Table Of ContentAQUATIC MITES: FROM GENES TO COMMUNITIES
AQUATIC MITES
FROM GENES TO COMMUNITIES
Editor
HEATHER C. PROCTOR
University ofA lberta, Edmonton, Canada
Reprinted from Experimental and Applied Acarology
Volume 34 Numbers 1-2, 2004
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Springer-Science+B usiness Media, B.V.
A C.I.P. Catalogue record for this book is available from the Library of Congress.
ISBN 978-90-481-6710-4 ISBN 978-94-017-0429-8 (eBook)
DOI 10.1007/978-94-017-0429-8
Prill ted 011 acid-Fee paper
All Rights Reserved
© 2004 Springer Science+ Business Media Dororecht
Originally published by Kluwer Academic Publishers in 2004
Softcover reprint of the hardcover 1s t edition 2004
No part of the material protected by this copyright notice may be reproduced or utilized
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permission from the copyright owner.
Cover photograph: With their often brilliant colours, water mites are strikingly con
spicuous compared to most other freshwater invertebrates. Many species, such as
Limnochares americana Lundblad (Limnocharidae), are bright red, and are also dis
tasteful to fish. The origin and function of this apparent aposematism are discussed in
Proctor & Garga in this volume. Photo by Heather Proctor; L. americana specimens
from Hasse Lake, Alberta, Canada.
TABLE OF CONTENTS
H.C. Proctor / Aquatic mites: from genes to communities - an
introduction 1-2
G. Sevik / The biology and life history of arctic populations of the
littoral mite Ameronothrus lineatus (Acari, Oribatida) 3-20
DJ. Marshall and P. Convey / Latitudinal variation in habitat
specificity of ameronothrid mites (Oribatida) 21-35
1. Bartsch / Geographical and ecological distribution of marine
halacarid genera and species (Acari: Halacaridae) 37-58
1. Rey, B.A. Dorda and A.G. Valdecasas / Traditional water mite
fixatives and their compatibility with later DNA studies 59-65
D.D. Edwards, D.E. Deatherage and B.R. Ernsting / Random
amplified polymorphic DNA analysis of kinship within host
associated populations of the symbiotic water mite Unionicola
Joili (Acari: Unionicolidae) 67-77
M.R. Forbes, K.E. Muma and B.P. Smith / Recapture of male and
female dragonflies in relation to parasitism by mites, time of
season, wing length and wing cell symmetry 79-93
P. Martin / Specificity of attachment sites of larval water mites
(Hydrachnidia, Acari) on their insect hosts (Chironomidae,
Diptera) - evidence from some stream-living species 95-112
B.P. Smith and J. Florentino / Communication via sex pheromones
within and among Arrenurus spp. mites (Acari: Hydrachnida;
Arrenuridae) 113-125
H.C. Proctor and N. Garga / Red, distasteful water mites: did fish
make them that way? 127-147
A. Boulton, M. Harvey and H. Proctor / Of spates and species:
responses by interstitial water mites to simulated spates in a
subtropical Australian river 149-169
T. Goldschmidt/Environmental parameters determining water mite
assemblages in Costa Rica 171-197
A. Di Sabatino, A. Boggero, F.P. Miccoli and B. Cicolani /
Diversity, distribution and ecology of water mites (Acari:
Hydrachnidia and Halacaridae) in high Alpine lakes (Central
Alps, Italy) 199-210
Experimental and Applied Acarology
© 2004 Kluwer Academic Publishers.
Aquatic mites: from genes to communities - an
introduction
Although chelicerates originated in the sea, the vast majority of extant species
are terrestrial. Since the invasion of land, only one species of spider has taken
up the old subaquatic life (Argyroneta aquatica), and there are a few intertidal
pseudoscorpions that can withstand inundation. It is among the mites that one
finds the greatest number of species, indeed, of entire superfamilies, that have
taken up a watery existence. Each aquatic taxon has close terrestrial relatives,
and sometimes displays rudiments of terrestrial adaptations (e.g. closed
stigmatal openings), and so it seems that invasion of the aquatic environment
by mites has occurred repeatedly. Approximately 7000 species from the Mes
ostigmata, Astigmata, Oribatida and especially the Prostigmata, now live in
marine and freshwater habitats. There are even a few ticks that parasitize
marine iguanas and sea snakes! In part because of their unusual lifestyle, and
likely also because few have any obvious agricultural or medical importance,
aquatic mites are usually relegated to habitat-specific journals together with
various other aquatic invertebrates, and have rarely appeared in the pages of
Experimental and Applied Acarology.
We hope that this special issue will help to bring aquatic mites into 'main
stream' Acarology. It is the product of a brief conversation between HP and
EAA-editor Jan Bruin at the poster session of the International Congress of
Acarology in Merida, Mexico, in September 2002. Jan asked Heather whether
she would be interested in acting as guest editor for a special issue of Experi
mental and Applied Acarology dealing with aquatic mites. Saying 'yes' required
but a moment's consideration; bringing the issue into fruition has been a rather
longer process! But the result was worth it. Here are 12 new papers that cover a
wide range of taxa and a great diversity of themes. Marine and freshwater
oribatids are featured in papers by Marshall and Convey and by S0vik, where
we learn that the Arctic species Ameronothrus lineatus takes 5 years to reach
maturity (twice as long as the average lifespan of many arctic rodents!). A
summary of current knowledge about the distribution and ecology of marine
halacarids is presented by Bartsch; this information will be extremely valuable
given the increasing transportation of marine taxa by shipping, and the
importance of assigning point of origin to animals found in ballast water. The
remaining papers deal with the most species-rich group of aquatic mites, the
'true' water mites (Prostigmata: Parasitengona). These range from studies of
genes (Rey et aI., Edwards et aI.), larval parasitism (Forbes et aI., Martin),
mating behaviour (Smith and Florentino), evolution of warning colouration
(Proctor and Garga), and relationship between environmental factors and
2
community composItion (Di Sabatino et aI., Boulton et aI., Goldschmidt).
These articles raise as many interesting questions as they answer, and should
provoke more studies of the biology of freshwater and marine Acari.
I thank the authors and the many excellent reviewers who helped to create
this special issue.
Heather Proctor
Department of Biological Sciences
University of Alberta
Edmonton
Alberta T6G 2E9
Canada
e-mail: [email protected]
... Experimental and Applied Acarology 34: 3-20, 2004.
" © 2004 Kluwer Academic Publishers. Printed in the Netherlands.
The biology and life history of arctic populations
of the littoral mite Ameronothrus lineatus
(Acari, Oribatida)
GULDBORG S0VIK
Institute of Marine Research, Tromso branch, P.O. Box 6404, N-9294, Tromso, Norway; (e-mail:
[email protected]; phone: + 47-77-60-9753; fax: +47-77-60-9701)
Key words: Latitudinal distribution, Life cycle, Microevolutionary adaptations, Multi-instar
aggregations, Population dynamics, Reproductive biology, Temperature
Abstract. The present study attempts to elucidate possible microevolutionary adaptations of life
history traits of high-latitude populations of the hoi arctic, littoral oribatid mite Ameronothrus
lineatus by comparing arctic and temperate populations. Additionally, the paper provides an
overview of the limited research on general ecology and population biology of arctic populations.
In the Arctic the larviparous A. lineatus has a 5-year life cycle (Iarva-to-Iarva), and adults survive a
further 2-3 years. High survival to maturity is consistent with a low lifetime reproductive output of
ca. 20 larvae. The life history can be regarded as an extreme version of the typical oribatid life
history. However, several life-history features suggest specific adaptations of arctic populations. In
particular, the pre-moult resting stage is synchronized with the warmest part of the arctic summer,
which shortens this vulnerable part of development. High reproductive investment by females at
relatively low temperatures may represent a physiological adaptation to the cool arctic summer.
Finally, prolonged cold exposure positively affects reproduction and survival the following sum
mer, suggesting adaptation of the species to the highly seasonal arctic environment. On the other
hand, the ability of all life-cycle stages to overwinter, and a flexible life history with the species
being able to take advantage of favourable climatic conditions to accelerate development and
larviposition, seem to be ancestral features. Thus, the success of A. lineatlls in arctic habitats is
probably attributable to a combination of derived and ancestral life-history traits. Studies of
conspecific temperate populations are required to elucidate further local adaptations of arctic
populations.
Introduction
The diversity of the arctic arthropod fauna is fairly well known (Danks 1981;
Coulson and Refseth 2004), but information on population biology and life
history strategies of component species is fragmentary (Danks 1981, 1999).
Central to our understanding of terrestrial polar arthropods is knowledge
about possible life-history adaptations to the harsh polar environment. Many
of the typical life-history traits of polar arthropods (e.g., extended life span and
reduced reproductive output) agree with predictions from adversity selected
life-history strategies (Convey 1996). However, many of these traits must be
viewed as plesiotypic (ancestral) characters, enabling successful colonization of
the arctic environment, or consequences of ecological or physiological limita
tions (Danks 1981; Norton 1994; Convey 1996).
4
Oribatid mites are usually among the most abundant and species-rich ar
thropods in arctic soils (Behan 1978; Danks 1981), however, little information
exists about their demography and life histories in cold regions. In the Ant
arctic only one species, Alaskozetes antarcticus (Michael), has been studied in
detail (Block and Convey 1995), whereas the holarctic Ameronothrus lineatus
(Thorell 1871) is the only oribatid mite inhabiting arctic regions, for which
detailed knowledge on life history and demography exists (S0vik et al. 2003;
S0vik and Leinaas 2003a, b). The latter species is found in littoral habitats
across a wide latitudinal gradient extending from warm-temperate to high
arctic regions. This makes it a good model organism for studies of local
adaptation, as population-specific differences in life history can be examined
across a correspondingly wide environmental gradient. Warm-temperate
coastal regions have a uniform climate with precipitation distributed evenly
throughout the year. Growing seasons are long whereas winters are short and
mild (average temperatures of 0-3 °C in December-February) (www.metof
fice.gov.uk; www.dwd.de). In comparison, summers in the high Arctic are
short (2-4 months) and cold with restricted time for reproduction and com
pletion of life cycles. The winter is long with low soil temperatures (0 to
-30°C) (Coulson et al. 1995, 2000), depending on air temperature and snow
cover. Arctic habitats are generally arid because of low precipitation and high
winds (Danks 1981, 1999).
The present study attempts to elucidate possible local life-history adapta
tions of A. lineatlls to the arctic environment by comparing arctic and tem
perate populations. Additionally, the paper provides an overview of published
research on the general biology of arctic populations of A. lineatus.
Taxonomy and distribution
Ameronothrus lineatus (Figure 1), first described from Svalbard as Eremaeus
lineatus (type species), belongs to Ameronothridae Willmann 1931, which
comprises species found in intertidal and terrestrial habitats in both hemi
spheres (Schulte and Weigmann 1977). Ameronothrus Berlese 1896 is pre
dominately northern. The 10 species (Gilyarov 1975; Schubart 1975;
Weigmann and Schulte 1975) are mainly aquatic or semiaquatic, being dis
tributed on arctic, temperate and subtropical coastlines (Schuster 1966, 1988;
Schulte et al. 1975; Schulte and WPoigmann 1977).
The latitudinal distribution of A. lineatus stretches from southern England to
the high Arctic, covering almost 30° of latitude (Coli off 1984). In North
America the equally wide distribution is shifted southwards into northern
California (Schulte et al. 1975; Schulte 1978). No reports exist from the
Canadian high Arctic (Danks 1981), probably due to limited sampling in those
inaccessible regions. Thus, A. lineatus has so far been found in Iceland, the
Faroe Islands, Greenland, Jan Mayen, Svalbard, Novaja Semlja, arctic Siberia,
5
Figure 1. Ameronothrus lineatlls. Lateral view of a female (from Colesbukta, Spitsbergen).
Fennoscandia, Great Britain, Ireland, Germany, Canada, Alaska, and Cal
ifornia (Hammer 1944; Strenzke 1955; Weigmann and Schulte 1975; Behan
1978; Danks 1981; Karppinen and Krivolutsky 1982; Colloff 1984; Coulson and
Refseth 2004). It is the only ameronothrid reported from the high-arctic Sval
bard (Coulson and Refseth 2004), where it has been sampled on Bear Island and
along the west and north coast of Spitsbergen (Summerhayes and Elton 1923,
1928; Karppinen 1967; S0vik et al. 2003). The life-history studies referred to in
the present work were carried out using specimens from Adventdalen (78°0'N
15°30'E) and Colesbukta (78°5'N 14°57'E) on West Spitsbergen.
Life history
Life-cycle stages
Oribatid mites have six life-cycle stages: pre-larva, larva, proto-, deuto-, tri
tonymph and adult. The pre-larva does not hatch from the egg (Norton 1994;
Walter and Proctor 1999), and in Ameronothrus is seen as a layer of cuticle
within the eggshell (Schubart 1975). The majority of oribatid species deposit
eggs, but some retain the progeny until the larval stage (Iarviparity) (Norton
1994). This seems to be a plesiotypic trait of the genus Ameronothrus (Schubart
1970, 1975; Weigmann and Schulte 1975; Pugh and King 1986; Tilrem 1994;
Biicking et al. 1998). Strictly speaking, A. lineatus is ovoviviparous as the
larvae hatch from the eggs some hours after deposition (Haq et al. 1991; S0vik
