**5. Spatial similarity pattern of oribatid mite communities (case study)**

Oribatid mites of the tropical regions had been almost completely unknown for science before 1958. With the general use of the Berlese-funnel, systematic collections started at that time, which resulted in the collection of hundreds or thousands of species. However, the majority of these samples are unprocessed till these days. Professor J. Balogh and his fellow-workers had to realize, that the description of all the species living there is an impossible task.

And even if the recognizable morphological kinds of the samples will be described with decades of monumental work by the practices of formal describing taxonomy and according to the rules of nomenclature, the biological and coenological information content of these would still remain hidden. However the material extracted with the Berlese-Tullgren device informs us not only about the presence and morphological diversity, but also about the species' abundance and dominance as well. Moreover it has become clear that this material together, as it was brought in front of us, contains a heap of such kind of information, which would be impossible to read from single species or from their constitutional characteristics. The samples collected this way are suitable for zoocoenological examinations. This observation led Humboldt to the recognition of "basic forms" (Grundformen) and later to the revelation of formations, which means the structure that can be found in plant associations without the exact knowledge of species. The emphasis here is on the "visibility" of the vegetation, because the recognition of biological communities began with the sight of the flora: vegetation is a "landscape element". Animal communities – apart from some exceptions – live hidden in the vegetation. But the Berlese-device concentrates and makes them visible.

In this present section of our examinations, the objective is to clarify the methodological possibilities of biological indication and the information content of the coenological data matrices by an appropriately chosen indication case study. We also set the aim to apply univariate indicators and to exploit the possibilities of multivariate coenological pattern analysis.

In this study series, there is need to introduce the main Oribatid sinusia of tropical areas as per climatic, vegetation and elevation zones. According to the holistic approach, we start from the whole and proceed towards the smaller parts. As a first step, we examined the similarities between the Oribatid sinusia of the Neotropical Region and the Notogaea. Stemming from the fact that the vertical stratification of Oribatid fauna follows the vegetation zones, and took samples for examination from 3 elevation zones:


190 Biodiversity Conservation and Utilization in a Diverse World

communities.

oribatid mites in the decomposition is the spreading of microbiota as they feed mainly on fungi and bacteria, and they are not in direct connection with the leaf litter input, accordingly (Maraun *et al*., 2001). This corresponds with the phenomenon observed by us, that the quantity of leaf litter may not have influenced the compositional changes of the

Our result that leaf litter and foerna substrates differ from moss was not interpreted by other literature yet, however, there are observations regarding other types of substrates. A common result is for example that the oribatid mite community living in the foliage of the trees differs significantly from the one living in the soil under the trees (e.g. Yoshida and Hijji, 2005; Karasawa and Hijii, 2008). Karasawa and Hijii (2004) showed that the substrate of oribatid mite communities living in the soil, in the foliage, on the bark of the tree trunks and on the remnants of algae accumulating on soil significantly differ from each other in seaside forests. In our study the community living in hypnum moss was simpler than the one living in the soil in the forest. Communities of moss and lichen are always relatively simple (Gjelstrup, 1979; Gjelstrup and Søchtig, 1979; Smrz, 1992; Smrz and Kocourková, 1999; Smrz, 2006). The three frequent genera found in moss turned up in the observations of others as well. *Zygoribatula exilis* is assumed to be a species living in moss (Gjelstrup, 1979), however, the *Zygoribatula* species found by us could be found on the forest ground as well. The genus *Tectocepheus* occurs everywhere from drier and more disturbed habitats to intact forests, accordingly it can also be found in moss in great quantities, especially because climatic fluctuation is larger in moss, what only some species can tolerate (Gjelstrup, 1979). A common epiphyte is *Eremaeus oblongus* (Smrz and Kocourková, 1999), in our study the genus

*Eremaeus* could be found only in moss (except for only one foerna sample).

oribatid mite communities living in other climatic zones show seasonal changes.

composition, changes and patterns are affected.

Therefore research showed that oribatid mite communities living in soil, leaf litter and hypnum moss, in Hungarian oak forests – similarly to those living in German beech forests – did not show seasonal changes. This result is important on the one hand because according to this, we are not bound to a season regarding sampling. However, besides that it would be important to detect the cause of the still occurring changes and patterns exactly. Furthermore it is unclear as well whether non-woody biocoenoses in the temperate zone or

Communities living in the soil differ from the oribatid mite community of moss living on tree trunks more significantly than from the community of the leaf litter. Oribatid mites in moss, especially the genus *Tectocepheus*, may be influenced by climatic factors to a large degree. In future it would be necessary to study oribatid mite communities of various microhabitats in order to detect exactly by which factors and to what degree their

**5. Spatial similarity pattern of oribatid mite communities (case study)** 

Oribatid mites of the tropical regions had been almost completely unknown for science before 1958. With the general use of the Berlese-funnel, systematic collections started at that time, which resulted in the collection of hundreds or thousands of species. However, the majority of In the study, we disregarded the mountain forest zone, which can be found between the zones of tropical rainforest and mossforest, because the determination of its borders is quite uncertain. To avoid transitions, Berlese-samples of tropical rainforest have been selected from 200 m elevation above sea level, close to the forest border. 2 ideal transects were set for representative sampling. The first one crosses Andes at Costa Rica, at the 10. degree of latitude, from coast to coast. The second one starts from Papua New-Guinea, from the valley of Fly River at the 4. degree of Southern latitude, and goes up to the 4000 m high ridge of Mt Wilhelm. As an amendment, samples were also taken near to the 23. degree of Southern

latitude in an additional transect, crossing Serra do Mar and Serra do Mantiguera. This transect has been set because 200-300 years ago there's been a belt of dense tropical rainforest – which even exceeded Amazonia in biodiversity – in the most densely populated area of Brazil, along the line marked by Sao Paulo and Rio de Janeiro. Almost 95 % of these forests have been devastated, but it could have been hoped that the rainforest spots reserved the original soil fauna – at least partially. 82 representative samples have been collected on 9 spots of the three transects. The spatial distribution of these samples and the abbreviation of individual sites can be seen in the next table:

Genus Lists of Oribatid Mites – A Unique Perspective of Climate Change Indication in Research 193

significant pattern-generating factor than geographical attribution. Rainforests, mossforests and paramos lying thousands of kilometers from each other are more similar than sites of other habitats at only a few kilometers distance. Following the results of these pattern analyses, exact examination of the observed phenomena with a regression model seems to

**Figure 3.** Similarity pattern of the examined sites in an NMDS ordination with the projection of the

We pointed out that dissimilarities of habitats caused even by their type and also by the continent they originate from can be recognized in the similarity pattern of genus lists of the examined habitats. But if we analyze the overall similarity pattern of all the individual samples, it is quite conspicuous that the type of habitat is a much more significant patterngenerating factor than the geographical location. Rainforest, mossforests and paramos located many thousand kilometers from each other are more similar to each other than sites

The most important result of our case study is that the list of Oribatid genera as a coenological indicator, primarily characterizes the present ecological effects of the habitat and its climatically determined type of vegetation; and represents the effect of zoocoenological past in a much lesser extent. Thus, it can be concluded that a meritable scientific faunagenetical analysis should not be based upon geohistorical, but climatological grounds. This is why the ecological indication based on Oribatid genus lists provides unique

hierarchical cluster analysis results, applying Euclidean distance.

of other kind of habitats in only a few kilometers away.

possibilities for the purpose of climate change research.

be practical.


#### **Table 2.**

The base table for our analyses was the 82 individual soil samples from the 9 examined habitat (3 habitat types of 3 areas) containing 111 Oribatid genera. For the various analyses, we created task-oriented assemblies from this base table.

Based on all of the individual samples, paired all samples with all other samples we created the similarity matrix of our data using multiple distance functions. In this current publication, Euclidean distance have been used.

Coenological similarity patterns can be analysed on multiple spatial scales (scale levels). We also analysed the similarity patterns of generic lists of different sites by NMDS with Euclidean distance, and hierarchical cluster-analysis. The two analytical results are shown projected onto each other (*Figure 3*). The multivariate similarity pattern of habitats' Oribatid community gave the expected picture. It can be stated that differences originating from the habitats and continents can also be recognized in the similarity patterns of the generic lists of the examined habitats. However, it can seem surprising that despite the vast geographical distances the pattern generating role of habitats does not disappear, it seems perhaps even more important. In accordance with the real ecological conditions, mossforest plays a transitional role between rainforest and paramo. However, mossforests are the most similar to each other and they are positioned in the middle of the similarity pattern, while the rest of the sites are separated radially. It is clearly visible that Brazilian sites are much more similar to each other than the Costa-Ricans.

Comparison of the sites can be fine-tuned if we also examine the similarity pattern of the individual soil samples considering every sample as different objects independent from the sites. We analysed this similarity pattern also by applying NMDS and Euclidean distance (*Figure 4*). Analysing the collective similarity pattern of every individual sample it can be stated that the groupings of elementary samples reflect their relations to the sites. This justifies the methodological decision by which sites are considered the basic objects of the examination. It can be stated furthermore, that habitat-type is unambiguously more significant pattern-generating factor than geographical attribution. Rainforests, mossforests and paramos lying thousands of kilometers from each other are more similar than sites of other habitats at only a few kilometers distance. Following the results of these pattern analyses, exact examination of the observed phenomena with a regression model seems to be practical.

192 Biodiversity Conservation and Utilization in a Diverse World

individual sites can be seen in the next table:

we created task-oriented assemblies from this base table.

publication, Euclidean distance have been used.

to each other than the Costa-Ricans.

**Table 2.**

latitude in an additional transect, crossing Serra do Mar and Serra do Mantiguera. This transect has been set because 200-300 years ago there's been a belt of dense tropical rainforest – which even exceeded Amazonia in biodiversity – in the most densely populated area of Brazil, along the line marked by Sao Paulo and Rio de Janeiro. Almost 95 % of these forests have been devastated, but it could have been hoped that the rainforest spots reserved the original soil fauna – at least partially. 82 representative samples have been collected on 9 spots of the three transects. The spatial distribution of these samples and the abbreviation of

Tropical rainforest Mossforest Paramo

Costa Rica RC: 10 MC: 10 PC: 10 30 Brazil RB: 7 MB: 6 PB: 10 23 New-Guinea RN: 10 MN: 9 PN: 10 29 27 25 30 82

The base table for our analyses was the 82 individual soil samples from the 9 examined habitat (3 habitat types of 3 areas) containing 111 Oribatid genera. For the various analyses,

Based on all of the individual samples, paired all samples with all other samples we created the similarity matrix of our data using multiple distance functions. In this current

Coenological similarity patterns can be analysed on multiple spatial scales (scale levels). We also analysed the similarity patterns of generic lists of different sites by NMDS with Euclidean distance, and hierarchical cluster-analysis. The two analytical results are shown projected onto each other (*Figure 3*). The multivariate similarity pattern of habitats' Oribatid community gave the expected picture. It can be stated that differences originating from the habitats and continents can also be recognized in the similarity patterns of the generic lists of the examined habitats. However, it can seem surprising that despite the vast geographical distances the pattern generating role of habitats does not disappear, it seems perhaps even more important. In accordance with the real ecological conditions, mossforest plays a transitional role between rainforest and paramo. However, mossforests are the most similar to each other and they are positioned in the middle of the similarity pattern, while the rest of the sites are separated radially. It is clearly visible that Brazilian sites are much more similar

Comparison of the sites can be fine-tuned if we also examine the similarity pattern of the individual soil samples considering every sample as different objects independent from the sites. We analysed this similarity pattern also by applying NMDS and Euclidean distance (*Figure 4*). Analysing the collective similarity pattern of every individual sample it can be stated that the groupings of elementary samples reflect their relations to the sites. This justifies the methodological decision by which sites are considered the basic objects of the examination. It can be stated furthermore, that habitat-type is unambiguously more

**Figure 3.** Similarity pattern of the examined sites in an NMDS ordination with the projection of the hierarchical cluster analysis results, applying Euclidean distance.

We pointed out that dissimilarities of habitats caused even by their type and also by the continent they originate from can be recognized in the similarity pattern of genus lists of the examined habitats. But if we analyze the overall similarity pattern of all the individual samples, it is quite conspicuous that the type of habitat is a much more significant patterngenerating factor than the geographical location. Rainforest, mossforests and paramos located many thousand kilometers from each other are more similar to each other than sites of other kind of habitats in only a few kilometers away.

The most important result of our case study is that the list of Oribatid genera as a coenological indicator, primarily characterizes the present ecological effects of the habitat and its climatically determined type of vegetation; and represents the effect of zoocoenological past in a much lesser extent. Thus, it can be concluded that a meritable scientific faunagenetical analysis should not be based upon geohistorical, but climatological grounds. This is why the ecological indication based on Oribatid genus lists provides unique possibilities for the purpose of climate change research.

Genus Lists of Oribatid Mites – A Unique Perspective of Climate Change Indication in Research 195

of Oribatid mites can be raised to genus- or family level, and this does not cause loss of data or sensitivity by multivariate methods. According to Osler (1999), it is possible that habitat preference is determined rather at family level. Furthermore it is also known that there is

At the same time, the standardization and quantification of current Oribatid-collection methods would be obviously necessary for the development of the bioindicational methodology in order to make the data from different authors comparable. The need for this is emphasized more and more in literature (Gulvik 2007). A criteria-system of classifying the collected individuals into genera (or into other optional morphological groups) can be considered as a part of the standardized method. The current identificational and taxonomical practice in oribatodology is almost completely unsuitable for the purposes of biological indication research. Till such methods are unavailable, case studies can be conducted only if the person doing field sample collection and taxonomical processing is identical, and furthermore, this is only true if the coenological matrices have been created

During our examinations (Balogh et al 2008), we pointed out that by advancing vertically upwards in the tropical high mountains (from rainforests towards the paramo vegetation), the average species number of genera present, and the extent of Holarctic and/or Antarctic relations of present genera grow, which can be well interpreted with the conception system of geographical analogy based on climatic similarities. This indication adequately supports former studies on the role of vegetational similarities and on genera as taxonomical units suitable for indication. Many authors (Andrew 2003, Melamud 2007 state that advancing upwards on the mountains, elevation above sea level and exposure have significant influence on the diversity of Oribatids. Besides, it is known that Oribatid diversity grows from the Boreal region towards warmer climates, but it does not grow further towards the

Furthermore, we pointed out that from the aspect of similarity of individual samples, among the examined habitat types the mossforest is the most homogeneous habitat, and paramo is the most heterogeneous. This latter phenomenon can be well interpreted if we consider the uniform moss cover prevalent in the mossforest, and the role of stable microclimate created by the moss cover. In the paramo, the observed proportion of species with low constancy level and also the heterogeneity of habitats (tussocks with different size)

The genus lists of oribatid mites provide a unique indication device for the research of

Provide uncomparably simple, fast and effective sample collection opportunity in case

are high, and the role of the resulting microclimatic variability is obvious.

of nearly any substrate type of nearly any habitat, all over the world.

climate change, based on their following characteristics:

strong relation between species-level and higher taxon level diversity indicators.

with the greatest care.

tropics (Maraun 2007).

**6. Conclusions** 

**Figure 4.** Similarity pattern of elementary samples in an NMDS ordination by applying Euclidean distance.

It became obvious that if we had chosen species and not genera as the basis of our examination, we would not get results that were interpretable from bioindicational aspect. This is because the majority of the described species would have appeared only as local specifica, and they do not provide any meritable information on similarity patterns – unless we are thinking in a very small grade of space. Species-level indication studies would be only rational in the case if we would deal only with cosmopolite species, but their number and detectability would not be sufficient for the majority of examinations for practical tasks. Thus, our important conclusion is that under current circumstances, the recommended taxonomical unit for indicational ecological studies can only be the genus. This statement is also important because many authors (primarily species describing taxonomists) consider a serious problem of indicational research that a number of researchers classify individuals only to genus level (Gulvik 2007). However, based on the work of Caruso (2006), we know that human contamination, intervention and disturbance can be better detected if we examine larger taxonomical units. It is also clear that rapid changes can not be detected on species-, but on community level. From the same work, it is also known that the species data of Oribatid mites can be raised to genus- or family level, and this does not cause loss of data or sensitivity by multivariate methods. According to Osler (1999), it is possible that habitat preference is determined rather at family level. Furthermore it is also known that there is strong relation between species-level and higher taxon level diversity indicators.

At the same time, the standardization and quantification of current Oribatid-collection methods would be obviously necessary for the development of the bioindicational methodology in order to make the data from different authors comparable. The need for this is emphasized more and more in literature (Gulvik 2007). A criteria-system of classifying the collected individuals into genera (or into other optional morphological groups) can be considered as a part of the standardized method. The current identificational and taxonomical practice in oribatodology is almost completely unsuitable for the purposes of biological indication research. Till such methods are unavailable, case studies can be conducted only if the person doing field sample collection and taxonomical processing is identical, and furthermore, this is only true if the coenological matrices have been created with the greatest care.

During our examinations (Balogh et al 2008), we pointed out that by advancing vertically upwards in the tropical high mountains (from rainforests towards the paramo vegetation), the average species number of genera present, and the extent of Holarctic and/or Antarctic relations of present genera grow, which can be well interpreted with the conception system of geographical analogy based on climatic similarities. This indication adequately supports former studies on the role of vegetational similarities and on genera as taxonomical units suitable for indication. Many authors (Andrew 2003, Melamud 2007 state that advancing upwards on the mountains, elevation above sea level and exposure have significant influence on the diversity of Oribatids. Besides, it is known that Oribatid diversity grows from the Boreal region towards warmer climates, but it does not grow further towards the tropics (Maraun 2007).

Furthermore, we pointed out that from the aspect of similarity of individual samples, among the examined habitat types the mossforest is the most homogeneous habitat, and paramo is the most heterogeneous. This latter phenomenon can be well interpreted if we consider the uniform moss cover prevalent in the mossforest, and the role of stable microclimate created by the moss cover. In the paramo, the observed proportion of species with low constancy level and also the heterogeneity of habitats (tussocks with different size) are high, and the role of the resulting microclimatic variability is obvious.

#### **6. Conclusions**

194 Biodiversity Conservation and Utilization in a Diverse World

distance.

**Figure 4.** Similarity pattern of elementary samples in an NMDS ordination by applying Euclidean

It became obvious that if we had chosen species and not genera as the basis of our examination, we would not get results that were interpretable from bioindicational aspect. This is because the majority of the described species would have appeared only as local specifica, and they do not provide any meritable information on similarity patterns – unless we are thinking in a very small grade of space. Species-level indication studies would be only rational in the case if we would deal only with cosmopolite species, but their number and detectability would not be sufficient for the majority of examinations for practical tasks. Thus, our important conclusion is that under current circumstances, the recommended taxonomical unit for indicational ecological studies can only be the genus. This statement is also important because many authors (primarily species describing taxonomists) consider a serious problem of indicational research that a number of researchers classify individuals only to genus level (Gulvik 2007). However, based on the work of Caruso (2006), we know that human contamination, intervention and disturbance can be better detected if we examine larger taxonomical units. It is also clear that rapid changes can not be detected on species-, but on community level. From the same work, it is also known that the species data

The genus lists of oribatid mites provide a unique indication device for the research of climate change, based on their following characteristics:

 Provide uncomparably simple, fast and effective sample collection opportunity in case of nearly any substrate type of nearly any habitat, all over the world.


Genus Lists of Oribatid Mites – A Unique Perspective of Climate Change Indication in Research 197

*Department of Mathematics and Informatics, Corvinus University of Budapest,* 

*"Adaptation to Climate Change" Research Group of the Hungarian Academy of Sciences,* 

We have to thank Prof. János Podani for his kind methodological help, Dr. Péter Balogh for his valuable professional help with the research of oribatid mites and for making the manuscripts from the Balogh-legacy available. We thank the "Adaptation to Climate Change" Research Group of the Hungarian Academy of Sciences and particularly the late Zsolt Harnos, who ensured the professional prerequisites of the research. Our research was supported by the Research Assistant Fellowship Support (Corvinus University of Budapest), "ALÖKI" Applied Ecological Research and Forensic Institute Ltd., the Landscape Architecture and Landscape Ecology PhD School of the Corvinus University of Budapest and the "Bolyai János" Research Fellowship (Hungarian Academy of Sciences, Council of Doctors). This work was supported by the research project of the National Development

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[2] Andres, P. (1999) Ecological risks of the use of sewage sludge as fertilizer in soil restoration effects on the soil microarthropod populations. – Land Degradation &

[3] Andres, P. (1999) Ecological risks of the use of sewage sludge as fertilizer in soil restoration effects on the soil microarthropod populations. Land Degradation &

[4] Andrew, N.R., Rodgerson, L., Dunlop, M. (2003) Variation in invertebrate-bryophyte community structure at different spatial scales along altitudinal gradients. – Journal of

[5] Arroyo, J., Iturrondobeitia, J.C. (2006) Differences in the diversity of oribatid mite communities in forests and agrosystems lands. – European Journal of Soil Biology 42:

[6] Askidis, M.D., Stamou, G.P. (1991) Spatial and temporal patterns of an oribatid mite community in an evergreen-sclerophyllous formation (Hortiatis, Greece). –

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L. Hufnagel

*Budapest, Hungary* 

**7. References** 

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**Acknowledgement** 

*Faculty of Horticulture, Budapest, Hungary* 

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Extremely high information content of the distance functions of genus lists.

Category orders as results of the analyses mostly met our expectations, first of all in case of genus lists, i.e. larger distances between genus lists correspond to larger spatial scales. However, time difference within a year could not be pointed out to a considerable extent either on genus or on family level. The most important differences in the orders are the followings:


The analysis on family level differs from that of genus level that family lists of the zoogeographic kingdoms resemble each other as much as family lists originating from the same types of substrate.

Thus, compiled a reference list with the help of which it can be expressed to what spatial distances the similarity – shown with distance functions – of the genus or family lists of two oribatid mite communities originating from samples of unknown quality corresponds. Disturbed and transformed habitats can be compared with the help of oribatid mite communities based on an existing reference list.
