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, November 2008, Pages
Symbiont microcosm in an ant society and the diversity of interspecific interactions, , , , , , , & Department Biologie II, Ludwig-Maximilians Universit&t M&nchen (LMU), Germany& Institute of Biological Sciences, University Malaya, MalaysiaColonies of the ponerine army ant Leptogenys distinguenda are regularly inhabited by a highly diverse symbiont fauna including insects, spiders, mites, crustaceans and even molluscs. Each of these myrmecophiles has adapted in a highly specific way to the lifestyle of its host. We studied this diverse myrmecophile fauna of L. distinguenda as a new model for multispecies parasitism to gain a better understanding of fundamental coevolutionary processes. Our study focused on behavioural and on chemical integration and exploitation strategies of the different symbiont species. In addition, we examined potential counterstrategies of the host ant. Myrmecophiles were studied both in large free-living L. distinguenda colonies and in more detail in parts of colonies separated for observation. We found that at least five myrmecophile species imposed cost on their host by exploiting its resources. Their impact varied considerably depending on both the type of resources exploited and their abundance. Myrmecophile species were well integrated into host societies either by chemical mimicry of host cuticular hydrocarbons or by remaining chemically insignificant, lacking most characteristic recognition cues. Despite these chemical integration strategies, host ants were able to recognize and kill the alien intruders to various degrees. This important finding demonstrates that symbiont populations are actively counter-regulated by the host. By constructing a host&parasite interaction network, we finally suggest that host defences can maintain myrmecophile diversity by keeping parasite populations small. This reduces interguild competition, comparable to top&down effects of predators on lower trophic levels in ecological food webs.Keywordsarmy ant; chemical integration; community ecology; host resistance; host&parasite interactions; Leptogenys distinguenda; migration; transmission mode
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.132 .132 scale
(Individual Differences in Attachment Preferences)S
.132 .132 scale
(Aurora Mendelsohn and Neal J. Pearlmutter, Northeastern University)S
.132 .132 scale
(Attachment Preferences)S
.132 .132 scale
(Relative Clause \(RC\) Attachment Ambiguity:)S
.132 .132 scale
(The crowds annoyed the chauffeur of the actor who wanted to )S
.132 .132 scale
(go home. )S
.132 .132 scale
(\245 English speakers prefer to attach the RC to the second NP \(N2\) )S
.132 .132 scale
(the actor )S
.132 .132 scale
(\(Cuetos and Mitchell, 1988\). )S
.132 .132 scale
(\245 Spanish speakers prefer to attach the RC to the first NP \(N1\) )S
.132 .132 scale
(the chauffeur
.132 .132 scale
(\(Cuetos and Mitchell, 1988\). )S
.132 .132 scale
(\245 Preferences tend to be weak, especially in on-line studies )S
.132 .132 scale
(\(Carreiras & Clifton, 1993\).)S
.132 .132 scale
234.95 6700.65 :M
(\245 Prepositions connecting the two potential attachment sites
.132 .132 scale
(\(e.g. )S
.132 .132 scale
(\) influence preferences. )S
(\(Gilboy et al., 1995\))S
.132 .132 scale
.132 .132 scale
(Individual Differences in Working Memory \(WM\))S
.132 .132 scale
(\245 Individual differences in WM have been found to influence )S
.132 .132 scale
(lexical and syntactic ambiguity resolution
(\(Miyake et al., 1994; )S
.132 .132 scale
(Pearlmutter and MacDonald, 1995\).)S
.132 .132 scale
(\245 Individual differences in WM have been considered differences )S
.132 .132 scale
(in experience )S
.132 .132 scale
(\(Pearlmutter & MacDonald, 1995\))S
.132 .132 scale
(\245 Exposure to one attachment pattern influenced preferences in )S
.132 .132 scale
500.217 9611.34 :M
(children )S
(\(Mitchell, 1994\))S
.132 .132 scale
WM differences might provide insight into this ambiguity)S
.132 .132 scale
.132 .132 scale
(References)S
.132 .132 scale
(Carreiras, M., & Clifton, C. \(1993\) Relative clause interpretation
preferences in Spanish and )S
.132 .132 scale
500.217 13499.84 :M
(English. )S
(Language & Speech)S
.132 .132 scale
.132 .132 scale
(, 353-372.)S
.132 .132 scale
(Cuetos, F., & Mitchell, D. C. \(1988\) Cross-linguistic differences in parsing: Restrictions on the use )S
.132 .132 scale
(of the Late Closure strategy in Spanish. )S
.132 .132 scale
(Cognition)S
.132 .132 scale
(, 73-105.)S
.132 .132 scale
(Cuetos, F.,
Mitchell, D. C, Corley, M. \(1996\) Parsing in different languages. In M. Carreiras, J. )S
.132 .132 scale
(Garcia-Albea & N. Sebastian-Galles \(Eds.\) Language
processing in Spanish \(pp.145-187\). )S
.132 .132 scale
(Mahwah, NJ, Lawrence Erlbaum.)S
.132 .132 scale
(Daneman, M., & Carpenter, P. A. \(1980\) Individual differences in working memory and reading.)S
.132 .132 scale
(Journal of Verbal Learning and Verbal Behavior)S
.132 .132 scale
(, 450-466.)S
.132 .132 scale
(Frazier, L. & Clifton, C. \(1996\) Construal. Cambridge, MA: MIT Press.)S
.132 .132 scale
(Gilboy, E., Sopena, J.-M., Clifton, C., & Frazier, L. \(1995\) Argument structure and association )S
.132 .132 scale
(preferences in Spanish and English
complex NPs. )S
(Cognition)S
.132 .132 scale
.132 .132 scale
(, 131-167.)S
.132 .132 scale
234.95 15379.66 :M
(Konieczny, L., Hemforth, B. Scheepers, C., & Strube, G. \(1997\) The role of lexical heads in )S
.132 .132 scale
(parsing: Evidence from German. )S
(Language and cognitive Processes)S
.132 .132 scale
.132 .132 scale
(, 307-348.)S
.132 .132 scale
(Mitchell, D. C. \(1994\) Sentence parsing. In M. A. Gernsbacher \(Ed.\),
Handbook of )S
.132 .132 scale
(psycholinguistics \(pp. 375-409\). San Diego, CA:
Academic Press.)S
.132 .132 scale
(Miyake, A., Just, M., & Carpenter, P. \(1994\) Working Memory constraints on the resolution of )S
.132 .132 scale
(lexical ambiguity: Maintaining multiple interpretations in neutral contexts. )S
(Journal of Memory )S
.132 .132 scale
500.217 16365.05 :M
(and Language, )S
.132 .132 scale
(, 175-202.)S
.132 .132 scale
(Pearlmutter, N. J., & MacDonald, M. C. \(1995\) Individual differences and probabilistic constraints )S
.132 .132 scale
(in syntactic ambiguity resolution.
(Journal of Memory and Language)S
.132 .132 scale
.132 .132 scale
(, 521-542.)S
.132 .132 scale
(Experiment 1)S
.132 .132 scale
(\245 Contrast two types of )S
.132 .132 scale
(\(Gilboy et al., 1995\) )S
.132 .132 scale
.132 .132 scale
Functional / Occupational)S
265 762 579 @c
.132 .132 scale
.132 .132 scale
(The crowds annoyed the chauffeur of the actor who wanted to )S
.132 .132 scale
(go home. )S
.132 .132 scale
Representational:)S
179 756 719 @c
.132 .132 scale
.132 .132 scale
(The artist recognized the sketch of the house that was mentioned )S
.132 .132 scale
(in the book.)S
.132 .132 scale
(\245 Free-choice questionnaire was used to ensure that subjects were )S
.132 .132 scale
(not alerted to alternatives, as opposed to forced-choice used in )S
.132 .132 scale
(previous studies.)S
.132 .132 scale
.132 .132 scale
(\245 10 functional, 10 representational sentences )S
.132 .132 scale
(\245 N1 and N2 were both animate or both inanimate and both )S
.132 .132 scale
(singular or both plural. )S
.132 .132 scale
(\245 N1 and N2 attachments were controlled for plausibility.)S
.132 .132 scale
(\245 Free-choice questionnaire: experimental items were followed by )S
.132 .132 scale
(a question about the attachment, with space for an answer:)S
.132 .132 scale
Who wanted to go home?_____________)S
.132 .132 scale
(\245 60 filler sentences \(each followed by a question\).)S
.132 .132 scale
(\245 Daneman and Carpenter \(1980\) reading span task classified )S
.132 .132 scale
(subjects as high span \(span >= 4\) or low span \(span < 4\).)S
.132 .132 scale
= Different from chance,)S
.132 .132 scale
( Error bars show 95% CI for cell mean differences)S
.132 .132 scale
.132 .132 scale
(Span Effect)S
121 759 2150 @c
.132 .132 scale
(: Low span subjects preferred N1 high )S
.132 .132 scale
(span subjects showed no preference. )S
.132 .132 scale
(\245 The )S
.132 .132 scale
( types did not differ and there was no interaction.)S
.132 .132 scale
(Experiment 2)S
.132 .132 scale
(\245 Examine )S
.132 .132 scale
( types along with other prepositions:)S
.132 .132 scale
(Locatives:)S
.132 .132 scale
(The tourists went to the store near the beach that was crowded )S
.132 .132 scale
(and hot. )S
.132 .132 scale
.132 .132 scale
(The personal assistant recommended the dress with the lace that )S
.132 .132 scale
(was imported from France.)S
.132 .132 scale
(\245 Locatives and )S
.132 .132 scale
( attach as adjuncts and not as arguments. )S
.132 .132 scale
(Theories )S
(\(e.g., Frazier & Clifton, 1996\))S
.132 .132 scale
( that use this distinction to )S
.132 .132 scale
(explain the difference between )S
.132 .132 scale
.132 .132 scale
( predict that locatives )S
.132 .132 scale
(will pattern with )S
.132 .132 scale
.132 .132 scale
(\245 10 locatives, 10 )S
items and 20 )S
.132 .132 scale
.132 .132 scale
( items from Exp 1.)S
.132 .132 scale
(\245 N1 and N2 attachments were controlled for plausibility.)S
.132 .132 scale
(\245 Same free-choice questionnaire method as Exp 1.)S
.132 .132 scale
(\245 61 filler sentences \(each followed by a question\).)S
.132 .132 scale
848.516 :M
= Different from chance)S
.132 .132 scale
121.393 :M
(Error bars show 95% CI for cell mean differences)S
.132 .132 scale
947.605 :M
(\245 Span Effect)S
.132 .132 scale
(: Low span subjects preferred N1 attachment more )S
.132 .132 scale
182.582 :M
(than high span subjects did. )S
.132 .132 scale
(\245 Preposition Effects:)S
.132 .132 scale
.132 .132 scale
( elicited N1 attachment preferences less )S
.132 .132 scale
690.437 :M
(than the other prepositions.)S
.132 .132 scale
963.314 :M
(\245 Interaction:)S
.132 .132 scale
963.314 :M
For High span subjects, functional )S
.132 .132 scale
963.314 :M
(f displayed the )S
.132 .132 scale
190.712 :M
(same low N1 attachment preference as )S
(. Low span subjects )S
.132 .132 scale
(did not show this effect.)S
854 1510 :M
currentpoint
currentpoint 3 -1 roll sub 557 div 3 1 roll exch sub 507 div exch
5 935 1926.5 @c
5 935 1891.5 @c
5 935 1856.5 @c
5 935 1820.5 @c
5 935 1784.5 @c
5 935 1749.5 @c
5 935 1714.5 @c
5 935 1678.5 @c
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5 935 1608.5 @c
5 935 1572.5 @c
355 940.5 1572 @w
371 940 1926.5 @c
371 940 1926.5 @c
.55 .56 .55 :F
.17 .16 .16 :F
9.5 74 227 rS
.55 .56 .55 :F
1.5 73 175 rS
.17 .16 .16 :F
8.5 73 228 rS
.913 .849 scale
.913 .849 scale
.913 .849 scale
993.202 2191.52 :M
.913 .849 scale
.913 .849 scale
.913 .849 scale
.913 .849 scale
.913 .849 scale
.913 .849 scale
.913 .849 scale
.913 .849 scale
:imsv 32 216 1 4 [29 0 0 182.692 859 1668] T T 1 T :f
:i FFF80000FFFFFFFE001FFFF
0FFFFFF80007FFF80007FFF80003FFFFFFF0001C
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0007FFF80007FFF80003FFFFFF80007FFFC01E0007FFF80007FFF0C0FF78C023FF00003DFE000
07FF000007FBCF00000%ADOeod
.913 .849 scale
.445(Functional)A
.913 .849 scale
.403(Representational)A
.913 .849 scale
7.95 .795(Preposition )J
.913 .849 scale
1.999(Type)A
.913 .849 scale
.17 .16 .16 :F
.034 .003(Experiment 1 Results)J
.17 .16 .16 :F
9.5 9 8 rS
.913 .849 scale
.55 .56 .55 :F
7.255 .725(Low Span \(N=57\))J
.55 .56 .55 :F
2.5 9 9 rS
.913 .849 scale
7.235 .724(High Span \(N=12\))J
np 939 1749 :M
np 939 1749 :M
939 1750 :L
941 1750 :L
941 1749 :L
939 1749 :L
np 943 1749 :M
943 1750 :L
944 1750 :L
944 1749 :L
943 1749 :L
np 946 1749 :M
946 1750 :L
948 1750 :L
948 1749 :L
946 1749 :L
np 950 1749 :M
950 1750 :L
952 1750 :L
952 1749 :L
950 1749 :L
np 954 1749 :M
954 1750 :L
955 1750 :L
955 1749 :L
954 1749 :L
np 957 1749 :M
957 1750 :L
959 1750 :L
959 1749 :L
957 1749 :L
np 961 1749 :M
961 1750 :L
963 1750 :L
963 1749 :L
961 1749 :L
np 964 1749 :M
964 1750 :L
966 1750 :L
966 1749 :L
964 1749 :L
np 968 1749 :M
968 1750 :L
970 1750 :L
970 1749 :L
968 1749 :L
np 972 1749 :M
972 1750 :L
974 1750 :L
974 1749 :L
972 1749 :L
np 975 1749 :M
975 1750 :L
977 1750 :L
977 1749 :L
975 1749 :L
np 979 1749 :M
979 1750 :L
981 1750 :L
981 1749 :L
979 1749 :L
np 983 1749 :M
983 1750 :L
985 1750 :L
985 1749 :L
983 1749 :L
np 986 1749 :M
986 1750 :L
988 1750 :L
988 1749 :L
986 1749 :L
np 990 1749 :M
990 1750 :L
992 1750 :L
992 1749 :L
990 1749 :L
np 994 1749 :M
994 1750 :L
996 1750 :L
996 1749 :L
994 1749 :L
np 997 1749 :M
997 1750 :L
999 1750 :L
999 1749 :L
997 1749 :L
.913 .849 scale
gS 5 129 rC
.913 .849 scale
.132 .132 scale
(Summary and Conclusions)S
.132 .132 scale
(\245 The difference between )S
.132 .132 scale
.132 .132 scale
.132 .132 scale
( cannot be explained by an )S
.132 .132 scale
(argument/ adjunct distinction because locatives \(also adjuncts\) )S
.132 .132 scale
(did not pattern with )S
.132 .132 scale
.132 .132 scale
(\245 High span subjects\325 lack of N1 preference in functional )S
.132 .132 scale
(s could )S
.132 .132 scale
(be due to an ability use the knowledge that in the N1 case, )S
.132 .132 scale
(another \(unambiguous\) genitive form \(e.g. )S
(the actor\325s )S
.132 .132 scale
(chauffeur)S
.132 .132 scale
(\) is available.)S
.132 .132 scale
Weak preferences in previous studies could be due to varying )S
.132 .132 scale
(ratios of high to low span subjects.)S
.132 .132 scale
.132 .132 scale
(Possible Explanations of Span Differences)S
.132 .132 scale
(\245 Tuning, Garden Path, Constural and Parameterized Head )S
.132 .132 scale
(Attachment )S
(\(Mitchell, 1994; Frazier and Clifton, 1996; Konieczny, et al., )S
.132 .132 scale
(1997\) )S
(do not predict this difference, but could be compatible, )S
.132 .132 scale
(given a mechanism to handle span differences. )S
.132 .132 scale
(\245 The Syntactic Prediction Locality Theory )S
(\(Gibson, 1998\) )S
.132 .132 scale
.132 .132 scale
(incompatible with these differences.)S
.132 .132 scale
(\245 A possible explanation for the span differences is that low span )S
.132 .132 scale
(subjects focus on the matrix verb and its arguments \(including )S
.132 .132 scale
(N1\) more than on elements deeper in the structure. So, N1 may )S
.132 .132 scale
(be more active than N2, and thus more likely to be chosen. )S
.132 .132 scale
(High span subjects may be able to keep all of the elements )S
.132 .132 scale
005.496 :M
(relatively active for attachment.)S
.132 .132 scale
285.953 :M
(\245 Planning on-line study to replicate findings and test above )S
.132 .132 scale
513.351 :M
(prediction.)S
.132 .132 scale
074.815 :M
(Plausibility Norming)S
.132 .132 scale
681.208 :M
(\245 N1 and N2 versions of the items used in both experiements)S
.132 .132 scale
075.364 :M
.132 .132 scale
(: the chauffeur who wanted to go home)S
.132 .132 scale
.132 .132 scale
(: the actor who wanted to go home)S
.132 .132 scale
250.252 :M
(\245 60 fillers, 62 subjects, scale:1-7 \(7= most plausible\))S
.132 .132 scale
636.828 :M
(\245 No significant differences in plausibility.)S
.132 .132 scale
220.482 :M
(Plausibility by Condition)S
.132 .132 scale
493.359 :M
(Item Type)S
.132 .132 scale
493.359 :M
.132 .132 scale
.132 .132 scale
773.816 :M
(Functional )S
.132 .132 scale
773.816 :M
(5.18 \(.47\))S
.132 .132 scale
(5.24 \(.82\))S
.132 .132 scale
046.693 :M
(Representational )S
.132 .132 scale
046.693 :M
(5.07 \(.49\))S
.132 .132 scale
(5.37 \(.43\))S
.132 .132 scale
327.151 :M
(Locative)S
.132 .132 scale
327.151 :M
(4.65 \(.85\))S
.132 .132 scale
327.151 :M
(5.05 \(.47\))S
.132 .132 scale
600.028 :M
.132 .132 scale
600.028 :M
(5.26 \(.43\))S
.132 .132 scale
600.028 :M
(5.06 \(.63\))S
.132 .132 scale
880.485 :M
(Note: standard deviations in brackets)S
.55 .56 .55 :F
.17 .16 .16 :F
.55 .56 .55 :F
.17 .16 .16 :F
.55 .56 .55 :F
.17 .16 .16 :F
.55 .56 .55 :F
.17 .16 .16 :F
.132 .132 scale
946.506 :M
.132 .132 scale
650.889 :M
.132 .132 scale
355.272 :M
.132 .132 scale
059.655 :M
.132 .132 scale
764.038 :M
.132 .132 scale
468.421 :M
.132 .132 scale
172.804 :M
.132 .132 scale
869.607 :M
.132 .132 scale
.132 .132 scale
278.373 :M
.132 .132 scale
:imsv 32 224 1 4 [37 0 0 238.126 ] T T 1 T :f
:i FFF80000FFFFFFFE001FFFF
0FFFFFF80007FFF80007FFF80003FFFFFFF0001C
F000FFF80000FFF000FFF80000FFFFC001FF3C00
0007FFF80007FFF80003FFFFFF80007FFFC01E0007FFF80007FFF0C0FF78C023FF00003DFE000
07FF000007FBCF%ADOeod
.132 .132 scale
(Functional)S
.132 .132 scale
075.364 :M
(Representational)S
.132 .132 scale
075.364 :M
(Locative)S
.132 .132 scale
075.364 :M
.132 .132 scale
287.602 :M
(Preposition Type)S
.132 .132 scale
.17 .16 .16 :F
(Experiment 2 Results)S
.17 .16 .16 :F
.132 .132 scale
194.994 :M
.132 .132 scale
194.994 :M
.55 .56 .55 :F
(ow Span \(N=53\))S
.55 .56 .55 :F
.132 .132 scale
210.154 :M
.132 .132 scale
210.154 :M
(igh Span \(N=20\))S
.132 .132 scale
172.804 :M
.132 .132 scale
923.216 :M
.132 .132 scale
043.945 :M
.132 .132 scale
665.499 :M
.132 .132 scale
975.726 :M
.132 .132 scale