PROFILE

Profile of Ilkka A. Hanski

atching birds and collecting largest of the Åland Islands in the northern butterflies in the field behind Baltic Sea, just off the southwestern corner Whis boyhood home, young of Finland. The 1,500-km2 landscape is Ilkka Hanski marveled at the a mosaic of surface bedrock, forest, small isolated habitats in which many insects live. villages, and a network of approximately Decades later, the Director of the Meta- 4,000 dry meadows that represent the population Research Group at the Uni- butterfly’s naturally fragmented habitat— versity of in Finland admits that ideal for Hanski’s studies. many of his most successful research The butterfly itself is a researcher- projects on the study of metapopulations, friendly organism, says Hanski; it is easy to or isolated yet interconnected populations breed, keep in the laboratory, and study in of the same species, were inspired by the field. In the case of the Glanville childhood adventures in his own back fritillary, the female lays eggs in large yard. Hanski, elected to the National Ilkka A. Hanski in the field. clutches. The caterpillars remain as a Academy of Sciences (NAS) in 2010, has group and spin a silken web around the helped predict how plant and animal community of dung beetles, small host plant. The rather conspicuous webs populations respond when their habitats resourceful insects that inhabit very iso- enable Hanski and his team to conduct are broken into fragments, whether by lated habitats: cattle dung pats. He was an accurate census of the large meta- urbanization, deforestation, climate population, which consists of tens of intrigued by the observation that most fl change, or entirely natural processes. His species of beetle clumped together, with thousands of butter ies in a network of studies have earned him the 2011 Cra- particular species common in some dung 4,000 meadows. To perform the census, foord Prize in Biosciences from the Royal Hanski employs an “army” of 70 under- pats but scarce or absent in others. “ Swedish Academy of Sciences, as well as During graduate school, Hanski be- graduates every autumn. It is a very big election to the Royal Society, the Finnish operation,” he says. came intrigued by mathematical popu- ’ Academy of Science and Letters, and most lation models and now had an interesting Hanski s studies remain a blend of recently, the NAS. laboratory experiments, modeling, and question to study: Could the spatial dis- fi Before the 1970s, ecologists paid little tribution of beetle species in the dung pats large-scale observational eldwork, pow- attention to the spatial structure of pop- on a pasture explain the coexistence of ered in large part by the students tasked ulations, whether they were continuously ecologically similar species within a com- with counting and sampling the larval distributed in space, assemblages of local families. Some of the experiments on munity? His models indicated that co- fl populations, or a mix of both. In 1969, existence was bolstered when individuals laboratory-reared butter ies are per- American population biologist Richard of the same species formed small clusters formed in large outdoor cages measuring fi — Levins rst introduced the concept of a throughout a landscape. Moreover, Han- up to 900 square meters about the same — metapopulation. However, it was Hanski ski realized that if individuals are not area as a single meadow. fi who has since led the eld of metapopu- extremely mobile and good at dispersing, Predicting Extinctions lation through the development of geographically separate populations may fi basic concepts, models, and informative lose the ability to interact. A physically The combination of eldwork and mod- empirical examples—who helped gauge the fragmented landscape at larger scale will eling led Hanski to craft some of his most fl prospects of long-term survival for species further amplify the spatial structure of in uential papers, including one published Nature living in fragmented landscapes. populations and lead to genuine meta- in in 2000 that described how Hanski’s mathematical models have re- populations, or networks of local pop- species respond to fragmentation (1). vealed how a collection of many small ulations, he mused. That pivotal study explained how a meta- local populations survive in a network of population can persist when a balance is habitat fragments. For the past 2 decades, Glanville Fritillary struck between local extinctions and re- “ Hanski has used the Glanville fritillary After receiving his Ph.D. from the Uni- colonization of vacant habitat. The butterfly as a study system to fuel his versity of Oxford in 1979, Hanski contin- question, then, is how the structure of the models and to probe the ecological, ge- habitat—the network of habitat patches, ued to develop simple mathematical — fl netic, and evolutionary consequences of models to predict the conditions for or fragments if you like in uences the habitat fragmentation. To date, Hanski’s processes of local extinction and coloni- metapopulation survival. By the late 1980s ” ’ models have been used to predict the dy- he was ready to test his predictions in the zation, he says. Hanski s model demon- namics of animal and plant populations field. Hanski struggled with the details— strated that landscape properties such as and to guide habitat conservation in the — the number and spatial distribution of which insect to study, and where until fl United States, Canada, Australia, Brazil, a fortuitous visit by renowned conserva- habitat patches in uence extinction and United Kingdom, Sweden, Finland, and colonization, ultimately determining tion and population biologist Paul Ehrlich ’ many other European countries. from Stanford University. During the a species viability. The model provides visit, Ehrlich described his research on a recipe for calculating the metapopu- fl Chasing Butter ies and Beetles Edith’s checkerspot butterfly(Euphydryas lation capacity, essentially the carrying Hanski says he cannot remember the editha). Moved by Ehrlich’s experience capacity of a fragmented landscape for a moment he first became interested in the ’ particular species. and Hanski s own teenage exploits, he ’ spatial structure of populations; it was selected the Glanville fritillary (Melitaea However, what makes Hanski s work so simply a part of life growing up in Tampere cinxia), a black, brown, and orange important is that it goes beyond in southern Finland and especially, during checkered relative of the Californian the summers, at his grandmother’s coun- checkerspot (Euphydryas editha bayensis). tryside home. So it was no surprise that, as Although the Glanville fritillary lives in fi ’ This is a Pro le of a recently elected member of the Na- a graduate student at the University of meadows across Europe, Hanski s meta- tional Academy of Sciences to accompany the member’s Oxford, his doctoral thesis focused on the population occupies a specific niche in the Inaugural Article on page 14397 in issue 35 of volume 108.

www.pnas.org/cgi/doi/10.1073/pnas.1117176108 PNAS | December 13, 2011 | vol. 108 | no. 50 | 19865–19866 Downloaded by guest on September 26, 2021 butterflies. Although his mathematical actions, he found himself guided back to of the butterfly, including their metabolic models were developed through fieldwork his former passion, the dung beetle. In rate during flight,” says Hanski. Among and experiments with the Glanville fritil- 2002, eager to rescue his long-lost pro- the best known is a gene that encodes an lary, the results can be tentatively applied ject, Hanski launched a large-scale study enzyme called phosphoglucose isomerase. to other species and landscapes. “What if of dung beetle evolution in Madagascar. In his Inaugural Article (5), Hanski re- human land use causes further fragmen- The large island off the southeastern veals that a variant of the phosphoglucose tation of a habitat? What if some frag- coastofAfrica,isolatedfromother isomerase gene can increase flight me- ments of the habitat are completely lost?” continents for an estimated 100 million tabolism by 25%, enabling a butterfly with he asks. “What are the likely con- years,ishometoahugenumberof the variant to fly twice as far as indi- sequences for the species?” Hanski’s unique plants and animals—including viduals that lack the variant. “Here is an models attempt to answer these ques- dung beetles—that have evolved in- example of a gene that has a big influence tions, and more. dependently from mainland species. on the performance of individuals in the Hanski and his students are currently field,” Hanski says. “Because flight ca- Predator and Prey attempting to reconstruct the evolution pacity is such a critical feature of life in While working on the Glanville fritillary of these species on an island where only fragmented landscapes, the gene variant in the 1990s, Hanski became interested in 10% of the natural forest cover remains. plays a key role in the establishment of a question that had stumped ecologists Because the beetles are largely forest new populations and, therefore, the dy- for decades. Every 4 or 5 years, the dwellers, knowledge of their ecology and namics of the entire metapopulation.” population densities of Arctic lemmings distribution is vital to establishing effec- As his team sequences the Glanville and boreal voles peak, then plummet. tive conservation measures. fritillary genome, they remain on the “The cyclical change in population Although the project differs dramati- lookout for genes like phosphoglucose numbers is striking,” Hanski says. “The cally from his studies of metapopulations, isomerase that impact the demographics question is, why?” Over the past 100 Hanski enjoys the opportunity to focus on of populations and the metapopulation. years, ecologists in North America, Eu- the spatial aspects of evolution, including Hanski’s team analyzes large numbers of rope, and Russia have proposed numer- how the species’ geographical ranges have butterflies collected from the field to ous mechanisms to explain the cyclic changed over time. His team uses mo- create a comprehensive genetic frame- nature of small mammal populations, lecular techniques to construct a family work for traits that influence mating suc- yet the phenomenon’struenature tree of the 300 or so species of beetles on cess, fertility and fecundity, lifespan, and remains elusive. the island (3). By searching for patterns in other aspects of a species’ fitness. Each Hanski proposes that the oscillating the evolution of these species, the re- butterfly is genotyped for thousands of populations reflect the interaction be- searchers also hope to learn whether— genes, as the researchers search for as- tween lemmings and voles and their nat- and how—evolution can be predicted. sociations between the genotype and ural predators. Much work has been Whereas his Finnish and Malagasy the phenotype. done on voles in Finland and Scandinavia, graduate students perform year-round Although Hanski considers himself an as well as in the high-Arctic tundra of fieldwork, Hanski cherishes his annual ecologist, the phosphoglucose isomerase eastern Greenland, where there is just one trip to the island where, for 2 or 3 weeks, revelation whet his appetite for pop- species of small mammal—the collared he camps in a rain forest to sample ulation genetics and evolution. “Re- lemming—and four species of predators. beetles and perform experiments. He searchers used to think that evolutionary Hanski developed a model to mirror what admits it is the most enjoyable part of changes occur on a very slow time scale he maintains is the simplest vertebrate his job. “No e-mail, no one telephoning; and demographic changes occur on a fast predator–prey community in existence. you are alone in the forest, eating rice time scale. That may not always be the His model revealed how the predator– and tracking beetles. That’savery case. They might occur on the same time prey interaction could trigger a waxing good time.” scale and influence each other,” he says. and waning of the populations every 4 To prove his point, Hanski continues to years, even when food is plentiful. Ecol- Ecology Meets Evolution develop models that integrate the genetic ogists later verified his predictions by These days Hanski is determined to un- and demographic changes in the Glanville analyzing decades of field data (2). ravel the underlying genetic aspects of fritillary (6), a topic that he elaborates on For Hanski, this work represents a satis- butterfly populations by sequencing the in his Inaugural Article (5). fying balance of mathematical modeling genome of the Glanville fritillary. Over the “These are very exciting days for and empirical work that he argues is ab- past 6 years, Hanski has been searching for population biologists, with so many lines solutely essential. “Without a first-hand the genetic roots of the life history char- of research coming together. The Glan- understanding of a species and its habitat,” acteristics of individual butterflies, in- ville fritillary in the Åland Islands used he says, “researchers might end up devel- cluding their reproductive potential, their to be a great system to study meta- oping misguided models that are neither lifespan, and their abilities to disperse and population ecology; now it is helping us interesting nor relevant because they lack establish new populations. Specifically, he integrate ecology and genomics in the essential features of the biological system.” has searched for a genetic explanation for spatial context. I can hardly wait to why one butterfly might be a better col- seewhatwewillhavelearnedinafew Madagascar onizer than another (4). “We have dis- years’ time!” Approximately a decade after Hanski covered a couple of genes that have a big began exploring predator–prey inter- influence on many aspects of the biology Bijal Trivedi, Freelance Science Writer

1. Hanski I, Ovaskainen O (2000) The metapopulation ca- 4. Hanski I, Saccheri I (2006) Molecular-level variation af- dispersal in the Glanville fritillary butterfly: Coupling of pacity of a fragmented landscape. Nature 404:755–758. fects population growth in a butterfly metapopulation. ecological and evolutionary dynamics. Philos Trans R 2. Gilg O, Hanski I, Sittler B (2003) Cyclic dynamics in a PLoS Biol 4:e129. Soc Lond B Biol Sci 364:1519–1532. simple vertebrate predator-prey community. Science 5. Hanski IA (2011) Eco-evolutionary spatial dynamics in 302:866–868. the Glanville fritillary butterfly. Proc Natl Acad Sci USA 3. Wirta H, Viljanen H, Orsini L, Montreuil O, Hanski I (2010) 108:14397–14404. Three parallel radiations of Canthonini dung beetles in 6. Zheng C, Ovaskainen O, Hanski I (2009) Modelling sin- Madagascar. Mol Phylogenet Evol 57:710–727. gle nucleotide effects in phosphoglucose isomerase on

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