Open access peer-reviewed chapter

Beneath the Surface: Comparative Diversity, Abundance, and Bio-Indicative Capabilities of Nematode Community Structure in Natural and Disturbed Habitats

Written By

Mohammad Asif and P. Fazul Rahaman

Submitted: 30 September 2023 Reviewed: 02 October 2023 Published: 28 February 2024

DOI: 10.5772/intechopen.1003686

From the Edited Volume

Nematodes - Ecology, Adaptation and Parasitism

Soumalya Mukherjee and Sajal Ray

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Abstract

This present study underscores the significance of soil nematodes as indicators of ecological health, particularly in areas affected by human activities like mining. Soil nematodes, classified into five trophic groups, play vital roles in nutrient cycling and provide insights into soil food web dynamics. Various nematode community indices, including the maturity index (MI) and trophic diversity index (TDI), are crucial for assessing soil food web diversity. Nematodes exhibit adaptability across diverse soil environments, from pristine to highly disturbed habitats, making them responsive indicators of environmental changes. Understanding nematode community structure enhances their potential as global indicators for assessing food resource availability and habitat characterization. This study compares three different zones based on anthropogenic disturbances in the coal mining region of Sonebhadra, Uttar Pradesh, representing undisturbed, moderately disturbed, and intensely disturbed sites. By analyzing nematode communities and trophic group abundance, the study assesses soil ecosystems across these zones. Human activities, especially mining, significantly impact soil nematode diversity and ecosystem health. Transition from natural forests to mining sites leads to shifts in nematode communities and species diversity, with intermediate disturbance fostering increased species diversity. Maturity index values reflect ecosystem maturity, with undisturbed and moderately disturbed zones indicating structured ecosystems, while highly disturbed zones represent degraded conditions. Faunal profiles mirror these findings, indicating shifts in decomposition pathways. This study highlights the potential of nematodes as indicators for environmental monitoring and quality assessment in coal mine areas. Further research on individual nematode species can inform biodiversity modeling and contribute to more effective ecological restoration efforts.

Keywords

  • soil nematodes
  • trophic groups
  • mining activities
  • faunal profiles
  • decomposition pathways
  • biodiversity modeling
  • ecological restoration

1. Introduction

Nematodes have been categorized into five primary trophic groups [1, 2]. They play a significant role in nutrient mineralization through energy decomposition pathways [3, 4, 5, 6] and offer valuable insights into the structure and function of soil food webs [1, 7, 8, 9]. Nematode community indices, such as the maturity index (MI) based on c-p scaling, the plant parasitic index (PPI) (weighted mean of c-p values of plant-parasitic nematodes), species richness, evenness, the ratio of bacterivore to fungivore nematodes, and the trophic diversity index (TDI), are routinely employed to assess soil food web diversity conditions [10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20]. Furthermore, various faunal food web indices for nematodes, such as enrichment (EI), structure (SI), and decomposition channel index (CI), based on the relative weighted abundance of nematode c-p guilds, provide information on enrichment, trophic connections, structure, and prevailing decomposition in a food web [1].

Nematodes are valuable indicators as they inhabit soils ranging from pristine to highly disturbed habitats and are represented in almost all trophic groups of the soil food web. They respond quickly and specifically to environmental changes. A comprehensive understanding of nematode community structure for assessing food resource availability and characterizing different habitats would enhance the indicative capability of nematodes on a global scale [1, 21].

Although we have limited knowledge about the influence of vegetation and abiotic parameters on nematode fauna, drawing trends can be challenging due to geological, climatological, and methodological differences among various sites or studies [22]. Analyzing the soil nematode community can reveal differences between habitats [19, 21].

In the present study, three different zones with varying degrees of anthropogenic disturbances in the coal mine region of district Sonebhadra, Uttar Pradesh, were compared. Various parameters of the nematode community were used to assess the prevailing soil ecosystem in these zones. Depending on the nature and frequency of disturbances (physical, chemical, or biological), active mining land and wild forests (potential future mining sites) represent two extremes in terms of ecosystem structure and stability. The three zones identified based on the degree of human interference were: undisturbed hillocks or potential future mining sites (Zone ‘A’), moderately disturbed and managed habitat represented by abandoned reclaimed land after mining (Zone ‘B’), and intensively disturbed habitat as the active mining site (Zone ‘C’).

The objectives of the study were to compare these three selected habitats using a combination of nematode community indices and trophic group abundance and to examine substrate characteristics through nematode community measures using multivariate analyses.

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2. Description of site and protocol of study

See Figure 1.

Figure 1.

A. Satellite images of the coal mine region. B. Topographic map of the study region showing three different zones. Zone ‘A’ = green color, Zone ‘B’ = light green color (samples were collected from median age coalmine spoils from this zone), Zone ‘C’ = dark blue color.

The coal mine region was divided broadly into three differentiating sites on gradient of anthropogenic disturbances.

Zone ‘A’: This area represented the unmined hills covered with wild vegetation and with least or no anthropogenic activities. Samples were collected on altitudinal gradients from foothill to top. The Zone lies between the geographic coordinates 24° 11′57.73″N, 82° 44′31.95″E to 24° 08′49.06″N, 82° 43′33.85″E (Figure 2A and B).

Figure 2.

A, B: Snapshots of Zone ‘A’.

Zone ‘B’: The area represented reclaimed overburden of about 21 years of age. For the proper management of spoils, anthropogenic inference occurs time to time. Samples were collected on an altitudinal gradient from foot to top of overburden. This Zone lies between the geographic coordinates 24° 10′32.99″N, 82° 45′51.29″E to 24° 07′40.38″N, 82° 48′32.51″E (Figure 3A and B).

Figure 3.

A, B: Snapshots of Zone ‘B’.

Zone ‘C’: The area represented the highly disturbed zone with an extremely nutrient-poor environment. Samples were collected on both altitudinal and horizontal basis. The samples were collected between the geographic coordinates 24° 10′57.92″N, 82° 45′07.83″E to 24° 07′47.38″N, 82° 43′24.89″E (Figure 4A and B). On horizontal basis, the active coal mining region was considered as source of disturbance, and the samples were collected in a circumference of 400 m diameter.

Figure 4.

A, B: Snapshots of Zone ‘C’.

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3. Materials and methods

Sampling for Zone ‘A’: Soil sampling was done on an altitudinal gradient from the foot to the top on undisturbed hills. Samples were taken 15–25 cm below the ground level.

Sampling for Zone ‘B’: Soil sampling was done from top to bottom and from the bench of the coal mine spoil.

Sampling for Zone ‘C’: Samples were taken at a 15 m distance along the altitudinal gradient starting from ground level (about 160 m above sea level) to the top (about 400 m high).

Sampling was done using steel corer of 5 cm diam. For each Zone 30 samples were collected with each sample being a composite of five samples. Samples were then stored in plastic bags and transported to the lab for chemical analysis and nematode identification. The climatic and edaphic information were also recorded (Table 1). Geographic data like geographic coordinates, altitude, and temperature were recorded via the navigation device GARMIN -eTREX- 20.

Nematode speciesFunctional guildMean ± SD (abundance)
BacterivoresZone ‘A’
Undisturbed forest
(site for advance quarry)
Zone ‘B’
Spoil of median age
Zone ‘C’
Active site of mining
Achromadora indicaBa210.56 ± 3.5428.24 ± 21.240.0 ± 0.0
Acrobeles cylindricusBa214.39 ± 16.5420.12 ± 15.540.0 ± 0.0
Acrobeloides conoidisBa210.34 ± 7.4512.45 ± 8.640.48 ± 0.75
Acrobeloides nanusBa218.15 ± 2.5615.10 ± 12.100.0 ± 0.0
Acrobelophis minimusBa25.24 ± 3.210.63 ± 2.250.0 ± 0.0
Alaimus primitivusBa318.35 ± 32.3030.34 ± 44.530.0 ± 0.0
Amphidelus sylvaticusBa316.3 ± 15.5313.54 ± 8.640.0 ± 0.0
Cephalobus cubaensisBa210.4 ± 6.247.52 ± 10.250.0 ± 0.0
Cephalobus parvusBa210.86 ± 7.257.81 ± 5.230.0 ± 0.0
Ceratoplectus armatusBa29.26 ± 4.2319.83 ± 25.30.0 ± 0.0
Chiloplacus symmetricusBa21.45 ± 7.590.25 ± 1.250.0 ± 0.0
Chiloplectus indicusBa20.50 ± 2.760.65 ± 2.540.0 ± 0.0
Eucephalobus oxyuroidesBa216.68 ± 18.8518.25 ± 12.240.0 ± 0.0
Geomonhystera glandulataBa212.24 ± 8.358.53 ± 12.530.0 ± 0.0
Halicephalobus gingivalisBa10.00 ± 0.000.94 ± 3.650.0 ± 0.0
Mesorhabditis vernalisBa113.51 ± 8.5622.45 ± 9.450.36 ± 0.67
Mesorhabditis minutaBa18.45 ± 6.4515.53 ± 12.670.48 ± 0.84
Monhystera gracilisBa211.84 ± 6.8515.28 ± 14.530.0 ± 0.0
Monhystrella kerryiBa212.54 ± 6.449.28 ± 5.680.0 ± 0.0
Oscheius vulvastriatus sp .n.Ba10.00 ± 0.008.22 ± 3.250.0 ± 0.0
Panagrellus dubiusBa13.15 ± 1.231.75 ± 0.570.0 ± 0.0
Panagrolaimus dendroctoniBa215.83 ± 6.868.75 ± 6.550.0 ± 0.0
Plectus geophilusBa216.84 ± 18.2418.49 ± 20.240.0 ± 0.0
Plectus parvusBa27.42 ± 4.245.77 ± 14.990.0 ± 0.0
Poikilolaimus oxycercusBa12.15 ± 0.240.55 ± 2.500.0 ± 0.0
Prismatolaimus intermediusBa312.46 ± 14.4816.86 ± 12.860.0 ± 0.0
Teratolobus regulusBa32.43 ± 1.751.25 ± 3.450.0 ± 0.0
Tylocephalus primitivusBa23.26 ± 2.363.86 ± 1.670.0 ± 0.0
Wilsonema bangaloreiensisBa23.35 ± 1.753.55 ± 5.430.0 ± 0.0
Zeldia punctataBa22.72 ± 2.240.74 ± 2.240.0 ± 0.0

Table 1.

Abundance of bacterivore species in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

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4. Results and observations

4.1 Nematode abundance

Zone ‘A’: Nematode abundance was found high in Zone ‘A’ and mean value of 1748.63 ± 163.18 (1576–1859) nematodes were found in 100 g of samples.

Zone ‘B’: Nematode abundance was low in Zone ‘B’ comparably to Zone ‘A’ and mean value of 983.8 ± 120.2 (748.57–1168.74) nematodes were found in 100 g of samples.

Zone ‘C’: Nematode abundance was negligible in Zone ‘C’, which was the active mining Zone and mean of 0.42 ± 0.37 (0.08–0.42) nematodes isolated from 100 g of samples.

4.2 Nematode species diversity

Zone ‘A’: A total of 119 species belonging to 107 genera, 40 families and eight orders were recorded from Zone ‘A’. The dominant species were Aphelenchoides composticola, Aphelenchus avenae, Dorylaimoides constrictus, Labronema baqrii, Labronema deoriaensis, Laimydorus baldus, Thornenema mauritianum, Tripyla glomerans, Tripyla arenicola, Mylonchulus contractus, Mylonchulus vasis, Sporonchulus ibitiensis, Aporcelaimellus tropicus.

Zone ‘B’: Zone ‘B’ recorded 105 species belonging to 96 genera, 39 families and eight orders. The number and dominance were quite comparable with Zone ‘A’.

Zone ‘C’: Only three species belonging to two genera, one family and one order were observed at Zone ‘C’ (Figure 5).

Figure 5.

Number of species, genera and individuals observed in three different zones of coal mine region.

4.3 Nematode trophic structure

In Zone ‘A’ there was much diversity of omnivores and predators in comparison to bacterivores and in Zone ‘B’ bacterivores and fungivores nematodes were the abundant group over omnivores and predators and Zone ‘C’ had only three bacterivore species.

4.3.1 Zone ‘A’

4.3.1.1 Trophic groups

Bacterivores (bacteriophagous): Of the 28 bacterivore species recorded (Table 2), five most abundant species were Acrobeloides nanus, Plectus geophilus, Eucephalobus oxyuroides, Amphidelus sylvaticus and Acrobeles cylindricus.

Nematode speciesFunctional guildMean ± SD (abundance)
FungivoresZone ‘A’
Undisturbed forest (Site for advance quarry)
Zone ‘B’
Spoil of Median Age
Zone ‘C’
Active site of mining
Aglenchus agricolaFu216.43 ± 6.0516.48 ± 8.450.0 ± 0.0
Aphelenchus avenaeFu232.04 ± 18.6412.6 ± 10.350.0 ± 0.0
Aphelenhoides besseyiFu236.13 ± 20.1318.74 ± 16.360.0 ± 0.0
Axonchium thoubalicumFu59.40 ± 3.562.25 ± 1.750.0 ± 0.0
Basiriotyleptus minimusFu28.95 ± 4.316.25 ± 4.210.0 ± 0.0
Belondira apiticaFu510.42 ± 4.105.40 ± 2.500.0 ± 0.0
Daladenus wilsoniFu23.67 ± 1.5813.43 ± 6.480.0 ± 0.0
Diptherophora alamiFu312.10 ± 4.204.61 ± 2.800.0 ± 0.0
Ditylenchus dipsaciFu23.26 ± 2.0511.4 ± 6.230.0 ± 0.0
Dorylaimellus indicusFu516.82 ± 6.428.64 ± 3.560.0 ± 0.0
Dorylaimoides constrictusFu239.27 ± 20.4613.45 ± 6.450.0 ± 0.0
Filenchus mircodorusFu222.30 ± 10.4616.43 ± 8.230.0 ± 0.0
Filenchus vulgarisFu218.20 ± 9.6515.43 ± 6.280.0 ± 0.0
Leptonchus granulosusFu24.85 ± 2.245.44 ± 3.240.0 ± 0.0
Roqueus indicusFu54.64 ± 3.415.53 ± 2.540.0 ± 0.0
Tylencholaimellus acutusFu428.41 ± 14.4212.34 ± 6.800.0 ± 0.0
Tyleptus projectusFu27.23 ± 3.5410.23 ± 4.250.0 ± 0.0

Table 2.

Abundance of fungivores species in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

Fungivores (mycophagous): Seventeen fungivore species were recorded (Table 3), among which the five most abundant ones were Dorylaimoides constrictus, Aphelenhoides composticola, Aphelenchus avenae, Tylencholaimellus acutus and Filenchus mircodorus.

Nematode speciesFunctional guildMean ± SD (abundance)
HerbivoresZone ‘A’
Undisturbed forest (site for advance quarry)
Zone ‘B’
Spoil of median age
Zone ‘C’
Active site of mining
Basiria abberansH216.80 ± 6.467.25 ± 4.230.0 ± 0.0
Boleodorus filiformisH29.35 ± 4.053.36 ± 2.350.0 ± 0.0
Coslenchus areolatusH26.87 ± 5.641.53 ± 2.500.0 ± 0.0
Helicotylenchus dihysteraH39.64 ± 4.3812.68 ± 5.850.0 ± 0.0
Hemicriconemoides communisH315.38 ± 8.366.86 ± 4.250.0 ± 0.0
Hemicycliophora corbettiH31.50 ± 2.351.02 ± 2.210.0 ± 0.0
Hoplolaimus indicusH32.30 ± 3.6610.25 ± 6.230.0 ± 0.0
Longidoroides longiurusH513.45 ± 8.362.52 ± 0.00.0 ± 0.0
Longidorus elongatusH516.84 ± 10.351.08 ± 2.500.0 ± 0.0
Malenchus nanellusH24.68 ± 3.451.24 ± 3.650.0 ± 0.0
Merlinius brevidensH33.87 ± 2.361.25 ± 0.580.0 ± 0.0
Neopsilenchus affinisH22.48 ± 2.011.20 ± 2.360.0 ± 0.0
Paratylenchus similisH29.85 ± 6.4612.26 ± 6.580.0 ± 0.0
Pratylenchus brachyurusH210.64 ± 6.680.0 ± 0.00.0 ± 0.0
Psilenchus mixusH34.65 ± 3.340.56 ± 1.140.0 ± 0.0
Rotylenchus mircrostylusH36.80 ± 4.880.26 ± 2.240.0 ± 0.0
Scutellonema brachyurumH31.54 ± 2.340.50 ± 1.540.0 ± 0.0
Scutylenchus hexincisusH32.38 ± 1.360.35 ± 2.250.0 ± 0.0
Trichodorus boraiH48.62 ± 4.381.24 ± 2.520.0 ± 0.0
Tylenchorynchus striatusH311.56 ± 8.4410.23 ± 6.450.0 ± 0.0
Tylenchus arcuatusH312.56 ± 6.355.86 ± 3.540.0 ± 0.0
Xiphinema americanumH56.24 ± 9.259.25 ± 5.240.0 ± 0.0

Table 3.

Abundance of herbivore species in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

Herbivores (plant parasites): Of the 22 herbivore species (Table 4), five most abundant ones were Longidorus elongates, Basiria abberans, Longidoroides longiurus, Tylenchus arcuatus and Tylenchorynchus striatus.

Nematode speciesFunctional guildMean ± SD (abundance)
OmnivoresZone ‘A’
Undisturbed forest (site for advance quarry)
Zone ‘B’
Spoil of median age
Zone ‘C’
Active site of mining
Amphidorylaimus flagellicaudaOm46.24 ± 4.36
Cephalodorylaimus papillatusOm46.20 ± 5.48
Crassolaibium gharwalienseOm416.84 ± 9.654.24 ± 2.35
Discomyctus cephalatusOm412.86 ± 9.806.33 ± 3.26
Dorylaimus stagnalisOm410.54 ± 5.461.56 ± 3.28
Enchodelus macrodorusOm48.64 ± 5.240.45 ± 1.56
Eudorylaimus silvaticusOm413.64 ± 8.305.26 ± 3.25
Eumenicus monhysteraOm48.64 ± 5.45
Ischiyodorylaimus navusOm43.58 ± 4.24
Labronema deoriaensisOm420.42 ± 12.9215.50 ± 11.54
Labronema baqriiOm424.56 ± 18.4518.35 ± 8.62
Allodorylaimus irritansOm431.54 ± 22.562.45 ± 3.56
Makatinus heynsiOm47.45 ± 6.52
Mesodorylaimus subtiloidesOm420.38 ± 11.36
Mesodorylaimus indicusOm420.38 ± 16.3623.56 ± 13.46
Microdorylaimus parvusOm418.45 ± 8.468.54 ± 5.64
Moshajia idioforaOm412.45 ± 6.758.65 ± 5.60
Opisthodorylaimus cavalcantiiOm416.45 ± 9.45
Oriverutus sundarusOm419.44 ± 12.46
Oxydirus ganagticusOm510.65 ± 6.54
Paraoxydirus gigasOm58.64 ± 4.84
Porodorylaimus sturhaniOm415.46 ± 12.24
Pungentella porosaOm49.86 ± 5.68
Pungentus angulatusOm412.6 ± 9.832.60 ± 1.35
Thonus cylindricusOm414.24 ± 8.3813.50 ± 8.45
Thornenema mauritianumOm422.54 ± 16.3815.68 ± 8.54

Table 4.

Abundance of omnivores species in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

Omnivores: Five most abundant species among the 25 omnivore species (Table 5) were Allodorylaimus irritans, Thornenema mauritianum, Labronema deoriaensis, Mesodorylaimus indicus and Mesodorylaimus subtiloides.

Nematode speciesFunctional guildMean ± SD (abundance)
PredatorsZone ‘A’
Undisturbed forest (site for advance quarry)
Zone ‘B’
Spoil of median age
Zone ‘C’
Active site of mining
Actinolaimus armatusCa512.22 ± 5.644.68 ± 2.52
Actus minutusCa411.22 ± 6.426.86 ± 6.36
Aporcelaimellus tropicusCa422.60 ± 14.3815.50 ± 8.46
Aporcelaimellus heynsiCa412.68 ± 8.42
Aquatides minutusCa54.23 ± 6.699.40 ± 6.53
Clarkus sheriCa49.68 ± 4.38
Discolaimodes bulbiferusCa516.46 ± 10.2614.84 ± 5.21
Discolaimus majorCa512.45 ± 8.6215.60 ± 8.64
Iotonchus monhysteraCa410.12 ± 5.366.18 ± 2.48
Iotonchus indicusCa415.99 ± 8.594.21 ± 3.26
Ironus dentifurcatusCa418.62 ± 9.968.64 ± 4.42
Laimydorus papillatusCa416.45 ± 4.66
Laimydorus baldusCa422.68 ± 6.6815.21 ± 8.62
Michonchus rectangularisCa414.44 ± 8.6411.64 ± 6.30
Mononchoides longicaudaCa10.25 ± 3.36
Mulveyellus jairiCa49.66 ± 3.361.89 ± 2.06
Mylonchulus contractusCa423.45 ± 14.6415.24 ± 11.42
Mylonchulus viasisCa426.20 ± 18.4616.54 ± 8.44
Neoactinolaimus thorneiCa56.48 ± 5.363.05 ± 2.58
Nygellus subclavatusCa416.28 ± 6.666.26 ± 5.88
Nygolaimus timmiCa58.35 ± 6.452.83 ± 1.26
Prionchulus muscorumCa48.32 ± 4.32
Solidedens bisexualisCa56.64 ± 7.384.21 ± 2.36
Sporonchulus ibitiensisCa425.68 ± 44.6110.58 ± 13.60
Sporonchulus vagabundusCa418.68 ± 12.685.36 ± 3.36
Tripyla glomeransCa324.35 ± 18.3832.26 ± 22.42
Tripyla arenicolaCa331.24 ± 24.2420.25 ± 12.24

Table 5.

Abundance of predator species in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

Predators: Of a total of 26 predator species (Table 6), the five most abundant ones were recorded as Tripyla arenicola Mylonchulus viasis Tripyla glomerans Mylonchulus contractus and Laimydorus baldus.

Type Area ‘A’
Non minning forest
Type Area ‘B’
Spoil of median age
Type Area ‘C’
Active site of mining
Genera (total count)1091042
Species (total count)1141103
Individuals1748.63 ± 163.18 (1576–1859)983.80 ± 120.24 (748.57–1168.74)0.42 ± 0.37 (0.08–0.42)
Margalef diversity16.17 ± 0.35 (15.58–16.79)15.11 ± 0.59 (14.23–16.23)
Menhinick diversity3.92 ± 0.35 (3.63–3.95)3.43 ± 0.43 (2.85–3.98)
Simpson dominance0.01 ± 0.00 (0.01–0.01)0.01 ± 0.00 (0.01–0.01)
Inverse Simpson dominance91.14 ± 3.25 (82.25–96.35)67.50 ± 7.33 (54.32–76.15)
Hill’s effective index98.23 ± 2.43 (97.24–99.89)75.64 ± 4.90 (65.18–77.89)
Shannon diversity4.58 ± 0.45 (3.97–5.65)4.32 ± 0.44 (3.54–4.88)
Simpson diversity0.99 ± 0.00 (0.99–0.99)0.99 ± 0.00 (0.99–0.99)
Brillouin diversity4.51 ± 0.65 (3.85–5.78)4.19 ± 0.34 (3.50–4.60)
Sheldon Index0.82 ± 0.00 (0.82–0.82)0.72 ± 0.00 (0.72–0.72)
Heip Index0.82 ± 0.00 (0.82–0.82)0.72 ± 0.00 (0.72–0.72)
Maturity Index (MI)3.46 ± 0.26 (3.36–3.87)2.96 ± 0.50 (2.87–3.12)
Plant Parasitic Index (PPI)0.78 ± 0.09 (0.50–1.25)3.26 ± 0.23 (2.00–3.65)
Trophic Diversity Index (TDI)1.17 ± 0.02 (1.06–1.32)1.09 ± 0.04 (1.04–1.25)1.00 ± 0.00 (1.04–1.25)
Nematode channel ratio (NCR %)0.40 ± 0.02 (0.35–0.72)0.68 ± 0.02 (0.62–0.73)
Structure Index (SI)68.23 ± 10.20 (58.30–77.43)45.1 5 ± 5.85 (40.68–52.68)
Enrichment Index (EI)12.62 ± 4.66 (7.65–32.25)22.62 ± 12.58 (16.53–64.65)9.54 ± 1.54 (9.50–10.56)
Basal Index (BI)41.28 ± 12.30 (32.04–62.05)57.29 ± 8.10 (44.24–67.35)74.46 ± 3.25 (70.13–78.16)
Channel Index (CI)0.31 ± 0.40 (0.25–0.84)0.54 ± 0.42 (0.50–1.24)

Table 6.

Ecological parameters of nematode population in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

4.3.2 Zone ‘B’

4.3.2.1 Trophic groups

Bacterivores (bacteriophagous): Of the 29 bacterivore species recorded (Table 2), five most abundant species were Alaimus primitivus, Achromadora indica, Mesorhabditis vernalis, Acrobeles cylindricus and Ceratoplectus armatus.

Fungivores (mycophagous): Seventeen fungivore species were recorded (Table 3), among which five most abundant ones were Aphelenhoides composticola, Aglenchus agricola, Filenchus mircodorus, Dorylaimoides constrictus and Aphelenchus avenae.

Herbivores (plant parasites): Of the 21 herbivore species (Table 4), five most abundant ones were Helicotylenchus dihystera, Paratylenchus similis, Hoplolaimus indicus, Tylenchorynchus striatus and Xiphinema americanum.

Omnivores: The five most abundant species among the 15 omnivore species (Table 5) were Mesodorylaimus indicus, Labronema baqrii, Thornenema mauritianum, Labronema deoriaensis and Thonus cylindricus.

Predators: Of a total of 22 predator species (Table 6), the five most abundant ones were recorded as Tripyla glomerans, Tripyla arenicola, Mylonchulus vasis, Discolaimus major and Aprocelaimellus tropicus.

4.3.3 Zone ‘C’

4.3.3.1 Trophic group

Bacterivores: In Zone ‘C’ only three species were reported from bacteriovores (Table 2) viz., Mesorhabditis vernalis, Mesorhabditis minuta and Acrobeloides nanus.

4.4 Biomass

The relative biomass of different trophic groups (Figure 6) varied in three zones and are as follows:

Figure 6.

Percentage biomass of the different trophic groups in the three different zones of coalfield region.

Zone ‘A’: The highest biomass was of predators, that is, about 36% of the total nematode biomass and the lowest,that is, 8% was constituted by bacterivores. Omnivore species showed 32% biomass followed by herbivores (13%) while fungivores constituted 11% of the total nematode biomass. The total nematode biomass in this Zone was calculated to be 6162.99 μg in 100 g. of soil.

Zone ‘B’: The nematode biomass demonstrated similar trends in Zone ‘B’ compared to Zone ‘A’. The highest biomass was of predators that is, 37% of the total nematode biomass while herbivores constituted the lowest 13%, omnivores constituted 21% followed by bacterivores (15%) and fungivores (14%) of the total nematode biomass. The total nematode biomass in this Zone was estimated to be 3438.93 μg in 100 g of soil.

Zone ‘C’: In Zone ‘C’ there were only three bacterivore species that constitute nematode biomass. The total nematode biomass in this Zone was estimated to be 2.37 μg in 100 g of soil.

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5. Diversity indices

Diversity indices and nematode maturity indices were calculated for each zone to assess the diversity of nematode species and maturity of soil ecosystem.

5.1 Margalef index (species richness)

In Zone ‘A’ the Margalef species richness index was found to be 16.17 ± 0.35 (15.58–16.79). Zone ‘B’ was calculated to be 15.11 ± 0.59 (14.23–16.23) and in Zone ‘C’, where only two species were recorded, the index was incalculable (Table 7 and Figure 7).

CharacteristicsMining sites
Zone ‘A’
Undisturbed forest (site for advance quarry)
Zone ‘B’
Spoil of median age
Zone ‘C’
Active site of mining
<2.0 mm (gravel)9.00 ± 0.9213.00 ± 1.8020.00 ± 2.15
2.–0.2 mm (sand)63.00 ± 3.5072.00 ± 5.2488.55 ± 6.54
0.2–0.1 mm (silt)13.00 ± 1.2310.00 ± 0.243.00 ± 1.26
<0.1 mm (clay)8.10 ± 1.457.80 ± 1.253.20 ± 1.56
pH6.54 ± 0.026.82 ± 0.025.45 ± 0.26
Bulk density1.25 ± 0.181.26 ± 0.081.75 ± 0.16
Natural moisture content (%)12.20 ± 2.2611.60 ± 0.286.2 ± 0.25
Porosity46.27 ± 2.2542.62 ± 2.6533.65 ± 2.66
Water holding capacity53.00 ± 0.9252.00 ± 2.6524.84 ± 3.25
Soil organic carbon (mg C/g)3.45 ± 0.0242.38 ± 0.028ND
Total soil N (μg N/g)2456.63 ± 148.34194.94 ± 25.46ND
Exchangeable K μg P/g spoil271.95 ± 6.7416.96 ± 2.35ND
Electrical conductivity ms/cm0.05 ± .010.33 ± 0.010.65 ± 0.01

Table 7.

Abiotic parameters in three zones of coal mine region of district Sonebhadra Uttar Pradesh.

Figure 7.

Comparison of species diversity indices in three selected zones.

5.2 Menhinick index (species richness)

Menhinick diversity, a measure to compare the species richness, was calculated to be 3.92 ± 0.35 (3.63–3.95) in Zone ‘A’, in Zone ‘B’ it was found to be 3.43 ± 0.43 (2.85–3.98) and the Menhinick index was incalculable for Zone ‘C’ (Table 7 and Figure 7).

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6. Evenness indices

The values for both the evenness indices (Sheldon and Heip indices) were observed to be the same for the two Zones ‘A’ and ‘B’ and calculated to be 0.82 ± 0.00 (0.82–0.82) and 0.72 ± 0.00 (0.72–0.72), respectively. For Zone ‘C’ the values were inestimable (Table 7).

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7. Maturity index

In Zone ‘A’ where there was the least disturbance, the MI was highest and a mean value of 3.46 ± 0.26 (3.36–3.87) was observed representing the structured soil ecosystem. Zone ‘B’ which represented coal mine-managed spoil of 21 years period, showed an MI equal to 2.96 ± 0.50 (2.87–3.12) while in Zone ‘C’ MI was incalculable (Table 7 and Figure 7).

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8. Plant parasitic index (PPI)

In Zone ‘A’ the PPI was calculated to be 0.78 ± 0.09 (0.50–1.25) whereas in Zone ‘B’ the PPI value was 3.26 ± 0.23 (2.00–3.65). The PPI and MI showed an inverse correlation (Table 7 and Figure 8).

Figure 8.

Comparison of MI, PPI, and TDI in three different zones of coal mine region.

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9. Trophic diversity index

The value of trophic diversity index of Zone ‘A’ was 1.17 ± 0.02 (1.06–1.32), while those of Zone ‘B’ and Zone ‘C’ were 1.09 ± 0.04 (1.04–1.25) and 1.00 ± 0.00 (1.04–1.25), respectively (Table 8 and Figure 8).

VariablesBac. SpFung. SpHerb. SpOmi. SpPred. sp.MIPPITDISIEIBICINCRTot. Indv.pHBDPorositySMWHCToCTSNAvail. KEC
Bac. Sp10.9960.9970.8720.9910.9490.9990.8980.9830.705−0.8610.8840.9820.8860.988−0.986−0.8500.9440.9960.9430.9320.534−0.804
Fung. Sp0.99611.0000.9100.9990.9720.9990.9320.9950.643−0.9010.9200.9630.9210.998−0.996−0.8910.9681.0000.9680.8990.603−0.851
Herb. Sp0.9971.00010.9050.9980.9691.0000.9270.9940.653−0.8960.9150.9670.9170.997−0.995−0.8850.9651.0000.9640.9040.593−0.844
Omi. Sp0.8720.9100.90510.9310.9820.8920.9990.9480.269−1.0001.0000.7661.0000.937−0.942−0.9990.9850.9140.9850.6360.879−0.992
Pred. sp.0.9910.9990.9980.93110.9830.9960.9500.9990.602−0.9230.9390.9480.9411.000−0.999−0.9140.9800.9990.9800.8740.644−0.877
MI0.9490.9720.9690.9820.98310.9610.9910.9910.445−0.9780.9860.8730.9870.986−0.988−0.9731.0000.9741.0000.7700.774−0.951
PPI0.9990.9991.0000.8920.9960.96110.9160.9900.675−0.8820.9030.9740.9050.994−0.992−0.8710.9570.9990.9560.9160.569−0.828
TDI0.8980.9320.9270.9990.9500.9910.91610.9640.321−0.9971.0000.8001.0000.955−0.959−0.9950.9930.9350.9930.6780.852−0.984
SI0.9830.9950.9940.9480.9990.9910.9900.96410.562−0.9400.9550.9310.9561.000−1.000−0.9330.9890.9960.9880.8490.681−0.900
EI0.7050.6430.6530.2690.6020.4450.6750.3210.5621−0.2470.2910.8250.2950.588−0.576−0.2260.4320.6360.4290.914−0.223−0.145
BI−0.861−0.901−0.896−1.000−0.923−0.978−0.882−0.997−0.940−0.2471−0.999−0.751−0.999−0.9290.9351.000−0.981−0.905−0.981−0.619−0.8890.995
CI0.8840.9200.9151.0000.9390.9860.9031.0000.9550.291−0.99910.7811.0000.945−0.950−0.9980.9890.9230.9890.6540.868−0.989
NCR0.9820.9630.9670.7660.9480.8730.9740.8000.9310.825−0.7510.78110.7830.942−0.937−0.7370.8660.9610.8640.9830.367−0.678
Tot. Indv.0.8860.9210.9171.0000.9410.9870.9051.0000.9560.295−0.9991.0000.78310.946−0.951−0.9970.9890.9250.9900.6580.866−0.988
pH0.9880.9980.9970.9371.0000.9860.9940.9551.0000.588−0.9290.9450.9420.9461−1.000−0.9210.9830.9980.9830.8660.657−0.886
BD−0.986−0.996−0.995−0.942−0.999−0.988−0.992−0.959−1.000−0.5760.935−0.950−0.937−0.951−1.00010.926−0.986−0.997−0.986−0.858−0.6690.892
Porosity−0.850−0.891−0.885−0.999−0.914−0.973−0.871−0.995−0.933−0.2261.000−0.998−0.737−0.997−0.9210.9261−0.976−0.895−0.977−0.602−0.8990.997
SM0.9440.9680.9650.9850.9801.0000.9570.9930.9890.432−0.9810.9890.8660.9890.983−0.986−0.97610.9711.0000.7600.783−0.955
WHC0.9961.0001.0000.9140.9990.9740.9990.9350.9960.636−0.9050.9230.9610.9250.998−0.997−0.8950.97110.9700.8940.610−0.856
ToC0.9430.9680.9640.9850.9801.0000.9560.9930.9880.429−0.9810.9890.8640.9900.983−0.986−0.9771.0000.97010.7590.785−0.956
TSN0.9320.8990.9040.6360.8740.7700.9160.6780.8490.914−0.6190.6540.9830.6580.866−0.858−0.6020.7600.8940.75910.192−0.534
Avail. K0.5340.6030.5930.8790.6440.7740.5690.8520.681−0.223−0.8890.8680.3670.8660.657−0.669−0.8990.7830.6100.7850.1921−0.932
EC−0.804−0.851−0.844−0.992−0.877−0.951−0.828−0.984−0.900−0.1450.995−0.989−0.678−0.988−0.8860.8920.997−0.955−0.856−0.956−0.534−0.9321

Table 8.

Correlation coefficient among the various variables used for nematode community analysis.

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10. Weighted faunal analysis

The indices calculated depended on weighted nematode indicator guilds [1, 12] to assess the level of organic enrichment (EI), the decomposition pathways (CI) (Table 8 and Figure 9), the basal/minimal level of resource utilization (BI) and the structured status or stability of ecosystem (SI). In Zone ‘A’ the structure index (SI) was highest i.e., 68.23 ± 10.20 (58.30–77.43), while in Zone ‘B’ the SI value was calculated to be 45.15 ± 5.85 (40.68–52.68) and in Zone ‘C’ the SI was zero. On the contrary, the enrichment index (EI) was calculated to be 12.62 ± 4.66 (7.65–32.25), 22.62 ± 12.58 (16.53–64.65) and 9.54 ± 1.54 (9.50–10.56) in Zone ‘A’, Zone ‘B’ and Zone ‘C’, respectively. The values of basal index (BI) for three different Zones were 74.46 ± 3.25 (70.13–78.16) in Zone ‘C’ followed by value of 57.29 ± 8.10 (44.24–67.35) in Zone ‘B’; lowest value of BI was observed in 41.28 ± 12.30 (32.04–62.05) in Zone ‘A’. The values of channel index (CI) in Zone ‘A’ and Zone ‘B’ were 0.31 ± 0.40 (0.25–0.84) and 0.54 ± 0.42 (0.50–1.24), respectively while CI was incalculable in Zone ‘C’ (Table 8 and Figure 10).

Figure 9.

Comparison of CI and NCR in three different zones of coal mine region.

Figure 10.

Comparison of SI, EI, and BI in three different zones of coal mine region.

In faunal ordination graph between structure index and enrichment index, it was observed that in Zone ‘A’ the mean values were placed in quadrat ‘C’ whereas the mean values for Zone ‘B’ were at the interjunction of quadrat ‘A’ and quadrat ‘C’. In Zone ‘C’ the soil had almost no vegetation, low nutrient, and very little organic carbon (Figure 11).

Figure 11.

Faunal profiles representing the food-web condition in relation to its structure (SI) and enrichment (EI) as indicated by the “weighted faunal analysis” (mean ± SE).

11. Correlations among different variables

The Principal component analysis (PCA) was carried out and a correlation circle was retrieved (Figure 12), which depicted the correlations among various variables. Structure index (SI) showed a positive correlation with number of bacterivores species (r = 0.98, p < 0.05), fungivores (r = 0.99, p < 0.05), and herbivores (r = 0.99, p < 0.05), omnivores (r = 0.94, <0.05) and predators (r = 0.99, p < 0.05). Almost all the trophic groups showed a positive relationship with SI. Structure index also showed a positive correlation with PPI (r = 0.99, p < 0.05), pH (r = 1.00, p < 0.05), TDI (r = 0.96, p < 0.05), NCR (r = 0.93, p < 0.05) and MI (r = 0.99, p < 0.05). SI also depicted low positive relationship EI (r = 0.56, p < 0.05) and strong negative correlation with CI (r = 0.99, p < 0.05) and BI (r = −0.94, p < 0.05). EI showed strong negative correlation with CI and BI all at (p < 0.05). All trophic groups showed a positive correlation among themselves (p < 0.05) and also with PPI, pH, and MI. Among the trophic groups, only bacterivores showed robust positive correlation with EI (r = 0.70, p < 0.05). EI did not show any correlation with PPI, nonsignificant correlation with SI and significant positive correlation with MI. Soil organic carbon was significantly positively correlated with all trophic groups (p < 0.05) and show a negative correlation with electrical conductivity, porosity, basal index and bulk density (Table 8 and Figure 13).

Figure 12.

Biplot correlation circle depicting the correlation among the different variables.

Figure 13.

Biplot ordination (PCA) of observation dates on trophic groups abundance in terms of species, total nematode abundance and various other variables with respect three different zones in coal mine region.

In Biplot ordination graph the three Zones can be clearly differentiated with respect to biotic and abiotic variables and can be superimposed on the anthropogenic disturbances to assess its impact.

12. Discussion

Human activities disturb the soil ecosystem and affect soil nematode diversity [8, 23, 24, 25]. The effect is well reflected in present study where there is a significant impact on below-ground soil biodiversity with special reference to nematodes. The transition from aboveground plant heterogeneity in natural forests to extreme loss of vegetation due to mining activities is reflected in the changed community characteristics of soil nematodes and in their species diversity.

In Zone ‘A’, the number of species (species richness) was greater, but species dominance was low while in Zone ‘B’, an intermediate value of species richness with high species dominance of bacterial-feeders was recorded. These species viz., Achromadora porosus sp. n., Acrobeloides nanus and Acrobeles cylindricus showed tolerance to environmental stress, seasonal anhydrobiosis and other anthropogenic disturbances. Further, Zone ‘B’ represented the reclaimed 21 years old coal mine spoil with some incorporation of amendments to enrich the soil thus another reason for abundance of bacterivores. In Zone ‘C’ only three species were found viz., Mesorhabditis tenuhystera sp. n., Mesorhabditis vernalis and Acrobeloides nanus. As, Zone ‘C’ was the active site for mining and with constant disturbances, the soil was almost lifeless with no nutrients and the area simply represented old exposed rocks mixed with coal dust having little organic carbon. The chemical residue from blasting and heavy metal contamination from mining process hindered the growth of plants and animals. The rhabditid species (cp-1) found in active mining Zone ‘C’, were enrichment opportunists and might have passively reached the active mining site via wind, water or through phoresis [26]. However, their sustenance showed their greater tolerance abilities towards mineral and industrial waste materials [27]. Thus Mesorhabditis spp. demonstrated greater endurance. In unfavorable periods, production of dauer larvae was the possible survival strategy for these enrichment opportunists. Zone ‘B’, the relatively less disturbed site, might have initially shown the growth of zymogenous (fast growing) microflora on fossil organic carbon that formed food for the microbivorous species, a condition also noticed by Háněl [28] who reported the abundant populations of genera Acrobeloides and Aphelenchoides. Zone ‘A’ consisting of natural wild forest, showed heterogenetic topography, and the litter accumulation due to undulating surfaces, was the reason for relatively high diversity and abundance of all trophic groups including bacterivores with values of both Simpson’s index and Shannon’s index comparable to any natural forest [23, 29, 30, 31, 32].

Although there was not a major difference between the total number of species of fungivores and herbivores in Zone ‘A’ and Zone ‘B’, the mean abundance of both the trophic groups was much high in Zone ‘A’. Despite the fact that stable ecosystems like Zone ‘A’ demonstrated species diversity and evenness, very often the species diversity has been reported to be greater in habitats subject to intermediate levels of disturbance because stochastic, intermediate (partial) elimination of resources by disturbance leads to species-specific mortality allowing the co-existence of competitively inferior species [33]. The combination of intermediate level of disturbance with intermediate productivity levels demonstrated a peak in species richness, not only due to periodic decreases of competitively dominant species but also to increased niche packing [34]. Structurally, complex environments thus provide more niches thereby increasing species diversity [35]. In conformity with the earlier reports on increased diversity at intermediate disturbance [36, 37, 38] and a positive relationship between habitat complexity and species diversity [39, 40, 41, 42, 43, 44], Zone B demonstrated similar status. It showed an intermediate level of disturbances and the species diversity too was somewhat closer to Zone A which represented undisturbed zone. Large mononchid and large dorylaimoid nematodes with higher c-p values were mostly missing from Zone ‘B’ (Table 5), except for Discolaimus major and Discolaimoides bulbiferous which constituted the predator group along with another low c-p value predator and disturbance or enrichment indicator Mononchoides longicauda. Zone ‘C’ simply represented bacterivore colonizers with c-p values 1 and 2. Thus the nematode community structure in the selected zones showed typical examples of maturity and species diversification with respect to anthropogenic disturbances.

The maturity index of Zone ‘A’ Zone ‘B’ did not show much difference and could well indicate the status of Zone ‘A’ as structured and mature type of ecosystem and Zone ‘B’ as moderately structured and maturing type. Zone ‘C’ was obviously a highly disturbed region because of mining activities. These results agree well with those reported by Háněl [28] on coal mines.

The faunal profile results also corresponded with MI values of Zone ‘A’ and Zone ‘B’ as both reflected the dominant fungal decomposition pathways whereas Zone ‘C’ with disturbed and degraded food web demonstrated bacterial decomposition channels.

Although the present study has provided some pertinent information related to nematodes found in coal mine areas and the differences in nematode communities and their gradual succession in different overburdens or spoils, a detailed discussion on the role of individual species could not be done due to space constraint. Nevertheless, it is clearly evident that nematode assemblages can be evaluated and can serve as excellent tools for environmental monitoring or environment quality assessment. The study has opened up avenues for more studies to be conducted in coal mine areas using nematodes as models. Also, future studies in this direction may further vouch for comparison and may indicate any specificity or association of nematode taxa to coal mine areas. Such information may also have predictive value about the specificity and occurrence of species in degraded habitats, early successional and late successional stages to undisturbed habitats and can be used in modeling and predicting future changes in biodiversity and species interactions with land use changes.

13. Conclusion

In conclusion, this study highlights the significant impact of human activities on soil nematode diversity, particularly in the context of mining activities. The transition from natural forests to mining sites results in notable changes in soil nematode communities and species diversity.

In Zone ‘A’, which represents undisturbed natural forests, there is a higher species richness and lower species dominance among nematodes. In Zone ‘B’, which is a reclaimed coal mine spoil with some soil enrichment measures, there is an intermediate level of species richness but a higher dominance of bacterial-feeders. These nematode species in Zone ‘B’ exhibit tolerance to environmental stress and anthropogenic disturbances. On the other hand, Zone ‘C’, an actively mined area with constant disturbances and little organic matter, contains only a few nematode species, primarily enrichment opportunists with a high tolerance for mineral and industrial waste materials.

The study also suggests that intermediate levels of disturbance, as seen in Zone ‘B’, can lead to increased species diversity due to periodic decreases of dominant species and increased niche packing. Habitat complexity, as observed in both Zone ‘A’ and Zone ‘B’, also contributes to higher species diversity.

The maturity index values of Zone ‘A’ and Zone ‘B’ indicate structured and mature ecosystems, while Zone ‘C’ represents a highly disturbed and degraded region due to mining activities. The faunal profile results align with the maturity index values, with Zone ‘A’ and Zone ‘B’ dominated by fungal decomposition pathways and Zone ‘C’ characterized by bacterial decomposition channels.

While this study provides valuable insights into nematode communities in coal mine areas, further research is needed to explore the roles of individual nematode species in these ecosystems. Nematodes can serve as excellent indicators for environmental monitoring and assessing environmental quality. Future studies in this area may reveal specific associations of nematode taxa with coal mine areas, aiding in biodiversity modeling and predicting how species interactions change with land use alterations. This research opens the door to a deeper understanding of the impact of human activities on soil ecosystems and the potential for ecological restoration efforts.

References

  1. 1. Neher DA, Williams KM, Lovell ST. Environmental indicators reflective of road design in a forested landscape. Ecosphere. 2017;8:e01734
  2. 2. Yeates GW, Bongers T, De Goede RGM, Freckman DW, Georgieva SS. Feeding habits in soil nematode families and genera-an outline for soil ecologists. Journal of Nematology. 1993;25:315-331
  3. 3. Costa SR, van der Putten WH, Kerry BR. Microbial ecology and nematode control in natural ecosystems. In: Davies K, Spiegel Y, editors. Biological Control of Plant-Parasitic Nematodes: Progress in Biological Control. Vol. 11. Dordrecht: Springer; 2011. pp. 39-64
  4. 4. Ferris H, Matute MM. Structural and functional succession in the nematode fauna of a soil food web. Applied Soil Ecology. 2003;23:93-110
  5. 5. Ferris H, Venette RC, Lau SS. Population energetics of bacterial feeding nematodes, stage specific development and fecundity rates. Soil Biology and Biochemistry. 1996;28:271-280
  6. 6. Ingham RE, Trofymow JA, Ingham ER, Coleman DC. Interactions of bacteria, fungi, and their nematode grazers: Effects on nutrient cycling and plant growth. Ecological Monographs. 1985;55:119-140
  7. 7. Ferris H, Bongers T, De Goede RGM. Nematode faunal indicators of soil food web condition. Journal of Nematology. 1999;31:534-535
  8. 8. Neher DA. Role of nematodes in soil health and their use as indicators. Journal of Nematology. 2001;33:161-168
  9. 9. Ritz K, TrudgilL DL. Utility of nematode community analysis as an integrated measure of the functional state of the soils: Perspectives and challenges. Plant and Soil. 1999;212:1-11
  10. 10. Bongers T. The maturity index: An ecological measure of an environmental disturbance based on nematode species composition. Oecologia. 1990;83:14-19
  11. 11. De Goede RGM, Georgieva SS, Verschoor BC, Kamerman JW. Changes in nematode community structure in a primary succession of blown-out areas in a drift sand landscape. Fundamental and Applied Nematology. 1993;16:501-513. DOI: 10.1016/S0169-5347(98)01583-3
  12. 12. Ferris H, Bongers T, De Goede RGM. A framework for soil food web diagnostics: Extension of nematode faunal analysis concept. Applied Soil Ecology. 2001;18:13-29
  13. 13. Hu J, Chen G, Hassan WM, Chen H, Li J, Du G. Fertilization influences the nematode community through changing the plant community in the Tibetan Plateau. European Journal of Soil Biology. 2017;78:7-16
  14. 14. Kudrin A. Effects of low quantities of added labile carbon on soil nematodes in intact forest soil microcosms. European Journal of Soil Biology. 2017;78:29-37. DOI: 10.1016/j.ejsobi.2016.11.002
  15. 15. Neher DA, Darby BJ. Computation and application of nematode community indices: General guidelines. In: Eyualem A, Traunspurger W, Andrassy I, editors. Freshwater Nematodes: Taxonomy and Ecology. Wallingford, UK: CAB International; 2006. pp. 211-222
  16. 16. Nethi S, Grewal PS, de Nardo EA, Stinner BR. Non-target effect of entomopathogenic nematodes on soil nematode community. Applied Soil Ecology. 2002;39:735-744
  17. 17. Schorpp Q, Schrader S. Dynamic of nematode communities in energy cropping systems. European Journal of Soil Biology. 2017;78:92-101. DOI: 10.1016/j.ejsobi.2016.12.002
  18. 18. Sohlenius B. Influence of cropping system and nitrogen input on soil fauna and microorganisms in a Swedish arable soil. Biology and Fertility of Soils. 1990;9:168-173
  19. 19. Yeates GW, Bongers T. Nematode diversity in agroecosystems. Agriculture Ecosystems and Environment. 1999;74:113-135
  20. 20. Zhao J, Li D, Fu S, He X, Fu Z, Zhang W, et al. Using the biomasses of soil nematode taxa as weighting factors for assessing soil food web conditions. Ecological Indicators. 2016;60:310-316. DOI: 10.1016/j.ecolind.2015.06.003
  21. 21. Neher DA, Wu J, Barbercheck ME, Anas O. Ecosystem type affects interpretation of soil nematode community measures. Applied Soil Ecology. 2005;30:47-64
  22. 22. Yeates GW. Soil nematodes in terrestrial ecosystems. Journal of Nematology. 1979;11:117-212
  23. 23. Freckman DW, Ettema CH. Assessing nematode communities in agroecosystem of varying human intervention. Agriculture, Ecosystems and Environment. 1993;45:239-261
  24. 24. Ito T, Araki M, Higashi T, Komatsuzaki M, Kaneko N, Ohta H. Responses of soil nematode community structure to soil carbon changes due to different tillage and cover crop management practices over a nine-year period in Kanto, Japan. Applied Soil Ecology. 2015;89:50-58. DOI: 10.1016/j.apsoil.2014.12.010
  25. 25. Zhao J, Neher D. Soil nematode genera that predict specific types of disturbance Appl. Soil Ecology. 2013;64:135-141
  26. 26. Rehfeld K, Sudhaus W. Vergleichende Untersuchung des Sexualverhaltens eines Zwillingsarten-Paares von Rhabditis (Nematoda) – Etude comparée du comportement sexuel de deux espèces sœurs de Rhabditis (Nematoda) – Comparative study of sexual behaviour of two sibling species of Rhabditis (Nematoda) Zoologische Jahrbucher. Abteilung für Systematik, Okologie und Geographie der Tiere. 1985;112(4):435-454
  27. 27. Dmowska E. Nematodes colonizing power plant ash dumps. I. Soil nematodes in ash dumps non-reclaimed and reclaimed by adding mineral soil and sowing grass. Polish Journal of Ecology. 2001;49:231-241
  28. 28. Háněl L. Succession of soil nematodes in pine forests on coal-mining sands near Cottbus, Germany. Applied Soil Ecology. 2001;16:23-34
  29. 29. Yeates GW, King KL. Soil nematodes as indicators of the effect of management on grasslands in the New England Tablelands (NSW): Comparison of native and improved grasslands. Pedobiologia. 1997;41:526-536
  30. 30. Pen-Mouratov S, Hu C, Hindin E, Steinberger Y. Soil microbial activity and a free-living nematode community in the playa and in the sandy biological crust of the Negev Desert. Biology and Fertility of Soils. 2011;47:363-375. DOI: 10.1007/s00374-011-0540-x
  31. 31. Liang W, Lavian I, Steinberger Y. Dynamics of nematode community composition in a potato field. Pedobiologia. 1999;43:459-469
  32. 32. Liang W, Lavian I, Steinberger Y. Effect of agriculture management on nematode communities in a Mediterranean agroecosystem. Journal of Nematology. 2001;33:208-213
  33. 33. Huston M. A general hypothesis of species diversity. American Naturalist. 1979;113:81-101. DOI: 10.1086/283366
  34. 34. Huston MA. Biological Diversity: The Coexistence of Species on Changing Landscapes. Cambridge: Cambridge University Press; 1994. 681 pp
  35. 35. MacArthur R, Wilson OE. The Theory of Island Biogeography. Princeton: Princeton University Press; 1967
  36. 36. Aronson RB, Precht WF. Landscape patterns of reef coral diversity: A test of the intermediate disturbance hypothesis. Journal of Experimental Marine Biology and Ecology. 1995;192:1-14. DOI: 10.1016/0022-0981(95)00052-S
  37. 37. Flöder S, Sommer U. Diversity in planktonic communities: An experimental test of the intermediate disturbance hypothesis. Limnology and Oceanography. 1999;44:1114-1119. DOI: 10.4319/lo.1999.44.4.1114
  38. 38. Huxham M, Roberts I, Bremner J. A field test of the intermediate disturbance hypothesis in the soft-bottom intertidal. International Review of Hydrobiology. 2000;85:379-394. DOI: 10.1002/1522-2632(200008)85:4<379::AID-IROH379>3.0.CO;2-X
  39. 39. Cramer MJ, Willig MR. Habitat heterogeneity, species diversity and null models. Oikos. 2005;108:209-218
  40. 40. Davidowitz G, Rosenzweig ML. The latitudinal gradient of species diversity among North American grasshoppers (Acrididae) within a single habitat: A test of the spatial heterogeneity hypothesis. Journal of Biogeography. 1998;25:553-560
  41. 41. French DD, Picozzi N. 'Functional Groups' of bird species, biodiversity and landscapes in Scotland. Journal of Biogeography. 2002;29:231-259. DOI: 10.1046/j.1365-2699.2002.00664.x
  42. 42. Hendrickx F, Maelfait JP, Van Wingerden W, Schweiger O, Speelmans M, Aviron S, et al. How landscape structure, land-use intensity and habitat diversity affect components of total arthropod diversity in agricultural landscapes. Journal of Applied Soil Ecology. 2007;44:340-351. DOI: 10.1111/j.1365-2664.2006.01270.x
  43. 43. Kerr JT, Packer L. Habitat heterogeneity as a determinant of mammal species richness in high-energy regions. Nature. 1997;385:252-254. DOI: 10.1038/385252a0
  44. 44. Tews J, Brose U, Grimm V, Tielborger K, Wichmann MC, Schwager M, et al. Animal species diversity driven by habitat heterogeneity/diversity: The importance of keystone structures. Journal of Biogeography. 2004;31:79-92

Written By

Mohammad Asif and P. Fazul Rahaman

Submitted: 30 September 2023 Reviewed: 02 October 2023 Published: 28 February 2024