Pawpaw Pollinator Watch
Project overview and information for
Spring 2021

UPDATE of 11 April 2021 by Connie Barlow

ASIMEN is the traditional Potawatomi term

Henceforth, "PAWPAW" and "ASIMEN" will be used interchangeably in this project.
Consult the Asimina triloba entry in Wikipedia.

PROJECT INTRODUCTION: Who, What, When, and Why


SOUTHERN MICHIGAN hosts the study sites for this joint citizen science / tribal observation project, which begins Spring 2021. Borrowing from the University of Michigan's Land Acknowledgment Statement:

"We acknowledge that the University of Michigan, named for Michigami, the world's largest freshwater system and located in the Huron River watershed, was formed and has grown through connections with the land stewarded by Niswi Ishkodewan Anishinaabeg: The Three Fires People who are Ojibwe, Odawa, and Potawatomi along with their neighbors the Seneca, Delaware, Shawnee, and Wyandot nations."


CONNIE BARLOW, citizen science volunteer and retired science writer, who wrote about pawpaw's loss of megafaunal seed distributors in her 2001 book, The Ghosts of Evolution: Nonsensical Fruit, Missing Partners, and Other Ecological Anachronisms, pp. 105-109.)

SHAWN SEVERANCE, naturalist, Washtenaw County Parks & Recreation Dept.

JEREMY JUBENVILLE, entomologist, Michigan State University Extension

SEEKING AN INDIGENOUS ADVISOR for communicating to collaborators and participants Traditional Ecological Knowledge (TEK) of reciprocal relations and responsibilities with asimen, while co-creating a fuller range of observational and interpretive practices toward better understanding of and relating with asimen. Note: Initial communications with JENNIFER KANINE, director, Dept. of Natural Resources, Pokagon Band of Potawatomi, are greatly appreciated.

NOTE ON TEK ENHANCINGING WESTERN SCIENCE (SEK): This webpage will embed excerpts from writings on TEK by Indigenous authors. To begin, this from Robin Wall Kimmerer, 2002:

"TEK is highly rational, empirical, and pragmatic, while simultaneously integrating cultural values and moral perspectives. With its worldview of respect, responsibility, and reciprocity with nature, TEK does not compete with science or detract from its power but extends the scope of science into human interactions with the natural world."

EMPIRICAL GOAL - IDENTIFY THE EFFECTIVE POLLINATORS. The primary shared goal of Spring 2021 fieldwork is to observe, distinguish, and document the effective pollinators of asimen at the study sites. Observers will be offered guidance in how to ascertain insect types and behaviors for distinguishing true pollinators from casual insect visitors.

INSPIRATION FOR LEARNING. Quotation from "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:

"... 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."
ANCILLARY GOALS. Outcomes from this pollinator study may be extended by the diverse collaborators in these possible ways:
CONNIE BARLOW - "As a practitioner of traditional "natural history" forms of field "work", I am keen on learning whether this form of qualitative, slow, and open observation might complement quantitative, single-focus forms of modern western science. I am also keen on learning whether this older form of western science might serve as a bridge for appreciating the efficacy and relational richness of TEK, "Traditional Ecological Knowledge." I shall also assess how the results of this study and collaboration might feed into my own advocacy for "assisted migration" of climate-threatened native trees, how Indigenous perspectives would reshape underlying worldviews (as well as suggested actions) in this endeavor, while striving for wider recognition of the Haudenosaunee phrasing "helping forests walk" for appreciating that native peoples in North America have by far the most experience in helping native plants move poleward. Clearly, they have been doing so for centuries (possibly for thousands of years), and asimen is likely one of the beneficiaries."

SHAWN SEVERANCE - Her original intention was to introduce participants to the "bioblitz" approach for rapid gain of biodiversity tabulation in an area, while amplifying citizen appreciation and natural history understanding of native asimen in Washtenaw County (and other) natural areas, possibly extending into suggesting new management options for improving reciprocal interactions with this being and its pollinators in wild or near-wild pawpaw patches in the county.
     April 8 update: Shawn cannot co-lead this effort Spring 2021 due to obligations with family members having medical needs and injuries.

JEREMY JUBENVILLE - advising artisinal and commercial cultivators of pawpaw in Michigan for achieving pollination goals, while amplifying regional appreciation of asimen as a seasonally available local fruit.

GENUS NAME DERIVES FROM INDIGENOUS: Linnaeus originally classified this plant as Annona triloba. Later, Micheaux renamed the species Orchidocarpum arietinum. Four years later, Persoon again transferred the species to Porcelia triloba. Finally, in 1817, Dunal reclassified the plant as Asimina triloba. This final genus name, Asimina, derives from indigenous naming. Asimen is the traditional Potawatomi term. Note that the settled scientific name, Asimina, is effectively a "latinized" version (add the suffix "a") to the spoken Potawatomi word. Henceforth, "PAWPAW" and "ASIMEN" will be used interchangeably in this project.

For more information on name derivation from indigenous languages, see p. 191 in Pawpaw: In Search of America's Forgotten Fruit, by Andrew Moore, 2015.

Table of Topics
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Study Sites Spring 2021

STUDY AIMS with Three Alternative Hypotheses

Pawpaw Flower Stages and Pollinator Types: An Overview

Details on Flower Stages and Wild Patch Characteristics

Details on Likely Pollinating Beetles

Guidance for the Spring 2021 Pollinator Watch

Appreciating Asimen: Geography and Taxonomic Relatives

Appreciating Asimen: Prevents Invasion by Japanese Stiltgrass

Appreciating Asimen: Ethnobotany

Toward Renewing and Expanding Reciprocity

New York State: Where Pawpaw is a "Threatened Species"

What About American Persimmon?

Appreciating Asimen: Original Instructions

Study Sites Spring 2021

PHOTOS: Marc Boone Farm by Connie Barlow 2020 (left) and Pawpaw patch in Draper-Houston Preserve by Dallas Ford 2020 (right).
LEFT: Boone Farm pawpaw rows are foreground. RIGHT: Along Saline River, Barlow is touching pawpaw stems.

SHAWN SEVERANCE and CONNIE BARLOW visited two contrasting study sites in Washtenaw County MI (near Ann Arbor) in autumn 2020:

MARC BOONE FARM: Photo above left early October. Pawpaw cultivars are planted in rows. The fully mature, short trees are in foreground, with autumn-yellow leaves. They are in full sun, with mowed lawn that prevents clonal proliferation of non-cultivar rootstock. The orchard is given no supplemental water(?) Marc reports sufficient wild pollination, though an early spring punctuated by severe frost in 2020 reduced fruiting. Because the fruits are large and heavy, he removes flowers toward the outer reaches of the branches(?) Note: American persimmon cultivars are also grown in full sun on this farm.

DRAPER-HOUSTON MEADOWS PRESERVE: Just west of the city of Milan, the preserve protects over one-half mile of Saline River shoreline, with a quarter mile of continuous pawpaw understory along the higher bank side. Notable large (and old) canopy trees include Swamp White Oak, Hackberry, Sycamore, Cottonwood, Black Ash, and Black Walnut. (Pawpaw, unlike many other subcanopy plants, is well adapted to grow beneath Black Walnut.) Following two site visits, Connie Barlow suggests that pawpaw here is a continuous patch that appears to have fully expanded to all contiguous sites accessible by lateral roots. Lower, swampy domains prevent further extensions.


ABOVE LEFT: Pawpaws nearly ripe at Boone Farm, early October 2020.

ABOVE RIGHT: Many stems at Draper-Houston show vertical cracks; maximal stem diameters about 1 foot, but roots may be centuries old. If such cracks predominate on the southwest side of the trunk, then it is possibly a freeze-thaw-freeze problem for this thin-barked tree, as offered in a comment to this very question: "Sunscald occurs on the SW side of the tree on a cold day. The sun heats the trunk up on the SW side and then passes behind a cloud or hill. The rapid temperature change causes the bark to separate from the tree causing a wound that usually doesn't heal."


ABOVE LEFT: Pawpaw is easily identified in winter by its flexible furry terminal leaf buds and its round furry flower buds.

ABOVE RIGHT: Stems last perhaps half a century maximum. However, the clone persists and expands via lateral roots extending and then sprouting new stems ("ramets"). Photo here was taken by Connie Barlow at Draper-Houston, autumn 2020.

STUDY AIMS with Three Alternative Hypotheses

• Discover why some pawpaw patches produce little or no fruit.

Co-leaders SHAWN SEVERANCE and JENNIFER KANINE each report large differences in fruit production (including complete absence of fruiting) among wild sites in their local areas of s. Michigan. As confirmed in a pawpaw study in Toledo, Ohio (see below), fruitless patches in the wild are not uncommon. Meanwhile, the Marc Boone Farm near Ann Arbor produces prolific fruit — and insects visiting Marc's pawpaw orchard in bloom have never been carefully observed and identified.
    After visiting a full-sun commercial orchard (October 2020) and then a fruitless wild patch near Ann Arbor MI with SHAWN SEVERANCE, CONNIE BARLOW posed several hypotheses for the lack of fruit production, beginning with the missing pollinator idea, drawn from the
pawpaw section of her 2001 book, The Ghosts of Evolution: Nonsensical Fruit, Missing Partners, and Other Ecological Anachronisms.

HYPOTHESIS #1. FEW POLLINATORS: Asimen is a classic beetle-pollinated flower, as has been confirmed through careful observation of four sister species of genus Asimina whose native ranges are much farther south. Only one of those sister species, A. parviflora has the same flower type as pawpaw: downward hanging, maroon color, and with an odor described as yeasty or fetid. A. parviflora, however, is smaller than A. triloba (its common name is Dwarf Pawpaw). Known beetle pollinators of genus Asimina require rotting hardwood standing or fallen trees (such as oak) for their larval stages. Thus, if our citizen watch detects differences in pollinator visitations between the fruiting and non-fruiting sites, a next step will be to determine if differences in rotting wood availability may be implicated — or whether agricultural spraying of neonicotinoid pesticides in the area may be implicated.

HYPOTHESIS #2. INADEQUATE SUNLIGHT: Pawpaw natively is a subcanopy tree of deciduous hardwood forests. It thus can be expected to self-limit flowering to the sunniest stems and branches within a clonal patch and may also halt development of successfully pollinated flowers owing to inadequate sunlight. Observers will be asked to offer their qualitative judgments as to whether access to sunlight seems to explain flowering sites and abundance in their watch area. Perhaps a publishable form of data gathering will be initiated in a follow-up stage of study.

HYPOTHESIS #3. LOW GENETIC DIVERSITY: Pawpaw reproduces vegetatively by sending forth root suckers. Over the course of decades and centuries a single genotype can expand its roots and stems ("ramets") throughout vast subcanopy areas. Scientific research has confirmed that wild patches tend to have very low genetic diversity, despite high diversity between patches. Because inbreeding can be detrimental, plants generally have ways to reduce "selfing" (self-pollination). The flower structure of asimen appears to discourage selfing.

Hypothesis #1 will be tested in this Pawpaw Pollinator Watch study. Hypothesis #2 will receive qualitative evaluation by comparing canopy light openings in flowering-rich v. poor microsites in the wild pawpaw patch. Hypothesis #3 will require genetic testing by professionals. Nonetheless, published papers suggest that all three potential causes may be at work.

This PAWPAW POLLINATOR WATCH is a mixture of both the pollination and low-genetic diversity hypotheses in this way: A 1985 paper (citation below) concluded:

"Given their strong protogyny [the female stage in flowers begins and ends before the pollen stage] and clonal habit [spread into large patches by underground roots], fertilization may require the arrival of pollen from a considerable distance."
This conclusion was based on wild patch fruiting distinctions that the team had documented:
Fruit production, however, varied strikingly among the stands. No fruits were produced by any of the 524 trees in the Toledo, Ohio stands. In the Cincinnati stands 11 of the 75 sample trees produced a total of 21 fruits.... Reasons for the total failure of fruit set at Toledo are unclear. Bowden and Miller (1951) concluded that the pawpaw in southeastern Ontario occurs in areas of low elevation that have "sufficiently long frost-free seasons" to permit the development of fruit. However, length of growing season does not seem to account for the failure of fruit set in our stands. At our Toledo sites nearly all flowers hung on the trees to maturity, withered, and fell. Few were aborted prematurely, and official daily minimum temperatures at Toledo's U.S. Weather Bureau during flowering were so far above 0°C that we do not believe frosts could have occurred in our stands. Subsequent visits to site T2 have revealed no fruit set in 1981 or 1982. Apparently the failure of fruiting in Toledo is an annual phenomenon independent of variations in the local climate regime. ... Protogyny combined with highly synchronous flowering and the need for outcrossing would make successful pollination a relatively rare event, especially in the Toledo stands, which are separated from other pawpaws by the distances of several to many km.
    Citation: "Geographic Variation in Size and Reproductive Success in the Pawpaw (Asimina triloba)", by Renee LaGrange and Elliot Tramer, 1985, Ohio Journal of Science.
LITERATURE SEARCH: Barlow conducted a literature search, which confirmed that Asimina triloba has not yet had published observations of floral insect visitors that adequately distinguish between casual visitors and effective pollinators. This paper, "Pollination and Evolution in Neotropical Annonaceae", by Gerhard Gottsberger (2008, Plant Species Biology) noted that while this ancient family of tropical flowering plants typically is pollinated by beetles, the flowers of Asimina triloba:
"... are of a dark maroon or purple color and exhale an unpleasant odor. Observations, however, are scarce and not well documented... Therefore, it is not yet clear whether this species is in fact pollinated by flies or by beetles, or by both."
• Discover the source(s) of POLLINATOR LARVAL FOOD at Marc Boone Farm.

Because the planted full-sun orchard at Marc Boone's farm produces fruit in abundance, we can expect to document species and abundance of truly effective pollinators (likely, beetles). Hence, an additional study element will be to locate the microsites at or near Boone's property that serve as home and food for the larval stage of the pollinators. As detailed in the next section, published sources indicate that pawpaw's pollinating beetles do not lay their eggs (and thus feed their larval stage) on carrion or dung. Rather the beetles seek out rotting hardwood trees and logs (or wood chips or leaf mulch thereof).

Pawpaw Flower Stages and Pollinator Types: An Overview

Photo by Eric Hunt, Flickr.

ABOVE: The flowers open and become receptive before the leaves are fully extended.

Photo by Mieke Janssens, Flickr.

ABOVE: Flowers prevent "selfing" by having the female portion of the central reproductive structure begin and end its several-day receptive stage (left) before the male (pollen-bearing anthers, right) begin theirs.

Photo credited to NatureServe, Flickr.

Photo by Robert Wolkowitz, Flickr.
   PHOTOS ABOVE & LEFT: Flies and even smaller insects (e.g., thrips) are the most common visitors to pawpaw flowers, but they are unlikely to be effective pollinators. Insects that have pollen adhering to their backs (see photo inset above) or none at all cannot pollinate the female tips of the central flower structure.

PHOTOS BELOW: True pollinators are too heavy to cling to the inner surface of a petal. Rather, they are likely beetles who fly under the downward-hanging flower and then flip, clinging to the female tip of the reproductive stalk and then walking upwards to the food-rich petal depths. During the male phase of pawpaw flowers, pollen would gather on the beetle's underside while walking upward. During the female-receptive stage, pollen would transfer immediately as the beetle landed, and no additional pollen would gather (or fall downward) as the beetle walked upward across the unripe anthers.

Overall, effective pollinators gather pollen on their undersides. Beetles with hair-like appendages on sides and underside are called "flower pollinating beetles" precisely for this reason. (See below photos.)

Photos from
Maryland Biodiversity Checklist

ABOVE: Published scientific papers have reported two genera of scarab beetles as effective pollinators of more southerly species of genus Asimina (whose flowers are smaller than the northern species, but have similar structure): Trichiotinus (left) and, to a lesser extent, Euphoria (middle and right). Notice that the photo on right shows Euphoria in the context of leaf litter. Flower pollinating beetles generally are neither carrion beetles nor dung beetles. That is, they lay their eggs not in rotting meat or dung but into rotting wood or thick leaf mulch, where larval grubs feed and eventually pupate. Key papers on Asimina species that identify beetle pollinators:

"Effects of Fire and Pollinator Visitation on the Reproductive Success of Asimina reticulata (Annonaceae), the Netted Pawpaw", by Louise K. Barton and Eric S. Menges, 2018, Castanea. (Note: Trichiotinus viridans, the "hairy flower scarab" is documented as the most prevalent visitor, and "low fruit set may be due to the lack of an effective pollinator, resulting in inbreeding depression.... Our study confirms that A. reticulata is self-compatible only if pollen is able to move among flowers, and we found that, in most cases, fruit was aborted prematurely and was found littering the ground beneath the plants.")

"How diverse are Annonaceae with regard to pollination?", by Gerhard Gottsberger, 2012, Botanical Journal of the Linnean Society. (Note: In this paper two Florida species, Asimina obovata and Asimina pygmaea are linked with pollinating beetle species in Trichiotinus and Euphoria genera. These flowers are categorized as "diurnal without thermogenesis.")

Photos from Maryland Biodiversity Checklist, Fritz Flohr Reynolds.

ABOVE: The American carrion beetle, Necrophilia americana, requires carrion for egg-laying and its larval stage, but as an adult it will happily feed on mushrooms — and fermenting pawpaw fruit. Note the lack of "hairs" on this beetle and thus its inability to serve as a pollinator.

A 2006 paper by Katherine R. Goodrich et al., "When Flowers Smell Fermented: The Chemistry and Ontogeny of Yeasty Floral scent in Pawpaw (Asimina triloba)", was published in International Journal of Plant Science. It offers evidence that carrion beetles are not involved in pawpaw pollination: "We did not detect the dimethyl oligosulfides that characterize the microbial decomposition of meat and universally constitute the odors of carrion-mimicking flowers." Instead, the authors reported a "yeastlike fragrance" in pawpaw flowers. However, "fermentation volatiles were largely absent from female-stage androgynoecia". (Many technical details in this paper may be useful for project participants.)

RECOMMENDED READING: In addition to various technical papers cited and linked here, a 296-page book on pawpaw is highly recommended: Pawpaw: In Search of America's Forgotten Fruit, by Andrew Moore, 2015, Chelsea Green.

Details on Flower Stages and Wild Patch Characteristics

Photo by Suzanne Cadwell, Flickr.

ABOVE: Flowers even on the same branch may be in different stages of development and receptivity.

TIMING OF FEMALE & MALE STAGES OF SEXUAL DEVELOPMENT: The definitive paper on the timing of stigma v anther receptivity and ripening also contains many color photos to aid observers with recognizing flower stages: "Pollen-pistil interaction in pawpaw (Asimina triloba), the northernmost species of the mainly tropical family Annonaceae", by Juan M. Losada et al., 2017, American Journal of Botany.

POLLINATION PROBLEMS: The only paper that deals centrally with pollination problems and effective pollinators of our northern-most Asimina species, pawpaw, was published four decades ago: "Pollinator limitation, fruit production, and floral display in Pawpaw (Asimina triloba)", by Mary F. Willson and Douglas W. Schemske, 1980, Bulletin of the Torrey Botanical Club. Effective pollinators at this study site in central Illinois were not detected for this reason: "Flowers on our study area were rarely visited [by insects]; several extended observation periods yielded no evidence of insect visitation." FLORAL STAGES are, however, well described in this paper, with excerpts as follows:

... Pawpaw was most common in old light gaps, some of which had virtually closed. Pawpaw clones extensively and forms such a dense shade that summer understory vegetation is much reduced. We do not know genetic relatedness among the stems we censused but suspect that most stems at a site may be genetically identical.

... Time from opening of the bud and exsertion of the stigma [female tip] from the clump of tightly-packed, closed stamens to petal-drop ranges from 5 days to > 19 days (in a sample of 18 flowers), with most flowers open 15 or 16 days. Petals are green when the flower first opens and gradually redden from pinkish to maroon. The stigma becomes shiny (probably receptive) after 5 to 10 days, remains so for 4 to 6 days, and then turns brown. Pollen usually becomes available when the stigma turns brown, but occasionally earlier. Pollen is available for about 2 days, and sometimes after the petals drop. Flowering times vary between years: in 1977 blooming began in mid-April, 1978 it began in early to mid May. We do not know if such annual differences are accompanied by adjustments of intrafloral phenology.

... There is a clear tendency for flowers on large stems, on high branches, and perhaps exposed to more light, to contain more ovaries.... The sites differed markedly in fruit production, ranging from 1 to 30% of flowering stems with fruit. Furthermore, fruit-bearing stems were not randomly distributed in each site. In two sites, almost all fruit-bearing stems were clumped in areas where the canopy was most open, and each site contained groups of flowering stems that produced no fruit. Despite the fact that flower production was often considerable, with many stems producing > 100 flowers, no stem produced > 4 fruits in 1976. In 1978, fruit production by some large stems reached at least 19. Only 0.41% of the 19,095 flowers censused in 1976 produced fruit.... Fruit production was clearly not associated with the numbers of flowers per site.... Fruiting stems are significantly larger than flowering stem.

... In sharp contrast is the relative success of hand pollinations. In 1978, 804 hand-pollinated flowers produced 135 fruit (17%); in 1977, 78 flowers bore four fruit (5%), and this figure is low because many flowers were not pollinated during the receptive phase. Because abortion of young fruits is rare (< 1%), these results suggest clearly that fruit production by pawpaw at Trelease is pollen-limited.... Pawpaw fruits are green until they are mature and potentially can provide photosynthate for their own growth.

... Flower-visitors are uncommon, which may select for a fairly long floral life and period of female receptivity.

... The fruiting success (and seeds per fruit) of hand-pollinated flowers was much greater than that of open-pollinated flowers and was unaccompanied by fruit abortion. We conclude that at least in our study area, fruit production was pollen-limited, yet it seems certain that a stem could neither mature nor support fruit from all ovaries and flowers produced. The tremendous flower output (up to 500 flowers/stem) may be related to problems of pollen-limitation. Since most flowers do not produce fruit, large floral displays, although perhaps energetically expensive, may increase pollinator visitation rates.

... The great variation in reproductive effort allocated to paternal and maternal functions of flowers is intriguing and worthy of further study. It is not difficult to understand low number of ovaries per flower, especially if local photosynthate is used to support nearby fruits on each branch. We never observed more than four fruits matured per flower on either hand- or naturally pollinated flowers at both Trelease and Mingo. However, the production of 9- and 10-ovaried flowers on some branches is less readily understood, inasmuch as it is virtually inconceivable that all could be matured.

CONFIRMATION OF NECTAR AT BASE OF THE INNER WHORL OF PETALS: "The mature flowers have an outer and inner whorl of three, maroon-colored, three-lobed petals (Fig. 7.1). The inner petals are smaller and fleshier, with a nectary band at the base." This statement appears on p 353 of "The North American Pawpaw: Botany and Horticulture", a paper published in 2005 in Horticultural Reviews by a pair of well-known pawpaw horticulturalists: Kirk W. Pomper (Kentucky State University) and Desmond R. Layne (Clemson University). The paper is rich in details for aiding cultivation of this species, so offers little assistance on understanding wild patches. One sentence simply recounts previous papers pointing to beetles and flies, though mention is also made that "Failure of trees to set fruit could be due to inadequate pollination." More broadly,
"The tropical Annonaceae relatives of the pawpaw, such as cherimoya, sweetsop (sugar apple), soursop, and atemoya also have low yields, due to low rates of natural pollination (Peterson 1991; George et al. 1992; Pena et al. 1999). In commercial plantings, these tropical pawpaw relatives are hand pollinated to increase yields (Peterson 1991; Pena et al. 1999). Low rates (<5%) of fruit set have also been noted in wild pawpaw patches (Willson and Schemske 1980; Lagrange and Tramer 1985).

Details on Likely Pollinating Beetles

"The diversity and evolution of pollination systems in Annonaceae", by Richard M.K. Saunders, 2012, Botanical Journal of the Linnean Society:

... There are two distinct floral forms within genus Asimina. One group (consisting of A. parviflora and A. triloba) possesses small maroon flowers that emit a foetid aroma; and the other group (consisting of all other species) possesses large, white, pink or yellow flowers that emit a more pleasant fragrance (Kral, 1960; Goodrich & Raguso, 2009). These differences clearly represent different pollination syndromes. Observations of floral visitors confirm that the foetid-smelling species are visited by small flies and/or small beetles (Willson & Schemske, 1980; Norman etal., 1992; Rogstad, 1993), whereas the fragrant species are typically visited by large beetles (Norman & Clayton, 1986; Norman et al., 1992) or rarely by flies (Uphof, 1933; Norman et al., 1992).
    ... Annonaceae flowers are visited by a taxonomically diverse range of insects: predominantly beetles (Coleoptera), but also thrips (Thysanoptera), flies (Diptera) and, rarely, bees (Hymenoptera) and cockroaches (Dictyoptera). Species are typically pollinated by only one of these groups, although there are several examples of species that are pollinated by more than one group, including: .... Asimina parviflora (Norman et al., 1992), A. pygmaea (Uphof, 1933; Norman & Clayton, 1986; Norman et al., 1992) and A. triloba (Willson & Schemske, 1980; Johnson & Willson, cited in Norman et al., 1992) are visited by flies and beetles.
    ... Despite the vast diversity of dipterans and their evident importance in pollination, there are few reports of fly pollination in Annonaceae. Flower visits by flies have only been reported for Annona (Webber, 1981b; as secondary floral visitors only), Asimina (Norman et al., 1992), Monodora (Gottsberger, 1985; Gottsberger et al., 2011), Pseuduvaria (Morawetz, 1988; Silberbauer-Gottsberger et al., 2003; Su et al., 2005) and Uvariopsis (Gottsberger et al., 2011). It should be noted, however, that Norman et al. (1992) did not observe any pollen attached to the common drosophilid flies that visited Asimina flowers (pollination resulted from visits by nitidulid beetles and rarer calliphorid flies), and that pollen transfer between flowers was not demonstrated in any of the other studies. ... To avoid confusion of stimuli, Calliphoridae (blowflies), which lay eggs in dung and carrion, favour yellow colours in the presence of sweet scents, but brown-purple colours in the presence of foetid scents (Kugler, 1956). The latter pollination system is observed in Asimina parviflora (Norman et al., 1992) and operates by DECEIT as the flies visit the flowers in order to lay eggs. Sapromyiophilous flowers, furthermore, often have partially enclosed pollination chambers that require the flies to crawl in through narrow openings (Fægri & van der Pijl, 1979).
"Phylogenetic Analysis of the North American Beetle Genus Trichiotinus (Coleoptera: Scarabaeidae: Trichiinae)", by T. Keith Philips et. al, 2016, Psyche: A Journal of Entomology.

... Adults are good fliers and forage on a variety of flowers while larvae are known to feed on various species of decaying hardwoods.... Trichiotinus is dependent upon decaying hardwoods as larval food including oak [6]. Jackson et al. [39] present evidence for a split in distribution of oaks (Quercus spp.) during the most recent glacial maximum on either side of the Mississippi drainage and may be indicative of the effects of earlier glacial maxima as well. P. A. Delcourt and H. R. Delcourt [40, 41] also postulated the presence of spruce (Picea glauca) forests in the Lower Mississippi Valley. This extension south of these more cool adapted forests all the way to the gulf coast that divided the hardwood forests into eastern and western blocks was thought to be due to glacial meltwater creating a cooler climate locally [42].

"Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Cetoniinae)", by Jesus Orozco, 2012, BioOne Complete.

"The North American Pawpaw: Botany and Horticulture", by Kirk W. Pomper (Kentucky State University) and Desmond R. Layne (Clemson University), 2005, in Horticultural Reviews. In this paper by horticulturalists, CARRION FLIES are mentioned as possible pollinators but also as not greatly effective (p. 371):

In the wild, pawpaw trees are usually found in the understory of hardwood forests. Low light levels in the understory likely result in reduced photosynthate partitioning to fruit that may cause low fruit set. Pawpaws in the wild often produce many root suckers that could potentially result in large clonal pawpaw patches contributing to poor fruit set because of self-incompatibility. Pollinator limitation can also cause low fruit set in wild pawpaw patches (Willson and Schemske 1980). Low pollinator activity is usually observed on cool, cloudy spring days. Since the pawpaw flowers are strongly protogynous (Willson and Schemske 1980), lack of pollen availability from other pawpaw genotypes could also limit pollination. Pawpaw growers report that placing carrion in buckets among pawpaw trees has resulted in improvements in fruit set (L. Sibley, pers. comm.) supporting the theory that pawpaw flowers may be pollinated by carrion flies. However, fruit set was 15 to 35% in KSU orchards in 1998 in nine-year-old seedlings where many pawpaw genotypes are in close proximity and flies are abundant due to nearby cattle. Pollinizer relationships between pawpaw cultivars need to be determined. Fruit set can be achieved by hand cross-pollination (Peterson 1997), and needs to be evaluated as a method to increase fruit set."
DISPUTING THE CARRION BEETLE HYPOTHESIS: A 2006 paper by Katherine R. Goodrich et al. published in International Journal of Plant Science offers evidence that carrion beetles are not involved in pawpaw pollination. "When Flowers Smell Fermented: The Chemistry and Ontogeny of Yeasty Floral scent in Pawpaw (Asimina triloba)" reported: "We did not detect the dimethyl oligosulfides that characterize the microbial decomposition of meat and universally constitute the odors of carrion-mimicking flowers." Instead, the authors reported a "yeastlike fragrance" in pawpaw flowers. However, "fermentation volatiles were largely absent from female-stage androgynoecia". (Many technical details in this paper may be useful for project participants.)

Guidance for the Spring 2021 Pollinator Watch

MODERN SCIENCE, NATURAL HISTORY, and TEK: We hope to have volunteers utilizing a spectrum of observational stances and techniques, ranging from the quantitative rigor of modern science (including citizen science), to the more qualitative emphases of the western tradition of natural history, and (if possible) Traditional Ecological Knowledge, TEK, as practiced by experienced or new learners of this Indigenous form of science and right relationship.

For those unfamiliar with the "Traditional Ecological Knowledge" (TEK), here is online access to and a lengthy excerpt from a 2002 article by ROBIN WALL KIMMERER, "Weaving Traditional Ecological Knowledge into Biological Education: A Call to Action", in BioScience:

"Traditional ecological knowledge is not unique to Native American culture but exists all over the world, independent of ethnicity. It is born of long intimacy and attentiveness to a homeland and can arise wherever people are materially and spiritually integrated with their landscape. TEK is rational and reliable knowledge that has been developed through generations of intimate contact by native peoples with their lands. TEK is being recognized as having equal status with scientific knowledge (UNEP 1998) and has been termed the 'intellectual twin to science' (DeLoria 1995). This long intellectual tradition exists in parallel to Western science, yet has been historically marginalized by the scientific community.
     Traditional knowledge has much in common with scientific ecological knowledge (SEK), which is not surprising since both traditions derive from the same source: systematic observations of nature. Both knowledge systems yield detailed empirical information of natural phenomena and relationships among ecosystem components. Both SEK and TEK have predictive power, and in both intellectual traditions, observations are interpreted within a particular cultural context.
     Traditional knowledge encompasses a wide range of biological information, which overlaps significantly with the content of a mainstream course in ecology or conservation biology. The scope of traditional ecological knowledge includes detailed empirical knowledge of population biology, resource assessment and monitoring, successional dynamics, patterns of fluctuation in climate and resources, species interactions, ethnotaxonomy, sustainable harvesting, and adaptive management and manipulation of disturbance regimes (Berkes 1999). Case histories of the utility of TEK in conservation biology span a range of biomes from the tundra to the tropical rainforest."
     Traditional ecological knowledge differs from scientific ecological knowledge in a number of important ways. TEK observations tend to be qualitative, and they create a diachronic database, that is, a record of observations from a single locale over a long time period. The National Science Foundation, in its support of the Long-Term Ecological Research program, has validated the importance of such continuous data. In TEK, the observers tend to be the resource users themselves, for example, hunters, fishers, and gatherers whose harvesting success is inextricably linked to the quality and reliability of their ecological observations. In contrast, scientific observations made by a small group of professionals tend to be quantitative and often represent synchronic data or simultaneous observations from a wide range of sites, which frequently lack the long-term perspective of TEK. Additional differences between scientific knowledge and traditional knowledge are described in Berkes (1993).
     Western science is conducted in an academic culture in which nature is viewed strictly objectively. In this aspect, TEK diverges significantly from Western science (Pierotti and Wildcat 2000). TEK is much more than the empirical information concerning ecological relationships. Unlike SEK, traditional knowledge is woven into and is inseparable from the social and spiritual context of the culture. Traditional knowledge can rival Western science as a body of empirical information, but traditional knowledge may also extend its explanatory power beyond the strictly empirical, where science cannot go. TEK is laden with associated values, while the scientific community prides itself on data that are "value free." TEK includes an ethic of reciprocal respect and obligations between humans and the nonhuman world. In indigenous science, nature is subject, not object. Such holistic ways of understanding the environment offer alternatives to the dominant consumptive values of Western societies (Berkes 1999, Hunn 1999). Embraced as an equal partner to the power of Western science, TEK offers not only important biological insights but a cultural framework for environmental problem solving that incorporates human values.
GUIDANCE from Western Science and Natural History Perspectives
    List prepared by science writer, Connie Barlow:

1. FIND AN OBSERVATIONAL SITE: As you enter the orchard of pawpaw at Marc Boone's farm or the wild patch along the Saline River, prepare yourself (in whatever way is natural for you), to be patient and grateful for this opportunity to experience, learn, and possibly contribute toward new understandings. Make sure you spend quality time at the beginning exploring the site as a whole in a receptive, observant way and with the intent on finding one or more specific locales for engaging in long flower-watching sessions. Ideal sites will likely include those with many flowers on branches low enough to touch and therefore to see details up close. Therefore, pay special attention to asimen trees that have low branches with multiple flowers in different stages of receptivity. Ideally, you might come upon a branch that has both a female receptive and a male receptive flower for viewing at the same time. You may find it helpful to have a magnifying lens handy — both for insect observation and for discerning the reproductive stage of flowers at your particular locale.

TEST FOR MALE RECEPTIVITY: Gently touch the anthers in a male-stage flower, and ensure that some pollen visibly sticks to your finger. Because pollen grains are released as stuck-together tetrads (4 grains together), they should be more visible than pollen from other plants.

TEST FOR FEMALE RECEPTIVITY: See if the green 3-pronged tip glistens with liquid. If unsure, gently test with your finger to feel if wet. If you can see detail, observe whether any pollen grains are already on it — and if those grains look the same as the grains on the ripe male anthers of another flower.

2. DOCUMENT SITE CONDITIONS: Make sure through notes, audios, and/or photos that each stem and branch where you watch for (and especially if you confirm) pollinator activity can be found again by you or another volunteer. This will enable additional visits in the months ahead for documenting whether fruits are actually developing. Pay special attention and make notes as to whether a flower-rich ramet or branch seems to be benefitting from canopy openings that enable more sunlight to reach the branch.

3. DOCUMENT FLOWER NUMBERS AND STAGE OF RECEPTIVITY at your observational site (through notes, audio, and/or photos).

4. GUIDANCE FOR OBSERVING INSECT TYPES AND THEIR BEHAVIOR: Even though the male (pollen) stage of an asimen flower follows the female stage, an effective pollinator must first visit a male-stage flower in order to pick up pollen before visiting a female-stage flower. Because the male-stage flower has both the outer and inner corolla of petals fully open, small insects (like thrips and flies) that are attracted to the smell and food value at the inner depths of the petals are likely to pick up pollen only incidentally, especially on their backs. Flies are likely to avoid the central reproductive structure ("receptacle") altogether and just land on the inside of an inner petal and then crawl upward toward the food, acquiring pollen on their backs (or none at all) while doing so. Because most insect visitors will likely be flies and because flies are not expected to be effective pollinators, it is crucial to remember what effective "flower beetles" look like and to focus your attention on those types. Crucially, WATCH FOR INSECT BEHAVIOR during your observations, with these questions in mind:

• FOR MALE RECEPTIVE-STAGE FLOWERS: Do potential pollinators land on the central reproductive stalk of the downward hanging flower, and then crawl upward over the pollen-rich anthers in order to reach their goal deep inside the flower? If so, do you see them acquiring pollen on their undersides, and do they spend a lot of time deep inside the flower (even out of sight)? In contrast, do the flies (being lighter) preference landing directly on the inner petals, thereby crawling deep into the flower? And do you notice whether those flies arrive and/or leave with yellow pollen grains on their backs?

• FOR FEMALE RECEPTIVE-STAGE FLOWERS: Do potential pollinators enter the flower by landing upside-down directly onto the tip of the central reproductive stalk? If they do, does it seem like they depart quickly, as if the female stage has an aroma that attracts a landing for belly pollen but the inner petals are still too tightly closed for beetles to access? And can you detect any pollen freshly stuck to the glistening wet flower tip right after the beetle departs? If you assess that pollinator landings on female-stage flowers are brief (perhaps even "accidental"), then make sure you pay a much closer watch of the female-stage flower than the male-stage (and alert other observers to do the same.)

5. IF YOU HAVE A SMART PHONE you may wish to instantly photo-document insect visitors for expert identification, by using the i-Naturalist online platform. Note: If photo-documentation is limited, then focus your photography on insect types that exhibit behavior likely to serve effectively for pollen gathering and transmittal.

6. DETAILED OBSERVATIONS OF INSECT BEHAVIOR: While the crucial observation on male-stage flowers is to discern whether an insect visitor acquires pollen, and where pollen sticks on its body, it will also be very interesting for observers to attempt to interpret what the insect pollinator gains during its visit. Possible gains are listed on p. 688 of "Visitor or vector?", by Thomas Sayers et al., 2019, in Arthropod-Plant Interactions. These benefits for insects include: pollen as food; nectar and/or floral tissue (inner thickened base of petals) as food; or conspecifics for potential mating. No gain (that is, deception) is also listed as a possibility, but other papers on Asimen clearly identify at least the inner petal base as a food source. However, the inner petal base is inaccessible to beetles during the phase of female receptivity, so it is possible that food reward occurs only during the male phase — which would account for only very brief beetle landings on early stage flowers, as photographs reveal that flowers of different stages occur even on the same branch. Thus, a possible outcome of close observation could be food reward during the male stage, but aroma-induced deception during the earlier female stage.

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.


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.

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).
   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.

PHOTO LEFT: Zebra Swallowtail laying an egg on a pawpaw stem.

There are many references to INDIGENOUS and EARLY SETTLER uses and appreciation of pawpaw fruit. For example:

"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.

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.

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."

   • BLACK ASH FIBER LOSS. 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)? And if so, would reciprocity entail helping Asimen to move northward into culturally significant woodlands in which Black Ash has been lost?

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."

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.

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?

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.

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."

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."

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