[Sphenophyllum].

The account of the Sphenophyllales given in the first volume[2] of this work must be extended and somewhat modified in the light of recent work on the fertile shoots of Sphenophyllum.

Sphenophyllostachys Dawsoni (Will.) was described as consisting of an axis bearing superposed whorls of bracts connate at the base in the form of a shallow funnel-shaped collar giving off from the upper surface and close to the axis of the cone two concentric series of sporangiophores. Occasionally there are three series, as represented in [fig. 112]. In another type of strobilus, Sphenophyllostachys Römeri[3] each sporangiophore terminates in two pendulous sporangia ([fig. 113], A; see also fig. 107, C, vol. I.). It has already been pointed out that the common occurrence of detached strobili necessitates their description under distinct specific names; it is only by a rare accident that we can assign fossil cones to their vegetative shoots. There are, however, reasons for believing that Sphenophyllostachys Dawsoni is the strobilus of the plant originally described by Sternberg[4] from impressions of foliage-shoots as Rotularia cuneifolia. Another difficulty presented by petrified material is that of determining, with certainty, whether two imperfect specimens, differing from one another in features which do not appear to be of sufficient importance to warrant specific separation, are forms of one species or portions of specifically distinct cones. It has been pointed out by Scott[5] that the strobilus known as Sphenophyllostachys Dawsoni probably includes two distinct species, one being the cone of Sphenophyllum cuneifolium Sternb., and the other the cone of S. myriophyllum Crép[6]. The stem of S. myriophyllum agrees anatomically with the type known as Sphenophyllum plurifoliatum Will. and Scott[7].

Fig. 112. Sketch of a radial longitudinal section of Sphenophyllostachys. There are usually two concentric series of sporangia on the sporophylls, not three as shown in the figure. The upper figure (after Zeiller) shows the linear bracts in surface-view.

In addition to the two types of cone already mentioned, Sphenophyllostachys Dawsoni and S. Römeri, others have been described by Kidston from carbonised impressions. One of these is the fertile branch of Sphenophyllum majus[8]. The basal portions of the bracts of each whorl form a narrow collar round the axis of the cone; the free portion of each bract consists of a lamina divided into two equal bifid lobes bearing on its upper surface one group, or possibly two groups, of four sessile sporangia between the narrow coherent bases of the laminae and the sinus between the terminal lobes ([fig. 113], C). Another characteristic feature is the greater length of the internodes; this renders the cone less compact and less sharply differentiated from the vegetative shoots than those of other species. A specimen in Dr Kidston’s collection illustrates the peculiar character of the fertile portion of this species; it consists of an axis bearing a succession of lax sporophylls succeeded above and below by whorls of sterile leaves. In this species, therefore, we cannot speak of a compact strobilus at the end of a shoot of limited growth, but of axes in which sterile and fertile leaves are borne alternately[9], a condition recalling the alternation of foliage leaves and sporophylls in Tmesipteris and in Lycopodium Selago.

Fig. 113.

  1. Sphenophyllostachys Römeri. (Solms-Laubach.)
  2. Sphenophyllum trichomatosum Stur.
  3. Sphenophyllum majus. Bronn. (A–C. After Kidston.)

Another form of cone, also from the Middle Coal Measures, is referred by Kidston to Sphenophyllum trichomatosum Stur[10] ([fig. 113], B): this is characterised by the more horizontal position of the bracts, which “do not appear to be so much or so suddenly bent upwards in their distal portion as in some other species of Sphenophyllum,” and by sessile sporangia borne singly on the upper face of each bract.