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14BIOLOGICAL SCIENCES
COMPARATIVE ANALYSIS OF THE DIVERSITY OF COLLEMBOLA AND ORIBATIDA GROUPS
IN AGROCENOSES OF SMALL POLISSIA
Merza S.
Postgraduate student, Lviv National Agrarian University
Kaprus I.
Prof., Dr. Sci.
Prof. of the Ecology Department, 1Lviv National Agrarian University,
2Ivan Franko National University of Lviv Sen. Res. of the Biosystematic and Evolution Department, 3State Natural History Museum, National Academy of Sciences of Ukraine
Abstract
The taxonomic structure of Collembola and Oribatida groups in the five most common types of Small Polissia agrocenoses, in particular, wheat, rape, soy, potato, and corn, has been analyzed. It has been revealed that the studied agrogroups of Collembola are, in general, characterized by sufficiently large species richness (at least 49 species from 35 genera and 12 families), as well as significant variability in density parameters. A total of 14 Oribatida species, belonging to 10 families, were identified. The studied cenotic faunas include from 2 to 6 species of oribatid mites and from 6 to 22 species of Collembola. In one soil sample, there are from 1 to 4 species of Oribatida and from 1 to 9 species of Collembola. In the studied agrocenoses by species richness among Oribatida, there are predominated the families Oppiidae, Mycobatidae, and Oribatulidae, each of which is represented by a total of 2-3 species. Among Collembola by species richness, there are predominated the families Isotomidae and Entomobryidae, each of which is represented by a total of 12 species. It has been revealed that in different types of Small Polissia agrocenoses, 10 species of Oribatida and 31 species of Collembola can potentially dominate. Due to the influence of agricultural land use for arable land, there are found the expansion of the mass forms of both Collembola and Oribatida in agricultural groups, compared with natural coenoses, the emergence of arable-specific dominants, as well as the emergence of superdominant species. The revealed features in the structure of the Oribatida group dominance, in general, are not characteristic of natural, little disturbed cenoses.
Keywords: biodiversity, Collembola, Oribatida, soil fauna, agrocenoses.
Soil microarthropods such as Collembola and Oribatida, due to their large numbers and biomass, are actively involved in the processes of transformation of matter and energy in soils and play an important role in soil functioning. Studying the structure of the diversity in groups of these pedobionts in different edaphic conditions, we can assess their biotic potential and functional role in ecosystems [36].
Agriculture is one of the important anthropogenic factors in reducing soil biodiversity. It significantly reduces the intensity of organic compound biodegradation, which determines the rate of soil fertility restoration in agrocenoses [1]. The study of the taxonomic and ecological structure of microarthropod groups in agro-cenoses, compared to the natural ecosystems, is an important step in the process of the arable fertility reproduction.
The parameters of Collembola diversity of Ukrainian arable lands have practically not been studied by specialists [11, 12]. Only a few works are known, which are devoted to the Collembola groups of man-made and urbanized landscapes of Ukraine [2, 14, 28, 31, 35]. It is revealed that in technogenic conditions, there is formed a taxonomically depleted fauna of Col-lembola with a low total number of biotope groups and a high level of the individual species dominance, as well as the representation of surface biomorphs.
Ecological and faunal studies of acarocomplexes in agrocenoses of Ukraine are also fragmentary and insufficient. In particular, the study of orchard oribatid
mites was conducted by S.G. Pohrebniak [26]. In total, he identified 14 species of Oribatida for the studied area. Some aspects on the impact of agricultural activities on the group of oribatid mites are also reflected in the works of T.F. Krutogolova and O. K. Furman [19]. The mentioned works concerned the features of the Ori-batida population in the conditions of seed treatment with microbial preparations and under the influence of fertilizers.
For the territory of Small Polissia, according to V.V. Melamud [23], only 136 species of oribatid mites are known. From 27 to 52 species of Oribatida have been found in some forest and meadow cenotic faunas [24]. Regarding Collembola, for Small Polissia, according to data of I.Ya. Kaprus [11], only 76 species of Collembola are known, while for the whole zone of deciduous forests of Ukraine - 303 species [11]. It is revealed that some forest and meadow cenotic fauna may include from 20 to 53 species of Collembola.
Although in the literature there are some data on the parameters of the group diversity of these microar-thropods in natural types of Small Polissia phytoceno-ses, the study of the structure of their groups' diversity in the agrocenoses of the region, as well as the main directions of population transformation of these pedo-bionts under the influence of various forms of arable land economic use remains relevant.
The purpose of the study was to describe the tax-onomic structure of Collembola and Oribatida groups in the main types of Small Polissia agrocenoses, as well
as to conduct a comparative analysis of the studied groups both among themselves and with their natural variants based on literature data.
Materials and methods. The study was conducted in Dubliany vicinity, Lviv region, on the experimental fields of LNAU (Lviv National Agrarian University) during the growing season (spring, summer, autumn) in 2017 in five main types of agrocenoses: 1) corn, 2) wheat, 3) rapeseed, 4) soy, and 5) potato. A total of ten agrocenoses were studied, two of each type. Each studied biotope is assigned a corresponding number: I, II - rapeseed; III, IV - soy; V, VI - wheat; VII, VIII - corn; IX, X - potato. It should also be noted that the model fields are characterized by evenly leveled terrain and soils, typical for Small Polissia.
The material was collected and processed according to the standard methods of soil and zoological studies [4, 27]. To collect the material, we used a metal drill with a volume of 577 cm3 (radius 3.5 cm, depth 15 cm). Soil sampling was performed by linear series every 10 m. In total, four similar sampling series were conducted: series 1 in June 2017; series 2 - in September 2017; series 3 - in November 2017, and series 4 - in April 2018. In each agrocenosis during the study period, 40 soil samples or 80 samples were taken for each of the five types of agrocenoses. A total of 400 soil samples were collected during the study period, and 296 collected Oribatida and 604 Collembola were identified.
Isolation of microarthropods from the substrate took place on Kempson thermophotoelectors. The collected zoological material was transferred into permanent micropreparations with Faure's medium for their further determination [27]. The species identification was performed using an Olympus BX52 microscope and using modern determinants.
The quantitative data obtained by us were extrapolated per unit area of 1 m2. For the comparative analysis of the population structure of the microarthropods in studied agrocenoses, we used not absolute, but relative (in% of the total number in the group) parameters of species density.
The standardized methods of quantitative analysis were used to assess the synecological structure of the
groups [22]. In particular, the structure of group dominance was determined by the approach of G. Stecker and A. Bergman [30]. The ecological affiliation of ori-batid mites was determined using data presented by G. Weigmann and L. Miko [32].
Statistical processing of the material was carried out using the program Past [6]. We used the program MS Excel as the main tool for the initial processing and display of data. Thus, the methodological approaches, used in the study, provided the necessary reliability and comparability of the obtained data.
Study results and discussion According to the study materials, a total of 49 species of Collembola and 14 species of Oribatida were identified. The identified Collembola belongs to 35 genera and 12 families (Table 1), which is on average 64.5% of local and 16.2% - zonal deciduous forest fauna [11, 13]. Oribatida belongs to 10 families (Table 2), which is 10% of the Small Polissia fauna [23]. During long-term, large-scale studies in agrocenoses, many more species can be identified (predicted to be at least 50 for Oribatida and 70 for Collembola). The studied cenotic faunas include from 2 to 6 species of oribatid mites. From 1 to 4 species of Oribatida occur in one soil sample. In the same agrocenoses, there are actually 3 times more species of Collembola, namely from 6 to 22 species. In one soil sample, there are from 1 to 9 types of Collembola.
In the studied agrocenoses, the average population density of Collembola and Oribatida varies in the twelve-fold (table 1) and thirty-fold (table 2) range of values, respectively. For Collembola, the density reaches the highest average level in wheat cenosis and the lowest - soy, and for Oribatida, the highest average level is in wheat and soy cenoses and the lowest - rape-seed. However, compared to the natural forest cenoses from the zone of deciduous forests [13], the maximum density of the studied agrocenoses Collembola is approximately 17-21 times lower, and to the meadows ce-noses, respectively, - 6-12 times. Regarding Oribatida, compared to the natural forest cenoses of deciduous forests [23], the maximum density of oribatid mites from the studied agrocenoses is approximately 218 times lower, and with meadow cenoses [10], respectively, 42 times.
Table 1.
Parameters of diversity of Collembola groups in studied agrocenoses of Small Polissia
Characteristics Agrocenoses
I II I IV I VI I VIII I X
1 2 3 4 5 6 7 8 9 10 11
HYPOGASTRURIDAE
Ceratophysellasuccinea (Gisin, 1949) 2,8 16,7 27,7 11,4
Hypogastruramanubrialis (Tullberg, 1869) 11,1 39,1 4,5 44,5 3,8 12,8 9,1
Willemiaintermedia (Mills, 1934) 1,6
BRACHYSTOMELLIDAE
Brachystomellaparvula (Schäffer, 1896) 2,8
TULLBERGIIDAE
Metaphoruraaffinis (Börner, 1902) 3,1 2,6 6,7
Mesaphoruracritica (Ellis, 1976) 1
Mesaphorura florae (Simonat al., 1994) 11,1
Mesaphoruramacrochaeta (Rusek, 1976) 13,9 14,1 4,5 1,6 20,5 33,3 11,1 4,5
Stenaphorura quadrispina 1,6 3,3
(Börner, 1901)
ONYCHIURIDAE
Onychiurus ambulans (Linnaeus, 1758) 2,6
Protaphorura cancellata (Gisin, 1956) 0,5
Protaphorura fimata (Gisin, 1952) 9,6
Protaphorura pannonica (Haybach, 1960) 1
Protaphorura subarmata (Gisin, 1957) 1,6 9,1 0,5 20 11,1 18,2
Agrophorura naglitshi (Gisin, 1960) 1,9
ISOTOMIDAE
Desoria fennica (Reuter, 1895) 3,3 4,5
Cryptopygus thermophilus (Axelson, 1900) 1,9
Proisotoma minuta (Tullberg, 1871) 1 5,8 5,1 9,1
Folsomia fimetaria (Linnaeus, 1758) 13,9 1,6 18,2
Folsomia lawrencei (Rusek, 1984) 0,5 2,3
Folsomia manolachei (Bagnal, 1939) 2,3
Folsomia spinosa (Kseneman, 1936) 5,6
Folsomides parvulus (Stach, 1922) 2,6 2,3
Isotoma anglicana (Lubbock, 1873) 2,8 4,5 0,5 5,8 2,6
Isotomiella minor (Schäffer, 1895) 16,7
Isotomodes productus (Axelson, 1906) 2,8 8,3 3,3
Parisotoma notabilis (Schäffer, 1896) 13,9 23,4 4,5 8,3 3,1 1,9 3,3 22,2 2,3
ENTOMOBRYIDAE
Entomobrya marginata (Tullberg, 1871) 7,7 7,7
Entomobrya sp 4,5
Sinella tenebricosa (Folsom, 1902) 13,6 3,8
Heteromurus nitidus (Templeton, 1835) 2,8 16,7 0,5 5,1 2,3
Orchesellaalbofasciata (Stach, 1960) 2,8 6,3 9,1 1 5,1
Orchesella multifasciata (Scherbakow, 1898) 15,4
Orchesella pseudobifasciata (Stach, 1960) 1,6
Lepidocyrtus cyaneus (Tullberg, 1871) 16,7 9,1 4,2 38,5 7,7 11,1 4,5
Lepidocyrtus paradoxus (Usel, 1890) 1,6
Pseudosinella alba (Packard, 1873) 4,7 13,6 2,6 3,8 5,1 16,7 33,3
Pseudosinella imparipunctata (Gisin, 1953) 2,8
Willowsia platani (Nicolet, 1842) 7,7
TOMOCERIDAE
Tomocerus vulgaris (Tullberg, 1871) 9,1
PARONELLIDAE
Cyphoderus albinus (Nicolet, 1842) 1,9
BOURLETIELLIDAE
Bourletiella arvalis (Fitch, 1863) 2,8 9,1 8,3 3,8 3,3
Bourletiella hortensis (Fitch, 1863) 2,1 2,6 6,7 6,8
Caprainea marginata (Schött, 1893) 2,8 0,5
Sminthurus maculatus (Tömösvary, 1883) 25
KATIANNIDAE
Sminthurinus aureus (Lubbock, 1862) 1,6 1 2,3
Sminthurinus elegans (Fitch, 1863) 0,5 5,1
ARRHOPALITIDAE
Arrhopalites caecus (Tullberg, 1871) 4,5
SMINTHURIDIDAE
Sphaeridia pumilis (Krausbauer, 1898) 1 1,9
Total species number 15 12 13 7 22 15 14 10 6 15
The fraction of the dominant species number (%) 75,1 87,6 100 100 76,4 86,5 89,6 100 100 86,2
Density, thousands of individuals/m2 0,23-0,42 0,08-0,14 0,34-1,24 0,19-0,25 0,06-0,29
Note. Agrocenoses: I,II - rapeseed; III,IV - soy;V,VI - wheat; VII,VIH - corn; IX,X - potato. The values of the relative number of Collembola dominant species are highlighted in gray color.
Considering Collembola in the studied agrocenoses, by species richness, the families Isotomidae and Entomobryidae predominated, each of which is represented by a total of 12 species (in some cenoses, there were from 1 to 7 species of Isotomidae, and Entomo-briidae - 1-6) (Table 1). By parameter of the relative number of Collembola families in most agrocenoses, Entomobryidae (6,8-49,9% of the total number of individuals, on average 30%), Isotomidae (5,1-41,0%, 21%), and Hypogastruridae (0-72,2%, 18,5%) dominate (Table 1, Image 1). The obtained data on the representation of families in cenotic fauna are generally consistent with the literature data for natural variants of cenoses in areas of deciduous and mixed forests of Ukraine [13, 21].
Studying Oribatida in agrocenoses by species richness, there is a domination of families Oppiidae, Myco-batidae, and Oribatulidae, each of which is represented
Analyzing the data on Oribatida, 10 species of Ori-batida can be dominant, the total fraction of which is 92.8-100% of the cenotic groups' number (Table 2). In some biotopes, they can be from 2 to 5 species. Among oribatid mites, there are dominated members of the families Tectocepheidae and Scheloribatidae, whose representatives occur in nine and six variants of ten agrocenoses (Table 2).
Among the dominant (otherwise mass) species of Collembola, there are found four eudominants (Н. ma-nubrialis, M. macrochaeta, L. cyaneus, P. alba), the relative number of each can reach up to 44,5 % of the total. Only in soy cenosis, they were not detected. In
relative number of Oribatida families in most agroce-noses, Tectocepheidae (11,0-80,0% of the total number of individuals, on average 46%) and Scheloribatidae (8,3-56,8%, 32%) dominate (Table 2, Image 2). The obtained data on the representation of families in cenotic fauna are generally consistent with the literature data for natural variants of cenoses in areas of deciduous and mixed forests of Ukraine [24].
It is revealed that in different types of studied ag-rocenoses of Small Polissia, 31 species of Collembola, the total fraction of which 75,1-100% of the cenotic groups' number can potentially are dominant (i.e, be eudominants, dominants or subdominants with a relative number greater than 3.2% of the total in the group) (Table 1). In some biotopes, they can be from 3 to 13 species. The most dominated members of the families Entomobryidae (9 forms), including two species of the genera Entomobrya and Orchesella, and Isotomidae (8)
addition to eudominants, 0-5 dominant and 0-11 subdominant species were identified in each agrocenosis. No species has been identified that would dominate in all studied agrocenoses simultaneously. Only seven of the ten studied cenoses were dominant Н. manubrialis, M. macrochaeta, and L. cyaneus, six - P. notabilis and P. alba. The rest of the mass forms, obviously, have certain ecological limitations and, therefore, dominated only in one - four biotopes. In particular, only in one of the agrocenoses, there were dominated 13 species, i.e. 42% of their total number, which was found in the studied variants of arable land.
by a total of 2-3 species (Table 2). By parameters of the - two species of the genus Folsomia (Table 1).
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О
X IX VIII VII VI V
IV
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dc
и III
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RELATIVE ABUNDANCE
80,0
100,0
□ hypogastruridae h isotomidae в entomobryidae 0 bourletiellidae qtullbergiidae □ onychiuridae □ katiannidae dtomoceridae
□ arrhopalitidae □ brachystomellidae к sminthurididae « paronellidae
Image 1. Correlation of Collembola families by number in the studied agrocenoses. Marking of agrocenoses I-X as in Table 1.
Table 2.
Parameters of diversity of Oribatida groups in studied agrocenoses
Characteristics Agrocenoses
I II I IV I VI I VIII I X
COSMOCHTHONIIDAE
Cosmochthonius reticulatus Grandjean, 1947 25,0
HYPOCHTHONIIDAE
Hypochthonius luteus Oudemans, 1917 2,4
EUPHTHIRACARIDAE
Acrotritia ardua (Koch, 1841) 13,3
TECTOCEPHEIDAE
Tectocepheus velatus (Michael, 1880) 60,0 75,0 80,0 11,0 56,1 43,9 21,6 25,0 46,7
OPPIIDAE
Oppia nitens Koch, 1835 6,7
Oppiella nova (Oudemans, 1902) 20,0 19,5 13,5 50,0 20,0
Ramusella cf. clavipectinata (Michael, 1885) 13,3
SUCTOBELBIDAE
Suctobelbella sp. 2,4
SCUTOVERTICIDAE
Scutovertex minutus (Koch, 1835) 2,4
MYCOBATIDAE
Punctoribates hexagonus Berlese, 1908 32,3 14,6 8,1 16,7 50
Punctoribatespunctum (Koch, 1839) 4,7 9,8
SCHELORIBATIDAE
Scheloribates pallidulus (Koch, 1841) 20,0 20,0 52,0 17,1 56,8 8,3 50,0
ORIBATULIDAE
Oribatula cf. exilis (Nicolet, 1855) 2,4
Oribatula cf. glabra (Michael, 1890) 29,3
Total species 3 2 2 4 6 5 4 4 5 2
The fraction of the dominant species number (%) 100 100 100 100 92,8 97,6 100 100 100 100
Density, thousands of individuals/m2 0,0130,016 0,0320,404 0,1300,131 0,0380,118 0,0130,048
Note. Agrocenoses: I, II - rapeseed; III, IV - soy; V, VI - wheat; VII, VIII - corn; IX, X - potato. The values of the relative number of Oribatida dominant species are highlighted in gray color.
Among the mass species of Oribatida, we found four eudominants (Tectocepheus velatus, Oppiella nova, Punctoribates hexagonus, Scheloribates pallidu-lus), the relative number of each can reach up to 80% of the total. In addition to eudominants, 0-3 dominant and 0-1 subdominant species were identified in each agrocenosis. No species has been identified that would dominate in all studied agrocenoses simultaneously.
Only nine and seven of the ten studied cenoses were dominant, respectively, Tectocepheus velatus and Scheloribates pallidulus. The rest of the mass forms, obviously, have certain ecological limitations and, therefore, dominated only in one - four biotopes. In particular, only in one of the agrocenoses, there were dominated 5 species, i.e. 36% of their total number, which was found in the studied variants of arable land.
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Relative Abundance
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:: Cosmochthoniidae
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Suctobelbidae ■ Scutoverticidae ^ Mycobatidae
ii Scheloribatidae c Oribatulidae
Image 2. Correlation of Oribatida families by number in the studied agrocenoses. Marking of agrocenoses I-X as in Table 2.
Thus, the agricultural use of land for arable land has a significant influence on the expansion of the mass forms of both Collembola and Oribatida in the composition of agricultural groups, compared with natural groups, due to the emergence of arable-specific dominants. (Collembola: I. productus, S. tenebricosa, H. ni-tidus,W. platanira, etc. Oribatida: C. reticulatus, A. ardua, O. nitens, R. cf. clavipectinata, etc.), as well as the emergence of eudominance (superdominance) of certain species. These features of the structure of the Collembola and Oribatida group dominance, in general, are not characteristic of natural, slightly disturbed cenoses [7, 8, 28]. The obtained data on the presence of a wide range of potential dominants, as well as the unpredictability and variability in the composition of Collembola mass species in agrocenoses, are consistent with the data obtained by specialists in urban biotopes of Uzhgorod and Lviv [5, 28].
Conclusions
Thus, the Collembola group of agrocenoses in the studied region, in general, is characterized by sufficiently high species richness (at least 49 species from 35 genera and 12 families), as well as significant variability in density (0,06-1,24 thousands of ind./m2). Species richness of Oribatida in the studied agrocenoses is characterized by a lower level and is 10% of the fauna in Small Polissia. A total of 14 Oribatida species, belonging to 10 genera, were identified. There is revealed the high variability on the density parameter of the Ori-batida group (variation in the thirty-fold range of values), as well as a decrease in the density parameter of approximately 218 and 42 times, compared to the zone of deciduous forests and meadow cenoses, respectively.
Due to the influence of agricultural land use for arable land, the divergent and often unpredictable changes in the synecological structure of Collembola cenotic groups have been recorded. In particular, there has been revealed the expansion of the Collembola mass forms in agricultural groups, compared to the natural cenoses, due to the emergence of arable-specific
dominants (I. productus, S. tenebricosa, H. nitidus, W. plataniTa, etc.), as well as the emergence of superdominant species. Similarly, for Oribatida we found four eudominants (Tectocepheus velatus, Oppiella nova, Punctoribates hexagonus, Scheloribates pallidulus), the relative number of each of them can reach up to 80% of the total.
It was revealed that by species richness in the studied agrocenoses, the families Oppiidae, Mycobatidae, and Oribatulidae were predominated. Considering Col-lembola in the studied agrocenoses by species richness, the families Isotomidae and Entomobryidae, each of which is represented by a total of 12 species, were predominated.
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