WINGS ESSAYS ON INVERTEBRATE CONSERVATION

THE XERCES SOCIETY FALL 2007 CONTENTS

Introduction: Invertebrates and Global Warming Scott Hoffman Black Page 3.

Wetland Invertebrates: What Happens When the Pond Is Half-Empty? John H. Matthews Water temperature and volume, both critical to the development and emergence of dragonfly larvae in wetlands, are changing as global warming increases. Page 5.

Bleached Out: Coral Reefs Under Stress David Obura Tropical coral reefs are one of the first ecosystems to show significant changes due to changing climate. Page 11.

Beer for Butterflies: Tracking Global Warming Matthew L. Forister As scientists study the impacts of global warming upon , patterns are emerging of both shifts in range and changes in the timing of life-history events. The consequences for these tiny creatures may be enormous. Page 15.

For Marine Worms, Timing Is Everything Andrew Lawrence Marine worms are a hugely important link in the food chain of intertidal areas. In- creasing water temperatures place their future breeding success at risk. Page 19.

Impacts of Global Warming on Pollinators David Inouye Climate change is altering the phenology of both plants and their visitors, breaking the link between the two and putting at risk the essential service of pol- lination. Page 24.

Xerces News Protecting native bumble bees and other pollinators, wetland conservation efforts, and a Butterfly-A-Thon; and new staff members join the Xerces Society. Page 28.

2 WINGS Introduction: Invertebrates and Global Warming

Scott Hoffman Black

Not long ago, as I was hiking along a collecting, and a prolonged drought roaring stream on my way up to Col- that was thought to be drying out the orado’s Red Cloud Peak, my mind wan- habitat, this butterfly had recently been dered back to the summer of 1995. I had listed as an . been hired by the Natural The lives Heritage Program to help organize a only in alpine meadows. Thought to search for the Uncompahgre fritillary have been broadly distributed near gla- ( improba acrocnema). With few cial margins during the last ice age, the known populations and multiple threats is now confined to small including sheep grazing, potential over- patches of habitat above thirteen thou-

The San Juan Mountains of Colorado are home to the Uncompahgre fritillary ( acrocnema), one of many alpine that are vulnerable to climate destabi- lization. Warming climate may spell extinction for many of these species. Photograph by Scott Hoffman Black.

FALL 2007 3 sand feet in the San Juan Mountains of but will it really be the same without a southwestern Colorado, where cool and species that has resided here for thou- wet environments have persisted to the sands of years? present. The Uncompahgre fritillary is not On my recent trip, I was heading up the only invertebrate facing a changing to see how things had changed in the environment. In this issue of Wings, we last twelve years. Although it was a lit- explore multiple aspects of climate de- tle late in the year, I hoped to see a few stabilization from the perspective of in- late-season stragglers. The area looked vertebrate conservation. Included are es- much the same as it did in 1995. Snow says on how warming temperatures are (the Uncompahgre fritillary’s lar- already leading to bleaching corals and val hostplant) was still abundant. There shifts in butterfly ranges. You will see were bumble bees, flies, and several spe- that global warming may lead to the ex- cies of butterflies including the arctic tinction of some butterflies and aquat- fritillary (), Uncompah- ic invertebrates. It may also lead to the gre’s sister species. loss of some marine worms that are an I carefully made my way through important food source for many birds the habitat hoping to spot an adult Un- and marine . In short, global compahgre. I thought about the com- warming is already having a profound plexities and uncertainties confronting impact on both terrestrial and marine us. Global warming will not lead to uni- ecosystems. form change across the landscape. Some It is interesting to note that the first places will be hotter and drier while species thought to have been driven ex- others may be wetter; in still others the tinct by global climate change was a weather may simply be more variable. snail. Does it matter that one snail goes It may affect plant communities and extinct or that the Uncompahgre fritil- cause seasonal shifts that put species out lary may follow? There has been talk of synchrony with their food sources. lately of taking action to protect some After a fruitless search, I decided to of the species that have been led to the hike the last few hundred feet to the brink of extinction by global warming. ridge to take in the magnificent view. Some scientists have floated the idea Just as I reached the very limit of the of assisted migration (helping species habitat, a familiar sight met my eyes. A move to habitat where they were never faded and battered Uncompahgre fritil- native). For most species this option is lary fluttered across the trail to land on neither feasible nor wise. What we need a rock a few feet away. I sat down next to do is to take heroic efforts as a soci- to the butterfly, noting that its life was ety to curb the impact of global warm- near an end. I then looked up and real- ing. We must find the societal and po- ized that this species really has nowhere litical will to take the actions that are to go as the climate changes. It cannot needed to curb this global threat. live in the rock and ice at the top of the To find out some changes that you peak. I thought about what future gen- can make as an individual, go to www. erations will see when they come up climatecrisis.net/takeaction/whatyou this mountain. It might look the same, cando/index6.html.

4 WINGS Wetland Invertebrates: What Happens When the Pond Is Half-Empty?

John H. Matthews

In May 2003, I stuffed my car with tween climate change and the aquatic camping and collecting equipment, said macroinvertebrates found in small wet- good-by to my wife, and began a twelve- lands. Given the diversity and abun- hundred-mile drive north from central dance of life they hold, these wetlands Texas. On the front seat next to me sat are both under-studied and beneath the a well-thumbed photocopy of a 1970 radar of the public and policy makers. Ph.D. dissertation about dragonflies, Most small wetlands are ephemeral written by a University of Toronto stu- aquatic habitats (that is, they dry up on dent named Robert Trottier. And the a regular basis), but even “permanent” topmost page had a fuzzy photograph wetlands swing wildly in their annual of a pond in Caledon, Ontario. water volume. Since many fish are in- I was headed to Caledon to begin a tolerant of large shifts in volume, these multi-year study of the relationship be- fish-free habitats are typically rich in in-

Common green darners (Anax junius) are found at lakes, ponds, and wetlands across much of North America. Male, photographed in Texas by John C. Abbott.

FALL 2007 5 When mating, the male dragonfly grasps the female behind her head, and she curls her abdomen around to accept sperm. Com- mon green darners, photographed in Texas by Giff Beaton. vertebrate diversity. Indeed, the top car- of the water in most small wetlands nivore positions held by fish in larger comes from precipitation. And in recent bodies of water tend to be occupied by decades, the timing, form, and amount dragonfly or beetle larvae in such wet- of precipitation have been changing. lands. But most of these wetlands are Less snow is falling. Globally, precip- viewed as insignificant and marginal itation amounts are increasing and ecosystems, and many countries offer individual precipitation events on av- them little or no protection. In my long erage are becoming more intense. But drive to Caledon I had time to ponder because air temperatures are also gener- the question of just how sensitive these ally warmer, evaporation rates are high- wetlands are to shifts in climate. er. The effect is even stronger in arid Climate change has been generally and semi-arid regions, where, for the viewed through the lens of air temp- most part, precipitation is decreasing erature, a bias reinforced by the term while evaporation rates are rising. As a “global warming.” However, in addition result, many places — even those receiv- to air temperature, many aspects of cli- ing more rain — are actually drier more mate such as wind speed, cloud cover, of the time. To make the matter worse, and relative humidity have been shift- the timing of precipitation is changing ing over the past century. The majority in many areas. As every gardener knows,

6 WINGS the timing of rain matters. Annual in- bodies of water such as rivers and lakes creases in mean precipitation are shift- tend to change volume (or temperature) ing at a monthly or seasonal scale in bi- slowly, even small changes in precip- ologically significant ways. And all of itation regime can radically alter the these changes are likely to intensify in volume of small wetlands, and thus coming decades. the temperature of their water. Less There are two major effects on small water volume means warmer water, wetlands that could come from shifts in since there is a smaller thermal mass. precipitation patterns. First, the timing And more water means cooler water, be- or seasonality during which ephemeral cause of the larger thermal mass. wetlands hold water (their hydroperiod) Temperature matters to all organ- is likely to alter. Such changes are im- isms, especially invertebrates. Metabo- portant because many wetland inverte- lism, development, activity level, re- brates depend on a regular and pre- production rate, the ability to find prey dictable hydroperiod for their life cycle. or escape predation, and many other If there’s no water, they might need to basic processes are influenced directly speed up their rate of development or by temperature. Thus, changes in water try to leave that wetland in search of an- volume that result in water temperature other with a longer hydroperiod. If they shifts should also alter behavior. can’t disperse very well, they might have At least, these were my thoughts as to survive harsh drying conditions, per- I pulled into Caledon with Trottier’s dis- haps by entering a resting state. Or they sertation. Research into biological im- might simply die, causing a local popu- pacts from climate change faces some lation to go extinct. For insects with basic hurdles. Many of these studies are aquatic larvae, such as stoneflies, dia- physiological, experimentally testing pause and death are the two most com- the tolerances of organisms to particu- mon responses to desiccation. lar environmental variables like tem- Second, water volume is also asso- perature or humidity. However, these ciated with temperature. Water is much studies have proven too difficult to gen- harder to heat or cool than air because eralize. Even closely related members of it has a lot of thermal mass. While large the same genus of butterfly or flower

Changes in water volume and temperature can impact the development and emergence of dragonfly larvae. Common green darner larva, photographed by John C. Abbott.

FALL 2007 7 can have very different physiological re- 1960s, Trottier had no awareness of sponses to the same stimulus. Moreover, anthropogenic climate change. He was outside of the laboratory, multiple cli- interested in untangling the develop- mate variables are changing simultane- mental cycle of the common green darn- ously, and there is no guarantee that the er (Anax junius), a species of dragonfly selected variable in a laboratory will be found from southern Canada to the Yu- very important in a natural setting. His- catan peninsula of Mexico whose larvae torical studies that compare current be- cannot survive outside of the small wet- havior to an earlier period are an effec- lands they favor. To explore their de- tive way to observe species’ responses velopment, Trottier tracked emergence, over time, which can then help us un- the process of metamorphosis from lar- derstand what further changes we can val stage to adult. When a green darner expect in coming decades. metamorphoses, the larva crawls from Unfortunately, historical studies the water onto some projection such as have been rather limited for inverte- a stick or cattail and then the adult brates. Although insects make up the emerges and flies away, leaving behind largest number of species on the planet the empty exoskeleton. (and have one of the largest biomasses), Trottier hired a young field assistant there are very few baseline datasets for named Mike McCormick, set up some studies on the impact of historical cli- fencing around the perimeter of a pond mate change. Which is why I had Trot- in Caledon as an emergence substrate, tier’s dissertation with me in Caledon. and waited. Trottier also brought some As a graduate student in the late green darners into his Toronto labora-

Due to their small size and seasonally variable conditions, ponds and wetlands may be impacted by changes in both rainfall and temperature. This is the Cale- don, Ontario, research site studied by the author. Photograph by John Matthews.

8 WINGS Many birds rely on adult aquatic insects for food. Tree swallows (Tachycineta bicolor), photographed in Quebec by Michel Bordeleau. tory where he found that he could alter of emergence had changed since the their rate of development by changing 1960s, and that these changes were the temperature of their ambient water. closely associated with decade-scale in- Developmental rates were quite plastic, creases and decreases in precipitation. varying by factors of two to three over We believe that changes in thermal small temperature ranges. mass and hydroperiod in this wetland I saw Trottier’s emergence data have significantly altered the timing of from the late 1960s on the timing of dragonfly emergence in Caledon over emergence as a potential baseline for just the past thirty-five years. comparison, particularly since his re- How important are these results search ended just before a period of rel- from a more general conservation per- atively rapid climate change began. spective? Ontario is a relatively wet re- Starting around 1974, many climate var- gion in the northern temperate zone, iables began to shift swiftly worldwide. and small shifts in dragonfly emergence Moreover, Trottier had documented the may not be critical there. Except during importance of temperature on the de- drought years, there are abundant wet- velopment rate of green darners. lands in eastern Canada of many sizes While in Caledon, I was especially and thermal masses. lucky to meet the grown-up Mike Mc- However, the implications of these Cormick, and he and his wife, Darleen, results should be of serious concern in provided access to a pond that was very regions where surface water is more lim- similar to the original site. With three ited. In semi-arid regions, small ephem- young and excited field assistants of my eral wetlands are often the most impor- own, I began four years of emergence tant and widespread type of freshwater and environmental monitoring at the habitat. For instance, across most of the McCormick pond. Following Trottier’s Great Plains and the Great Basin of method, we set up an emergence sub- North America, small wetlands — often strate. We found both that the timing called prairie potholes or playa lakes at

FALL 2007 9 the northern and southern ends of the tion. In the United States, small wet- Great Plains, respectively — are critical lands are under growing pressure be- habitat and contain a different inverte- cause recent judicial reviews of the brate fauna from nearby reservoirs and Clean Water Act have eroded the abili- rivers. Many climate models suggest ty of federal agencies to protect non- these regions will become drier in com- flowing freshwater systems. ing decades. Second, there are grounds for hope Therefore, in these regions changes for small wetlands and the species they in hydroperiod and thermal mass are house if we can commit to their pro- likely to have a strong effect, and not tection. These wetlands should be ex- just on dragonflies. Precipitation-driven cellent candidates for protective legisla- changes in habitat are also likely to af- tion and active resource management. fect mayflies, stoneflies, caddisflies, dip- Particularly in regions projected to be- terans (such as mosquitoes and midges), come drier in future decades, relatively and many other groups. The impacts on small modifications to the shape and these species will not be limited to fresh- size of small wetlands can alter their water taxa. Many bird species, for in- surface-to-volume ratio, reducing the stance, specialize on the adult terrestri- rate of evaporation and shifting the al stage of aquatic insect species. David thermal regime. We need to give these Winkler of Cornell University believes species some more time to adapt during that continental-scale changes in the a period of intensifying climate change. timing of egg laying in tree swallows In some cases, more small wetlands may (Tachycineta bicolor) is an indirect effect need to be created at higher latitudes so of shifting climate on the emergence that the ranges of species can move as timing of aquatic dipterans, their pri- their preferred climate “envelope” shifts mary prey. Given that many species are spatially. sensitive to the timing of behaviors in These wetlands are little worlds in other species — think of the importance themselves, full of life and activity. I of the coordination between flowering find them beautiful and compelling. time and a specialist pollinator — even Our care for them and their inhabitants minor changes can alter complex inter- may tell us something about how pre- species relationships. I fear that ephem- pared we are to help organisms in more eral small wetlands may be one of the complex habitats respond to our plan- types of habitats most endangered by et’s emerging new climate. climate change. For the next decade, there are prob- ably two broader lessons to keep in John Matthews is a freshwater climate mind about global warming and fresh- change specialist with the World Wildlife water invertebrates in small wetlands. Fund in Corvallis, Oregon. He continues to First, few studies have focused on how work on issues related to aquatic inverte- climate change is altering wetlands, so brates, though occasionally he has to spend more research should be conducted on time on vertebrates as well. John thanks this topic while more formal protection his Caledon field assistants, Nathan Bur- is enacted to reduce wetland destruc- nett, Kieran Samuk, and Marie Riddell.

10 WINGS Bleached Out: Coral Reefs Under Stress

David Obura

Objects of great beauty in themselves, this skeleton that gives stony (or scler- coral reefs are home to immeasurably actinian) corals their name. A few stony diverse communities of marine life. corals are solitary polyps that may reach Often described as “tropical forests of ten inches (twenty-five centimeters) the sea,” or “gardens of paradise,” reefs across, but the great majority are tiny support a tourism industry worth a bil- polyps, usually less than an eighth of an lion dollars, and tens of millions of peo- inch (three millimeters) across, and live ple living along tropical coasts are de- in vast, reef-building colonies. pendent directly on them for transport Stony corals cannot successfully live networks, building materials, fisheries, on their own. For survival they rely on and the protection of coastlines. a symbiotic relationship with zooxan- Corals are closely related to sea thellae, single-cell algae that live with- anemones. The major difference is that in the polyps. The relationship is clear- many corals produce a cup-shaped skel- ly mutually beneficial. Indeed, it has eton of calcium carbonate, within which evolved to the point of being effectively the anemone-like coral polyp lives. It is obligate in nature for the coral: the

Healthy corals form reefs of many different colors and shapes. This coral garden was photographed in Papua New Guinea by Tammy Peluso.

FALL 2007 11 The purple areas of this Pocillopora coral are normal, while the pink and white areas are in successive stages of bleaching. The brown areas are dead. Photographed in the Mariana Islands by David Burdick; courtesy the National Oceanic and Atmospheric Administration. two can survive apart in controlled degrees Celsius), so reefs are almost ex- laboratory conditions but in the real clusively restricted to tropical seas less world, coral species that are normally than 150 feet (50 meters) deep. The part- symbiotic are incapable of maintaining nership between corals and zooxan- themselves alone against competition thellae is one of the most successful on and infection. the planet; this relationship has existed Through photosynthesis, zooxan- for more than four hundred million thellae capture energy into chemical years and the resulting reefs are the only form, a large part of which they pass to biogenic structures visible from space. the coral polyp. This provides the fuel Over the years, a picture has devel- for cellular functions within the polyp, oped of coral reefs as highly sensitive including the creation of its calcium car- networks of species in a delicate equi- bonate skeleton, which leads to the ac- librium. Cold temperatures have long cretion of reef structures as new corals been known to be inimical to reef sur- grow on the skeletons of previous gen- vival, but in the 1970s evidence accu- erations. In return, the zooxanthellae mulated that showed corals have upper are physically protected by living with- temperature and light limits as well as in the polyp, and the reef structure pro- lower limits. By the 1980s, scientists vides them with access to light, which were witnessing large-scale “bleaching” they require to survive. They also re- and mortality of corals believed related quire water temperatures greater than to warming seas associated with in- sixty-eight degrees Fahrenheit (twenty creasing levels of greenhouse gases in

12 WINGS the atmosphere and perturbation of and peak fluctuations are being pushed ocean-atmosphere oscillations such as higher than ever. Corals in all regions the El Niño and La Niña cycles. Since appear to be adapted to a late-summer that time, bleaching and mortality have peak of temperature and light when the increased rapidly. sun is at its local zenith. However, any It is now widely accepted among increment above seasonal norms push- coral reef scientists that coral bleaching es corals and zooxanthellae over their represents a last-ditch attempt by corals adaptive limit — and around the world and zooxanthellae to resist stressful con- temperature and/or light are beginning ditions. A wide range of stress causes to exceed their historic summertime them to separate, and the departure of maxima. In 1998 alone, during the hot- the pigmented zooxanthellae from the test year on record, coinciding with an coral tissue leaves the tissue transparent, El Niño peak in the Pacific and a similar revealing the bright white skeleton be- climate peak in the Indian Ocean, 16 neath. Under not-too-stressful condi- percent of the world’s reefs were judged tions, the partners can often rejoin and to be severely degraded. continue growing; however, when con- The implications of continuing ditions are too severe, bleaching is usu- stress are dire for coral reefs. Although ally followed by mortality. few species on a reef rely directly on Excessive temperature and light in- stony corals for food or energy, the tensities are the main causes of stress in structure that corals provide enables the the polyp-zooxanthellae symbiosis, and rest of the reef community to exist. The these two factors are increasing globally. decline in coral diversity and cover on Background levels of both are rising, some reefs has resulted in lower diver-

Completely bleached corals are white; as they bleach they can take on fluorescent hues of purple, blue, green, or yellow. Acropora coral, photographed in the Mariana Islands by David Burdick; courtesy the National Oceanic and Atmospheric Administration.

FALL 2007 13 sity and abundance of obligate associ- search on coral and zooxanthellae evo- ates of coral, those crustaceans and fish lution and physiology as well as reef dy- that live in spaces among branching namics, and it is clear that with two part- corals, or fish that feed only on corals. ners interacting, each with their own If coral growth does not resume, the genomes, there are levels of adaptation framework of the reef itself erodes, re- and acclimation available to corals that sulting in still lower diversity and abun- we don’t yet understand. Coral reef sci- dance of fish, including many fisheries entists are split between those who have species of economic importance. In the measured very limited capacity for ad- Galapagos, where only small reefs ever aptation, and those who offer evidence build up, near-total death of corals in for considerable capacity for adaptation. the El Niño of 1983 resulted in complete In my view there will be many sur- erosion of some local reef frameworks prises, but it is hard to imagine that an within the next twenty years. ecosystem knocked down to just 10 per- Quantifying the potential loss of cent of its historic extent by conditions biodiversity from declining reef health that are predicted to become increas- is probably impossible. What would it ingly hostile, can have much chance of mean if the area of healthy reefs de- functional or widespread survival. Per- clines to less than 10 percent of what it haps a few locations will retain func- was a few decades ago? What is the like- tional coral reefs, and in fifty years we’ll lihood of coral species going extinct? have to travel to South Africa or Japan The majority of coral reef scientists, to see the corals that make it farther managers, and enthusiasts, including from the equator as seas continue to the scientists on the United Nation’s warm. People may debate the details of Intergovernmental Panel on Climate coral declines but, for those who de- Change and the team that put togeth- pend on coral reefs to earn enough er last year’s Stern Review for the British money to support their families, it is not government, believe that coral reef de- fair to say “no problem, no action need- cline is a harbinger of decline in other ed,” when we really don’t know. There ecosystems. Others who minimize the are many challenges that need our at- significance of coral reef declines point tention and that will benefit from a pre- to the coral species that bounce back cautionary rather than a laissez-faire ap- quickly after bleaching. These species, proach to climate change. however, are mainly widespread gener- alists and not representative of all spe- cies on a reef, much as weeds and cock- David Obura coordinates CORDIO East roaches are not representative of all Africa, supporting activities in mainland plants or insects. Their ability to survive Africa and the island states, including re- changing conditions masks the impact search, monitoring, and capacity building of such change on global coral diversity. of coral reefs and coastal ecosystems. Of course no one can precisely pre- He has studied sediment stress and coral dict the future for corals, but some out- bleaching and life-history strategies, and comes are more likely than others. The has a primary research interest in climate climate-change debate is fueling new re- change and the resilience of coral reefs.

14 WINGS Beer for Butterflies: Tracking Global Warming

Matthew L. Forister

The cabbage white butterfly (Pieris ra- Dr. Shapiro’s lab, biologists were com- pae) is so common and plain as to be of ing to a consensus that contemporary little interest to most butterfly enthusi- changes in our climate were having asts. However, under certain conditions equally contemporary and dramatic a cabbage white butterfly can be cashed changes on the plants and animals in for a pitcher of beer. For more than around us. For butterflies, much of the thirty years, Dr. Art Shapiro of the Uni- discussion was coming from northern versity of California, Davis, has held a Europe (England in particular), while we “Beer for a Butterfly” contest. A pitcher knew relatively little about the same of beer goes to the person who nets the phenomena at lower latitudes. This ge- first cabbage white within California’s ographical disparity in our knowledge Sacramento, Solano, or Yolo counties inspired us to examine the results from on or after the first of the New Year. the “Beer for a Butterfly” contest. We Since he has the ability to think like a found that the beer had been handed butterfly, Dr. Shapiro usually wins. I out earlier and earlier over the course of tried, but never came close. the previous three decades. In fact, the Around the turn of this century, as first cabbage white of the year was I was pursuing my graduate studies in now appearing on average almost three

The cabbage white (Pieris rapae) is a European species naturalized in North America. Because of warming temperatures it now appears nearly three weeks earlier in the spring in some areas. Photographed in California by Henk Wallays.

FALL 2007 15 weeks earlier, as were many of its fellow perature controlled. Consider a butter- butterflies in the same region. These fly that spends the winter as a caterpil- changes appeared to be related to warm- lar in arrested development. In the heart ing and drying climatic conditions in of winter, a succession of cold days caus- the region. es the caterpillar to “break” dormancy Results like these from the cabbage and resume normal metabolic activity white and its neighbors are all too com- in preparation for spring. After that, mon. Similar effects are turning up on subsequent growth and emergence may every continent and with many differ- be dramatically affected by temperature: ent kinds of creatures. with a very warm spring, the caterpillars Of species’ varied responses to cli- will grow more rapidly and the first mate change that have been observed, adult butterflies will be seen earlier than shifts in geographic range and in the in a year with a cold spring. timing of life-history events (phenol- The other side of the coin is move- ogy) are of fundamental importance. ment in space. Studies by scientists, Both kinds of changes follow from the including Camille Parmesan of the Uni- fact that organisms depend on cues versity of Texas and Jane Hill of the Uni- from their environment to determine versity of York, have shown that but- when, where, and how to live out their terflies across Europe are shifting north- lives. When conditions become warmer ward. In particular, a study of thirty-five or drier (or colder or wetter), the envi- non-migratory species found that, dur- ronmental cues change and the biology ing the twentieth century, the ranges of plants and animals may or may not of nearly two-thirds of the species ex- change in response. It has become very panded northward by between 35 and clear (at least in temperate regions) that 240 kilometers (22 and 150 miles). In the phenology of butterflies is changing some but not all cases, the southern edge in predictable ways. In particular, the of the range has also shifted northward. date when the first butterfly of many An instructive example of similar species is observed is coming earlier in movement comes from the western the season. United States, and the work of Lisa Cro- Why does this happen? In regions zier of the National Oceanic and Atmos- with strongly seasonal climates, most pheric Administration. The sachem butterflies spend at least part of the skipper (Atalopedes campestris) has re- year in a dormant phase. This part of cently expanded the northern edge of the year is often winter but in some its range from northern California to cases, it’s the heat of summer. Butterflies southern Washington. Through exper- have a variety of ways of spending these iments in which caterpillars were ex- dormant times: as different life-history posed to a range of environmental con- stages and in different states of inactiv- ditions, Crozier was able to show that ity. For many butterflies, entry into dor- winter temperatures were the primary mancy is controlled by photoperiod factor affecting the recent expansion. (day length), and may or may not be in- Pacific Northwest winters warmed by fluenced by temperature, while emer- roughly three degrees Celsius (five and gence from dormancy is largely tem- a half degrees Fahrenheit) in the second

16 WINGS The Sachem skipper (Atalopedes campestris) is found across the United States. On the West Coast its range has recently expanded northward into Washington state. Photographed in Virginia by Preston Scott Justis. half of the twentieth century. Before the suitable higher-elevation habitat, but- 1990s, winter temperatures in Wash- terflies colonize the higher locations ington were generally below a critical and establish new, viable populations. survival threshold for overwintering Robert Wilson of Madrid’s Universidad caterpillars; the threshold appeared to Rey Juan Carlos reports similar behav- be average January minimum temper- ior among numerous species in the atures of minus four degrees Celsius mountains of central Spain. (twenty-five degrees Fahrenheit). Once So what’s the problem with all this temperatures on average warmed above shifting around in space and time? that point in the 1990s, the skipper was For changes in space, the consequences able to rapidly expand its range and is may be painfully obvious. Montane now a regular member of the Washing- populations can be pushed only so far ton fauna. upslope or poleward along a mountain A comparable phenomenon is the chain: if warming continues, they may upslope movement of butterflies on run out of mountain. The consequences mountains in response to warming of shifting in time are more subtle. The conditions. In France, populations of a life cycles of many butterflies, particu- montane butterfly, the Apollo (Parnas- larly those with a single spring genera- sius apollo), have gone extinct at lower tion, are tightly timed to the phenolo- elevations but, as described by Henri gy of their hostplants, the ones that the Descimon at the Université de Pro- caterpillars depend on for food. Cater- vence, in Marseille, they persist on pla- pillars may, for example, need to eat teaus at higher elevations. Where the newly unfolded leaves, which are not lower-elevation populations are close to only tender but also relatively free of the

FALL 2007 17 toxins that accumulate as the leaf de- lutionary process, but there are daunt- velops. If the butterfly’s flight period is ing issues of chance and time that all shifted either forward or backward, the adapting populations face. For species adults may lay eggs but caterpillars will with nowhere to go it seems likely that not find optimal foliage. It is also possi- many will simply go locally extinct, in- ble that the phenology of plants may cluding not only species on mountains change, but there is no particular reason — such as the Apollo butterflies trapped to think that a shift in plant phenology on warming plateaus — but also species will perfectly match a shift in butterfly in extremely localized or fragmented phenology. Of course, climate change habitats. does not result in uniform warming. Given the possibility of extinction Many areas are receiving increased win- in some cases, and phenological or spa- ter precipitation, which may delay de- tial shifts in others, it’s interesting to velopment or even increase mortality consider what your favorite butterfly through mold or disease; even if not community, in your backyard or local killed outright, stressed larvae become park, will look like in ten, twenty, or less healthy adults. fifty years. It is quite possible that you If changes in range and phenology will see one or two species that you have can lead to such problems, why don’t never seen before as they colonize from organisms adapt to changing conditions lower latitudes, and you might even and just stay put? After all, it is clear that spot a few tropical gems. It’s equally butterflies have adapted to almost all possible that some species may be gone, manner of environments because we either locally or globally, and that new find them in the hottest deserts and on ecological interactions may arise as the highest mountains. Although the plant and predator communities are fact is that we know relatively little also shifting around the butterflies. Out- about the evolutionary responses of comes will also depend on many fac- butterflies to climate change, examples tors not related to climate change, such are accumulating of one intriguing pos- as regional habitat destruction. In any sibility: dispersal facilitated by adap- event, the coming decades may be both tation. Populations at northern range intriguing and disheartening for ob- boundaries may evolve greater dispersal servers of butterflies. Perhaps we should ability (in response to newly available all have a “Beer for a Butterfly” contest, habitat), and, in the case of the brown and raise a glass or two for the butterflies argus (Aricia agestis), populations at a as they come and go in space and time. northern range boundary have colo- nized a novel hostplant, which subse- quently facilitated range expansion, as Matthew Forister is a research assistant described by Chris Thomas at the Uni- professor in the Department of Natural Re- versity of Leeds. sources and Environmental Science at the But what if there is nowhere to University of Nevada, Reno. His research go? Is adapting to new conditions pos- involves the ecology and evolution of her- sible? The answer is certainly yes: we bivorous insects, particlarly issues of speci- know of few true constraints on the evo- ation and ecological genetics in butterflies.

18 WINGS For Marine Worms, Timing Is Everything

Andrew Lawrence

Let me begin by asking you to remove you will die the next day, do you think your watch, close your calendar, turn that your genes would have been passed off your TV or radio, throw away your on to the next generation? newspaper, and not speak to anyone. This is the challenge facing many How long do you think that it would be invertebrates. Lucky then for the is- before you lost track of time — the pre- landers of Samoa and Fiji that the palo- cise time of day, day of the week, or lo worm has managed to solve this puz- month of the year? And, if you were zle. Each year, millions of worms release asked to keep track of time, how might their epitokes, their rear sections full of you try to do this? sperm or eggs, which swarm at the sea What if I said that you have the surface to reproduce. The islanders have chance to meet the man or woman of been able to predict the date and time your dreams at 7:00 in the evening of of day of this phenomenon for cen- October nineteenth and that, if you turies. At dawn on the day before and meet, you will enjoy a night of passion then on the actual day of the moon’s leading to the birth of your only child? last quarter in October or November, Do you think that you would be able to people row or wade out to sea with nets make the meeting? And, if I said that ready to catch the animals for the brief

During their annual swarm, palolo worms (Palola siciliensis) release epitokes, rear sections filled with eggs or sperm. The male epitokes are tan, those of females a darker gray-green. Photographed in American Samoa by Stassia Samuels; courtesy the National Park Service.

FALL 2007 19 period that they are available. The palo- ternal clock mechanism, which in turn los are considered a delicacy and eaten controls the ’s annual cycle of raw or cooked, and their arrival is often activity or physiological processes (bio- followed by festivals. rhythm). Temperature is a secondary The palolo worm is one of the most control. While it also follows seasonal dramatic examples of an animal that cycles, it can show significant variation coordinates its reproductive cycle to on a yearly basis and may cause onset of lunar periodicity. However, spawning certain phases of development at in- is only the final stage in a complex appropriate times if it is used as the process of gamete (eggs or sperm) de- main cue. velopment and maturation, and in Cueing to environmental cycles in order for the worms to be able to spawn, order to synchronize reproduction is the gametes have to be ready at the ap- widespread in invertebrates, but some propriate time. This too requires precise of the best examples are seen in the ma- synchronization. rine worms in the class Polychaeta. The most commonly used cues to Ragworms (genus Nereis; also called regulate breeding are environmental: sandworms or clamworms in North photoperiod and temperature. Of these, America) are widespread in intertidal photoperiod (length of day) is the most areas across the Northern Hemisphere. accurate predictor of time of year and it Up to twelve inches (thirty centimeters) is often used to entrain an animal’s in- long, they are found mostly in muddy

Intertidal mudflats may appear featureless and barren, but under the surface is a rich community of invertebrates waiting for the returning tide. England’s Humber Estuary, photographed by Alan Clements.

20 WINGS Ragworms in the genus Nereis are agile predators when the tide covers their mudflats. Photographed in the North Sea by Edgar Donkervliet. estuarine areas and may live for one to trolled, not only within the individual three years in temperate waters. When but also across the population. the tide goes out they retreat to U- or J- The two most important factors shaped burrows in the sand or mud. Dur- controlling the development of eggs ing high tide they emerge from their bur- and sperm in ragworms are water tem- rows as active, fast-swimming predators. perature and photoperiod. In the king Ragworms are among the most im- ragworm (Nereis virens), for example, egg portant food sources for shorebirds and development is initiated by the switch are preferred prey in Britain for birds to short winter days — a day length of such as dunlin, bar-tailed godwit, grey eight hours is essential to promote egg plover, and redshank. With burrow den- growth — but it is temperature that con- sities that may exceed several hundred trols the speed of egg development. Low per square yard, ragworms are a huge temperatures speed up development so food resource, and consequently some that eggs starting to develop later in the estuaries are of national or internation- winter grow faster at the lower temper- al importance as migratory stopovers for atures they experience and catch up to birds. The Humber Estuary, for example, those eggs that began development dur- on the east coast of England, is one of ing the milder conditions earlier in the Europe’s key sites for wintering wildfowl season. All eggs are ready for spawning and may host more than 175,000 birds. in early spring. Its importance is recognized worldwide. Temperature has also been linked Ragworms are different from many with spawning. For example, the rag- other polychaetes in that they are sem- worm Nereis diversicolor spawns in the elparous — that is, breeding only once, spring after a period of low winter tem- after which the adult dies. Synchroniz- peratures. In this species it seems that ing the timing of reproduction is there- the rise in temperature from winter fore essential and must be tightly con- to early spring is highly important in

FALL 2007 21 aiding the synchronization of gamete but only in those animals that have ex- maturation and spawning. Further syn- perienced at least fifty short days. chrony is imposed as mature animals re- Scaleworms and ragworms, along lease their gametes only with the spring with many other marine creatures, rely tides, which occur on the semi-lunar on a delicate balance of day length cycle of 14.8 days and coincide with the and temperature to control breeding. If full and new phases of the moon. one of these two environmental cues Scaleworms (genus Harmothoe) are fails them, the worms may not be able another group of polychaete worms to breed successfully. If an individual that utilize photoperiod and water tem- misses the spawning period, its indi- perature to synchronize reproduction. vidual genes may be lost from the pop- Like ragworms, they are widely distrib- ulation forever. If a population misses uted across the Northern Hemisphere the spawning period, then the conse- and are an important food source for quences will be more severe, not just for shorebirds, but they differ from rag- the worms themselves but also for the worms in behavior, habitat, and life other animals that are reliant on them cycle. They are sit-and-wait predators of as a seasonal food source. Global warm- rocky shores, using cryptic coloration to ing makes this more than just a possi- hide in crevices or under stones or sea- bility. The United Nations’ Intergovern- weed holdfasts, and are iteroparous, mental Panel on Climate Change pre- which means they reproduce once a dicts higher surface temperatures, with year and may live for several years. the change 50 percent higher during One species, Harmothoe imbricata, winter in northern temperate regions actually produces two batches of eggs than elsewhere, so it is likely that the each year, with the spawning of these relationship between photoperiod and occurring just weeks apart. The process temperature will be broken or fall out begins in autumn when the first batch of phase. of eggs grows during the winter before For the species discussed here, the being spawned in March. After this, the impact of climate change could be dra- second batch of eggs develops rapidly matic. The photoperiod cues will remain and is spawned approximately three the same, so gamete development will weeks later. Spawning is linked to cal- be initiated and spawning will occur at endar date and H. imbricata exhibits an the same time, but higher winter tem- annual reproductive cycle in which peratures will slow the rate of develop- photoperiod sets the clock. The animals ment, leading to a significant reduction must be exposed to at least six weeks of in the number and quality of gametes continuous short days (thirteen hours ready to spawn. Dissynchronous spawn- or less of daylight) for successful egg de- ing between individuals has been seen velopment. Falling temperatures during in populations of Nereis diversicolor this period also help to synchronize ga- maintained in constant high tempera- mete progression to maturity. A final tures. It may also lead to the time of rapid growth phase is accelerated by low spawning becoming uncoupled from temperatures combined with longer day the best time of year for larval survival. lengths as winter transitions to spring, While this may not ultimately lead to

22 WINGS species extinctions, it could lead to the vertebrates is likely to go beyond just local extinction of some populations or the individual species. Because poly- the significant collapse of populations chaetes are one of the most important from some areas. Their survival will de- components of the estuarine sediment pend on the speed at which they can community, loss of these animals may adapt to climate change. have a significant impact on the ecol- Recent events in the seas around ogy of these systems. For some estuar- northwest Europe have shown that ies, any significant loss of invertebrate changes in environmental conditions biomass could result in the loss of bird during the winter can have significant populations. impacts on marine invertebrate repro- So finally, going back to your big duction. Ecological responses to the night on October nineteenth, would climatic effects of the North Atlantic you have made the meeting? Do you Oscillation have included changes in think that you would have noticed that the timing of reproduction, population this is the night of the third quarter of dynamics, abundance, and spatial dis- the moon in North America? I suspect tribution of marine invertebrates. In ad- that most of us would miss the date. dition, unseasonable climatic events in the Baltic Sea have lead to high losses of shearwaters due to a collapse in their Andrew Lawrence has been studying ma- food supplies. rine worms and impacts of climate change The structure and function of ma- for more than twenty years. He is a senior rine ecosystems are intimately linked lecturer in marine ecology at the University to the atmosphere, and the impact of of Hull, a couple of miles from the north climate change on these and other in- shore of the Humber Estuary.

Invertebrates of intertidal areas provide a vital food resource for mi- grating or overwintering shorebirds. Photographed at the Humber Estuary, England, by Paul Glendell; courtesy Natural England.

FALL 2007 23 Impacts of Global Warming on Pollinators

David Inouye

The global climate is changing. Al- one degree Celsius (one and three-quar- though many of the finer details have ters degrees Fahrenheit) could advance yet to be revealed, the overall picture is first and peak appearance of most but- one that portends trials and changes for terflies by two to ten days. This sort of invertebrate species generally and for change is not limited to butterflies. Al- pollinators specifically. Pollinators will though they are not pollinators, Britain’s face multiple challenges to their sur- dragonflies and damselflies have been vival — many of them have complex life emerging about three and one-third cycles that include dependence on dif- days earlier for every degree Celsius of ferent resources at different stages of warming in temperature since 1960. growth, some have life spans of multi- It also appears that the general ar- ple years, and for others migration is a rival of spring has advanced in recent factor as well. Pollinators seem most decades. Where detailed studies have likely to respond to a warming envi- been conducted on more local scales, ronment in two ways, through changes earlier flowering is a common pattern in phenology (the timing of life-cycle for many species. A danger for pollina- events), and through changes in their tors comes from the loss of synchrony geographic distribution. between their life cycles and those of Phenology of both pollinators and the plants upon which they depend, ei- the plants on which they feed is an im- ther as hostplants or nectar and pollen portant ecological factor that is chang- resources. ing in response to global, regional, and My own research has been focused local climate changes. Key phenological primarily on montane and alpine en- events in pollination, including polli- vironments, particularly in Colorado, nator emergence or arrival and timing where I have worked for almost four of flowering plants, generally appear to decades at the Rocky Mountain Biolog- be happening earlier than in previous ical Laboratory. Here the emergence of years in temperate regions of the world. Milbert’s tortoiseshell (Nymphalis mil- Some of the clearest examples come berti), a butterfly that overwinters as an from Britain. Data from the United King- adult, has not been changing in concert dom’s Butterfly Monitoring Scheme with the flowering of early spring wild- show that the first appearances of most flowers, suggesting a loss of synchrony butterflies have advanced in the last two between their need for nectar and the decades. These changes were strongly availability of their customary floral related to earlier peak appearances and, resources. If this pattern turns out to for multi-brooded species, longer flight be more widespread — if overwintering periods. Models using these data suggest bumble bee queens or emerging solitary that climate warming on the order of bees, beetles, or flies respond differently

24 WINGS than do spring wildflowers to changes One ongoing research project at the in environmental cues (such as snow- Rocky Mountain Biological Laboratory melt dates and warming soil tempera- is an investigation of potential shifts tures in temperate habitats) — it raises in the altitudinal distribution of bum- the possibility that changes to many ble bee (Bombus) species. In 1974 and historical plant-pollinator relationships 1975 Graham Pyke and I were graduate are on the way. students at the Laboratory and both of A second prediction of the conse- us looked at the distribution of these quences of global warming is that some bumble bees along altitudinal transects species will move their ranges to high- that spanned up to about 1,500 verti- er latitudes in the Northern Hemisphere cal feet (460 meters). The bees typically or to higher altitudes in mountains. sorted out by a combination of pro- There is now evidence that some species boscis length (short, medium, long) and of butterflies have already undergone altitude (high, medium, low). Consis- such latitudinal changes, which match- tent with the predictions for global es the observation that some species of warming, we have anecdotal evidence birds, fish, other insects, and plants that species in this habitat are moving have also expanded their latitudinal their ranges up in altitude. This summer distributions. It would be surprising if we brought Dr. Pyke back to the Labo- other pollinators don’t also respond to ratory from Australia, where he now warming temperatures by moving to- works, to help relocate his original tran- ward the poles; there is already evidence sects from 1974. The students who then available of bumble bees moving to re-surveyed the transects last summer higher mountain elevations. observed at least one of the bumble bee

Global warming is changing the timing of flower bloom and but- terfly emergence, in some cases breaking synchrony between species. Milbert’s tortoiseshell (Nymphalis milberti), photographed in Wisconsin by Mike Reese.

FALL 2007 25 species about fifteen hundred feet high- also face problems if gaps develop in the er than we had reason to expect to find temporal sequence of resources they it. The results of our recent census will need. For example, bumble bee colonies provide both insight into any signifi- are started early in the spring by queens cant changes in distribution and an- emerging from overwintering under other baseline for future studies. ground, but new reproductives aren’t Pollinators are dependent on plant produced until the end of the summer, species for nectar and pollen as sources so a whole series of plant species must of carbohydrate and protein, so it makes be available to sustain this long period sense to consider how their interactions of colony development. with plants may be affected by climate In addition to changes caused by in- change. In some cases these are very spe- creasing global temperatures, the effects cific relationships, with insects visiting of increasing carbon dioxide could have only a single species of plant, or perhaps implications for pollinators. In an ex- a group of related species, while other perimental study in Britain, elevated pollinators may be much more catholic carbon dioxide levels had an effect on in their diets. Perhaps the former strat- flowering phenology and nectar pro- egy is the one most at risk under sce- duction of some butterfly hostplants. In narios of climate change, for if the one another experimental study, a change species of plant a pollinator depends on of food-plant preference was found for shifts its distribution, or if its abundance caterpillars under elevated carbon diox- decreases, the pollinator would have to ide, and development took longer with duplicate the range shift or face a cer- increasing levels of carbon dioxide. tain decline. Pollinators dependent on Yet another prediction of some cli- a suite of flowering plant species may mate-change scenarios is that there will

Pollinators are key components of terrestrial ecosystems, yet little research has been done on the impacts of global warming on bees, flies, and bee- tles. A bee fly on valerian, photographed in Colorado by David Inouye.

26 WINGS be increased variability in precipitation. At Stanford University’s Jasper Ridge Biological Station, a nearly four-decade- long study of bay checkerspot butterflies (Euphydryas editha bayensis) found evi- dence that such variability may have amplified population fluctuations, lead- ing to their local extinction. Modeling studies also suggest that rapid extinc- tion could result from precipitation- amplified fluctuations in butterfly pop- ulations. Precipitation is also important for both the abundance of flowering and the abundance of nectar; in drought years plants may lose flower buds and the flowers that are produced may not produce nectar. Over the last three decades the range of at least one species of bumble bee in the To date, butterflies have been the Colorado Rockies has moved fifteen hun- focus of most climate-change and pol- dred feet higher in elevation. Bombus flavi- linator research for a variety of reasons. frons on lupine (Lupinus argenteus), pho- They are easy to observe, there are often tographed in Colorado by David Inouye. good historical records of distribution and abundance, and they seem to res- habitat for pollinator species. If we pro- onate with the public more than many tect habitat where pollinators occur other invertebrate species. But they are today, what will happen in the future as not necessarily representative of all pol- their ranges change? The growing real- linators; it is likely that many less mo- ization of the economic and ecological bile invertebrate species could face even significance of pollinators may help to greater challenges in responding to fu- spur the research that is needed to in- ture changes in temperature, precipita- crease our understanding of the changes tion, and phenology. they are facing and provide better strate- The recent report on the status of gies for protecting these vital creatures. pollinators in North America sponsored by the National Research Council point- ed out that little is known about the David Inouye is a professor of biology at ecology, the population biology, or even the University of Maryland, College Park. the distribution of most pollinators. He has worked at the Rocky Mountain Bi- This lack of information hampers both ological Laboratory since 1971, studying our ability to forecast the changes that hummingbirds, pollination biology, ant- are undoubtedly ongoing and our abil- plant mutualisms, bumble bees, and the ity to plan for conservation. The kinds phenology and demography of wildflowers. of changes predicted in phenology or Inouye was a member of the National Re- range due to climate change present a search Council committee on the status of challenge for the goal of protecting pollinators in North America.

FALL 2007 27 XERCESFEATURE NEWS

Conserving Our Most Important Native Pollinators Seven years ago, bee taxonomists began tion. This coincidental timing suggests to notice a decline in the abundance that an exotic disease is the cause for the and distribution of several very com- widespread loss, although this hypoth- mon bumble bee species across the esis is still in need of validation. United States. Three bee species, Bombus The Xerces Society is collaborating occidentalis (the western bumble bee), B. with a bumble bee expert, Dr. Robbin affinis (the rusty-patched bumble bee), Thorp, to learn the former and current and B. terricola (the brown-tipped bum- ranges of these species. We are soliciting ble bee), were once very common and information from the scientific litera- important crop pollinators across North ture, museum collections, and experts America, but are now only rarely found. across the country to document the A fourth species, B. franklini (Franklin’s bees’ former range and abundance; and bumble bee) may already be extinct. we are conducting background research The decline of these species is cor- on each species’ biology, ecology, and related with a crash in the laboratory role in crop pollination. This informa- populations of commercially raised tion will help determine the best way to bumble bees that were distributed across protect the populations that remain. North America for greenhouse pollina- We are also working to raise public

An introduced disease carried by commercially raised bumble bees may be the cause of decline in native spe- cies, including Franklin’s bumble bee (Bombus franklini). Photographed in Oregon by Dr. Peter C. Schroeder.

28 WINGS awareness about these species, their role of these bees and to solicit information in pollination of crops and natural about their current distribution. areas, and the threats they face. We are In addition to protecting the popu- creating a series of “Wanted” posters to lations of these declining bumble bees, send to bee researchers and entomolo- we hope to prevent the unregulated gy and ecology departments across the transport of bumble bees (and their dis- country, as well as to annual meetings eases) in order to prevent future declines of groups such as the Entomological So- in other bumble bee species. ciety of America and the Society for Funding for this project comes from Conservation Biology. The purpose of the CS Fund. To find out more about the posters is to educate the public and this effort please visit www.xerces.org/ scientific community about the decline bumblebees.

Partnership with the Natural Resources Conservation Service For five years we have been building a We are working with NRCS offices program focused on pollinator conser- across the country to provide pollinator vation in agricultural landscapes. On conservation and management training July 20, 2007, the Xerces Society signed and to help NRCS staff incorporate pol- a memorandum of understanding with linator conservation into existing farm the Natural Resources Conservation Ser- bill programs. This effort has been fund- vice that is helping us to expand our ed by Xerces members together with the outreach to a broader segment of the Turner Foundation and other private agricultural community. foundations.

Xerces Works to Understand and Protect Wetlands In the past century, millions of acres of protected migratory bird species. wetlands in North America have been In the Pacific Northwest, many wet- lost due to agriculture or urbanization. lands are severely compromised but few Many of the remaining wetlands suffer have been studied thoroughly and little from a variety of human impacts, in- information is available on their bio- cluding pollution, changes in water logical integrity (their ability to support flow, and invasive species. life). While there are a variety of tech- Wetlands are important elements of niques available to assess the biological watersheds, providing such valuable condition of rivers and streams, evalua- ecosystem services as water filtration, tion of the biological attributes of wet- flood control, erosion control, and crit- lands and their responses to human dis- ical habitat for animals and plants. turbances has lagged behind. An in- Quality wetland habitat is essential to creasing number of watershed councils native and endangered species; about in the Northwest are taking on wetland half of all federally listed endangered restoration projects, but lack the tools animals rely on wetlands for survival, to assess wetland quality and evaluate including over 50 percent of America’s restoration success.

FALL 2007 29 Sarina Jepsen, Xerces Society senior conservation associate, uses a dip net to collect aquatic invertebrates as part of our effort to develop a bi- ological assessment tool for wetlands. Photograph by Logan Lauvray.

The Xerces Society is working to de- tions drawn from background research velop an invertebrate-based biological and our pilot study have been incorpo- assessment tool that can be used to as- rated into a CD-ROM guide to wetland sess wetland quality. With funding from invertebrate identification in the Pacific the Environmental Protection Agency Northwest. The guide has family-level and the Mountaineers Foundation, we keys for aquatic invertebrates encoun- completed a pilot project to identify the tered in Northwest wetlands, and gen- invertebrates in one class of wetland and eral information on invertebrate mon- have begun developing this assessment itoring in still water habitats. To pur- tool for the Pacific Northwest. Next year chase the CD-ROM Aquatic Invertebrates we hope to expand the project to in- in Pacific Northwest Freshwater Wetlands: clude additional classes of wetland. An Identification Guide and Educational The information and recommenda- Resource, please go to www.xerces.org.

Support Butterfly Conservation: Pledge the Butterfly-A-Thon Throughout 2008, noted lepidopterist Swallowtail Seasons: The First Butterfly Big and writer Robert Michael Pyle will be Year. The project is also a fundraiser, undertaking a historic journey to find, and you can participate in this effort by experience, and identify as many spe- making a pledge for every butterfly cies of butterflies as possible in the Unit- species that Bob encounters, as part of a ed States and Canada. The literary fruits “Butterfly-A-Thon.” All proceeds from of this project will be published by the Butterfly-A-Thon will go towards Houghton Mifflin as a book entitled the Xerces Society’s work to preserve

30 WINGS and protect rare and endangered but- Bob Pyle founded the Xerces Soci- terfly species. ety in 1971. He has published fifteen During his travels, Bob will be seek- books, including such butterfly classics ing to find as many as he can of the as the Audubon Society Field Guide to nearly eight hundred butterfly species North American Butterflies, the Handbook recorded in North America north of for Butterfly Watchers, and The Butterflies Mexico, although the numbers will be of Cascadia, as well as award-winning lit- secondary to his actual encounters with erary works, including Wintergreen and the butterfly fauna. To update partici- Sky Time in Gray's River. pants on his progress, he will be posting To make a pledge or to find out regular updates from the road on a But- more about the Butterfly-A-Thon, please terfly-A-Thon blog. visit www.xerces.org.

New Staff Members at the Xerces Society We are delighted to have two new staff veloping assessment tools for wetlands members at Xerces. of the Pacific Northwest. Celeste Mazzacano joins us as a Lisa Schonberg is our new conser- conservation associate. Celeste holds a vation assistant. She recently complet- Ph.D. in entomology from the Univer- ed her M.S. in entomology at Evergreen sity of Minnesota. She has more than State College, researching ants in the twelve years of experience in research, cloud forests and lowland forests of education, and conservation, and has Costa Rica. Lisa will be working on a va- worked extensively with aquatic inver- riety of pollinator conservation and en- tebrates. At Xerces she is currently de- dangered species projects.

WINGS, Fall 2007 Volume 30, Number 2 Wings is published twice a year by the Xerces Society, an international, non- profit organization dedicated to protecting the diversity of life through the conservation of invertebrates and their habitat. A Xerces Society member- ship costs $25 per year (tax-deductible) and includes a subscription to Wings. Copyright © 2007 by the Xerces Society. All rights reserved. Xerces Exec- utive Director: Scott Hoffman Black; Editors: Scott Hoffman Black, Adair Law, Matthew Shepherd, and Mace Vaughan; Design and Production: John Laursen. Printed on recycled paper. For information about membership and our conservation programs for native pollinators, endangered species, and aquatic invertebrates, contact us: THE XERCES SOCIETY FOR INVERTEBRATE CONSERVATION 4828 Southeast Hawthorne Boulevard, Portland, OR 97215 telephone 503-232-6639 fax 503-233-6794 [email protected] www.xerces.org

FALL 2007 31 Individual coral polyps are usually tiny, but in colonies can form massive stony formations, such as this brain coral (Oulophyllia crispa). Coral reefs are among the first ecosystems to display signs of significant damage due to global warming. Photographed in Tanzania’s Songo Songo archipelago by David Obura.

THE XERCES SOCIETY FOR INVERTEBRATE CONSERVATION 4828 Southeast Hawthorne Boulevard, Portland, OR 97215

Board of Directors Michael J. Bean Piotr Naskrecki May R. Berenbaum Scott E. Miller Paul A. Opler President Robert Michael Pyle Counselors J. Kathy Parker Charles L. Remington Paul R. Ehrlich Vice President Michael Samways Claire Kremen Linda Craig Cheryl Schultz John Losey Treasurer Thomas Lovejoy Scientific Advisors Sacha Spector Jerrold Meinwald Thomas Eisner Secretary Michael G. Morris E. O. Wilson

A $25 per year Xerces Society membership includes a subscription to Wings.

Our cover photograph shows an Apollo butterfly (Parnassius apollo). Butterflies are one of the best studied insects with respect to the impacts of climate change; the range of this species has shifted due to global warming. Photographed in Italy by Angie Sharp.