Pinus radiata plantation in New Zealand; photo by Jon Sullivan
As countries and conservation organizations ramp up tree planting as one solution to climate change, I worry that many of the plantings will use species not native to the region – with the risk of promoting more bioinvasions. My second fear is that inadequate attention will be paid to ensuring that the propagules thrive.
Warning from New Zealand
New Zealand has adopted a major afforestation initiative (“One Billion Trees”). This program is ostensibly governed by a policy of “right tree, right place, right purpose”. However, Bellingham et al. (2022) [full citation at end of blog] say the program will probably increase the already extensive area of radiata pine plantations and thus the likelihood of exacerbated invasion. They say the species’ potential invasiveness and its effects in natural ecosystems have not been considered.
Bellingham et al. set out to raise the alarm by evaluating the current status of radiata, or Monterrey, pine (Pinus radiata) in the country. They note that the species already occupies ~1.6 M ha; the species makes up 90% of the country’s planted forests. Despite the species having been detected as spreading outside plantations in 1904, it is generally thought not to have invaded widely.
The authors contend that, to the contrary, radiata pine has already invaded several grasslands and shrublands, including three classes of ecosystems that are naturally uncommon. These are geothermal ecosystems, gumlands (infertile soils that formerly supported forests dominated by the endemic and threatened kauri tree Agathis australis), and inland cliffs. Invasions by pines – including radiata pine – are also affecting primary succession on volcanic substrates, landslides on New Zealand’s steep, erosion-prone terrain, and coastal sand dunes. Finally, pine invasions are overtopping native Myrtaceae shrubs during secondary succession. Bellingham et al. describe the situation as a pervasive and ongoing invasion resulting primarily from spread from plantations to relatively nearby areas.
kauri; photo by Natalia Volna, iTravelNZ
The New Zealanders cite data from South America and South Africa on the damaging effects of invasions by various pine species, especially with respect to fire regimes.
Furthermore, their modelling indicates that up to 76% of New Zealand’s land area is climatically capable of supporting radiata pine — most of the country except areas above 1000 m in elevation or receiving more than 2000 mm of rainfall per year. That is, all but the center and west of the South Island. This model is based on current climate; a warmer/drier climate would probably increase the area suitable to radiata pine.
These invasions by radiata pine have probably been overlooked because the focus has been on montane grasslands (which are invaded by other species of North American conifers). [See below — surveys of knowledge of invasive plants’ impacts.]
Bellingham et al. recognize the economic importance of radiata pine. They believe that early detection of spread from plantations and rapid deployment of containment programs would be the most effective management strategy. They therefore recommend
1) taxing new plantations of non-indigenous conifers to offset the costs of managing invasions, and
2) regulating these plantations more strictly to protect vulnerable ecosystems.
They also note several areas where additional research on the species’ invasiveness, dispersal, and impacts is needed.
Survey of Awareness of Invasive Plants
A few months later a separate group of New Zealand scientists published a study examining tourists’ understanding of invasive plant impacts and willingness to support eradication programs (Lovelock et al.; full citation at end of the blog). One of the invasive plant groups included in the study are conifers introduced from North America and Europe. These conifers are invading montane grasslands, so they are not the specific topic of the earlier article. The other is a beautiful flowering plant, Russell lupine. These authors say that both plant groups have profound ecological, economic, and environmental impacts. However, the conifers and lupines are also highly visible at places valued by tourists. Lovelock et al. explored whether the plants’ familiarity – and beauty – might affect how people reacted to descriptions of their ecosystem impacts.
Visitors from elsewhere in New Zealand were more aware of invasive plants’ impacts and more willing to support eradication programs for these species specifically. Asian visitors had lower awareness and willingness to support eradication of the invasives than tourists from the United Kingdom, Europe, or North America. This pattern remained after the tourists were informed about the plants’ ecological impacts. All groups were less willing to support eradication of the attractive Russell lupine than the conifers.
Conifers invading montane grasslands are perhaps the most publicized invasive plants in New Zealand [as noted above]. Lovelock et al. report that New Zealand authorities have spent an estimated $NZ166 million to eradicate non-native conifers over large tracts of land on the South Island. Still, only about half the New Zealand visitors surveyed were aware of the ecological problems caused by wild conifers.
invasive lupines in New Zealand; photo by Michael Button via Flickr
Russell lupine (Lupinus × russellii) is invading braided river systems, modifying river flows, reducing nesting site availability for several endangered birds, and provides cover for invasive predators. While initially planted in gardens, the lupines were soon being deliberately spread along the roads to ‘beautify’ the landscape. Foreign tourists often specifically seek river valley invaded by the lupine because pictures of the floral display appear in both official tourism promotional material & tourist-related social media. It is not surprising, then, that even among New Zealanders, only a third were aware of the lupines’ environmental impacts.
The oldest participants (those over 60) had the lowest acceptance of wild conifers. Participants 50–59 years old were most aware of ecological problems caused by wild conifers. Participants 30–39 years old showed the highest acceptance of wild conifers and lowest awareness of ecological issues.
Female participants showed a higher preference for the landscape with wild conifers (45.90%) than males (36.89%). Female participants were also half as aware of ecological problems (25.62% v. 46.12% among male participants).
Nearly all survey participants (96.1%) preferred the landscape with flowering lupine; only 19.4% were aware of associated ecological problems. New Zealand domestic visitors were more aware. After the impacts of lupines were explained, half decided to support eradication. However, the same proportion of all survey participants (42.5%) still wanted to see lupines in the landscape.
Once again, participants older than 50 were more aware of ecological problems arising from lupine invasions. Both men and women greatly preferred the landscape with Russell lupins.
While the authors do not explore the ramifications of the finding that younger people are less aware of invasive species impacts, I think they bode ill for future protection of the country’s unique flora and fauna. They did note that respondents had a high level of acceptance overall for these species on the New Zealand landscapes.
While the study supported use of simple environmental messaging to influence attitudes about invasive species, also showed that need to consider such social attributes as nationality and ethnicity. So Lovelock et al. call for investigation of how and why place of origin and ethnicity are important in shaping attitudes towards invasives. Conveying conservation messages will be more difficult because tourist materials often contain photographs of the lupines. Much of this information comes from informal media such as social media, which are beyond the control of invasive species managers.
SOURCES
Bellingham, P.J., E.A. Arnst, B.D. Clarkson, T.R. Etherington, L.J. Forester, W.B. Shaw, R. Sprague, S.K. Wiser, and D.A. Peltzer. 2022. The right tree in the right place? A major economic tree species poses major ecological threats. Biol Invasions Vol.: (0123456789) https://doi.org/10.1007/s10530-022-02892-6
Lovelock B., Y. Ji, A. Carr, and C-J. Blye. 2022. Should tourists care more about invasive species? International and domestic visitors’ perceptions of invasive plants and their control in New Zealand. Biological Invasions (2022) 24:3905–3918 https://doi.org/10.1007/s10530-022-02890-8
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
Burmese python in Everglades National Park; photo by Bob Reed, US FWS
Scientists continue to apply data collected in an international database (InvaCost; see “methods” section of Cuthbert et al.; full citation at end of this blog) to estimate the economic costs associated with invasive alien species (IAS). These sources reported $22.24 billion in economic costs of bioinvasion in protected areas over the 35-year period 1975 – 2020. Because the data has significant gaps, no doubt invasions really cost much more.
Moodley et al. 2022 (full citation at end of this blog) attempt to apply these data to analyze economic costs in protected areas. As they note, protected areas are a pillar of global biodiversity conservation. So it is important to understand the extent to which bioinvasion threatens this purpose.
Unfortunately, the data are still too scant to support any conclusions. Such distortions are acknowledged by Moodley et al. I will discuss the data gaps below a summary of the study’s findings.
The Details
Of the estimated $22.24 billion, only 4% were observed costs; 96% were “potential” costs (= extrapolated or predicted based on models). Both had generally increased in more recent years, especially “potential” costs after 1995. As is true in other analyses of InvaCost data, the great majority (73%) of observed costs covered management efforts rather than losses due to impacts. The 24% of total costs ascribed to losses, or damage, exceeded the authors’ expectation. They had thought that the minimal presence of human infrastructure inside protected areas would result in low records of “economic” damages.
The great majority (83%) of reported management costs were reactive, that is, undertaken after the invasion had occurred. In terrestrial environments, there were significantly higher bioinvasion costs inside protected areas than outside (although this varied by continent). However, when considering predicted or modelled costs, the importance was reversed: expected management costs represented only 5% while these “potential” damages were 94%.
Higher expenditures were reported in more developed countries – which have more resources to allocate and are better able to carry out research documenting both damage and effort.
More than 80% of management costs were shouldered by governmental services and/or official organizations (e.g. conservation agencies, forest services, or associations). The “agriculture” and “public and social welfare” sectors sustained 60% of observed “damage” and 89% of “mixed damage and management” costs respectively. The “environmental” and “public and social welfare” sectors together accounted for 94% of all the “potential” costs (predicted based on models) generated by invasive species in protected areas; 99% of damage costs. With the partial exception of the agricultural sector, the economic sectors that contribute the most to movement to invasive species are spared from carrying the resulting costs.
Lord Howe Island, Australia; threatened by myrtle rust; photo by Robert Whyte, via Flickr
Invasive plants dominated by numbers of published reports – 64% of reports of observed costs, 79% of reports of “potential”. However, both actual and “potential” costs allotted to plant invasions were much lower than for vertebrates and invertebrates. Mammals and insects dominated observed animal costs.
It is often asserted that protected areas are less vulnerable to bioinvasion because of the relative absence of human activity. Moodley et al. suggest the contrary: that protected areas might be more vulnerable to bioinvasion because they often host a larger proportion of native, endemic and threatened species less adapted to anthropogenic disturbances. Of course, no place on Earth is free of anthropogenic influences; this was true even before climate change became an overriding threat. Plenty of U.S. National parks and wilderness areas have suffered invasion by species that are causing significant change (see, for example, here, here, and here).
Despite Best Efforts, Data are Scant and Skewed
Economic data on invasive species in protected areas were available for only a tiny proportion of these sites — 55 out of 266,561 protected areas.
As Moodley et al. state, their study was hampered by several data gaps:
Taxonomic bias – plants are both more frequently studied and managed in protected areas, but their reported observed costs are substantially lower than those of either mammals or insects.
The data relate to economic rather than ecological effects. The costliest species economically might not cause the greatest ecological harm.
Geographical bias – studies are more plentiful in the Americas and Pacific Islands. However, studies from Europe, Africa and South America more often report observed costs. The South African attention to invasive species (see blogs here, here, and here), and economic importance of tourism to the Galápagos Islands exacerbate these data biases.
Methodological bias – although reporting bioinvasion costs has steadily increased, it is still erratic and dominated by “potential” costs = predictions, models or simulations.
I note that, in addition, individual examples of high-cost invasive species are not representative. The highest costs reported pertained to one agricultural pest (mango beetle) and one human health threat (mosquitoes).
Great Smokey Mountains National Park; threatened by mammals (pigs), forest pests, worms, invasive plants … Photo by Domenico Convertini via Flickr
As these weaknesses demonstrate, a significant need remains for increased attention to the economic aspects of bioinvasion – especially since political leaders pay so much greater attention to economics than to other metrics. However, the reported costs – $22.24 billion over 35 years, and growing! – are sufficient in the view of Moodley et al. to support advocating investment of more resources in invasive species management in protected areas, including – or especially – it is not quite clear — preventative measures.
SOURCES
Cuthbert, R.N., C Diagne, E.J. Hudgins, A. Turbelin, D.A. Ahmed, C. Albert, T.W. Bodey, E. Briski, F. Essl, P.J. Haubrock, R.E. Gozlan, N. Kirichenko, M. Kourantidou, A.M. Kramer, F. Courchamp. 2022. Bioinvasion cost reveals insufficient proactive management worldwide. Science of The Total Environment Volume 819, 1 May, 2022, 153404
Moodley, D., E. Angulo, R.N. Cuthbert, B. Leung, A. Turbelin, A. Novoa, M. Kourantidou, G. Heringer, P.J. Haubrock, D. Renault, M. Robuchon, J. Fantle-Lepczyk, F. Courchamp, C. Diagne. 2022. Surprisingly high economic costs of bioinvasions in protected areas. Biol Invasions. https://doi.org/10.1007/s10530-022-02732-7
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
APHIS is seeking stakeholder input to its new strategic plan to guide the agency’s work over the next 5 years.
The strategic plan framework is a summary of the draft plan; it provides highlights including the mission and vision statements, core values, strategic goals and objectives, and trends or signals of change we expect to influence the agency’s work in the future. APHIS is seeking input on the following questions:
Are your interests represented in the plan?
Are there opportunities for APHIS to partner with others to achieve the goals and objectives?
Are there other trends for which the agency should be preparing?
Are there additional items APHIS should consider for the plan?
range of American beech – should APHIS be doing more to protect it from 3 non-native pests?
Comments must be received by July 1, 2022, 11:59pm (EST).
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
monarch butterfly on swamp milkweed; photo by Jim Hudgins, USFWS
I have been impressed recently by two groups of scientists who are trying to broaden understanding of the impacts of invasive plants by examining the interactions of those plants with insects. As they note, herbivorous insects are key players in terrestrial food webs; they transfer energy captured by plants through photosynthesis to other trophic levels. This importance has been recognized since Elton first established the basic premises of food webs (1927) [Burghardt et al.; full citation at end of blog] Arthropods comprise significant members of nearly every trophic level and are especially important as pollinators. If introduced plants cause changes to herbivore communities, there will probably be effects on predators, parasites, and other wildlife through multitrophic interactions [Lalk et al.; Tallamy, Narango and Mitchell].
[I briefly summarize the findings of a third group of scientists at the end of this blog. The third group looks at the interaction between agriculture – that is, planting of non-native plants! – and climate change.]
One approach to studying this issue, taken by Douglas Tallamy of the University of Delaware and colleagues, is to look at the response of herbivorous insects to NIS woody plants fairly generally. They integrate their studies with growing concern about the global decline in insect populations and diversity. They note that scientists have focused on light pollution, development, industrial agriculture, and pesticides as causes of these declines. They decry the lack of attention to disruption of specialized evolutionary relationships between insect herbivores and their native host plants due to widespread domination by non-indigenous plants [Richard, Tallamy and Mitchell].
In their studies, Tallamy and colleagues consider not just invasive plants, but also non-native plants deliberately planted as crops or ornamentals, or in forestry. They point out that such introduced plants have completely transformed the composition of plant communities in both natural and human-dominated ecosystems around the globe. At least 25% of the world’s planted forests are composed of tree species not native to their locale. At least one-sixth of the globe is highly vulnerable to plant invasions, including biodiversity hotspots [Richard, Tallamy and Mitchell].
A different approach, taken by Lalk and colleagues, is more closely linked to concern about impacts of the plants themselves. They have chosen to pursue knowledge about relationships between individual species of invasive woody plants and the full range of arthropod feeding guilds – pollinators, herbivores, twig and stem borers, leaf litter and soil organisms. In so doing, they decry the general absence of data.
Both teams agree that:
Invasive plants are altering ecosystems across broad swaths of North America and the impacts are insufficiently understood.
The invasive plant problem will get worse because non-native species continue to be imported and planted. (Reminder: the Tallamy team considers impacts of deliberate planting as well as bioinvasion.)
Plant-insect interactions are the foundation of food webs, so changes to them will have repercussions throughout ecosystems.
Tallamy team
Non-native plants have replaced native plant communities to a greater or lesser extent in every North American biome – including anthropogenic landscapes [Burghardt]. The first trophic level in suburban and urban ecosystems throughout the U.S. is dominated by plant species that evolved in Southeast Asia, Europe, and South America [Tallamy and Shropshire]. Abundant non-native plants not only dominate plant biomass; they also reduce native plant taxonomic, functional and phylogenetic diversity and heterogeneity. Thus, they indirectly alter the abundance of native insects [Burghardt; Richard, Tallamy and Mitchell].
I think these articles might actually underestimate the extent of these impacts. While Richard, Tallamy and Mitchell report that more than 3,300 species of non-native plants are established in continental U.S., years ago Rod Randall said that more than 9,700 non-native plant species were naturalized in the U.S. (probably includes Hawai` i. The Tallamy team cites USDA Forest Service data showing 9% of forests in the southeast are invaded by just 33 common invasive plant species [Richard, Tallamy and Mitchell], I have cited that and other sources showing even greater extents of plant invasion in the east and here; other regions and here.
The Tallamy team has conducted several field experiments that demonstrate that the presence of non-native plants suppress numbers and diversity of native lepidopteran caterpillars. These non-native woody plants have not replaced the ecological functions of the native plants that used to support insect populations. This is true whether or not the non-native plants are deliberately planted or are invading various ecosystems on their own. [Richard, Tallamy and Mitchell]. (Of course, they expect plant invasions to grow; they note that some of the many ornamental species that are not yet invasive will become so.)
The result is disruption of the ecological services delivered by native plant communities, including the focus of their studies: plants’ most fundamental contribution to ecosystem function: generation of food for other organisms [Burghardt].
They note that plants’ relationship to insects differs depending on the insects’ feeding guilds — folivores, wood eaters, detritivores, pollinators, frugivores, and seed-eaters; and among herbivores with different mouthparts — chewing or sucking; and as host plant specialists or generalists. They decry studies that fail to recognize these differences [Tallamy, Narango, and Mitchell].
The Tallamy team explores why insect populations decline among non-native plants. That is,
1) Do insects directly requiring plant resources have lower fitness when using non-native plants; do they not recognize them as viable host plants; or do they avoid them altogether?
2) Are reductions in numbers of specialist herbivores mitigated by generalists? A decade of research shows that both specialists and generalists decline.
The team’s studies focus on lepidopteran larvae (caterpillars). Insect herbivores are both the largest taxon of primary consumers and extremely important in transferring energy captured by plants through photosynthesis to other trophic levels [Burghardt]. In addition, insects with chewing mouthparts are typically more susceptible to defensive secondary metabolites contained in leaves than are insects with sucking mouthparts that tap into poorly defended xylem or phloem fluids [Tallamy, Narango and Mitchell].
A study by Burghardt et al. found that 75% of all lepidopteran species and 93% of specialist species were found exclusively on native plant species. Non-native plants that were in the same genus as a native plant often supports a lepidopteran community that is a similar but depauperate subset of the community found on its native congener. In fact, the insect abundance and species richness supported by non-native congeners of native species was reduced by 68%.
A meta-analysis of 76 studies by other scientists found that, with few exceptions, caterpillars had higher survival and were larger when reared on native host plants. Plant communities invaded by non-native species had significantly fewer Lepidoptera and less species richness. In three of eight cases examined, non-native plants functioned as ecological traps, inducing females to lay eggs on plants that did not support successful larval development. Richard, Tallamy and Mitchell cite as an example the target of many conservation efforts, monarch butterflies (Danaus plexxipus), which fail to reproduce when they use nonnative swallowworts (Vincetoxicum species.) instead of related milkweeds (Asclepias species.).
Tallamy and Shropshire ranked 1,385 plant genera that occur in the mid-Atlantic region by their ability to support lepidopteran species richness. They found that introduced ornamentals are not the ecological equivalents of native ornamentals. This means that solar energy harnessed by introduced plants is largely unavailable to native specialist insect herbivores.
Tallamy, Narango, and Mitchell describe the following patterns:
1) Insects with chewing mouthparts are typically more susceptible to defensive secondary metabolites contained in leaves than are insects with sucking mouthparts that tap into poorly defended xylem or phloem fluids. As a result, sucking insects find novel non-indigenous plants to be acceptable hosts more often. However, there are more than 4.5 times as many chewing (mandibulate) insect herbivores than sucking (haustellate) species. It follows that the largest guild of insect herbivores is also the most vulnerable to non-native plants as well as being the most valuable to insectivores.
native azalea Rhododendron periclymenoides; photo by F.T. Campbell
2) Woody native species, on average, support more species of phytophagous insects than herbaceous species.
3) Although insects are more likely to accept non-native congeners or con-familial species as novel hosts, non-native congeners still reduced insect abundance and species richness by 68%.
4) Host plant specialists are less likely to develop on evolutionarily novel non-indigenous plants than are insects with a broader diet. There are far more specialist species than generalists, so generalists will not prevent serious declines in species richness and abundance when native plants are replaced by non-indigenous plants. In addition, non-native plants cause significant reductions in species richness and abundance even of generalists. In fact, generalists are often locally specialized on particular plant lineages and thus may function more like specialists than expected.
5) Any reduction in the abundance and diversity of insect herbivores will probably cause a concomitant reduction in the insect predators and parasitoids of those herbivores – although few studies have attempted to measure this impact beyond spiders, which are abundant generalists. The vast majority of parasitoids are highly specialized on particular host lineages.
6) Studies comparing native to non-native plants must avoid using native species that support very few phytophagous insects as their baseline, e.g., in the mid-Atlantic region tulip poplar trees (Liriodendron tulipifera) and Yellowwood (Cladrastus kentuckea).
7) Insects that feed on well-defended living tissues such as leaves, buds, and seeds are less likely to be able to include non-native plants in their diets than are insects that develop on undefended tissues like wood, fruits, and nectar. Although this hypothesis has never been formally tested, they note the ease with which introduced wood borers – emerald ash borer, Asian longhorned beetle, polyphagous and Kuroshio shot-hole borers, redbay ambrosia beetle, Sirex woodwasp (all described in profiles posted here — have become established in the US.
palamedes swallowtail Papilio palamedes; photo by Vincent P. Lucas; this butterfly depends on redbay, a tree decimated by laurel wilt disease vectored by the redbay ambrosia beetle
Lalk and Colleagues
As noted, Lalk and colleagues have a different frame; they focus on individual introduced plant species rather than starting from insects. They also limit their study to invasive plants. The authors say there is considerable knowledge about interactions between invasive herbaceous plants and arthropod communities, but less re: complex interactions between invasive woody plants and arthropod communities, including mutualists (e.g., pollinators), herbivores, twig- and stem-borers, leaf-litter and soil-dwelling arthropods, and other arthropod groups.
They ask why this knowledge gap persists when invasive shrubs and trees are so widespread and causing considerable ecological damage. They suggest the answer is that woody invaders rarely encroach on high-value agricultural systems and some are perceived as contributing ecosystem services, including supporting some pollinators and wildlife.
Lalk and colleagues seek to jump-start additional research by summarizing what is currently known about invasive woody plants’ interactions with insects. They found sufficient data about 11 species – although even these data are minimal. They note that all have been cultivated and sold in the U.S. for more than 100 years. All but one (mimosa) are listed as a noxious weed by at least one state; two states (Rhode Island and Georgia) don’t have a noxious weed list. None of the 11 is listed under the federal noxious weed statute.
Ailanthus altissima
Illustrations of how minimal the existing information is:
Tree-of-heaven (Ailanthus altissima) is noted to be supporting expanded populations of the Ailanthus webworm moth (Atteva aurea), which is native to Central America; and to be the principal reproductive host for SLF (Lycorma delicatua) https://www.dontmovefirewood.org/pest_pathogen/spotted-lanternfly-html/
Chinese tallow (Triadica sebifera) is thought to benefit both native generalist bee species and non-indigenous European honeybees (Apis mellifera).
Chinese privet (Ligustrum sinense) appears to suppress populations of butterflies, bees, and beetles.
Lalk and colleagues then review what is known about interactions between individual invasive plant species in various feeding guilds. They point out that existing data on these relationships are scarce and sometimes contradictory.
They believe this is because interactions vary depending on phylogenetic relationships, trophic guild, and behavior (e.g., specialized v. generalist pollinator). Arthropods can be “passengers” of a plant invasion. That is, they can be affected by that invasion, with follow-on effects to other arthropods in the community. Also, arthropods can be “drivers” of invasion, increasing the success of the invasive plants.
They then summarize the available information on various interactions. For example, they note that introduced plants can compete with native plants in attracting pollinators, causing cascading effects. Or they can increase pollination services to native plants by attracting additional pollinators.
They note that herbivore pressure on invasive plants can have important impacts on growth, spread, and placement within food webs. They note that these cases support the “enemy release hypothesis”, although they think there are probably additional driving mechanisms.
Lalk and colleagues note that most native twig- and stem-borers (Coleoptera: Buprestidae, Curculionidae, Cerambycidae; Hymenoptera: Siricidae) are not considered primary pests but that some of our most damaging insect species are wood borers (see above).
Some of these borers are decomposers; in that role, they are critical in nutrient cycling.
Arthropods in leaf litter and soil also serve important roles in the decomposition and cycling of nutrients, which affects soil biota, pH, soil nutrients, and soil moisture. They act as a trophic base in many ecosystems. Lalk and colleagues suggest these arthropod communities probably change with plant species due to differences in leaf phytochemistry. They cite one study that found litter community composition differed significantly between litter beneath tree-of-heaven, honeysuckle (Lonicera maackii), and buckthorn (Rhamnus cathartica) compared to litter underneath surrounding native trees.
Recommendations
Both the Tallamy and Lalk teams call for ending widespread planting of non-native plants. Lalk and colleagues discuss briefly the roles of
The nursery industry (including retailers); they produce what sells.
Scientists and educators have not sufficiently informed home and land owners about which species are invasive or about native alternatives.
Private citizens buy and plant what their neighbors have, what they consider aesthetically pleasing, or what is being promoted.
States have not prohibited sale of most invasive woody plants. Regulatory actions are not a straightforward matter; they require considerable time, supporting information, and compromise.
Tallamy team calls for restoration ecologists in the eastern U.S. to consider the number of Lepidopterans hosted by a plant species when deciding what to plant. For example, oaks (Quercus), willows (Salix), native cherries (Prunus)and birches (Betula) host orders of magnitude more lepidopteran species in the mid-Atlantic region than tulip poplar.(Those lepidopteran in turn support breeding birds and other insectivorous organisms.) [Tallamy & Shropshire]
Lalk and colleagues focused on identifying several key knowledge gaps:
How invasive woody plants affect biodiversity and ecosystem functioning
How they themselves function in different habitats.
Do non-native plants drive shifts in insect community composition, and if so, what is that shift, and how does it affect other trophic levels?
How do IAS woody plants affect pollinators?
The authors do not minimize the difficulty of separating such possible plant impacts from other factors, including climate change and urbanization.
Outhwaite et al. (full citation at end of this blog) note that past studies have shown that insect biodiversity changes are driven primarily by land-use change (which is another way of saying planting of non-native species – as Dr. Tallamy and colleagues describe it) and increasingly by climate change. They south to examine whether these drivers interact. They found that the combination of climate warming and intensive agriculture is associated with reductions of almost 50% in the abundance and 27% in the number of species within insect assemblages relative to levels in less-disturbed habitats with lower rates of historical climate warming. These patterns were particularly clear in the tropics (perhaps partially because of the longer history of intensive agriculture in temperate zones). They found that high availability of nearby natural habitat (that is, native plants) can mitigate these reductions — but only in low-intensity agricultural systems.
Outhwaite et al. reiterate the importance of insect species in ecosystem functioning, citing pollination, pest control, soil quality regulation & decomposition. To prevent loss of these important ecosystem services, they call for strong efforts to mitigate climate change and implementation of land-management strategies that increase the availability of natural habitats.
SOURCES
Burghardt, K. T., D. W. Tallamy, C. Philips, and K. J. Shropshire. 2010. Non-native plants reduce abundance, richness, and host specialization in lepidopteran communities. Ecosphere 1(5):art11. doi:10.1890/ES10-00032.
Lalk, S. J. Hartshorn, and D.R. Coyle. 2021. IAS Woody Plants and Their Effects on Arthropods in the US: Challenges and Opportunities. Annals of the Entomological Society of America, 114(2), 2021, 192–205 doi: 10.1093/aesa/saaa054
Richard, M. D.W. Tallamy and A.B. Mitchell. 2019. Introduced plants reduce species interactions. Biol Invasions
Tallamy, D.W., D.L. Narango and A.B. Mitchell. 2020. Ecological Entomology (2020), DOI: 10.1111/een.12973 Do Non-native plants contribute to insect declines?
Tallamy, D.W. and K.J. Shropshire. 2009. Ranking Lepidopteran Use of Native Versus Introduced Plants Conservation Biology, Volume 23, No. 4, 941–947 2009 Society for Conservation Biology DOI: 10.1111/j.1523-1739.2009.01202.x
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
I post here photos from two creek valleys in northern Virginia.
The Accotink creek valley is completely overrun by invasive plants … the herbaceous layer is made up of lesser celandine (Ficaria verna Huds; Ranunculus ficaria L.) and – in some places — Leucojum.
Neighboring Pohick creek valley still supports native hebaceous plants – skunk cabbage, spring beauties, trout lillies.
They both flow through wealthier suburbs in Fairfax County.
?????
P.S. In a ditch connecting to Pohick creek I have found this aquatic plant:
Plant is rooted, but leaves float on the water surface. In March the leaves were wide with scalloped edges; by April they are longer – lanceolate? I have seen it nowhere else. Anyone know what it is? Local authorities say it is not water chestnut (Trapa).
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
black berry eating hawthorn berries; photo by Paul D. Vitucci
Articles by Evan Fricke and colleagues remind us to look more broadly at bioinvasion to consider the impact on ecosystem function and evolution. They focus on animal interactions with plants in the shared environment, especially animals’ role as seed dispersers.
The authors also remind us that natural barriers explain why there are different species in different areas and thus how evolution and speciation follow different paths in different places. Think of Galapagos finches evolving in isolation from a few ancestors that somehow made it over the ocean from mainland South America.
These points are made in two recent articles.
In the first, Fricke and Svenning 2020 (full citation at end of this blog) note that about half of all plant species depend on animals to disperse their seeds. Animal seed dispersal is influenced by several drivers of global change, including local or generalized extinction (= defaunation); bioinvasion; and habitat fragmentation. The decline of large vertebrates has a particularly important role in these interactions.
Their study focused on fleshy-fruited plants that are dispersed by animals. (The study does not include nuts, e.g., acorns, which are presumably subject to some of the same pressures.) They expect evolution of the affected plants and animals to proceed differently as a result of the new partnerships, but they did not study any such interactions.
Their study covered animal seed-dispersal interactions with plants at 410 locations. The data encompassed 24,455 unique animal-plant pairs involving 1,631 animal and 3,208 plant species. Three quarters of the animals were birds; most of the rest were mammals, primarily bats and primates. Only 1% were in other animal groups – lizards, tortoises, or fish.
fruit bats on Luzon, Philippines; photo by Francesco Vernonesi; Flickr.com
They found that introduced plants and animals are twice as likely as native species to interact with introduced partners. The resulting interactions are likely to amplify biotic homogenization in future ecosystems. Already, introduced species have largely replaced missing native frugivore species in some places. In fact, mutualisms in which either or both the plant and animal is an introduced species are now about seven times higher than decades ago.
These mutual-benefit interactions of introduced species are even more prevalent in areas where human modification of the environment is greater. The proportion of introduced species and of novel interactions caused by introduced plant or animal species was higher for oceanic island systems than for continental bioregions. This finding adds a new dimension to the already recognized heightened susceptibility of remote islands to invasion and their loss of native species. Continental bioregions’ networks typically had few introduced animals and a greater prevalence of intro plants than animals.
Fricke and colleagues think plant-frugivore networks are likely to increasingly favor a relatively few introduced generalists over many native species, reducing the uniqueness of future biotas. The result might be to reduce resilience of terrestrial ecosystems by, first, allowing perturbations to propagate more quickly; and, second, by exposing disparate ecosystems to similar drivers. They called for giving higher priority to managing increasing ecological homogenization.
In the second article, Fricke, Ordonez, Rogers, and Svenning (2022) note that climate change requires many plant species to shift their populations hundreds of meters to tens of kilometers per year to track their climatic niche. Earth is also experiencing the formation of novel communities as species introductions and shifting ranges result in co-occurrence of species that do not share co-evolutionary history. They conclude that the novel mutualistic interaction networks will influence whether certain plant species persist and spread.
These authors examined four scenarios to assess how current long-distance dispersal has been affected by past defaunation and invasion and how it is threatened by species endangerment. These scenarios are as follows:
1st scenario (current scenario) = natural and introduced ranges of extant species today.
2nd scenario (natural scenario) = mammal and bird ranges as they would be if unaffected by extinctions, range contractions, or introductions.
3rd scenario (extinction scenario) = those bird and mammal species listed as vulnerable or endangered by the IUCN go extinct.
4th scenario (extirpation of introduced species scenario) = introduced species are extirpated.
Fricke and colleagues estimate that extinction of at least local populations of seed-dispersing mammals and birds has already reduced the capacity of plants to track climate change by 60% globally. The effect is strongest in temperate regions and regions with little topographic complexity. Two examples are eastern North America and Europe. These regions face a double threat: rapid climate change and loss of large mammals that provided long-distance dispersal.
The extinction scenario is most evident in Southeast Asia and Madagascar. The remaining animal seed dispersers are already threatened or endangered. Fricke and colleagues project that future loss of vulnerable and endangered species from their current ranges would result in a further reduction of 15% in the capacity of plants to track climate change.
The contrary situation is found on islands which have few native mammals. Introduced species are now important long-distance seed dispersers. In some cases, the introduced animals are dispersing invasive plant seeds, e.g., on Hawai`i feral hogs are spreading the invasive plant strawberry guava (Psidium cattleianum).
strawberry guava on Maui; photo by Forest and Kim Starr
People’s actions have resulted in ecoregions disproportionately losing the species that provide long-distance seed dispersal function, i.e., large mammals. In other words, human activities have caused not only rapid climate change—requiring broad-scale range shifts by plants—but also defaunation of the birds and mammals needed by plants to do so. Habitat fragmentation and other land-use changes will likely amplify existing constraints on plant range shifts.
Fricke and colleagues say their findings emphasize the importance of not only promoting habitat connectivity to maximize the functional potential of current seed dispersers but also restoring biotic connectivity through the recovery of large-bodied animals to increase the resilience of vegetation communities under climate change.
SOURCES
Fricke, E. C., & Svenning, J. C. (2020). Accelerating homogenization of the global plant–frugivore meta-network. Nature, 585(7823), 74-78. https://www.nature.com/articles/s41586-020-2640-y
Fricke, E. C., Ordonez, A., Rogers, H. S., & Svenning, J. C. (2022). The effects of defaunation on plants’ capacity to track climate change. Science, 375(6577), 210-214. https://www.science.org/doi/full/10.1126/science.abk3510
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
brown tree snake Boiga irregularis; via Wikimedia; one of the species on which the most money is spent on preventive efforts
In recent years a group of scientists have attempted to determine how much invasive species are costing worldwide. See Daigne et al. 2020 here.
Some of these scientists have now gone further in evaluating these data. Cuthbert et al. (2022) [full citation at end of blog] see management of steadily increasing numbers of invasive, alien species as a major societal challenge for the 21st Century. They undertook their study of invasive species-related costs and expenditures because rising numbers and impacts of bioinvasions are placing growing pressure on the management of ecological and economic systems and they expect this burden to continue to rise (citing Seebens et al., 2021; full citation at end of blog).
They relied on a database of economic costs (InvaCost; see “methods” section of Cuthbert et al.) It is the best there is but Cuthbert et al. note several gaps:
Only 83 countries reported management costs; of those, only 24 reported costs specifically associated with pre-invasion (prevention) efforts.
Data comparing regional costs do not incorporate consideration of varying purchasing power of the reporting countries’ currencies.
Data available are patchy so global management costs are probably substantially underestimated. For example, forest insects and pathogens account for less than 1% of the records in the InvaCost database, but constitute 25% of total annual costs ($43.4 billion) (Williams et al., in prep.) .
Still, their findings fit widespread expectations.
These data point to a total cost associated with invasive species – including both realized damage and management costs – of about $1.5 trillion since 1960. North America and Oceania spent by far the greatest amount of all global money countering bioinvasions. North America spent 54% of the total expenditure of $95.3 billion; Oceania spent 30%. The remaining regions each spent less than $5 billion.
Cuthbert et al. set out to compare management expenditures to losses/damage; to compare management expenditures pre-invasion (prevention) to post-invasion (control); and to determine potential savings if management had been more timely.
Economic Data Show Global Efforts Could Be – But Aren’t — Cost-Effective
The authors conclude that countries are making insufficient investments in invasive species management — particularly preventive management. This failure is demonstrated by the fact thatreported management expenditures ($95.3 billion) are only 8% of total damage costs from invasions ($1.13 trillion). While both cost or losses and management expenditures have risen over time, even in recent decades, losses were more than ten times larger than reported management expenditures. This discrepancy was true across all regions except the Antarctic-Subantarctic. The discrepancy was especially noteworthy in Asia, where damages were 77-times higher than management expenditures.
Furthermore, only a tiny fraction of overall management spending goes to prevention. Of the $95.3 billion in total spending on management, only $2.8 billion – less than 3% – has been spent on pre-invasion management. Again, this pattern is true for all geographic regions except the Antarctic-Subantarctic. The divergence is greatest in Africa, where post-introduction control is funded at more than 1400 times preventive efforts. It is also significant for Asia and South America.
Even in North America – where preventative actions were most generously funded – post-introduction management is funded at 16 times that of prevention.
Cuthbert et al. worry particularly about the low level of funding for prevention in the Global South. They note that these conservation managers operate under severe budgetary constraints. At least some of the bioinvasion-caused losses suffered by resources under their stewardship could have been avoided if the invaders’ introduction and establishment had been successfully prevented.
While in the body of the article Cuthbert et al. seem uncertain about why funding for preventive actions is so low, in their conclusions they offer a convincing (to me) explanation. They note that people are intrinsically inclined to react when impact becomes apparent. It is therefore difficult to motivate proactive investment when impacts are seemingly absent in the short-term, incurred by other sectors, or in different regions, and when other demands on limited funds may seem more pressing. Plus efficient proactive management will prevent any impact, paradoxically undermining evidence of the value of this action!
Aedes aegypti mosquito; one of the species on which the most money is spent for post-introduction control; photo by James Gathany; via Flickr
Delay Costs Money
The reports contained in the InvaCost database indicate that management is delayed an average of 11 years after damage was first been reported. Cuthbert et al. estimate that these delays have caused an additional cost of about $1.2 trillion worldwide. Each $1 of management was estimated to reduce damage by $53.5 in this study. This finding, they argue, supports the value of timely invasive species management.
They point out that the Supplementary Materials contain many examples of bioinvasions that entail large and sustained late-stage expenditures that would have been avoided had management interventions begun earlier.
Although Cuthbert et al. are not as clear as I would wish, they seem to recognize also that stakeholders’ varying perceptions of whether an introduced species is causing a detrimental “impact” might also complicate reporting – not just whether any management action is taken
Cuthbert et al. are encouraged by two recent trends: growing investments in preventative actions and research, and shrinking delays in initiating management. However, these hopeful trends are unequal among the geographic regions.
Which Taxonomic Groups Get the Most Money?
About 42% of management costs ($39.9 billion) were spent on diverse or unspecified taxonomic groups. Of the costs that were taxonomically defined, 58% ($32.1 billion) was spent on invertebrates [see above re: forest pests]; 27% ($14.8 billion) on plants; 12% ($6.7 billion) on vertebrates; and 3% ($1.8 billion) on “other” taxa, i.e. fungi, chromists, and pathogens. For all of these defined taxonomic groups, post-invasion management dominated over pre-invasion management.
When considering the invaded habitats, 69% of overall management spending was on terrestrial species ($66.1 billion); 7% on semi-aquatic species ($6.7 billion); 2% on aquatic species ($2.0 billion); the remainder was “diverse/unspecified”. For pre-invasion management (prevention programs), terrestrial species were still highest ($840.4 million). However, a relatively large share of investments was allocated to aquatic invaders ($624.2 million).
Considering costs attributed to individual species, the top 10 targetted for preventive efforts were four insects, three mammals, two reptiles, and one alga. Top expenditures for post-invasion investments went to eight insects [including Asian longhorned beetle], one mammal, and one bird.
Asian longhorned beetle
Just two of the costliest species were in both categories: insects red imported fire ant(Solenopsis invicta) and Mediterranean fruitfly (Ceratitis capitate). None of the species with the highest pre-invasion investment was among the top 10 costliest invaders in terms of damages. However, note the lack of data on fungi, chromists, and pathogens. (I wrote about this problem in an earlier blog.)
Discussion and Recommendations
Cuthbert et al. conclude that damage costs and post-invasion spending are probably growing substantially faster than pre-invasion investment. Therefore, they call for a stronger commitment to enhancing biosecurity and for more reliance on regional efforts rather than ones by individual countries. Their examples of opportunities come from Europe.
Drawing parallels to climate action, the authors also call for greater emphasis on during decision-making to act collectively and proactively to solve a growing global and inter-generational problem.
Cuthbert et al. focus many of their recommendations on improving reporting. One point I found particularly interesting: given the uneven and rapidly changing nature of invasive species data, they think it likely that future invasions could involve a new suite of geographic origins, pathways or vectors, taxonomic groups, and habitats. These could require different management approaches than those in use today.
As regards data and reporting, Cuthbert et al. recommend:
1) reducing bias in cost data by increasing funding for reporting of underreported taxa and regions;
2) addressing ambiguities in data by adopting a harmonized framework for reporting expenditures. For example, agriculture and public health officials refer to “pest species” without differentiating introduced from native species. (An earlier blog also discussed the challenge arising from these fields’ different purposes and cultures.)
3) urging colleagues to try harder to collect and integrate cost information, especially across sectors;
4) urging countries to report separately costs and expenditures associated with different categories, i.e., prevention separately from post-invasion management; damage separately from management efforts; and.
5) creating a formal repository for information about the efficacy of management expenditures.
While the InvaCost database is incomplete (a result of poor accounting by the countries, not lack of effort by the compilers!), analysis of these data points to some obvious ways to improve global efforts to contain bioinvasion. I hope countries will adjust their efforts based on these findings.
Seebens, H. S. Bacher, T.M. Blackburn, C. Capinha, W. Dawson, S. Dullinger, P. Genovesi, P.E. Hulme, M.van Kleunen, I. Kühn, J.M. Jeschke, B. Lenzner, A.M. Liebhold, Z. Pattison, J. Perg, P. Pyšek, M. Winter, F. Essl. 2021. Projecting the continental accumulation of alien species through to 2050. Glob Change Biol. 2021;27:970-982.
Williams, G.M., M.D. Ginzel, Z. Ma, D.C. Adams, F.T. Campbell, G.M. Lovett, M. Belén Pildain, K.F. Raffa, K.J.K. Gandhi, A. Santini, R.A. Sniezko, M.J. Wingfield, and P. Bonello 2022. The Global Forest Health Crisis: A Public Good Social Dilemma in Need of International Collective Action. submitted
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
Hakalau Forest, Hawai“i; nearly 90% of Hawaiian flora is unique to the Islands
A recent article by Yang et al. 2021 (full citation at the end of this blog) seeks to determine the extent to which introduced plants reduce the uniqueness of regional floras. They analyzed data from 658 regions covering about 65.7% of the Earth’s ice-free land surface and about 62.3% of the planet’s known plant species.
They found strong homogenization of plant species’ taxonomic and phylogenetic diversity results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.
There are two aspects of a region’s floral uniqueness. One is the number of species that it shares with other regions. This is taxonomic uniqueness. The other is the distinctiveness of the evolutionary history of the region. When several species are endemic to a region’s flora, and lack close relatives in other regions, that equals phylogenetic uniqueness.
The effect of a species introduction differs depending on which of these aspects one focuses on. Thus, naturalization of a species closely related to native species (e.g., a congeneric species) will have less impact on the phylogenetic floristic uniqueness of the region than naturalization by a distantly related species. Taxonomic uniqueness, however, will be affected to the same degree, irrespective of the phylogenetic distance between the naturalized and native species.
Yang et al. found strong homogenization of plant diversity. They found that species introductions increased the taxonomic similarity in 90.7% of all regional pairs and phylogenetic similarity in 77.2% of all region pairs. Most homogenization results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.
This loss of regional biotic uniqueness or distinctiveness changes biotic interactions and species assemblages. These, in turn, have ecological and evolutionary consequences at larger scales and higher levels.
The degree of homogenization between regions’ floras depends on three factors:
1) The distance between the donor and recipient regions. Since nearby regions share more species, an introduction from a more distant origin is more likely to be a novel species and so contribute to homogenization of “donor” and “receiving” floras.
2) Climatic similarity, especially temperature. A plant species introduced from a climatically similar but geographically distant place is more likely to establish than a species from a different climatic zone. As a result, the recipient area’s flora is changed to more closely resemble the flora of the donor region with which it shares climatic conditions – regardless of the distance between them.
3) The level of exchange of goods and people between two regions. The higher the rate of exchange between two regions, the greater the chance that a species will be introduced and become established. Yang et al. used the existence of current or past administrative relationships (e.g., colonial relationship) between two regions as a proxy for intensity of trade and transport between donor and recipient regions. They found that floras of regions with current or past administrative links have taxonomically become more similar to each other than the floras of regions with no such links.
flora of the Cape Floral Kingdom – South Africa; photo from Michael Wingfield
Establishment of introduced species can increase floristic similarity of the donor and recipient regions (= floristic homogenization) when the species is native to one of the two regions and naturalizes in the other, or when it is not native to both regions and naturalizes in both. On the other hand, a species introduction can decrease the floristic similarity of the two regions (i.e., enhance floristic differentiation) when the species is not native to both regions but naturalized in only one.
Homogenization hotspots differed slightly depending on whether one focused on taxonomic or phylogenetic aspects.
The regions with the greatest average increase in taxonomic similarity with other regions due to naturalized alien species were New Zealand, portions of Australia, and many oceanic islands. The Australasian situation probably reflects its long biogeographic isolation from other parts of the globe and its highly unique native flora. As a result, nearly all non-native plants introduced to Australasia strongly increase levels of its floristic similarity to the rest of the world. Oceanic islands have species-poor floras with large proportions of unique endemics. They have also received high numbers of naturalized alien plants.
Hotspots of phylogenetic homogenization on continents are the same as those for taxonomic homogenization, but this is not true for islands. Yang et al. think this is because islands’ native floras were established by natural colonization from nearby continental floras so – despite subsequent speciation – they retain their phylogenetic relationship to the donor areas’ floras.
Yang et al. concede that they lacked high-quality data on native and naturalized alien species lists for a third of Earth’s ice-free terrestrial surface, especially Africa, Eastern Europe, and tropical Asia. They believe, however, that data from these regions are unlikely to change the overall finding. (Scientists are beginning to compile lists of forest pests in Africa). link to blog
Yang et al. note that introduction and naturalization of alien species are likely to increase in the future, thusaccelerating floristic homogenization. The ecological, evolutionary and socioeconomic consequences are largely unknown.They call for stronger biosecurity regulations of trade and transport and other measures to protect native vegetation.
SOURCE
Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290. https://doi.org/10.1038/s41467-021-27603-y
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
It’s everywhere! multiflora rose (photo by Famartin)
The United States is overrun with introduced plants. Five years ago, Rod Randall’s database listed more than 9,700 non-native plant species as naturalized in the U.S. Not all of these species were “invasive”.
At that time, regional invasive plant groups listed the following numbers of invasive species in their regions:
Southeast Exotic Plant Pest Council – approximately 400 invasive plants
Mid-Atlantic Invasive Plants Council – 285 invasive plants
Midwest Invasive Plants Network reported that state agencies or state-level invasive plant councils in its region listed more than 270 plant species as invasive, noxious, or pest species
California Invasive Plants Council listed 208 species.
Texas Invasives reported more than 800 non-native plant species in the state, of which 20 were considered invasive.
Species – Rankings and Extents
We know that these invaders are affecting wide swaths of many ecosystems. A recent study based on Forest Inventory and Analysis (FIA) data (explained here) showed that nation-wide, 39% of forested plotssampled contained at least one invasive species. Hawai`i was first, at 70%. Eastern forests were second, at 46%. In the West overall, 11% of plots contained at least one invasive species. Plots in both Alaska and the Intermountain states were at 6% of plots invaded. A different study (Barrett and Robertson 2021; full citation at end of blog) reported the proportion of Western forest covered by invasive plants. This approach resulted in different numbers, but the same general ranking: Hawai`i again “led” at 46%; Pacific Coast states at 3.3%; Rocky Mountain states at 0.75%; coastal Alaska at 0.01%.
In more arid regions, data from the Bureau of Land Management showed that invaded acreage had more than doubled between 2009 and 2015.
buffelgrass removal in Tucson; Photo by Julia Rowe, Arizona Sonora Desert Museum
The situation is expected to get worse: a study of just one small portion of U.S. naturalized plants found that non-native plant species were more widely distributed than native species and that the average invasive plant inhabited only about 50% of its expected range. Furthermore, human actions were more important in facilitating spread than the species’ biological attributes.
Most of the detailed studies have been conducted in the Northeast – by both Forest Service and National Park Service scientists. The USFS’ Northern Region (Region 9) contains 24 states, from Maine to Minnesota, from Delaware to Missouri. A review of forest inventory (FIA) data (Oswaltet al. 2015) provided details on 50 plant species. (Unfortunately, the Southern Region [Region 8] has chosen to report in different formats, so it is hard to get an overall picture of invasive plants throughout the forests of the entire East. This is especially annoying to those of us who live in Mid-Atlantic states, which are divided between the two regions.)
Oswalt et al. (2015) provided data on the percentage of FIA plots in each state that were reported to have at least one invasive plant species. The northern Midwest ranked highest – e.g., one state (Ohio) at 93%; one state (Iowa) at 81%; two states (Indiana and Illinois) above 70%. Parts of the Mid-Atlantic region were almost as invaded – West Virginia at 79% and Maryland at 65%. The Northern plains states ranked lowest in invasions – North Dakota at 29% and South Dakota at 15%.
A study by the National Park Service of part of the Northeast (from Virginia and West Virginia to Maine) found a situation similar to that found by USFS researchers. In 35 of 39 park units, more than half of the plots had at least one invasive plant species when the 2015-2018 survey began. In 10 parks (a quarter of those surveyed), every plot had at least one. Invasions are worsening: 80% of the park units showed there was a significant increase in at least one trend measuring abundance.
Japanese stiltgrass in Shenandoah National Park; Photo by J. Hughes
The USFS and NPS report different species to be most widespread. In the National Park Service-managed units, Japanese stiltgrass (Microstegium vimineum) was found on 30% of all plots, in more than 75% of all NPS-managed units in the study. This magnitude comes despite the species not being found north of 41o N latitude. In forest plots inventoried by the USDA Forest Service, Japanese stiltgrass was the 14th most widespread species in the Northern region. I speculate that the species might not be common in the upper Midwest, which was not included in the NPS study. Oswalt et al. (2015) noted that Japanese stiltgrass was the 5th most common invasive plant in the Southern region.
Both studies agreed that garlic mustard (Alliaria petiolata) is widespread. The NPS study found it to be the most frequently detected non-grass herbaceous species, detected in 20% of plots throughout the study area (Virginia and West Virginia to Maine). On forest plots monitored by the USFS, garlic mustard was the 3rd most frequently detected species, on 4.5% of the surveyed plots. The species is reported to be present in 36 states & 5 provinces.
Why do Studies Ignore Deliberate Planting as a Factor?
Both USFS & NPS found shrubs and vines to be highly widespread. NPS specified Japanese barberry (Berberis thunbergii), Japanese honeysuckle (Lonicera japonica), multiflora rose (Rosa multiflora), and wineberry (Rubus phoenicolasius). USFS FIA data showed multiflora rose to be the most frequently recorded invasive plant, present on 16.6% of surveyed plots. It is otherwise recorded in 39 states and 5 provinces. Multiflora rose is almost ubiquitous in some states; in Ohio it is recorded on 85% of the plots. “Roses” were reported to be the 3rd most common invasive plant in the Southern Region. Other shrubs also dominated FIA plot detections: European buckthorn was 4th most frequently detected species, present on 4.4% of survey plots; or in 34 states and 8 provinces. Its presence is highest in New York, at 16.8% of plots. If the plots invaded by the various bush honeysuckle species do not overlap, these shrubs occupy 9.5% of all surveyed plots – second to multiflora rose. The vine Japanese honeysuckle ranked 6th – present on 3.6% of survey plots across the region. Japanese honeysuckle is reported to be the most common invasive plant in the Southern region. Other shrubs ranking 12th or above included Autumn olive and Japanese barberry
Tree-of-heaven (Ailanthus altissima) was the most common invasive tree found in National parks, again, despite not growing north of 41o N latitude. It is found in 9% of plots.
Ailanthus
I will say that I find it extremely annoying that the scientists carrying out these studies never mention that virtually all these shrub species had been deliberately planted in forests or nearby lands! Instead, they focus on such factors as histories of agriculture and other disturbances and fragmentation. It is well documented (e.g., Lehan et al. 2013) that the vast majority of shrub species introduced to the U.S. were introduced deliberately. Furthermore, more than 500 plant species invasive in some region are being sold on-line globally.
Deliberate planting of species that turn out to be invasive is also rarely recognized in the West, e.g., Pearson et. al. There, the motivation for planting might be livestock forage or erosion control rather than wildlife habitat “enhancement” or ornamental horticulture.
I am pleased that the most recent study (Barrett and Robertson 2021) differs somewhat by noting (sometimes) both invasions by forage grasses and the appearance in the mesic forests of Pacific states such planted species as Armenian blackberry. However, while this report notes the potential that pathogens might be transported to new areas by restoration planting and “assisted migration”, it does not mention the concomitant risk of introducing plant species that might prove invasive in the naïve ecosystems.
English ivy invading forest in Washington State; photo from Washington Noxious Weed Board
[Go to the earlier blogs linked here and the Western forests report for discussions of management strategies.]
Barrett and Robertson (2021) state that although invasive plants are increasing in extent and intensity in Western forests, they are usually considered to be contributing factors rather than as proximate causes. However, they note two caveats: 1) determining the ultimate causes and resulting implications of these recent increases is more difficult; and 2) data are particularly poor on plant species’ presence. Indeed, the FIA survey process link is ineffective for early detection and tactical monitoring [that is, identifying particular species in specific habitats of concern] of plant invasions.
Of the 23.4 M ha of forested lands that have experienced a disturbance over a five-year window (the time frame for FIA), only 600,000 ha was affected by the combined categories of geologic, vegetation, and other disturbances. (This is 10% of the area affected by either insects or pathogens.) Cheatgrass (Bromus tectorum) was by far the most abundant species in Western forests, covering 480,000 ha, or about 0.49%cover of all forested land in the conterminous Western United States. Because of the difficulties of surveying, Barrett and Robertson (2021) conclude that the area covered by IAS plants on the Pacific Coast and Rocky Mountains could be twice recorded values.
FIA surveys detected the highest number of non-native plant species in the forests of the continental Pacific states — 259 species. Many were grasses (although different species than in the Rockies), but shrubs and other forbs were also present. In the Rocky Mountain states the surveys detected a total of 195 non-native species, primarily grasses. FIA surveys in Hawai`i detected 136 non-native species. The most abundant was strawberry guava, which was detected on 9% of the forested area in the state. Surveys of FIA plots in coastal Alaska detected only 8 non-native plant species; common dandelion was the most abundant. Except in Hawai`i, the plants were expected to have substantially lower impacts than in eastern forests.
I note that the US Geological Service (Simpson and Eyler, 2018) reports there are approximately 1,754 non-native plants in Hawai`i and 424 in Alaska. Not all are necessarily invasive. And the USGS study covered all of Alaska, not just the southeastern coastal region.
Barrett and Robertson (2021) found that plant invasions are less extensive in older forest stands, mesic stands in contrast to drier areas and those with sparse or open tree canopies, and farther from roads. Thus, invasive plant cover was higher in hardwood and low-elevation and dry conifer forest types than in high-elevation and moist conifer types. In Hawai`i, mean plant cover was more than 40 % in all forest types except cloud forest, where it was 7.8 %. Again, proximity to roads was mentioned in the context of the likelihood of disturbance but no mention was made of the fact that households and businesses (e.g., tourist facilities, even agency facilities!) might deliberately introduce plants – e.g., horticulture.
Barrett and Robertson (2021) expect the impacts of NIS plants on forest lands to increase in the future, due to both additional introductions (despite efforts to prevent such) and spread of established species. They note that every disturbance creates an opportunity for the many ruderal and graminoid species to establish – facilitated by their abundance nearby. They note the significant challenge presented by secondary invaders, which often respond to space made available by “weed control” projects better than natives.
I welcome their concern about shade-tolerant plants apparently increasing in wetter areas of the Pacific coast states. They note that the presence of non-native plants in a forest is less obvious, and the impacts might be more subtle, perhaps primarily affecting tree regeneration through competition or other effects (e.g., promoting fire). Barrett and Robertson (2021) note that many of the shade-tolerant non-native species abundant in temperate Eastern U.S. forests (e.g., garlic mustard) are present in the West and are likely to become important.
SOURCES
Barrett, T.M. and G.C. Robertson, Editors. 2021. Disturbance and Sustainability in Forests of the Western US. USDA Forest Service Pacific Northwest Research Station. General Technical Report PNW-GTR-992
March 2021
Simpson, A., and Eyler, M.C., 2018, First comprehensive list of non-native species established in three major regions of the United States: U.S. Geological Survey Open-File Report 2018-1156, 15 p., https://doi.org/10.3133/ofr20181156.
ISSN 2331-1258 (online)
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed tree-killing pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm (These reports do not discuss invasive plants.)
In February the USFS published a lengthy analysis of invasive species: Invasive Species in Forests and Rangelands of the United States. A Comprehensive Science Synthesis for the US Forest Sector (Poland et al. 2021; full citation at the end of the blog). More than 100 people contributed to the book; I helped write the chapters on legislation and regulations and international cooperation. The book is available for download at no cost here.
Chapters address impacts in terrestrial and aquatic systems; impacts on ecosystem processes; impacts on various sectors of the economy and cultural resources; interactions with climate change and other disturbances; management strategies for species and landscapes; tools for inventory and management. Each chapter evaluates the current status of knowledge about the topic and suggests research needs. There are also summaries of the invasive species situation in eight regions.
Miconia – one of many invasive plants damaging ecosystems in Hawai`i
I greatly appreciate the effort. Authors first met in 2015, and most chapters were essentially written in 2016. The long delay in its appearance came largely from negotiations with the publisher. The delay means some of the information is out of date. I am particularly aware that several experts – e.g., Potter, Guo, and Fei – have published about forest pests since the Aukema source cited. I wonder whether inclusion of their findings might change some of the conclusions about the proportion of introduced pests that cause noticeable impacts.
Since the report’s publication in February I have struggled with how to describe and evaluate this book. What is its purpose? Who is its audience? The Executive Summary says the report is a sector-wide scientific assessment of the current state of invasive species science and research in the U.S.
However, the Introduction states a somewhat different purpose. It says the report documents invasive species impacts that affect ecosystem processes and a wide range of economic sectors. This would imply an intention to enhance efforts to counter such effects– not just to shape research but also to change management. Indeed, the Conclusion of the Executive Summary (pp. xvi-xvii) is titled “An Imperative for Action”.
Tom Vilsack, Secretary of Agriculture
I am not the author to evaluate how effectively the book sets out research agendas. Regarding its usefulness in prompting policy-makers to do more, I regretfully conclude that it falls short.
Getting the balance right between an issue’s status and what needs to be done is difficult, perhaps impossible. I appreciate that the report makes clear how complex bioinvasion and ecosystem management and restoration are. Its length and density highlight the difficulty of making progress. This daunting complexity might well discourage agency leadership from prioritizing invasive species management.
On the other hand, summary sections sometimes oversimplify or bury important subtleties and caveats. The question of whether some key questions can ever be resolved by science is hinted at – but in detailed sections that few will read. The same is true regarding the restrictions imposed by funding shortfalls.
The Report Would Have Benefitted from Another Round of Editing
Editing this tome was a Herculean task. I feel like a curmudgeon suggesting that the editors do more! Nevertheless, I think the report would have been improved by the effort. One more round of editing – perhaps involving a wider range of authors – could have pulled together the most vital points to make them more accessible to policymakers. It could also have tightened the ecosystem-based descriptions of impacts, which are currently overwhelmed by too much information.
A precis for policymakers
A precis focused on information pertinent to policymakers (which the current Executive Summary does not) should contain the statement that the continued absence of a comprehensive investigation of invasive species’ impacts hampers research, management, and policy (mentioned only in §16.5, on p. 332). It should note situations in which insufficient funding is blocking recommended action. I note three examples: programs aimed at breeding trees resistant to non-native pests (resource issues discussed only in §§8.3.1 and 8.3.2, p. 195); sustaining “rapid response” programs (§6.4.3, p. 125); costs of ecosystem restoration, especially for landscape-level restoration (§16.4). I am sure there are additional under-funded activities that should be included!
cross-bred ash seedlings being tested for vulnerability to EAB; photo courtesy of Jennifer Koch
Other important information that should be highlighted in such a precis includes the statement that many ecosystems have already reached a point where healthy functions are in a more tenuous balance due to invasive species (p. 51). Effective carbon storage and maintaining sustainable nutrient and water balance are at risk. Second, costs and losses caused by invasive forest pests generally fall disproportionately on a few economic sectors and households. They cannot be equated to governmental expenditures alone (p. 305). Third, even a brief estimate of overall numbers of invasive species appears only in §7.4. Information about individual species is scattered because it is used as example of particular topic (e.g., impacts on forest or grassland ecosystems, or on ecosystem services, or on cultural values).
Ecosystem Impacts Overwhelmed
As noted above, the report laments the absence of a comprehensive investigation of invasive species’ impacts. Perhaps the editors intended this report to partially fill this gap. To be fair, I have long wished for a “crown to root zone” description of invasive species’ impacts at a site or in a biome. Concise descriptions of individual invasive species and their impacts are not provided by this report, but they can be found elsewhere. (The regional summaries partially address the problem of too much information – but they do not provide perspective on organisms that have invaded more than one region, e.g., emerald ash borer or white pine blister rust.) Another round of editing might have resulted in a more focused presentation that would be more easily applied by policymakers.
Welcome Straightforward Discussion of Conceptual Difficulties
I applaud the report’s openness about some important overarching concepts that science cannot yet formulate. If supportable theories could be conceived, they would assist in the development of policies:
Despite decades of effort, scientists have not established a clear paradigm to explain an ecosystem’s susceptibility to invasion (p. 85). Invasibility is complex: it results from a dynamic interplay between ecosystem condition and ecological properties of the potential invader, especially local propagule pressure.
Scientists cannot predict how climate warming will change distributions of invasive species [see Chapter 4] and alter pathways. This inability hampers efforts to develop effective prevention, control, and restoration strategies (p. xi). Climate change and invasive species need to be studied together as interactive drivers of global environmental change with evolutionary consequences.
The Report’s Recommendations
Policy-oriented recommendations are scattered throughout the report. I note here some I find particularly important:
Measures of progress should be based on the degree to which people, cultures, and natural resources are protected from the harmful effects of invasive species.
Managers should assess the efficacy of all prevention, control, and management activities and their effect upon the environment. Such an evaluation should be based on a clear statement of the goals of the policy or action. [I wish the report explicitly recognized that both setting goals and measuring efficacy are difficult when contemplating action against a new invader that is new to science or when the impacts are poorly understood. Early detection / rapid response efforts are already undermined by an insistence on gathering information on possible impacts before acting; that delay can doom prospects for success.]
Risk assessment should both better incorporate uncertainty and evaluate the interactions among multiple taxa. Risk assessment tools should be used to evaluate and prioritize management efforts and strategies beyond prevention and early detection/rapid response.
Economic analyses aimed at exploring tradeoffs need better tools for measuring returns on invasive species management investments (§16.5).
Actions that might be understood as “restoration” aim at a range of goals along the gradient between being restored to a known historic state and being rehabilitated to a defined desired state. The report stresses building ecosystem resilience to create resistance to future invasions, but I am skeptical that this will work re: forest insects and disease pathogens.
Propagule pressure is a key determinant of invasion success. Devising methods to reduce propagule pressure is the most promising to approach to prevent future invasions (p. 115). This includes investing in quarantine capacity building in other countries can contribute significantly to preventing new invasions to the US.
Resource managers need additional studies of how invasive species spread through domestic trade, and how policies may differ between foreign and domestic sources of risk.
I appreciate the report’s attention to such often-ignored aspects as non-native earthworms and soil chemistry. I also praise the report’s emphasis on social aspects of bioinvasion and the essential role of engaging the public. However, I think the authors could have made greater use of surveys conducted by the Wisconsin Department of Natural Resources and The Nature Conservancy’s Don’t Move Firewood program.
Lost Opportunities
I am glad that the report makes reference to the “rule of 25” rather than “rule of 10s”. I would have appreciated a discussion of this topic, which is a current issue in bioinvasion theory. As noted at the beginning of this blog, the long time between when the report was written and when it was published might have hampered such a discussion
Also, I wish the report had explored how scientists and managers should deal with the “black swan” problem of infrequent introductions that have extremely high impacts. The report addresses this issue only through long discussions of data gaps, and ways to improve models of introduction and spread.
I wish the section on the Northwest Region included a discussion of why an area with so many characteristics favoring bioinvasion has so few damaging forest pests. Admittedly, those present are highly damaging: white pine blister rust, sudden oak death, Port-Orford cedar root disease, balsam woolly adelgid, and larch casebearer. The report also notes the constant threat that Asian and European gypsy moths will be introduced. (The Entomological Society of America has decided to coin a new common name for these insects; they currently to be called by the Latin binomial Lymatria dispar).
And I wish the section on the Southeast and Caribbean discussed introduced forest pests on the Caribbean islands. I suspect this reflects a dearth of research effort rather than the biological situation. I indulge my disagreement with the conclusion that introduced tree species have “enriched” the islands’ flora.
SOURCE
Poland, T.M., P. Patel-Weynand, D.M Finch, C.F. Miniat, D.C. Hayes, V.M Lopez, editors. 2021. Invasive species in Forests and Rangelands of the United States. A Comprehensive Science Synthesis for the US Forest Sector. Springer
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
