Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review

*Sigmund Hågvar*

## **Abstract**

Long-term studies on mites in Norwegian coniferous forests are summarized. In podzol soil with raw humus, mite densities could pass 1 million per m2 , with 48 species of Oribatida and 12 species of Mesostigmata. Field and laboratory experiments with liming and artificial acid rain showed that soil pH affected the structure of the mite community. Certain species of mites and springtails typical for acid soils did, however, show preference for a higher pH in monoculture. We hypothesized that competition could be a strong regulating factor in microarthropod communities. Several oribatid species were flexible regarding soil type, vegetation, substrate, and decomposition stage. The genus *Carabodes* showed examples on specialists: two species were grazers on *Cladonia* lichens in dry pine forests, while three were decomposers in dead polypore fungi. Another three oribatid species from different genera were unique in excavating spruce needles, producing slowly decomposing excrements, and probably contributing to stable, carbon-storing humus. In microcosms, predatory Gamasina mites were seen to regulate microarthropod numbers. Mites were able to adjust both their vertical and horizontal distribution in soil according to environmental change. A local and temporary burst of fungal activity could rapidly attract opportunistic fungal feeders. Several mites were active under snow, often feeding. Some even penetrated into the snow layer.

**Keywords:** Acari, coniferous forest, ecology, mites, Norway, Oribatida, review, soil pH

## **1. Introduction**

Nowhere else, in nature, organisms are so densely packed as in soil. Combined with a huge number of species, "biodiversity in the dark" has fascinated biologists for long. In concert, soil organisms play a key role in terrestrial ecosystems, being of fundamental importance for plant growth, sustainable crop production, and biogeochemical cycling of nutrients. At the same time, soil biodiversity is vulnerable to human disturbance of different kinds. There is a critical need for understanding soil processes, how soil organisms respond to global change, and to take measures for long-term protection of soil biodiversity [1].

Mites (Acari) represent one of the species rich and abundant soil animal groups. Oribatid mites alone cover five feeding guilds, including the ability to digest chitin [2], and they represent four trophic levels in the decomposition process [3].

Another mite group, Mesostigmata, contains a multitude of predator species which control other microarthropod populations, both in the soil and in vegetation [4, 5]. Forest habitats, especially old forests with a well-developed litter layer, tend to have a high mite density, often with a species-rich fauna of oribatids [6–8].

Norwegian coniferous forests represent the western outpost of the Eurasian taiga. This giant forest belt, which is dominated by Norway spruce (*Picea abies* (L.) H. Karst.) and Scots pine (*Pinus silvestris* L.), typically contains a well-developed raw humus layer which represents a considerable global carbon storage. The slowly decomposing needles, cones, and other litter items in the forest floor create a fungus rich and sometimes deep, humus world, in which several mite groups thrive, including many oribatid species.

The present review is a synthesis of mite studies in coniferous forest soils of Southern Norway, published over a 40-year period [9–26]. In the 1970s, extensive studies on soil microarthropods were initiated as part of a large project, "Effects of acid precipitation on forest and soil," and certain subjects were followed up long after the project was ended. In addition to summing up field and laboratory experiments with liming and artificial acid rain, spotlights will be given on the following topics: density and species numbers of mites, their horizontal and vertical distribution, effects of different pH, vegetation types, soils and substrates, succession in the mite fauna during decomposition, whether mites can influence the humification process, how species within one genus may differ in habitat use, an experiment on the predatory effect of Gamasina mites, and mite activity beneath and within snow.

## **2. Material and methods**

## **2.1 Study areas**

## *2.1.1 Main study area: Nordmoen*

This was a spruce forest with *Vaccinium myrtillus* L. vegetation, situated on a flat plain of glaciofluvial sandy deposits, about 45 km N of Oslo. On clearcut areas, *Deschampsia flexuosa* (L.) dominated*.* The soil was a stone-free iron podzol with a 3 cm thick organic layer and a correspondingly bleached layer below. Experiments with artificial acidification and liming and decomposition experiments with litter bags were performed here, partly in a young spruce stand, and partly on a clearcut area [14].

## *2.1.2 Two study areas covering the range of coniferous forest types: Ås (A) and Skrukkelia (B)*

Two study areas were chosen for soil sampling in natural forest, each area with a gradient in vegetation types from the poorest pine forest to the richest spruce forest [15, 27]. Area A near Ås, about 30 km south of Oslo, had a cover of marine sediments. In area B in Skrukkelia, NW of lake Hurdalssjøen and about 60 km north of Oslo, the soil was mainly morainic deposits. In both study areas, spruce forest with *Vaccinium myrtillus* dominated. Listed after increasing soil fertility based on plant associations, the vegetation types were short named as follows:

1.*Cladonia* sp.: pine forest on iron podzol soil, with a dense cover of *Cladonia* lichens. Due to a thin soil layer, conditions were dry, and trees grew slowly (**Figure 1**).

**83**

**Figure 1.**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

2.*Calluna vulgaris:* pine forest with less *Cladonia,* and a field layer dominated by *Calluna vulgaris* (L.) Hull. The soil was shallow peat in area A and iron podzol

3.*Vaccinium sp.:* pine forest on iron podzol soil, with a dense cover of *Vaccinium myrtillus* or *Vaccinium vitis-idaea* L.*,* but also containing some *Cladonia* lichens.

4.*Vaccinium myrtillus:* spruce forest with *Vaccinium myrtillus*. Brown earth-like

5. *Small ferns:* spruce forest with small ferns, *Dryopteris phegopteris* (L.) C. Chr. and *Dryopteris linnaeana* C. Chr. Brown earth in area A and iron podzol in area B.

6.*Small herbs:* spruce forest on brown earth, with small herbs like *Carex digitata*

7.*Tall herbs:* spruce forest on brown earth, with tall herbs like *Filipendula ulmaria* (L.) Maxim., *Athyrium filix-femina* (L.) Roth., and *Aconitum septentrionale* Koelle.

Dead sporocarps of different wood-living polypore fungi were sampled in an old

Activity under snow was studied in an old spruce forest with *Vaccinium myrtillus* vegetation near Veggli in Numedal valley, about 150 km NW of Oslo. Here, at 850 m above the sea level, a snow cover of 1–2 m is common [23]. In the main study area,

Each vegetation type in areas A and B was sampled twice, in autumn 1977

*The poorest coniferous forest type: slow-growing pines on a thin soil layer dominated by Cladonia lichens.* 

*Certain drought-tolerant, lichen-feeding mites were abundant here. Photo: S. Hågvar.*

, 20 soil cores were taken both

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

soil in area A and iron podzol in area B.

*2.1.3 Study area for mites in decomposing sporocarps*

*2.1.4 Study areas for mite activity under and within snow*

and in spring 1978. Using a soil corer of 10 cm<sup>2</sup>

**2.2 Methods for field studies**

*2.2.1 Soil sampling*

L., *Melampyrum silvaticum* L., and *Fragaria vesca* L.

spruce forest in the Østmarka area, about 20 km east of Oslo [24, 25].

Nordmoen, mite activity was studied both under and within snow [11].

in area B.

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*


Dead sporocarps of different wood-living polypore fungi were sampled in an old spruce forest in the Østmarka area, about 20 km east of Oslo [24, 25].

#### *2.1.4 Study areas for mite activity under and within snow*

Activity under snow was studied in an old spruce forest with *Vaccinium myrtillus* vegetation near Veggli in Numedal valley, about 150 km NW of Oslo. Here, at 850 m above the sea level, a snow cover of 1–2 m is common [23]. In the main study area, Nordmoen, mite activity was studied both under and within snow [11].

## **2.2 Methods for field studies**

#### *2.2.1 Soil sampling*

*Pests Control and Acarology*

including many oribatid species.

and within snow.

**2.1 Study areas**

area [14].

*Skrukkelia (B)*

(**Figure 1**).

**2. Material and methods**

*2.1.1 Main study area: Nordmoen*

Another mite group, Mesostigmata, contains a multitude of predator species which control other microarthropod populations, both in the soil and in vegetation [4, 5]. Forest habitats, especially old forests with a well-developed litter layer, tend to have

Norwegian coniferous forests represent the western outpost of the Eurasian taiga. This giant forest belt, which is dominated by Norway spruce (*Picea abies* (L.) H. Karst.) and Scots pine (*Pinus silvestris* L.), typically contains a well-developed raw humus layer which represents a considerable global carbon storage. The slowly decomposing needles, cones, and other litter items in the forest floor create a fungus rich and sometimes deep, humus world, in which several mite groups thrive,

The present review is a synthesis of mite studies in coniferous forest soils of Southern Norway, published over a 40-year period [9–26]. In the 1970s, extensive studies on soil microarthropods were initiated as part of a large project, "Effects of acid precipitation on forest and soil," and certain subjects were followed up long after the project was ended. In addition to summing up field and laboratory experiments with liming and artificial acid rain, spotlights will be given on the following topics: density and species numbers of mites, their horizontal and vertical distribution, effects of different pH, vegetation types, soils and substrates, succession in the mite fauna during decomposition, whether mites can influence the humification process, how species within one genus may differ in habitat use, an experiment on the predatory effect of Gamasina mites, and mite activity beneath

This was a spruce forest with *Vaccinium myrtillus* L. vegetation, situated on a flat plain of glaciofluvial sandy deposits, about 45 km N of Oslo. On clearcut areas, *Deschampsia flexuosa* (L.) dominated*.* The soil was a stone-free iron podzol with a 3 cm thick organic layer and a correspondingly bleached layer below. Experiments with artificial acidification and liming and decomposition experiments with litter bags were performed here, partly in a young spruce stand, and partly on a clearcut

Two study areas were chosen for soil sampling in natural forest, each area with a gradient in vegetation types from the poorest pine forest to the richest spruce forest [15, 27]. Area A near Ås, about 30 km south of Oslo, had a cover of marine sediments. In area B in Skrukkelia, NW of lake Hurdalssjøen and about 60 km north of Oslo, the soil was mainly morainic deposits. In both study areas, spruce forest with *Vaccinium myrtillus* dominated. Listed after increasing soil fertility based on plant

1.*Cladonia* sp.: pine forest on iron podzol soil, with a dense cover of *Cladonia* lichens. Due to a thin soil layer, conditions were dry, and trees grew slowly

*2.1.2 Two study areas covering the range of coniferous forest types: Ås (A) and* 

associations, the vegetation types were short named as follows:

a high mite density, often with a species-rich fauna of oribatids [6–8].

**82**

Each vegetation type in areas A and B was sampled twice, in autumn 1977 and in spring 1978. Using a soil corer of 10 cm<sup>2</sup> , 20 soil cores were taken both

#### **Figure 1.**

*The poorest coniferous forest type: slow-growing pines on a thin soil layer dominated by Cladonia lichens. Certain drought-tolerant, lichen-feeding mites were abundant here. Photo: S. Hågvar.*

during spring and autumn in each vegetation type. The cores were divided into 0–3 and 3–6 cm depth. In the main study area at Nordmoen, the same sampling method was used. Here, a clearcut area with 0.5 m high *Picea abies* seedlings was chosen for intense studies. Eight random replicates were established, each 4 × 4 m. Density of mites per replicate was based on 10 soil cores, each 5.3 cm<sup>2</sup> and 6 cm deep.

## *2.2.2 Artificial acidification and liming*

Lime was applied as crushed CaCO3 (3000 kg Ca0 ha<sup>−</sup><sup>1</sup> ), and 50 mm of artificial acid rain was applied monthly by adding sulfuric acid to ground water (**Figure 2**). Treatments were no watering, pH 6 (control), pH 4, pH 3, pH 2.5, and pH 2. The natural pH in the organic layer (upper 3 cm) was 3.9. Liming increased pH about 2 units, and the strongest acid reduced pH about 0.5 units. Only application of acid rain with pH 3 or stronger lowered the pH in the organic layer [14].

## *2.2.3 Litter bag studies on succession*

The clearcut area in the main study area was used to study the mite succession during decomposition of spruce needles [19] and birch leaves [12, 13]. Cylindrical litter bags, 3 cm high and with a diameter of 3.4 cm, were filled with 4.2 g (dry weight) of naturally shed spruce needles. The litter bags were then inserted into holes made in the raw humus layer, which had a corresponding depth. This is not a natural position of the litter, but it allowed to study the preference among mites for different decomposition phases. While the litter bags stood in this fixed position, in contact with various depth levels of the organic horizon, all species had a continuous access to the needles. With a mesh size of 0.6 mm, migration to and from the bags was easy for all microarthropods.

Succession in decomposing birch leaves was studied in a similar way in the same site. Cylindrical litter bags with a mesh size of 1 mm, 3 cm high and with a diameter of 6.5 cm, were each filled with 6.85 g (dry weight) of naturally shed birch leaves. These bags also received artificial rain of pH 6, 4, 3, and 2.

**85**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

In the main study area at Nordmoen, naturally shed spruce needles were sampled on snow and dried. Later, needles were stuck into fine-meshed nylon strips, which were placed on the ground of a 10–20 m high spruce stand. Gradually, needles were covered by new litter in a natural way. Strips with needles were recov-

Dead sporocarps were brought to the laboratory, carefully fragmented, and

Specially designed pitfall traps were used [23]. The mechanism allowed sam-

This was a greenhouse experiment, where forest soil was kept in large plastic boxes [10]. Microarthropods (and microflora as well) had the opportunity to colonize sterilized soil (raw humus, poor mull, and rich mull) which had been adjusted to three different pH levels. Cylindrical litter bags with a mesh size of 1 mm, 3 cm high and with a diameter of 6.5 cm, were used. The design can be characterized as a preference experiment, where also the ability to reproduce during the four-month

Small microcosms were used, consisting of a cylindrical, open litter bag which was inserted into a lidded plastic container. The litter bag was 3 cm high, 3.4 cm in diameter, and made from a nylon cloth with 0.6 mm mesh size. Holes drilled in the plastic container were covered with nylon cloth of 5-μm mesh size. Before adding microarthropods to sterilized soil, microflora was introduced partly by soil water sieved through 5-μm pores and partly by allowing soil fungi to grow in through corresponding pores for 1–2 months. Then animals were added, either from monocultures or from ordinary soil samples [17]. Raw humus adjusted to different pH levels was used in the microcosms. About 25 microcosms were extracted after 3, 6, and 12 months, respectively. This setup allowed for studying the effect of soil pH on population growth in monocultures of selected species. An interesting by-product was the effect of predatory Gamasina mites, which survived in some microcosms,

**3.1 The coniferous forest floor: a high density and species rich habitat for mites**

Podzol soil with vegetation type 4 in the main study area contained 48 species of Oribatida and 12 species of Mesostigmata (**Table 1**). The density of mites was high.

mites were extracted in funnels, using heat from a light bulb [24, 25].

pling without disturbing the subnivean air space near the traps.

period influenced the establishment of each species.

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

*2.2.5 Sporocarp sampling and extraction*

*2.2.6 Sampling mites active under snow*

**2.3 Methods for laboratory studies**

*2.3.1 A "preference" experiment*

*2.3.2 Microcosm studies*

but went extinct in others [21].

**3. Results and discussion**

*2.2.4 Mites living inside decomposing spruce needles*

ered after 4, 12, 16, 24, 35, 38, 40, and 52 months [22].

**Figure 2.** *Artificial acid rain is applied on a 4 × 4 m experimental plot with small pine trees.*

## *2.2.4 Mites living inside decomposing spruce needles*

In the main study area at Nordmoen, naturally shed spruce needles were sampled on snow and dried. Later, needles were stuck into fine-meshed nylon strips, which were placed on the ground of a 10–20 m high spruce stand. Gradually, needles were covered by new litter in a natural way. Strips with needles were recovered after 4, 12, 16, 24, 35, 38, 40, and 52 months [22].

## *2.2.5 Sporocarp sampling and extraction*

*Pests Control and Acarology*

and 6 cm deep.

*2.2.2 Artificial acidification and liming*

*2.2.3 Litter bag studies on succession*

bags was easy for all microarthropods.

These bags also received artificial rain of pH 6, 4, 3, and 2.

*Artificial acid rain is applied on a 4 × 4 m experimental plot with small pine trees.*

Lime was applied as crushed CaCO3 (3000 kg Ca0 ha<sup>−</sup><sup>1</sup>

rain with pH 3 or stronger lowered the pH in the organic layer [14].

during spring and autumn in each vegetation type. The cores were divided into 0–3 and 3–6 cm depth. In the main study area at Nordmoen, the same sampling method was used. Here, a clearcut area with 0.5 m high *Picea abies* seedlings was chosen for intense studies. Eight random replicates were established, each 4 × 4 m. Density of mites per replicate was based on 10 soil cores, each 5.3 cm<sup>2</sup>

acid rain was applied monthly by adding sulfuric acid to ground water (**Figure 2**). Treatments were no watering, pH 6 (control), pH 4, pH 3, pH 2.5, and pH 2. The natural pH in the organic layer (upper 3 cm) was 3.9. Liming increased pH about 2 units, and the strongest acid reduced pH about 0.5 units. Only application of acid

The clearcut area in the main study area was used to study the mite succession during decomposition of spruce needles [19] and birch leaves [12, 13]. Cylindrical litter bags, 3 cm high and with a diameter of 3.4 cm, were filled with 4.2 g (dry weight) of naturally shed spruce needles. The litter bags were then inserted into holes made in the raw humus layer, which had a corresponding depth. This is not a natural position of the litter, but it allowed to study the preference among mites for different decomposition phases. While the litter bags stood in this fixed position, in contact with various depth levels of the organic horizon, all species had a continuous access to the needles. With a mesh size of 0.6 mm, migration to and from the

Succession in decomposing birch leaves was studied in a similar way in the same site. Cylindrical litter bags with a mesh size of 1 mm, 3 cm high and with a diameter of 6.5 cm, were each filled with 6.85 g (dry weight) of naturally shed birch leaves.

), and 50 mm of artificial

**84**

**Figure 2.**

Dead sporocarps were brought to the laboratory, carefully fragmented, and mites were extracted in funnels, using heat from a light bulb [24, 25].

## *2.2.6 Sampling mites active under snow*

Specially designed pitfall traps were used [23]. The mechanism allowed sampling without disturbing the subnivean air space near the traps.

## **2.3 Methods for laboratory studies**

## *2.3.1 A "preference" experiment*

This was a greenhouse experiment, where forest soil was kept in large plastic boxes [10]. Microarthropods (and microflora as well) had the opportunity to colonize sterilized soil (raw humus, poor mull, and rich mull) which had been adjusted to three different pH levels. Cylindrical litter bags with a mesh size of 1 mm, 3 cm high and with a diameter of 6.5 cm, were used. The design can be characterized as a preference experiment, where also the ability to reproduce during the four-month period influenced the establishment of each species.

## *2.3.2 Microcosm studies*

Small microcosms were used, consisting of a cylindrical, open litter bag which was inserted into a lidded plastic container. The litter bag was 3 cm high, 3.4 cm in diameter, and made from a nylon cloth with 0.6 mm mesh size. Holes drilled in the plastic container were covered with nylon cloth of 5-μm mesh size. Before adding microarthropods to sterilized soil, microflora was introduced partly by soil water sieved through 5-μm pores and partly by allowing soil fungi to grow in through corresponding pores for 1–2 months. Then animals were added, either from monocultures or from ordinary soil samples [17]. Raw humus adjusted to different pH levels was used in the microcosms. About 25 microcosms were extracted after 3, 6, and 12 months, respectively. This setup allowed for studying the effect of soil pH on population growth in monocultures of selected species. An interesting by-product was the effect of predatory Gamasina mites, which survived in some microcosms, but went extinct in others [21].

## **3. Results and discussion**

## **3.1 The coniferous forest floor: a high density and species rich habitat for mites**

Podzol soil with vegetation type 4 in the main study area contained 48 species of Oribatida and 12 species of Mesostigmata (**Table 1**). The density of mites was high.

In the upper 6-cm soil, the mean numbers per m<sup>2</sup> , based on eight replicates, were: Prostigmata (Actinedida) 490,000, Oribatida 220,000, and Astigmata (Acaridida) 10,000. The total mite density was 720,000 per m2 . The highest total density in one replicate amounted to 1.2 million mites per m2 [14].

Comparable data exist from Finland and Sweden. In southern and central parts of Finland, mites were studied in four coniferous forest sites [28]. The localities corresponded to vegetation types 2 and 4 in the present study. The densities of oribatids, 186,000–351,000 per m<sup>2</sup> , were in the same order of magnitude as in


#### **Table 1.**

*In the clearcut area of the main study site Nordmoen, 48 species/taxa of Oribatida were recorded, and 12 of Mesostigmata.*

**87**

**Table 2.**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

the present study for vegetation type 4. However, their Prostigmata densities,

taxa were recorded in a Finnish spruce site with vegetation type 4. In another Finnish study of spruce forest soil, 35 taxa of oribatids were recorded and a

In an old Swedish pine forest of vegetation type 1–2, 52 oribatid species were

found, which surpasses both the Norwegian and Finnish densities mentioned above.

number of 720,000. We can conclude that Nordic coniferous forest soils with raw humus have a very rich mite fauna, both in oribatid species and in total mite numbers.

The main study area had very homogeneous soil conditions over a large area. It was a flat plain with stone-free, sandy soil, without visible variations in moisture conditions or vegetation. Still, as shown in **Table 2**, the horizontal distribution of

In another experiment, litter bags with birch leaves were placed in the humus layer of four random blocks. The mite fauna which colonized the litter varied significantly between blocks [12]. The Astigmata species *Tyrophagus* cf. *fungivorus* (Oudemans) colonized heavily in Blocks 1 and 2, while *Oppia ornata* occurred mainly in the other two. Actinedida mites were especially numerous in litter bags of Block 4, while the same litter bags had the lowest number of *Autogneta trägårdhi*. Block 1 had high numbers of *Oribatula tibialis*, while *Chamobates incisus* had its

The study of vertical distribution in mites was restricted to the upper 6 cm. *Carabodes* species only rarely occurred in the 3–6 cm layer and were to a large degree

*and homogeneous forest area. O = Oribatida and M = Mesostigmata. Mite density in a given plot was the mean* 

*.*

**Species Group Densities** *Parazercon sarekensis* M 1.7–5.2 *Veigaia nemorensis* M 0.1–1.7 *Tectocepheus velatus* O 20–110 *Nothrus silvestris* O 2–95 *Brachychochthonius zelawaiensis* O 2–100 *Oppia obsoleta* O 0–5.5 *Oppia nova* O 0–4.5 *Paulonothrus longisetosus* O 0–3.7 Brachychthoniidae O 20–200 Total Oribatida 80–360 Astigmata (Acaridida) 3–30 Prostigmata (Actinedida) 230–850 Total Acari 400–1200

recorded and very high densities [30]. As much as 425,000 oribatids per m2

, were only about one tenth of ours. As much as 62 oribatid

[29].

) were between Norwegian and Finnish

) approached the high Norwegian

*) in eight random study plots (each 4 × 4 m) on a flat* 

were

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

Prostigmata numbers (210,000 per m2

**3.2 Horizontal and vertical distribution**

highest numbers in Blocks 2 and 3 (**Table 3**).

*Lowest and highest density of various mites (1000 per m2*

*of 10 soil cores, 6 cm deep and with a surface area of 5.3 cm<sup>2</sup>*

relatively low density, only 70,000 oribatids per m<sup>2</sup>

numbers, and total mite numbers (684,000 per m2

many species showed considerable local variations [14, 15].

34,000–80,000 per m<sup>2</sup>

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

the present study for vegetation type 4. However, their Prostigmata densities, 34,000–80,000 per m<sup>2</sup> , were only about one tenth of ours. As much as 62 oribatid taxa were recorded in a Finnish spruce site with vegetation type 4. In another Finnish study of spruce forest soil, 35 taxa of oribatids were recorded and a relatively low density, only 70,000 oribatids per m<sup>2</sup> [29].

In an old Swedish pine forest of vegetation type 1–2, 52 oribatid species were recorded and very high densities [30]. As much as 425,000 oribatids per m2 were found, which surpasses both the Norwegian and Finnish densities mentioned above. Prostigmata numbers (210,000 per m2 ) were between Norwegian and Finnish numbers, and total mite numbers (684,000 per m2 ) approached the high Norwegian number of 720,000. We can conclude that Nordic coniferous forest soils with raw humus have a very rich mite fauna, both in oribatid species and in total mite numbers.

#### **3.2 Horizontal and vertical distribution**

*Pests Control and Acarology*

In the upper 6-cm soil, the mean numbers per m<sup>2</sup>

10,000. The total mite density was 720,000 per m2

**Oribatida Oribatida (continued)** *Adoristes poppei* (Oudemans) *Oppia subpectinata* Willmann *Autogneta parva* Forsslund *Oppia unicarinata* (Paoli) *Autogneta trägårdhi* Forsslund *Oppia nova* (Oudemans) *Belba* cf. *compta* Kulczynski *Oribatula tibialis* (Nicolet)

*Brachychochthonius zelawaiensis* (Sellnick) *Palaeacarus* sp.

*Carabodes femoralis* (Nicolet) *Phthiracarus* sp.

*Carabodes subarcticus* Trägårdh *Steganacarus* sp.

*Chamobatidae* sp. **Mesostigmata** *Eueremaeus silvestris (*Forsslund) *Eviphis ostrinus* (Koch)

*Nothrus silvestris Nicolet Trachytes* sp.

*Oppia* cf. *translamellata* (Willmann) *Veigaia cerva* (Kramer) *Oppia obsoleta (*Paoli) *Veigaia nemorensis* C. L. Koch

*Caleremaeus monolipes* (Michael) *Parachipteria* cf. *willmanni* (V. D. Hammen)

*Camisia biurus* (C. L. Koch) *Paraleius* cf. *leontonycha* (Berlese) *Camisia* cf. *lapponica* Trägårdh *Paulonothrus longisetosus* (Willmann) *Camisia spinifer* (C. L. Koch) *Pergalumna nervosus* (Berlese)

*Carabodes forsslundi* Sellnick *Platynothrus peltifer* (C. L. Koch) *Carabodes labyrinthicus* (Michael) *Porobelba spinosa* (Sellnick) *Carabodes marginatus* (Michael) *Scheloribates laevigatus* (C. L. Koch)

*Cepheus cepheiformis* (Nicolet) *Suctobelba subcornigera* (Forsslund) *Ceratozetes* sp*. Tectocepheus velatus* (Michael)

*Eupelops duplex* (Berlese) *Gamasellus montanus* (Willmann) *Eupelops geminus* (Berlese) *Hypoaspis forcipata* Willmann *Euphthiracaridae Leioseius bicolor* Berlese *Hemileius initialis* Berlese *Parazerkon sarekensis* Willmann *Hypochthonius rufulus* C. L. Koch *Pergamasus* cf. *lapponicus* Trägårdh *Liacarus* cf. *coracinus* (C. L. Koch) *Pergamasus parrunciger* Bhattacharyya *Licneremaeus licnophorus* (Michael) *Pergamasus robustus* Oudemans *Nanhermannia* cf. *forsslundi* Karppinen *Prozercon kochi* Sellnick

*Chamobates incisus* (V. D. Hammen) *Zygoribatula* cf. *trigonella* Bulanova & Zachvatkina

*In the clearcut area of the main study site Nordmoen, 48 species/taxa of Oribatida were recorded, and 12 of* 

replicate amounted to 1.2 million mites per m2

oribatids, 186,000–351,000 per m<sup>2</sup>

Prostigmata (Actinedida) 490,000, Oribatida 220,000, and Astigmata (Acaridida)

Comparable data exist from Finland and Sweden. In southern and central parts of Finland, mites were studied in four coniferous forest sites [28]. The localities corresponded to vegetation types 2 and 4 in the present study. The densities of

[14].

, were in the same order of magnitude as in

, based on eight replicates, were:

. The highest total density in one

**86**

**Table 1.**

*Mesostigmata.*

*Oppia ornata* (Oudemans)

The main study area had very homogeneous soil conditions over a large area. It was a flat plain with stone-free, sandy soil, without visible variations in moisture conditions or vegetation. Still, as shown in **Table 2**, the horizontal distribution of many species showed considerable local variations [14, 15].

In another experiment, litter bags with birch leaves were placed in the humus layer of four random blocks. The mite fauna which colonized the litter varied significantly between blocks [12]. The Astigmata species *Tyrophagus* cf. *fungivorus* (Oudemans) colonized heavily in Blocks 1 and 2, while *Oppia ornata* occurred mainly in the other two. Actinedida mites were especially numerous in litter bags of Block 4, while the same litter bags had the lowest number of *Autogneta trägårdhi*. Block 1 had high numbers of *Oribatula tibialis*, while *Chamobates incisus* had its highest numbers in Blocks 2 and 3 (**Table 3**).

The study of vertical distribution in mites was restricted to the upper 6 cm. *Carabodes* species only rarely occurred in the 3–6 cm layer and were to a large degree


#### **Table 2.**

*Lowest and highest density of various mites (1000 per m2 ) in eight random study plots (each 4 × 4 m) on a flat and homogeneous forest area. O = Oribatida and M = Mesostigmata. Mite density in a given plot was the mean of 10 soil cores, 6 cm deep and with a surface area of 5.3 cm<sup>2</sup> .*


#### **Table 3.**

*Examples of how the number of mites per litter bag with birch leaves may vary between four blocks in a flat and apparently homogeneous forest floor [12].*

living in close connection with *Cladonia* lichens on the surface [25]. In the main study area, there was no sharp change in the mite fauna between the organic layer (0–3 cm) and the bleached mineral layer (3–6 cm). For instance, the large *Nothrus silvestris* was equally abundant in the two layers. However, the addition of strong doses of lime or artificial acid rain was apparently stressful for several mites, forcing animals to deeper layers. After treatment, the following oribatids moved significantly deeper, shifting from living mainly in the organic layer, to live mainly in the mineral layer: *Nothrus silvestris, Suctobelba* sp., *Brachychochthonius zelawaiensis*, and total oribatids. However, Prostigmata mites showed a shift upwards in the soil profile [14]. A frequent natural stress factor in soil is drought. In a Finnish forest, *Nothrus silvestris* was seen to migrate into deeper layers during warm periods [31].

In the comparative study between different vegetation types and soils, all the six selected mites showed variations in depth distribution, not only between habitats, but also between seasons [15]. On the average, the following percentages of the populations occurred in the upper 3 cm compared to 3–6 cm depth: 85% in *Tectocepheus velatus*, 65% in *Parazercon sarekensis*, 60% in *Schwiebea* cf. *cavernicola* Vitzthum, 54% in *Brachychochthonius zelawaiensis*, 52% in *Nothrus silvestris*, and 51% in *Schwiebea* cf. *nova* (Oudemans). The somewhat deeper distribution of *Nothrus silvestris* compared to *Tectocepheus velatus* has been confirmed by other studies [32–34].

#### **3.3 Effect of ground vegetation and soil type**

Eight mite species were studied systematically with respect to vegetation types and soil characteristics [15, 25]. Five belonged to the oribatids, two belonged to Acaridida, and one to Mesostigmata (**Table 4**). Most species preferred poor and acidic podzol soils with raw humus (up to vegetation type 4), but *S.* cf. *cavernicola* had the highest density in a poor brown earth (type 6). None of the eight species were abundant in the richest soil, a brown earth with mull humus (type 7). The non-*Carabodes* species in **Table 4** were tested for correlation between population size and soil chemical parameters. Soil pH, and the accompanying parameters base saturation and calcium content, turned out to be the strongest explanatory factor.

Some other *Carabodes* species were so rare in all soils that they have been excluded from **Table 4**, but further mentioned under the next point.

Comparable data from Finland and Sweden confirm that *Nothrus silvestris* and *Tectocepheus velatus* occur in many different plant communities of coniferous forest, but typically in acid raw humus, and with low densities in richer soils [31, 33, 35, 36]. Although preferences exist, it has been concluded on a general basis that many oribatid species are able to persist in a wide range of humus forms and vegetation types [37].

**89**

unknown preferences.

**Table 4.**

*Abundance (1000 per m2*

*vegetation types in coniferous forest.*

*3.5.1 General results*

occurred mainly at certain pH levels [15].

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

**Species Group Vegetation type**

*Carabodes willmanni* Oribatida 37.1 43.1 4.7

*Carabodes subarcticus* Oribatida 15.2 0.5 2.5 0.02

*Parazercon sarekensis* Mesostigmata 3.4 1.5 1.8 4.0 2.9 1.3

*Tectocepheus velatus* Oribatida 175.3 66.8 99.1 47.2 7.4 11.5 0.7 *Brachychochthonius zelawaiensis* Oribatida 0.9 1.5 27.7 38.8 12.2 1.6 0.5 *Nothrus silvestris* Oribatida 1.8 3.5 14.8 22.0 3.7 7.3 1.9 *Schwiebea* cf. *cavernicola* Acaridida 0.7 1.3 7.1 4.0 7.8 11.3 2.6 *Schwiebea* cf. *nova* Acaridida 0.4 0.9 1.8 12.2 3.4 0.4 0.5 *Numbers are mean value from two localities, each sampled during spring and autumn. Vegetation types 1–7 are described in Material and Methods. Soil fertility increased from left to right. For complete vegetation data, see [27].*

**1 2 3 4 5 6 7**

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

**3.4 Carabodes: a genus with different life forms**

**3.5 Effect of soil acidity on mites—natural and manipulated**

Three approaches were used to test whether soil pH was an important environmental factor for mites. First, a "preference experiment" was arranged in the laboratory [10]. Here, mites were allowed to colonize soils adjusted to different pH levels. Second, we studied responses to artificial pH changes in soil through liming and artificial acid rain, both in the field and in the laboratory [13, 14]. Third, mites were sampled in natural soils of varying pH, to check if there were species that

**Table 6** gives the most consistent results from the first two approaches. Clear

responses were found in three oribatid species, in total Oribatida, and in the Acaridida species *Schwiebea* cf*. nova.* Raised pH due to liming reduced densities of these taxa, while acidification usually led to higher densities. The third approach from natural soils of different pH supported the pattern: species which increased

The combined study of mites in different coniferous forest types and mites in decomposing polypore fungi illustrated that closely related species within a genus (*Carabodes*) can fill quite different niches in the forest ecosystem [25]. The most common *Carabodes* species in soil were rare in sporocarps and vice versa. The first two species in **Table 5** were considered *Cladonia*-feeders on the ground and were able to live in a dry forest floor. The third species on the list is also a lichen-feeder, which often climbs tree stems. Then, we have three fungal feeders which decompose dead sporocarps and may achieve high densities in these patchy and temporary habitats. Their relative numbers were rather similar in dead sporocarps of five different fungal species, including annual and perennial sporocarps, soft and hard. Although being tolerant to different fungal species, these specialists were considered vulnerable in forests with little dead wood and few sporocarps [25]. The five lower species have been found in low numbers, both in sporocarps, in dead wood, and in soil. They are either generalists or have

 *in the upper 6 cm soil layer) of some common mite species in seven different* 


#### *Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

*Numbers are mean value from two localities, each sampled during spring and autumn. Vegetation types 1–7 are described in Material and Methods. Soil fertility increased from left to right. For complete vegetation data, see [27].*

#### **Table 4.**

*Pests Control and Acarology*

**Table 3.**

**Species Sample** 

*and apparently homogeneous forest floor [12].*

**No**

*Autogneta trägårdhi* I 56.6 45.1 55.8 21.4

living in close connection with *Cladonia* lichens on the surface [25]. In the main study area, there was no sharp change in the mite fauna between the organic layer (0–3 cm) and the bleached mineral layer (3–6 cm). For instance, the large *Nothrus silvestris* was equally abundant in the two layers. However, the addition of strong doses of lime or artificial acid rain was apparently stressful for several mites, forcing animals to deeper layers. After treatment, the following oribatids moved significantly deeper, shifting from living mainly in the organic layer, to live mainly in the mineral layer: *Nothrus silvestris, Suctobelba* sp., *Brachychochthonius zelawaiensis*, and total oribatids. However, Prostigmata mites showed a shift upwards in the soil profile [14]. A frequent natural stress factor in soil is drought. In a Finnish forest, *Nothrus silvestris* was seen to migrate into deeper layers during warm periods [31]. In the comparative study between different vegetation types and soils, all the six selected mites showed variations in depth distribution, not only between habitats, but also between seasons [15]. On the average, the following percentages of the populations occurred in the upper 3 cm compared to 3–6 cm depth: 85% in *Tectocepheus velatus*, 65% in *Parazercon sarekensis*, 60% in *Schwiebea* cf. *cavernicola* Vitzthum, 54% in *Brachychochthonius zelawaiensis*, 52% in *Nothrus silvestris*, and 51% in *Schwiebea* cf. *nova* (Oudemans). The somewhat deeper distribution of *Nothrus silvestris* compared

*Examples of how the number of mites per litter bag with birch leaves may vary between four blocks in a flat* 

**Block numbers Significance**

**B 1 B 2 B 3 B 4**

*Tyrophagus* cf. *fungivorus* I 533.5 735.8 13.5 2.3 B3 & B4 < B1 & B2 *Oppia ornata* III 0 0 21.7 6.3 B3 > B1, B2 & B4 Prostigmata (Actinedida) II 46.8 77.5 211.0 343.8 B4 > B1 & B2

*Oribatula tibialis* II 208.5 92.7 44.9 28.6 B1 > B2, B3 & B4 *Chamobates incisus* II 0 2.0 4.6 0.6 B3 > B1 & B4

to *Tectocepheus velatus* has been confirmed by other studies [32–34].

excluded from **Table 4**, but further mentioned under the next point.

Comparable data from Finland and Sweden confirm that *Nothrus silvestris* and *Tectocepheus velatus* occur in many different plant communities of coniferous forest, but typically in acid raw humus, and with low densities in richer soils [31, 33, 35, 36]. Although preferences exist, it has been concluded on a general basis that many oribatid species are able to persist in a wide range of humus forms and vegetation types [37].

Eight mite species were studied systematically with respect to vegetation types and soil characteristics [15, 25]. Five belonged to the oribatids, two belonged to Acaridida, and one to Mesostigmata (**Table 4**). Most species preferred poor and acidic podzol soils with raw humus (up to vegetation type 4), but *S.* cf. *cavernicola* had the highest density in a poor brown earth (type 6). None of the eight species were abundant in the richest soil, a brown earth with mull humus (type 7). The non-*Carabodes* species in **Table 4** were tested for correlation between population size and soil chemical parameters. Soil pH, and the accompanying parameters base saturation and calcium content, turned out to be the strongest explanatory factor. Some other *Carabodes* species were so rare in all soils that they have been

**3.3 Effect of ground vegetation and soil type**

**88**

*Abundance (1000 per m2 in the upper 6 cm soil layer) of some common mite species in seven different vegetation types in coniferous forest.*

## **3.4 Carabodes: a genus with different life forms**

The combined study of mites in different coniferous forest types and mites in decomposing polypore fungi illustrated that closely related species within a genus (*Carabodes*) can fill quite different niches in the forest ecosystem [25]. The most common *Carabodes* species in soil were rare in sporocarps and vice versa. The first two species in **Table 5** were considered *Cladonia*-feeders on the ground and were able to live in a dry forest floor. The third species on the list is also a lichen-feeder, which often climbs tree stems. Then, we have three fungal feeders which decompose dead sporocarps and may achieve high densities in these patchy and temporary habitats. Their relative numbers were rather similar in dead sporocarps of five different fungal species, including annual and perennial sporocarps, soft and hard. Although being tolerant to different fungal species, these specialists were considered vulnerable in forests with little dead wood and few sporocarps [25]. The five lower species have been found in low numbers, both in sporocarps, in dead wood, and in soil. They are either generalists or have unknown preferences.

#### **3.5 Effect of soil acidity on mites—natural and manipulated**

#### *3.5.1 General results*

Three approaches were used to test whether soil pH was an important environmental factor for mites. First, a "preference experiment" was arranged in the laboratory [10]. Here, mites were allowed to colonize soils adjusted to different pH levels. Second, we studied responses to artificial pH changes in soil through liming and artificial acid rain, both in the field and in the laboratory [13, 14]. Third, mites were sampled in natural soils of varying pH, to check if there were species that occurred mainly at certain pH levels [15].

**Table 6** gives the most consistent results from the first two approaches. Clear responses were found in three oribatid species, in total Oribatida, and in the Acaridida species *Schwiebea* cf*. nova.* Raised pH due to liming reduced densities of these taxa, while acidification usually led to higher densities. The third approach from natural soils of different pH supported the pattern: species which increased


*Very high abundance is subjectively indicated by ++++ and very low abundance by (+). Short remarks are given for some species.*

#### **Table 5.**

*Simplified overview on the occurrence of various Carabodes species in different forest habitats, compiled from several sources. From [25].*


#### **Table 6.**

*Significant effects of liming and acidification on mite densities. Compiled from several studies.*

in numbers during artificial acidification were often numerous in naturally acid soils [15]. It was concluded that soil pH was a highly relevant environmental factor for certain mites. Among them was the rather large oribatid species *Nothrus silvestris* (**Figure 3**).

In field experiments with application of artificial rain, the structure of the mite community changed in a characteristic way. **Figure 4** shows how the dominance structure was influenced by liming and application of "rain" with pH 2.5 and 2. Watering with pH 6 was considered as control. The dominance of Oribatida increased with increased acidification. Changes were mainly due to reactions in the sensitive species from **Table 6**.

Finnish [38] and Swedish [39] experiments conformed well with these data, as well as other studies referred to in [14].

Soil acidity is, of course, only one of many factors that modify the abundance of these species, and the relation is not absolute. Even if the pH level is favorable, other limiting factors, for instance drought, may depress populations. The

**91**

be limiting [16].

*Acaridida = Astigmata.*

**Figure 4.**

**Figure 3.**

*3.5.2 Is competition a key factor?*

Soil pH is a measure of the H+

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

experiments support the following conclusion: a high abundance in certain species can only be achieved within a certain pH interval (and only if other factors are not limiting), while within another pH interval, high abundance cannot be achieved. In soils of the latter pH interval, the acidity level (or correlated factors) seems to

*Effect of liming and acidification (the two most extreme treatments, pH 2.5 and 2) on the relative dominance* 

*among mites. Watering with pH 6 was considered as "control." The dominance of Oribatida mites, indicated by double arrows, increased with increased acidification [14]. Actinedida = Prostigmata and* 

*Nothrus silvestris is an oribatid species that is typical for acid raw humus and declined after liming. Photo by* 

Relations between abundance of mites and soil acidity are difficult to explain.

a direct importance for the water-living part of the soil fauna (such as Protozoa and Rotifera), and to other groups living in contact with the soil solution, as Nematoda [40, 41]. Both Enchytraeidae and Lumbricidae prefer relatively high moisture in the soil [42]. The survival of the Enchytraeidae species *Cognettia sphagnetorum* Vejdovsky decreased rapidly when the animals were submerged in diluted sulfuric acid of pH below 4 [9]. Many Enchytraeidae species show distinct relations to soil pH, both in experiments and in the field [10, 39, 43, 44]. The dependence of

Lumbricidae species upon soil pH is well documented [45–47].

activity of the soil solution. This parameter may have

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

*courtesy of SNSB—Zoologische Staatssammlung München.*

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

#### **Figure 3.**

*Pests Control and Acarology*

**90**

(**Figure 3**).

**Table 6.**

*Brachychochthonius zelawaiensis*

*some species.*

*several sources. From [25].*

**Table 5.**

sensitive species from **Table 6**.

well as other studies referred to in [14].

in numbers during artificial acidification were often numerous in naturally acid soils [15]. It was concluded that soil pH was a highly relevant environmental factor for certain mites. Among them was the rather large oribatid species *Nothrus silvestris*

*Tectocepheus velatus* — — + + + +

*Very high abundance is subjectively indicated by ++++ and very low abundance by (+). Short remarks are given for* 

*Simplified overview on the occurrence of various Carabodes species in different forest habitats, compiled from* 

*Total Oribatida* — + + + *Schwiebea* cf*. nova* — + +

*Significant effects of liming and acidification on mite densities. Compiled from several studies.*

**Species Effect of liming Effect of acidification**

**Species In sporocarps In dead wood In soil Remark**

*C. willmanni* Bernini (+) ++++ *Cladonia*-feeder on the ground *C. subarcticus* Trägårdh (+) + ++ *Cladonia*-feeder on the ground? *C. labyrinthicus* (Michael) +(+) + +(+) Lichen-feeder, common on tree stems

*C. femoralis* (Nicolet) ++++ ++ + Polypore specialist *C. areolatus* Berlese +++ ++ (+) Polypore specialist *C. reticulatus* Berlese +++ + Polypore specialist

**Field** 

**experiment [14]**

**Colonization experiment [10]**

— — + + —

**Field** 

**experiment [14]**

**Birch leaves [13]**

**Field Greenhouse**

**Colonization experiment [10]**

*C. marginatus* (Michael) (+) + + *C. forsslundi* Sellnick + + + *C. rugosior* Berlese + + (+) *C. tenuis* Forsslund + + (+) *C. coriaceus* Koch + + (+)

*Nothrus silvestris* — — +

In field experiments with application of artificial rain, the structure of the mite community changed in a characteristic way. **Figure 4** shows how the dominance structure was influenced by liming and application of "rain" with pH 2.5 and 2. Watering with pH 6 was considered as control. The dominance of Oribatida increased with increased acidification. Changes were mainly due to reactions in the

Finnish [38] and Swedish [39] experiments conformed well with these data, as

Soil acidity is, of course, only one of many factors that modify the abundance

of these species, and the relation is not absolute. Even if the pH level is favorable, other limiting factors, for instance drought, may depress populations. The *Nothrus silvestris is an oribatid species that is typical for acid raw humus and declined after liming. Photo by courtesy of SNSB—Zoologische Staatssammlung München.*

#### **Figure 4.**

*Effect of liming and acidification (the two most extreme treatments, pH 2.5 and 2) on the relative dominance among mites. Watering with pH 6 was considered as "control." The dominance of Oribatida mites, indicated by double arrows, increased with increased acidification [14]. Actinedida = Prostigmata and Acaridida = Astigmata.*

experiments support the following conclusion: a high abundance in certain species can only be achieved within a certain pH interval (and only if other factors are not limiting), while within another pH interval, high abundance cannot be achieved. In soils of the latter pH interval, the acidity level (or correlated factors) seems to be limiting [16].

### *3.5.2 Is competition a key factor?*

Relations between abundance of mites and soil acidity are difficult to explain. Soil pH is a measure of the H+ activity of the soil solution. This parameter may have a direct importance for the water-living part of the soil fauna (such as Protozoa and Rotifera), and to other groups living in contact with the soil solution, as Nematoda [40, 41]. Both Enchytraeidae and Lumbricidae prefer relatively high moisture in the soil [42]. The survival of the Enchytraeidae species *Cognettia sphagnetorum* Vejdovsky decreased rapidly when the animals were submerged in diluted sulfuric acid of pH below 4 [9]. Many Enchytraeidae species show distinct relations to soil pH, both in experiments and in the field [10, 39, 43, 44]. The dependence of Lumbricidae species upon soil pH is well documented [45–47].

Microarthropods, on the other hand, have a hydrophobic cuticula and are restricted to the air-filled pore spaces of the soil. The relations described are probably indirect. Several possibilities have been discussed [16]: changes in ground vegetation due to artificial acid rain, direct effects of lime or sulfuric acid, various factors correlated to soil pH, changed predation pressure, availability of fungal hyphae as food, or fecundity. After having refuted several hypotheses, the following laboratory experiment pointed toward competition as a possible explanation [18].

Some microcosms were added a full soil fauna, while others were monocultures of selected species. The acidophilic Acaridid mite *Schwiebea* cf. *nova* (later named *S.* cf. *lebruni* Fain) thrived in monocultures. Starting with 30 specimens, populations increased to around 2000. Surprisingly, population growth in monoculture was lowest in the most acid soil. In the "full fauna" microcosms, however, the species revealed its typical acidophilic character and achieved the highest populations in the most acid soil. Quite parallel results were achieved for the acidophilic springtail *Mesaphorura yosii* (Rusek) [18]. These species have an optimum at a high pH when being alone. However, by some reason, they seem to be good competitors at low pH. They were winners both in natural soils with a low pH and in various experiments with artificial acid rain. Also the acidophilic oribatid *Nothrus silvestris* reproduced best in limed soil when alone [18].

For Collembola, other laboratory studies on population growth, with or without other species present, have illustrated that competition occurs [48, 49]. In most cases, the presence of another species reduced population growth. The most common mechanism was disturbance during oviposition. A classic study about competition among oribatid mites was performed in microcosms with natural soil. Two species with overlapping niches, *Hermaniella granulata* (Nicolet) and *Nothrus silvestris*, were first bred in monocultures. When put together, both species underwent significant shifts in their use of space and food. Their vertical distribution changed so that *Hermaniella* moved upwards into the litter layers, while the *Nothrus* population increased in the deeper fermentation layer [50].

Competition may attain many forms, and the topic is not easy to disentangle. However, since species live so densely packed in soil, one can imagine that disturbance or limited space or food may have an influence. If competition is a key factor regulating population size in soil, a general study of competition in microarthropods might be rewarding. Although a species may have its set of preferences, the key quality may be its ability to compete under suboptimal conditions.

#### **3.6 The effect of predatory Mesostigmata mites**

While the function of soil mites is often focused on their role in decomposition, predatory Mesostigmata mites have the potential to control the density of little sclerotized prey of various taxa. The evolution of strongly sclerotized bodies in many oribatid species obviously has an antipredator role.

The microcosm experiment described above illustrated the predatory effect of large Gamasina mites. At the start, 96% of the cultures contained predatory Gamasina mites, mainly *Veigaia nemorensis*. This percentage was reduced to 73% after 3 months, 62% after 6 months, and 50% after 12 months. The local extinction of these predators often resulted in very high densities of springtails or mites. For instance, after 1 year, the number of *Schwiebea* mites in certain predator-free microcosms could amount to several hundred, while predator-containing cultures usually had numbers below 30. Also for springtails, the highest populations were recorded in cultures where predatory Gamasina mites had gone extinct [21].

**93**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

less sclerotized oribatids, as well as Collembola and Protura [5].

From the literature, another laboratory experiment illustrated well this topdown control of microarthropods. The addition of predatory mites to isolated soil cores containing a natural microarthropod fauna reduced the density of small and

In agroecosystems, edaphic Mesostigmata have been shown to be important predators of Collembola and Nematoda, and those living on plants may efficiently

In the main study area at Nordmoen, the clearcut area was used for litter bag studies, as described above. Litter bags with birch leaves were placed out in July 1975. There were four samplings: September 1975, April 1976, September 1976, and November 1978. The number of leaf-containing litter bags harvested at each

Litter bags with spruce needles were placed out in September 1977, and samplings were made after 7 weeks, 8 months, 1 year, 2 years, 5 years, and 10 years. All samplings, except for the second one, were taken at the same time of the year. There were four replication sites, and 5–15 litter bags were harvested from each replication at a given sampling. Detailed results were given for birch leaves [12] and for needles

In both litter types, a gradual change in the mite community was observed during the decomposition process. However, the succession pattern differed in spruce needles and birch leaves. It means that mites in the surrounding soil were selective about which litter they colonized, at which rate, and at which decomposition stage. For instance, two oribatid species which were common in the soil, *Tectocepheus velatus* and *Nothrus silvestris*, never became abundant in litter bags. On the other hand, certain low-density species in soil could achieve very high densities in the bags. In such cases, a high density was only seen in one of the litter types. Examples in spruce needle bags were high density of *Eremaeus* sp. after 1 year, *Steganacarus* sp.

A considerable number of spruce needles were decomposed from the inside by certain specialized oribatid mites [22, 26]. Smaller, deeper-living species became abundant after 5–10 years, when the needles had been more or less fragmented. The fragmentation created new microhabitats and perhaps allowed for a more intense

While colonization of needle litter was slow, and no species or group achieved its maximum abundance within 8 months, colonization of birch leaves was much faster. Here, certain mites, which had a low density in the surrounding soil,

appeared very numerous already after 7 weeks. Examples were three oribatid mites: *Oribatula tibialis, Eupelops duplex*, and *Autogneta trägårdhi*, and one Acaridida (Astigmata): *Tyrophagus* cf. *fungivorus.* Studies of the gut contents of these four species revealed a mixture of fungal spores and hyphae, and some guts contained mainly spores. This indicated an intense grazing, probably due to a temporal "flush" of fungal activity. The same was seen for certain springtail species [12]. It is, of course, important for soil microarthropods to detect such spatial and temporal food sources, and it is reasonable to assume that animals were attracted from surroundings by smell. Also other studies have documented a rapid migration of microarthropods into decomposing deciduous leaves [51–53]. Such species can be characterized as mobile opportunists. An abundant food source may allow a high number of species and specimens to coexist in a substrate with a low structural diversity. The body of *Eupelops duplex*, but also other species, was often covered by

**3.7 Succession in the mite community during decomposition of spruce** 

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

control pests like spider mites [4].

**needles and birch leaves**

sampling was 32, 68, 128, and 78, respectively.

after 5 years, and *Oppiella nova* after 10 years.

microfloral colonization.

[19]. Here, the main trends shall be presented and compared.

From the literature, another laboratory experiment illustrated well this topdown control of microarthropods. The addition of predatory mites to isolated soil cores containing a natural microarthropod fauna reduced the density of small and less sclerotized oribatids, as well as Collembola and Protura [5].

In agroecosystems, edaphic Mesostigmata have been shown to be important predators of Collembola and Nematoda, and those living on plants may efficiently control pests like spider mites [4].

## **3.7 Succession in the mite community during decomposition of spruce needles and birch leaves**

In the main study area at Nordmoen, the clearcut area was used for litter bag studies, as described above. Litter bags with birch leaves were placed out in July 1975. There were four samplings: September 1975, April 1976, September 1976, and November 1978. The number of leaf-containing litter bags harvested at each sampling was 32, 68, 128, and 78, respectively.

Litter bags with spruce needles were placed out in September 1977, and samplings were made after 7 weeks, 8 months, 1 year, 2 years, 5 years, and 10 years. All samplings, except for the second one, were taken at the same time of the year. There were four replication sites, and 5–15 litter bags were harvested from each replication at a given sampling. Detailed results were given for birch leaves [12] and for needles [19]. Here, the main trends shall be presented and compared.

In both litter types, a gradual change in the mite community was observed during the decomposition process. However, the succession pattern differed in spruce needles and birch leaves. It means that mites in the surrounding soil were selective about which litter they colonized, at which rate, and at which decomposition stage. For instance, two oribatid species which were common in the soil, *Tectocepheus velatus* and *Nothrus silvestris*, never became abundant in litter bags. On the other hand, certain low-density species in soil could achieve very high densities in the bags. In such cases, a high density was only seen in one of the litter types. Examples in spruce needle bags were high density of *Eremaeus* sp. after 1 year, *Steganacarus* sp. after 5 years, and *Oppiella nova* after 10 years.

A considerable number of spruce needles were decomposed from the inside by certain specialized oribatid mites [22, 26]. Smaller, deeper-living species became abundant after 5–10 years, when the needles had been more or less fragmented. The fragmentation created new microhabitats and perhaps allowed for a more intense microfloral colonization.

While colonization of needle litter was slow, and no species or group achieved its maximum abundance within 8 months, colonization of birch leaves was much faster. Here, certain mites, which had a low density in the surrounding soil, appeared very numerous already after 7 weeks. Examples were three oribatid mites: *Oribatula tibialis, Eupelops duplex*, and *Autogneta trägårdhi*, and one Acaridida (Astigmata): *Tyrophagus* cf. *fungivorus.* Studies of the gut contents of these four species revealed a mixture of fungal spores and hyphae, and some guts contained mainly spores. This indicated an intense grazing, probably due to a temporal "flush" of fungal activity. The same was seen for certain springtail species [12]. It is, of course, important for soil microarthropods to detect such spatial and temporal food sources, and it is reasonable to assume that animals were attracted from surroundings by smell. Also other studies have documented a rapid migration of microarthropods into decomposing deciduous leaves [51–53]. Such species can be characterized as mobile opportunists. An abundant food source may allow a high number of species and specimens to coexist in a substrate with a low structural diversity. The body of *Eupelops duplex*, but also other species, was often covered by

*Pests Control and Acarology*

explanation [18].

reproduced best in limed soil when alone [18].

population increased in the deeper fermentation layer [50].

**3.6 The effect of predatory Mesostigmata mites**

many oribatid species obviously has an antipredator role.

in cultures where predatory Gamasina mites had gone extinct [21].

Microarthropods, on the other hand, have a hydrophobic cuticula and are restricted to the air-filled pore spaces of the soil. The relations described are probably indirect. Several possibilities have been discussed [16]: changes in ground vegetation due to artificial acid rain, direct effects of lime or sulfuric acid, various factors correlated to soil pH, changed predation pressure, availability of fungal hyphae as food, or fecundity. After having refuted several hypotheses, the following laboratory experiment pointed toward competition as a possible

Some microcosms were added a full soil fauna, while others were monocultures of selected species. The acidophilic Acaridid mite *Schwiebea* cf. *nova* (later named *S.* cf. *lebruni* Fain) thrived in monocultures. Starting with 30 specimens, populations increased to around 2000. Surprisingly, population growth in monoculture was lowest in the most acid soil. In the "full fauna" microcosms, however, the species revealed its typical acidophilic character and achieved the highest populations in the most acid soil. Quite parallel results were achieved for the acidophilic springtail *Mesaphorura yosii* (Rusek) [18]. These species have an optimum at a high pH when being alone. However, by some reason, they seem to be good competitors at low pH. They were winners both in natural soils with a low pH and in various experiments with artificial acid rain. Also the acidophilic oribatid *Nothrus silvestris*

For Collembola, other laboratory studies on population growth, with or without other species present, have illustrated that competition occurs [48, 49]. In most cases, the presence of another species reduced population growth. The most common mechanism was disturbance during oviposition. A classic study about competition among oribatid mites was performed in microcosms with natural soil. Two species with overlapping niches, *Hermaniella granulata* (Nicolet) and *Nothrus silvestris*, were first bred in monocultures. When put together, both species underwent significant shifts in their use of space and food. Their vertical distribution changed so that *Hermaniella* moved upwards into the litter layers, while the *Nothrus*

Competition may attain many forms, and the topic is not easy to disentangle.

While the function of soil mites is often focused on their role in decomposition,

predatory Mesostigmata mites have the potential to control the density of little sclerotized prey of various taxa. The evolution of strongly sclerotized bodies in

The microcosm experiment described above illustrated the predatory effect of large Gamasina mites. At the start, 96% of the cultures contained predatory Gamasina mites, mainly *Veigaia nemorensis*. This percentage was reduced to 73% after 3 months, 62% after 6 months, and 50% after 12 months. The local extinction of these predators often resulted in very high densities of springtails or mites. For instance, after 1 year, the number of *Schwiebea* mites in certain predator-free microcosms could amount to several hundred, while predator-containing cultures usually had numbers below 30. Also for springtails, the highest populations were recorded

However, since species live so densely packed in soil, one can imagine that disturbance or limited space or food may have an influence. If competition is a key factor regulating population size in soil, a general study of competition in microarthropods might be rewarding. Although a species may have its set of preferences, the key quality may be its ability to compete under suboptimal

**92**

conditions.

fungal spores or hyphae, promoting the spread of microflora to all parts of the litter. The study also indicated that several species did not reproduce in the substrate, but only visited it during the adult stage for feeding purpose.

**Table 2** shows that litter-dwelling pioneer mites in birch litter had a very uneven horizontal distribution, within 20–50 m. It meant that the succession pattern in the early decomposition phase varied widely, even within an apparently homogeneous forest floor. In later decomposition stages, however, the microarthropod community was less variable and more predictable.

In both litter types, large, surface-living species were among the early colonizers, while smaller, usually deeper-living species, took over the dominance in later decomposition stages. Since the litter bags had continuously contact with the whole organic layer in the actual soil, the succession studies confirmed that deeper-living, and often small species, preferred a more decomposed material.

While this experiment demonstrated that species often had different preferences for litter type or decomposition stage, it also showed that many species had wide tolerances and could survive, sometimes in low densities, under rather different circumstances. In an English study of oribatid mites in decomposing leaves of beech and chestnut, the 12 most abundant species were present in the litter bags throughout the 20-month study period. During this time, species were able to remain by changing their feeding habits [51]. Another example of high tolerance among oribatid species to different decomposition stages of leaf litter is from Central Amazonas. During the one-year long study, there was no successional changes in the species composition [54].

Few decomposition studies last long enough to describe the late stages of the microarthropod succession. For instance, in a study of root litter decomposition, it was found that oribatid mites showed a preference for the late stages of decomposition [55]. A general challenge in litter bag studies is how to simulate natural conditions. And even if natural conditions are achieved, the result may only have local value. Anyhow, due to a high species number and an ecological flexibility in many species, mites do in several ways contribute in transforming litter to humus. This is exemplified in the next chapter.

## **3.8 From litter to humus: can mites influence the process and the products?**

Juveniles of certain specialized mites excavated cavities in about 40% of newly fallen spruce needles. Their activity reduced the decomposition rate of the actual needles, at least temporarily, probably because their excrements decomposed slowly [22, 26]. The adult mites, which hatched after about 2 years, attacked other needles from the outside and fragmented these (**Figure 5**). Their "inert" excrement pellets

#### **Figure 5.**

*Two spruce needles that have been fragmented by adult "box mites" (Steganacarus cf. striculus) kept in culture. Two ellipsoide-shaped animals are seen. Excrement pellets are numerous. Photo: S. Hågvar.*

**95**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

may contribute to a stable humus layer and perhaps to carbon sequestration [26]. Also other studies have pointed to the fact that fecal pellets of oribatids decompose slowly and may contribute significantly to humus production [56, 57]. Even pine

Individual spruce needles may show quite different decomposition patterns, even if situated close to each other in soil. While some are heavily transformed to excrement pellets, others remain morphologically intact for years. Needles which happen to come in close contact with fine roots may be rapidly "dissolved." These "individual fates" of needles may explain the heterogeneous structure of deep humus [26].

Norwegian coniferous forest is covered by snow for several months each year. When the snow layer exceeds about 20 cm, the temperature at the soil surface stabilizes around 0°C [58]. At this temperature, several surface-living invertebrates are active in the subnivean air space and even feeding [23]. Among these are several species of springtails and mites. During two winters, pitfall traps were operated under 30–150 cm snow in a high altitude spruce forest with bilberry vegetation in Southern Norway. Traps were emptied and replaced at least monthly during the

Twelve taxa of Oribatida were trapped and 10 of Mesostigmata. A number of Prostigmata were also taken (**Table 7**). The Oribatida material was dominated by one species, *Platynothrus capillatus*. All developmental stages of this species were active under snow, and fungal hyphae and spores in their guts proved winter feeding. It was assumed that they were grazing on certain fungi known to decompose litter beneath snow (snow molds) [23]. Also other species of Oribatida, as well as

In the main study area at Nordmoen, microarthropod activity both beneath and within snow was studied [11]. Most surface-living springtails were winter active and even migrated up into the snow layers. Among mites, four predacious Mesostigmata mites and one oribatid species (*Adoristes poppei* Oudemans) were taken in small numbers in pitfall traps, together with numerous Prostigmata. Mites were also found within the snow layers: some Prostigmata, seven taxa of predacious Mesostigmata, and six taxa of oribatids, of which *Adoristes poppei* was the most numerous. It was suggested that microarthropods went into snow to escape possible

Several mite species showed a high tolerance for different plant communities, soils, humus types, litter type, and succession phase. Both birch leaves and spruce needles in litter bags were colonized by a high number of oribatid species. Several of them occurred in both substrates, although colonization was much slower in needle litter. Birch leaves represented an uncommon substrate at the actual site, but probably offered a flush of fungal food. Furthermore, at least some individuals of most species participated in various decomposition phases, where the substrate underwent significant changes. Except for pH, mites seemed to have few strong

Each mite species continually adjusts its vertical position, as far as narrow pores allow, to optimize its survival, food access, and reproductive ability. Such changes were seen also in the horizontal distribution. A more fixed vertical or horizontal position of each species could reduce interspecific competition but would be a disadvantage as soon as adverse or favorable conditions developed in certain layers or sites.

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

needles can be tunneled by phthiracarid mites [57].

**3.9 Mite activity beneath and within snow**

some Prostigmata, had visible gut content.

relations to soil chemical parameters [15].

snow-covered period from October/November to April/May.

harmful water logging or ice formation in late winter [11].

**3.10 Remarks on ecological flexibility and vulnerability**

may contribute to a stable humus layer and perhaps to carbon sequestration [26]. Also other studies have pointed to the fact that fecal pellets of oribatids decompose slowly and may contribute significantly to humus production [56, 57]. Even pine needles can be tunneled by phthiracarid mites [57].

Individual spruce needles may show quite different decomposition patterns, even if situated close to each other in soil. While some are heavily transformed to excrement pellets, others remain morphologically intact for years. Needles which happen to come in close contact with fine roots may be rapidly "dissolved." These "individual fates" of needles may explain the heterogeneous structure of deep humus [26].

#### **3.9 Mite activity beneath and within snow**

*Pests Control and Acarology*

composition [54].

exemplified in the next chapter.

fungal spores or hyphae, promoting the spread of microflora to all parts of the litter. The study also indicated that several species did not reproduce in the substrate, but

**Table 2** shows that litter-dwelling pioneer mites in birch litter had a very uneven horizontal distribution, within 20–50 m. It meant that the succession pattern in the early decomposition phase varied widely, even within an apparently homogeneous forest floor. In later decomposition stages, however, the microarthropod community

In both litter types, large, surface-living species were among the early colonizers, while smaller, usually deeper-living species, took over the dominance in later decomposition stages. Since the litter bags had continuously contact with the whole organic layer in the actual soil, the succession studies confirmed that deeper-living,

While this experiment demonstrated that species often had different preferences for litter type or decomposition stage, it also showed that many species had wide tolerances and could survive, sometimes in low densities, under rather different circumstances. In an English study of oribatid mites in decomposing leaves of beech and chestnut, the 12 most abundant species were present in the litter bags throughout the 20-month study period. During this time, species were able to remain by changing their feeding habits [51]. Another example of high tolerance among oribatid species to different decomposition stages of leaf litter is from Central Amazonas. During the one-year long study, there was no successional changes in the species

Few decomposition studies last long enough to describe the late stages of the microarthropod succession. For instance, in a study of root litter decomposition, it was found that oribatid mites showed a preference for the late stages of decomposition [55]. A general challenge in litter bag studies is how to simulate natural conditions. And even if natural conditions are achieved, the result may only have local value. Anyhow, due to a high species number and an ecological flexibility in many species, mites do in several ways contribute in transforming litter to humus. This is

**3.8 From litter to humus: can mites influence the process and the products?**

Juveniles of certain specialized mites excavated cavities in about 40% of newly fallen spruce needles. Their activity reduced the decomposition rate of the actual needles, at least temporarily, probably because their excrements decomposed slowly [22, 26]. The adult mites, which hatched after about 2 years, attacked other needles from the outside and fragmented these (**Figure 5**). Their "inert" excrement pellets

*Two spruce needles that have been fragmented by adult "box mites" (Steganacarus cf. striculus) kept in culture.* 

*Two ellipsoide-shaped animals are seen. Excrement pellets are numerous. Photo: S. Hågvar.*

only visited it during the adult stage for feeding purpose.

and often small species, preferred a more decomposed material.

was less variable and more predictable.

**94**

**Figure 5.**

Norwegian coniferous forest is covered by snow for several months each year. When the snow layer exceeds about 20 cm, the temperature at the soil surface stabilizes around 0°C [58]. At this temperature, several surface-living invertebrates are active in the subnivean air space and even feeding [23]. Among these are several species of springtails and mites. During two winters, pitfall traps were operated under 30–150 cm snow in a high altitude spruce forest with bilberry vegetation in Southern Norway. Traps were emptied and replaced at least monthly during the snow-covered period from October/November to April/May.

Twelve taxa of Oribatida were trapped and 10 of Mesostigmata. A number of Prostigmata were also taken (**Table 7**). The Oribatida material was dominated by one species, *Platynothrus capillatus*. All developmental stages of this species were active under snow, and fungal hyphae and spores in their guts proved winter feeding. It was assumed that they were grazing on certain fungi known to decompose litter beneath snow (snow molds) [23]. Also other species of Oribatida, as well as some Prostigmata, had visible gut content.

In the main study area at Nordmoen, microarthropod activity both beneath and within snow was studied [11]. Most surface-living springtails were winter active and even migrated up into the snow layers. Among mites, four predacious Mesostigmata mites and one oribatid species (*Adoristes poppei* Oudemans) were taken in small numbers in pitfall traps, together with numerous Prostigmata. Mites were also found within the snow layers: some Prostigmata, seven taxa of predacious Mesostigmata, and six taxa of oribatids, of which *Adoristes poppei* was the most numerous. It was suggested that microarthropods went into snow to escape possible harmful water logging or ice formation in late winter [11].

#### **3.10 Remarks on ecological flexibility and vulnerability**

Several mite species showed a high tolerance for different plant communities, soils, humus types, litter type, and succession phase. Both birch leaves and spruce needles in litter bags were colonized by a high number of oribatid species. Several of them occurred in both substrates, although colonization was much slower in needle litter. Birch leaves represented an uncommon substrate at the actual site, but probably offered a flush of fungal food. Furthermore, at least some individuals of most species participated in various decomposition phases, where the substrate underwent significant changes. Except for pH, mites seemed to have few strong relations to soil chemical parameters [15].

Each mite species continually adjusts its vertical position, as far as narrow pores allow, to optimize its survival, food access, and reproductive ability. Such changes were seen also in the horizontal distribution. A more fixed vertical or horizontal position of each species could reduce interspecific competition but would be a disadvantage as soon as adverse or favorable conditions developed in certain layers or sites.


#### **Table 7.**

*Mites (Acari) caught in pitfall traps under snow during two winter seasons in a high altitude spruce forest, central South Norway. Modified from [23].*

**97**

extinct.

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

The present documentation [12] showing that many springtails and mites change their food habits through the different successional stages is in good accor-

species are relatively rare. A high tolerance for various habitat or nutritional factors, often combined with asexual reproduction, may keep species going on in low numbers. However, when special conditions are created locally, rare species may act as opportunists and flourish temporarily. They also represent an important resource if the ecosystem has to adapt to a new situation, for instance due to

Within both springtail and mite communities, it is a general pattern that most

Although single species may show tolerance to different environmental conditions, the mite community as a whole can be vulnerable to various types of human disturbance. For instance, in New York, the diversity of oribatid mites decreased along a gradient of land use types in the order from forests, via abandoned fields and willow, to corn [7]. A European review on mites as indicators of soil biodiversity and land use monitoring illustrated how sensitive mite communities can be to various types of soil disturbance [59]. Changes in the dominance structure of mite communities were suggested to be an "early warning criterion" for stressed mite communities. The author concluded that residual natural and semi-natural habitats (such as old woodlands, riparian ecosystems, old hedges, and grasslands) with species-rich mite communities found in rural and urban landscapes should be

Coniferous forests are rich in mites: a podzol soil with acid raw humus may con-

Flexible vertical and horizontal distribution: mites can adjust both their depth in the soil profile and their horizontal distribution, either to escape stress or to

Opportunism as a successful strategy: several litter-dwelling mite species rapidly colonized birch leaves in an early decomposition phase, in order to feed on a tempo-

Substrate flexibility: decomposition of spruce needles and birch leaves followed quite different succession patterns, but several mite species participated in both. Closely related species may differ widely in habitat choice and life forms: this was

Predacious Gamasina mites matter: microcosm studies showed high population growth of certain mites and springtails if predatory Gamasina mites went

Oribatids matter in the decomposition process from litter to humus: specialized oribatids excavate spruce needles and produce slowly decomposable excrements. Soil acidity matters: colonization experiments and population studies in monocultures showed that soil pH affected population size in certain species. This led to

Successful competition under suboptimal conditions: surprisingly, certain mites common in acid soils thrived best in less acid soil when being alone (in monocul-

Mites are winter active: several mites are active under snow, often feeding. Some

. This includes a species-rich oribatid fauna.

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

preserved as refuges for dispersion of soil fauna.

aggregate in a patchy and temporary food source.

rary and patchy flush of fungal hyphae and spores.

predictable changes in the community structure of mites.

ture). However, in acid soil, they were good competitors.

**4. Conclusions: spotlights in short**

tain more than a million mites per m2

exemplified in the genus *Carabodes*.

even penetrate into the snow layer.

dance with other observations [51].

climate change.

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

The present documentation [12] showing that many springtails and mites change their food habits through the different successional stages is in good accordance with other observations [51].

Within both springtail and mite communities, it is a general pattern that most species are relatively rare. A high tolerance for various habitat or nutritional factors, often combined with asexual reproduction, may keep species going on in low numbers. However, when special conditions are created locally, rare species may act as opportunists and flourish temporarily. They also represent an important resource if the ecosystem has to adapt to a new situation, for instance due to climate change.

Although single species may show tolerance to different environmental conditions, the mite community as a whole can be vulnerable to various types of human disturbance. For instance, in New York, the diversity of oribatid mites decreased along a gradient of land use types in the order from forests, via abandoned fields and willow, to corn [7]. A European review on mites as indicators of soil biodiversity and land use monitoring illustrated how sensitive mite communities can be to various types of soil disturbance [59]. Changes in the dominance structure of mite communities were suggested to be an "early warning criterion" for stressed mite communities. The author concluded that residual natural and semi-natural habitats (such as old woodlands, riparian ecosystems, old hedges, and grasslands) with species-rich mite communities found in rural and urban landscapes should be preserved as refuges for dispersion of soil fauna.

### **4. Conclusions: spotlights in short**

Coniferous forests are rich in mites: a podzol soil with acid raw humus may contain more than a million mites per m2 . This includes a species-rich oribatid fauna.

Flexible vertical and horizontal distribution: mites can adjust both their depth in the soil profile and their horizontal distribution, either to escape stress or to aggregate in a patchy and temporary food source.

Opportunism as a successful strategy: several litter-dwelling mite species rapidly colonized birch leaves in an early decomposition phase, in order to feed on a temporary and patchy flush of fungal hyphae and spores.

Substrate flexibility: decomposition of spruce needles and birch leaves followed quite different succession patterns, but several mite species participated in both. Closely related species may differ widely in habitat choice and life forms: this was exemplified in the genus *Carabodes*.

Predacious Gamasina mites matter: microcosm studies showed high population growth of certain mites and springtails if predatory Gamasina mites went extinct.

Oribatids matter in the decomposition process from litter to humus: specialized oribatids excavate spruce needles and produce slowly decomposable excrements.

Soil acidity matters: colonization experiments and population studies in monocultures showed that soil pH affected population size in certain species. This led to predictable changes in the community structure of mites.

Successful competition under suboptimal conditions: surprisingly, certain mites common in acid soils thrived best in less acid soil when being alone (in monoculture). However, in acid soil, they were good competitors.

Mites are winter active: several mites are active under snow, often feeding. Some even penetrate into the snow layer.

*Pests Control and Acarology*

MESOSTIGMATA

**ORIBATIDA Stage Number** 

*Camisia biurus* Ad 3 No

*Carabodes labyrinthicus* Ad 2 Yes *Carabodes marginatus* Ad 1 No *Carabodes* sp. T 1 No *Chamobates pusillus* (Berlese) Ad 1 No *Eobrachychthonius borealis* Forsslund Ad 3 Yes *Oppiella neerlandica* (Oudemans) Ad 12 Yes *Oppiella* sp. Ad 1 Yes *Oribatella calcarata* (C.L. Koch) Ad 1 No

*Platynothrus capillatus* (Berlese) Ad 10 Yes

*Steganacarus* sp. Ad 1 Yes *Belba* sp.? Ad 2 No

PROSTIGMATA (ACTINEDIDA) 115 Yes

*Mites (Acari) caught in pitfall traps under snow during two winter seasons in a high altitude spruce forest,* 

*Numbers per 12 functioning traps. Only periods with a continuous snow cover are included. Ad = adults,* 

*Mixozercon serlachii* Lehtinen 1 *Zercon curiosus* Trägårdh 1 *Zercon colligans* Berlese 2 *Holoparasitus* sp. 1 *Lysigamasus lapponicus* (Trägårdh) 4 *Vulgarogamasus kraepelini* (Berlese) 14 *Veigaia nemorensis* 6 *Trachytes aegrota* (C.L. Koch) 2 *Urodiaspis tecta* (Kramer) 1 *Uropodina* sp., nymph 2

TOTAL 244

*T = tritonymphs, D = deuteronymphs, P = protonymphs, and L= larvae.*

*central South Norway. Modified from [23].*

**trapped**

T 2 Yes P 3 No L 2 No

D 7 Yes P 5 Yes

T 9 Yes D 14 Yes P 8 Yes L 2 Yes

T/juv 5 Yes

**Gut contents observed?**

**96**

**Table 7.**

## **5. Final remarks**

There is an increasing awareness for preserving the huge biodiversity of soils [1, 60, 61]. Fragmentation and various management practices of forests may affect even these tiny animals. Some microarthropod species are confined to local soil types, for instance under dry or wet conditions. Furthermore, a forest contains various microhabitats in addition to soils. Examples are moss or lichen vegetation on certain trees, suspended soils in birds´ nests, mold in old, hollow trees, decomposing wood, or fruiting bodies of various fungi. To preserve the species, diversity of microarthropods may demand a relatively large forest area, covering a variety of vegetation types, soils, humus types, and microhabitats.

Due to their long life span, low fecundity, slow development, and low dispersion ability, oribatid mites have been suggested as suitable indicators of soil biodiversity and land use monitoring. In this respect, there is a need to develop standardized procedures for sampling and data analysis [59].

## **Acknowledgements**

I am grateful for being allowed to reuse **Figure 4** from Oikos, **Table 5** from Scandinavian Journal of Forest Research, and **Table 7** from Soil Organisms. Zoologische Staatssammlung München gave permission to use the photo of *Nothrus silvestris.* Ole Wiggo Røstad kindly helped with some figures.

## **Author details**

Sigmund Hågvar

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, Ås, Norway

\*Address all correspondence to: sigmund.hagvar@nmbu.no

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

**99**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

SNSF-project; Internal Report 32/77; 1977. pp. 1-47. (In Norwegian, English

[11] Leinaas HP. Activity of Arthropoda in snow within a coniferous forest, with special reference to Collembola. Holarctic Ecology. 1981;**4**:127-138

[12] Hågvar S, Kjøndal BR. Succession,

microarthropods in decomposing birch leaves. Pedobiologia. 1981;**22**:385-408

microarthropod fauna in decomposing

[14] Hågvar S, Amundsen T. Effects of liming and artificial acid rain on the mite (Acari) fauna in coniferous forest.

[15] Hågvar S. Six common mite species (Acari) in Norwegian coniferous forest soils: Relations to vegetation types and soil characteristics. Pedobiologia.

[16] Hågvar S. Why do collemboles and mites react to changes in soil acidity? Entomologiske Meddelelser.

[17] Hågvar S. Decomposition studies in an easily-constructed microcosm: Effects of microarthropods and varying soil pH. Pedobiologia. 1988;**31**:293-303

[18] Hågvar S. Reactions to soil acidification in microarthropods: Is competition a key factor? Biology and Fertility of Soils. 1990;**9**:178-181

diversity and feeding habits of

[13] Hågvar S, Kjøndal BR. Effects of artificial acid rain on the

birch leaves. Pedobiologia.

1981;**22**:409-422

Oikos. 1981;**37**:7-20

1984;**27**:355-364

1987;**55**:115-119

[10] Hågvar S, Abrahamsen G. Colonisation by Enchytraeidae, Collembola and Acari in sterile soil samples with adjusted pH levels. Oikos.

summary)

1980;**34**:245-258

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

[1] Wall DH, Bardgett RD, Kelly EF. Biodiversity in the dark. Nature Geoscience. 2010;**3**:297-298

[2] Siepel H, Maaskamp F. Mites of different feeding guilds affect decomposition of organic matter. Soil Biology and Biochemistry.

[3] Schneider K, Migge S, Norton RA, Scheu S, Langel R, Reineking A, et al. Trophic niche differentiation in soil microarthropods (Oribatida, Acari): Evidence from stable isotope ratios (15N/14N). Soil Biology and Biochemistry. 2004;**36**:1769-1774

[4] Koehler HH. Mesostigmata (Gamasina, Uropodina), efficient predators in agroecosystems. Agriculture, Ecosystems and Environment. 1997;**62**:105-117

Soil Biology and Biochemistry.

[6] Siepel H. Biodiversity of soil microarthropods: The filtering of species. Biodiversity and Conservation.

of soil mites (Acari: Oribatida,

Ecology. 2007;**35**:140-153

[9] Hågvar S, Abrahamsen G.

Eksperimentelle forsuringsforsøk i skog 5. Jordbunnszoologiske undersøkelser.

[7] Minor MA, Cianciolo JM. Diversity

Mesostigmata) along a gradient of land use types in New York. Applied Soil

[8] Erdmann G, Scheu S, Maraun M. Regional factors rather than forest type drive the community structure of soil living oribatid mites (Acari, Oribatida). Experimental & Applied Acarology.

2009;**41**:170-175

1996;**5**:251-260

2012;**57**:157-169

[5] Schneider K, Maraun M. Top-down control of soil microarthropods – Evidence from a laboratory experiment.

1994;**26**:1389-1394

**References**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

## **References**

*Pests Control and Acarology*

**Acknowledgements**

**5. Final remarks**

**98**

**Author details**

Sigmund Hågvar

provided the original work is properly cited.

University of Life Sciences, Ås, Norway

\*Address all correspondence to: sigmund.hagvar@nmbu.no

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

Faculty of Environmental Sciences and Natural Resource Management, Norwegian

There is an increasing awareness for preserving the huge biodiversity of soils [1, 60, 61]. Fragmentation and various management practices of forests may affect even these tiny animals. Some microarthropod species are confined to local soil types, for instance under dry or wet conditions. Furthermore, a forest contains various microhabitats in addition to soils. Examples are moss or lichen vegetation on certain trees, suspended soils in birds´ nests, mold in old, hollow trees, decomposing wood, or fruiting bodies of various fungi. To preserve the species, diversity of microarthropods may demand a relatively large forest area, covering a variety of

Due to their long life span, low fecundity, slow development, and low dispersion ability, oribatid mites have been suggested as suitable indicators of soil biodiversity and land use monitoring. In this respect, there is a need to develop standardized

I am grateful for being allowed to reuse **Figure 4** from Oikos, **Table 5** from Scandinavian Journal of Forest Research, and **Table 7** from Soil Organisms.

Zoologische Staatssammlung München gave permission to use the photo of *Nothrus* 

vegetation types, soils, humus types, and microhabitats.

*silvestris.* Ole Wiggo Røstad kindly helped with some figures.

procedures for sampling and data analysis [59].

[1] Wall DH, Bardgett RD, Kelly EF. Biodiversity in the dark. Nature Geoscience. 2010;**3**:297-298

[2] Siepel H, Maaskamp F. Mites of different feeding guilds affect decomposition of organic matter. Soil Biology and Biochemistry. 1994;**26**:1389-1394

[3] Schneider K, Migge S, Norton RA, Scheu S, Langel R, Reineking A, et al. Trophic niche differentiation in soil microarthropods (Oribatida, Acari): Evidence from stable isotope ratios (15N/14N). Soil Biology and Biochemistry. 2004;**36**:1769-1774

[4] Koehler HH. Mesostigmata (Gamasina, Uropodina), efficient predators in agroecosystems. Agriculture, Ecosystems and Environment. 1997;**62**:105-117

[5] Schneider K, Maraun M. Top-down control of soil microarthropods – Evidence from a laboratory experiment. Soil Biology and Biochemistry. 2009;**41**:170-175

[6] Siepel H. Biodiversity of soil microarthropods: The filtering of species. Biodiversity and Conservation. 1996;**5**:251-260

[7] Minor MA, Cianciolo JM. Diversity of soil mites (Acari: Oribatida, Mesostigmata) along a gradient of land use types in New York. Applied Soil Ecology. 2007;**35**:140-153

[8] Erdmann G, Scheu S, Maraun M. Regional factors rather than forest type drive the community structure of soil living oribatid mites (Acari, Oribatida). Experimental & Applied Acarology. 2012;**57**:157-169

[9] Hågvar S, Abrahamsen G. Eksperimentelle forsuringsforsøk i skog 5. Jordbunnszoologiske undersøkelser.

SNSF-project; Internal Report 32/77; 1977. pp. 1-47. (In Norwegian, English summary)

[10] Hågvar S, Abrahamsen G. Colonisation by Enchytraeidae, Collembola and Acari in sterile soil samples with adjusted pH levels. Oikos. 1980;**34**:245-258

[11] Leinaas HP. Activity of Arthropoda in snow within a coniferous forest, with special reference to Collembola. Holarctic Ecology. 1981;**4**:127-138

[12] Hågvar S, Kjøndal BR. Succession, diversity and feeding habits of microarthropods in decomposing birch leaves. Pedobiologia. 1981;**22**:385-408

[13] Hågvar S, Kjøndal BR. Effects of artificial acid rain on the microarthropod fauna in decomposing birch leaves. Pedobiologia. 1981;**22**:409-422

[14] Hågvar S, Amundsen T. Effects of liming and artificial acid rain on the mite (Acari) fauna in coniferous forest. Oikos. 1981;**37**:7-20

[15] Hågvar S. Six common mite species (Acari) in Norwegian coniferous forest soils: Relations to vegetation types and soil characteristics. Pedobiologia. 1984;**27**:355-364

[16] Hågvar S. Why do collemboles and mites react to changes in soil acidity? Entomologiske Meddelelser. 1987;**55**:115-119

[17] Hågvar S. Decomposition studies in an easily-constructed microcosm: Effects of microarthropods and varying soil pH. Pedobiologia. 1988;**31**:293-303

[18] Hågvar S. Reactions to soil acidification in microarthropods: Is competition a key factor? Biology and Fertility of Soils. 1990;**9**:178-181

[19] Hågvar S. Succession of microarthropods in decomposing spruce needles: A litter bag study over ten years. In: Veeresh GK, Rajagopal D, Viraktamath CA, editors. Advances in Management and Conservation of Soil Fauna. Proceedings from the 10th International Soil Zoology Colloquium, Bangalore 7-13 August 1988. New Delhi, India: Oxford & IBH Publishing Co. PVT. LTD; 1991. pp. 485-489

[20] Hågvar S. Soil animals and soil acidity. In: Abrahamsen G, Stuanes AO, Tveite B, editors. Long-Term Experiments with Acid Rain in Norwegian Forest Ecosystems. Ecological Studies. Vol. 104. Berlin: Springer-Verlag; 1993. pp. 101-121

[21] Hågvar S. Instability in small, isolated microarthropod communities. Polskie Pismo Entomologiczne. 1995;**64**:123-133

[22] Hågvar S. Mites (Acari) developing inside decomposing spruce needles: Biology and effect on decomposition rate. Pedobiologia. 1998;**42**:358-377

[23] Hågvar S, Hågvar EB. Invertebrate activity under snow in a south-Norwegian spruce forest. Soil Organisms. 2011;**83**(2):187-209

[24] Hågvar S, Steen R. Succession of beetles (genus *Cis*) and oribatid mites (genus *Carabodes*) in dead sporocarps of the red-banded polypore fungus *Fomitopsis pinicola*. Scandinavian Journal of Forest Research. 2013;**28**:436-444. DOI: 10.1080/02827581.2012.755562

[25] Hågvar S, Amundsen T, Økland B. Mites of the genus *Carabodes* (Acari, Oribatida) in Norwegian coniferous forests: Occurrence in different soils, vegetation types and polypore hosts. Scandinavian Journal of Forest Research. 2014;**29**:629-638. DOI: 10.1080/02827581.2014.965195

[26] Hågvar S. From litter to humus in a Norwegian spruce forest: Long-term studies on the decomposition of needles and cones. Forests. 2016;**7**:186. DOI: 10.3390/f7090186

[27] Hågvar S. Collembola in Norwegian coniferous forest soils I. Relations to plant communities and soil fertility. Pedobiologia. 1982;**24**:255-296

[28] Huhta V, Hyvönen R, Kaasalainen P, Koskenniemi A, Muona J, Mäkelä I, et al. Soil fauna of Finnish coniferous forests. Annales Zoologici Fennici. 1986;**23**:345-360

[29] Siira-Pietikäinen A, Penttinen R, Huhta V. Oribatid mites (Acari: Oribatida) in boreal forest floor and decaying wood. Pedobiologia. 2008;**52**:111-118

[30] Persson T, Bååth E, Clarholm M, Lundkvist H, Söderström BE, Sohlenius B. Trophic structure, biomass dynamics and carbon metabolism of soil organisms in a scots pine forest. Ecological Bulletins. 1980;**32**:419-459

[31] Karppinen E. Ecological and transect survey studies on Finnish Camisiids. Annales Zoologici Societatis Zoologicae Botanicae Fennicae "Vanamo". 1955;**17**(2):1-80

[32] Drift JVD. Analysis of the animal community in a beech forest floor. Tijdschrift voor Entomologie. 1951;**94**:1-168

[33] Karppinen E. Über die Oribatiden (Acar.) der Finnischen Waldboden. Annales Zoologici Societatis Zoologicae Botanicae Fennicae "Vanamo". 1958;**19**(1):1-43

[34] Lebrun P. Écologie et biocénotique de quelques peuplements d'arthropodes édaphiques. Mémoires Institut Royal des Scienses Naturelles de Belgique. 1971;**165**:1-203

**101**

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review*

[43] Abrahamsen G. Ecological study of Enchytraeidae (Oligochaeta) in Norwegian coniferous forest soils. Pedobiologia. 1972;**12**:26-82

[44] Abrahamsen G. Effects of lime and artificial acid rain on the enchytraeid (Oligochaeta) fauna in coniferous forest. Holarctic Ecology. 1983;**6**:247-254

DKMCE, editor. Soil Zoology. London: Butterworths; 1955. pp. 180-201

[46] Abrahamsen G. Ecological study of Lumbricidae (Oligochaeta) in Norwegian coniferous forest soils. Pedobiologia. 1972;**12**:267-281

[47] Nordström S, Rundgren S. Environmental factors and lumbricid associations in southern Sweden. Pedobiologia. 1974;**14**:1-27

[48] Christiansen K. Competition between collembolan species in culture jars. Revue d'écologie et de biologie du

[49] Nygard J, Solberg J. Laboratory study on competition between four soilliving species of Collembola. Thesis, University of Oslo (in Norwegian); 1985

[50] Anderson JM. Competition between two unrelated species of soil cryptostigmata (Acari) in experimental microcosms. Journal of Animal Ecology.

[51] Anderson JM. Succession, diversity and trophic relationships of some soil animals in decomposing leaf litter. Journal of Animal Ecology.

[52] Mignolet R, Lebrun P. Colonisation par les microarthropodes du sol de cinq types de litière en decomposition. In: Vanek J, editor. Progress in Soil Zoology. The Hague, The Netherlands: Dr. W. Junk, B.V; 1975. pp. 261-281

sol. 1967;**4**:439-462

1978;**47**:787-803

1975;**44**:475-495

[45] Satchell JE. Some aspects of earthworm ecology. In: Kevan

*DOI: http://dx.doi.org/10.5772/intechopen.83478*

[35] Forsslund KH. Studier över det lägre djurlivet i nordsvensk skogsmark. Meddelanden från statens skogforsöksanstalt. 1944;**34**(1):1-1283

[36] Persson T. Markarthropodernas abundans, biomassa och respiration i ett äldre tallbestånd på Ivantjärnsheden, Gästrikland - en första studie. Swedish Coniferous Forest Project, Internal Report; 1975. vol. 31. pp. 1-35

[37] Osler GHR, Cole L, Keith AM. Changes in oribatid mite

2006;**50**(4):323-330

1980;**20**:85-100

Pedobiologia. 1961;**1**:6-24

Report 74/80. pp. 1-23

1971;**11**:417-424

community structure associated with the succession from heather (*Calluna vulgaris*) moorland to birch (*Betula pubescens*) woodland. Pedobiologia.

[38] Huhta V, Hyvönen R, Koskenniemi A, Vilkamaa P. Role of pH in the effect of fertilization on Nematoda, Oligochaeta and microarthropods. In: Lebrun P, editor. New Trends in Soil Biology. Proceedings of the VIII. Int. Colloq. of Soil Zoology, Louvain-la-Neuve (Belgium). 1983. pp. 61-73

[39] Bååth E, Berg B, Lohm U, Lundgren B, Lundkvist H, Rosswall T, et al. Effects of experimental acidification and liming on soil organisms and decomposition in a scots pine forest. Pedobiologia.

[40] Bonnet L. Caractères généraux des populations thécamoebiennes endogées.

[41] Stachurska-Hagen T. Acidification experiments in conifer forest. 8. Effects of acidification and liming on some soil animals: Protozoa, Rotifera and Nematoda. SNSF-project. 1980;Internal

[42] Abrahamsen G. The influence of temperature and soil moisture on the population density of

*Cognettia sphagnetorum* (Oligochaeta: Enchytraeidae) in cultures with

homogenized raw humus. Pedobiologia.

*Ecological Spotlights on Mites (Acari) in Norwegian Conifer Forests: A Review DOI: http://dx.doi.org/10.5772/intechopen.83478*

[35] Forsslund KH. Studier över det lägre djurlivet i nordsvensk skogsmark. Meddelanden från statens skogforsöksanstalt. 1944;**34**(1):1-1283

*Pests Control and Acarology*

pp. 485-489

1995;**64**:123-133

[19] Hågvar S. Succession of microarthropods in decomposing spruce needles: A litter bag study over ten years. In: Veeresh GK, Rajagopal D, Viraktamath CA, editors. Advances in Management and Conservation of Soil Fauna. Proceedings from the 10th International Soil Zoology Colloquium, Bangalore 7-13 August 1988. New Delhi, India: Oxford & IBH Publishing Co. PVT. LTD; 1991.

[26] Hågvar S. From litter to humus in a Norwegian spruce forest: Long-term studies on the decomposition of needles and cones. Forests. 2016;**7**:186. DOI:

[27] Hågvar S. Collembola in Norwegian coniferous forest soils I. Relations to plant communities and soil fertility. Pedobiologia. 1982;**24**:255-296

[28] Huhta V, Hyvönen R, Kaasalainen P, Koskenniemi A, Muona J, Mäkelä I, et al. Soil fauna of Finnish coniferous forests. Annales Zoologici Fennici.

[29] Siira-Pietikäinen A, Penttinen R, Huhta V. Oribatid mites (Acari: Oribatida) in boreal forest floor and decaying wood. Pedobiologia.

[30] Persson T, Bååth E, Clarholm M, Lundkvist H, Söderström BE, Sohlenius B. Trophic structure, biomass dynamics and carbon metabolism of soil organisms in a scots pine forest. Ecological Bulletins.

[31] Karppinen E. Ecological and transect survey studies on Finnish Camisiids. Annales Zoologici Societatis

Zoologicae Botanicae Fennicae "Vanamo". 1955;**17**(2):1-80

[32] Drift JVD. Analysis of the animal community in a beech forest floor. Tijdschrift voor Entomologie.

Botanicae Fennicae "Vanamo".

[33] Karppinen E. Über die Oribatiden (Acar.) der Finnischen Waldboden. Annales Zoologici Societatis Zoologicae

[34] Lebrun P. Écologie et biocénotique de quelques peuplements d'arthropodes édaphiques. Mémoires Institut Royal des Scienses Naturelles de Belgique.

10.3390/f7090186

1986;**23**:345-360

2008;**52**:111-118

1980;**32**:419-459

1951;**94**:1-168

1958;**19**(1):1-43

1971;**165**:1-203

[20] Hågvar S. Soil animals and soil acidity. In: Abrahamsen G, Stuanes AO, Tveite B, editors. Long-Term Experiments with Acid Rain in Norwegian Forest Ecosystems. Ecological Studies. Vol. 104. Berlin: Springer-Verlag; 1993. pp. 101-121

[21] Hågvar S. Instability in small, isolated microarthropod communities. Polskie Pismo Entomologiczne.

[22] Hågvar S. Mites (Acari) developing inside decomposing spruce needles: Biology and effect on decomposition rate. Pedobiologia. 1998;**42**:358-377

[23] Hågvar S, Hågvar EB. Invertebrate

activity under snow in a south-Norwegian spruce forest. Soil Organisms. 2011;**83**(2):187-209

[24] Hågvar S, Steen R. Succession of beetles (genus *Cis*) and oribatid mites (genus *Carabodes*) in dead sporocarps of the red-banded polypore fungus *Fomitopsis pinicola*. Scandinavian Journal of Forest Research. 2013;**28**:436-444. DOI: 10.1080/02827581.2012.755562

[25] Hågvar S, Amundsen T, Økland B. Mites of the genus *Carabodes* (Acari, Oribatida) in Norwegian coniferous forests: Occurrence in different soils, vegetation types and polypore hosts. Scandinavian Journal of Forest Research. 2014;**29**:629-638. DOI: 10.1080/02827581.2014.965195

**100**

[36] Persson T. Markarthropodernas abundans, biomassa och respiration i ett äldre tallbestånd på Ivantjärnsheden, Gästrikland - en första studie. Swedish Coniferous Forest Project, Internal Report; 1975. vol. 31. pp. 1-35

[37] Osler GHR, Cole L, Keith AM. Changes in oribatid mite community structure associated with the succession from heather (*Calluna vulgaris*) moorland to birch (*Betula pubescens*) woodland. Pedobiologia. 2006;**50**(4):323-330

[38] Huhta V, Hyvönen R, Koskenniemi A, Vilkamaa P. Role of pH in the effect of fertilization on Nematoda, Oligochaeta and microarthropods. In: Lebrun P, editor. New Trends in Soil Biology. Proceedings of the VIII. Int. Colloq. of Soil Zoology, Louvain-la-Neuve (Belgium). 1983. pp. 61-73

[39] Bååth E, Berg B, Lohm U, Lundgren B, Lundkvist H, Rosswall T, et al. Effects of experimental acidification and liming on soil organisms and decomposition in a scots pine forest. Pedobiologia. 1980;**20**:85-100

[40] Bonnet L. Caractères généraux des populations thécamoebiennes endogées. Pedobiologia. 1961;**1**:6-24

[41] Stachurska-Hagen T. Acidification experiments in conifer forest. 8. Effects of acidification and liming on some soil animals: Protozoa, Rotifera and Nematoda. SNSF-project. 1980;Internal Report 74/80. pp. 1-23

[42] Abrahamsen G. The influence of temperature and soil moisture on the population density of *Cognettia sphagnetorum* (Oligochaeta: Enchytraeidae) in cultures with homogenized raw humus. Pedobiologia. 1971;**11**:417-424

[43] Abrahamsen G. Ecological study of Enchytraeidae (Oligochaeta) in Norwegian coniferous forest soils. Pedobiologia. 1972;**12**:26-82

[44] Abrahamsen G. Effects of lime and artificial acid rain on the enchytraeid (Oligochaeta) fauna in coniferous forest. Holarctic Ecology. 1983;**6**:247-254

[45] Satchell JE. Some aspects of earthworm ecology. In: Kevan DKMCE, editor. Soil Zoology. London: Butterworths; 1955. pp. 180-201

[46] Abrahamsen G. Ecological study of Lumbricidae (Oligochaeta) in Norwegian coniferous forest soils. Pedobiologia. 1972;**12**:267-281

[47] Nordström S, Rundgren S. Environmental factors and lumbricid associations in southern Sweden. Pedobiologia. 1974;**14**:1-27

[48] Christiansen K. Competition between collembolan species in culture jars. Revue d'écologie et de biologie du sol. 1967;**4**:439-462

[49] Nygard J, Solberg J. Laboratory study on competition between four soilliving species of Collembola. Thesis, University of Oslo (in Norwegian); 1985

[50] Anderson JM. Competition between two unrelated species of soil cryptostigmata (Acari) in experimental microcosms. Journal of Animal Ecology. 1978;**47**:787-803

[51] Anderson JM. Succession, diversity and trophic relationships of some soil animals in decomposing leaf litter. Journal of Animal Ecology. 1975;**44**:475-495

[52] Mignolet R, Lebrun P. Colonisation par les microarthropodes du sol de cinq types de litière en decomposition. In: Vanek J, editor. Progress in Soil Zoology. The Hague, The Netherlands: Dr. W. Junk, B.V; 1975. pp. 261-281

[53] Mignolet R. Écologie des microarthropodes associés à la decomposition des litières de forêt. Thesis, Univ. Catholique de Louvain. 1975. 136 pp

[54] Franklin E, Hayek T, Fagundes EP, Silva LL. Oribatid mite (Acari: Oribatida) contribution to decomposition dynamic of leaf litter in primary forest, second growth, and polyculture in the Central Amazon. Brazilian Journal of Biology. 2004;**64**. DOI: 10.1590/ S1519-69842004000100008

[55] Fujii S, Takeda H. Succession of soil microarthropod communities during the aboveground and belowground litter decomposition processes. Soil Biology and Biochemistry. 2017;**110**:95-102

[56] Pande YD, Berthet P. Studies on the food and feeding habits of soil Oribatei in a black pine plantation. Oecologia. 1973;**12**:413-426

[57] Ponge J-F. Succession of fungi and fauna during decomposition of needles in a small area of scots pine litter. Plant and Soil. 1991;**138**(1):99-113

[58] Coulianos C-C, Johnels AG. Note on the subnivean environment of small mammals. Arkiv för Zoologi. 1962;**Ser.2**:363-370

[59] Gulvik M. Mites (Acari) as indicators of soil biodiversity and land use monitoring: A review. Polish Journal of Ecology. 2007;**55**:415-440

[60] Hågvar S. The relevance of the Rio-convention on biodiversity to conserving the biodiversity of soils. Applied Soil Ecology. 1998;**9**:1-7

[61] Hågvar S. The Huge Biodiversity of Soils: Can it be Saved? Agricultural University of Iceland, Hvanneyri. AUI Publication. 2006;**9**:61-66

**103**

**Chapter 7**

**Abstract**

**1. Introduction**

confusion about ecological concepts [4].

Invasive Mite Species in the

*Carlos Vásquez and Yelitza Colmenárez*

Americas: Bioecology and Impact

Invasive species represent one of the most relevant threats for biodiversity in many ecosystems, mainly in those so-called agroecosystems due to which they exhibit reduced biodiversity and simplified trophic interactions. These two factors make many niches unoccupied, thus increasing the risk that invasive species especially arthropod pests occupy these niches or compete with native species. In spite of potential impact of invasive species, our understanding of their ecological consequences is developing slowly. In the last years, more attention is being paid on phytophagous mites because several noneconomic species have become severe pests on many crops as a consequence of irrational use of agrochemicals. Also, due to the small size of the mites, they can be transported throughout the world and established in new areas where favorable conditions and the absence of efficient natural enemies favor their development. Thus, phytophagous mites are feasible to become invasive species since they are able to provoke severe damage to plants. Since 2004, *Steneotarsonemus spinki*, *Schizotetranychus hindustanicus,* and *Raoiella indica* have been introduced in the Neotropical region. Information about pest status, seasonal

trends, and natural enemies in invaded areas is provided for these species.

Natural environments are continuously submitted to severe transformations, including movement of species beyond the limits of their native geographic ranges into areas where they do not naturally occur and where they can inflict substantial changes [1]. Thus, considering changes inflicted by alien species to the properties of an ecosystem, an increasing number of studies that consider the environmental impacts have been published [2]. However, according to Ricciardi [3], a predictive understanding of the ecological impacts of invasive species has developed slowly, owing largely to an apparent lacking of clearly defined hypotheses and of a broad theoretical framework. In this regard, confusion about terminology used for the designation of nonindigenous species, which alternatively have been called "exotic," "introduced," "invasive," and "naturalized," is particularly acute, which leads to

Another term needing delimitation of definition is referred to concept of impact. On an ecological basis, an impact is defined as a measurable change to the properties of an ecosystem by a nonnative species, which is considered to provoke a positive or negative impact simply by becoming integrated into the system [3].

**Keywords:** phytophagous mites, invasive species, Neotropical region, *Steneotarsonemus spinki*, *Schizotetranychus hindustanicus*, *Raoiella indica*

## **Chapter 7**

*Pests Control and Acarology*

1975. 136 pp

[53] Mignolet R. Écologie des microarthropodes associés à la decomposition des litières de forêt. Thesis, Univ. Catholique de Louvain.

[54] Franklin E, Hayek T, Fagundes

[55] Fujii S, Takeda H. Succession of soil microarthropod communities during the aboveground and belowground litter decomposition processes. Soil Biology and Biochemistry. 2017;**110**:95-102

[56] Pande YD, Berthet P. Studies on the food and feeding habits of soil Oribatei in a black pine plantation. Oecologia.

[57] Ponge J-F. Succession of fungi and fauna during decomposition of needles in a small area of scots pine litter. Plant

[58] Coulianos C-C, Johnels AG. Note on the subnivean environment of small mammals. Arkiv för Zoologi.

indicators of soil biodiversity and land use monitoring: A review. Polish Journal

[60] Hågvar S. The relevance of the Rio-convention on biodiversity to conserving the biodiversity of soils. Applied Soil Ecology. 1998;**9**:1-7

[61] Hågvar S. The Huge Biodiversity of Soils: Can it be Saved? Agricultural University of Iceland, Hvanneyri. AUI

and Soil. 1991;**138**(1):99-113

[59] Gulvik M. Mites (Acari) as

of Ecology. 2007;**55**:415-440

Publication. 2006;**9**:61-66

1973;**12**:413-426

1962;**Ser.2**:363-370

EP, Silva LL. Oribatid mite (Acari: Oribatida) contribution to decomposition dynamic of leaf litter in primary forest, second growth, and polyculture in the Central Amazon. Brazilian Journal of Biology. 2004;**64**. DOI: 10.1590/ S1519-69842004000100008

**102**

## Invasive Mite Species in the Americas: Bioecology and Impact

*Carlos Vásquez and Yelitza Colmenárez*

## **Abstract**

Invasive species represent one of the most relevant threats for biodiversity in many ecosystems, mainly in those so-called agroecosystems due to which they exhibit reduced biodiversity and simplified trophic interactions. These two factors make many niches unoccupied, thus increasing the risk that invasive species especially arthropod pests occupy these niches or compete with native species. In spite of potential impact of invasive species, our understanding of their ecological consequences is developing slowly. In the last years, more attention is being paid on phytophagous mites because several noneconomic species have become severe pests on many crops as a consequence of irrational use of agrochemicals. Also, due to the small size of the mites, they can be transported throughout the world and established in new areas where favorable conditions and the absence of efficient natural enemies favor their development. Thus, phytophagous mites are feasible to become invasive species since they are able to provoke severe damage to plants. Since 2004, *Steneotarsonemus spinki*, *Schizotetranychus hindustanicus,* and *Raoiella indica* have been introduced in the Neotropical region. Information about pest status, seasonal trends, and natural enemies in invaded areas is provided for these species.

**Keywords:** phytophagous mites, invasive species, Neotropical region, *Steneotarsonemus spinki*, *Schizotetranychus hindustanicus*, *Raoiella indica*

#### **1. Introduction**

Natural environments are continuously submitted to severe transformations, including movement of species beyond the limits of their native geographic ranges into areas where they do not naturally occur and where they can inflict substantial changes [1]. Thus, considering changes inflicted by alien species to the properties of an ecosystem, an increasing number of studies that consider the environmental impacts have been published [2]. However, according to Ricciardi [3], a predictive understanding of the ecological impacts of invasive species has developed slowly, owing largely to an apparent lacking of clearly defined hypotheses and of a broad theoretical framework. In this regard, confusion about terminology used for the designation of nonindigenous species, which alternatively have been called "exotic," "introduced," "invasive," and "naturalized," is particularly acute, which leads to confusion about ecological concepts [4].

Another term needing delimitation of definition is referred to concept of impact. On an ecological basis, an impact is defined as a measurable change to the properties of an ecosystem by a nonnative species, which is considered to provoke a positive or negative impact simply by becoming integrated into the system [3].

Various studies have shown that nonnative species can promote extinction of native species, and also they can provoke changes in genetic composition of native populations, behavior patterns, species richness and abundance, phylogenetic and taxonomic diversity, etc.

Finally, when considering invasive species in an agricultural ambit, it is strongly recommended to define invasion threat, which is conceptualized as the likelihood of a particular pest or pathogen arriving in a new location as well as the establishment likelihood considered as the chances of those pests or pathogens to establish in a new location [5].

Plant and animal species have been transported by humans for millennia; even a well-defined period in biological invasions dates as far back as 1500 AD, a period associated with the birth of colonialism and the start of radical changes in patterns of human demography, agriculture, trade, and industry [6]. However, more recently, increasing globalization and world trade have augmented the possibility of arrival of invasive species to geographic regions in which they were previously absent, making necessary to quantify impact of invasive species and develop effective biosecurity policy [4].

Since the end of the twentieth century, more attention is being paid on phytophagous mites because several noneconomic species have become severe pests on many crops as a consequence of irrational use of agrichemicals. Also, due to the small size of the mites, they can be transported throughout the world and set up in new areas, in which favorable conditions and the lack of efficient natural enemies favor their development, resulting in economic losses [7]. There are various examples of introductions of phytophagous mites in new areas such as the cassava green mite [*Mononychellus tanajoa* (Bondar, 1938)], the coconut mite [*Aceria guerreronis* (Keifer, 1965)], and the tomato spider mite [*Tetranychus evansi* Baker and Pritchard, 1960] [7, 8]. Both *M. tanajoa* and *A. guerreronis* were introduced into Africa, while *T. evansi* has been introduced in Africa in the Mediterranean Basin.

Similarly, some phytophagous mite species have been introduced in the Neotropical region, i.e., *Steneotarsonemus spinki* (Smiley, 1967), *Schizotetranychus hindustanicus* (Hirst, 1924), and *Raoiella indica* (Hirst, 1924). In the present review, information about recent phytophagous mite on pest status, seasonal trends, and natural enemies in invaded areas is provided for these species. Because invasive species may evolve during the invasion process, comparison of behavior, and damage and management options between native and invaded areas for these species will be useful for understanding the invader's success and their ability to colonize new regions.

#### **2. Some concepts related to invasive species**

The introduction of species beyond their native range as a direct or indirect result of human action causes changes in the ecosystems to which they are introduced [9]. Moreover, these biological invasions are causing tremendous damages to ecosystems and economic activities [10]. Many important terms related to the invasion ecology, such as "invasive," "weed," or "transient," can be susceptible to subjective interpretation, consequently causing a lack of consensus about terms used to define nonindigenous species [4]. Thus, some terms such as "noxious" and "nuisance" are generally used to indicate direct or indirect adverse effects on humans; however, according to Colautti and MacIsaac [4], interactions have three important implications:

a.It is necessary to define if invasive species cause aesthetical displeasing effects and are vectors for serious human diseases.

**105**

*Invasive Mite Species in the Americas: Bioecology and Impact*

b.Species might be erroneously considered as an environmental threat (or weedy, invasive, etc.) in areas where they have little or no impact only based on these species had been identified as a disturbance elsewhere, disregarding thus the

*Harmonia axyridis* (Pallas, 1773) (Coleoptera, Coccinellidae) is a well-known predator widely used as a classical biological control agent of aphids around the world; however, this species has provoked the displacement of native aphid

c.A particular species can have both beneficial and detrimental effects.

a.An alien species whose introduction does or is likely to cause economic or

b.A species that is nonnative to the ecosystem under consideration and whose introduction causes or is likely to cause economic or environmental harm or

Invasion consists of a series of steps that an organism must undergo to become a successful invader, and thus, it can inflict an ecological damage [12]. According to

a.*Large-scale geographical barriers*: species from a geographical area is supposed to overcome a geographical barrier (mountain range, ocean, or other physical barriers) to arrive in a new area where it does not previously occur (so-called alien species, nonnative species). This movement is often mediated by human

b.*Survival barriers*: these often refer to environmental barriers such as environmental conditions that let the nonnative organism to survive and develop in its new location. Other survival barriers may include host plants, competitor

c.*Establishment barriers*: depending on the survival abilities of the alien species, it will be able to form a self-sustaining population and does not need a re-

d.*Dispersal and spread barriers*: once established, alien species must disperse and spread relatively fast from their site of establishment. However, this movement or spread alone does not necessarily make this nonnative species an invasive species.

Although bioinvasions have occurred for many years, most documented cases have been reported in recent decades and even invasive or adventive mites have gained attention only in the last few years when they have been the target of research to determine their potential distribution [7]. After colonization, it is fundamental to determine the intrinsic properties of the invasive population, the genetic structure of populations, and the response to environmental factors to

species in Brazil and other South American countries [11].

Beck et al. [13], successful invasion is preceded by the following stages:

Some definitions of invasive species are the following:

environmental harm or harm to human health.

activities, either deliberately or unintentionally.

organisms, predators, and pathogens.

introduction to support a population base.

**3. Mite invasions in the Neotropical region**

developing strategies and policies of management [7].

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

ecological phenomenon.

harm to human health

*Pests Control and Acarology*

taxonomic diversity, etc.

tive biosecurity policy [4].

a new location [5].

Various studies have shown that nonnative species can promote extinction of native species, and also they can provoke changes in genetic composition of native populations, behavior patterns, species richness and abundance, phylogenetic and

Finally, when considering invasive species in an agricultural ambit, it is strongly recommended to define invasion threat, which is conceptualized as the likelihood of a particular pest or pathogen arriving in a new location as well as the establishment likelihood considered as the chances of those pests or pathogens to establish in

Plant and animal species have been transported by humans for millennia; even a well-defined period in biological invasions dates as far back as 1500 AD, a period associated with the birth of colonialism and the start of radical changes in patterns of human demography, agriculture, trade, and industry [6]. However, more recently, increasing globalization and world trade have augmented the possibility of arrival of invasive species to geographic regions in which they were previously absent, making necessary to quantify impact of invasive species and develop effec-

Since the end of the twentieth century, more attention is being paid on phytophagous mites because several noneconomic species have become severe pests on many crops as a consequence of irrational use of agrichemicals. Also, due to the small size of the mites, they can be transported throughout the world and set up in new areas, in which favorable conditions and the lack of efficient natural enemies favor their development, resulting in economic losses [7]. There are various examples of introductions of phytophagous mites in new areas such as the cassava green mite [*Mononychellus tanajoa* (Bondar, 1938)], the coconut mite [*Aceria guerreronis* (Keifer, 1965)], and the tomato spider mite [*Tetranychus evansi* Baker and Pritchard, 1960] [7, 8]. Both *M. tanajoa* and *A. guerreronis* were introduced into Africa, while

*T. evansi* has been introduced in Africa in the Mediterranean Basin.

**2. Some concepts related to invasive species**

and are vectors for serious human diseases.

Similarly, some phytophagous mite species have been introduced in the Neotropical region, i.e., *Steneotarsonemus spinki* (Smiley, 1967), *Schizotetranychus hindustanicus* (Hirst, 1924), and *Raoiella indica* (Hirst, 1924). In the present review, information about recent phytophagous mite on pest status, seasonal trends, and natural enemies in invaded areas is provided for these species. Because invasive species may evolve during the invasion process, comparison of behavior, and damage and management options between native and invaded areas for these species will be useful for understanding the invader's success and their ability to colonize new regions.

The introduction of species beyond their native range as a direct or indirect result of human action causes changes in the ecosystems to which they are introduced [9]. Moreover, these biological invasions are causing tremendous damages to ecosystems and economic activities [10]. Many important terms related to the invasion ecology, such as "invasive," "weed," or "transient," can be susceptible to subjective interpretation, consequently causing a lack of consensus about terms used to define nonindigenous species [4]. Thus, some terms such as "noxious" and "nuisance" are generally used to indicate direct or indirect adverse effects on humans; however, according to Colautti and MacIsaac [4], interactions have three

a.It is necessary to define if invasive species cause aesthetical displeasing effects

**104**

important implications:


Some definitions of invasive species are the following:


Invasion consists of a series of steps that an organism must undergo to become a successful invader, and thus, it can inflict an ecological damage [12]. According to Beck et al. [13], successful invasion is preceded by the following stages:


## **3. Mite invasions in the Neotropical region**

Although bioinvasions have occurred for many years, most documented cases have been reported in recent decades and even invasive or adventive mites have gained attention only in the last few years when they have been the target of research to determine their potential distribution [7]. After colonization, it is fundamental to determine the intrinsic properties of the invasive population, the genetic structure of populations, and the response to environmental factors to developing strategies and policies of management [7].

In the Neotropical region, several mite species have recently invaded the agricultural landscapes in Latin America, for example, the citrus Hindu mite, *S. hindustanicus* (Tetranychidae), the rice mite, *S. spinki* (Tarsonemidae), and the red palm mite, *R. indica* (Tenuipalpidae).

## **3.1** *Schizotetranychus hindustanicus*

The genus *Schizotetranychus* includes 114 species; however, information about economic importance of most of the species is still scarce [14, 15] (**Table 1**). Most of the species occur in Asia and CIS, and only 20 species (17.5%) are in the Neotropical region, including *S. hindustanicus*.

The citrus Hindu mite, *S. hindustanicus*, was originally described from citrus from southern India (Hirst, 1924), and its occurrence had been reported in this country for almost 80 years; however, in 2005, this species was surprisingly found in the northwestern Venezuela [16] and soon after in Colombia and Brazil [17].

*S. hindustanicus* had only been reported on four host plant species in India (see **Table 1**); however, posteriorly, it was found on *Acacia* sp., *Melia azedarach* L. and various *citrus* species (**Figure 1**). Symptoms of mite feeding first appear on the upper leaf surface, along the main rib, later extending to the entire leaf; while when feeding on fruits the females webs over concavities or depressions in the rind; attacked fruits become uniformly silvered and hard under severe infestation [18]. Návia and Marsaro [17] reported that although this damage by mite feeding is supposed to affect the commercial value of infested fruits, nothing has been published about the resulting economic impact.

In Venezuela, *S. hindustanicus* has been observed forming colonies in several *citrus* species and/or varieties such as *C. latifolia* (Tanaka ex Yu. Tanaka), *C. aurantifolia* (Chistm) *C. reticulata* Blanco, *C. limon* (L.), and *C. sinensis* (L.) Osbeck [19, 20].

In Colombia, this tetranychid mite species was first reported in the northern coast in Dibulla (Guajira) and Magdalena [21]. After that, ICA (Agropecuary Colombian Institute) carried out samplings in departments of Atlantico, Bolivar, Guajira, Magdalena, and Vichada as shown in **Table 2**. Similarly, presence of circular whitish spots on leaves and fruits of "tahiti" and "galeguinho" lemon trees in urban areas of Roraima (Brazil) is alerted to the Brazilian plant protection authorities as this country is the largest *citrus* producer [17]. According to these authors, dispersion of *S. hindustanicus* could cause high economic impact and/or commercial restrictions due to sanitary.

Since these tetranychid mite species can be the pest on *citrus* spp., some studies have been carried out in Venezuela. Niedstaedt and Marcano [15] observed the effect that the developmental time of *S. hindustanicus* varied from 30.12 to 31.10 days on sweet orange or Persian lime, respectively, at 25°C. Additionally, population studies on Persian lime, lemon, sweet orange, and tangerine showed that number of individuals was relatively low in two peaks: the first peak during June 2005 with 24.17, 21.67, and 12 individuals was observed on tangerine, sweet orange, and Persian lime, respectively, while the second peak with lightly higher number of mites developed during April 2006 was observed with 69.17, 31.2, and 20.2 mites on sweet orange, tangerine, and Persian lime, respectively [22].

Field observations on different citrus species have demonstrated that *S. hindustanicus* can colonize which seems to be verifying the entire canopy so far economic impact have not been evaluated in the neotropical areas. There are some studies on this genus, associated mainly with grasses such as rice and bamboo and some fruit trees [23, 24].

**107**

*Invasive Mite Species in the Americas: Bioecology and Impact*

*siamensis Gamble*

*Pteridophyta*

Kurz

*Stipa eminens* Cav.

Zeyher ex Steudel

**Species Host plants Distribution**

*Andropogon annulatus* Forssk.*, Chloris incomplete* Roth*, Dichanthium annulatum* (Forssk.) Stapf*, Oryza sativa* L.

*Bambusa vulgaris* Schrad. ex J.C. Wendl.*, Thyrsostachys* 

*Acacia horrida* (L.) Willd.*, A. longifolia* (Andrews) Willd*., Ananas* sp., *Aspalathus* sp., *Asparagus* sp., *A. africanus* Lam.*, A. officinalis* L.*, A. plumosus* Baker*, A. setaceus* (Kunth) Jessop*, A. sprengeri* Regel*, A. suaveolens* Burch.*, Protasparagus capensis* (L.) Oberm*., P. compactus* (T.M. Salter) Oberm.*, P. laricinus* (Burch.) Oberm.*,* 

*Mundulea pungens* R. Vig.*, Tephrosia striata* Ecklon &

*Citrus grandis* (L.) Osbeck*, C. madurensis* Lour.*, C.* 

*Arundinaria* sp. *Phyllostachys* sp., *P. nigra* (*Lodd. ex Lindl.*) *Munro, P. reticulata* (Rupr.) K. Koch

*Cajanus cajan* (L.) Huth*, Cassia* sp., *C. siamea* Lam.*, Colocasia esculenta* (L.) Schott*, Pterocarpus macrocarpus*

*Bouteloua rothrockii* Vasey*¸Commelina dianthifolia* L.*,* 

*medica* L.*, C. sinensis* (L.) Osbeck

*Spiraea* sp. CIS

*Calamagrostis* sp., *Dactylis* sp., *Helictotrichon* sp. CIS

*Brachypodium silvaticum* (Huds.) P. Beauv. CIS

*Quercus* sp., *Q. glauca* Thunb. Japan

*C. cajan* India

*Agropyron desertorum* (Fisch. ex Link) Schult. The USA

*Euclea crispa* (Thunb.) Gürke South Africa

CIS, India, Mexico, Pakistan, and Thailand

Australia, Germany, Hawaii, Israel, Morocco, Portugal, Puerto Rico, South Africa, the Netherlands, and the USA

Madagascar

Thailand

Mexico and the USA

Burma, China, Hong Kong, India, Philippines, Taiwan, and Thailand

CIS, China, Hainan Island, Japan, and Korea

Malaysia

*Alnus* sp. CIS

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

*S. agropyron* (Tuttle and Baker, 1976)

*S. andropogoni* (Hirst, 1926)

*S. approximatus* (Ehara, 1988)

*S. arcuatus* (Meyer, 1974)

*S. asparagi* (Oudemans, 1928)

*S. australis* (Gutierrez, 1968)

*S. avetjanae* (Bagdasarian, 1954)

*S. baltazari* (Rimando, 1962)

*S. bambusae* (Reck, 1941)

*S. beckeri* (Wainstein, 1958)

*bhandhufalcki* (Ehara and Wongsiri, 1975)

*S. boutelouae* (Tuttle and Baker, 1968)

*S. brachypodii* (Livshits and Mitrofanov, 1968)

*S. brevisetosus* (Ehara, 1989)

*S. cajani* (Gupta, 1976)

*S.* 

*S. alni* (Beglyarov and Mitrofanov, 1973)

*Pests Control and Acarology*

mite, *R. indica* (Tenuipalpidae).

**3.1** *Schizotetranychus hindustanicus*

region, including *S. hindustanicus*.

about the resulting economic impact.

Brazil [17].

Osbeck [19, 20].

In the Neotropical region, several mite species have recently invaded the agricultural landscapes in Latin America, for example, the citrus Hindu mite, *S. hindustanicus* (Tetranychidae), the rice mite, *S. spinki* (Tarsonemidae), and the red palm

The genus *Schizotetranychus* includes 114 species; however, information about economic importance of most of the species is still scarce [14, 15] (**Table 1**). Most of the species occur in Asia and CIS, and only 20 species (17.5%) are in the Neotropical

The citrus Hindu mite, *S. hindustanicus*, was originally described from citrus from southern India (Hirst, 1924), and its occurrence had been reported in this country for almost 80 years; however, in 2005, this species was surprisingly found in the northwestern Venezuela [16] and soon after in Colombia and

*S. hindustanicus* had only been reported on four host plant species in India (see **Table 1**); however, posteriorly, it was found on *Acacia* sp., *Melia azedarach* L. and various *citrus* species (**Figure 1**). Symptoms of mite feeding first appear on the upper leaf surface, along the main rib, later extending to the entire leaf; while when feeding on fruits the females webs over concavities or depressions in the rind; attacked fruits become uniformly silvered and hard under severe infestation [18]. Návia and Marsaro [17] reported that although this damage by mite feeding is supposed to affect the commercial value of infested fruits, nothing has been published

In Venezuela, *S. hindustanicus* has been observed forming colonies in several

In Colombia, this tetranychid mite species was first reported in the northern coast in Dibulla (Guajira) and Magdalena [21]. After that, ICA (Agropecuary Colombian Institute) carried out samplings in departments of Atlantico, Bolivar, Guajira, Magdalena, and Vichada as shown in **Table 2**. Similarly, presence of circular whitish spots on leaves and fruits of "tahiti" and "galeguinho" lemon trees in urban areas of Roraima (Brazil) is alerted to the Brazilian plant protection authorities as this country is the largest *citrus* producer [17]. According to these authors, dispersion of *S. hindustanicus* could cause high economic impact and/or commercial restrictions due to sanitary. Since these tetranychid mite species can be the pest on *citrus* spp., some studies have been carried out in Venezuela. Niedstaedt and Marcano [15] observed the effect that the developmental time of *S. hindustanicus* varied from 30.12 to 31.10 days on sweet orange or Persian lime, respectively, at 25°C. Additionally, population studies on Persian lime, lemon, sweet orange, and tangerine showed that number of individuals was relatively low in two peaks: the first peak during June 2005 with 24.17, 21.67, and 12 individuals was observed on tangerine, sweet orange, and Persian lime, respectively, while the second peak with lightly higher number of mites developed during April 2006 was observed with 69.17, 31.2, and 20.2 mites on

Field observations on different citrus species have demonstrated that *S. hindustanicus* can colonize which seems to be verifying the entire canopy so far economic impact have not been evaluated in the neotropical areas. There are some studies on this genus, associated mainly with grasses such as rice and bamboo and some fruit trees [23, 24].

*citrus* species and/or varieties such as *C. latifolia* (Tanaka ex Yu. Tanaka), *C. aurantifolia* (Chistm) *C. reticulata* Blanco, *C. limon* (L.), and *C. sinensis* (L.)

sweet orange, tangerine, and Persian lime, respectively [22].

**106**



**109**

*Invasive Mite Species in the Americas: Bioecology and Impact*

G. Benn. & Seem.

Sm.*, S. tristis* Aiton

Scribn.

W.R.B. Oliv.

*Sorghum bicolor* (L.) Moench

**Species Host plants Distribution**

*Aster filifolius* Vent. South Africa

*O. sativa* Costa Rica

*Gahnia aspera* Spreng. Australia

Unknown Zaire

Poaceae CIS

*Molinia caerulea* Milk. France and the

*Cassia nictitans* L. Guatemala

*Halimodendron halodendron* (Pall.) Druce CIS

*Quercus* sp. CIS

*Imperata* sp. China

*Quercus* sp. CIS

Bambusaceae Malaysia

*A. adscensionis, Hilaria rigida* (Thurb.) Benth. ex

*Calopogonium mucunoides* Desv.*, Cordyline kaspar*

*Azadirachta indica* A. Juss.*, Citrus* sp., *Cocos nucifera* L.*,* 

*Arundo formosana* Hack.*, Bambusa* sp., *B. spinosa* Roxb. Philippines, Taiwan

New Caledonia

India, USA

Iran, Poland, Switzerland,

and the USA

Netherlands

USA

India

Hainan Island and New

Zealand

*Ficus edulis* Burm. f.*, F. fraseri* Miq.*, F. habrophylla*

*A. adscensionis, C. cajan, Epicampes rigens* Benth.*,* 

*Acer* sp., *Populus tremula* L.*, Quercus* sp., *Q. robur* L.*, Salix* sp., *S. caprea* L.*, S. humilis* Marshall*, S. petiolaris*

*Muhlenbergia rigens* (Benth.) Hitchc.

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

*S. fauveli* (Gutierrez, 1978)

*S. filifolius* (Meyer, 1974)

*S. floresi* (Rimando, 1972)

*S. fluvialis* (McGregor, 1928)

*S. freitezi* (Ochoa, Gray and von Lind., 1990)

*S. gahniae* (Davis, 1969)

*S. garmani* (Pritchard and Baker, 1955)

*S. gausus* (Baker and Pritchard, 1960)

*S. glabrisetus* (Ugarov and Nikolskii, 1937)

*S. graminicola* (Goux, 1949)

*S. guatemalaenovae* (Stoll, 1886)

*S. halimodendri* (Waistein, 1958)

*S. hindustanicus* (Hirst, 1924)

*S. hilariae* (Tuttle and Baker, 1968)

*S. ibericus* (Reck, 1947)

*S. imperatae* (Wang, 1983)

*S. jachontovi* (Reck, 1953)

*S. kaspari* (Manson, 1967)

*S. kochummeni* (Ehara, 1988)


*Pests Control and Acarology*

*S. camur* (Pritchard and Baker, 1955)

*S. celarius* (Banks, 1917)

*S. celtidis* (Tuttle and Baker, 1968)

*S.* 

*S. cercidiphylli* (Ehara, 1973)

*chiangmaiensis* (Ehara and Wongsiri, 1975)

*S. colocasiae* (Ehara, 1988)

*S. cornus* (Pritchard and Baker, 1955)

*S. cynodontis* (McGregor, 1950)

*S. dalbergia* (Meyer, 1974)

*S. denmarki* (Baker and Tuttle, 1994)

*S. echinulatus* (Mitrafanov, 1978)

*S. elongates* (Wang and Cui, 1991)

*S. elymus* (McGregor, 1950)

*S. emeiensis* (Wang, 1983)

*S. eremophilus* (McGregor, 1950)

*S. euphorbiae* (Livshits and Mitrofanov, 1968)

Hitchc.

**Species Host plants Distribution**

*Arundinaria hindsii* Munro*, Bambusa* sp., *Ficus stipulata* Thunb.*, Miscanthus sinensis* Andersson*, Oryza* sp., *Phyllostachys* sp., *P. makinoi* Hayata*, P. nigra* (Lodd. ex Lindl.) Munro*, P. reticulata* (Rupr.) K. Koch

*Celtis reticulata* Torr*., Leptochloa uninervia* (J. Presl) Hitchc. & Chase*, Sporobolus flexuosus* (Thurb. ex Vasey)

*Dysoxylum spectabile* Hook. f.*, Elaeocarpus dentatus*

(J.R. Forst. and G. Forst.) Vahl

Rydb., *Tridens pulchellus* (Kunth) Hitchc.

*Arundinaria* sp. USA

*Cercidiphyllum japonicum* Siebold & Zucc. Japan

*Calotropis gigantea* (L.) W.T. Aiton Thailand

*Colocasia* sp. Malaysia

*Agrostis* sp., *Cynodon dactylon* (L.) Pers. USA

Poaceae USA

*Spiraea* sp. CIS

Bambusaceae China

Bambusaceae China

*Euphorbia amygdaloides* L. CIS

*Agropyron* sp., *Agrostis* sp., *Aristida adscensionis* L.*, Bouteloua hirsuta* Lag.*, C. dactylon, Distichlis stricta* (Torr.) Rydb., *Elymus* sp., *E. trachycaulus* (Link) Gould*, Hordeum* sp., *Malva parviflora* L.*, Panicum obtusum* Kunth*, P. scribnerianum* Nash*, Stipa ichu* (Ruiz & Pav.) Kunth*, Tridens pulchellus* (Kunth) Hitchc.*, Triticum aestivum* L.*, Typha latifolia* L.*, Vicia pulchella* Kunth

*Aristida adscensionis* L.*, A. glabrata* (Vasey) Hitchc.*, Bothriochloa saccharoides* (Sw.) Rydb*., Bouteloua* sp., *B. barbata* Lag.*, C. dactylon, Distichlis stricta* (Torr.) Rydb.*, Lycurus phleoides* Kunth*, Tridens pulchellus* (Kunth)

*Dalbergia melanoxylon* Guill. & Perr. Zimbabwe

Australia, China, France, Hawaii, Hong Kong, Japan, Korea, Okinawa Island, Taiwan, the Netherlands, and the USA

Mexico and the USA

New Zealand

Mexico and the USA

Mexico and the USA

**108**


**111**

*Invasive Mite Species in the Americas: Bioecology and Impact*

Flüggé*, Vitis* sp.

**Species Host plants Distribution**

*Vaccinium uliginosum* L. CIS

Bambusaceae Brazil

*Bambusa* sp. Brazil

*Lebeckia linearifolia* E. Mey. Namibia

*Prosopis juliflora* (Sw.) DC. Mexico

*Cliffortia linearifolia* Eckl. & Zeyh.*, C. repens* Schltr. South Africa

*S. senanensis* Japan

Unknown Zaire

*Lomandra multiflora* Britten Australia

*C. dactylon* CIS

*Saccharum officinarum* L. Brazil

*O. sativa* Costa Rica and Panamá

*Grewia* sp. Comoro Island and Zaire

*Rhynchospora* sp. Argentina and Brazil

*C. dactylon, Dactylis glomerata* L.*, Distichlis spicata* (L.) Greene*, D. stricta* (Torr.) Rydb., *Paspalum notatum*

*O. sativa, Panicum maximum* Jacq. Argentina, Brazil,

*O. sativa, P. maximum* Colombia and Venezuela

Colombia, Surinam, and

Brazil, Colombia, Poland,

and the USA

Venezuela

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

*S. oryzae* (Rossi de Simons, 1966)

*S. oudemansi* (Reck, 1948)

*S. papillatus* (Flechtmann, 1995)

*S. paraelymus* (Feres and Flechtmann, 1995)

*S. parasemus* (Pritchard and Baker, 1955)

*pennamontanus* (Meyer, 1987)

*S. prosopis* (Tuttle, Baker and Abbatiello, 1976)

*S. protectus* (Meyer, 1975)

*S. recki* (Ehara, 1957)

*S. reticulatus* (Baker and Pritchard, 19600

*S. rhodanus* (Baker and Pritchard, 1960)

*S. rhynosperus* (Flechtmann and Baker, 1970)

*S. russeus* (Davis, 1969)

*S. saba-sulchani* (Reck, 1956)

*S. saccharum* (Flechtmann and Baker, 1975)

*S. pseudolycurus* Ochoa, (Gray and von Lind., 1990)

*S.* 

*S. paezi* (Alvarado and Freitez, 1976)


*Pests Control and Acarology*

*S. laevidorsatus* (Ehara, 1988)

*S. lanyuensis* (Tseng, 1975)

*S. lechrius* (Rimando, 1962)

*S. lespedeza* (Beglyarov and Mitrofanov, 1973)

*S. levinensis* (Manson, 1967)

*S. longirostrus* (Feres and Flechtmann, 1995)

*S. longus* (Saito, 1990)

*S. luculentus* (Tseng, 1990)

*S. lushanensis* (Dongsheng, 1994)

*S. lycurus* (Tuttle and Baker, 1964)

*S. malayanus* (Ehara, 1988)

*S. malkovskii* (Waistein, 1956)

*S. mansoni* (Gupta, 1980)

*S. minutus* (Wang, 1985)

*S. miscanthi* (Saito, 1990)

*S. miyatahus* (Meyer, 1974)

*S. montanae* (Tuttle and Baker, 1968)

*S. nanjingensis* (Ma and Yuan, 1980)

*S. nesbitti* (Meyer, 1965)

*S. nugax* (Pritchard and Baker, 1955)

**Species Host plants Distribution**

*Cassia siamea* Lam.*, Citrus* sp., *C. esculenta, Glycine max* (L.) Merr., *Pterocarpus indicus* Willd.*, P. vidalianus* Rolfe

Bambusaceae, *Gigantochloa levis* (Blanco) Merr. Malaysia

*Unknown* Taiwan

*Bauhinia* sp., *Desmodium* sp., *Lespedeza* sp., *L. bicolor* CIS, China, Japan, Korea,

Poaceae New Zealand

*Bambusa* sp. Brazil

*Sasa senanensis* (Franch. & Sav.) Rehder Japan

*Diospyros* sp. Taiwan

*Cinnamomum camphora* Meisn. China

*Calamagrostis* sp. CIS

*Oryza* sp. India

Bambusaceae China

*Miscanthus* sp., *M. sinensis* Andersson Japan

*Phyllostachys* sp. China

Poaceae South Africa

*Hilaria mutica* (Buckley) Benth.*,* Poaceae Mexico and the USA

*Muhlenbergia montana* (Nutt.) Hitchc., *Pappophorum* 

*Pterocarpus rotundifolius* (Sond.) Druce South Africa

*Manihot* sp. Indonesia and Malaysia

*Leersia oryzoides* (L.) Sw., *Lycurus phleoides* Kunth*,* 

*Setaria macrostachya* Kunth

*mucronulatum* Nees

Indonesia, Philippines,

and Taiwan

and Taiwan

Mexico and the USA

Mexico and the USA

**110**


**113**

**Figure 1.**

**3.2** *Steneotarsonemus spinki*

The rice mite, *S. spinki*, is the origin of southeastern Asia, where it has been reported causing damage to rice crops varying from 30 to 90% in China and 20–60% in Taiwan [25]. Presently, it is considered as a serious pest of rice in Tropical Asia and Caribbean [26]. Other than rice*, S. spinki* is associated to more than 70 plant species including weeds growing near rice fields, such as wild rice:

*Schizotetranychus hindustanicus colony on citrus leaves (a) and citrus leaves showing characteristic symptoms* 

*for S. hindustanicus feeding (courtesy of Dr.). (b) Mario Cermelli and Pedro Morales.*

*Invasive Mite Species in the Americas: Bioecology and Impact*

**Species Host plants Distribution**

*Acacia nilotica* (L.) Willd. ex Delile*, Beaucarnea stricta*

Lem*., Jasminum grandiflorum* L.

*Alhagi pseudalhagi* (M. Bieb.) Desv. ex B. Keller & Shap. CIS

*Acacia* sp. Zimbabwe

Poaceae Thailand

*Citrus medica* L.*, C. paradisi* Macfad. Taiwan

*Quercus gilliana* Rehder & E.H. Wilson China

*Salix* sp. China

*O. sativa* China and Thailand

India and Mexico

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

*S. ugarovi* (Wainstein, 1960)

*S. umtaliensis* (Meyer, 1974)

*S. undulates* (Beer and Lang, 1958)

*S. vermiculatus* (Ehara and Wongsiri, 1975)

*S. yoshimekii* (Ehara and Wongsiri, 1975)

*S. youngi* (Tseng, 1975)

*S. zhangi* (Wang and Cui, 1992)

*zhongdianensis* (Wang and Cui, 1992)

*Species names that are nonvalid according to the MOBOT.*

*Worldwide Schizotetranychus species (from [14]).*

*S.* 

*1*

**Table 1.**



#### **Table 1.**

*Pests Control and Acarology*

*S. sacrales* (Baker and Pritchard, 1960)

*S. sagatus* (Davis, 1969)

*S. saitoi* (Ehara, 1968)

*S. sayedi* (Attiah, 1967)

*S. schizopus* (Zacher, 1913)

*S. setariae* (Meyer, 1987)

*S. smirnovi* (Waistein, 1954)

*S. spicules* (Baker and Pritchard, 1960)

*S. spiraefolia* (Garman, 1940)

*S. taquarae* (Paschoal, 1971)

*S. tbilisiensis* (Reck, 1959)

*S. tephrosiae* (Gutierrez, 1968)

*S. textor* (Waisntein, 1954)

*S. triquetrus* (Meyer, 1987)

*S. tuberculatus* (Ugarov and Nikolskii, 1937)

*S. tumidus* (Wang, 1981)

*S. tuminicus* (Ma and Yuan, 1982)

*S. tuttlei* (Zacher, Gomaa and El-Enany, 1982)

**Species Host plants Distribution**

Fabaceae Zaire

*Themeda australis* (R. Br.) Stapf Australia

*B. vulgaris* Malaysia

*Ficus carica* L. Egypt

*Citrus* sp., *Murraya koenigii* (L.) Spreng. India and Kenya

CIS, China, Germany, Hungary, Japan, Poland, Switzerland, the Netherlands, the UK, and

China, India, Poland, and

India, Madagascar, and South Africa

the USA

CIS

the USA

CIS

Egypt

South Africa

*Populus* sp., *P. tremula, Salix* sp., *S. alba* L.*, S. aurita* L.*, S. balsamifera* (Hook.) Barratt ex Andersson*, S. bicolor* Willd.*, S. caprea* L.*, S. daphnoides* Vill., *S. elegantissima* K. Koch*, S. fragilis* L.*, S. nigra* Marshall*, S. purpurea* L.*, S. subfragilis* Andersson*, S. viminalis* L.*, Vaccinium* 

*Setaria sphacelata* (Schumach.) Stapf & C.E. Hubb. ex

*C. cajan, S. officinarum, Spiraea* sp., *S. latifolia* (Aiton) Borkh.*, S. pubescens* Turcz*., S. salicifolia* L.*, S. trilobata* L.

*Agrostemma githago* L.*, Bromus* sp., *Elytrigia repens* (L.)

*Balanites pedicellaris* Mildbr. & Schltr*., Eriobotrya japonica* (Thunb.) Lindl., *Mikania cordata* (Burm. f.) B.L. Rob.*, Mundulea pungens* R. Vig., *M. sericea* Hook. & Arn., *Tephrosia striata* Ecklon & Zeyher ex Steudel

*B. vulgaris* Brazil

*Elaeagnus angustifolia* L.*, Lonicera* sp. CIS

*Morus* sp. CIS

*Melia radula* (nonvalid name)<sup>1</sup> China

*Bridelia monoica* (Lour.) Merr. China

*Arundo donax* L.*, Cuscuta planiflora* Ten.*, Mentha* 

*pulegium* L.*, O. sativa*

*Pentzia incana* (Thunb.) Kuntze South Africa

*Juglans regia* L.*, Malus domestica* (Suckow) Borkh.,

*Morus alba* L.*, Prunus armeniaca* L.

Desv. ex Nevski*, Marrubium* sp.

*uliginosum* L.

M.B. Moss

**112**

*Worldwide Schizotetranychus species (from [14]).*

#### **Figure 1.**

*Schizotetranychus hindustanicus colony on citrus leaves (a) and citrus leaves showing characteristic symptoms for S. hindustanicus feeding (courtesy of Dr.). (b) Mario Cermelli and Pedro Morales.*

#### **3.2** *Steneotarsonemus spinki*

The rice mite, *S. spinki*, is the origin of southeastern Asia, where it has been reported causing damage to rice crops varying from 30 to 90% in China and 20–60% in Taiwan [25]. Presently, it is considered as a serious pest of rice in Tropical Asia and Caribbean [26]. Other than rice*, S. spinki* is associated to more than 70 plant species including weeds growing near rice fields, such as wild rice:


#### **Table 2.**

*Surveyed localities in the northern Colombia to detect occurrence of S. hindustanicus (from ICA, 2012).*

*O. latifolia*, *C. dactylon* (Poaceae), *Cyperus articulatus* L., *Cyperus iria* L., and *Oxycaryum* sp. (Cyperaceae) [26, 27].

The rice mite feeds on the adaxial surface of leaf sheaths and developing kernels evidenced by brown lesions and consequently reducing photosynthesis and having a negative effect on fertility [26]. Damage also results in sterile grain syndrome, which is characterized by losing and brown discoloration of the flag leaf sheath, twisted panicle neck, and impaired grain development with empty or incompletely filled grains with brown spots and panicles standing erect. The damage to grains showing sterility and malformed curved appearance is referred to as "parrot-beak" [28]. On the other hand, Shikata et al. [29] found for the first time virus-like particles associated with the tarsonemid mites in the rice plants; the spherical virus-like particles were isolated from the rice plants infected with rice ragged stunt, dwarf, black-streaked dwarf, grassy stunt viruses, as well as from the "healthy" plants, which were not inoculated with those viruses, and in addition, the same particles were also found in the dip preparations of the rice tarsonemid mites and eggs.

*S. spinki* was first reported in North America in 1960 on *Tagosodes orizicolus* (Muir, 1926) in Louisiana, USA. Several years after, the rice mite was found causing damage in rice crops (*O. sativa*) in Cuba in 1997 [30]. Subsequently, this tarsonemid mite spreads over all the Caribbean and Central America: Dominican Republic [31], Costa Rica [32], Haiti [33], Panama [34], Guatemala, Honduras [35], and Mexico [36]. In South America, it has been reported in Colombia [37] and Venezuela [38].

After being introduced in Cuba, outbreaks were registered from 1997 to 1998 when an increase in vain grains of 15–20% and a loss of 2 t/ha were recorded [25]. At the end of 1998, *S. spinki* was also found in Dominican Republic and Haiti, causing about 30% of yield loss; however, less intense damage was verified as compared to Cuba [34].

#### **3.3** *Raoiella indica*

The red palm mite, *R. indica*, is of Asian origin, and it is widely distributed in India, Pakistan, Russia, Iran, Israel, Oman, Pakistan, Egypt, Sudan, and Mauritius [39]. Since 2004, *R. indica* was reported from several Caribbean islands, including

**115**

**Figure 2.**

*coconut leaves (C).*

*Invasive Mite Species in the Americas: Bioecology and Impact*

Martinique [40], Saint Lucia and Dominica [41], Guadeloupe and Saint Martin [42], Puerto Rico and Culebra Island [43], and Jamaica [44] (Welbourn, 2007). More recently, it has also been found in Venezuela [45], Colombia [46], and Brazil [47]. *Raoiella indica* can cause severe damage not only to Arecaceae, especially coconut (*C. nucifera*), but also to Musaceae and other plant families [40, 42, 48] (**Figure 2**). Infested plants exhibit a characteristic "yellowing" as a result of mites feeding on the nutrient-rich layers of the leaves' mesophyll tissues [40]. *Raoiella* species inflict damage by introducing the infrastratum through the stomatal opening to feed on the underlying mesophyll cells [49]. Therefore, the distribution of the stomata on the leaf surface could have a greater influence on the feeding capacity of *R. indica* on the host plant [50], and the severity of the feeding damage by the red palm mite

After RPM was reported in the New World, little was known about bioecology of this Red palm mite. Regarding host plant, only coconut and *Adonidia merrillii* (Becc.) Becc. had been recorded as host plants to this mite [41, 48]. After RPM occurrence in South America in the coastal Sucre state of Venezuela, Vásquez et al. [52] registered higher population levels of RPM on coconut, banana (*Musa* spp.), ornamental plants, and weeds in the northern Venezuela (**Figure 3** and **Table 3**). These authors observed all RPM stages only on eight arecaceous, one musaceous,

*Coconut and plantain trees showing symptoms of R. indica feeding (A, B) and a colony of the red palm mite on* 

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

increases in the young plants [43, 51].

#### *Invasive Mite Species in the Americas: Bioecology and Impact DOI: http://dx.doi.org/10.5772/intechopen.86127*

*Pests Control and Acarology*

*O. latifolia*, *C. dactylon* (Poaceae), *Cyperus articulatus* L., *Cyperus iria* L., and

*Surveyed localities in the northern Colombia to detect occurrence of S. hindustanicus (from ICA, 2012).*

**Department Municipality Presence/absence Host plant**

Polonuevo − Baraona −

Santa Ana − Ciénaga − Zona Bananera −

San Sebastián − San Zenón −

Atlántico Luruaco −

La Guajira Dibulla + Magdalena Santa Marta +

El Banco

Vichada Puerto Carreño −

The rice mite feeds on the adaxial surface of leaf sheaths and developing kernels evidenced by brown lesions and consequently reducing photosynthesis and having a negative effect on fertility [26]. Damage also results in sterile grain syndrome, which is characterized by losing and brown discoloration of the flag leaf sheath, twisted panicle neck, and impaired grain development with empty or incompletely filled grains with brown spots and panicles standing erect. The damage to grains showing sterility and malformed curved appearance is referred to as "parrot-beak" [28]. On the other hand, Shikata et al. [29] found for the first time virus-like particles associated with the tarsonemid mites in the rice plants; the spherical virus-like particles were isolated from the rice plants infected with rice ragged stunt, dwarf, black-streaked dwarf, grassy stunt viruses, as well as from the "healthy" plants, which were not inoculated with those viruses, and in addition, the same particles were also found in the dip preparations of the rice tarsonemid mites and eggs. *S. spinki* was first reported in North America in 1960 on *Tagosodes orizicolus* (Muir, 1926) in Louisiana, USA. Several years after, the rice mite was found causing damage in rice crops (*O. sativa*) in Cuba in 1997 [30]. Subsequently, this tarsonemid mite spreads over all the Caribbean and Central America: Dominican Republic [31], Costa Rica [32], Haiti [33], Panama [34], Guatemala, Honduras [35], and Mexico [36]. In South America, it has been reported in Colombia [37] and Venezuela [38]. After being introduced in Cuba, outbreaks were registered from 1997 to 1998 when an increase in vain grains of 15–20% and a loss of 2 t/ha were recorded [25]. At the end of 1998, *S. spinki* was also found in Dominican Republic and Haiti, causing about 30% of yield loss; however, less intense damage was verified as compared to Cuba [34].

Guamal + *C. sinensis*

*C. limon*

The red palm mite, *R. indica*, is of Asian origin, and it is widely distributed in India, Pakistan, Russia, Iran, Israel, Oman, Pakistan, Egypt, Sudan, and Mauritius [39]. Since 2004, *R. indica* was reported from several Caribbean islands, including

*Oxycaryum* sp. (Cyperaceae) [26, 27].

**Table 2.**

**114**

**3.3** *Raoiella indica*

Martinique [40], Saint Lucia and Dominica [41], Guadeloupe and Saint Martin [42], Puerto Rico and Culebra Island [43], and Jamaica [44] (Welbourn, 2007). More recently, it has also been found in Venezuela [45], Colombia [46], and Brazil [47].

*Raoiella indica* can cause severe damage not only to Arecaceae, especially coconut (*C. nucifera*), but also to Musaceae and other plant families [40, 42, 48] (**Figure 2**). Infested plants exhibit a characteristic "yellowing" as a result of mites feeding on the nutrient-rich layers of the leaves' mesophyll tissues [40]. *Raoiella* species inflict damage by introducing the infrastratum through the stomatal opening to feed on the underlying mesophyll cells [49]. Therefore, the distribution of the stomata on the leaf surface could have a greater influence on the feeding capacity of *R. indica* on the host plant [50], and the severity of the feeding damage by the red palm mite increases in the young plants [43, 51].

After RPM was reported in the New World, little was known about bioecology of this Red palm mite. Regarding host plant, only coconut and *Adonidia merrillii* (Becc.) Becc. had been recorded as host plants to this mite [41, 48]. After RPM occurrence in South America in the coastal Sucre state of Venezuela, Vásquez et al. [52] registered higher population levels of RPM on coconut, banana (*Musa* spp.), ornamental plants, and weeds in the northern Venezuela (**Figure 3** and **Table 3**). These authors observed all RPM stages only on eight arecaceous, one musaceous,

#### **Figure 2.**

*Coconut and plantain trees showing symptoms of R. indica feeding (A, B) and a colony of the red palm mite on coconut leaves (C).*

*Distribution of R. indica in Venezuela based on collection records (from 2008 to 2012) [52].*


#### **Table 3.**

*Plant species onto which Raoiella indica was found in the northern Venezuela (from [52]).*

and one streliziaceous species, indicating that the pest developed and reproduced only on these plants, while specimens found on weeds were considered spurious events. Later, the list of host plants increased including 73 species of Arecaceae, six of Musaceae, five of Heliconiaceae, four of Zingiberaceae, and two each of Pandanaceae and Strelitziaceae [53].

**117**

*Invasive Mite Species in the Americas: Bioecology and Impact*

Due to the potential impact of *R. indica*, in 2007, the Brazilian Ministry of Agriculture added the RPM to the list of quarantine pests so that an extensive survey was initiated in the state of Roraima, a Brazilian state bordering Venezuela [54]. Only after 2 years, in July 2009, the red palm mite was found in samples of coconut and banana leaves in urban areas of Boa Vista (Roraima) [47]. Despite quarantine efforts, this mite became established in South America inasmuch as in 2011; it was reported in the urban areas of Manaus occurring not only on coconut plants but also on dwarf royal palm (*Veitchia merrillii* (Becc.) H. E. Moore) and fishtail palm tree (*Caryota mitis* Lour) [55]. Recently, in May 2015, the RPM was found in the urban area of Dracena (state of São Paulo), about 2300 km southeast of Manaus, on several arecaceous plants such as *C. nucifera*, *Phoenix roebelenii* O'Brien, and *Rhapis* 

Discovery of *R. indica* in several countries in South America suggests that this region exhibits climate conditions, which, along with the wide diversity of host plant species, stimulate its development, representing an imminent threat to the economy of those countries where coconut palm and banana are grown as crops of

Biological invasions have increased greatly in the last century due to the intensification in international trade, thus representing one of the most relevant threats for biodiversity in agroecosystems. Over several decades, scientists are more interested in phytophagous mites since some noneconomic species have become severe pests on many crops due to wrong pest management strategies. Thus,

phytophagous mites from the Neotropical region such as the cassava green mite, the coconut mite, and the tomato spider mite have been introduced in the Old World. As expected, some mite species have also been introduced in the Neotropical region, i.e., *S. spinki*, *S. hindustanicus*, and *R. indica* with remarkable economic impact on agriculture. These biological invasions in the New World require the participation of several public institutions (Universities, Government Agricultural Institutions) and farmers in order to mitigate the current impact on production of rice, citrus, coconut, and Musaceous crops. Most of the research has been focused on geographical distribution, host plant range, and natural enemies associated, but few studies

The authors would like to acknowledge the Dirección de Investigación y Desarrollo (DIDE) of Universidad Técnica de Ambato and the Centre for Agricultural Bioscience

International (CABI) for the financial support to this research.

The authors have declared no conflict of interests.

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

*excelsa* (Thunb.) A. Henry [54].

have dealt with management strategies.

**Acknowledgements**

**Conflict of interest**

economic importance.

**4. Conclusions**

Due to the potential impact of *R. indica*, in 2007, the Brazilian Ministry of Agriculture added the RPM to the list of quarantine pests so that an extensive survey was initiated in the state of Roraima, a Brazilian state bordering Venezuela [54].

Only after 2 years, in July 2009, the red palm mite was found in samples of coconut and banana leaves in urban areas of Boa Vista (Roraima) [47]. Despite quarantine efforts, this mite became established in South America inasmuch as in 2011; it was reported in the urban areas of Manaus occurring not only on coconut plants but also on dwarf royal palm (*Veitchia merrillii* (Becc.) H. E. Moore) and fishtail palm tree (*Caryota mitis* Lour) [55]. Recently, in May 2015, the RPM was found in the urban area of Dracena (state of São Paulo), about 2300 km southeast of Manaus, on several arecaceous plants such as *C. nucifera*, *Phoenix roebelenii* O'Brien, and *Rhapis excelsa* (Thunb.) A. Henry [54].

Discovery of *R. indica* in several countries in South America suggests that this region exhibits climate conditions, which, along with the wide diversity of host plant species, stimulate its development, representing an imminent threat to the economy of those countries where coconut palm and banana are grown as crops of economic importance.

## **4. Conclusions**

*Pests Control and Acarology*

**116**

**Table 3.**

**Figure 3.**

Pandanaceae and Strelitziaceae [53].

**Plant family Species** Apocynaceae *Rauvolfia viridis* Arecaceae *Adonidia merrillii*

Musaceae *Musa* sp*.* Sterculiaceae *Sterculia* sp. Strelitziaceae *Strelitzia* sp*.*

and one streliziaceous species, indicating that the pest developed and reproduced only on these plants, while specimens found on weeds were considered spurious events. Later, the list of host plants increased including 73 species of Arecaceae, six of Musaceae, five of Heliconiaceae, four of Zingiberaceae, and two each of

*Plant species onto which Raoiella indica was found in the northern Venezuela (from [52]).*

*Cocos nucifera Roystonea oleracea Pritchardia pacifica Ptychosperma macarthurii*

*Distribution of R. indica in Venezuela based on collection records (from 2008 to 2012) [52].*

*Roystonea regia Washingtonia* sp*. Washingtonia robusta Phaseolus* sp*.*

Biological invasions have increased greatly in the last century due to the intensification in international trade, thus representing one of the most relevant threats for biodiversity in agroecosystems. Over several decades, scientists are more interested in phytophagous mites since some noneconomic species have become severe pests on many crops due to wrong pest management strategies. Thus, phytophagous mites from the Neotropical region such as the cassava green mite, the coconut mite, and the tomato spider mite have been introduced in the Old World. As expected, some mite species have also been introduced in the Neotropical region, i.e., *S. spinki*, *S. hindustanicus*, and *R. indica* with remarkable economic impact on agriculture. These biological invasions in the New World require the participation of several public institutions (Universities, Government Agricultural Institutions) and farmers in order to mitigate the current impact on production of rice, citrus, coconut, and Musaceous crops. Most of the research has been focused on geographical distribution, host plant range, and natural enemies associated, but few studies have dealt with management strategies.

## **Acknowledgements**

The authors would like to acknowledge the Dirección de Investigación y Desarrollo (DIDE) of Universidad Técnica de Ambato and the Centre for Agricultural Bioscience International (CABI) for the financial support to this research.

### **Conflict of interest**

The authors have declared no conflict of interests.

*Pests Control and Acarology*

## **Author details**

Carlos Vásquez1 \* and Yelitza Colmenárez<sup>2</sup>

1 Faculty of Agronomy Sciences, Technical University of Ambato, Ambato, Ecuador

2 CABI Brazil, UNESP- Fazenda Experimental Lageado, Fundação de Estudos e Pesquisas Agrícolas e Florestais, Botucatu-São Paulo, Brazil

\*Address all correspondence to: ca.vasquez@uta.edu.ec

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

**119**

*Invasive Mite Species in the Americas: Bioecology and Impact*

alien species: Quarantine procedures. In: Proceedings of the XI International Congress of Acarology. Merida: Universidad Nacional Autonoma de

[9] Jeschke JM, Bacher S, Blackburn TM, Dick JT, Essl F, Evans T, et al. Defining the impact of non-native species. Conservation Biology. 2015;**28**(5):1188-1194. DOI: 10.1111/

[10] Courtois P, Figuieres C, Mulier C, Weill J. A cost-benefit approach for prioritizing invasive species. Ecological Economics. 2018;**146**:607-620. DOI: 10.1016/j.ecolecon.2017.11.037

[11] Martins CBC, Almeida LM, Zontade-Carvalho RC, Castro CF, Pereira RA. *Harmonia axyridis*: A threat to Brazilian Coccinellidae? Revista Brasileira de Entomologia. 2009;**53**(4):663-671. DOI: 10.1590/S0085-56262009000400018

[12] Richardson DM, Pysek P, Rejmánek M, Barbour MG, Panetta FD, West CJ. Naturalization and invasion of alien plants: Concepts and definitions. Diversity

[13] Beck KG, Zimmerman K, Schardt JD, Stone J, Lukens RR, Reichard S, et al. Invasive species defined in a policy context: Recommendations from the federal invasive species advisory committee. Invasive Plant Science and Management. 2008;**1**:414-421. DOI:

[14] Bolland H, Gutiérrez J, Fletchmann C. World Catalogue of the Spider Mites Family (Acari: Tetranychidae). Leiden: Koninklijke Brill NV; 1998. p. 392

[15] Nienstaedt B, Marcano R. Estudio de la biología del ácaro hindú de los cítricos *Schizotetranychus hindustanicus* (Hirst, 1924) (Acari: Tetranychidae), en tres tipos de alimentos. Entomotropica.

and Distributions. 2000;**6**:93-107

10.1614/IPSM-08-089.1

2009;**24**(2):51-56

Mexico; 2007. pp. 307-316

cobi.12299

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

[1] Blackburn TM, Essl F, Evans T, Hulme PE, Jeschke JM, Kühn I, et al. A unified classification of alien species based on the magnitude of their environmental impacts. PLoS Biology. 2014;**12**(5). DOI: 10.1371/journal.

[2] Pysek P, Jarosik V, Hulme P, Pergl J, Hejda M, Schaffner U, et al. A global assessment of invasive plant impacts on resident species, communities and ecosystems: The interaction of impact measures, invading species' traits and environment. Global Change Biology. 2012;**18**:1725-1737. DOI: 10.1111/j.1365-2486.2011.02636.x

[3] Ricciardi A, Hoopes MF, Marchetti MP, Lockwood JL. Progress toward understanding the ecological impacts of nonnative species. Ecological Monographs. 2013;**83**(3):263-282

[4] Colautti RI, MacIsaac HJ. A neutral terminology to define 'invasive' species. Diversity and Distributions.

[5] Paini DR, Sheppard AW, Cook DC, De Barro PJ, Worner SP, Thomas MB. Global threat to agriculture from invasive species. Proceedings of the National Academy of Sciences of the United States of America. 2016;**113**(27):7575-7579. DOI: 10.1073/

[6] Preston CD, Pearman DA, Hall AR. Archaeophytes in Britain. Botanical

[7] Ferragut F, Navia D, Ochoa R. New mite invasions in citrus in the early years of the 21st century. Experimental & Applied Acarology. 2012;**59**(1-2):145-164.

[8] Navia D, de Moraes GJ, Flechtmann CHW. Phytophagous mites as invasive

Journal of the Linnean Society.

DOI: 10.1007/s10493-012-9635-9

2004;**10**:135-141

pnas.1602205113

2004;**145**:257-294

pbio.1001850

**References**

*Invasive Mite Species in the Americas: Bioecology and Impact DOI: http://dx.doi.org/10.5772/intechopen.86127*

## **References**

*Pests Control and Acarology*

**118**

**Author details**

Carlos Vásquez1

provided the original work is properly cited.

\* and Yelitza Colmenárez<sup>2</sup>

Pesquisas Agrícolas e Florestais, Botucatu-São Paulo, Brazil

\*Address all correspondence to: ca.vasquez@uta.edu.ec

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

1 Faculty of Agronomy Sciences, Technical University of Ambato, Ambato, Ecuador

2 CABI Brazil, UNESP- Fazenda Experimental Lageado, Fundação de Estudos e

[1] Blackburn TM, Essl F, Evans T, Hulme PE, Jeschke JM, Kühn I, et al. A unified classification of alien species based on the magnitude of their environmental impacts. PLoS Biology. 2014;**12**(5). DOI: 10.1371/journal. pbio.1001850

[2] Pysek P, Jarosik V, Hulme P, Pergl J, Hejda M, Schaffner U, et al. A global assessment of invasive plant impacts on resident species, communities and ecosystems: The interaction of impact measures, invading species' traits and environment. Global Change Biology. 2012;**18**:1725-1737. DOI: 10.1111/j.1365-2486.2011.02636.x

[3] Ricciardi A, Hoopes MF, Marchetti MP, Lockwood JL. Progress toward understanding the ecological impacts of nonnative species. Ecological Monographs. 2013;**83**(3):263-282

[4] Colautti RI, MacIsaac HJ. A neutral terminology to define 'invasive' species. Diversity and Distributions. 2004;**10**:135-141

[5] Paini DR, Sheppard AW, Cook DC, De Barro PJ, Worner SP, Thomas MB. Global threat to agriculture from invasive species. Proceedings of the National Academy of Sciences of the United States of America. 2016;**113**(27):7575-7579. DOI: 10.1073/ pnas.1602205113

[6] Preston CD, Pearman DA, Hall AR. Archaeophytes in Britain. Botanical Journal of the Linnean Society. 2004;**145**:257-294

[7] Ferragut F, Navia D, Ochoa R. New mite invasions in citrus in the early years of the 21st century. Experimental & Applied Acarology. 2012;**59**(1-2):145-164. DOI: 10.1007/s10493-012-9635-9

[8] Navia D, de Moraes GJ, Flechtmann CHW. Phytophagous mites as invasive

alien species: Quarantine procedures. In: Proceedings of the XI International Congress of Acarology. Merida: Universidad Nacional Autonoma de Mexico; 2007. pp. 307-316

[9] Jeschke JM, Bacher S, Blackburn TM, Dick JT, Essl F, Evans T, et al. Defining the impact of non-native species. Conservation Biology. 2015;**28**(5):1188-1194. DOI: 10.1111/ cobi.12299

[10] Courtois P, Figuieres C, Mulier C, Weill J. A cost-benefit approach for prioritizing invasive species. Ecological Economics. 2018;**146**:607-620. DOI: 10.1016/j.ecolecon.2017.11.037

[11] Martins CBC, Almeida LM, Zontade-Carvalho RC, Castro CF, Pereira RA. *Harmonia axyridis*: A threat to Brazilian Coccinellidae? Revista Brasileira de Entomologia. 2009;**53**(4):663-671. DOI: 10.1590/S0085-56262009000400018

[12] Richardson DM, Pysek P, Rejmánek M, Barbour MG, Panetta FD, West CJ. Naturalization and invasion of alien plants: Concepts and definitions. Diversity and Distributions. 2000;**6**:93-107

[13] Beck KG, Zimmerman K, Schardt JD, Stone J, Lukens RR, Reichard S, et al. Invasive species defined in a policy context: Recommendations from the federal invasive species advisory committee. Invasive Plant Science and Management. 2008;**1**:414-421. DOI: 10.1614/IPSM-08-089.1

[14] Bolland H, Gutiérrez J, Fletchmann C. World Catalogue of the Spider Mites Family (Acari: Tetranychidae). Leiden: Koninklijke Brill NV; 1998. p. 392

[15] Nienstaedt B, Marcano R. Estudio de la biología del ácaro hindú de los cítricos *Schizotetranychus hindustanicus* (Hirst, 1924) (Acari: Tetranychidae), en tres tipos de alimentos. Entomotropica. 2009;**24**(2):51-56

[16] Quirós M, Geraud-Pouey F. *Schizotetranychus hindustanicus* (Hirst) (Acari: Tetranychidae), new spider mite pest damaging citrus in Venezuela, South America. In: Proceedings of the XI International Congress of Acarology; 8-13 September 2002. Mérida: Universidad Nacional Autónoma de México; 2008. pp. 255-256

[17] Návia D, Marsaro AL Jr. First report of the citrus Hindu Mite, *Schizotetranychus hindustanicus* (Hirst) (Prostigmata: Tetranychidae), in Brazil. Neotropical Entomology. 2010;**39**(1):140-143

[18] Quirós M, Geraud-Pouey F. *Schizotetranychus hindustanicus* (Hirst) (Acari: Tetranychidae), new spider mite pest damaging citrus in Venezuela, South America. In: International Congress of Acarology. Mérida, México; 8-13 September 2002; Program and Abstract Book; 2002. pp. 255-256

[19] Nienstaedt BM. Estudio de algunos aspectos biológicos y ecológicos del ácaro hindú de los cítricos *Schizotetranychus hindustanicus* (Hirst, 1924) (Acari: Tetranychidae) en Maracay, Venezuela [thesis]. Maracay: Universidad Central de Venezuela; 2007

[20] Quirós M, Dorado I. Eficiencia de tres productos comerciales en el control del ácaro hindú de las cítricas *Schizotetranychus hindustanicus* (Hirst), en el laboratorio. In: Congreso Venezolano de Entomología; 4-7 July 2005; Maracaibo. Maracay: Entomotropica; 2005. pp. 184-185

[21] Mesa NC. Ácaros Asociados a Cítricos en Colombia [Internet]. 2010. Available from: http:// www.comfenalcoantioquia.com/ boletinempresas/MEMORIAS\_ CITRICOS/CONFERENCISTAS/ NACIONALES/Nohora%20Cristina%20 Mesa%20-%20Colombia/Acaros%20 en%20citricos%20-%20Congreso.pdf

[22] Nienstaedt B, Marcano R. Fluctuación poblacional y distribución vertical del ácaro *Schizotetranychus hindustanicus* (Hirst, 1924), sobre especies de *Citrus*. Entomotropica. 2009b;**24**(2):57-63

[23] Alvarado G, Fréitez F. *Schizotetranychus paezi* sp. n. y *Schizotetranychus oryzae* (Acarina: Tetranychidae) atacando arroz en Venezuela. Agronomía Tropical. 1976;**26**(2):159-165

[24] Zhang Z, Zhang Y, Lin J. Mites of *Schizotetranychus* (Acari: Tetranychidae) from moso bamboo in Funjian, China. Systematic and Applied Acarology Special Publications. 2000;**4**:19-35. DOI: 10.11158/sasp.4.1.5

[25] Almaguel L, Cabrera I, Hernández J, Ramos M, Sandoval I. Etiología, Biología, Ecología y Manejo Integrado del Vaneado de la Panícula y Pudrición de la Vaina del Arroz en Cuba. Instituto de Sanidad Vegetal, Instituto de Investigaciones del Arroz, Centro Nacional de Sanidad Agropecuaria: La Habana; 2002

[26] Chandrasena GDSN, Jayawardane JDKM, Umange SD, Gunawardana ADBU. Host range of panicle rice mite *Steneotarsonemus spinki* Smiley (Acari: Tarsonemidae) in Sri Lanka. Universal Journal of Agricultural Research. 2016;**4**(1):21-24. DOI: 10.13189/ ujar.2016.040104

[27] Rao J, Prakash A. Paddy field weed, *Schoenoplectus articulatus* (Linn.) Palla (Cyperaceae): A new host of tarsonemid mite, *Steneotarsonemus spinki* Smiley and panicle thrips, *Haplothrips ganglbaureri* Schmutz. Journal of Applied Zoological Researches. 2002;**13**:174-175

[28] Hummel NA, Castro BA, McDonald EM, Pellerano MA, Ochoa R. The panicle rice mite, *Steneotarsonemus spinki* Smiley, a re-discovered pest of rice in the United States. Crop

**121**

*Invasive Mite Species in the Americas: Bioecology and Impact*

February-1 March 2006; Texas: The

[36] Arriaga JT. Detection of the Rice Tarsonemid Mite (*Steneotarsonemus spinki* Smiley) in Palizada, Campeche, Mexico [Internet]. 2007. Available from: http://www.pestalert.org/oprDetail.

[37] ICA (Instituto Colombiano

[38] Aguilar H, Murillo P. Nuevos hospederos y registros de ácaros fitófagos para Costa Rica: Periodo 2002-2008. Agronomica Costarricense.

10.13140/RG.2.1.1194.0886

DOI: 10.11158/saa.9.1.16

et%20al.pdf

[39] Mendonça RS, Návia D, Flechtmann CHW. *Raoiella indica* Hirst (Acari: Prostigamata: Tenuipalpidae), o ácaro vermelho das palmeiras: Uma ameaça para as Américas. Documento 146. Brasilia: EMBRAPA; 2005. 37 p. DOI:

[40] Flechtmann CHW, Etienne J. The red palm mite, *Raoiella indica* Hirst, a threat to palms in the Americas (Acari: Prostigmata: Tenuipalpidae). Systematic and Applied Acarology. 2004;**9**:109-110.

[41] Kane E, Ochoa R, Mathurin G, Erbe E. *Raoiella indica* Hirst (Acari: Tenuipalpidae): An island-hopping mite pest in the Caribbean [Internet]. 2005. Available from: www.sel.barc.usda.gov/ acari/PDF/Raoiella%20indica-Kane%20

[42] Etienne J, Flechtmann CHW. First record of *Raoiella indica* (Hirst, 1924) (Acari: Tenuipalpidae) in Guadeloupe and Saint Martin, West Indies. International Journal of

Agropecuario). Resolución No. 001195 de 2005. Diario Oficial, Edición 45.892. 27 abril de 2005 [Internet]. 2005. Available from: https://normograma. info/invima/docs/pdf/resolucion\_

Woodlands. pp. 97-98

cfm%3FoprID¼268

ica\_1195\_2005.pdf

2008;**32**(2):7-28

*DOI: http://dx.doi.org/10.5772/intechopen.86127*

Protection. 2009:1-14. DOI: 10.1016/j.

[29] Shikata E, Kawano S, Senboku T, Tiongco ER, Miyajima K. Small viruslike particles isolated from the leaf sheath tissues of rice plants and from the rice tarsonemid mites, *Steneotarsonemus spinki* Smiley (Acarina, Tarsonemidae). Annals of the Phytopathological Society

cropro.2009.03.011

of Japan. 1984;**50**:368-374

1997;**13**(1):25-28

Vegetal. 2001;**16**:6-9

Costa Rica; 2005. 16 p

[33] Herrera LAR. Ácaro del Vaneamiento del Arroz-*Steneotarsonemus spinki* Smiley

foro-agosto-pdf05/acaro.pdf

[34] Almaguel L, Botta E. Manejo Integrado de *Steneotarsonemus spinki* Smiley: Resultados de Cuba y Transferencia Para la Región de Latinoamérica y el Caribe [Internet]. 2005. Available from: http://www. inisav.cu/OtrasPub/Curso%20 acarolog%C3%ADa.pdf

[35] Castro BA, Ochoa R, Cuevas FE. The threat of the panicle rice mite, *Steneotarsonemus spinki* Smiley, to rice production in the United States. In: Proceedings of the Thirty First Rice Technical Working Group; 26

[30] Ramos M, Rodríguez H.

*Steneotarsonemus spinki* Smiley (Acari: Tarsonemidae): Nuevo informe para Cuba. Revista de Protección Vegetal.

[31] Ramos M, Gómez C, Cabrera RI. Presencia de *Steneotarsonemus spinki* (Acari: Tarsonemidae) en cuatro variedades de arroz en la República Dominicana. Revista de Protección

[32] Sanabria C, Aguilar H. El Ácaro del Vaneo del Arroz (*Steneotarsonemus spinki* L: Tarsonemidae). Ministerio de Agricultura y Ganaderia: San José de

(Prostigmata: Tarsonemidae) [Internet]. 2005. Available from: www.flar.org/pdf/

*Invasive Mite Species in the Americas: Bioecology and Impact DOI: http://dx.doi.org/10.5772/intechopen.86127*

Protection. 2009:1-14. DOI: 10.1016/j. cropro.2009.03.011

*Pests Control and Acarology*

[16] Quirós M, Geraud-Pouey F. *Schizotetranychus hindustanicus* (Hirst) (Acari: Tetranychidae), new spider mite pest damaging citrus in Venezuela, South America. In: Proceedings of the XI International Congress of Acarology; [22] Nienstaedt B, Marcano R. Fluctuación poblacional y distribución vertical del ácaro

[23] Alvarado G, Fréitez F. *Schizotetranychus paezi* sp. n. y *Schizotetranychus oryzae* (Acarina: Tetranychidae) atacando arroz en Venezuela. Agronomía Tropical.

1976;**26**(2):159-165

10.11158/sasp.4.1.5

Habana; 2002

ujar.2016.040104

*Schizotetranychus hindustanicus* (Hirst, 1924), sobre especies de *Citrus*. Entomotropica. 2009b;**24**(2):57-63

[24] Zhang Z, Zhang Y, Lin J. Mites of *Schizotetranychus* (Acari: Tetranychidae) from moso bamboo in Funjian, China. Systematic and Applied Acarology Special Publications. 2000;**4**:19-35. DOI:

[25] Almaguel L, Cabrera I, Hernández J, Ramos M, Sandoval I. Etiología, Biología, Ecología y Manejo Integrado del Vaneado de la Panícula y Pudrición de la Vaina del Arroz en Cuba. Instituto

[26] Chandrasena GDSN, Jayawardane JDKM, Umange SD, Gunawardana ADBU. Host range of panicle rice mite *Steneotarsonemus spinki* Smiley (Acari: Tarsonemidae) in Sri Lanka. Universal Journal of Agricultural Research. 2016;**4**(1):21-24. DOI: 10.13189/

[27] Rao J, Prakash A. Paddy field weed, *Schoenoplectus articulatus* (Linn.) Palla (Cyperaceae): A new host of tarsonemid mite, *Steneotarsonemus spinki* Smiley and panicle thrips, *Haplothrips ganglbaureri* Schmutz. Journal of Applied Zoological

[28] Hummel NA, Castro BA, McDonald EM, Pellerano MA, Ochoa R. The panicle rice mite, *Steneotarsonemus spinki* Smiley, a re-discovered pest of rice in the United States. Crop

Researches. 2002;**13**:174-175

de Sanidad Vegetal, Instituto de Investigaciones del Arroz, Centro Nacional de Sanidad Agropecuaria: La

8-13 September 2002. Mérida: Universidad Nacional Autónoma de

[17] Návia D, Marsaro AL Jr. First report of the citrus Hindu Mite, *Schizotetranychus hindustanicus* (Hirst) (Prostigmata: Tetranychidae), in Brazil. Neotropical Entomology.

[18] Quirós M, Geraud-Pouey F. *Schizotetranychus hindustanicus* (Hirst) (Acari: Tetranychidae), new spider mite pest damaging citrus in Venezuela, South America. In: International

[19] Nienstaedt BM. Estudio de algunos aspectos biológicos y ecológicos del ácaro hindú de los cítricos *Schizotetranychus hindustanicus* (Hirst, 1924) (Acari: Tetranychidae) en Maracay, Venezuela [thesis]. Maracay: Universidad Central de

[20] Quirós M, Dorado I. Eficiencia de tres productos comerciales en el control del ácaro hindú de las cítricas *Schizotetranychus hindustanicus*

(Hirst), en el laboratorio. In: Congreso Venezolano de Entomología; 4-7 July 2005; Maracaibo. Maracay: Entomotropica; 2005. pp. 184-185

[21] Mesa NC. Ácaros Asociados a Cítricos en Colombia [Internet]. 2010. Available from: http:// www.comfenalcoantioquia.com/ boletinempresas/MEMORIAS\_ CITRICOS/CONFERENCISTAS/

NACIONALES/Nohora%20Cristina%20 Mesa%20-%20Colombia/Acaros%20 en%20citricos%20-%20Congreso.pdf

Congress of Acarology. Mérida, México; 8-13 September 2002; Program and Abstract Book; 2002. pp. 255-256

México; 2008. pp. 255-256

2010;**39**(1):140-143

Venezuela; 2007

**120**

[29] Shikata E, Kawano S, Senboku T, Tiongco ER, Miyajima K. Small viruslike particles isolated from the leaf sheath tissues of rice plants and from the rice tarsonemid mites, *Steneotarsonemus spinki* Smiley (Acarina, Tarsonemidae). Annals of the Phytopathological Society of Japan. 1984;**50**:368-374

[30] Ramos M, Rodríguez H. *Steneotarsonemus spinki* Smiley (Acari: Tarsonemidae): Nuevo informe para Cuba. Revista de Protección Vegetal. 1997;**13**(1):25-28

[31] Ramos M, Gómez C, Cabrera RI. Presencia de *Steneotarsonemus spinki* (Acari: Tarsonemidae) en cuatro variedades de arroz en la República Dominicana. Revista de Protección Vegetal. 2001;**16**:6-9

[32] Sanabria C, Aguilar H. El Ácaro del Vaneo del Arroz (*Steneotarsonemus spinki* L: Tarsonemidae). Ministerio de Agricultura y Ganaderia: San José de Costa Rica; 2005. 16 p

[33] Herrera LAR. Ácaro del Vaneamiento del Arroz-*Steneotarsonemus spinki* Smiley (Prostigmata: Tarsonemidae) [Internet]. 2005. Available from: www.flar.org/pdf/ foro-agosto-pdf05/acaro.pdf

[34] Almaguel L, Botta E. Manejo Integrado de *Steneotarsonemus spinki* Smiley: Resultados de Cuba y Transferencia Para la Región de Latinoamérica y el Caribe [Internet]. 2005. Available from: http://www. inisav.cu/OtrasPub/Curso%20 acarolog%C3%ADa.pdf

[35] Castro BA, Ochoa R, Cuevas FE. The threat of the panicle rice mite, *Steneotarsonemus spinki* Smiley, to rice production in the United States. In: Proceedings of the Thirty First Rice Technical Working Group; 26

February-1 March 2006; Texas: The Woodlands. pp. 97-98

[36] Arriaga JT. Detection of the Rice Tarsonemid Mite (*Steneotarsonemus spinki* Smiley) in Palizada, Campeche, Mexico [Internet]. 2007. Available from: http://www.pestalert.org/oprDetail. cfm%3FoprID¼268

[37] ICA (Instituto Colombiano Agropecuario). Resolución No. 001195 de 2005. Diario Oficial, Edición 45.892. 27 abril de 2005 [Internet]. 2005. Available from: https://normograma. info/invima/docs/pdf/resolucion\_ ica\_1195\_2005.pdf

[38] Aguilar H, Murillo P. Nuevos hospederos y registros de ácaros fitófagos para Costa Rica: Periodo 2002-2008. Agronomica Costarricense. 2008;**32**(2):7-28

[39] Mendonça RS, Návia D, Flechtmann CHW. *Raoiella indica* Hirst (Acari: Prostigamata: Tenuipalpidae), o ácaro vermelho das palmeiras: Uma ameaça para as Américas. Documento 146. Brasilia: EMBRAPA; 2005. 37 p. DOI: 10.13140/RG.2.1.1194.0886

[40] Flechtmann CHW, Etienne J. The red palm mite, *Raoiella indica* Hirst, a threat to palms in the Americas (Acari: Prostigmata: Tenuipalpidae). Systematic and Applied Acarology. 2004;**9**:109-110. DOI: 10.11158/saa.9.1.16

[41] Kane E, Ochoa R, Mathurin G, Erbe E. *Raoiella indica* Hirst (Acari: Tenuipalpidae): An island-hopping mite pest in the Caribbean [Internet]. 2005. Available from: www.sel.barc.usda.gov/ acari/PDF/Raoiella%20indica-Kane%20 et%20al.pdf

[42] Etienne J, Flechtmann CHW. First record of *Raoiella indica* (Hirst, 1924) (Acari: Tenuipalpidae) in Guadeloupe and Saint Martin, West Indies. International Journal of

Acarology. 2006;**32**:331-332. DOI: 10.1080/01647950608684476

[43] Rodrigues JCV, Ochoa R, Kane E. First report of *Raoiella indica* Hirst (Acari: Tenuipalpidae) and its damage to coconut palms in Puerto Rico and Culebra Island. International Journal of Acarology. 2007;**33**:3-5. DOI: 10.1080/01647950708684493

[44] Welbourn C. Red Palm Mite *Raoiella indica* Hirst (Acari: Tenuipalpidae) [Internet]. 2007. Available from: http:// www.doacs.state.fl.us/pi/enpp/ento/r. indica.html

[45] Vásquez C, Quirós M, Aponte O, Sandoval MF. First report of *Raoiella indica* Hirst (Acari: Tenuipalpidae) in South America. Neotropical Entomology. 2008;**37**(6):739-740. DOI: 10.1590/S1519-566X2008000600019

[46] Carrillo D, Návia D, Ferragut F, Peña JE. First report of *Raoiella indica* (Acari: Tenuipalpidae) in Colombia. Florida Entomologist. 2011;**94**(2):370-371. DOI: 10.1653/024.094.0241

[47] Návia D, Marsaro AL Jr, da Silva FR, Gondim MGC Jr, de Moraes GJ. First report of the red palm mite, *Raoiella indica* Hirst (Acari: Tenuipalpidae), in Brazil. Neotropical Entomology. 2011;**40**(3):409-411. DOI: 10.1590/ S1519-566X2011000300018

[48] Flechtmann CHW, Etienne J. Un nouvel acarien ravageur des palmiers en Martinique, premier signalement de *Raoiella indica* pour les Caraïbes. Phytoma. 2005;**548**:10-11

[49] Beard JJ, Ochoa R, Bauchan GR, Welbourn WC, Pooley C, Dowling APG. External mouthpart morphology in the Tenuipalpidae (Tetranychoidea): *Raoiella* a case study. Experimental & Applied Acarology. 2012;**57**:227-255. DOI: 10.1007/s10493-012-9540-2

[50] Vásquez C, Egurrola Z, Valera R, Sanabria ME, Colmenárez Y. Anatomía y química foliar en especies ornamentales de Arecaceae: Posibles barreras a la alimentación de *Raoiella indica* Hirst (Acari: Tenuipalpidae). Gayana Botánica. 2015;**72**(2):256-264. DOI: 10.4067/S0717-66432015000200012

[51] Roda A, Dowling A, Welbourn C, Peña J, Rodrigues JC, Hoy M, et al. Red palm mite situation in the Caribbean and Florida. Proceedings of the Caribbean Food Crops Society. 2008;**44**:80-87

[52] Vásquez C, de Moraes GJ. Geographic distribution and host plants of Raoiella indica and associated mite species in northern Venezuela. Experimental & Applied Acarology. 2013;**60**:73-82. DOI: 10.1007/ s10493-012-9623-0

[53] Carrillo D, Amalin D, Hosein F, Roda A, Duncan RE, Peña JE. Host plant range of *Raoiella indica* (Acari: Tenuipalpidae) in areas of invasion of the new world. Experimental & Applied Acarology. 2012;**57**:271-289. DOI: 10.1007/s10493-011-9487-8

[54] Oliveira DC, Prado EP, de Moraes GJ, de Morais EGF, Chagas EA, Gondim CMG Jr, et al. First report of *Raoiella indica* (Acari: Tenuipalpidae) in southeastern Brazil. Florida Entomologist. 2016;**99**(1):123-125. DOI: 10.1653/024.099.0124

[55] Rodrigues JCV, Antony LMK. First report of *Raoiella indica* (Acari: Tenuipalpidae) in Amazonas state, Brazil. Florida Entomologist. 2011;**94**(4):1073-1074. DOI: 10.1653/024.094.0452

**123**

**Chapter 8**

**Abstract**

Perception

*Muhammad Sarwar*

ing creatures for pests controlling.

have been greatly encouraged [29–31].

**1. Introduction**

**Keywords:** arthropod, Acari, mite, pest, Phytoseiidae, control

Biology and Ecology of Some

Predaceous and Herbivorous Mites

Mites are numerous species of minute arthropods, members of class Arachnida subclass Acari or Acarina and pests of many economic prominence living in a wide range of habitats. Mites are predators and parasites, performing crucial means of biological control, essential herbivores and detritivores, acting fungivorous and saprophytic, vectors of diseases, and play vital role in soil formation. These live on plants and animals, in the depths of ocean, in soil and fresh or brackish water, in lungs of birds and animals, in stored grains and stored products, on leaves of rainforest, and in human clothes and bedding. In spite of magnificent diversity of predaceous, phytophagous and granary mites found on plants and stored grains, these are often overlooked, and even skilled zoologists may be unaware of their importance. This chapter aims to provide an updated analysis of their biology, life history, reproduction and ecology to fill gap in our understanding of these fascinat-

Mite complex is worldwide in its distribution in all regions of globe and more prominent in tropical a well as subtropical climates. Mites can be either inflicting damage to humans and animals [1, 2], or pestilent that feed on plants [3] and stored commodities [4–23], otherwise predacious which are the carnivorous of leaffeeding mites and other pests [24]. All harmful types of mites are able to devastate agricultural crops, fruits and vegetables [25, 26]. During the previous few decades, owing to increasing concerns over health, environment and pest resistance risks accompanying with chemical control, and the use of alternate pest management strategies has received considerable attention [27, 28]. In this context, the uses of generalist predators that can perform as a broad spectrum fighters against pests

Currently, mites belonging to the family Phytoseiidae (Arachnida:

Mesostigmata) are economically important predators of some phytophagous mites and insects in greenhouses or field crops. Amongst others predators, mass reared phytoseiid mites are commercially available and used, against spider mites, thrips

Important from the Agricultural

## **Chapter 8**

*Pests Control and Acarology*

indica.html

Acarology. 2006;**32**:331-332. DOI: 10.1080/01647950608684476

[50] Vásquez C, Egurrola Z, Valera R, Sanabria ME, Colmenárez Y. Anatomía y química foliar en especies ornamentales de Arecaceae: Posibles barreras a la alimentación de *Raoiella indica* Hirst (Acari: Tenuipalpidae). Gayana Botánica. 2015;**72**(2):256-264. DOI: 10.4067/S0717-66432015000200012

[51] Roda A, Dowling A, Welbourn C, Peña J, Rodrigues JC, Hoy M, et al. Red palm mite situation in the Caribbean and Florida. Proceedings of the Caribbean Food Crops Society.

[53] Carrillo D, Amalin D, Hosein F, Roda A, Duncan RE, Peña JE. Host plant range of *Raoiella indica* (Acari: Tenuipalpidae) in areas of invasion of the new world. Experimental & Applied Acarology. 2012;**57**:271-289. DOI: 10.1007/s10493-011-9487-8

[54] Oliveira DC, Prado EP, de Moraes GJ, de Morais EGF, Chagas EA, Gondim CMG Jr, et al. First report of *Raoiella indica* (Acari: Tenuipalpidae) in southeastern Brazil. Florida

Entomologist. 2016;**99**(1):123-125. DOI:

[55] Rodrigues JCV, Antony LMK. First report of *Raoiella indica* (Acari: Tenuipalpidae) in Amazonas state, Brazil. Florida Entomologist. 2011;**94**(4):1073-1074. DOI: 10.1653/024.094.0452

2008;**44**:80-87

s10493-012-9623-0

10.1653/024.099.0124

[52] Vásquez C, de Moraes GJ. Geographic distribution and host plants of Raoiella indica and associated mite species in northern Venezuela. Experimental & Applied Acarology. 2013;**60**:73-82. DOI: 10.1007/

[43] Rodrigues JCV, Ochoa R, Kane E. First report of *Raoiella indica* Hirst (Acari: Tenuipalpidae) and its damage to coconut palms in Puerto Rico and Culebra Island. International Journal of Acarology. 2007;**33**:3-5. DOI: 10.1080/01647950708684493

[44] Welbourn C. Red Palm Mite *Raoiella indica* Hirst (Acari: Tenuipalpidae) [Internet]. 2007. Available from: http:// www.doacs.state.fl.us/pi/enpp/ento/r.

[45] Vásquez C, Quirós M, Aponte O, Sandoval MF. First report of *Raoiella indica* Hirst (Acari: Tenuipalpidae) in South America. Neotropical

Entomology. 2008;**37**(6):739-740. DOI: 10.1590/S1519-566X2008000600019

[46] Carrillo D, Návia D, Ferragut F, Peña JE. First report of *Raoiella indica* (Acari: Tenuipalpidae) in Colombia. Florida Entomologist. 2011;**94**(2):370-371. DOI:

[47] Návia D, Marsaro AL Jr, da Silva FR, Gondim MGC Jr, de Moraes GJ. First report of the red palm mite, *Raoiella indica* Hirst (Acari: Tenuipalpidae), in Brazil. Neotropical Entomology. 2011;**40**(3):409-411. DOI: 10.1590/ S1519-566X2011000300018

[48] Flechtmann CHW, Etienne J. Un nouvel acarien ravageur des palmiers en Martinique, premier signalement de *Raoiella indica* pour les Caraïbes.

[49] Beard JJ, Ochoa R, Bauchan GR, Welbourn WC, Pooley C, Dowling APG. External mouthpart morphology in the Tenuipalpidae (Tetranychoidea): *Raoiella* a case study. Experimental & Applied Acarology. 2012;**57**:227-255. DOI: 10.1007/s10493-012-9540-2

Phytoma. 2005;**548**:10-11

10.1653/024.094.0241

**122**
