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And Community Structure studet reasol food , and ir comp fauna and S the re and I classr other unde paras Th simp SpeciesInteractions this r inter and CommunityStructure short from Ti Questions concerning the nature and importance of species interac- vari< tions have become increasingly controversial. In the mid-1960sit was tive thought that interspecific competition was the major organizing factor ond in many communities. Through the pioneering work of the late Robert and MacArthur and his students,many aspectsof the structure and dynam- com ics of bird communities seemedbest explained by competitive interac- ied r tions, which could be describedin a generalway by the Lotka-Volterra spec competition equations. This work, or at least its generality, has been ofn challengedon three levels. First, there is the question of how important species interactions are in general and whether the interactions are competitive, parasitic, predatory, or mutualistic. For example, Connell whi (1975)noted that a significantfraction of, but by no means all, commu- nities are physically rather than biotically controlled. In addition, there As has been a tendency in the last few years to concentrate on single- dyn species demography. While interactions between species can still be the considered in the guise of age-specificfecundity and mortality effects, (va the emphasisis away from a coevolutionary perspective and toward a hos individualistic concept of communities. the Second, the question of which type of interaction is most important sor in structuring a particular community is very much open. Many ecol- tha ogists have pointed out that the birds studied by MacArthur and his wit SPECIESINTERACTIONS 97 studentswere at or near the top of the food chain. There is no a priori reasonto expect that competitionwill also be important lower in the food chain. Thus Connell arguesthat predationis a more widespread and important interaction.Furthermore, some of the best examplesof competition, such as character displacementin the Gal6pagosavi- fauna,are probablynot examplesof competitionat all (Strong,Szyska, and Simberloff 1979).Until very recently the debate revolved around the relative importanceof competitionand predation.However, Risch and Boucher(1976) and Price (1980)have forcefully arguedthat whole classesof interactionshave been ignored.Risch and Boucher, among others, have argued that mutualistic interactionsmay be the key to understandingcommunity structure, while hice arguesthe same for parasitism. Third, there has been widespreaddisillusionment with the utility of simple mathematicalmodels to describespecies interactions. In part, this disillusionmentstems from the very real difficulties in measuring interaction coefficients and from the failure of almost all suggestecl shortcutsto measuringinteraction coefficients.But in part it comes from the failure of the modelsto predict observedpatterns (Neill 1974). Thesethree questionswill be consideredthroughout this chapter in l- variousways. The first sectionreviews what is known about the rela- IS tive importanceof differentinteractions in cavecommunities. The sec- )r ond sectionconsiders in somedepth the beetle-cricket egginteraction, rt and the third sectionreviews in detail the interactionsin cave stream :l- communitiesin the southernAppalachians, the most thoroughly stud- :- ied case. The questions 'a concludingsection suggeststhat most about speciesinteractions are ill posedand arguesfor the central importance n of models. It 'e ll l- Which lnteractions Are lmportant? 'e As Price (1980)points out, the role of parasitesin the structure and dynamicsof most communitieshas been greatly underestimated. Even e the inventory of parasites of cave organisms is very incomplete ;, (Vandel 1964),and there are almost no data on the percentageof the a host populationinfected or the effect of the parasiteon the host. Even the meager data available suggestthat parasites may be important in It some communities.Keith (1975)found that almost all Pseudanoph- thalmustenuis beetles in Murray SpringCave in Indianawere infected S with the fungal parasiteLaboulbenia subterranea, with an averageof CAVELIFE up to fifteen infestations per beetle. The effect on the beetles of these Table 6-1 symbionts on the integument is not known, and in fact they may be .,nship (,1' ' commensals rather than parasites. Extremes of specializalion, even for !]ns()r; - the sma parasites, occur in caves. One of the most spectacularexamples are l)irta frofi the Temnocephala, parasitic platyhelminth worms intermediate in morphology between turbellarians and trematodes, which parasitize European cave shrimp. Matja5id (1958)reported that sevenspecies and several genera of Temnocephala are found only on the cave shrimp \ pecies Trogoc'uris sc'hmittti, with each species specializing on a particular t' (()llecte region of the body. For example, Srthtelsortictperiunalis is found i rtertrtis around the anus of T. schmitlti. 'tr(otle(I( A similar level of ignorance obtains for mutualistic interactions. No re stii free-living mutualists have been reported from caves, but ectosym- '.,uttb(tru! bionts that are probably mutualistic or commensal are known. Hobbs t:.-LIDtburtt! (1973,1975)studied the entocytherid ostracodsthat live on the exoskel- etons of cave crayfish. They feed on microorganisms and on detritus that accumulateson the host exoskeleton and are unable to complete their life cycle away from their host. Crayfish probably derive some ad- vantagefrom the cleaning activity of the ostracods, but the main effect spe of the interaction is benefit to the ostracods. Hobbs compared the os- spe tracod symbionts of the cave-limited Orc'onectesinermis to those of det the facultative cave dweller Cumborus luey,is.Almost all the ostracods dis onO. inermis were Sugittocytherebarri , which is rarely found on other the speciesin Hobbs'study area. In contrast, C. luevis commonly har- rat bored three species.Infestation and reinfestationoccur when the hosts fec copulate, when ostracod eggsbecome attached to newly hatched cray- sio fish carried under the abdomen and, following a molt, when the exu- col viae are eaten by the crayfish. Levels of infestation are lower in the sul cave-limited species, but this is complicated by the strong effect of lio crayfish size on the number of ostracod infestations(Table 6- 1). Since SO the cave-limited speciesare smaller, l'ewer ostracods per crayfish are int expected. Regressionanalysis indicates that a larger minimum size is an required for infestation of the cave-limited species than of the faculta- VC tive cave dwellers, but that the rate of infestation increasesmore rap- ca idly with size in troglobitic species.This may be a consequenceof the OC specializationof S. borri onOrconectes inermis, but the adaptive sig- nificance, if any, is not clear. m Christiansen and Bullion (1978)attempted to assessthe importance pt of competition and predation for terrestrial cave fauna in the Haute- m Garonne and Aridge regions of France. Their basic procedure was to fa visually census for 100 minutes the populations of about fifty terrestrial si SPECIESINTERACTIONS oo these Table 6-1 Numbers of entocytherid ostracodsinhabiting various crayfish, and the rela- ':onship ay be (y -- o * bx) between crayfish carapace length (x) and number of ostracod infesta- 3nfor ::ons ()'); b is the slope and a is the intercept of the regressionequation. Minimum size )s are . the smallest crayfish expected to harbor ostracods, based on the regression analysis. .te in [)ata from Hobbs 1973, 1975.) sitize t ('r 95% s and Ecological confidence Minimum rrimp \ pecles Habitat range interval) a b size (mm) cular )r(()ltectes + -28 bund inermis Cave Cave-limited 18.8 3.6 1.95 15 trtertnis )r( onectes itterntis Cave Cave-limited 20.2 + 13.6 -45 2.80 16 r. No I t'stii 'sym- ' tntburus lueyis Cave Facultative 26.5 + 3.7 6.2 0.60 0 .obbs cave-dweller skel- I Lutlborusbartonii Surface Occasionally 119 + 17.5 Lritus ln caves plete e ad- :ffect species and to estimate various environmental parameters, such as dry e os- speleothems, calcareous clay, guano, and standing crop of organic se of debris, for fifty-eight caves. Some classification of caves was made, cods distinguishingunderground rivers, vertical sinks, and so on, as well as rther the aphotic and entrance zones. Each environmental parameter was har- rated on a scale of I to 6. They then attempted to determine what af- IOSTS fected the abundance of various species by stepwise multiple regres- :ray- sion techniques. Their study was almost exclusively a between-cave exu- comparison and did not detect all interactions, such as competition re- t the sulting in microhabitat separationwithin a cave. Christiansenand Bul- lt of lion themselvespointed out some statisticalweaknesses and noted that lince some climatic variables were not measured, but their study did provide r are insights into the importance of species interactions. There were ex- ze is amples of apparent competitive exclusion between troglobitic omni- ulta- vore and troglophilic carnivore beetles but not among troglobitic rap- carnivores or troglophilic omnivores. That is, competitive exclusion I the occurred but was infrequent. sig- Collembola specieswere analyzedmore completely. Table 6-2 sum- marizes the results for three cave-limited Collembola: Tomocerus tnce problematicus, Pseudosinella theodoridesi, and P. r,rrer. Most of the ute- major negative correlates are other species rather than physical sto factors, and most are probably competitors rather than predators.
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