Oribatid mites in a

changing world

Arguitxu de la Riva Caballero

Dissertation for the degree of philosophiae doctor (PhD) University of Bergen, Norway 2011

Front drawing: Suctobelbella longicuspis by Arguitxu de la Riva-Caballero To Adriana and Matteo

Supervisors:

Main supervisor: Torstein Solhøy

Co-supervisor: Hilary H. Birks

Co-supervisor: Lita G. Jensen

Contents

Abstract ...... 3

Acknowledgments ...... 5

List of papers ...... 7

Declaration ...... 9

Synthesis Oribatid mites in a changing world

Introduction ...... 11 Archives and Proxies ...... 11 What are oribatid mites? ...... 13 Oribatids in palaeoecology ...... 15

Aims of this thesis ...... 18

Study Areas ...... 19

Methods ...... 20 Field sampling ...... 21 Laboratory methods ...... 22 Data analysis ...... 23

Outline of main results in the four papers ...... 24 Background fauna characterisation ...... 24 Paper I - Oribatid mite communities in western Norway ...... 24 Oribatid mite representation in lake sediments and best coring point ...... 26 Paper II - Oribatid mite assemblages across the tree-line in western Norway and their representation in lake sediments ...... 26 Palaeolimnological and archaeological reconstructions using oribatid mites ...... 27 Paper III: Holocene environmental and climate history of Trettetjørn, a low-alpine lake in western Norway, based on subfossil pollen, diatoms, oribatid mites, and plant macrofossils ...... 27 Paper IV: Neolithic impact on local vegetation at Kvitevoll, Halsnøy Island, western Norway reconstructed from oribatid mites and pollen analysis ...... 28

Conclusions, problems, and further work ...... 31

Acknowledgements ...... 32

References ...... 32

Oribatid Mites in a Changing World

De la Riva-Caballero, A. 2011. Oribatid Mites in a Changing World. PhD thesis, Bergen Museum and Faculty of Mathematics and Natural Sciences, University of Bergen, Norway

new records for Norway. The species Chamobates borealis, Oppiella nova, Abstract Moritzoppia neerlandica, and The main scope of this thesis is to Rhinoppia subpectinata characterised illustrate the validity of oribatid mites as the oribatid communities of Betula, tools for palaeoecological mixed, and Picea forest subsets. reconstructions. Palaeoecology studies Deciduous forest oribatid communities the responses of past organisms to past were characterised by Achipteria environmental changes. This can be coleoptrata, Acrotritria ardua, accomplished through the use of Ceratozetes gracilis, and Oribatella biological proxies, which are indicators calcarata. Hemileius initialis, of past conditions. The search for Nanhermannia dorsalis, C. borealis, additional means of distinguishing Tectocepheus velatus, and Atropacarus climate change has only recently led to striculus characterised wet habitats. In the use of other commonly found water-logged habitats, Limnozetes biological proxies such as tiny oribatid ciliatus, Mucronothrus nasalis, and mites known as moss-mites. Oribatid Trimalaconothrus glaber dominated. mites are among the most numerous Carabodes labyrinthicus, C. biological remains in anoxic sediments, marginatus, Melanozetes mollicomus, yet until now oribatids have not been and T. velatus characterised the oribatid widely used due to the uncertainties community of the lichen and moss about their present distribution and the subset. The tree-line ecotone was lack of expertise to identify them to dominated by the euryceous species H. species level. This thesis contains four initialis, T. velatus, and Oribatula papers which provide evidence about tibialis. This study represents a how oribatid mites, when they are thorough survey of oribatid properly identified to species level and communities in western Norway, and their background distribution is the insights it gives are an important adequately known, can give useful tool for habitat reconstructions, as they additional and supporting information provide the background knowledge for reconstructing past habitat and about modern oribatid fauna needed to environmental conditions. identify the type of past plant Paper I studied oribatid community and past environments preferences and ecology in different represented in Quaternary sediments. habitats, mainly forested, in western Paper II studied the oribatid Norway. One hundred and ninety two communities at the tree-line in western species were found of which 64 were Norway and compared them with the

U oribatid fossil assemblages found in Mires began to replace forested areas on Lake Trettetjørn. The modern oribatid the landscape as a more oceanic climate assemblage provided a guide to the began to prevail. All proxies indicated reliability of the fossil assemblages to increasingly intensive human land-use reconstruct ecological and as the Upsete settlement grew to environmental changes and, in addition, accommodate the construction of the to find the most favourable coring point Bergen-Oslo railway. within the small lake. Results showed In Paper IV, oribatid mites and that the core retrieved from the middle pollen were used to reconstruct the local of Lake Trettetjørn basin represented habitat at an archaeological excavation. the oribatid fauna from the catchment The study aimed to identify the start of area. Aquatic oribatids were the best cereal cultivation at Kvitevoll farm, on group represented in the lake sediments, Halsnøy island, western Norway. The followed by oribatids from the habitats high number of oribatid remains adjacent to the lake. This constitutes identified to species level and the close good evidence that oribatids are match to pollen stratigraphy led to a excellent indicators of local habitats. detailed palaeoenvironmental Comparison of the oribatid fauna found reconstruction. Oribatids and pollen in the lake traps with the oribatid indicated the development of a moist assemblages from paper III illustrated forest followed by vegetation openings the importance of identifying the mites and mire expansion over the site. At the to species level, as this increased the top of the sequence, the presence of ecological indicator value and, oribatids such as Tectocepheus velatus therefore, the reliability of the and the increase in members of the palaeoreconstructions. family Oppiidae indicated a higher In Paper III, sub-fossil oribatid degree of disturbance, probably from mites, pollen, plant macrofossils, and grazing. Pollen of Cerealia indicated the diatoms from a lake sediment core from start of cultivation around the same western Norway were studied. This time. multi-proxy study attempted to reconstruct tree-line fluctuations and their impact on Lake Trettetjørn’s environment. Evidence from pollen, plant macrofossils, and oribatids complemented and corroborated each other in the reconstruction of the vegetational development. A semi-open grassland developed into forest.

V Acknowledgments It is difficult to thank all the people who have helped me with this thesis. I am certain I will forget mentioning some, but please do not be offended. I am grateful to all of you.

Torstein Solhøy deserves a special thanks, not only because he has been my supervisor, but also because he was the one who introduced me to oribatid mites and gave me the chance to fall in love with them during that first summer of 2001. He is an extraordinary person in many ways. He knows when he is really needed, and then he is there. John Birks also especially deserves to be thanked. He has helped me through the process, and not only with the numerical analyses! Hilary Birks has contributed, especially in the final steps, to make this thesis a reality. She is the perfectionist I needed at my side to show me how to do things properly. Lita G. Jensen, my supervisor at Bergen Museum, has always been there. Thank you all!!

In addition, I would like to thank Jorunn Larsen, Anne E. Bjune, and Lene S. Halvorsen, all fellow authors in one of the papers comprised in this thesis, for their encouragement and support through the process. Working with you has been wonderful.

My colleagues at the Department of Biology: Marianne P. Heggen, Ingelinn Aarnes, Kristine Fjordheim, Linn C. Kruger, Bjørn Arild Hatteland, and Arild Breistøl. It has been great to share so many coffee breaks, conversations about theses and other matters, parties, etc., with you. Your support through the process has been especially invaluable.

Marianne P. Heggen through the years became not only the best colleague one could dream of, but also a good friend. She has been encouraging, enlightening, patient, funny and serious, helping me any time I needed it. Thanks Marianne! I look forward to the next time we share an office!

I am in debt to Anna Seniczak, with whom I shared my first summer working with mites at Acarilaben. She is an exceptional teacher and was able to make the task of learning to identify those wonderful but tricky creatures funny, challenging, and fascinating. Definitively, I could not have started my PhD thesis without that first summer.

Luis S. Subías has been an invaluable help. He was able to pass on his love for oribatid mites, especially for oppiids, and I suddenly discovered that I was completely fascinated by those golden little creatures. He has shared knowledge about not only mites but also life in our coffee breaks and meetings whenever I visited Madrid, and he patiently listened to my thesis or everyday life problems and tried to help me out. In other words, he became not only a guide in the oribatid world but also a friend. Thank you Luis!!

At Bergen Museum I would especially like to thank Ingrid Herø, who is now retired. She has been the most effective and helpful woman in administration that I have ever met. Moreover, she was always in a good mood and smiling regardless of the “strange” situations I came to explain in my funny Norwegian.

W The last year and a half I have been living in Milan. I do not have working colleagues there, but I have a wonderful mother-in-law, mamma Vale, who has made my everyday work a pleasure. She has not only invited me into her house, but also cooked for me! She has been my coffee break “colleague” and has listened patiently to all of my achievements and complaints. In addition, she has always helped at home not only with cooking and cleaning but also, and most importantly, by playing with Adriana. Thank you very much Vale!

Thank you to Hanneke, Usman, Danyal, and Ishaq, unconditional friends despite the distance. They are our Norwegian family, and they always made our weekends something to look forward to, always supported me, and have offered not only their house whenever I was in Bergen during this last year and a half, but also made me feel like one of their family.

Susi, thank you very much, for being more than a friend, no matter the distance, for your enthusiasm, your good mood, the coffees, the laughs, dinners, lunches, vacations, and your help when needed.

My family, mamá, papá, Pilar, Fernando, María, Isabel, and Txema, though not always close physically, are always close to me. Their support has been crucial through this process. They have always understood me and guided me. Thank you very much! You are probably the strangest but the best family in the world. I love you all.

And last but not least I am extremely grateful to my husband, Matteo, and my daughter, Adriana. Matteo is, without a doubt, my better half. He understands me and supports me unconditionally. Adriana is still too young to understand what mummy was doing but her laughs, her first words, her first steps, her hugs, and her kisses made this last year much easier. I do not think that I will ever have words to tell you how grateful I am and how much I love you both.

X List of papers This thesis is based on four individual papers, which will be referred to by their Roman numbers in the synthesis.

Paper I: de la Riva-Caballero, A. Oribatid mite communities in western Norway. To be submitted to Norwegian Journal of Entomology

Paper II: de la Riva-Caballero, A., H. J. B. Birks, A. E. Bjune, H. H. Birks, and T. Solhøy. Oribatid mites assemblages across the tree-line in western Norway and their representation in lake sediments. Journal of Paleolimnology 44: 361-374

Paper III: Larsen, J., A. E. Bjune, and A. de la Riva-Caballero. 2006 Holocene Environmental and Climate History of Trettetjørn, a Low-alpine Lake in Western Norway, Based on Subfossil Pollen, Diatoms, Oribatid Mites, and Plant Macrofossils. Arctic, Antarctic and Alpine Research 38:571-583.

Paper IV: de la Riva-Caballero, A. and L. S. Halvorsen. Neolithic impact on local vegetation at Kvitevoll, Halsnøy Island, western Norway reconstructed from oribatid mites and pollen analysis. To be submitted to Journal of Archaeological Science.

All reprints have been made with the permission of the editors.

Journal of Paleolimnology is published under Springer (http://s100.copyright.com/CustomerAdmin/PLF.jsp?lID=2011090_1314958885973)

Arctic, Antarctic, and alpine research is published under University of Colorado

Order Detail ID: 56131228 Arctic, antarctic, and alpine research by University of Colorado, Boulder.Institute of Arctic and Alpine Research Copyright 2006 Reproduced with permission of INSTITUTION OF ARCTIC & ALPINE RESEARCH, UNIVERSITY OF COLORADO - BIOONE in the format Dissertation via Copyright Clearance Center.

Y Z Declaration Papers II to IV are co-authored, and the nature of the contributions by the different authors are listed below.

Paper II de la Riva-Caballero, A., H. J. B. Birks, A. E. Bjune, H. H. Birks, and T. Solhøy. Oribatid mites assemblages across the tree-line in western Norway and their representation in lake sediments. Journal of Paleolimnology 44: 361-374 A. de la Riva-Caballero: Acari identification and analyses, numerical analyses, writing. HJB. Birks: Numerical analyses, editing. AE. Bjune: Experimental design, field sampling, editing. HH. Birks: Project management, experimental design, field sampling, editing. T. Solhøy: Acari identification, editing.

Paper III Larsen, J., A. E. Bjune, and A. de la Riva-Caballero. 2006 Holocene Environmental and Climate History of Trettetjørn, a Low-alpine Lake in Western Norway, Based on Subfossil Pollen, Diatoms, Oribatid Mites, and Plant Macrofossils. Arctic, Antarctic and Alpine Research 38:571-583. J. Larsen: Project management, field sampling, diatom analysis, writing AE. Bjune: Field sampling, pollen and plant macrofossils analyses, writing. A. de la Riva-Caballero: Oribatid mites analysis, writing.

Paper IV de la Riva-Caballero, A. and L. S. Halvorsen. Neolithic impact on local vegetation at Kvitevoll, Halsnøy Island, western Norway reconstructed from oribatid mites and pollen analysis. Submitted to Journal of Archaeological Science. A. de la Riva-Caballero: Field sampling, oribatid mite analysis, numerical analysis, writing. LS. Halvorsen: Field sampling, pollen analysis, writing

[ SR Oribatid Mites in a Changing World

De la Riva-Caballero, A. 2011. Oribatid Mites in a Changing World. PhD thesis, Bergen Museum and Faculty of Mathematics and Natural Sciences, University of Bergen, Norway

and other anoxic locations, which become archives of past environments. Introduction One can reconstruct the development of the environment in and around the lake Archives and Proxies or mire using continuous sequences of Palaeoecology is the study of responses the sediments and the biological proxies of past organisms to past environments. contained in them. The use of only one This includes the study of past climatic proxy in a palaeoenvironmental changes which are reflected in the reconstruction will always be less change of abundance and occurrence of satisfactory than reconstructions in different biological proxy indicators which several proxies are combined, as through time. Palaeoecological the strengths and weaknesses of the reconstructions enable the comparison individual proxies are then of past and present ecosystems, which counterbalanced (Birks and Wright can reveal possible causes and 2000, Birks and Birks 2006). mechanisms of biological change over In palaeoecology one can time (Birks and Birks 1980, Berglund reconstruct the organism assemblages as 1986, Willis and Birks 2006, Douglas presence or absence data, populations as 2007, Gamble 2007). numbers of individuals of the taxa, and A biological proxy is a communities as a combination of the biological indicator of past assemblages and the individual environmental conditions (Smol et al. abundances. This thesis concentrates on 2001, Douglas 2007). Among a wide the reconstruction of communities, variety of biological proxies, the most which allows one to draw inferences widely used are pollen, plant about the environment and habitat of macrofossils, chironomids (non-biting past ecosystems (Birks and Birks 1980). midges) (Fig 1), and diatoms (Douglas Each species in a community has an 2007). The search for additional means optimum and a niche within its range of of distinguishing climate change has environmental tolerance. There is also a only recently lead to the use of other community niche, which is the range of also commonly found biological proxies environmental factors common to all such as the tiny oribatid mites known as the species that comprise the moss-mites (Solhøy and Solhøy 2000, community. Identifying a complete past Erickson and Platt 2007). Remains of community of oribatid mites is almost animals and plants are deposited and impossible because life assemblages are preserved in sediments of lakes, mires modified by taphonomic processes.

SS These are the processes the deceased analysis of fossil data to identify mites go through from the moment they possible gradients that reflect die until they are finally deposited into environmental change; and (3) analysis the sediments. They include of the abiotic component of the transportation from the source to the sediment (Birks and Birks 1980, Birks lake or mire, predation, preservation, 2003). In this study, the first and second and possible redeposition (Birks and approaches are used. These approaches Birks 1980, Erickson 1988, Erickson assume that little or no change in the and Platt 2007). There are three main ecological requirements of the species approaches used to reconstruct past found in Holocene sediments has habitats: (1) modern analogue, which occurred since the deposition of the compares modern communities with the fossil assemblage. fossil assemblage; (2) numerical

Figure 1. Some examples of biological proxies. 1. Oribatid mites. a. Tectocepheus velatus. b. Dometorina plantivaga. c. Cosmochthonius lanatus (Picture 1 A. De la Riva-Caballero, pictures 2 and 3. J. Arroyo). 2. Chironomids a. Chaetocladius cf. piger b. Limnophyes sp. (pictures G. Velle) 3. Pollen a. Pinus. b. Valeriana sambucifolia. (pictures J. Berge).

ST Oribatid mites are tiny, flightless What are oribatid mites? arthropods belonging to the subphylum The word oribatid comes from Chelicerata with sizes ranging from the Greek words oreos – mountain and 0.13 to 1.2 mm (Walter and Proctor batein – travel (i.e. those that travel 1999, Norton and Behan-Pelletier around in the mountains). They are also 2009). As chelicerates they have two known as moss or box mites. Oribatids pairs of chelicerae and palpi in the oral are among the most abundant region, a body divided in two parts, inhabitants of the soil system (Travé et prodorsum and notogaster (Fig. 2), and al. 1996, Walter and Proctor 1999). three pairs of legs in the larval stages There are ca. 11000 known species, and and four in the nymphal and adult their local diversity can be high, for stages (Fig. 3). Except for some example with up to 150 species may primitive families, oribatids have a coexist in a small area of temperate hard, darkly pigmented (chitin) forest floor (Hansen and Colleman exoskeleton as adults but are usually 1998, Solhøy and Solhøy 2000, Norton soft-bodied as immatures (Norton and 2007, Schatz and Behan-Pelletier 2008, Behan-Pelletier 2009). Norton and Behan-Pelletier 2009).

Figure 2. Mite morphology: prodorsum, notogaster, and legs are indicated on a picture of Xenillus tegeocranus (left picture, by Stefan Hewig – X. tegeocranus average length 720-1100µm, Weigmann 2006), chelicerae and palpi are indicated on a SEM picture of an Hydrozetes sp. juvenile (right picture, by Anna Seniczak); blue line on the right indicates the measure scale for Hydrozetes sp. (20µm).

Among oribatids, there are both usually takes one or two years in sexual and asexual species. Generally, temperate to boreal regions (Norton and ancestral groups have asexual Behan-Pelletier 2009), but it can take up reproduction through thelytoky to seven years in extreme habitats (production of females, from females, (Søvik et al. 2003). Their full keeping the diploid condition), while developmental sequence is typical of most derivative groups tend to show Chelicerata, with egg, pre-larva sexual reproduction instead (Travé et al. (immovable), larva, three nymphal 1996, Walter and Proctor 1999, Norton stages (proto-nymph, deuto-nymph and and Behan-Pelletier 2009). These trito-nymph) and the adult, which is the minute creatures have a life cycle that reproductive stage. They do not have SU parental care, but they carefully choose (Norton 1990), which makes them ideal where to lay their eggs (Norton 1990). palaeo-indicators of such habitats. Oribatids are mostly particle- Moss mites are generally not feeding detritivores (i.e. obtain nutrients considered to be phoretic organisms; from decomposing organic matter) and they do not use other organisms for mycophages (i.e. feed on members of transportation mean. Both Berthet the fungus kingdom) that live in soil (1964) and Hammer (1965) state that and litter (Luxton 1972, Walter and oribatids do not show long dispersal, in Proctor 1999, Norton 2007). They play part because of their flightless condition an active role in ecosystems, for and gregarious nature. However, several example in the decomposition of investigators have shown that dispersal organic matter, nutrient cycling, and over more than the few meters that soil formation. In addition, they are oribatids can cover in their lifespans is good indicators of moisture and pH in possible through some kind of phoresy both terrestrial and aquatic ecosystems on insects (Norton 1980, Coulson (Behan-Pelletier 1999). 2009), mammals (Miko and Stanko Oribatid mite community 1991), and even on birds (Lebedeva and composition changes with vegetation Krivolutsky 2003, Lebedeva and composition. The composition of the Lebedev 2008). However, none of these mite community is controlled partly by studies discusses whether this is food availability, soil type, and moisture common within oribatid mites. To date, (Walter and Proctor 1999, Solhøy and only one family (Mesoplophora) has Solhøy 2000). Certain oribatids have been shown to have body modifications rather narrow niches and are obligatory that enable attachment to other or facultative inhabitants of substrates organisms (Norton 1980). like trees, rocks, lichens, and mosses

Figure 3. Adult (left) and larva (right) of Hydrozetes sp. (pictures by Anna Seniczak). Blue lines on the right of both pictures indicate a scale of 200µm for the adult (left) and 80µm for the larva (right).

SV Oligocene and the Pleistocene. Oribatids in palaeoecology Labandeira et al. (1997) gives an Brief history - Oribatid mites are overview of pre-Pleistocene records. known from the early Ordovician Krivolutsky and Drouk (1986) state that (Bernini et al. 2001) and the Devonian the major trends in the evolution of (Norton et al. 1988), with some other oribatids were established before the evolved genera dating back to the Jurassic, and they suggest that oribatid Jurassic (Krivolutsky and Drouk 1986) mites have very low morphogenetic (Fig. 4). rates. I, therefore, assume that oribatid According to Hammer (1965) fossils from the Holocene do not differ and Hammer and Wallwork (1979), in their morphology and behaviour from most extant species had already arisen those of today. sometime between the Eocene –

Figure 5. Oribatid subfossils. a. general view of the washed out oribatid fragments from a sample b. Liacarus keretinus (average measure for L. coracinus 650-1100µm (Weigmann 2006)). c. Oribatula tibialis (average measure 410-530µm (Weigmann 2006)) (pictures A. de la Riva-Caballero).

Their flightless condition, hard did not interpret them ecologically. One chitinous exoskeleton (well preserved in of the main problems they faced was anoxic sediments) and narrow niches that the sediments/layers were not make oribatid mites very suitable radiocarbon dated, as the technique was indicators of local past habitats not developed until the 1950s by Libby (Erickson 1988, Drouk 1997, Solhøy (Gosse 2007). During the second half of and Solhøy 2000, Solhøy 2001, the 20th century, the use of fossil Erickson and Platt 2007). However, the oribatids started to expand. In Europe, use of fossil oribatid mites as palaeo- Knülle (1957a), Karppinen et al. (1979), indicators is a relatively new discipline and Golosova et al. (1985) presented (Erickson 1988, Solhøy and Solhøy records of fossil oribatids from northern 2000, Solhøy 2001, Erickson and Platt Europe, Greenland, and northern 2007, Elias 2010). Nordenskiöld (1901) Siberia. Schelvis (1989) was the first was the first author who identified fossil researcher to use fossil oribatid mites as oribatid mites in his work. After him, indicators of past ecosystems, many others in the first half of the 20th interpreting the fossil oribatid century mentioned fossil oribatid mites assemblages found in archaeological and listed them in their works, but they sediments from an ecological point of

SW

view and even implementing a semi- water run-off and streams or by falling quantitative method (Schelvis 1989, from the nearby vegetation can also be Schelvis and van Geel 1989, Schelvis found. Taphonomic processes filter the 1990a, Schelvis 1990b, Schelvis 1992, species and number of oribatids that are Schelvis 1997a, Schelvis 1997b). The finally incorporated into the sediments. most recent works published using

Million Years Eras Mayor Divisions Important Events fossil oribatid mites from lake Ago sediments are from Solhøy and Solhøy 0.01… Holocene

(2000), Hodgson and Convey (2005), 1.64 Pleistocene ICE AGES Quaternary

Larsen et al. (2006), and Heggen et al. 5.2 Pliocene

(2010). The work from Solhøy and 23.3 Miocene

Mesostigmatans in Solhøy (2000) is outstanding because of 35.4 Oligocene amber the high resolution of the sediment core, Tertiary 56.5 Eocene Fossil ticks abundant radiocarbon dates, 65 Palaeocene End of dinosaurs identification to species level of all the 145.6 Cretaceous adults found, and good documentation KrivolutskyEvolved fossil and oribatid Druk genera 208 Jurassic 1986– Krivolutsky and Drouk 1986 of the species’ present-day distributions Mesozoic 245 Triassic Mass extintion and habitat requirements (Solhøy 2001). 290 Permian Recently, oribatids have been used in 362.5 Carboniferous palaeolimnology and in archaeology, Fossil endeostigmatans and oribatids – Norton

408.5Upper Paleozoic Devonian for example by Schatz et al. (2002), et al 1988, Norton et al Arroyo et al. (2007), Chepstow-Lusty et 1989 438.1 Silurian al. (2007), and Wild et al. (2007). In Fossil oribatid mites from 505 Ordovician Bernini et al 2001 North America, Erickson (Erickson Sweden – Bernini et al. 2001 First potential 545 Cambrian 1988, Erickson 1997, Erickson et al. Lower Palaeozoic chelicerate 2003, Erickson and Solod 2008) was the 2500 Proterozoic 3500 Archaean first researcher to use fossil oribatids Precambrian retrieved from lake sediments to infer climatic/habitat changes. Figure 4. Acarocentric point of view of the history of life. Modified after Walter and Sampling sites and identification- The Proctor 1999. Only references related to oribatid use of oribatid mites as indicators of mites are indicated. past habitats relies on their good preservation in anoxic sediments (Fig. Identification to species level 5). Thus, sediments from lakes and and a good knowledge of their present- mires and other waterlogged day ecology is of extreme importance environments are usually optimal. for obtaining a reliable reconstruction of Figure 6 illustrates the process of the past environment. This knowledge oribatid deposition in a lake. The allows oribatid species to be grouped oribatids most commonly found in lake according to their present-day ecology. sediments are aquatic and wetland As with plant macrofossils (Birks species living on aquatic plants and 2001), oribatids are usually used to other vegetation on the lake shore. reconstruct local habitat changes as However, terrestrial oribatids that their occurrence and distribution are reached the lake mainly through surface highly dependent on local substrate SX type. In addition, oribatid remains are Erickson and Solod 2008). Oribatid normally not transported far from their mite analysis requires patience and area of origin (Fig. 6) as, in contrast to good knowledge of not only mite pollen, they are not wind transported. identification from fragments but also In general, knowledge about the the ecology of the species found broad-scale geographical distribution of (Schelvis 1997a, Solhøy 2001, Erickson oribatid mites is imperfect (Drouk 1997, and Platt 2007). It is far more Behan-Pelletier 1999, Weigmann 2006). informative to find a set of species This limits detailed reconstruction of indicating the same type of vegetation past climate using only fossil oribatid than to find many individuals of the assemblages. Oribatid distribution same species (Schelvis 1997a). In some patterns do not correspond with instances there are very few remains of biogeographic boundaries and are not mites found in the sediments or too limited by known climatic factors, and many representatives of generalist oribatid’s species composition’s within- species, restricting the interpretation of zone variability is as high as its among- the past environment and vegetation zone variability (Drouk 1997). These (Wild et al. 2007). It is, therefore, factors limit the use of oribatid mites to optimal to combine the study of fossil qualitative studies. Nonetheless, mites with other proxies that will aid in Schelvis (1989, 1990b) and Erickson obtaining a comprehensive and Platt (2007) have demonstrated the understanding of the palaeo- application of oribatid mites in semi- environment (Birks and Birks 2006). quantitative methods (Schelvis 1990a,

Figure 6. Processes leading to the deposition of terrestrial and aquatic oribatid mites in lake sediments.

SY oribatid fossil assemblages found in the Holocene sediments of Lake Trettetjørn, Aims of this thesis a small lake situated at the present tree- My work deals with extant and past line. The modern oribatid fauna from oribatid communities. The study of different vegetation zones along an extant oribatid communities improves elevational gradient across the tree-line the knowledge on the ecology and was characterised as a basis for biology of oribatid mites, which is of comparison with the fossil oribatid extreme importance in order to infer assemblages found in sediments from a habitat changes through oribatid mites. lake at tree-line within the altitudinal The main scope of the entire thesis is to gradient. The ecological information demonstrate the usefulness of oribatid acquired, in addition to that from Paper mites as tools for inferences about past I, will also be of use in future habitats in paleoecology and investigations. The modern assemblages archaeology and to address questions provide a guide to the reliability of the related to the use of oribatid mites as fossil assemblages to reconstruct proxies in palaeoecology and ecological and environmental changes archaeology. This has resulted in 4 and, in addition, to finding the most papers that address different research favourable point to take a sediment core questions: within the small lake. Paper I - Are there distinct Paper III – Does the mite fauna oribatid mite communities characteristic in lake sediments of Trettetjørn reflect of different vegetation types in western Holocene environmental changes? What Norway? Can we detect characteristic does the addition of oribatid mites to a oribatid communities in the tree-line multiproxy study add to the overall ecotone? The oribatid communities environmental interpretations? In this associated with mixed (coniferous and paper I use oribatid mites as proxies to deciduous), birch, pine, and deciduous reconstruct the Holocene environmental forests, lichens and mosses, wet history of Trettetjørn, the low-alpine habitats, and the tree-line ecotone are (800 m a.s.l.) small lake in Paper II. characterized. This type of study is This is a multi-proxy study combining invaluable for palaeoecological mites with pollen, plant macrofossils, reconstructions, as it helps in and diatoms analyses in an attempt to identifying the type of plant community reconstruct the fluctuations in tree-line and environment the fossil oribatid and their impact on the lake assemblages might reflect. environment. Paper II - Which oribatid Paper IV - Can oribatid mites ecological groups are best represented be used to detect Neolithic or later in lake sediments? Which is the best impact on the surroundings of the coring point within Lake Trettetjørn settlement? In this study I use oribatid basin? In this paper I study the mites in an archaeological context. Mite representation of oribatid communities data are combined with pollen data to at the tree-line ecotone in western reconstruct the environment Norway and compare it with the surrounding a human settlement from SZ the Norwegian Bronze Age. The study basement rocks date from the also aimed to identify the start of cereal Precambrian and are basically cultivation at Kvitevoll farm, on unaffected by later events. They are Halsnøy Island, western Norway. mainly gneiss, granite, and quartzite and are found in northern and southern Norway. The Caledonian orogeny Study Areas developed during the Silurian and The climate in Norway is determined by Devonian periods. The resulting its northerly location in Europe on the mountain chain stretches southwest to east side of the Atlantic Ocean and the northeast in Scandinavia, and in marked relief of the country (Moen Norway from Rogaland to Finnmark. In 1999). The world climate is divided in 5 addition to the bedrock, the superficial major regions based on precipitation deposits are also important for the and temperature, and Norway falls into vegetation cover that will eventually 3 of them: the arctic and alpine climate, develop. These deposits vary greatly found in the mountains and the areas far over Norway. In western Norway there north; the cool temperate climate, found are mainly three types: till, deposited by in most of the land below the tree-line; glaciers; mass-movement deposits, and the warm temperate climate, found which are rocks, stones, and earth that in the lowlands in southern and western have slid down a slope and accumulated Norway (Moen 1999). at its foot; and exposed bedrock with a The study areas of this thesis are thin cover of superficial deposits (Moen mainly situated in western Norway, in 1999). The third type dominates the the counties of Hordaland and Sogn og study area of this thesis. The type of soil Fjordane (papers I, II, III, IV). Thus, I that develops on top of the superficial focus on the climate of western deposits depends on climate and Norway. According to Moen (1999), bedrock. In western Norway, which has coastal regions of western Norway have an oceanic climate, podsolic soils are an oceanic climate influenced by the sea typical in forests producing acid litter. and characterised by relatively low Peaty soils are also common in areas summer (12ºC in average) and high with high ground water and where the winter (>0ºC) temperatures and high production of plant material exceeds its precipitation and atmospheric humidity decomposition. (eKlima 2009). Both precipitation and Elevation above sea level of the temperature vary widely not only study area varies in each of the studies geographically but also annually. In comprising this thesis. Thus, in paper I general, coastal districts receive most of there are different transects which vary the precipitation during autumn and in elevation from 300 to over 1000 m early winter, while inland precipitation a.s.l. Lake Trettetjønn is situated at 810 is higher during summer (Moen 1999). m a.s.l. (paper II and III), but the study The geology in Norway is area in paper II ranges from 663 m a.s.l. complex, but Moen (1999) divides it to 1120 m a.s.l. Kvitevoll farm (paper into two main categories: basement rock IV) is situated at 55 m a.s.l. Figure 7 and Caledonian mountain chain. The shows the position of the study sites. S[ The study sites are situated in swallow soils and dry areas, while the boreonemoral, southern boreal, Norway spruce (Picea abies) grows in middle boreal, northern boreal, and low richer soils. Downy birch (Betula alpine zones (Moen 1999). Coastal pubescens) is a constant element of the areas and areas along the fjords are in woodlands in this part of the country. the boreonemoral and southern boreal Usually, the woodland field layer is zones, which are characterised by dominated by members of the coniferous and broad-leaved woodlands. Ericaceae, such as Calluna, Empetrum, Ombrotrophic bogs and dry grasslands and Vaccinium. At higher altitudes, the are common along the coast and inland field layer is often dominated by along the fjords (Fremstad 1997). Scots bilberry (Vaccinium myrtillus) pine (Pinus sylvestris) is common on (Fremstad 1997).

Figure 7. Map of the study areas. Numbers I- II-III-IV on the upper left corner correspond to the different papers. Triangle marks on map I correspond to the sampling sites in paper I from Hardanger northwards to Sognefjord. Scale of map II-III is about 1:100000, see paper II.

Mires are a consistent the groundwater level is periodically component of the landscape in western high, swamp woodlands might be Norway. They are frequently found. There are several swamp types in ombrotrophic bogs with dwarf shrubs western Norway. Species-poor swamp such as Calluna. The tree layer is not woodland and grey alder – willow common in bogs unless they have dried woodland are common in the lowlands out. The ground layer is usually (Fremstad 1997, Moen 1999). dominated by a carpet of Sphagnum Soligenous mires, often dominated by species (Fremstad 1997). In areas where Cyperaceae, and lake-margin fens

TR dominated by Cyperaceae and communities in different habitats in Equisetum, are also common. western Norway. Soil samples of 25 x 25 x 5 cm were collected from different vegetation types in the selected habitats. Methods After collection, the samples were extracted using Berlese-Tullgren Field sampling funnels (Fig. 8) until they were Different sampling methods were completely dry (ca. one week) and the selected to match the purpose and scope extracted material was stored in 70% of the different projects. ethanol until sorted and identified. Paper I – This project studied the composition of oribatid mite

Aluminium Light funnel bulb

Aluminium Sample funnel b Grid

a Jar with ethanol + drops of detergent Figure 8. a. Berlese-Tullgren funnel as used at the Department of Biology, UiB. b. Inverted sample placed inside the Berlese-Tullgren funnel, viewed from above.

Paper II – In order to study the inlet, one in the middle, and one near plant and animal debris that is deposited the outlet. These traps were large plastic on the ground and potentially containers weighted down with a large transported into a lake, terrestrial traps rock and attached to a float to mark were set out in the vegetation types their exact location (Birks and Bjune along the elevation gradient. A 2010). terrestrial trap consisted of a plastic Paper III – The sampling AstroTurf© doormat, 50x80 cm, which followed the standard methodology of was laid out in the field for a season, palaeoecological studies (Douglas winter (September – June) or summer 2007). During winter 2001 a sediment (June - September), and then collected core from the deepest part of the lake and replaced to collect another season. (Trettetjørn) was retrieved, using a 110- The sampling was done during four mm modified piston corer (Nesje 1992). seasons (two years). In addition, to Paper IV – The sampling evaluate which oribatid ecological followed the standard methodology in groups were best represented in the archaeology and in the study of modern lake sediments, three traps were Quaternary insects in general (Elias set in the lake (Trettetjørn), one at the 1994, Elias 2010), in which the size of TS the samples analysed tends to be much a subsampling method which allows for larger than in palaeolimnology. Samples extrapolation. The methodology was at different depth intervals (every 3 cm) developed by Southwood (1978) and in the archeological trench were taken modified by Johannessen (2002) and is and sealed in labelled plastic bags. In described in detail in paper I. addition, a peat block was retrieved Some oribatid species can be from a trench dug in the mire beside the identified to species under a stereo archaeological excavation. The block microscope, but most need to be was wrapped in heavy-gauge aluminium examined under a compound light foil in order to preserve it as well as microscope, following treatment in possible until it was sub-sampled in the concentrated lactic acid that renders the laboratory. chitinous cuticle of most oribatids clear. Dorsal and ventral characteristics of Laboratory methods oribatid mites are important to identify Preparation of samples – Depending them to species. Therefore, they are on the nature of the project and where mounted on non-permanent cavity the samples were taken, different slides in some drops of concentrated techniques were needed to prepare the lactic acid which allows them to be samples so that they could be examined turned so that the desired character can under the stereo microscope to extract be observed at the correct angle. the oribatid specimens. Therefore, I Paper II – The samples describe briefly what was done in each discussed in this paper were collected in study. Mites were identified following doormats or submerged boxes. The mainly, Gilyarov (1975), Subías and doormats were washed using a shower Arillo (2001), and Weigmann (2006). In head and the water was passed through addition, special papers and reference a sieve of 125 µm mesh. The residues collections were used in some cases. were stored in water with a little phenol Specialist were consulted when needed. to prevent fungal growth. The rest of the Nomenclature was, afterwards, laboratory procedure was similar to standardized to Subías (2004, 2010). many palaeo-studies. Mites were picked Paper I – After collection, the out under a stereo microscope and samples were transported to the identified to species under the stereo Department of Biology at UiB. Mites microscope and compound microscope. were extracted from the soil samples Because these samples were not palaeo- using modified Berlese-Tullgren samples, most oribatids still retained funnels into 70% ethanol with a few their hard chitinous cuticles. These drops of detergent (as illustrated in specimens were treated with figure 8). After extraction, which concentrated lactic acid to make their usually took one week, the sample was cuticles clear. inspected for mites under a stereo Paper III and IV – These two microscope. Soil samples might contain papers provide practical examples of more than a thousand individuals of how oribatids can be used in palaeo- oribatid mites, resulting in tedious and reconstructions. In paper III, a sediment time consuming lab work. Thus, I used core (350 cm long) was retrieved from TT Trettetjørn under a water depth of 6.8 concentrated lactic acid in cavity slides m, using a 110-mm modified piston and examining them under a compound corer (Nesje 1992). Samples for plant light microscope. The identified mites macrofossil analysis were taken from 26 were counted and the numbers levels. The residues of those samples normalised to a constant volume (100 were used for mite analysis. cm3) as concentrations. In paper IV, samples were cut every three cm from the peat-block Data analysis (83.6 cm long and 16 cm wide) Depending on the research questions retrieved from a trench excavated in the addresses different analytical methods mire next to the archaeological were used. excavation. Paper I – TWINSPAN (Two-way Displacement of water in a Indicator Species Analysis, Hill 1979) measuring cylinder was used to estimate and DISCRIM (simple discriminant the volume of each sediment sample. In analysis, ter Braak 1986) were used to both papers, a sample was usually ca. explore the large and noisy data-set and 3 30 cm . It is also possible to measure uncover patterns in oribatid distribution. the weight instead of the volume in Canonical Correspondance Analysis order to estimate the total number of (CCA) was used to organise the data in oribatid mites found, but I find it less relation to the environmental variables. time consuming to measure the volume. Forward selection and Monte Carlo Palaeo-samples are usually permutation test were also performed to disaggregated in dilute sodium detect those variables that significantly pyrophosphate or, if necessary, in 10% explain, in a statistical sense, the KOH (Birks 2001, Solhøy 2001, Elias variance in the data. Variation 2010) and rinsed through two sieves, partitioning was used to assess the 125 µm and 250 µm mesh size, in order relative importance of the spatial to divide the sample into two fractions. variables and habitat conditions in This facilitates finding the mites when influencing the oribatid communities. examining the samples under the stereo The program CANOCO 4.5 for microscope. Some authors who work in Windows (ter Braak and Šmilauer palaeo-studies use flotation techniques 2002) was used to perform CCA and to separate the mites from the palaeo- Variation partitioning. samples (see Erickson and Platt 2007 Paper II – Diagrams were made using for examples). However, afterwards the TILIA and TGView (Grimm 1993, residues should also be inspected for 2004). CANOCO 4.5 for Windows (ter mites under the stereo microscope. Braak and Šmilauer 2002) was used to Therefore, I found it less time perform Principal Components Analysis consuming to pick out the mites from (PCA) to summarise the patterns within the complete sieved samples under the the modern assemblages, to explore the stereo microscope. relationship to the different habitats Detailed identification of mite along the transect, and to compare the species is done by placing the modern oribatid fauna along the specimens in some drops of altitudinal gradient to the fossil oribatid TU assemblages in the Trettetjørn This work comprises the most sediments. A simple percentage exhaustive study of oribatid mites in approach was used to assess how well is Norway since Karppinen’s work in the oribatid fauna of the area 1971. A total of 64 new oribatid records represented in the Trettetjørn sediments. for Norway were documented. Numbers Paper III – TILIA and TGView of single species vary between several (Grimm 1993, 2004) were used to make thousand to a few hundred individuals, the stratigraphic diagrams. CANOCO and total sample numbers are always 4.5 for Windows (ter Braak and several thousand individuals. Šmilauer 2002) was used to perform a Chamobates borealis, Oppiella Detrended Correspondence Analysis nova, Moritzoppia neerlandica, and (DCA) to summarise the main trends in Rhinoppia subpectinata characterised the mite data through the core. the oribatid communities from Betula, Paper IV – TILIA and TGView mixed, and Picea forest. Achipteria (Grimm 1993, 2004) were used to make coleptrata, Acrotritria ardua, the stratigraphic diagrams. CANOCO Ceratozetes gracilis, and Oribatella 4.5 for Windows (ter Braak and calcarata were the most common Šmilauer 2002) was used to perform a species in deciduous forest. PCA to summarise the main trends in Hemileius initialis, the oribatid data in that part of the Nanhermannia coronata, C. borealis, sequence where they were found. Tectocepheus velatus, and Atropacarus striculus were the most common species in wet habitats. Limnozetes ciliatus, Outline of main results in the Mucronothrus nasalis, and four papers Trimalaconothrus glaber appeared only in waterlogged habitats, which confirms Background fauna characterisation their status as true aquatic species. The lichen and moss habitat was characterised by Carabodes Paper I - Oribatid mite communities labyrinthicus, C. marginatus, in western Norway Melanozetes mollicomus, and T. velatus. Distinct oribatid mite communities are The forest species Parachipteria characteristic of different vegetation punctata (Schatz 1983, Weigmann types in western Norway. The 2006) was rather common in this TWINSPAN (Hill 1979) and DISCRIM habitat, which can be explained by the (ter Braak 1986) analysis (Fig. 9 and frequent presence of moss in the Paper I), and also the CCA, samples from this habitat type. conveniently divided the whole dataset Can we detect characteristic into subsets according to the main oribatid communities in the tree-line habitat. This facilitated the ecotone? The reconstruction of the tree- characterisation of the oribatid line is a recurrent topic in communities found in each of the palaeoecology (Bjune 2005, Larsen et modern habitat types included in the al. 2006, Birks and Bjune 2010). The study. characterisation of the fauna inhabiting TV this ecotone may help in identifying this Several authors have underlined ecotone in Holocene sediments, and this the importance of a good knowledge of study of the oribatid communities in this the modern ecology of the oribatid ecotone in western Norway addresses fauna when using them in palaeo- this need. Generalist species such as T. studies (Schelvis 1990a, Solhøy 2001, velatus, H. initialis, and Oribatula Erickson and Platt 2007,). Therefore, tibialis dominated this habitat. studies like this one, in which oribatid However, forest species such as communities from different habitats are Camisia biurus (Schatz 1983, Gjelstrup characterised, are extremely useful in and Solhøy 1994), and lichen associated palaeoecology. In particular, the study species such as Carabodes marginatus provides thorough knowledge of the (Seyd and Seaward 1984) were also oribatid species and the communities found at the tree-line ecotone, which that occur in western Norway today. exemplifies the boundary character of this habitat.

Figure 9. TWINSPAN-DISCRIM diagram. The line on the left indicate the level of division. Indicator environmental variables are on the right of the class. Indicator species are on the left. The abbreviations at the end of each nodule correspond to the samples as indicated in table ESM1 from paper I.

TW disturbed and the longest, most Oribatid mite representation in lake complete sequence (Birks and Birks sediments and best coring point 1980, Berglund 1986, Glew and Last 2001). However, it seems that for Paper II - Oribatid mite assemblages oribatid mite analysis this is not always across the tree-line in western the optimal location (Erickson 1988, Norway and their representation in Solhøy and Solhøy 2000). The results of lake sediments this paper show that within Lake Trettetjørn basin a core retrieved from The best represented oribatid ecological the middle of the basin adequately group in the lake traps were aquatic, reflects the oribatid fauna of the followed by oribatids from the habitats catchment area. adjacent to the lake (Fig. 10). Similar The comparison of the Holocene results have been obtained in other oribatid assemblages from paper III and studies using lake sediments (Solhøy the oribatid fauna found in the lake and Solhøy 2000, Larsen et al. 2006, traps from paper II showed the Heggen et al. 2010), providing good importance of identifying the oribatid evidence that oribatids are excellent individuals to species level. This proxies of local habitats. increases the ecological indicator value In a multidisciplinary approach and consequently the reliability of any the same sampling point might not be palaeo-reconstruction done using optimal for all the proxies analysed. In oribatid mites. palaeolimnology, cores are usually retrieved from the deepest part of the basin in order to obtain the least

es peci

ted s cia o es ss es a s peci peci t s s ita s b ted a s cie specie ocia et h e c ss p ti id-w t s qua ire a res chen associated

TRAPS o i A M Hum F L Montane species Generalists ZONE S02 Northern boreal S03 S04 Sub-alpine S05 Tree-line S06 Low-alpine S07 S08 Mid-alpine INL MDL Lake OUL 20 40 60 20 40 60 20 40 60 20 20 40 60 20 40 ind/m 2of trap

Figure 10. Diagram showing the different ecological groups found along the altitudinal gradient and in the Trettetjørn traps. The horizontal axis represents the calculated number of oribatid individuals found per m2 of doormat trap over the collection period of 2 years. From left to right: aquatic species, mire associated species, moist-habitats associated species, forest species, lichen associated species, montane species and generalists. The vertical axis represents the different traps along the altitudinal gradients (S02 to S08) and in Trettetjørn (INL – inlet, MDL – middle, OUL – outlet). The dashed line through S05 indicates that this trap site is situated at the tree-line.

TX recovered. After deglaciation most Palaeolimnological and archaeological probable few oribatids populated the reconstructions using oribatid mites landscape, and therefore few or no The use of oribatid mites as indicators oribatid is likely to be incorporated into of past habitats, as stated above, is a the sediments. The unstable landscape rather recent approach. The fact that might also have an stronger physical many aspects of oribatids’ biology and effect on the fossil mites during ecology are unknown makes it difficult deposition, i.e. in the processes of being to implement quantitative techniques washed from the catchment. using oribatid fossil data. However, this Aquatic oribatid species became is not a problem when reconstructing abundant in the second zone with the past habitats, as outstanding studies development of aquatic and shore using oribatid mites have been vegetation after 8578 cal yr BP. This published, including one from Kråkenes might also indicate a wetter climate, in western Norway (Solhøy and Solhøy with a higher number of oribatids being 2000). This thesis is also comprised of washed into the basin. The oribatid two reconstructions of past habitats assemblage was a combination of open using oribatid mites as habitat habitat and woodland species such as indicators, paper III and paper IV. Liebstadia similis and Dometorina plantivaga (Grandjean 1950, Travè Paper III: Holocene environmental 1963, Solhøy 1976, Solhøy 1979, and climate history of Trettetjørn, a Maraun and Scheu 2000) during the low-alpine lake in western Norway, second zone (8270–7760 cal. yr. BP). based on subfossil pollen, diatoms, The macrofossil record indicated oribatid mites, and plant macrofossils that woodland of birch and pine became closed around Trettetjørn during the This study combined data on oribatids third phase (7760–5200 cal. yr. BP) with pollen, plant macrofossil, and associated with the occurrence of the diatom data. The results obtained from oribatid species Ophidiothricus tectus the fossil oribatid analysis added and D. plantivaga that are typical reliability to this study, as they point to woodland species (Weigmann 2006). the same direction of environmental During the fourth phase (5200– change as the other proxies involved in 4175 cal. yr. BP), there was a retreat of this project. In addition, the weaknesses the forest probably due to a cooling of and strengths of the different proxies are the climate. No oribatid species counterbalanced. associated with woodland were found. The Holocene oribatid mite This corresponded to the increase in sequence was divided into 6 phases, oribatids indicating more open habitats following the pollen zones (Figures 3 such as Hypochthonius rufulus. This and 4 in paper III) with a time span species is also associated with wet to between 8500 and 1000 cal. yr. BP. In soaked habitats (Solhøy 1979). the first or pioneer pollen zone (8575– According to the pollen record it seems 8270 cal. yr. BP) no mites were that mires developed locally during this phase, but judging from the oribatid TY record, mire formation had already number of oribatid species decreases started in the previous phase, in which with human disturbances. However, the oribatid species Trimalaconothrus members of the Oppiidae family seem spp., Malaconothrus sp., Moritzoppia not to suffer as much as other oribatid neerlandica, associated with wet taxa and are still present after habitat habitats (Solhøy 1979, Weigmann disturbance. Therefore, even if the 2006), increased. remains found at Trettetjørn were not Trees around Trettetjørn identified to species level, they could be disappeared in the fifth phase (4175– an indication of a higher disturbance 1555 cal. yr. BP). Forest oribatids were level in the area by sheep and goat still absent from the sediments, and grazing around the lake. there was a slight increase of oribatids characteristic of mires and grassland Paper IV: Neolithic impact on local such as Nanhermannia dorsalis (as N. vegetation at Kvitevoll, Halsnøy coronata in the paper) and Banksinoma Island, western Norway lanceolata (Karppinen 1955, Tarras- reconstructed from oribatid mites Wahlberg 1961, Solhøy 1976, Solhøy and pollen analysis 1979). During this phase, members of the family Camisiidae, an oribatid taxon This work is the first study in Norway often associated with heath (Solhøy in which oribatids are used as indicators 1979), were also found. This increase of of past environments in an terrestrial oribatids might be a result of archaeological context. A peat sequence increased water transport to the lake by was analysed and the preservation of increased precipitation. This might also oribatid remains was exceptional. have resulted in the increase in fern Therefore, it was possible to identify the spores as soil was washed in from the oribatids to species level. This, in catchment. addition to the high abundance of During the last phase (1555 cal. oribatid remains, increased the yr. BP–AD 2000) there were few reliability and robustness of the oribatid remains. The pollen record palaeoenvironmental reconstruction. indicated a more intense use of the The time span ranges from the landscape through grazing and summer late Mesolithic to the Late Neolithic; farming, which would result in an 7725 14C yr BP (6300 cal. yr. BP) at the artificial lowering of the local tree-line. bottom to 3130 14C yr BP (1300 cal. yr. There was an increase in microscopic BP) at the top of the sequence. Zonation charcoal, which is further evidence for of the sequence followed major changes human impact in the area. Aquatic in the mite abundances (Fig. 11) and plants disappeared and also aquatic pollen followed the same zonation. mites, only oribatids associated with The bottom of the sequence was mires and few others indicating heath devoid of oribatid remains and pollen were found. There were several remains was scarce. It seemed that at this stage of oribatids from the Oppiidae family. there was running water through the According to Behan-Pelletier (1999) site, which was an open landscape and Iturrondobeitia et al. (2005) the dominated by shrubs and wet areas. TZ This was followed by a rapid increase in drier. Oribatid species tolerating drier trees such as Betula and Salix, and an conditions such as Malaconothrus increase in Sphagnum moss, indicating monodactylus and Nanhermannia the onset of mire succession (Fig.4 in dorsalis (Knülle 1957b, Popp 1962, Paper IV). Markkula 1986) replaced Liebstadia Fossil oribatids do reflect lagenula, which prefers wetter changes in the surroundings of the conditions. Mites species typical of settlement. The oribatid sequence more open and moist biotopes such as showed the steadily opening of the Scheloribates laevigatus, Liebstadia woodland around the site (Zone II Fig. similis, Lauroppia fallax, Moritzoppia 11). The oribatid assemblage was neerlandica, and Banksinoma dominated by forest species such as lanceolata (Knülle 1957b, Wunderle et Caleremaeus monilipes and Micreremus al. 1990, Subías and Arillo 2001, brevipes, Cultroribula juncta (Schatz Weigmann 2006) indicated the further 1989, Subías and Arillo 2001, T. Solhøy opening of the woodland (Zone IV, pers. comm.), throughout the peat Fig.11a). However, the mite signal still sequence (Fig. 11), but steadily species showed an important presence of trees associated with more open habitats in the area (Fig. 11b). Species such as appeared, indicating the opening of the Damaeobelba minutissima, woodland (Fig. 11a). Mire became a Odontocepheus elongatus, and more dominant element of the Ophidiothricus vindobonensis indicated landscape. The oribatid fossil the presence of well-established assemblage reflected the typical forest- (deciduous) forest (Pérez-Íñigo 1997, mire oribatid community (Zone III, Fig. Weigmann 2006, Schatz and Fischer 11a,b), as suggested by the presence of 2007). Oribatids cannot directly indicate the oribatids Fuscozetes fuscipes, human disturbances, but the increase in Moritzoppia neerlandica, and members of the family Oppiidae, Liebstadia lagenula all considered peat considered common in disturbed areas soil dwellers (Knülle 1957b, Weigmann (Behan-Pelletier 1999, Clapperton et al. and Miko 2002, Schatz 2004, 2002, Iturrondobeitía et al. 2005), as Weigmann 2006, Schatz and Fischer well as the generalist species 2007, Schatz 2008) and forest species Tectocepheus velatus, during zone IV, such as Liacarus splendens and indicated higher disturbances in the Scheloribates pallidulus (Subías and area, although not necessarily Arillo 2001, Weigmann 2006). Towards anthropogenic. the end of the sequence the mire got

T[ Mire species Moist habitat species

s u yl ct lis ca s da a i tu rs lata no o a iga l is nu ceo erland ia d an al us mon age l allax iti ia ne n a l ma s i nothr i o pp u ibates laev o ad o ei r t tz l lo lac aherman ri e Depth (cm) bs mi 14C yr BP a e nksin o e M Nan Li Ba LauroppiaM f FuscozetesH fuscipes LiebstadiaSch similis Zone 3130±50 46 Lithology 51 56 61 IV - Anthropogenic signal 66 71 3905±90 76 81 III - Mire 86 91 96 101 II - Forest - mire development 106 111 20 20 40 60 20 200 400 600 800 20 20 40 60 7725±145 no. mites in 100 cc a

Forest species

is ns us s one u gat s e dob a dul lon ip in tsi li v v g erat al se oi p s monilipes hu um e chus h es lba minutissima es va at ep mus bre i maeu toc re tad re re diotri

Depth (cm) bs 14C yr BP ale hamobate helorib DameobeCultroribulaLiacarusC juncta splendens Odon Mic OphiC Li Sc Zone 3130±50 46 Lithology 51 56 61 IV - Anthropogenic signal 66 71 3905±90 76 81 III - Mire 86 91 96 101 II - Forest - mire development 106 111 20 40 60 20 20 40 60 80 7725±145 no. mites in 100 cc b

UR Lichen associated species Generalist

s eta ekensi orum s lis c nata ubpectinataor parmeliae exi a lu s a es es sara n nova ul hi a oppi oppia lani ell bat i Depth (cm) 14C yr BP 14C yr ri haul ult MycobatMycobatO P M RhinoppiaDissor Oppi Tectocepheus velatus Zone 3130±50 46 Lithology 51 56 61 IV - Anthropogenic signal 66 71 3905±90 76 81 III - Mire 86 91 96 101 II - Forest - mire development 106 111 20 20 20 20 200 400 600 800 50 100 150 7725±145 no. mites in 100 cc c

Limus humosus 3 Detritus lignosus 1Detritus herbosus 1 Limus humosus 1 Detritus lignosus 3

Limus humosus 2 Detritus lignosus 2 Limus humosus 3 Limus humosus 4 Argilla granosa 1 Grana arenosa 4

Figure 11. a. Mite concentrations from the Kvitevoll peat sequence showing mire species, moist-habitat associated species, and lichen associated species. b. Mite concentrations from Kvitevoll showing forest species. c. Mite concentrations from Kvitevoll peat sequence showing lichen associated species and generalists. The data are shown on a depth basis with the three radiocarbon dates (7725±145, 3905±90, 3130±50) on the left-hand side on all diagrams. Lithology symbols follow of all diagrams follow top legend.

other works, there is still a long way to go before the usefulness of oribatids in Conclusions, problems, and palaeoecological studies is fully further work developed. Among the main reasons are the lack of knowledge about oribatids’ The primary aim of this thesis is to biology and ecology, about the investigate and evaluate the usefulness taphonomical processes controlling of oribatid mites as past habitat which species are incorporated into the indicators and to characterise some sediments, and about oribatid mite’s present day oribatid communities taxonomy. The four papers comprised relevant for palaeoecological in this thesis contribute to overcome reconstructions. The results indicate that these problems. Paper I does not fossil oribatid mites are valuable and provide new aspects of their biology, useful indicators of past habitats. but it characterises the oribatid Despite the results shown in this and communities found in common US vegetation types in western Norway. able to reflect habitat changes in the This knowledge is of vital importance lake catchment. when using oribatids as proxies in Scientific knowledge is not built palaeoecological studies, especially in quickly. It may take years, decades, western Norway. Further meticulously even centuries before a new concept or planned studies of community structure, paradigm is accepted. Much work is habitat choice of oribatid mites, and still needed in the palaeo-acarological changes due to anthropogenic world, but it is, after all, only a century disturbances will be of invaluable use in since oribatids were first discovered as palaeoecology; they are definitely fossils. Thus far, their use in needed. palaeoecological contexts has The abundance of oribatid demonstrated that they can provide remains found, especially in lake unique palaeoenvironmental sediments, also constitutes an interpretations. These generally cannot interesting problem. There are many stand alone, but need to be seen in a aspects of oribatid taphonomy that are multi-proxy context where they still not known, yet they affect which enhance habitat and environmental species are incorporated and preserved reconstructions made from the other in the sediments. It could be proxies. hypothesised that better preserved and higher numbers of mites might be retrieved from sediment cores taken Acknowledgements closer to the lakeshore. However, paper I am grateful to Torstein Solhøy, John II shows that this is not always Birks, Hilary Birks,Lita G. Jensen and necessarily true. The problem is less Matteo Pedercini for their helpful acute when analysing mire-peat comments on earlier versions of this sediments, but in this case, species synthesis. I would like to thank Brooke associated with mires will dominate the Wilkerson for proof-reading and for her sequence. Paper IV shows that peat valuable suggestions to improve this sequences may result in robust habitat synthesis. reconstructions, and oribatids can indicate the changes in the surrounding local environment. References There is no doubt that reliability Arroyo, J., A. de la Riva-Caballero, J. C. and robustness of any palaeo- Iturrondobeitia, J. M. B. Castro, E. Carbonell, J. L. Arsuaga, and C. Diez. reconstruction using oribatid mites 2007. Primera aproximación a la increases with increasing taxonomic paleoentomología de los yacimientos de la resolution. This is because within the sierra de Atapuerca (Burgos, España): la same genus there might be species fauna subfósil de Oribatidos (Acari, Oribatida). Graellsia, 63: 27-34. living in completely different habitats. Paper III shows that even if the mite Behan-Pelletier, V. 1999. Oribatid mite remains are identified at a low level of biodiversity in agroecosystems: role for detail, this taxonomic resolution is also bioindication. Agriculture, Ecosystems and Environment, 74: 411-423.

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