There seem to be three main concerns in studying ecological restoration: the first, one should have an idea of which state one intends to restore and why it is the best choice (i.e. the target should be defined and justified); the second concern is to exclude obviously invasive and/or recent introductions from the area, that might or might not have been at the root of the need to restore things (clear the area of what is certainly unwanted, whether mechanically or using a longer term perspective, eg. planting shade trees); the third is to allow to propagate or, if locally extinct, bring back from as nearby as possible, the original vegetation. The insects and other dependents will follow automatically.

Here in the Himalaya, a belt of 90 year old tea plantation (i.e. a hillside cleared of everything except for tea bushes and some fruit trees in 1860) was fenced off and permitted to re-grow in 1951. Today, it is barely distinguishable from any well protected broadleaf forest in the area, with tea bushes still growing under the deep shade of the undisturbed canopy. So for restoration, all that was required was protection, not plantation. Conditions will differ in different areas, but it is always better not to get bogged down in trivia, define and address the major challenges and proceed with the matter in hand. The tea bushes, although exotic (they were imported from China in 1860s) are doing well, having acclimatized over the last 150 years. The goal of enabling water to percolate into the ground to recharge groundwater has been achieved, which was one of the targets in 1951.

Good luck!

Interesting system and perspective. I work in North America (tallgrass prairie, serpentine grasslands, and now the longleaf pine savanna) and these particular systems are way beyond the 'remove the disruptive activity (agriculture)' and they are very few (if any) native systems within a reasonable distance for natural recolonization. We have to take a more proactive approach, but you are correct in that once the natural dynamics are allowed to return, key features of the ecosystem will recover nicely. I am a plant molecular ecologist, but I am specifically selecting species for my next experimental restoration field plots that have a high probability of really cool plant insect interactions (butterflies, moths, parasitiod wasps, etc...). Thanks, Danny

Good question and discussion! I have been working recently in a transformed area (drawdown zone of a major Chinese reservoir created in 2009) and the authorities are seeking re-greening of the muddy littoral zone. "Restoration" is impossible of course, because it was previously always above water, but is now seasonally flooded. Natural succession is happening quickly, and the adventitious species include some weedy exotics. So the challenge is to try to get ahead of the less desirable species and introduce relatively desirable native species (especially any that are rare or extirpated, of which we have some). In a context like this, it's a matter of setting priorities, and to me it seems obvious that it is better to put effort into taking action rather than wringing hands. There are other situations where the opposite is the case, and where just leaving it alone will produce equally good results. The Himalayan tea plantation case is a good one in that regard (I like it!).

While I think the word "restoration" does not apply in the situation in which I am working, there are those who are using the term nonetheless because it has become catchy. "Ecological engineering" is a suitable descriptor.

In summary, don't listen to purists, judge each situation on its merits, don't let language (e.g. "restoration") distort desirable outcomes. In your case, if there are no local seed sources, or if the seed source has too narrow a genetic base, bring some seeds from further away.

Where do you want to restore tall grass prairie? If that place was covered with ice 10,000 years ago, then everything that was there 1000 years ago, migrated there relatively recently and the system is still young and therefore plastic. If you are south of the ice-age ice front, then the situation could be different. I live north of the ice-front and the ecological transition is still very obvious. There are plenty of lichen-covered bare rocks that were scoured by receding glaciers 10,000 years ago, as well as many 'erratic' lichen-encrusted boulders in among the characteristic mixed forest, which I can see out of the window as I type.

I have written more than I planned ... hope it makes sense!

Hello Martin,

Interesting system and opportunity. There is a push in some circles to restore ecological services rather than a historical community, which would fit nicely with your project. Katherine Nash Suding (UC Berkely I think) had a paper in TREE or Ecology Letters articulating this approach. The point in time (10,000 years ago vs 1000 years) is valid, but clearly an academic just wanting to have an argument (it takes one to know one - haha). Typically in North America - pre-European settlement is often the reference, but we all acknowledge that the Native Americans did alter the system dynamics. I have been doing this ecotypic variation / molecular ecology research in grasslands for awhile and will be starting in the Longleaf Pine Savanna (herbaceous understory) - closer to the University and I want to give back to where I live / raising the family. Some of us population genetic minded people have been talking about numbers of sources. Plant out 1000 individuals and let sexual recombination and selection drive the dynamics. It is difficult to explain why having 1/2 of your plants die in the first few years us OK...Science vs. General Public perspective (managers too). Thanks for the note. Danny

Hello Danny --- thanks --- restoring ecological services is what we are trying to do. It's popular now in China and fits with the Chinese non-dogmatic mind-set.

Following your advice, I found the 2008 paper "Restoration through reassembly: plant traits and invasion resistance" in TREE by Katherine Suding and others. Nice paper and I may cite it. Is that the one you mean?

Martin

Yep,

Also look for papers by Tom Jones - he has been pushing this position for 10+ years. Tom is a grass breeder by training and works in Utah. Danny

Danny -- great --- found this one, thanks again!

http://www.bioone.org/doi/abs/10.1525/bio.2013.63.3.9

”Restoration professionals always ask 'can I use seeds collected from __km/miles away?'”

the closest, the best

because is local flora

OK, of course in case You do not take all the seeds from only one plant (individual)...

Zsolt,

I agree in cases when you have a similar donor community in close proximity, but what do you do when you do not have a sufficient seed supply / source close by. In the US, this could mean 600 miles ~ 1000 km distance between the seed production plots and the proposed restoration site. Another issue it the time constraints set by the agency or property owner. I like the idea of using a local (often small) remnant source, plant a small section of the proposed site, and then use these to help seed the rest of the site over a series of years. Reasonable?

Local sources of any particular species may be better adapted, but are not necessarily so. For this reason, it is wise to think about the objective of the restoration project. If the objective is to restore the pre-existing community, then the seed source should be as local as possible. If the objective is to restore ecological functions in a badly damaged ecosystem, then the local source is only better if it is (a) plentiful, and (b) effective. This is the message I got from the Tom Jones paper (see ref. above), and it makes complete sense to me.

In Nova Scotia (eastern Canada) we have an interesting case of a large mammal. The moose (Alces alces) had been extirpated from a large island (Cape Breton Island). The species was re-introduced in 1947, but rather than taking the local sub-species which still existed nearby, animals were brought from Western Canada. The population on Cape Breton Island is now vigorous and individuals now migrate from the island to the mainland. By contrast, the mainland population of the local sub-species is very unhealthy and listed as endangered. This sub-species appears to be less able to co-exist with white-tailed deer (due to a helminth which is lethal for the moose but not for the deer). The deer arrived by natural and facilitated migration in the 1890s. They quickly displaced caribou (Rangifer), which became locally extinct, and progressively pushed back moose. This was not due to aggressive interaction, just incompatibility. We don't know what the future holds, but this example shows that we should not assume that a local variety is better adapted when the broader ecological conditions have changed.

”what do you do when you do not have a sufficient seed supply / source close by. In the US, this could mean 600 miles ~ 1000 km distance between the seed production plots and the proposed restoration site”

well, that distance seems to be a little bit too big to be considered in case of ordinary restoration...

Practically, about which (concrete) plant-species are You talking about?

P.S. Because I do not really believe/convinced that people from USA are moving hundreds of species from one corner of the country into the opposite corner...

Martin,

Do you have any reference for the apparent inbreeding depression with the moose population? That is exactly the unintentional negative consequence we should all be avoiding. It may all come down to numbers (pop gen bias), but selecting those that have undergone some selection (presumably) would be a jump start. Now there is an issue of outbreeding depression. The problem with that is that studies that found no negative effects of crossing populations from X distances away (no outbreeding) do not seem to get published. Negative results in this case are hugely important for noting a low probability of outbreeding depression, which would help the restoration & conservation community greatly. I am guessing that problem is universal...

Dear Zsolt,

Thank you for your reply. In fact, this is one of those issues of scale. We do move seeds of native species up to 1000 miles in some cases and we are still within the species native range. It is a blessing of abundance in terms of scale, but we have done our very best to convert almost all of the native systems into agriculture system (universal problem). Matching ecological conditions / setting is desirable (similar selection), but that is often not possible.

According to Your post: moving of seeds in the area representing the range of the respective species is a current activity.

OK... so what is the problem?

P.S. I mean : related to ecological restoration...

Danny ... regarding Nova Scotia moose ... quite a lot has been written about this, but not (as far as I know) in terms of inbreeding depression. The problem is that the local moose subspecies was (as far as we know) in good condition, and abundant, until white-tailed deer arrived in the 1890s. So the decline was driven not by inbreeding depression but by environmental change.

Here's a helpful webpage:

http://www.novascotia.ca/natr/wildlife/large-mammals/mmoosefaq.asp

Here's a recent page about the brain worm

http://atlantic.ccwhc.ca/?p=1222

Here's an old paper identifying the disease agent:

http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1696956/pdf/canvetj00501-0013.pdf

Here's a 2003 status report:

http://novascotia.ca/NATR/wildlife/biodiversity/pdf/statusreports/StatusReportMooseNSComplete.pdf

There's a molecular genetic study of relevance by Broders (he's on RG):

http://www.smu.ca/faculty/hughbroders/documents/molecol99_000.pdf

There has long been discussion and doubt about the cause of "moose sickness"

http://www.nrcresearchpress.com/doi/abs/10.1139/z94-111#.UwDBEfldVAo

There is at least one substantial review (by T. Snaith) but I can't find it just now. I should have it somewhere in my old computer and will post it here if I can find it. Several papers have been published in the journal Alces but it is not open access. Karen Beazley, the lead author of these papers, is on RG and you will find some of the relevant papers on her page.

My description in the previous post is a personal opinion derived from following this story from the periphery of those involved in it. There are disparate opinions, though I think my opinion (as outlined in the earlier post) is close to mainstream. My conservation views are a bit different from some engaged in this issue in that I think the potential for out-breeding with Alberta moose is a good thing and could enhance the fitness of moose in this region, while some others worry about the purity of the regional subspecies.

I guess one could liken this to those who like pure breeds of dogs versus mutts. I'm in the mutt camp.

Danny ... found the Snaith paper. This is the pre-publication version, the text as published is the same but layout is different. It's a peer-reviewed work. This edition of the journal has not been digitized yet and so the paper is not available for download (see: http://nsis.chebucto.org/articles-available-for-download/)

Zsolt,

The practical problem is that there are almost never a seed source within a reasonable proximity (50-100 miles), if at all. Some people have started moving species from one side of their range to the other side (different abiotic drivers due to prec and temp patterns on a large scale). Concerns are 1) introduction of maladapted genotypes could have reduce the likelihood of successful establishment (wasted $$ and time), 2) distance source become established but decouple local gene complexes (outbreeding depression), 3) some native grasses have been developed as forage crops and erosion control - these can be aggressive and outcompete any non-dominant grass (polyculture of grasses rather than a species diverse grassland community, 4) plant genotype * soil microbial community interactions decoupled with non-native genotypes introduced, and 5) any seed production process has the potential of reducing genetic variation (unintentionally) / seeds from local remnants may have little genetic variation = both cases resulting in founder effects. If you are most interested in ecosystem functions, then you may be of the mind that plant X that can grow there would be just fine. Thanks, Danny

Martin,

Thanks for the references and information. I also think that mixing genes makes a lot of sense in many cases. Would one consider the panda breeding program in zoos to be functionally equivalent to a large naturally occurring population in nature. I (plant guy) talk about gardening (zoos) versus functional communities (nature preserves). Thanks, Danny

Danny, I agree. As a gardener I try to use as much diversity as possible (lots of different small fruits and vegetables) and to provide ecosystem services (berry bushes for birds), but the natural park next to my house is richer in diversity, biomass, and so on. Gardening involves compromises for human-centered purposes. Breeding pandas in zoos may eventually be useful in restoring depleted populations in the wild (it is being attempted in China), but this is not preferable to maintaining the wild populations in situ, and outside of western China (where the restoration program is being attempted), the pandas in zoos have little conservation relevance.

By the way ... I thought about the inbreeding depression issue. If we consider that the founding population of the moose in Cape Breton was necessarily small, it must be an "inbred" population, but it appears to be displaying greater fitness just now.

Martin,

I work in the highly modified sagebrush-steppe rangelands west of the Rockies. Because the ecosystem is in such poor shape and semi-arid conditions make it marginal to restore anyway, I emphasize performance and genetic diversity in the plant materials I develop. But, as you can imagine, this is very controversial (actually anathema to many conservation botanists). I find my work dovetails quite nicely with the work of Richard Hobbs and his collaborators on novel ecosystems.

Here are three more publications that may be of interest.

Jones, T. A. Plant materials for novel ecosystems. Chapter 26 (p. 212-227) In R.J. Hobbs, E. S. Higgs, C. M. Hall (eds.) Novel ecosystems: intervening in the new ecological world order. Wiley-Blackwell Press. 2013.

Jones, T. A. and T. A. Monaco. A role for assisted evolution in designing native plant materials for domesticated landscapes. Front. Ecol. Environ. 7:541-547. 2009.

Jones, T. A. When local isn’t best. Evolutionary Applications 6:1109-1118. 2013.

Thanks,

Tom J.

Thanks Tom. This has been a useful discussion for me, and has helped me to appreciate the values of RG.

The author of the questions seems to be about moving native PLANT genetic material, and also, in general, what are other concerns when doing ecological restoration. Here are my list of ecological restoration concerns and comments about native plant movement:

1.) Not using the proper local genetic plant material is the number one cause of long term failure when trying to restore an ecosystem. If you use the correct species, and replant it somewhere within its natural range, the US Forest Service discovered probably a century ago, that the long term survival of those trees was not assured, so Tree and Shrub Seed Zone were drawn up.

Tree seeds are harvested from a particular zone to be replanted back in to that same zone. In Oregon and California seed zone maps are being created for wildflower and native grass planting.

In California, we have widespread native grass and wildflower species, that are able to live in various habitats because they are genetically fixed ecotypes, and a lot of work was done 50 years ago on ecotypes at Stanford University, and you can see a bibliography of plant ecotype studies from 1880-1996 at http://www.ecoseeds.com/juicy.gossip.three.html.

Ultimately, once Ecological Restoration becomes a common practice in a particular country, ecotype maps and seed zone maps would need to be drawn, so that practitioners know how far they can move genetic material successfully.

2.) Quick methods to deal with weeds, like 100% control within 18 months or less.

3.) Quick methods to establish the local ecosystem, like 100% local native cover within 18 months or less.

4.) Methods to determine the local native species needed to replant a self-sustaining ecosystem, how many species in what percentages, like a cake mix recipe.

5.) No patented or licensed technologies being offered to conduct the ecological restoration process is a big concern. Without patented and licensed technologies, that means that the business of Ecological Restoration is still an academic experimental exercise, and has not fully made it way as a business with solid performance standards for its work ofered.

In the southeastern US local sources of native seed are very limited, and the majority of seed suppliers don't supply information on the origin of the seed source. On the bright side, there are many more suppliers than there were 10 years ago. Many plant species are sensitive to the length of day, and so if moved more than a few degrees in latitude will flower out of sync with local individuals. Here is my list, most of which have already been mentioned above:

1) Better information from seed suppliers on the collection location, including approximate coordinates, ecosystem classification, and soil type. This would help practitioners to select the most appropriate material available.

2) Better ways to control non-native species, particularly forage grasses.

3) More information on interactions between plant species and soil microbial populations.

4) Criteria with which to select plant species for restoration. We will never get there by using trial and error to select plant species. There are so many possibilities, but it takes years to develop a commercial seed supply. Failures are rarely publicized. Where to start? What are some of the characteristics of species that a) establish reliably on restoration sites and b) promote the development of a diverse native ecosystem?

Hello Jennifer,

I could not agree more. Having done conservation genetics / restoration genetics since the late 1990's, my more frustrating issues relate to getting information of 'where did you get your seed for your restoration'. I do understand that professions making a living doing restoration want to keep their information close to the chest (so they can make money), but it would be so much better if I had the whole story when looking at the genetic marker results. One thing pushed back in the 1990's was the idea of a regional approach to seed sources. There is fairly logical reasons for that approach and most of my fellow scientists see the value of increasing the numbers of sexually reproducing individuals so that we can have maximum genetic variation in which to plant into historical sites (restoration). That way selection can act of a range of genotypes rather than a few (limited sources / limited through nursery production). But there will always be those who point to the potential for outbreeding depression as a reason to not do anything (fear of disrupting coadapted gene complexes w/o any data indicating there are coadapted gene complexes). The rub on that very real and important concern is that studies that show a lack of outbreeding depression are not likely to be published (negative results are extremely important, but negative results tend to not be published). Some of the outbreeding depression species papers that have been published (100 to 1000 miles scales) actually have other studies with these same species showing an optimal crossing distance in native grasslands between 10 and 100 meters (Cassia fasiculata). This requires us to think of population genetics within the context of gene flow and selection at the population / community / and landscape levels. Then we need to convey that complex information to policy people and well meaning land managers (private/government) in a way we can start moving forward with recovering some of these natural areas in the modern landscape matrix. Thank you for your response (first from North America). Danny

Tom Jones,

Thanks for the information and contributing to the discussion. We do agree on most of the issues, but where we disagree likely reflect scale and ecological system (things are so much bigger out west). Thanks again. Danny

Martin,

I agree on the founder effects and lack of genetic variation without a decrease in fitness. It also follows with the Northern Elephant Seals too. Danny

Hello Craig,

Thanks for the response and there is a rich history of ecotypic variation research done with the Stanford and the amazing serpentine grasslands of California too. Seed transfer zones work great with economically important plants (trees and shrub especially), but the matrix species of many systems are not characterized in the same way. My research has shown spatial genetic patterns reflecting gene flow patterns (historical and current) and differential plant performance - ecotypic variation. I have also looked at the genetics in restored grasslands and found that the diversity and genetic signature follows the number of sources you used and where you got those seeds. One study I did looked at a grassland established for about 25 years with non-local and local dominant grass sources (same species). Even after 25 years, you could walk out in the field and see the difference between the two adjacent fields. Genetic markers of the material plants were constantly local and non-local, while the seedlings from these material plants were a hybrid mix. There were differences between the two sources (phenology, insect damage - amazingly consistent) - but they both survived for 25 years and apparently the offspring did not recruit from seed into these grasslands after establishment. Whether the lack of recruitment was do to maladapted hybrids (outbreeding depression) or that seedling recruitment is rare in established tallgrass prairies is unclear. That study was started because the Goose Lake Prairie manager heard someone was starting to do prairie grass genetic research and he had noticed that there were differences between his NE and IL bigbluestem (2001 Prairie Conference paper).

I agree that the lack of some type of standards is a real issue for restoration ecological research. When a restoration project fails because of the restoration professionals lack of understanding of the system, the public only sees a failure in restoration. It would be great to remove that false division between academics and restoration professionals, but I do understand where the friction comes into play. I am on the academic research side, but some of my dear friends are actually the ones in the field trying to make a difference. I can give them X reasons why you should approach a particular project in a specific way and they counter with 'where do I get the material' and 'who will be restoring the natural disturbance regime'. If we establish a regional perspective of native seed sources for some systems, then we would have material for establishing / restoring communities. Potential problems with this could be 1) who is responsible for maintaining these sources (seed production plots or regional plant communities established specifically to provide seed for future planting within the region, 2) would there be sufficient genetic variation to account for potential edaphic or small scale ecotypic variation and could we seed / plant enough seeds to let natural selection decide the winners and losers, and 3) how would this influence the current native seed producers and restoration professionals. I think the selection of species for planting likely reflect historical records (what was there), but what if the habitat has been so altered that the historical 'ecotypes' are not the best adapted (see Tom Jone's post and papers above).

Clearly there is no one right answer for every situation, but hopefully we can start progressing towards a more thoughtful and larger scale (spatial & temporal) approach to reestablishing and restoring native communities. Thank you again for your thoughtful comments. Danny

I wanted to comment on Jennifer's item No. 4)

Criteria with which to select plant species for restoration. We will never get there by using trial and error to select plant species. There are so many possibilities, but it takes years to develop a commercial seed supply. Failures are rarely publicized. Where to start? What are some of the characteristics of species that a) establish reliably on restoration sites and b) promote the development of a diverse native ecosystem?

Telling the story from the California perspective, where the ecosystem understory is 99.99% gone, replaced by exotics, you have to be an ecosystem archaeologist of sorts. Fortunately some artists accurately painted the understory, like "California spring" 1875 by Albert Bierstadt, or there are old post cards like those from the wildflower festival in Arvin near Bakersfield from 1910-1940 that you can get on Ebay.

Otherwise you have to start from scratch like I am in the hills of Palo Alto right now, at http://www.ecoseeds.com/arastradero.html.

1.) First step is to make a list of potential local species, using relic stands as models or herbarium specimens or data from local floras.

Potentially we had about 100-150 in our native California grasslands in any one spot, but I usually start with 20-25 that I know can fight the weeds, and be able to harvest quantities of seeds when they mature. For example I would choose a fast growing native grass over a violet, but perhaps plug in the violet at a later date.

2.) Second step is to find relic stand and harvest seeds locally. I like to stay within a watershed, and from plus or minus 1,000 ft. elevation, from a similar soil type, and within 20 inches of rainfall. Within a 50 miles radius of my project, we have normally from 8 to 80 inches annually, supporting from redwood forests to deserts with horned toads.

3.) Then grow the teaspoons of local native seeds into seedlings, and at the same time, start getting the planting area 100% weed-free, using native grass mulch for the annuals. We have over 1,000 species of weeds in California, covering about 99.99% of the State so that is where most project fail, in not inventing a method to get the area 100% weed-free.

4.) Planting the seedlings is the fun and creative part. This is where you have to figure out how close and what species should be planted next to each other, at what densities, like a cake mix. If you want to get fancy, you can write a computer program where you can input into fields the percentage cover of each species, and it will crunch the numbers and spit out the cake mix recipe.

5.) Harvest stock seeds from your first planting. If you harvest one pound of stock seeds, and give it to a commercial seed grower who has the specialized equipment to reproduce native seeds, it is usually a one to one hundred turn around in one season.

6.) Once you get bulk local native seeds, then the next step is to do small scale test plots, about one square meter each, to develop you seeding rates, and to experiment with mixing species, find out what fertilizers you need to add, how much mulch of what kind, etc.

7.) Once you have your bulk native seeds, you know what the cake mix is, how much seed, how much fertilizer, mulch, etc., then start sowing!

8.) Expect to budget a ton of money for these steps. I am estimating one million dollars for the first acre in the arid Western USA or Southwest and $70,000 for the next acre up to 70 acres. After that, when you have enough seed bulked up, the costs for seeds and materials may drop to a more reasonable $50,000 per acre.

You can see how valuable an intact non-riparian, grassland ecosystem or forest understory is, when you try and rebuild one from scratch.

Craig,

I appreciate your real world view, which is often missing in our academic fields. We are taking what I call the observant gardener approach to trying to propagate an endangered hemiparasite in our greenhouse. We not only have been able to grow plants from seed for about 6 months (without host plants) under greenhouse conditions, but I think we may be able to get these darn things up to flowering (knock on wood). That will be a game changer for this endangered species, but I will have to start looking for the correct venue for the paper. Any suggestions? Danny

I have published several times in the past in Ecological Restoration and there is a more academic journal Restoration Ecology, and also a newer journal Native Plants. Probably easiest to get published in Native Plants, more difficult in Ecological Restoration and probably very difficult in Restoration Ecology.

My test plot in the Palo Alto, California park has a dozen blooms today, and by mid-April hope for about 2,000 like last April. It is right next to the parking lot, and anyone can visit during the day sunrise to sunset. The rest of the 70 acre hill around the planting is 100% exotic weed cover introduced from Europe.

I have actually had an amazing impact with a small paper in Native Plants - lead to a push for the State of Minnesota to develop seed transfer zones...did not become law. Test plot next to the public access is a good idea. The more you show people what could be possible, the more they will help make it so. Thanks to the Twilight movies, may be I can sell the establishment of an endangered vampire plant rather than endangered hemiparasitic Schwalbea americana. marketing gold or another dumb idea...time will tell (haha). Thanks for the suggestions. Danny