Pawpaw Ecological Survey in Michigan
Part 2: Restoring Reciprocity

Most recent UPDATE: 2023 by Connie Barlow
[email protected]

ASIMEN is the traditional Potawatomi term Henceforth, "PAWPAW" and "ASIMEN" will be used interchangeably in this project. Consult the Asimina triloba entry in Wikipedia. Linked Table of Topics (on this webpage) PART 2: RESTORING RECIPROCITY • Appreciating Asimen: Geography and Taxonomic Relatives • Appreciating Asimen: Fully resists deer browsing • Appreciating Asimen: Prevents Invasion by Japanese Stiltgrass • Appreciating Asimen: Ethnobotany • Toward Renewing and Expanding Reciprocity • Could pawpaw help compensate for the loss of Black Ash as a fiber source? • Could pawpaw restore the subcanopy where deer are overpopulated? • Appreciating Asimen: Original Instructions • New York State: Where Pawpaw is a "Threatened Species" • What About American Persimmon? Companion Pawpaw Webpages PART 1: Who Are the Pollinators? • Conclusions & Recommendations for Attracting Pollinators • Video Documentation of Pawpaw Pollinator Fieldwork • Collecting and Distributing Seeds • Other Anatomical Features of Pawpaw • Basic Elements of the Pollinator Study • Sites (Fruited and Fruitless) for Field Observations and Comparisons • STUDY AIMS with Three Alternative Hypotheses re POLLINATION • Pawpaw as an ancient flower type (CRUCIAL READING FOR SERIOUS OBSERVERS) • Pawpaw Flower Stages • Food for Pollinators • Pawpaw Pollination: Knowns and Unknowns • Probable Pollinators of Spring 2021 - Nitulidid Beetles • APPENDIX: PHOTOS of Spring 2021 surveillance at sites (insects, pawpaw stages, ecology) • South African Cycad Paper Distinguishes Casual Visitors from Effective Pollinators • Ecological Interpretations of 2021 Field Experience • BACKGROUND PAPERS for Ascertaining Ecological Interpretations • RECOMMENDATIONS for ORCHARD HABITAT MANAGEMENT • FURTHER QUESTIONS and SUGGESTIONS FOR EXPERIMENTS • Guidance for the Future Pollinator Watches • Additional POLLINATOR Information from Technical Papers PART 3: WILD-PLANTING EXPERIMENTS • BASELINE DOCUMENTATION of a Wild Patch (Saline River, Michigan, 2020 & 2021) • Learning about Germination (2023) • Helping Forests Walk: Assisted Migration of Pawpaw • Helping Forests Walk 2025 wild-planting success along HURON RIVER, Michigan • Planting Thornapple River Slope (2022) Part 2 RESTORING RECIPROCITY Appreciating Asimen: Geography and Taxonomic Relatives    PHOTO LEFT: Annona muricata is a tropical relative of American Pawpaw. Asimina triloba is the northern-most species of a huge tropical fruit family: Annonaceae. This family entails 108 genera and some 2,400 known species, including the fruits known as cherimoya, custard apple, soursop (photo left), and annona. About 900 species are Neotropical, 450 are Afrotropical, and the remaining are Indomalayan. Learning more about asimen's ability to thrive at its poleward extreme (and perhaps beyond) is thus an important aspect of this project. Annonaceae is the largest taxonomic family within one of the most ancient orders of flowering plants: Magnoliales. Genus Asimina bears a classic magnolia type flower. Opening downward keeps this long-lasting flower viable despite episodes of rain, during which it becomes an attractive rain shelter for insects. As with its presumed early ancestors, magnolia type flowers specialize in attracting beetles as pollinators. This deep-time, planetary understanding (and wonderment!) of Asimen's place within its vast and ancient family are gifts from the sciences of paleoecology and evolutionary biology. The concept of elder, drawn from Indigenous thinking and experience, amplifies regard for Family Annonaceae and its member species. From a now-classic TEK paper, "Traditional Ecological Knowledge" (published in 2000 in the journal Ecological Applications), Raymond Pierotti and Daniel Wildcat wrote, "In nearly all native stories animal- and plant-persons existed before human-persons. Thus, these kin exist as our elders and, much as do human elders, function as our teachers and as respected members of our community. Acknowledging nonhumans as teachers and elders requires that we pay careful attention to their lives, and recognize that these lives have meaning on their own terms."    Another tree within the Magnoliales order that bears a similar and very large flower is also at its northern-most range in southern Michigan. Liriodendron tulipifera is not yet widespread here, but does have the capacity to become the tallest tree of all. LEFT: Native range of Liriodendron tulipifera. Its common name, Tulip Tree, is appropriately descriptive. (See photo below.) However, another common name for this tall, fast-growing canopy species is Yellow Poplar. Perhaps this tree was called "poplar" by the settler population from England because its large leaves jostle in the wind like members of genus Populus: cottonwoods and aspens. Because this name suggests faulty affinities, it is important to call this species Tulip Tree — or its lovely genus name, Liriodendron. Photo left by Robert Miller; right by Connie Barlow This genus has only a single species in North America, and one in Asia. A European species was wiped out during glacial times, as were many other plant species that were geographically blocked from moving southward by Europe's rugged E-W mountain ranges and the Mediterranean Sea. Hence, observing wild Tulip Tree is an opportunity to reflect with gratitude on the north-south orientation of our gentle Appalachian mountains — and the major river systems that quickly delivered seeds southward as the glaciers advanced.   MAPS SOURCE: TorreyaGuardians.org Tulip Tree has thus far not been harmed by any of the insects translocated from another continent (unlike other canopy dominant trees, such as Ash, Beech, and Elm). Moreover, although American Chestnut was the canopy giant in the 19th century in much of eastern North America, Tulip Tree took the lead when chestnut blight made its way through the forests. As well, given continuing climate disruption, there will be opportunity for Tulip Tree to be welcomed into more forests of southern Michigan and increasingly northward beyond its current range. THE FRUIT OF ASIMEN IS PEST-FREE.    Because the leaves and twigs are not browsed by any native vertebrate, deer are no threat to seedlings and saplings. See "Intensive Selective Deer Browsing Favors Success of Asimina triloba (Paw Paw), a Native Tree Species", by Mitchell Slater and Roger Anderson, 2014, in Natural Areas Journal. Hence asimen is an ideal native fruit for organic farming and the localizing movement in southern Michigan. Although pawpaw is the exclusive host to the caterpillar stage of our Zebra Swallowtail butterfly, finding this native in a pawpaw patch is a rare delight. Eurytides marcellus is the only member in the temperate zone of the otherwise large tropical group of swallowtail butterflies known as the "kite swallowtails" (tribe Leptocircini). PHOTO LEFT: Zebra Swallowtail laying an egg on a pawpaw stem. • RETURN TO TOP Appreciating Asimen: Fully resists deer browsing Land use in southern Michigan offers ideal habitat for uncontrollable population growth by deer: Our patchwork of small woodlots and forested river edges amidst farm fields, large rural properties with lawns and gardens, and abundant roads provide edge habitats favored by deer. Dwellings and people are interspersed such that opportunities even for bow-hunting (much less rifles) are difficult to arrange. As well, once the sound of rifles begins, deer can be expected to retreat to hunting-free safe zones that they have experienced in the past. Overall, so long as traditional carnivores are missing and hunting is not a year-round activity, winter starvation, disease, and collisions with cars are what controls the deer population. Meanwhile, the native palatable plants are all but lost and the deer-resistant non-natives take over. Most worrisome for foresters is that canopy trees will not be able to replace themselves wherever deer are overpopulated: their seedlings and saplings are killed by perpetual browsing.    LEFT: Invasive Amur honeysuckle, Lonicera maackii from Asia is now a dominant subcanopy woody tree/shrub in many recovering forested zones in southern Michigan. This photo, taken in early spring, along the Huron River near Ypsilanti, shows a dense patch of the mature honeysuckle. The shade is now too dense for the fallen canopy tree in the background to be replaced by native canopy trees. According to scholarly papers, there are only three native subcanopy trees in Michigan that can, to a large extent, grow and maintain despite overpopulated deer: PAWPAW, striped maple, and American hornbeam. RESOURCES: • News article, 2023, "Few good options for shrinking Michigan's problem deer herds". EXCERPT: "... Beyond risk of crashes and disease, deer can destroy forest ecosystems, suburban hosta gardens and farmers' crops. One study found that in Presque Isle County alone, deer consumed a tenth of farmers' soybeans. In overbrowsed forests, saplings disappear and mature trees have a manicured look from deer nibbling branches as high as their necks can reach. That eliminates habitat for ground-nesting songbirds and other animals..." • "Impacts of White-Tailed Deer Overabundance in Forest Ecosystems: An Overview", 2008, by Thomas J. Rawinski, USFS. EXCERPTS: "Deer have successfully exploited the human-altered environment, feeding in agricultural fields, orchards, roadsides, lawns, and gardens... The forest floor is presently dominated by unpalatable plant species, such as black huckleberry, Japanese barberry, Pennsylvania sedge, and various woodland grasses. As trees mature and die, or topple over during storms, gaps in the canopy become larger and more numerous. There are no young trees to fill the gaps.... Some plants resist deer herbivory, owing to chemical or morphological defenses or low digestible content. Among tree species, black cherry would be classified as fairly resistant. Horsley and others (2003) demonstrated a trend toward black cherry dominance in forests impacted by deer. At Rock Creek Park in Washington, D.C., Rossell and others (2007) found all major woody species to be impacted by deer, with the exception of American beech and spicebush. At very high population densities deer will ultimately browse black cherry, American beech, spicebush, mountain laurel, and many other less palatable species....Native herb populations are often decimated by deer, but some species are consistently avoided. Examples [of native species avoided by deer] include white snakeroot, black bugbane, mayapple, blue cohosh, Pennsylvania sedge, and eastern hayscented fern. A forest understory dominated by deer-resistant species is usually diagnostic of a forest afflicted by overly abundant deer. Foresters are especially concerned that eastern hayscented fern has become too abundant in some areas, preventing or impeding tree seedling establishment (Horsley and others 2003)." • "Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks", 2023, by Kathryn M. Miller et al., Ecological Applications. EXCERPT: "... Many of these browse-tolerant species, like paw paw (Asimina triloba), American holly (Ilex opaca), and American hornbeam (Carpinus caroliniana), are short-stature trees that are unable to grow to current canopy height..." • "Compounding human stressors cause major regeneration debt in over half of eastern US forests", 2019, by Kathryn M. Miller and Brian J. McGill, Journal of Applied Ecology EXCERPT: "... For example, several low canopy, browse-resistant species (e.g. striped maple and pawpaw) have shown increases in areas with high deer densities, which can further suppress regeneration of canopy species and greatly alter forest structure (Kain et al., 2011; Rossell et al., 2005)." • Anecdotal report by Laurie Potteiger, West Virginia Native Plant Society: "Does anyone have experience removing/controlling pawpaw trees? We won't get to planting trees or shrubs until this fall or next year and in the meantime they have/are taking over. What should I do in the meantime? More background: I live in an area (Jefferson County, near Harpers Ferry at about 700' elevation) with an extremely high deer population. The deer eat virtually every other native tree and shrub. Even spicebush. There are virtually no saplings of trees other than pawpaw, although the overstory contains tulip poplars, maples, oaks, hickories, etc. The seedlings for those other native species don't get more than an inch or two high before the deer eat them. We have an acre of mostly wooded land (on the top of the ridge and downslope facing NW). Pawpaw are already dominant in some areas, and popping up where they are not yet dominant, but grow way faster than anything else. We have been planting some native forbs, and have planted a few shrubs and trees, but have to cage or fence everything I plant to protect them for the deer. (When I find a tiny tree seedling other than pawpaw I try to cage it before the deer get to it.) The goal is to eventually put an 8' deer fence around a significant portion of the property rather than having all the individual-plant cages and the multiple deer exclosures, which still leave most of the area unprotected. Also: most of my outdoor time in spring the last two or three years has been pulling garlic mustard (along with several other invasive non-native plants). But making significant progress. I've already started cutting down some of the young [pawpaw] trees (leaving a stump so I know where they were) but I've read that only stimulates more growth. And I've read painting stumps with glyphosate in fall doesn't really help kill the tree the way it does with many other trees and shrubs. What to do?" • RETURN TO TOP Appreciating Asimen: Prevents Invasion by Japanese Stiltgrass In 2017, Michigan DNR reported first detection of invasive Japanese Stiltgrass in this state. Resembling a small bamboo, Microstegium vimineum was spotted in Washtenaw County. Wikipedia reports that it arrived first in Tennessee, as packing material for porcelain from China, in 1919.    Japanese stiltgrass is similar to and often grows along with the North American grass Leersia virginica. A silver stripe running down the center of its leaf (PHOTO LEFT) is diagnostic of this non-native grass. Also, Leersia flowers one to two months earlier than the stiltgrass, which flowers in late summer. Connie Barlow reports seeing dense patches of a similar-looking grass while walking the pawpaw patch at Draper-Houston. But she quickly learned by observation that if such a grassy patch is present, there will be no pawpaw stems there. Fortunately, a 2004 report confirmed that in Tennessee, "Dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy.... Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer." Ref: "Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests", by Patrice Cole and Jake Weltzin, 2005, in Biological Invasions. Appreciating Asimen: Ethnobotany   Photos by Connie Barlow ABOVE: Connie Barlow purchased this box of native pawpaw and persimmon at the U-pick orchard of Marc Boone, near Ann Arbor MI, early October 2020. ASIMEN is not only the largest fruit in America, it also excels in taste and nutrition, as in this characterization by Sheri Beth Crabtree, in her master's thesis at the University of Kentucky, 2004, titled "Sexual and Asexual Reproductive Characteristics of the North American Pawpaw": The fruit is very sweet, with a strong aroma when ripe and soft custard-like flesh. The flavor of the pawpaw fruit is unique, having been described as resembling banana, mango, pineapple, or melon. The fruit are also highly nutritious, containing double the vitamin C of apples, peaches, or grapes, amounts of vitamin A similar to apples and grapes, higher levels of many minerals, including potassium, phosphorus, magnesium, calcium, and iron, than apples, peaches, or grapes; and high levels of several essential amino acids (Peterson et al., 1982). There are many references to INDIGENOUS and EARLY SETTLER uses and appreciation of pawpaw fruit. For example: "Some Native American tribes cultivated the pawpaw for fruit and are responsible for its widespread range today. The Cherokee and many other tribes used the pawpaw fruit for food. The fruit, which is the largest edible fruit native to America, is high in amino acids. The Iroquois used the mashed fruit to make small cakes that were dried and stored. The dried cakes were soaked in water and cooked to make a sauce or relish that was served with corn bread. Raw and cooked fruits were dried by the sun or on a fire. These were stored for use in the future or taken on hunts. The Cherokee used the inner bark to make cordage. By twisting the bark, they made string and strong ropes." Excerpted from "USDA Plant Guide: Pawpaw". "The pawpaw has a rich history as an important food source to Native Americans, early European explorers, and early settlers of North America (Pickering, 1879). Native Americans likely aided in the distribution of pawpaw across North America, with evidence of the Iroquois tribe bringing pawpaws into central New York, where they were not found previously (Keener and Kuhns, 1997). The pawpaw was first formally noted in 1541, by the De Soto exploration group in the Mississippi River valley (Sargent, 1890). Lewis and Clark wrote in their journal that when rations were low, their exploring party subsisted mostly on pawpaws, and enjoyed the taste (Pomper and Layne, 2005). Early settlers depended partially on pawpaw to sustain them in times of crop failure (Peterson, 1991)." Excerpted from "Sexual and Asexual Reproductive Characteristics of the North American Pawpaw [Asimina triloba]", master's thesis by Sheri Beth Crabtree, 2004, University of Kentucky. "There is evidence that humans played a role in pawpaw dispersal as well. 'Natives in the eastern half of the country have always used pawpaws,' said Dr Devon Mihesuah, a citizen of the Choctaw Nation who holds the Cora Lee Beers Price professorship in International Cultural Understanding at the University of Kansas. 'Iroquois reportedly mashed pawpaws and made the flesh into cakes and then dried them in the sun. They were used as a travel food or mixed with water into cornbread.'" Excerpted from "The revival of a forgotten American fruit", by Jonathan Shipley, 2022, BBC Travel. Connie Barlow queried a friend, Lamar Marshall, who documented the traditional trail system of the Cherokee, using archived texts and maps followed by field investigations of each. His work was sponsored and is owned by the Eastern Band tribe of Cherokee in western North Carolina. LAMAR MARSHALL replied to Connie 2 April 2021: "... Regarding pawpaws, we don't have near as many in the Southeast as the central and northeast states. They occurred down through Alabama but they rarely fruit there and here in the Blue Ridge. I was told by old timers down in Alabama that pawpaw was used for firemaking and cordage. We used it extensively for hand drills and fire boards. One of the best and softest for getting coals fast. As far as primary sources, very few in my possession and I haven't checked the archaeological research from those recovering seeds and food remains in fire pits. U.T and Chapel Hill folks have done a lot. I attached two references you likely have. One from the encyclopedic Native American Botany by Moerman and the other Plants of the Cherokee by William Banks. The latter is from a thesis done about 1950 with the author actually gathering first-hand data from Cherokees. It includes the Cherokee name. I have attached excerpts from both books. If I find anything further, I will get it to you." John Ambrose, a retired botanist in southern Ontario Province, wrote in an email (4 August 2021) to Barlow: "I remember thinking that the pawpaws in s. Ontario must have arrived by canoe; all are delicious and the ones in the US are sometimes said to be insipid, so some human selection likely has taken place, plus almost all Ontario populations occur in river floodplains." An excellent academic paper that aggregates previous studies in listing pawpaw among some 20 native fruit and nut trees planted by the Indigenous prior to European colonization is: "Native Americans as active and passive promoters of mast and fruit trees in the eastern USA", by Marc D. Abrams and Gregory J. Nowacki, 2008, The Holocene: Fleshy tree fruits were used for complex carbohydrates, vitamins and minerals and at least 20 fruit- and berry-producing trees native to the eastern USA were commonly consumed by Native Americans (Table 1). In the southeast USA, persimmon (Diospyros virginiana), elderberry (Sambucus), hackberry and sugarberry (Celtis), hawthorne (Crataegus) and pawpaw (Asimina triloba) were most accessible (Fogelson, 2004: 62). They eliminated the astringent taste of persimmon by leaving the fruit on the tree until after the first frost (Briand, 2005). Fruits and berries could be eaten in season or dried and used throughout the year (Lieberman, 1984). Cherokees girdled trees and used fire to clear areas to stimulate production of woody shrub and vines important to their diet, such as raspberry, elderberry, blackberry, blueberry, huckleberry and grapes. They ate wild cherries (Prunus), pawpaw, mulberry (Morus), serviceberry (Amelanchier) and persimmon. Creek Indians ate fruits of plum (Prunus), mulberry, persimmon and honey locust pods (Fogelson, 2004: 375-76). Seminole Indians in northern Florida were reported to have large groves of wild sour orange (Citrus aurantium; introduced by the Spanish) near their villages, and their diet also included native fruits of wild plum, wild cherry, pawpaw and the berries of many shrubs (Fogelson, 2004: 342, 431, 456).     ... The northern migration of pawpaw to Ohio and New York was thought to have been facilitated by Iroquois transport and planting (Keener and Kuhns, 1997), but it was later argued that the distribution of the species could be explained by mammal dispersal (Murphy, 2001). Tim MacWelch, published this survivalist advice 27 August 2021 in Outdoor Life magazine: 9 Reasons the Pawpaw Is the Ultimate Tree for Survivalists. EXCERPT: "7. Make a Friction Fire: Pawpaw gave me one of the best results I have ever had in front of a class with a field-built bow drill set. While taking a class to look for friction fire equipment along a river's edge, a small pawpaw grove just kept giving and giving. I found some great branches for a drill and board that were both dead and dry, but not rotten. I was able to find another branch that was flexible enough for a bow. I stripped off some very rotten fibrous bark for tinder, and less rotten bark for cordage on the bow. The only part that wasn't pawpaw was my handhold block, which was a local piece soapstone that I drilled out with a small chunk of harder stone. The whole kit came together in roughly 30 minutes, but the most astounding part of the demo was the part where it worked on the first try. Field-built kits often need tweaking and part substitutions before they start working, but that sweet little kit worked right away (which, as any instructor will tell you, usually doesn't happen when people are watching). Pawpaw can give you great wood for friction fire components like drills and boards. As we've already discussed, the rotten bark can also be found in stages of decomposition that allow it to be both tinder and cordage material. You could even use branches as bows and chunks of wood for handhold blocks." • RETURN TO TOP    Pawpaw: In Search of America's Forgotten Fruit This 296-page book by Andrew Moore was published in 2015 by Chelsea Green. It surveys past and current uses and appreciation of asimen by indigenous cultures and early settler cultures. Chapters 13-19 survey past and present human relationships with asimen region by region. "Pawpaw is ancient and belongs to an earlier culture." (p. 86) Pages 9-10, 155: brief survey of indigenous ways of preparing and preserving the FRUIT of asimen, including drying the pulp for later use, and its nutritional values. Pages 11, 172, 223: Indigenous uses of fibrous INNER BARK for rope, nets, and footwear were widespread in the eastern USA. Page 83 (and other pages): Asimen is ideally suited for wild gathering in that its foliage is not browsed by deer and its fruit hosts no insect larvae. The beginning chapters of Moore's book reveal that the pawpaw cultivars now sold by nurseries and planted in commercial orchards were selected by the dominant culture barely a century ago from wild patches. Very little, if any, subsequent breeding ensued. (Nurseries propagate cultivars not from seed but by grafting cut stems onto rootstock; this ensures stable fruit qualities.) Because these cultivars trend from northern states (like Indiana, Illinois, Ohio, and Michigan), evidence is strong that native Americans not only greatly assisted pawpaw in migrating as far north as climate warming enabled, but also selected seeds from their preferred patches. The first peoples thus engaged in horticultural selection of pawpaw for human consumption, just as they did for traditional crops of squashes, beans, and maize. In 2021, a photo-rich essay summarizes the horticultural history of pawpaw from the standpoint of colonial, settler culture: "The Mad Scientist of Pawpaws", by Bill Heavey, February 2021, Garden and Gun Magazine. Toward Renewing and Expanding Reciprocity PRACTICAL OUTCOMES: Might this field study yield results that help recover reciprocal interactions for assisting asimen in the wild, along with renewed gathering opportunities for humans? Q: If the wild patch is missing pollinators, will we find ways to encourage pollinators to return? Q: If the wild patch lacks genetic diversity and thus cannot self-fertilize, will we offer the patch seeds from elsewhere? Q: If the patch is too shaded to bring fertilized flowers into fruit, will we seek out where and how to offer canopy breaks? Q: But if this ancient wild patch regards seeds as a necessity only following disturbance, how will we hear that and how will we respond? And just who is WE? And by what process will discernment be undertaken? If and when these questions arise from the results of our field studies, surely this is where participants immersed in western science and natural history worldviews will choose to hear from (and defer to) participants and advisors grounded in Traditional Ecological Knowledge (TEK). Botanist Robin Wall Kimmerer offers encouragement in "Restoration and Reciprocity: The Contributions of Traditional Ecological Knowledge," chapter 18 in Human Dimensions of Ecological Restoration: Integrating Science, Nature, and Culture, 2011, edited by Dave Egan et al. "The idea of reciprocity with land is fundamental to many indigenous belief systems. Indeed, such beliefs serve as the foundation for what have been described as cultures of gratitude. In such cultures, people have a responsibility not only to be grateful for the gifts provided by Mother Earth, they are also responsible for playing a positive and active role in the well-being of the land. They are called not to be passive consumers, but to sustain the land that sustains them. Responsibilities to the more-than-human world are simultaneously material and spiritual, and, in fact, the two are inseparable. Ecological restoration can be viewed as an act of reciprocity, where humans exercise their care-giving responsibility for ecosystems." • REVITALIZING HUMAN RECIPROCITY WITH THE FRUITS OF ASIMEN. There is little debate that humans in North America have been crucial dispersal agents for helping Asimen track the warming climate northward as the glaciers retreated. Whether or not seeds were intentionally selected, carried, and planted, it is indisputable that, "As the ice retreated from its last southward advance, which peaked about twenty thousand years ago, four of the five above-mentioned anachronistic trees of eastern and central North America would have been helped to reclaim former territory by newly arriving humans. Pawpaw and persimmon fruits would have been carried back to camp, their seeds removed or spit out at the time of eating." — Connie Barlow, 2001, "Anachronistic Fruits and the Ghosts Who Haunt Them", Arnoldia • WIKIPEDIA "Asimina triloba": "The natural distribution of the common pawpaw in North America, prior to the ice ages and lasting until roughly 10,000 years ago, was done by certain megafauna until they became extinct during the Quaternary extinction event. After the arrival of humans and the subsequent extinction of megafauna that were distributing A. triloba, the probable distribution of these large fruit-bearing plants has been by humans." • "Ecological niche modelling and phylogeography reveal range shifts of pawpaw, a North American understorey tree", by Graham E Wyatt et al., 12 January 2021, in Journal of Biogeography. ABSTRACT: Predictions of species' responses to accelerating global climate change require an understanding of historical range shifts. However, large-scale phylogeographical studies of Eastern North American understorey plant taxa are relatively scarce. Here we employ ecological niche modelling and genetic analyses for inference of optimal pawpaw habitat in the past and future.... Models suggest that 22,000 YBP A. triloba was restricted to two major refugia in narrow bands of mostly now-submerged habitat and possibly several small inland refugia. Molecular data are consistent suggesting that the eastern refugia expanded to give rise to the eastern cluster which is characterized by higher genetic diversity. The Gulf of Mexico refugium likely gave rise to populations in the western cluster, which is characterized by lower genetic diversity.   22,000 years ago                          Year 2070 projected range ABOVE: Green signifies the geographic range of the most suitable habitats past and future. LEFT: Refuges for pawpaw during last glacial maximum, 22,000 YBP (low sea level at that time enabled retreat to coastal areas now underwater). RIGHT: Projected ideal range in 2070 (using IPCC high emissions scenario). "Given the low fruit-set observed in populations at the northern edge of the current range (Wyatt, unpublished data) and limited vagility of contemporary seed dispersal vectors (gravity and small mammals), A. triloba is unlikely to expand its range rapidly and track environmental changes in lockstep.... Although our models indicate that optimal habitat will expand significantly, between the possible loss of southern populations and environmental change that outpaces the rate of dispersal and colonization at the northern edge, the realized distribution of A. triloba may diminish by 2070." • RETURN TO TOP Could pawpaw help compensate for the loss of Black Ash as a fiber source? BLACK ASH PAWPAW    Note: The WIKIPEDIA page for Black Ash states: "Black ash is unique among all trees in North America in that it does not have fibers connecting the growth rings to each other. This is a useful property for basket makers. By pounding on the wood with a mallet, the weaker spring wood layer is crushed, allowing the tougher and darker summer wood layer to be peeled off in long strips." Could the fiber qualities of asimen inner bark help compensate for the immense cultural loss of fiber wrought by the Emerald Ash Borer's impact on Black Ash trees (Fraxinus nigra)? If so, would reciprocity entail helping asimen to move northward into culturally significant woodlands in which Black Ash has been lost? One woman in northern MICHIGAN seems to have initiated that experiment. The Little Traverse Bay Bands of Odawa Indians are already planting tree species onto their lands whose northern-most range is several hundred miles southward. One of those trees is pawpaw: • "As northern Michigan warms, scientists bring tree seedlings from the south", by Kelly House, 3 May 2021, in Bridge Michigan. "RENEE DILLARD is a member of the Little Traverse Bay Bands and a renowned Anishinaabe natural fiber artist known for her intricate black ash baskets. She had to stop harvesting ash in northern Michigan's swamps after the emerald ash borer decimated the population. She switched to other fiber, including cedar, dogbane, basswood and bullrush. But climate change threatens to push some of those species northward, too, and Dillard believes they'll eventually become scarce near the Little Traverse Bay Bands' land in the Northwestern Lower Peninsula.      So she is experimenting with substitutes from further south. She keeps a stash of pawpaw seeds, and occasionally buries a few near her home in Harbor Springs, hoping they'll grow into the deciduous tree nicknamed 'the Indiana banana' for its custardy, tropical-tasting fruit. But it's the fibrous bark Dillard is after...." "...NOAH JANSEN's tree migration project is aimed at facilitating this kind of cultural resilience. He has specifically chosen plants that could preserve tribal members ability to hunt, gather and carry on cultural traditions: There's sassafras, a species with medicinal properties. And American plum, which can be harvested by humans or grazed by deer.      As foliage on the forest canopy matures, tribal species managers hope to continue their efforts by filling in the understory with plants that can survive in the future climate. Similar experiments are taking place or planned in communities throughout the state." • RETURN TO TOP Could pawpaw restore the subcanopy where deer are overpopulated?   ABOVE: Connie Barlow took these photos November 2021 while planting pawpaw seeds onto the forested slopes along the Huron River where it flows through collapsed industrial landscapes of Ypsilanti, Michigan. All green leaves in these photos are of a non-native honeysuckle: Lonicera maackii. This exotic Amur honeysuckle is the dominant subcanopy woody plant in these photos. Notice the distinct longitudinal-ridged bark of the foreground stem in the left photo and the opposite leaves with red berries in the right. Uniquely, here in Michigan this invasive deciduous plant retains bright green leaves that simply fall off in December without browning or yellowing. HYPOTHESIS: Might the foreign honeysuckle express so invasively because overpopulated deer eat virtually all of the seedlings of our native trees that attempt to grow? Yet, because pawpaw is poisonous to deer, might our native subcanopy tree be able to effectively compete with (possibly even wrest control from) the now-dominant subcanopy invasive? SCIENTIFIC REPORTS OF PAWPAW THRIVING WHERE DEER ARE OVERPOPULATED: • "In recent decades, naturalists have noted the expansion of pawpaw from well-drained, lowland habitats into drier, upland forests. This phenomenon appears to be driven, at least in part, by patterns of deer browse. Deer find pawpaw foliage unpalatable and, therefore, avoid browsing pawpaw seedlings and saplings. Instead, they preferentially browse species such as spicebush (Lindera benzoin), oaks (Quercus spp.), red maple (Acer rubrum), and blackgum (Nyssa sylvatica).... This deer behavior benefits pawpaw in two ways. First, small pawpaws don’t need to allocate energy to recovering from browse, and instead can put that energy towards growth and reproduction. Second, frequent deer browse on sapling and shrub species preferred by deer suppresses the growth of these species, clearing the way for pawpaw. As a result, we might expect to see pawpaw becoming more common in forest understories that are heavily impacted by deer browse (which describes most NCR forests).... Another potential contributor to the success of pawpaw is the suppression of fires that were an important part of the disturbance regime in many eastern forests before European settlement. Pawpaw are not strongly fire-adapted (unlike other common canopy dominates, such as oaks), and they likely benefit from the lack of fire in contemporary forests." National Park Service webpage: "Pawpaw: Small Tree, Big Impact". • "Indiana state parks historically did not allow hunting. But by the 1990s, white-tailed deer populations in parks had swelled to such size that many species of native wildflowers such as trillium and lilies largely disappeared, replaced by wild ginger and exotic species such as garlic mustard and Japanese stiltgrass, plants not favored by deer. Oak and ash tree seedlings gave way to highly deer-resistant or unpalatable trees such as pawpaw." 2014, "Hunting gives deer-damaged forests in state parks a shot at recovery". • "Although white-tailed deer are generalist herbivores, they can have significant effects on species composition and abundance of forest trees, especially when deer densities are high and most plant species are heavily browsed but a few are selectively avoided as browse. We evaluated effects of selective deer browsing on tree species abundance in an old-growth mesic/wet-mesic forest in central Illinois.... The study site has high deer density (75 deer km2) during winter months, and initial observations indicated that pawpaw (Asimina triloba (L.) Dunal) was strongly avoided as browse. Paw paw density increased in the seedling stratum between the two sample periods. However, nearly all other tree species declined in density" 2014, "Intensive Selective Deer Browsing Favors Success of Asimina triloba (Paw Paw) a Native Tree Species".   ABOVE: In contrast to the invasive Amur honeysuckle, a pawpaw subcanopy welcomes native spring ephemeral herbs. (Photos by Connie Barlow, 2 May 2021, Draper-Houston Meadows Preserve.) WHERE TO PLANT PAWPAW SEEDS FOR FOREST HEALTH AND REWILDING: Pawpaw likely has a much greater habitat capability than just the strips of sloping and raised areas near a river. Rivers likely became the dominant means for pawpaw seed dispersal when the megafauna disappeared — and even moreso when black bears retreated as Euro-American colonizers arrived and began farming and grazing operations. Below is a natural history observation about a wild, fruit-producing pawpaw patch in an upland forest in southwestern Virginia. The email came from Sharon Mohney, sent on 17 December 2021: "... This year was my best pawpaw season so far, and I missed probably half of them due to having to go out of town. You'd think a lot of wildlife would eat them, but they tend to just rot on the ground if I don't get them, although wasps and bees get some of them, and I found a box turtle eating one once. It's surprising to me that the deer, bear, possums, and raccoons around here don't scoop up all the pawpaws before I get a chance at them, but they don't.      In recent years I've noticed a lot of pawpaws showing up on higher drier sites on our place. I didn't think much about it until I noticed that my "Flora of Virginia" phone app also makes note of this development: "In recent decades, this species has expanded into dry-mesic or even dry upland habitats and increased in abundance in many areas. The reasons for these changes are not clear but could include fire exclusion and the plant's unpalatability to white-tailed deer (and the deer's selective browsing on competition.)"      I've had good luck planting pawpaw seeds soon after I clean the fruit. I understand they don't do well if allowed to dry..." • VIDEO: "Helping Subcanopy Trees Migrate" - 50 minutes - published November 2021    "Helping Subcanopy Trees Migrate" features two subcanopy species of the eastern USA. Pawpaw, while having a long north-south reach in its historic range, can benefit from "assisted range expansion" northward. Florida Torreya is an endangered glacial relict for which citizens, including Connie, have done what the official recovery program implementers have been unwilling to do — restore its health by nothing more difficult than planting seeds well to the north. Indigenous values are advocated as well as the "natural history" style of observation and interpretation. • WIKIPEDIA "Asimina triloba: Habitat Restoration": "Pawpaws are sometimes included in ecological restoration plantings, as they have many characteristics that make them ideal for repair of riparian ecosystems. The tree's fondness of wet soil and tendency to multiply clonally to form dense and well-rooted thickets can protect against erosion and runoff. As a native species, pawpaw can be planted on river slopes for erosion control, as introduced species formerly used in the eastern United States for this purpose (such as bamboo and Amur honeysuckle) are now discouraged or prohibited because of their invasiveness. The nonexistent commercial demand of pawpaw timber also protects trees used for ecological reasons from potential future harvest.      In the eastern United States, where large predators are almost entirely lacking, pawpaw is one of the few native subcanopy trees whose bark and leaves are too poisonous for deer to browse. It is therefore a viable species for forest understory restoration in areas where fragmented landscapes, dwellings, and parks status preclude hunting as a population control." • RETURN TO TOP Appreciating Asimen: Original Instructions • TO DO: We seek to recruit a tribal person to guide and contribute to this section (and to ensure that the previous sections are complete and the language appropriate.) Until an indigenous author for this section arises, the below content (contributed by Connie Barlow), will need to suffice. • "Let Our Indigenous Voices Be Heard", a 2017 document coauthored by Robin Kimmerer (Potawatomi), Ph.D., Rosalyn LaPier (Blackfeet/Métis), Ph.D., Melissa Nelson (Anishinaabe), Ph.D., and Kyle Whyte (Potawatomi), Ph.D. As original peoples, we have long memories, centuries old wisdom and deep knowledge of this land and the importance of empirical, scientific inquiry as fundamental to the well-being of people and planet.     Let us remember that long before Western science came to these shores, there were Indigenous scientists here. Native astronomers, agronomists, geneticists, ecologists, engineers, botanists, zoologists, watershed hydrologists, pharmacologists, physicians and more—all engaged in the creation and application of knowledge which promoted the flourishing of both human societies and the beings with whom we share the planet. We give gratitude for all their contributions to knowledge. Native science supported indigenous culture, governance and decision making for a sustainable future — the same needs which bring us together today.     ... Indigenous science provides a wealth of knowledge and a powerful alternative paradigm by which we understand the natural world and our relation to it. Embedded in cultural frameworks of respect, reciprocity, responsibility and reverence for the earth, Indigenous science lies within a worldview where knowledge is coupled to responsibility and human activity is aligned with ecological principles and natural law, rather than against them. We need both ways of knowing if we are to advance knowledge and sustainability.     ... While Indigenous science is an ancient and dynamic body of knowledge, embedded in sophisticated cultural epistemologies, it has long been marginalized by the institutions of contemporary Western science. However, traditional knowledge is increasingly recognized as a source of concepts, models, philosophies and practices which can inform the design of new sustainability solutions. It is both ancient and urgent.      Indigenous science offers both key insights and philosophical frameworks for problem solving that includes human values, which are much needed as we face challenges such as climate change, sustainable resource management, health disparities and the need for healing the ecological damage we have done...." • "Mishkos Kenomagwen, the Lessons of Grass: Restoring Reciprocity with the Good Green Earth", by Robin Wall Kimmerer, chapter 3 in Traditional Ecological Knowledge, edited by Melissa K. Nelson and Dan Shilling, 2018. "When we look about us on the earth, what we see is colored by our worldview and the languages that we use to describe our observations. A landscape of streams and lakes, mountains and rich valleys, shared by thousands of species of plants and animals, is understood through the lens of the western materialist worldview as a wealth of ecosystem services or natural resources. In contrast, through the lens of traditional Indigenous philosophy the living world is understood, not as a collection of exploitable resources, but as a set of relationships and responsibilities. We inhabit a landscape of gifts peopled by nonhuman relatives, the sovereign beings who sustain us, including the plants.      "... Reciprocal restoration is the mutually reinforcing restoration of land and culture, such that the repair of ecosystem services contributes to cultural revitalization, and renewal of culture promotes restoration of ecological integrity. In Indigenous communities, these reciprocal relationships may include the return of subsistence activities, the practice of traditional resource management, the restoration of traditional diets, language revitalization, and the exercise of spiritual/ethical responsibility. Concepts of reciprocal restoration also apply to mainstream society by re-engaging people with land, renewing place-based connections, and supporting cultural practices that sustain the land. Integration of TEK can support this new direction in restoration ecology, as a model for restoration of reciprocal relationships.      "... While our fluency with plant knowledge is diminishing, in both Native and non-Native communities, I have been taught that the knowledge itself is not lost. Humans may have forgotten, but the knowledge is resident in the land itself. Thus, knowledge revitalization depends as much on gaining the skills for learning from the land as it does on transmitting specific information. We need to ensure that we are educating people with the capacity to learn from the land again, to retrieve the knowledge that is held for us by the plants.... When plants are understood as teachers, it is an act of reciprocity to be an attentive student and to pass on the teachings of the plants." • "Indigenous Knowledge for Earth Healing", 2018, excerpt of AUDIO interview of Robin Wall Kimmerer, May 2016, on For the Wild Podcast, with timecodes. (27:35) "I think about Traditional Ecological Knowledge (TEK) as a way of being in a web of relationships. While nature may change, while the players may change, the stage may change, I think it's the relationship to place that endures and can guide us. So these relationships which are based in reciprocity, for example, I think will be a guide for adaptation in rapidly changing times.      "And, you know, often times we say that, as the land changes, traditional knowledge is lost. We hear this from our relatives in the north, who see their traditional knowledge threatened by climate change and by melting ice, and the disasters that are befalling them. But I've also been taught that the knowledge itself may disappear from the people, but it's not lost because that knowledge is actually resident in the land. It's the land and the rivers and the plants and the animals that are going to teach us. And it's our relationship with them that is going to enable us to learn.      "So while it's super important to maintain and revitalize traditional knowledge, I think that an element that we also have to focus on is revitalizing our ability to learn from the land — to be better students of the land again." (39:22) "Renewing our subjective relationship with place I think is at the heart of renewing that relationship of reciprocity: to see the world as made of gifts and not natural resources." • The Great Lakes Indian Fish & Wildlife Commission (GLIFWC) was established in 1984, as an agency of eleven Ojibwe tribes in Michigan, Wisconsin, and Minnesota. Although these tribes may all be associated with "ceded territory" northward of Asimen's current native range, continuing climate change could make those geographic locales welcome range expansions for Asimen in the decades ahead. Robin Wall Kimmerer 2011, pages 257-276 "Reciprocal restoration also offers the opportunity for an immigrant culture to start becoming 'indigenous to place' by healing relationships with land and history." "This does not mean appropriating the culture of indigenous people, but generating an authentic new relationship. It means throwing off the mindset of the immigrant, including the frontier mindset of 'take what you can get and move on.' It means becoming involved with the 'language' and dynamics of the place you live — learning its landforms, weather patterns, animals, plants, waterways, and seasons." "Being indigenous to place means to live as if we'll be here for the long haul, as if our children's future mattered. It means taking care of the land as if our lives, both spiritual and material, depended on it. It involves entering into a covenant of reciprocity with the land, which includes restoration. That's what it means to become indigenous to place." • RETURN TO TOP Where Pawpaw is a "Threatened Species" State of New York Washtenaw County MICHIGAN is the source and focal location where the idea for this Pawpaw Pollinator Watch project began. Opportunities for applying results are direct and obvious for indigenous peoples and thus there is great potential for enhancing reciprocity on tribal lands in this state. Study results may also benefit commercial growers of pawpaw in any state. It is probably a stretch, however, for managers of natural areas "preserved/protected" by dominant culture to consider "intervening" for the purpose of encouraging fruit production in fruitless wild pawpaw patches — whether that action be introducing genetic diversity, creating canopy gaps, or (least interventionist) enhancing habitat for nurturing crucial pollinators. Exceptions may include where tribal groups maintain gathering rights on "ceded" lands and where such groups advocate for management interventions of public lands toward that end. Therefore, only in a state where pawpaw is regarded as imperiled could managers of nature preserves easily draw upon arguments from the "conservation" worldview and practice for intervening in ways that could augment the wellbeing and future viability of this species. NEW YORK is such a state. This, from the pawpaw entry of wikipedia: Fortunately, basic research toward this end has already been completed and published for the northwestern region of New York where pawpaw is present, but only spottily and rare. The map below appears in a 2021 paper by Stephen Tulowiecki, a professor of geography at State University of New York, Geneseo: "Modeling the geographic distribution of pawpaw (Asimina triloba [L.] Dunal) in a portion of its northern range limits, western New York State", by Stephen J. Tulowiecki, 2021, in Plant Ecology. EXCERPT: "... Potential pawpaw habitat occurs on or near three reservations: Cattaraugus, Tonawanda, and Tuscarora Indian Reservations. Conservation or introduction of pawpaw on tribal lands may therefore present an opportunity for reciprocal restoration, described as a 'positive feedback relationship between cultural revitalization and land restoration' (Kimmerer 2011)." Previous research by the same author (Tulowiecki, 2015) offers additional background by having mapped for one county in northwestern New York archeological, historic, and forest ecological evidence of the fullness of indigenous occupation and effects on the landscape. Although the focus was on mast-bearing trees (oak, hickory, chestnut) and thus pawpaw was not evaluated, this research can be used (in combination with the 2021 mapping) for identifying places to begin pawpaw restoration and enhancement in Chautauqua County: "Native American impact on past forest composition inferred from species distribution models, Chautauqua County, New York", by Stephen J. Tulowiecki and Chris P.S. Larsen, 2015, in Ecological Monographs. NOTE: Connie notified Prof. Tulowiecki, SUNY Geneseo about this Pawpaw Pollinator Watch project. While he found it "valuable and fascinating," several factors made it impossible for him to join in Spring 2021 fieldwork: (1) pawpaw patches in New York are few; (2) they are mostly on private property, (3) far from his campus, and (4) "pawpaw typically bloom in mid-May in western New York, after the end of the school year." Also, he mentioned that this being a "threatened species" in the State of New York, there may be requirements for gaining state authorization to work with this species. Nonetheless, he wrote, "I do have a few Seneca contacts that might be interested in collaborating, on the restoration side of things." Connie will post on this page if any collaborations with the Seneca in New York State (or elsewhere) do develop. • RETURN TO TOP What About American Persimmon?       PHOTOS: Native range of American persimmon and its downward-facing flower. Michigan is beyond the northern-most range of American persimmon. Even so, Marc Boone successfully grows persimmon cultivars next to his orchard of pawpaw in southeastern Michigan. QUESTIONS: What pollinators visit the flower of persimmon, and when does the bloom occur? Overall, is there some advantage for pawpaw pollination success in having these two native fruits (both of tropical plant families) planted together? A publication of MSU Extension reports that "persimmon pollinators are unknown insects," Unusual Fruit Plants for Gardens in the North Central Region. Finally, in a "helping forests walk" way, American persimmon (Diospyros virginiana) could be wild-planted now even in southern Michigan. Unlike pawpaw, persimmon requires full sun. Therefore, abandoned farm fields or full-sun edges of woodlots growing alongside farm or grazing fields would be the kinds of sites to look for. And even in the more southerly states, folks who value native persimmon know that the fruit is inedible before a frost ripens it. So Michigan frosts could assist the ripening process. A 1909 book, Trees Every Child Should Know, reports: "Persimmons improve, the longer they hang upon the trees. As late as January or February, little trees scarcely a dozen feet high ... are found to be hung with fruits exceptionally large and fine." This 1953 paper, "The Distribution of Diospyros virginiana", by Harry R. Skallerup (in Annals of the Missouri Botanical Garden), is helpful not only for learning the natural history of this species. Several distributional findings in this paper suggest that indigenous peoples managed lands in ways that favored persimmon. But colonizing settler takeovers of these sites meant that human assistance in preventing overtopping of sun-dependent persimmons by taller trees was lost: "... Evidently, persimmon was more of a dominant tree (as were others) in the primeval forest than it is in the second growth timber of today... Although persimmon once was known to occur in pure, dense stands, more recent reports indicate that this is now not the case. Throughout its range persimmon is reported as a minor species in older association." • RETURN TO TOP Table of Topics for Parts 1 & 3 (on separate webpages) PART 1: Who Are the Pollinators? • Conclusions & Recommendations for Attracting Pollinators • Video Documentation of Pawpaw Pollinator Fieldwork • Collecting and Distributing Seeds • Other Anatomical Features of Pawpaw • Basic Elements of the Pollinator Study • Sites (Fruited and Fruitless) for Field Observations and Comparisons • STUDY AIMS with Three Alternative Hypotheses re POLLINATION • Pawpaw as an ancient flower type (CRUCIAL READING FOR SERIOUS OBSERVERS) • Pawpaw Flower Stages • Food for Pollinators • Pawpaw Pollination: Knowns and Unknowns • Probable Pollinators of Spring 2021 - Nitulidid Beetles • APPENDIX: PHOTOS of Spring 2021 surveillance at sites (insects, pawpaw stages, ecology) • South African Cycad Paper Distinguishes Casual Visitors from Effective Pollinators • Ecological Interpretations of 2021 Field Experience • BACKGROUND PAPERS for Ascertaining Ecological Interpretations • RECOMMENDATIONS for ORCHARD HABITAT MANAGEMENT • FURTHER QUESTIONS and SUGGESTIONS FOR EXPERIMENTS • Guidance for the Future Pollinator Watches • Additional POLLINATOR Information from Technical Papers PART 3: WILD-PLANTING EXPERIMENTS • BASELINE DOCUMENTATION of a Wild Patch (Saline River, Michigan, 2020 & 2021) • Learning about Germination (2023) • Helping Forests Walk: Assisted Migration of Pawpaw • Helping Forests Walk 2025 wild-planting success along HURON RIVER, Michigan • Planting Thornapple River Slope (2022) Return to linked Table of Topics Bio and Contact Info of Connie Barlow