Colonization and development of oribatid mite communities (Acari: Oribatida) on post-industrial dumps

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Colonization and development of oribatid mite communities

(Acari: Oribatida)

on post-industrial dumps

Piotr Skubała

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Colonization and development of oribatid mite communities

(Acari: Oribatida) on post-industrial dumps

Honour species Pay attention to small organisms

Wa l t e r & Pr o c t o r (1 9 9 9 )

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PRACE NAUKOWE

UNIWERSYTETU ŚLĄSKIEGO W KATOWICACH

NR 2219

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Colonization and development of oribatid mite communities

(Acari: Oribatida)

on post-industrial dumps

Piotr Skubała

Wydawnictwo Uniwersytetu Śląskiego Katowice 2004

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E ditor o f Series: Biologia

Paweł Migula

R eview ers

Mieczysław Górny Wojciech Niedbała

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4. Physical and chemical properties of the upper (A) and lower (B) sections of soil samples from Chorzów (4a), Katowice Wełnowiec (4b), Zabrze Biskupice (4c), Zabrze Makoszowy (4d), Katowice Murcki (dump) (4e), Katowice Murcki (sedimentation tank) (4f) and Brzeszcze (4g), from which oribatid mites were extracted

5. Check-list of plant species on the study dumps and in the neighbouring biotopes

6. Check-list of oribatid species on the study dumps and in the neighbouring biotopes

7. General view of the study sites at seven localities (34 colour photos) 8. SEM-photos of selected oribatid species

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We know more about the m ovem ent o f celestial bodies than about the soil u n

derfoot.

Le o n a r d o Da Vi n c i, c i r c a 1 5 0 0 ’s

Preface - soil as the basis for civilization

For centuries soil and the life in it were o f little interest to humankind. Two centuries ago, in 1788, Gilbert White called our attention to life in the soil. He discussed seven functions o f earth

worms: “by boring, perforating, and loosening the soil, and render

ing it pervious to rains and the fibres o f plants, by drawing straws and stalks of leaves and twigs into it; and, most of all, by throwing up such infinite numbers of lumps o f earth called worm-casts [...]”

(A l l e n , 1900). The value of earthworms in the soil system is better known from Charles Darwin’s classic book The form ation o f vegetable mould through the action o f worms with observations on their habits.

However, soil biology began to develop after the Second World War.

There was no comprehensive publication on the soil fauna until 1950 when two books with the same title “Bodenbiologie” w ritten by Kuhnelt and Franz, summarised the knowledge of soil fauna up to that time (V e e r e s h & R a ja g o p a l, 1988). The situation has not changed much since. There are only a few academic books on soil biology, and these are not new ones; and soil science is rarely taught at universities, at least in Poland. The true enigma is that although decomposition is the equal o f photosynthesis in ecosystem impor

tance, and half or more o f terrestrial biodiversity may be tied to the soil-litter system, the study of soil biology has been neglected ( W a l t e r

& P r o c t o r , 1999).

The soil is a living organism o f fabulous complexity. Soil systems contain some of the most species-rich communities in nature. Most authors describe soil communities as being am ongst the most species-rich components o f terrestrial ecosystems (A n d e rs o n , 1975a, 1978; G h ila r o v , 1977; S t a n t o n , 1979). Well-developed tem perate woodland soils may contain up to a thousand species of soil fauna

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(A n d e r s o n , 1975a). The calculation made by F itta k a u & K lin g e (1973) is even more impressive; they estimated that 80% of the total animal biomass in the Amazonian rainforest is soil fauna. U s h e r et al. (1979) used impressive words to describe soil communities as “the poor m an’s tropical rainforest” . It is noteworthy that only a proportion of all the soil animal species has been described and very little is known about their role, community structure and dynamics.

Research concerning soil is not purely an academic subject. The soil is the very basis of earth’s productivity. It is fundamental to agriculture and forestry, water purification and biogeochem ical cycling, and is the grounding for civilization ( B e h a n - P e l l e t i e r &

N e w to n , 1999). This is particularly true where human activity tends to induce irreversible disturbances (L e b ru n , 1979). At a time when demographic pressure is too high, and when the needs o f human population are intense and immense, it is wise to realize that the soil is central to human survival. Meanwhile, soil biology has fallen somewhat behind advances in the understanding of other types of com m unities ( G i l l e r , 1996). Soils are still the least understood habitats on Earth, while also being among the most biologically diverse ( B e h a n - P e lle t ie r & N e w t o n , 1999).

From both theoretical and applied perspectives, this state of affairs is surprising from three points of view ( G i l l e r , 1996).

1. Firstly, in terms of the importance of soils to global biodiversity.

2. Secondly, in terms o f the ecosystem processes, in particular those that occur in the soil. The soil performs a fundamental role as the location where 60% to 90% of terrestrial primary production is decomposed. Soil fauna appears to be the major regulatory agent of soil processes affecting the physical and chemical fertility o f soils.

Moreover, a full understanding of above-ground ecosystem processes is not possible without consideration of processes occurring in the soil (M a y, 1997; O s l e r & B e a t t ie , 2001).

3. Thirdly, soil fauna can offer a suite o f bioindicators for clas

sification of soils and detection of disturbances and pollution.

The place of mites, which are the subject o f this study, within the soil, is important. Their quantitative and qualitative roles in energy flow are not fundamental; however, their interactions with other members o f soil biota are of great ecological meaning. The contribution o f Acarology to the understanding of our “black box” , which may be a good description o f the soil system, could be immense.

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Acknowledgements

The author would like to thank Mrs Ewa Skipirzepa, Ms Sabina Słomian and Dr. Aleksander Stodółka for excellent technical assis

tance. I must especially acknowledge Ms Krystyna Pilarczyk for her great effort in sampling, extracting and sorting the microarthropods.

I wish to express my grateful thanks to Prof. Wojciech Niedbała (A. Mickiewicz University in Poznań) and Prof. Stanisław Seniczak (University o f Technology and Agriculture in Bydgoszcz) for confirm

ing identifications of Phthiracaridae and juvenile forms, respectively.

I am also grateful to Dr. Zbigniew W ilczek and Dr. Gabriela Woźniak (University o f Silesia) for making phytosociological surveys o f the study sites. Many thanks to Dr. Tomasz Zaleski (Agriculture Academy in Cracow) for undertaking chemical and physical analy

ses of soil and Dr. Ryszard Kulik (University o f Silesia) for taking excellent photos of the study sites. I wish to thank Dr. hab. Jerzy Błoszyk (A. Mickiewicz University in Poznań) and Dr. Ritva Niemi (University of Turku, Finland) for making SEM-photos of selected oribatid species.

I w arm ly thank Dr. John Parker (Forest Research, Alice Holt Lodge, United Kingdom) for checking the manuscript. Last but not least I thank my wife Elvira and daughter Kaja for their forbear

ance in the period o f my scientific inquiries and preparation of the book.

The studies were supported by grant project No. 6 P04F 035 18 from the Polish State Committee for Scientific Research and by the grant projects “Colonization of post-industrial dumps by saprophagous oribatid mites (Acari: Oribatida)” (1998) and “Studies on succession o f oribatid communities (Acari: Oribatida) on post-industrial dumps” (1999) from the University o f Silesia.

I dedicate this book to those scientists whom I have met in Poland and abroad and who have supported me in my scientific career by interesting discussions and helpful comments. Special thanks to Prof. Paweł Migula (University o f Silesia) and Prof. Henryk Sko

limowski (Ann Arbor University in Michigan, U.S.A., Department of Ecological Philosophy at Technical University in Łódź) who have always shown personal interest in my scientific and private life and have supported me in the fields o f ecology and environm ental philosophy.

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M ites (Acari) are representatives o f the taxonom ic dilem m a facin g researchers who study soil ecosystem processes.

Be h a n- Pe l l e t ie r & Ne w t o n ( 1 9 9 9 )

Chapter i . Introduction

1.1. Oribatid mites - life hidden in the soil

Oribatida or moss mites are small, chelicerate arthropods, im portant representatives o f mites (Acari). Mites are ubiquitous and, with the exception o f the open oceans, they exist in every sort of terrestrial, aquatic, arboreal and parasitic habitat ( W a l t e r & P r o c t o r , 1999). They are found at eveiy elevation and every latitude, from the Arctic to the Antarctic. Mites have a diversity o f function in the ecosystem, as shown by the range o f feeding guilds to which they belong (M o o r e et al., 1988). They include predators, parasites, fun

gal feeders, root feeders, bacterial feeders, omnivores, and scaven

gers (K r a n t z , 1978). Ignoring mites, however, is a mistake. They are not passive inhabitants o f ecosystems; rather they are strong interactors, important indicators o f disturbance in ecosystems and major components of biological diversity ( W a l t e r & P r o c t o r , 1999).

More than any other habitat, the soil litter stratum is the province of mites. Two-thirds o f the mite fauna occur in this habitat ( W a l t e r

et al., 1996). Any true understanding o f the soil system must in

clude knowledge o f the mite fauna.

Oribatid mites have successfully invaded all compartments of the biosphere (B e rn in i, 1986). They constitute the main component of acarine populations in the soil. They are not confined to the soil, however, and may occur in considerable numbers in the above

ground parts o f vegetation, among aquatic plants, in stored food, in the marine littoral zone, and in house dust. Temperate forests with well-developed surface organic layers and a predominance of fungal over bacterial decomposition are home to the highest diver

sities of oribatids. Oribatid mites can comprise about 50% o f the

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t o t a l m ic r o a r t h r o p o d fa u n a (Gonzalez & Se as te d t, 2000). D e n s it ie s o f 50 000-250 000 o r m o r e m ite s p e r s q u a r e m e t r e in t h e u p p e r

10 c m o f s o il a r e c o m m o n ly r e p o r t e d (Pe te r s e n, 1982). Rajsk i (1961)

r e c o r d e d in th e P r im a e v a l B ia ł o w ie ż a F o r e s t a c o u n t o f 1 m illio n o r ib a t id m it e s p e r s q u a r e m e t r e . B u t e v e n t h e d r ie s t , h o t t e s t o r c o ld e s t o f s o ils a re d o m in a t e d b y A c a r i, a n d its m o s t c o n s p ic u o u s r e p r e s e n t a t iv e s - o r ib a t id m it e s (Wa lte r & Pr o c t o r, 1999). N u m e r o u s p e c u lia r , e p h e m e r a l a n d s m a ll h a b it a t s , s u c h a s d u n g , b ir d n e s ts , lic h e n t h a llu s , m o s s e s , fu n g a l m y c e lia , m u s h r o o m s , th e in s id e o f c o n ife r n e e d le s , fo o d p r o d u c t s , e tc ., a re c o lo n iz e d b y o r ib a t id s (Lebrun & Van Str a ale n, 1995). E v e n m o r e p e c u lia r m ic r o h a b it a t s c a n p r o v id e a h o m e fo r o r ib a t id s , e .g . lu m b r ic id g a lle r ie s (Le b r u n &

Wa u t h y, 1981), a e r ia l r o o t s o f o r c h id s (Denm ark & Wo o d r in g, 1965),

c a v it ie s o f c u r c u lio n id b e e t le e ly t r a s (Gre ssit e t a l., 1966), o r a n t n e s t s (Aoki e t a l., 1994). E v e n th e m a n - m a d e e n v ir o n m e n t h a s b e e n in v a d e d b y s o m e e n d e m ic s p e c ie s b e lo n g in g to th e d o m e s t ic g e n u s

Cosmochthonius o r s p e c ie s o f Trimalaconothrus liv in g in e n v ir o n m e n t s s u c h a s s w im m in g p o o ls (Tagam i e t a l., 1992). Olsza n o w sk i

(1996) fo u n d o r ib a t id s in a n a q u a r iu m . O r ib a tid s a re a ls o fr e q u e n t ly a d o m in a t in g g r o u p a m o n g m ic r o a r t h r o p o d s o n p o s t - i n d u s t r i a l w a s t e la n d s (Da v is, 1963; Sk u b a ła, 1995). N o in fo r m a t io n is a v a ila b le c o n c e r n in g th e g r o u n d - w a t e r s u b s y s t e m , a lt h o u g h o r ib a t id s c o u ld liv e in u n d e r g r o u n d s t r e a m s a n d la k e s , a s s u g g e s t e d b y t h e ir p r e s e n c e in c o ld s p r in g s (Lebrun & Van Str a a le n, 1995). In c o n c lu s io n w e c a n s a y t h a t a ll n ic h e s c o n t a in in g o r g a n ic m a t t e r c o n t e n t a r e c o lo n iz e d b y o r ib a t id s , u s u a lly in h ig h n u m b e r s .

Oribatid mites constitute the order richest in species in the subclass Acari. Together with the Actinedida (Prostigmata), Acaridida (Astigmata) and Endeostigmata they constitute a group o f mites of common origin called the Acariform es (Za c h v a t k in, 1947) or Actinotrichida (Ha m m e n, 1972). Their evolutionary history is appar

ently a long one. They have the richest fossil record o f any mite group, dating back to the Devonian, or 420-430 million years ago

(Coleman & Cr o s sle y, 1996). The origin and phylogeny of the mites and oribatids is not clear yet. And the systematics o f the Oribatida is therefore not finally accepted by the society of acarologists. The Oribatida are grouped in two supercohorts: Oribatida Inferiores or

“lower oribatid mites” and Oribatida Superiores or “higher oribatid m ites” (Gr a n d je a n, 1954, 1969). Lower oribatid mites include five cohorts - the Palaeosomata, Enarthronota, Parhyposomata, Mixo- nom ata and Desm onom ata (Gr a n d j e a n, 1954, 1969). O ribatida Inferiores may be broadly characterized as species with contiguous genital and anal shields occupying the entire length of the genitoanal field, and with leg tibiae and genua of uniform length and shape

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(K r a n t z , 1978). The great majority o f described Oribatida belongs to Oribatida Superiores. The supercohort may be broadly character

ized as aptychoid species with rounded, generally well separated genital and anal fields on a distinct ventral shield, and with leg tibiae distinctly longer and o f a different shape than the adjacent genua.

It comprises two cohorts - the Brachypylina and the Poronota.

Cohortal separation is based on presence or absence o f dorsal pores and pteromorphae (K r a n t z , 1978).

Oribatids are a highly morphologically diverse group of mites. The species diversity and the variability within species greatly differ in separate morpho-ecological types. K r iv o lu t s k y (1965, 1968, and 1995) divided oribatids into 6 groups, including 16 m orpho-ecological types. These groups are: inhabitants o f the soil surface; small dwellers in narrow soil pores; deep-soil weakly sclerotized oribatids;

inhabitants of the substrate, able to widen the pores; non-special- ized forms; and inhabitants of wet habitats and aquatic systems.

Oribatid mites in general have conservative life histories. N o r t o n

(1994) claimed that a low metabolic rate is the “driving force” o f the oribatid life cycle. A low metabolism results in slow development, low reproductive output, limited body size and energy storage, and a long adult life. Iteroparity is common in oribatids and “X-selected”

species prevailed because of the energy costs of maintenance require

ments and dispersal (Fig. 1). All these characteristics may result in

low metabolism

slow develoDment low fertility limited body size

and energy storage

long life stable populations

1

_ rapid increase in abundance long development

and long life

survival during food shortage Fig. 1. Oribatid mite life strategy

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r a t h e r s ta b le p o p u la t io n s a s d e m o n s t r a t e d b y m a n y a u t h o r s (Lebrun,

1964; M it c h e ll, 1977; S c h a tz, 1985). T h e d e v e lo p m e n t tim e o f o r i b a tid s is g e n e r a lly lo n g a n d is w o r t h y o f r e g is t r a t io n in th e Guinness Book o f Records. T h e r e c o r d in t h is c o n t e x t is h e ld b y th e b o r e a l in h a b it a n t Ceratozetes kananaskis. Its m e a n d e v e lo p m e n t tim e is 770

d a y s , w h ile th e a d u lt life s p a n a p p r o a c h e s 4 y e a r s (M it c h e ll, 1977).

L o n g d e v e lo p m e n t a n d lo n g life r e s u lt s in a lo w c a p a c it y fo r p o p u la tio n in c re a s e . O r ib a tid s a re a b le to s u r v iv e fo r a lo n g p e r io d d u r in g fo o d s h o r ta g e . T h e s e c o n s t r a in e d “ K a t t r ib u t e s ” a r e p le s io t y p ic fo r o r ib a t id s a n d h a v e o fte n p la y e d a r o le in p r e a d a p t a t io n o f s p e c ie s in v a d in g e x t r e m e h a b it a t s (B ü ck in g e t a l., 1998). M o s t o f t h e s e c h a r a c t e r is t ic s a re u n u s u a l a n d illu s t r a t e w h y O r ib a t id a a r e a n e x c e p tio n w ith in th e a c a r o lo g ic a l w o r ld (Lebrun & Van S tra a len , 1995).

Thelytokous parthenogenesis is again peculiar to oribatids. Many hundreds of species of oribatids are parthenogenetic, and never pro

duce functional male offspring (Walter & Proctor, 1999). Approximately 10% of the species studied exhibit this clonal mode of gene-pool trans

mission and maintenance. Perhaps 1% of known insect species and 0.1% of known members of the animal kingdom are obligated parthenogens. In this respect, oribatid mites present a striking anomaly

(Lebrun & Van Straalen, 1995; Norton & Palm er, 1991).

Surprisingly, knowledge of feeding biology o f oribatid mites is poor and the available information is in part contradictory. Unlike the vast majority of other Arachnida, which adopted a carnivorous mode o f life, oribatids are, for the most part, vegetarian. Gut content analyses indicate that most oribatid mites ingest a wide range of food materials including spores of various fungal species, living and dead plant material, moss, lichens, conifer pollen and carrion (Behan- Pelletier & Hil l, 1983). Dietary features include poor nutritive value o f ingested food, relatively low ingestion rates, and consequent minimal assimilation rates (Be r t h e t, 1964a; Lu x to n, 1972, 1979;

Wa l l w o r k, 1983), which constrain their life-history param eters

(No r t o n, 1994).

Mites link together many components of soil food webs and a study of their behaviour can shed light on our understanding of ecosystem functioning (Wa lte r & Pr o c t o r, 1999). Unfortunately, rather little information is available on their relationships with other soil biota in comparison with other soil arthropods (Wa llw o rk, 1983).

Let’s consider the present knowledge on the role o f oribatid mites in the soil system.

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The dormant microbes require the “k iss”

o f arthropod “Prince C h arm in gs” to awaken.

Wa l t e r & Pr o c t o r ( 1 9 9 9 )

1.2. Oribatids perform an important

“ecological service”

The sheer numbers of oribatids, at least in most soil systems, suggest that they play key, but still virtually unexplored, roles in their environment (soil). It is generally accepted today that oribatid mites are responsible for the indispensable process of antiphotosynthesis, achieving this by small, multiple and comple

mentary stages that are a direct result of the species diversity of this group (W a l l w o r k , 1976). Mites play an essential part in the biological fertility of the soil and they affect soil energetics. Their activity contributes greatly to organic decomposition, the synthesis o f humus, the restitution of biogenic elements, and the stimulation of fungal and bacterial metabolism (C r o s s le y , 1977a; L e b ru n , 1979;

N o r t o n , 1986; R u sek , 1975). It is noteworthy that oribatids are the most important group o f arachnids from the standpoint o f direct and indirect effects in the development and maintenance o f soil struc

ture in organic horizons ( M o o r e et al., 1988; N o r t o n , 1986). Many species sequester calcium and other minerals in their thickened cuticle (N o r t o n & B e h a n - P e lle t ie r , 1991). Thus, their bodies may form important “sinks” for nutrients, especially in nutrient-limited envi

ronments (C r o s s le y , 1977b).

F o u r m a jo r p r in c ip le s c a n b e t a k e n in to a c c o u n t c o n c e r n in g th e p a r t ic u la r fu n c tio n s o f o r ib a t id m ite s .

• The result o f mechanical breakdown and fragm entation is to increase the active surface o f litter, enhancing its colonization by microorganisms, especially saprophytic fungi. The oribatid activ

ity facilitates and accelerates the leaching o f hydrosoluble ele

ments and hydration of the organic matter (B e h a n - P e lle t ie r & H i l l ,

1983; C u r r y , 1969; W itkam p, 1971).

• The digestive transit ensures physical and chemical change as well as biological breakdown. This is accompanied by a m ixing of mineral and organic elements and microorganisms (L eb ru n , 1979).

• The production of faecal pellets creates a highly fertile environ

ment. It facilities growth of roots and the germination o f seeds

(H a a r lö v , 1960; L e b r u n , 1979; P a n d e & B e r t h e t , 1973). The con

sumption o f dead roots by saprophagous oribatids is also of great importance because it considerably increases soil porosity and the development o f humus (L e b ru n , 1979).

2 - Colonization. 17

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• T h e g r a z in g o f m it e s s t im u la t e s m ic r o f lo r a l a c t iv it y . T h is f u n c t io n m a y b e o f m a jo r im p o r t a n c e . T h e y a id t h e d i s p e r s a l o f b a c t e r ia a n d fu n g i, b o t h e x t e r n a lly o n t h e ir b o d y s u r fa c e a n d b y in g e s t in g s p o r e s t h a t s u r v iv e p a s s a g e t h r o u g h t h e ir a l im e n t a r y t r a c t (Be h an- Pe lle tie r & Wa l t e r, 2 0 0 0 ). M ic r o b e s h a v e l im it e d a b ilit ie s to m o v e fr o m o n e r e s o u r c e p a t c h to a n o th e r . O n c e th e e n e r g y in a p a r t ic u la r p a t c h h a s b e e n e x p e n d e d , m ic r o b a l b io m a s s s h u t s d o w n a n d r e m a in s d o r m a n t u n t il n e w r e s o u r c e s b e c o m e a v a ila b le . Lavelle (1 9 9 7 ) d e s c r ib e d t h is p h e n o m e n o n a s th e “ S le e p in g B e a u t y P a r a d o x ” . Walte r 8s Pr o cto r (1 9 9 9 ) c la im e d t h a t “ d o r m a n t m ic r o b e s r e q u ir e th e »k is s « o f a r t h r o p o d »P r in c e C h a r m in g s « to a w a k e n ” . T h u s , m ic r o b ia l p r o p a g u le s a r e m ix e d w it h fr e s h r e s o u r c e s in th e fa e c a l p e lle t s a n d a r e t r a n s p o r t e d to n e w s ite s .

S o m e d a t a s u p p o r t th e v ie w th a t o r ib a t id s a r e e s s e n t ia l fo r e f fic ie n t d e c o m p o s it io n a n d n u t r ie n t c y c lin g . N e a r ly 5 6 % o f th e n e t p r o d u c t io n o f fu n g i is c o n s u m e d b y m y c o p h a g o u s m it e s p e c ie s (M c B r a y e r e t a l., 1 9 7 4 ). B u t c h e r e t al. (1 9 7 1 ) e s t im a t e d t h a t a d u lt o r ib a t id m ite s d ir e c t ly m e t a b o lis e o n ly 1 .8 % o f th e e n e r g y in fo r e s t litte r, e v e n t h o u g h in o n e y e a r t h e y in g e s t a n a m o u n t o f m a t e r ia l e q u a l to a b o u t 5 0 % o f a n n u a l l e a f fa ll. G h ila r o v (1 9 6 3 ) p r o v e d t h a t o r g a n ic d e c o m p o s it io n is fiv e tim e s fa s t e r w h e n m ic r o o r g a n is m s a n d m ite s w o r k t o g e t h e r th a n b y m ic r o o r g a n is m s a lo n e . E v e n t h o u g h t h e s e g e n e r a l c h a r a c t e r is t ic s o f o r ib a t id m ite s a re k n o w n , t h e r o le o f m o s t m ite s p e c ie s in e c o s y s t e m fu n c t io n in g is u n c le a r , a s is th e c a s e w it h o t h e r g r o u p s o f s o il o r g a n is m s (B e h a n - P e l l e t i e r & N e w to n ,

19 9 9 ).

In v ie w o f t h is , a n d s in c e t h e y n o r m a l ly e x c e e d m o s t o t h e r a r t h r o p o d s in a b u n d a n c e a n d d iv e r s it y (s e e n e x t p a r t o f th is c h a p te r ), o r ib a t id m ite s s h o u ld b e r e g a r d e d as “ k e y in d u s t r y ” a n im a ls in d e c o m p o s e r fo o d c h a in s (W a llw o r k , 1 9 8 3 ).

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We must take it for granted that a large part o f the mite fauna o f the world will rem ain unsam pled, unnam ed, and u n classified (not to m ention unwept, unhonoured, and unsung) for decades to come.

Ma y ( 1 9 7 8 )

1.3. Contribution of oribatids to global biodiversity

Exploring the diversity o f life will be one of the essential topics in 21st century ecology. The importance of biodiversity arises from the fact that the world depends on self-sustaining biological sys

tems that include many kinds o f organisms. Knowledge of biodiversity is required to understand the natural and artificial changes it may undergo. Furthermore, such knowledge permits the wise use and m anagement o f ecosystems, both as elements of natural heritage and as reservoirs of actual and potential resources.

Biological diversity has been used to refer to almost any measure (taxonomic, numerical, genetic, etc.) of the variety o f organisms that live in a particular place. Three dominant themes can be distin

guished in this field, namely:

• a c c o u n t in g fo r th e d iv e r s it y ,

• d e t e r m in in g h o w th is d iv e r s it y is m a in ta in e d ,

• e n u m e r a t in g p r in c ip a l fu n c tio n s p r o v id e d b y th e d iv e r s it y o f life.

M o n it o r in g th e v it a lit y o f s o il b io t a is o n e o f th e a c c e p t e d is s u e s to b e fo u n d in th e G lo b a l D iv e r s ity A s s e s s m e n t (Heywood & Ga rd n e r, 1 9 9 5 ). T h e r a t io n a le fo r in c lu d in g a r t h r o p o d s in b io d iv e r s it y s t u d ie s h a s b e e n e s t a b lis h e d (Didham e t a l., 19 9 6 ; Eh r lic h, 1 9 8 8 ; Wil so n, 19 8 8 ; Verhoef & Br u ssard, 1 9 9 0 ; Win c h e ste r, 1 9 9 7 ). T h e c o n tr ib u t io n o f m ite s to g lo b a l b io d iv e r s it y is fa r fr o m b e in g a p p r e c i

a te d . M it e a n d o r ib a t id b io d iv e r s it y m a y b e o n e o f th e r ic h e s t r e s e r v o ir s o f s p e c ie s in th e w o r ld . Walter & Proctor (1 9 9 9 ) p r e s e n te d th e h y p o t h e s is a b o u t lik e ly m ite m e g a d iv e r s it y . T h e r e a r e a p p r o x i

m a t e ly 4 5 0 0 0 n a m e d s p e c ie s o f A c a r i (T a b le 1). R e c e n t e s tim a te s o f g lo b a l a c a r in e d iv e r s it y r a n g e fr o m b e tw e e n h a lf to o v e r 1 m illio n s p e c ie s (Walter & Pr o ctor, 1 9 99 ). So th e n u m b e r o f s p e c ie s o f m ite s d e s c r ib e d so fa r is e s t im a t e d to r e p r e s e n t b e t w e e n 4 % a n d 8 % o f t o t a l m ite d iv e r s it y . As r e g a r d s o r ib a t id m ite s , 1 1 ,0 0 0 s p e c ie s in m o r e t h a n 1 1 0 0 g e n e r a h a v e b e e n d e s c r ib e d ; t h e ir t o t a l s p e c ie s r ic h n e s s c o u ld b e 3 to 10 t im e s h ig h e r . In t e m p e r a t e r e g io n s , th e a c a r o fa u n a s a r e c e r t a in ly d iv e r s e , b u t n o t e x c e p t io n a lly so . If th e A c a r i a r e a h y p e r d iv e r s e g r o u p , t h e n th e m a jo r it y o f t h e m w a it to b e e x p lo r e d in t h e a c a r o l o g i c a l l y u n e x p lo r e d t r o p ic s (Wa l t e r &

Pr o c to r, 1 9 9 9 ).

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T a b l e 1 Current and expected diversity in m ites (Acari)

Families Genera Species Species estimates minimum maximum

Opilioacariform es 1 9 17 85 170

Parasitiform es

Holothyrida 3 9 32 160 320

Ixodida 3 12 880 1 000 1 200

M esostigm ata 73 567 11632 97 520 200 500

Total 79 588 12 544 98 680 202 020

Acariform es

Endeostigm ata 11 25 120 1 200 2 400

Sarcoptiformes

Oribatida 150 1 100 11 000 33 000 110 000

Astigm ata 70 627 4 500 90 000 180 000

T rom bidiform es 120 1 323 17 050 317 250 637 500

Total 351 3 075 32 670 441 450 929 900 Total Acari

Percentage of species described to date

431 3 672 45 231 540 215

8,4

1 132 090

4,0 S o u rce : WALTER & PROCTOR (1999).

The future o f oribatids with respect to biodiversity does not look very optimistic. Numbers o f described taxa increase annually, but the Oribatida although fascinating and diverse, is not widely stud

ied by taxonomists. Only eight scientists work with oribatids in Poland. Data on mites and on oribatid diversity in tropical ecosys

tems is especially rare (Wa lte r & Pr o cto r, 1999). The reasons for this neglect are the minute size of individuals (0.1 to 3 mm in length), difficulty of identification, their cryptic habits, and their relative lack of economic importance.

C o n t in u in g b io d iv e r s it y lo s s e s a n d th e e v e r - w o r s e n in g q u a lit y o f th e e n v ir o n m e n t a r e w e ll- d o c u m e n t e d fa c ts . S c ie n t is t s d if fe r in e s tim a tio n o f th is p r o c e s s . A c c o r d in g to c o n s e r v a t iv e c a lc u la t io n s m o r e t h a n 13 s p e c ie s b e c o m e ‘e x tin c t e a c h d a y . A c c o r d in g to p e s s im is tic e s tim a tio n s , it is p o s s ib le t h a t 410 s p e c ie s d is a p p e a r e v e r y d a y (Goo dland, 19 9 1 ). L e g a l e n v ir o n m e n ta l c o n s t r a in t s to s to p lo s s e s o f b io d iv e r s it y a re s till la r g e ly u n s a tis fa c t o r y . T h e s c ie n t ific c o m m u n it y d o e s in s u ffic ie n t w o r k o n th is p r o b le m . Lebrun 8& Van Straalen

(1995) u n d e r lin e d th e o b s t in a c y o f s c ie n t is t s w h o w is h to u n d e r s t a n d c o m p le t e ly a ll th e m e c h a n is m s in v o lv e d b e fo r e s o u n d in g th e a la r m b e lls . Walter 8s Proctor (1999) w r it e a b o u t th e p r e s e n t w o r ld fa s c in a t e d b y m o le c u la r m a n ia , b u t w h e r e lit tle a t t e n t io n is g iv e n to d im in is h in g d iv e r s it y a n d th e d e s t r u c t io n o f n a tu r e . A s r e g a r d s a n im a l b io d iv e r s it y , a v e r t e b r o c e n t r ic v ie w is r a r e ly q u e s t io n e d , w h e r e a s in v e r t e b r a t e s m a k e u p 95% o f a n im a l b io d iv e r s it y . Do w e

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also have a significant decrease in oribatid mite species in the world?

Will species seen only under microscope disappear and will we not even record this lost? There are no answers to these questions. We do not have any calculations or suspicions. It is also one o f the reasons why acarology is a fascinating discipline.

1.4. Oribatids in biomonitoring studies

The use of bioindicators to derive biological information on certain environmental variables has a long tradition, especially in botany.

The potential indicator value o f soil invertebrates has often been emphasized by authors (V an S t r a a le n & V e r h o e f , 1997). As for the arthropod fauna, field studies have revealed correlations between groups of species and certain soil factors (H ä g v a r & A braham sen, 1984;

P o n g e , 1993), but no system for bioindication has been proposed

(Van S t r a a le n & V e r h o e f , 1997). Living in almost all biosphere m i

crohabitats, oribatid mites are certainly an appropriate group for surveying all the environmental compartments to detect any struc

tural or functional damage. This is particularly true for the soil where oribatids display high densities and diversities and so are well suited for biomonitoring studies (L e b ru n & V a n S t r a a le n , 1995; W a l t e r &

P r o c t o r , 1999).

Oribatid species and their communities offer several advantages for assessing the quality of terrestrial ecosystems ( B e h a n - P e lle t ie r ,

1999; L e b r u n & V a n S t r a a le n , 1995). Their high densities and diver

sities have been noted previously. They are easily sampled and they can be sampled at all seasons. It is noteworthy that oribatids are in close contact with defined microenvironmental conditions. When sudden changes occur, oribatids are unable to escape: they are sedentary or slow moving, lack marked dispersal mechanisms, and are therefore subjected directly to conditions o f stress. In addition, oribatids are directly exposed to toxicants, by contact, by direct ingestion o f soil particles and soil water, and through food-chain transfer (Fig. 2). In general, oribatids fulfil most criteria listed by

C ra n s to n (1990) in assessing the suitability of taxa for biomonitoring.

However, there are also negative aspects o f the use of oribatids in bioindication studies, e.g. their small size, difficulties in identifica

tion, the abundance o f individuals often encountered, some diffi

culties in standardising sampling and extraction, and time taken in sorting (H u n t, 1994).

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in direct contact with rapid reaction microenvironmental conditions to stress factors

OBSTACLES

difficulties in standarising lack of taxonomic knowledge sampling and extraction (especially juvenile forms) Fig. 2. Oribatids in biom onitoring studies

Mites are becoming increasingly used as bioindicators in soil.

Oribatids may demonstrate the impact of various human activities:

• a i r p o l l u t i o n ( An d r£ & Le b r u n, 1984; Be r in a e t a l . , 1989; Se n ic z a k e t a l . , 1997, 1998; St e i n e r, 1995),

• acid rain ( H ä g v a r & A m u n d s e n , 1981; H e n e g h a m 8s B o l g e r , 1996),

• cultivation (D e k k e r s et al., 1994; E h r n s b e r g e r 8sB u t z - S t r a z n y , 1993;

F r a n c h in i 8s R o c k e t , 1996),

• burning ( A^{n t o n y}, 2001; N^{o b l e} et al., 1989),

• use o f fertilizers ( G a t i l o v a , 1970; K o s k e n ie m i 8s H u h t a , 1986;

Ż y r o m s k a - R u d z k a , 1976),

• use o f herbicides and insecticides ( B h a t t a c h a r y a 8s J o y , 1980;

E d w a r d s 8s L o f t y , 1969; H o y , 1990; K o e h l e r , 1987; M u e l l e r et al., 1990),

• r a d i o a c t i v e p o l l u t i o n (Kr iv o l u t s k y, 1979; Kr iv o l u t s k y 8s Po k a r z h e v s k i,

1991),

• forest harvesting ( An t o n y, 2001; Hu h t a, 1976),

• reclam ation o f post-industrial dumps ( C r o s s 8s W ilm a n , 1982;

D u n g e r et al., 2001),

• soil contamination (D e n n e m a n 8sV a n S t r a a l e n , 1991; L u d w ig et al., 1991; S a lm in e n 8s S u lk a m , 1997; S i e p e l , 1995a; V a n S t r a a l e n et al., 1989),

• sewage water irrigation ( Di n d a l, 1977),

• trampling ( B o r c a r d 8s M a t t h e y , 1995; S c h a t z , 1983a).

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L in d en et al. (1994) list the different properties of soil animals, which can be potentially used as indicators of soil quality.

• Use o f single organism characteristics. There have been nu

merous studies on the ecological and reproductive response of oribatid species in the past decade (B e h a n - P e lle t ie r , 1999). S ie p e l

(1996) has shown how effective this approach can be in predict

ing species loss. Nevertheless, this method will become more precise, useful and widely adopted as knowledge of taxonomy and ecology o f species improves. No datasets exist to illustrate our knowledge on habitat and niche profiles of oribatid species (B eh a n - P e l l e t i e r , 1999).

• Use o f community level characteristics. This approach is more advanced. However, knowledge o f the taxonomy and ecology of species is still needed in order to use this method. Total mite density, species richness, dominance or constancy o f certain species, as well as various biodiversity indexes can be used to determine the degree of disturbance. From this dataset, appro

priate indicators can be selected once the stress has been defined

(L in den et al., 1994).

• Use o f biological process level. This method is still at a prelimi

nary stage. R u s e k (1986) presents an interesting example o f this approach. He studied the structure o f oribatid faeces and its contribution to soil structure.

Several authors have discussed the advantages of using oribatids in bioassay work and as ecological indicators, and the possibility o f using the above mentioned approaches, e.g. B e c k , 1994a; B e h a n - P e l l e t i e r , 1999; H u n t, 1994; It u r r o n d o b ie t ia et al., 1997; K r iv o lu t s k y ,

1970; L e b ru n & V a n S t r a a le n , 1995; N ie d b a ła , 1983; V a n S t r a a le n ,

1988.

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P o st-in d u s tria l d u m p s are a n “e x p e r i

m en ta l ra n g e ” to stu d y m a n y e c o lo g i

cal an d even ev o lu tio n a ry processes.

To k a r s k a- Gu z ik & Ro s t a ń s k i ( 2 0 0 1 )

chapter 2 ^. Save our dumps (SOD) as an entity

2.1. Post-industrial dumps as a “unique”

experiment for ecologists

Post-industrial wastelands are constantly increasing in area;

some o f them may be dangerous to human health (Fig. 3). These facts provide the major stimulus for the reclamation and restora

tion o f large devastated areas. The reclamation o f dumps, especially of toxic dumps, is a challenge and is a severe problem o f today. For this purpose, a knowledge and understanding o f soil development - mainly induced by soil organisms - is an important prerequisite for any form o f soil management on dumps ( Wa n n e r et al., 1998).

From the viewpoint o f reclamation, a practice very poorly developed at this time, a knowledge o f soil fauna development is required as an indicator for artificial cultivation techniques ( Du n g e r et al., 2001).

The restoration o f disturbed ecosystems has recently received increasing attention from ecologists and land-use managers. This is due, in a large part, to more stringent government regulations, which mandate extensive reclamation procedures followed by bio

logical monitoring efforts ( Pa r m e n t e r & Ma cMa h o n, 1987). It seems obvious that dumps should at first be managed biologically. In this way the negative influence o f dumps on the natural environment may be limited, and the beauty o f the landscape can be improved.

Nevertheless, vegetation cover on dumps may develop naturally.

Hitherto, reclamation measures taken on dumps have appeared unsatisfactory in many cases despite the high expenditure o f money and effort. Spontaneous succession o f plants is m ore desirable

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Fig. 4. The youngest part o f the coal-m ine dump at Zabrze Biskupice

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c&rojMiib

Fig. 5. The initial plant assem blage w ith Corynephorus canescens at site 1 on the coal-m ine dump at Zabrze Makoszowy

Fig. 6. The plant assem blage w ith Calamagrostis epigejos at site 2 on the coal-m ine dump at Zabrze Makoszowy

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Fig. 7. The plant assem blage with Betula pendula - Calluna vulgaris at site 3 on the coal-mine dump at Zabrze Makoszowy

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(W eidem an n et al., 1982), but unfortunately the spontaneous process of colonization and succession is a long-term process.

Much of the research on land restoration has been devoted to the reestablishment of vegetation and vertebrates. Soil fauna has been treated marginally in many restoration practices and studies, whereas we can obtain useful results for derelict land management, acquiring knowledge of all processes during succession on waste

lands. Studies on recolonization of arthropods on wastelands are few (Hawkins & C r o s s , 1982; M a je r , 1985; M a j e r et al., 1982; Neumann,

1971; N ic h o ls & B u r r o w s , 1985; U s h e r , 1979), whereas arthropods can drastically influence revegetation efforts via herbivory, seed predation, litter decomposition, pollination, and soil aeration (M a je r ,

1989a). More empirical studies of these processes on different kinds of dumps are urgently needed to better understand the factors affecting soil fauna developm ent in a post-industrial landscape

(F r o u z et al., 2001).

The growing area o f dumps is a real disaster for humankind, although from the viewpoint of soil ecology dumps are not “waste land” . They offer a tremendous experimental field in which to study the colonization by animals and the development o f their commu

nities in this hostile environment. A new habitat for many new inhabitants is created as a result of spoilheap construction. A dump is a “land” lacking plants and animals, initially with a complete absence of soil. Then for a long period the “soil” that is present lacks stratification, has insufficient organic matter, few nutrients and an inadequate water content. Additionally, what makes post-industrial dumps an excellent polygon on which to test ecological hypotheses is the simplified and variable relationships between living components.

Furthermore, they are usually not under threat by prospectors.

The variability o f spoil heaps and the range o f environmental conditions established on post-industrial dumps result in a hetero

geneous environment for soil biota (D u n g e r , 1991; S c h n e id e r et al., 1995; Topp et al., 1992). Furthermore, different reclamation tech

nologies used on dumps also create varied conditions. So ecologists get an experiment with many differentiated environmental variables.

2.2. A chance to test successional theories

The origin and progress o f successional theory are based on our knowledge about primary and secondary succession o f plant com

munities, which is well documented (P u llin e n , 1986). Our knowledge o f succession o f soil organisms, particularly soil arthropods, asso

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ciated with basic biological parameter changes is significantly poorer

(Beckm ann & S c h r i e f e r , 1989; H u h ta et al., 1979; P i ż l , 1992; S c h e u

& S c h u lz , 1996; S t a r y , 1999; S t r e i t et al., 1985; S t r ü v e - K u s e n b e r g ,

1982; T a jo v s k Y , 1990). Only a small part o f research on succession deals with degraded post-industrial biotopes (e.g. C h r is tia n , 1993;

D u n g e r, 1968; H u tso n , 1980a; H u ts o n & L u f f , 1978; P a r r , 1978; V o g e l

& D u n g e r , 1991). The first worldwide review o f the succession o f soil animals on reclaimed areas was edited by M a j e r (1989a).

Several different pathways o f succession are described in the literature ( M a j e r , 1989b).

1. The facilitation pathway

2. The initial floristic composition pathway 3. The tolerance pathway

4. The inhibition pathway

5. The chronic disturbance pathway

These pathways are not mutually exclusive and elements o f more than one type may be recognized within a particular succession. The evidence for these pathways has been derived from the study o f plant communities ( C o n n e l l 8& S l a t y e r , 1977). Possibly they are not all applicable to heteromorphic succession because of, for instance, the mobility of animals ( M a je r , 1989b).

The facilitation model assumes that colonization by later suc

cessional species depends upon the early species preparing favourable environmental conditions (C le m e n ts , 1916; O dum , 1969).

The inhibition model assumes that irrespective of which species colonizes first, the primary colonist will adversely affect the colo

nization o f other species by using up resources (B e g o n et al., 1986).

The tolerance model (C o n n e l l & S l a t y e r , 1977) states that later invaders simply tolerate the conditions at the site and eventually outcompete the pioneers. Competition and mutualistic relationships are assumed to be o f minor importance in this model. E g l e r (1954) (initial floristic composition pathway) claimed that representa

tives o f all serai stages may be present very soon after disturbance, and that variations in life-history characteristics among the groups of species determine the succession. The chronic disturbance model

(H o r n , 1976) pertains to areas suffering chronic disturbance in small patches. It proposes that any species is likely to invade an opening and that locally a dominant species may be replaced by any num ber o f other species.

Testing the successional theory is one o f the m ost im portant trends in the study of ecosystems at present, and research on dumps can contribute a lot to this approach (S ta r Y , 1999). Dumps seem to be an excellent tool for the study o f biological communities in relation to environmental gradients. We may wish to know as much

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as possible about colonization by flora and fauna in order to direct or accelerate development o f modified ecosystems (Da v is, 1986).

Oribatid mites are especially useful in successional research; they are sensitive to small differences in environmental factors (only a small percentage of oribatids are ubiquitous) and their species pool is large (Verschoor & Kr e b s, 1995).

2.3. Oribatids on dumps

If th e s u c c e s s io n o f v e g e t a t io n o n p o s t - in d u s t r ia l w a s t e la n d s is to b e d e s c r ib e d th e n s o m e u n d e r s t a n d in g o f th e fa u n a a s s o c ia te d w ith p la n t s a n d o f d e c o m p o s it io n in th e s o il s e e m s im p e r a t iv e . T h e d e v e lo p m e n t o f t e r r e s t r ia l p o p u la t io n s in a r e a s w h e r e s u c h p o p u la tio n s d id n o t p r e v io u s ly e x is t o r h a d b e e n e lim in a t e d h a s b e e n s t u d ie d b y a n u m b e r o f w o r k e r s . H o w e v e r , o r ib a t id m ite c o m m u n itie s , w h ic h p o s s ib ly a ls o p la y a fu n d a m e n t a l r o le o n w a s te tip s , h a v e b e e n n e g le c t e d b y e c o lo g is ts . Dunger (1968) u n d e r t o o k w o r k o n th e d e v e lo p m e n t o f s o il fa u n a o n p o s t - in d u s t r ia l d u m p s . F o r th e fin d in g s o f D u n g e r a n d h is c o -w o r k e r s se e Beck (1994b). H is r e s e a rc h o n d u m p s in E a s t G e r m a n y s till c o n tin u e s . U n fo r t u n a t e ly , o r ib a t id s h a v e b e e n lo o k e d a t in d e t a il o n ly in s o m e o f D u n g e r ’s s t u d ie s . M o r e o v e r , h is in v e s t ig a t io n s d e a l w it h o n e ty p e o f p o s t - in d u s t r ia l d u m p , th e o p e n - c a s t m in in g a re a s . B a s e lin e d a t a c o n c e r n in g th e d e v e lo p m e n t o f m ite c o m m u n itie s o n d u m p s a re s till la c k in g . F e w s tu d ie s d e a l w ith o r ib a tid s in d e ta il (T a b le 2 a n d 3). Q u a lita tiv e d a t a o n o r ib a tid s c o u ld b e o n ly fo u n d in h a lf o f th e p a p e r s , e.g. Ba be n k o,

1980; Beckmann, 1988; Bielsk a, 1982a, b, 1995; Bielska & Pasze w sk a,

1995; Frouz e t a l., 2001; Hu b e r t, 2001; Lu x to n, 1982; Stebaeva &

An d rievskii, 1997; Żbiko w ska- Zd u n, 1988. I h a v e p e r s o n a lly s t u d ie d a b r o a d s e le c tio n o f d iffe r e n t d u m p s a n d h a v e p u b lis h e d r e s u lts in v a r io u s p a p e r s (Table 3). These a r tic le s c o v e r 10 d iffe r e n t ty p e s o f d u m p s , e.g. c o a l-m in e d u m p s , ir o n m e t a llu r g ic d u m p s , z in c d u m p s , d o lo m itic d u m p s , m in e s e d im e n ta tio n ta n k , d u m p s o f c h e m ic a l p la n ts, g a le n a - c a la m in e w a s t e la n d s a n d o th e r s . P r e c is e q u a n t it a t iv e a n d q u a lita tiv e d a t a o n o r ib a tid s w e r e a n a ly s e d in a ll o f th e s e s tu d ie s .

With regard to the acronym in the heading of this chapter - SOD - which means Save Our Dumps, it seems to be proven enough.

Some other scientists have put forward such proposals in the past.

It was not only the important value o f the vegetation that inspired them (To k a r s k a- Gu z ik & Ro s t a ń s k i, 2001; Wie r z b ic k a & Ro s t a ń s k i,

2002). Historical conditioning and widely understood touristic and

Colonization and development of oribatid mite communities (Acari: Oribatida) on post-industrial dumps

Colonization and development of oribatid mite communities

(Acari: Oribatida)

on post-industrial dumps

Piotr Skubała

Colonization and development of oribatid mite communities

(Acari: Oribatida) on post-industrial dumps

Honour species Pay attention to small organisms

Colonization and development of oribatid mite communities

(Acari: Oribatida)

on post-industrial dumps

Piotr Skubała

Contents

Preface - soil as the basis for civilization

Acknowledgements

Chapter i . Introduction

1.1. Oribatid mites - life hidden in the soil

1

1.2. Oribatids perform an important

“ecological service”

1.3. Contribution of oribatids to global biodiversity

1.4. Oribatids in biomonitoring studies

chapter 2 . Save our dumps (SOD) as an entity

2.1. Post-industrial dumps as a “unique”

experiment for ecologists

2.2. A chance to test successional theories

2.3. Oribatids on dumps

chapter 2 ^. Save our dumps (SOD) as an entity