The influence of spatial scale by thaiv2012


									Ecological Entomology (2003) 28, 252–256


The influence of spatial scale on quantifying insect
dispersal: an analysis of butterfly data
CHRISTINE SCHNEIDER                                   Swedish University of Agricultural Sciences, Department of Landscape
Planning, Alnarp, Sweden

                                      Abstract. 1. A linear relationship between mean movement distances recorded
                                      and the size of the study area was found in a comparison of five mark–release–
                                      recapture studies of the meadow brown butterfly.
                                        2. The scale impact on mean butterfly movement distances was also apparent
                                      when comparing the results for different butterfly species reported in 21 mark–
                                      release–recapture studies.
                                      Key words. Dispersal distance, landscape scale, Maniola jurtina, mark–release–
                                      recapture, movement.

Introduction                                                                                                       `
                                                                     underestimation of butterfly mobility (Neve et al., 1996;
                                                                     Hanski et al., 2000). Koenig et al. (1996) addressed the
Dispersal is seen as crucial for species conservation at the         problem of movement truncation due to small study areas
landscape level (Baudry & Burel, 1997), therefore it is an           for vertebrates. Mason et al. (1995) showed for a grasshopper,
issue that has been studied quite intensively in recent years        as did Wilson and Thomas (2002) for butterflies, that
in a wide range of taxa. Butterfly dispersal is especially well      mark–release–recapture data can be biased and how this
studied because butterflies are of great interest for conser-        leads to underestimation of dispersal ability.
vationists and have become a key group of species in                    The aim of the work reported here was to contribute to
metapopulation research (Harrison et al., 1988; Hanski &             the debate by reviewing the impact of the dimensions of the
Gilpin, 1991).                                                       study area on reported mean distances moved by the butterfly
   One of the main methods used to study dispersal is the            species under investigation. The meadow brown was used
mark–release–recapture technique. Based on mark–release–             as an example to illustrate the point that studies at
recapture experiments, dispersal parameters such as mean             different scales can arrive at quite contrasting results. A
and maximum distances moved can be derived. The mean                 comparison of several mark–release–recapture studies of
distance in butterfly mark–release–recapture studies has             different species demonstrated the influence of study area
often been used to evaluate the dispersal ability of a species.      size on mean movement distances recorded.
The results of several mark–release–recapture studies with
butterflies show that mean distances moved by butterflies
are quite low and it has been assumed that the studied               Methods
species are sedentary (Brakefield, 1982; Thomas, 1985;
Warrington & Brayford, 1995; Fischer, 1998; Fjellstad,               The meadow brown Maniola jurtina L. was chosen as an
1998; Fischer et al., 1999). It is known that mark–release–          example to examine the influence of study area dimensions,
recapture studies underestimate dispersal (Shreeve, 1992),           because it is one of the best-studied butterfly species. Five
but it was first with the results of large-scale studies that this   studies were compared in relation to the size of the study
became more obvious (overview by Hanski & Kuussaari,                 area and the mean distances calculated for Maniola jurtina
1995; Brunzel & Reich, 1996; Neve et al., 1996; Mousson              (Brakefield, 1982; Dover et al., 1992; Lortscher et al., 1997;
et al., 1999; Baguette et al., 2000; Petit et al., 2001). In         Ouin, 2000; Schneider et al., 2003). The approximate size of
addition, observations of colonies of species classified             the study area of Lortscher et al. (1997) was reconstructed
previously as very sedentary have provided evidence of an            on the basis of a map published in that article. The mean
                                                                     distances were given for both sexes in all studies except for
                                                                     that of Ouin (2000). Thus mean distances for male move-
  Correspondence: Christine Schneider, Swedish University of         ments were used in the comparison, apart from the data of
Agricultural Sciences, Department of Landscape Planning, PO Box      Ouin (2000), in which the mean distance moved was ana-
58, 230 53 Alnarp, Sweden. E-mail:   lysed for both sexes together.

252                                                                                        #   2003 The Royal Entomological Society
                                                                                       Influence of spatial scale on dispersal   253

   Additionally, the mean distances of 21 species reported in
21 mark–release–recapture studies were compared according
to the size of the study area and the mean distance moved
by the butterfly. The analysis of butterfly data was confined
to European studies. Considered were studies known to the
author and studies found in the BIOSIS database (years
1992–2002) under the key words butterfly and mark-release-
recapture, and Lepidoptera and mark-release-recapture. The
studies included in the analysis were chosen because they
fulfilled the requirement that the mean distance moved and
the size of the study area were recorded. Mean distances
were given for both sexes in most but not all studies, there-
fore overall means for both sexes were calculated by taking
                                                                      Fig. 1. Recorded mean distance moved by the meadow brown
the sex ratio into account. This was possible for all studies         (males) in relation to the size of the study area.
apart from that of Fischer et al. (1999) for which rougher
estimates had to be used. Surprisingly few studies named
                                                                         It is also interesting to note that some of the studies that
explicitly the size or extent of their study area, but all studies
                                                                      investigated several species in the same habitat network,
where the approximate size could be calculated from maps
                                                                      arrived at similar results for the different species. Even if
were included. Maximum distances recorded are given as
                                                                      the differences turned out to be significant in these studies,
additional information.
                                                                      they seem to be minor in a larger context. Compare,
                                                                      for example, the results for Maniola jurtina (mean 398 m)
                                                                      and Pyronia tithonus (mean 395 m) studied by Dover
                                                                      et al. (1992), Lysandra coridon (mean 130 m), Melanargia
Results and discussion
                                                                      galathea (mean 105 m), and Maniola jurtina (mean
                                                                      94 m) studied by Lortscher et al. (1997), Maniola jurtina
Despite the fact that the meadow brown in the five reports
                                                                      (mean 323 m) and Lycaena virgaureae (mean 271 m) studied
reviewed was studied in different types of landscape,
                                                                      by Schneider et al. (2003), and Erynnis tages (mean 85 m)
ranging from continuous habitat patches to very frag-
                                                                      and Zygaena filipendulae (mean 33 m) studied by Gutierrez  ´
mented landscapes (Table 1), the overriding factor influen-
                                                                      et al. (2001). The differences between the same species
cing the recorded mean distances seemed to be the
                                                                      studied at different scales were, on the other hand,
dimensions of the study area. There was a linear relationship
                                                                      much larger, as the example of Maniola jurtina shows.
between mean distance recorded for the meadow brown and
                                                                      This might also show that the spatial configuration of
the size of the study area in which the experiment was carried
                                                                      the mark–release–recapture areas has an important impact
out (R2 ¼ 0.95, F1,7 ¼ 138.0, P < 0.001; Fig. 1).
                                                                      on the results derived from mark–release–recapture studies.
   The comparison of butterfly mark–release–recapture
studies (Table 2, Fig. 2) showed (1) that the scale impact
on butterfly movement was also apparent when comparing
different species (R2 ¼ 0.81, F1,25 ¼ 106.6, P < 0.001, without       Conclusion
the Maniola studies); (2) that there does not seem to be a
clear relationship between observed mean distance and size            This review illustrates the fact that the scale of the study
of study area for the studies carried out in areas of 50 ha          area has a major impact on the results of mark–release–
(R2 ¼ 0.007, F1,14 ¼ 0.1, P ¼ 0.75). Thus it could be expected
that more dispersal studies at a larger scale would make the
differences between species clearer.

Table 1. Size, nature of the study areas, and scale used in mark–
release–recapture experiments on Maniola jurtina.

Size (ha)          Nature of the site       Author (year)
1.4–3 (3 sites)    Mainly meadows           Brakefield (1982)
9                  Alpine grassland         Lortscher et al. (1997)
40–110 (3 sites)   Mosaic grassland,        Ouin (2000)
172                Mosaic of grassland,     Schneider et al.
                   arable and settlement    (2003)
300                Arable farmland          Dover et al. (1992)       Fig. 2. Relationship between mean distances recorded in mark–
                                                                      release–recapture studies and the size of the study area.

#   2003 The Royal Entomological Society, Ecological Entomology, 28, 252–256
254 Christine Schneider

Table 2. Comparison of mean and maximum distances observed between recaptures in mark–release–recapture studies and the size of the
study area.

                                                  Mean distance between
                               Size of study      recaptures (males and         Maximum
Species                        area (ha)y         females) (m)z                 distance (m)      Author
Aphantopus hyperantus           37                 92                             460             Sutcliffe et al. (1997)
Boloria aquilonaris            262                205§                           1270             Mousson et al. (1999)
Erynnis tages                   28                 85                             660                         ´
                                                                                                  Gutierrez et al. (1999, 2001)
Euphydryas aurinia               4                 70                             486             Munguira et al. (1997)
Euphydryas aurinia             400                630                            1300             Wahlberg et al. (2002)
Euphydryas maturna             400                224                             640             Wahlberg et al. (2002)
Lycaena helle                    6                 44{                            280             Fischer et al. (1999)
Lycaena helle                   13                 44{                            380             Fischer et al. (1999)
Lycaena hippothoe               12.5               53                             575             Fischer (1998)
Lycaena virgaureae               3.4               57                           >1000             Douwes (1975)
Lycaena virgaureae              40                 61                            1424             Fjellstad (1998)
Lycaena virgaureae             172                271                            1460             Schneider et al. (2003)
Lysandra coridon                 9                130                             712             Lortscher et al. (1997)
Maniola jurtina                  1.4               45                                             Brakefield (1982)
Maniola jurtina                  1.6               68                                             Brakefield (1982)
Maniola jurtina                  3                 59                                             Brakefield (1982)
Maniola jurtina                  9                 94                              377            Lortscher et al. (1997)
Maniola jurtina                 40                 80yy                                           Ouin (2000)
Maniola jurtina                 53                150yy                                           Ouin (2000)
Maniola jurtina                110                180yy                                           Ouin (2000)
Maniola jurtina                172                323                             2100            Schneider et al. (2003)
Maniola jurtina                300                398                             1770            Dover et al. (1992)
Melanargia galathea              9                105                              433            Lortscher et al. (1997)
Melitaea athalia                15                101zz                            787            Schwarzwalder et al. (1997)
Melitaea athalia               400                505                             1676            Wahlberg et al. (2002)
Melitaea cinxia                160                353§§                           1150            Kuussaari et al. (1996)
Melitaea cinxia                400                480                             1700            Wahlberg et al. (2002)
Melitaea diamina               400                441                             1370            Wahlberg et al. (2002)
Parnassius apollo              350                341                             1840            Brommer & Fred (1999)
Parnassius mnemosyne            75                177                             2250                              ˇ
                                                                                                  Konvicka & Kuras (1999)
Plebejus argus                  50                 23                              395            Lewis et al. (1997),
                                                                                                  Gutierrez et al. (2001)
Proclossiana eunomia            30                 85                              920            Baguette & Neve (1994)`
Pseudophylotes baton            50                102                             1490            Vaisanen et al. (1994)
                                                                                                          ¨ ¨
Pyronia tithonus               300                395                             1060            Dover et al. (1992)
Thymelicus sylvestris            1                 70{{                                           Warrington & Brayford (1995)
Zygaena filipendulae            28                 33                              533                            ´
                                                                                                  Gutierrez et al. (2001)

yNumbers in italics indicate that the size of the study area was calculated on the basis of maps.
zNumber in italics indicates that the mean distance moved was calculated from the means of males and the means of females given the sex
ratio; if there were means from several years, the highest mean was chosen.
§Data from area Grande Fange.
{Mean distance moved was calculated as mean of mean male distance and mean female distance as sex ratio was not given.
yyNumbers were derived from graph.
zzMean distance was only given between first and last recapture.
§§Release experiments where individuals were released in an empty habitat network.
{{Number is estimated roughly from a table.

recapture investigations. To get a more accurate picture of            Acknowledgements
a butterfly’s dispersal ability, large-scale studies are recom-
mended. Additionally, if a species shows sedentary                     I am especially thankful to Gary Fry, who has given
behaviour when studied in small habitat fragments, this                valuable comments on earlier drafts of the manuscript.
can also be misleading, because this might be a result of              I also thank Chris Thomas for encouraging me to carry out
the inability of the species to cross an unfavourable matrix           this study. Mary McAfee has kindly improved the English.
rather than a lack of movement ability (Shreeve, 1995).                This study was financed by the Department of Landscape

                                                          #   2003 The Royal Entomological Society, Ecological Entomology, 28, 252–256
                                                                                           Influence of spatial scale on dispersal        255

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                                                                        Konvicka, M. & Kuras, T. (1999) Population structure, behaviour
                                                                              and selection of oviposition sites of an endangered butterfly,
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                                                            #   2003 The Royal Entomological Society, Ecological Entomology, 28, 252–256

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