structure and dynamics of an arboreal-nesting termite community...
TRANSCRIPT
Structure and dynamicsStructure and dynamicsof an arboreal-nestingof an arboreal-nestingtermite community intermite community in
New-GuineaNew-Guineacoconut plantationscoconut plantations
Why a semi-natural habitat ?Why a semi-natural habitat ?
New-Guinea coconut plantationsNew-Guinea coconut plantations
largely occupiedlargely occupiedby arboreal-by arboreal-nesting termitesnesting termites
interspecificinterspecificexclusionexclusion
variousvariousplantationplantationpatternspatterns
Community compositionCommunity compositionhalf of the trees are occupied by termiteshalf of the trees are occupied by termites
M. biroi M. biroi : : N. princeps N. princeps : : N. novarum-N. novarum-hebridarum hebridarum = = 8080 : : 1414 : : 66
Nasutitermes novarum-hebridarum
Nasutitermesprinceps
Microcerotermes biroi
% trees
no termites
termites
Ressources exploitedRessources exploited
All 3 species useAll 3 species usethe coconutthe coconuttree as nestingtree as nestingsite and foodsite and foodsourcesource
dead palm leave covered with galleries
Goals:Goals:
Community organization:Community organization:
Which ecological factors affect theWhich ecological factors affect thedistribution and abundance of termites ?distribution and abundance of termites ?
How do coexist agonistic colonies ?How do coexist agonistic colonies ?
What is the territoriality of theseWhat is the territoriality of thesearboreal-nesting termites ?arboreal-nesting termites ?
OverviewOverview
Biology of the 3 speciesBiology of the 3 species
Response to the physical environmentResponse to the physical environment
Interspecific interactions betweenInterspecific interactions betweencoloniescolonies
Intraspecific interactions betweenIntraspecific interactions betweencoloniescolonies
PredationPredation
Biology of the 3 speciesBiology of the 3 species
mandibulate soldiers, 3% of mandibulate soldiers, 3% ofthe castesthe castes
29 nests with reproductives on29 nests with reproductives on1 ha1 ha
trees with: 1 nest (66%), 2trees with: 1 nest (66%), 2nests (27%), 3 nests (7%)nests (27%), 3 nests (7%)
1 colony extends on 1-3 trees1 colony extends on 1-3 trees
only 1 nest with reproductives,only 1 nest with reproductives,other nests = satellite nestsother nests = satellite nests
N. princeps galleries onlyN. princeps nest and galleriesroyal cell without reproductivesroyal cell with reproductivesM. biroi galleries onlyM. biroi nest and galleries
Biology of Biology of Microcerotermes biroiMicrocerotermes biroi:morphology, colony organizationmorphology, colony organization
0 10 20 30 40 50 60 70 80 90 100
ME
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100North
METERS
84% of colonies84% of coloniesheaded byheaded bydealated imagosdealated imagos
orphaningorphaningexperiments:experiments:nymphoidsnymphoids(neotenics arised(neotenics arisedfrom nymphs),from nymphs),ergatoidsergatoids(neotenics arised(neotenics arisedfrom workers),from workers),NEVER adultoidsNEVER adultoids(alates)(alates)
Reproductives found in field coloniesReproductives found in field colonies
Biology of Microcerotermes biroi
Royal cell containing 1 imaginal queen
16% of colonies16% of coloniesheaded byheaded bypolygynouspolygynousassociations ofassociations ofneotenics:neotenics:nymphoids,nymphoids,ergatoidsergatoids(replacement /(replacement /budding)budding)
reproduction:reproduction:mainly by mainly by nuptialnuptialflightsflights, buddingbuddingpossible but rarepossible but rare
nymph
queen
king
worker
5 mm
Nymphoids
Reproductives found in field coloniesReproductives found in field colonies
Biology of Microcerotermes biroi
NestNestcasemate-like structurecasemate-like structure
Biology of Microcerotermes biroi
5 mm
chambers communicate by small orifices10 cm
worker
nasute soldiersnasute soldiers(>10% of the(>10% of thecastes)castes)
pheromonal actionpheromonal actionof the soldiersof the soldiersdefensivedefensivesecretionsecretion
open nest structureopen nest structure
Biology of Biology of Nasutitermes princepsNasutitermes princeps
1 mm
60% of colonies60% of coloniespolygynouspolygynous
neotenics:neotenics:adultoids oradultoids orpseudimagospseudimagos
long and shortlong and shortwinged alateswinged alates(microimagos)(microimagos)
reproduction byreproduction bybuddingbudding
Biology of Biology of Nasutitermes princepsNasutitermes princeps
5 mm
microimago
adultoid queens
nasute soldiersnasute soldiers
no pheromonalno pheromonalaction of soldiersaction of soldiersdefensive secretiondefensive secretion
Biology of Biology of NasutitermesNasutitermesnovarumhebridarumnovarumhebridarum
workers and soldiers
often on deadoften on deadtreestrees
open nestopen neststructurestructure
neotenics:neotenics:ergatoidsergatoids
colonies alwayscolonies alwaysmonogynousmonogynous
reproduction byreproduction byswarming flightsswarming flights
Biology of Biology of NasutitermesNasutitermesnovarumhebridarumnovarumhebridarum
imaginal queen
Environmental influencesEnvironmental influences
Environmental influences:Environmental influences:1. How community composition vary among 3 types of plantations ?1. How community composition vary among 3 types of plantations ?
OPENOPEN STANDARDSTANDARD SESEDENDEN
PlantationPlantation squaresquare sq. / trianglesq. / trianglepatternpattern 10 x 10 m10 x 10 m 8 - 9 m8 - 9 m
Tree heightTree height 25 -30 m25 -30 m +/- 13 m+/- 13 m 13 - 13 - 25 m25 m
Tree densityTree density < 100< 100
variablevariable
100 - 180100 - 180 180180 - 3 - 30000(tre(trees/ha)es/ha)
shade, food and nesting sites availability increases
Proportion of trees occupied byProportion of trees occupied bytermitestermites
Environmental influences:Environmental influences:1.How community composition vary among 3 types of plantations ?1.How community composition vary among 3 types of plantations ?
TREES / HECTARE
0 100 200 3000.0
0.2
0.4
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0.8
1.0
AR
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BY
TYPE OF COCONUT PLANTATION:DENSESTANDARDOPEN
Proportion of trees occupied by each termite speciesProportion of trees occupied by each termite species
Environmental influences: Environmental influences: 1. How community composition vary among 3 types of plantations ?1. How community composition vary among 3 types of plantations ?
OP
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Environmental influence on nest volumeEnvironmental influence on nest volume
Environmental influences: Environmental influences: 1. How community composition vary among 3 types of plantations ?1. How community composition vary among 3 types of plantations ?
Influence of direct sunlight on nest temperatureInfluence of direct sunlight on nest temperatureEnvironmental influences: Environmental influences: 2. Effect of environmental variables on termites ?2. Effect of environmental variables on termites ?
Microcerotermes biroi nestfullsun
pale
clouds
sun
HOUR OF THE DAY
0 6 12 18 0 6 12 18 0 6 12 18 0
TE
MP
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CE
LS
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)
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cloudsrainheavy
0 6 12 18 0 6 12 18 0 6 12 18 0TE
MP
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20
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sunAPR 28 APR 29 APR 30 1994
APR 21 APR 22 APR 23 1994
nestNasutitermes princeps
Tree heightTree height
Tree inclinationTree inclination
Tree conditionTree condition-- N. novarumhebridarum N. novarumhebridarum is 4 times moreis 4 times more
abundant on dead trees than on live onesabundant on dead trees than on live ones
RaintracksRaintracks-- M. biroi M. biroi tends to build its nests outside oftends to build its nests outside of
raintracksraintracks
Environmental influences: Environmental influences: 2. Effect of environmental variables on termites ?2. Effect of environmental variables on termites ?
Other variablesOther variables
Interspecific interactionsInterspecific interactions
Colony sizesColony sizes
100METERS
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ME
TE
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+
North
N. novarum-hebridarumN. princepsM. biroi
Interspecific interactionsInterspecific interactions
Interspecific encountersInterspecific encounters
AlwaysAlwaysaggressiveaggressivebehavioursbehaviours
Interspecific interactionsInterspecific interactions
N. princepsN. princepssoldiersoldier
N. novarum-N. novarum-hebridarumhebridarumsoldiersoldier
N. princepsN. princepsworkerworker
N. novarum-N. novarum-hebridarumhebridarumworkerworker
ME
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+
1990 54 trees with M. biroi218 trees with N. princeps 5 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi1. Natural expansion of 1. Natural expansion of N. princeps N. princeps coloniescolonies
ME
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+
1991 42 trees with M. biroi246 trees with N. princeps 5 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi1. Natural expansion of 1. Natural expansion of N. princeps N. princeps coloniescolonies
ME
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1992 15 trees with M. biroi291 trees with N. princeps 6 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi1. Natural expansion of 1. Natural expansion of N. princeps N. princeps coloniescolonies
ME
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+
+
1990 54 trees with M. biroi218 trees with N. princeps 5 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi1. Natural expansion of 1. Natural expansion of N. princeps N. princeps coloniescolonies
150
1990: current and past occupation by M. biroi
Plain symbols = trees occupied in May 1990Open symbols = trees occupied before May 1990
ME
TR
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50
100
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi1. Natural expansion of 1. Natural expansion of N. princeps N. princeps coloniescolonies
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198432 trees with M. biroi68 trees with N. princeps02 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
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198965 trees with M. biroi30 trees with N. princeps02 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
0 10 20 30 40 50 60 70 80 90 100
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199058 trees with M. biroi17 trees with N. princeps01 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
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199177 trees with M. biroi16 trees with N. princeps01 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
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199287 trees with M. biroi24 trees with N. princeps00 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
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+199394 trees with M. biroi29 trees with N. princeps00 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
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+ 1994107trees with M. biroi30 trees with N. princeps00 trees with N. novarum- hebridarum
Large symbols = nestsSmall symbols = galleries
Dynamics of territorial interactions between Dynamics of territorial interactions between N. princeps N. princeps and and M. biroiM. biroi2. Experimental weakening of 2. Experimental weakening of N. princeps N. princeps coloniescolonies
Occurence of direct interspecific contestsOccurence of direct interspecific contests
Offensive strategy of Offensive strategy of N. princepsN. princeps
M. biroi gallery under siege
N. princepsN. princeps
M. biroiM. biroi
M. biroiM. biroi: dynamics of colonization: dynamics of colonizationM
ET
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area
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1990
ME
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area
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1992
10 % of the trees occupied by M. biroi
M. biroiM. biroi: dynamics of colonization: dynamics of colonization
ME
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area
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1993
40 % of the trees occupied by M. biroi
M. biroiM. biroi: dynamics of colonization: dynamics of colonization
ME
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area
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1994
63 % of the trees occupied by M. biroiM. biroi is a pionneer species
M. biroiM. biroi: dynamics of colonization: dynamics of colonization
Intraspecific interactionsIntraspecific interactions
agonism in 66-79% ofagonism in 66-79% ofpaired combinationspaired combinationstestedtested
group of 112 anagonistgroup of 112 anagonistN. princeps N. princeps nests:nests:supercolonysupercolony
no seasonal variationno seasonal variationof agonismof agonism
100 150
0
50
100
150A2
ANAGONISMANAGONISM
Intraspecific interactions
LaboratoryLaboratorybioassaysbioassays
AA BB
AA BB
50 cm
results inresults insemi-naturalsemi-naturalconditionsconditionsconsistentconsistentwith resultswith resultsin laboratoryin laboratory
Intraspecific interactions
Confrontation ofConfrontation ofentire nestentire nestpopulationspopulations
1 m
AA BB
50 cm
trailtraildivergencedivergencebetweenbetweenagonistagonistcoloniescolonies
trail fusiontrail fusionbetweenbetweenanagonistanagonistcolonies incolonies inlaboratorylaboratory
Intraspecific interactions
PredatorsPredators
Inquiline antsInquiline ants-- occasional predatorsoccasional predators
-- in 10% of in 10% of M. biroi M. biroi nestsnests
-- in 5% of in 5% of N. princeps N. princeps nestsnests
-- may coexist several years with the termitesmay coexist several years with the termites
Arboreal-nesting antsArboreal-nesting ants-- In spots heavily occupied byIn spots heavily occupied by
CrematogasterCrematogaster sp. A: termites 2x less sp. A: termites 2x lessabundantabundant
-- Coconut plantation: marginal habitat forCoconut plantation: marginal habitat forCrematogasterCrematogaster sp. A sp. A
Predators
ConclusionsConclusions
Coexistence of the 3 speciesCoexistence of the 3 species
Similar feeding habits and nesting sitesSimilar feeding habits and nesting sites
Asymmetries between the reproductiveAsymmetries between the reproductiveand defensive mode of the 3 species:and defensive mode of the 3 species:
M. biroi M. biroi invests in alate production, longinvests in alate production, longdistance colonizerdistance colonizer
N. princeps N. princeps invests in aggressivity, shortinvests in aggressivity, shortdistance colonizerdistance colonizer
N. novarumhebridarum N. novarumhebridarum specialized inspecialized inthe exploitation of dead trees, effectivethe exploitation of dead trees, effectivelong distance colonizerlong distance colonizer ./../.
Why does N. princeps rarely becomes theWhy does N. princeps rarely becomes themost abundant species in the habitat ?most abundant species in the habitat ?
BuddingBudding
Requires dense habitat to develop largeRequires dense habitat to develop largecoloniescolonies
Coconut trees: unsuitable support forCoconut trees: unsuitable support forthe very large neststhe very large nests
Habitat preemption by Habitat preemption by M. biroiM. biroi
Coexistence of the 3 speciesCoexistence of the 3 species
Factors affecting termiteFactors affecting termitedistribution and abundancedistribution and abundance
Habitat structureHabitat structure
Arboreal antsArboreal ants
Interspecific competitionInterspecific competition
Intraspecific competitionIntraspecific competition
Inquiline antsInquiline ants
Factors affecting termiteFactors affecting termitedistribution and abundancedistribution and abundance
Habitat structureHabitat structure
Interspecific competitionInterspecific competition
Intraspecific competitionIntraspecific competition
Arboreal antsArboreal ants
Inquiline antsInquiline ants
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
5050M
ETR
ESM
ETR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
5050M
ETR
ESM
ETR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo
Small territorySmall territory
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
5050M
ETR
ESM
ETR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo
Small territorySmall territory
N. princepsN. princepsPolygynousPolygynous FrequentFrequent LL MobileMobile YesYes
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
5050M
ETR
ESM
ETR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo
Small territorySmall territory
N. princepsN. princepsPolygynousPolygynous FrequentFrequent LL MobileMobile YesYes
Large territoryLarge territory
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
5050M
ETR
ESM
ETR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo
Small territorySmall territory
N. princepsN. princepsPolygynousPolygynous FrequentFrequent LL MobileMobile YesYes
Large territoryLarge territoryXLXL
SupercoloniesSupercolonies
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
0
50
ME
TR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo**
idem: idem: N. ephratae, N. ephratae, * variable composition* variable composition Small territorySmall territory
N. princepsN. princepsPolygynousPolygynous FrequentFrequent LL MobileMobile YesYes**
idem: idem: N. costalisN. costalis, * stable composition, * stable composition Large territoryLarge territoryXLXL
idem:idem: N. polygynus, N. corniger N. polygynus, N. corniger SupercoloniesSupercolonies
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
0
50
ME
TR
ES
Ter
ritor
ialit
yT
errit
oria
lity
M. biroiM. biroiMonogynousMonogynous RareRare S / MS / M StaticStatic ----
Small territories, iSmall territories, interdigitatenterdigitate
N. novarumhebridarumN. novarumhebridarumMonogynousMonogynous NeverNever MM MobileMobile NoNo**
idem: idem: N. ephratae, N. ephratae, * variable composition* variable composition Small territorySmall territory
N. princepsN. princepsPolygynousPolygynous FrequentFrequent LL MobileMobile YesYes**
idem: idem: N. costalisN. costalis, * stable composition, * stable composition Large territoryLarge territoryXLXL
idem:idem: N. polygynus, N. corniger N. polygynus, N. corniger SupercoloniesSupercolonies
Territory: related to colony sizeTerritory: related to colony size+ to reproductive and defensive strategies+ to reproductive and defensive strategies
QueenQueennumbernumber
BuddingBudding ColonyColonysizesize
WarfareWarfare PheromonalPheromonalaction of S FGSaction of S FGS
0
50
ME
TR
ES
