title: A geographic cline induced by negative frequency-dependent selection.
abstract: BACKGROUND
  Establishment of geographic morph frequency clines is difficult to explain in organisms with limited gene flow. Balancing selection, such as negative frequency-dependent selection (NFDS), is instead suggested to establish a morph frequency cline on a geographic scale at least theoretically. Here we tested whether a large-scale smooth cline in morph frequency is established by NFDS in the female-dimorphic damselfly, Ischnura senegalensis, where andromorphs and gynomorphs are maintained by NFDS.
  
  
  RESULTS
  We found a large-scale latitudinal cline in the morph frequency: andromorph frequency ranged from 0.05 (South) to 0.79 (North). Based on the empirical data on the numbers of eggs, the number of ovariole, abdomen length and latitude, the potential fitness of andromorphs was estimated to be lower than that of gynomorphs in the south, and higher in the north, suggesting the gene-by-environment interaction. From the morph-specific latitudinal cline in potential fitness, the frequency of andromorphs was expected to shift from 0 to 1 without NFDS, because a morph with higher potential fitness wins completely and the two morphs will switch at some point. In contrast, NFDS led to the coexistence of two morphs with different potential fitness in a certain geographic range along latitude due to rare morph advantage, and resulted in a smooth geographic cline of morph frequency.
  
  
  CONCLUSIONS
  Our results provide suggestive evidence that the combination of NFDS and gene-by-environment interaction, i.e., multi-selection pressure on color morphs, can explain the geographic cline in morph frequency in the current system.
authors.affiliation: Division of Ecology and Evolutionary Biology, Graduate School of Life Sciences, Tohoku University, 6-3, Aoba, Aramaki, Aoba, Sendai, Miyagi 980-8578, Japan. takahashi.yum@gmail.com
authors.forename: Yuma
authors.initials: Y
authors.lastname: Takahashi
authors.forename: Satoru
authors.initials: S
authors.lastname: Morita
authors.forename: Jin
authors.initials: J
authors.lastname: Yoshimura
authors.forename: Mamoru
authors.initials: M
authors.lastname: Watanabe
mesh_terms: D000818:Animals; D049628:Body Size; D001837:Body Weights and Measures; D005260:Female; D005843:Geography; D007313:Insecta / Q000235:genetics* / Q000502:physiology*; D007564:Japan; D016014:Linear Models; D010063:Ovum / Q000166:cytology; D010858:Pigmentation / Q000235:genetics; D012641:Selection, Genetic; D012727:Sex Characteristics