Professor Arthur Shapiro – Conservation Sense and Nonsense

Monarch Mysteries Update

“I am sick to death of being told you must use natives, especially if a butterfly has no more interest in it than a fire hydrant.” –Professor Arthur Shapiro, Bay Nature, June 2022

Monarch butterfly populations are studied and quantified during the winter, when they are roosting in the shelter of trees, and during the summer breeding season in warmer climates. These studies tell different stories. The breeding population in North America seems to be holding steady since the 1990s in many parts of the country, but the over-wintering population has been steadily dwindling during the same period. As an academic ecologist recently told the New York Times, “’So it’s not really a production problem,’ said Dr. Davis, an author of the new paper. ‘We don’t have fewer monarchs. We have fewer monarchs reaching the wintering colonies.’”

The most recent study of the breeding population of monarchs is based on a huge data set of 135,000 observations in 403 different sites in North America, partly collected by volunteers of the North American Butterfly Association (NABA) annual summer butterfly count since 1993. The analysis of current population trends reveals interesting clues about the future of monarchs and probably many other butterfly species:

The study “used federal data to estimate how much glyphosate was being used in the area around each survey site. They found that in some regions, especially in parts of the Midwest, glyphosate use was associated with declines in abundance.”
“But they also documented a countervailing force: climate change. In the northern part of the United States, increasing temperatures were correlated with increases in monarch abundance. This effect was especially pronounced in the Midwest, suggesting that the warming climate might have partly offset the effects of glyphosate in that region.”

The study of monarch breeding populations in North America found that the Southwest was one of the regions in the US where monarch population declines were greatest. That finding is consistent with the study of academic entomologists, Matt Forister and Arthur Shapiro, of butterfly populations in the West. They analyzed data from over 40 years of counting butterfly populations (including NABA data) to learn that 450 butterfly species in western states have declined 1.6% per year in the past 20 years, for a cumulative total of 25% fewer butterflies. Although there are several factors—such as habitat loss and pesticides—their study determined that the strongest factor was climate change, particularly warmer temperatures in the fall.

Professor Shapiro explained during an interview on KALW why extreme heat is harmful to butterflies, although the reasons have not been proven yet. Monarchs are one of the butterfly species that is dormant during winter months. They breed in spring when temperatures begin to warm and days become longer. Warmer winter temperatures are reducing the length of dormancy, which increases their need for year-around food and weakens them if there is inadequate food. Extreme heat and drought have an impact on plants, reducing available food for all butterflies.

Studies of Migrating Monarchs

How do studies of migrating monarchs compare to studies of breeding populations in North America? There are two major migrations of monarchs in North America. The migration east of the Rocky Mountains spends the winter in Mexico and the migration west of the Rockys spends the winter on the coast of California. Both of the overwintering populations have plummeted since the 1990s until the winter of 2022 when the population stabilized in Mexico and increased substantially in California.

The increase in the California monarch migration was described by Jessica Griffiths in an article published by the Sierra Club’s national magazine. That article is significant for several reasons. The particular roosting site where the population increase was greatest was a eucalyptus grove in Pismo Beach, California: “We are standing in a eucalyptus grove on a small patch of undeveloped land bordered by farms near the town of Pismo Beach, on the central California coast. The air smells faintly of brussels sprouts and compost, with an overlay of something like Vicks VapoRub—the distinct scent of eucalyptus. Griffiths gazes up at the branches and smiles. There are so many butterflies.” The irony is that Jessica Griffiths is the author of a deeply flawed study that claims that monarchs prefer native conifer trees to eucalyptus trees for their winter roost. One wonders if Jessica Griffiths experienced cognitive dissonance as she counted 17,845 monarch butterflies roosting in a eucalyptus grove where only nine monarchs roosted the previous year.

Jessica Griffiths provides an important clue to changes in the monarch migration in the Sierra Club article. She says monarchs roost in the trees until the temperature rises to about 55⁰ Fahrenheit, when their body temperatures rise enough that they can actively seek the nectar they need to survive. She says, “They are basically solar powered,” which is another way of saying they are cold-blooded animals that require the heat of the sun to be active. In the eucalyptus groves that monarchs prefer as their winter roost in California, nectar is close at hand because eucalyptus blooms during winter months, at a time when little else is blooming.

Pismo Beach, November 2021 Source: https://youtu.be/Su2Ma2lUWFY

When the climate changes, entire ecosystems change with it

When days become shorter in the fall, monarchs in California stop breeding and begin their migration to the coast. Breeding resumes when days become longer in the spring. But hours of daylight are not the only determinant of the monarch breeding season. Warmer temperatures at night are triggering the monarch breeding season earlier than in the past. In fact, some entomologists hypothesize that many monarchs are now breeding year around. The presence or absence of milkweed does not trigger the breeding season, which is determined by hours of daylight and temperature.

If the warming climate enables monarchs to breed year around, why would we object? The more monarchs, the merrier, right? Unfortunately, hobbyist naturalists DO object to altering the timing and location of the breeding of monarchs. This is a Letter to the Editor of the Yodeler, the newsletter of the San Francisco Bay Chapter of the Sierra Club:

The author of the letter to the editor of the Yodeler asks us NOT to plant milkweed near the coast or monarch overwintering sites, presumably because she doesn’t want the monarch’s breeding season to begin when and where it has not occurred in the past. The fact is, the climate has changed and monarchs are responding to those changes. Who are we to argue with monarchs about what they need to do to survive?

Bay Nature has published an article about monarchs seen in Marin County during their breeding season, where they have over-wintered in the past, but not bred historically. The warming climate and the availability of perennial tropical milkweed is making Marin County suitable breeding habitat: “A lot of people have this feeling that without the migration, the monarch is nothing,” says James. “That’s not necessarily true. If we got rid of the migration, the butterflies could still continue. For humans, that would be a pity. But in the ecology of things…it’s not that bad.” The author of the article welcomes monarchs to Marin County, “A new Bay Area neighbor, adapting to a changing world, making do with what is available, as we all must.”

The monarch migration is not sacrosanct. Monarch butterflies also live in Central and South America, in the Caribbean, in Australia, and even in parts of Europe and New Guinea. But all of these monarch populations are sedentary, meaning they stay in one place and don’t migrate. If changes in climate enable monarchs to live and breed year around, why would we want to prevent them from doing so?

If monarchs can find what they need year around, why should they be forced to migrate? Migration is physically demanding, depleting the physical resources of an animal. If survival of a species doesn’t require migration, more physical resources are available for other functions, such as increased reproduction or less need for food to fuel the migration. Images of struggling human migrants come to mind. Wouldn’t they all be better off if circumstances at home would enable them to stay home?

A comparable change has occurred in the life cycle of Anise Swallowtail butterflies. Prior to the introduction of non-native fennel to California, Anise Swallowtails bred only once each year because its native host plant—closely related to non-native fennel–was not available during most of the year. Non-native fennel is a perennial plant that is available year-around, making it possible for Anise Swallowtails to breed throughout the year. Thanks to non-native fennel, we enjoy the company of many more Anise Swallowtails. We should not think of the life cycles of plant and animal species as immutable. Rather, they are constantly changing to adapt to changes in their environment and adaptation is what will ensure their survival.

Native vs. Non-native Milkweed?

Hobbyist naturalists also ask that we plant only native milkweed, the host plant for monarch caterpillars. Such restrictive advice is not beneficial to the survival of monarch populations. Although a popular opinion among hobbyists, advice against planting non-native milkweed for monarchs is contradicted by scientific sources:

“…there is little evidence to support the idea that planting Tropical Milkweeds will weaken Monarch populations and NO evidence to support the idea that Tropical Milkweeds are “trapping” Monarchs and stopping them from migrating…” A merican Butterflies, magazine of the North American Butterfly Association
A study of lifespan of monarchs breeding on non-native milkweed compared to native milkweed found that monarchs raised on tropical milkweed (A. curassavica) lived as long or longer than monarchs raised on other species of milkweed. They were less likely to be infected, and once infected, tolerated the infection well. (Leiling Tao et.al., “Disease ecology across soil boundaries: effects of below-ground fungi on above-ground host—parasite interactions,” Proceedings of Royal Society of Britain, 282: 2015.1993.)
An article from the UC Davis Bug Squad says they plant tropical milkweed and two species of native milkweed. Monarchs have a strong preference for tropical milkweed: “In July, we collected 11 caterpillars from the narrowleaf [native] milkweed; we rear them to adulthood and release them into the neighborhood. But in the numbers game, the tropical milkweed won. From July through today, we have collected a whopping 43 eggs or caterpillars from [non-native] A. curassavica. How many from [native] A. speciosa? Sadly, none.”

Hobbyists theorize that tropical milkweed harbors more parasites than native milkweed because tropical milkweed is a perennial plant, which suggests that parasites could accumulate from one year to the next. If gardeners are concerned about the potential for accumulation of parasites, they are advised to cut tropical milkweed back during winter months. Because tropical milkweed is a perennial, it is available for monarch breeding earlier in the spring than annual native milkweed. If monarchs breed earlier in the spring, tropical milkweed accommodates earlier breeding.

How to help monarchs

The future of monarchs is uncertain, just as the future of all life in our changing climate is uncertain. I am betting that monarchs have a future partly because they have survived many changes in the environment for some 50 million years since butterflies evolved from moths. We can best help monarchs by staying out of their way. They would also probably benefit if we would stop destroying their habitat, particularly eucalyptus trees and tropical milkweed.

Vegetation changes as the climate changes and animals follow the vegetation they need as they must to survive. Breeding season of butterflies and other wildlife is also likely to change with the climate. The rebounding monarch population is probably another case of animals moving to find what they need. We should not stand in their way. They know what they need better than we do.

Doug Tallamy’s Blame Game

The fact that insect populations are declining in many places around the world is well known, but the reasons for the decline are not well known. Where there is uncertainty, there is speculation and where there is speculation, there is debate.

Doug Tallamy recently stepped into that debate by publishing a review article about insects and their use of plants. The article is a mind-numbing list of studies that find both positive and negative relationships between insects and non-native plants.

Tallamy contends those studies add up to support for his belief that non-native plants are bad for insects and native plants are good for insects. He suggests that declining populations of native plants should be considered one of the reasons for declining populations of insects, but then he goes one step further. Tallamy suggests that non-native plants are responsible for declining populations of native plants. It follows that Tallamy blames non-native plants for the disappearance of insects.

My interpretation of the studies in Tallamy’s review is different. The studies tell me that there is too much variation in insect-plant relationships to generalize about the relative value of native vs. non-native plants to insects. A more accurate conclusion would be that sometimes insects make a successful transition from a native to a non-native plant—especially in the absence of a native in the same lineage—and sometimes they don’t…or at least they haven’t yet.

Anise swallowtail butterfly is one of many insects that have made a successful transition from a disappearing native plant to an introduced non-native plant in the same lineage. Prior to that transition, swallowtails were able to lay eggs only once a year, when the native was available. The introduced non-native is available year around, which enables the swallowtail to lay its eggs year around. Courtesy urbanwildlife.org

Since evolution is a process and not a historical event, these insect/plant relationships will continue to change. There are many studies that document such transitions and Tallamy cites some of them in his review. Tallamy assumes insects will be forever handicapped, if not killed, by whatever deficiencies there are in the non-native substitute. I assume the insect is more likely to adapt and eventually evolve to cope with those deficiencies. Both our assumptions are just guesses. Tallamy considers nature immutable, while I consider it dynamic. Where Tallamy sees doom and gloom, I see opportunity.

Professor Art Shapiro’s (Distinguished Professor of Evolution and Ecology, UC Davis) assessment of Tallamy’s review article is less equivocal than mine. Keep in mind when reading his assessment that he is far more knowledgeable than I am:

“There is little evidence known to me of alien plants (‘invasives’) competitively displacing natives in ‘communities’ except in highly disturbed environments, except in the case of ‘ecological engineer’ species like Japanese honeysuckle, Himalayan Blackberry, climbing fern in Florida, Purple Loosestrife, etc. — things that drastically alter the ground rules for structuring the vegetation by smothering or prompting fire.
“The use of natives and non-natives by insects has a long and venerable history, going back to T.R.E. Southwood and his comparisons of insect faunas on British trees to Godwin’s history of the British flora, Azevedo’s student study at SF State, etc. — demonstrating overall that enemies accumulate in time on naturalized aliens, but it may be a very slow process if there is no phylogenetic or chemical bridge to their colonization. Experiments using haphazardly-selected species to examine acceptability are basically silly, and very easy to ‘stack’ if one knows one’s phytochemistry.
“As I have repeatedly pointed out, ‘weed’ eradication would lead rapidly to the extirpation of nearly all of the non-tree-feeding urban and suburban butterfly fauna in lowland California (and many other places).”

Why are insect populations declining?

A 2017 study revealed a shocking 76 percent decline in the biomass of flying insects over 27 years in protected areas in Germany. The German study does not offer specific explanations for the significant decline in insects, but it speculates about probable cause: “Agricultural intensification (e.g. pesticide usage, year-round tillage, increased use of fertilizers and frequency of agronomic measures) that we could not incorporate in our analyses, may form a plausible cause. The reserves in which the traps were placed are of limited size in this typical fragmented West-European landscape, and almost all locations (94%) are enclosed by agricultural fields. Part of the explanation could therefore be that the protected areas (serving as insect sources) are affected and drained by the agricultural fields in the broader surroundings (serving as sinks or even as ecological traps). Increased agricultural intensification may have aggravated this reduction in insect abundance in the protected areas over the last few decades.” Presumably “protected areas” in Germany are not landscaped with non-native plants, rendering the use of this study to corroborate Tallamy’s hypothesis irrelevant.

A comprehensive review of 73 reports of declining insect populations around the globe was published in 2019. These studies report the reasons for declining populations: “The main drivers of species declines appear to be in order of importance: i) habitat loss and conversion to intensive agriculture and urbanization; ii) pollution, mainly that by synthetic pesticides and fertilisers; iii) biological factors, including pathogens and introduced species; and iv) climate change. The latter factor is particularly important in tropical regions, but only affects a minority of species in colder climes and mountain settings of temperate zones.” The “introduced species” are usually insects rather than plants.

In a Yale e360 article about Tallamy’s review, one commenter offers his opinion that the over-population of deer and their preference for eating native vegetation is likely a greater threat to native plants than the existence of non-native plants that provide an alternative source of food for deer, thereby reducing predation of native plants. Tallamy seems to agree that deer are a problem for native plants, while rejecting deer as a greater threat to native plants than the existence of non-native plants.

The list of reasons for declining insect populations is long and will probably get longer as more research is done. If the existence of non-native plants is on that list, it is unlikely to be higher on a prioritized list than the pesticides that are being used to eradicate non-native plants. The more herbicide that is used to eradicate non-native plants, the more harm is done to insects.

EPA Biological Evaluation of glyphosate is a black eye for native plant “restorations” that use herbicide

The Environmental Protection Agency has finally published its Biological Evaluation (BE) of the impact of glyphosate products (all registered formulations of glyphosate products were studied) on endangered animals (mammals, birds, amphibians, reptiles, fish, invertebrates) and plants. The BE reports that 1,676 endangered species are “likely adversely affected” by glyphosate products. That is 93% of the total of 1,795 endangered species evaluated by the study. Of the total of 792 critical habitats of endangered species, 759 (96%) were “likely adversely affected” by glyphosate products. Most of those critical habitats probably contain predominantly native plants that are clearly not benefiting from herbicides used to kill their competitors.

Both agricultural and non-agricultural uses of glyphosate products were evaluated by the BE. Although only endangered plants and animals were evaluated by the BE, we should assume that all other plants and animals are likewise harmed by glyphosate because the botanical and physiological functions of plants and animals are the same, whether or not they are endangered. Herbicides, specifically glyphosate products, are used by the majority of projects that attempt to eradicate non-native plants. As a result, the crusade against non-native plants is undoubtedly a far more important factor in the decline of insect populations than their mere existence.

Why are native plant populations declining?

There are many reasons why native plant populations are declining, but there is little evidence that non-native plants are the cause of declining populations of native plants. Many of the causes of declining insect populations are also causes of declining populations of native plants. A recent study reports that 65 taxa of native plants in the US and Canada are thought to be extinct. The study did not report a single case in which the extinction was caused by the existence of non-native plants. Sixty-four percent of extinct plants were single-site endemics. The same drivers cited by recent insect studies appear on the list of causes of plant extinctions. Nearly half of the extinctions occurred more than 100 years ago, long before introduced plants were considered an issue.

Butterfly bush is a host plant of Variable checkerspot butterflies. It is also an important source of nectar for butterflies and bees. It is being eradicated on public land because it is not a native plant. butterflybush.com

My New Year’s Wish

Nature is too complex to be reduced to a single cause for changes in the environment. Human knowledge is insufficient to identify all of the causes. That’s why we make many mistakes when trying to fix a perceived problem in nature. Our own priorities influence our evaluation of changes in the environment. We should not automatically assume that a change is a problem or that it must be reversed.

The existence of novel ecosystems is a case in point. They can as easily be seen as positive as negative. If a native plant or animal is no longer adapted to changes in the environment, such as climate change, we should be grateful that a non-native substitute is capable of tolerating the change. Where some see enemies, others see friends.

I wish you all a very happy New Year in 2021. I can’t wish 2020 a fond farewell. I can only say good riddance! I am hopeful for a more peaceful year, one in which we befriend our enemies and work together for a better world for nature and for humanity. I am grateful for your readership.

Eradicating non-native plants does NOT benefit insects

We briefly reactivate the Million Trees blog to publish an interesting and important debate between Jake Sigg and Professor Art Shapiro about the relationship between insects and native plants. Their debate was initiated by this statement published in Jake Sigg’s Nature News on April 26, 2019:

“Did you know that 90 percent of insects can only eat the native plant species with which they’ve co-evolved?”

Jake Sigg has been the acknowledged leader of the native plant movement in the San Francisco Bay Area for 30 years. He is a retired gardener for the Recreation and Parks Department in San Francisco. Art Shapiro is Distinguished Professor of Ecology and Evolution at UC Davis. He has studied the butterflies of Central California for 50 years.

Jake and Art are both passionately committed to the preservation of nature, but their divergent viewpoints reflect their different experiences. Jake’s viewpoint is based on his personal interpretation of his observations. As a gardener, his top priority is the preservation of plants rather than the animals that need plants. As a scientist, Art’s viewpoint is based on empirical data, in particular, his records of plant and butterfly interactions over a period of 47 years as he walked his research transects about 250 days per year. The survival of butterflies is Art’s top priority.

Although their discussion is informative, it does not resolve the questions it raises because Jake and Art “agree to disagree.” Therefore, Million Trees will step into the vacuum their discussion creates to state definitively that it is patently false to say that “90% of insects can only eat native plants.” That statement grossly exaggerates the degree of specialization of insects and underestimates the speed of adaptation and evolution.

There are several reasons why insects do not benefit from the eradication of non-native plants:

Insects use both native and non-native plants.
Pesticides used to eradicate non-native plants are harmful to both plants and insects as well as the entire environment.
There is no evidence that insects are being harmed by the existence of non-native plants.

Insects use both native and non-native plants

This statement was recently made in an article published by Bay Nature magazine about Jake Sigg: “More than 90 percent of all insects sampled associate with just one or two plant families.” (7,500 insect species were sampled by the cited study. There are millions of insect species and their food preferences are largely unknown.) This exaggerated description of specialization of insects seems the likely origin of the subsequent, inappropriate extrapolation to the statement that specialized insects require native plants.

Anise Swallowtail butterfly in non-native fennel. Courtesy urbanwildness.org

There are over 600 plant families and thousands of plant species within those families. Most plant families include both native and non-native plant species. An insect that uses one or two plant families, is therefore capable of using both native and non-native plant species.

We will use the Oxalidaceae plant family to illustrate that insects can and do use both native and non-native plants. Oxalidaceae is a small family of about 5 genera and 600 plant species. We choose that family as an example because Jake Sigg’s highest priority for eradication is a member of that plant family, Oxalis pes-caprae (Bermuda buttercup is the usual common name). In a recent Nature News (April 9, 2019), Jake explained why: “Oxalis is not just another weed; this bugger has a great impact on the present and it will determine the future of the landscapes it invades.”

Five members of the Oxalis genus in the Oxalidaceae family are California natives. An insect that uses native oxalis can probably also use the hated Bermuda buttercup oxalis because they are chemically similar.

Honeybee on oxalis flower, another non-native plant being eradicated with herbicide.

The consequences of eradicating non-native plants

Partly because of Jake’s commitment to eradicating non-native oxalis, San Francisco’s Recreation and Parks Department has been spraying it with herbicide for 20 years. Garlon (triclopyr) is the herbicide that is used for that purpose because it is a selective herbicide that does not kill grasses in which oxalis usually grows. Garlon is one of the most toxic herbicides available on the market. More is known about Round Up (glyphosate) because it is the most widely used of all herbicides. However, according to a survey of land managers conducted by California Invasive Plant Council in 2014, Garlon is the second-most commonly used herbicide to eradicate non-native plants.

Garlon is toxic to bees, birds, and fish. It is an endocrine-disrupter that poses reproductive and developmental risks to female applicators. It damages the soil by killing mycorrhizal fungi that are essential to plant health by facilitating the transfer of nutrients and moisture from the soil to plant roots.

A recent article in the quarterly newsletter of Beyond Pesticides explains that insecticides are not the only killers of insects: “Insecticides kill insects, often indiscriminately and with devastating consequences for biodiversity, ecosystem stability, and critical ecosystem services. Herbicides and chemical fertilizers extinguish invaluable habitat and forage critical to insect survival. Taken together, insecticides, fungicides, herbicides and chemical fertilizers make large and growing swaths of land unlivable for vast numbers of insect species and the plants and animals they sustain.” The loss of insects where herbicides are used to kill non-native plants are undoubtedly contributing to the failure of attempts to “restore” native plants which require pollinators and insect predator control as much as non-native plants.

In other words, eradicating non-native oxalis is damaging the environment and the animals that live in the environment. Furthermore, after twenty years of trying to eradicate it, Jake Sigg admits that there is more of it now than there was when this crusade began: “Maybe you’ve noticed that there’s more and more of it every year, and fewer and fewer other plants. That is unlikely to reverse.” (Nature News, April 9, 2019).

coyote in oxalis field. Copyright Janet Kessler

In fact, local failure of eradication efforts mirrors global failures of similar attempts: “…despite international policies aimed at mitigating biological invasions, the implementation of national- and regional-scale measures to prevent or control alien species has done little to slow the increase in extent of invasions and the magnitude of impacts.” (1)

Update: The California Invasive Plant Council has published “Land Manager’s Guide to Developing an Invasive Plant Management Plan.” It says very little about the disadvantages of using herbicides to eradicate plants they consider “invasive” other than a vague reference to “unintended consequences,” without discussion of what they are or how to avoid them.

However, it does give us another clue about why eradication efforts are often unsuccessful. When herbicides are used repeatedly, as they have been in the past 20 years, weeds develop resistance to them: “The International Survey of Herbicide Resistant Weeds (2018) reports there are currently 496 unique cases (species x site of action) of herbicide-resistant weeds globally, with 255 species…Further, weeds have evolved resistance to 23 of the 26 known herbicide sites of action and to 163 different herbicides.” The Guide therefore recommends that land managers rotate herbicides so that the “invasive” plants do not develop resistance to any particular herbicide. The Guide gives only generic advice to use “herbicide X” initially and “herbicide Y or Z” for subsequent applications.

In other words, the California Invasive Plant Council continues to promote the use of herbicides to kill plants they consider “invasive.” They give advice about ensuring the effectiveness of herbicides, but they do not give advice about how to avoid damaging the soil, killing insects, and harming the health of the public and the workers who apply the herbicides.

Do insects benefit from eradicating non-native plants?

There is no question that insects are essential members of every ecosystem. They are the primary food of birds and other members of wildland communities. They perform many vital functions in the environment, such as consuming much of our waste that would otherwise accumulate.

The Economist magazine has reported the considerable evidence of declining populations of insects in many places all over the world. (However, the Economist points out that the evidence does not include large regions where insect populations have not been studied. The Economist is therefore unwilling to conclude that the “insect apocalypse” is a global phenomenon.) The report includes the meta-analysis of 73 individual studies that describe declines of 50% and more over decades. The meta-analysis concluded that there are four primary reasons for those declines, in order of their importance: habitat loss, intensive farming, pesticide use, and spread of diseases and parasites. The existence of non-native plants is conspicuously absent from this list of threats to insect populations.

In other words, although the preservation of insects is extremely important, there is no evidence that the eradication of non-native plants would benefit insects. In fact, eradication efforts are detrimental to insects because of the toxic chemicals that are used and the loss of the food the plants are providing to insects.

Jake Sigg and Art Shapiro discuss insects and native plants

The discussion begins on April 26, 2019, with this statement published in Jake’s Nature News:

“Did you know that 90 percent of insects can only eat the native plant species with which they’ve co-evolved?”

On April 26, 2019, Arthur Shapiro wrote:

“No, I didn’t know 90% of insects can only eat the native plants with which they’ve co-evolved. I’ve only been studying insect-plant relationships and teaching about them for 50 years and that’s news to me, especially since on a global basis we don’t know what the vast majority of insects species eat, period! That’s even true for butterflies and moths, which are probably the best-studied group. And it’s even true here in California, one of the best-studied places on the planet (though way behind the U.K. and Japan). Where on earth did that bit of non-information come from?”

Jake Sigg responds:

“Art, I did my best to run down source for that statement. As I suspected, it may lack academic precision. That kind of precision is hard come by, and what exists is not entirely relevant. Most of the information comes from Doug Tallamy. But the statement is not accurate; it should have read “…90 percent of plant-eating insects eat only the native plants they evolved with”. Whether that is true or not I don’t know, but it accords with my understanding and I am willing to go along with it, even if proof is lacking. If you wait for scientific proof on everything you may wait a long time and lose a lot of biodiversity. I have had too much field experience to think that exotic plants can provide the sustenance that natives do.

I expect you will be unhappy with this response.”

On May 2, 2019, Art Shapiro replies:

“If Tallamy said “90% of the plant-eating insects that I have studied…” or “90% of the plant-eating insects that have been studied in Delaware…” or some such formulation I might take him more seriously. The phenomenon of “ecological fitting,” as described by Dan Janzen, is widespread if not ubiquitous. “Ecological fitting” occurs when two species with no history of coevolution or even sympatry (co-occurrence) are thrown together and “click.” A.J.Thorsteinson summed up some 60 years ago what is needed for an insect to switch onto a new host plant: the new plant must be nutritionally adequate, possess the requisite chemical signals to trigger egg-laying and feeding, not possess any repellents or antifeedants and not be toxic. That set of circumstances is met very frequently. To those of us who study it, it seems to happen every other Tuesday. As we showed, the urban-suburban California butterfly fauna is now overwhelmingly dependent on non-native plants. The weedy mallows (Malva) and annual vetches (Vicia) are fed upon by multiple native butterfly species and are overall the most important butterfly hosts in urban lowland California. . Within the past decade, our Variable Checkerspot has begun breeding spontaneously and successfully on Butterfly Bush (Buddleia davidii). The chemical bridge allowing this is iridoid glycosides. When I was still back East I published that the Wild Indigo Dusky Wing skipper, Erynnis baptisiae, had switched onto the naturalized European crown vetch (Coronilla varia) which had converted it from a scarce and local pine-barrens endemic to a widespread and common species breeding on freeway embankments. And the hitherto obscure skipper Poanes viator, the Broad-Winged Skipper, went from being a rare and local wetland species best collected from a boat to becoming the most abundant early-summer butterfly in the New York metropolitan area by switching from emergent aquatic grasses and sedges to the naturalized Mesopotamian strain of Common Reed, Phragmites australis. I can go on, and on, and on. If you find a sponsor for me to give a lecture about this in the Bay Area, I’ll gladly do it. If you promise to come!

I won’t snow you under with pdfs. Here’s just one, a serendipitous one that resulted from my walking near Ohlone Park in Berkeley. And one from the high Andes in Argentina. That paper cites one of mine in Spanish demonstrating that the southernmost butterfly fauna in the world, in Tierra del Fuego and on the mainland shore of the Straits of Magellan, is breeding successfully on exotic weeds.-! Copy on request.”

On May 2, 2019, Jake Sigg published his last reply:

“I believe many of your statements, Art, and many of these cases I am familiar with. A conspicuous local example is the native Anise Swallowtail butterfly that still lays eggs on native members of the Umbelliferae, the parsley family, but which also breeds on the exotic fennel, which is an extremely aggressive weed that in only a few years can transform a healthy and diverse grassland supporting much wildlife into a plant monoculture—that, btw, won’t even support the butterfly, which shuns laying eggs where its larval food plant is too numerous and easy target for a predator, like yellow jackets.

What puzzles me is why you can keep your equanimity at the prospect of losing acres of very diverse habitat to a monoculture of fennel. You live in the heart of the world’s breadbasket where for hundreds of miles both north and south there are almost no native plants except those planted by humans. That would tend to distort one’s view. I don’t mean to be flip, but it is not normal for even an academic to be indifferent about a loss of this magnitude. I have worked hands-on on the land (I was raised on a ranch) all my life and still work every Wednesday maintaining our natural habitat in San Francisco—a task that hundreds of citizens pitch in on because they value the quality and diversity of the areas. And why do you remain indifferent, are you just a contrarian? You cite examples to bolster your view, but the examples are too small a percentage to be meaningful and wouldn’t stand up against a representative presentation.

I got my view from life. I type this in my second-floor sunroom, which looks into a coast live oak growing from an acorn I planted in the late 1960s, about 50 years ago and which is immediately on the other side of the window. It is alive with birds of many different species—flocks of bushtits, chickadees, juncos every day (plus individuals of other species), which species-number balloons in the migratory season. What I can’t figure out is how the tree can be so productive as to stand up to this constant raiding. I will take instances of this sort as my guide rather than the product of academic lucubrations. And I will throw in Doug Tallamy; the world he portrays is one I recognize and love.

I think our battle lines are drawn. This discussion could go on, as we have not even scratched the surface of a deep and complex subject. But will either of us change our minds? No.”

“Jake Sigg: N.B. Art responded with another long epistle, not for posting. It clarified some of the points that were contentious and seemed to divide us. We differ, but not as much as would appear from the above discussion.”

(1) “A four-component classification of uncertainties in biological invasions: implications for management,” G. LATOMBE , S. CANAVAN, H. HIRSCH,1 C. HUI, S. KUMSCHICK,1,3 M. M. NSIKANI, L. J. POTGIETER, T. B. ROBINSON, W.-C. SAUL, S. C. TURNER, J. R. U. WILSON, F. A. YANNELLI, AND D. M. RICHARDSON, Ecosphere, April 2019.

Dr. Arthur Shapiro: Composition of ecological communities is dynamic

We are republishing an article from the San Francisco Forest Alliance with permission. This excellent summary of Professor Shapiro’s presentation was approved by him. Professor Shapiro is an expert on the butterflies of California. We have had the great pleasure of attending some of his lectures at UC Davis. He is a gifted educator with a deep and profound knowledge of ecology.

Recently, UC Davis Professor Art Shapiro gave a talk at the Commonwealth Club. It was a tour-de-force. He described it as a very quick resume of a course he’s been teaching for 40 years at UC Davis.

The takeaway: The conventional wisdom about ecology is often wrong.

[You can listen to the one-hour audio recording of his talk HERE.]

SPECIES THROWN TOGETHER

Nativists idealize an ecosystem as a community of plants, animals, fungi, and other organisms that have evolved together over many thousands of years in a particular place so that they fit like a complicated jigsaw – the balance of Nature. (We’ve heard them use phrases such as “lock and key” to describe the effect of this co-evolution.) When non-native and invasive species enter, nativists believe, they destroy this intricate mechanism, resulting in an impoverished and simplified ecosystem with fewer species and no natural balance – and even the dire possibility of ecosystem collapse. They talk in terms of plants and animals that “shouldn’t be there” – usually, immigrant species brought in by humans.

But it’s not often true. What the scientific data show is that “communities” of that interdependent kind are unusual. Instead, most ecosystems are groups of plants and animals that happen to be in a place where they can thrive. When they interact, it’s usually because of “ecological fitting” – they can use the other plants and animals in that area to help them survive. Depending on how ancient they are, communities may include tightly co-evolved mutually interdependent multispecies systems. But these make up only a fraction of the community as a whole.

Anise swallowtail butterfly breeds on fennel

Here’s the evidence against the concept of tight-knit interdependent “communities”:

1. There’s no functional difference between a heritage ecosystem and one with exotic species. If there was, scientists should be able to tell an undisturbed “community” from an invaded one without knowing its history. In fact, they can’t. There are no consistent functional differences once an “invading” species has been established. Some ‘invaders’ can drastically transform the systems they enter – an example is cheatgrass in the Western deserts, which greatly amplified fire risk there. But most do nothing of the sort.

2. Species recolonize open land at different rates. “Species move, communities don’t.” If a landscape is wiped clean – say by glaciers or a volcanic eruption – nature begins to move back in almost immediately. The pollen record allows scientists to understand which species of trees arrived at which time. It shows that tree-species move individually, not as communities.

3. Species that now don’t exist in the same place did so in the past, which would not be true if plants and animals normally lived in fixed communities. One example: the wood turtle and the southern toad are not found in the same areas now – but the fossil record shows that in the past, the ranges did overlap. This couldn’t have happened if they needed to be part of different communities. Vast areas were occupied in the past by “no-analogue” communities – ones that simply don’t exist anywhere at all today.

He ended by pointing out that we – humans – are an invasive species. So are most things, at least at one time.

Read on for detailed notes from Professor Shapiro’s talk at the Commonwealth Club.

Again, you can listen to an audio-recording of the whole talk HERE (on the Commonwealth Club website).

————————————————————————

NOTES FROM ART SHAPIRO’S TALK:

ECOLOGICAL COMMUNITIES AND THE MARCH OF TIME

(The talk was dedicated to Prof Shapiro’s late neighbor, Steven Warnock.)

commonwealth club motto — Commonwealth Club motto

The talk was in three parts: The first laid out the historic context for two opposing schools of thought about ecology. The second examined the data, and concluded that the evidence supports Gleason. The third part looked at the future, which includes climate change.

ECOLOGICAL “COMMUNITIES” OR “ASSEMBLAGES” ?

Here’s the conventional wisdom about ecology, associated with Frederic Clements: Plants, animals, insects, fungi and microscopic creatures form interdependent groups, or “communities.” The process by which this happens is “co-evolution” (sometimes described as evolving a “lock and key”), leading to an ecology where all the species fit together like a jigsaw puzzle. (“Co-evolution” is associated with Peter Raven and Paul Ehrlich, who described it in butterflies and plants that evolved together.) If an area is disturbed, it will go through a predictable process of “natural succession” that will lead to a stable “climax” situation, with all its species again interacting as a community.

This stable ecosystem is sometimes called “the balance of nature.” Tamper with it, this theory says, and you could destabilize the whole community, even leading to ecological collapse.

The opposing view, associated with Henry Gleason, is that plants and animals do not necessarily form ecological communities. Instead, groupings or “assemblages” of plants and animals occur mainly by accident. They happen to arrive in that space at that time, and find conditions that allow them to survive and thrive. The species in such an assemblage will interact, not because they co-evolved, but because they find an opportunity to do so.

These theories about how species fit within an ecosystem were quantified when several ecologists – including the famous Robert McArthur – introduced mathematical models that looked at populations of plants and animals and their interactions. They used these models to look at Species Packing – i.e., how many different species of plants and animals could live in a particular ecosystem. Assembly rules says that the distribution of plant and animals species in a given area isn’t random: both competition and cooperation between plants and animals affect what you find. Competing species can’t all live in the same area, but their niches can overlap. Where they do overlap, the two species may evolve more differences (“character displacement“) so they compete less. These mathematical models assumed a condition of equilibrium, i.e. stability. Opponents have argued that ecological niches are seldom stable because the physical environment is not stable for very long.

R.H. Whittaker introduced the idea that the levels of dependency could vary within communities. For any two species, you could assign a number: +1 meant that the species needed each other to survive; -1 meant that they could not live in the same space. He speculated that these relationships tended to be distributed in a bell curve – meaning that most species in a group didn’t depend on the presence or absence of another species. But some subsets of the community were tightly integrated.

How adaptable are living things? They can evolve, but only in certain ways. Niche conservatism is the idea that most species cannot change very much or very fast in response to changes in their ecological niches.

The idea of co-evolution was fine-tuned with John Thompson’s concept of Geographic Mosaics. Co-evolution between two species can happen differently in different geographic areas. So, for example, a plant in one place might depend totally on one insect for pollination, but elsewhere, the same species of plant might find alternative pollinators available. Such Fine-Tuning is the opposite of Niche Conservatism – and both occur in Nature.

TESTING THESE IDEAS

Cladistics (i.e., the system of showing how related species evolved from common ancestors) provided a way to test the Ehrlich-Raven co-evolution hypothesis. If one kind of animal or plant developed into a separate species (“speciation”) then did the plants and animals depending on it also co-evolve into a separate species? There was no evidence that this happened. Co-evolution was a lot sloppier and more unpredictable than that!

Every time you see two organisms working together, it doesn’t necessarily mean they are co-evolved. Dan Janzen, a great tropical ecologis, pointed out that organisms could be taking advantage of niches and resources that appear through Ecological fitting, with no history of coevolution at all. We see this happen when introduced pests attack native plants, and native insects attack introduced plants, forming brand-new associations. It happens all the time.

pacific reed grass under eucalyptus — Pacific reed grass thrives in eucalyptus fog drip

Is there really a difference between “intact” ecosystems and ones that are disturbed or invaded? Mark Sagoff pointed out that if there really is a functional difference between “invaded” and “co-evolved” ecosystems, then scientists should be able to tell them apart without knowing their history.

“The theory that evolutionary processes structure ecosystems and endow them with a mathematical organization (e.g., rule-governed patterns that ecologists can study) has the following implication….scientists should be able to tell by observation whether a given ecosystem is heavily invaded or remains in mint condition…

“In fact, once non-native species have become established, which may take only a short time, ecologists are unable by observing a system to tell whether or not a given site has been heavily invaded. Invaded and heirloom systems do not differ in pattern or process, structure or function, in any general ways.”

There’s more evidence against the idea of stable interdependent communities as the norm in nature. For example: pollen core data shows that trees recolonizing lands after glaciation don’t move in “communities.” The tree species migrate at different rates. Only those species that have mutualistic relationships move together (for example, mycorrhizae and trees).

“Communities” are like still shots from a movie. They show a set of relationships at a particular point in time. That doesn’t mean the relationships are stable or unchanging. Many species show different sets of relationships in the past. For instance: at present, the wood turtle and the southern toad have completely different ranges – but in the past, those ranges overlapped in places. In the UK, workers building Hadrian’s Wall nearly 2000 years ago left middens that have remains of beetles of species now found only in Lapland. When an event that wipes out an ecosystem and recolonization starts, it takes trees 50-100 years to leave a pollen signature. Bufo_terrestris public domain Norman Benton sm Beetles that can fly get there in months to a few years.

If communities were stable groupings of interdependent co-evolved species, then we would expect to see the same communities repeated in similar conditions. But in fact, we often see different groupings in similar conditions.

THE FUTURE

Decisions about what ecosystems should look like are subject to human preferences. For most people, what they grew up with is “normal.” But the world has changed. The climate is changing. The pool of available species has increased enormously. In terms of trying to “restore” an earlier ecosystem – there’s no going back.

What is “biodiversity?”

We are republishing an article from the San Francisco Forest Alliance with their permission and the permission of the author of the article, Professor Arthur Shapiro, UC Davis.

There’s been a lot of talk of ‘biodiversity’ in San Francisco recently. The city’s ‘Recreation and Open Space Element’ (ROSE) mentions it without clearly defining it. The Natural Areas Program claims to preserve it. There’s a new position, the Biodiversity Coordinator (currently Peter Brastow, formerly of Nature in the City) within San Francisco’s Department of the Environment.

One of our readers, puzzled by all the discussion, asked a simple question of UC Davis Professor Arthur Shapiro, who gave a talk at the Commonwealth Club a few days ago. Instead of the two-line answer they expected, he sent this detailed response — which he kindly permitted us to publish.

WHAT IS BIODIVERSITY? BY ARTHUR M.SHAPIRO
A buzzword. Biodiversity means whatever you want it to mean. I hate the word. Here’s why.

The following is from the introductory biology textbook we use at U.C. Davis, Life: The Science of Biology, (10th edition, Sadava et al., p.1229 — yes, I said p. 1229!):

“…the term BIODIVERSITY, a contraction of ‘biological diversity,’ has multiple definitions. We may speak of biodiversity as the degree of genetic variation within a species….Biodiversity can also be defined in terms of species richness in a particular community. At a larger scale, biodiversity also embraces ecosystem diversity–particularly the complex interactions within and between ecosystems….One conspicuous manifestation of biodiversity loss is species extinction…”

Got that?

The glossary at the back of the book defines “biodiversity hot spot” (itself ambiguous, conflating numbers of species and degree of endemism), but NOT biodiversity itself. One can see why.

Where did this verbal monstrosity come from?

Heliconius mimicry. Creative Commons Generic 2.0

The raw number of species in a defined area or system – what many of us call “species richness“–is a useful number. There are more species of butterflies in Brazil than in California, and more in California than in Alaska. That is true even if we pro-rate species number by area, and it is not trivial to ask why.

But there is more to biodiversity than mere numbers of species. Ecologists are also interested in how individuals are divided among species, that is, the distribution of commonness and rarity among species. You can have a “community” consisting of exactly two species. It could have, say, 50 individuals of each species, or it could have 99 of one and 1 of the other–or any ratio in between. Does this matter? Why? What can those numbers tell us?

QUANTIFYING DIVERSITY – A DIVERSITY INDEX

A century ago a Danish plant ecologist named Christen Raunkiaer observed that there was a statistical regularity to this; he called it the “law of frequency.” In subsequent years it was found to hold for bird censuses and moths collected at lights, as well as for old-field plants. A whole series of mathematical models developed over the years attempted to account for this regularity by means of assumptions about how species interacted–competing for resources, for example. These exercises were at the core of community ecology for several decades, and were seen as immensely important.

During World War II an applied mathematician named Claude Shannon, working on war-related communications problems at Bell Labs, developed a formula that concisely expressed the information content of a message. Ecologists discovered the Shannon formula in the 1960s and realized it could easily be adapted to give a single number that combined the number of species in a community and their relative abundances.

Thus whole communities could be compared efficiently, a potentially informative and useful tactic in trying to understand how multispecies systems worked. The number generated by the Shannon formula came to be called diversity, and the formula became the first and most widely-used of several diversity indices. I learned it in high school and I still use it in teaching. Diversity had two components, then:

Species richness and
“Equitability,” (the difference between a 50:50 and a 99:1 community).

And we were off and running. Now everything could be quantified with a diversity index: “foliage height diversity” in a forest canopy, or “aspect diversity” in moth faunas (how many wing shape-pattern themes could be recognized?). The number of uses and abuses of the term multiplied like rabbits. By 1971 things had gotten so bad that a paper was published caustically titled “The nonconcept of species diversity.” It was widely applauded for its candor. Unfortunately, the author ended up inventing his own new measure of diversity–one he thought was better than the old ones.

MORE LEVELS OF ‘BIODIVERSITY’

But things could get worse. And they did. With the passage of the Endangered Species Act, which opened the door to protection of endangered subspecies (keep in mind that there is no concept of the subspecies; a subspecies is whatever some taxonomist says it is) and even “distinct population segments” (no one knows what that means), genetics got in on the diversity game. Now we would not be content with diversity at the species level; we needed to
get inside species.

In the scramble to define what might be protectable, a search was launched for “evolutionarily significant units.” With modern molecular-genetic tools, we quickly learned that taxonomic subspecies may be genomically nearly identical, while organisms indistinguishable by the naked eye may be wildly different. Defining diversity at the genetic level is still, well, challenging.

One very useful dimension of biodiversity is known as alpha, beta and gamma diversity:

Alpha diversity is species richness at the local level.
Beta diversity is a measure of how much the biota of different localities within a region differ among themselves–that is, how quickly species composition “turns over” in space [i.e. when you have many different little ecosystems next to each other].
Gamma diversity is at a large spatial scale.

The Bay Area has phenomenally high beta diversity in almost everything.

THE BOTTOM LINE

So what is biodiversity? It’s species richness, plus the distribution of abundance and rarity, plus the geography of all that, plus the amount of genetic variation in selected species of interest, plus whatever you please.

Somehow or other concepts of “quality” have gotten mixed in, too. When you clear-cut a redwood forest (which has very low species richness), the early-successional communities that develop on the site, which may be dominated by “invasive weeds,” will have both much higher species numbers and a richer distribution of species abundances than the forest they replaced. But early-successional communities don’t get any respect despite being more diverse and despite the supposition that biodiversity is good. Because they’re made up of the ‘wrong’ species–whatever that means.

Because biodiversity, after all, is only a buzzword.

Dunnigan Test Plot, Augusst 2011. The result of an eight-year effort to restore native grassland. Does it look "biodiverse?" ecoseed.com. — Dunnigan Test Plot, August 2011. The result of an eight-year effort to restore native grassland. Does it look “biodiverse?” ecoseed.com.

Addendum: We have recently learned that Peter Brastow, San Francisco’s Biodiversity Program Coordinator, has applied for an Urban Greening Planning Grant for the City of San Francisco to fund the creation of a Biodiversity and Ecology Master Plan. This grant application was submitted to the State of California’s Strategic Growth Council for $250,000.

This Master Plan would “identify land owners at the parcel level” (including your backyard; see below*) for “consolidating ownership and/or management of wild lands and natural areas into as few departments as possible in order to facilitate coherent and higher quality habitat restoration and management.” The Natural Areas Program “serves as a model for extending this work beyond Recreation and Park Department lands.” (Quotes are directly from the grant application.) In other words, the Biodiversity and Ecology Master Plan will extend the work of the Natural Areas Program to all open space in San Francisco, regardless of the current ownership of the land.

The complete document is available here: Biodiversity Action Plan – grant application questions.final. We find this a horrifying prospect. The Natural Areas Program is extremely controversial because it destroys existing habitat, uses large quantities of toxic herbicides, and restricts access to designated trails. Applying these policies into all open space in the city is a bad idea

– Million Trees

*The grant application says, “We will review the potential of rear yard open space, green roofs, green walls, landscaping, street trees, mini and pocket parks, and other urban design potential to enhance biodiversity in the urban landscape.”