News of the Lepidopterists’ Society Volume 52, Number 1 Conservation Matters: Contributions from the Conservation Committee Are in Trouble? If So, Why?

Arthur M. Shapiro, Center for Population Biology, University of California, Davis, CA 95616 USA [email protected]

I sometimes think of the story of P eter Bamm, who was on a lovely island where he met all kinds of people, good and bad. He dreamt in a nightmare that a bomb might come and destroy everything, and the first thing that occurred to him was what a pity it would be for the butterflies. Dietrich Bonhoeffer , Letters and Papers from Prison

I’ve been hearing the same mantra all Park Inn, with its hand-painted sign specialty of military intelligence. I my adult life: “There were more picturing eastern Red Cedar (Juniperus relied on city planning documents to butterflies when I was a kid.” And for virginiana). The tree had once been track the course of urbanization. Even most of those years, in my capacity as a common here; now it was extinct in that was ahead of its time. The first guru to the public and the Philadelphia County, and with it the papers of this sort that I know about media, I’ve pooh-poohed the idea. I Olive Hairstreak (Mitoura gryneus), appeared decades later! certainly didn’t see it happening—and if which I had to go deep into the country But those losses were local, their cause anyone should see it I should, since I’m to find. Even in the seemingly intact was transparent – loss due to in the field at least 200 days a year cool woodsiness of the Wissahickon development – and I regarded such looking at butterflies. The ubiquity of Ravine, which seemed as if it could not losses as regrettable but inevitable, and the perception, from New York to San have changed appreciably since Ben I wasn’t thinking on larger scales. And Francisco to Buenos Aires, led me to Franklin’s day, the Falcate Orange-Tip at the same time, in southwest Philly , hypothesize that its roots were ( midea) and Bates’ the Eastwick urban-renewal project had psychological. Butterflies, I reasoned, Crescent ( batesii) vanished leveled many blocks of run-down are more prominent objects in a little on my watch. (The latter seems to be housing, creating an exuberant swath kid’s landscape than in an adult’s. As we extinct in most of its historic range.) As of old-field succession that was absolute get bigger and older we notice them less, a senior at the University of butterfly heaven. (There were outbreak not because there are fewer of them but Pennsylvania I used this history as my populations of the Checkered White, because they are smaller relative to term paper in Jack McCormick’s Pontia protodice, there; it is reputed to us…and we get preoccupied with other community ecology class. He gave me be extinct in Pennsylvania now.) So sorts of things, like football and an “A” and said with a little tweaking losses might be reversible, at least in consumer electronics and sex. A neat it might be publishable. (One specific part: the bulldozer taketh away, but it explanation; perhaps even a correct one. tweak: to drop the expression “to go also giveth. And yet…I had the 19th- extinct,” which he regarded as slangy . Of course, I knew butterflies can go and early 20th-Century records of It is normal in the scientific literature downhill. My experience in this regard Henry Skinner, Eugene Aaron, Frank today.) But I never tweaked it. It’s a is anything but unique: I watched the Haimbach, Philip Laurent, J .U.D. pity; it was 40 years ahead of its time – butterfly fauna of my childhood Pleasants…; I knew that the Mulberry not dissimilar, if more mature, papers neighborhood in Philadelphia, West Oak Wing ( massasoit) and the are appearing all the time now. In 1965 Lane, shrink as the city encroached Silver-Bordered Fritillary ( Boloria Geographic Information Systems farther and farther into what had been selene myrina) had once occurred in hadn’t been invented yet and aerial woodlots and old fields. Down the road Fairmount Park, and now they were photointerpretation was still largely a from my house was the ancient Cedar gone. I saw one of the last Regal

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Fritillaries (Speyeria idalia) recorded in When I initiated my project in 1972, no to our transect across north-central southeastern Pennsylvania with my one was talking about global warming. California. The degree to which they own eyes (near Devon, Chester County, In fact, some were talking about global can be generalized elsewhere remains to in 1966; David Wright says the very last cooling, and the possibility that we were be seen. They are, however, broadly was in 1975— at least until 1990 and heading for a new Ice Age. My goals consistent with data on other 1992, also in Chester County!). Despite were short-term. To use some applied- taxonomic groups and on butterflies in all that, I remained an optimist. I also math jargon, I was not looking for a other places, as I’ll discuss a bit later . headed West. signal; I was trying to identify biotic 1. Butterfly faunas near sea level are Now I know better . I don’t think responses to noise—the short-term deteriorating rapidly, especially in the butterflies are in trouble. I know it. But weather fluctuations that a 5-year last decade. But the deterioration is not I didn’t in 1971. study would focus on. The data were adequately explained by climate change. not collected to identify biotic When I was hired at the University of The most important factor appears to consequences of any long-term trend. be habitat loss (as documented by land- California, Davis, I finally had the But when we had about 30 years’ worth opportunity to do what I had been use statistics at the County level, a the mere amount of data was daunting, more-refined use of the same technique dreaming of doing: I wanted to monitor and my research group, led by then- a butterfly fauna and use multivariate I applied in 1965!). We suspect that doctoral student Matt F orister, more sophisticated analysis using statistical methods to identify what convinced me that it was high time we environmental variables exerted the Geographic Information Systems will started analyzing the data. (I had tried reveal that loss of habitat connectivity most influence on its seasonality to get money from the National Science (phenology). My undergrad adviser had is more important than absolute Foundation years before to do this. habitat area. been Robert MacArthur, the great They were willing to fund data ecologist, and he had encouraged me to collection, but not the analysis unless 2. Butterfly faunas at mid-elevation on think of life-history phenomena as I could explain in some detail the both slopes of the Sierra Nevada are “adaptive strategies” fashioned by statistical methods to be used. I either holding their own or Darwinian evolution. My project was couldn’t, because some of them hadn’t deteriorating slowly. Here there has conceived as running about five years. been invented yet! When we did get been no significant habitat loss, and In a Mediterranean climate, with high funded, that was one of our highest changes, such as they are, are inferred interyear variance, that should be priorities—to figure out how to do what to be climate-driven. enough to give meaningful statistics. we needed to do. They teach you in grad 3. At our highest (tree-line) site, overall Besides, it was the time frame for school that all research should be butterfly richness is increasing, as more learning whether or not I’d get tenure; designed with the analytical procedures and more lower-elevation follow I might have to move on after that. fully-defined in advance. They rarely warming uphill. However, most of them I did get tenure, and the data were so teach you that the idiosyncrasies of real- cannot establish as breeding residents exciting that the project just kept going. world research routinely trump such because their essential resources, It’s now in year 39. It expanded to ten notions. They do.) especially larval host plants, are not sites (and ten faunas) from sea level to So we took all those data and began available; plants, which cannot fly , tree line, embracing both slopes of the taking them apart and asking questions respond to climate change much more Sierra Nevada and 159 species and whose answers were by and large slowly than butterflies, which can. At subspecies of butterflies (so far). The inapparent on inspection but could be the same time, 3 of the 4 most only comparably large butterfly teased out with statistical analyses. characteristic butterfly species of the database is in the United Kingdom, and You have probably read about our alpine zone at Castle P eak (not is of similar age but very differently results in the press. The study came out necessarily globally) are becoming less organized. I collect all my own data in mid-January 2010 and is the first of common. (that’s why I’m afield 200 days a year); several projected papers in various 4. The most surprising finding—we the British use a network of many stages of completion. Now, by 2010 were totally unprepared for this!—was cooperative observers who monitor some things were glaringly apparent on that the common ruderal (“weedy”), their local faunas. They have fewer the Philadelphia model—that is, visible multiple-brooded species, which some than 60 species in the entire country , to the naked eye—, but other things collectors take for granted and and substantially less topographic and were not. Here is a summary of what sometimes refer to derisively as “junk climatic diversity than exists on my we found.Keep in mind as you read this species,” are actually declining faster transect. The two projects are that we had no axe to grind; we were than the ecological specialists. These complementary, and both are designed letting the data tell us their own story . species regularly colonize upslope in for data mining. And they are being Keep in mind also that the data and the summer but cannot overwinter at high mined. inferences from them apply strictly only elevations. We expected to find them

Volume 52, Number 1 11 News of the Lepidopterists’ Society Volume 52, Number 1 becoming more common as the climate Let me qualify all of this: I know that used to be. We tend to think of whatever warmed, but in fact the reverse is folks who are concerned about we grew up with as “normal.” Within happening! This is apparently due to pesticides, air pollution, genetically- our own threescore and ten, we see loss of their preferred weedy at modified organisms, introduced change as something alarming, low elevations (as they are replaced by biological-control agents, and so forth something deviant. But Nature as we sterile residential subdivisions, are going to ask how we can be so sure see it is a freeze-frame from a very long business parks and such), which their particular bête noire isn’t involved movie. Change is the normal state of reduces their populations and thus the in these declines. (They always do.) The affairs: it’s stasis that is abnormal and number of individuals available to answer in a nutshell is that we can’t. requires explanation. There is disperse and colonize upslope. (One For some of these factors no useful data controversy over whether human “junk species” that is not declining is is available. For others—pesticides— activity is driving the current episode the European Cabbage Butterfly, Pieris too much data is available, and we have of climate change; there is no rapae, which has benefitted from the no idea how to prioritize them for controversy that the change is spread of the invasive weed P erennial analysis. Agriculture in the Central happening. There is also no Peppergrass (Lepidium latifolium) at Valley is a remarkably complex spatio- controversy that land-use change, low elevations. This weed is now temporal mosaic. The crops planted and which is apparently driving our low- marching upslope, is established at the pesticides used on them within the elevation butterfly decline in California, 5000’ and starting to show up at 7000’.) relevant geographic areas change is human-caused! In the Philadelphia of my youth, the constantly, particularly as a So when some geezer my age says to me Common Sooty Wing ( Pholisora consequence of yearly variance in “There were a lot more butterflies when catullus) was a “junk species.” When I rainfall, economics, and the actions of I was a kid,” I’m a lot more willing than came to California, it was one here too; regulatory agencies. The observed I used to be to take him seriously . patterns of butterfly decline do not by I could find larvae within ten minutes’ WANT TO LEARN MORE? walk from my lab, and I had it in my and large suggest pesticides as an garden every year . Now it is important factor, but as of now we just There’s a lot of professional literature approaching regional extinction; I don’t know. (And even in “eco- on butterfly declines, climate change know one active population in my conscious” Davis a fair number of people and related subjects. By and large it is county (Yolo) and one in adjacent have their lawns chemically treated for unknown to amateurs because it Sacramento County. It breeds on weeds and pests, but that hasn’t dented appears in scientific journals not Amaranth pigweeds—not exactly the abundance of the ubiquitous Fiery focused on per se. Here is endangered plants. The Large Marble (Hylephila phyleus) – at least some suggested reading. This is NOT ( ausonides) was common not yet. Nor have garden pesticides an attempt at an exhaustive throughout this region in the 1970s, prevented the spectacular recrudescence bibliography! of the Gulf Fritillary, Agraulis vanillae, breeding on naturalized mustards THE GRINNELL PROJECT is an (Brassica) and wild radish (Raphanus). in this region in the past few years after a 40-year absence, an event that has attempt to resurvey the altitudinal Now it appears to be regionally extinct. distributions of mammals studied in Also on a regional basis, the entire drawn lots of media attention. Of course, both of these almost completely detail by Berkeley zoologist Joseph macrolepidopteran fauna of willows Grinnell a century ago. Because his (Salix) in riparian habitat is in dire urban species are of subtropical origin… Last year the Western Tiger notes and voucher specimens and site straits on the floor of the Sacramento photographs are lovingly preserved at Valley for no obvious reason— Swallowtail, Papilio rutulus, which unaccountably went extinct in Davis— the Museum of Vertebrate Zoology, UC Lorquin’s Admiral (Limenitis lorquini), Berkeley, it is possible to revisit nearly the Mourning Cloak ( Nymphalis but not elsewhere in the region—a decade before, reappeared all over the all of his collection sites and see how antiopa), the Sylvan Hairstreak much the distributions have changed. (Satyrium sylvinus) and the once- city in extraordinary numbers. There’s a lot we don’t understand. ) You can read about the project at http:/ abundant diurnal Sheep Moth /mvz.berkeley.edu/Grinnell/index.html (Hemileuca eglanterina), all extinct in There is no doubt that climate is and from it you can download the major many former localities and hanging on changing. Climate is always changing. publication to emerge thus far: C. perilously only here and there. The At UC Davis I teach about Moritz et al., 2008. Impact of a century habitats of these species appear paleoclimates and paleovegetation. I tell of climate change on small-mammal unchanged; we infer that the cause of the students that our imaginations are communities in Yosemite National the declines is on a larger-than-local hamstrung by the temporal scale of a Park, USA. Science 322: 261-264. The scale. Most of the changes are less human life. Let’s harken back to my mammalian patterns the Berkeley crew striking and more subtle—but none the hypothesis about why people think is finding are quite similar to ours in less real. there are fewer butterflies than there butterflies.

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BIOTIC RESPONSES TO years on a tropical mountain. climate and land-use change on the CLIMATE CHANGE are documented Proceedings of the National Academy of global diversity of . PLoS Biology in hundreds of papers, with more Sciences of the USA 106:1479-1483. 5:1211-1219. coming out weekly—as an on-line Conrad, K.F. et al. 2006. Rapid declines Opdam,P. and D . Wascher. 2004. search will quickly show! Here are a few of common, widespread British moths Climate change meets habitat important ones. provide evidence of an fragmentation: linking landscape and Bale,J.S. et al. 2002. Herbivory in biodiversity crisis. Biological biogeographical scale levels in research global climate change research: direct Conservation 132:279-291. and conservation. Biological effects of rising temperature on insect Dennis, R.L.H. and T.H. Sparks. 2007. Conservation 117:285-297. herbivores. Global Change Biology 8:1- Climate signals are reflected in an 89- vanDyck,H. et al. 2009. Declines in 16. year series of British Lepidoptera common, widespread butterflies in a Hickling,R. et al. 2006. The records. European Journal of landscape under intense human use. distributions of a wide range of Entomology 104:763-767. Conservation Biology 23:957-965. taxonomic groups are expanding Morecroft,M.D. et al.2009. The UK vanSwaay,C., M. W arren and G . polewards. Global Change Biology Environmental Change Network: Lois.2006. Biotope use and trends of 12:450-455. Emerging trends in the composition of European butterflies. Journal of Insect Menendez, R. et al. 2006. Species plant and communities and the Conservation 10:189-209. richness changes lag behind climate physical environment. Biological Warren, M.S. et al. 2001. R apid change. Proceedings of the Royal Conservation 142:2814-2832. responses of British butterflies to Society B (Biological Sciences) Parmesan, C. et al. 1999. P oleward opposing forces of climate and habitat 273:1465-1470. shifts in geographical ranges of change. Nature 414:65-69. Parmesan, C. 2006. Ecological and butterfly species associated with White, P. and J .T.Kerr.2006. evolutionary responses to recent regional warming. Nature 399:579-583. Contrasting spatial and temporal global climate change. Annual Review of Pollard,E. and B.C.Eversham. 1995. change impacts on butterfly species Ecology, Evolution and Systematics Butterfly monitoring 2—interpreting richness during the 20th Century . 37:637-669. the changes. In A.Pullin, ed. Ecology Ecography 29:908-918. Parmesan, C. and G. Yohe. 2003. A and Conservation of Butterflies. One of the first papers spotlighting globally coherent fingerprint of climate- Chapman & Hall. Pp.23-26. urban butterfly ecology was by Bob change impacts across natural systems. Poyry,J. et al. 2009. Species traits (R.M.) Pyle, 1983: Urbanization and Nature 421:37-42. explain recent range shifts in Finnish endangered insect populations, Ch.15 Root, T.R. 2003. Fingerprints of global butterflies. Global Change Biology in G.Frankie and C.S.Koehler, eds., warming on wild and plants. 15:732-743. Urban Entomology: Interdisciplinary Nature 398:611-615. Roy, D.B. and T .H. Sparks. 2000. Perspectives, pp. 367-394. Praeger Walther, G.R. et al. 2002. Ecological Phenology of British butterflies and Scientific, New York. This paper was responses to recent climate change. climate change. Global Change Biology far enough ahead of its time that even Nature 416:389-395. 6:407-416. I forgot about it for years, and it is hard to find. It should be reprinted accessibly Wilson,R.J. et al. 2005. Changes to the IMPACTS OF LAND USE AND for Lepidopterists. elevational limits and extent of species INTERACTIONS WITH CLIMATE range associated with climate change. are increasingly well-documented, …and finally, our own paper is Ecology Letters 8:1138-1146. sometimes with Leps, e.g.: Forister, M.L. et al. 2010. Compounded IMPACTS ON LEPIDOPTERA are Brook, B.W., N.S. Sodhi and effects of climate change and habitat documented in a few dozen papers so C.J.A.Bradshaw. 2008. Synergies alteration shift patterns of butterfly far, many of them from Europe and the among extinction drivers under global diversity.Proceedings of the National British Isles, some from the Tropics! change. Trends in Ecology and Academy of Sciences of the USA. DOI: Examples: Evolution 23:453-460. 10.1073/pnas.0909686107. Altermatt, F. 2009. Climatic warming Clark, P.J., J.M. Reed and F.S. Chew. And our Web site, with details on our increases voltinism in European 2007. Effects of urbanization on project and summary data, is http:// butterflies and moths. Proceedings of butterfly species richness, guild butterfly.ucdavis.edu. Come visit us! the Royal Society B (Biological structure, and rarity . Urban Sciences): DOI: 10.1098/rspb.2009.1910 Ecosystems 10:321-337. X Chen, I.C. et al. 2009. Elevation Jetz,W., D.S. Wilcove and increases in moth assemblages over 42 A.P.Dobson.2007. Projected impacts of

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Are Butterflies in Trouble?

2 Fig.1) Riparian habitat in the Sacramento V alley looks fine, though reduced by an estimated 99-95% since the 19th Century . It is losing species faster than any other habitat type on our transect. This is a scene in the North Sacramento study site. Fig.2) The Mourning Cloak, Nymphalis antiopa, shown here visiting Rabbitbrush, Chrysothamnus nauseosus, at Donner P ass in the Sierra Nevada, has undergone a catastrophic decline near sea level on our transect in the past ten years, but its troubles may be related to its rhythm of annual altitudinal migration. Fig. 3) The familiar Acmon Blue, Plebejus acmon, shown on a Smartweed ( Polygonum) flower in a drainage ditch at our W est Sacramento site, is one of the “weedy” species that seem to be suffering from loss of habitat at low elevations—leading to a decrease in occurrence in the mountains, where it is an immigrant. Fig. 4) Here at tree-line on Castle P eak in the Sierra Nevada, more and more lower- elevation species are turning up as strays. Meanwhile, the true high- altitude species, such as the Ivallda Arctic ( Oeneis chryxus ivallda), found in this rubble-strewn rock garden, appear to be in decline.

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