the structure. X45. 
 
 Fig. 2. Longitudinal section showing the general character of the medulla, with 
 trumpet-hyphae, and on the extreme left of the field a portion of a medullary spot. 
 X45- 
 
 Fig. 3. I-ongitudinal section showing the occasional exposure of the tubular 
 cells for a great length, trumpet-hyphae, and two medullary spots on the left. 
 X4S' 
 
 Fig. 4. Transverse section showing a cell of the medulla with, at the centre of 
 the figure, three branches. The darker area of the field on the upper side is the 
 side of a medullary spot, x 1 70. 
 
 Fig. 5. Longitudinal section showing trumpet-hyphae at centre and left side, 
 with crystals of silica occupying the spaces between the large tubular cells, x 170. 
 
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