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Diversity and Distribution of Ant Communities in Puerto Rico1

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Diversity and Distribution of Ant Communities in Puerto Rico1 Diversityand Distributionof Ant Communities in Puerto Rico1 Juan A. Torres Departamentode Biologia, Colegio UniversitarioTecnol6gico de Bayam6n, Bayam6n, Puerto Rico 00620, U.S.A. ABSTRACT I studiedants in upland tropicalforest, grassland and agriculturalland in San Lorenzo,Puerto Rico, to uncoverfactors responsible forthe distributionand numberof speciesin thesecommunities. Observations, laboratory studies and fieldexperiments were used. Microclimateinfluenced the distributionsof Pheidolefallax, Solenopsisgeminata and Monomoriumebeninum; forest ants had lower toleranceto high temperaturesthan ants fromnon-forested areas. In addition,Ponerinae ants did not toleratehigh temperatures as well as ants in the Formicinae,Myrmicinae and Dolichoderinae.The largernumber of speciesin non-forestedareas is correlated withlarger food overlaps,tighter guilds, a more variablemicroclimate, and higherlevels of aggressiveness. RESUMEN Estudielas hormigasen areasadyacentes de bosque tropical,pradera y zonas agricolasen San Lorenzo,Puerto Rico. Observaciones, estudios en el laboratorioy experimentosde campo fueronrealizados para tratarde descubrirlos factoresresponsables de la distribuci6ny el numerode especiesen estas comunidades.Encontre que las distribucionesde Pheidolefallax, Solenopsisgeminata y Monomoriumebeninum son altamenteinfluenciadas por el microclima.Las hormigasdel bosque toleranaltas temperaturasmenos que las hormigasde areasno forestales.Ademas, las hormigasde la subfamiliaPonerinae toleran altas temperaturasmenos que las que pertenecena las subfamiliasFormicinae, Myrmicinae y Dolichoderinae.El numeromayor de especiesde hormigasen areas no forestalesesta relacionadoal mayorsolapamiento alimenticio, gremios mas compactos,un microclimamas variabley nivelesmas altos de agresividad. Two PROBLEMS THAT ECOLOGISTS ADDRESS are the identifi- overlap,and microclimatediversity to determinethe con- cationof thosefactors that determine the distribution and tributionof thesefactors to thenumber of speciesin three numberof speciesin a community.Species distributions adjacent ant communities.Also, ant toleranceto high have been found to be affectedby climate,nutritional temperatureis analyzedto assess its contributionto ant requirements,predators, competitors and dispersalbar- speciesnumbers. A companionpaper discusses the subject riers(Andrewartha and Birch1954, Krebs 1972). Several of ant coexistencein thesecommunities (Torres 1984). factorshave been found to accountfor differences in species numbers.Paine (1966) and Harper (1969) found that predationon superiorcompetitors increases diversity by HABITATS AND ANTS keeping the weak competitoron the system.Connell (1978) suggestedthat disturbancesintermediate in fre- FromJuly 1978 to September1980, I studiedants in a quencyand intensityshould accountfor high diversity in subtropicalwet forest,grassland dominated by Chloris coral reefsand tropicalrain foresttrees. Increase in area radiata and agriculturalcrops in San Lorenzo, Puerto resultsin higherdiversity due to its effectin reducing Rico. Cropsconsisted of coffee(Coffea arabica), plantains extinctionrates (MacArthur and Wilson 1967). Greater (Musa paradisiaca), and yams (Dioscorea alata). These resourcediversity, smaller niche breadth,greater mean habitatswere located near Route 7740 (forest:Km 4.4; nicheoverlap, compact guilds, through their influence on grasslandand crops:Km 1.1-2.5). The tightforest can- speciespacking (Inger and Colwell 1977) and stochastic opy produceda microclimatecooler and wetterthan in phenomena(Sale 1977, 1982; Caswell 1978) have been thegrassland and plantain-yamscrops. Coffee groves and foundto be relatedto diversitypatterns. foresthad similarmicroclimates. The forestcontained 20 Througha seriesof experimentalfield studies, I tried ant species,the grassland37, and agriculturalland 38 to determinethe influenceof food competitionand mi- (see accompanyingpaper for a listof species).Thirty-four crodimateon the distributionof Pheidolefallax, Mono- species were shared by grasslandand agriculturalland, moriumebeninum and Solenopsisgeminata. In addition,I and ten were foundin all threehabitats. The same ants conductedanalyses of food diversity,food breadthand tendedto be foundin coffeegroves and forest.Excluding the coffee-grovesspecies, agricultural land species were I Received 3 February1983, final revision9 January1984, similarto thosefound in grassland.Iridomyrmex melleus accepted 16 February1984. and Myrmelachistaramulorum were found once on grass- 296 BIOTROPICA 16(4): 296-303 1984 This content downloaded from 166.4.81.212 on Thu, 25 Feb 2016 16:07:15 UTC All use subject to JSTOR Terms and Conditions land and mighthave been transientmembers of thiscom- I transferredfive additional nests from grassland to grass- munitysince they were usually found in shaded areas. land approximately250 m fromthe forestto controlfor Due to the insulareffect, the faunain thesehabitats movingeffects. Observations were made as in Monomo- is depauperate,and ants are an example.Ants foundon riumebeninum's experiments. The experimentended on the island are generallysmall. Representativesof larger April27, 1980 whenthe fruitswere opened to checkfor Camponotiof the maculatusand abdominalisgroups are live ants. The experimentwas repeatedfrom April 3 to absent(Wheeler 1908). Neotropicalgenera such as Atta, April27, 1980. The reasoningbehind these experiments Azteca and Eciton are also absent and Crematogaster,was the same as forthe M. ebeninumexperiments. Pseudomyrmexand Pheidole,well developed in tropical The ground-nestingPheidole fallax was almost im- mainland,are scarcein PuertoRico (Wheeler 1908). possibleto transplant.I constructeda shade in situ over P. fallax nestssimulating the forestcanopy. The shade produceda coolerand wettermicroclimate compared to controlunshaded nests. I drovefour wooden stakesat a METHODS distanceof 18 cm fromnest openings and fixeda 28 x FIELD EXPERIMENTS.-Monomortiumebeninum, Solenopsis 50 cm wire net coveredwith cloth to the stakes. Rain geminataand Pheidolefallax are grasslandand agricul- waterpercolation was normalthrough the shade and light turalland ants found in forestgrass-covered dirt roads intensitywas approximately50 percentof unshadedareas but not withinthe forest.I conductedexperiments to and approximately25 percenthigher than in the forest. determinethe factorsthat barredthese species from the Controlshad fourwooden stakesdriven at the same dis- forest. tancefrom the nestwith dothless wire nets. The experi- On July3, 1980 I introducedto the forestnine logs mentswere performed three times involving four nests as containingMonomortium ebeninum colonies to whichI fed experimentalunits and fouras controls,and were con- honeyand tuna fishevery other day. Six othernests were ductedin 1980 on 9 Feb-22 March, 19 Feb-27 April, introducedbut not fed.I transferredsix nestsfrom grass- and 5 July-26 Sept. Observationswere made weekly. land to anothergrassland 200 m fromthe forest as control formoving effects. I fed tuna fishand honeyto six ad- TOLERANCE TO 45?C.-Surface temperaturesof 450C and ditionalnests in the grasslandto determinetheir nutri- higherwere experienced by antsthat foraged during mid- tionaladequacy as ant food. I isolatedall fed nestsfrom day, and few ant speciesforaged in these temperatures. interspecificinterference by hangingthese fromtrees or Since 45?C representsa realistic,but potentiallylimiting fenceposts and applyingautomotive grease to thestrings. temperature,I studied ant toleranceto 45?C in an envi- Grease was applied to unfednests but in a mannerthat ronmentalchamber (Environator Model WL221). The allowed antsto leave and re-enter.I observednests every chamber'scover was replacedwith glass to enable obser- threedays and placed food nearunfed nests to checkfor vationof ants. I placed a minimumof ten ants at 45?C ant presenceand removedit when activitywas observed. in vials withperforated caps to allow air circulation.The The experimentended on September25, 1980 whenlogs timeit took 50 percentof the antsto assumea crouched weresplit open to searchfor ants. Presenceor absenceof positionwas taken as a measureof toleranceto 450C. I ant coloniesand theirconditions were recorded. If micro- replicatedeach experimentwith individuals from ten col- dimate is the factorbarring these species from the forest, onies formost species. fedand unfednests in the forestwould disappear;on the otherhand, if competitionfor food is the limitingfactor FIELD TEMPERATURES AND NICHE METRICS.-I measured fed nestsshould remainand prosperwhile unfed nests ground and vegetationtemperatures under shade and should die, disappearor not prosper. sunlightat randomlychosen times of day for the three Finally,four M. ebeninumnests were introducedto habitatsusing YSI telethermometersand with the help the forestand fed but not isolatedto studyinterspecific of two assistants.Information on food habits was ob- interactions.I brought three Solenopsis B (mainlya forest tainedby observingant nestsfor five minutes each hour. species) neststo the grasslandon a sunnyday and put Food that ants were carryingwas removedwith forceps themnext to threeM. ebeninumnests. Food was placed or a piece of adhesive tape and later identifiedin the betweenthe neststo studyinterspecific aggression. laboratory.In addition,gastric distention, which occurred On March 23, 1980 I introducedto the forestten when liquid food was brought,was recorded.To study coconutfruits containing Solenopsis geminata nests. Five microhabitatutilization, I put baits
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