Systematic conservation planning has a substantial theoretical underpinning that allows optimization of tradeoffs between biodiversity conservation and other socioeconomic goals. However, thistheory assumes perfect spatial information about the locations of biodiversity features (e.g., species distributions). In practice, planners represent well-known taxa and other biodiversity ''surrogates'' inprotected area systems, hoping that unmapped species will also be conserved. However, empirical research finds that surrogates predict species presence imperfectly, and sometimes rather poorly, at scales relevant toplanning, and existing theory provides no further guidance. We developed new theory, explicitly incorporating aspects of spatial scale, for the representation problem when the locations of speciesdistributions are unknown. Using probability theory and simulated and real species distributions, we found that the probability of adequately representing an unmapped species in a protected area system willbe low unless the total fraction of the region being protected is larger than the species representation target. Furthermore, successful conservation depended critically on the relative sizes of the species distributionand of the individual protected areas; fewer, larger protected areas allowed the entire species distribution to fall into an unprotected gap. This scale-dependence varied with the configuration of the protected areasystem, with the conservation objective most likely to be attained if the individual protected areas were hyperdispersed (evenly spaced across the planning region). Using these results, we developed three designprinciples for representing unmapped species in protected areas: (1) The fraction of the region placed in protected areas should be substantially larger than the species-level representation target; (2) Individualprotected areas must be at least one to two orders of magnitude smaller than the unmapped species' distribution; and (3) Protected areas should be evenly dispersed over geographic space. We also performedpreliminary investigations of the effects of surrogates and socio-economic cost data on the probability of adequately representing unmapped species, finding that the primary effect of surrogates may simply be topromote hyperdispersion of protected areas across the planning region, and that seeking to minimize opportunity costs gives poorer conservation results than random protected area placement.