Is it enemy‐free space? The evidence for terrestrial insects and freshwater arthropods

Abstract. 1 Enemy‐free space (EFS) was defined by Jeffries & Lawton (1984) as ‘ways of living that reduce or eliminate a species’ vulnerability to one or more species of natural enemies’. EFS has emerged in the literature as a significant niche‐moulding factor. However, the lack of consistency among the empirical studies as to how EFS should be defined, and what hypotheses should be tested in order to evaluate its relative importance, prompted us to review the literature and to propose a working definition that results in a general set of testable hypotheses. 2 To test the relative importance of EFS in structuring the communities of organisms, we propose a set of three falsifiable null hypotheses that must be tested sequentially and rejected. Ho1: The fitness of the organism in an original habit (e.g. on an original host plant) in the presence of natural enemies is equal to the fitness of the organism in that habit in the absence of natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in the presence of natural enemies is less than in the absence of natural enemies is necessary to demonstrate the importance of natural enemies. Ho2: The fitness of the organism in an alternative habit with natural enemies is equal to the fitness of the organism in the original habit with natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in the alternative habit with natural enemies is greater than that in the original habit with natural enemies is necessary to demonstrate that the alternative habit provides EFS. Ho3: The fitness of the organism in an alternative habit without natural enemies equals the fitness of the organism in the original habit without natural enemies. Acceptance of the alternative hypothesis that the fitness of the organism in an alternative habit without natural enemies is less than in the original habit without natural enemies is necessary to demonstrate the relative importance of EFS compared with other co‐occurring niche‐moulding factors such as competition or host nutritional quality. 3 We searched the literature and evaluated fifty‐three references (nineteen references to seventeen different terrestrial systems and thirty‐four references to twenty‐four different freshwater systems) to test our hypotheses. 4 Of the forty‐one systems examined, nineteen (46%) tested only for differences in vulnerability of the prey or host species between EFS and non‐EFS options (our Ho2); sixteen (39%) tested for the importance of natural enemies and the effectiveness of the alternative habit in providing EFS (our Ho1 and Ho2); and only ten systems (24%) tested for Ho1, Ho2 and the relative importance of EFS in the system as measured by fitness (our Ho3). 5 Of the systems that tested for EFS, sixteen of nineteen (84%), thirteen of sixteen (81%) and seven of ten (70%) showed evidence in support of the existence of EFS according to hypothesis Ho2 only, hypotheses Ho1 and Ho2, and our three working hypotheses, respectively. 6 These results indicate that very few studies have actually tested for the existence of EFS. Nevertheless, results from this limited number of natural systems suggest that EFS may be important in moulding the niches of arthropods. Because of the large number of claims for EFS in systems where none of the basic hypotheses were investigated, we suggest that authors test for EFS experimentally, be judicious in selecting articles to cite in support of EFS, and exert care in attributing it as a selective force in the evolution of arthropods in specific systems.