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Impact of Human Activity

on Soil ArthropodsAlana Frampton

40100918

Environmental Biology

Dr Tancredi Caruso

Word Count: 3, 728

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1. Abstract

Soil arthropods carry out a wide range of functions including respiration and decomposition.

Anthropogenic disturbances alter soil biota communities, which results in the community

function changing. There are many factors that affect soil communities which can be

categorised into three main areas; agriculture, pollution and urbanisation. These categories will

be discussed in this review. Through agriculture, lower plant diversity can cause lower microbial

diversity, which lowers the diversity of the whole food web. High input management systems

can cause waterlogging and increased salinity, which reduces diversity. Invasive species can

lower abundance of soil arthropods. Moderate grazing and maintenance can increase

abundance and diversity through creating many microhabitats. Pollution in the form of

pesticides, herbicides, heavy metals and other sources causes a reduction in abundance and can

take many years to recover. However, it is suggested that toxic chemicals naturally become

unavailable to be absorbed by organisms over time, so the effects of these compounds could be

exaggerated. CO2 levels are continually increasing which is having a detrimental effect on soil

communities. During urbanisation many communities have changed due to loss of habitat and

invasive species. Green roofs and open gardens provide new habitats for soil arthropods. Some

urban area communities could benefit from increased temperatures and moderate CO2 levels.

This review clearly demonstrates that soil arthropods respond to all these factors. However,

there is a lack of research to have a mechanistic understanding of these responses. Future

research should focus on soil arthropods to better understand them, so that they could be used

as soil health indicators.

2. Introduction

Nutrient rich and well structured soil leads to the growth of healthy crops (Anwar et al., 2014).

Nitrogen, Phosphorus and Potassium are among those nutrients essential for increased crop yield

(Parmar, 2014). Poor soil fertility and higher numbers of weeds are linked and contributes to

lower production (Birkett et al., 2014). Pea-barley intercrops in Western Europe suppressed the

weeds more effectively when there was higher soil Nitrogen availability: the increased leaf areas

made them more competitive for light than the weeds (Ambus et al., 2011).

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Soil arthropods assist to form productive nutrient rich soil (Eggleton & Stork, 1992). Bottinelli et

al. (2015) discuss how earthworms, ants and termites are good indicators of soil quality and how

these macro-invertebrates produce and maintain biostructures that contribute to the variety and

health of the soil. Decomposer biota including earthworms are considered to be vital to maintain

the agricultural ecosystems in the tropics through altering the soil (Beare, 1997). These larger soil

fauna stabilize the soil structure, the linings of biopores and smaller aggregates (Oades, 1993).

However, to help to incorporate the nutrients needed into the soil, the smaller soil fauna are also

an essential part of the ecosystem (Caruso, 2015).

These smaller biota break down leaf litter and other debris which transforms the nutrients into a

form that is easily used by plants (Capinera, 2008). The abundance and species richness as part of

a healthy ecosystem can be illustrated in a small way through Oribatid mites. This is explained by

Coleman & Hansen (1998) through an experiment on a forested area in the North Carolina

mountains in the USA. Plots of pure birch, oak and maple litter were named simple litter

treatments. Plots of these three leaf litters and seven other leaf litters mixed were the complex

litter treatments. The complex litter treatments contained significantly more species of Oribatid

mites and a significantly greater variety of microhabitats than the simple litter treatments. This is

shown in Figure 1.

Microhabitat Variety among Litter Treatments at Four Depths along a 5cm Transect

Figure 1 (Coleman & Hansen, 1998)

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The activities of humans are changing the soil ecosystems (Bender et al., 2014). Agriculture,

pollution and urbanisation are three factors that show a clear change in the soil biota

communities. When the soil environment is changed, the soil biota have to adapt to new roles to

survive (Mellino & Ulgiati, 2015) (Coviella et al., 2015). However, for many species the changes

are too rapid to adapt to and the ecosystem is out of balance (Akrami et al., 2015). This review

discusses how these three human activities affect the soil and how this influences the soil

communities.

3. Humans Activities Impacting Soil Arthropods

3.1 Agriculture

Large areas of crops contain lower densities of soil arthropods than in overgrown and natural

areas. Microbial biomass is higher in overgrown areas. Microbes are consumed by nematodes

which are in turn consumed by soil micro-arthropods. Fewer microbes in a food web leads to the

reduction of micro-arthropods and meso-arthropods since they are an essential food source

(Heidemann et al., 2014; Frouz et al., 2015).

To elaborate, microbe biomass can be reduced when agriculture causes lower plant diversity.

Grandy et al. (2014) states that the soil microbe biomass increased significantly when a

monoculture was rotated with another crop or an additional crop was introduced to the

monoculture. This would affect the organisms higher in the food chain such as the Oribatid mites

and Collembola. In addition, the management method can effect soil arthropods as well as the

plant variety.

Low input agriculture management methods conserve soil biodiversity. High input methods (large

amounts of fertilisers and environmental disturbance) reduce diversity. Bardgett & Cook (1998)

observe that high input methods encourage bacterial decomposition. From the previous

discussion it is logical to assume that this would increase the abundance of the nematodes,

therefore, the Oribatid mites and other groups would increase in numbers. However, it was

discovered that is not the case. Increase in bacteria increases the abundance of other

opportunistic bacterial feeding organisms. These compete with other groups and reduce

biodiversity.

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Livestock and defoliation are specific aspects of high input methods that can have a significant

effect on the soil biota. As aerobic organisms, well aerated soil is the most suitable habitat.

Vegetation cover is needed for the food chain to be healthy and productive. When large grazers

trample the ground and reduce vegetation cover, the soil becomes waterlogged. Soil salinity is

increased by defoliation and increased to greater levels by defoliation and compaction. When

waterlogged, micro-arthropods showed a fourfold decrease in abundance and compaction caused

smaller average body size (Bakker et al., 2015). The anaerobic decomposition of large quantities of

fertilisers could reduce the oxygen levels of the soil. This could be another reason why the

Oribatid mites could not compete with the opportunistic fauna discussed previously. Excessive

fertilisation causes waterlogged soil which increases the crop production for the short term but

damages the soil communities and the soil itself. This leads to soil not being able to sustain itself

and continued damage to the soil arthropods (Chen et al., 2009).

However, moderate grazing and trampling by livestock can increase the number of microhabitats

which would increase the abundance and diversity of soil arthropod species. A study by Benoist et

al. (2015) stated that moderate cattle grazing increased the variety of soil and vegetation. The

impact of grazing was sampled at three different levels of cattle pressure; low, medium and high.

Invasive species can be encouraged through agricultural disturbance, this affects the species

richness and abundance of soil arthropods. Forty to fifty percent of available land has been

converted to agricultural and urban areas. This has influenced some species to utilise these

habitats and outcompete other species. The level of vulnerability to invasive species is usually

affected more by the characteristics of the native and invasive species than the species richness

and availability of vacant niches. Disturbance and other factors are affected by humans and

increase the invasion of exotic species. Figure 2 displays how humans interact with the

environment (Chapin FS et al., 2000). When Bromus tectorum was introduced to native perennial

grass (Hilaria jamesii) the abundance of nematodes and microarthropods dropped significantly

(Belnap et al., 2005). Anthropogenic changes through agriculture alter the food chain completely

and invasive species further alter the food chain.

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Humans Interacting with the Ecosystem

Cattle grazing and invasive species can be linked. The disturbance can encourage new species to

invade the habitat and the two factors combined alter the soil community. Borer et al. (2015)

states that the biovolume of arthropods was 79% higher in the grazed plots. However, predatory

arthropods were 13% higher in the ungrazed plots. Where non-native grasses were grazed, the

volume of the arthropods increased as native plants replaced them. This displays how many

factors are linked through agriculture to affect soil arthropods.

Agriculture would benefit from soil arthropods properly carrying out their natural functions.

Neher (1999) states that these organisms are essential in many ecosystem functions such as

decomposition; maintaining stability of soil; increasing plant productivity; enhancing water

relations. If management methods more closely resembled natural ecosystems then the soil

community could become stable and healthier crops would result (Abawi & Widmer, 2000).

Bender & Heijden (2015) studied how soil biota affected nutrient-use efficiency, nutrient leaching

and plant performance. Crop rotation caused a significant increase in the crop yield, Nitrogen and

Phosphorus uptakes and reduced leaching losses of Nitrogen. The crop rotation had increased the

soil life which led to these results.

3.2 Pollution

Figure 2 (Chapin FS et al., 2000)

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Chemicals can influence the species richness and abundance of soil inhabiting mites. An example

is when they are removed by pesticides as non-target organisms (European Commission, 2013;

Bauda et al., 2015). When two biofungicides were applied to plots on fields of cucumber in Egypt,

the community composition of the Oribatid mite community changed. Some species numbers

decreased while another species numbers increased resulting in the same overall total abundance

of mites as before the experiment. Many Oribatid mites are fungal feeders so these species would

decrease as their food source is removed by the biofungicides (Ageba et al., 2014).

In addition, some groups such as the Oribatid mites recover more slowly than Collembola from

pollutants including fertilisers and pesticides. They have lower fecundity rates, slower

development and lower metabolic rates than the Collembola. Due to this life history they cannot

respond rapidly when large numbers are removed from the population. Astigmata have a very

different life history which allows them to respond quickly after exposure to toxic compounds

(Behan-Pelletier, 1999). This demonstrates that the soil community does not return to its original

condition swiftly since some species need more time, therefore, the soil ecosystem needs time to

function properly again. Malkomes & Wohler (1983) explains that inhibitory effects lasted several

months under laboratory conditions.

To add, Stegeman (1964) stated that Collembola took longer than mites to recover from treatment

of the Carbamate Insecticide Cararyl. As in the previously discussed experiment, it took several

months for the soil community to recover based on the Collembola and mite data. Despite a

couple of these experiments being older the biota responds the same way to chemicals now as in

the past.

Pollution can come from unexpected sources, for example, olive mill wastewater. The waste

water is applied to the soil in olive orchards. It was discovered that Oribatida populations were

restricted and this resulted in a community shift. The abundance of Collembola increased (Bruhl

et al., 2015). The diversity of pollutants and species causes each study to produce different

results. However, the overall picture displays that the soil communities are changed.

Other groups are affected by pollution apart from Oribatid mites and Collembola, Enchytraeid

communities were confirmed to be reduced in species richness and abundance through heavy

metal pollution (Zinc and Lead). The data was compared to results from unpolluted areas and the

abundance of Enchyraeid species was generally lower in the polluted areas as shown in Figure 3.

The closer to the smelter, the less healthy the community structure was. Interestingly,

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Enchytraeids are not affected by the natural soil properties but were affected by soil pH (Kapusta

& Sobczyk, 2015). Other arthropods such as Collembola are more sensitive to pH than the

pollutant. Folsomia candida Collembola were proven to react to the acidity of the soil and not the

Lead. Collembolan mortality, avoidance and lower reproduction were all significantly greater in

acidic soil (Gestel et al., 2014a).

Abundance and Diversity of Enchytraeids Decreases during Heavy Metal Pollution

However, it is suggested that the effect of toxic compounds on soil biota is exaggerated. As

organic compounds age they become less available to be absorbed by organisms. Therefore, their

toxic effects reduce over time (Alexander M, 2000). In previous studies this has not been

discussed. It is possible that the effect of pollution on these organisms is not as severe as at first

appearance.

Other factors combined with pollution can have different effects on soil biota. On a permanent

grassland the effects of fertilizer addition (Nitrogen, phosphorus and Potassium), cutting frequency

and herbicides on soil microorganisms were measured. Microbial biomass decreased when cut

once a year but not three times a year when fertilizer was applied during both experiments. This

suggests that cutting reduces the negative effect of applying fertilisers. Earthworms and beetles

increased in abundance when fertiliser was applied but grass cutting was detrimental. This shows

that polluting substances can be detrimental but the effects are complicated when comparing

short and long term effects and other management processes (Lemanski & Scheu, 2015).

Pollutants can be absorbed from the atmosphere, they are not always toxic chemicals or

fertilisers. CO2 levels will become more available to soil biota over time since CO2 levels are

Figure 3 (Chapin FS et al., 2000)

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continually increasing. In a mesocosm study involving Collembola, Oribatid mites (Acari) and

Enchytraeidae, the organisms were collected after five and eleven weeks as the CO2 levels

increased. The CO2 increase did not affect the plants but the predatory mite and two Collembola

species reduced in abundance (Audisio et al., 2015).

Cold CO2 producing gas vents provide the ideal environment to research how soil biota will

respond to the continuing increase in CO2. The top few centimetres of the soil contained

microhabitats of 100% CO2 and <2% CO2. Collembola and nematodes declined in abundance as

CO2 levels increased. However, nematode densities were not affected until 62% CO2 levels.

Collembola developed viable populations up until 20% CO2. The mofettophilous species had

denser populations in the higher CO2 areas due to either a lack of competition or a more suitable

food supply. This suggests that as atmospheric CO2 levels increase, the soil microarthropod

abundance and species diversity will decrease and be replaced by others (Balkenhol et al., 2015).

3.3 Urbanisation

Urbanisation creates the ideal physical environment to fulfil the needs of humans only. The

species that are able to adapt well to urban environments are multiplying since they are able to

outcompete other species (McKinney, 2006). It was expected that the soil community

composition would change following the same logic as previously discussed. However, a study in

Italy disagrees. Urban and suburban holm oak in a small city and a large city were sampled. There

was no significant difference between anthropogenic factors resulting from urbanisation, and

arthropod density and variety. This data is displayed in Table 1 (Agamennone et al., 2014).

Over time biodiversity could show different results. A study specifically examining Collembola

community composition displayed differences over time. Native species diversity reduced, the lost

species being replaced by more generalised and invasive species. This resulted in a species poor

community. The more resistant Collembola species persist and less stressed urban grass areas

contain more diversity. The changes over time suggest that the response is influenced by multiple

processes. This information is displayed in Figure 4 (Rzeszowski & Sterzyriska, 2015).

Collembola Species Accumulation for Whole Warsaw Area using Old (Longer Time after

Urbanisation) and New (Shorter Time after Urbanisation) Data

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Interestingly, in urban areas new environments are being created that soil communities are taking

possession of. In Germany green roofs were sampled during 2002, ‘old roofs’ were built between

1990 and 1994, ‘young roofs’ were built between 1998 and 1999. In ‘old roofs’ there was

improved niche specialisation of the collembolan and a more stable community. ‘Young roofs’ had

less advanced soil formation and the soil arthropod community was not as stable (Boning &

Schrader, 2006). This shows that while habitats are being removed by urbanisation, new ones are

being created within urban areas as the environment adapts. However, the number of species

was very similar in the ‘old’ and ‘new roofs which suggests that the communities will not develop

more than they already have. Even if green roofing was extensive it would still not replace the

natural environments.

In Naples, Italy it was discovered that agriculture has more of an effect on soil properties than

urbanisation. It is possible that it is more important to create new habitats in agricultural areas

than urban areas. The agricultural soil was dominated by very few species, mainly Proisotoma

minuta and Entomobrya multifasciata. These two species are adapted to environments of high

disturbance. The urban soil community more closely resembled the forest community (Cortet et

al., 2015). These results could vary from country to country but this evidence suggests that the

level of disturbance in an urbanised area will affect how much the soil community changes. It is

Figure 4 (Rzeszowski & Sterzyriska, 2015)

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possible that the level of pollution in urban areas is what reduces the species diversity and

abundance in urban areas. Another study in Naples, Italy stated that collembolan density

decreased as pollution increased. The higher the organic content of the soil the greater the

species richness and diversity (Arena et al., 2014). This provides more evidence that the

urbanisation itself does not cause changes in soil biota but the human activities in those areas do

change the community.

How human activities within urbanisation change soil communities is emphasised through

different types of soil communities being found in different types of garden. Research was

conducted on lawns, open beds and closed canopy beds in Dunedin, New Zealand. Collembola

were most frequently found in lawns. Detritivores were found in higher numbers in gardens that

had experienced no pesticides (Barratt et al., 2015). Considering studies discussed previously, it

suggests that lawn soils were less disturbed than beds that would be turned over regularly for

maintenance of the plants.

To add, some soil biota could benefit from urbanisation due to plants benefiting. One paper

highlights that within urbanised areas increased temperatures and high CO2 levels increase the

productivity of plants (Vodyanitskii, 2015). It has been previously discussed that increased soil CO2

levels are detrimental to the soil community. However, if the CO2 was not too high and used

primarily by the plants without large amounts being absorbed by the soil, the soil community

could resemble forest arthropod communities.

3.4 Additional Discussion

It is difficult to predict how these organisms respond to human activity. One paper states that due

to Oribatid mites having generalised diets, they are less affected by environmental changes than

other groups. These mites do not suffer from a lack of leaf litter because it only contributes to

22% of their diet (Gan et al., 2014). However, there are many more factors affecting this group

and it does not explore any of the other arthropods.

An example of one of these factors could be the introduction of invasive species. Acacia dealbata

(Australian tree legume) produces allelopathic compounds that reduce microbial biomass and

reduce the diversity of the native plants around it. The arthropods reacted differently depending

on the ecosystem (Lorenzo et al., 2013). This further displays that many factors affect them

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depending on how anthropogenic activity alters the environment, plant species and other biota

surrounding them.

Gestel et al. (2014b) emphasises the importance of the time of year of sampling. In the autumn

higher temperatures and lower precipitation were recorded than in the spring. In both seasons

the most abundant taxa were Collembola and Acarina but the relative abundances were affected

differently by seasonality and the metal contamination in the area. Therefore, for accurate future

sampling, time needs to be taken into consideration.

Urbanisation and pollution are closely linked while affecting soil arthropods over time. One study

states that there was no significant difference between the toxicity of soil in different forested

areas except in different seasons. Different forested areas were sampled along an urbanisation

gradient and are displayed in Figure 3. Significantly higher levels of pollutants were recorded in

the autumn than in the spring (Baranyai et al., 2016). This provides more evidence suggesting that

sampling over time is a good idea for future studies.

Areas of Forest Sampled Along an Urbanisation Gradient in Debrecen in Hungary

Figure 5 (Baranyai et al., 2016)

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Mechanistic processes could be studied by observing how soil arthropods are affected over time.

Therefore, dynamical studies would give more insight into the processes within soil arthropod

communities. In Egypt two fields were sampled from October to March 2015. The effects of

changing from conventional to organic management were studied. Over time, the organic matter

built up which increased Oribatid mite abundance (Al-Assiuty et al., 2015). If these fields had not

been sampled over a period of time, the relationship between organic matter and mites would not

have been observed.

4. Conclusion

Most studies have been conducted at the community level. Due to all the complexities discussed

above, it is difficult to obtain a mechanistic understanding of processes at that level. Integrating

studies of species, populations and communities would provide useful information on how

different factors affect soil arthropods.

A useful method to distinguish between species, population and community level effects would be

to examine studies on ecotoxicology that focus on soil invertebrate species. Gestel et al. (2012)

examined the quality of urban soil through bioassays with Eisenia Andrei, Folsomia candida and

Enchytraeus crypticus. Metal bioaccumulation was the highest in E. crypticus and was more

sensitive than the other two species. Evidence suggested that the reasons for this species being

more sensitive was related to soil properties and the metal contamination. Therefore, this

information can be used to assess how this species interacts with the ecosystem and provides a

mechanistic understanding.

If the factors affecting soil biota could be more clearly understood then it would be a useful way to

measure the health of soil. In 1998 a conference was held entitled ‘Soil Health: Managing the

Biological Component of Soil Quality’ to emphasise how important soil organisms are as measures

of soil quality and health. This could assist in the understanding of how to manage soil properly so

it can function properly and humans can utilise it in a sustainable way (Doran & Zeiss, 2000).

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