ON SOME SEEDS FROM THE COAL-BEARING DEPOSITS OF THE KUZBASS

M. D. PARFENOVA

The present paper gives a description of several seeds from the Balakhonka and Kolchugino series of the Kuzbass; in most cases we sought to study not only their morphology, but also the cellular structure of their integument, and, when the available material permitted, to study their individual variability.

The genera of fossil seeds are distinguished by purely morphological characters, and, although these morphological genera are fairly well differentiated from one another, in a number of cases they are possibly composite and in the future will be divided according to the anatomical structure of the epidermis. Thus, many seeds of the genus Cordaicarpus that have lost the sarcotesta are externally very similar to seed nucelli of other genera; therefore, when the sarcotesta is lost during burial—which is quite possible—the latter acquire a great morphological resemblance to seeds of the genus Carpolithes. In this case the study of cellular structure may be of decisive importance (see below, p. 187).

Key for determining the morphological genera of seeds

1. Wing absent . . . . . . . . . . . . . . . . . . . . . 2
   — Wing present: Samaropsis Goeppert.

2. The seed has a sarcotesta, which on impressions surrounds the nucellus in the form of a ridge, keel, leathery or fleshy covering . . . 3
   — Sarcotesta absent; the surface of the seed smooth or finely wrinkled: Carpolithes Sternberg.

3. Sarcotesta in the form of a small ridge or keel. The seed nucellus is clearly visible . . . . . . . . . . . . . . . . . 4
   — Sarcotesta is a wrinkled fleshy or leathery covering; the wrinkles of the sarcotesta almost obscure and mask the outlines of the nucellus: Angarocarpus Radczenko.

4. Seeds bulb-shaped, bud-like, or heart-shaped in outline, with an elongated, almost pointed apex and a heart-shaped, notched base: Cardiocarpus Brongniart.
   — Seeds oval or rounded-oval, narrowly rounded or slightly pointed at the apex; the trace of attachment of the seed to the stalk in the form of a small notch: Cordaicarpus Geinitz.

The study of the anatomical structure of the epidermis helps to establish more precise diagnoses of species; at times morphologically identical seeds have different anatomical structure, which we have had to encoun-

...be dealt with in the present work. But, since the anatomical structure of the epidermis is not always preserved, the morphological species must be retained, even if they are composite.

In the course of studying the anatomical structure of the epidermis of seeds, it became clear that its character in different species shows considerable diversity. The different types of cells of which the epidermis consists are sometimes repeated in different species. Therefore, in order to avoid repetition in the descriptions of species, we consider it advisable to give descriptions of the cell types, which we distinguish with regard to absolute dimensions, shape, combination and mutual orientation of the cells, and also the thickness and character of the cell wall. For the present we distinguish seven types (Fig. 1).

Fig. 1. Types of cells. Explanations in the text.

1. Polygonal cells of medium size, more often quadrangular–pentagonal. The cell wall is of medium thickness (3–4 μ), more rarely thin (1–2 μ). The cells closely adjoin one another and in most cases are oriented along the length of the seed, forming parallel rows, more or less distinct. The dimensions of cells of this type are 24.9–30.3 × 9.5–12.2 μ. Usually such cells occur with type 3, not infrequently forming gradual transitions to this type by an increase in size and the loss of the quadrangular outline. Most often cells of this type are situated in the middle part of the seed or near its apex.

2. Cells long and narrow, generally quadrangular, sometimes spindle-shaped with pointed ends. The width and length of the cells vary considerably in different species of seeds. The dimensions of the cells are 10.1–20.1 × 57.2–112.4 μ. Such cells usually occur along the margin of the lateral part of the seed. The walls of the cells are parallel or nonparallel.

3. Polygonal cells, rather large, with a cell wall of medium thickness (3–4 μ). The cells closely adjoin one another, without orientation or slightly oriented; their dimensions are 14.3–38.1 × 32.9–73 μ.

4. Cells very close to those just described, but distinguished by being more often rounded and forming intercellular spaces. They resemble broken links of a chain. The dimensions of such cells are 23.3–42.4 × 13.8–24.4 μ.

5. Cells large, polygonal and rounded, with a thin and medium-thick cell wall. Rounded cells form intercellular spaces. Dimensions 73.1–126.1 × 37.1–73.1 μ.

6. Cells quadrangular, sometimes rhombic, with straight walls equal to one another, closely adjoining one another. Dimensions 30.7–53 × 24.4–60.4 μ.

7. Elongated polygonal cells with a thick cell wall (5–8 μ). The cell walls are somewhat sinuous and closely adjoin one another. Combinations of three cells are often seen: two other, longer cells adjoin the narrow side of one cell (Table IV, fig. 3). Dimensions of the cells 11.7–57 × 27.6–116.6 μ.

The cell types described occur in different combinations in different species of seeds. Here the types with the most distinct charac-

by the signs of the cells, but there are impressions whose cells cannot be confidently assigned to any definite type, either because of imperfect preservation or because of the transitional character of the cells (see p. 181). In most seeds the cells of the episperm and of the nucellus are different, and, as a rule, the cells of the nucellus are better preserved, whereas the cells of the episperm are satisfactorily preserved only exceptionally rarely.

Of course, the types listed do not exhaust the entire diversity of cells. It is evident that, with further study, it will become possible to distinguish new types.

Without giving a review of all the stratigraphic schemes of the Kuzbass, we shall very briefly discuss the scheme proposed by S. G. Gorelova, since it is this scheme that we followed in our work.

S. G. Gorelova, in the bank section along the Tom River, divides the Erunakovskaya Formation into 2 members according to seam 60. The lower member includes three horizons: “a”—from seam 19 to seam 38, corresponding approximately to the Uskat subformation; “b”—from seam 38 to seam 49, and “v”—from seam 49 to seam 60—together corresponding to the Leninskaya subformation. The upper member includes two horizons: “g”—from seam 60 to seam 69, and “d”—from seam 69 and higher [2].

In studying individual variability, as the principal variants we took: \(L\)—the total length of the seed, and \(a\)—its greatest width. Determination of the arithmetic mean, standard deviation, coefficient of variation, and coefficient of correlation was carried out according to the formulas already presented by the author in a published work [1].

The present work was carried out in the Department of Historical Geology of the Tomsk Polytechnic Institute.

Genus Angarocarpus Radczenko, 1955.

1. Angarocarpus tychtensis (Zalessky) Radczenko.

Pl. I, figs. 1, 2.

1. Samaropsis tychtensis Zalessky. Contrib. a la flore perm. du bassin de Kousnetzk, p. 135, fig. 17.
2. Samaropsis tychtensis Neuburg. Upper Paleozoic flora of the Kuznetsk Basin, pp. 287, 283, pl. XXIII, figs. 20a—d, 21.
3. Angarocarpus tychtensis Radczenko. Atlas of guide forms of the fossil fauna and flora of Western Siberia, p. 150, pl. XIV, fig. 11.

Holotype originates from the Kuznetsk Basin, from the left bank of the Tykhta River near the village of Sokolovaya, Erunakovskaya Formation. Pl. 1, fig. 1.

Diagnosis. Seeds of obovate outline or bulb-shaped, nearly flat. The apex of the seed is drawn out into a long, sharp beak; at the base of the seed there is a deep notch—the scar from attachment to the fruit stalk. On the seed the central part—the nucellus—and the surrounding peripheral border—the sarcotesta—are quite clearly distinguished. The surface of the seed is covered with fine concentric wrinkles.

Dimensions: length 4.5—9 mm, width of seeds—5—14 mm (in one case—18 mm).

Variability: in our collection there are 31 specimens of this species. In the overwhelming majority the width of the seeds exceeds their length; seeds in which the length is equal to the width or slightly exceeds it are very rare. The greatest number of seeds have dimensions within the limits of 5.5—8.5 mm in length and 8—12 mm in width; in general, with increasing seed size, its width increases more rapidly than its length.

The ratio \(\frac{a}{L}\) varies from 1 to 2.3; in general, with an increase in the absolute dimensions of the seed, the ratio of its width \((a)\) to its length \((L; \text{fig. } 2)\) regularly increases. The greatest number of seeds has a ratio

of width to length, equal to 1.3; this is clearly visible in Fig. 3; apparently, the additional zigzags of the curve shown in Fig. 3 are explained by the limited number of specimens of this species available to us.

As the graphs show, all the seed specimens of this species available to us, with respect to their dimensions, form a compact group whose members are connected by gradual transitions; the exception is a specimen (museum No. 137—715), which differs from the rest already by its unusually large dimensions \((a = 18\ \mathrm{mm})\); possibly in this case we are dealing with some special form.

Fig. 2. Change in the value \(\dfrac{a}{L}\) in seeds of Angarocarpus tychtensis depending on the change in seed width \((a)\).

Fig. 3. Angarocarpus tychtensis. Variation curve.

The arithmetic mean is \(a = 5.74\), for \(L = 9.74\). The mean square deviation, which is equal to \(a = 1.4\) and to \(L = 2.6\), as well as the coefficient of variability, equal to \(a = 20.8\%\) and to \(L — 26.7\%\), po-

show that the greatest degree of variation in the seeds of this species is possessed by seed length. The value of the correlation coefficient for these characters (it is equal to 0.48) indicates the presence of a partial, but moderate direct correlation between them.

No definite dependence between the ratio $\dfrac{a}{L}$ and the distribution of the seeds through the section (Fig. 4) can be detected: seeds with different values of the ratio $\dfrac{a}{L}$ occur at one and the same stratigraphic level, and conversely. But, by paying attention to the maximum values of this ratio, one can note (Fig. 4) the appearance of increasingly broad seeds in the higher stratigraphic beds.

Fig. 4. Distribution of seeds of Angarocarpus tychtensis in the section of the Erunakovskaya suite.

higher stratigraphic beds. This is especially emphasized by a comparison of seeds at three levels: at the level of the group of beds 33–38, beds 45–50, and beds 59–65. For each of these levels we have about ten seeds; the maximum values of $\dfrac{a}{L}$ at these levels, in ascending order, are as follows: 1.6; 1.9; 2.0–2.3. Despite the limited number of specimens, this can hardly be explained only by chance.

not accidental: apparently, a certain direction in the evolution of this group of plants is discernible here. This is also confirmed by the fact that the widest seeds appear at the moment of the greatest quantitative development of the species A. tychtensis, which occurs precisely at the level of seams 63–65, above which this species disappears entirely (Fig. 4).

Comparisons. The species Angarocarpus tychtensis does not differ morphologically from Angarocarpus princeps sp. n. and from the majority of the seeds of Angarocarpus variabilis sp. n. We retain the name Angarocarpus tychtensis for seeds that have not been studied anatomically, whereas the species Angarocarpus princeps and Angarocarpus variabilis are distinguished by the structure of the epidermis.

It differs from Angarocarpus dubius and Angarocarpus trigonalis in its kidney-shaped outline and in the position of the greatest width in the middle of the seed, whereas in the above-mentioned species it is displaced to varying degrees toward the apex of the seed.

Distribution. Upper part of the Ilyinskaya Formation; in the Erunakovskaya Formation—horizons “a,” “b,” “v,” “g”; in the Beregovoi section it appears at the level of seam 19 and disappears at the level of seam 65, reaching its greatest development at the level of seam 63.

Localities. Everywhere in the areas of development of the deposits of the Ilyinskaya and Erunakovskaya formations.

2. Angarocarpus dubius sp. n.

Table 1, figs. 3, 8.

Holotype. Collection of the Tomsk Polytechnic Institute. Specimen No. 85–213. Kuznetsk Basin, Nikitinskoye locality; lower part of the Erunakovskaya Formation. Table 1, fig. 3.

Diagnosis. Seeds flat, with rounded-triangular outlines. The widest part of the seed is situated near the apex. The ratio
\[ \frac{b^{1)}}{c} \]
is 0.3–0.75. With a strong displacement of the greatest width of the seed toward the apex, the seed often acquires the shape of a triangle with rounded angles. The apex of the seed is slightly drawn out in the form of a sharp papilla, the length of which reaches 1 mm. The nucellus is almost the same shape as the seed as a whole, but noticeably smaller. The sarcotesta reaches a width of 3 mm, and at the place of attachment of the seed, at the base of the nucellus, it is absent. The surface of the seeds is covered with longitudinal wrinkles. On many seeds a small rib is clearly visible along the middle of the seed; it increases in thickness from the apex toward the base, reaching a width of 1 mm.

Dimensions: seed length 5.5–13.5 mm, seed width 6.5–14 mm.

Comparisons. Angarocarpus dubius has some similarity to Angarocarpus tychtensis (Zal.) Radcz., but is easily distinguished from it by the considerable displacement of the greatest width of the seed toward its apex, in connection with which the outlines of the seed often acquire a rounded-triangular character. There are eight specimens; the ratio of the seed width to its length changes quite gradually within the limits from 1 to 1.85.

Remarks. The anatomical structure of the epidermis was not studied.

Distribution. Erunakovskaya Formation, from the tops of horizon “a” to the lower parts of horizon “d.”

Localities. Belovskoye, Plotnikovskoye, Leninskoye, Nikitinskoye localities, Erunakovsky district (seams 35–50).

^1) \(b\)—distance from the apex of the seed to the line of greatest width; \(c\)—distance from the line of greatest width to the base of the seed.

3. Angarocarpus trigonalis sp. n.

Plate 1, figs. 4, 9, 14.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 24—82. Kuznetsk Basin, Raspadskoye locality, above the 9th coal seam. Plate 1, fig. 4.

Diagnosis. Seed triangular in outline, flattened. The apex is slightly broadly drawn out and pointed. The seed is attached to the stalk by a broad base, leaving a trace of attachment in the form of a broad strip or a slightly arc-shaped notch. The broadest part of the seed is situated at the very apex. The sarcotesta on the impressions surrounds the nucellus as a narrow border. In the places of greatest seed width it is somewhat broader (up to 2 mm), and then, gradually narrowing toward the apex and base, almost disappears. The nucellus is of the same shape as the seed and is poorly visible. On the surface of the seed, arc-shaped longitudinal wrinkles are visible.

Structure of the epidermis. The cells are polygonal, predominantly small, more rarely of medium size, and closely adjoin one another. The cell wall is thick; the cells are not oriented (type 3, but the dimensions are smaller than usual for this type). The character of the cells is the same over the entire surface of the seed.

Dimensions: seed length 10.5 mm, seed width 17 mm, length of the seed apex 2.5 mm.

Comparisons. Morphologically, the described species is somewhat similar to Angarocarpus dubius sp. n., but differs in its triangular outline, the almost straight apical side of the seed, and its larger dimensions.

In the structure of the epidermis this species is very similar to Carpolithes virgatus sp. n., but differs in the absence of longitudinal bands of elongated and very narrow cells, which are so characteristic of Carpolithes virgatus.

Remarks: this species was established by us on the basis of a single specimen, possessing very characteristic morphological and anatomical features that distinguish it from all other species.

Distribution. The very lowest parts of the Erunakovskaya suite, horizon “a”.

Localities. Raspadskoye locality, 9th coal seam.

4. Angarocarpus variabilis sp. n.

Plate 1, figs. 7, 10, 11, 12, 13, 15, 16, 17.

Holotype. Collection of the West Siberian Geological Administration, specimen 15/30—50. Kuznetsk Basin, Erunakovsky district, above the 42nd coal seam. Plate 1, fig. 12.

Diagnosis: seeds flat, bud-shaped or inversely bulb-shaped, with the apex drawn out in the form of a small papilla. At the base, at the place where the seed was attached to the stalk, there is an arc-shaped notch. The greatest width of the seed is situated in the middle or is shifted considerably toward the apex. The leathery and wrinkled sarcotesta covers the nucellus with a thin layer. The nucellus is poorly visible and has the same outlines as the seed.

Structure of the epidermis. The cells are rather large, rounded, and form intercellular spaces. The cell wall is of medium thickness. An indistinct orientation of the cells along the length of the seed is apparent (type 4). Near the lateral margins of the seed the cells are elongated (type 2); the cells become quite narrow and long in the apical part (2.7 × 19.2 μ), where their rows converge in a bundle toward the strongly drawn-out papilla.

Dimensions: seed length 5—9 mm, seed width 5.5—11.5 mm, length of apex 1.5—2.5 mm.

Comparisons. Angarocarpus variabilis has some similarity in the structure of the integument to Angarocarpus psinceps sp. n., but is readily distinguished from it by the character of the epidermis in the apical part, where

rows of narrow and long cells converge in a bundle toward a strongly elongated papilla, which is entirely absent in Angarocarpus princeps.

Remarks. The species is distinguished by the character of the anatomical structure of the epidermis. In terms of morphological characters it unites specimens similar to Angarocarpus tychtensis (Zal.) Radcz. and Angarocarpus dubius sp. n. Variability has not been traced because of the insufficient quantity of material.

Distribution. The Erunakovo suite, from the very top of horizon “a” to the lower part of horizon “г”.

Localities. Erunakovo district, from bed 37 to bed 63; Plotnikovo district; Leninsk district, Brusnitsyn and Veretenovo seams.

5. Angarocarpus princeps sp. n.

Table II, figs. 1, 6, 9, 11.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 11—55 [1]. Kuznetsk Basin, Raspadskaya locality, below seam 13. Table II, fig. 9.

Diagnosis. Morphologically the seeds of this species do not differ from Angarocarpus tychtensis (Zal.) Radcz. (p. 167).

Structure of the epidermis. The cells have a rather thick membrane, adjoin one another closely, and are oriented along the seed, but the shape and size of the cells in different parts of the seed are not the same. In the near-papillar part the cells are four- and five-sided, from small to medium-sized (type 1). In the central part and at the base of the seed the cells are rather large, polygonal (more often hexagonal), not distinctly oriented (type 3). Along the margin of the lateral part of the seed the cells are long and narrow, quadrangular, sometimes spindle-shaped, with pointed ends (type 2).

All types of cells pass into one another gradually, without sharp changes.

Dimensions: seed length—5–17 mm, seed width—5–20 mm.

Variability. We studied 56 specimens of this species. Most seeds have a length within the limits of 6–9 mm, a width of 8–13 mm. Shi-

Fig. 5. Change in the value \(\frac{a}{L}\) in seeds of Angarocarpus princeps as a function of change in seed width \((a)\).

The width of the seeds almost always exceeds the length and in the rarest cases is equal to it. The ratio $\frac{a}{L}$ varies continuously from 1 to 2.2.

In general, with an increase in seed size the value of the ratio $\frac{a}{L}$ increases, i.e., the relative width of the seeds increases; only one specimen (museum No. 37/672) sharply departs from this rule, in which, at maximum dimensions, the width only slightly exceeds the length (Fig. 5). This sharp deviation may characterize a special form (variety?). The dependence of the ratio $\frac{a}{L}$ and the number of seeds (Fig. 6) is expressed by a broken curve, which at first rises rather rapidly, at a ratio $\frac{a}{L}=1.2—1.4$ reaches a maximum, and then decreases stepwise to a minimum at points with $\frac{a}{L}=2—2.2$. The greatest number of seeds has the ratio $\frac{a}{L}=1.2—1.4$, i.e., on average—1.3. There is not observed, as in the species Ang. tychtensis (Fig. 4), any definite dependence between the distribution of seeds in the section and the change in the ratio $\frac{a}{L}$ (Fig. 7).

Fig. 6. Angarocarpus princeps. Variation curve.

The occurrence, noted for Ang. tychtensis (p. 170), of broader seeds in higher stratigraphic beds is here disrupted by two specimens from the Leninsk district; they have high values of $\frac{a}{L}$ (respectively 2.0 and 1.8), but come from the lower part of the Yerunakovo suite. If one takes into account the considerable remoteness of the Leninsk district from the Berezovo section and the still imperfect parallelization of the sections of these two districts, then perhaps decisive significance should not be attached to these specimens. And if they are excluded from consideration, then the graphs presented in Fig. 3 and Fig. 6 and characterizing the dis—

the distribution of seeds of A. tychtensis and A. princeps in the shoreline section gives a picture of complete coincidence: on them three maxima of quantitative development of both species are distinguished (bed intervals 33—38, 44—50 and 63—66), the disappearance of both species above the level of beds 65—66, and the regular appearance of broader seeds upward in the section. Let us recall that both these species have the same morphology, but for the former the anatomical structure of the epidermis is unknown. A pairwise comparison of Figs. 4 and 7, Figs. 3 and 6, Figs. 2 and 5 may speak in favor of the assumption that these are one species. If this assumption were confirmed by subsequent observations on the structure of the epidermis in seeds from the collections of M. D. Zalessky, in particular in the holotype, these species should be united, abolishing the name Ang. princeps.

The arithmetic means for the characters \(a\) and \(L\) are 5.44 and 3.78. The standard deviation, equal for \(a\) to 3.02 and for \(L\) to 2.04, and the coefficient of variability, which is equal for \(a\) to 26.9%, and for \(L\) to 26.4%, show that in this species the width of the seeds possesses a somewhat greater degree of variation than their length. The correlation coefficient for these characters is 0.44, which indicates the presence of a partial, but moderate, direct correlation between these characters.

Fig. 7. Distribution of seeds of Angarosarpus princeps in the section of the Erunakovo suite.

Comparisons. Morphologically, Angarocarpus princeps does not differ from Angarocarpus tychtensis (Zal.) Radcz. and from the seeds of Angarocarpus variabilis sp. n., which have the widest part in the middle of the seed. Angarocarpus princeps is readily distinguished from all other species by the anatomical structure of the epidermis characteristic of it.

Distribution. Erunakovskaya Formation, horizons “a,” “b,” “v,” “g” up to coal seam 66.

Localities. Erunakovskii District, seams 37—66; Belovskii District; Leninskii District, Maksimovskii, Veretenovskii, Polenovskii, Nadbaikaimskii 1, Poddyagilevskii, Boldyrevskii, Emel’yanovskii, Brusnitsynskii seams; Raspadskii District, seam 13.

6. Angarocarpus sagittarius sp. n.

Plate II, figs. 7, 8, 10, 13.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 21/272. Kuznetsk Basin, Plotnikovskii District, Erunakovskaya Formation, seam XXIII. Plate II, fig. 10.

Diagnosis. Seed small, generally oval, but the drawn-out sharp apex and the acute-angled notch at the base give it, to some degree, sagittate outlines. The greatest width passes approximately through the middle of the seed. The seed nucellus is oval. The sarcotesta on impressions is 1 mm wide. On the surface of the seed, slight longitudinal wrinkling is noticeable.

Structure of the epidermis. The epidermal cells within the contours of the nucellus are polygonal, elongated, closely adjoining one another, oriented along the length of the nucellus. The cell wall is thin or of medium thickness (type 2 with broader cells). Toward the apex and base of the nucellus the cells become shorter and more polygonal (type 3), and toward the very ends of the nucellus, decreasing in size, the cells approach type 1. The epidermal cells of the sarcotesta have not been preserved.

Dimensions: seed length 6—6.5 mm, seed width 3—3.5 mm, nucellus length 4—4.5 mm, nucellus width 2—2.5 mm.

Comparisons. The seeds of the described species are morphologically similar to the Balakhonka species Angarocarpus ungensis (Zal.) Radcz., but differ in their smaller dimensions and in the fact that the nucellus is located in the center of the seed, whereas in Angarocarpus ungensis it is shifted toward the base [3, p. 276]; moreover, in the described species there is no straight rib running from the base of the notch to the apex of the nucellus.

Distribution. Erunakovskaya Formation, seams 37 and 48.

Localities. Erunakovskii District, seams 37, 48. Plotnikovskii District, seam XXIII.

7. Angarocarpus proprius sp. n.

Plate I, figs. 5, 6.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 178—1009. Kuzbass, Chumyshskii District, Nizhnebalakhonskaya Formation. Plate I, fig. 5.

Diagnosis. Seeds of medium and rather large size, of triangular outline. The angle at the base is truncated and has a small notch. The upper side of the seed is strongly concave, as a result of which the bluntly rounded upper corners seem drawn upward. The apex of the seed is situated in the center of the concave side, slightly drawn out and pointed. The greatest width of the seed is located near the apex. The margin at the base is slightly convex. The nucellus has the same outlines, but its apex is still more drawn out and pointed. The nucellus is covered with faint longitudinal wrinkling.

Dimensions: seed length 10.5—13 mm, seed width 15.5—23 mm, base width 2—6 mm.

Comparisons. Angarocarpus proprius has a considerable resemblance to A. trigonalis sp. n. from the Erunakovo suite. It differs from it in having the angles at the apex of the seed more broadly rounded and drawn upward, convex margins, and a considerable width of the sarcotesta at the base, whereas in A. trigonalis the sarcotesta at the base tapers off to nothing.

For comparisons with Samaropsis stila sp. n. and S. ampla sp. n., see below, in the descriptions of the corresponding species.

Remarks. There are three specimens of this species, two of them of good preservation.

Distribution. Lower Balakhonka suite.

Locality. Chumysh district.

8. Angarocarpus enodatus sp. n.

Plate II, figs. 2, 3.

Holotype. Collection of the West-Siberian Geological Administration, specimen No. 22/256. Kuzbass, Zmeinogorsk district, Tomsk II seam, upper part of the Alykaevo subsuite. Plate II, fig. 3.

Diagnosis. Seeds of medium size, oval in shape. The sarcotesta, almost equal in width to the border, surrounds the nucellus, forming only at the apex a small acute-angled notch. The nucellus is oval, with somewhat pointed ends. The greatest width of the seed is situated in the middle. A distinct longitudinal groove runs through the whole seed along the middle. The entire seed is covered with coarse longitudinal wrinkles.

Dimensions: seed length — 10–19 mm; seed width 7.5–10 mm; nucellus length 11–12 mm, nucellus width 5.5–7 mm.

Epidermis consists of hexagonal, regular, more rarely elongated cells with a thin cell wall, oriented along the length of the seed.

Comparisons. Angarocarpus enodatus has a remote resemblance to A. tyrganensis (Zal.) [3, p. 285]. It differs from it in the presence of an acute-angled notch at the apex and in the much larger dimensions of the nucellus.

Remarks. Of five specimens, macroscopically of fairly good preservation, only in one was it possible to observe some elements of the anatomical structure of the epidermis. Hexagonal, more often regular, more rarely elongated cells are visible, oriented along the length of the seed, with a thin cell wall (Plate II, fig. 2).

Distribution. Upper part of the Alykaevo subsuite.

Localities. Zmeinogorsk district, Tomsk II and Opolznevyi seams.

Cardiocarpus Brongniart, 1828

9. Cardiocarpus corculum sp. n.

Plate III, figs. 6, 7, 11, 14, 15.

Holotype. Collection of the West-Siberian Geological Administration, specimen No. 12/126. Kuznetsk Basin. Erunakovo district, above seam 28. Plate III, figs. 14, 15.

Diagnosis. Seed of heart-shaped outline, with a somewhat pointed apex and a slightly notched base. The nucellus is of the same outline, but the apex is more pointed, even drawn out in the form of a sharp papilla. On the impressions, a narrow (0.8 mm), uniform border of sarcotesta surrounds the nucellus. Longitudinal wrinkling is noticeable. The widest part of the seed is closer to the base.

Structure of the epidermis. For this species, differentiation of the epidermal cells in different parts of the seed is characteristic. The cells of the nucellus are polygonal, rather large, with a cell wall of thin or medium thickness. The type of cells is close to 5. Near the lateral sides of the seed the cells become narrower and therefore approach type 2. Such same

cells are located along the edge of the cordate notch at the base. Toward the apex of the seed the cells rather rapidly shorten and become characteristic of type 3.

Dimensions: seed length 5 mm, seed width 5 mm, apex length 3.5 mm.

Comparisons: The species Cardiocarpus corculum differs from the most closely related Cardiocarpus minutus Radcz. in its less attenuated and slightly pointed apex. The anatomical structure of the epidermis in our species is very characteristic; in Cardiocarpus minutus Radcz. it is unknown.

Remarks. The integument of the sarcotesta of the seed consists of cells of another type. The boundary between the sarcotesta and the nucellus is clearly visible under the microscope, but the preservation of the specimen does not permit the morphology of the sarcotesta cells to be described; apparently they are close to type 1.

Distribution. Erunakovskaya Suite, horizon “a”.

Localities. Erunakovskii district, bed 28.

10. Cardiocarpus strictus sp. n.

Plate III, fig. 12.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 22/223. Kuzbass, Zmeinskii district, Zmeinskii II site. Lower part of the Alykaevskaya Subsuite. Plate III, fig. 12.

Diagnosis. Seeds of medium size, rounded-cordate. Apex slightly pointed; at the base a small cordate notch is noticeable.

Dimensions: seed length 7.5–10.5 mm, seed width 6.5–10 mm.

Comparisons. This species has some similarity to Cardiocarpus corculum sp. n., but differs from it in a less pronounced cordate outline and in the absence of a sarcotesta.

Remarks. The species is described from the two specimens available to us. Because of the absence of a sarcotesta, it should be assigned to the genus Carpolithes, but, taking into account the cordate outlines of the seeds, so characteristic of the genus Cardiocarpus, and the fact that the sarcotesta may often have been lost in the process of burial, we refer our species to the latter genus.

Distribution. Alykaevskaya Subsuite.

Localities. Zmeinskii district, Zmeinskii II bed; Chumyshskii district.

Genus Samaropsis Goeppert, 1865

11. Samaropsis angularis sp. n.

Plate II, fig. 12.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen 206—1136. Kuznetsk Basin, Leninskoye locality, above the Boldyrevsky bed. Plate II, fig. 12.

Diagnosis. Seed of medium size, outline close to quadrangular. A clearly defined elongate-ovoid nucellus with a pointed apex is distinguished. The wing is very broad—on each side of the nucellus its width exceeds the width of the nucellus, but it tapers to nothing at the base and forms a shallow cordate notch at the apex of the seed.

Dimensions: seed length 10 mm, seed width 8 mm, nucellus length 8 mm, nucellus width 4 mm.

Comparisons. The species described is quite readily distinguished from all others by its quadrangular outline, the wing characteristic of it, and the acute apex of the nucellus.

Remarks. The structure of the epidermis has not been described because of its poor preservation.

Distribution. Erunakovskaya Suite, horizon “a”.

Locality. Leninskoye locality, above the Boldyrevsky bed.

12. Samaropsis stila sp. n.

Plate II, figs. 4, 5.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 22/220. Kuzbass, Zmeinskii district, Pervomaiskii seam IIIa. Ishanovskaia subformation. Plate III, figs. 4, 5.

Diagnosis. Seed of medium size, somewhat triangular in outline. The greatest width of the seed is situated in its uppermost part. The apex and the upper corners of the seed are drawn out, so that the upper margin of the seed is twice concave. The other two sides are slightly convex. The base is not broad, arcuately concave; from it there extends a short stalk. The seed nucellus is round and is not quite distinctly differentiated against the background of the wing.

Dimensions: seed length 8 mm, seed width 14 mm, nucellus diameter 6.5 mm.

Comparisons. The described species has some resemblance to Samaropsis ampla sp. n. and Angarocarpus proprius sp. n. It differs from the first in the slightly pointed upper corners of the seed, drawn upward, the convex margins at the base, and the round nucellus; from the second, in the pointed upper corners of the seed, the round nucellus, and the absence of a wing.

Remarks. The species is described from a single specimen; the anatomical structure of its epidermis has not been studied, since imprints of the cells were not preserved. Macroscopically, the preservation of the specimen is quite satisfactory and makes it possible to give a complete morphological description of the species.

Distribution. Ishanovskaia subformation.

Localities. Zmeinskii district, Poryvaikii seam IIIa.

13. Samaropsis ampla sp. n.

Plate II, fig. 14.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 22/140. Kuznetsk Basin. Zmeinskii district, Opolznevoi seam. Upper part of the Alykaevskaia subformation. Plate II, fig. 14.

Diagnosis. Seed of rather large size, triangular in outline, with wing-like wings. The greatest width of the seed lies almost at its very apex. The apex is slightly drawn out and pointed. The corners of the upper part of the seed are bluntly rounded. The margins at the base are slightly concave. The base is arcuately concave. The nucellus is spindle-shaped, poorly visible.

Dimensions: seed length 11 mm, seed width 21 mm, apex length 3.5 mm, base width ~3.5 mm.

Comparisons. See the description of the species S. stila sp. n. In the outline of the seed there is a certain resemblance to Angarocarpus proprius sp. n., from which the species described differs in that in the latter the corners in the upper part of the seed are drawn upward, the margins at the base are somewhat convex, and the nucellus is triangular in outline, whereas in the described species the corners at the apex of the seeds are not drawn out, the margins at the base are slightly concave, and the nucellus is spindle-shaped.

Remarks. There is only one specimen, on which the description of the species is based. Despite the fairly good preservation of this specimen, its microstructure has not been preserved.

Distribution. Upper part of the Alykaevskaia subformation.

Localities. Zmeinskii district, Opolznevoi seam.

14. Samaropsis acutula sp. n.

Plate III, figs. 9, 10, 13, 16.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen 1164, Kuzbass, Kiselevskii district, area—field of mine No. 6. Balakhonskaia series. Plate III, fig. 16.

Diagnosis. Seeds elongated, of medium size, with a somewhat narrowed upper end. At the base of the seed there is a heart-shaped

notch; in the apical part the notch is deeply arcuate, so that the ends of the upper part of the seed appear strongly drawn out. The nucellus is oval in shape, its ends are pointed, and the basal part is even slightly drawn out. The seed is covered with barely noticeable fine longitudinal wrinkles, which are more visible on the nucellus and less so on the wing.

Structure of the epidermis of the nucellus and wing differs in the shape of the cells: the epidermis of the nucellus consists of rounded-polygonal cells of medium and small size, closely adjoining one another, with a cell wall of medium thickness. The cells of the wing are elongated, narrow, and well oriented along the seed.

Dimensions: seed length 14–16 mm, seed width 9–9.5 mm, nucellus length 11.5–12 mm, nucellus width 6 mm.

Comparisons. In the outline of the seed, the present species resembles S. skokii Neub. It differs from it in the presence of a heart-shaped notch at the base and in the straight longitudinal grooves of the nucellus, whereas in S. skokii the grooves of the nucellus are sinuous, usually joined in loops.

Remarks. This species is represented in our material by three specimens of varying preservation. The best of them is taken as the holotype. The microstructure of the epidermis is, as a rule, poorly preserved. Only in specimen No. 1164 can some elements of its structure be seen. The epidermis of the nucellus consists of rounded-polygonal cells, of medium and small size, compact, with a cell wall of medium thickness. Narrow longitudinal strips of two, more rarely three, rows of elongated narrow cells are visible, by which, in the main, a certain orientation of the cells along the seed is created. The cells of the wing are elongated, narrow, more often quadrangular, and well oriented along the seed. The boundary between the cells of the nucellus and of the wing is quite clearly visible.

Distribution. Balakhon Series.

Localities. Shushtulenskiy district, Ananyinskiy tract. Kiselevskiy district, area of mine field No. 6.

Carpolithes Sternberg, 1823.

1. Carpolithes simplex sp. n.

Plate III, fig. 4.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 12/16. Kuzbass. Erunakovskiy district, seam 35. Plate III, fig. 4.

Diagnosis. Seeds flat, round, 4–5 mm in diameter; at the base a trace of attachment is noticeable. The surface of the seed is slightly wrinkled.

Comparisons. The seeds of the described species somewhat resemble Carpolithes minuta Radcz. [4, pl. X, figs. 7, 8], but differ from it in that in Carpolithes minuta the apex is short and slightly drawn out, whereas in Carpolithes simplex the apex is not differentiated, and, with poor preservation, it is difficult to determine the position of the trace of attachment.

In Carpolithes rostriformis (Neub.) the apex is drawn out in the form of a thin, long, beak-like process, by which it differs sharply from the described species.

Remarks. This species is most likely a composite one. It has been distinguished on morphological characters.

Distribution. Erunakovskaya Formation, horizons “a” and “b” from seams 31 to 43. Balakhon Series, Mazurovskaya and Alykaevskaya subformations.

Localities. Erunakovskiy district, seams 31, 35, 39, 42, 43. Zmeinskiy district, Kolokoltsevskiy seam; Chumyshskiy district, Pervyi and Podapanasovskiy seams.

16. Carpolithes ineditus sp. n.

Plate III, fig. 3.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 12/32. Kuzbass, Erunakovskii district, seam 31. Plate III, fig. 3.

Diagnosis. Seeds small, somewhat elongated, overall oval in shape. The apex and base are slightly pointed. Sometimes only the apex is pointed, or else the apex and base are blunt, and then the seed has an oval shape. Longitudinal wrinkling is noticeable, and some seeds have a distinct longitudinal groove along the middle of the seed.

Dimensions: seed length 4–6 mm, seed width 3–5 mm.

Comparison. The species described has some resemblance to Carpolithes minuta Radcz. [4, pl. X, figs. 7 and 8], but it is not difficult to distinguish it by the character of the apex; in Carpolithes minuta the apex is short and slightly drawn out. In Carpolithes ineditus the apex is never drawn out; it is either pointed or bluntly rounded.

From Carpolithes simplex sp. n., which includes seeds of rounded form, the species described differs in its slightly elongated, oval outlines.

Variability. In our collection there are 36 specimens of this species. The absolute dimensions of the seeds fluctuate within narrow limits. The greatest number of seeds have a length from 5 to 6 mm and a width from 3 to 4.5 mm. The length of the seed exceeds the width, and only in exceptional cases is equal to it; the ratio

\[ \frac{a^{1)}}{L} \]

varies from 0.6 to 1, generally increasing somewhat with increase in seed size; almost all seeds have the ratio

\[ \frac{a}{L} \]

within the limits 0.6–0.8.

The arithmetic mean for \(a = 5.44\) and for \(L = 3.78\). The values of the standard deviation and especially of the coefficient of variation for these characters of the given species are very close. Consequently, the degree of variation of these characters is almost the same. Their correlation coefficient is 0.42, which indicates the presence of a partial, but moderate, direct correlation between these characters.

Fig. 8. Distribution of seeds of Carpolithes ineditus in the section of the Erunakovskaya suite.

Stratigraphically, the seeds of this species are distributed from the 12th to the 59th coal seam (fig. 8). The curve expressing the quantita—

\(^{1)}\) \(a\) — seed width. \(L\) — seed length.

distribution of seeds through the section forms two peaks at the level of seams 31 and 38.

Remarks. Under the name Carpolithes ineditus we have combined seeds with unknown anatomical structure. It is quite possible that this species is composite: below are described morphologically similar species differing from it in the cellular structure of the integument. The possibility is not excluded that among the specimens assigned to this species there are even seeds of the genus Cordaicarpus Gein, in which the sarcotesta was lost in the process of burial (see below, p. 188).

Distribution. Upper part of the Ilyinskaya Formation and the lower part of the Erunakovo Formation. In the indicated formations this species is widely distributed within a very narrow stratigraphic interval (approximately seams 330—40), within which it gives two maxima of quantitative development (seams 31 and 38). Above and below the indicated interval this species occurs in single specimens, disappearing completely below seam 12 and above seam 59 (Fig. 8). Entirely similar seeds also occur in the Balakhon Series (Mazurovo and Ishanovo subformations).

Localities. Plotnikovo district. Erunakovo district, seams 12—59. Kondoma district, seams 9, 7, 21. Zmeinka district, seams Opolznevy, Tomsky II, Molodezhny, Nizhny, Poryvaevsky, Chumysh district, seam Apanasovsky I.

17. Carpolithes levis sp. n.

Plate III, figs. 2, 5, 8.

Holotype. Collection of the West Siberian Geological Administration, specimen No. K-12/28-50. Kuznetsk Basin, Erunakovo district, above the 38th coal seam. Plate III, fig. 5.

Diagnosis. Seed flat, round or slightly elongate. The ratio of width to length in most cases is equal to 1 and rarely reaches 1.5. At the base a trace of attachment is visible in the form of a small scar or the corresponding notch. The seed is covered with longitudinal wrinkles.

Structure of the epidermis. Cells polygonal, slightly elongate, from rather large to medium and small. The cells closely adjoin one another and are oriented along the seed. The cell wall is of medium thickness. The cell type may be defined as transitional between 1 and 3, since cells of both of the indicated types occur here. The character of the cells over the entire surface of the seed is the same.

Dimensions: seed length 3.5—6 mm, seed width 3.5—6 mm.

Comparison. Morphologically, this species has great similarity to a number of other species of the genus Carpolithes Sternb., except for the species Carpolithes rostriformis (Neub.) and Carpolithes minuta Radcz., from which it differs by its undifferentiated apex. In the structure of the epidermis, Carpolithes levis differs from other species by the type of cells, which is defined as transitional between 1 and 3, and by the fact that the character of the cells over the entire surface of the seed is the same.

Distribution. Erunakovo Formation, horizons “a,” “b.”

Localities. Erunakovo district, seams 28, 38, 48.

18. Carpolithes pulchella sp. n.

Plate III, fig. 1; Plate IV, figs. 3, 7.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 111-918. Kuznetsk Basin. Erunakovo district, above seam 38. Plate 4, fig. 7.

Diagnosis. Seed flat, round or slightly elongated; the surface of the seed is covered with longitudinal wrinkles; sometimes a longitudinal groove is observed along the middle of the seed.

Epidermal structure. The cells are polygonal, often curved, some with a slightly sinuous wall, large, and closely adjoining one another. Sometimes characteristic groups of three cells stand out: two others adjoin the smaller side of one cell (type 7, Plate IV, fig. 3). There is no orientation of the cells. Toward the apex and base the cells gradually become smaller, the cell walls become less sinuous, the number of angles in the outlines of the cells decreases, and the cells acquire a certain orientation along the seed. Elongating and decreasing in width, the cells seem to converge in a bundle toward the base and apex.

Dimensions: seed length 4–5 mm, seed width 3.5–5 mm.

Comparisons. Morphologically this species is similar to Carpolithes simplex, Carpolithes levis, Carpolithes lucidus, and others, but is readily distinguished from all these species by the structure of the epidermis that is typical for it.

Remarks. In one specimen, in contrast to the others, the cell walls of the bud do not form sinuations and are usually straight. Possibly this indicates differences in the conditions of existence, since it is known that with frequent changes in temperature and humidity the cell walls become more sinuous than under comparatively constant values of these factors.

Distribution. Erunakovo Formation, horizon “b.”

Localities. Erunakovo district, bed 38; Plotnikov district, bed XXV;

19. Carpolithes lucidus sp. n.

Plate IV, figs. 2, 6, 9.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 29-59. Kuznetsk Basin. Raspadskoye deposit, below bed 3. Plate 4, fig. 6.

Diagnosis. Seed rounded-oval, flat, smooth; the width slightly exceeds the length (ratio of width to length equals 1.2). The broadest part is in the middle of the seed.

Epidermal structure. The cells are rather diverse both in size and in shape. Here there occur elongate polygonal cells (type 2), and small polygonal cells close to type 1, and large cells of polygonal form with blunted angles (type 5a). The cell envelope is rather thick. No definite orientation of the cells is noticeable, but cells of different form and size alternate irregularly with one another in inconstant combinations, forming many intercellular spaces. In one place at the margin of the seed the cells are larger than usual (type 5a). Evidently this is the apex or the base, but since no traces of attachment are visible, it is difficult to decide whether this increase in cell size will characterize the proximal or distal part of the seed.

Dimensions: seed length 4.5 mm, seed width 5.2 mm.

Comparisons. Morphologically Carpolithes lucidus has great similarity to Carpolithes simplex, Carpolithes pulchella, Carpolithes levis. It differs from all the indicated species by the peculiar anatomical structure of the epidermis: the considerable diversity of cell types and the character of their combinations (Plate IV, figs. 2, 9).

Distribution. Erunakovo Formation, lower unit, lower part of horizon “a.”

Locality. Raspadskoye deposit; bed 3.

20. Carpolithes virgatus sp. n.

Plate IV, figs. 1, 5.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 182-1014. Kuznetsk Basin. Nikitinskoye deposit. Plate IV, fig. 5.

Diagnosis. The seed is round, most likely lentil-shaped. The greatest width is in the middle part of the seed. At the base, at the place where the seed was attached to the pedicel, there is a small arc-shaped notch. The entire seed is covered with fine longitudinal grooves.

Structure of the epidermis. Cells of medium size, polygonal (type 3). The cell wall is thin. The cells are weakly oriented along the seed, but the principal orientation is due to narrow bands that extend over the entire length of the seed and sometimes anastomose with one another. These bands are situated at varying distances from one another and consist of several rows of very narrow and elongated cells (type 2).

Dimensions: seed length 6 mm, seed width 6.5 mm.

Comparisons. Morphologically, Carpolithes virgatus can easily be confused with Carpolithes lucidus, Carpolithes simplex, Carpolithes pulchella, and Carpolithes levis, but it is readily distinguished from all these species by the structure of the epidermis: characteristic of this species are narrow longitudinal, anastomosing bands, which consist of several rows of very narrow, elongated cells.

Distribution. Lower part of the Erunakovskaya Formation.

Localities. Nikitinskoye deposit.

21. Carpolithes ineptus, sp. n.

Plate IV, figs. 4, 8, 10.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 21/299. Kuznetsk Basin, Plotnikovskoye deposit, below coal seam XVIII. Plate IV, fig. 8.

Diagnosis. Seeds slightly elongated, with pointed ends; surface slightly wrinkled. At the base, a trace of attachment is visible.

Structure of the epidermis. Cells somewhat elongated along the length of the seed, polygonal (type 1). Such cells are located near the lateral sides of the seed. In the center of the seed, rounded, irregularly polygonal cells of different sizes are less clearly visible, but they are larger than those along the sides of the seed.

Dimensions: seed length 4–5.5 mm, seed width 3.5 mm.

Comparisons. Morphologically, Carpolithes ineptus does not differ from Carpolithes ineditus and Carpolithes procerulus. The described species differs in the nature of the epidermal structure—namely, no other species has, near the lateral sides of the seed, cells of this type (see diagnosis); equally characteristic of it are the cells of the central part of the seed.

Distribution. Erunakovskaya Formation, lower unit, horizon “a”.

Localities. Plotnikovsky district, seam XVIII; Erunakovsky district, seam 30.

22. Carpolithes procerulus, sp. n.

Plate V, figs. 1, 4, 7, 9, 11, 12.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 96-921. Kuznetsk Basin, Erunakovsky district, near the village of Markina. Plate V, fig. 12.

Diagnosis. Small, slightly elongated or rounded seeds with pointed ends, usually with traces of attachment at the base; slightly longitudinally wrinkled.

Structure of the epidermis. In the central part of the seed the cells are of medium size, polygonal, with a cell wall of medium thickness (type 3). Toward the apex of the seed the cells are compact, elongated along the length of the seed (type 2); along the margin of the lateral side the cells are very large (type 5), polygonal, elongated lengthwise. Toward the base of the seed the cells are also polygonal and fairly large, but no longer elongated lengthwise.

Dimensions: seed length 4.5–6 mm, seed width 3.5–4 mm.

Variability. The collection contains 21 specimens of this species. The dimensions of the seeds, as in other representatives of this genus, vary within narrow limits (see dimensions); most seeds are 5.5–6 mm long and 3.5–4 mm wide. The length always exceeds the width and only in exceptionally rare cases is equal to it. The ratio $\frac{a}{L}$ varies from 0.5 to 1; the greatest number of seeds has a ratio $\frac{a}{L} = 0.7–0.8$ (Fig. 9).

Stratigraphically, seeds of this species occur only within coal seams 25–39. The greatest number of specimens is found in coal seam 38; there also occurs the greatest variety in the dimensions of these seeds. In the remaining indicated horizons, seeds of C. procerulus are distributed insignificantly and approximately uniformly (Fig. 10).

Comparisons. Carpolithes procerulus does not differ in external outline from Carpolithes ineditus sp. n. and Carpolithes ineptus sp. n., but is readily distinguished by the anatomical structure of the epidermis; it is characterized by different types of cells in the central, apical, basal, and lateral parts of the seed.

Fig. 9. Carpolithes procerulus. Variation curve.

Fig. 10. Distribution of seeds of Carpolithes procerulus in the section of the Erunakovskaya suite.

Distribution. Erunakovskaya suite, horizon “a” and the lower part of horizon “b.”

Localities. Erunakovskii district, seams 25, 28, 30, 35, 37, 38, 39.

23. Carpolithes rostriformis (Neuburg)

Plate V, fig. 3.

1. Samaropsis [?] rostriformis Neuburg. Upper Paleozoic flora of the Kuznetsk Basin, p. 286, pl. XXIII, fig. 19.

Holotype. Collection of the Institute of Geological Sciences, Academy of Sciences of the USSR, specimen No. 2276-86. Kuznetsk Basin, Prokopyevsk, Tserkovnyi log, roof of the Moshchnyi seam. Upper Balakhonka Series. Pl. XXIII, fig. 19.

Diagnosis. Seed oval in outline, with an apex sharply drawn out into a beak-like appendage.

Dimensions: length of the seed without the appendage 7 mm and more, width of the seed 5–7 mm, length of the appendage 11–13 mm, width of the appendage 0.75–2 mm.

Comparisons. By its characteristic beak-like elongated apex, this species is readily distinguished from all other species of fossil seeds from the Kuzbass.

Remarks. At present only two specimens of this species are known: one from the materials of M. F. Neuburg (see synonymy); the other was found by us in the Shushtulep district, in the lower horizons of the Balakhonka Series. In our specimen the main part of the seed has not been preserved, but on the basis of the characteristic apex we consider it possible to assign our specimen to this species.

M. F. Neuburg, pointing to the possible presence of a wing in such seeds, tentatively assigned this species to the genus Samaropsis. The specimen found by us shows the absence of a wing in these seeds. On the basis of the morphological character, we assign seeds of this type to the genus Carpolithes Sternb.

Distribution. Balakhonka Series.

Localities. Shushtulep district, Ananyinskii area; Prokopyevsk, Tserkovnyi log, roof of the Moshchnyi seam.

24. Carpolithes agitabilis, sp. n.

Pl. V, figs. 2, 8.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 1153. Kuzbass, Shushtulep district, Ananyinskii area, Afanasovskii seam. Lower Balakhonka Formation.

Diagnosis. Seeds small, oval or broadly oval; at the base they have a barely noticeable trace of attachment in the form of a very small notch. Seeds smooth; only some show weak wrinkling near the base; arranged on a twig in the form of a raceme.

Dimensions: length of the seed 2.5–4 mm, width of the seed 2–3 mm.

Comparisons. In outline, the seeds of the species described have some similarity to the above-described C. ineditus sp. n. and to C. simplex sp. n. They differ from both species in their very small size.

Remarks. Seeds of similar outline and size occur rather often in the lower horizons of the Balakhonka Series, but, as a rule, scattered in groups of several seeds. On one specimen a twig has been preserved, and on it several seeds. Evidently such seeds grew on a twig in the form of a raceme and could easily fall off.

Distribution. Lower Balakhonka Formation.

Locality. Shushtulep district, Ananyinskii area, Afanasovskii seam.

25. Carpolithes novicus sp. n.

Pl. V, figs. 6, 5, 10, 14, 15.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 22/268. Kuzbass, Emeinskii district, Korshnyi seam. Alykaevka Subformation. Pl. V, fig. 4.

Diagnosis. Seed round, with a slightly noticeably drawn-out apex. A longitudinal groove passes along the middle of the seed.

Epidermis consists of cells of four types: along the very margin of the seed the cells are narrow and long; at the base (?) the cells are of medium size, polygonal, closely adjoining one another, some with a curved wall. At the apex (?) the cells are large, polygonal, more often quadrangular, with straight cell walls, closely adjoining one another. In the center of the seed the cells are also rather large, many-

angular, rounded or slightly elongated, represent a transitional type between the cells of the apex and the base.

All cells of the seed fit closely to one another and have a thin cell wall.

Dimensions. Seed length 7.5 mm, seed width 8 mm.

Comparisons. In outline the seed has a great similarity to Carpolithes simplex sp. n. It differs from it in the presence of a longitudinal furrow, a slightly drawn-out apex, and also in the distinctive structure of the epidermis (see diagnosis).

Remarks. This species is represented in our collection by only a single specimen. Taking into account the good preservation, which makes it possible to characterize the anatomical structure of the epidermis, we establish a new species on the basis of one specimen, despite its rather simple morphology.

Distribution. Alykaevskaya subformation.

Locality. Zmeinsky district, Korshunny seam.

Genus Cordaicarpus Geinitz, 1861.

26. Cordaicarpus amictus sp. n.

Plate V, fig. 13.

Holotype. Collection of the West-Siberian Geological Administration, specimen No. 20/130. Kuznetsk Basin. Erunakovsky district, above seam 65. Plate V, fig. 13.

Diagnosis. Seeds of small size, oval; on impressions the sarcotesta, in the form of a rim (3 mm wide), surrounds the nucellus. The apex and base of the seed are drawn out and pointed, the base being drawn out and pointed more strongly than the apex. The sarcotesta in these places reaches its maximum width. The nucellus is oval; at the base it has a trace of attachment in the form of a dot, from which a barely noticeable furrow runs toward the base of the seed through the sarcotesta.

Dimensions: seed length 5—8.5 mm, seed width 3—6.5 mm.

Comparisons. The species described is, by its form, fairly easily distinguished from the others. It has some similarity to Cordaicarpus ellipticus Radcz. and Cordaicarpus iljinskiensis Radcz., but differs from both in its drawn-out and pointed base, where the sarcotesta reaches its greatest width, whereas in the former the base is slightly drawn out in the form of a short stalk, and the sarcotesta is interrupted at this place; in the latter the base is bluntly rounded.

Remarks. The species has been distinguished on morphological characters. The structure of the epidermis has not been studied. Most likely, this species is composite, and it is quite possible that anatomically unstudied representatives of the species Cordaicarpus notulus sp. n., Cordaicarpus fascis sp. n., Carpolithes procerulus sp. n., which have similar morphology, belong to it.

Distribution. Erunakovskaya suite, upper part of horizon “a,” horizons “b,” “v,” and the lower part of horizon “g.” Balakhonskaya series, middle subformation of the Prokopyevsk district.

Localities. Erunakovsky district, seams 30, 31, 49, 65. Prokopyevsk-Kiselevsk district.

27. Cordaicarpus notulus sp. n.

Plate VI, figs. 1, 3, 5, 6, 7, 8.

Holotype. Collection of the Tomsk Polytechnic Institute, specimen No. 204-1128. Kuznetsk Basin, Nikitinskoye locality. Erunakovskaya suite. Plate VI, fig. 6.

Diagnosis. Seeds of oval or rounded outline, of small size. On impressions the sarcotesta surrounds the nucellus as a narrow rim (0.5 mm) and is not always preserved. At the apex the sarcotesta is somewhat

widens and reaches 1 mm in width. The nucellus is oval, covered with fine longitudinal wrinkles.

Structure of the epidermis. The cells are rather large, with thick walls, quadrangular, closely adjoining one another (type 6). In microscopic preparations, round bright points of undetermined nature are visible in the cell wall, more often at the cell corners. The orientation of the cells is varied, to some extent intersecting: the cells are often arranged in parallel rows, with several such rows occurring at different angles (Pl. VI, figs. 3, 8). In the parallel rows of cells the transverse walls are thicker and therefore more clearly visible. In general the cells are the same over the entire surface of the seed.

Dimensions: seed length 5.5–7 mm, seed width 5.5 mm, nucellus length 5.5 mm, nucellus width 3.8 mm.

Remarks. The species is distinguished on the basis of two specimens that have the same anatomical structure of the epidermis. In one specimen (No. 18/8) the sarcotesta is absent; evidently it was lost in the course of burial. On the basis of the identical anatomical structure, we assign these specimens to one species.

Comparisons. Morphologically, if the sarcotesta has not been preserved, the described species can easily be confused with some representatives of the genus Carpolithes Sternb., and also with the species Cordaiсarpus fascis sp. n., but by the structure of the epidermis Cordaiсarpus notulus is readily distinguished from all other species.

Distribution. Lower horizons of the Erunakovskaya Formation.

Localities. Kasminsky district and the Nikitinskoye locality.

28. Cordaiсarpus fascis sp. n.

Pl. VI, figs. 2, 4, 9.

Holotype. Collection of the West Siberian Geological Administration, specimen No. 12/49. Kuznetsk Basin, Erunakovsky district, below seam 37. Pl. VI, fig. 9.

Diagnosis. Seeds of small size, slightly oval, almost round. Apex and base slightly pointed. Nucellus of the same outline. The sarcotesta on the impressions surrounds the nucellus as a narrow rim (up to 2.2 mm wide); it is not always preserved. The surface of the seed is covered with slightly noticeable longitudinal wrinkles.

Structure of the epidermis. The cells were apparently elongated along the seed, since striae are visible in that direction; most likely these are remnants of cell walls. Near the margins the cells are larger (close to type 2). The cell envelope is rather thick. Toward the apex and base, rows of cells converge in a bundle.

Dimensions: seed length 4.5–6 mm, seed width 3.5–4.5 mm.

Comparisons. Morphologically the described species can easily be confused with some species of the genus Cordaiсarpus, and in the absence of the sarcotesta with species of the genus Carpolithes, but Cordaiсarpus fascis is determined without difficulty microscopically, especially by the bundles, characteristic of this species, formed by rows of narrow cells at the base and at the apex of the seed.

Remarks. Not all specimens of this species have a sarcotesta; evidently it was easily lost in the process of burial (see above, the description of the preceding species). The species is distinguished by the anatomical structure of the epidermis.

Distribution. Erunakovskaya Formation, lower unit.

Localities. Erunakovsky district, seams 37, 49, 55.

References Cited

1. Artamontseva, M. D. Cordaites and seeds of the Erunakovo Formation. Problems of the Geology of Kuzbass, vol. 1, Moscow, 1956.

2. Gorelova, S. G. Flora and stratigraphy of the Erunakovo Formation of Kuzbass. Author’s abstract. Tomsk, 1953.

3. Neiburg, M. F. Upper Paleozoic flora of Kuzbass. Paleontology of the USSR, vol. XII, issue 2, 1948.

4. Radchenko, G. P. Guiding forms of the Upper Paleozoic flora of the Sayan-Altai region. Atlas of Guiding Forms of the Fossil Fauna and Flora of Western Siberia, vol. II, Moscow, 1955.

Tomsk Polytechnic Institute
named after S. M. Kirov

EXPLANATIONS OF THE PLATES

Plate I

Fig. 1. Angarocarpus tychtensis (Zal.) Radcz. Left bank of the Tychtа River near the village of Sokolovaya. Holotype (Zalessky 1937, p. 135, fig. 17).

Fig. 2. Angarocarpus tychtensis (Zal.) Radcz. Specimen (1266), Erunakovo district, bed 65.

Fig. 3. Angarocarpus dubius sp. n., specimen no. 85—213. Holotype. Nikitino locality, lower part of the Erunakovo Formation.

Fig. 8. Angarocarpus dubius sp. n. Specimen no. 184—1060. Nikitino locality, lower part of the Erunakovo Formation.

Figs. 4, 9, 14. Angarocarpus trigonalis sp. n. Specimen no. 24—82, holotype. Raspadskaya locality. Erunakovo Formation, bed 9. Fig. 14—anatomical structure of the epidermis.

Figs. 7, 12, 13, 15. Angarocarpus variabilis sp. n. Specimen 12/30—50. Holotype. Erunakovo district, bed 42. Fig. 15—structure of the epidermis in the central part of the seed. Fig. 7—structure of the epidermis in the apical part of the seed.

Figs. 10, 11, 16, 17. Angarocarpus variabilis sp. n. Specimen 12/404. Erunakovo district, bed 44. Fig. 17—structure of the epidermis in the central part of the seed. Fig. 16—structure of the epidermis near the lateral margins of the seed.

Figs. 5, 6. Angarocarpus proprius sp. n., specimen no. 178—1009. Holotype. Chumysh district, Balakhonka Series.

Plate II

Figs. 1, 6, 9, 11. Angarocarpus princeps sp. n. Specimen 11—55 (1). Holotype. Raspadskaya locality. Erunakovo Formation, below bed 13. Fig. 1—structure of the epidermis along the margin of the lateral part of the seed. Fig. 6—structure of the epidermis in the micropylar part of the seed. Fig. 11—structure of the epidermis in the central and basal part of the seed.

Figs. 7, 8, 10, 13. Angarocarpus sagittarius sp. n. Specimen 21/272. Holotype. Plotnikovsky district. Erunakovo Formation, bed XXIII. Fig. 7—structure of the epidermis in the central part of the seed. Fig. 8—cells on the surface of the nucellus near its apex and base. Fig. 13—cells on the surface of the nucellus at the very apex and base.

Figs. 2, 3. Angarocarpus enodatus sp. n. Specimen no. 22/256. Holotype. Zmeinsky district. Balakhonka Series, Tomsky II bed. Fig. 2—anatomical structure of the epidermis.

Fig. 12. Samaropsis angularis sp. n. Specimen no. 206—1136. Holotype. Leninsky district. Erunakovo Formation, above the Boldyrevsky bed.

Figs. 4, 5. Samaropsis stila sp. n. Specimen no. 22/220. Holotype. Zmeinsky district. Balakhonka Series, Pervomaisky bed.

Fig. 14. Samaropsis ampla sp. n. Specimen no. 22/140. Holotype. Zmeinsky district. Balakhonka Series, Opolznevyi bed.

Plate III

Figs. 6, 7, 11, 14, 15. Cardiocarpus corculum sp. n. Specimen no. 12/126. Holotype. Erunakovo district, above bed 28. Fig. 7—structure of the epidermis of the central part. Fig. 11—structure of the epidermis at the apex of the seed. Fig. 6—structure of the epidermis near the lateral sides of the seed and along the margin of the cordate notch at the base.

Fig. 12. Cardiocarpus strictus sp. n. Specimen no. 22/223. Holotype. Zmeinsky district. Balakhonka Series, Zmeinsky II bed.

Figs. 9, 10, 13, 16. Samaropsis acutula sp. n. Specimen no. 1164. Holotype. Kiselevsk district, field area of mine no. 6. Balakhonka Series. Figs. 9—10—anatomical structure of the epidermis of the wing. Fig. 13—anatomical structure of the surface within the nucellus.

Fig. 4. Carpolithes simplex sp. n. Specimen No. 12/16. Holotype. Erunakovskii district, bed 35. Erunakovskaya suite.

Fig. 3. Carpolithes ineditus sp. n. Specimen No. 12/32. Holotype. Erunakovskii district, bed 31. Erunakovskaya suite.

Figs. 2, 5, 8. Carpolithes levis sp. n. Specimen No. 12/28—50. Holotype. Erunakovskii district, bed 38. Figs. 2, 8—anatomical structure of the epidermis.

Fig. 1. Corpolithes pulchella sp. n. Specimen No. 111—918. Holotype. Erunakovskii district, bed 38. Erunakovskaya suite. Structure of the epidermis at the apex and base of the seed.

Table IV

Figs. 3, 7. Carpolithes pulchella sp. n. Specimen No. 111—918. Holotype. Erunakovskii district, bed 38. Erunakovskaya suite. Fig. 3—structure of the epidermis in the central and lateral parts of the seed.

Figs. 2, 6, 9. Carpolithes lucidus sp. n. Specimen No. 29—59. Holotype. Raspadskoe deposit. Erunakovskaya suite, bed 3. Figs. 2, 9—anatomical structure of the epidermis, with Fig. 9 showing the structure of the epidermis at the edge of the seed.

Figs. 1, 5. Carpolithes virgatus sp. n. Specimen No. 182—1014. Holotype. Nikitinskoe deposit, lower part of the Erunakovskaya suite. Fig. 1—anatomical structure of the epidermis.

Figs. 4, 8, 10. Carpolithes ineptus sp. n. Specimen No. 21/299. Holotype. Plotnikovskii district. Erunakovskaya suite, below bed XVIII. Fig. 4—structure of the epidermis near the lateral sides of the seed. Fig. 10—structure of the epidermis in the central part of the seed.

Table V

Figs. 1, 4, 7, 9, 11, 12. Carpolithes procerulus sp. n. Specimen No. 95—921. Holotype. Erunakovskii district, near the village of Markina. Fig. 4—structure of the epidermis near the lateral sides of the seed. Fig. 7—structure of the epidermis at the base of the seed. Figs. 1, 9—structure of the epidermis in the apical part of the seed. Fig. 7—structure of the epidermis at the base of the seed.

Fig. 3. Carpolithes rostriformis (Neub.) specimen 1153a. Shushtulepskii district, Anan’inskii area. Balakhonskaya series, Apanasovskii bed.

Figs. 2, 8. Carpolithes agitabilis sp. n. Holotype, specimen No. 1153. Shushtulepskii district, Anan’inskii area. Balakhonskaya series, Apanasovskii bed.

Figs. 6, 5, 10, 14, 15. Carpolithes novicus sp. n. specimen 22/268. Holotype. Zmeinskii district. Balakhonskaya series, roof of the Korshinnoye bed. Figs. 5, 10, 13, 14—anatomical structure of the epidermis: figs. 5, 13—at the apex and base of the seed; fig. 10—at the edge of the seed; fig. 14 in the central part of the seed.

Fig. 13. Cordaicarpus amictus sp. n. specimen No. 20/130. Holotype. Erunakovskii district, bed 65.

Table VI

Figs. 6, 7, 8. Cordaicarpus notulus sp. n. Specimen No. 204—1128. Holotype. Nikitinskoe deposit, lower part of the Erunakovskaya suite. Figs. 7, 8—anatomical structure of the epidermis.

Figs. 1, 3, 5. Cordaicarpus notulus sp. n. Specimen No. 18/8. Kasminskii district. Erunakovskaya suite. Anatomical structure of the epidermis.

Figs. 2, 4, 9. Cordaicarpus fascis sp. n. Specimen No. 12/49. Erunakovskii district, bed 37. Figs. 2, 4—anatomical structure of the epidermis, with fig. 2 showing the structure of the epidermis at the apex and base of the seed.

Plate I

1 — ×180
2 — ×2
3 — ×2
4 — ×2
5
6
7
8 — ×2
9
10 — ×2
11
12
13
14 — ×100
15 — ×100
16 — ×100
17 — ×100

Plate II

1 ×180
2 ×180
3
4
5 ×2
6 ×180
7 ×180
8 ×180
9 ×2
10 ×2
11 ×180
12 ×2
13 ×180
14

Plate III

1 ×180
2 ×180
3
4
5
6 ×180
7 ×180
8 ×180
9 ×180
10 ×180
11 ×180
12
13 ×180
14 ×2
15
16 ×2

Plate IV

Table V

1 — ×180
2
3 — ×2
4 — ×180
5 — ×180
6
7 — ×180
8 — ×2
9 — ×180
10 — ×180
11 — ×180
12 — ×2
13
14 — ×180
15 — ×180

Plate VI

1 — ×180
2 — ×180
3 — ×180
4 — ×180
5 — ×180
6 — ×1
7 — ×180
8 — ×180
9 — ×2