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Section 4. Geology
DOI: http://dx.doi.org/10.20534/ESR-17-1.2-29-32
Evgeny Eugenievich Sukhov, Doctor of Geology and Mineralogy Kazan Federal University,
Kazan City E-mail: evgeny.suchov@yandex.ru
New data on the phylogenetic development of the Permian Ichtyolaria
Abstract: The article deals with problems of the genus Ichtyolaria, which is one of the most important and numerous in the Permian system. This genus has a significant stratigraphic significance. The predominant number of species is presented by characteristic or principal species. The phylogeny of the genus Ichtyolaria is studied for the first time on the basis of representative material collected by the author. The study reveals the complex mechanism of speciation, which is influenced by a significant number of factors: paleo-ecological crisis, induced mutations, changes in the genotype, a significant exchange of genes. Much attention is paid to the habitat.
Keywords: Phylogenesis, foraminifera, Permian system, age, branch, tree, population, shell, stratotype, complex, species, genus, the Ufimian, century.
Introduction
Synonymies of the genus Ichtyolaria [1] is controversial for more than eighty years. At different times under this generic name micropaleontologists described representatives of the following genera: Spandelina [2, 3-9]; Frondiculinita [10]; Frondicularia [11] and etc. The main reason for unclear systematic position is that the author of the species R. Wedekind [12] gave practically no description and did not substantiate the detection of the genus.
The genus Frondicularia was redescribed as Ichtyolaria by Sivrye and Desovazhi in 1965. But the diagnosis almost duplicated the description of the genera Spandelina and Frondicularia. Later, the validity of the genus Ichtyolaria was presented in the work of A. V. Fursenko [13]. However, up to the present time the question concerning the essence of the genera Spandelina, Frondicularia and Ichtyolaria remains open.
Among micropaleontologists, who deal with the study of late Paleozoic foraminifera, there are three points of view:
1. Some believe that the name of the genus Spandelina is identical to the genus Ichtyolaria [1] and by the right of priority, preference should be given to the genus Spandelina, because it was detected earlier.
2. A. A. Gerke classified in his numerous works all species, which were described by K. V. Miklukho-Maklay as the genus Spandelina, to the genus Frondicularia.
3. V. M. Igonin [14] thought that shells of spindle-like shape with a compressed shell, calcite wall and little embracing loculi should be attributed to the genus Ichtyolaria.
4. Foreign researchers think that the genus Frondicularia consists of two and maybe more genera. For example, the Australian specialist V. Palmieri [15] isolated new genus Howchinella from the genus Frondicularia, within the framework of which he described four species. Some old species, that were regarded by V. Khovchin, I. Krespin as the genus Frondicularia, were attributed by him to the representatives of the genus Howchinella. However, he also singles out the genus Ichtyolaria.
In our opinion, it is most precise to name the genus Frondicularia as Ichtyolaria due to the fact that it was described in detail by Sivrye and Desovazhi. One should not exclude the possibility that Ichtyolaria covers several genera, but such a claim requires more substantial arguments, for example, extensive use of electronic microscope for study of internal and external morphology of a shell: aperture, septa, pre-sutural thickening; the latest methods should be applied for ultrastructural study of the shell walls [16-19].
Results. Phylogeny of Permian Ichtyolaria
From calcareous Permian foraminifera, along with the genus Nodosaria [20, 21], Ichtyolaria should be regarded as the most polytypic genus, which numbers more than fifty species, and which originated apparently from the genus Perunia [22]. During Permian age the genus Ichtyolaria underwent the prosperity. Evolution of the genus Ichtyolaria can be seen on the example of phylogenetic tree, where there are all three classes: leaves, nodes and root. Ichtyolaria prima (Gerke) should be referred to as the most ancient species, standing at the base of the root, which appeared in the early Saranin-sky age, had considerable self-regulation, created a sustainable bio-coenosis and had a life area within all basin of Bjarmaland [23; 24]. Shell is small, leaf-shaped, with thin septa, one- and two-layer shell. With clear characteristics of sustainability of generic features.
Until the middle of Saraninsky age, there was a stable biogeocoe-nosis (climactic), after which evolutionary development took place: the rooted trunk was divided into two large branches A and B, which later continued to evolve on their own until the Ufimian age. To the first branch A we should attribute forms, which inherited from their ancestor a small shell, not sunken thin septa, some extension in the middle part, often a developed aperture. To the second branch B we should attribute larger species, which had from the ancestor the callosity in the second half ofa shell; low septa; extension of the ultimate loculus with a significant deepening of seams. Both branches evolved in the direction ofincrease ofshells, extending the number ofloculi, complication of apertural apparatus, thickening of walls, as well as a more complex joining of septa with the inner surface of shells [23; 25; 26].
At the baseline of the branch A there is an ancestral form Ich-tyolaria amygdaleformis (Gerke, (principal species of middle Sara-ninsky age), which has small loculi and a quite elongate aperture. It constituted a large population, which was spread in normal salty environments and existed until the Ufimian age because of universality of genomes and their ability to adapt to changing conditions. The species is especially widespread in Pechora province and in Taymyr-Kolyma sub-region. Conditions for existence of the species in Sarginsky age were very favorable. At the same time, within the population there were forms with significant changes of genotype (expansion of shell, considerable elongation of ultimate locu-lus), what led to the formation of phenotype. For example, at a later development stage the complication of aperture occurred in many specimens. This feature is dominant and manifested in the whole phylogenesis. Since the middle Saraninsky age there was a strong modification variability, which was adaptive to the changing living conditions, what further led to the formation of new species.
Icht. extera (Zolot.) appeared in the early Filippovsky age. It was similar to the ancestral species due to the structure of aperture and cassideous ultimate loculus. The species Icht. extera (Zolot.) is principal for Filippovsky age. A significant population is observed in the East-European water area (the species is rare in Taymyr-Kolyma sub-region). On the boundary of Saraninsky and Filippovsky ages there was a significant shallowing of the Eastern-European sea. There were intense induced mutations, associated directly with the change of ecosystem, what in turn led to the emergence ofnew species. There was another important evolutionary transformation: two species emerged from the node of Ichtyolaria extera (Zolot.): Icht. soloduch-owi Suchov, which was principal for Filippovsky age, and Icht. caseyi (Crespin), which was previously found in Notalnaya area. Allopatric speciation played an important role, which increased during Irensky and Solikamsky ages due to the greatest isolation of basins.
During early Irensky age the branch Icht. soloduchowi Suchov gave a rise to 2 species — Icht. mica (Gerke), which existed until Solikamsky age (the species inherited from the ancestor a similar shell, low structure of the aperture), and also Icht. reliqua (Gerke) (from the ancestral form it inherited ultimate loculus, not sunken seams, thin septa). The species Icht. soloduchowi Suchov is very numerous in the stratotype, — huge populations emerged in water areas, in which new species developed, which are very similar to the original species, what is typical for sympatric speciation. The species Icht. reliqua (Gerke) was formed as a result of epigenetic imprinting (change ofthe phenotype). This species also existed until the first half of Solikamsky age, when it along with many other species became extinct as a result of environmental changes.
The species Icht. mica (Gerke) was quite numerous, and during late Irensky age it created a huge population of related forms. As a result of shallowing of the basin, a persistent transformation of the genotype (mutation) occurred; new species dedifferentiated from the node during Solikamsky age, including: Icht. sectorialis (Voron.), Icht. propria (Voron.), Icht. pseudotriangularis (Gerke), Icht. abies (Schleif.). These types can be designated as leaves of the rooted tree.
During early Solikamsky age a whole set of new species differentiated from the node Ichtyolaria reliqua (Gerke), which considerably inherited from the ancestor the shape of a shell aperture, among which are Icht. bephana Suchov, Icht. vesta Suchov, Icht. dilemma (Gerke) (including different varieties), Icht. longissima (K. M.-Macl.), Icht. valeevae Igon., Icht. acutancula Igon., Icht. petschorica Suchov, Icht. bajcirica (Zolot.). The species are similar to each other and inherit from ancestral forms in a different degree a spear-shaped shell, high thin septa, ultimate loculus, apertures, shape of initial loculus, what serves as proof that there is a polyphyletic group at this level.
Some principal species of early Solikamsky age, such as Ichtyolaria valeevae Igon., Icht. acutancula Igon., existed until late Solikamsky age, in which foraminifera were not diverse and numerous. In fact, the complex of late Solikamsky age represents a depleted population of early Solikamsky age, what is associated with a significant reduction of sea area and with deterioration of living conditions. In the complex there are different species of some taxa, for example, Icht. dilemma (Gerke)). However, there are some new species that were able to adapt to changing conditions, including Icht. zavodovskyi (A. M.-Macl.) (typical for Kazanian age), which derived from Icht. valeevae Igon. and had a common shell shape and aperture structure.
During late Solikamsky age there was a decline of the taxonom-ic group, characterized by a decrease in the number of specimens, narrowing of habitat and reduction in species numbers. Among small populations there was a constant transfer of gene flow, which reduced genetic differences between them, and which also significantly affected the speciation. During late Solikamsky age a significant number of species became extinct, which were unable to pass through a paleoecological crisis, among which there were Ichtyolaria acutancula Igon., Icht. dilemma (Gerke), Icht. fallax (Gerke).
Branch B starts from the species Ichtyolaria inflata (Gerke), which emerged in middle Saraninsky age, which was very favorable for progressive development of taxonomic group (broad water areas, warm climate). The population belongs to another adaptive zone, which is characterized by large species, what serves as an evidence of arogenesis. Numerous foraminiferal complexes faced an enhanced diversification, and the late Saraninsky age was characterized by the following evolutionary transformations: two branches branched off the nodes of Ichtyolaria inflata (Gerke): the first branch B 1 with the species Ichtyolaria hemiinflata (Gerke), which existed until late Kun-gurian age, and the second branch B 2 with the species Ichtyolaria carinatostata (Gerke), which became extinct in the early Ufimian age.
The branch B 1: the species Ichtyolaria hemiinflata (Gerke) resembles the ancestral form due to the shell structure (it is more narrow in the first half of ontogenesis; very similar to some initial loculi). All this demonstrates evolutionary change in the initial stage of speciation (archallaxis), what just emphasizes the relationship with Ichtyolaria inflata (Gerke). The species also inherited from the ancestor a three-layer structure of the wall, as well as the manner of joining of septa, which are marked by a considerable thickening.
In the early Filippovsky age, the species Icht. bojana Suchov differentiated from Icht. hemiinflata (Gerke), which was principal for that geo-chronological level. The species differs from the ancestor by a higher ultimate loculus and also by a long apertural elevation with elongate apertural tube. Formation of this species was connected with sympatric speciation, when during Filippovsky age there were considerable changes of the paleoecological environment. In the early Filippovsky age the Artinskian complex (with a low apertural elevation) was replaced with another one — with a long aperture and high elevation. In the early Irensky age Icht. kruga Suchov, which existed until the beginning of the Ufimian age, branched off Icht. hemiinflata (Gerke). From the ancestral forms it inherited low loculi and low aperture with a small apertural elevation (the species is principal for Irensky age). Obviously, the reason for their extinction was a selective elimination — the species could not withstand competition with the most highly organized forms under the changed paleoecological conditions (shallowing and salinization of the basin, aridization of the climate).
The Ufimian complex of Ikhtiolaria, which has many specialized species (only some of them survived until the late Solikamsky age),
differs significantly from Kungurian complex. However, paleobio-coenosis of Ikhtiolaria of early Solikamsky age undoubtedly has a high taxonomic diversity, is located in adaptive area and has dozens of new species, what serves as an evidence of biological progress. Moreover, the morpho-physiological component is dominant (for many species, adaptation occurs within one regional paleo-basin). The most typical species are Icht. belboga Suchov, Icht. daniilowi Suchov, Icht. ra Suchov, Icht. trojana Suchov.
In early Kungurian age Icht. planilata (Gerke) differentiated from Icht. hemiinflata (Gerke), which existed until the end of early Solikamsky age and which had the thickened loculus as well as other representatives of this branch.
In the course of phylogenetic development there was an exchange of genes between populations of the branches A and B, which existed in parallel, what allows us to speak about a common gene pool of the genus. For example, in the beginning of the Ufimian age the species Icht. daniilowi Suchov emerged, which was much similar to Icht. reliqua (Gerke) (branch A) (because of ray structure of the aperture, low loculi), and the species Icht.planilata (Gerke) (because of a large number ofloculi, a broader structure of the shell). And in early Solikamsky age, the exchange of genes only increased due to reduction of water areas. Polyphyletism played a big role in Solikamsky age.
In early Solikamsky age the populations of Ikhtiolaria evolved at a lower pace than the external environment, what led to the biological regression. The water areas significantly reduced, the total number of species decreased, the complexes became taxonomi-cally impoverished. The complex of Ikhtiolaria seemed oppressed. Since lower Solikamsky age the population underwent mutations, which lead to morphological regression and then to the extinction of species. A significant number of species, which existed until the late Solikamsky age, became extinct, including Icht. planilata Gerke, Icht. Icht.plana (Gerke), Icht. miranda (Gerke).
The branch B 2, at the baseline ofwhich there is Icht. carinatosta-ta (Gerke) (beginning of Saraninsky age), underwent considerable morpho-physiological changes (aromorphoses), which significantly determine arogenesis: longitudinal undulate formations on the shell surface (sculpture) appeared in all group of Ikhtiolaria. The
sculpture survived throughout the whole historical and geological development of the branch. The external transformation is associated primarily with a change in the paleo-ecological environment.
At the beginning of Saraninsky age Icht. bella (Gerke) separated from Icht. carinatostata (Gerke) (the species survived until the end of Irensky age), whose shell resembled the ancestor's shell; besides, the structure of loculi was a dominant factor. At the beginning of Filippovsky age Icht. costiferella (Gerke) differentiated from the species Icht. carinatostata (Gerke) (it inherited from the ancestral species the shape of ultimate loculus, as well as the aperture), which existed until the beginning of the Solikamsky age. Both species have undulated relief, typical for the branch B 2. On the boundary of Irensky and Solikamsky ages Icht. petschorica Suchov differentiated from the branch Icht. carinatostata (Gerke), which had in common with the ancestor an ultimate loculus and ray aperture with elevation.
On the boundary of Kungurian and Ufimian ages there was active exchange of genes in two genetically close populations B1 and B 2, what led to the formation of modified population with common characteristics, resistant to abiotic factors. Active speciation occurred within the population. Typical representatives of mutational possibilities in the population are such species as Icht. Ibri-gita Suchov and Icht. subtilistriata (Gerke). For example, Icht. brigita Suchov with a large number of loculi, inherited from Icht. hemiinflata (Gerke) (branch B 1), and a pronounced undulated relief, inherited from Ichtyolaria carinatostata (Gerke) (branch B 2).
At the turn of early and late Solikamsky ages due to the shallowing ofwater areas and the deterioration ofpaleo-climate, and as a result of narrowing of populations' habitat, the complex of Ikhtiolaria experienced a general adaptive syndrome, in which most species could not adapt to the changed conditions [23; 24]. Then degeneration process occurred (shells reduced in size, acquired ugly features, their internal structure simplified), what further led to the biological regression. The latest species of ribbed Ikhtiolaria were discovered in the beginning of early Solikamsky age.
It should be noted that the species with relief on the shell surface, likely, represent a new species, according to morphological criteria and internal structure of loculi.
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DOI: http://dx.doi.org/10.20534/ESR-17-1.2-32-34
Evgeny Eugenievich Sukhov, Doctor of Geology and Mineralogy Kazan Federal University, Kazan City E-mail: evgeny.suchov@yandex.ru
New data on the phylogenetic development of the genus Nodosaria in the Pre-Ural period (Permian) in the Eastern-European sub-region
Abstract: Phylogenetic research of small foraminifera is an important aspect of their study. It allows to represent in a more comprehensive manner the picture of their development during geological time; to track changes of species and to understand the role of foraminifera in a geological history of the Earth; it allows to define the stability of morphological modifications, to identify genetic relationships of genera and other taxa, to find out the course of development of the studied group of fauna. Phylogeny has a great importance in solving the issues of detailed biostratigraphy. The knowledge of its laws allows us to define phylozones, i. e. the layers, where there are taxa, which characterize the segment of the specific phylogenetic line. The order Nodosariida is most interesting from the viewpoint of studying the systematization of foraminifera as the general course of phylogenesis. Its species are ancestral forms of many genera of the calcareous foraminifera, including such important forms for stratigraphy as Lingulonodosaria, Dentalina, Pseudonodisaria, Protonodosaria. It is assumed that the first Nodosariidae appeared as early as in the Ordovician period, however, they are known for certain only since the middle Devonian period. The family Nodosariidae developed by change of the morphology of a shell, its sculpture as well as by complication of the internal structure — appearance of multi-layered walls, thickened septal walls, complication of the apertural apparatus. The wall of Nodosariidae is secretory, calcareous, fine-pored, vitreous, radial fibrous, one-layered or complex multi-layered. Each of the listed properties is very important for systematization of the order.
Keywords: phylogeny, species, genus, genetic relationship, population, characteristics, Nodosariidae, shell, branch, aperture, wall, Foraminifera.
Results. The genus Nodosaria is most interesting for phylogenetic study, which was common during the Upper Paleozoic and Mesozoic eras, and which originated numerous genera [1]. Phylogeny of the genus Nodosaria can be traced in detail in the Permian Pechora sections, which are distinguished from other provinces of the Eastern-European sub-region through more representative Permian deposits. Significant formation of species Nodosaria took place in Permian period, in Artinskian age. The basin of Artinskian age in the Pechora Cisurals was represented by a relatively narrow meridional intracontinental sea (in fact — by lagoon) [2]. In gen-
eral, its fauna was presented predominantly by Nodosaria and Ichtyolaria. The most ancient species in Pechora province is Nodosaria shikanica Lip. (It is found in Carbonic period and passes to Artinskian age). In the early Irginsky age, the population of the species Nodosaria shikanica Lip., which had a vast habitat, underwent significant changes, connected with dwelling in different environments, in which the divergence of characteristics and the formation of new variations, subspecies and species took place: in some species — shells with high loculi developed, in other species — a slight curve in the middle part of a shell appeared. Approximately
