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BARTONIA 


JOURNAL OF THE 
PHILADELPHIA BOTANICAL CLUB 
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0. 62 


LEWIS AND CLARK BICENTENNIAL ts 
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BARTONIA 
Journal of the Philadelphia Botanical Club 


Since its founding in 1891, the Philadelphia Botanical Club has offered outstandin: programs, field is and other 

opportunities for those with an interest in plants to meet and exchange information. Monthly meetings feature 

speakers from various botanical backgrounds. They are held at 7:30 p.m. on the fourth Thursday a the month in 

September, October, and January through May i the third Thursday in November and December, at the Marvin 

Comisky Conference — One Logan Square (0 (one block east of the Academy of ‘Natural eryeiise of 
Phil 1. ih . . Each year A 3 CY . MS A | 


region and occasionally nae in North America or overseas. 


Officers, 2004-2005 Past officers, some pore a 2003-2004 
President: 


urt 
ee ig Secretary: No soe oe 
ula: Roger Latham Recording ct Robert W. McCombe 

Members of the Executi Editor, Bartonia: Roger Latham 


Endowment Fund Director: John C. O’Herron I 
a ! Committee Chair. Ann F. Rhoads 
Constitution Committee sae William H. Roberts 


Bartonia No. 62: 1-24, 2004 


On the Paper Trail in the Lewis and Clark Herbarium 


RICHARD M. MCCOURT’’ AND EARLE E. SPAMER*** 
"Department of Botany *Archives, Ewell Sale Stewart Libra 
The Academy of Natural ato of Philadelphia, 1900 Benjamin Franklin Parkway, 
elphia, Pennsylvania 19103 
3Current address: 0 Cresheim Road, Philadelphia, Pennsylvania 19118 
*Current address: 1 Locust Court, Maple Shade, New Jersey 08052 


ABSTRACT. The Lewis and Clark expedition produced a series of journals of more than one million 
words, and in the two centuries since the expedition in 1803 to 1806 a flood of books and articles has 
detailed their travels and travails. In this paper we explore a neglected portion of the expedition: the 
labels and notes associated with the Lewis and Clark Herbarium, a collection of 222 dried, pressed 
plants housed in the Academy of Natural Sciences of Philadelphia herbarium (PH). We also comment 
on an additional 10 specimens of Lewis that reside at the Royal Botanic Gardens at Kew. This residue 
of written records includes notes by Meriwether Lewis (primary plant collector on the expedition), 
Frederick Pursh (German botanist who first studied the plants), Thomas Meehan (Academy of Natural 
Sciences botanist who re-discovered the plants in Philadelphia after nearly a century of oblivion in 
storage), and several other nineteenth, twentieth, and twenty-first century botanists and other 
researchers who have found occasion to annotate the specimens. The notes reveal insights into the 
collection, curation, taxonomy, and research uses of the Lewis and Clark Herbarium during the 200 
years since Lewis made his gatherings. Much remains to be learned from a closer study of this little- 
examined part of the paper trail of Lewis and Clark. 


INTRODUCTION 
“all eaten!” 


With this annotation (Fig. 1), botanist Thomas Meehan in 1897 described some of the 
remaining, ostensibly missing, plant specimens gathered by Meriwether Lewis and William 
Clark along the Columbia River on 14 April 1806. In fact, his was probably a note of irony. 
Just years after Lewis and Clark returned from the West, horticulturist Frederick Pursh had 
copied or paraphrased Lewis’s original field label, now lost: “An umbelliferous plant of which 
the natives don’t eat the root.” Yet Meehan found just shreds, identifiable only as some kind 
of Lomatium, a wild carrot, which had been a feast to insects. In any other herbarium this 
would have been a disaster, and the specimens summarily discarded. But in the Lewis and 
Clark Herbarium they are as informative to a historian as are shards of broken pottery to 
an archaeologist. The two-century chatter of labels and annotations throughout the 
herbarium tells stories, too, a cyclorama of travel, botanical discovery, ecological record, and 
taxonomic opinion about the first-ever scientific exploration of the American Northwest. 
Not so many of Lewis and Clark’s plants were eaten, actually, and 222 herbarium sheets 
now in the Academy of Natural Sciences of Philadelphia offer a “feast” for the Lewis and 


*The order of authors was determined by coin toss. Manuscript submitted 7 November 2003, revised 
25 October 2004 


1 MISSOURI BOTANICAL 


MAY 0 2 2095 
GARDEN LIBRARY 


Z BARTONIA 


LEWIS AND CLARK elie 
PH-LC 131: Lomatium sp. 


James L. Reveal (MARY), ALFRED E. SCHUYLER (PH) Jun 19 


Figure 1. Tag (all eaten!) by Thomas Meehan (lower right) and plant fragments. The omy point 
normally indicates confirmation in botanical annotation, although in this case it may also signify 
dismay and surprise that the insects have had their way with the specimen. (Lomatium sp., PH-LC 
131.) 


Clark aficionado. There also are ten sheets at the Royal Botanic Gardens at Kew, which we 
mention herein as needed; but our focus will be on the main body of Lewis and Clark’s 
plants in the Academy of Natural Sciences of Philadelphia. 


THE LEWIS AND CLARK EXPEDITION 


The Lewis and Clark Expedition of 1804 to 1806 is, at least for the moment of its 
bicentennial, so well rediscovered by the American media that it is superfluous to introduce 
its historical significance at length. The expedition traveled through a largely unknown 
territory of the upper Missouri River, the Rocky Mountain divide, and the Columbia River 

to the Pacific Ocean—and back. The definitive documentaries are two: Gary Moulton’s 
(1986-2001) comprehensive 13-volume edition of the journals of Lewis, Clark, a1 and others of 
the expedition whose writings survive; and Donald Jackson’s (1978) edition of Letters of the 
Lewis and Clark Expedition with Relates Documents, 1 Lage which transcribes all of the 
to the expedition, its planning, 
and results. A raaeuctaal Lg See and literary account of the printed record of the 
expedition by Beckham et al. (2003) documents the post-expedition history in a handsomely 
designed and illustrated volume. 

In addition to the manuscript residue of the expedition, the explorers shipped back boxes, 
barrels, and cages full of natural history specimens and ethnological collections to the 
“civilized world” of what today is the Northeast Corridor of the United States. A few 
artifacts and relics were kept by the explorers or sent to their relatives; a few more were 
retained by their first recipient, Thomas Jefferson. He sent the larger prize en masse to the 
American Philosophical Society in Philadelphia, from whence they were promptly dispersed 


THE PAPER TRAIL OF LEWIS AND CLARK 3 


to scholars and learned men in many disciplines. Never again would the entire shipment 
reside in one place. 

Some specimens were displayed in Charles Willson Peale’s public museum in Philadelphia. 
Accident and loss contributed insidious wear on the collections as decades passed. Zoologists, 
botanists, and geologists studied, corresponded about, and published papers based upon 
various individual specimens and small subsets of the collections; once these were completed, 
the material was overlooked. The novelty of these natural curiosities waned as the expanding 
American frontier filled the blank and imaginative areas of the North American map. In 
time, more expansive and accessible collections were assembled from geographically broader 
and ecologically more diverse regions than those of the narrow routes of the explorers. 
Forgetfulness, too, cast a pall that obscured specimens from the pool of commonly known 
references (Cutright 1969; Spamer and McCourt 2002b); and the deaths of principal early 
workers sealed the fate of Lewis and Clark’s collections because later researchers went on 
to work with their own, fresher materials. 

Paul Cutright (1969), sixteen decades after the fact, succeeded in publishing the first 
scholarly treatise on the bounty of the expedition’s natural history collections, Lewis and 
Clark: Pioneering Naturalists. It remains the most concise and complete work on the subject. 
Such a book was intended to have appeared much earlier, edited by botanist and all-around 
naturalist and ethnologist Benjamin Smith Barton, who served as the “principal investigator” 
of the expedition’s biological, geological, and ethnological collections. The volume was to 
accompany Nicholas Biddle’s (1814) two-volume ghost-written paraphrase of the journals of 
Lewis and Clark. Barton died in 1815, having never worked on it, and apparently no one 
else had the wherewithal to bring it into being. 

The pressed plants from the expedition have always been the most well-studied portion 
of the scientific materials brought back from the West. Frederick Pursh’s (1813) inclusion 
of them in his pioneering Flora Americae Septentrionalis is the benchmark for all botanical 
studies of the expedition. His was the first such flora to span the North American continent, 
one that named dozens of new taxa based on the gatherings of Lewis and Clark. Remark- 
ably, after Pursh, few new taxa were erected based upon the expedition’s plants. The 
historical perspective of these plants has been covered well by Cutright (1969), and the 
significance of Pursh’s work by Joseph Ewan (1979) in his introduction to a facsimile 
reprinting of Pursh’s Flora; but more recently this suite of botanical specimens has been 
reinvestigated with modern eyes. Reveal et al.’s (1999) taxonomic overview of the Lewis and 
Clark vascular plants was the first since Thomas Meehan’s (1898) cursory summary, which 
in its day followed only the descriptions and notes scattered throughout Pursh’s Flora. The 
work by Reveal et al. also brought up to date much of the geographical data associated with 
the specimens. 

Moulton (1999) and Spamer and McCourt (2002a,b) presented historical perspectives about 
the plants of the Lewis and Clark expedition. Moulton illustrated all of the Lewis and Clark 
plants for the first time with photographs of entire herbarium sheets. Spamer and McCourt 
(2002a) showed every specimen, individually, at greater detail in digital format on CD-ROM; 
this is, in effect, an illustrated edition of Reveal et al.’s (1999) comprehensive taxonomy. The 
extensive references cited in these recent works open into the deep mine of additional 
resources, beginning with digital facsimiles of Meehan’s (1898) and Coues’ (1898) papers, 
presented in Spamer and McCourt’s CD. Spamer and McCourt’s (2002b) history is 
complemented by papers on the conservation of the Lewis and Clark Herbarium (McCourt 
et al. 2002) and modern biogeochemical investigations of some of the herbarium’s preserved 


4 BARTONIA 


specimens (Teece et al. 2002). 

With this so noted, we proceed on a new trail—the paper trail in the Lewis and Clark 
Herbarium in the Academy of Natural Sciences. Many sheets are cluttered with labels and 
annotations from the nineteenth, twentieth, and now the twenty-first centuries. Paramount 
among these are the annotations written on blotting papers by Meriwether Lewis in the 
field, of which now just 34 survive. Frederick Pursh copied, edited, and seemingly threw 
away many of Lewis’s originals—we know because in a few cases we have both Lewis’s and 
Pursh’s corresponding labels together—but Pursh left 225 labels now distributed to 210 
herbarium sheets, which proxy for those of Lewis’s that are now absent. And 170 
annotations in 1897 by Thomas Meehan, curator of botany in the Academy, document the 
recovery of the plants and their reuniting after a century. In a sense, herbarium sheets are 
akin to web pages for plant species and the labels are links to judgments and knowledge of 
earlier botanists. Some of those links are followed here; others remain mysteries to be solved 
by future researchers. 


MERIWETHER LEWIS, BOTANIZER 


The journals written by members of the Lewis and Clark expedition (Moulton 1986-2001) 
contain frequent references to plants, from casual observations to careful descriptions of 
morphology, ecology, and ethnobotanical uses. Interestingly enough, few references to the 
plants occur on the very days that label notations indicate were the gathering dates. This 
discrepancy offers some insight into the ways and means of the expedition as it traveled to 
the Pacific Ocean and back. Journal entries were made at various times, often on the day of 
observation or when the group stopped for a time (such as during the winter at Fort 
Clatsop). By contrast many of the plants were taken as matters of opportunity—when and 
where the men and Sacagawea, the only woman and Native American in their group, found 
them. For most of the gatherings, a comparison of the day’s events and activities, as 
registered in the journals, to the day’s collection of plants seem to have no consistent 
correlation. Meriwether Lewis gathered in good weather and bad, on days of ease and labor 
alike. He very well may have opted to gather better or more mature, or flowering, specimens 
for those plants that were commonly seen, regardless of when he wrote about them in his 
journals. He may even have elected to discard previously collected plants in favor of better 
ones found on later days; but this is speculation. 

Meriwether Lewis was the naturalist of the trip, by virtue of his quick, intensive courses 
of study received in May and June 1803, tutored by the most notable men in their fields in 
Philadelphia and Lancaster, Pennsylvania. He was not a formal botanist but he was an astute 
observer, capable of carefully following his instructions and comparing and describing his 
finds against the published references that he brought with him on the expedition. What he 
did not already know of plants as a farmer and from his herbalist mother, Lewis learned 
from Benjamin Smith Barton in Philadelphia. As far as we know, he never made pressed 
plant collections prior to the expedition nor did he do so in the few remaining years of his 
life after its return. Lewis may well have eerreeed some pressing mnethiods as the 
expedition crossed to the Pacific. Surely, the various d him with 
numerous problems of pressing and drying. Unfortunately, the journals and correspondences 
provide not a clue to how all this was done. All we ~_ are 7a pressed plants and Lewis’ s 
annotations. They indicate mostly “where” and “when,” 
note. And that is precisely what his job was—to eae record, and communicate back to 


THE PAPER TRAIL OF LEWIS AND CLARK 5 


Thomas Jefferson. In turn, Jefferson dispatched specimens and scribblings to the American 
Philosophical Society (he was its president at the time), the premier American “think tank.” 
The “what” would be added by his Philadelphia brain trust. 

Lewis did not maintain a separate register of gatherings by place and date, so his field 
annotations are collectively one of the most important pieces of documentation for the 
gathered specimens. The fact that he dated most of these annotations are what save the 
detailed precision of the locality data we have. Because historians have documented precisely 
where the explorers were on given dates, we are assured that almost all of the plants in the 
Lewis and Clark Herbarium have precise geographical data. Even though just 34 of Lewis’s 
original annotations are present now in the herbarium, Frederick Pursh’s 225 transcriptions 
document most of the rest. Fortunately, they include the dates that establish the geographical 
provenance for each. 

Labels are the modus operandi of museum collections, although they are ancillary to the 
specimens. As a rule, original labels are never discarded; they corroborate original 
provenance and taxonomy. Later labels record changing opinions about these data by 
scientists and other scholars after the fact. Many labels are tags of paper pasted to the 
herbarium sheet; others are in the form of annotations written directly on the sheet. In the 
Lewis and Clark Herbarium, there are six general categories of labels and annotations: 
(1) Meriwether Lewis’s annotations, which are about gathering and ethnobotanical uses; 
(2) Frederick Pursh’s transcriptions of Lewis annotations, somewhat edited, with taxonomic 
information added; (3) Aylmer Lambert’s provenance annotations, written after 1812 when 
a set of herbarium sheets was prepared in London for the specimens separated by Pursh from 
the original collection in Philadelphia; (4) Thomas Meehan’s tags and a few by J. M. 
Greenman of the Gray Herbarium at Harvard, providing revised taxonomic data upon the 
restudy of the collection as newly recovered in 1897; (5) American Philosophical Society 
ownership labels, affixed to the set of herbarium sheets that were probably prepared in 1921 
at the Academy of Natural Sciences, 115 years after the expedition returned from the field; 
and (6) later labels and annotations of the twentieth and twenty-first centuries, contributed 
by systematists, historians, curators, and collection management personnel. The clamor these 
six types of labels represent testifies that this is an actively used collection, hardly a quietly 
sequestered icon of historical reflection. The kinds and numbers of historical labels are 
summarized in Table 1. Together, all of these annotations and tags provide a chronology and 
critical history of the thoughts of many professionals as to the identity, source, and scholarly 
uses of these plants. 


WHAT BECAME OF LEWIS AND CLARK’S PLANTS? 


Today, 232 herbarium sheets hold plants known to survive from the Lewis and Clark 
Expedition. Ten sheets are at the Royal Botanic Gardens at Kew (K), near London; 222 
sheets are in the Academy of Natural Sciences of Philadelphia (PH), where the collection 
comprises the Lewis and Clark Herbarium (PH-LC). Several other sheets attributed to Lewis 
and Clark at PH and K are now considered to be plants grown from seeds brought back by 
Lewis (and thus from field-collected plants) or gatherings by Thomas Nuttall that were 
mistakenly attributed to Lewis and Clark (Reveal et al. 1999). Thorough histories of Lewis 
and Clark’s plants and Frederick Pursh’s connection with them have been presented by 
Cutright (1969), Ewan (1979), Moulton (1999), and Spamer and McCourt (2002b). An 
analytical study of the seed-grown materials has yet to be written. 


6 BARTONIA 


Table 1. Summary of historical annotations, tags, and labels in the Lewis and Clark Herbarium at the 
Academy of Natural Sciences of Philadelphia and attributed to Lewis and Clark at the Royal Botanic 
Gardens at Kew. 


Academy of Natural Sciences of Philadelphia 
Total sheets: 222 


A.P.S. deposit 

Lambert Herbarium 43 (6 circumstantial?) 
Lewis annotations cut from blotting papers 34 on 33 sheets 
Pursh annotation tags 225 on 210 sheets 
Lambert annotations on reverse of sheet 15 

Lambert annotations moved to obverse of sheet 29 on 26 sheets 
Meehan annotation tags 144 on 139 sheets 
Meehan annotations on Lambert sheets 26 

Greenman annotation tags 62 on 58 sheets 


A.P.S. ownership labels 


Royal Botanic Gardens at Kew 
Total sheets: 9-10* 


Sheets with Lambert Herbarium designation 11 
Sheets with Pursh annotation tags 10 
*Two sheets (L illus Pursh and El ernh. ex Rydb.) are likely Thomas Nuttall collections 


deg os Reveal, personal c communication, 2003), although pote research may reveal otherwise. G. E. Moulton 
Geivonal communication, 2003) concedes that L. pusillus is a Nuttall gathering. Thus, it seems most accurate to say 
that “there are nine or ten Lewis specimens at Kew from the Lewis ee Clark expedition of the eleven that have 
traditionally been thought to come from that endeavor” (G. E. Moulton, personal communication, 2003). 
“Traditionally” refers to the listing of Lewis specimens at Kew eR Ewan (1979) in his introduction to the 
facsimile reprinting of Pursh’s (1813) Flora Americae Septentrionalis. 


With regard to the paper trail discussed here, it is important to recall that a quarter of the 
collection had been taken by Pursh to London, where in 1812 or after they were prepared 
onto herbarium sheets. They went into the herbarium of Aylmer Bourke Lambert, a well-to- 
do botanist and Fellow of the Linnaean Society. In 1842, following Lambert’s death, a small 
portion of his herbarium was bought at estate auction by an American lichenologist, Edward 
Tuckerman. Contained therein were 47 sheets from Lewis and Clark. The ten sheets now 
at Kew were in another auction lot that was acquired by the British Museum. In 1856, 
Tuckerman gave his share of Lambert’s herbarium to the Academy as a gesture of 
professional reciprocity for the use of the Academy’s collection of cryptogams. The 
remaining two-thirds of the collection had remained in Philadelphia, transferred back to the 
American Philosophical Society (A-P.S.) after the 1815 death of Benjamin Smith Barton. 
Thomas Meehan recovered them in 1897 and published the first comprehensive taxonomy 
of the whole Lewis and Clark herbarium (Meehan 1898). The specimens were transferred on 
deposit to the Academy in 1897, but herbarium sheets for them were not prepared until 
1921, at which time it seems that the surviving original annotations by Meriwether Lewis 
were salvaged from the blotting papers and affixed to the herbarium sheets. The 179 sheets 
from A.P.S. remain the property of that institution. Note that 47 Lambert plants, plus 179 
from A.P.S., total 226, four more sheets than are now accepted as Lewis material in PH. 
This diserinaiay-4 is accounted for by four sheets previously attributed to Lewis and Clark 
that are probably gatherings by Thomas Nuttall in 1811 (Reveal et al. 1999). The four 


THE PAPER TRAIL OF LEWIS AND CLARK 7 


Figure 2. Labels by Meriwether Lewis (dark hued) and Frederick Pursh (light-colored). (Ericameria 
nauseosa (Pall. ex Pursh) G. L. Nesom & Baird var. graveolens (Nutt.) Reveal & Schuyler, PH-LC 51.) 


are retained in the Lewis and Clark Herbarium at the suggestion of Reveal et al., in the 
event that new research corroborates the original claim. In 2000, the Academy received a 
Save America’s i ceagures grant to conserve the Lewis and Clark Herbarium. Specially 

tructed containers and cabinets were made for the sheets, which now are stored in a new 
facility with optimum deed controls (McCourt et al. 2002). 


THE PAPER TRAIL: 1804-2004 AND ON 


To botanists, the specimens of the Lewis and Clark Herbarium are less important for their 
famous collectors than they are for their taxonomic role; 76 of them are type specimens, 71 
of them named by Pursh in his Flova in 1813 (Reveal et al. 1999). In the context of this 
paper, the plants play secondary roles. It is the labels and annotations that reveal data about 
the travels, not of Lewis and Clark, but of the plants themselves during and since the 
expedition, a time span of 200 years. 


Meriwether Lewis’s Annotations (1804-1806) 


Thirty-six annotations written by Meriwether Lewis in the field are known to survive, 
35 of which accompanied plants (Moulton 1999; Spamer and McCourt 2002a) and the other, 
a fossil fish jaw (Spamer et al. 2000). Properly speaking, the herbarium annotations are not 
labels although they have that appearance now (Fig. 2). All but five had been cut out from 
a larger blotting paper, which presumably had been folded and in which the finished pressed 
plant was stored for the journey back from the West. The five other annotations were 
written on more conventional writing paper; why, we do not know. Of the 35 plant 
annotations by Lewis, 34 are in the Lewis and Clark Herbarium, and one, not associated 
with any specimens now known from the expedition and obtained from a private collector, 
is owned now by Gary Moulton (Moulton 1987) 

The blotting paper is generally similar to one kind used in two bound volumes of plant 
specimens assembled by Benjamin Smith Barton in 1795, an exsiccata he called Herbarium 
Americanum. Today these volumes are in the care of the Academy’s herbarium, having come 
from the American Philosophical Society in 1897 (Cutright 1969; Spamer and McCourt 


g BARTONIA 


2002b). One herbarium sheet (Nicotiana quadrivalvis, PH-LC 146) has two Lewis annotations 
(one each on blotting paper and writing paper), thus there are 33 discrete sheets that hold 
Lewis’s field notes. We have not done a close comparison of these labels with the paper used 
in the original journals, or for that matter with paper used in correspondences sent back East 
from the field. By the circumstantial evidence of Frederick Pursh’s labels as well as logical 
conjecture, many more Lewis annotations once existed. 


Dates and Selective Survival of Lewis’s Annotations 


Dates for Lewis’s surviving annotations range from 10 August 1804 to 14 September 1806, 
which spans nearly the entire duration of the expedition from St. Louis to the Pacific Ocean 
and back. The distribution of dates is far from uniform, however, with most of the 
annotations bearing dates in the autumn of 1804, with a few from 1805 and 1806. Because 
we do not know why all the other Lewis labels were destroyed or lost, the chronological 
clustering of surviving labels in the autumn of 1804 has no explanation other than the 
obvious one, that these were part of the first shipment sent back East from Fort Mandan. 
That shipment was forwarded by Thomas Jefferson to the American Philosophical Society, 
which redistributed its contents for study. Benjamin Smith Barton received the plants, and 
it is he who presented them to Frederick Pursh in 1807 when Pursh was hired to write up 
the taxonomy of the collection. Yet why the Lewis annotations survived preferentially with 
the Fort Mandan shipment, and not the remainder of the collections received by the A.P.S. 
in 1806, has not been explained. This is especially bewildering given that Pursh studied the 
entire collection in Philadelphia and it seems that most if not all of his own annotations, 
borrowing from Lewis’s annotations, were prepared before he left Philadelphia (as we discuss 
later). 

Those Lewis original annotations that did survive apparently benefited from the several 
decades of oversight and disuse in storage, after which they were cut out and mounted on 
the herbarium sheets along with the pressed plants. This may have been at the direction of 
Thomas Meehan, when plans were made for the mounting of loose specimens received from 
A.P.S. (thus between 1897 and Meehan’s death in 1901), or in 1921, when Academy curator 
Francis W. Pennell supervised John M. Fogg, Jr. in the preparation of the Lewis and Clark 
herbarium sheets (Fogg 1982; Spamer and McCourt 2002b). In the Academy’s herbarium, the 
intervening time between Meehan’s death and Pennell’s arrival at the Academy in 1920 
seems to have been one of great activity in the curation of collections; for unexplained 
reasons, however, this work did not address the relatively few Lewis and Clark plants 
(Spamer and McCourt 2002b). 

One Lewis label of blotting paper, no longer associated with plant specimens, is in the 
private collection of Gary Moulton (Moulton 1987); it may have been associated with one 
of the 30 plant specimens somehow lost in Philadelphia between 1805 and 1807 (Moulton 
1999: 9, n. 7), thus also another survivor from the Fort Mandan shipment. 

Yet another Lewis label, this one on writing paper, is associated with a frag y fossil 
fish jaw, the type specimen of Saurocephalus lanciformis Harlan 1824, which had been 
collected by expedition sergeant Patrick Gass. It, too, was a part of the Fort Mandan 
shipment. The specimen is in the Vertebrate Paleontology collection of the Academy 
(Spamer et al. 1995: 91-92; Spamer et al. 2000: 51, Fig. 4). The label, which had been affixed 
to the specimen by a small spot of glue on one end, became detached ca. 1990 and was 
misplaced, as reported by Spamer and McCourt (2002b: 34-35, m. 122). 


THE PAPER TRAIL OF LEWIS AND CLARK 9 
The Mystery of the Blotting Papers and Lewis’s Plant Press 


Of the 34 Lewis annotations that are affixed to herbarium sheets, 30 are written on the 
same red-purple blotting papers between which the finished plant pressings seem to have 
been kept for their journey back East. Four other labels are written on light-colored writing 
paper. When Thomas Meehan rediscovered many of the plants at A.P.S. in 1897, they were 
still wrapped “in the original packages as presented many years ago” (Meehan 1898: 14). We 
have inferred (Spamer and McCourt 2002b) that these bundles were composed entirely of 
blotting papers, very likely folded in half, in which pressed plants were stored. Although it 
is common knowledge that Lewis wrote his annotations directly on these blotting papers, 
there are unanswered questions about where he got them and how he used them in the field. 
No written records survive from anyone affiliated with the expedition, nor from those who 
studied the specimens later, that could inform us specifically about the materials and methods 
used to press these plants in the field. Using the only evidence we have of this portion of 
Lewis and Clark’s paper trail—the trimmed fragments of blotting papers written on by Lewis 
himself—we here address these questions to see how the collection’s artifacts might provide 
some clues. 

As meticulous as Lewis was in itemizing the supplies he purchased in Philadelphia and 
elsewhere before the expedition embarked in 1803, the crucial blotting papers seem to have 
been lumped under some larger, all-encompassing category. If Lewis followed his mentor’s 
example, he may even have bought the blotting papers from Barton’s supplier. 

Lewis’s annotations, now serving the purpose of labels pasted to the herbarium sheets, are 
wide enough for us to infer that they were written across a folded piece of blotting paper 
sized to fit a plant press. We have it on Meehan’s (1898: 15) authority that, in their original 
state in A.P.S. in 1897, each of Lewis’s annotations was “written wholly across the sheet 
containing the specimens.” The fact that he noted, “containing,” lends credence to the idea 
that the loose plant specimens were held within something, like a fold, rather than 
interleaved between loose, open sheets. This offers a clue toward how the sheets were used 
in the field, and what constraints Lewis had in pressing his specimens and protecting them 
for transport back East. 

Keep in mind that storage space would have been at a premium during the expedition’s 
laborious travels over thousands of miles in boats and canoes and on foot and horseback; any 
economy of size would have been beneficial, though not likely a deciding factor between one 
kind of blotting paper or another. Given the incredible amount of gear that the Lewis and 
Clark expedition carried even during the laborious overland passages, and given that the 
mission of exploration included collecting large numbers of bulky and unwieldy specimens 
and artifacts of all kinds (not to mention living animals!), we rather doubt that “ultralight” 
contraptions would have been designed and pressed into service solely for the botanical 
collections. 

Close examinations of specimens throughout the herbarium (Spamer and McCourt 2002a) 
show clear evidence, not surprisingly, that some of them had been folded before being placed 
in the press. Barton’s own Herbarium Americanum, the example of pressings that Lewis 
almost surely had seen when we was tutored by Barton, has reasonably small specimens on 
page sizes that are smaller (13 x 7% inches) than the more standard American herbarium 
sheets we are used to today (16% x 11% inches). Lewis, if he had been restrained to this 
model, would have had to fold many of his specimens, which he did; and his blotting papers 
would have been proportioned accordingly, which they were, as we show here. 


10 BARTONIA 


We do not know the original manufacturing sizes of blotting papers used by Barton and 
Lewis. Blotting papers were probably manufactured at the same sizes, using the same kinds 
of frames, as were used for regular printing papers. This is important if we are to understand 
the economy of paper sizes and how they are cut down for use. Unlike some printing 
papers, though, blotting papers were probably cut to size by the manufacturer rather than 
the buyer; the papers would have been sold in bundles for specific purposes like letter- 
writing. But the needs of specialists like botanists may have impelled them to procure sheets 
that were larger than those usually stocked for use by writers. Here we hypothesize based 
on the traditional methods of manufacture and use of printing papers. 

One likely unit of measurement for production-sized paper is the conventional unit of 
measurement, “folio,” a size from which many printed works are produced. Regardless of 
the size of the finished work, most begin with a very large sheet. These single, large, 
unfolded papers (the so-called double-elephant folio sheet) are about 38 x 28 inches. Folded 
once, these produce the traditional “folio” pages of large-format books, about 28 x 19 inches; 
folded twice produces “quarto”-format pages, which for a book are gathered into signatures 
and their outside edges trimmed off; and so forth to ever greater numbers of pages printed 
on a single sheet for ever smaller formats. 

If we assume that the blotting sheets that Barton used had been cut from a large sheet into 
four manageable pieces, doubling the dimensions of the blotting sheets in Herbarium 
Americanum yields an original sheet size of about 26 x 15% inches, or reasonably similar 
to that of a folio sheet. Paper sizes were not standardized; variations were the result of 
materials and methods unique to a manufacturer at a given time. Yet for the sake of 
economy in manufacturing, large pages were all similarly large. Thus, smaller, folded pages 
would all be proportionately small, closely in agreement with the figures measured and 
estimated here. 

Because we do not know from records the original sizes of blotting paper used either by 
Barton in Philadelphia or by Lewis in the field, we shall make some observations in the 
manner of a working hypothesis. We note that a folio-sized sheet as seems to have been used 
by Barton (about 26 x 15% inches), folded in half, is about 16 x 13 inches. Thus, this 16 
x 13-inch sheet could be cut in half to produce two sheets the size of those in Herbarium 
Americanum—or it could be folded in half to serve as a holder roughly 13 x 8 inches in size 
for plants pressed in the field (and upon which Lewis could write his annotations across the 
sheet). This helps to substantiate the supposition that Lewis had seen this unique exsiccata 
when he was tutored by Barton in 1803. If the folded sheet was the size of Lewis’s blotting 
papers, his plant press was not much larger. A test of this hypothesis is still preserved in a 
sample of Lewis’s own, folded large specimens, too large to have otherwise fit into the press. 

Sheet PH-LC 3 is a specimen of the large-leafed maple, Acer macrophyllum. The leaf is 
folded inward from each side, presumably to have fit the dimensions of the plant press and 
storage container. PH-LC 3 was part of the A.P.S. component of plants that was mounted 
on modern herbarium sheets at the Academy in 1921, by which time it was too fragile to 
be unfolded. In this folded form, the leaf’s “footprint” on the sheet is about 12 x 8 inches, 
within the one-fold blotting-paper storage sheet of our hypothetical original in Lewis’s 
press—and the page size of Herbarium Americanum. 

In comparison, sheet PH-LC 4, also Acer macrophyllum, is a specimen that was mounted 
fully extended. It occupies the width of the herbarium sheet, although the specimen’s tips 
were trimmed off evenly with the sheet edges, perhaps at a later time. This specimen, like 
its counterpart in PH-LC 3, had been folded; the creases are visible. It was one of those 


THE PAPER TRAIL OF LEWIS AND CLARK 11 


taken to England by Pursh, the herbarium sheets for which were prepared soon after 1812. 
Just a few years after its gathering, the specimen was apparently still pliable enough to 
unfold it for mounting. Still, its original folds indicate that spatial economy was important 
when it was pressed. 

Another example is shown by a single large leaf of Veratrum californicum (PH-LC 195), 
which measures 11% x 5% inches. Not folded, this specimen clearly fit in Lewis’s press. 

A fourth example is expressed with the specimens of wild rice, Zizania palustris, on sheet 
PH-LC 226. Folded, each specimen occupies a “footprint” of about 10 x 6 inches, much 
smaller than the size of modern herbarium sheets but a comfortable fit within Barton’s 
Herbarium Americanum model. 

Ironically, historians do not know just how the finished, pressed plants were dried and 
carried by the expedition. Drying must have taken place in camp, when conditions favored 
it, probably by a fire. Oiled cloth was purchased in bulk when Lewis was obtaining 
expedition supplies in Philadelphia, which would have been used to create water-resistant 
wraps for all kinds of materials collected along the way that needed to be protected from the 
elements. It is a remarkable testament to Lewis’s planning (and luck) that as much made it 
back East as did, and in fine condition. 


William Clark’s Copy of a Receipt (1806) 


The single example of William Clark’s hand in the Lewis and Clark Herbarium (Fig. 3) 
has nothing to do with plants. It is an unsigned copy of a receipt for wages paid by Lewis 
to Toussaint Charbonneau, the French fur-trapper who helped the members of the 
expedition communicate with the Native Americans they met. On the reverse of the receipt 
is a botanical annotation by Frederick Pursh, occupying a much smaller area than that 
covered by Clark’s writing. This is one of the mysteries of the Lewis and Clark Herbarium. 
Perhaps Clark had “lost” the receipt amongst Lewis’s pressed plants. When Pursh found the 
large piece of paper, blank on one side, he wrote his annotation on it but cast it aside later. 
(See more below about Pursh’s labels.) The receipt was recovered when the herbarium sheet 
(now PH-LC 66) was prepared in 1921, and for some reason this document was pasted 
down—historic, non-botanical side up—to accompany the specimen for which Pursh had 
originally written his annotation. Perhaps this was because there was also a second tag from 
Pursh with the specimen, which provides better information. 

Even though the Clark receipt is pasted to the herbarium sheet, the ink from Pursh’s 
annotation bled through the paper. Pursh’s writing can be read (in reverse) with close 
examination or more easily by placing the sheet on a light-table—or by reversing and 
enhancing a digital image of the herbarium sheet. The discarded annotation on the reverse 
of Clark’s receipt reads, “Specimen from White river Cleome A new species”. The second 
Pursh tag on the sheet updates the evaluation: “Cleome Serrulata var rosea Nova Species.” 
Pursh (1813) did indeed publish this name as the new species, Cleome serrulata. 

We can only suppose that the receipt was affixed to the herbarium sheet as a historical 
relic. Or, we can just as easily opine that the receipt was pasted down in this fashion by 
mistake. The botanical annotation might have been intended to be cut out and dutifully 
pasted to the sheet with the other annotation—sacrificing its out-of-scope (rion-botanical) 
“reverse” side! 


12 BARTONIA 


Figure 3. Copy of a receipt written by William Clark for Toussaint Charbonneau (top). On the reverse 
of this receipt, visible more faintly (below; image digitally reversed), is er annotation by Frederick 
Pursh associated with this specimen. (Cleome serrulata Pursh, PH-LC 6 


THE PAPER TRAIL OF LEWIS AND CLARK 13 


RIUM OF 


Figure 4. Tags by Frederick Pursh with plant stems passed through parallel slits. These are the only 
remaining examples in this collection of how Pursh attached tags to specimens. Thomas Meehan has 
annotated me upper right tag (“Pursh’s writing”). His annotation under the lower tag reads “Lewis coll. 
label.” Here Meehan may have erred in attributing the label to Lewis, or he might after all have 
recognized Pursh’s labels as transcriptions of some of Lewis’s collecting data. (Equisetum arvense L., 
PH-LC 80.) 


Frederick Pursh’s Labels (1807 and after) 


When Frederick Pursh was assigned the task of identifying the botanical gatherings from 
the Lewis and Clark expedition, the intention was that these results would be compiled for 
publication in a volume of the scientific results of the expedition. Of course, that never came 
to be, and as we now know Pursh left Philadelphia, taking a quarter of the collection with 
him to London. His tags, however, are found throughout the expedition’s gatherings, both 
in those left in Philadelphia and those he had taken to England. Of the latter group, his tags 
now are found in the Lambert Herbarium component of the Lewis and Clark Herbarium 
at the Academy and with the Lambert sheets that are at Kew. Thus it seems very likely that 
most if not all of the tags were written while he worked on the collection in Philadelphia. 

Pursh’s tags (Figs. 2, 4) seem to be transcriptions of the information that Lewis ha 
written on the blotting papers in which the loose, pressed plants were returned to the East. 
This is corroborated by 27 sheets that preserve both Lewis’s original annotations and Pursh’s 
labels (seven sheets bear Lewis labels without Pursh counterparts). Pursh was rather faithful 
to Lewis’s original text, as we see in these examples, although he did take some editorial 
liberties, such as correcting Lewis’s inconsistent spelling, or omitting “superfluous,” non- 
scientific observations like some ethnobotanical uses. Fortunately, he was not consistent in 
such omissions, and some important data survive. In once instance, Pursh even corrected the 
year of the date of collection, which in context was obvious to him. 


14 BARTONIA 


Lewis’s field notations are written across a broader area than that used by Pursh in his 
tags; that is most likely because Lewis had a folded sheet of blotting paper upon which to 
write whereas Pursh was creating tags to be physically affixed to the specimens—tags which, 
incidentally, would have more legible penmanship than that of Lewis’s. In other words, 
Pursh was curating the collection. His tags would be attached to a stem, usually by cutting 
two parallel slits in the paper through which the stem would be interleaved to make it more 
difficult to disassociate the tag from the specimen. The tags were removed prior to mounting 
in most cases, although two specimens still bear the tag affixed to the stem (Fig. 4). So 
affixing labels is good curatorial practice for storage, and the procedure can also be seen as 
preparing them to travel without losing their data. 

Of course, the principal reason for having created these tags was to apply taxonomic 
identifications to the loose specimens, which Lewis, not being formally trained in botany, 
could not have made. This fact by itself led Thomas Meehan to think at first that these were 
the original field labels. On 17 August 1897, he wrote in a letter to Gray Herbarium’s 
Benjamin L. Robinson (Gray Herbarium Archives), “The writer in many cases knew 
something of botany, and the letters were German. I thought they must have had a German 
botanist with them. Before I got through I was sure the labels were by Pursh! But where did 
he get the notes!” Meehan meticulously identified every existing label he found (“Lewis” or 
“Pursh” written in pencil). Somehow, he apparently never did come to the conclusion—so 
far as we can tell—that each of Pursh’s notes was actually a transcription of Lewis’s 
annotation, with the logical inference that Pursh had probably discarded most of Lewis’s 
originals (but see also Fig. 4). 

Nearly every sheet in the Lewis and Clark Herbarium contains a Frederick Pursh tag; just 
12 sheets do not. Thirteen sheets contain two Pursh tags, and one sheet has three, yielding 
a total of 225 Pursh labels in the collection. (Of the ten sheets at Kew, all contain tags 
written by Pursh.) If nothing else, this is adequate testimony that Pursh did carefully study 
the whole collection. 


Aylmer Lambert’s Annotations (after 1812) 


Provenance data for the plants mounted on herbarium sheets in Aylmer Lambert’s 
collections were written on the reverse side of the sheets (Fig. 5), a custom then used by the 
British Museum. Lambert indicated on most sheets, “Herb. Lewis & Clark N. America Fred. 
Pursh,” or some similar indication. Many of these verso annotations were later cut off—when 
and by whom are not known—and pasted onto the fronts of their respective sheets. In some 
instances, cutting them out would have interfered with another label or the specimen on the 
front of the sheet, so only part of the annotation was cut out and transferred to the front. 


Figure 5. Portion of herbarium sheet bearing Aylmer Lambert’s handwriting. Lambert annotated Lewis 
and Clark specimen sheets on the verso of the top edge. Later, someone cut out some of these 
annotations and pasted them to the front of the respective sheets. (Ericameria nauseosa (Pall. ex Pursh] 
G. L. Nesom & Baird var. graveolens [Nutt.] Reveal & Schuyler, PH-LC 52.) 


THE PAPER TRAIL OF LEWIS AND CLARK 15 


The purpose was obvious—to place the provenance data on the front of the sheet with the 
rest of the annotations, although in the process this aesthetically damaged the sheets, 
sometimes very unsightfully so. Spamer and McCourt (2002a) for the first time illustrated 
all of the verso annotations. 


Academy of Natural Sciences of Philadelphia Ownership Labels 
(Mid-nineteenth Century) 


As might be expected in any large museum collection, many kinds of official labels will 
reflect design changes and other peculiar needs over time. Among the Lewis and Clark 
herbarium sheets are those that had been a part of the Lambert Herbarium, which were 
given to the Academy by Edward Tuckerman in 1856. All of these have Academy herbarium 
labels of a design from the mid-nineteenth century (Fig. 6). They are not especially 
remarkable and the handwriting on them has not been attributed, so far as we know. 


HERBARIUM OF 


ACADEMY OP NATURAL SCIRNORS, PHILADBLPHAA, 


Figure 6. Label of “Herbarium of Academy of Natural Sciences, Philadelphia.” These labels were 
attached to the Lambert specimens that had been bought by Edward Tuckerman. The “ANS PHILA” 
punch, seen here and in Figs. 3a and 7, is a twentieth century collection identification stamp applied 
to all of the Academy’s herbarium sheets. (Pediomelum argophyllum [Pursh] J. W. Grimes, PH-LC 157.) 


Thomas Meehan’s Labels (1897) 


When Thomas Meehan went to see if the American Philosophical Society retained any 
of Frederick Pursh’s study material, first in 1896, he was looking for Lewis and Clark’s 
plants as well as the lode of Pursh’s reference specimens. Once he found Lewis and Clark’s 
gatherings among the bundles he saw the opportunity to study the whole collection for the 
first time in nearly 90 years. They had last been used by Pursh while he was in Philadelphia 
and the information from them was included in his 1813 Flora. Meehan, in correspondence 
with B. L. Robinson of the Gray Herbarium, mentioned that even as an A.P.S. member he 
had to be assertive in order to get permission to hunt for Pursh’ s flora (see Spamer and 
McCourt 2002b). After Meehan made his find, some t and procedural 
entanglements ensued between the administrations of the: Academy and A.P. S. Meehan 
received permission to send the specimens whose identities were not clear to him to Harvard 
for proper identification; but he decided, seemingly in a fit both of professional despair and 
bureaucratic defiance, that he was not sure of any of them and sent the whole lot to 
Massachusetts, where Robinson and Greenman passed their pronouncements on some of the 
materials, including the historical, insect-eaten fragments. 


16 BARTONIA 


Figure 7. Tag by Thomas Meehan (right, initialed “T.M”), confirming species identification. Also visible 
in this photograph are a Meriwether Lewis label (lower, dark), Pursh transcription (upper left, light- 
colored), and perforated initials of the Academy of Natural Sciences Herbarium. (Rosa arkansana 
Porter, PH-LC 195.) 


The entire collection was studied and Meehan finished an updated taxonomy for them 
between August and November 1897—rather short order for such a large and important 
collection (see Spamer and McCourt 2002b). Meehan’s annotations occur as hastily written 
notes in pencil on small slips of common paper (Fig. 7); there are 144 of them pasted now 
to 139 herbarium sheets. These are his own identifications as well as those of the Harvard 
botanists; almost every one of these tags is signed distinctively, “T.M” (with one period). 
Possibly, Meehan wrote on larger sheets that were laid in with the appropriate specimens 
and later his annotations were cut out when the herbarium sheets were prepared. This helps 
corroborate the story that the A.P.S. sheets were prepared in 1921 because Meehan also 
annotated 26 of the herbarium sheets that had come from the Lambert Herbarium by 
writing directly on the herbarium sheet. Had sheets already existed for the A.P.S. specimens, 
surely he would have annotated them directly. 

In addition to his tags noting “all eaten” or “all gone,” Meehan also wrote notes next to 
Lewis’s annotations on the blotting sheets, most often simply corroborating the identity of 
the writer, “Lewis.” Whoever it was that drew outlines in pen around Lewis’s handwriting 
on the larger blotting papers also was careful to draw around Meehan’s adjacent handwriting 
(Spamer and McCourt 2002a,b; see Fig. 7, where portions of the outline can be seen along 
the cut edges). Meehan, as part of curatorial projects to mount loose specimens, may have 
outlined everything himself some time after having annotated the labels in pencil but we 
believe it is more likely that the Lewis-Meehan annotations were delineated by Academy 
curator Francis W. Pennell, who supervised John M. Fogg, Jr. in the mounting of A.P.S.’s 


THE PAPER TRAIL OF LEWIS AND CLARK 17 


Lewis and Clark plants in 1921. Elsewhere we have discussed in more detail the points 
regarding curation of the collection (Spamer and McCourt 2002b). 


J. M. Greenman’s Labels (1897) 


When Thomas Meehan in his bureaucratic snit sent the entire A.P.S. component of the 
Lewis and Clark herbarium off to Harvard University in 1897, he was soliciting help in 
confirming or identifying the taxa represented by what sometimes were mere twigs and leaf 
fragments. J. M. Greenman replied with a number of identifications—62 of his tags are 
preserved on 58 of the sheets (Fig. 8). All of them accompany specimens that had been found 
at A.P.S., another indication that Meehan had not sent away any of the Lambert Herbarium 
sheets then known to him. In fact, although he mentions the Lambert specimens (expressing 
ignorance of how they arrived at PH), he does not include them in his table comparing the 
Pursh material with published comments from Pursh’s Flora (Meehan 1898). This suggests 
that Meehan found the Lambert specimens in PH after sending the Pursh find from A.P.S. 
to the Harvard botanists. 


(4 - al fn 


Figure 8. Tag by Jesse M. Greenman with his species identification. (Artemisia longifolia Nutt., PH-LC 
27.) 


American Philosophical Society Ownership Labels (1897?) 


All of the herbarium sheets that were prepared from the specimens found in 1897 at 
A.P.S. hold printed labels with the banner, “American Philosophical Society. Lewis & Clark, 
Herbarium. From the Atlantic to the Pacific.” The ownership labels (Fig. 9) are very unlike 
other labels in the Academy’s collections, even for the thousands of other sheets that at one 
time had belonged to A.P.S., which have Academy labels with handwritten provenance data 
crediting A.P.S. So it is reasonable to assume that the “Atlantic to the Pacific” labels were 
printed at the behest of (or by) A.P.S., specifically as a condition of the deposit, document- 
ing the ownership of these very important plants. Each of these labels has been placed in the 
lower right corner of the sheets from the A.P.$. component of the Lewis and Clark 
Herbarium, a style so systematic that it clearly demonstrates that the labels were present 
when the sheets were prepared; they were not added to preexisting sheets. 

Although the A.P.S. ownership labels contain printed lines for “Locality,” “No.”, and 
“Date,” almost none of the 179 labels have these data filled out. Most labels contain only the 


18 BARTONIA 


AMERICAN PHILOSOPHICAL SOCIET 
LEWIS & CLARK, HERBARIUM. 


FROM THE ATLANTIC To THE PACIFIC. 


wat a lpsonnsanbetnrttecastenieemnattntitnaaann 
Pets Contes Dea a 


his 


Figure 9. Label identifying specimen belonging to American Philosophical Society, with species name 
written by Thomas Meehan. (Astragalus missouriensis Nutt., PH-LC 33 


taxonomic identifications, and on the few labels where the geographic data have been added 
the handwriting is different from that of the person who wrote the taxonomic identifica- 
tions. Until now, no one seems to have mentioned whose hand this is for the taxonomic 
data. We have compared the handwriting to Thomas Meehan’s and believe it is his 
handwriting. This effectively dates when the labels were printed to between 1897 and 1901. 
Notably, this was when a great number of unprocessed botanical collections were being 
curated in the Academy’s botany department (see Spamer and McCourt 2002b). However, 
many jobs were interrupted by Meehan’s death in 1901, including, it seems, the mounting 
of A.P.S.’s Lewis and Clark plants. We do not know who added the few annotations of 
geographic data or when this was done. 

As for the labels indicating “From the Atlantic to the Pacific,” this is a generalization of 
the historical origins of the expedition on the eastern seaboard, perhaps also to specifically 
acknowledge A.P.S.’s role in establishing the collection. The easternmost specimens in the 
herbarium were collected in Missouri. Inasmuch as the Delaware and Schuylkill Rivers in 
Philadelphia are tidewaters of the Atlantic Ocean, the notation of “Atlantic” may be 
defended as literary, if not geographical, license. 


Twentieth- and Twenty-first-century Labels and Annotations 


As with any working collection of botanical specimens, a herbarium sheet continues to 
accrue labels and annotations. They reflect taxonomic revision, changes of information 
associated with the specimens on the sheet, or comments about work performed with the 
specimens or the herbarium sheet itself. In every respect they indicate (in the parlance of the 
explorers’ journals) how science and history have “proceeded on” in the study and use of the 


THE PAPER TRAIL OF LEWIS AND CLARK 19 


Figure 10. Annotations of three late-twentieth-century researchers. (Lower) label of Academy of rons 
Sciences of Philadelphia intern Erica Armstrong; (middle) label of Reveal et al. (1999); ( 
annotation of Earle E. Spamer indicating photographic plate number in Moulton (1999). Zieudenus 
elegans Pursh, PH-LC 225.) 


collections. Numerous sheets contain the usual taxonomic chatter of botanists who confirm 
or contradict previous identifications or who make comments on typification or the 
reliability of putative geographic data that accompany the specimens. These all are typical 

inds of annotations found in herbaria everywhere. Every label and annotation on the Lewis 
and Clark Herbarium sheets has been transcribed, and their writers identified, by Moulton 
(1999: 13-55). 

Unfortunately, some annotations were made with archivally unsuitable materials, such as 
ballpoint pen, which over time will bear watching in a historically important collection such 
as this. One peculiar mark requires an explanation. On every sheet, there is a small green 
dot; they seem to have been drawn with a felt- or other soft-tipped pen. These were marked 
during a project around 1980 that photographed the Academy’s entire Type Collection and 
other special collections of flowering plants and ferns. The green dots were the photogra- 
pher’s key to assure that all herbarium sheets were photographed. These photographs were 
reproduced on microfiche, with a large, folio volume that served as an introduction and 
index (Mears 1984). The set of hundreds of 60-frame fiche of negative images was available 
commercially. The Academy’s Archives holds a set of these fiche; and interestingly, the 
image subset for the Lewis and Clark Herbarium is a “sample copy” of positive images. 

Unique to the Lewis and Clark Herbarium are labels and annotations that relate to 
specialized work done with these sheets, either by Academy curatorial staff or by 
conservationists and visiting researchers. Rather noticeable throughout the herbarium are 
laser-printed labels in a Courier font by Erica Armstrong (Fig. 10); they appear on 174 
sheets. Armstrong was an intern in the Academy’s Department of Botany who in 1993 and 
1994 worked on a project to update the taxonomy of the Lewis and Clark plants. Only 
those sheets that had not been recently studied by a taxonomic botanist were updated; thus 
her annotations do not appear on every sheet. 

n 1997, Mark A. Teece, then of the Carnegie Institution of Washington’s Geophysical 
Laboratory, inquired to the Botany Department about the utility and likelihood of the Lewis 


20 BARTONIA 


Oe re aed 


LL RIS LASS ht 


Figure 11. Label of Mark A. Teece indicating samples taken for biogeochemical analysis. (Zizania 
palustris L., PH-LC 226.) 


and Clark plants being available for biogeochemical analysis (Teece et al. 2002). His research 
relates to carbon-based fatty acid and stable-isotope compositions of leaf tissues. Specimens 
in older collections that are accompanied by well-documented date and locality information 
can be used toward interpreting regional atmospheric chemistry, which can be applied to 
studies on larger geographic scales. Since many old collections lack the precision of date and 
locality among their specimens, they are not suitable for these analyses. The Lewis and Clark 
Herbarium, however, is unusual in this regard. Historians have very well documented the 
daily locations of the Lewis and Clark expedition from 1803 to 1806. The fact that Lewis 
dated almost all of his botanical gatherings thus allows us to locate with geographic and 
temporal precision the provenance of each specimen, which is precisely the kind of 
resolution required in Teece’s work. Because the Lewis and Clark Herbarium predates the 
Industrial Revolution, which instigated dramatic changes in the chemistry of the world 
atmosphere, this herbarium has the potential to provide information on the atmospheric 
chemistry of the American Northwest from a time before the introduction of gases of 
factory effluents and internal combustion engines. 

Teece did obtain the permission necessary to sample some specimens in the Lewis and 
Clark Herbarium. Some of the results of the analyses were published by Teece et al. (2002). 
At the time when the samples were taken from the sheets, on 18 December 1997, labels 
documenting the act were affixed to each sheet so sampled (Fig. 11). Eleven sheets were 
sampled. The size of each sample was not large, approximately a centimeter square, and care 
was taken that the sampling did not take proportionally too much of the specimen nor that 
it aesthetically damaged the specimen. Normally, aesthetics is not a concern in sampling a 
herbarium sheet but given that this collection is a public treasure of immense historical 
importance, it is accorded this special criterion. The selection and sampling process was 
documented by Teece et al. (2002). 

In 1999, Reveal et al. published the first comprehensive taxonomic review of the Lewis 
and Clark Herbarium in a century. Their review assigned a “PH-LC” number, an acronym 
indicating the Lewis and Clark Herbarium in the Academy of Natural Sciences Herbarium, 
which is housed in the Types and Special Collections room of PH. As a result of that 
publication, every sheet bears the authors’ printed labels indicating the PH-LC number; most 
up-to-date name for each specimen sheet; author, publication, and date for the name; type 
status (lectotype, duplicate, etc.); and the names of the determiners, J. L. Reveal and A. E. 
Schuyler (see in Fig. 10). These labels were printed on 100% rag bond, acid-free paper using 
a laser printer. A Reveal and Schuyler label appears on every sheet in the Lewis and Clark 
Herbarium, including the seaweed, moss, and liverwort sheets not included in their review 
(Reveal et al. 1999) but included in Spamer and McCourt’s (2002a) CD-ROM (see below). 

In 1999, the Academy of Natural Sciences was awarded a grant from the Save America’s 
Treasures program administered by the Institute for Museum and Library Services and the 


THE PAPER TRAIL OF LEWIS AND CLARK pat 


Conservation work on this sheet done as part of 
Save America’s Treasures grant to the Academy 
of Natural Sciences. . 
Catharine Hawks, Conservator 9 Aug 2000 


Figure 12. Label of Catharine Hawks indicating conservation work performed under the auspices of 
the Save America’s Treasures program. (Atriplex nuttallii S. Watson, PH-LC 34.) 


U.S. Department of the Interior, National Park Service. Matching funds from private-sector 
sources were received from the Lattner Foundation and The Bay Foundation, Inc. The 
purpose of the grant was to re-house the entire Lewis and Clark Herbarium in new, 
archivally long-lasting housings, stored in new cabinetry in a climate-controlled room 
designed specifically for it. Sheets and specimens requiring special conservation measures 
were evaluated and the work performed by conservator Catharine A. Hawks. For the 31 
sheets so affected, dated labels were affixed to document the acts (Fig. 12). In addition, the 
special containers, modified from an original design first funded by the National Geographic 
Society in 1998, were constructed to hold each sheet within a separate stiff, closable holder. 
These holders also are used to display the specimens when needed, thus greatly reducing the 
amount of handling of the specimens while leaving them easily available for botanical and 
historical researchers. For a description and documentation of the conservation program, see 
McCourt et al. (2002). 

When one of us (Spamer) was Collection Manager in the Academy’s botany department, 
he made digital images of the entire Lewis and Clark Herbarium between 1999 and 2001. 
This was for the purpose of conservational documentation of the plant specimens. Rather 
than imaging whole sheets, each specimen on every sheet and sometimes specimen groups 
were digitized with an accompanying scale for size comparison. The images were recorded 
in grayscale format rather than color, and usually at a resolution of 300 dpi, to conserve the 
disk space necessary to store the more than 500 images (e.g., Fig. 13). Only a couple of 
specimens that retain relict coloration were imaged in color. This collection of digital images 
was later used to produce a CD-ROM study set for the Lewis and Clark Herbarium (Spamer 
and McCourt 2002a), which is in part an illustrated edition of the taxonomic revision of the 
herbarium made by Reveal et al. (1999). In the process of creating the digital images, Spamer 
also annotated the reverse of the herbarium sheets as a means to keep track of the images 
in an unobtrusive manner. These annotations (Fig. 14) have no significance to the history 
or taxonomy of the specimens; they were a matter of temporary record-keeping only. 
Spamer annotated the front of every sheet, as close as possible to the PH-LC label, to 
reference the sheet’s corresponding plate number as published by Moulton (1999; Fig. 10). 

Finally, in 2000, whole-sheet, high-density, color digital images of the entire Lewis and 
Clark Herbarium were made by intern Sarah Rice. The TIFF-format images, prepared under 
controlled lighting, presently occupy 27 CD-ROM disks (about 15 GB total). A copy of this 
set is in the Archives of the Academy of Natural Sciences. Migration to newer media will 
significantly reduce the physical storage space required, and eventually the entire image 
collection should be made available on a single disk or other storage medium. Copies on 
DVD were also stored in the Department of Botany. In 2003, McCourt prepared medium- 
resolution JPEG-format images of the full sheets, which can be used in more widely focused 


22 BARTONIA 


‘wt fst 


eee | | 


9 lod je. Vege 4 
eb ie eke sake 


Diimitcs bE ohe 


Figure 13. Black and white digital scan of specimen from CD-ROM on the Lewis and Clark 
Herbarium, with ruler for scale Spamer and McCourt 2002a). Visible are portions of tags by Meehan 
(left) and Pursh (below). (Fritillaria pudica (Pursh) Spreng., PH-LC 95.) 


THE PAPER TRAIL OF LEWIS AND CLARK 23 


Figure 14. Code number on reverse of herbarium 
sheet added by Spamer during digital imaging of the 
Lewis and Clark Herbarium (Spamer and McCourt 
2002a). (Berberis nervosa Pursh, PH-LC 39.) 


public venues such as web sites and future technologies when they become available. These 
digital images were made possible by the Academy’s Albert M. Greenfield Center for Digital 
Imaging of Collections and a grant from the Save America’s Treasures program of the 
National Park Service. 


Proceeding On Along the Paper Trail 


The history to be gleaned from the paper trail of Lewis and Clark is not complete. The 
labels offer data for testing historical hypotheses such as those proposed above. Each label 
and annotation attached at a particular date become traits of the specimen and contain 
unique information—some of which its author intended and some he or she did not. For 
example, the 34 Lewis labels are found only on specimens that remained in Philadelphia at 
the American Philosophical Society; none is found on the specimens Pursh took to London. 
This falsifies the hypothesis that surviving Lewis labels are remnants of a random process. 
But the finding begs further questions. Why did only 34 Lewis labels survive with the plants? 
Why this particular set of 34? And what of the 35th plant label, now owned by Gary 
Moulton, which may have been associated with the so-called “missing 30 plants” that were 
lost between 1805 and 1807? The labels are the desiderata of the plant collections in which 
to search for patterns that can lead to new insights into the botany of Lewis and Clark. 
Clearly, much remains to be learned from the paper trail of the Lewis and Clark Herbarium. 


ACKNOWLEDGMENTS 


We thank Drs. Gary E. Moulton (University of Nebraska), James L. Reveal (University 
of Maryland, Emeritus), and A. E. Schuyler (Academy of Natural Sciences of Philadelphia, 
Emeritus) for helpful comments and answers to queries during the writing of the manuscript. 


LITERATURE CITED 


BECKHAM, S. D., D. ERICKSON, J. SKINNER AND P. MERCHANT. 2003. The Literature of the Lewis and 
Clark Expedition: a Bibliography and Essays. Lewis and Clark College, Portland, Oregon. 315 pp. 

(BIDDLE, N.] 1814. History of the Expedition Under the Command of Captains Lewis and Clarke, to the 
Sources of the Missouri, Thence Across the Rocky Mountains and Down the River Columbia to the Pacific 
Ocean. Performed During the Years 1804-5-6. By Order of the Government of the United States. 
Bradford and Inskeep, Seyg te and Amb. H. Inskeep, New York. 2 vols. 

COUES, E. 1898. Notes on Mr. Thomas Meehan’s paper on the plants of Lewis and Clark’s expedition 
across the continent, 1804-1806. Probst of the Academy of Natural Sciences of Philadelphia 50: 
291-315. 


24 BARTONIA 


CUTRIGHT, P. 1969. Lewis and Clark: Pioneering Naturalists. University of Illinois Press, Urbana. 506 
pp. [Reprinted 1989, University of Nebraska Press, Lincoln and London.] 

EWAN, J. A. 1979. Introduction to the facsimile reprint of Frederick Pursh’s Flora America 
Septentrionalis (1814). Pp. 7-117 in F. T. Pursh, Flora Americae Septentrionalis (facsimile reprint, J. 
Cramer, Vaduz). 

FOGG, J. M., JR. 1982. Reminiscences of a Botanist. Harrowood Books, Newtown Square, Pennsylvania. 


P- 

JACKSON, D. (ed.). 1978. Letters of the Lewis and Clark Expedition with Related Documents, 1783-1854. 
2nd edition, with additional cs and notes. University of Illinois Press, Urbana. 2 vols. 
McCourt, R. M., C. HAWKS AND E. E. SPAMER. 2002. The Lewis and Clark Herbarium of the 
Academy of Natural Scena. se 2. Saving an American treasure: preservation of the herbarium 

on the bicentennial of the expedition. Notulae Naturae, no. 476. 16 pp. 

MEARS, J. A. 1984. Indices to the Microfiche of the Types and Special Collections (Flowering Plants and 
Ferns) of the Herbarium of the Academy of Natural Sciences of Philadelphia. Meckler Publishing Co., 
Westport and London. 274 

MEEHAN, T. 1898. The plants of Lewis and Clark’s expedition across the continent, 1804-1806. 
Proceedings of the Academy of eT Sciences of Philadelphia 50: 12-49. 

MOULTON, G. 1987. New documents of Meriwether Lewis. We Proceeded On 13(4 [November]): 4-8. 

MOULTON, G. (ed.). 1986-2001. The Journals of the Lewis and Clark Expedition. University of Nebraska 
Press, Lincoln and London. ols. 

MOULTON, G. (ed.). 1999. The bast of the Lewis and Clark Expedition, vol. 12: Herbarium of Lewis 
and Clark. University of Nebraska Press, Lincoln and London. 359 pp. 

PURSH, F. 1813. Flora Americae Septentrionalis. White, Cochrane and Co., London. 2 vols. [Title-page 
dates indicate 1814; documented as available ai og 1813. 

REVEAL, J. L., G. E. MOULTON AND A. E. SCHUYLER. 1999. The Lewis and Clark collections of 
vacned plants: Names, types, and comments. be nae of the Academy of Natural Sciences of 
Philadelphia 149: 1-64. 

SPAMER, E. E. AND R. M. MCCOuRT. 2002a. The Lewis and Clark Herbarium, Academy of Natural 
Sciences of Philadelphia (PH-LC): Digital aed study set. Academy of Natural Sciences of 
Philadelphia Special Publication 19. CD-RO 

SPAMER, E. E. AND R. M. MCCourT. 2002b. mi Lewis and Clark Herbarium of the Academy of 
Natural Sciences. Part 1. History. Notulae Naturae, no. 475. 46 pp. 

SPAMER, E. E., E. DAESCHLER AND L. G. VOSTREYS-SHAPIRO. 1995. A Study of Fossil Vertebrate Types 
in the Academy of Natural Sciences of Philadelphia: Taxonomic, Systematic, and Historical Perspectives. 
Academy of Natural Sciences of Philadelphia Special Publication 16. 434 pp. 

SPAMER, E. E., R. M. MCCOURT, R. MIDDLETON, E. GILMORE AND S. B. DURAN. 2000. A national 
treasure: accounting for the natural history specimens from the Lewis and Clark expedition (western 
North America, 1803-1806) in the Academy of Natural Sciences of Philadelphia. Proceedings of the 
Academy of Natural Sciences of Philadelphia 150: 47-58. 

TEECE, M. A., M. L. FOGEL, N. TUROSS, R. M. MCCourRT AND E. E. SPAMER. 2002. The Lewis and 
Clark Herbarium of the Academy of Natural Sciences. Part 3. Modern environmental applications 
of a historic nineteenth century botanical collection. Notulae Naturae, no. 477. 20 pp. 


Bartonia No. 62: 25-43, 2004 


Effect of Herbivore Exclosure Caging on the Invasive Plant 
Aliaria petiolata in Three Southeastern New York Forests 


T A. MORRISON AND LEONE BROWN! 
Department of Biology, The College of New Jersey, P.O. Box 7718, Ewing, New Jersey 08628 


‘Current address: Department of Ecology and Evolution, State University of New York at Stony Brook, 
650 Life Sciences Building, Stony Brook, New York 11794-5245 


ABSTRACT. We examined effects of herbivore exclosures on non-native Alliaria petiolata (Bieb.) 
Cavara and Grande, and on the native herb layer, to determine if selective herbivory by mammals 
(particularly white-tailed deer) may facilitate A. petiolata invasion. The study was done from 199% to 
2000, in one urban forest without deer (New York Botanical Garden Forest), and two suburban forests 
(Kitchawan Preserve and Mt. Holly Sanctuary), both in a region with > 50 deer km~. Each forest had 
four pairs of 4-m? plots, with one of each pair caged to exclude deer. No significant differences 
developed in percent cover of native plants between uncaged and caged plots. At Mt. Holly, A 
petiolata cover in caged plots averaged nearly twice its cover in uncaged plots, with a similar trend at 
Kitchawan but not at NYBG. Individual A. petiolata size in caged plots at Mt. Holly averaged more 
than three times that in uncaged plots. Herbivory on A. petiolata was 30 to 40 times more frequent 
in uncaged plots in both forests with deer, but only one plant showed herbivory at NYBG. We 
attribute A. petiolata cover and size differences between caged and uncaged plots to deer herbivory, 
noting that Mt. Holly appeared most heavily browsed. We suggest that interactions between deer and 
invasive species could change as densities of both increase, and that these interactions should be 
considered in forest management. 


INTRODUCTION 


Alliaria petiolata (Bieb.) Cavara and Grande (Brassicaceae), commonly known as garlic 
mustard, is one of the most important non-native herbaceous plants threatening native 
woodland plants in eastern deciduous forests of North America (McCarthy 1997; Nuzzo 
1993a; Schwartz and Heim 1996; Yost et al. 1991). In southeastern New York, where this 
study was done, A. petiolata is a biennial, with germination and growth of a basal rosette 
of leaves in the first spring and summer, persistent foliage and some new growth throughout 
the winter, followed by further growth, flowering, and fruiting the subsequent spring and 
summer. It grows under closed forest canopy and along forest edges, and exhibits wide 
ecological amplitude for light levels and soil moisture, which may be due to high phenotypic 
plasticity (Byers and Quinn 1998; J. A. Morrison, unpublished data). Alliaria petiolata has 
been a component of the North American flora at least since it was first recorded on Long 
Island in 1868, and it has spread exponentially since then. It is now found throughout 30 
northeastern states and southeastern Canada (Cavers et al. 1979; Nuzzo 1993b). It can spread 


Submitted 7 January 2002; revised 9 December 2002, 7 October 2004 


25 


26 BARTONIA 


rapidly once introduced to a site, with apparent displacement of native herbaceous species 
within ten years (Anderson et al. 1996, Nuzzo 1994). 

Researchers focus mostly on attributes of the plant itself to understand why A. petiolata 
is invasive. Its high reproductive capacity, autogamy, and competitive ability, for example, 
have been addressed by various studies (Anderson et al. 1996; Baskin and Baskin 1992; Byers 
and Quinn 1998; Cavers et al. 1979; Cruden et al. 1996; McCarthy 1997; McCarthy and 
Hanson 1998; Nuzzo 1991, 1993a; Nuzzo et al. 1991; Yost et al. 1991). Underlying 
ecological changes that may promote its spread have received little attention, however, even 
though there is growing recognition of the need for broader ecosystem understanding for 
management of plant invasions (Hobbs and Humphries 1995). 

One ecological factor that may be important in A. petiolata invasion, but has not been 
investigated experimentally in the field, is herbivory by mammals, particularly white-tailed 
deer (Odocoileus virginianus Zimmerman). White-tailed deer have 1 dd ically since 
the early twentieth century in the successional, fragmented forests of the northeastern 
United States (Anderson 1997; McShea et al. 1997; Porter and Underwood 1999). For 
example, Knox (1997) estimates pre-colonial density of fewer than 4 deer km” in the eastern 
United States, in contrast with densities of 6 to 12 deer km~ in much of Virginia by 1988. 
Alverson et al. (1988) suggest pre-colonial estimates of 4 deer km™ in northern Wisconsin, 
compared with up to 9 deer km” in 1988. Deer density can be much greater in some areas. 
In northern Westchester County, New York, where we conducted part of our study, density 
is estimated at more than 50 deer km™ (Glenn Cole, Regional Wildlife Manager, New York 
State Department of Environmental Conservation, Region 3, personal communication). 

White-tailed deer at their current densities have become a keystone species, significantly 
altering the composition of forest ecosystems and leading to alternate stable states in many 
forests of eastern North America (Alverson and Waller 1997; DeCalesta 1997; Healy 1997; 
Schmitz and Sinclair 1997; Stromayer and Warren 1997; Waller and Alverson 1997). Deer 
herbivory can affect plant growth, fitness, and competitive ability of food plants, especially 
where deer densities are high due to a lack of natural predators, where active management 
increases deer populations, or where hunting is limited because of proximity to towns 
(McShea et al. 1997; Philips and Maun 1996). Selective herbivory by dense deer herds 
damages trees of certain species, potentially leading to long-term changes in forest 
composition (Horsley and Marquis 1983; Stewart and Burrows 1989; Strole and Anderson 
1992; Tilghman 1989). In addition, deer eat more-palatable herbaceous flora (Williams et al. 
2000), allowing less-palatable herb species to increase (Alverson et al. 1988; Waller and 
Alverson 1997). 

Whether or not a plant species is eaten or avoided by selective herbivores like deer can 
be very important for its success, and can be considered a potential influence on the 
invasiveness of a species. Several studies suggest that deer avoid A. petiolata in favor of more 
palatable species, and it is tempting to attribute the invasiveness of A. petiolata, in part, to 
this selective herbivory (Anderson et al. 1996; Williams 1996). A lack of herbivory is often 
suggested as an explanation for why certain non-native species become invasive. This “enemy 
release hypothesis” (Keane and Crawley 2002) primarily concerns herbivory by insects and 
pathogens that specialize on a host plant in its native range and help to regulate its 
population size. When a host plant is introduced to a new region it is possible for these 
specialists to be left behind, resulting in plant population release, and invasiveness. The 
hypothesis also predicts that generalist herbivores will have greater effects on native rather 
than non-native plants, but there is no obvious reason why even selective generalists like 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 27, 


deer should avoid non-native species per se. Because food plant choice by deer is partly 
frequency-dependent (Brown and Doucet 1991), it is even plausible that deer may switch to 
an invasive species if it becomes very abundant while native species become scarce. Field 
experiments are needed to explore whether non-native species are avoided by generalist 
herbivores, and whether preference for native food plants is frequency-dependent. 

We investigated how A. petiolata responded to protection from herbivory by comparing 
caged and uncaged plots of vegetation dominated by A. petiolata in three southeastern New 
York forests (two with deer, one without). We compared changes in percent cover of A. 
petiolata and other vegetation over four years in caged and uncaged plots, and also measured 
differences in A. petiolata size and herbivory rate. We hypothesized that, within a forest, 
there would be no difference between caged and uncaged plots for A. petiolata cover, size, 
or herbivory rate if deer did not eat A. petiolata. Alternatively, if deer did eat A. petiolata, 
then cover would be less, plants would be smaller, and herbivory greater in uncaged plots 
relative to caged plots, but only in the forests with deer. 

Our hypotheses address direct effects of herbivores on A. petiolata. It is also possible for 
herbivores to have indirect effects on plant cover and size (Damhoureyeh and Hartnett 
1997). For example, smaller size and lower cover of A. petiolata in caged plots could be 
caused by increased competition from native plants released from herbivory. In addition, 
herbivory is only one of the complex ways that deer can affect plant cover and size and lead 
to differences in caged and uncaged plots (Waller and Alverson 1997). Trampling can damage 
plants, but may also create disturbed microsites for recruitment. Nutrient addition from deer 
scat may favor certain plant species, particularly invaders (Stohlgren et al. 1999). Our 
experiment does not directly assess all deer affects, but our results can be interpreted in 
reference to them. 


STUDY AREAS 


We conducted the caging experiment in three forests. The 16-ha New York Botanical 
Garden (NYBG) Forest, located in the Bronx, New York City, is an old-growth, never 
clearcut forest remnant, completely surrounded by a highly urbanized landscape. The latest 
published vegetation survey, from 1985, showed that canopy dominants were, in order of 
importance, Tsuga canadensis (L.) Carriere, Quercus rubra L., Acer rubrum L., Betula lenta 
L., Fagus grandifolia Ehrh., Liquidambar styraciflua L., Prunus serotina Ehrh., Liriodendron 
tulipifera L., and Fraxinus americana L. (Rudnicky and McDonnell 1989). There has been 
high mortality of T: canadensis since that survey due to the hemlock woolly adelgid (J. A. 
Morrison, unpublished data). Deer are not present in the NYBG Forest. 

The other two forests are Westchester County’s 84-ha Kitchawan Preserve, in Kitchawan, 
New York, and The Nature Conservancy’s 86-ha Mt. Holly Sanctuary, near Cross River, 
New York. Both preserves consist of second-growth deciduous forest with closed canopy, 
located in northern Westchester County. Kitchawan is 42 km north of NYBG and Mt. 
Holly is 15 km northeast of Kitchawan. Both forests are contiguous with tracts of privately 
held forest fragments embedded in a suburban matrix of mixed land use, including houses, 
lawns, and forest. No formal canopy study has been done at either Westchester site, but 
common canopy trees at Kitchawan are A. saccharum, B. lenta, Fraxinus pennsylvanica 
Marshall, P. serotina, and Quercus spp. Common trees at Mt. Holly are A. rubrum, B. lenta, 
P. serotina, and Quercus spp. (J. A. Morrison, personal observation). Common herb layer 
species in the three forests are shown in Table 1. White-tailed deer are not present in the 


28 BARTONIA 


NYBG Forest, but Kitchawan and Mt. Holly fall within the northern Westchester region, 
with estimated density of more than 50 deer km™. Mt. Holly in particular has the 
appearance of a forest strongly impacted by deer, with a clearly defined browse line and a 
nearly barren herb layer in many places (J. A. Morrison, personal observation). 


METHODS 


At NYBG and Kitchawan, in July 1996, we established four pairs of 4-m? plots, with each 
plot surrounded by an additional 0.5-m walkway. We originally established four pairs at Mt. 
Holly also, but two pairs had to be eliminated due to a new property line demarcation. We 
established two new pairs at Mt. Holly in May 1997; initial data collected from the two 
older pairs were dropped from the study. We chose locations for plots by searching each 
forest for four areas where A. petiolata occurred in stands large enough to accommodate the 
plots. A plot pair was situated within each stand so that each plot had similar A. petiolata 
densities. One plot plus walkway, per pair, was randomly assigned to a caging treatment 
(described below). Distances between the outer edge of the walkways of caged and uncaged 
plots, within a pair, ranged from 0.5 to 2 m. Distances between the four stands within a 
forest ranged from 10 to 500 m. 

We censused the herb layer vegetation in all plots, prior to cage installation, between July 
10 and 25, 1996, except in the two new plots at Mt. Holly, which we initially censused 
between May 28 and June 10, 1997. We divided each 4-m? plot into sixteen 0,25-m? subplots 
with a quadrat frame, and by careful visual estimates assigned a percent cover interval score 
for each species in each subplot, as follows: < 5%, 5-10%, 11-20%, 21-30%, 31-40%, 41-50%, 
51-60%, 61-70%, 71-80%, 81-95%, > 95%. This method allowed us to census all plots within 
a short enough time interval to avoid large phenological differences from one plot to 
another. Dividing plots into 16 smaller subplots allowed us to be more accurate in our visual 
rankings. We were able to stand in the walkway on all sides of the 4-m’ plot and lean over 
each subplot to make estimates, thus avoiding trampling the estimated vegetation. To obtain 
total percent cover per species per 4-m? plot, we converted interval scores to interval 
midpoints, summed across all 16 subplots, and divided by 16. Total percent cover for all 
native species combined was calculated by adding the values for all native species. In a few 
cases this resulted in more than 100% cover in a plot due to overlapping layers of foliage. 
Species names and native status were assigned according to Gleason and Cronquist (1991). 
Specimens of species not readily identified in the field were brought to the lab for 
identification and are stored at The College of New Jersey. Specimens of some uncommon 
species lacked sufficient characters for identification; they were assumed to be native. 

We installed the cages after the initial censuses in July 1996 or June 1997 (for the two plot 
pairs added at Mt. Holly). Cages were square, with an open top, made with flexible plastic 
fencing stapled to 2-m cedar posts at each corner. The fencing at Kitchawan and Mt. Holly 
was strong black polypropylene netting, with filaments 1 to 2 mm wide and a 36-cm’ 
opening between filaments (manufactured by Deerbusters, Inc., Frederick, Maryland). The 
fencing was staked along the cage bottom; small animals such as voles, birds, or chipmunks 
could move in and out of the plots through the open top or through the fencing at the 
forest floor. The netting used at NYBG was a finer plastic of 0.5 to 1-mm width, with a 4 
cm’ opening. It was loosely staked at the bottom so that small animals could enter the plot 
under the netting as well as through the open top. Stronger netting was required in the 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 29 
Westchester County sites where deer are abundant but its extra expense was not justified at 
NYBG 


The cages excluded larger animals, such as white-tailed deer or eastern cottontail rabbits 
(Sylvilagus floridanus [Allen]). In another study, rabbits were able to chew gates through 
similar fencing material (J. Courteau, personal communication), but we observed no such 
gates in any of our cages. Our study focused on deer exclusion because of the overabundance 
of deer in suburban forests but we have observed rabbits on the NYBG grounds near the 
forest and presumably they are present in the other two forests also (but were not observed 
by us). Differences between caged and uncaged plots in the NYBG Forest could be 
attributed to exclusion of rabbits, while differences in the two suburban forests could be 
attributed to deer or rabbit exclusion. 

The very thin filaments of the caging material allowed free movement of air and no 
appreciable shading and neither type of fencing was considered likely to alter microsite 
conditions inside cages compared to uncaged plots. We documented light and temperature 
in the plots. Using an AccuPAR 2000 (Decagon Devices, Inc., Pullman, Washington), we 
measured photosynthetically active radiation (PAR) in caged and uncaged plots. Measure- 
ments were taken in all pairs of plots at Mt. Holly and Kitchawan on September 17, 2000. 
At NYBG, two pairs of plots were measured on September 20, 2000 (the two other pairs 
of plots had been eliminated by treefall and flooding during the previous year). The 
AccuPAR was configured to first read full-sun PAR from a nearby light gap or forest edge, 
and then, at the plot in the forest, collect and average measurements from 10 points along 
a 1-m probe over 30 seconds. The probe was held over a plot at waist height at four 
regularly spaced positions, ensuring that no shade was cast by the operator. We used the 
averages of the four probe positions to obtain percent of full-sun PAR transmitted to the 
plot and found no significant difference between caged and uncaged plots (mean percent [SE]: 
caged, 5.01 [0.023]; uncaged, 2.97 [0.009]; one-tailed t-test for paired comparisons, t = 1.21, 
P = 0.13, N =10). We measured one-time temperature in caged and uncaged plots at Mt. 
Holly and Kitchawan at the same time PAR data were collected and found no difference 
(mean degrees C [SE]: caged, 17.87 [0.895]; uncaged, 17.81 [0.886]; one-tailed t-test for paired 
comparisons, tf = 1.00, P = 0.18, N = 8). 

We censused the plots twice more, from May 28 to June 3, 1998, and May 21 to May 29, 
2000, following the same procedure described above. Alliaria petiolata is a biennial, so plants 
were either in rosette form or flowering form during the censuses; we combined both forms 
when estimating cover. We did not quantify rosette and adult forms separately, but field 
notes indicate that most of the A. petiolata in 1996, 1998, and 2000 was in flowering form 
in all plots except for one uncaged plot at NYBG that was dominated by rosettes. Censusing 
the same plots every two years allowed us to see A. petiolata stands at the same life history 
stage during each census, with the exception of the two new plot pairs at Mt. Holly 
established in 1997. The May/June census dates in 1998 and 2000 were earlier in the growing 
season than the July 1996 census but A. petiolata rosettes have largely finished their spring 
growth by the end of May and do not change size appreciably though July. Flowering adults 
have cauline leaves, but these leaves are present and fully expanded by late May and are 
retained through July (Anderson et al. 1996) at our sites (J. A. Morrison, personal 
observation). Therefore we were confident that plots in different treatments were not 
affected differently by the census dates. 

We analyzed percent cover of A. petiolata and total percent cover of all native plant 
species with repeated measures analysis of variance (von Ende 1993) using PROC GLM of 


30 BARTONIA 


SAS v. 6 (SAS Institute 1990). There were a few other non-native species present, but they 
contributed little to total percent cover in these plots, and so were not included in the 
present study (the exception was Polygonum cuspidatum Sieb. and Zucc., but it occurred in 
only one set of plots at NYBG). Between-plot effects were caging and stand, and within-plot 
effects were time, time x caging, and time x stand. 

The percent cover measures were analyzed separately for each forest because of differences 
in the duration and timing of data collection, However, because the experiments tested the 
same hypotheses we determined the significance of F statistics with the Simes-Hochberg 
method (Simes 1986; Hochberg 1988), a sequential Bonferroni correction. At Kitchawan, 
four pairs of plots were in the experiment from 1996 through 2000. At Mt. Holly, two pairs 
of plots were added one year later than the others. At NYBG, repeated measures analysis 
of all four pairs could be done only through 1998 because one pair of plots was destroyed 
by flooding and another by a fallen tree in 1999. 

We assigned all pre-caging measurements a 1996 date in the Mt. Holly analysis, even 
though two plot pairs were initially measured in 1997. Combining the 1996/1997 initial 
census dates is reasonable ecologically because the data from both years describe the 
vegetation before the caging treatment was begun. Our focus is the comparison of percent 
cover change between caged and uncaged plots, and the 1997 plots were equally divided 
between caged and uncaged treatments. 

We also measured size of adult, flowering A. petiolata individuals inside and outside of 
cages during the June 1998 census. Ideally, we wanted a size measurement that would 
capture individual biomass, since our observations of uncaged plants suggested that they 
were smaller overall, with both shorter stems and smaller leaves. We could not measure 
destructively, however, so we opted to measure stem length of adult plants. We measured 
the degree to which stem length reflects plant biomass by destructively sampling additional 
A. petiolata plants along transects in each forest (81 plants total) and correlating stem length 
with aboveground dry mass, obtained after harvesting and drying plants to constant weight 
at 60° C (Pearson’s r = 0.79, P < 0.01). 

We measured size of all adult A. petiolata individuals within each plot, or up to 32 plants 
per plot, sampling in a systematic manner by dividing the plot into an 8 x 4 grid and 
measuring the plant closest to each grid intersection point. We collected size data from all 
four pairs of plots at Mt. Holly and from three pairs at Kitchawan (one was inadvertently 
not sampled) and at NYBG (one had only rosettes at the 1998 census). We tested for 
difference in plant size between caged and uncaged plots in each forest with t-tests for paired 
comparisons, using the mean plant size per plot as the tested variable to avoid pseudoreplica- 
tion (sample sizes in Table 2). 

In September 2000, we scored A. petiolata plants for presence or absence of herbivory 
inside and outside of cages. We observed all plants in the plots and in the 0.5-m walkway 
surrounding the plots, noting whether rosettes had any bitten petioles with missing leaves, 
or no bitten petioles and all leaves present. We used G-tests for independence (Sokal and 
Rohlf 1981) for each forest data set to determine if the frequency of bitten plants inside 
cages differed from the frequency outside of cages. 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 31 


Table 1. Mean percent cover estimates of 10 most abundant herb layer plant species in Alliaria- 
dominated 4-m/? plots (N = 8) at start of experiment before caging. Plots were censused in July 1996, 
except for four at Mt. Holly that were added and censused in June 1997. SE = standard error. 


mean 
% cover SE 
Mt. Holly 
Alliaria petiolata (Bieb.) Cavara & Grande 29.92 3.82 
Eupatorium rugosum Houttuyn. i Fe 4 | 2.21 
Carex sp. 1.68 0.86 
Oxalis sp. 1.53 0.81 
Berberis thunbergii DC 0.59 0.47 
Fraxinus americana L 0.43 0.20 
Arisaema ih pe 2 (L) Schott 0.40 0.30 
Picket 0.40 0.18 
m “ 0.27 0.10 
por orbiculatus Thunb. 0.19 0.10 
Kitchawan 
Alliaria petiolata (Bieb.) Cavara & Grande 15.86 1.84 
Acer saccharum Marshall 4.73 1.26 
Fraxinus americana L. 4.65 1.84 
Galium sp. 2.19 1.52 
Parthenocissus quinquefolia (L.) Planchon 1.91 0.92 
Polystichum acrostichoides (Michx.) Schott 1.52 0.91 
Carex spp. 1.50 0.87 
Lindera benzoin (L.) Blume 1.29 0.39 
Arisaema triphyllum (L.) Schott 0.94 0.49 
fern 0.94 0.94 
NYBG 
Alliaria petiolata (Bieb.) Cavara & Snes 44.31 6.46 
Polygonum eaverrt Sieb. & Zuc 14.15 8.36 
Circea lutetiana 2.62 1.97 
Impatiens capensis elt 1.88 1.34 
Phellondendron amurense Maxim. 1.67 1.05 
Acer negund: io 1.57 
Solanum sp Pe 1.13 
Fraxinus americana L. 1.06 0.93 
Commelina communis L. 0.90 0.81 
Viola sp. 0.90 0.56 


RESULTS 
Percent Cover 


Over the course of the experiment at Mt. Holly, percent cover of A. petiolata decreased 
significantly in uncaged plots relative to caged plots (Fig. 1). This is shown by the significant 


32 BARTONIA 


Table 2. Number of plant size measurements of Alliaria petiolata used to calulate average plant size 
per plot in caged and uncaged plots. 


plot number of plots number of plant size 
Forest treatment measured measurements in each plot 

Mt. Holly caged 4 $2,52,-21;51 

uncaged 4 16, 14, 17, 23 
Kitchawan caged 3 32, 28, 32 

uncaged 3 a4, 33 
NYBG caged 3 32, 32,32 

uncaged 3 32; 32,12 


caging effect, and in the marginally significant time x caging interaction in the repeated 
measures analysis (Table 3A). Before caging, those plots assigned to the caging treatment had, 
by chance, somewhat lower average A. petiolata percent cover compared to plots assigned 
to the uncaged treatment, but over four years the caged plots ended up with significantly 
higher percent cover (Fig. 1). In Kitchawan Preserve (Table 3B) and in the NYBG Forest 
(Table 3C), percent cover of A. petiolata was not significantly different in caged and uncaged 
plots, although at Kitchawan the trend was toward lower cover in uncaged plots (Fig. 2), 
and at NYBG there was no consistent trend across the four years (Fig. 3). 

Native plant percent cover showed no significant differences between caged and uncaged 
plots at Mt. Holly (Table 3A, Fig. 1), Kitchawan (Table 3B, Fig. 2), or NYBG (Table 3C, 
Fig. 3). No consistent trend of differences in native plant cover between caged and uncaged 
plots was evident. 


Alliaria petiolata Size 


Individual A. petiolata plants were significantly larger in caged plots than in uncaged plots 
at Mt. Holly (Fig. 4; t-test for paired comparisons: Mt. Holly, t = 3.19, df = 3, P = 0.05). 
At Kitchawan, sizes were similar in caged and uncaged plots (Fig. 4; t = 0.68, df = 2, 
P = 0.57). At NYBG, they were more variable and not significantly different (Fig. 4; 
t = 1.18, df = 2, P = 0.36). 


Herbivory 


Herbivory on A. petiolata plants occurred with significantly greater frequency in uncaged 
plots, compared to caged plots, at both Mt. Holly (Fig. 5; G.y = 132.13, df = 1, P < 0.001) 
and Kitchawan (Fig. 5, Gy = 46.87, df= 1, P < 0.001). At Mt. Holly, 27% of the 255 
uncaged plants and 0.82% of the 488 caged plants observed had bitten petioles. At 
Kitchawan, bitten petioles were present in 8% of the 562 uncaged plants and 0.21% of the 
460 caged plants. The NYBG plants did not experience herbivory either inside or outside 
of cages, except for one out of 332 uncaged plants observed (the G-test of independence for 
NYBG data was not needed, or possible, because the frequency of observations for one level 
of the herbivory factor was so low across both levels of the caging factor). 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 


% cover 


% cover 


|A. petiolata 


2 
wi 
‘ ® 
3 
|native plants 
n®@ 
2G "Sr ‘98 


(before caging (N= 4) 
N= 2) 


a 


‘00 


(N= 4) 


Figure 1. Mt. Holly Sanctuary: estimated percent cover (mean + SE; small error bars are hidden by 


the symbol) in 4-m? plots that were caged to prevent herbivory (squares) or uncaged (circles). 


% cover 


o 


Oo 


—- NO WwW 
Oo © 


|A. petiolata 


os 


native tebe : 


a? 


'96 ' 


(before caging (N= 4) 
N= 4) 


Figure 2. Kitchawan Preserve (see Fig. 1 caption). 


’ 


34 BARTONIA 


j * 
99 | A. petiolata nee 
80 | 
. 60 
© a 
Sone @ 
se 40 
30 +. 
20 
10 4 
5 20 native plants 
B sok Mod | 
ee 
'96 '98 ‘00 
(before caging (N = 4) (N = 4) 
N=4) 


Figure 3. NYBG Forest (see Fig. 1 caption). 


a. Mt. Holly Kitchawan NYBG 

Ee 

° 140 4 

= 

4 120 sat. 
100 eo 
L 

® 

© 

& 

Q 

“a. 0 


Figure 4. Mean (+ SE) across plots of average A. petiolata size in plots that were uncaged (circles) or 
caged (squares) for two years (one year for two plots at Mt. Holly) to prevent hebivory (N = 4; small 
error bars are hidden by the symbol). Size is the sum of stem lengths measured on adult flowering 
plants. 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 35 


Mt. Holly Kitchawan NYBG 


® 500 ; 

i mi he erbivory 

= 400 | | Gino herbivory}... 

& 

& 300 | 

5s) 

rs 

900 Foe 

ro) 

s cS a 

= 

e foe oe an 
caged uncaged caged icilae: caged uncaged 


Figure 5. Frequency of A. petiolata plants with herbivory (bitten pitioles and missing leaves), inside 
and outside of cages. Plants were censused in September 2000. 


DISCUSSION 


We expected that protection from herbivory would have little effect on A. petiolata but 
would have a dramatic effect on the native herb layer in the forests with deer. We had three 
major grounds for our expectation: the prevalent idea that non-native species gain an 
advantage, in part, because they lack a suite of herbivores that utilize them as food (Baker 
1974; Mack 1985; Keane and Crawley 2002), comments in the literature that deer do not eat 
A. petiolata (Nuzzo 2000; Tilghman 1989), and our observations of thriving A. petiolata in 
forests with deer herds. However, our results were not consistent with this expectation. 

We found that native vegetation did not respond to protection from herbivores after four 
years, but A. petiolata did respond under certain circumstances. The following evidence 
supports this conclusion: (1) there was no difference in percent cover of native vegetation 
inside and outside of cages over the four years of the experiment; (2) A. petiolata cover was 
higher inside of cages, but only in the two forests with deer, and significantly higher only 
at Mt Holly, where the native vegetation is especially denuded, potentially providing little 
food for mammalian herbivores, particularly in the winter; (3) individual A. petiolata plants 
were larger inside cages only in one of the forests with deer, Mt. Holly; and (4) there were 
much greater herbivory rates on uncaged A. petiolata in the forests with deer, especially at 
Mt. Holly, and hardly any herbivory at NYBG, the forest without deer. 

The lack of response to caging by native vegetation at NYBG can be explained by the 
absence of deer in the forest and appears to indicate that there is also little herbivore 
pressure from rabbits. In the forests with deer, however, reasons for the lack of native plant 
response are less clear. It is possible that some single-species percent cover responses 
occurred, but if so they were not great enough to affect the overall cover of the native 
community (a future paper will explore responses of individual species). There was some 
indication of native cover increase in the 2000 data at Kitchawan (Fig. 2) indicating that a 
longer time of protection from herbivory may allow natives to recover, but that trend was 
not observed at Mt. Holly. We did not directly measure herbivory on native species as we 


a6 BARTONIA 


did for A. petiolata, but it is unlikely that native vegetation would be avoided by mammalian 
herbivores; the browse line and barren appearance of the herb layer strongly suggest 
otherwise, especially at Mt. Holly. We hypothesize that the lack of response by the native 
community compared to A. petiolata may be explained by the fact that the native 
community was so severely reduced to begin with, while the A. petiolata population was 
comparatively vigorous, with many successfully reproducing individuals. The native species 
were sparse and small, and perhaps had little resources to draw on for growth after release 
from herbivory. In addition, many species may have a depleted seed bank due to a history 
of chronic overbrowsing, in which case recruitment could remain very low even when pro- 


Table 3. Repeated measures analyses of variance for percent cover of Alliaria petiolata and native 
species in caged and uncaged 4-m? plots situated in four A. petiolata stands in each of four sites. 
Asterisks denote significance based on the Simes-Hochberg sequential Bonferroni procedure, which 
provides critical values of aw’ for three tests across the three sites (? = <0.10;* = <0.05; ** = <0.01). 
The “adjusted P” values given for within-plot effects are conservative tests that account for departures 
from sphericity in the variance-covariance matrix in repeated measures data. (A) Mt. Holly: 
measurements were made shortly before cages were installed in 1996 or 1997 (treated as one date) and 
also in 1998 and 2000. (B) Kitchawan: measurements were made shortly before cages were installed in 
1996 and also in 1998 and 2000. (C) NYBG: measurements were made shortly before cages were 
installed in 1996 and again in 1998 (adjusting P values is unnecessary when there are only two repeated 
measures). 


(Bonferroni- Greenhouse Huynh- 
adjusted signif- -Geisser Feldt 
Source of variation df MS F icance level) adjusted P adjusted P 


(A) Mt. Holly Sanctuary 


Alliaria petiolata 
Between-plot effects 
Caging 


1 5.272 23.85 0.016* 
tan 3 0.956 4.33 0.130 

Error (= caging x stand) 3 0.221 

Within-plot effects (€=0.55) (€= 1.81) 
Time 2 16.807 14.74 0.005* 0.025* 0.005* 
Lad x Acaging 2 7.616 6.68 0.030? 0.073 0.030* 
Tim and 6 0.705 0.62 0.713 0.657 0.713 
Error 6 1.140 


Native Species 


Between-plot effects 
Cagin 


1 0.062 0.18 0.702 

Coane 3 13.704 39.47 0.007* 
Error (= caging x stand) 3 0.275 

ree effects 6=0.575 =1.944 
Tim 2 3.066 25.70 0.001** odio 0. Bote 
Time x ep 2 0.113 0.94 0.440 0.411 0.440 
Tim and 6 2.822 23.65 0.001** 0,009* 0.001** 
Error 6 051 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 37 
Table 3 (cont'd) 
3 
(Bonferroni- Greenhouse Huynh- 
adjusted signif- -Geisser eldt 
Source of variation df MS F icance level) adjusted P adjusted P 
(B) Kitchawan Preserve 
Alliaria petiolata 
Between-plot effects 
Caging 1 0.328 0.31 0.618 
an i) 1.970 1.84 0.314 
Error (= caging x stand) 3 1.069 
Within-plot effects (€=0.65) (€=2.50) 
ime 2 2.479 9.49 0.014* 0.035* 0.0147 
Time x caging 2 0.405 1.55 0.287 0.298 0.287 
Time x stand 6 2.5514 8.99 0.009* 0.029 0.009* 
Error 6 
Native Species 
Between-plot effects 
Caging 1 0 0.01 0.935 
Stand 3 3.064 2.70 0.219 
Error (= caging x stand) 3 1 
——— effects (€=0.694) (€=2.821) 
Tim Zz 2.390 14.82 0.005* 0.015* 0.005* 
Time x Shin ne 2 0.777 4.82 0.057 0.087 0.057 
Time and 6 0.299 1.85 0.236 0.278 0.236 
Error 6 0.161 
(C) NYBG Forest 
Alliaria petiolata 
Between-plot effects 
Caging 1 0.603 1.10 0.372 
tan J 17.469 Lao 0.009* 
Error (= caging x stand) 3 0.550 
Within-plot effects 
ime 1 1.899 0.87 0.420 
Time x caging 1 1.186 0.54 0.515 
Time x stand 3 19.52 8.95 0.052 
Error 3 oe 
Native Species 
Between-plot effects 
Caging 1 0.129 0.47 0.543 
Stand 3 4.849 17.60 0.021* 
Error (= caging x stand) 3 0.275 
Mipingles effects 
Tim 1 0.268 5.20 0.107 
Time x x caging 1 0.241 4.65 0.120 
Tim and 3 2.986 5/77 0.004** 
+ pe 7 0.051 


38 BARTONIA 


tected from herbivory. It is possible, however, that the native community did respond to 
caging with increased recruitment, but our percent cover measure did not detect it. Percent 
cover is very useful for making accurate yet rapid estimates of biomass per species, which 
is important in a study like ours in order to avoid large phenological differences between 
sampling sites. It does not measure numbers of plants, however, so if new individuals 
recruited but contributed little new biomass, the percent cover of the species may not show 
any change. 

We attribute the differences in A. petiolata cover and size between caging treatments to 
protection from a direct effect of herbivory rather than any indirect herbivory effect or 
some other cage effect. Indirect effects of herbivory could be lower cover and size in caged 
plots because of increased competition from plants released from herbivory, or higher cover 
and size in uncaged plots due to decreased competition from plants subject to herbivory. 
However, A. petiolata cover and size showed the opposite pattern — higher in caged plots 
relative to uncaged plots — indicating direct herbivory. Disturbance of the herb layer by 
trampling could be a second direct effect of mammals in our study, but the strikingly lower 
herbivory rate inside cages suggests that herbivory differences were of primary importance. 
If there was another cage effect not attributable to mammal exclusion, we would expect to 
detect it in all three forests. However, differences inside and outside of cages were seen only 
in the two forests with deer and were more pronounced at Mt. Holly, where deer density 
was probably highest. We chose the thin filament mesh for use as caging material to 
minimize any effect on the plant community and, as expected, microsite measurements of 
light and temperature were no different inside and outside of cages. 

We did not detect the animal species responsible for the observed herbivory on A. 
petiolata. It makes sense to attribute the herbivory to deer, because it best explains our 
results and because of the high density of deer in northern Westchester County (>50 km”), 
but it is possible that leaves could also have been taken by rabbits (S. floridanus) or voles (for 
example, Microtus pinetorum [Le Conte]). However, if the herbivory we observed was due 
only to small mammals such as voles, then we should have consistently seen little difference 
inside and outside of cages because small animals could easily access the caged plots through 
or under the mesh. There was some herbivory inside cages at Mt. Holly and Kitchawan 
indicating the presence of small herbivores, but there was significantly more herbivory 
outside of cages. If the difference was due to rabbits, that would not explain why herbivory 
on A, petiolata was nearly nonexistent at NYBG but common at Kitchawan and Mt. Holly, 
because we know that rabbits were present at NYBG. 

We do have reason to think that deer were browsing more heavily at Mt. Holly. There 
was a noticeable deer browse line at Mt. Holly and at every visit over four years we 
observed deer in the forest. Native plant cover measured before the experiment was lower 
at Mt. Holly than at Kitchawan (Figs. 1 and 2), which is consistent with the almost barren 
herb layer throughout much of the Mt. Holly forest. Kitchawan did not have a distinct 
browse line and the herb layer appeared more abundant (J. A. Morrison, personal 
observation). We did sight deer at Kitchawan but less frequently than at Mt. Holly, even 
though these forests are not far from each other and are in similar landscapes. We 
hypothesize that deer may be less prevalent at Kitchawan because many area residents take 
dogs there, usually allowing them to run unleashed. We saw deer at every visit to Mt. Holly 

e saw dogs at nearly every visit to Kitchawan. Our evidence that deer are the animals 
responsible for herbivory on A. petiolata is indirect but compelling. It would be useful to 
investigate this problem more closely with hand-lens examination of bitten petioles at 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 39 


regular intervals throughout the year, in order to distinguish the shredded bites of deer from 
the clipped and nibbled bites of rabbits and voles (Strole and Anderson 1992). Experimental 
feeding trials would also be helpful. 

A key reason for successful invasion by a non-native plant species is commonly thought 
to be a relative lack of herbivory because of escape from herbivore species found in its 
native range (Baker 1974; Mack 1985; Keane and Crawley 2002). In fact, the enemy release 
hypothesis is the premise upon which the scientific discipline of biological control is based 
(Debach and Rosen 1991; Guretzky and Louda 1997) and a biological control program is 
being developed for A. petiolata (Blossey et al. 2001). The idea applies especially to feeding 
by highly specialist insect herbivores but it may not apply to herbivores with a broader diet. 
White-tailed deer have diet preferences leading to avoidance of relatively unpalatable plant 
species as long as preferred species are available (Alverson et al. 1988; Longhurst et al. 1968; 
McCullough 1985; Nudds 1980; Short 1975; Strole and Anderson 1992; Vangilder et al. 
1982); however, they do not rely on any tightly co-evolved genetic relationship with their 
host plants. The fact that A. petiolata is non-native is probably of little importance in 
whether it is eaten by deer compared to the fact that it is a member of the mustard family 
(Brassicaceae) and so contains a suite of secondary chemicals (Chew 1988; Cole 1975; Larsen 
et al. 1983; Van Etten and Tookey 1979). These bitter compounds can make mustards less 
palatable to vertebrate herbivores, although cows in Ontario are reputed to eat A. petiolata 
leaves in autumn and spring-(Cavers et al. 1979). Anecdotally, A. petiolata is considered 
unappealing to deer (Tilghman 1989). 

A, petiolata’s life history, on the other hand, could encourage herbivory, especially during 
the winter, by animals that otherwise would avoid such a chemically defended plant. It 
germinates in early spring and spends the following winter as a basal rosette of green leaves, 
even growing new leaf tissue in the winter months (Anderson et al. 1996). In addition, it 
begins spring growth before nearly all other understory plants and shrubs (J. A. Morrison, 
personal observation). Fresh A. petiolata leaf tissue is thus available to herbivores throughout 
the winter, when most other foliage is unavailable. If deer make frequency-dependent food 
choices, a plant species that occurs at very low frequency may be relatively ignored, but may 
become a primary food as its proportional representation in the flora increases (Brown and 
Doucet 1991). In addition, deer are predicted to shift to a more generalist diet during winter 
(Nudds 1980). Whether or not A. petiolata has become a primary food for deer at Mt. Holly 
we cannot say, but our results are consistent with deer including A. petiolata in their diet 
because of a lack of other forage plants below the browse line. If deer do feed on A. petiolata 
at some sites, it is possible that biological control in those sites will have little additional 
effect because deer already may be suppressing the A. petiolata population to a substantial 
degree. 

It appeared that deer ate A. petiolata at lower rates at Kitchawan and had less effect on 
its size and cover than at Mt. Holly. The 8% of uncaged plants with herbivory at Kitchawan 
was lower than the 27% at Mt. Holly, and this was just a one-time look at herbivory in the 
fall while there were still other species available as food. It would be of interest to measure 
and compare herbivory on native species and A. petiolata over the course of the year, 
especially in the winter when most other species have senesced or are perennating 
underground, unavailable to herbivores. There is so little plant food available to deer at Mt. 
Holly in the winter that the green foliage of A. petiolata rosettes may be their only choice, 
while at Kitchawan, where the woody vegetation is not as denuded, other foods are 
available. Another possible explanation for different herbivory rates among sites is 


40 BARTONIA 


population variation in defensive chemistry. Recent evidence indicates that different 
populations of A. petiolata express different levels of chemical defenses (Haribal and Renwick 
2001), which can be due to differences in site environmental quality (Cipollini 2002). 

Our results for A. petiolata in metropolitan forests near New York City are likely to be 
relevant to other similar areas where deer herds cause overbrowsing and there are 
populations of invasive plant species. Forest fragments adjacent to human use and habitation 
are increasing. These forests have a high edge-to-interior ratio and can have a high rate of 
immigration of invasive plant species (Brothers and Spingarn 1992; Hill 1985). They often 
support large deer herds that have virtually no natural predators and are subject to very 
limited or no hunting. Lands that face this dual challenge of an overabundance of deer and 
invasion by non-native plants may also be the repositories for much of an entire 
metropolitan region’s biological diversity. We need to understand the relationship between 
deer abundance and the spread of non-native plants in the urbanizing landscape if our goal 
is to maintain native plant biodiversity in these areas. 

Ecosystem management should become the tool of choice for limiting plant invasions 
(DeCalesta 1997; Hobbs and Humphries 1995), but we urge caution in assuming that 
management of deer in an ecosystem will necessarily be effective in reducing the abundance 
of invasive plants and favoring recovery of the native plant community (Alverson et al. 
1988). In a forest such as Mt. Holly, with high deer density, very denuded vegetation, and 
herbivory on the A. petiolata population, deer herd reduction could potentially cause further 
population growth of A. petiolata before there is time for the native community to recover. 
We did not measure reproductive effort of A. petiolata directly, but it is likely that the 
observed larger size of plants protected from herbivory would result in more seeds because 
there is a correlation between seed number and plant size in this species (Byers and Quinn 
1998). Such forests may be particularly appropriate for deer management combined with a 
program of invasive species biological control, perhaps augmented with native species 
restoration efforts. When planning management of metropolitan forests to preserve and 
increase biodiversity, attention should be paid to if and how deer affect invasive plant 
population growth, and how the effect may alter as densities of both change over time. 


ACKNOWLEDGMENTS 


Forest research at the New York Botanical Garden was supported by funding from the 
Baldwin Foundation, with additional science funding at NYBG from The LuEsther T. Mertz 
Charitable Trust, the Andrew W. Mellon Foundation, Bristol-Myers Squibb Company and 
Bristol Myers Squibb Foundation, Inc., Mr. and Mrs. Thomas J. Hubbard, and the Texaco 
Foundation. Much of the work was done under a grant from the FIRSL/SOSA Committee 
of The College of New Jersey to J. A. Morrison, for release time from teaching for research. 
For field assistance and data entry, thanks go to C. Bui, R. Cardeiro, R. Ford, M. Love, J. 
McLaughlin, D. Melman, J. Roth, S. Sullivan, and S. Swahla. Don Cipollini, Jacqueline 
Courteau, Faith Kostel, Victoria Nuzzo, Scott Ruhr, Charles Williams, and two anonymous 
reviewers made very helpful suggestions on the manuscript. Thanks to Westchester County 
Department of Parks, Recreation, and Conservation for permission to conduct research at 
Kitchawan Preserve. Thanks to the Lower Hudson Chapter of The Nature Conservancy for 
permission to conduct research at Mt. Holly Sanctuary. 


EFFECT OF HERBIVORE EXCLOSURE ON ALLIARIA PETIOLATA 41 
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Bartonia No. 62: 45-54, 2004 


The Genus Ophioglossum in Pennsylvania 


BONNIE L. ISAAC,’ CARL F. CHUEY? AND JOSEPH A. ISAAC? 
'>Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213 
Youngstown State University, One University Plaza, Youngstown, Ohio 44555 


ABSTRACT. Three species of Ophioglossum are known from Pennsylvania. The distribution of 
Ophioglossum vulgatum Linnaeus and Ophioglossum pusillum Rafinesque is related to the glacial 
boundary in Pennsylvania. Ophioglossum engelmannii Prantl is known from only one county. Until 
this report, O. vulgatum was considered extirpated in the state. The status of O. pusillum should be 
reassessed, and O. engelmannii is endangered. A key to the species is provided. 


INTRODUCTION 


Efforts to identify specimens of Ophioglossum collected at three different sites discovered 
in Greene County in 1993 and 1996 led to the need to understand the differences among the 
species occurring in Pennsylvania. The Greene County specimens were determined to be O. 
vulgatum, which, at that time, was a species listed as extirpated in the state by the 
Pennsylvania Natural Heritage Program. In this paper, we provide a key to the species of 
Ophioglossum in Pennsylvania, a discussion of the results of our investigation, and a review 
of the legal status of each species. 


METHODS 


Nomenclature follows the Flora of North America (Wagner and Wagner 1993). Other 
names have been used in recent floras. The Southern adder’s-tongue, O. vulgatum, has 
also been known as O. pycnostichum (Fernald) A. Love & D. Léve and O. vulgatum L. 
var. pycnostichum Fernald. Northern adder’s-tongue, O. pusillum has been referred to as 
O. vulgatum L. var. pseudopodum (S.F. Blake) Farwell. No synonymy for O. engelmannii, 
the Limestone adder’s-tongue, was found. 

We investigated the occurrences of Ophioglossum species to find and map historic 
localities in Pennsylvania. Based on characteristics published by Wagner and Wagner 
(1993) we found that several of the specimens at the Carnegie Museum of Natural 
History (CM) herbarium were misidentified. After determining these specimens, 
Ophioglossum vulgatum was discovered to be more widespread than previously believed. 
In addition to the CM specimens, loans of Pennsylvania specimens of Ophioglossum were 
requested from the Cleveland Museum of Natural History (CLM), Ohio State University 
(OS), Penn State University (PAC), Academy of Natural Sciences of Philadelphia (PH), 
Shippensburg University, and West Virginia University (WVA). Specimens were also 
examined at the Missouri Botanical Garden (MO) and Youngstown State University 
(YUO). Herbarium specimens were also requested from the northern panhandle of West 
Virginia and Ohio counties adjacent to Pennsylvania for examination. The following key 
uses the characters that the authors found to be reliable for identification. 


Submitted 27 June 2003, revised 5 October 2004 


45 


46 BARTONIA 


KEY TO OPHIOGLOSSUM IN PENNSYLVANIA 


A Blade with apiculate apex, veins forming larger heavier areoles enclosing smaller 
areoles. Plants from thinly vegetated limestone habitats 
Sipe Us ee oe eo Ophioglossum engelmanii Prant| 


A Blade rounded at apex, veins forming areoles that enclose free included veinlets, not 
forming small areoles enclosed by larger areoles. Plants from non-calcareous sites . B 


B Frond widest near base and tapering abruptly to the stipe; dark colored leathery 
persistent sheath present at base of stipe; sporangia oblong and closely crowded. 
Plants from humus-rich woodland areas ..... Ophioglossum vulgatum Linnaeus 


B Frond widest near middle, gradually tapering to the base; sheath at base of stipe if 
present, membranous and papery; sporangia globular and more widely spaced. 
Plants from moist midly acid open areas and meadows 


pita cota has ies my Sa abies gor iar Wit ie tae W a ar gies tome Ophioglossum pusillum Rafinesque 
RESULTS 


A total of 173 herbarium sheets were examined (Appendix 1). Several sheets had more 
than one mounted specimen. Of the Pennsylvania specimens, 74 proved to be O. pusillum; 
the oldest was collected in 1863 from Chester County, the most recent in 1965 from Erie 
County. Eighty-four were determined to be O. vulgatum, the oldest of which was collected 
in Chester County circa 1840 and the most recent in Greene County in 2003. Localities were 
found from 20 Pennsylvania counties for O. pusillum and 26 counties for O. vulgatum. 
Ophioglossum engelmannii was found in Franklin County in 1990 and collected there again 
in 2001. All but two of the Ophioglossum collections north of the glacial boundary are of O. 
pusillum and the glaciated region accounts for more than half of this species’ recorded 
occurrences in the state (Fig. 1). 

Specimens were requested from the four Ohio counties adjacent to Pennsylvania and the 
northern panhandle of West Virginia. Seven specimens from two Ohio counties were 
examined. O. pusillum was found from both Ashtabula and Mahoning Counties. 
Ophioglossum vulgatum was found from Ashtabula County only. No specimens were seen 
from the northern panhandle of West Virginia. Two sheets appeared to be mixed specimens 
and two sheets were too poor to determine to species level. (Appendix 1) 


DISCUSSION 


Keys tend to rely on the presence or absence of a basal sheath to differentiate O. vulgatum 
and O. pusillum. Many specimens examined did not have roots and in most of these cases 
the basal sheath was lacking. McAlpin (1971) assessed the taxonomic usefulness of the 
developmental and morphological nature of the sheaths in separating these species, which 
he considered varieties of Ophioglossum vulgatum. “Regardless of the mechanism, the value 
of the sheath as a character to separate var. pycnostichum from var. pseudopodum tis in the 
persistence of the sheath and not in whether the sheath is present or absent.” Wagner (1971) 


OPHIOGLOSSUM IN PENNSYLVANIA 47 


Figure 1. Localities of Ophioglossum specimens examined from Pennsylvania. 

& = O. engelmanii @ = O. pusillum @ = O. vulgatum © = O. vulgatum without precise locality 
information. The thick line is the maximum southern advance of the most recent (Wisconsinan) 
glaciation. 


pointed out that the “differences are not confined to sheath development alone.” Thus, other 
characters were searched for to help determine the speci species. In combination with 
other characters, frond shape proved to be the most readily available, useful character found 
on herbarium sheets. The blade of the frond when pressed occasionally folds backs upon 
itself and makes it difficult to determine the widest point of the frond. Mature fronds are 
more readily distinguishable. The frond shape by itself, however, is not always diagnostic. 
Blake (1913) noted “the plants, which grew in two adjacent bits of sphagnous meadowland, 
usually in the open but occasionally on the edges of thickets, show great variation in size, 
shape, and position of leaf, size of spike, and number of fronds, sufficient to constitute half 
a dozen ‘species’ if brought back by collectors from as many regions.” 

We found, as did Wagner (1971), that the separation of these two species is, “...strongly 
warranted on the basis of a number of average differences.” Wagner also found that, “...a 
correlation can be demonstrated between the two taxa and the southern boundary of 
Wisconsin{[an] glaciation...”, although this correlation seems to break down near the coastal 
plain. Unique ecological habitats or hybridization (as suggested by Wagner and Wagner 1966) 
may explain this phenomenon. 

The shape of the sporangia is useful before they mature, split, and release spores. Once 
the sporangia split, the shape is difficult to distinguish. Ophioglossum pusillum (Fig. 2) has 
very globular sporangia, which are not as closely arranged as the transversely oblon 
sporangia of O. vulgatum (Fig. 3). The color of the frond can also be helpful. Ophioglossum 
pusillum fronds are duller and paler green than those of O. vulgatum. This character can be 
very difficult to interpret on dried herbarium specimens because of variation in drying 


BARTONIA 


Figure 2. Ophioglossum pusillum (scale bar = 1 cm). 


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OPHIOGLOSSUM IN PENNSYLVANIA 49 


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50 BARTONIA 


techniques. Rapid i ae of specimens aids in color retention. Wagner (1971) also noted a 
difference in spore s 

Ophioglossum illo tends to prefer open mesic sites. McMaster (1994) suggested that 
O. pusillum “...is an early-successional species that frequently occurs in small, isolated habitat 
patches subject to rapid succession.” Ophioglossum vulgatum on the other hand, is most often 
found in rich mesic wooded areas. Ophioglossum vulgatum also generally tends to emerge and 
senesce earlier in the year than O. pusillum. Cranfill (1980) conveniently summarized these 
differences discussed by Fernald (1939, pp. 494-499) Wagner (1971) and Wherry (1961). 

The status of O. vulgatum in Pennsylvania was listed as tentatively undetermined (TU) 
in 1986, “It was listed as TU based on the lack of field investigation concerning verification 
of historical records” (Rare Plant Forum, 1986, unpublished notes). This species then had its 
status changed from TU to extirpated in P ia (PX), following a proposal at the 1991 
Pennsylvania Rare Plant Forum (Rare Plant Forum, 1991 anpublaned notes). At that time, 
O. pusillum was assumed to be a more widely distributed taxon, not in need of protection. 
Parks and Montgomery (2000) noted that O. pusillum is “...uncommon in wet meadows and 
moist woods; throughout.” O. pusillum has never had any special protective status in 
Pennsylvania. However, because this species has not been collected in Pennsylvania since 
1965, its conservation status should be reassessed. The authors are unaware of any fieldwork 
for this species, and several other authors have noted the ease at which Ophioglossum is 
overlooked. Clute (1901) wrote, “It is safe to say that the adder’s-tongue ... is much better 
known to the collector from pictures and herbarium specimens than it is from experience 
in the field. ... All who have once found it testify to the ease with which they subsequently 
find other stations for it, and incline to the belief that its single leaf is often passed under 
the impression that it is the leaf of some flowering plant. ... It seems a plant that one must 
first discover by accident before he can find it by intention.” Cranfill (1980) stated, “because 
of their manifestly unfern-like appearance, together with their proclivity for unfern-like 
habitats ... they are inconspicuous and often passed over.” Wagner and Wagner (1966) noted, 
“The plants are probably more common than our few records ... indicate, but they are 
notoriously easy to overlook (and indeed, many field botanists have never seen the species 
alive and consider it a great rarity).” 

Ophioglossum engelmannii was first listed as endangered in Pennsylvania (PE) in 1991. This 
species is at the northern edge of its range and has a limited, specialized habitat. It is unlikely 
that many more sites for this species will be found in the state. Thus it is likely to retain its 
endangered status in Pennsylvania. 


ACKNOWLEDGMENTS 


The authors wish to thank the curators and collection managers of the cited herbaria for 
the use of their specimens. We would also like to thank Jane Hyland, scientific illustrator 
at Carnegie Museum of Natural History, for completing the artwork on short notice. 


LITERATURE CITED 


BLAKE, S. F. 1913. Forms of Ophioglossum vulgatum in eastern North America. Rhodora 15: 86-88. 

CLUTE, W. N. 1901. Our Ferns in Their Haunts. Frederick A. Stokes Co., New York. 332 pp. 

CRANFILL, R. 1980. Ferns and fern allies of Kentucky. Kentucky Nature Preserves Commission 
Scientific and Technical Series, No. 1: 77-83. 

FERNALD, M. L. 1939. Last survivors in the flora of Tidewater Virginia. Rhodora 41: 465-504. 


OPHIOGLOSSUM IN PENNSYLVANIA 51 


HIOLMGREN, P. K., W. KEUKEN AND E. K. SCHOFIELD. 1981. Index Herbariorum, Part 1. The Herbaria 
of the World, 7th ed. Regnum Vegatabile 106. 452 pp. 

MCALPIN, B. W. 1971. Ophioglossum leaf sheaths: development and morphological nature. Bulletin of 
the Torrey Botanical Club 98: 194-199. 

MCMASTER, R. T. 1994. Ecology, reproductive biology and population genetics of Ophioglossum 
vulgatum (Ophioglossaceae) in Massachusetts. Rhodora 96: 259-286. 

PARKS, J. C. AND J. D. MONTGOMERY. 2000. Ferns. Pp. 71-106 in A. F. Rhoads and T. A. Block, The 
Plants of Pennsylvania: an Illustrated Manual, University of Pennsylvania Press, Philadelphia. 

WAGNER, W. H. 1971. The southeastern adder’s-tongue, il ea vulgatum var. pycnostichum 
found for the first time in Michigan. Michigan Botanist 10: 6 

WAGNER, W. H. JR. AND F. S. WAGNER. 1966. saranda e the Mountain Lake Area, Giles 
County, Virginia: biosystematic studies, 1964-1965. Castanea 31: 121-140. 

WAGNER, W. H. JR. AND F. S. WAGNER. 1993. Ophioglossaceae. Pp. 85-106 in Flora of North America, 
Oxford Press, New York. 

WHERRY, E. T. 1961. The Fern Guide: Northeastern and Midland United States and Adjacent Canada. 
Doubleday and Co., Garden City, New York. 318 pp. 


APPENDIX 1: SPECIMENS EXAMINED 


Standard herbarium acronyms follow Holmgren et al. (1981). 


Ophioglossum engelmannii Prantl 


PENNSYLVANIA: Franklin Co.: 0.5 mi. S of Williamson, between Conococheague Creek & Rt. 955, 
under power line, successional red-cedar woods on limestone, 13 June 1990, J. Walck & L. Klotz 446 
(PH, Shippensburg); near Williamson, near abandoned dirt road, limestone prairie, 22 June 2001, 
D. Laughlin 943 (PAC). 


Ophioglossum pusillum Rafinesque 


OHIO: Ashtabula Co.: East Conneaut, 17 May 1933, L. E. Hicks s.n., (OS); East Conneaut, 17 June 
1933, L. E. Hicks s.n., (OS); Farnham, 17 June 1933, L. E. Hicks & F. B. Chapman s.n., (PH). 
Mahoning Co.: North Jackson, 9 June 1909, E. W. Vickers s.n. (OS). 

PENNSYLVANIA: Beaver Co.: bluff above Potato Garden Run and Raccoon Creek, 23 July 1944, 
L. K. Henry s.n. (CM). Berks Co.: 1 mi. N of Moselem Springs, 18 August 1915, W. H. Leibelsperger 
340 (PH). Bradford Co.: Sayre, swampy ground, August 1905, W. C. Barbour s.n. (PH). Bucks Co.: 
Monroe, July 1885, J. A. Ruth & H. F. Ruth s.n. (PH); near Monroe, July 1885, J. A. Ruth & H. F. 
Ruth s.n. (PH); near Monroe, July 1889, R. Brothers s.n. (PH); The Hedge, Lower Solebury Twp., 
low damp woods, 4 July 1896, A. B. Williams s.n. (PH); near Langhorne, 30 May 1904, L. Sowden 
s.n. (PH); below Woodburne, 30 May 1904, A. Jahn, s.n. (PH); Finland, 5 July 1913, J. R. Mumbauer 
5 (PH); SE tributary to Ridge Valley Creek, Finland, boggy swale on streamlet, 29 June 1921, B 
one | neo (PH); ca. 0.75 mi. W of Monroe, boggy swale, 17 June 1933, B. Long 40559 (PH); 0.5 

3 Geryville, marshy spring-run, 5 June 1938, E. 7. Wherry s.n. (PH). Chester Co.: [no 
ae Sd aed 1863, W. M. Canby s.n.( PH). Crawford Co.: Shelmadine Springs, old yuaetl 11 
September 1939, J. Murdock s.n. (CM); 4 mi. ENE of Meadville, wet depression at woods mar 
7 June 1942, E. T. Wherry s.n. (PH); 4 mi. ENE of Meadville, wet pasture and thickets, 7 June 1942, 
W. H. Wagner 848 (PH). Delaware Co.: 1 mi. NW of Lima, Upper Middletown Twp., 17 June 
1906, F. Pennell s.n. (PH); near Lima, swamp, 6 July 1907, W. A. Poyser s.n. (CM); Prats Swamp, 
N of Lima, among tall grass, 6 July 1907, W. A. Poyser s.n. (PH); Pratts Swamp, Lima, 6 July 1907, 
W. A. Poyser 11153 (PH); Pratts Swamp, Lima, between hummocks, 6 July 1907, W. A. Poyser s.n. 


BARTONIA 


(PH); Pratts Swamp near Lima, 6 July 1907, W. A. Poyser s.n. (PH); Lima, among tall grass, 6 July 
1907, W. A. Poyser sn. (MO). Erie Co.: Presque Isle, 8-9 June 1906, O. E. Jennings s.n. (CM, PH); 
near E end of Presque Isle, July 1926, O. E. Jennings s.n. (CM); Presque Isle, 16 July 1927, J. Bright, 
sn. (CM); 4 mi. SE of Wattsburg, bog, 1 July 1950, W. E. Buker sn. (CM); 3 mi. N of Edinboro, 
off Route 99, Edinboro Bog, among Sphagnum, 17 July 1962, L. K. Henry s.n. (CM); 1 mi. W of 
Corry, calcareous bog, 14 August 1962, C. Hand s.n. (CM); 1 mi. NE of Union City, behind Union 
City Fish Hatchery, bog, 2 July 1963, C. Hand & J. Stull s.n. (CM); 1 mi. E of Union City on Rt. 
6, Sphagnum bog, 26 June 1965, W. E. Buker s.n. (CM); 1 mi. E of Waterford, 26 June 1965, J. Stull 
& D. Stull s.n. (CM). Lackawanna Co.: E of Baylors Lake, Fleetville, damp bramble thicket, in 
clearing, 18 July 1946, S. L. Glowenke 7736 (PAC, PH). Lancaster Co.: Mountville, June 1864, A. 
P. Garber s.n. (PH); West Hempfield Twp., A. Garbers swamp, 5 June 1865, A. P. Garber s.n. (PH). 
Lehigh Co.: opposite Duck Farm Hotel, ~SW of Allentown, knoll in meadow, 16 June 1911, H. 
W. Pretz, 3507 (PH); along Cedar Creek, Griesemersville, marshy meadow, 27 July 1912, H. W. 
Pretz 4853a (PH); S of trolley tracks just SW of Trexlertown, meadows, 27 July 1913, H. W. Pretz 
5927 (PH); about 1.5 mi. S by SW of Lanark P.O., vicinity of springy slope (just E of road) on 
gneissic hillside, 12 September 1915, H. W. Pretz, 7911 (PH); about 1.5 mi. S by SW of Lanark P.O., 
open springy marshy slope, just E of road, 4 July 1917, H. W. Pretz 8863 (PH); Saucon Creek, ca. 
5/8 mi. S by slightly SE of Friedensville Crossroads, in low open meadows beside (S side-W of 
road), 8 June 1919, H. W. Pretz 9690 (PH); just E of Sigmund (Hampton Furnace), in open marshy 
meadow along streamlet, 15 June 1919, H. W. Pretz 9738 (PH); along Little Lehigh River, on S side 
of stream about 3.25 mi. SW by S of Centre Square, Allentown, open (calcareous) marshy meadow, 
15 August 1920, H. W. Pretz 10352 (PH); along S side of Cedar Creek about 2.12 to 2.25 mi. W by 
SW of Centre Square, Allentown, open meadows, 30 May 1924, H. W. Pretz 12083 (PH); 0.25 mi. 
S of Allentown, along Cedar Creek, in meadow, 14 September 1930, C. E. Mobr s.n. (PH); along 
Little Lehigh River, on S side of stream about 3.25 mi. SW by S of Centre Square, Allentown, open 
marshy place, 23 May 1948, H. W. Pretz 13971 (PH). Luzerne Co.: Lily Lake, 29 July 1889, A. A. 
Heller s.n. (PH); Lily Lake, 15-16 August 1889, J. K. Small s.n. (PH). Lycoming Co.: Williamsport, 
25 May 1920, j. P. Young s.n. (CM). Monroe Co.: Tannersville, 4 July 1896, J. Albrecht s.n. (CM); 
Tannersville, 4 July 1896, S. Brown s.n. (PH); Pocono, 4 July 1896, C. D. Fretz s.n. (PH); near 
Tannersville, peat bog, 4 July 1896, 7: C. Porter s.n. (PH); Henryville, 18 August 1906, B. Long s.n. 
(PH). Montgomery Co.: SE tributary to Ridge Valley Creek, Finland, boggy swale on streamlet, 
16 June 1920, B. Long 23301 (PH); Zieglersville, wet meadows, 5 August 1943, J. R. Mumbauer s.n. 
(PH). Northampton Co.: Mount Bethel W + 0.5 mi. S of RR, 1 July 1908, C. C. Bachman s.n. 
(PH); Mount Bethel, +0.5 mi. W, S of RR, 2 August 1908, S. S. Van Pelt & C.C. Bachman 11122 
(PH); 1 mi. W of Wassergass, meadow, 15 August 1946, R. L. Schaeffer, Jr. 24381 (PH). Susquehanna 
Co.: Mud Pond, Ararat region, moist muddy pastured margin, 26 June 1936, E. T. Wherry s.n. : 
Tioga Co.: Wellsboro, 7 July 1869, A. P. Garber s.n. (PH). Warren Co.: near Donaldson, open 
grassy patch on wooded hillside, 29 May 1933, A. N. Leeds 543 (PH); 3 mi. W of Tidioute, 
sphagnous area at edge of maple-hemlock woods, 8 June 1942, W. H. Wagner 879 (PH); 3 mi. N of 
Tidioute, in sphagnous areas at edge of maple-beech hemlock woods, 8 June 1942, CE. Wood 2267 
(PH); near Bear Lake, 22 August 1962, C. Hand s.n. (CM). Wayne Co.: South Sterling, 17 June 1906, 
B. Long s.n. (PH); vicinity of White Oak Pond, 24 August 1920, O. E. Jennings, G. K. Jennings & 
E. M. Gress s.n. (CM); E of Weigh Lake, Preston Twp., swamp, 24 August 1921, H. B. Meredith s.n. 


Ophioglossum vulgatum Linnaeus 


OHIO: Ashtabula Co.: Jefferson, maple grove, July 1917, R. J. Sim s.n. (OS); Farnham, 17 June 1933, 


L. E. Hicks s.n. (OS). 


PENNSYLVANIA: Beaver Co.: southern Independence Township, Witherow, 10 June 1947, E. Mason 


s.n. (CM); 1 mi. from Rt. 30 crossing of Raccoon Creek, 14 July 1951, M. Henrici s.n. (CM); 100 
yards E of Hwy 30, 0.5 mi. N of E end, Raccoon Creek State Park, meadow, 20 July 1964, J. T. 


OPHIOGLOSSUM IN PENNSYLVANIA 55 


Laitsch s.n. (PH). Bedford Co.: 2.5 mi. SE of Alum Bank, moist woods, 1120 ft., 5 July 1952, D. 
Berkheimer 13866 (CM, PH). Berks Co.: Hamburg, 17 May 1891, J. Crawford, s.n. (PH); Hamburg, 
11 July 1892, S. Brown s.n. (PH); Blue Ridge, Hamburg, no date, W. Stone 34 (CM); foot hills, Blue 
Mountains, above Hawley [probably Hamburg], 11 June 1892, J. Crawford s.n. (PH); Hamburg, 15 
July 1891 & 11 June 1892, J. Crawford s.n. (PH); Hamburg, 11 June 1892, B. Heritage s.n. (PH); 1.4 
mi. NW of Shartlesville, rich moist woods, 31 July 1938, W. C. Brumbach 3076 (PH); 1.25 mi. 
NNW of Shartlesville, rich woods, 800 ft., 20 July 1941, D. Berkheimer 2836 (PH); 1.5 mi. NE of 
Bernharts, moist flat in woods on E side of stream (above abandoned buildings), 23 July 1947, E. 
T. Wherry s.n. (PH); 2.25 mi. NE of Bernharts, damp woods, 23 July 1947, E. T. Wherry s.n. (PH); 
1.37 mi. WSW of Hopewell, moist woods, 540 ft., 24 July 1948, D. Berkheimer 10041 (PH); 1.5 mi. 
WSW of Hopewell Furnace, damp soil in low woods, 19 July 1950, W. C. Brumbach 4318 (PH); 1 
mi. NW of Eckville, swamp, 22 July 1953, R. L. Schaeffer, Jr. 44239 (PH). Bucks Co.: Riegelsville, 
1882, J. F. Ruth s.n. (PH); Buckwampum Mtn. near Springtown, 6 August 1903, W. D. Witte s.n. 
(PH); near Gerharts Mill, 4 July 1918, F. Ball s.n. (PH); along Fork of Jericho Creek, 1 mi. ENE 
of Pineville, rich woods, 21 May 1953, B Long 76504 (PH). Butler Co.: NW side of 
Connoquenessing Creek, NW of Zelienople, 4 August 1985, F. Lochner s.n. (CM). Centre Co.: 2 

i. N of Port Matilda, Worth Township, 19 June 1976, W. Harpster s.n. (CM). Chester Co.: West 
Chester, ca. 1840, J. Wolle herbarium s.n. (CM). Crawford Co.: 3 mi. SE of Meadville, soggy humus, 
7 July 1970, R. C. Leberman s.n. (CM). Dauphin Co.: 0.5 mi. S of Manada Gap, 9 mi. N of 
Hummelstown, in swamp, 14 June 1936, E. T. Wherry s.n. (PH); 2 mi. NNE of Dauphin, moist 
woods, 420 ft., 20 May 1952, D. Berkheimer 12563 (PH). Delaware Co.: near Darby, no date, Leidy 
s.n. (PH); Darby Creek, above Bonsalls Mill, June, G. Miller s.n. (PH); Middletown, 4 July 1906, W 
A. Poyser s.n. (PH). Franklin Co.: 4 mi. NE of Ft. Loudon, wet woods along stream, 22 May 1963, 
D. L. Emory s.n. (PH). Greene Co.: ca. 2.5 mi. WNW of Hunters Cave on T-581, Morris Township, 
mixed hardwood forested hillside, 17 June 1993, J. A. Isaac 4363 (CM); ca. 1.5 mi. WNW of 
Triumph, State Game Lands 179, Jackson Township, mixed deciduous woods, dry hillside, 19 June 
1993, J. A. Isaac 4504 (CM); ca. 3 mi. SW of Deep Valley, along Knob Run, mature deciduous forest, 
39°43’N, 80°30’W, 26 May 1996, B. L. Isaac & J. A. Isaac 8912 (CM); 0.3 km WNW of Crabapple, 
dry young forest, 39°55’13"N, 80°28'42"W, 5 June 2002, J. A. Isaac &, M. Takacs 14409 (CM); 1.8 
km NW of Durbin, wooded hillside, 39°55’49"N, 80°29°54"W, 7 June 2002, J. A. Isaac &, M. Takacs 
14421 (CM); just E of Crows Mills, mesic forest near head of small run , 39°55’ 40”N, 80°30°02”W, 
14 June 2002, J. A. Isaac & M. Takacs 14489 (CM); ca. 1.4 km E of Crows Mills, mixed deciduous 
woods, 39°55’45”N, 80°29’22”W, 14 June 2002, J. A. Isaac & M. Takacs 14490 (CM); ca. 1.3 km N 
of Aleppo, Lindera thicket in regenerating forest, 39°49’59"N, 80°26’21”W, 24 April 2003, J.A. Isaac 
15876 (CM); 2.3 km NNE of Aleppo, regenerating forest on old pastures, 39°50’36”N, 80°27°05"W, 
24 April 2003, J. A. Isaac 15878 (CM); ca. 2 km NE of Ryerson Station, oak-maple forested slopes, 
39°54’05”N, 80°27°16”W, 1 May 2003, J. A. Isaac, R. Coxe & S. Ernst 15891 (CM); ca. 1.5 km W of 
Wind Ridge, 39°52’49”N, 80°27’07”W, 1 May 2003, J. A. Isaac, R. Coxe & S. Ernst 15893 (CM); 1.9 
km E of Bryan, 39°52’38”N, 80°27'20"W, 1 May 2003, J. A. Isaac, R. Coxe & S. Ernst 15894 (CM); 

. 2.6 km ENE of Bryan, 39°52’52”N, 80°26’57”W, 1 May 2003, j. A. Isaac, R. Coxe & S. Ernst 
15895 (CM); ca. 2.8 km E of Bryan, 39°53’36"N, 80°26’31”W, 1 May 2003, J. A. Isaac, R. Coxe & 
S. Ernst 15896 (CM). Huntingdon Co.: Martin Gap, Rothrock State Forest, between stream and 
road, red oak-mixed hardwood riparian forest, 44°33’59"N, 77°50’56"W, 880 ft., B. Brokaw & A. 
Weber 2002-1 (CM). Lancaster Co.: Conestoga above Petersville, June 1862, E. B. Weaver s.n. (PH); 
Little Britain Twp., 5 August 1881, gift of Harlan Gatchell s.n. (PH); Haines Station, 4 July 1934, 
M. E. Groff s.n. (PH); 0.5 mi. SE of Haines Station, near S margin of county, moist woods, 7 June 
1936, E. T. Wherry s.n. (PH); 1.5 mi. NW of Hopeland, near Segloch Run, woods, 5 May 1938, L. 
F. A. Tanger s.n. (PH). Lebanon Co.: Mt. Gretna, 2 July 1927, H. A. Ward sn. (PH); Mt. Gretna, 
a) dry woods at SE corner post of camp meeting grounds b) swamp across highway NW of village, 
28 July 1934, E. 7. Wherry s.n. (PH). Lehigh Co.: along Cedar Creek, about opposite the Duck 
Farm Hotel, about on a line with Hamilton St., meadows, 27 July 1912, H. W. Pretz 4853 (PH); 
along Cedar Creek between L.V.R.R. branch & Duck Farm Hotel, Griesemersville, vicinity of slight 


BARTONIA 


rise in marshy meadow, 27 July 1912, H. W. Pretz 4853 (PH); about 3/8 mi. SW by S of 
Crackersport Crossroads, lightly wooded edge of an (at times marshy, or dry) mud-hole or 
depression in woods by streamlet, 21 May 1922, H. W. Pretz 11294 (PH). Mifflin Co.: no further 
locality, July 1850, 7: C. Porter s.n. (MO). Montgomery Co.: near Haverford College, June, Charles 
E. Smith Herbarium s.n. (PH). Northampton Co.: 0.75 mi. SW of Johnsonville, swamp, 24 August 
1950, R. L. Schaeffer, Jr. 34671 (PH). Perry Co.: Hemlock State Forest Park, 12 July 1936, N. B 
Kimber & E. W. Evans s.n. (PH). Philadelphia Co.: on Judge Peters place, near old inclined place 
(now the Park), no date, 1 Burk s.n. (PH); Tacony, Philadelphia, July 1860, 1 Burk s.n. (PH). 
Schuylkill Co.: 0.25-0.5 mi. SSE of Schuylkill Haven, right bank of Schuylkill River, low rich 
woods, 6 July 1938, P. R. Wagner 7471 (PH); S side of river, opposite (SE of) Schuylkill Haven, 
damp woods at base of slope, 8 August 1938, E. T. Wherry s.n. (PH); 1.37 mi. ENE of Auburn, rich 
moist woods, 16 July 1944, D. Berkheimer 5173 (PH); 0.5 mi. ENE of Port Clinton, moist woods, 
480 ft., 20 June 1946, D. Berkheimer 7439 (PH). Snyder Co.: 3-3.5 mi. E of Beavertown, moist 
woods, 21 June 1939, E. C. Earle 2092 (PH); 3 to 3.5 mi. E of Beavertown, damp rich woods, 21 
June 1939, P. R. Wagner 8006 (PAC). Venango Co.: 5 mi. NW of Franklin, springy slope, maple 
woods, 7 June 1942, W. H. Wagner 826 (PH); 5 mi. NW of Franklin, maple woods, 7 June 1942, 
C. E. Wood 2184 (PH); Cranberry Twp., S of U.S. 322 on Whip-poor-will Rd., in loam and clay 
of average to dry moisture in an open exposure, twice a year mowed lawn, 17 July 1980, C. F. 
Chuey 1586 (YUO); Westmoreland Co.: 0.5 mi. S of Laughlintown, 9 July 1934, E. M. Gress & H. 
B. Kirk s.n. (PH); Powdermill Nature Reserve, under Crataegus and Pyrus, edge of swamp from Iron 
Spring, 30 May 1966, L. K. Henry & R. Leberman s.n. (CM); Powdermill Nature Reserve, edge of 
swamp, 20 July 1966, N. D. Richmond s.n. (CM); adjacent to Phoebe Run, near the corner of 
Wilcoxs property, Powdermill Nature Reserve, woods, 27 May 1984, R. C. Leberman & R. S. 
Mulvihill s.n. (CM); ca. 5.3 km E of Stahlstown, mixed deciduous woods, 11 July 2002, B. L. Isaac, 
J. A. Isaac, T. Pearson & D. Byers 14480 (CM). York Co.: Stephenstown, Neffs Hill, 25 June 1935, 
H. B. Kirk & A.B. Champlain s.n. (PH). 


Bartonia No. 62: 55-62, 2004 


The Current Status of Two Rare Species of Ruellia 
(Acanthaceae) in Pennsylvania 


ERIN A. TRIPP 
Department of Biology, Duke ge bel Science .. Box 90338, Durham, NC 27708 
tripp@duke.ed: 


ABSTRACT. Ruellia is a plant genus of conservation concern in Pennsylvania. I examined the current 
status of R. humilis Nutt. and R. strepens L. in the state by conducting surveys to assess the stability 
of populations since their 1984 listing by the Pennsylvania Natural Diversity Inventory. Roughly half 
of the historically occurring populations are extant and half are extirpated or are no longer apparent. 
I attempted to characterize genetic diversity levels within and between these populations with allozyme 
assays, but results were inconclusive. 


INTRODUCTION 


Ruellia, with approximately 250 species, is the second largest genus in the monophyletic 
family Acanthaceae sensu stricto (McDade et al. 2000). Though primarily pan-tropical in 
distribution, the potentially monophylletic Rwellia (E. Tripp, unpublished data) extends into 
temperate latitudes including the United States. The family and genus are most easily 
recognized by having fruits with internal retinacula (hooked structures aiding in ballistic seed 
dispersal), opposite leaves with swollen internodes, and, except in Acantheae, cystoliths 
(calcium crystal deposits visible as short streaks on leaf surfaces with a hand lens or 
dissecting scope). Additionally, members of Ruellia possess a complex and interesting floral 
feature termed the filament curtain. Four fertile didynamous stamens form a partition that 
divides the corolla longitudinally into two compartments. This structure is presumably 
linked to pollinator relationships and switches (Manktelow 2000). 

Acanthaceae have been of great botanical interest because of their widespread distribution, 
diverse habits (herbs, vines, shrubs, and trees), and varied pollinator association. Flowers of 
Ruellia, for example, vary from purple to red, yellow, white, green, and even black, and are 
pollinated by bees, hummingbirds, butterflies, hawkmoths, and bats. The major economic 
value of the family is horticultural; their stunning floral and vegetative morphologies have 
attracted a crowd of breeders and growers. Species of Ruellia and other genera such as 
Acanthus, Aphelandra, Barleria, Eranthemum, Fittonia, Hypoestes, Odontonema, Pachystachys, 
Sanchezia, and Thunbergia are found in the ornamental trade. 

In the Western Hemisphere, species of Ruellia occupy a diversity of habitats extending 
from 43° N in Wisconsin to 35° S in Argentina (Ezcurra 1993). There are roughly 20 species 
of Ruellia in the United States, most of which occur across the Southeast and Texas 
(NatureServe 2003). A few species of Ruellia occur in the arid Southwest (Daniel 1984), and 
at least 30 species occur in Mexico. In the U.S. Ruellia species extend from Pennsylvania 


Manuscript submitted 30 September 2003, revised 12 December 2003. 


55 


56 BARTONIA 


northwest to Minnesota, south to Nebraska, southwest to Arizona, and east to Florida 
(Tharp and Barkley 1949). Unlike most other Acanthaceae, many Ruwellia bear cleistogamous 
as well as chasmogamous flowers. The evolution, genetic basis, and developmental pathway 
of cleistogamy in Ruwellia and other plants remain unresolved (but see Long 1977 and Lord 
1981 for discussions). 

Ruellia humilis Nutt. is currently listed as endangered (S1) and R. strepens L. as threatened 
(S2) in Pennsylvania by the Pennsylvania Natural Heritage Program (P.N.H.P., formerly 
Pennsylvania Natural Diversity Inventory). Ruellia humilis is an $1 (critically imperiled) 
species in Maryland, Michigan, and North Carolina; in Wisconsin it is classified $2 
(imperiled). Rwellia strepens is ranked $1 in Maryland, Michigan, North Carolina, and the 
Washington, D.C. area and $2 in Nebraska (NatureServe 2003). An imperative component 
of state and federal endangered species programs is the periodic re-evaluation of listed species. 
Since P.N.H.P. listed R. humilis and R. strepens in 1984, some new populations have been 
discovered; however, other sites have not been thoroughly checked (J. Kunsman, personal 
communication, 2003). The purpose of this study of Ruellia in Pennsylvania is to ascertain 
any changes in population status and contribute to the effectiveness of our Natural Heritage 
Program. 

In the Northeast, Ruellia caroliniensis (Walt.) Steud., R. humilis, and R. strepens reach their 
northern limit of distribution in Pennsylvania and New Jersey. They are the three most 
widespread Ruellia species in the United States. Ruellia pedunculata Torr. ex Gray, a 
southern species, has also been reported from Pennsylvania, but in only one location near 
a residential area. There are no herbarium records to verify this species in Pennsylvania 
(Rhoads and Block 2003), thus it probably escaped from a nearby garden. R. humilis and R. 
strepens reach north to Minnesota and Michigan, southwest to Nebraska, south to Texas, and 
east to Florida. Ruellia caroliniensis is extirpated from Pennsylvania (Rhoads and Block 
2000), but still occurs in New Jersey and Maryland. It occupies the southeastern U.S., Texas 
and Oklahoma, and its Midwestern distribution terminates in Illinois. 

Ruellia caroliniensis, extirpated in the state, is represented by only two known herbarium 
specimens, both of which I have confirmed as correct identifications. One was collected at 
McCalls Ferry in York County along the Susquehanna River in the late 1800s (Herbarium 
of the Academy of Natural Sciences of Philadelphia, PH). I recently discovered a second 
collection at the University of Wisconsin Herbarium (WIS) in Madison. Collected by 
Thomas Porter (1822-1901), the label contains no data other than “Pennsylvania.” Both R. 
humilis and R. strepens are perennials and grow in the Ridge and Valley physiographic 
province in Cumberland and Franklin counties in south-central Pennsylvania. The latter 
additionally occurs in the Pittsburgh Plateau province in Greene and Washington counties 
(Rhoads and Klein 1993). For this study, only the south-central populations were surveyed. 

Ruellia humilis, a partial sun-loving species, is found chiefly in one site and is abundant 
there. The site is a privately owned, active limestone quarry (approximately 2 km? in area) 
in Franklin County (same location as R. strepens population number 3, Fig. 1). The quarry 
is semi-forested with Juniperus virginiana L. and Cercis canadensis L. and a non-native 
herbaceous dominant, Bromus imermis Leyss. Klotz and Walck (1993) designate the 
community type for R. humilis as “successional redcedar woodland” (see their 1993 
publication for a detailed description of geologic affinities and community ecology of Ruellia 
in Pennsylvania). This population appears to be relatively stable over time. Some herbarium 
specimens document its existence just outside of this quarry, but there are no records in 
Pennsylvania of its occurrence outside of Franklin County. In preparing this manuscript, 


RUELLIA IN PENNSYLVANIA 57 


Susquehanna River yf" 


| Franklin 


{ eee Adam ¢ | 

S ay 

. : t\ 

2 ee Cree 

= Ww E 
0 20 i Kilometers S 


%* Plants found 
4 No plants found 
? No access to site 


Figure 1. Map of extant Ruwellia strepens localities in south-central Pennsylvania in 2002; not shown: 
Cherry Run (Franklin County), Safe Harbor (Lancaster County), and sites not surveyed by the author 
(see text). 


I learned of a second major population of substantial size at Baker Caverns, Franklin 
County. Unfortunately, because it was discovered after the completion of fieldwork, it was 
not included in this study. Ruellia humilis additionally occurs near the Williamson 
Community Center, but I did not survey this population. 

Ruellia strepens, a floodplain, shade-tolerant species, historically grew along the Conestoga 
River in Lancaster and Dauphin counties, but there is no documentation of its presence in 
Dauphin County in a half-century, and the Lancaster population was eradicated in 2001 by 
lawn mowing (J. Parks, personal communication, 2001). In Franklin County, it occurs on 
the banks and floodplains of the Conococheague Creek and its West Branch, both of which 
have headwaters in Franklin County (in Michaux and Buchanan State Forests) and empty 
into the Potomac River in northern Maryland (Fig. 1). In Cumberland County, it occurs 


58 BARTONIA 


along the Conodoguinet Creek, which begins near the Franklin/Fulton County border on 
the southwestern slopes of Kittatinny Mountain. The Conodoguinet flows east through 
Cumberland County and into the Susquehanna near Harrisburg. The community type for 
R. strepens is “xeric to mesic, calcareous, upland forest” sensu Klotz and Walck (1993). Based 
on specimen data (Rhoads and Block 2003; unpublished data, PH), there is an indication of 
a decline in the range and stability of R. strepens in Pennsylvania. It no longer grows in two 
previously occupied counties, and, based on this survey, no longer occupies many former 
sites in Franklin and Cumberland counties. From this survey, populations of R. strepens vary 
in size from one to approximately 55 individuals, but half of them contain 12 or fewer 
individuals. These smaller populations do not always appear each year (J. Parks, personal 
communication, 2001). Thus, a thorough update of its status is needed. 

After completing my field surveys, I learned of seven new populations of R. strepens 
discovered by others between 2002 and 2003 (L. Klotz and J. Kunsman, personal 
communication, 2003). All seven populations are distributed in Franklin County, southwest 
of Chambersburg, along either the Conococheague Creek (five sites) or the West Branch 
Conococheague (two sites). One of these is among the most southern of R. strepens 
populations in Pennsylvania. It occurs about one minute of longitude north of the Maryland 
border and slightly south of the West Branch junction along the Conococheague. This site 
has been estimated to contain ca. 560 plants, perhaps the largest population in the state. It 
should be a top priority for monitoring and conservation as it may be the ancestral source 
population for other Pennsylvania populations. Threats to this population include logging, 
clearing, and invasive plant spread. The remainder of this paper addresses only the 
populations that I surveyed in 2002 (Table 1). 

Two questions are addressed in this paper: (1) What is the current population status and 
distribution of Ruellia humilis and R. strepens in Pennsylvania? (2) Does this reassessment 
have conservation implications? 


METHODS 


John Kunsman, botanist for the Nature Conservancy in Pennsylvania, provided previous 
field form data for 25 R. strepens sites across Cumberland, Franklin, and Lancaster counties 
and two sites for R. humilis in Franklin County (Table 1). He provided specific locality data 
for all sites except number 10, Cherry Run. Because he was not able to locate the field form 
data for this site, it was subsequently removed from this study. However, it occurs near the 
extreme southwestern border of Franklin County. With its exclusion, the combination of 
sites 13 and 20 due to proximity, and the addition of a newly discovered population 
(number 26, which I term Bernheisel Bridge South), 26 total sites were to be surveyed (24 
R. strepens plus 2 R. humilis populations). Attempts were made to visit all 26 of these sites 
between 23 August and 14 September 2002. Permits were obtained from the Pennsylvania 
Department of Conservation and Natural Resources (D.C.N.R.) to collect voucher 
specimens. A voucher specimen was prepared from all populations except those with very 
few individuals (< 3); these have been deposited at the Academy of Natural Sciences of 
Philadelphia (PH). Between 45 and 90 minutes were spent surveying each site. Grounds were 
walked in a consistent, grid-like manner. Threats to populations were visually assessed and 
populations were designated as having immediate threats, minor threats, or no obvious 
threats. A full report of this survey was submitted to the Pennsylvania Science Office of The 
Nature Conservancy in Middletown. 


Table 1. Recently reported (since 1984) s 


RUELLIA IN PENNSYLVANIA 


59 


outh-central Pennsylvania Ruellia populations that the author 


surveyed or attempted to survey in 2002 
USGS. Approximate 
topographic number of 
Site number, name County quadrangle individuals Current threats to habitat 
R. strepens populations 
1 Baker Caverns Franklin Williamson 0 Mowing 
2 Rockdale Woods Franklin Williamson 0 Exotic plant spread, herbicide 
use 
3 Williamson Quarry Franklin Williamson 0 None 
4 Fort Louden Franklin McConnellsburg Not examined Probably detroyed by fishery 
5 Siberia Franklin Chambersburg 0 None, habitat intact 
6 Licking Creek Franklin Mercersburg None, habitat intact 
7 Martins Mill Bridge Franklin Williamson Not examined _ Private property, but habitat 
appears to be intact 
8 North Welsh Run Franklin Williamson 12 None, habitat intact 
9 Mercersburg Woods Franklin Mercersburg Minor garbage dumping 
10 Licking Creek 2 Franklin Cherry Run Not examined Not examine 
11 Conococheague Franklin Williamson Minor foot traffic 
Bridge 
12 Hampden School Cumberland Harrisburg West 25 
13 Lambs Gap Road/ Cumberland Wertzville 50 Deer browsing 
/20 LGR West 
14 Cave Hill Cumberland Carlisle 0 None, habitat intact 
15 Carlisle Woods Cumberland Carlisle 12 Minor foot traffic 
16 Willow Grove/ Cumberland Plainfield Not examined —_ Not examined 
Opossum Creek 
17 Bloserville Hill Cumberland Plainfield Not examined Not examined 
18 Bridge Road Cumberland Plainfield 1 Foot traffic, c rs 
19 Mountain Road Cumberland Newburg 0 Some exotic plant spread 
(Polygonum cuspi 
21 Howard Lane Cumberland Wertzville 17 None, habitat intact 
22 Bernheisel Bridge Cumberland Wertzville 25 Minor Appalachian Trail foot 
traffic 
23 Sample Bridge Cumberland Wertzville 0 one 
24 Willow’s Mill Cumberland Wertzville 0 Mowing, foot traffic 
25 Owl Bridge Lancaster § Safe Harbor ) Mowing 
26 Bernheisel Bridge Cumberland Wertzville 10 Minor Appalachian Trail foot 
Sout tr affic 
umilis populations 
27 Valley Quarry Franklin Williamson None, habitat intact 
28 Johnston Run Franklin Mercersburg Not examined = Not examine 


In an attempt to identify the consequences of rarity in Pennsylvania Ruellia, while field- 
surveying R. strepens populations, samples were gathered for use in an allozyme-based study 
of genetic diversity. Many enzyme systems were assayed, following stain recipes from 
Wendel and Weeden (1989). 


60 BARTONIA 


RESULTS 


Because this project was initiated late in the summer, R. strepens was not seen in open 
flower; it blooms primarily between May and July. Some cleistogamous flowers were visible 
in late August persisting on the beaks or distal portions of the capsules. Ruellia humilis 
produced chasmogamous and cleistogamous flowers concurrently through mid-September. 
The open flowers were visited by small fritillary butterflies as well as large and small bees, 
the smaller of which seemed to be less effective pollinators because they were able to bypass 
the anthers and stigma lobes to access the nectar (personal observation, 2002). All flowering 
had ceased in this species by 29 September 2002 at site 27. 

Twenty-one of the 26 sites were surveyed (Table 1). The remaining five sites, numbers 4, 
7, 16, 17, and 28, could not be surveyed because they were on private property and heavily 
posted, and the owners could not be located. Site 4 was owned by Mt. Parnell Fisheries. The 
manager would not grant permission for me to access his site because he was concerned any 
rare plant finds might deny him the opportunity to expand his fisheries in the future. 
Moreover, he noted that the habitat for the historic R. strepens population there had since 
been converted to a fishery. 

Of the 21 sites surveyed, plants were found at 11 sites (Table 1). An estimated total of 209 
R. strepens and 750 R. humilis individuals in Pennsylvania was observed. Thus, of the 26 total 
historic sites, population reassessment was not permitted at 6 sites (22%), extant populations 
were found at 11 sites (41%), and no plants were found at 10 sites (37%). Of the 21 sites 
successfully surveyed, 7 were under immediate levels of threat from a combination of deer 
browsing and habitat degradation (Table 1). Six other sites were moderately threatened by 
the above factors and the remaining eight were under no obvious threat to population vigor. 
Fifteen of the 26 sites were on private property and 11 were on public lands or lands not 
posted as private property. 

Results from the allozyne-based study were inconclusive. No banding activity occurred 
in many assayed enzyme systems (AAT, ACO, ADH, CAT, GDH, HEX, IDH, ME, and 
RBC). Some systems showed activity at least once (GPI, MPI, PGD, PGM, SKD, and SOD), 
but bands were often faint and a definitive difference in enzyme mobility was difficult to 
detect (GPI, MPI, and SKD). When banding patterns were clear, PGM and SOD were 
consistently invariable, though PGD did show some variation. It is suspected that enzymatic 
activity was low in these samples due to prolonged freezer storage (-80° C for up to seven 
months) after field collection. Efforts are now underway to assess genetic diversity and gene 
migration patterns in these populations using amplified fragment length polymorphisms 
(AFLP), a technique that can utilize dried plant material and may detect variation more 
readily than allozymes. 


DISCUSSION 


Nearly half of the historic populations of Ruellia strepens are no longer in existence. These 
data reflect a decrease in the number of extant R. strepens populations in Pennsylvania. Most 
of this decrease is likely attributable to a combination of deer browsing and habitat 
degradation including mowing, foot traffic, exotic plant spread, and various types of 
development. Possible inaccuracies in these population decrease estimates include the 
surveyor failing to find plants at the 10 non-extant sites, the lack of plants at the time of 


RUELLIA IN PENNSYLVANIA 61 


survey, and the unknown status of populations in the western part of the state (Greene and 
Washington counties). It is possible these populations may have died back before the survey 
or, as James Parks noted, they may not have appeared this season. No management strategies 
were developed for either species because most of the sites were privately owned. Though 
there is no evidence of large populations persisting on the inaccessible, privately owned sites, 
it would be worthwhile to attempt to contact the owners and look for plants. Most private 
property and neighboring-property owners of Ruellia sites who were contacted for this 
study were not aware of an endangered or threatened plant on their land. An attempt was 
made to inform them of this by showing them pressed specimens of Ruellia. 

Recommendations to P.N.H.P. were made to change the status of R. strepens from a 
threatened to an endangered plant in Pennsylvania. The one R. humilis population on the 
limestone quarry site is under no immediate threat, as the plants are widespread across the 
glade and active mining is minimal and localized. However, future mining expansion could 
dramatically alter the current stability of the major Pennsylvania population. Ruellia humilis 
- is already listed as an endangered species and no recommendations for status change were 
made. The changes in the populations of R. strepens in Pennsylvania indicated by this field 
study affirm the importance of periodically re-evaluating rare species, both for the benefit 
of the species and the credibility of state and federal endangered species programs. However, 
because I learned of additional, substantial R. humilis and R. strepens populations after 
preparing this manuscript, P.N.H.P. might choose to reconsider my recommended status 
change for R. strepens. 

Identifying both the causes (e.g., natural or anthropogenic) and consequences (e.g., loss 
of genetic diversity) of rarity in plants is necessary for properly conserving biological 
diversity (Fiedler 2001). Species of Ruellia may be both naturally rare and further diminished 
by human disturbance. Rabinowitz et al. (1981) identified seven forms of rarity in plants, 
including plants that are broadly distributed but never abundant, narrowly distributed but 
abundant where found, and narrowly distributed and never abundant. Rwellia humilis and 
R. strepens are currently widespread across the eastern United States. Because species of 
Ruellia are relatively long-lived (at least a decade), have very high seed viability, and can 
mature in a single growing season (B. Lamack, personal communication, 2003), it may be 
that they have narrow ecological or habitat affinities that contribute to their rarity in 
particular areas. The aim of my ongoing research is to better understand these factors as well 
as the evolutionary history and flora diversification of this remarkable genus. 


ACKNOWLEDGMENTS 


I thank Tim Block and Ann Rhoads (Morris Arboretum of the University of Pennsylva- 
nia) for their support and sharing their knowledge of the state flora. Special thanks to 
Lucinda McDade (Academy of Natural Sciences of Philadelphia) for her mentoring and 
expertise on Acanthaceae. Larry Klotz (Shippensburg University) and an anonymous 
reviewer offered helpful comments to improve this manuscript. Chris Spolsky and Thomas 
Uzzell (Academy of Natural Sciences) welcomed my participation in their laboratory. John 
Kunsman (The Nature Conservancy) provided population locality data. Bill Lamack 
(Bowman’s Hill Wildflower Preserve) provided information on plant viability and longevity. 
The late James C. Parks (Millersville University) contributed information on the natural 
history of local Ruellia populations. Funding was provided by the Morris Arboretum and 
the Academy of Natural Sciences of Philadelphia. 


62 BARTONIA 


REFERENCES CITED 


Daniel, T. F. 1984. The Acanthaceae of the southwestern United States. Desert Plants 5: 162-179. 

Ezeurra, C. 1993. SRN: sf Ruellia (Acanthaceae) in southern South America. Annals of the 
Missouri Botanical Garden 80: 787-845. 

Fiedler, P. L. 2001. Rarity in ae plants. Pp. 2-3 in J. P. Smith, K. Berg, and L. May (eds.), 
Inventory of Rare and Endangered Vascular Plants of California, California Native Plant Society 
Special Publication No. 1, 6th edition, Sacramento, 336 pp. 

Klotz, L. H. and J. L. Walck. 1993. Rare vascular plants associated with limestone in southwestern 
Franklin County, Pennsylvania. Bartonia 57 Supplement: 16-41. 

Long, R. W. 1977. Artificial induction of obligate sett in species-hybrids in Rwellia 
(Acanthaceae). Bulletin of the Torrey Botanical Club 1 -56. 

Lord, E. M. 1981. Cleistogamy: a tool for the study of floral cease function, and evolution. 
Botanical Review 47: 421-449. 

Manktelow, M. 2000. The filament curtain: a structure important to systematics and pollination 
biology in the Acanthaceae. Botanical Journal of the Linnaean Society 133: 129-160 

McDade, L. A., S. E. Masta, M. L. Moody, and E. Waters. 2000. Phylogenetic relationships among 
Acanthaceae: evidence from two genomes. Systematic Botany 25: 106-121. 

NatureServe. 2003. NatureServe Explorer web site. NatureServe, Arlington, Virginia. 
www.natureserve.org/explorer/ (accessed 29 September 2003). 

Rabinowitz, D., $. Cairns and T. Dillon. 1981. Seven forms of rarity. fe 182-204 in M. Soul€é (ed.), 
The Science of Scarcity and Diversity. Sinauer. Sunderland, Massachuse 

Rhoads, A. F. and T. A. Block. 2000. The Plants of Pennsylvania: an ‘Bsn Manual. University of 
Pennsylvania Press, Philadelphia. 1,061 pp. 

Rhoads, A. F. and T. A. Block. 2003. The flora of Pennsylvania database. Pennsylvania Flora Project, 
Morris Arboretum, University of Pennsylvania, Philadelphia. www.paflora.org (accessed 29 
September 2003). 

, A. F. and W. M. Klein. 1993. The Vascular Flora of Pennsylvania: Annotated Checklist and 
Atlas. American Philosophical Society, Philadelphia. 636 pp. 
io B. C. and F. A. Barkley. 1949. The genus Ruellia in Texas. American Midland Naturalist 42: 
-86. 


Wende, J. F. and N. F. Weeden. 1989. Visualization and interpretation of plant isozymes. Pp. 5-45 
in D.E. Soltis and P.S. Soltis (eds.), sozymes in Plant Biology, Dioscorides Press, Portland, Oregon. 


Bartonia No. 62: 63-93, 2004 


An Ethnobotanical and Medical Research Literature Update 
on the Plant Species Collected in the Lewis and Clark 
Expedition of 1803-1806 


DARRALL HEATON AND ARA DERMARDEROSIAN 
Philadelphia College of Pharmacy, University of the Sciences in arog 
600 South 43rd Street, Philadelphia, Pennsylvania 19104-4 
darrallheaton@comcast.net, a. dermar@usip.edu 


ABSTRACT. We compiled information on the medicinal and pharmacological activity of each of the 
177 Lewis and Clark Herbarium (LCH) plant species, representing 129 genera, from ethnobotanical 
and current biomedical research literature. Medicinal uses by Native American groups have been 
documented for 123 species (approximately 69%) in 83 genera. We found research studies reporting 
pharmacological activity for only 26 (15%) of the species, although we also found reports of 
pharmacological activity for other species worldwide in 80 of the genera | in the LCH. The 
disparity between the number of LCH species used in traditional American Indian medicine and the 
number that have undergone modern pharmacological evaluation cedic a future research may 
turn up additional compounds of use to modern medicine from Lewis and Clark’s discoveries. 


INTRODUCTION 


Woolly mammoths, Peruvian llamas, blue-eyed, Welsh-speaking Indians — depictions of 
the land, creatures, and native peoples in the West often came from the imaginations of men 
who had never been there. Reports told of western terrain spotted with wondrous creatures: 
unicorns, seven-foot-tall beavers, and friendly, slim-waisted buffalo. In 1803, such myths 
defined the uncharted West; however, the Lewis and Clark expedition later dispelled such 
speculations (Burns 2004). 

Thomas Jefferson took the oath of office as the third President of the United States on 
March 4, 1801. Even before Jefferson became president, he dreamed of exploring the land 
west of the Mississippi but his previous attempts to organize an expedition failed. After 
Jefferson was elected president, however, he was even more convinced that the future of the 
United States was tied to an expansion to the West. 

On February 28, 1803, President Jefferson won approval for a congressional grant of 
$2,500 to fund an expeditionary group. Founded as an interdisciplinary effort, it would 
include commercial, geographical, political, and scientific interests. Meriwether Lewis and 
William Clark would lead the expedition, and their mission was to explore the unknown 


West. 
Submitted 3 January 2000; revised 12 January 2001, 29 January 2003, 20 October 2004 


63 


64 BARTONIA 


On the 6,000-mile expedition, the explorers mapped and described mountains, lakes, and 
rivers and documented 80 plant species that were sources of botanicals then unknown to 
science. Lewis and Clark collected, described, packed and sent East plant, animal, and 
mineral specimens, including over 200 plant specimens. This collection is now in the Lewis 
and Clark Herbarium at the Academy of Natural Sciences of Philadelphia (Johnston 1998; 
McCourt and Schuyler 2000). 

Of the $2,500 allocated for the entire expedition, Lewis budgeted $55 for medicine and 
$696 for presents to give to Indians along the way. As the journey unfolded, however, the 
medicine was more important than the presents in securing beneficial relationships with the 
northwest natives (Loge 1996). Overall, however, Johnston (1998) believes that the Lewis 
and Clark expedition and botanists of the Western world owe a deep debt to those Native 
American tribes who shared their wealth of knowledge with the explorers, in some cases 
saving them from starvation and illness (Johnston 1998). 

Reveal et al. (1999) document a total of 226 plant specimens in the Lewis and Clark 
Herbarium (LCH). They represent 177 plant species in 129 genera (Table 1). The purpose 
of this study is to provide answers to the following questions for each of the LCH plant 
species. What type of pharmacological activity is associated with each of the LCH plant 
species? Is the medicinal use of a plant supported by modern medical research and 
ethnobotanical data? How many of the plants are considered poisonous or toxic? How many 
of the plants lack documented modern medical research? 


METHODS 


Moerman’s (1998) landmark book, Native American Ethnobotany, was our primary 
resource for the ethnobotanical information. It documents Native American use of 4,029 
plants; more than half were used medicinally. The breadth of Native American plant 
knowledge is shown by plant uses for food, fiber, dye, ceremonial and magical items, 
cleaning agents, containers, fertilizers, fuels, incense and fragrance, insecticides, jewelry, 
lubricants, musical instruments, preservatives, smoking, soap, waterproofing, tools, toys, and 
weapons. For each of the 177 LCH plant species, we indicated which of Moerman’s 79 
categories, from abortifacient to witchcraft medicine, describe its medicinal use and tallied 
the number of tribal groups for which each use has been documented. 

Our source for research studies reporting pharmacological activity was the biomedical 
literature search system PubMed. It contains over 12 million citations from approximately 
4,800 biomedical journals published in the United States and 70 other countries (United 
States Library of Medicine 2004). 


RESULTS 


Medicinal uses by Native American groups have been documented for 123 species 
(approximately 69%) in 83 genera (Table 1). We found research studies reporting 
pharmacological activity for only about 28 (16%) of the species (Table 1), although we also 
found reports of pharmacological activity for other species worldwide in 80 of the genera 
represented in the LCH 

Antibacterial activity is present in Alnus rubra, Arctostaphylos uva-ursi, Balsamorhiza 
sagittata, Ceanothus velutinus, and Veratrum californicum. Antifungal compounds have been 
found in Alnus rubra, Artemisia ludoviciana, and Balsamorhiza sagittata. Artemisia ludoviciana 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 65 


has anti-malarial properties, and research has shown anti-viral capacity in Amelanchier 
alnifolia, Amorpha fruticosa, Ipomopsis aggregata, Shepherdia argentea, and Vaccinium myrtillus. 

Anti-inflammatory properties have been discovered in Achillea millefolium, Arctostaphylos 
uva-ursi, Artemisia ludoviciana, Juniperus communis, and Matricaria matricarioides. Anticancer 
activity has been shown for extracts of Amorpha fruticosa, Ipomopsis aggregata, Iris 
missouriensis, Liatris py hya, Maclura pomifera, Oenothera cespitosa, Polanisia dodecandra, 
Rhus trilobata, and Vaccinium myrtillus. Compounds in Artemisia dracunculus inhibit certain 
immune responses while substances in Euphorbia marginata stimulate others. 

LCH species with other pharmacological effects include Achillea millefolium (used to treat 
gastrointestinal diseases; antispermatogenic), Alnus rubra (promotes glucose metabolism), 
Arctostaphylos uva-ursi (used to treat urinary tract diseases; inhibits melanin biosynthesis), 
Equisetum arvense (used to prevent and treat kidney stones), and Vaccinium myrtillus (used 
to treat ulcers; used for treatment of ischaemia reperfusion injury; reduces LDL-cholesterol 
levels; affects night-vision). 

Toxicity has been observed in Equisetum arvense (ingestion can cause dermatitis), 
Euphorbia marginata (sap causes contact dermatitis, conjunctivitis, and keratitis), Pinus 
ponderosa (toxic to livestock), Rubus spectabilis (causes epidermal necrolysis when ingested), 
and Veratrum californicum (teratogenic, i.e., causes limb deformities in fetuses when ingested 
by pregnant females). 


ee 


DISCUSSION 


Prescription drugs prolong and improve the quality of life. They also frequently reduce 
or replace more expensive forms of medical treatment such as hospitalization, nursing care, 
and surgery. With the great potential for continued pharmaceutical breakthroughs, 
Prescription drugs will continue to play an important role in containing costs, even as 
overall healthcare expenditures increase (Balick and Cox 1996). According to Pharmaceutical 
Research and Manufacturers of America (Anonymous 2004), it typically takes 10 to15 years 
and over $800 million to advance a potential new medicine from a research idea to a 
treatment approved by the Food and Drug Administration. In the past, the discovery of a 
drug was a process of trial and error and serendipitous discovery; it has now become more 
systematic through the use of increasingly sophisticated technology (Anonymous 2004). 
However, does discovery of a drug always rest only on “solid” science, such as structural 
chemistry and pharmacology, or can traditional usage, woods lore, and knowledge handed 
down over many generations play a role (Balick and Cox 1996)? We favor a rational 
combined approach utilizing both ethnobotanical data and the results of modern laboratory 
and clinical research as appropriate means to identify areas of research potential. _ 

This paper is the first medical literature summary for the 177 species of the Lewis and 
Clark Herbarium collection. If you were to walk into any pharmacy in the United States, 
Canada, or Western Europe and pick any prescription medicine at random, there is a one 
in four chance that the medicine you choose has an active ingredient derived from a plant 
(Balick and Cox 1996). Most of these plant-derived drugs were originally discovered through 
the study of the folk knowledge and traditional cures of indigenous peoples — much the 
same ethnobotanical approach (Balick and Cox 1996) practiced in 1803 to 1806 by Lewis and 
Clark in the American West. o 

Approximately one-fifth of the LCH species used medicinally by Native Americans have 
been subjected to modern research methods and found to have pharmacological activity. The 


66 BARTONIA 


disparity between the number of LCH species used in traditional American Indian medicine 
and the number that have undergone modern pharmacological evaluation suggests that 
future research may turn up additional compounds of use to modern medicine from Lewis 
and Clark’s discoveries. 


REFERENCES CITED 


ANONYMOUS. 2004. entice Industry Profile. Pharmaceutical Research and Manufacturers of 
America, Washington, D.C. 61 pp. 

BALICK, M. J. AND P. A. Cox. eens Plants, People, and Culture: The Science of Ethnobotany. W. H. 
Freeman, New York. 228 pp. 

BURNS, K. 2004. Lewis and Clark, the journey of the corps of discovery. www.pbs.org/lewisandclark 
(accessed 10 October 2004). 

JOHNSTON, B. A. 1998. Botanical discoveries of Lewis and Clark. HerbalGram 44: 30-32, 49-51. 

LOGE, R. V. 1996. Two dozes of barks and opium: Lewis and Clark as physicians. Pharos Alpha Omega 
Alpha Honor Medical Society 59: 26-31. 

McCourt, R. M. AND A. E. SCHUYLER. 2004. The Lewis and Clark herbarium. www.acnatsci.org/ 
lewis&clark/index.html (accessed 10 October 2004). 

MOERMAN, D. E. 1998. Native American Ethnobotany. Timber Press, Portland, Oregon. 927 pp. 

REVEAL, J. L., G. E. MOULTON AND A. E. SCHUYLER. 1999. The Lewis and Clark collections of 
vascular plants: names, types, and comments. Proceedings of the Academy of Natural Sciences of 
Philadelphia 149: 1-64. 

SPAMER, E. E. AND R. M. MCCourRT. 2002. The Lewis and Clark Herbarium, Academy of Natural 
Sciences of Philadelphia (PH-LC): Digital imagery study set. Academy of Natural Sciences of 
Philadelphia Special Ans oe 19. CD-ROM. 

UNITED STATES LIBRA F MEDICINE. 2004. Scientific research: PubMed www.ncbi.nlm.nih.gov/ 
entrez/query.fcgi rae 06 October 2004). 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 67 


Table 1. Ethnobotanical (Moerman 1998) and biomedical information on the Lewis and Clark 
Herbarium (LCH) plant species. Subjects of biomedical studies were dervied from research 
publications and abstracts compiled using the PubMed biomedical literature search system; primary 
sources are numbered (see endnotes on pages 89-93). Plant nomenclature follows Spamer and 
McCourt (2002). 


Uses by Native Americans: number of tribal Subject matter of selected biomedical 
LCH plant species groups for which use is documented studies 


Acer circinatum Pursh — Antidiarrheal, love medicine, miscellaneous * 
disease remedy (treatment for polio): 1 


% 


Acer macrophyllum Dermatological aid, tonic, tuberculosis remedy: 
rsh 1 


— millefolium L. Anthelmintic, anticonvulsive, antirh icused Isolation of anti-inflammatory 
nulosa (Nutt.) internally, breast treatment, ceremonial medi- _ principles! 
eg cine, dietary aid, diuretic, ear medicine, emetic, 


ies ag of chronic hyposecretory 


dicine, h h , h 
heart medicine, hemorrhoid remedy, hemostat, f° EA ECAR TO 
on angiocholitis* 


internal medicine, Staite poultice, sedative, 
witchcraft medicin 


: : eee 
Antispermatogenic effect in mice 
Antiemetic, antihemorrhagic, burn dressing, ca- tispermatogenic eff 


thartic, disinfectant, herbal steam, other, repro- 
ductive aid, snakebite remedy, unspecified, 
urinary aid, venereal aid, veterinary aid: 2 


pei medicine, cough medicine, in a aid, 
imulant, tonic, tuberculosis remedy: 


Kidney aid, liver aid: 4 
Diaphoretic: 5 
Antidiarrheal, eye medicine, miscellaneous 


disease remedy (treatments for grippe, mumps, 
influenza and the flu), orthopedic aid, pediatric 
aid: 6 


Gynecological aid, panacea, throat aid, tooth- 
ache remedy: 7 


Antirheumatic used externally: 8 
Gastrointestinal aid: 9 
Febrifuge: 10 

Analgesic: 14 

Cold remedy: 17 
Dermatological aid: 27 


Allium geyeri $. Watson Used as food. Not utilized as a drug. 


* Non-LCH species in the genus investigated and found to have Lpheneenes properties. 
+ No pharmacological study was found of any species in the ge 


68 


Table 1 (cont’d) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Alnus rubra Bong. 


Amelanchier — 

a Nutt. ¢ 

er var. mine 
k) Cc 


nfl se 


Amphora fruticosa L. 


Ampelopsis cordata 
Michx. 


Amsinckia menziesii 
ore A. Nelson & 
. Macbr. var 
retrorsa apa Reveal 
& Schuy] 


Anemone canadensis L. 


Anemone piperi Britt. 
ex Rydb. 


Lenore cold remedy, cough medi- 
cine, emetic, internal medicine, miscellaneous 
disease ak pane. for internal ail- 

ments), respiratory aid, toothache remedy: 1 


Antidiarrheal, cathartic, orthopedic aid, pul- 
onary aid, tonic: 2 

Gastrointestinal aid: 3 

Tuberculosis remedy, unspecified: 4 

Analgesic: 5 

Dermatological aid: 8 


Venereal aid: 1 


Primarily used for fiber. Other uses include 
hunting and fishing items and cooking tools. 
Not utilized as a 


Urinary aid: 1 


Analgesic, anthelmintic, ceremonial medicine, 
dermatological aid, eye medicine, saci or- 
thopedic aid, throat aid, witchcraft medicin 


Panacea: 2 


0 


Contains enzymes of glucose meta- 
olism* 


Antibacterial activity® 


fig meer against nine 
fungal sp 

According to interviews with two 
elder Salishan women, the bark 
was used as medicine for digestive 
tract porn 


Extracts acuiye against an enteric 
coronavirus* 


Strong cera effects on Ep- 
stein-Barr v 

Active antitumor compounds, 
which include eight cytotoxic 


isoflavones"! 


bs 


* Plant toxins (alkaloids) in 
Amsinckia pose a toxicity hazard” 


= 


* Non-LCH species in the genus investigated and found to have ig os properties. 
t No pharmacological study was found of any species in the gen 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 69 


Subject matter of selected biomedi- 
cal studies 


Angelica arguta Nutt. 
in Torr. & A. Gray 


Arbutus menziesii 
Pursh 


Arctostaphylos uva-urst 
(L.) Spreng. 


Antidiarrheal, antiemetic, antihelmintic, carmi- 
native, cold remedy, eye medicine, febrifuge, 
gynecological aid, love medicine, pediatric aid, 
psychological aid, respiratory aid, sedative, 
snakebite remedy, venereal aid: 1 


Dermatological aid, orthopedic aid: 2 
Analgesic, gastrointestinal aid: 4 
Burn dressing, ceremonial medicine, dietary 


aid, love medicine, miscellaneous disease reme- 
dy (treatment for diabetes), veterinary aid: 


Emetic: 2 
Cold remedy, throat aid: 3 


Dermatological aid, gastrointestinal aid: 4 


a adjuvant, antidiarrheal, antirheu- 


tic used externally, burn dressing, cold 
iy cough medicine, dietary aid, diuretic, 
ear medicine, emetic, gynecological aid, hunt- 
ya medicine, laxative, panacea, psychological 


Analgesic, blood medicine, ton medi- 
cine, narcotic, orthopedic aid, tonic: 


Antihemorrhagic, dermatological aid, eye 
medicine, oral aid, pediatric aid: 


Kidney aid, unspecified, urinary aid: 4 


* 


According to interviews with two 
elder Salishan women, respiratory, 
digestive and gynecological prob- 
lems were treated with the bark’ 


Disinfectant action for urolithiasis” 


Antimicrobial activity of extracts 
most active against Escherichia coli 
and Proteus vulgaris“ 


Investigations of iridoid substances, 

disinfective effect on the urinary 

tract’? 

Increases effect on antiallergetic 
anti-i matory activities of 

dexamethasone ointment 

Arbutin, isolated from its leaves, 

may increase the inhibitory action 

of indomethacin ” 

Combined effects of arbutin and 

prednisolone on immuno-inflam- 

mation™ 

Inhibitatory effects on melanin 

biosynthesis” 

Anti-tyrosinase activity” 


(continued) 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


70 
Table 1 (cont'd) 


LCH plant species 


BARTONIA 


Uses by Native Americans: sev ah of tribal 
groups for which use is documen 


Subject matter of selected biomedi- 
cal studies 


Arctostaphylos uva-ursi 
(L.) Spreng, 
(cont'd) 


Argentina anserina 
(Lehm.) L. var. grandis 
(Torr. & A. Gray) 
Rydb. 


Artemisia cana Pursh 


Artemisia dracunculus 
i 


Artemisia frigida Willd. 


Analgesic, diuretic, emetic, pediatric aid: 1 
Antidiarrheal, dermatological aid: 2 


Dermatological aid, dietary aid, tonic: 1 
Unspecified: 2 


Analgesic, antidiarrheal, eye medicine, gastro- 
intestinal aid, stimulant, tonic, unspecified, 
ing 4 aid, veterinary aid, witchcraft medi- 
cine: 


Cold remedy, gynecological aid: 2 
Dermatological aid, pediatric aid: 3 
Antirheumatic used externally: 4 


Analgesic, anticonvulsive, antihemmorhagic, 
cancer treatment, carminative, dermatological 


monary 
losis remedy, unspecified, venereal aid, 
veterinary aid: 1 


Ceremonial medicine, febrifuge, gynecological 
aid: 2 
Hemostat, stimulant, disease 


remedy (treatments fet a mountain fever, diabe- 
tes, flu): 3 


Cold remedy, cough medicine, gastrointestinal 
aid: 4 


Abortifacient: 5 


A field study documents some 
traditional and contemporary 
knowledge of the medicinal use of 
plants by the Carrier people, 
which includes A. uva-ursi” 


Reduced hyperphagia and polydip- 
sia associated with streptozotocin 


* Artemisia species frequently util- 
ized for treating malaria, hepatitis, 
cancer, inflammation and infections 
by fungi, bacteria and virus 


Natural inhibitor of complement™* 


Oil has genotoxic properties” 


Source of fall allergic symptoms, 
particularly in western United 
tates”® 


Field study documents medicinal 
uses of plant by Carrier people, an 

kan-speaking people of 
British Columbia” 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 71 


Table 1 (cont’d) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Artemisia longifolia 
Nutt. 


Artemisia ludoviciana 
Nutt. var. latiloba 
utt. 


Aster eatonii (A. Gray) 
Howell 


Aster oblongifolius 
Nutt. 


Astragalus canadensis L. 


Astragalus missouriensis 
Nutt. 


Atriplex canescens 
(Pursh) Nutt. 


Analgesic, antidiarrheal, antirheumatic used 
minative, cathartic, ceremonial 


influenza), nose 
medicine, orthopedic aid, other, pediatric aid, 
pulmonary aid, strengthener, tuberculosis 
remedy: 1 


Disinfectant, eye medicine, gastrointestinal aid, 
panacea, respiratory aid: 


Throat aid, cold remedy: 3 
Veterinary aid: 4 
Unspecified: 5 


Dermatological aid: 8 


Oo 


Witchcraft medicine: 1 
Analgesic, cough medicine, dermatological aid, 
ebrifuge: 1 


Antihemorrhagic, pediatric aid, pulmonary 
aid: 2 


Analgesic, cathartic, cough medicine, hunting 
miscellaneous disease remedy (treat- 


medicine, Anat stimulant, 
toothache remedy, veterinary ai 


Ceremonial medicine, emetic, gastrointestinal 
aid: 2 


Dermatological aid: 5 


Ethanolic leaf extracts exhibit anti- 
inflammatory activit 


Antimalaric effect of an alcoholic 
extract in a rodent malaria model” 


Antifungal activity® 


* Lysosomal storage diseases in- 
by ingestion of various 
Astragalus species” 


* Astragalus, containing the toxin 

swainsonine, causes severe adverse 
effects on podantive function in 
livestock*? 


Phytochemical studies of nucleo- 
tide sequence of CDNA” 


+ Excretion of selenium (in 
Atriplex species) via milk is impor- 
tant in the deficiency state, but 
when in — may cause toxicity 
to offsprin 


* Non-LCH species in the genus investigated and found to have Aion ime a properties. 
t No pharmacological study was found of any species in the ge 


72 BARTONIA 


Table 1 (cont’d) 


LCH plant species Uses by Native Americans: number of tribal Subject matter of selected biomedi- 
groups for which use is documented cal studies 

Atriplex nuttallii S. Used as food. Not utilized as a drug. 

Wats. 

Balsamorhiza sagittata Antidiarrheal, cathartic, cold remedy, dietary Antibacterial thiophene isolated™* 

(Pursh) Nutt. aid, eye medicine, febrifuge, gynecological aid, oF ” 
hemostat, oral aid, urinary aid, panacea, pul- Root extracts exhibited antifungal 
monary sedative, throat aid, toothache EY 
remedy: 
Antirheumatic used internally, burn dressing, 
diaphoretic, disinfectant, gastrointestinal aid, 
tuberculosis remedy, venereal aid: 
Analgesic: 4 
Dermatological aid: 7 

Bazzania trilobata (L.) | Used asa dye. Not utilized as a drug. Antifungal activity® 

S.F. Gray 

Berberis aquifolium Q * Berbamine, a constituent of 

rsh Berberis species, exhibits leuko- 


genic, anti-arrhythmic, chs ae 
tensive, anti-inflammatory and anti 
tumor activity*®”8 

Berberis nervosa Pursh 0 

Blechnum spicant (L.) Antidiarrheal, cancer treatment, dermatologi- : 

Sm. 


cal aid, orthopedic aid, panacea, pulmonary 
aid: 1 


Gastrointestinal aid: 2 


Calochortus elegans 0 t 
Pursh 
Calypso bulbosa L. Anticonvulsive: 1 t 
Oakes var. occidentalis 
(Holz.) Boivin 

Sia Qua. Gynecological aid: 1 ef 
(Pursh) Greene 
Camassonia subacaulis 0 + 


(Pursh) Raven 
Cardamine nuttallu 0 * 


Greene 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS vo 


Subject matter of selected biomedi- 
cal studies 


Ceanothus sanguineus 
Pursh 


Ceanothus velutinus 
Dougl. ex Hoo 


Cerastium arvense L. 
Chrysothamnus viscidi- 
florus (Hook.) Nutt. 
ssp. viscidiflorus 
Cirsium edule Nutt. 
Clarkia pulchella Pursh 
Claytonia lanceolata 
Pursh 

Claytonia parviflora 
Douglas ex Hook. 


Claytonia perfoliata 
Donn ex Willd. 


Claytonia siberica L. 


Clematis hirsutissima 
Pursh 


Cleome serrulata Pursh 


Burn dressing, dermatological aid: 1 


Antidiarrheal, antirheumatic used externally, 
antirheumatic used internally, cancer treat- 
ment, ceremonial medicine, cough medicine, 
dietary aid, febrifuge, miscellaneous disease 
remedy, other, panacea, pediatric aid, venereal 
aid: 1 


Analgesic, dermatological aid, orthopedic aid, 
unspecified: 2 
Dermatological aid, gynecological aid: 1 
Antirheumatic used externally, cold remedy, 
cough medicine, dermatological aid, diaphoret- 
ic, miscellaneous disease nee (treatment for 
influenza), toothache remedy: 
Used as food. Not utilized as a drug. 
0 
Used as food. Not utilized as a drug. 
Used as food and in toys and games. Not 

a drug. 


utilized as a 


Analgesic, antirheumatic used externally, eye 
medicine: 1 


Eye medicine, gynecological aid, tonic, urinary 
aid, venereal aid: 1 


Throat aid: 2 
Dermatological aid: 4 


Respiratory aid, veterinary aid, witchcraft 
medicine: 1 


Analgesic: 2 


Blood medicine, ceremonial medicine, derma- 
* gical aid, eye medicine, febrifuge, gastroin- 
estinal aid, ce aid: 1 


* Alkaloids — from 
Ceanothus species 


Antimicrobial properties” 


—- 


T 


Blistering agent isolated and used 
by the Nez Perce and Teton Sioux 
nations as a horse stimulant*** 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
t No pharmacological study was found of any species in the gen 


74 
Table 1 (cont'd) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Collinsia seers 
dl. var. grandiflora 

(Lindl.) iota & 

Krause 

Collomia linearis Nutt. 

Coreopsis tinctoria 


Nutt. var. atkinsoniana 


(Dougl. ex Lindl.) 
.M. Parker 


Cornus canadensis L. 


Crataegus douglasti 
Lindl. 


Dalea purpurea Vent. 
Dasiphora fruticosa (L.) 
Rydb. 

Delphinium menziesii 


C. var. pyramidale 
(Ewan) C.L. Hitchce. 


Dodecatheon poeticum 
L.F. Hend. 


Dryopteris carthusiana 
(Vill.) H.P. Fuchs 


\gregia menziesii 
a urn.) Aresch. 


Dermatological aid, veterinary aid: 1 


Dermatological aid: 1 


Antidiarrheal: 1 


Cathartic, dermatological aid, febrifuge, gastro- 
intestinal aid, gynecological aid, orthopedic 
aid, pediatric aid, tuberculosis remedy, unspec- 
ified: 1 


Anticonvulsive, cold remedy, eye medicine, 
tonic: 2 


Analgesic: 3 

Antirheumatic, oral aid, panacea, pediatric aid: 
1 

Antidiarrheal, gastrointestinal aid: 2 
Dermatological aid: 3 

Dermatological aid: 1 


0 


Dermatological aid, love medicine, poison, 
pecified: 1 


Antidote: 1 


Used as food, fertilizer, and for hunting and 
fishing. Not utilized as a drug. 


t 


* Alcohol esters from Coreposis 
species and synthetic derivatives 


examined as lipoxygenase inhibi- 
tors and as LDL (low density 
lipoprotein) ing agents” 


* Crataegus extract is used in cardi- 
ology in Germany for the treat- 
ment a mild to moderate heart 
failure 


* Larkspurs (Delphinium spp.) are 

toxic plants that contain numerous 
diterpenoid alkaloids and are asso- 
ciated with poisoning in livestock“ 


+ 


* Non-LCH species in the genus investigated and found to have — properties. 
+ No pharmacological study was found of any species in the gen 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 75 


Subject matter of selected biomedi- 
cal studies 


Eleagnus commutata 
Bernh. ex Rydb 


Equisetum arvense L. 


Ericameria nauseosa 
(Pall. ex Pursh) G.L. 
Nesom & Baird 


Ericameria nauseosa 
(Pall. ex Pursh) G.L. 
Nesom & Baird var. 
graveolens (Nutt.) 
Reveal & Schuyler 
Erigeron compositus 
Pursh 

Eriophyllum lanatum 
(Pursh) Forbes var. 
lanatum 


Erysimum capitatum 


Erythronium grandi- 
florum Pursh 


Euphorbia cyathophora 
Murr. 


Dermatological aid, pediatric aid, venereal aid: 
1 


Blood medicine, laxative, pediatric aid, stimu- 
lant, toothache remedy, venereal aid: 1 


Analgesic, antirheumatic used internally, di- 
uretic: 2 


Kidney aid, urinary aid, veterinary aid: 3 
Orthopedic aid: 4 
Dermatological aid: 5 


Dermatological aid, orthopedic aid: 1 


Dermatological aid, love medicine: 1 


Analgesic, antirheumatic used externally, 
ceremonial medicine, gynecological aid, respi- 
ratory aid, toothache remedy, tuberculosis 
remedy: 1 


Emetic: 2 


Dermatological aid, cold remedy: 1 


0 


Sterols from E. arvense® 


ona 2 sg to prevent 
and trea ne formation 
with urolithiasis risk factors (citrat- 
uria, calciuria, phosphaturia, pH, 
and diuresis)” 


When mixed with a cholesterol 

diet at 4%, caused —— at the 

neck, head and back in about 
20-60% of rats* 


Seborrhoeic a. induced by 
nicotine of horsetails” 


T 


Contains 2- = Sra api Soa buty- 
rolactone, which is 

Native American food and medici- 
nal plants* 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


76 
Table 1 (cont'd) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Euphorbia marginata 
Pursh 


Festuca idahoensis 
Elmer 


Frangula purshiana 
(DC.) Cooper 


Frasera fastigiata 
(Pursh) Heller 


Fritillaria affinis 
Gehl & Schultes f.) 
Fritillaria pudica 
(Pursh) Spreng. 


Gaillardia aristata 
Pursh 


earache used externally, gynecological 
aid, poison: 1 


Used for fiber. Not utilized as a drug. 


Adjuvant, analgesic, ee antihel- 
mintic, antirheumatic us rnally, blood 
medicine, disinfectant, “et rue tonic, unspeci- 

fied: 1 


Emetic, panacea, poison, venereal aid: 2 
Dermatological aid: 3 


Gastrointestinal aid: 5 


Cathartic: 7 
Laxative: 21 
@) 


Used as food, ar piper and seasonal indica- 
tor. Not utilized 


Breast treatment, cancer ea dermato- 
logical aid, eye medicine, gastrointestinal aid, 
aid, miscellaneous disease pane 
(treatment for mumps), nose medicine, tuber- 
culosis remedy, venereal aid, veterinary aid: 1 


Analgesic, orthopedic aid: 2 


Irritant contact dermatitis‘? 


Lectin is strongly mitogenic for 
human T-lymphocytes and induces 
the release of interleukin-1 beta 
and tumor necrosis factor-alpha 
from cultured mononuclear cells*! 


Sap may Cane acute conjunctivitis 
and keratitis 


* 


* Review article: anthranoid laxa- 
tives and ny Foc carcino- 
genic effects® 


* Non-LCH species in the genus investigated and found to have permease properties. 
+ No pharmacological study was found of any species in the gen 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Gaultheria shallon 
Pursh 


Geum triflorum Pursh 
var. ciliatum (Pursh) 
Fassett 


Grindelia squarrosa 
(Pursh) Dunal 


Gutierrezia sarothrae 


(Pursh) Britt. & Rusby 


Hesperostipa comata 
(Trin. & Rupr.) 
Barkworth 


Antidiarrheal, burn dressing, cough <a 
oral aid, other, reproductive aid, tonic 


ee aH gastrointestinal aid, tuber- 
culosis reme 


Blood medicine, eye medicine, gastrointestinal 
aid, stimulant: 1 


Veterinary aid: 2 
Abortifacient, analgesic, antihemorrhagic, 


emetic, gynecological aid, liver aid, miscella- 
neous disease remedy, orthopedic aid, pediatric 
1 


ee: eye medicine, kidney aid, respira- 
tory aid, tuberculosis remedy, urinary aid, and 
veterinary ai 


Cold remedy, dermatological aid, disinfectant, 

gastrointestinal aid, pulmonary aid, venereal 
aid: 3 

Cough medicine: 5 


Antidote, cathartic, , cough medicine, diuretic, 


tive, strengthener, venereal aid, vertigo medi- 
cine: 1 


Antidiarrheal, cold remedy, diaphoretic, febri- 
fuge, miscellaneous disease remedy (treatment 
- measles and gastric influenza), urinary ai 


Analgesic, antirheumatic used externally, 
ceremonial medicine, gastrointestinal aid, 
gynecological aid, snakebite remedy: 3 


Disinfectant, veterinary aid: 4 
Dermatological aid: 5 


0 


ao 


* Triterpenoid sapogenins in the 
genus Grindelia® 


* Snakeweeds are toxic and cause 
abortions in cattle, sheep, and 
goats™ 


* Diets containing as little as 10% 

snakeweed will induce early em- 
onic toxicosis and abortion in 

Sprague-Dawley rats” 


Isolation of an antitumor protein- 
aceous substance* 


* Non-LCH species in the genus investigated and found to have pecner ioe properties. 
Tt No pharmacological study was found of any species in the genus 


78 
Table 1 (cont’d) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documente 


Subject matter of selected biomedi- 
cal studies 


Holodiscus discolor 
(Pursh) Maxim 


Hordeum jubatum L. 


Hypnum oreganum 
Sull. 


Ipomopsis aggregata 
(Pursh) V. Grant 


Tris missouriensis Nutt. 


Juniperus communis L. 


var. depressa Pursh 


Antidiarrheal, blood medicine, burn dressing, 
dermatological aid, eye medicine, miscella- 
neous disease remedy (infusion of seeds taken 
for smallpox, black measles, and chickenpox), 
oral aid, orthopedic aid, tonic, unspecified: 1 
Miscellaneous disease remedy: 2 


Eye medicine, poison, unspecified: 1 


0 


Blood medicine: 1 


Antirheumatic used externally, burn dressing, 
ceremonial medicine, kidney aid: 


Analgesic, dermatological aid, ear medicine, 
gastrointestinal aid, urinary aid, venereal aid: 2 


Emetic, toothache remedy: 3 


Antirheumatic used externally, ceremonial 
medicine, diaphoretic, disinfectant, emetic, 
heart medicine, herbal steam, hypotensive, 
love medicine, miscellaneous disease remedy 
(taken to prevent flu and used also for treat- 
ment of flu), orthopedic aid, other, pediatric 
aid, sedative, unspecified, urinary aid, venereal 
aid: 1 


Analgesic, antirheumatic used internally, blood 
medicine, cathartic, eye medicine, febrifuge, 
gastrointestinal aid, | eran throat aid, 
toothache remedy: 2 


Antidiarrheal, gynecological aid, pulmonary 
aid: 3 


Dermatological aid, respiratory aid, tuberculo- 
sis remedy: 5 

Cold remedy, kidney aid: 6 

Cough medicine: 7 

Tonic: 8 


Potential anticancer agents” 


An extract demonstrated activity 
against parainfluenza virus type II’ 


Triterpenes, plant anticancer 
agents 


Field study on medicinal uses of 

plant by Carrier people, an Atha- 
party peaking people of British 
Columbia” 


Hiypesiycemic activity of juniper 
berries”® 


Used in Swedish traditional medi- 
cine to treat inflammatory diseases 
and/or wounds exhibiting inhibi- 
tory activity on prostaglandin bio- 
synthesis and platelet activating 
factor (PAF)# 


* Non-LCH species in the genus investigated and found to have A i a properties. 
+ No pharmacological study was found of any species in 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 79 


Subject matter of selected biomedi- 
al studies 


Juniperus horizontalis 
Moench 


Juniperus scopulorum 
Sarg. 


Koeleria macrantha 


(Ledeb.) J.A. Schultes 


Lewitsia rediviva Pursh 


Lewisia triphylla (S. 
Wats.) B.L. Robins. 


Leymus mollis (Trin.) 
Hara 


Luatris aspera Michx. 


Liatris pycnostachya 
ichx. 


Cold remedy, cough medicine, febrifuge 
gynecological aid, herbal steam, love conden 
sedative, throat aid, veterinary aid: 1 


Ceremonial medicine, kidney aid: 2 
Antidiarrheal, antiemetic, diaphoretic, dietary 
aid, diuretic, heart medicine, herbal steam 


love medicine, poison, sedative, a éa. 
tonic, venereal aid, witchcraft medicine: 1 


Analgesic, cough medicine, disinfectant, kid- 
ney aid, other, panacea, tuberculosis remedy, 
unspecified, urinary aid, veterinary aid: 2 


Antihemorrhagic, gastrointestinal aid, gyneco- 
logical aid: 3 


Dermatological aid: 4 


Antirheumatic used ow Pee ceremonial 
medicine, pulmonary ai 


Febrifuge, miscellaneous disease remedy (treat- 
ment for sugar diabetes, flu and colds, cholera, 
kidney trouble, black measles or chickenpox): 
6 

Cold remedy: 7 


Ceremonial medicine, dermatological aid, 
stimulant: 1 

Blood medicine, breast treatment, dermatologi- 
cal aid, heart medicine, miscellaneous disease 
remedy (treatment for diabetes), pulmonary 
aid, throat aid, witchcraft medicine: 1 
Gynecological aid: 2 


0 


Strengthener: 1 
Unspecified: 2 
0 


) 


+ 


Contains PICHON, an antitumor 
constituent® 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


80 
Table 1 (cont’d) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
al studie 


Linum lewisii Pursh 


Lomatium cous (S. 
Wats.) Coult. & Rose 


Lomatium cuspidatum 
Mathias & Constance 


Lomatium dissectum 
(Nutt.) Mathias & 
Constance var 
multifidum (Nutt.) 
Mathias & Constance 


Lomatium nudicaule 
(Pursh) Coult. & Rose 


Lomatium Raf. 


Lomatium triternatum 
(Pursh) Coult. & Rose 


Analgesic, carminative, disinfectant, liver aid, 
poultice, unspecified 

Gastrointestinal aid, pediatric aid: 2 

Eye medicine: 3 

Dermatological aid: 5 


0 


Dietary aid, eye medicine, gastrointestinal aid, 
other, pediatric aid, poison, stimulant, unspeci- 
fied: 1 


Analgesic, antirheumatic used externally, 
ceremonial medicine, tonic: 2 


Disinfectant, herbal steam, panacea, pulmo- 
nary aid, throat aid, tuberculosis remedy, 
venereal aid: 3 


Cold remedy, cough medicine, miscellaneous 
disease remedy (treatment of colds and flu, 
influenza, smallpox), orthopedic aid, stein! 
tory aid: 4 


Veterinary aid: 6 
Dermatological aid: 7 

Analgesic, antirheumatic used exter 
ceremonial icine, cough see. aie 
tological aid, diaphotétic, febrifuge, gastroin- 
testinal aid, gynecological aid, berhak steam, 
hunting medicine, internal medicine, laxative, 
orthopedic aid, panacea: 1 


Throat aid: 4 
Cold remedy: 6 


Antirheumatic me roam dermatological 
aid, love medicin 


, panacea, pulmonary aid, 
strengthener, ‘throat aid: 1 


+ 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
t No pharmacological study was found of any species in the genus. 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 81 


Table 1 (cont'd) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Lonicera ciliosa (Pursh) 
ir. ex DC. 


Lonicera involucrata 
(Richardson) Banks ex 
Spreng. 


Lonicera utahensis S. 
Watson in C. King 


Lupinus argenteus 
Pursh 
Lupinus sericeus Pursh 


Machaeranthera 
canescens (Pursh) A. 
Gray 


Machaeranthera 
Pinnatifida (Hook.) 
Shinners 


Maclura pomifera (Raf.) 
Schneid. 


Anticonvulsive, cold remedy, contraceptive, 
reproductive aid, sedative, throat aid, tubercu- 
losis remedy, unspecified: 1 


Dermatological aid, gynecological aid, tonic: 2 


Antidote, antirheumatic used externally, burn 
dressing, cathartic, ceremonial medicine, di- 
etary aid, disinfectant, gastrointestinal aid, 
herbal steam, love medicine, oral aid, psycho- 
logical aid, pulmonary aid, stimulant, throat 
aid, urinary aid: 1 

Analgesic, cough medicine, venereal aid: 2 
Eye medicine, orthopedic aid: 3 

Emetic, gynecological aid: 4 

Poison: 5 

Dermatological aid: 9 

Blood medicine, dermatological aid, hunting 


medicine, laxative: 1 


Dermatological aid: 1 


Eye medicine: 1 


Emetic, nose medicine, throat aid, witchcraft 
medicine: 1 


Analgesic: 1 


Eye medicine: 1 


* Excretion of selenium via milk 


ciency, but in excess may cause 
toxicity to offspring” 


M. pomifera may have limited 
usefulness in diagnosing pulmonary 
neoplasms because it binds to all 


cinoma, and small-cell carcinoma” 


* Non-LCH species in the genus investigated and found to oat enema properties. 
t No pharmacological study was found of any species in the 


82 
Table 1 (cont’d) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Matricaria matricar 


ioides (Less.) Porter 
Mimulus guttatus DC. 


Mirabilis nyctaginea 
(Michx.) MacM. 


Nicotiana quadrivalvuis 
Pursh 


Oecnothera cespitosa 
Nutt. 


ocarpus tenuifolius 
(Pursh) Benth. 


Osmorhiza Raf. 


Oxytropis besseyt 
(Rydb. fF Blank 


Paxistima myrsinites 


(Pursh) Raf 


pre ik: Siceiis 
folia R 


Pedicularis groenlandica 
Retz. 


Analgesic, dermatological aid, gastrointestinal 
aid, herbal steam, orthopedic aid: 1 


Anthelmintic, burn dressing, febrifuge, gyne- 
cological aid, oral aid, pediatric aid, urinary 
aid: 1 

Orthopedic aid: 3 

Dermatological aid: 4 


Used as a aa item and smoke plant. 
Not utilized as a 


Ceremonial aid, gynecological aid, unspecified: 
1 


Dermatological aid: 3 


ie) 


Analgesic, antidiarrheal, blood medicine, ca- 
thartic, dermatological aid, emetic, febrifuge, 

unting medicine, laxative, love medicine, 
pediatric aid, pulmonary aid, snakebite reme- 
dy, tonic, unspecified, venereal aid, veterinary 
aid: 1 


Analgesic, ceremonial medicine, cold remedy, 


dermatological aid, emetic, internal aid, kidney 


aid, orthopedic aid, unspecified: 1 
Tuberculosis remedy: 2 


oO 


Cough medicine: 1 


Contains helianol, a triterpene 
alcohol Pte anti-inflamma- 
tory activity 


i 2 


O. cespitosa contains antineoplastic 
agents 


* Oxytropis species cause severe 
adverse effects on a Prapagtieg 
function in livestock’) 


T 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 83 


Table 1 (cont’d) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Pediomelum mare 
lum (Pursh) J.W 
Grimes 


Pediomelum esculentum 


(Pursh) Rydb. 


Penstemon fruticosus 
(Pursh) Greene 


Penstemon wilcoxii 
Rydb 

Phacelia heterophylla 
Pursh 


Phacelia linearis 
(Pursh) Holz. 


Philadelphus lewisii 
Pursh 


Phlox speciosa Pursh 


Pinus ponderosa P. & 
C. Lawson 


Dermatological aid, febrifuge, laxative, unspec- 
ified: 1 


Veterinary aid: 2 


Antidiarrheal, antirheumatic used yea 
burn dressing, ear medicine, eye medicin 
gastrointestinal aid, orthopedic aid, ohne 
aid, pulmonary aid, throat aid, toothache 
remedy: 1 

Unspecified: 2 

Analgesic, antirheumatic used externally, cold 
remedy, dermatological aid, emetic, eye medi- 
cine, gynecological aid, kidney aid, love medi- 
cine, miscellaneous disease remedy (treatment 
for flu), orthopedic aid, toothache remedy, 
unspecified, urinary aid: 1 


Gastrointestinal aid, veterinary aid: 2 


0 


Dermatological aid: 1 


Cold remedy, unspecified: 1 


Antirheumatic used externally, breast treat- 
ment, cathartic, dermatological aid, hemor- 
rhoid remedy, pulmonary aid: 

0 


Abortifacient, analgesic, antihemorrhagic, 
cough medicin 


tive, stimulant, veterinary aid, witchcraft 
medicine: 1 


Antirheumatic used externally, ceremonial aid, 


febrifuge, pediatric aid: 2 
Eye medicine: 3 
Dermatological aid: 6 


—- 


— 


* Contact allergies/dermatitis 
associated with various Phacelia 
species®”>”! 


T 


Ponderosa pine and broom snake- 

weed: poisonous plants that affect 

livestock* 

spe of feeding ponderosa plant 
eedle extracts their residues to 

eal cattle”? 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


84 
Table 1 (cont’d) 


LCH plant species 


BARTONIA 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Plagiobothrys tenellus 
(Nutt. ex Hook.) Gray 


Poa canbyi (Scribn.) 
Howell 


Polanisia dodecandra 
(L.) DC. ssp. trachy- 
sperma (Torr. & Gray) 
Iltis 


Polemonium pulcher- 
rimum Hook. 


Polygala alba Nutt. 


Polygonum bistortoides 
Pursh 


Populus balsamifera L. 
ssp. trichocarpa (Torr. 
& Gray ex Hook.) 
Brayshaw 


Populus deltoides Bartr. 
arsh. ssp. 

monilifera (Ait.) 

Eckenwalde: 


Prunus emarginata 


(Dougl. ex Hook.) 
Walp. 


o 


oO 


Used for fiber and as a ceremonial item and 
smoke plant. Not utilized as a 


Dermatological aid: 1 


Ear medicine: 1 


Dermatological aid: 1 


Antirheumatic used externally, burn oii 
ceremonial medicine, cold remedy, ear me 
cine, gynecological aid, love medicine, pulmo 
nary aid, respiratory aid, veterinary ai 


Disinfectant, orthopedic aid, throat aid, tuber- 


culosis remedy, venereal aid: 2 
Unspecified: 4 
Dermatological aid: 12 


Ceremonial medicine, dermatological aid: 1 


Cancer treatment, dermatological aid, dietary 
aid, eye medicine, gastrointestinal aid, hemo- 
stat, oral aid, orthopedic aid, pediatric aid, 
preventive ite Eid psychological aid, un- 
specified: 1 


Heart medicine, laxative, tuberculosis remedy: 
2 


Blood medicine, panacea: 3 
Cold remedy: 4 
Gynecological aid: 5 


* IgE-binding trypsin inhibitors in 
plant gotten extracts of various Poa 
species 


Isolation of antitumor agents (cyto- 
toxic and antimitotic flavonols and 
cytotoxic triterpenes)”*”> 


According to interviews with two 


ems were treated with the bark’ 


* Non-LCH species in the genus investigated and found to have peeled properties. 
+ No pharmacological study was found of any species in the ge 


Table 1 (cont’d) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 85 


Subject matter of selected biomedi- 
cal studies 


Prunus virginiana L. 
var. melanocarpa (A. 
Nelson) Sarg. 


Prunus virginiana L. 
var. virginiana 
Pseudoroegneria spicata 
(Pursh) A. Love 


Psoralidium lanceol- 
atum (Pursh) Rydb. 


Psoralidium tenui- 
florum (Pursh) Rydb. 


Purshia tridentata 


(Pursh) DC 


ercus garryana 
Dougl. ex Hook. 


Quercus macrocarpa 
Michx. 


Cough medicine: 1 
Unspecified: 2 


Antidiarrheal: 1 


Ceremonial medicine: 5 


Antirheumatic used externally: 1 


Gynecological aid, y eee aid, venereal aid, 
witchcraft medicin 


Analgesic, dermatological aid, gastrointestinal 
aid: 2 


Ceremonial medicine: 3 


Analgesic, disinfectant, miscellaneous disease 
remedy cn for ei tuberculosis 
remedy, veterinary ai 


Analgesic, anthelmintic, ceremonial medicine, 
cold remedy, cough medicine, disinfectant, 
febrifuge, gastrointestinal aid, hunting medi- 
cine, liver aid, other, respiratory aid, tubercu- 
losis remedy: 1 


Antihemmorhagic, dermatological aid, gyneco- 


logical aid, laxative, miscellaneous disease 
remedy (treatment for smallpox, chickenpox, 
measles and rashes), tonic, Saoeres! aid: 2 


Cathartic, pulmonary aid: 3 
Emetic: 5 


Gynecological aid, tuberculosis remedy: 1 


Abortifacient, analgesic, anthelmintic, antidiar- 


rheal, antidote, gastrointestinal aid, heart 
medicine, orthopedic aid, pulmonary aid: 1 


Dermatological aid: 2 


According to interviews with two 
elder Salishan women, respiratory 
ailments were treated with the 
bark’ 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
+ No pharmacological study was found of any species in the genus. 


86 BARTONIA 


Table 1 (cont’d) 


LCH plant species Uses by Native Americans: number of tribal Subject matter of selected biomedi- 
groups for which use is documented cal studies 

Rhus trilobata Nutt. Analgesic, burn dressing, ceremonial medicine, | Antineoplastic agent” 

var. trilobata contraceptive, dietary aid, diuretic, emetic, 


hemostat, reproductive aid, toothache remedy, 
tuberculosis remedy, unspecified, veterinary 
aid: 1 


Cold remedy, gynecological aid, oral aid: 2 


Gastrointestinal aid, miscellaneous disease 
remedy (treatment for smallpox pustules and 
grippe): 5 


Dermatological aid: 6 
Ribes aureum Pursh Dermatological aid, unspecified: 1 t 
Orthopedic aid: 2 
Ribes divaricatum Cold remedy, dermatological aid, other, psy- t 
Dougl. ch i 


ological aid, tuberculosis remedy, venereal 
aid: 1 


Eye medicine, throat aid: 3 


Ribes sanguineum Used for fiber. Not utilized as a drug. t 
Pursh 
Ribes viscosissimum Used as food. Not utilized as a drug. + 
Pursh 


Rosa arkansana Porter ogee ks eye medicine, hemostat, stimu- t+ 
Jant, ton 


Rubus parviflorus Nutt. Alternative, antidiarrheal, antiemetic, anti- 
hemorrhagic, blood medicine, burn dressing, 
dietary aid, gynecological aid, miscellaneous 
disease remedy (young sprouts considered a 
valuable antiscorbutic), internal medicine, 
pediatric aid, pulmonary aid, tonic: 1 
Gastrointestinal aid: 2 
Dermatological aid: 4 
Rubus spectabilis Pursh — Disinfectant, gastrointestinal aid, gynecological Stevens-Johnson syndrome (bullous 
aid, pediatric aid, toothache remedy: 1 form of erythema multiform) 
: ae 
Ainpite 3 secondary to ingestion 
(continued) 


* Non-LCH species in the genus investigated and found to have — properties. 
+ No pharmacological study was found of any species in the 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 87 


Table 1 (cont’d) 


LCH plant species 


Uses by Native Americans: number of tribal 
groups for which use is documented 


Subject matter of selected biomedi- 
cal studies 


Rubus spectabilis Pursh 
(cont’d) 


Salvia reflexa Hornem. 


Sarcobatus vermiculatus 
) Torr 


Scutellaria angustifolia 
Pursh 

Sedum lanceolatum 
Torr 

Sedum stenopetalum 
Pursh 


Shepherdia argentea 
(Pursh) 


Solidago rigida L. 
Sorbus scopulina 
Greene 


Sphaeralcea coccinea 


(Nutt.) Rydb 


Synthyris missurica 


(Raf.) Pennell 


Trifolium macro- 
cephalum (Pursh) Poir. 


Trifolium micro- 


cephalum Pursh 


Burn dressing, dermatological aid: 3 


Oo 


Antidiarrheal, antihemorrhagic, blood medi- 
cine, emetic, gastrointestinal aid, toothache 
remedy, veterinary medicine: 1 


Ceremonial medicine, dermatological aid: 2 


Eye medicine: 1 
Gynecological aid, laxative: 1 
Venereal aid: 1 


Ceremonial medicine, war ae gastrointesti- 
nal aid, laxative, unspecified: 1 


Cathartic, dermatological aid, diuretic: 1 
Febrifuge, pediatric aid, unspecified, urinary 
aid: 1 


etary aid, ppmerie other, panacea, 
ielipibhen 


Ceremonial medicine, dermatological aid: 2 


0 


Used as food. Not utilized as a drug. 


According to interviews with two 


complaints were treated with the 
bark’ 


Leaf extract exhibited cnet 
activity apie t human immu: 
ficiency virus (HIV)-1 reverse 


ears 


* 


* Non-LCH species in the genus investigated and found to have re properties. 
t No pharmacological study was found of any species in the gen 


88 BARTONIA 
Table 1 (cont’d) 


LCH plant species Uses by Native Americans: number of tribal Subject matter of selected biomedi- 
groups for which use is documented cal studies 


Trillium ovatum Pursh Love medicine, poison: 1 * 
Dermatological aid: 2 


Eye medicine: 4 


Trillium petiolatum Used as food. Not utilized as a drug. : 

Pursh 

Triteleia grandiflora Adjuvant, poison, unspecified: 1 . 

Lindl. 

Uropappus lindleyi 0 . 

(DC.) Nutt. 

Vaccinium myrtillus L. Used as food. Not utilized as a drug. Therapeutic value of anthocyan- 
ees in ee oo medicine 
depart 
Anthocyanosides reduce vascular 
impair ments due to ischemia reper- 
fusion in 


ie effective in pro- 
moting and enhancing arteriolar 
rhythmic diameter changes that 
play a role in the redistribution of 
microvascular blood flow and 
interstitial fluid formation™ 


Activity of inp ums extracts 
n night vision® 


Leaves potentially — for treat- 
ment of dyslipidemia® 


Extract exerts potent pROnraee 
action on LDL particles 


Extracts exhibit antiviral action in 
experimental tick-borne encephali- 
7085 

tis 


Antiulcer activity, probably by 
potentiating the defensive barriers 
of the gastrointestinal mucosa® 


Anticancer activity of fruit extracts 
from Vaccinium species 


Vaccinium ovatum Gynecological aid, miscellaneous disease 
Pursh remedy (treatment for diabetes): 1 


* Non-LCH species in the genus investigated and found to have OS ie a properties. 
t No pharmacological study was found of any species in the gen 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 89 


Table 1 (cont’d) 


LCH plant species Uses by Native Americans: number of tribal Subject matter of selected biomedi- 
groups for which use is documented cal studies 
Veratrum californicum Blood medicine, burn dressing, emetic, febri- Congenital deformities in lambs, 
r fuge, orthopedic aid, panacea, poison, respir a- calves and goats resulting from 
tory aid, toothache remedy, veterinary aid: 1 maternal ingestion™ 
Cold remedy, disinfectant, gland medicine, Antimicrobial activity of Veratrum 
oids” 


gynecological aid, snakebite remedy, throat 
aid, venereal aid: 


Contraceptive: 3 


Antirheumatic used externally, dermatological 
aid: 4 


Xerophyllum tenax Hemostat, orthopedic aid: 1 T 


ursh) Nutt. 
deem Dermatological aid: 2 


Zigadenus elegans Analgesic, antirheumatic used externally, * Significant toxicity can result 
Pursh iin logat aid, diaphoretic, svensthenes: from ingestion of certain species of 
veterinary aid: 1 Zigadenus’”*” 
Poison: 4 
Zizania palustris L. Used as food. Not utilized as a drug. T 


var. interior (Fassett) 
Dore 


* Non-LCH species in the genus investigated and found to have pharmacological properties. 
t No pharmacological study was found of any species in the genus 


Notes 


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* Krivenko, V. V., G. P. Potebnia and V. V. Loiko. 1989. Experience in treating digestive organ 
diseases with medicinal plants. Vrachebnoe Delo 3: 76-78. 

> Montanari, T., J. E. de Carvalho and H. Dolder. 1998. Antispermatogenic effect of Achillea 
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* Lopez, M. F. and J. G. Torrey. 1985. Enzymes of glucose metabolism in Frankia sp. Journal of 
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> McCutcheon, A. R., S. M. Ellis, R. E. Hancock and G. H. Towers. 1992. Antibiotic screening of 
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® McCutcheon, A. R., S. M. Ellis, R. E. Hancock and G. H. Towers. 1994. Antifungal screening of 
medicinal plants of ear Columbian native peoples. Journal of Ethnopharmacology 44: 157-169. 

’ Turner, N. J. and R. J. Hebda. 1990. Contemporary use of bark for medicine by two Salishan native 
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’ McCutcheon, A. R., T. E. Roberts, E. Gibbons, S. M. Ellis, L. A. Babiuk, R. E. Hancock and G. H. 
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logy 49: 101-110. 
* Konoshima, T., H. Terada, M. Kokumai, M. Kozuka, H. Tokuda, J. R. Estes, L. Li, H. K. Wang and 


90 BARTONIA 


K. H. Lee. 1993. Studies on inhibitors of skin tumor promotion, XII. Rotenoids from Amorpha 
fruticosa. Journal of Natural Products 56: 843-848. 

0 Li, L., H. K. Wang, J. J. Chang, A. T. McPhail, D. R. McPhail, H. Terada, T. Konoshima, M. 
Kokumai, M. Kozuka and J. R. Estes. 1993. Antitumor agents, 138. Rotenoids and isoflavones as 
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'! Wang, H. K., Y. Xia, Z. Y. Yang, S. L. Natschke and K. H. Lee. 1998. Recent advances in the 
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2 Panter, K. E. and L. F. James. 1990. Natural plant toxicants in milk: a review. Journal of Animal 


3 Grases, F., G. Melero, A. Costa-Bauza, R. Prieto and J. G. March. 1994. Urolithiasis and 
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5 James, L. F., R. J. Molyneux, K. E. Panter, D. R. Gardner and B. L. Stegelmeier. 1994. Effect of 
feeding ponderosa pine needle extracts and their residues to pregnant cattle. Cornell Veterinarian 
84: 33-39. 

‘6 Matsuda, H., $. Nakamura, T. Tanaka and M. Kubo. 1992. Pharmacological studies on leaf of 
Arctostaphylos uva-ursi (L. ) Spreng. V. Effect of water extract from Arctostaphylos uva-ursi (L. ) 
Spreng, (bearberry leaf) on the antiallergic and antiinflammatory activities of dexamethasone 
ointment. Yakugaku Zasshi 112: 673-677 

” Matsuda, H., T. Tanaka and M. Kubo. 1991. Pharmacological studies on leaf of Arctostaphylos uva- 
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Yakugaku Zasshi 111: 253-258. 

'® Matsuda, H., H. Nakata, T. Tanaka and M. Kubo. 1990. Pharmacological study on Arctostaphylos 
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immuno-inflammation. Yakugaku Zasshi 110: 68-76 

x ado H., M. Higashino, Y. Nakai, M. Iinuma, M, Kubo and Fk Lang. 1996. Studies of cuticle 
drugs from natural sources. IV. loisbaory effects of some Arctostaphylos plants on melanin 
biceyaihena Biological and Pharmaceutical Bulletin 19: 153-156. 

© Matsuo, K., M. Kobayashi, Y. Takuno, H. Kuwajima, H. Ito and T. Yoshida. 1997. Anti-tyrosinase 

activity constituents of i uva-ursi. Yakugaku Zasshi 117: 1028-1032. 

*! Ritch-Kre, E. M., S. Thomas, N. J. Turner and G. H. Towers. 1996. Carrier herbal medicine: 
traditional and eeuaianee sala use. Pra of Ethnopharmacology 52: 85-94. 

* Swanston-Flatt, $. K., C. Day, C. J. B and P. R. Flatt. 1989. Evaluation of traditional plant 
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Tan, R. X., W. F. Zheng and H. es Tang. 1998. Biologically active substances from the genus 

emisia. Planta Medica 64: 295-302 

** Gancevici, G. G. and C. Popescu. 1987. Natural inhibitors of complement. II. Inactivation of the 
complement cascade in vitro by vegetal spices (Ocimum basilicum, Artemisia dracunculus and Thymus 
vulgaris). Archives Roumaines de Pathologie Experimentales et de Microbiologie 46: 321-331. 

* Zani, F., G. Massimo, S. Benvenuti, A. Bianchi, A. Albasini, M. Melegari, G. Vampa, A. Bellotti and 
P. Mazza. 1991. Studies on the genotoxic properties of essential oils with Bacillus subtilis rec-assay 
and Salmonella/microsome pe assay. Planta Medica 57: 237-241. 

6 Katial, R. K., F. L. Lin, W. W. Stafford, R. A. Ledoux, C. R. Westley and R. W. Weber. 1997. 
Mugwort and sage (As eetiiste) polls cross-reactivity: ELISA inhibition and immunoblot evaluation. 
Annals of Allergy, Asthma and Immunology 79: 340-346. 

7 Bork, P. M., M. L. Schmitz, M. Kuhnt, C. Escher and M. Heinrich. 1997. Sesquiterpene lactone 
containing Mexican Indian medicinal plants and pure sesquiterpene lactones as potent inhibitors of 
transcription factor NF-kappaB. FEBS Letters 402: 85-90. 

8 Malagon, F., J. Vazquez, G. Delgado and A. Ruiz. 1997, Antimalaric effect of an alcoholic extract 
of Artemisia ludoviciana mexicana in a rodent ae model. Parasitologia 39: 3-7. 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 91 


* de Balogh, K. K., A. P. Dimande, J. J. van der Lugt, R. J. Molyneux, T. W. Naude and W. G. 
Welman. 1999. A lysosomal storage disease induced by Jpomoea carnea in goats in Mozambique. 
Journal of Veterinary Diagnostic Investigation 11: 266-273. 

* Panter, K. E., L. F. James, B. L. Stegelmeier, M. H. Ralphs and J. A. Pfister. 1999. Locoweeds: 
effects on reproduction in livestock. Journal of Natural Toxins 8: 53-62. 

* Stegelmeier, B. L., L. F. James, K. E. Panter, M. H. Ralphs, D. R. Gardner, R. J. Molyneux and J. 
A. Pfister. 1999. The pathogenesis and toxicokinetics of locoweed (Astragalus and Oxytropis spp.) 
poisoning in lives stock. Journal of Natural Toxins 8: 35-45. 

* Cairney, J., R. J. Newton, E. A. Funkhouser, S. Chang and D. eae 1995; Nucleotide sequence 
of acDNA for a water channel protein (aquaporin) h (Pursh.) Nutt. 
Plant Physiology 108: 1291-1292. 

* Cairney, J., R. J. Newton, E. A. Funkhouser and $. Chang. 1995. Nucleotide sequence of a CDNA 
from Atriplex canescens (Pursh. ) Nutt. A homolog of a jasmonate-induced protein from barley. 
Plant Physiology 108: 1289-1290. 

* Matsuura, H., G. Saxena, $. W. Farmer, R. E. Hancock and G. H. Towers. 1996. An antibacterial 
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* Scher, J.M., J. B. Speakman, J. Zapp and H. Becker. 2004. Bioactivity guided isolation of antifungal 
compounds from the liverwort Bazzania trilobata (L.) S. F. Gray. Phytochemistry 65: 2583-2588. 

*© Fukuda, K., Y. Hibiya, M. Mutoh, M. Koshiji, $. Akao and H. Fujiwara. 1999. Inhibition of 
activator protein 1 activity by berberine in human hepatoma cells. Planta Medica 65: 381-382. 

” Ivanovska, N. and S. Philipov. 1996. Study on the anti-inflammatory action of Berberis vulgaris root 
extract, alkaloid fractions and pure alkaloids. International Journal of Immunopharmacology 18: 553- 
561 


* Luo, C. N., X. Lin, W. K. Li, F. Pu, L. W. Wang, S. S. Xie and P. G. Xiao. 1998. Effect of 
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Journal of Ethnopharmacology 59: 211-215. 

* Klein, F. K. and H. Rapoport. 1968. Ceanothus alkaloids. Americine. Journal of the American 
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Rose, S. L., C. Y. Liand A. §. Hutchins. 1980. A streptomycete antagonist to Phellinus weirii, Fomes 
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*! Kern, J. R. and J. H. Cardellina. 1983. Native American medicinal mec Anemonin from the horse 
stimulant Clematis hirsutissima. Journal of Ethnopharmacology 8: 121- 

” Morgan, G. R. 1981, "Sugar wig siete hirsutissima): a horse restorative of the Nez Perces. 
Journal of Ethnopharmacology 4: 

® Muller, A., W. Linke and W. ae 1999. Crataegus extract blocks potassium currents in guinea pig 
ventricular cardiac myocytes. Planta Medica 65: 335-339. 

* Pfister, J. A., D. R. Gardner, K. E. Panter, G. D. Manners, M. H. Ralphs, B. L. Stegelmeier and T. 
K. Schoch. 1999, Larkspur (Delphinium spp..) poisoning in livestock. Journal of Natural Toxins 8: 


81-94 

* D’Agostino, M., A. Dini, C. Pizza, F. Senatore and R. Aquino. Sterols from Eguisetum arvense. 
Bollettino - Societa Italiana Biologia Sperimentale 60: 2241-2245. 

*° Maeda, H., K. Miyamoto and T. Sano. 1997. Occurrence of dermatitis in rats fed a cholesterol diet 
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553-563. 

*” Sudan, B. J. 1985. Seborrhoeic dermatitis induced by nicotine of horsetails (Equisetum arvense L.). 
Contact Dermatitis 13: 201-202. 

‘8 Diamond, K. B., G. R. Warren and J. H. Cardellina. 1985. Native American food and medicinal 
plants. 3. alpha-Methylene butyrolactone from Erythronium grandiflorum Pursh. Journal of 
Ethnopharmacology 14: 99-101. 

* Pinedo, J. M., V. Saavedra, F. map  areg and P. Llamas. 1985. Irritant dermatitis due to 
Euphorbia marginata. Contact Dermatitis 

° Urushibata, O. and K. Kase. 1991. sige 4" contact dermatitis from Euphorbia marginata. Contact 


92 BARTONIA 


Dermatitis 24: 155-156. 
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2 Szafran, L. 1967. Acute conjunctivitis and keratitis due to the sap of Euphorbia marginata. Polski 
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3 Siegers, C. P., E. von Hertzberg-Lottin, M. Otte and B. Schneider. 1993. Anthranoid laxative 
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* van Gorkom, B. A., E. G. de Vries, A. Karrenbeld oid J. H. Kleibeuker. 1999. Review article: 
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Therapeutics 13: 443-452. 

55 Kreutzer, S., O. Schimmer and R. Waibel. 1990. Triterpenoid sapogenins in the genus Grindelia. 
Planta Medica 56: 392-394. 

% Gardner, D. R., L. F. James, K. E. Panter, J. A. Pfister, M. H. Ralphs and B. L. Stegelmeier. 1999. 
Ponderosa pine and broom snakeweed: poisonous plants that affect livestock. Journal of Natural 
Toxins 8: 27-34. 

7 Staley, E. C., G. S. Smith and J. A. Greenberg. 1996. Decreased reproductive effects from snakeweed 
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eterinary and Human Toxicology 38: 259-264. 

8 Ulubelen, A., M. E. Caldwell and J. R. Cole. 1965. Isolation of an antitumor proteinaceous 
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*° Arisawa, M., S. Funayama, J. M. Pezzuto, A. D. Kinghorn, G. A. Cordell and N. R. Farnsworth. 
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°° Wong, S. M., Y. Oshima, J. M. Pezzuto, H. H. Fong and N. R. Farnsworth. 1986. Plant anticancer 
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6! Sanchez de Medina, F., M. J. Gamez, I. Jimenez, J. Jimenez, J. I. Osuna and A. Zarzuelo. 1994. 
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® Tunon, H., C. Olavsdotter and L. Bohlin. 1995. Evaluation of anti-inflammatory activity of some 
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°° Herz, W. and P. Sharma. 1976. Dienolide a seco-germacradienolide from Liatris pycnostachya and 
other antitumor ge of Liatris species. The Journal of Organic Chemistry 41: 1248-1253. 

 Sarker, A. B., T. R. Koirala, Aftabuddin, H. J. Jeon and I. Murakami. 1994. Lectin histochemistry 
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6 Akihisa, T., K. Yasukawa, H. Oinuma, Y. Kasahara, $. Yamanouchi, M. Takido, K. Kumaki and 
T. Tamura. 1996. Triterpene alcohols from the flowers of compositae and their anti-inflammatory 
effects. Phytochemistry 43: 1255-1260. 

% Pettit, G. R., P. M. Traxler and C. P. Pase. 1973. Antineoplastic agents 31. Oenothera caespitosa. 
Lloydia 36: 202-203. 

*? Ralphs, M. H. and L. F. James. 1999. Locoweed grazing. Journal of Natural Toxins 8: 47-51. 

*8 Stegelmeier, B. L., L. F. James, K. E. Panter, M. H. Ralphs, D. R. Gardner, R. J. Molyneux and J. 
A. Pfister. 1999. The pathogenesis and toxicokinetics of locoweed (Astragalus and Oxytropis spp.) 
poisoning in lives stock. Journal of Natural Toxins 8: 35-45. 

® Ippen, H. 1990. Contact allergy to Phacelia spp. (Hydrophyllaceae). Dermatologic Clinics 8:67-71. 

” Reynolds, G., W. Epstein, D. Terry and E. Rodriguez. 1980. A potent contact allergen of Phacelia 
(Hydrophyllaceae). Contact Dermatitis 6: 272-274. 

7! Reynolds, G. W. and E. Rodriguez. 1981. oe. _— that cause contact dermatitis from 
trichomes of Phacelia ixodes. Planta Medica 43: 187-1 

7? James, L. F., R. J. Molyneux, K. E. Panter, D. R. pith and B. L. Stegelmeier. 1994. Effect of 
feeding ponderosa pine needle extracts and their residues to pregnant cattle. Cornell Veterinarian 


MEDICINAL RELEVANCE OF LEWIS AND CLARK PLANTS 93 


84: 33-39. 

” Berrens, L. and F. Maranon. 1995. IgE-binding trypsin inhibitors in plant pollen extracts. Experientia 
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* Shi, Q., K. Chen, T. Fujioka, Y. Kashiwada, J. J. Chang, M. Kozuka, J. R. Estes, A. T. McPhail, D. 
R. McPhail and K. H. Lee. 1992. Antitumor agents, 135. Structure and stereochemistry of 
polacandrin, a new cytotoxic Cas from Polanisia dodecandra. Journal of Natural Products 55: 

8-1497. 


” Shi, Q., K. Chen, L. Li, J. J. Chang, C. Autry, M. Kozuka, T. Konoshima, J. R. Estes, C. M. Lin 
and E. Hamel. 1995. Antitumor agents, 154. Cytotoxic and antimitotic flavonols from Polanisia 
dodecandra. Journal of Natural Products 58: 475-482. 

” Pettit, G. R., E. I. Saldana and E. Lehto. 1974. Antineoplastic agents 35. Rhus trilobata. Lloydia 37: 

539-540 

” Steiner, G. C., R. W. Arnold, R. R. Roth and J. S. Ice. 1991. Stevens-Johnson syndrome secondary 
to ingestion af salmon berries. Alaska Medicine 33: 57-59. 

78 Yoshida, T., H. Ito, T. Hatano, M. Kurata, T. Nakanishi, A. Inada, H. Murata, Y. Inatomi, N. 
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Chemical and Pharmaceutical Bulletin 44: 1436-1439. 

a suri ini M. 1972. Therapeutic value of Vaccinium myrtillus anthocyanosides in an internal 
medicine department. Therapeutique 48: 579-581. 

7 Saris S., S. Malandrino and A. Colantuoni. 1995. Effect of Vaccinium myrtillus anthocyanosides 
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*! Colantuoni, A., S. Bertuglia, M. J. Magistretti and L. Donato. 1991. Effects of Vaccinium myrtillus 
anthocyanosides on arterial vasomotion. beim gigas 41: 905-909. 

® Jayle, E., M. Aubry, H. Gavini, G. Braccini and C. De la Baume. 1965. Study concerning the 
action a anthocyanoside extracts of ae siete on night vision. Annales de Oculistique 198: 
556-562. 

*® Cignarella, A., M. Nastasi, E. Cavalli and L. Puglisi. 1996. Novel lipid- lowering properties of 
Vaccinium wonjveilis L. leaves, a traditional antidiabetic treatment, in several models of rat 
dyslipidaemia: a comparison with ciprofibrate. Thrombosis Research 84: 311-322. 

* Laplaud, P. M., A. Lelubre and M. J. Chapman. 1997. Antioxidant action of Vaccinium myrtillus 
extract on human low density lipoproteins in vitro: initial observations. Fundamental and Clinical 
Pharmacology 11: 35-40. 

5 Eokina, G. L, V. M. Roikhel’, M. P. Frolova, T. V. Frolova and V. V. Pogodina. 1993. The antiviral 
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*° Magistretti, M. J., M. Conti and A. Cristoni. 1988. Antiulcer activity of an anthocyanidin from 
Vaccinium myrtillus. Arzneimittelforschung 38: 686-690. 

*” Bomser, J., D. L. Madhavi, K. Singletary and M. A. Smith. In vitro anticancer activity of fruit 
extracts from Vaccinium species. Planta Medica 62: 212-216. 

8 Binns, W., R. F. Keeler and L. D. Balls. 1972. Congenital deformities in lambs, calves and goats 
resulting trou maternal ingestion of Veratrum californicum: hare lip, cleft palate, ataxia and 
hypoplasia of metacarpal and metatarsal bones. Clinical Toxicology 5: 245-261. 

*® Keeler, R. F. and L. D. Stuart. 1987. The nature of congenital limb defects induced in lambs by 

maternal ingestion of Veratrum californicum. Journal of Toxicology. Clinical Toxicology 25: 273-286. 

bs Wolters, B. 1970. Antimicrobial activity of Veratrum alkaloids. Planta Medica. 19: 189-196. 

*! Heilpern, K. L. 1995. Zigadenus poisoning. Annals of Emergency Medicine 25: 259-262. 

* Spoerke, D. G. and S. E. Spoerke. 1979. Three cases of Zigadenus (death camus) poisoning. Veterinary 
and Human Toxicology 21: 346-347. 

> Wagstaff, D. J. and A. A. Case. 1987. Human poisoning by Zigadenus. Journal of Toxicology. Clinical 
Toxicology 25: 361-367. 


ve 


Bartonia No. 62: 95-102, 2004 


The Second Two Hundred Years Are Better: 
The Endearing, Enduring Plants of Lewis and Clark 


EARLE E. SPAMER’? AND RICHARD M. MCCourRT?* 
"Archives, Ewell Sale Stewart Libary, *Department of Botan 
The Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway, 
Philadelphia, Pennsylvania 19103 
*Current address: 1 Locust Court, Maple Shade, New Jersey 08052 
‘Current address: 7801 Cresheim Road, Philadelphia, Pennsylvania 19118 


THIS is a story about a Prussian pilferer, his proclivities for gathering North American 
plants, and his putatively inebriant state that ushered along his work on the first 
comprehensive flora for North America. Not a one of the specimens in question was 
actually collected by him, but some of them accumulated 14,000 miles of travel by 
horseback, canoe, and sailing ship before rejoining their long-forgotten companions nearly 
a century later. They had been guests in the White House and captives in London. Others 
lay quietly in a Philadelphia attic for three-quarters of a century, completely forgotten. 
Some plants were eaten by bugs, others were eaten by humans. Researchers in the twentieth 
century would even go so far as to incinerate parts of some of them. And now everyone 
wants to see them. 


THE AMERICAN ZEPHYR 


Meriwether Lewis and William Clark are among the lesser known botanical collectors, 
even though as explorers they have been objects of infatuation ever since they tramped to 
the Pacific Ocean and back in the earliest part of the nineteenth century. Scholars and 
students alike have studied the expedition, its labors, and its booty. Later, the explorers’ 
travails became the focus of cinematographers and editors. But the plants they collected are 
hardly ever seen against the dazzle of travel and discovery. The 222 specimen sheets of the 
Lewis and Clark Herbarium, now in the Academy of Natural Sciences of Philadelphia, are 
likewise eclipsed by the larger, longer story told by the Academy’s entire herbarium—a 
million more dried, pressed plant specimens from around the globe and a century farther yet 
into the past. Even though the Lewis and Clark Herbarium represents just two one-hun- 
dredths of a percent of the plant specimens curated at the Academy, they nonetheless stand 
out as the most publicly recognized symbol of the unique historical treasures in the entire 
herbarium. 

It is easy to gloat when you have such a treasure. Yet some historians glibly point out 
that in one way Lewis and Clark were dismal failures. They had not achieved the principal 
objective of the expedition conceived by President Thomas Jefferson and secretively funded 
by Congress. The ever elusive quest for a mostly water-borne passage between the Atlantic 


Manuscript submitted 28 May 2004 


96 BARTONIA 


and Pacific Oceans was, again, a disappointment. Ultimately the passage did not cross the 
Great Divide of the American West. Indeed, the divide was not the single, thin range of 
mountains imagined to lie in the way of industrious pioneers and merchant loads. The quest 
for a waterway eventually would have to be won, a century later, by digging and blasting 
through Panama. Yet the efforts of Lewis and Clark never were overshadowed by the 
geography that denied them that one goal. Their every word has been pored over, 
rewritten, edited, and interpreted in print and film for the consumption and entertainment 
of the studious and curious alike. Perhaps the greatest measure of their success was reached 
within the realm of bibliophiles, who have ballooned the cost of the earliest publications 
about the expedition to levels attained by few other printed works. 

The bounty of the expedition and the legacy of the plants play out the themes of 
American spirit and intellectual freedom on an international stage. And an “abcedarium” 
of taxa is represented in the Lewis and Clark Herbarium, from Acer to Zizania, half of 
which were new to science when they were named in 1813. The last time that all of the 
botanical gatherings of Lewis and Clark were in one place was in 1806, in Philadelphia. 
Today, 96 percent of the pressed plants that survive are still in this town. It was not always 
that way. Portions of the collection were scattered to the winds, and only by happenstance 
were nearly all of the surviving specimens reunited. 


TO LONDON, TO LONDON 


The Lewis and Clark Herbarium has a long, curious history. Its story began in 
Philadelphia, where in 1803 Meriwether Lewis went to obtain instruction in botany and 
other sciences and to shop for supplies for the long journey. Trained by the University of 
Pennsylvania’s eminent professor of botany and medicine, Benjamin Smith Barton, Lewis 
searched for and gathered new and potentially useful plant species that he encountered from 
St. Louis through 10 future states, an 8,000-mile round trip. He was dismayed when he lost 
many plant specimens that he had stored in caches, intending to retrieve them on the 
journey back home rather than carry them across the continent and back; the caches were 
water-soaked by winter and spring floods. More of his dried, pressed plants were 
successfully returned to Thomas Jefferson in Washington, D.C., who forwarded them to the 
American Philosophical Society in Philadelphia. By the time the entire collection was 
grouped together around 1807, upon Lewis’s arrival in Philadelphia with the balance of the 
expedition’s collections, some 30 plants had been inexplicably, utterly lost. 

Barton was assigned the task of coordinating the scientific examination of the collections 
from the expedition; of course, as a botanist he got the plants. But he dallied at that work, 
finding the pressures of his “day job” always in the way. Seven years or so later, he fell ill, 
and he planned to go to Europe for his health—the Academy’s archives hold Thomas 
Jefferson’s letter of introduction for Barton’s visit to an Italian scholar. But Barton died in 
New York in 1815. He left a detailed will, written over a period of days shortly before his 
death, precisely dispersing his effects, but there is no mention of the Lewis and Clark plants 
that were in his custody in Philadelphia. He never had gotten around to working on them 
himself. 

What Barton never knew was that nearly a quarter of the Lewis and Clark collection was 
missing, taken from right under his nose. The specimens weren’t even in the country 
anymore. When Barton had dallied from the outset, he soon turned to his friend William 
Hamilton for help. Hamilton, who was as much of a landed country squire as the new 


THE PLANTS OF LEWIS AND CLARK 97 


American national culture would tolerate, had a lush estate, The Woodlands, which survives 
greatly reduced in size and planted with the stones and monuments of a cemetery in the 
West Philadelphia part of the city. It was a horticulturist’s paradise. Hamilton had gotten 
seeds from the Lewis and Clark expedition, too, and was growing plants that were hitherto 
unknown to the East, or for that matter even in the cultured gardens of Europe. His able 
gardener and classically trained botanist was Frederick Pursh, a Prussian of great horticultur- 
al talents, and it was Pursh to whom the Lewis and Clark plants were given for identifica- 
tion and study. 

Barton already knew Pursh and had hired him to collect plants. Barton also knew of 
Pursh’s proclivity for the “toddy” and that it sometimes interfered with his collecting work. 
But he trusted in Pursh’s botanical acumen to find the nuggets in Lewis’s collection, and 
Pursh took to the task with relish. New species were soon recognized as well, as detailed 
in Pursh’s own notes that survive with the plant specimens. The job, partly paid for by 
Meriwether Lewis, went ably and successfully enough, but by 1808 Pursh had to look for 
other work. He left the plants to Hamilton, Barton, and their horticulturist friends. Pursh 
went to New York and eventually left the country for England, arriving there in 1811. 
What no one knew was that in Pursh’s bags were samples of Lewis and Clark’s gatherings, 
amounting to a quarter of the entire collection. These were specimens that he understood 
to be new and interesting to botany and which he intended to illustrate or better describe 
in a much-needed comprehensive flora for North America. 

Some historians and botanists have quibbled over the meaning of Pursh’s purloined plants. 
Of course he did not have permission to take them. But did he need it? Gracious and 
gratuitous sharing was de rigeur among scholars of the day, often with implied permission. 
And after all, Pursh had selected only a portion of every scrap of a species represented in the 
collection. Only in a couple of instances did he seem to have taken all of what remained 
of the booty of a particular species. Perhaps he knew all too well that Barton wasn’t going 
to use them, and who else, then, was there? Historical hindsight answers for us: no one. 
In fact, not only did no one ever inquire about the missing material brought to England or 
how Pursh had gotten the specimens in order to publish his findings in 1813, but no one 
ever again inquired about the whole lot of Lewis and Clark’s plants after the ostensibly 
intact collection was bundled off to the American Philosophical Society. They were in 
safekeeping and utterly forgotten. But for Pursh’s petty larceny, Lewis and Clark’s botanical 
legacy would be a footnote to science. 

The plants that remained in Philadelphia lay unremembered and unvisited. After Barton 
died, they were sent back again to the American Philosphical Society, where they remained, 
wrapped in their bundles with Meriwether Lewis’s own field notes written on blotting 
papers [see McCourt and Spamer, page 1 in this issue —Ed.]. After better and more 
extensive gatherings from the Great Plains and the American Northwest began to be 
available, no one bothered to inquire about the original collections from Lewis and Clark. 
So they sat at the Society, unstudied and unnoticed. Not until 1897 would Thomas Meehan, 
a botanist with the Academy and a member of the American Philosophical Society, 


rediscover them. 
ACROSS THE ATLANTIC—AGAIN 


Meanwhile, the London residuum of the collection had been mounted on herbarium 
sheets in the British Museum, the workplace of Aylmer Bourke Lambert, who hired the 


98 BARTONIA 


wandering Prussian, Pursh. However, Pursh’s propensity for drink worked peculiarly to 
Lambert’s advantage. Pursh’s fondness became known to his employers, as there is some 
discrete correspondence to the effect, which said either to pare down the consumption or 
to water it down, lest the line be crossed from good work to no work! Pursh finally 
published his North American flora in December of 1813, which took a while yet to come 
to America because of the unsettled times of the War of 1812 between England and the 
United States. Pursh then left to Canada, where he died in 1820 without ever again doing 
any substantive work. 

Lambert lived on until 1842, after which his rich estate was auctioned by Sotheby’s. 
Serendipitously, an American, Edward Tuckerman, was present and bought some of the 
“dregs” of the day’s auctions; the largest and best collections had already been taken by 
British institutions. Little did Tuckerman know at the time that his “miscellaneous” 
American plants included numerous that were the types of Frederick Pursh’s new 
species, plus virtually all of the plants pusloinell from the set of Lewis and Clark’s gatherings 
that were in bundled storage in Philadelphia. Interestingly and fortuitously for us, Lambert 
seems not to have absorbed into his main collections the bulk of Pursh’s voucher specimens 
from the landmark flora that Pursh published some 30 years earlier. Otherwise, a much 
larger portion of this uniquely American expedition’s botany may likely have remained in 
British collections. 

In 1856, Tuckerman sent his won collection from Massachusetts to the Academy in 
Philadelphia in exchange for the liberty of access to what then was the most comprehensive 
collection of American lichens, which had become Tuckerman’s own field of expertise. So 
Lewis and Clark’s plants came home again. What Tuckerman had not gotten at auction 
were just 10 more of Lewis and Clark’s plants that were mixed in with much larger 
collections purchased by the British Museum. Those 10 plants are kept today at the Royal 
Botanic Gardens at Kew, the only Lewis and Clark specimens known outside of Philadel- 
phia. 

Forty years later, the Academy’s Thomas Meehan was clued to the possibility that Pursh’s 
voucher specimens—Lewis and Clark’s specimens included—might still repose at the 
American Philosophical Society, of which Meehan was a member also. In his subsequent 
fight with administrators to get Lewis and Clark’s plants out of the attic and off to proper 
study at the Academy with the help of botanists at Harvard, he unwittingly had again 
consolidated almost all of the extant botanical materials from the expedition. For some 
reason, however, the bundled plants that had remained in Philadelphia and were deposited 
at the Academy in 1897, still were not mounted on herbarium sheets until 1921. 

The trove from the Society’s attic brought grim and giddy comments from the observers. 
Bugs had sometime gotten to a few specimens, feasting on the dried, nutritious plant tissue. 
Thomas Meehan wrote on one label, “all gone!” Frederick Pursh left a label while he was 
in Philadelphia, which is a transcription of Meriwether Lewis’s own observations of the 
plant’s food use by Native Americans. The bug-eaten fragments that remained in the bundle 
brought Meehan to wag on another label, “all eaten!” 


LEWISIA AND CLARKIA REDUX 


Lewis and Clark were not wholly anonymous collectors. When Frederick Pursh had 
written up the scientific botany of the expedition as part of his landmark flora for North 
America, he named several species for Lewis (recently deceased in 1809), such as the syringa 


THE PLANTS OF LEWIS AND CLARK 99 


Philadelphus lewisii, and he named the genera Lewisia and Clarkia to recognize the two 
explorers’ contributions to botany. 

The botanical collections from Lewis and Clark have been reappraised, coincidentally, 
about once each century. First there was Pursh’s 1813 flora, then Meehan’s review of the 
refound plants in 1898, and a completely updated taxonomy by Reveal, Moulton, and 
Schuyler in 1999. The specimens are a working collection for modern botanists and 
environmental scientists. Of the 199 species and varieties known to have been collected by 
Lewis and Clark—178 species are in the herbarium today—nearly half were new to science 
when Pursh published them in 1813. Nurserymen of the time were keen to get their hands 
on the new finds from the West. Peter Hatch, in 2003, recorded that Thomas Jefferson had 
more than two dozen species in cultivation at Monticello, including several currants, corn, 
and tobacco. 

Perhaps the most remarkable use of the Lewis and Clark Herbarium to date has been as 
the source of material for molecular studies of carbon isotopes in the pre-Industrial 
Revolution atmosphere at the beginning nineteenth century. This required the judicious, 
yet heart-pausing, removal and incineration of tiny pieces of leaf material in order to perform 
chemical analyses of the gases given off. Scanning electron microscopy of key tissue 
structures in those specimens was also performed. Future uses of these plants could include 
DNA matching of present-day plant populations against the specimens of two centuries ago, 
among other reasons to corroborate the root stock and provenance data that accompany the 
specimens. The inspiration is greater still. Much as Lewis and Clark, or even the polymath 
Thomas Jefferson, could not have imagined studies of DNA and molecular chemistry, we 
likely cannot imagine what investigatory techniques will be available and what scientific or 
ecological worth will come of these flowers, leaves, and twigs between now and 2204. 

With the marking of the 200th anniversary of the Lewis and Clark expedition, renewed 
interest in the plants led the curatorial staff of the Academy of Natural Sciences of 
Philadelphia Botany Department to successfully apply for a grant from the Save America’s 
Treasures program administered by the National Park Service and the National Trust for 
Historic Preservation. The grant funded repair of damaged specimens and herbarium papers, 
construction of a custom archival housing for each of the 222 specimens, custom metal 
cabinets, and remodeling of a new climate-controlled, secure room for long-term storage. 
The grant also funded a CD-ROM, which was the first illustrated scientific guide to the 
entire Lewis and Clark Herbarium. 


“OCIAN IN VIEW!” William Clark, November 7, 1805 


The Lewis and Clark specimens are traveling again—nearly to the Pacific and back. When 
William Clark rejoiced in his journal upon seeing the Pacific Ocean, surely the farthest thing 
from his mind was that some of the plants he carried would ever come this way again. It 
is for botany as momentous a trip as when Leonardo da Vinci’s Mona Lisa and Michelange- 
lo’s Pieta came to New York from France and Italy in the 1960s, the likes of which may 
never happen again. During the bicentennial years, selected plants will be shown at venues 
of the Lewis and Clark Bicentennial National Exhibition, in St. Louis, Philadelphia, Denver, 
Portland (Oregon), and Washington, D.C., among others. Other exhibitions also feature 
some of Lewis and Clark’s plants, at Jefferson’s Virginia home, Monticello, and in 
Richmond, Virginia; Tacoma, Washington; Boise, Idaho; Helena, Montana; and Topeka, 
Kansas. Altogether, some three dozen specimens will travel thousands of miles to allow the 


100 BARTONIA 


public to see these precious items collected by Meriwether Lewis 200 years ago. Of course, 
they will travel more safely and securely than they ever did on Lewis’s original trek, in the 
care of fine-arts shippers by truck and airplane. No oilskin bags or pirogues here! 

After the bicentennial hoopla dies down, the specimens will return to the relative quiet 
of the special collections room of the Academy of Natural Sciences of Philadelphia. Some 
of them will by then have traveled far enough in two centuries to have circumnavigated the 
globe. Scholars will continue to use them in their studies of vegetational distributions, 
taxonomy, and climate. Much as with the standardization of spelling and the proper 
teaching of spelling since Lewis and Clark’s time—prolifically shown by the ingenious, 
multiple spellings of the same words throughout the journals of the expedition—so too are 
the physical and biological sciences ever growing and adjusting to new standards of 
understanding and utility. 

When the quadricentennial rolls around, the explorers and their botanical legacy we hope 
will be re-discovered again with who knows what tools and methods. The plants will be 
waiting for these future explorers. The measures we took in conserving them two centuries 
earlier should have proven their worth, and we presume that the plant species will be 
thriving, too, in some vestige of the natural world from which Lewis and Clark picked 
them. None of the plant species collected by Lewis and Clark are either extinct or 
endangered. Their natural ranges may have shifted, but that is another story altogether. 
When the Lewis and Clark Herbarium has reached the end of its fourth century, we 
optimistically expect that from our efforts the second 200 years will be seen to have been 
less taxing than the first 200. 


BIBLIOGRAPHICAL NOTES AND FURTHER READING 


Frederick Pursh’s seminal flora for North America was published in London in December 
1813 with the ponderous, Latin title, Flora Americae Septentrionalis. As might be expected, 
the two volumes are now scarce and often found only in the rare-book collections of 
academic libraries. A facsimile reprint 16 decades later (1979) is likewise infrequently met 
except in specialized libraries. In either case, Pursh’s English and Latin text is as dry as were 
the plants he described, but the reprint was accompanied by a scholarly but readable, book- 
length introduction by Joseph Ewan (1979). 

The history of the Lewis and Clark Herbarium can be read, however, in modern 
literature, too. First, the adventures and achievements in botany were covered by Gary 
Moulton (1999). After the first specimens were returned to Thomas Jefferson and sent on 
to the American Philosophical Society in 1804, the history of the plants twists and turns 
mostly in correspondence. In and between the lines alike, one can read of innuendo and 
despair, as summarized by Earle Spamer and Richard McCourt (2002, Notulae Naturae No. 
475, a publication of the Academy of Natural Sciences of Philadelphia). 

Frederick Pursh’s own life story and accomplishments are amply outlined by Joseph 
Ewan’s introductory volume accompanying the 1979 reprinting of Pursh’s Flora. Thomas 
Meehan’s report on the first examination of the Lewis and Clark plants in a century was 
published in the Proceedings of the Academy of Natural Sciences of Philadelphia (1898). Further 
information on the geography of the collections appeared later that year in an article by 
Elliott Coues in the Proceedings (1898). A century later, a completely revised taxonomy was 
prepared for the same journal by James Reveal, Gary Moulton, and A. E. Schuyler (1999). 
The new taxonomy subsequently was used as the foundation for the first-ever illustrated 


THE PLANTS OF LEWIS AND CLARK 101 
taxonomic study of the Lewis and Clark Herbarium, a CD-ROM by Spamer and McCourt 
(2002) 


The Lewis and Clark Herbarium was awarded a significant conservation grant in the Save 
America’s Treasures program in 1999. When implemented, the conservational procedures 
were documented and later described by Richard McCourt, Catharine Hawks, and Earle 
Spamer (2002, Notulae Naturae No. 476). Mark Teece’s landmark study of the carbon 
isotopes contained in Lewis and Clark’s plants was published in Notulae Naturae No. 477 
(2002). William Clark’s joyous journal entry may be read in its original context in Gary 
Moulton’s Journals of the Lewis and Clark Expedition (1990, vol. 6, p. 58 and the correspond- 
ing narratives on pp. 30-33). Peter Hatch’s article on the cultivation of Lewis and Clark’s 
seed trove is reported in Monticello’s Twinleaf Journal (January 2003). 

For those who are more attuned to the ways and means of the internet, information on 
the Lewis and Clark Herbarium can be found at http://www.acnatsci.org/lewisclark. Living 
examples of Lewis and Clark’s plants can also be studied in a web site composed by James 
L. Reveal, at http://www.life.umd.edu/emeritus/reveal/pbio/LnC/LnCpublic.html. For 
those who prefer books, the modern plants also appear in four recent works by Susan H. 
Munger and Charlotte Staub Thomas (2003), H. Wayne Phillips (2003), A. Scott Earle and 
James L. Reveal (2003), and Richard McCourt and Earle Spamer (2004). 


REFERENCES CITED 


COUES, E. 1898. Notes on Mr. Thomas Meehan’s paper on the plants of Lewis and Clark’s expedition 
across i continent, 1804-1806. Proceedings of the Academy of Natural Sciences of Philadelphia 50: 
291- 

EARLE, “ $i AND J. L. REVEAL. 2003. Lewis and Clark’s Green World: the Expedition and its Plants. 
Farcountry Press, Helena, Montana. 256 pp. 

EWAN, J. A. 1979. Introduction to the facsimile reprint of Frederick Pursh’s Flora America 
Septentrionalis (1814). Pp. 7-117 in F. T. Pursh, Flora Americae Septentrionalis (facsimile reprint, J. 


Cramer, Vaduz). 
HATCH, P. J. 2003. “Public treasures”: Thomas ooamine and the garden Say of Lewis and pote 
Twinleaf Journal (January 2003). Also posted on the web at articles/ 


(accessed 2004-12-02). 

McCourt, R. AND E. SPAMER. 2004. Jefferson’s Botanists: Lewis and Clark Discover the Plants of the 
West. Academy of Natural Sciences, Philadelphia. 25 pp 

McCourt, R. M., C. HAWKS AND E. E. SPAMER. 2002. The Lewis and Clark Herbarium of the 
Academy of Natural Sciences. Part 2. Saving an American treasure: preservation of the herbarium 
on the bicentennial of the expedition. Notulae Naturae, no. 476. 16 pp. 

MEEHAN, T. 1898. The plants of Lewis and Clark’s expedition across the continent, 1804-1806. 
Proceedings of the Academy of Natural Sciences of Philadelphia 50: 12-49. 

MOULTON, G. E. (ed.). 1990. The Journals of the Lewis and Clark Expedition, vol. 6: November 2, 1805- 
March 22, 1806. University of Nebraska Press, Lincoln and London. 531 pp. 

MOULTON, G. E. (ed.). 1999. The Journals of the Lewis and Clark Expedition, vol. 12: Herbarium of 
Lewis and Clark. University of Nebraska Press, Lincoln and London. 359 pp. 

MUNGER, S. H. AND C. S$. THOMAS. 2003. Common to This Country: Botanical Discoveries of Lewis and 
Clark. Artisan, New York. 114 pp. 

PHILLIPS, H. W. 2003. Plants of the Lewis and Clark Expedition. Mountain Press Publishing Company, 
Missoula, Montana. 288 pp. 

PURSH, F. 1813. Flora Americae Septentrionalis (facsimile reprint, J. A. Ewan, ed., 1979, J. Cramer, 
Vaduz). 868 pp. 

REVEAL, J. L., G. E. MOULTON AND A. E. SCHUYLER. 1999. The Lewis and Clark collections of 


102 BARTONIA 


vascular plants: Names, types, and comments. Proceedings of the Academy of Natural Sciences of 
Philadelphia 149: 1-64. 

SPAMER, E. E. AND R. M. MCCourtT. 2002a. The Lewis and Clark Herbarium, Academy of Natural 
Sciences of Philadelphia (PH-LC): Digital i ees study set. Academy of Natural Sciences of 
Philadelphia Special Publication 19. CD-R 

SPAMER, E. E. AND R. M. MCCourrT. ne we Lewis and Clark Herbarium of the Academy of 
Natural Sciences. Part 1. History. Notulae Naturae, no. 475. 46 pp. 

TEECE, M. A., M. L. FOGEL, N. TUROSS, R. M. MCCOURT AND E. E. SPAMER. 2002. The Lewis and 
Clark Herbarium of the Academy of Natural Sciences. Part 3. Modern environmental applications 
of a historic nineteenth century botanical collection. Notulae Naturae, no. 477. 20 pp. 


Bartonia No. 62: 103-105, 2004 


BOOK REVIEWS 


Profiles of Rafinesque, by Charles Boewe (ed.). 2003. University of Tennessee Press, 
Knoxville. xli + 411 pp. $45 (cloth). 


Constantine Samuel Rafinesque (1783-1840) considered himself a botanist, naturalist, 
geologist, geographer, historian, poet, philosopher, economist, philanthropist, traveler, 
merchant, manufacturer, collector, improver, professor, teacher, surveyor, draftsman, 
architect, engineer, author, and editor. We can add artist, realtor, banker, philologist, 
anthropologist, and who knows how many more. To a Boston psychiatrist, he was a 
paranoid neurotic genius. His overzealous demeanor did not ingratiate him with others, and 
he was often the subject of ridicule. He was gullible and devious, although the magnitude 
of the latter is subject to debate. At the end of his description of Rafinesquia (1841, 
Transactions of the American Philosophical Society 7: 429), Thomas Nuttall wrote “Dedicated 
to the memory of an almost insane enthusiast in natural history; sometimes an accurate 
observer, but whose unfortunate monomania was that of giving innumerable names to all 
objects of nature, and particularly to plants.” Rafinesque continues to attract the attention 
of botanists because of the thousands of plant names he authored, many of which, along 
with his extant specimens, still need evaluation. His controversial accomplishments and his 
eccentricities make him a colorful historical figure. His life and work are unending subjects 
of interest; hence, the book reviewed here. 

In his introduction, Charles Boewe states that the book was “assembled in the belief that 
it will in some measure supply the lack of the authoritative biography C. S. Rafinesque 
deserves but has not yet received.” There are 17 articles taken or revised from other 
publications and three written specially for this volume by the editor. The introduction, 
“Evolution of the Rafinesque Biography,” provides useful, sometimes bizarre, background 
information and an overview of the articles. 

The first and longest article is Francis Pennell’s “The Life and Work of Rafinesque.” This 
article and those on Rafinesque’s genealogy (Georges Reynaud), last days (Boewe), and 
portraits (Boewe) deal with Rafinesque “The Man.” The last two clear up misconceptions 
about the circumstances of his death and a “Rafinesque” portrait now thought to be someone 
else. Articles on his eccentricity (John James Audubon) and the occupant of his tomb 
(Boewe) deal with Rafinesque “The Legend.” Missing is an article on the organisms named 
for Rafinesque that continually bring him to our attention as a species of eternity. A tabular 
chronology of the major events of his life also is lacking. 

Seven articles (three by Boewe, and one each by Arthur J. Cain, Ronald L. Stuckey, Leon 
Croizat under the alias Henricus Quatre, and Elmer D. Merrill), dealing with Rafinesque 
“The Naturalist,” contain evaluations of his published work, his personality, and his 
interactions with others. Some of the diverse opinions expressed in these articles are as 
eccentric as those of Rafinesque are. Here the concern is with people and their idiosyncrasies 
as much as it is with natural history. 

Less familiar to botanists and naturalists is the work of Rafinesque discussed in articles 
on linguistic activity (Vilen V. Belyi), the Walam Olum hoax (David M. Oestreicher), Indian 


103 


104 BARTONIA 


languages (Boewe), and Mayan hieroglyphics (George E. Stuart). In the case of the Walam 
Olum, opinions differ on whether Rafinesque was the hoaxer or the gullible victim of a 
hoax. 

Finally, there are articles on Rafinesque’s Medical Flora (Michael A. Flannery), sentimental 
botany (Beverly Seaton), and fugitive publications (Boewe). Finding Rafinesque’s “fugitive” 
publications is an ongoing activity of bibliophiles. Missing is an article on the status of 
Rafinesque’s lost, extant, and “fugitive” collections, a subject of interest to many. 

The thing that disappoints me most about the book is the lack of documentation for 
many literature citations. This includes articles in the book previously published elsewhere. 
For the convenience of users, I prefer to see notes and references at the end of a book. In 
this book we have to look for notes in 16 different places and references in 7 places, where 
they are at the end of the articles. 

The diverse articles in this book give the reader a much more comprehensive overview 
of Rafinesque’s life than previously available. It will appeal to historians, naturalists, and 
those who cannot let go of Rafinesque. It is a major rung on the ladder toward an 
“authoritative biography.” 


ALFRED E. SCHUYLER, Curator Emeritus 
The Academy of Natural Sciences, Philadelphia 
November 2003 


Shrubs and Vines of New Jersey and the Mid-Atlantic States: a Field Identification and Natural 
History Guide, by Christopher T. Martine, illustrated by Rachel A. Figley. 2002. New Jersey 
Forest Service, Trenton. 114 pp. $10 (paper). (Available from the Forest Resource Education 
Center, 370 East Veterans Highway, Jackson, NJ 08527, 732-928-0987.) 


This new companion guide to the Trees of New Jersey and the Mid-Atlantic States, also 
written by Christopher Martine, provides useful aids to identifying the often-underap- 
preciated shrubs and vines. The shrubs and vines are important members of the diverse 
habitats found in the mid-Atlantic region, especially in successional communities. New Jersey 
has a rich and complex flora due primarily to the geology but compounded by its climate, 
which together allow many species to persist that are at the northern or southern edge of 
their range. Some of the most difficult groups taxonomically are shrubs (e.g., Crataegus, Rosa, 
Rubus, Salix, Viburnum), and thus providing a treatment for an audience of naturalists and 
amateur botanists is a challenge. Many of the difficult groups are simplified, with some of 
their diversity noted but not treated. In Crataegus, for example, C. uniflora Muench is treated 
but C. intricata Lange, which is reasonably common in the state, is only noted under 
“similar species.” Further, the notes indicate that 13 species of hawthorn exist in the state, 
11 of which are small trees, and none are treated in the Trees of New Jersey. A reference to 
hawthorns can be found in the “similar species” section under flowering crabapple (Malus 
spp.), but the information given is vague and in some cases incorrect. This may limit the use 
of these guides by the knowledgeable botanist. 


BOOK REVIEWS 105 


The text provides a clear description of each plant and highlights possible confusion with 
similar species. Illustrations provided are simple and clearly show the characters needed to 
identify the plant. I like the use of the entire page to provide text and illustrations, although 
the text could be slightly larger for those who have a vision impairment. Perhaps a second 
edition could use a slightly larger page format. 

The reader is also informed whether the plants are rare, native, exotic or invasive. 
Certainly in the case of invasive species the more knowledge we provide to nature 
enthusiasts about these plants, and especially how to tell them apart from their native 
relatives, the better. Although only a generalized locator key is provided at the beginning, 
supplemental keys to more difficult groups are provided, which in many cases are produced 
with the advice of experienced field botanists in the region. 

In summary, although Shrubs and Vines of New Jersey and the Mid-Atlantic States cannot 
be considered comprehensive, it does provide enough detail and interesting anecdotes for 
naturalists with beginner to intermediate botanical savvy. 


JAMES MACKLIN, Collection Manager 
The Academy of Natural Sciences, Philadelphia 
October 2004 


26 odd of | 
Mi TinistssO 
aden 


fe eee a Pees: Hy was 
ae iS te 


Bartonia No. 62: 107-111, 2004 


OBITUARIES 


Roy Linden Hill, Jr. (1913-2001) 


Roy L. Hill died after a short illness on 3 June 2001 in 
Newport, Vermont. Roy was born in Philadelphia on 16 
May 1913, the son of Mabel and Roy Hill. He lived his 
boyhood and teenage years in Philadelphia, Wilmington, 
and Tamaqua. He graduated from the Wilmington Friends 
School in 1931. Many summers of his early years were spent 
at the Newlin Grist Mill outside Media, which was operated 
by his great-grandfather and grandfather. He received 
degrees in chemistry from the University of Delaware and 
in business from the Wharton School of the University of 
Pennsylvania. His professional career was spent at the Rohm 
and Haas Company in Philadelphia. Roy and his wife 
Margaret (Peg) Elizabeth Thatcher Hill became members of 
the Philadelphia Botanical Club in 1963. He was for some years a volunteer in the Botany 
Department of the Academy of Natural Sciences, during which time he completed numerous 
projects for the staff. He served as treasurer of the Club from 1983 until 1994. It was 
through Roy’s endeavors that grants were obtained and funds raised in support of the Club’s 
centennial programs in 1991. Excess monies from those events became the basis of the Club’s 
Endowment Fund. Roy was made an Honorary Member of the Philadelphia Botanical Club 
in 1999, 

In the 1980s, upon Roy’s completion of the course of study in botany and horticulture 
at the Arboretum of the Barnes Foundation, he served as a volunteer there. I was privileged 
to work with him for several years as he engraved plant labels (a tedious chore) and placed 
them on trees and shrubs throughout the Arboretum grounds. One was left breathless in 
trying to match his energy and devotion to the tasks he carried out. 

Roy’s volunteer service to diverse groups was awe-inspiring. The well-known slogan of 
the U.S. Postal Service, “neither snow, nor rain, nor heat ... stays these couriers from ... their 
appointed rounds” could have been applied to Roy. He was punctilious in meeting his 
schedules at various organizations. He was an Arboretum Assistant at the Scott Arboretum 
of Swarthmore College from 1987 to 1999. He was involved with many civic groups in the 
Lansdowne area where he lived—the Lansdowne Allied Youth Council, the Lansdowne 
Street Tree Committee, the Boy Scouts, the park system, and other groups. Roy had a strong 
commitment to the preservation of historic trees. 

Roy and Peg had a deep and abiding passion for Vermont, where eventually they built a 
small vacation house in Barton. He was an enthusiastic student of the local flora in the 
Northeast Kingdom. He and his wife enticed a number of friends from the Delaware Valley 
to visit the area. My husband and I went there twice to be led down trails and up mountains 


107 


108 BARTONIA 


in pursuit of particular plants that Roy wanted to show us. I particularly remember being 
taken by him to large patches of Cornus canadensis and Linnaea borealis. Roy and Peg, who 
predeceased him, were most generous in sharing their world with others. 

Roy is survived by two children, Richard W. Hill of East Lansing, Michigan, and Janet 
L. Hill of Barton, Vermont. He is also survived by two grandchildren, David Hill and 
Christine Hill. 

ELIZABETH B. FARLEY 
October 2003 


OBITUARIES 109 


James C. Parks (1942-2002) 


The passing of James C. Parks 
of Millersville, Pennsylvania, 
just prior to Christmas, 2002, 
came as sad news for students 
he mentored and his colleagues. 
His ability to contribute to 
plant systematics and his will- 
ingness to share knowledge 
with students, colleagues, and 
the lay public is a genuine loss 
for botanical science. Jim cher- 
ished plants early in life and 
the botanical realm was woven 
into the very fabric of his soul; 
even his daughters have botani- 
cal names: Holly and Heather. 

Due to retire from Millers- 
ville University in the spring of 
2003, Jim acknowledged, “Far 
more than my several research 
papers, you, my students, are 
my legacy.” However, he will 
influence students beyond the 
classroom through his contri- 
bution (along with Jim Mont- 
gomery) of the fern section to 
The Plants of oo an 
Illustrated Manual by 

Fowler Rhoads and Timothy 
A. Block. At the time of his death, he was working on a taxonomic treatment of Melanthera 
(Asteraceae) for the Flora of North America and had planned to co-author a Field Guide to the 
Fern Flora of Pennsylvania with Jim Montgomery. Second to his passion for family, friends, 
and students, botany was an integral part of Jim’s life. It was not unusual for Jim to devote 
vacation time to visit field sites. Jim generously shared his knowledge with others and 
contributed frequently to local organizations such as the Muhlenberg Botanical Society and 
the Native Plants in the Landscape Conference at Millersville. His neighbors appreciated his 
concern for conservation of our natural heritage and welcomed his efforts in combating 
establishment of industrial farms and developments. 

Jim was born on 9 August 1942 in Altoona, Pennsylvania, and grew up on a small farm 
at the edge of the mountains on the Allegheny Front. Just beyond the back forty of the 
family farm, Jim often hiked in the mountains for solace and recreation; he was introduced 
to botany through first-hand experience at an early age. Later in life, he worked hard to have 
his students share that same sense of discovery. While it may be difficult to document the 
initiation of a career, Jim’s professional focus was clearly noted in one of his papers from 


110 BARTONIA 


grade 6 where he stated that he wanted to be a biology teacher. He has devoted most of his 
professional career to this goal. 

With limited resources and a desire to be a teacher, he used his skill in animal husbandry 
to enable him “to ride a pig to college every fall”—sale of a hog each fall paid his college 
tuition. In addition, he used the family combine to custom-cut small grains for neighboring 
farmers. Jim found that keeping a small tractor-pulled combine in one piece on small, 
unfamiliar fields was a real challenge. Proceeds from custom harvesting also helped to pay 
his college tuition. 

After earning his undergraduate B.S. degree from Shippensburg State Teachers College in 
Pennsylvania, Jim taught science for one year in Seneca Falls, New York. Then he proceeded 
to Vanderbilt University in Nashville, Tennessee for a Ph.D. Under the guidance of Robert 
Kral he revised the genus Melanthera Rohr (Asteraceae) for his thesis; this work was 
published in 1973 (Rhodora 75: 169-210). Jim joined the faculty at Millersville University in 
1968 where he taught for 34 years. As a cornerstone in the Concepts of Botany course at 
Millersville, he fostered student interest in other courses he taught such as Lower Plants, 
Plant Systematics, and Dendrology. Moreover, he developed a Methods of Teaching 
Controversial Issues seminar to help students preparing to become teachers wrestle with 
socially sensitive issues such as evolution and biotechnology. Early on Jim recognized the 
role of mentors in learning and took pride in being a mentor to students with a serious 
interest in botany. Second to mentorship, Jim recognized field experience as a primary 
educational tool. He delighted in having his students learn from the paradigms evident in the 
field; this was reflected in his enthusiasm for nature, which he learned while walking in the 
mountains on the Allegheny Front as a youth. 

Although Jim’s professional life was grounded in teaching and introducing students to the 
field, he devoted considerable effort to the infrastructure of the University that would have 
direct impact on his efforts as a teacher and scientist. He was active in developing the 
glasshouse and herbarium facilities at Millersville University. As the sole curator of the 
Millersville University Herbarium, he expanded the collection from a few cabinets to 
thousands of specimens in a room of compactors. It is fitting that the Herbarium will be 
named in his honor. 

Over the past few decades, Jim became interested in pteridophytes. This interest may have 
been the result of Jim’s participation in a field course given in 1978 by Don Farrar at the 
University of Virginia’s Mountain Lake Biological Station. Shortly after the field course, Jim 
worked with Don on locating fern gametophytes in the Northeast. They published (with 
Bruce McAlpin) the first findings of Vittaria and Trichomanes gametophytes in Pennsylvania 
(Rhodora 85: 83-92). In 1984, they reported the most northern record for Vittaria 
gametophytes (Rhodora 86: 421-423). In a subsequent publication (Rhodora 91: 201-206), Jim 
presented an intensive study of the distributions of Vittaria and Trichomanes including many 
new records. In addition, Jim used allozyme analysis to study the largest genets known (1.1 
km? in spatial extent) for bracken fern in Virginia. This work was published in 1993 with 
Charlie Werth in the American Journal of Botany (80: 537-544). Later that year, Jim and his 
wife Vicki traveled to the U.K. to work with Adrian Dyer and Stuart Lindsay at the Royal 
Botanic Gardens in Edinburgh, Scotland. There he used allozymes and frond and spore 
morphology in Cystopteris to determine if C. dickieana was, in fact, a separate species; it was 
not (Edinburgh Journal of Botany 57: 83-105). During this sabbatical leave, Jim traveled with 
Adrian to the tenth Simposio Nacional de Botanica Criptogamica at La Laguna, Tenerife, 
to collect material and talk to several Spanish pteridologists. Back in the States, Jim worked 


OBITUARIES 111 


with Kate Moser, an undergraduate at Millersville, on a study of Cystopteris tennesseensis 
(Pennsylvania Academy of Sciences 71: 78-83); this species had never before been reported in 
Pennsylvania. We will miss the unwritten chapters of Jim’s forays into pteridophytes. 

Throughout his career, Jim was concerned with the loss of biodiversity and generously 
gave time and wisdom to a variety of agencies. In 1994, Jim went to Edinburgh to participate 
in a meeting on the Ecology and Conservation of Scotland’s Rare Ferns. In Pennsylvania, 
he was an active member of the Vascular Plant Technical Committee of the Pennsylvania 
Biological Survey, which advises the Pennsylvania Department of Conservation and Natural 
Resources on the status of rare and endangered plants in the state. As a member of the 
Shenks Ferry Wildflower Preserve Advisory Committee, he developed educational materials 
and helped with the management of this valuable resource. His legacy will be evident in the 
natural heritage of southeastern Pennsylvania. 

Students and colleagues will be indebted for his insights on humanity and the natural 
world. We will miss his thought-provoking comments, historic perspective, and wry sense 
of humor. He will rest easily knowing that his students are now mentors for the next 
generation of botanists. 

Guy L. STEUCEK 
Department of Biology 
Millersville University 
Millersville, PA 17551 


Bartonia No. 62: 113-128, 2004 


2000-2002 FIELD TRIPS 


by TED GORDON, Southampton, New Jersey 
except where otherwise indicated 


2000 Field Trips 


6 May: Bowmans Hill Wildflower Preserve, Bucks County, Pennsylvania. This trip to the 
wildflower preserve was during the height of the spring wildflower bloom. We walked 
various trails at the preserve and observed the following species: Mertensia virginica, Trillium 
grandiflorum, Trillium cernuum, Trillium cuneatum, Trillium erectum, Trillium luteum, 
Trillium sessile, Viola sororia, Viola striata, Uvularia sessilifolia, Antennaria plantaginifolia, 
Antennaria solitaria, Packera aurea, Caulophyllum thalictroides, Jeffersonia diphylla, 
Podophyllum peltatum, Arabis laevigata, Cardamine concatenata, Silene virginica, Stellaria 
pubera, Sedum ternatum, Carex pensylvanica, Carex prasina, Euphorbia purpurea, Cercis 
canadensis, Lupinus perennis, Adlumia fungosa, Dicentra cucullaria, Geranium maculatum, 
Hydrophyllum canadense, Hydrophyllum virginianum, Meehania cordata, Erythronium 
americanum, Maianthemum canadense, Maianthemum racemosum, Polygonatum biflorum, 
Polygonatum pubescens, Veratrum viride, Floerkea proserpinacoides, Sanguinaria canadensis, 
Stylophorum diphyllum, Phlox stolonifera, Polemonium reptans, Claytonia virginica, Actaea 
pachypoda, Aquilegia canadensis, Caltha palustris, Hepatica nobilis var. obtusa, Thalictrum 
thalictroides, Houstonia caerulea, Mitella diphylla, Tiarella cordifolia, and Dirca palustris. Report 
by leader: Bill Olson. 


3 June: Pine Barrens of Atsion and vicinity, Burlington County, New Jersey. Joint trip with 
the New Jersey Audubon Society. The morning was spent along Quaker Bridge Road near 
the Atsion Ranger Station, where a mix of habitats produced the expected species of 
Gaylussacia and Vaccinium, as well as Quercus stellata and Q. marilandica. Lowland woods 
along the Wesickaman Creek had extensive stands of Osmunda cinnamomea, O. regalis, 
Thelypteris palustris, and Woodwardia areolata. Sedges seen here included Carex intumescens, 
C stricta, C. stipata, C. striata, C. vulpinoidea, C. folliculata, and C. canescens. A variety of 
grasses was found, including Danthonia sericea, Panicum scoparium, and the distinctive 
Piptochaetium avenaceum (= Stipa avenacea). Also seen were Polygonatum biflorum, 
Chimaphila maculata, and Akebia quinata (chocolate-vine), this last species on a tree near the 
ranger station where it has been for years without spreading. After lunch, the group moved 
south a few hundred yards to explore some grassy fields south of the Mullica River. Here 
we found large patches of Minuartia caroliniana (= Arenaria c.) in bloom, as well as Krigia 
virginica, Linaria canadensis, and Opuntia humifusa. A large patch of Iris prismatica, still in 
reasonably good bloom on the east side of Route 206, was much admired. Explorations along 
the west side of Route 206 were not particularly productive, with some areas flooded, other 
areas desiccated, and still others mowed. We did find Utricularia subulata in bloom in a wet 
swale with Proserpinaca pectinata and Drosera intermedia. Attendance: 18. Leader: Karl 
Anderson. 


113 


114 BARTONIA 


10 June: Lonely Road Meadow, Sellersville, Buck County, Pennsylvania. Fifteen hardy souls 
braved unseasonably hot weather to visit a floodplain meadow along the East Branch 
Perkiomen Creek near Sellersville. They were treated to the sight of a large population of 
slender blue iris (Jris prismatica) in full bloom. Iris prismatica, which has a coastal 
distribution ranging from Nova Scotia to Georgia, is currently known from only three sites 
in Pennsylvania and is classified as an endangered species in the state. Other rarities seen on 
the trip included downy phlox (Phlox pilosa) and brown sedge (Carex buxbaumit), both with 
a recommended status of endangered, and cloud sedge (Carex haydenit), recommended as 
endangered. Mead’s sedge (Carex meadii), another plant that has been recommended for 
endangered status that was not previously known from the site, was also documented. 
Altogether 13 species of Carex were identified in the meadow. Annual mowing by the 
owners maintains this diverse herbaceous community that was formerly a hayfield. Report 


by leader: Ann Rhoads. 


17 June: Tacony Creek Park, Philadelphia County. We entered the park close to the 
junction of Cheltenham Avenue, Crescentville Road, and Tacony Parkway near the 
Montgomery/Philadelphia county line. On the west side of Tacony Creek there were many 
non-native trees (including Acer platanoides, Ailanthus altissima, Paulownia tomentosa, Prunus 
avium) and shrubs (including Aralia elata, Ligustrum obtusifolium). The ground cover was 
primarily Alliaria petiolata and Lonicera japonica. On the east side of the creek, the forest 
was dominated by native trees (Fagus grandifolia, Quercus rubra, Q. velutina), and shrubs 
(Cornus florida, Corylus amricanus, Viburnum acerifolium, Rhododendron periclymenoides). 
Many more native vines and herbs that were not encountered on the west side included 
Amphicarpa bracteata, Anemone quinquefolia, Apocynum cannabinum, Eurybia divaricata (= 
Aster divaricatus), Athyrium felix-femina, Carex laxiculmis, C. laxiflora, C. pensylvanica, 
Cimicifuga racemosa, Circaea lutetiana, Collinsonia canadensis, Cypripedium calceolus, 
Danthonia spicata, Desmodium paniculatum, D. perplexum, Dioscorea villosa, Elymus hystrix, 
Eupatorium fistulosum, Festuca obtusa, Geranium maculatum, Heuchera americana, Lonicera 
sempervirens, Luzula multiflora, Lysimachia ciliata, Monotropa uniflora, Osmorhiza longistylis, 
Panicum cladestinum, Phegopteris hexagonoptera, Podophyllum peltatum, Polygonum 
virginianum, Sanguinaria canadensis, Sanicula canadensis, Scutellaria elliptica, Silene stellata, 
Smilacina racemosa, Smilax glauca, S. rotundifolia, Solidago caesia, and Thelypteris novabora- 
censis. The striking floristic differences between the east and west sides of the creek suggest 
that plants in late-successional forests can outcompete most non-native plants. Also there was 
less evidence of deer browsing here than we have seen in other parts of Fairmount Park, 
although some leaves of Podophyllum were browsed. Report by leader: Ernie Schuyler. 


18-22 June: Sage College, Albany, Albany County, New York. Joint field meeting of the 
Philadelphia Botanical Club, the Torrey Botanical Society, and the Northeast Section of the 
Botanical Society of America. Monday field trips began at the Albany Pine Bush Preserve, 
a 2,600-acre protected remnant of a pitch pine-scrub oak community that covered tens of 
thousands of acres in colonial times. Carefully controlled burns now attempt to duplicate 
the historic “natural” conditions and maintain the area’s unique and diverse flora. In the 
afternoon, a trip to the escarpment overlooking the Hudson Valley and Albany gave an 
opportunity for about half of the group to find lime-loving plants among fossils of the 
Devonian limestones and shales, while the other half joined special trips for ferns, lichens, 
or graminoids. Tuesday was spent at the famed Ice Meadows along the northern Hudson 


2000-2002 FIELD TRIPS 115 


River, Warren County. Ice buildup on the shores over the winter and floodwaters of early 
spring create natural meadow conditions here, which support a very diverse plant 
community. Various orchids, lilies, grasses, and sedges that are rare or missing in the rest of 
New York were observed here. Trips on Wednesday were to a 300-acre nature preserve 
owned by Skidmore College, Saratoga County. It has a rich and varied flora, due to a 
combination of limestone ledges, limy soils, and non-limy glacial deposits. This preserve has 
a large population of Hydrastis canadensis, unusual in that there are thousands of healthy 
plants. This plant is rarely observed in eastern upstate New York. Fern experts in the group 
added two unusual Botrychium species, so the Skidmore fern list now stands at 30, not 
counting sterile hybrids. On the way back to Albany a brief stop was made to see 500- 
million- year-old fossilized blue-green alga formations. The last stop was at a town park with 
a very healthy stand of lupine and a highly visible host of small blue butterflies, probably 
the rare and federally-protected Karner blue. Participants: 42. Chairperson: Edward Miller. 
Report by Karl Anderson. 


15 July: Pleasant Mills and Batsto back-country, Wharton State Forest, Atlantic and 
Burlington Counties, New Jersey. At Nescochague Bog, an open, savanna-like swale of 
hummocks between the creek and West Mill Road, about 1.75 miles northwest of Pleasant 
Mills Church, we observed a large Narthecium americanum population, discovered by the 
leader on 16 August 1996. Whether this occurrence is identical to the “Pleasant Mills” 
collections by J. Darrach in 1861 and C. F. Parker in 1877 can never truly be known. 
Among the associated species were Carex exilis, Sphagnum pulchrum, Schizaea pusilla, Scleria 
muhlenbergii, Lobelia canbyi*, Mublenbergia torreyana, Calamovilfa brevipilis, Rhynchospora 
cephalantha, R. gracilenta, R. alba, R. fusca, Carex livida, Lophiola aurea*, Sabatia difformis*, 
Pogonia ophioglossoides*, and Utricularia striata*. Although the 29 July 1997 wildfire has 
slowed shrub and cedar sapling invasion, it may have expunged a fine occurrence of 
Rhynchospora oligantha, discovered here in 1996. In a streamside swale dominated by 
Mublenbergia torreyana, ca. 0.2 mi. to the northwest, we saw five plants of Asclepias rubra’, 
Lobelia canbyi*, Carex livida, Hypericum densiflorum™, H. denticulatum*, H. canadense*, 
Schoenoplectus pungens var. pungens, Pontederia cordata”*, Dioscorea villosa, Iris prismatica, and 
a single specimen of Platanthera blephariglottis about to bloom. We stopped at Forge Pond 
to the south to examine club-mosses: Lycopodiella alopecuroides, L. appressa, L. caroliniana var. 
caroliniana (= Pseudolycopodiella caroliniana), and L. xcopelandit (L. alopecuroides x L. 
appressa). In the afternoon we visited “Batsto Oxbow,” a sphagnous savanna of hummocks 
on the east bank of the Batsto River about 1.5 miles north of the village. Among scattered 
clusters of cedar and hardwood, the site contained some five pockets of bog asphodel, an 
occurrence reported in 1996 by the leader to the New Jersey Heritage Program. There is no 
doubt that this savanna is the site of Chrysler’s 1930 collection of Narthecium designated as 
“frequent,” and likely that of earlier botanists in the second half of the nineteenth century. 
At its downstream border, a backwater cove enables portions of the savanna to flood 
periodically. This flushing appears to have held in check invasion by trees and shrubs and 
has helped to maintain habitat stability over many decades. All species listed above for 
Nescochague Bog were seen here as well. In addition, we saw Carex collinsi, Juncus 
caesariensis, Rhynchospora oligantha, Tofieldia racemosa”, Eleocharis tuberculosa*, Utricularia 
cornuta*, U. purpurea”, U. geminiscapa*, and U. juncea*. Thanks to William F. Standaert for 
maintaining a comprehensive list of species seen. (* = plants in anthesis.) Attendance: 12. 


Leader: Ted Gordon. 


116 BARTONIA 


22 July: Old Mine Road, Sussex and Warren Counties, New Jersey. The morning of this trip 
was spent in the vicinity of the seventeenth-century copper mine at Pequaharry and along 
the shore of the Delaware River nearby. Some interesting plants included Thelypteris 
phegopteris, Phyrma leptostachya, Acer spicatum growing next to Acer pensylvanicum, Sium 
suave, Equisitum fluviatile, and Lycopodium hickeyi (= L. obscurum var. isophyllum) growing 
near L. obscurum var. obscurum. After lunch, we drove north to the Flatbrook, where we 
found Cystopteris bulbifera in profusion, as well as Asplenium rhizophyllum, Thelypteris 
hexagonoptera, Equisetum hyemale, and several other ferns and fern allies. Sedges seen here 
included Carex hystericina, C. lupulina, C. intumescens, C. lurida, C. argyrantha, and C. stricta. 
In bloom were Cimicifuga racemosa, Myosotis scorpioides, Mimulus ringens, and Alisma 
subcordatum. The final stop was at a rocky hillside about four miles north of the Flatbrook, 
where we found Cystopteris fragilis, C. protrusa, Matteucia struthiopteris, and Athyrium 
thelypteroides. We also puzzled over the wide-leaved, glaucous, basal rosettes of Carex 
platyphylla and took note of Heuchera americana, Ulmus rubra, and Staphylea trifolia. 
Attendance: 5. Leader: Karl Anderson. 


29 July: Pine Barrens at Hog Wallow, Rutgers Experimental Station, and Buck Run Vicinity, 
Burlington County, New Jersey. Attracted by the white spikes of several tall Platanthera 
blephariglottis, we stopped to examine the road shoulder on the east side of Route 563 
between Hog Wallow and Pineworth. A species of a bulrush rare in the Pine Barrens, Scirpus 
georgianus (= S. atrovirens var. georgianus) has persisted here for many years. We noted in 
flower Chamaecrista nictitans, Desmodium ciliare, Drosera rotundifolia, Eupatorium pilosum, 
Euthamia graminifolia var. graminifolia, Hypericum canadense, Erigeron annuus, Hypochoris 
radicata, Lachnanthes caroliniana, Solidago odora, and, about to flower, Juncus scirpoides, J. 
acuminatus, and J. debilis var. debilis. After visiting the Rutgers University laboratory and 
greenhouse facilities at the Phillip E. Marucci Cranberry/Blueberry Research Center on 
Oswego Lake Road (= Penn Place Road), we botanized the border of a reservoir and canal 
adjacent to the experimental cranberry bogs. Here we saw a few Platanthera blephariglottis 
and several P. cristata with deep orange flowers and an isolated cluster bearing light yellow 
flowers. A single orchid appeared to be P. xcanbyi (P. cristata x P. blephariglottis), bearing 
a whitish yellow plume and a spur about as long as the ovary. Also in flower were 
Cephalanthus occidentalis, Hypericum crux-andreae (= H. stans), H. hypericoides, H. gentianoides, 
Cuscuta gronovii, Apocynum cannabinum, Decodon verticillatus, Drosera intermedia, Lobelia 
nuttallii, and Ilex laevigata in immature fruit. At Harrisville in the Wharton State Forest, we 
saw the golden rays of a long-established occurrence of Pityopsis falcata (= Chrysopsis f.) in 
bare stretches of sand. Other dry sites had Scleria triglomerata, Danthonia spicata, D. sericea 
and Euphorbia ipecacuanhae. Northeast of Martha Furnace in the savannas and cedar swamps 
associated with Buck Run, we saw—already past flowering—Orontium aquaticum, Pogonia 
ophioglossoides, Carex livida, Trientalis borealis, Narthecium americanum, Tofieldia racemosa, 
Danthonia epilis, and Eriocaulon compressum. In flower were our three Drosera, Eriocaulon 
decangulare, Platanthera clavellata, Xyris difformis var. difformis, Utricularia cornuta, U. 
geminiscapa, U. striata, Sarracenia purpurea, Polygala brevifolia, P. cruciata, Sabatia difformis, 
and a few Rhynchospora alba, R. chalarocephala, and R. gracilenta. Also noted were Juncus 
caesariensis, Scirpus cyperinus, Schoenoplectus subterminalis (= Scirpus s.), Oclemena nemoralis 
(= Aster n.), Smilax laurifolia, and Schizaea pusilla. We recorded two significant herpetologi- 
cal sightings: a mature northern pine snake (Pituophis mel ) in the water 
of a cedar-hardwood swamp above Buck Run and a timber rattlesnake (Crotalus horridus 


2000-2002 FIELD TRIPS 117 


horridus) crossing the sand road south of Martha. The latter was only observed (and 
photographed) by Dan Jassby and Boel Denne-Hinnov. Thanks to William F. Standaert for 
compiling a comprehensive list of species observed. Attendance: 31. Leader: Ted Gordon. 


30 July: McCarthy’s Lake, Franklin Township, Gloucester County and Cedar Lake, Buena 
Vista Township, Atlantic County, New Jersey. We met in the parking lot east of Piney 
Hollow Road in anticipation of walking the dry pond bottoms of the old cranberry bogs 
that are McCarthys Lake. Since all lakes in the area were filled to capacity, most of us 
elected to focus on the edges of this impoundment. Here we noted Eleocharis tuberculosa, 
Rhexia virginica, Proserpinaca palustris, Myriophyllum humile, Wolffia columbiana, Lemna 
minor, Lindernia dubia var. anagallidea, and many common Pine Barrens plants. In the 
afternoon, we examined Cedar Lake Wildlife Management Area. The lake was full of water 
and only the tips of Juncus militaris were exposed. The dense mats of Rhynchospora scirpoides 
seen last year were reduced to scattered individuals, hidden by the lush, emergent rush flora. 
A colony of Eupatorium resinosum seen last year was relocated, but it too was considerably 
less abundant. These dramatic fluctuations in population size over the past two years were 
clearly related to the cyclic events of severe drought and flooding. Other species observed 
included Cephalanthus occidentalis, Galium tinctorum, Gratiola aurea, Lachnanthes caroliniana, 
Ludwigia sphaerocarpa, Potamogeton natans, Lysimachia terrestris, Eleocharis acicularis, 
robbinsii, Rhynchospora scirpoides, R. chalarocephala, R. fusca, R. macrostachia, R. capitellata, 
R. alba, Utricularia purpurea, and U. geminiscapa. Leader: Joe Arsenault. 


26 August: Bowmans Hill Wildflower Preserve, Bucks County, Pennsylvania. A nice 
collection of ferns is featured at the wildflower preserve. The group saw Onoclea sensibilis, 
Osmunda claytoniana, Osmunda cinnamonea, Osmunda regalis var. spectabilis, Thelypteris 
noveboracensis, Thelypteris palustris var. pubescens, Thelypteris simulata, Athyrium filix-femina, 
Diplazium pycnocarpon, Deparia acrostichoides, Polystichum acrostichoides, Polystichum braunii, 
Dryopteris campyloptera, Dryopteris marginalis, Dryopteris carthusiana, Dryopteris intermedia, 
Dryopteris x triploidea, Cystopteris tenuis, Cystopteris protusa, Cystopteris bulbifera, Adiantum 
pedatum, Woodwardia areolata and Woodwardia virginica. Report by leader: Bill Olson. 


27 August: Great Bay Wildlife Management Area, Ocean County, New Jersey. Joint trip 
with the New Jersey Audubon Society. At Tip Seaman County Park in Tuckerton, we 
found Tripsacum dactyloides, Juncus acuminatus, J. pelocarpus, and Schoenoplectus pungens (= 
Scirpus p.). At the salt marshes along Great Bay Boulevard, we found Schoenoplectus robustus 
(= Scirpus r.), Fimbristylis caroliniana or F. castanea, Spartina cynosuroides, Amaranthus 
cannabinus, Sabatia stellaris, Symphyotrichum tenuifolium (= Aster tenuifolius), Typha 
angustifolia, Samolus floribundus, Eleocharis rostellata, and Pluchea odorata, in a small pocket 
of high marsh. Continuing south, we examined a lush growth of Toxicodendron radicans on 
a Native American shell mound, and looked briefly but unsuccessfully for Polygonatum 
biflorum, noted on this mound in previous years. We did find Salicornia bigelovit, S. europaea, 
and S. virginica here, as well as the ever-present Spartina alterniflora, S. patens, and Distichlis 
spicata. As we neared the shore of Great Bay, plants of beach and strand included Ammophila 
breviligulata, Salsola kali, Suaeda linearis, Bassia hirsuta, Cakile edentula, Cyperus filicinus, C. 
esculentus, Eupatorium hyssopifolium, E. album, and Strophostyles helvola. On our final stop 
we visited nearby Stafford Forge Wildlife Management Area. Here Eupatorium leucolepis, 
Polygala cruciata, Chrysopsis mariana, Gratiola aurea, Xyris difformis, and Oclemena nemoralis 


118 BARTONIA 


(= Aster n.) were in bloom, and Juncus biflorus, Vaccinium macrocarpon, Drosera intermedia, 
Sarracenia purpurea, and other species typical of the Pine Barrens were found. Attendance: 
20. Leader: Karl Anderson. 


30 September: Monmouth County exploration, New Jersey. We visited three sites in Howell 
Township. A wetland area off Louise Drive was a pitch pine lowland including Scleria 
minor, Rhynchospora torreyanum, and Sphagnum compactum. At Shark River Station the 
group saw Scleria triglomerata, Rhynchospora torreyanum, Aletris farinosa, and Sphagnum 
pylaesit After lunch, we took a short walk on a the trail around the Manasquan Reservior. 

capitellata, and 


F ye 7 


Solidago odora. ~ at leader: Bill Olson. 


2001 Field Trips 


20 January: Winter Botany Walk in the Pinelands of New Jersey. Cancelled due to snow. 
Leader: Bill Olson. 


5 May: Nockamixon State Park, Bucks County, Pennsylvania. A small group of six 
participants gathered to explore the south shore of Lake Nockamixon in Nockamixon State 
Park, Bucks County. The group traversed the slope along the lake looking at large stands 
of native yew (Taxus canadensis). Unfortunately the decline of eastern hemlock (Tsuga 
canadensis) in the canopy due to infestation by hemlock woolly adelgid is raising some 
question as to whether the Taxus will survive at this site. A trek along the lakeshore led to 
a stand of nodding trillium (Trillium cernuum) in full bloom. On the way we stopped to 
admire blunt-leaved hepatica (Hepatica nobilis var. obtusa), bloodroot (Sanguinaria canadensis), 
narrow-leaved toothwort (Cardamine angustata), yellow fumewort (Corydalis flavula), false 
mermaid (Floerkia proserpinacoides), wild-ginger (Asarum canadense), wild sarsaparilla (Avalia 
nudicaulis), doll’s-eyes (Actaea pachypoda), creeping phlox (Phlox subulata), plantain-leaved 
pussytoes (Antennaria plantaginifolia), and other spring wildflowers. Several sedges were also 
identified on the wooded slopes including Carex pensylvanica, C. albicans, C. laxiflora, and 
C. albursina. At the edge of the parking lot white ash (Fraxinus americana) and hackberry 
(Celtis occidentalis) were in full glorious bloom. Report by leader: Ann Rhoads. 


26 May: Menantico and Peaslee Wildlife Management Areas (W.M.A.), Cumberland County, 
New Jersey. The trip began at the intersection of Route 49 and Union Road (Route 671) 
adjacent to Cumberland Pond. The group walked a few hundred yards to the west along 
Route 49 and botanized along the roadside. Here the group saw a mixture of oak species: 
Quercus alba, Q. coccinea, Q. falcata, Q. ilicifolia, Q. marilandica, Q. montana (= Q. prinus), 
Q. stellata, and Q. velutina. The three pines of the pine barrens, Pinus rigida, P. echinata, and 
P. virginiana, were also noted. The highlight along the roadside was a specimen of 
Chionanthus virginiana in full bloom. The group then visited Menantico Ponds W.M.A. This 
small preserve (349 acres) is situated south of Route 49 in the Menantico Creek watershed 
adjacent to railroad tracks. Much of it is comprised of abandoned sand mining pits. On both 
sides of the tracks were vast expanses of exposed sand, and here large stands of Hudsonia 
tomentosa, some in bloom, were noted. Mixed in were stands of H. ericoides, and some 
material appeared intermediate between the two species, suggesting hybridization. Just east 


2000-2002 FIELD TRIPS 119 


of Route 55, a sandy, open field was botanized. Here the group also noted good cushions 
of the two Hudsonia species, as well as Carex pensylvanica, Carya pallida, Helianthemum 
canadense, H. propinquum, Mirabilis nyctaginea, Panicum addisonit, P. meridionale, Petrorhagia 
prolifer, and Quercus prinoides. In dry pine woods nearby, Polygonatum biflorum an 
Uvularia sessilifolia were noted. We studied the differences between Toxicodendron radicans 
(poison-ivy) and T. pubescens (poison-oak), both growing along the railroad tracks. Just west 
of the Menantico Creek crossing was a stand of Lomnicera sempervirens in bloom. At 
Cumberland Pond we saw a few specimens of Quercus michauxii. In a bog in back of the 
pond along the Lawrens Branch were excellent stands of Arethusa bulbosa and Eriocaulon 
compressum in bloom. In the adjacent oak-pine woods we saw stump sprouts of Castanea 
dentata. Our last stop was an Atlantic white-cedar swamp in the headwater of the Middle 
Branch in the Peaslee W.M.A., a preserve that now totals more than 25,000 acres. Here we 
saw the state’s largest specimen of Chamaecyparis thyoides, with a circumference of over 10 
feet.. Other species noted were Carex collinsii, C. atlantica ssp. capillacea (= C. howe), 
Chionanthus virginicus, Ilex laevigata, Smilax laurifolia, and Trientalis borealis. Attendance: 
9. Leader: Gerry Moore. 


2 June: Fairmount Park, southwest of the Recycling Center and sorthwest of Chamounix 
Mansion, Philadelphia County. Our field survey produced the following list of species: Carya 
cordiformis, Phellodendron levallei, Poa triviale, Poa sp., Athyrium filix-femina, Ilex crenata, 
Parthenocissus quingucfolia, Viburnum obiegromiech si srangeareseit racemosum ssp. racemosum 
(= Smilacina racemosa), P. phy nolia acuminata, L it , Galium 
aperine, Cedrela sinensis (= Toona s.), Liguidambar | sntifia. Impatiens capensis, Symplocarpa 
foetidus, Circaea quadrisulcata, Rubus allegheniensis, R. phoenicolasius, Prenanthes altissima, 
Eupatorium pupureum, Osmorhiza longistylis, Parmelia sulcata, Physcia tenella, Vitis vulpina, 
Liriodendron tulipifera, Carex blanda, C. radiata, Allium vineale, Solidago canadensis, Rosa 
multiflora, Dioscorea villosa, Ailanthus altissima, Polygonum virginianum, Duchesnia indica, 
Sanicula canadense, Sambucus canadensis, Acer negundo, Pilea pumila, Philadelphus coronarius, 
aster sp., Eurybia divaricata (= Aster divaricatus), Tilia sp., Oxalis stricta., Prunus avium, 
Bromus arvensis, Hedera helix, Arisaema triphyllum, Rumex crispus, Gymnocladus dioica, 
Ranunculus ficaria, Dactylis glomerata, Lepidium virginicum, Artemesia vulgaris, Polygonum 
perfoliatum, Geum canadense, Celastrus orbiculatus, Polygonum cuspidatum, Leersia virginica, 
Quercus rubra, Ligustrum obtusifolium, Viburnum dentatum, V. acerifolium , V. prunifolium, 
V. plicatum, Cystopteris fragilis, Platanus hybrida, Acer psuedoplatanus, Onoclea sensibilis, Viola 
cucullata, Hydrangea paniculata, Juncus tenuis, Poa trivialis, Hamamelis virginiana, Phytolacca 
americana, Boehmeria cylindrica, Thelypteris novaboracensis, Dennstaedtia punctilobula, Pinus 
strobus, Polygonum perfoliatum, Betula lenta, Castanea dentata, Lindera benzoin, Nyssa 
sylvatica, and Toxicodendron radicans. The field trip was followed by an open house of the 
Botany Department of the Academy of Natural Sciences for a chance to meet department 
personnel and to tour the Academy’s herbarium. Leader: David Hewitt. 


16 June: Miller Farm, Chester County, Pennsylvania. We braved wind and intermittent rain 
from tropical storm Allison to explore fields and meadows of this Brandywine Conservancy- 
owned property. After a brief talk by Thom Larson on the history of the property, we 
walked down through an unmown hayfield to the wetland corridor draining the eastern edge 
of the site. Here we looked at wetland plants and discussed how to tell various species of 
Carex apart. Larson also described the methods employed in keeping the various field and 


120 BARTONIA 


wetland habitats from succeding to shrubland and woodland. Unfortunately, colonies of 
gaywings, Polygala paucifolia, had already bloomed, but we were able to see them in fruit. 
In addition, we were rewarded by a blossoming display of a tremendous population of 
Hypoxis hirsuta. Other notable plants seen were Spiraea tomentosa, Dryopteris cristata, 
Eupatorium pilosum, Andropogon gyrans, Rubus hispidus, which in some spots was the 
dominant ground cover, and Stellaria alsine. We also found a new addition for the property, 
Gratiola neglecta. Carices seen included Carex albicans, C. amphibola, C. annectens, C. blanda, 
C. debilis var. debilis, C. digitalis, C. glaucodea, C. gracilescens, C. hirsutella, C. lurida, C. 
pensylvanica, C. radiata, C. scoparia, C. spicata, C. swanii, C. stricta, C. styloflexa, C. vestita, 
and C. vulpinoidea. Due to impending inclement weather, the trip concluded at noon. 
Attendance: 6. Leaders: Jack Holt and Janet Ebert. 


24-28 June: Wesley College, Dover, Kent County, Delaware. Joint field meeting with the 
Northeast Section of the Botanical Society of America and the Torrey Botanical Society. Dr. 
William Kroen of the Biology Department at Wesley College provided assistance as the local 
host. William McAvoy of the Delaware Natural Heritage Program planned the itinerary for 
the three days of field trips. They included sites in central Delaware (Killens Pond State 
Park, Cape Henlopen State Park, Blackbird State Forest) and eastern Maryland (Adkins 
Arboretum at Tuckahoe State Park, Big Marsh at Echo Outdoor School). The plant 
communities included examples of upland forest, swamp forest, shrub swamp, seasonal pond 
(Delmarva bay), fresh marsh, salt marsh, dune, and beach. William McAvoy led a field trip 
at one of the sites each day and provided the species lists and maps distributed to the 
participants. The other trip leaders were Jack Holt and Janet Ebert (Botanical Consultants, 
Chadds Ford, Pennsylvania), Keith Clancy (Delaware Native Plant Society), and Brent Steury 
(National Park Service, Washington, D.C.). Evening lectures were given by William McAvoy 
and Keith Clancy, by Robert Naczi and Susan Yost (Claude E. Phillips Herbarium, Delaware 
State University), and by Victor Soukup (University of Cincinnati and Ohio Native Plant 
Society). In addition, Arthur Tucker (Delaware State University) hosted a tour of the new 
Phillips Herbarium building and James McClements of Dover invited the participants to 
examine his cultivated collection of American and Eurasian forest perennial herb species. 
Chairmen: Tim Draude and Larry Klotz. Attendance: 65, representing 11 northeastern states 
plus Florida, California, and the District of Columbia. Report by Karl Anderson. 


11-20 July: Newfoundland, Canada. Joint meeting with the New Jersey Audubon Society 
and the Torrey Botanical Society. This tour visited areas on Newfoundland’s west coast, 
from Gros Morne National Park to Saint Anthony. Habitats studied included boreal forest, 
fens, bogs, serpentine exposures, coastal limestone barrens, ponds, shores, and roadsides. 
Over 300 plant species were included in a partial list of species seen; of these about 250 were 
species that are non-existent or rare in the Philadelphia area. Of seventeen species of orchids 
seen, fifteen were in bloom, including Cypripedium reginae, Orchis rotundifolia, Platanthera 
orbiculata, and the endemic Platanthera straminea. We saw four species of saxifrages, of 
which Saxifraga aizoon, S. aizoides, and S. cespitosa were in bloom, and a good variety of low 
arctic specialties such as Bartsia alpina, Potentilla crantzii, P. usticapensis, Primula egalikensis, 
Tofieldia pusilla, Epilobium latifolium, Dryas integrifolia, Lesquerella purshii, and Arnica 
terranovae. In addition to plants, participants enjoyed good weather, lovely scenery, and 
good looks at moose, humpback whales, and other wildlife. Visits were made to the Norse 
settlement site at L’Anse aux Meadows and to the visitor center at Port au Choix, site of 


2000-2002 FIELD TRIPS 121 


ongoing archaeological investigations into Maritime Archaic and Paleo-Eskimo cultures. 
Attendance: 8. Leader: Karl Anderson. 


28 July: Pancake Turfcut near Waretown and Lochiel Creek near Barnegat, Ocean County, 
New Jersey. In the early 1970s, co-leader Ted Gordon introduced the term “turfcut” to 
designate a man-made plant community in early succession harboring a diverse assemblage 
of showy or rare pioneer species. These persistent communities were created by the 
removal—often down to the mineral soil—of the low shrub layer or turf (e.g., sheep laurel, 
teaberry, dwarf huckleberry) from the moist areas that border Atlantic white-cedar swamps 
or pitch pine lowlands. The chunks of turf were used by charcoal burners to cover their pits, 
by road builders to stabilize steep slopes, and by farmers to stabilize dikes around their 
cranberry bogs. Our first stop was at Pancake turfcut between the Garden State Parkway 
and the abandoned Tuckerton Railroad right-of-way about 2 miles northwest of Waretown. 
We rediscovered two small stands of the federally threatened Rhynchospora knieskernii along 
a damp sand road that bisects the site. The globally rare Narthecium americanum was mostly 
concentrated in three stands along this road with a few isolated clumps further in the 
interior of the tract. Only about 20 severely retarded plants were in bloom, which suggested 
that habitat conditions were not optimal for sexual reproduction. The turfcut also contained 
thousands of plants of the globally rare Schizaea pusilla; the occurrence may be the largest 
in the Pine Barrens. Additional wetland plants included Rhynchospora alba, R. capitellata, R. 
gracilenta, R. fusca, Carex exilis, C. striata, Cladium mariscoides, Eleocharis tuberculosa, 
Andropogon glomeratus, Muhlenbergia uniflora, Eriophorum virginicum, Drosera filiformis, D. 
rotundifolia, Vaccinium macrocarpon, Xyris difformis, Lobelia nuttallii, Polygala cruciata, 
Pogonia ophioglossoides, Calopogon tuberosus, Lycopodiella alopecuroides, L. appressa, and 
Pseudolycopodiella caroliniana. Although impacted by the regeneration of Pinus rigida and 
Chamaecyparis thyoides, this turfcut, created in the early 1950s, continues to produce a 
remarkable herb flora. We next visited a Helonias bullata occurrence along Lochiel Creek 0.7 
mile west of the stream’s junction with Route 9 and 1.5 miles north of Barnegat. Over the 
past several years, a portion of this swamp-pink population has shown a substantial decline 
in flowering. This is most likely a result of water deprivation related to upstream 
construction of a series of detention basins linked to housing development at Rose Hill 
Estates. In contrast, the flowering of a fine second colony of swamp-pink below the dam of 
an abandoned cranberry bog was not impeded; these plants were receiving a steady flow 
from two small tributaries of Lochiel Creek. In the adjacent bog that has regenerated to a 
sphagnous cedar swamp with scattered openings, we saw in flower Rhexia virginica, Sabatia 
difformis, Xyris torta, Polygala brevifolia, P. cruciata, Utricularia striata, U. subulata, Drosera 
intermedia, D. rotundifolia, D. filiformis, and Platanthera clavellata. Also recorded here were 
Rhynchospora alba, R. capitellata, R. gracilenta, Eleocharis tuberculosa, Carex bullata, C. 
collinsii, C. striata, C. atlantica, Dulichium arundinaceum, Schoenoplectus pungens, Andropogon 
glomeratus, Sarracenia purpurea, Orontium aquaticum, Eriocaulon aquaticum, Sparganium 
americanum, Triantalis borealis, Hypericum canadense, Eupatorium perfoliatum, Thelypteris 
simulata, Decodon verticillatus, Uvularia sessilifolia, Viburnum nudum var. nudum, Lindera 
benzoin, Xerophyllum asphodeloides, Juncus caesariensis, Utricularia purpurea, Schizaea pusilla, 
Sphagnum papillosum, S. cuspidatum, S. tenerum, S. pulchrum, and S. magellanicum. Our most 
significant discovery was a few clumps of the state endangered Eleocharis tortilis on the upper 
edge of sphagnous depressions in the bog. It appears that this sedge, with its peculiar, spirally 
twisted culms, was last collected in Ocean County by Bayard Long in 1915 in swampy 


122 BARTONIA 


woods nearby to the east. We were unsuccessful in finding more plants of swamp-pink in 
a wider search of the bog and the swamp upstream of it. Attendance: 18. Leaders: Alfred E. 
Schuyler and Ted Gordon. 


11 August: Atsion, Burlington County and Rockwood-West Mill Tract of Wharton State 
Forest, Atlantic County, New Jersey. At Atsion near the abandoned Jersey Central railroad 
tracks, we saw Fimbristylis puberula, Rhynchospora torreyana, Juncus biflorus, and Croton 
willdenowti (= Crotonopsis elliptica). What once was a vigorous Gentiana autumnalis 
population here has been reduced to a few scattered, sterile and budding individuals wilted 
by an extended period of drought. We traveled in 4-wheel-drive vehicles to the iron-ore 
swales and pitch pine lowland southeast of Atsion and east of Dutchtown to see an 
occurrence of Xyris caroliniana, discovered by the leader in 1998 on a relatively dry patch 
of sand. This state-endangered yellow-eyed grass has never been known from more than two 
or three sites in the barrens. A couple of the solitary plants examined had the characteristic 
deeply set, lustrous brown sheaths below, dilated to form an elongated, bulb-like base, a 
feature that makes this Xyris readily distinguishable from other members of the genus. Also 
seen nearby were Calamovilfa brevipilis, Aristida virgata (= Aristida purpurascens var. 
virgata), Agalinis purpurea var. racemulosa (= A. virgata), Carex livida, Leiophyllum 
buxifolium, and additional colonies of both Gentiana autumnalis and Fimbristylis puberula. 
Via a narrow, potholed trail we reached an extensive, peaty, iron-ore swale along Gun 
Branch just south of Rockwood. The diverse species assemblage of this shallow depression 
included Xyris difformis, X. torta, Eriocaulon decangulare, Carex striata, Dulichium 
arundinaceum, Eleocharis tuberculosa, E. tenuis, E. olivacea, Rhynchospora capitellata, R. 
cephalantha, R. chalarocephala, R. alba, R. fusca, R. gracilenta, Cyperus dentatus, Cladium 
mariscoides, Mublenbergia torreyana, Erianthus giganteus, Panicum verrucosum, P. virgatum, 
P. longifolium, Andropogon glomeratus, Glyceria obtusa, Juncus pelocarpus, J. canadensis, J. 
scirpoides, J. effusus, Sagittaria englemanniana, Iris prismatica, Lobelia canbyi, Lophiola aurea, 
Lachnanthes caroliniana, Hypericum canadense, H. denticulatum, Triadenum virginicum, Rhexia 
virginica, Woodwardia virginica, Chamaedaphne calyculata, and Vaccinium macrocarpon. By 
trails only recently improved by the Forest Fire Service, we drove southeast toward Pleasant 
Mills to observe the impacts of the 29 July 1997 wildfire on 1,900 acres of the landscape 
between Sleeper Branch of Bear Swamp and the Noscochague Creek. Everywhere in 
profusion were Amphicarpum purshii, Calamovilfa brevipilis, and Cyperus dentatus; 
Mublenberia torreyana and Rhynchospora cephalantha were noted in several bog-ore basins. A 
couple of these swales also contained small pockets of Rhynchospora knieskernii. Attendance: 
20. Leader: Ted Gordon. 


19 August: Stafford Forge Wildlife Management Area (W.M.A.), Ocean County, New 
Jersey. We visited three impoundments in the southern section of this W.M.A. Water levels 
were very low, but this did not appear to have a negative effect on the flora. Some of the 
plants seen in bloom were Eupatorium leucolepis, Polygala cruciata, Chrysopsis mariana, 
Gratiola aurea, Xyris difformis, X. torta, X. smalliana, Rhexia virginica, Oclemena nemoralis 
(= Aster n.), Eurybia compacta (= Aster gracilis), Utricularia cornuta, U. purpurea, 
Rhynchospora alba, R. capitellata, R. chalarocephala, Juncus biflorus, Vaccinium macrocarpon, 
Drosera filiformis, D. intermedia, D. rotundifolia, and Sarracenia purpurea. A small savanna 
near the northernmost pond was explored, and Schizaea pusilla was found, as were 
Eriophorum virginicum, Lycopodiella appressa, L. caroliniana, and abundant foliage and fruit 


2000-2002 FIELD TRIPS 123 


of Pogonia ophioglossoides. Ted Gordon pointed out Xyris fimbriata, a new plant for most of 
the participants. The day ended with a side trip to the edge of a salt marsh north of New 
Gretna, where Lythrum lineare, Sabatia dodecandra, Agalinis purpurea, Kosteletzkya virginica, 
and Schoenoplectus americanus (= Scirpus a.) were found. Attendance: 12. Leader: Karl 
Anderson. 


31 August: Thompson Park, Jamesbury, Middlesex County, New Jersey. A large number 
of coastal plain species was found in a wet spring area in the park, including Andropogon 
glomeratus, Clethra alnifolia, Eupatorium dubium, Eupatorium hyssopifolium, Eupatorium 
pilosum, Hypericum gentianoides, Lechea pulchella, Lycopodiella appressa, Panicum verrucosum, 
and Rhynchospora capitellata. The dredge material from Manalapan Lake in the park, 
deposited adjacent to this wetland area, had an interesting flora including Andropogon 
gerardii, Bidens aristosa, Echinacea purpurea, Helenium autumnale, Helenium flexuosum, 
Panicum virgatum, Tripsacum dactyloides, and Verbesina alternifolia. Some of the group 
wandered into the wooded area of the wetlands near the spring and found Botrychium 
dissectum, Lycopodium digitatum, Onoclea sensibilis, Osmunda cinnamomea, Thelypteris 
noveboracensis, Thelypteris palustris var. pubescens, and Woodwardia areolata. Report by leader: 
Bill Olson. 


29-30 September: Maurice River Cove and Delaware Bay, Cumberland County, New Jersey. 
Joint trip with the Torrey Botanical Society. The trip began on a warm, sunny day at the 
intersection of Route 47 and the Maurice River Causeway just east of Maurice River Station. 
Here along the roadside and pine-oak woods adjacent to the railroad tracks, we noted several 
species of oak: Quercus alba, Q. coccinea, Q. marilandica, Q montana (= Q. prinus), Q. 
stellata, and Q. velutina. Most interesting was Q. xsaulii, a cross between Q. alba and Q. 
montana. Mac Alford, a student of Dioscorea from Cornell University, identified material 
from here as D. hirticaulis. We also noted five taxa of Eupatorium in bloom: E. album, E. 
hyssopifolium, E. pilosum, E. rotundifolium var. rotundifolium, and E. rotundifolium var. 
ovatum. In an open field at the intersection of Route 47 and Whitney Point Road, fall herbs 
were in excellent bloom: Agalinis purpurea, Doellingeria umbellata, Lobelia puberula, 
Spiranthes cernua, Symphiotrichum dumosum, S. lateriflorum, and Veronica noveboracensis. 
Adjacent to the field, the group had an excellent opportunity to distinguish between Morella 
caroliniensis (= Myrica pensylvanica) and M. cerifera, as both were growing together. In the 
adjoining woods, a few specimens of Quercus michauxit were noted. The group then visited 
the Moores Beach area and walked all the way out to the beach. Species that were noted in 
the salt marsh and beach habitats included Ambrosia artemisiifolia, Chamaesyce polygonifolia, 
Chenopodium ambrosiodes, C. berlandieri, Cycdoloma atriplicifolium, Kochia scoparia, 
Limonium carolinianum, Polygonum prolificum, Suaeda calceoliformis, Solidago sempervirens, 
Symphiotrichum subulatum, and Trichostema dichotomum. Along Moores Beach, a good 
number of monarch butterflies (Danaus plexippus) were noted, many of them on the blooms 
of Solidago sempervirens. On our last stop at the edge of a salt marsh along Thompsons Beach 
Road, we noted an excellent stand of Euthamia minor (= Solidago microcephala), along with 
Cyperus filicinus, Suaeda linearis, and Pluchea odorata. 


The second day of the trip was quite cool and rainy as we traveled to Bivalve and noted a 
good stand of Setaria magna growing in the tidal marshes. This grass was quite distinctive, 
as it was taller than the Phragmites australis also present in the marsh. Other plants noted 


124 BARTONIA 


here included Eleocharis parvula, Hibiscus moscheutos, Polygonum cespitosum, P. lapathifolinm, 
P. pensylvanicum, Ruppia maritima, Schoenoplectus robustus, Setaria faberi, and Typha latifolia. 
The group then visited the woods and tidal marshes along Hansey Creek Road. Here we 
focused on Carya, Pinus, and Quercus, noting C. glabra, C. pallida, C. tomentosa (= C. alba), 
P. rigida, P. echinata, P. taeda, P. virginiana, Q. alba, Q. coccinea, Q. falcata, Q. marilandica, 
Q. nigra, Q. phellos, Q. montana, and Q. velutina. Linda Kelly discovered a population of 
Pyrrhopappus carolinianus in this area. Attendance: 14. Leader: Gerry Moore. 


2002 Field Trips 


22 June: Martha Furnace and Oswego River Savannas, Wharton State Forest, Burlington 
County, New Jersey. At the town site of old Martha, a cellar hole, scattered fragments of 
furnace slag, a tailrace, and the presence of Catalpa bignonioides, Juglans nigra, Ulmus 
americana, and Robinia pseudoacacia were vivid reminders of former human habitation. Here 
a small glade of less than an acre of man-altered soil has produced what may be the most 
diverse fern flora in all of the Pine Barrens. We recorded 12 species of ferns: Asplenium 
hel Botrichium dissectum (both forma obliguum and forma dissectum), B. virgin- 

m, Dryopteris carthusiana, D. cristata, Ophioglossum pusilum (= O. vulgatum var. 
jesudipediand Osmunda cinnamomea, O. regalis var. spectabilis, Thelypteris palustris var. 
pubescens, Onoclea senstbilis, Polytrichum acrostichoides, and Woodwardia areolata. Seen here 
in former years, Dennstaedtia punctilobula, Athyrium felix-femina, and Dryopteris neveborac- 
ensis eluded us. However, Pteridium aquilinum var. latiusculum and Schizaea pusilla were 
observed nearby. At Martha we also saw Vitis aestivalis var. aestivalis, Triodanis perfoliata, 
Apocynum xfloribundum, Holcus lanatus, Poa compressa, and in the oak-pine forest 
overlooking the Oswego River, Panicum columbianum, Danthonia sericea, and Piptochaetium 
evanaceum (= Stipa evanacea). Upstream of the furnace site in sphagnous seeps, muck flats, 
and savannas that stretch along the river from the foot of Calico Ridge to just north of 
Cutts Pumphouse, we observed several bright yellow bands of Narthecium americanum, the 
fuzzy stars of Lophiola aurea, massive goblets of Sarracenia purpurea, the white buttons of 
Eriocaulon compressum and E. decangulare, scattered culms of Danthonia epilis, and flowering 
patches of Utricularia cornuta, U. striata, and U. subulata (including forma cleistogama). 
Several lovely pink blossoms of Calopogon tuberosus and Pogonia ophioglossoides dotted the 
landscape. A highlight was the rediscovery of a population of the rare Utricularia resupinata, 
occurring as numerous delicate, tiny, violet flecks on black muck. Unable to locate a known 
occurrence of viscid asphodel here, we went north to Buck Run Savanna where we noted 
two specimens of this lily just starting to flower. The Flora of North America (Vol. 23, 2002) 
has noted that all specimens of Triantha (= Tofieldia) collected in Burlington County have 
been annotated Triantha glutinosa x T. racemosa, representing “a surviving disjunct remnant 
with attributes of both species.” It appears that 7. racemosa does not occur in the state. 
Thanks to William Standaert for maintaining a list of species observed. Attendance: 23. 
Leader: Ted Gordon. 


29 June: Nescopeck State Park, Luzerne County, Pennsylvania. Ann Rhoads and Tim Block 
of the Morris Arboretum, who have conducted an inventory of the Nescopeck Valley for 
the Pennsylvania Bureau of State Parks, led the trip. Although Ann and Tim had compiled 
a list of over 600 species of vascular plants from the area, club members were able to add 


2000~2002 FIELD TRIPS 125 


several new records. The first stop was in the acidic oak-heath forest typical of south-facing 
slopes on Wisconsinan glacial till, where the globally rare variable sedge (Carex polymorpha) 
is abundant. Other species noted in this area were climbing fern (Lygodium palmatum), 
which is abundant in low areas along Nescopeck Creek, fly-poison (Amianthium 
muscaetoxicum), and beaked and American hazelnut (Corylus cornuta and C. americana), 
which grow in mixed populations along woods roads and trails. Several species with a more 
northern distribution in the state were noted such as bush-honeysuckle (Diervilla lonicera), 
dewdrop (Dalibarda repens), and northern wood-sorrel (Oxalis acetosella). 


The unglaciated lower slope of Mt. Yaeger on the south side of the valley was the target for 
the afternoon. Unlike the south-facing slopes on the opposite side of the valley, the lower 
slopes of Mt. Yaeger support a northern hardwood forest characterized by sugar maple (Acer 
saccharum), basswood (Tilia americana), and beech (Fagus grandifolia). Here the group saw 
mountain maple (Acer spicatum), fly-honeysuckle (Lonicera canadensis), purple-flowering 
raspberry (Rubus odoratus), dwarf raspberry (Rubus pubesens), Canada violet (Viola 
canadensis), and barrens strawberry (Waldsteinia fragarioides). The find of the day was a small 
population of ginseng (Panax quinquefolius), first spotted by David Lauer. Report by Ann 
oads. 


13 July: Intermittent ponds—Decou Pond and Sykes Branch, Woodland Township, 
Burlington County, New Jersey; Micajas and Hidden Ponds, Stafford Township, Ocean 
County, New Jersey. We visited two intermittent ponds in pitch pine lowland about 1.5 
miles west of Coyle Airfield in the West (Upper) Plains and about 1 mile south of Route 
72. Ringed primarily by Chamaedaphne calyculata, Vaccinium corymbosum (including var. 
caesariensis), and Pinus rigida, Decou Pond (= Deacon Pond), largest of at least seven open 
ponds in the vicinity, was completely devoid of water. Readily observed on its sandy bottom 
interspersed with muckier zones were Eleocharis olivacea, Drosera intermedia, Carex striata, 
Sphagnum cuspidatum, immature Juncus pelocarpus, young Rhynchospora alba, Xyris difformis, 
Panicum verrucosum, Eleocharis robbinsii, Rhexia virginica, Lachnanthes caroliniana, Eriocaulon 
aquaticum, and a few leaves of Nymphoides cordata prostrate on muck. In a much smaller, 
though similar, dried-up pond associated with an intermittent feeder of Pope Branch about 
260 yards to the north of Decou Pond, we found only the first eight species listed above, 
along with numerous Acer rubrum seedlings. Eleocharis robbinsit, common in Decou Pond, 
appeared to be missing here. About 1 mile to the southeast, we surveyed the desiccated 
headwater of Sykes Branch, a tire-marred, intermittent stream corridor bisecting an open 
lowland pitch pine swale on the state-owned West Plains Natural Area. Here we relocated 
a natural occurrence of the globally threatened Rhynchospora knieskernii, discovered by the 
leader in August 1992. A mere dozen culms of this beaked-rush were noted on peaty sand 
in deep tire ruts in association with Rhynchospora capitellata, Amphicarpum purshi, Lobelia 
nuttallii, and Calamovilfa brevipilis. It appears that periodic wildfire, light vehicular traffic 
by hunters, and inundation following precipitation have helped to retard shrub succession. 
This in turn has helped to maintain the pioneer conditions so essential for sustaining this 
and perhaps similar populations of Knieskern’s beaked-rush. After eating lunch by the bridge 
over the upper Oswego at historic Cedar Bridge Inn, a hostelry well known to early 
Philadelphia botanists, we headed about 3.5 miles southeast of the fire house in Warren 
Grove to Micajas Pond on the edge of the East (Lower) Plains within the Stafford Forge 
Wildlife Management Area. During the past decade, this headwater pond, situated in the 


126 BARTONIA 


intermittent west prong of Long Branch of Cedar Run, has been known to produce severely 
fluctuating populations of Rhynchospora knieskernit. During severe drought in 1995, an 
estimated 10,000 fruiting culms of this sedge were seen; on this occasion of even more 
devastating drought, we found only about 150 culms, all confined to the upper third of the 
desiccated basin. Among the associated species were Rhynchospora fusca, R. gracilenta, R. 
capitellata, R. alba, R. torreyana, Gratiola aurea, Hypericum canadense, Eleocharis olivacea, 
Dichanthelium wrightianum (= Panicum w.), and Scleria reticularis. 


At Hidden Pond, a natural, open, intermittent pond in the east prong of Long Branch some 
400 yards to the northeast, we noted about 300 culms of Rhynchospora knieskernii, far below 
the 9,000 tufts recorded upon discovery in 1996. Underlain by Downer loamy sand and 
nestled between two gentle slopes occupied by pine-oak forest, Hidden Pond was, as all of 
the other sites visited, completely desiccated. All species observed in Micajas Pond were also 
seen here with the exception of Rhynchospora torreyana. Additional species recorded were 
Mublenbergia torreyana, Panicum virgatum, Xyris difformis, and Lobelia nuttallii. Thanks to 
Bill McLaughlin for assistance with this report. Attendance: 13. Leader: Ted Gordon. 


20 July: Ker-Feal (country home of Albert and Laura Barnes), West Pikeland Township, 
Chester County, Pennsylvania. On this Barnes Foundation property, we identified numerous 
cultivated trees and shrubs around the house, barn, and long driveway. These included Abies 
concolor, Acer ginnala, Acer griseum, Acer mandshuricum, Acer tataricum, Berberis tricanth- 
ophora, Cercidophyllum japonicum, Cotoneaster acutifolia, Eleagnus umbellata, Exochorda 
giraldii, Gymnocladus dioica, Hovenia dulcis, Hydrangea petiolaris, Ilex crenata, Lonicera 
pileata, Metasequoia glyptostroboides, Neilia sinensis, Platanus hybrida, Pseudolarix amabilis, 
Pyracantha coccinea, Spiraea douglasii, Syringa vulgaris, and Tetradium daniellii (Euodia d.). 
The driveway was lined with Ulmus rubra, most of which were dying. We also recorded 
wild plants in the woods and fields on the northwest portion of the property where there 
was a mixture of natives and exotics. Some of the common woody exotics were Acer 
platanoides, Berberis thunbergii, Celastrus scandens, Euonymus alatus, Lonicera japonica, Malus 
sp., Morus alba, Phellodendron sp., Prunus avium, Rhodotypos scandens, and Viburnum 
dilatatum. Of the 200+ plant species recorded, over 70 were aliens. A comprehensive plant 
list, compiled by Jack Holt and Alfred E. Schuyler, is available from the latter. Leader: 
Alfred E. Schuyler 


24 August: Bar-hopping along the Brandywine, Pennsylvania. Exploration of the floodplain 
of the Brandywine Creek near Chadds Ford was cut short at noon by rain. From the 

parking lot of the Brandywine Conservancy, we ventured into some older floodplain woods 
south of Route 1 where te — se tg riparius, Solidago flexicaulis, Tilia americana, and an 
uncommon i lium oleraceum, distinguished from A. vineale by its later fruiting 
period and the extremely fe sheath clasping the head of bulblets. After going under Route 
1, we went onto a muddy gravel | bar on the cupswreane side of an old dam. On the recently 
exposed mud grew L pyll icatum, Zosterella dubia, 
Heteranthera sana and Potomogeton nodosus and on recently-barred, gravelly flats, 
Lindernia dubia, Penthorum sedoides, Ludwigia palustris, Veronica anagallis-aquatica, Rorippa 
palustris, Alisma subcordatum, and Eragrostis hypnoides. Growing at an elevation just a few 
inches higher than the above-listed species were Amaranthus blitum, A. spinosus, Epilobium 
coloratum, Bidens frondosa, Mimulus ringens, Echinochloa muricata, and Chenopodium 


2000-2002 FIELD TRIPS 127 


ambrosioides. The prize species seen during this part of the walk was a small population of 
the state rarity Rotala ramosior growing on a muddy shore just north of the bar. We crossed 
a powerline cut dominated by aliens into an extensive shrub swamp and open marsh 
community. Because the drought had drawn down the water table, we were able to cross 
areas that normally would have been at least knee-deep in water and mud. The vegetative 
zones created by fluctuating water tables were also quite evident here. The highest and driest 
parts of the wetland were dominated by trees, principally willow and ash, with extensive 
colonies of Crataegus crus-galli (cockspur hawthorn). These gave way along the edges and in 
the higher parts of the open marsh to large colonies of Cephalanthus occidentalis, occasionally 
mixed with Rosa palustris and enormous colonies of the sedge Cyperus esculentus mixed with 
Eragrostis cilianensis. Plants of Panicum rigidulum, Scirpus cyperinus, and Carex lupulina 
dotted the edges of the open drawdown. In the center of the drawdown, the mud was bare 
and extensively cracked; here the only common species was Nuphar advena. Attendance: 6. 
Leaders: Janet Ebert and Jack Holt. 


21-22 September: Maurice River Watershed and Delaware Bay, Cumberland County, New 
Jersey. Joint trip with the Torrey Botanical Society. The group entered the Menantico Ponds 
Wildlife Management Area from Orange Street in the Millville Industrial Park and walked 
along the railroad tracks toward Menantico Creek. Along the railroad we noted Toxicoden- 
dron pubescens and T. radicans in fruit. In some tidal ponds adjacent to the creek, Flatine 
americana and Eriocaulon parkeri were noted. During this drought year, populations of these 
two species were considerably smaller and the water was more brackish than in previous 
years. It was the first year the leader had noted blue crabs (Calinetes sapidus) in these ponds. 
A vernal pond just off Route 47 south of Brickboro was then visited. The pond, devoid . 
water, was dominated by Gratiola aurea in bloom. Associated species were 

canadense, H. mutilum, Lycopodiella appressa (= Lycopodium appressum), Panicum lonedolicia, 
Rhynchospora capitellata, Scleria reticularis, Vaccinium macrocarpon, and Xyris difformis. The 
group then walked west through pine-oak woods and botanized abandoned sand mining pits. 
Here large stands of Eriocaulon aquaticum, Myriophyllum humile, and Schoenoplectus 
subterminalis were seen. The group continued westward and then north along abandoned 
railroad tracks through a salt marsh adjacent to the Maurice River. At times, this site was 
nearly impenetrable due to large thickets of Baccharis halimifolia. On the following day, the 
group visited the woods and tidal marsh along Hansey Creek Road. Many of the species 
noted in 2001 were again seen this year, among them Pyrrhopappus carolinianus in bloom and 
excellent stands of Agalinis purpurea, Lechea pulchella (= L. legettii) and Quercus nigra. The 
group then headed to the Thompson Beach area and botanized along the road leading into 
the salt marsh. All of the typical salt marsh species seen last year at Moores Beach were also 
noted here. Growing right in the sand in the road, we found a small population of the rare 
Sesuvium maritimum in bloom. Attendance: 15. Leader: Gerry Moore. 


28 September: Fall flowers at the Jenkins Arboretum, Chester County, Pennsylvania. 
Jenkins Arboretum is a combination of a remnant of the once continuous eastern hardwood 
forest and a built naturalistic landscape using predominately flora native to eastern North 
America. As a new botanical garden, which opened to the public in 1976, Jenkins was not 
a converted estate garden but a carefully planned and planted arboretum. Due to the large 
numbers of ericaceous plants indigenous to the site such as Rhododendron periclymenoides, 
Vaccinium palidum, and Kalmia latifolia, Ericaceae became the Arboretum’s area of 


128 BARTONIA 


specialization, Specifically, species and hybrid rhododendrons from all over the world are the 
majority of accessions, which number in the range of 4 to 5 thousand plants. In addition to 
Club members, we were joined by on anne herbaceous peices class from Temple 
University. The field trip in late S usual eupatoriums, fall asters, and 
goldenrods but it was the richness ‘and diversity of deciduious'and evergreen rhododendrons 
that made the experience unique. Jenkins Arboretum’s goal of creating a garden-like feel in 
concert with nature was a major highlight. Also, where else can one go botanizing so easily 
and most of the plants are already identified and labeled? Report by leader: Harold E. 
Sweetman, Executive Director. 


Bartonia No. 62: 129-130, 2004 


PROGRAM OF MEETINGS 
September 2002-May 2005 


Date Subject Speaker 
2002 
26 Sep Members’ Reports on Summer Botanizin 
24 Oct Fifty-six Orchids of New Jersey: a Video Film Presentation ........ David Snyder 
21 Nov __ Spatial and Temporal Views of tidal Freshwater Plants: Their Seed Banks 

ond Cpertoistion Ronieey 02.0. rc ee Se ea i tee ks 5 Mary Leck 
19 Dec What Can Mites Tell Us About the Systematics of Western 

HMeehisphere Pircher Plante? 6.0 226 vs oe 0 Oo SF eG Robert F. C. Naczi 
2003 
23 Jan Saving an American Treasure—the Lewis and Clark Herbarium Durin 

tne Dieses G00 Nears. 5 oe ks ae FA Richard M. McCourt 
27 Feb Between a Rock and a Soft Place: Plant Habitats Scrutinized ......... Rick Mellon 
27 Mar Pollination, Breeding system, and Cushion Structure of the Alpine 

Forget-me-not (Eritrichium nanum) ... 2... c cee eens Heinrich Zoller 
24 Apr Around the World: 80 Days and 80 Plants, with Stops in Zimbabwe, 

China, Breen Taies, and the Americas... oc he. oe ee Harold Sweetman 
22 May Philadelphia Botany and Horticulture in the Time of Lewis and Clark . . . Joel T. Fry 
25 Sep Members’ Reports on Summer Botanizin 
23 Oct Mosses and the Conservation of Natural Communities ........... Terry O’Brien 
20 Nov The Fores Pera is ae ee Ee 8 Ann F. Rhoads 
18 Dec The Lost Worlds of Venezuela: Flora of the Tepuis and Tabletop 

Wii i cr i a wy oe as 0 ok ee es Ce ae Lena Struwe 
2004 
22 Jan The Bryophytes of New Jersey... 6 5 bee eee tee ee ee Bill Olson 
26 Feb The Lichter: oF New dere i ee ys eee James Lendemer 
25 Mar Plant Diversity and Exotic Species Invasion in Southern Appalachian 

Fir eaes YSU os ie oe ee a wens eon * Rebecca L. Brown 
22 Apr The Flora of Coastal Plain Seasonal Ponds on the Delmarva 

Penintie ss Eos Fe ee a CG: oe os William McAvoy 
27 May Exotic Trees in Our Landscapes, and mat od Urban Forestry Program . John Kuser 
23 Sep Members’ Reports on Summer Botanizin 
28 Oct eee Exploration in South Africa ie Ne ieee. Ted Gordon 
18 Nov ter Collinson and His Philadelphia Friends ...........-. Elizabeth P. McLean 
19 Dec a Bee of the Benjamin Smith Barton Historical Marker 


Philadelphia Chapter, Lewis & Clark Trail Heritage Foundation 


ie ee Res a ee eT 


129 


130 
2005 
27 Jan 
24 Feb 


24 Mar 
28 Apr 


26 May 


BARTONIA 


Highlights of the Madagascar Flora 
Ectomycorrhiza Underground Networking, or Fungi and the 

WO IE, WE a es oe ot a see te es oe 4 
Cedar Glades: History, Ecology, and Conservation 
Flora of - Warren Grove Gunnery Range, Burlington County, 

lal <5 > SPOS 20a sere paren rae ariia ae ONe rhe ae 
teks Hoi Ecology of Eastern Oak Forests: Past, Present, and 
Fut 


es ea Se ee ee ee ee a eo 8 ee ee ee ee ee ee ee eee ee eee ee 


Bartonia No. 62: 131-138, 2004 


2004-2005 Membership List 


Honorary Member 

FARLEY, PERO: B. — 319 Bala Ave., Bala Cynwyd, PA 19004, 610-667-0625 

NEWBOLD, ANN — 411 N, Middletown Rd, Apt. E-111, Media, PA 19063, 610-754-7573 

SCHUYLER, ALFRED E. — Academy of Nasal Sciences, 1900 Benjamin Franklin Pkwy., 
Philadelphia, PA 19103, 215-299-1193, schuyler@acnatsci.org 


Life Members 

ARSENAULT, JOE — 961 Clark Ave., Franklinville, NJ 08322, njplants@aol.com 

BIEN, WALTER — 144 Summit Ave., Langhorne, PA 19047, 215-752-3762, walter.bien@verizon.net 

GREENLAND, CHRISTINE MANVILLE — 790 E. Street Rd., Warminster, PA 18974, 215-322-4105 

HOLT, ROBERT J. — 3032 Taft Rd., Norristown, PA 19401, 610-584-5578 

IRETON, MARY LOU — 213 4th Ave., Haddon Heights, NJ 08035, 856-547-1118 

LAUER, DAVID — 49 Cornell Ave., Churchville, PA 18966, 215-357-2646, dml100@aol.com, 
dml100@att.net 

MCLEAN, ELIZABETH — He Cherry Ln., Wynnewood, PA 19096-1208, 610-642-4196, 
epmclean@worldlynx.ne 

MCLEAN, WILLIAM II — bs Cherry Ln., Wynnewood, PA 19096-1208, 610-642-4196, 
epmclean@worldlynx.ne 

MOoRrE, GERRY — Brooklyn Botanic Gardens, 1000 Washington Ave., Brooklyn, NY 11225, 718- 
623-7332, gerrymoore@bbg.org 

O’HERRON, JOHN — 220 Washington St., Mount Holly, NJ 08060, 609-261-0711, 
joheerron@voicenet.com 

OLSON, WILLIAM — 1005 Lakewood-Farmingdale Rd., Howell, NJ 07727, 732-938-3187, 
wolson3@optonline.com 

PATRICK, RUTH — 750 Thomas Rd., Philadelphia, PA 1911 

ROBERTS, WILLIAM H. — 1922 Rittenhouse Sq; Philadelphia, PA 19103, 215-569-5632, 
roberts@blankrome.com 

RYAN, NANCY — 2355 Oakdale Ave., Glenside, PA 19038-4220 

STAILEY, HELEN M. — 8701 Macon St., Philadelphia, PA 19152, 215-673-8163 

STEVENS, CHARLES E. — 615 Preston Pl., Charlottesville, VA 22903, 804-293-8658 

THOMPSON, SUE — 129 E. Sycamore St., Pittsburgh, PA 15211-1719, 412-622-3295 

TREADWAY, SUSAN — 1509 Monk Rd., Gladwyne, PA 19035, 610-642-8050, sptreadway@aol.com 


Sponsoring Members 
RICK, JULIA — Blair 213, 1400 Waverly Rd., Gladwyne, PA 19035-1265, 610-645-8863 

HAMILTON, MARSHALL — 18 Lakewood Dr., Media, PA 19063, mhamil2741@aol.com 

HENRY FOUNDATION FOR BOTANICAL RESEARCH — 801 Stony Ln., Gladwyne, PA 19035-0007, 
610-525-2037 

JUELG, RUSSELL J. — 79 Grassy Lake Rd., Shamong, NJ 08088, russelljuelg@comcast.net, 
russell@pinelandsalliance.org 

LIBBY, VALENCIA — 607 Cloverly Ave., Jenkintown, PA 19046, 215-576-5725 

KIMELMAN, GAY — 2212 St. James Pl., Philadelphia, PA 19103, 215-563-0285 

PINELANDS PRESERVATION ALLIANCE — 17 Pemberton Rd., Southampton, NJ 08088, 609-859- 
8860, ppa@pinelandsalliance.org 


131 


132 BARTONIA 


SCHNEIDER, WILLIAM — 340 Sugartown Rd., Apt. C93, Devon, PA 19333-2346, 610-431-2449, 
schndrw@aol.com 

SWEETMAN, HAROLD — Jenkins Arboretum, 631 Berwyn Baptist Rd., Devon, PA 19333, 610-647- 
8870, harold@jenkinsarboretum.org 


Regular Member 

ADAMS, Mone’ eet Olive Natural Heritage Society, Inc., 212 High Point Road, West 
Shokan, NY 1249 

ALDHAM, ALBERT — fie Hemlock Farms, Hawley, PA 18428 

AMOS, SANDRA — 41 Laurel Rd., Clementon, NJ 08021, 856-346-2242 

ANDERSON, KARL — Rancocas Nature Center, 46 North Childs St., Woodbury, NJ 08060, 609-267- 
2195, rancoc@bellatlantic.net 

BALDWIN, DON — 4240 Fairview Ave., Newtown Square, PA 19073, 610-353-1550 

BAUCHSPIES, JAMES — 4320 Chetwin Terr., Easton, PA 18045-4908, 610-253-8925 

BENJAMIN, JESSIE — 366 Chatham Rd., West Grove, PA 19390, 610-869-4285, 
jessie@taprootnativedesign.com 

BIDDLE, DORRELL — 701 Washington Ave., Palmyra, NJ 08065, 856-829-0748, dell2733@aol.com 

BLOCK, TIMOTHY — 203 Juniper Ct., Zieglerville, PA 19492-9724, 215-234-0645, 
block@pobox.upenn.edu 

BOWELL, MICHAEL — 2148 Bodine Rd., Malvern, PA 19355, 610-827-1268, mwb@createascene.com 

BOYD, ANNA — 181 Highland Ave., Montclair, NJ 07042, 973-796-5817 

BRAM, MARGOT — 312 Bethlehem Pike, Ft. Washington, PA 19034 

BRINTON, EDWARD — 424 Crosslands Dr., Kennett Square, PA 19348 

BROTHERSON, ROBERT — P.O. Box 179, Revere, PA 18953, 610-847-1005, nellsboy@epix.net 

BUCK, WILLIAM — New York Botanical Garden, Bronx, NY 10458-5126, 718-817-8624, 
bbuck@nybg.org 

CAIAZZA, NICHOLAS — 5 Dorothea Terr., Lawrenceville, NJ 08648, ncaiazza@gateway.net 

CARR, DEBBIE — 314 E. eng Ave., Philadelphia, PA 19118, 215-242-0734 
wechslercarr@earthlink.ne 

CASEY, CINDY — 7901 Henry Ave., B101, Philadelphia, PA 19128, 215-508-0699, 
ccdakota@hotmail.com 

CHELSVIG, GORDANA — 308 Woodbine Ave., Narberth, PA 19072, 610-667-2581, 
mainlinenutrition@yahoo.com 

CONNOR, J. — 35 Clarks Landing Rd., Port Republic, NJ 08241, 604-652-5143, 
jconnor@stockton.edu 

COOK, BUD — H.C. 1, Box 1117, Blakeslee, PA 18610, rcook@tnc.org 

COONEY, PATRICK — 221 Mt. Hope Blvd., Hastings-on-Hudson, NY 10706, 914-478-1803, 
plcooney@aol.com 

COURTNEY, JOHN — 439 Gladstone Ave., Haddonfield, NJ 08033, 856-429-4987 

CRICHTON, pil = Be Loveville Rd., Ctg. 5, Hockessin, DE 19707, 302-235-0571, 
crichton@magpage.co 

DANZENBAKER, ore 607 Cloverly Ave., Jenkintown, PA 19046, 215-576-5725, 
vlibby@temple.edu 

DAVIS, CHARLES — 1510 Bellona Ave., Lutherville, MD 21093-5525, 410-252-4154, cadavis@bcpl.net 

DAVIS, THOMAS P. — 6010 Canon Hill Rd., Ft. Washington, PA 19034 

DECASTRO, LINDA — 1100 Concord Dr., Bridgewaue, NJ 08807, I.decastro@worldnet.att.net 

DENNE-HINNOV, BOEL — 1-D Hibben Apts., Faculty Rd., Princeton, NJ 08540-5508, 609-921-1272, 
hinnov2001@yahoo.com 

DENNY, GUY — po Mt. Gilead Rd., Fredericktown, OH 43019-9513, 740-694-6087, 
guydenny@ecr.ne 

DIEDRICH, panes — 502 Highland Terr., Pitman, NJ 08071-1524, 856-589-8455 


MEMBERSHIP LIST 133 


DODDS, JILL — 56 Tumble Falls Rd., Stockton, NJ 08559, 908-996-3214, 
jsdodds@biostarassociates.com 

DOUGLAS, KIM — 109 Penarth Rd., Bala Cynwyd, PA 19004, 610-667-3997, kdouglas@olinptp.com 

DORN, RUTH — 800 Trenton Rd., Apt. 244, Langhorne, PA 19047, 732-932-8165 ext. 303, 
rdorn@aesop.rutgers.edu 

DRAUDE, TIMOTHY — 415 Poplar St., Lancaster, PA 17603, 717-393-7233 

DUGGAN, CLARE — 316 Wright St., Philadelphia, PA 19128, 215-487-7515, 
moira.duggan@verizon.net 

DumMIc, MARK J. — 310 Stanton Court, Glen Mills, PA 19342, 484-840-1327, 
eet choca hihi edu 

EBERT, JANET — 1611 Smith Bridge Rd., Chadds Ford, PA 19317-9170, 610-459-0585 

EDINGER, GREG — 442 Ryan Rd., Greenwich, NJ 12834, 518-692-1725, gedinger@tnc.org 

EELLS, CAROLINE — 2001 Ridley Creek Rd., Media, PA 19063, 610-566-2534, ceels@hotmail.com 

EIGENRAUCH, JANE — P.O. Box 85, Red Bank, NJ 07701, 732-842-0690 

EVANS, WILLIAM — 5 Foxwood Ln., Media, PA 19063, 610-627-9288 

EVANS, JANET — Library, Pennsylvania Historical Society, 100 N. 20th St., Philadelphia, PA 
19103, 215-988-8779, jevans@pennhort.org 

FALLON, HENRY — 243 Vernon Rd., Monroe, NJ 08831, 609-395-1909, henry. fallon@verizon.net 

FARNON, CHRISTA — 1418 Surrey Ln., Wynnewood, PA 19096, 610-649-2668, 
christa.farnon@drexel.edu 

FEDERICI, ANTONIO — Golder Associates, 24 Commerce St., Suite 430, Newark, NJ 07102, 201- 
864-8992, tony_federici@golder.com 

FERREN JR., WAYNE R. — 67 Main St., Vincentown, NJ 08088, 609-859-8682, wrfjrl@comcast.net 

FIELD, STEPHEN — 5 Evelyn Ave., Vineland, NJ 08360, 856-691-5868 

FIELD, THERESA — 5 Evelyn Ave., Vineland, NJ 08360, 856-691-5868 

FINE, NORMAN — 73 Nature Ln., Sewell, NJ 08080-2145, 856-218-4809, handnfine@comcast.net 

FINE, HELEN — 73 Nature Ln., Sewell, NJ 08080-2145, 856-218-4809, handnfine@gateway.net 

FLANIGAN, TONI — 662 W. Johnson St., Philadelphia, PA 19144, 215-951-9211, 
toniann@philadelphiagardens.com 

FLISSER, DANIEL — Dept. of ar sn County College, P.O. Box 200, Blackwood, NJ, 
08012, 856-309-1499, dflisser@camd 

FOGARASI, KASIA — 317 bode pie Philadelphia, PA 19128, 215-482-3835, 
kasiankal@aol.com 

FORD, JOANNE — 729 Westview Ave., Philadelphia, PA 19119, 215-844-8054, jb.ford@verizon.net 

FORD, CARIK — 532 W. Springfield Ave., Philadelphia, PA 19118, 215-248-2783, 
franklincl@andropogon.com 

FUSSELL, CATHARINE — 179 Kendal Dr., Kennett Square, PA 19348, 610-388-6523 

GARBACK, MARY — 3839 Janice St., Philadelphia, PA 19114-2826, 215-332-7105 

GLASS, AMELIA — 135 Washington Ave., Pitman, NJ 08071, 856-589-6435 

GODDARD, PAUL — 127 N. Van Pelt St., Philadelphia, PA 19103, 215-557-0187 

GOFF, ELINOR — 791 College Ave., Apt. 1, Haverford, PA 19041, 215-247-5777 

Goop, NORMA — 745 Redman Ave., Haddonfield, NJ 08033, 856-428-1396 

GORDON, PAT — 31 Burrs Mill Rd., Southampton, NJ 08088, 609-859-3566 

GORDON, TED — 31 Burrs Mill Rd., Southampton, NJ 08088, 609-859-3566 

GRACIE, CAROL — 19 . Lake Circle, South Salem, NY 10590, 914-763-5938, 
ts geet is 

DENNIS — 668 Piieciog Ave., Collingswood, NJ 08108, 856- 858-6642, 
pratt rutgers.edu 

CRRENOWIEE DUANE — 100 Motor Rd., Pine Beach, NJ 08741, 732-349-0364, 
polarstar@adelphia.net 

GRIMES, BRUCE G. — P.O. Box 222, Sumneytown, PA 18084, 215-234-8424, taxkwaxak@cs.com 


134 BARTONIA 


Gross, MICHAEL F. — Biology Dept., Georgian Court College, 900 Lakewood Ave., Lakewood, 
NJ 08701-2697, 732-364-2200, gross@georgian.edu 
RUND, STEVE — Western Pennsylvania Conservancy, 209 Fourth Ave., Pittsburgh, PA 15222, 
412-288-2777, wpc@paconserve.org 
GYER, JOHN — P.O. Box 185, Clarksboro, NJ 08020, 856-423-3889, ee com 
HALLIWELL, TOM — 19 Kings Rd., Netcong, NJ 07857, 973-347-6071, tbhalliwell@att. 
HAMMERSTROM, FREDERICA — 542 W. Montgomery Ave., Haverford, PA 19041- 1409, “610-649 
3811 
HARDESTY, GAIL — 488 Big Oak Rd., Morrisville, PA 19067, 215-295-4734, evergreenl1@aol.com 
HARLAN, MAGGIE — Pennsylvania Native Plant Society, 166 Zachary Rd., Coburn, PA 16832, 814- 
349-5029, harlanmm_1999@yahoo.com 
HARPEL, WILLIAM — 150 E. Winchester Ave., Langhorne, PA 19047, 215-752-0802 
HARRIS, JESSIE M. — 4401 W St. N.W., Washington, DC 20007, 202-338-9083, jes405@hotmail.com 
HART, ROBIN L. — 22 Watson Rd., Poughkeepsie, NY 12603-3121, 941-351-1554 
HASSEL, LLOYD V. — 2001 Harrisburg Pike, Lancaster, PA 17601-2641, 717-569-2368 
HAUPT, ANDREA — 639 W. Ellet St., Philadelphia, PA 19119-3428, 215-438-8175, reh215@aol.com 
HAWK, JEFFREY — 16 Andrea Ln., Hamilton, NJ 08619-2222 
HECKSCHER, STEVENS — Natural Lands Trust, 1030 E. Lancaster Ave., Apt. 314, Rosemont, PA 
19010, 610-527-6324, heckscher@natlands.org 
HELD, MICHAEL — 285 Penn Estates, East Stroudsburg, PA 18301-9023, 201-915-9187 
HERMELY, ALAN — 671 Moore Rd., King of Prussia, PA 19046-1318, 610-337-3666, 
alan_hermely@urscorp.com 
HIRST, FRANKLIN S. — 5004 Little Mill Rd., Stockton, MD 21864-2234, 410-632-1362 
HOFFMAN, ROBERT — Fairweather Gardend P.O. Box 330, priacen ty NJ 08323, 856-451-6261 
HOCHNER, DIANA — 1024 Cemetery Road, Schwenksville, 9473 
HOWE, NATALIE — 4820 Cedar Ave., Philadelphia, PA nero, 617-435-4101, 
nataliemhowe@yahoo.com 
HUGHES, STUART W. — 721 Butler Pike, Maple Glen, PA 19002, 215-646-3873, stuhu@aol.com 
HUMBERT, KAREN C. — 2401 Pennsylvania Ave., 7A6, Philadelphia, PA 19130, 215-232-0295 
HUNT, LYNN F. — P.O. Box 906, Tuckerton, NJ 08087, 609-296-8022 
HUTCHEON, DAVID J. — 25 Caledonia Dr., Warminster, PA 18974, 215-957-0976 
INSKEEP, MICHAEL — 412 N Wayne Ave., Apt. 204, Wayne, PA 19087-3243, 610-896-0777, 
mike@gentlehelpers.com 
JANOSKI, JEFFREY F. — 1801 Buttonwood St., #1610, Philadelphia, PA 19130, 215-569-1949, 
jjfjeh@aol.com 
JEss, ROBIN — 55 Lahiere Ave., Edison, NJ 08817, 732-572-5928, rjess5928@aol.com 
JOHNSON, ALAN A. — 63 Central Ave., Audubon, NJ 08106, 856-547-3498, aajohnson@comcast.net 
JOHNSSON, FANNY M. — 7422 Ridge Rd., Frederick, MD 21702-3518, 301-371-5215, 
johnsson@starpower.net 
KAISER, GEOFFREY D. — P.O. Box 222, Sumneytown, PA 18084, 215-234-8424, 
taxkwaxak@enter.net 
KAPLAN, PAULA WEST — Little ong Farm, N653 County Rd. T, Brodhead, WI 53520, 608-897- 
3483, Tatlebhaflarm@hbtinnllics 
KELLER, ELIZABETH — 114 Wyomissing Blvd., Wyomissing, PA 19610, 610-374-3458, 
zab12345@aol.com 
KELLY, LINDA — 60 Forest Dr., Lakewood, NJ 88701, 732-363-1266, kellylcO1@aol.com 
KENDIG, JAMES W. — 1212 S. Prince St., Palmyra, PA 17078, 717-832-3899, kkendlarix@aol.com 
KLIGMAN, DOUGLAS — 1198 Rossiter Ln., Radnor, PA 19087, 610-688-7790, kligahm@earthlink.net 
KLOTZ, LARRY — Biology Dept., Shippensburg University, 305 N. wigs St., Apt. C, 
Shippensburg, PA 17257, 717-477-1402, larry.klotz@hotmail.co 
KOBLER, EVELYN — 118 21st St., Apt. 613, Philadelphia, PA 19103-4431, 215-563-4712, 
koblerev@earthlink.net 


MEMBERSHIP LIST 135 


KOERBER, WALTER A. 7 — 1380 Valley Green Rd., Etters, PA 17319, 717-938-9618, 
wakoerber@yahoo.co. 

KOERBER, DIANA — ‘aio Valley Green Rd., Etters, PA 17319, 717-938-9618, 
dbkoerber@hotmail.com 

KOLAGA, VAL — 186 Dilworthtown Rd., West Chester, PA 19382-8369, 610-399-3136, 
sfox@axs2000. net 

KRAIMAN, CLAIRE T. — 7129 N. Via de Paesia, Scottsdale, AZ 85258-3706, 610-775-9737 

KUNKLE, KARL — 507 Nina Ln., Bear, DE 19701, 302-834-2802, kkkunkle@aol.com 

KUNZ, DAVID — 300 Parsippany Rd., Apt. 24M, Parsippany, NJ 07054, 201-417-8082, 
dkunz@ekmail.com 

LADEN, MILTON — 1919 Chestnut St., #1407, Philadelphia, PA 19103, 215-568-6599 

LADIG, KIM — 712 Wood Lane, Haddonfield, NJ 08033, 856-354-3296, krladig@bellatlantic.net 

LANYON, HONOR — 1900 J. F. Kennedy Blvd., Apt. 1704, Philadelphia, PA 19103, 215-567-5543 

LAMONT, ERIC — 717 Sound Shore Rd., Riverhead, NY 11901, 631-722-5542, 
elamont@optonline.net 

LATHAM, ROGER — Continental Conservation, P.O. Box 57, Rose Valley, PA 19086-0057, 610-565- 
3405, rel@continentalconservation.us 

LEA, CHRISTOPHER — P.O. Box 457, Pine, CO 80470-0457, 410-641-1443, chrisleal5@yahoo.com 

LECK, MARY — 105 Kendall Rd., Kendall Park, NJ 08824-1246, 732-821-8310, leck@rider.edu 

LECK, CHARLES — 105 Kendall Rd., Kendall Park, NJ 08824-1246, 732-821-8310 

LEFEVRE, WILLIAM M. — 13 Hickory Dr., Doylestown, PA 18901-4746, 267-880-0860, 
wmilefevre@aol.com 

LEVIN, MICHAEL H. — 414 Mill Rd., Havertown, PA 19083, 610-449-7400, erainemhI@aol.com 

LEWANDOWSKI, RICK J. — Mt. Cuba Center, P.O. Box 3570, Greenville, DE 19807-0570, 302-239- 
7980 


LIGHTY, RICHARD — 501 oe a Mill Rd., Kennett Square, PA 19348, 610-444-2987, 
taraxicum@earthlink.ne 

LOEFFLER, CAROL — Best of anes Dickinson College, P.O. Box 1773, Carlisle, PA 17013, 
717-245-1360, loeffler@dickinson.e 

LOFURNO, MICHAEL — 2028 Pins eaidi St., Philadelphia, PA 19146, 215-732-0849, 
mlfcomp@aol.com 

MACKLIN, JAMES — Academy of Natural oa 1900 Benjamin Franklin Pkwy., Philadelphia, 
PA 19103, 215-299-1192, macklin@acnatsci 

MARTIN, HARRIS — 5100 Bay Rd., Saashiess PA 19020, 215-639-3686, martin68@atc-enviro.com 

MAURICE, KEITH R. — 63 Union Ave., New Holland, PA 17557-1343, kmaurice@normandeau.com 

McAvoy, WILLIAM A. — 4876 Haypoint Landing Rd., Smyrna, DE 19977, 302-653-2880, 
william.mcavoy@state.de.us 

MCELWEE, KAREN — 300 W. Vernon Ave., Linwood, NJ 08221, 609-653-6332 

MCCoMBE, ROBERT — 445 E. Main St., Moorestown, NJ 08057, 609-313-5000, 
rob.mccombe@att.net 

McCourt, RICHARD — Academy of Natural Sciences, was: ae Franklin Pkwy., 
Philadelphia, PA 19103, 215-299-1157, mccourt@acnatsci 

MCCREA, ESTHER J. — 70 Overhill Rd., Bala Cynwyd, PA "18004, 610-664-1642 

McDaDE, LUCINDA — Academy of Natural Sciences, 1900 Benjamin Franklin Pkwy., Philadelphia, 
PA 19103, mcdade@acnatsci.org 

MCLAUGHLIN, WILLIAM — 5 Oak Dr., Tabernacle, NJ 08088, 609-268-1054, wmcl268@aol.com 

MELENDEZ, MARISA — 1005 lakewood Farmingdale Rd., Howell, NJ 07731, 732-566-0297, 
marisabel626@hotmail.com 

MELLON, RICHARD — 200 Flint Cr. S., Yardley, PA 19067, 215-493-0697, rmellon@voicenet.com 

MICKLE, ANN — Dept. of Biology, La Salle University, 1900 W. Olney Ave., Philadelphia, PA 
19141, 215-951-1254, mickle@lasalle.edu 

MITCHELL, JOHN — 56 Upper Creek Rd., Stockton, NJ 08559 


136 BARTONIA 


agp none — 305 Westtown Circle, West Chester, PA 19382-7653, 610-344-0860, 
martinmilner@earthlink.net 

BP 0 SEs HARRIET — 176B Lark Ln., Cherry Hill, NJ 08003, 856-428-2342, 
v.r.monshaw@iece.org 

MONTGOMERY, JAMES — Ecology I, 804 Salem Blvd., Berwick, PA 18603, 570-542-2191, 
jdm@sunlink.net 

MOoBERRY, F.M. — 111 Spotswood Ln., Kennett Square, PA 19348-1725, 610-444-5495 

MorrIs, MELISSA — 234 Broughton Ln., Villanova, PA 19085, 610-525-6659, 
mhmorris@ucdavis.edu 

Moss, MIRIAM — 8120 Brookside Rd., Elkins Park, PA 19027, 215-635-0176 

NACZI, ROBERT — Herbarium, aor of Agriculture, Delaware State University, Dover, DE 19901- 
a 302-857- — rnaczi@desu.edu 

NEWSTEAD, EDWIN — 270 Roseland Ave., Essex Fells, NJ 07021, 973-226-7651 

Sa ce — 270 Rosléad Ave., Essex Fells, NJ 07021, 973-226-7651 

NICHOLS, RAY — 412 Federal City Rd., Pennington, NJ 08534-4209, 609-737-7442, 
nichols@nj1.aae.comr 

NOVAC, JANET — 19 Willow Grove Ave., Philadelphia, PA 19118, 215-248-2642, janet@indri.org 

O’BRIEN, TERRY — Dept. of Biological Sciences, Rowan University, Glassboro, NJ 08028, 856-256- 
4500, obrien@rowan.edu 

Pag Lage 9 fo nese Bridge Rd., Newtown Square, PA 19073-1211, 610-359-9887, 


age@ms 
Beene oe. "1518 Millstone River Rd., Hillsborough, NJ 08844, 908-359-2073, 
mipalmer@eden.rutgers.edu 
PAYNE, ELISE — 643 Fernfield Circle, Wayne, PA 19087, 610-688-4377, aparece com 
, D. — 418 Crosslands Dr., Kennett Square, PA 19348, dotp77@ 
PREUCEL, RUTH — 1147 Norsam Rd., Gladwyne, PA 19035 
QUIGLEY, PATRICIA — 1080 Quarry Hall Rd., Norristown, PA 19403, 610-584-1829, 
paqinc@ao .com 
RADIS, RICHARD — 69 Ogden Ave., Rockaway, NJ 07866, 973-586-0845, isotria@bellatlantic.net 
RHOADS, ANN — 3 Blythewood Rd., Doylestown, PA 18901, 215-348-8139, 
rhoadsaf@pobox.upenn.edu 
RHODES, CHARLES — 107 Stonycreek Ave., Lansdale, PA 19446-5259, 215-368-9591, 
carhodes@earthlink. net 
RISKA, MICHAEL — Delaware Nature Society, Box 700, Hockessin, DE 19707, 302-239-2334, 
ike@dnsashland.or 
ROCHE, LESLIE — 73 Hull Ave., Freehold, NJ 07728, 732-780-0121 
ROM , GABRIELLA — 4240 Fairview Ave., Newtown Square, PA 19073, 610-353-1550 
ROSENBERG, ANN — 5 Westview Rd., Bryn Mawr, PA 19010, 610-525-8683 
ROSENBERG, DICK — 5 Westview Rd., Bryn Mawr, PA 19010, 610-525-8683 
ROWAN, JANE — Schnabel Engineering Associates, Inc., 510 E. Gay St., West Chester, PA 19380, 
610-696-6066, jrowan@schnabel-eng.com 
RUCH, PAMELA — 542 Liberty St., Emmaus, PA 18049, 610-966-2635, pamruch@hotmail.com 
RUDYJ, ERICH — 1021 Irwins Choice, Bel Air, MD 21014, 410-420-8973, 
erich.s.rudyj@aphis.usda.gov 
RUE, MARGARET — 209 Arden Rd., Gulph Mills, ae a 
RYAN, JOEY — 521 S. Narberth Ave., Merion, PA 1 
SCAGNELLI, RENEE — 226 N. Lincoln Ave., sirens = 08361, 816-305-3238, 
pbfiddler@yahoo.com 
SCHNEIDER, GEORGE W. — 345 Nursery Rd., Wellsville, PA 17365, 717-292-4035 
SCHUBERT, PAUL — 315A Dickinson Ave., Swarthmore, PA 19081-2002, 610-543-9450, 
pauls@depersico.com 


MEMBERSHIP LIST 137 


SCOTT, JOHN — 55 pa School Rd., Mertztown, PA 19539, 610-682-2809, 
johndscott@mindspring.co 

SEAGER, KEITH — 278 Fishing Creek Rd., Cape May, NJ 08204-4422 

SEIFFERT, JUDITH — 1135 E. Oxford St., Philadelphia, PA 19125, 215-426-4201 

SEIPLER, MARYJANE — 4200 Tamarack Ln., Murrysville, PA 15668-9353, 724-325-3571, 
grosbeaker@aol.com 

SETTLEMYER, KENNETH — 219 Maple St., Jersey Shore, PA 17740, 570-398-2546, 
ksettlem@uplink.net 

SHERMAN, LYNN — P.O. Box 84, Barstow, CA 92312-0084, 518-756-5028, shermanL@capital.net 

SLANE, JOSEPH — 1207 Shackamaxon St., Philadelphia, PA 19125, 215-634-6332, 
jslane@avantgardens.net 

SLATER, MIKE — 4411 New Holland Road, Mohnton, PA 19540, 610-775-3757 

SMITH, CRAIG — 728 Seymour Rd., Bear, DE 19701, 302-324-9486, csmith6666@aol.com 

SPEEDY, LOREE — 279 Orr Rd., West Newton, PA 15089, 724-872-5232, yoree@stargate.net 

ST. JOHN, MICHAEL — 4 Perry St., New York, NY 10014, misaintjon@aol.com 

STAHL, STANLEY — 104 Harvard Ave., Lancaster, PA 17603-1704, 717-293-0292, 
joselyns@fine_hotels.com 

STALTER, RICHARD — Dept. of Biology, St. John’s University, Jamaica, NY 11439, 718-990-6269 

STANDAERT, WILLIAM — 45 Maltbie Ave., Apt. 19B, Midland Park, NJ 07432, 201-612-9069, 


wtszzz@aol.com 

STECKEL, DAVID — 423 N. 27th St., Allentown, PA 18104, 610-740-0141 

STECKEL, CLAUDIA ~— 423 N. 27th St., Allentown, PA 18104, 610-740-0141 

STUCKEY, RONALD ~— 1315 Kinnear Rd., Columbus, OH 43212-1192, 614-292-6095 

SU AN, MAURA — 164 York St., Lambertville, NJ 08530, 609-397-0467, 
mauraesullivan@earthlink.net 

SWEENEY, BILL — Lockhouse, 393 Island Park Rd., Easton, PA 18042, 610-253-7053 

SZURA, BRIAN — 216 Volkert St., Highland Park, NJ 08904, 732-393-1950, flyin.brian@verizon.net 

TEESE, PAUL — Bowmans Hill Wildflower Preserve, 1183 Apple Rd., Quakertown, PA 18951, 215- 
536-3719, teese@bhwp.org 

TETI, ANDREA — 31 Boulder Dr., Sellersville, PA 18960 

THOMPSON, BRAD — 235 W. Winona St., Philadelphia, PA 19144, 215-951-2092, 
thompsonw@philau.edu 

TOFFEY, WILLIAM — 9001 Verree Rd., Philadelphia, PA 19115, 215-969-6789, wtoffey@prodigy.net 

TREXLER, LARRY — 7501 Germantown Ave., Philadelphia, PA 19119, 215-242-1309 

TREXLER, NANCY — 7501 Germantown Ave., Philadelphia, PA 19119, 215-242-1309 

TUCKER, ARTHUR — Dept. of Agriculture & Seal Resources, Delaware State University, 
Dover, DE 19901, 302-857-6402, atucker@desc.edu 

TYNDALL, WAYNE — 15245 Oakland Rd., Goldsboro, MD 21636, wtyndall@intercom.net 

UDELL, VAL — 2746 Yost Rd., Perkiomenville, PA 18074, 610-754-7163 

VINCENT, MICHAEL — Dept. of Botany, Miami University, Oxford, OH 45056, 513-529-2755, 
vincent@myohio.edu 

VOLLMER, JOHN — 42 Burrs Mill Rd., Southampton, NJ 08088, 609-859-2805 

WECHSLER, DEBORAH CARR — 314 E. Highland Ave., Philadelphia, PA 19118, 215-242-0734 

WHITING, GEORGE — 116 Spring Mount Rd., Schwenksville, PA 19473, 610-287-6397, 215-283- 
1297, gwhiting@temple.edu 

WIEBOLDT, THOMAS — 155 Shady Grove Ln., Newport, VA 24128, 540-544-7967, wieboldt@vt.edu 

WILEN, ELLEN — 143 Ridge Rd., Southampton, NJ 08088, 609-859-8685 

WILHELM, DONNA C. — 6115 McCallum Street, Philadelphia, PA 19144 

WILLIAMS, CARL — Box 347, Hereford, PA 18056, 215-234-0545 

WILLIAMS, DAVID — 61 Coppermine Rd., Princeton, NJ 08540, 732-297-0642, idwill@superlink.net 

WILLIG, SARAH — 190 Sycamore Ln., Phoenixville, PA 19460, 610-933-3539, 
sallywillig@earthlink.cnet 


138 BARTONIA 


WILSON, RONALD — 3740 Ridge Rd., Snow Hill, MD 21863, 410-632-3892, rmwilson@comcast.net 

WINDISCH, ANDREW — P.O. Box 312, Chatsworth, NJ 08019, 609-726-9054, windisch2@erols.com 

WITMAN, DEANNA — 214 3rd St., East Greenville, PA 18041, 215-541-1789, d.witman@verizon.net 

WOLFF, EMILY ~— 411 N. Middlevsuti Rd., # E-202, Media, PA 19063-4404, 610-566-6387 

WoOoD, HOWARD — 3300 Darby Rd., C-802, Haverford, PA 19041, 610-642-9963, 
howardpagewood@aol.com 

WOODFORD, JEANNE — Wildlife Advocates, 2 Sawmill Rd., Medford, NJ 08055, 856-983-1617, 
jeannewoodford@earthlink. net 

YOUNG, MARK — 2800 Forest View Ave., Baltimore, MD 21214-3109, 410-254-8425, 
amersfoort@msn.com 

= ROBERT — 12 Concord Dr., Shamong, NJ 08088, 609-268-3363, 

ampella@njpines.statenj.us 
Poesy HEINRICH — 411 N. Middletown Rd., Apt. E-111, Media, PA 19063, 610-754-7573 


BARTONIA TT 


Instructions to Authors 


Types of Articles Published oe 

Research papers communicate original research in plant ecology, cla cos on biology, plant 
systematics, and related topics. Other contributed papers convey the results. of srudien in n floristics, 
distribution, methods, biography, bibliography, t 


book reviews, and field trip reports. The focus is on he mid gi 
Delaware, Maryland, New York, Virginia, and West Virginia), hae contri f interest to Barto 4 
readers from farther afield are welcomed. : : lee 


Manuscript Style 
Write in simple, clear sentences. Use the active voice where possible. Avoid | 
gipsieed conforms to the Council of Biology E 
t recent issues of Bartonia for style of main headings, subheadings, literature. 
oe captions, and tables. 
uble-space all text, including tables. 
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