·arno ~aVolume 61 · Number 2 - · 2001

Arnoldia (ISSN 004-2633; USPS 866-100) ispubhshed quarterly by the Arnold Arboretum ofHarvard University. Second-class postage paid atBoston, Massachusetts.

Subscriptions are $20.00 per calendar year domestic,$25 00 foreign, payable m advance. Smgle copies ofmost issues are $5.00; the exceptions are 58/4-59/1(Metasecluoia After Fifty Years) and 54/4 (A Source-book of Cultmar Names), which are $10 00. Remit-tances may be made m U.S. dollars, by check drawnon a U.S bank; by international money order; orby Visa or Mastercard. Send orders, remittances,change-of-address notices, and all other subscription-related communications to’ Circulation Manager,Arnoldm, The Arnold Arboretum, 125 Arborway,Jamaica Plam, Massachusetts 02130-3500.Telephone 617.524.1718; facsimile 617.524.1418;e-mail arnoldia@arnarb.harvard edu.

Postmaster: Send address changes toArnoldia Circulation ManagerThe Arnold Arboretum125 ArborwayJamaica Plam, MA 02130-3500

Karen Madsen, EditorMary Jane Kaplan, CopyeditorAndy Wmther, Designer

Editonnl Committee

Phylhs AndersenEllen S. BennettRobert E. CookPeter Del Tredici

Gary KollerStephen A. SpongbergKim E. Tnpp

Copyright © 2001. The President and Fellows ofHarvard College

Page2 Plant Hunting on the Rooftop of the World

Susan Kelley

14 Anachronistic Fruits and the GhostsWho Haunt Them

Conme Barlow

22 Pastures of Plenty: A Case Studyin Field BiologyP. L. Marks

26 Henry David Thoreau and the Yankee ElmThomas J. Campanella

Front e·J back cover The barley harvest m southeast-ern Tibet photographed by Susan Kelley m July 2000.Barley is the only crop that grows at the highelevations of the Tibetan Plateau.

Inside front cover E. H Wilson photographed the"Knight" elm, near Newburyport, Massachusetts, m1927. He measured its girth at 18 1/2 feet four feetfrom the ground and 46 feet at ground level. From theArchives of the Arnold Arboretum.

Inside back cover. The "Howard" elm m Ware,Massachusetts, measured over 19 feet m circumferenceat three feet from the ground and 100 feet through thecrown when E. A. Richardson photographed it m1904. From the Archives of the Arnold Arboretum.

Anachronistic fruits of North America. Photographby Conme Barlow.

Plant Hunting on the Rooftop of the World

Susan Kelley

lant exploration has always played animportant role in shaping the ArnoldJL Arboretum’s collections and has been

the driving force behind the many Arboretum-sponsored trips to the Far East and within NorthAmerica. Living plants grown from seeds gath-ered on these expeditions grow on the groundsof the Arnold Arboretum, in the collections ofother botanical institutions in North Americaand abroad, in the stock inventories of nurseriesacross the country, as well as m our own home

gardens. Although new plant material fromexpeditions is added to the living collectionseach year, the main goal of the majority ofArboretum-sponsored fieldwork is the creationof botanical inventories of eastern and south-eastern Asia in the form of herbarium vouchers.In fact, few people familiar with the Arbo-retum’s collections in Jamaica Plain are aware ofthe institution’s collection of approximately 1.4million herbarium specimens housed in theHarvard University Herbaria in Cambridge,Massachusetts (http://www.huh.harvard.edu/).

In 1997, under the auspices of the BioticSurveys and Inventory program of the NationalScience Foundation (NSF), the Arnold Arbore-tum began a three-year collaborative effort toinventory the plant and fungal diversity in theHengduan Mountains of south-central China,one of the unique biological regions of theworld. Lying at the eastern end of the Himalayasbetween the edge of the Qinghai-Xizang(Tibetan) Plateau and the central plain of

China, these spectacular north-south trendingridges contain the most diverse vascular plantflora of any region of comparable size in thetemperate zone. Identified as one of twenty-five biodiversity "hotspots" on earth,’ this vastregion, covering an area of approximately300,000 square miles (500,000 sq km), containsover 12,000 species of vascular plants, with

almost 3,500 endemic species and at least 20endemic genera.Although some botanical exploration has pre-

viously been carried out in the HengduanMountains, the region has never been fullyinventoried because of the sensitive politicalatmosphere in Tibet and because the ruggedterrain makes much of the area extremelydifficult to traverse. Elevations range from 3,300feet (1,000 m) to over 25,000 feet (7,556 m/ at thesummit of Gongga Shan in western Sichuan.Average elevation is 10,000 to 13,000 feet (3,000to 4,000 m) with precipitous drop-offs of 1,000to 3,000 feet (300 to 900 m) not uncommon.No one, however, has yet identified the fullextent of the geography and plant life of thisparticular "hotspot."The term "hotspot," coined in 1988 by British

ecologist Norman Myers, is used to designateareas that have a high number of endemic spe-cies (those whose distribution is limited to asingle region) and that are under severe threat ofdestruction because of human activities. Thesethreatened regions cover less than two percentof the earth’s land area, but are home to morethan sixty-five percent of all vascular plant spe-cies. Of the twenty-five designated hotspots, theHengduan Mountains and the California Floris-tic Province are the only two located in theNorthern Hemisphere. All other hotspots, withthe exception of central Chile, the Cape Prov-ince of South Africa, and southwestern Austra-lia, are located in the tropics.The Hengduan Mountain region, as currently

defined, constitutes only five percent of China’sland area, occupying portions of southeasternXizang (Tibet), western Sichuan, and northernYunnan, but it contams almost half the totalnumber of all Chinese flowering plant species.The extremes in climate and topography almostcertainly contribute to the diversity of plant life

The upper reaches of the Mekong River and one of its tributaries.

4

The Hengduan Mountain region, one of the twenty-five biodiversityhotspots of the world.

there. More than a quarter of the world’s Rhodo-dendron, Primula, Corydahs, Delphinium,Anaphalis, Gentiana, Saussurea, and Sorbusspecies and over half the species of Ligularia,Cremanthodium, Cotoneaster, and Pedicularishave been recorded here. In addition, there maybe as many as fifty species of endemic mosses.A few numbers will illustrate the extent of

some particularly species-rich groups in thespectacularly diverse Hengduan region (approxi-mate numbers of species worldwide in paren-

thesesZ: Rhododendron-224 (850);Androsace-28 ( 100/; Pmmula-113

(400) ; Gentiana-117 (350/; Saussurea-101 (300); Impatiens-45 (850);Pedicularis-250 (350+); Aconitum-104 (100+); Delphinium-71 (250/;Arisaema-39 (150) ; Cotoneaster-41(50) ; Astragalus-98 (2,000) ; Ilex-44(400) ; Corydalis-85 (300); Sorbus-36(85/; Anaphalis-33 /100/.The table on the opposite page com-

pares the levels of diversity and ende-mism in the Hengduan region withthose of other nontropical areas, provid-ing another indication of its richness.3Four of the great rivers of Asia, the

Yangtze (Jinsha Jiang), the Mekong(Lancang Jiang), the Salween (Nu Jiang),and the Brahmaputra (Yarlung ZangboJiang), flow through the valleys of thesedramatic mountains. All of these rivers

origmate on the 16,500-foot (5,000-m) highQinghai-Xizang (Tibetan) plateau, and, far

downstream, all are of great economic impor-tance to the people who live along them. Therapidly increasing human impact on the regionthreatens not only the diversity of plants andanimals there, but also the survival of indig-enous cultures that define much of eastern andsoutheast Asia.

In the summer of 2000, the third year of theNSF project, fieldwork was conducted for two

--. - - ~~ - ------

Three species of Saussurea, a member ofthe aster family: above, a cushion formgrowing on scree slopes at 17,000 feet,left and center, two species grow amongboulders at 15,000 feet

months in southeast Xizang (Tibet) by a team offour American, one Tibetan, and four Chinesebotanists. Dave Boufford, assistant director ofthe Harvard University Herbaria and an authorof the NSF grant proposal, headed the Americanteam. I was fortunate to be part of that team,along with Rick Ree, who received his Ph.D.from Harvard this year and works on the genusPedicularis (lousewort), and Brian Perry, a doc-toral student at Harvard in mycology. FourJapanese botanists traveled and collected withthe group as well, but were not working underthe auspices of the NSF grant.The logistics of the expedition were coordi-

nated by Wu Sugong of the Chinese Academy ofScience Institute of Botany in Kunming inYunnan Province. Professor Wu had done field-work in the area in the 1970s and 1980s andhelped compile the two-volume checklist Vas-cular Plants of the Hengduan Mountains.’ Hisposition was not an enviable one, smce effortsto arrange permits, lodging, and rations werecontinually complicated and delayed by land-slides, broken-down vehicles, massive roadway

construction projects, obstinate local officials,and inclement weather.

The expedition team assembled at the KunmingBotanical Garden on June 28, 2000. There wehanded over our passports to a young Tibetan

woman, Yang Zhen, who was to fly them toLhasa to obtam permits for travel in the TibetAutonomous Region. While waiting for herreturn, we drew up the final itinerary and gath-ered equipment: plant presses, ventilators, por-table dryers, kerosene burners, a fifty-gallondrum of kerosene, food, camping gear, and evenan extra pair of springs for the rather rickety busthat was to transport much of our equipment.Little did I suspect how valuable these springswould prove to be weeks later down the incred-

ibly rough road.Finally, on July 4, with three SUV’s and a

small bus, we began our journey north. Thefully loaded truck remained m Kunming to waitfor our passports and the all-important permitsto arrive from Lhasa. For the next several dayswe headed north, stopping in the towns of Dali,

6

Wu Sugong and Dave BoufroW Li

Zhongdian, and Deqen. In Dali, a popular Chi-nese tourist destination situated next to Lake

Erhai, we encountered a few Tibetan vendorswhose wares included fruiting bodies of theCordyceps fungus, a dried tiger penis ("For yourhealth!"), and the antlers of an unidentifiedantelope. We took advantage of the town’sinternet cafe, which would be the last we sawuntil we reached Lhasa on August 17.During the eighth and ninth centuries, Dali

was the capital of a separate kingdom, Nanzhou.The local Bai people-who ruled from 902 until1252, when Kublai Khan conquered the area-renamed it the kingdom of Dali. Diancang Shan,an uplifted mountain of granite and marble,rises 13,500 feet (4,100 m) just west of the city.Since most of the remainder of the province of

Yunnan is limestone, the flora of

Diancang Shan is distinctive and inter-esting in and of itself. In 1984 DaveBoufford spent six weeks collectingherbarium specimens there; on thistrip we could spend only a day on themountain.

Heading north we saw fields of

tobacco, corn, rape seed (used for cook-ing oil), cabbage, beans, and peas. LargePopulus yunnanensis grew along theroadsides, but most of the land wasstripped completely of woody vegeta-tion. Joseph Rock, the Viennese-bornbotanist, ethnologist, and linguist,made his home in Lijiang for somethirty years. Between 1924 and 1927,he collected hundreds of herbarium

specimens for the Arnold Arboretumand sent back seed of many new plantsfor its living collections.s

For more than an hour the road fol-lowed the Yangtze River (Jinsha Jiang), abroad, muddy expanse. The weatherwas warm, but clouds prevented usfrom seeing Yulongxue Shan (SnowMountain), at 18,467 feet (5,596 m) thehighest in Yunnan. Growing on theroadside were species of Philadelphus(mock orange), Sambucus (elderberry),Pyracantha, Indigofera (indigo), juni-per, and an evergreen oak with a dense

covering of yellow-brown hairs on theunderside of the leaves. By the time we reachedZhongdian early in the evening, we had gonefrom an elevation of 7,000 feet (2,100 m/ in Dali,to about 10,500 feet (3,200 m).Zhongdian, close to the Tibetan border, was

the first town in which we saw a sizable popula-tion of Tibetans. Some of the shops had Tibetangoods for sale, such as silver bracelets, coral andturquoise necklaces, daggers, and clothing. Thearchitecture of the town, however, was typicalHan Chinese white tile bmldings. Alongside themodern food shops, banks, restaurants, hotels,and CD and DVD shops, pigs ran in the muddystreets and vendors in open-air markets offeredhousewares, horse blankets and saddles, freshyak meat, live chickens and fish, vegetables,brooms, baskets, pots and pans. One vendor had

7

Rheum nobile, a type of rhubarb (also seen in closeup at right), and several species of Pmmula grow nearTiensi Lake m northern Yunnan.

stacks of fresh yak butter, which I had tasted thatmorning m a bowl of tea. In the outlying areas,we saw the first signs of Tibetan architecture,square, two-story stucco structures, paintedwhite but with colorful decorations around the

doorways and windows. Buddhist prayer flagsflew from the rooftops.We spent two days in Zhongdian, waiting for

Yang Zhen to arrive with our passports and thepermits needed to cross the provincial borderinto Tibet. We spent the time exploring the veg-etation in the mountains outside Zhongdian.Lake Tiensi, at 13,500 feet (4,100 m), is a beau-tiful alpine lake about two hours fromZhongdian. Growing there among grazing yaks,we found Rhododendron wardii, a tall rhodo-dendron with pale yellow flowers named forBritish botanist Frank Kingdon-Ward. Between1909 and 1957, Kingdon-Ward introduced hun-dreds of new species into cultivation (rhododen-drons, primroses, gentians, the Himalayan bluepoppy) from here and other parts of westernChina and from upper Burma and FrenchIndochina. We also found the distinctive Rheumnobile, a rhubarb with large, pale yellow bracts;purple and yellow species of Primula; and a yel-low Mecanopsis, the Himalayan poppy. Twocurious Tibetan boys, perhaps nine or ten yearsold, appeared seemingly out of nowhere.

Although initially quite shy, they followed us ata distance and eventually helped our mycologistfind several interesting fungi.

Our second day in the Zhongdian area wasspent in drizzling rain at Shudu Hu, anotheralpine lake, this one surrounded by heavilygrazed meadows with patches of cut-over spruceforests. Scattered m the meadows were

primroses, asters, gentians, and Stellera

chamaejasme, a member of the thyme familywhose corolla color varies from yellow to pinkacross its geographic range and even m smglepopulations.From Zhongdian we drove toward Deqen,

closer to the Tibetan border. In the early after-noon we reached a 14,000-foot (4,250-m) passwhere for the first time we saw Tibetan nomadtents woven of black yak hair, with largebranches of juniper propped against the doors forinsulation. A few specimens of Pinus armandii,almost 60 feet (18 m) high, with bright green,pendulous cones, were growing there alongwith a few specimens of P. yunnanensis and P.densata, but most of the woody vegetation hadbeen cut for fuel or lumber. (There is a nice 65-foot [20-m] specimen of P. armandm on PetersHill that was grown from seed collected in 1909by William Purdom m Shaanxi Province, to thenortheast of the Hengduan Mountains.)On July 11 we finally crossed into the prov-

ince of Xizang (Tibet), descending from almost11,000 feet (3,265 m/ in Deqen to 7,000 feet(2,100 m/ on the banks of the upper MekongRiver. The landscape here was dry and sparselycovered with scrubby vegetation, with a few

9

low, cut-over shrubs and many herba-ceous plants growing along the road.Following the road up from the

Mekong, we came to the village ofYenyng where we had lunch in a low,dark, wooden dwelling (complete withsatellite dish on the roof~, with almostunbearable smoke flowing from thecooking area.Within fifteen minutes of leaving

town, we climbed another 1,000 feet(300 m) and soon began seeing mixedbroadleaf deciduous and conifer forestsof Salix, Ables, Populus, Cornus, andQuercus, along with many herbaceousalpine plants growing by the roadside.After two hours we reached yet another

14,000-foot (4,250-m) pass, Hong La,and began the descent to the town ofMarkam. The fifty-four-mile journeyfrom our lunch spot had taken morethan three-and-a-half hours over rough,narrow roads that required our driversto negotiate numerous hairpin turnsand the remains of recent landslides

overlooking precipitous drops of morethan 1,000 feet (300 m/.Markam, lymg in a long, wide valley

created by a tnbutary of the Mekong, istruly a Tibetan outpost, a primitive,extremely poor town with red mudeverywhere and a wild west look to it.It seemed strange amidst the squalor to hearchants emanating from the walls of a Buddhisttemple. While most Tibetan men wore tradi-tional long, fur-lined coats with one sleevehangmg off the shoulder, a few wore ChicagoBulls ~ackets. The women, in traditional dressescalled chhubas, stared at us as we walked downthe street. A large group of local people followedus to our Chinese guesthouse. It was a bit

unnervmg to have five or six of them watchingfrom the doorway, long silver daggers hangingfrom their belts, as we pressed plants andentered data into our laptop computer. Anotherguesthouse-a government-sponsored temple-restoration pro~ect-and the early-morningradio news that blared out from speakers ontall poles were the only signs of non-Tibetan

Dave Boufford bargaining with Tibetan men m Markam.

influence. In fact, this would be the only townin which we saw vestiges of the Tibetan culture,albeit in shambles.The following day we retraced the road to

Hong La (Pass), south of Markam, and madeninety-six collections of vascular plants andmosses in a mixed broadleaf deciduous andconifer forest. Two government officials accom-panied us, but they seemed quite uninterestedin our work. For two more days we collected mdifferent habitats around Markam. Heavilygrazed meadows of grass and Kobresia, a type ofalpme carex, were dotted with scrubby rhodo-dendrons and remnants of spruce and juniperforests. Many genera were very familiar toWestern eyes: Gentiana, Ligulana, Lomcera,Berberis, Carex, Clematis, Rubus, Anemone,

A small Tibetan village In a valley near Markam.

10

Flowers of Clematis chmbmg on shrubs at a forest’sedge near Nyingchi m southeastern Tibet.

Trollius, Potentilla, Pedicularis, Picea, Cam-panula, Cerastium, Polygonum.

Three days and fifty-some collections later, weheaded northwest, climbing another pass abovethe Mekong River. Apricots were ripening above12,000 feet (3,665 m), as well as dark blue Del-phinium, tall Thalictrum, and several ferns. Thelandscape was vast, and although a few scatteredpine, spruce, and juniper remained, there wasmuch evidence of clear cutting. Our sense of thestrangeness of this land was intensified by thesight of three monks on the road in long, heavycarmine robes, making their pilgrimage to Lhasa,some 400 miles (250 km) away.

A species of Meconopsis, theHimalayan blue poppy, growingat 17,000 feet at Dongda La.

It was at DongdaLa, south of Zogong,that we hiked to

17,500 feet (5,300 m~to collect alpineperennials from thescree slopes. In thelevel areas of the

glacial cirques, wecollected two Hima-

layan poppies (oneblue and one an

intense yellow), a

creeping willow, sev-eral species of rhodo-dendron, and dozensof tiny herbaceous

plants. The diversityin this barren land-

scape was amazing:

in the three days that we spent around the townof Zogong, we collected 57 flowering plants andover 140 mosses.While we collected in the field, Wu Sugong

drove to Changdu (Qamdo) to obtain permits fortraveling west to Lhasa across the northern roadof the Plateau. The local officials in Zogong hadinformed us that a major bridge had washed outwest of the town of Bomi, so the southern routewould be impossible to negotiate. On July 19the entire team set out for Changdu, a 45-mile(75-km) trip that would take six hours becauseof the now familiar delays caused by road con-struction, landslides, and generally rough roads.Changdu, the second largest town in Tibet,

lies on the banks of the muddy Mekong and wasonce a thriving population center. Its large mon-astery complex, dating to 1444, formerly housedmore than 5,000 monks. Only a few hundredmonks remain today, but the Chinese govern-ment is providing funds to restore their livingquarters and the many temples of the lamasery.Elsewhere in the town, however, all traces of Ti-betan culture are being destroyed to make roomfor more modern Chinese architecture and

goods. Although Changdu lies on the main roadfrom Sichuan and attracts many tourists fromthat province, its best hotel can offer hot run-ning water for only one-and-one-half hours eachnight, and even that isn’t guaranteed. Massiveconstruction projects throughout the city oftendisrupt the basic services that Westerners takefor granted.Our group was able to spend only one day

collecting along the Mekong River south ofChangdu. The dry slopes and ravines harbor axeric shrub vegetation dominated by herbs andgrasses. Among the thirty-four species we col-lected were Salweenia wardii, an endemicmember of the pea family, and Tribulusterrestris (devil’s thorn), which grew for manyyears m California as a noxious weed that hasonly recently been eradicated.Most of the expedition’s fieldwork had been

scheduled to take place around the southerntownships of Bomi, Yigong, and Nyingchi. Thenews about bad road conditions west of Bominow made a long stay there unfeasible if we wereto arrive in Lhasa on schedule, but we were deter-mined to spend at least a few days in the area.

11 I

The trip to Bomi from Changdu took ten days,but along the way we collected in a number ofinteresting habitats. On the outskirts of Banda,a tiny outpost consisting of two restaurants, afew shops, a primitive guesthouse, and a

Chinese army base, we collected for two days.We spent the first day along a steep, gravellymountain slope (15,700 feet; 4,760 m~ and atthe crest of a limestone ridge in a Kobresiameadow with Potentilla, Sibiraea, Rhododen-dron, and Salix shrubs interspersed with

species of Corydahs, Lonicera, Caragana, Spi-raea, Paraquilegia, Lepisorus, Cryptogramma,Pedicularis, and Draba. On the second daywe ventured east of Banda to a dry ravine andslope just above the Mekong River. There weencountered a family of Tibetan children col-lectmg firewood, and throughout the day wecould hear them laughing, talking, and singing.The woody plants had all been cut by the localpeople, but a rich flora remamed. Among thefifty taxa we collected that day were Gentiana,Geranium, ten species of Pedicularis, Ranuncu-lus, Stipa, Rheum, Allium, Artemisia, Silene,and Astragalus.On July 24 we began our serious push west to

Bomi. Our caravan and the other vehicles trav-

eling along this road were forced to negotiatearound numerous landslides and road construc-tion projects. Indeed, the sixty-mile journey tothe town of Baxio took almost ten hours. Hav-

ing crossed several more mountain passes, wewere now in the Salween River valley. TheSalween is as muddy a river as the Yangtse or

the Mekong, but the land around it is as dry asany desert, and the vegetation is sparse.

In the small village of Rawu we saw evidencethat deforestation was occurring to the west.

Large trucks were unloading logs up to threefeet (one m/ in diameter in the local lumberyard.Our accommodations in Rawu were in the mili-

tary compound, where we took our meals withmembers of the Chinese army and watchedthem perform their early morning drills. Wespent four and a half days in and around thisbeautiful valley: on the moist, open slopes alongRawu Lake; in alpine meadows; on grazed slopesdotted with Juniperus, Rhododendron, Salix,Sorbus, and Potentilla glabra; on a boulder-strewn mountain slope along the PalongzangRiver; and m a broad, gravelly floodplain domi-nated by small specimens of Hippophie. (Thisgenus, a member of Elaeagnaceae, containsthree species. The Arboretum has made severalattempts to grow H. rhamnoides (sea buck-thorn) and H. salicifolia, but conditions in theBoston area do not appear to be ideal for thesetaxa.) One particularly cold, miserable day, wewere invited into the tent of a Tibetan familyto sit by an open fire and drink fresh, hot yakmilk and eat tsampa, a mixture of ground,roasted barley and warm yak milk. We watchedwith fascination as the family made fresh yakcheese and accepted their gracious offer to shareit with us.The region around Bomi, which we finally

reached on July 30, supports mixed semi-humidbroadleaf forests of deciduous species such as

Two species of gentian, both growmg in thm mountain soils The blue flowers, m photo at mght, are scarcelylarger than one centimeter across

12

A hospitable Tibetan family boiling fresh yak milk mside them tent.

Betula, Alnus, and Sorbus. Conifer forests athigher elevations are dominated by Abies,Tsuga, and Pinus densata. Other familiar taxawe saw included Rosa, Populus, Lonicera,Ribes, Rubus, Pmmula, Gentiana, Rhododen-dron, Cornus, Potentilla, Sambucus, Vibur-num, Berbens, Rhus, Elaeagnus, Quercus,Philadelphus, Clematis, Prunus, and Daphne.One nice surprise was finding Linderaobtusiloba growing wild. (A 100-year-old speci-men of this species, grown from seed collectedin 1892 in Japan by Charles Sargent, standsacross from the lilacs in the Arboretum.) Unfor-tunately, an unwelcoming attitude on the partof local officials forced us to cut short our timein Bomi; nevertheless, we collected about 125taxa in the course of our three days there.

Fruit of a Viburnum growing at the edge of a forestnear Bomi.

On August 4, we retraced our stepstoward Banda and then headed north

again to Changdu, where we remamedfor two days while Wu Sugong againmet with local officials and made plansfor the difficult road ahead. The five-

day, 750-mile (465-km) journey fromChangdu to Lhasa featured several ofthe by-now-all-too-familiar hazards ofbad roads and washed-out bridges.Twice (once in the dark) we were forcedto ford rivers so deep that the waterpoured over the hood of the vehiclesand then rose up through the floor-boards. After one night in Lhasa weheaded southeast, back toward theBomi region, to spend four final daysin the field.

The more than 6,700 specimens (18,883 sheets)collected over the course of this three-year NSFproject will no doubt help to define more pre-cisely this critical biodiversity hotspot. All ofthe collection and locality data, as well as theimages from these trips and others in the region,are linked to a geographic information system(GIS) and are available over the worldwide web/http://maen.huh.harvard.edu:8080/china/.Specimens collected earlier in the region andnow housed in the Harvard University Herbariawill be entered into a database and linked to thewebsite in the nearfuture. By providingtraining and com-

puter equipment forAmerican and Chi-nese students and pro-fessionals, the projectlaid the foundationfor future long-termresearch projects onChina’s biodiversity.In addition, the re-search may havepaved the way forconservation effortsand for detailed analy-ses of biogeographicpatterns and pro-cesses of diversifica-

Frmt on stump sprouts of aSorbus species growmg on

grazed slopes near Rawn

13

---------------------

The author collectmg in a meadow outside Markam.

tion in the region. Conservation International(http://www.conservation.org/xp/CIWEB/ /home), an organization that is currently work-ing closely with Chinese botanists and govern-ment officials in western Sichuan to prioritizeareas for conservation, has sought the expertiseof Dr. Boufford, who has over twenty years offield experience in China.Now, after a year’s work, we have finished

sorting these specimens and dividing them intosets for distribution to over a dozen otherbotanical institutions. The hardships of the tripare long forgotten, and the magic and richness ofthis remote, exotic land once known as Shangri-La beckons again. Who knows what otherbotanical treasures are still to be discovered onthe rooftop of the world?

Notes

1 N. Meyers, 1988, Threatened biotas: ’Hot spots’in tropical forests, Envmonmentahst 8: 187-208;

E O. Wilson, 1992, The Dmersity of Life, HarvardUniversity Press (Cambridge).

2 From D. J. Mabberley, 1987, The Plant Book,Cambridge University Press.

3 P. H. Raven and D. I. Axelrod, 1978, Origin andrelationships of the California flora, University ofCahforma Pubhcatlons m Botany 72.

4 W. T. Wang, S G. Wu, K. Y. Lang, P Q Li, F. T. Pu, andS. K Chen, eds., Vascular Plants of the HengduanMountams, 1993, Vol. 1, Ptendophyta, Gymno-spermae, Dicotyledoneae (Saururaceae to Cornaceae~;1994, Vol. 2, Dicotyledoneae (Diapensiaceae to

Asteraceae) to Monocotyledoneae (Typhaceae toOrchidaceae).

5 S. B. Sutton, 1974, In Chma’s Border Provmces theTurbulent Career of Joseph Rock, Hastings House,(New York).

Susan Kelley is an associate curator and responsible formanagmg and developing the Arnold Arboretum’s digitalmapping system, which tracks the ca 15,000 plants mthe living collections. Her previous experience plant-hunting m Asia was m the central mountams of Taiwan

Anachronistic Fruits andthe Ghosts Who Haunt Them

Connie Barlow

hirteen thousand years

ago, the Age of GreatMammals came crashing

to a close in the Western Hemi-

sphere. Lost were the giants of theelephant clan: the mammoths,mastodons, and gomphotheres,which had maintained a presencein North America for twenty mil-lion years. The native horses, atall camel, and all but one speciesof pronghorn-each group from alineage thought to have originatedin North America-vanished fromthe plains. Gone, too, were thestrange beasts that had evolved inSouth America during millions of years of con-tinental isolation: ground sloths as massive aselephants, hippo-like toxodons, and lumbering,spike-tailed glyptodonts, which looked uncan-nily like the ankylosaurs that had shared theCretaceous landscape with T. rex.These large herbivores (along with the biggest

bear, the biggest canid, and several big cats, allof which depended upon the plant eaters) disap-peared in a geological instant. Evidence is

mounting that newly arrived humans with for-midable stone-tipped spears of Clovis designwere to blame.’ This "extinction of the mas-sive" that marks the end of the Pleistocene

epoch ravaged the megafauna. What happenedto the plants?We cannot be sure. Leaves and flowers and

seeds are not preserved as readily as bone. Plantlineages restricted to upland habitats may comeand go without a trace. Bogs and lakes, ofcourse, receive a shower of pollen, often fromvast distances. But detectable quantities of pol-len preserved in sediments are restricted towind-pollinated plants-notably, grasses, coni-

Reumted: osage orange and mastodon. Shown here is surely the firsttime m thirteen thousand years that the fruit of osage orange,Maclura pomifera, has touched a molar of its missmg partner inevolutlon, Mammut amencanum.

fers, and many deciduous trees of our temperateforests. (And of these, some produce pollen thatcan be distinguished only at the genus or evenfamily level.) Indeed, fossil pollen from strati-fied bog and lake sediments is the primary evi-dence used to reconstruct the vegetational shiftsthat accompanied the repeated coolings andwarmings of the Pleistocene epoch.The Pleistocene pollen record shows only one

tree extinction (a species of spruce) in NorthAmerica near the end of the epoch.z Is there rea-son to suspect that plants pollinated by insects,birds, or bats-that is, plants with little or nopollen record-might have been more vulner-able to extinction?The answer is yes, but vulnerability to extinc-

tion in this case has nothing to do with the modeof pollination. Many plants that are pollinated byanimals rather than by wind produce fleshy fruitswhose seeds are dispersed by other, larger ani-mals. Plants dependent on megafauna for dis-persal would indeed have been vulnerable torange reduction or even outright extinction whentheir partners in evolution and ecology vamshed.

15 s

Riddle of the Rotting FruitEcologist Dan Janzen speculates that althoughfossil evidence is lacking, some fruit-bearingplants probably did follow the megafauna intoextinction. These plants would have begun todecline when their megafaunal seed dispersersvanished. Some might have gone extinct rela-tively soon after losing their partners. Others,especially the long-lived and those that regener-ate clonally from their roots as well as fromseed, still survive but may ultimately be ontrack for extinction. Still others have been

regenerated as domesticated cultivars byhumans fond of their fruit or other botanical

qualities. But by and large Janzen thinks thatthose that are still here today suffered signifi-cant reductions in range when important mem-bers of their disperser guilds were extinguished.The narrowly restricted ranges of some of

today’s big-fruited plants suggest that Janzenmay be right: plant extinctions may, in fact,have stemmed from the animal extinc-tions. An indicator that something isamiss is evident in the case of treeswhose fleshy fruits fall and rot beneaththe canopy of the parent. It was this"riddle of the rotting fruit" that cap-tured Janzen’s attention twenty-fiveyears ago while he was studying theecology of Costa Rican plants. It madeno sense for plants to waste energy bybuilding pulp that attracted few if anydispersers. Worse, in the case of manyfleshy fruits, when the pulp rots, theembedded seeds are killed as well. Whatwas going on here?

In 1982 Dan Janzen, with paleoecolo-gist Paul Martin, published a paper inthe journal Science titled "NeotropicalAnachronisms: The Fruits the

Gomphotheres Ate."’ Janzen had car-ried out field studies in Costa Rica tolearn whether introduced livestock(horses and cattle) served as surrogatePleistocene megafauna for the bereftplants. The result: a list of some thirtyspecies of trees and vines of the CostaRican dry forest whose fruits bear thephysical and ecological characteristicsof "ecological anachronisms."’ These

plants are living in a time warp; they are adaptedfor a lost world. Their missing animal partnersare "the ghosts of evolution."Anachronisms and ghosts caught the early

attention of Robert E. Cook, who is the directorof the Arnold Arboretum. Citing Janzen andMartin, Cook published an essay in 1982 inNatural History that described the avocado,Persea americana, as an ecological anachromsmthat has been stunningly successful in attract-ing a replacement dispersal agent: us.~ Withinthe past few hundred years, avocado has beentaken from the New World tropics to orchardsin Florida, California, northern Mexico, and farbeyond this hemisphere. Wild elephants whoraid village fruit trees in Africa are now "plant-ing" American avocado on that continent.’

In their landmark paper, Dan Janzen and PaulMartin concentrated on Costa Rican plants. Butin the final paragraph they extrapolated theanachronism concept to large-fruited plants of

Strategies of the megafaunal dispersal syndrome. Domesticatedvaneties of three tropical fruits native to the New Worlddemonstrate a range of pulp attractions and seed defenses Ripepapaya frmt, Canca papaya, is soft enough to mash rather thanchew, so the tmy seeds require no physical protection Amammal that madvertently crushes a papaya seed is, however,deterred by a sharp, peppery flavor-and thus the toxms sosignaled. Pmmate fruit thieves (hke us) can eat around anddiscard the concentration of seeds, thus foiling the papaya’s ’s

mtent Avocado, Persea americana, produces a shppery anddense seed, whose potent toxms taste bitter to mammals Incontrast, the seeds of the camstel tree, Poutena campechmana,have a mild flavor and are protected mstead by a tough coatmg.All but the biggest frugmores could be expected to eat aroundor spit the seeds of avocado and camstel

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Honey locust and her new partner m evolution. Strung alongthe top are the usual number of seeds m a honey locust pod,such as the one restmg on the author’s arm.

The most anachromstic legume m North Amemca. The mpe podof kentucky coffee tree is toughened byresms, and the seeds aremvulnerable to msect attack The green pulp is sweet butreputedly poisonous to humans Early colomsts m Kentuckyroasted and ground the seeds to make a coffeehke brew. Theground "cof fee" shown here is courtesy of Carl Mehling.

the eastern and central United States: kentuckycoffee tree (Gymnocladus dioicusJ, honeylocust (Gleditsia triacanthos), pawpaw (Asiminatriloba), persimmon (Diospyros virginiana), andosage orange (Maclura pomi fera). ).To a plant lover, the notion that ghosts may

be haunting some of the most magnificentnative fruits of one’s homeland is a revelation. Itwas quite a surprise, therefore, when I began thelibrary research four years ago for my book TheGhosts of Evolution (2001) to discover that vir-tually no effort had been made to test these fivetemperate plants or to identify other possible

anachronisms in the forests and fieldsof North America. Indeed, scientistscurrently breeding honey locust trees toincrease the fodder value of their pods,and those developing improved pawpawcultivars and promoting the use of theirfruit, have been doing so unaware thatthe fruits are anachronistic.

Osage Orange-An ExtremeAnachronism

Recently, I spoke with an archeologist,Frank Schambach, who felt frustratedbecause nowhere in the published lit-erature could he find the informationhe needed to solve the ecological puzzleof osage orange-information crucialfor validating his thesis that before set-tlers began rearranging the landscape,the wood of this tree, highly valued formaking hunting bows, may have beentraded far and wide in North Americaunder the complete control of a singleindigenous tribe.8 Schambach suspectedthat osage orange (also known as boisd’arc, "wood of the bow") occupied avery constricted range that could in factbe claimed by a single tribe.Osage orange would more appropri-

ately have been named osage breadfruit.A close relative to Maclura, of America,and to its sister genus Cudrama, ofeastern Asia, is the breadfruit genus,Artocarpus. All are linked through themulberry family, a largely tropicalfamily dominated by the figs andstriking for its members’ compound

fruits and the white latex some exude whencut. Osage orange was named after the OsageIndians of Missouri, who first introduced whitetraders to this strange fruit-the color of a glow-green tennis ball and about the size and firm-ness of a softball.

Inhabitants of the plains and prairie statesknow this fruit by another name: hedge apple.Until the invention of barbed wire in 1874,there was no more effective or economical wayto fence free-ranging livestock out of one’s veg-etable garden or cornfield than to plant a hedgeof thorny osage orange stems, later interweav-

17 7

An elephant fruit m a land without elephants The bnght greenfruit of osage orange was shaped by the now extinct megafaunaof North Amenca.

ing the abundant root suckers that the treessend up in response to severe pruning. Thus

osage orange gained a replacement dispersalagent, and its range expanded rapidly.Maclura pomlfera, a wind-pollinated tree, is

known from pollen samples to have been widerangmg in North America during earlier ice-freephases of Cenozoic history. But by the time ofEuropean contact, its range had shrunk consid-erably : just before its transformation into a com-mon hedge plant, it inhabited only a smallstretch of the Red River watershed near the

junction of Texas, Oklahoma, and Arkansas.Frank Schambach suspects that at its nadir therange was even more restricted-possiblylimited to the Bois d’Arc tributary of the RedRiver in Texas and nearby creeks of the adjacentBlackland Prairie. Such isolation would explamhow a single tribe-the Spiroans of Mississipianculture-could have controlled the entire bowwood trade. And this, in turn, would explain thearcheological evidence of extraordinary wealthaccumulated by this people in the centuriesprior to European contact.What ecological information did Schambach

need to solve his archeological puzzle? Hewanted to know whether horses spread osageorange-that is, do horses eat the fruit anddefecate viable seeds? If so, then by the timenaturalists got around to documentmg the geo-graphic reach of osage orange, its renaissancehad already been initiated by horses, which had

been reintroduced into North Americain the sixteenth century. Surely ama-teur naturalists and people living wher-ever osage orange and ranch horsesco-exist know the answer to the ques-tion-it seems to be yes. But the matterisn’t discussed in the published litera-ture. Thus a core contention in

Schambach’s "Spiroan trader" theoryrests on what he has been able to gleanfrom Red River ranchers and his owncasual observations.The search continues. A year after

publication of his "Spiroan trader"theory, Schambach obtained crucialanecdotal information: "A volunteer at

my dig this summer lives on a ranch onthe Blackland Prairie in east Texas,"

Schambach wrote me. "She has a small herd ofhorses which, she assures me, routinely eatosage orange fruits on their own. Furthermore,she knows for a fact that horses spread the treevia their manure because when she and her hus-band acquired their property there was no osageorange growing on the upland (prairie) parts ofit, only in the bottomlands along the creeks. Butsoon after they began pasturing horses on theland, the osage orange began to migrate out ofthe bottoms, and it is now growing all over theirprairie areas, to their dismay." "

Here’s another intriguing story that came myway while I was writing this article-this onefrom Robert M. Timm, professor of ecology andevolutionary biology at the Umversity of Kan-sas. Timm has horses and one mule on his farmin Kansas. They all "love the fruits" of osageorange, he wrote me. "The mule is the best at

locating them, but if you are experienced withmules that wouldn’t be a surprise. They arealways much more curious and more explor-atory than horses. I’d say of all the natural foodsaround here, osage orange fruits are the mule’sfavorite. He seems to remember them from one

year to the next, but that too is typical of mules.I have no doubt that Pleistocene horses wouldhave long-term memory of favorite trees to feedat every fall."

II

Timm is very familiar with Dan Janzen’sanachronism theory and has been casuallyobserving mice, rabbits, and tree squirrels feed-

18 8

ing on the pulp and seeds. "They don’t cacheosage orange," he observes, "they eat it on thespot." Most intriguing is his discovery of largequantities of shredded, freshly fallen fruit in thestomachs of deer. All these mammals great andsmall are indeed eating the fruit, but are theydispersing the seed? Timm concludes no.Timm acquires buckets of osage orange fruits

from neighbors and spreads them on the

unpastured sections of his own property assupplemental feed for wildlife. Nevertheless, hehas encountered no seedling trees. "I picked upanother five gallon bucket of fruits this morn-ing," Timm wrote me as this article went topress, "and I’ll check here in a few weeks tosee if the seeds make it intact through a mule’sdigestive tract. I’ll pull the mule and horsesoff the pasture later this month and keep themin a paddock for the winter and give the mulefruits where it will be easy for me to retrievethe seeds."

Overall, anachronism theory seems to beanecdotally well supported for horses as dis-persal agents of osage orange. Are there anyother plausible ghosts to pair with this nativefruit besides Pleistocene horses?

Paul Martin and I had a chance to test osageorange vicariously on a much bigger Pleistocenesurrogate: African elephants at the BrookfieldZoo in Chicago. Almost surely this was the firsttime since the Pleistocene that the fruit of

An extreme anachromsm. Osage orange is one of NorthAmerica’s most anachromstic frmts. Freshly sliced fruit oozesa white latex, which has been wiped clean from the slice onthe left. Honey locust seed (1 cm) for scale

Maclura pomifera had met the molar of a pro-boscidean. I sent a box of freshly fallen fruit toMartin, which he in turn forwarded to theBrookfield Zoo. After a search of the literature,zoo staff decided that there was no danger inoffering the elephants a few fruits.The results were inconclusive. At first, the

youngest two elephants didn’t want to eventouch the fruits offered by their keepers. Finally,each curled a trunk around a sphere and hurledthe offensive object out of their habitat. Thematriarch, however, chose to sample the firstfruit offered, chewing and swallowing. But shewould then accept no more.

Herbivores are known to be wary of novelfoods. Cultural knowledge of gastronomic pos-sibilities, passed from one generation to thenext in social animals, would not be available tozoo elephants. Then too, well-fed captive am-mals will often turn up their noses at foods thattheir wild counterparts would happily consume.The case is therefore still to be made that Pleis-tocene mastodons and mammoths would have

joined horses in dispersing osage orange fruits inNorth America. But it will be a most interestinghypothesis to pursue-for anyone excited by thetheory and who has access to elephants!

Powers of Persistence

It is perhaps no coincidence that the five speciesof temperate American trees judged anachronis-

tic by Dan Janzen and Paul Martinare all prodigious cloners. Vegetativemeans of reproduction would havehelped these trees persist for the thir-teen thousand years that sexual repro-duction has been disrupted for want ofadequate seed dispersal.Kentucky coffee tree, honey locust,

pawpaw, persimmon, and osage orangeall send up root suckers-prolifically sowhen the main stem is pruned or other-wise damaged. Kentucky coffee andpawpaw are extraordinarily skilled ingrowing lateral root runners that sproutnew stems many meters from the elder

stem, supported by photosynthates sup-plied by the parent. Indeed, a few yearsago when a pawpaw tree "died" of old

age in the Arnold Arboretum, more

19

than a hundred fresh stems popped up almostimmediately from a vast network of root run-ners. And although the Arboretum isolates itsmature kentucky coffee specimens by encir-cling the trees with a wide buffer of mowedlawn, grounds staff must periodically rid neigh-boring beds of the vigorous fresh stems emanat-mg from hidden root runners.As the ice retreated from its last southward

advance, which peaked about twenty thousandyears ago, four of the five above-mentionedanachronistic trees of eastern and central NorthAmerica would have been helped to reclaimformer territory by newly arriving humans.Pawpaw and persimmon fruits would have beencarried back to camp, their seeds removed or

spit out at the time of eating. Honey locust podswould have been opened and licked for theirsweet matrix, the hard seeds discarded. Ken-tucky coffee trees were valued not for their fruitbut for their large, nearly spherical seeds, whichtook a lovely polish and were used for gamingtokens. Although the wood of osage orange washighly prized and known to have been tradedacross great distances in the time just prior toEuropean contact, the fruits held little if anyvalue. Does this perhaps explain why the rangeof osage orange became so constricted?

Is the Endangered Torreya TreeAnachronistic?

Coming to terms with the likelihood that nativehorses almost certainly and elephants probablywere effective seed dispersers of osage orangeduring the Pleistocene and for several tens ofmillions of years before that provides freshinsights into how to rescue from extinction aseverely endangered American tree, Torreyataxifolia (florida torreya). Torreya is a conifer ofthe plum yew family, Cephalotaxaceae, andthus bears no fruit as botanically defined.Nevertheless, the fleshy design of its diaspore-ecologically a fruit-is an obvious lure for am-mals. Like a yew or a gmkgo, torreya producessingle large seeds enveloped in what ought to beviewed as fruits by vertebrate dispersers.The proximate cause of Torreya taxifoha’s

imminent extinction, and thus the cause thatgets all the attention, is disease. Some thirtypathogens are known to infest it, but no single

disease seems to be the culprit.9 Once commonin the rich soils of the Apalachicola River ofnorthern Florida, adult specimens growing inthe wild suddenly began to die in the 1950s, andnone remam today. Like the American chestnutthat was destroyed by (an imported) blight dur-ing the early years of the twentieth century,torreya survives only because new stems keepsprouting from the same rootstock. Sadly, eachfresh sprout of torreya is doomed to die before itis old enough to produce pollen or ova. Energystored in the roots will eventually give out,since new starts in the shady forest may con-sume more photosynthate than they can returnto the roots before their demise.The genus Torreya was once distributed

throughout the Northern Hemisphere. Rangefragmentation has created distinct species ineastern China, patches of the Coast Range andthe Sierras of California, and the Apalachicolaof Florida. The geographic range of the Floridaspecies is today restncted to the cool ravinesalong the east side-only the east side-of a 22-mile (35-km) stretch of the Apalachicola Riverin northern Florida. During the coldest timesof the Pleistocene, the Apalachicola, with itsmoderate climate and rich soils, was a refuge forthe trees and forbs that now enrich the CoveHardwood forest of Great Smoky MountainsNational Park, 375 miles (600 km) to the north.After the ice retreated, most of the plantshitched rides from wind and animals and movedback north to their pre-glacial home. Torreyaseems to have been left behind.Some experts confirm that the tree’s troubles

may have begun for want of a disperser. 10 Glo-bal or regional extinction of an animal partner(or partners) may be the root cause of the tree’scurrent distress. Torreya is probably not ideallysuited for the warmth and humidity of today’sApalachicola region. It wants to head north, butit hasn’t found a vehicle.That florida torreya may be haunted by the

ghosts of extinct dispersers is suggested by ahost of clues. First, the diaspore of all species ofTorreya is distasteful or toxic to many (possiblyall) mammals who normally consume fruits.The pulp has a high terpene content and itleaves a sticky residue on one’s skin. Squirrelstreat the fruit as they treat gmkgo fruit m New

20

A florida torreya photographed well north of its "native"grounds, near Philadelphia, Pennsylvama, at the Henry BotamcGarden, near Gladwyne.

York City’s parks: they discard the flesh andsteal the seeds. Squirrels that fed on torreyaseeds on the east side of the river would beunable to carry them across water to the westside, and if the rich soils of the Apalachicola areisolated from rich soils to the north by a barrierof sandy soils, then the squirrels would also beunable to disperse the seeds farther north. Squir-rels may thus be a disperser, but they apparentlyare not the right disperser for helping this treereclaim its pre-glacial range. This explanationwould account for the seemingly paradoxicalfact that until the 1950s, florida torreya was theseventh most abundant tree species in an aston-ishingly small patch of "native" habitat.

Perhaps the best evidence that floridatorreya may be suffering from an mabil-ity to track climate change is thatbefore the blight took hold, this treewas planted hundreds of miles north ofits Florida habitat in the mountains ofNorth Carolina, near Asheville. There,on the Biltmore Estate, the torreyas arethriving, and the females produce abun-dant seeds. "Flower beds often aboundwith seedlings ’planted’ by squirrels,"reports Bill Alexander, landscape histo-rian at the Biltmore. During his 23 yearsthere, Alexander has watched the

torreyas stand up well to a five-yeardrought. And in the winter of 1985 thethermometer plunged to minus-20

degrees Fahrenheit, yet "our trees

smiled right through," he told me.For a number of years, Alexander had

been thinking that "florida" torreyareally belonged back in North Carolina.So he was delighted to hear of the lost-disperser theory. A megafaunal ghost?If so, the ghost may well be a largeextinct tortoise, I suggested, as reptilesare far more tolerant of plant terpenesthan are mammals, and as the thin"shell" protecting the large single seedof this conifer offers scant protectionagainst molars.One must not, however, ponder the

plight of the florida torreya in isolationfrom its sister species. In contrast toTorreya taxifolia, California’s torreya

(Torreya californica) is maintaining its popula-tion, as are the several Asian species of

Torreya-all of which bear nearly identicalpropagules. Nevertheless, all occupy restrictedgeographic ranges. What if the entire genus lostits key dispersers and now depends on the localactivities of squirrels?

Bill Alexander and I easily came up with twoplausible explanations for the differences inendangerment, based strictly on geographicdifferences. In eastern North America, the cli-matic effects of the Ice Sheet reached muchfarther south than was the case in either west-ern North America or eastern Asia, forcing theAppalachian species to take refuge at a lower

21

latitude. Perhaps even more significant isthat latitudinal migration was the only optionfor florida torreya as the climate warmed. In

contrast, torreya species in California andin Asia could head upslope. These torreyas arenative to mountainous regions, where altitudi-nal gain facilitated by nothing more than squir-rels could help the trees keep pace with awarming climate.Such unsubstantiated and untested leaps of

speculation are normally not well receivedwithin the scientific commumty-but these arenot normal times. Without some drastic break-

through in the management of Florida’s wildpopulation of torreya trees, Torreya taxlfoliawill, within fifty years, almost surely be extinctoutside of botanical gardens. Perhaps it is timeto help this torreya gain rootholds of wild popu-lations in the mountams of North Carolina.Such is not, alas, how things are done with

endangered species-the exception being therecent return of the california condor to itsPleistocene home near the Grand Canyon.Native territory is regarded as the last best placeto be. But what is "native"? How far might wejustifiably reach back in time for a benchmark?

In a study of endangered species published in2000, Rob Channell and Mark Lomolino con-cluded that "most species examined persist inthe periphery of their historical geographicranges."" If habitat at the periphery of histori-cal range is adequate but not ideal, then the lastplace a troubled species is found may not, infact, be the best place to assist its recovery.

Transplantation across great distances is anuncommon and controversial technique forbiodiversity conservation today. But as thegreenhouse effect ratchets up temperaturesand reroutes ramfall, and as botanical preservesbecome even more isolated islands in a seaof human development, long-distance trans-plantation will become the norm. If gardeninga few local patches of endangered plants is

tough today, it’s gomg to get a lot tougher when,like it or not, we become gardeners of theplanet.l2 Helping plants track climate changefrom one patch of habitat to another will bea routine tactic for conserving biodiversitydecades hence. Is it too early to begin now withflorida torreya?

Notes

1 In the June 8, 2001, issue of Science, two papersconfirmed the "overkill hypothesis" of end-Pleistocene extinctions, one for Australia and theother for North America. The North American paperis John Alroy, 2001, "A multispecies overkillsimulation of the end-Pleistocene megafaunal massextinction," 292 1893-1896. A news report by LeighDayton pubhshed in the same issue (p. 1819), "Massextinctions pmned on ice age hunters," suggests thatthis view is now the majority position.

2 Stephen T. Jackson and Chengyu Weng, 1999, "LateQuaternary extinction of a tree species in easternNorth America," Proceedings of the National

Academy of Scsence 96. 13847-14852.3 Dan Janzen’s suggestion that fleshy-fruited plantsmay have gone extinct smce the end of thePleistocene is m Connie Barlow, 2001, The Ghosts ofEvolution (New York: Basic Books~, 88

4 D H. Janzen and P. S Martin, 1982, "Neotropicalanachromsms: The fruits the gomphotheres ate,"Science 215: 19-27.

5 For the story of how Janzen and Martin developedtheir ideas and conducted their fieldwork, see chapters1-3 of the author’s Ghosts of Evoluuon, op cit.

6 Robert E. Cook, 1982, "Attractions of the flesh,"Natural History 91 / 1 20-24.

~ Martrn N. Tchamba and Prosper M. Seme, 1993,"Diet and feedmg behavior of the forest elephant mthe Santchou Reserve, Cameroon," Afncan Journal ofEcology 31: 165-171.

8 The theory that osage orange, prior to Europeancolonization, occupied a very restricted range ispresented in Frank F. Schambach, 2000, "Spiroantraders, the Sanders Site, and the Plams InteractionSphere," Plains Anthropologist 45 7-33.

9 Mark W Schwartz, Sharon M. Hermann, and Philip J.Van Mantgem, 1999, "Population persistence inFlorida Torreya," Conservation Biology 14. 1023-1033.

lo See p. 229 of The Ghosts of Evolution.l Rob Channell and Mark V. Lomolino, 2000,"Dynamic biogeography and conservation ofendangered species," Nature 403: 84-86

’z D. H. Janzen, 1998, "Gardemfication of wrldlandnature and the human footprint," Science 279: 1312-1313.

AcknowledgmentsI wish to thank Dan Janzen, Paul Martm, Bill Alexander,Robert Timm, and Frank Schambach for helpfulcritiques, comments, and anecdotes.

Connie Barlow is a science writer and author of TheGhosts of Evolution Nonsensical Frmts, MissingPartners, and Other Ecological Anachromsms,published in 2001 by Basic Books.

Pastures of Plenty: A Case Study in Field BiologyP. L. Marks

ow pastures are taken for granted int rural parts of the northeastern UnitedStates, where they are common. Wedon’t think of them as special habitats, yet theyare. Cow pastures have been present in thenortheastern landscape since the earliest farmfamilies first carved agricultural lands from theforests. The best land in terms of soil fertilityand drainage generally became arable fields,whereas more marginal lands went into pas-tures. Other marginal lands, often wetter,steeper, or more remote from the main property,were left as forest. But historically, every dairyfarm had at least one pasture.

Field biologists work to discover patterns innature-usually by observation-and then tounderstand the causal influences behind those

patterns. More often than not, the explanationsfor patterns in nature are complex, a result ofmultiple factors interacting in various waysover time. Often it is not possible by observa-tion alone to fully understand the cause of agiven pattern.Cow pastures are interesting from a scientific

point of view because one factor-the presenceof many large, hungry cows-is of overridingimportance in determining the abundance andkinds of plants that occur. Whether a pasturebegan as a natural meadow or as planted forage,the grazing and trampling activities of cows, car-ried out over years or decades, produce a habitatof closely cropped edible plants interspersedwith taller plants that are avoided by the cowsdue to their physical or chemical properties.

Pastures can be surprisingly beautiful, per-haps because they take on some of the featuresof landscaped yards and public gardens: largeareas of mowed green grass with scattereddrifts of taller herbs and small flowering trees-sometimes dramatically pruned into unusualshapes. How does this happen?

In active pastures, the most conspicuousplants are those not eaten by cows. A strikingfeature of many of these plants is the presence

of thorns or their equivalent. Thistles (Circiumspp.) are common in many pastures. And amongwoody plants that tend to catch our attentionvisually, wild or volunteer (not planted) appletrees (Malus spp.) and hawthorns (Crateagusspp.) stand out. Apple trees do not have truethorns, but their short side branches are sharp atthe tips and function as thorns. These distinc-tive thorny woody plants allow many pasturesto be recognized as such as we drive or walkalong country roads. Pastures with hawthornsare especially easy to recognize at a distancebecause the hawthorn bark is light gray and thebranches are horizontal in orientation, produc-ing a distinctive growth form. Because theypersist for many decades after a pasture is aban-doned, the apples and hawthorns also provide adiagnostic legacy of former land use.We now have a pattern to explain. Why are

thorny woody plants so common in cow pas-tures ? An obvious and important first thought isthat the thorns prevent or reduce cows’ feedingon these plants. This is consistent with otherrelated observations. One is that a variety ofnon-thorny woody plants invade abandoned pas-tures, but not pastures where cows are still

present. Another observation is that the applesand hawthorns that commonly invade pasturesdo so when the pastures are active. However,other thorny plants, such as multiflora rose(Rosa multiflora), invade a variety of open habi-tats, not just active pastures.The implication is that pasture cows are a

dominant influence. In their presence thornyplants hold an advantage, whereas on the sameland without cows non-thorny plants have theedge. Presumably, plants without thorns cannotinvade active pastures because cows eat them. In

pastures abandoned for decades one can some-times see both thorny and non-thorny woodyplants. There are dense patches of thorny scrubwhere plants that started when the pasture wasactive have grown larger and taller; here fewother woody plants have been able to invade.

23

An active cow pasture in spring, showing cows, closely cropped grass, and a mixture of apple and hawthorn trees. The appletrees range from closely pruned cones to larger trees In flower.

Other areas, where no apples or hawthorns werepresent at the time of abandonment, now havea variety of non-thorny trees and shrubs suchas white ash (Fraxinus amencana) and gray dog-wood (Cornus racemosa).A careful accounting of plant ages should

reveal differences between the groups: the

thorny plants that invaded when the pasturewas active should be ten or more years olderthan the non-thorny trees that invaded after itsabandonment. But inferring plant ages fromplant sizes can be misleading. Decades afterpasture abandonment, ash trees, which growfaster than apples, will be taller but younger.Although thorns reduce the feeding pressure

from cows, they do not prevent feeding alto-

gether. Indeed, the closely pruned hawthorn andapple plants are perhaps the most spectacularaspects of an active pasture. From a distance, itis difficult to believe these are wild plants, soexquisitely shaped are their crowns. Many looklike inverted bowls or cones, whereas otherslook like old-fashioned hourglass timers. Closerinspection reveals that these geometrical shapesare made up of a proliferation of branchescaused by the repeated release of side branches,which, in turn, is caused by cows eatmg thebranch tips year after year. The resulting protec-tive matrix of short, interwoven branches is sodense and rigid that cows can feed only on theplant bits that protrude from it. Conversely,leaves that are near the exterior of the matrix

24

-- ~..._._-_.__._._~....~.,-~-- ,

A typically cone-shaped, closely pruned apple tree, photographed inwinter, resides m an active cow pasture. The glove m the foregroundpromdes scale.

~1iJ¡ ~~~~_~ . .

The top pctrt or thrs apple tree, now beyonci the reach of hungrycows, shows a spurt of new growth, the tree takes on a characteristichourglass shape

but rccessed from the edge are protected fromcows while still exposed to sunlight. The denselycropped apples and hawthorns and the cowsachieve a kind of equilibrium, with the cowsunable to penetrate the "shell" and the plantsunable to grow beyond it. One of my studentsproposed the term "bovine bonsai" to describethis lovely piece of pasture horticulture.The balance between the growth of plants and

the cropping by cows sometimes tips in favor ofthe plants-though just barely. Some thornywoody plants in active pastures expand the vol-ume of their crowns slightly each year, despite

the bovine pruners. Much of thisgrowth occurs in the lower half of thecrown so that after twenty or thirtyyears of modest annual gains plantsattain an inverted cone shape. Eventu-ally, this slow lateral expansion reachesa point where cows can no longer reachthe tops. This marks an important tran-sition in the lives of these plants. Afterdecades of tortuously slow growth,plants whose tops are beyond the reachof cows suddenly escape much of thebovine influence. Rapid shoot growthbegins atop these plants and, in subse-quent years, lateral growth increasesonce it is above the reach of the cows.Eventually a bizarre new growth formis achieved: the shape approximatesan hourglass, consisting of a lowerinverted cone whose closely croppedouter surface is still maintained by thecows, an upper upright cone that hasrapid growth on the inside where thecows cannot reach, and a smoothedlower outer surface where protrudingtwigs are removed by the cows.

I have walked through a number ofpastures, making observations and try-mg to understand more about the issuesdiscussed above. In active pastures in

early spring I have seen seeds from non-thorny trees like white ash scatteredwithin the close cropped turf. Yet seed-lings of the same non-thorny species arenotably absent, suggestmg that thebottleneck for the establishment ofthese plants is at the seedling stage. Incontrast, seedlings of apples and haw-

thorns must at least occasionally become estab-lished in pastures in order to account for thelarger plants that are so conspicuous later on.Interestmgly, hawthorn and apple seedlings lackthorns. Why then can these seedlings survive inactive pastures while other non-thorny seed-lings cannot? Why doesn’t eating and tramplingby cows eliminate apple and hawthorn seed-lings, as it apparently does the seedlings of non-thorny species? We don’t know all the answers,but our understanding of how young plants getstarted in pastures is more complete for applesthan for hawthorns.

, 25

Many pastures have apple trees around theedges or a few large, old apple trees within thepasture proper. In autumn, the fruits ripen andcows eat the apples that fall to the ground. Theseeds pass through the cow unharmed and aredeposited in "cow pies," where they spend thewinter. Come spring, these seeds germinate intoseedlings that grow directly from the cow pies.The rotting manure provides a locally fertileenvironment for apple seedlings in cow pas-tures. In addition, cows typically do not feed onplants growing in or near cow pies. This reprievefrom feeding damage, which may last throughthe first growing season, allows apple seedlingsto grow and create energy reserves so that theyare more likely to recover the first time they areeaten or trampled upon by cows.

Since not all pastures have appletrees, and since only those cow piesproduced when apples are ripe containseeds, most cow pies do not have appleseedlings. But examining many cowpies in the spring should yield somewith apple seedlings, which are recog-nizable by their two large rounded seedleaves, or cotyledons. One study con-ducted by Cornell thirty years agofound as many as 250 apple seedlings ina single cow pie!

I once harvested a closely croppedapple tree about five feet 1.5 ml tall inorder to estimate its age by counting theannual growth rings on a cross sectionof the trunk. (Gaining access was noteasy-I had to saw a hole in the side of thedense, rigid, sharply tipped branch matrix beforeI could reach the trunk.) The tree was aboutthirty years old, which was consistent with theages reported in the Cornell study. However,even more interesting was the new dimensionof pasture ecology it revealed: inside the protec-tive branch shell of this apple tree grew severalnon-thorny woody plants.

I have since found a number of non-thornywoody species, including white ash, honey-suckle (Lonicera spp.), and chokecherry (PrunusvmgmianaJ, growing inside the protective shellof several apple and hawthorn trees in pastures.These species of trees and shrubs are among thesame species that invade abandoned arablefields and pasture. In active pastures, species

intolerant of cattle grazing can generally growonly within the refuge provided by prunedapples and hawthorns.

This essay focuses on cow pastures in centralNew York State, but the issues raised hereapply, to some extent, to cow pastures aroundthe world. Elsewhere there are species equiva-lent to our hawthorns and apples, such as juni-pers. There are other kinds of pastures as well,and what is true of cow pastures is not necessar-

ily true of sheep or horse pastures. Both thenumber of animals and the continuity of the

grazing can influence the numbers and kinds ofplants that are able to invade. Finally, the func-tion of cow pastures seems to be changing inthe Northeast as dairy farming becomes more

Apple seedlmbs emerynd m sprmg from cow manure depositedthe preceding fall.

intensive. Historically, pastures provided foragefor dairy cows during the warm parts of the year.On today’s large dairy farms, cows may not bepastured at all, and since many dairy cows nowreceive all their food indoors, cows that are pas-tured may not crop apples and hawthorns asclosely as they did in the past. Even so, evidenceof former pastures still remains in the largeplants left behind.

Peter L. Marks is a professor m the department ofecology and evolutionary biology at Cornell University.

This article is number six in the author’s series

"Reading the Landscape" (another, on primary vssecondary forests, appeared in Arnoldia Vol 55, No. 3.)It is reprinted from Cornell Plantauons Magazme,Summer 2001, courtesy of Cornell Plantations,607.255.3020 or www.plantations.cornell.edu.

HENRY DAVID THOREAU ANDTHE YANKEE ELM

Thomas j. Campanella

HE American elm (Ulmus americana) has always occupied a hallowed place inTthe pantheon of our native trees, and nowhere was its presence greater than inNew England. Planted by the thousands on streets and town commons throughout theregion, the elm was a defining symbol of Yankee life. Charles Joseph Latrobe, a Britonwho toured New England m the 1830s, summed it up best when he chmstened theYankee elm "the glory of New England."’ Indeed, elms moved pens. Paeans to the treeand its beauty abound in the literature of nineteenth-century New England.Hawthorne, Lowell, Longfellow, Holmes, and Henry James all penned eloquent trib-utes to the tree. So did literary visitors such as Trollope and Dickens.

But no wnter who lifted pen to elm did so with more grace and insight than thebearded scnbe of Walden Pond. Henry David Thoreau was the poet laureate of the Yan-kee elm. A prolific writer, Thoreau’s pen roved far beyond his celebrated sojourn atWalden. His journals record nearly twenty-five years of observation and bristle withexacting detail. These books map the geography of Thoreau’s intellect, a terram welltimbered with elms.To Thoreau, elms consecrated the landscape. Mircea Eliade has written m The

Sacred and the Profane that we "found" the world by investmg it with religiousmeaning, and fix sacred space with signs and totems.z In the traditional New Englandlandscape, writes John Stilgoe, meetinghouse spires and church steeples functioned asarchitectural exclamation points that "made the cellular countryside intelligible toresident and traveler ahlce."3 Thoreau’s leafy steeples were rougher hewn, but they toolinked earth and soul. For in Thoreau’s church of nature, elms were the beacons of

moral and spatial order.If they rose more leisurely to the sky, once there the elms challenged the lofty white

spires themselves. "Some are so lifted up m the horizon," he wrote of elm crowns spiedfrom afar, "that they seem like portions of the earth detached and floating off by them-selves into space." Below, Thoreau imagined harmony and spatial order, for where elmsbrushed the sky, below lay a town or home. "When I see their magnificent domes,miles away m the homzon, over intervening valleys and forests," wrote Thoreau, "theysuggest a village, a community, there." Spying the "dark little dome" of a far-off elm,Thoreau was reminded of the "rural and domestic life passing beneath it."

"

Homestead telegraphs to homestead through these distant elms seen from hill-tops. I fancy I hear the house-dog’s bark and lowmg of the cows asking admit-tance to their yard beneath it. The tea-table is spread; the master and mistressand the hired men now have just sat down m their shirtsleeves.

In summer the great canopies cast shelter over house and town, harboring a chorusof insects whose gentle hum mduced in Thoreau "contemplation and philosophicthoughts." "

The "Washington" elm (a misnomer; see Sheila Connor, New England Natives, page 111) onCambridge Common, Massachusetts, and the "Whitfield" elm on the right, photographed in1860. From the Archives of the Arnold Arboretum.

28

But Thoreau’s tree was no mere totem of pastoral retreat from the "great world." Forhim, elms objectified a range of human values. He saw m the elm stoicism, persever-ance m the face of adversity. Elms "adjourn not mght nor day," he wrote, "they standfor magnificence; they take the brunt of the tempest; they attract the lightning thatwould smite our roofs, leaving only a few rotten members scattered over the highway." "

Thoreau was hardly a misanthrope-he attended dinner parties even during his Waldenso~ourn-but he did favor the company of trees.When comparing elms with men, however, Thoreau could not help but see the

former as more worthy. "I have seen many a collection of stately elms," he confidedto his Journal in January 1856, "which better deserved to be represented at theGeneral Court than the mamkins beneath." Elms may have been set out by villagers,but they towered-literally and figuratively-above mere mortals. "I find that mtomy idea of the village," he wrote, "has entered more of the elm than of the humanbeing." Indeed, the elms were "worth many a political borough," and certainly morethan most politicians.

The poor human representative of his party sent out from beneath their shadewill not suggest a tithe of the dignity, the true nobleness and comprehensive-ness of view, the sturdiness and mdependence, and the serene beneficence that[the elms] do. They look from township to township. A fragment of their barkis worth the backs of all the politicians m the umon.

Thoreau then extended his metaphor far beyond the environs of Concord. Elmsbecame a medmm through which he channeled his outrage over slavery, particularlyover the Fugitive Slave Law, which authorized federal agents to return to their ownersslaves who had escaped to north of the Mason-Dixon line, and the Kansas-NebraskaAct, which cleared the way for extending slavery into territories not yet granted state-hood. Passage m 1854 of the Kansas-Nebraska Act prompted outbreaks of violenceacross Kansas and drew an impassioned response from Thoreau-"Slavery in Massa-chusetts"-which he delivered to a convention of abolitionists.4 .4

In Thoreau’s wntings, the Concord elms are a metaphor for the abolition movement,particularly for the Free Soil Party, which had been formed m response to the indeci-siveness of the Whigs and Democrats on the expansion of slavery. In a remarkable pas-sage, Thoreau implores the freedom-lovers to remain true to their principles and toseek equilibrium between old and new, past and progress, conservative and radical. Theelms, writes Thoreau, "are free-soilers in their own broad sense." "

They send their roots north and south and east and west mto many aconservative’s Kansas and Carolina, who does not suspect such undergroundrailroads-they improve the subsoil he has never disturbed-and many timestheir length, if the support of their principles requires it. They battle with thetempests of a century. See what scars they bear, what limbs they lost before wewere born! Yet they never adjourn; they steadily vote for their principles, andsend their roots further and wider from the same centre. They die at their posts,and they leave a tough butt for the choppers to exercise themselves about, anda stump which serves for their monument.

Writing of the relationship between the living sapwood (alburnum~ and the deadheartwood (duramen), Thoreau contmues to develop his metaphor. The elms "combinea true radicalism with a true conservatism." Yet,

29

Their radicalism is not cutting away of roots, but an infinite multiplication andextension of them under all surrounding institutions. They take a firmer holdon the earth that they may nse higher mto the heavens. Their conservativeheartwood, m which no sap longer flows, does not impoverish their growth, butis a firm column to support it; and when their expandmg trunks no longerrequire it, it utterly decays. Their conservatism is a dead but solid heart-wood,which is the pivot and firm column of support to all this growth, appropriatmgnothmg to itself, but forever by its support assistmg to extend the area of theirradicalism. Half a century after they are dead at the core, they are preserved byradical reforms. They do not, like men, from radicals turn conservative. Theirconservative part dies out first; their radical and growmg part survives.

In the end the elms (and Free Sellers) will "acquire new States and Territories, whilethe old dominions"-the slaveholdmg South-"decay, and become the habitation ofbears and owls and coons."On most days, however, the elms of Concord were simply Thoreau’s companions,

soul mates he loved and mourned. In January of 1856 Thoreau witnessed the felling ofa great elm. The tree, which measured more than fifteen feet in circumference, hadmade creaking noises in a recent storm, frightening neighboring homeowners intobelieving that the great mass of wood was about to crash onto their roofs. For Thoreau,the destruction of this tree represented the slaughter of a priceless witness to the his-tory of the commumty.

"I have attended the felling," he wrote m his Journal, "and, so to speak, the funeralof this old citizen of the town ... I have not known a fitter occasion for a sermon of

late." Having "taken the measure of his grandeur," Thoreau spoke "a few words ofeulogy at his grave, remembering the maxim de mortms nil nisi bonum (in this casemagnum)" [speak nothing but good of the dead (m this case the best/]. But only thewoodchoppers and passersby heard his words. The shattered tree had hardly come torest than the "axe-boys had chmbed upon it like ants" and begun hacking at its limbs."How have the mighty fallen!" eulogrzed Thoreau, "Methmks its fall marks an epochm the history of the town."

How much of old Concord falls with it! The town clerk will not chromcle its

fall. I will, for it is of greater moment to the town than that of many a humaninhabitant would be. Instead of erecting a monument to it, we take all pos-sible pams to obliterate its stump, the only monument of a tree which is com-monly allowed to stand. How much of old Concord was cut away with it!A few such elms would alone constitute a township. They might claim tosend a representative to the General Court to look after their mterests, if afit one could be found, a native American one m a true and worthy sense, withcatholic principles.

"Our town," concluded Thoreau, "has lost one of its venerables." The woodcutters"have laid the axe ... to one of the king-posts of the town." "Is it not sacrilege," he

asked, "to cut down the tree which has so long looked over Concord beneficently?" "

Once the tree had been felled, its great size awed even Thoreau. His fellow

Concordians seemed oblivious to the glory of the living elm and would not appreciatethis magnificent specimen of plant life until it lay prostrate on the earth. At a dinnerparty shortly before the woodchoppers laid low the tree, he tried to convey the senseof wonder that the behemoth stirred m him. "I surprised some the other day," he wrotem his journal, recountmg the mcident, "by saying that when its trunk should lie pros-

30

trate it would be higher than the head of the tallest man in the town, and that two suchtrunks could not stand in the chamber we were then in, which was fifteen feet across;that there would be ample room for a double bedstead on the trunk, nay, that the verydinner-table we were sitting at, with our whole party of seven, chairs and all, aroundit, might be set there." "

Just as he had plumbed the depths of Walden Pond, Thoreau recorded the girths ofthe elms of Concord.~ These records were for him "the quantitative expression of hisimmersion in Nature," writes Alfred Kazin, "proof positive that he touched Nature onevery side."’ In the winter of 1846, Thoreau had measured the waters of Walden Pond,in part to prove that local wisdom regarding its depth was utterly in error. "It is remark-able how long men will believe in the bottomlessness of a pond," he wrote, "withouttaking the trouble to sound it. "8With its annual rings exposed by the axe, the trunk of the great elm offered a precise

record of the tree’s journey in time. Yet it seems that only Thoreau possessed the imagi-nation to count them. "Men have been talking now for a week at the post office aboutthe age of the great elm," he wrote, "as a matter interesting but impossible to be deter-mined." Even the very men who felled the tree "stood upon its prostrate trunk and

speculated upon its age, as if it were a profound mystery." He endeavored to show themthat in fact there was no mystery at all, but his words fell on deaf ears.By Thoreau’s measure, the elm had lived 127 years, but no one took seriously his

method or his result. Some villagers insisted that the tree was 200 years old; one main-tained it was closer to 150 years, having spent 50 years growing, another 50 standmgstill, and a final 50 dying. ("Wonder what portion of his career he stood still!" Thoreauwrote.) As with the storied depths of Walden Pond, Thoreau was incredulous that menwould choose to remain ignorant when the truth lay within easy reach, requiringbut a simple act of measurement to bring forth. "Truly they love darkness rather thanlight," he wrote in his journal.

They dwell within an mtegument of prejudice thicker than the bark of the cork-tree, but it is valuable chiefly to stop bottles with. Tied to their buoyant preju-

~

dices, they keep themselves afloat when honest swimmers smk.

Mary Emerson alone offered this rational poet a measure of consolation: "It was notthe fashion to be so original when I was young," she told Thoreau, before agreeing thatmdeed the fallen elm had revealed its truths only to him.The autumnal elm, with its "early and golden maturity," made a particularly strong

impression on the Concord naturalist. "It would be worth the while," he wrote inAutumnal Tints, "to set out these trees, if only for their autumnal value." "

Think of these great yellow canopies or parasols held over our heads and housesby the mile together, making the village all one and compact-an ulmanum,which is at the same time a nursery of men! And then how gently and unob-

"

"

served they drop their burden and let m the sun when it is wanted, their leaves "

not heard when they fall on our roofs and m our streets; and thus the villageparasol is shut up and put away’

Thoreau saw autumn as symbolic of the glorious destiny he predicted for the UnitedStates. It was an appropriate metaphor, for the vibrant colors of the northern decidu-ous forests rank among North America’s greatest natural wonders. The seasons ofhuman progress had brought the American to these autumnal shores, where a richbounty would be harvested as its people grew m wisdom and maturity.9

31

The great fall of elm leaves in October transformed Concord mto "a scene of a greatharvest-home," its paths and walks strewn with the summer’s spent array. In theirform and color, the great yellow masses remmded Thoreau of sheaves of wheat; it was"as if the harvest had indeed come to the village itself." Now, he suggested, "we mightexpect to find some maturity and flavor in the thoughts of the villagers at last." Wouldthere be, he wondered, an "answering ripeness" in the lives of the men who livedbeneath these glorious domes. He found it untenable that such beauty could accom-pany mean and illiberal thoughts. "Under those bright rustling yellow piles ..." wroteThoreau, "how can any crudity or greenness of thought or act prevail?" As he watcheda farmer disappear beneath the village elms, his wagon creaking beneath the fruits ofhis summer, Thoreau was tempted to follow the man to the granary, perchance to wit-ness a "husking of thoughts, now dry and ripe, and ready to be separated from theirinteguments ... " But he turned away, knowmg it would be "chiefly husks and littlethought ... for, as you sow, so shall you reap." "

For all his seriousness of purpose, Thoreau never lost his playful, rhapsodic voice.Standing beneath a cluster of October elms, "warm from their September oven," heimagined that he stood "within a ripe pumpkin-rind"; "I feel as mellow as if I were thepulp," he quipped, "though I may be somewhat stringy and seedy withal." For thisman, walking beneath the October elms of Concord was itself a harvest-of seeds andthoughts he had sown himself-and which many generations would reap.

NOTES

Unless otherwise noted, quotations from Henry DavidThoreau can be found m the Riverside editions ofThe Journals of Henry David Thoreau, VIII (Nov. 1,1855-Aug. 15, 1856). 117, 130-132; 139-142; X (Aug. 8,1857-June 29, 1858): 89 (edited by Bradford Torrey andFrancis H Allen, Boston: Houghton Mifflin, 1906);and "Autumnal Tmts" in Excursions (Houghton Miffhn,1883), pages 233-234.

1 Charles Joseph Latrobe, The Rambler m NorthAmemca, vol. 1 (New York: Harper & Brothers, 1835), (,43-44.

z Mircea Eliade, The Sacred and the Profane (NewYork: Harcourt Brace, 1959), 20-31.

3 John R. Stilgoe, Common Landscape of Amenca(New Haven: Yale University Press, 1982), 57.

4 William Howarth, ed. Walden and Other Wntmgs(New York: Modern Library, 1981), xxiv

5 Robert L. Gale, A Cultural Encyclopedia of the 1850sm Amenca (Londom Greenwood Press, 1993), 139-140 The Free Soilers, established m 1848, "adopted aplatform with three mam planks. no slavery m theterritories, no slavery in any new state, and freehomesteadmg rights for public domam settlers.The spirited slogan of the new party-’Free Soil,Free Speech, Free Labor, Free Men’-gave the partyits name." "

6 Thoreau made the followmg entry on 23 January,1856: "Holbrook’s elm measured to-day 11 feet 4

mches m circumference at six feet from ground, thesize of one of the branches of the Davis elm ...

Cheney’s largest m front of Mr. Frost’s, 12 feet 4mches, at six feet, 16 feet 6 mches, at one foot. Thegreat elm opposite Keyes’s land, near by (call it theJone’s elm): 17 feet 6 mches, at two behmd and oneplus before; 15 feet 10 mches, at four, 15 feet 5 inches,at six; 16 feet at seven and a half, or spike on westside. At the smallest place between the ground andbranches, this is a httle bigger than the Davis elm,but it is not so big at or near the ground, nor is it sohigh to the branching ... nor are the branches so big,but it is much sounder, and its top broader, fuller, andhandsomer." See Thoreau, /ournal, 135-136.

7 Alfred Kazm, A Wmter’s America Landscape mLiterature (New York: Alfred A. Knopf, 1988), 61.

8 Henry David Thoreau, "The Pond in Winter," inWalden and Other Wmtmgs, William Howarth, ed.(New York: Modern Library, 1981), 256.

9 William Howarth, xxvn.

Thomas J. Campanella is an urbamst and historian.His books mclude Cities from the Sky (PnncetonArchitectural Press, 2001) and Repubhc of Shade (YaleUniversity Press, forthcoming). He teaches m thedepartment of urban studies and planning at MIT, and isa Mercer Fellow at the Arnold Arboretum

32

t

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1. Publication Title Arnoldia 2. Publication No: 0004-2633. 3. Fihng Date: 31 October2001. 4. Issue Frequency Quarterly 5 No. of Issues Published Annually. 4. 6. Annual Sub-scription Price: $20.00 domestic, $25 00 foreign 7 Complete Mailing Address of KnownOffice of Publication: Arnold Arboretum, 125 Arborway, Jamaica Plam, Suffolk County,MA 02130-3500. 8. Complete Mailing Address of Headquarters of General Busmess Officeof Pubhsher: Arnold Arboretum, 125 Arborway, Jamaica Plain, Suffolk County, MA 02130-3500. 9. Full Names and Complete Mailing Address of Pubhsher, Editor, and ManagmgEditor: Arnold Arboretum, 125 Arborway, Jamaica Plam, Suffolk County, MA 02130-3500,pubhsher; Karen Madsen, Arnold Arboretum, 125 Arborway, Jamaica Plain, MA 02130-3500, editor. 10. Owner: The Arnold Arboretum of Harvard University, 125 Arborway,Jamaica Plam, Suffolk County, MA 02130-3500. 11 Known Bondholders, Mortgagees, andOther Security Holders Owning or Holdmg 1 Percent or More of Total Amount of Bonds,Mortgages, or Other Securities: none 12 The purpose, function, and nonprofit status ofthis organization and the exempt status for federal income tax purposes have not changedduring the preceding 12 months. 13 Publication Name: Arnoldia 14. Issue Date for Cir-

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’

The Arnold oretum, , / ~, N E W 5 . 2 0 0 1

Projects, Programs, and Research: Planning for the FutureRobert E. Cook, Director

Rising out of the ground belowthe Dana Greenhouses is a visible

harbinger of change at the ArnoldArboretum. On a sloping, three-acre site, massive fieldstone walls

that were constructed over the

summer have formed a series of

terraces to hold a new collection of

sun-loving shrubs and vines Soona steel-framed, wood-covered

pavilion will be erected to providea shady resting place overlookingthe collection beds and the greatlawn that runs down the center of

the garden. This project is one ofseveral major landscape improve-ments that will be completed overthe next two years.

These physical transformationswill be accompanied by other sig-nificant changes in the programsand operations of the Arboretum.New goals, consistent with ourtraditional mission of research and

education, are emerging from the

long-range planning process thathave engaged the staff over thepast ten months. One goal willinclude a renewed commitment

to improve the physical settingand, therefore, the image of theArboretum in the eyes of the pub-lic. The shrub and vine gardendescribed above will set a new

standard of excellence to be fol-

lowed in renewing the older partsof the grounds-the gates andwalls, the benches and pathways,location and directional signage.

We are also contemplatinga significant expansion of the

Arboretum’s public and profes-sional education programs. The

Radcliffe Institute (formerlyRadcliffe College) has approachedus about the possibility of merg-ing their programs in landscapedesign and landscape design his-tory with our existing adult edu-cation programs. Through a seriesof rigorous courses and studios,the Radcliffe programs have for

35 years offered part-time and

continuing education students acurriculum that leads to a certifi-

cate in landscape design and land-scape design history. Providing ahome at the Arboretum for the

programs would greatly enhanceour educational acumues.

Finally, the planning processhas revealed significant dissatisfac-tion with the future prospects for

scholarly work at the Arboretum,which currently includes researchon the lmmg collections inJamaica Plain, herbarium and

library research in Cambridge, andresearch conducted in foreign set-tings such as the tropical forests ofSoutheast Asia. It is unclear that

the existing criteria for research

appointments and the related sup-

port systems truly serves the long-term interest and international

reputation of the institution.

Alternative configurations ofthe research model are not easyto envision because of the compli-cated relations between the Arbo-

retum and academic units at

Harvard At the very least, the

long-range plan will include acall for change in the model andan outline of the benefits that

such a change might bring toour programs of research and edu-

cation on woody plants aroundthe world.

Mary Gibson HenryMedal HonorsCarroll Wood

Carroll E. Wood, Jr., professor ofbiology emeritus at Harvard Uni-versity, is among the first recipi-ents of the Mary Gibson HenryMedal, presented on September 7at the Henry Botanic Garden inGladwyne, Pennsylvania.

Dr. Wood has enjoyed a longassociation with Harvard Univer-

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©

. from page 1

sity and the Arnold Arboretum.

He received a master of arts from

Harvard in 1947 and a doctoratein 1949. After serving as a biol-

ogy instructor for the Faculty ofArts and Sciences from 1949 to

1951, Dr. Wood became an asso-

ciate curator for the Arboretum

in 1954 and a curator in 1970, a

position he held until his retire-ment in 1988. He lectured on

taxonomy at Harvard between

1964 and 1972, when he wasnamed a professor of biology.Dr. Wood continues to focus his

energies on the completion of hisGenerac Flora of the SoutheasternUnited States. Smce its begmmngin 1954, 162 papers of this pro-digious work have been pub-lished, both in the Journal of the

Arnold Arboretum and the Harvard

Papers zn Botany.While a student at the Umver-

sity of Pennsylvania in 1942, Dr.Wood became acquainted withMary Gibson Henry during a col-lecting trip for the University’sFlora of Pennsylvania. Their com-mon interest in native plants keptthem in contact over the years,

beginning with a gift of Arizonayuccas and cacti he sent to Mrs.

Henry during the war years. Later,he provided scientific support forsome of her plant discoveries,including writing the Latindescription for Lzlrum zrzdollae,the pot-of-gold lily. (See MaryHarrison’s article "Mary Gibson

Henry, Plantswoman Extraor-dinaire" in Arnoldia, vol 60, no. 1.)Around the time of Mrs. Henry’s

death in 1967, Dr. Wood becamea member of the Board of the

Henry Foundation for BotanicalResearch and has been instrumen-

tal in the acquisition of additionalland and of nonprofit status forthe Foundation’s gardens.

The Henry Botanic Gardenhonored Dr. Wood and three

other medal recipients with a cer-

emony and reception at the

Henry Foundation for BotanicalResearch, where, on 50 acres, awide variety of plants grow innaturalistic woodland and hill-

side settings and research gardenspromote genetic diversity. Dr.Wood, who contributed wild-collected plant material to thegarden’s collections, was cited forhis "extraordinary contributionsto botanical research."

Albert J. Fordham, 1911 -2000Al Fordham, who retired in 1976 as head propagatorand research horticulturist of the Arnold Arboretum,passed away last December at the age of 89 inNorwood, Massachusetts. He had been affiliatedwith the Arboretum for nearly 50 years. Al will beremembered for his many contributions to horticul-

ture, including the many new cultivars of dwarfconifers he developed and his work on woody plantseed germination.

Al began his career at the Arboretum in 1929 as astudent intern to propagator William H. Judd. In1936 he spent a year studying European horticulturalpractices at the Royal Botanic Garden in Kew. Afterthree years of service in the U.S. Army Infantry duringWorld War II, Al returned to the Arboretum as assis-tant superintendent of grounds. From 1958 to 1976,he was the sole plant propagator at Harvard Univer-sity, and it was during this period that the propaga-tion department of the Arboretum was reorgamzedand the Dana Greenhouses were designed and built.

Al’s work was distinguished by a willingness totest new methods of propagation and by his fastidi-ous recording of experimental results. In addition toteaching and lecturing extensively on propagation,he published more than 100 research papers between1958 and 1975, many in the pages of this magazine.His research topics included the barriers that hinder

germination of woody plant seeds, the role of physi-ological juvemlity in plant propagation, and meth-ods of seed dispersal. He also studied microclimatesas they relate to plant survival.

In 1963 Al began collecting and experimentingwith seed from witches’ brooms, which resulted inthe introduction of several new cultivars of Pznus

bankstana, P. strobus, and P. resznosa. He gave his cul-tivars native American names such as "Merrimack,"

v v contznued on page 3

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IMLS Grant Integrates Past andPresent CollectionsThe Arnold Arboretum was recently awarded a$49,000 conservation support grant from the federalgovernment’s Institute of Museum and Library Ser-vices in Washington, D.C. As outlined in the pro-posal, written by Jianhua Li and Peter Del Tredici,the grant will be used to finish integrating data fromthe Arboretum’s herbarium specimens into the liv-

ing collections database (BG-Base).The project will proceed in three interrelated

steps. The first is to enter information into the data-base from each sheet (voucher) of specimens thathave been collected on the Arnold Arboretum’s

grounds. The staff will then be able to generate a listof plants that have not been vouchered or whosespecimens do not include diagnostic structures(mostly flowers and fruits), leading to the secondstep: specimen collection, preparation, and dataentry for those plants. In the third step, data will beentered for existing herbarium specimens represent-ing plants no longer growing on the grounds.

At its most basic level, the project will allowthe staff to learn which plants now growing on thegrounds are not adequately represented by specimensin the herbarium. In addition, it will expand thedatabase to include not only plants currently livingon the grounds, but also those that have ever grownhere and are represented by an herbarium specimen.Our services to botanical and horticultural commu-

nities at home and abroad will also be enhanced sincethe database will be one of the most comprehensivevoucher systems for large living collections in theworld, a crucial component of meaningful scientific

Botanical drawing of Salisburia adiantbifolia (Ginkgobiloba) from Siebold and Zuccarini’s Flora Japonica,courtesy of the Arnold Arboretum Library.

research. The Arboretum staff have been workingslowly and deliberately to achieve this goal for overten years, but the IMLS grant will allow us to incor-

porate the last 24,000 herbarium specimens into theliving collections database in timely fashion.

The project team, to be coordinated by JianhuaLi, will include Kyle Port, Susan Kelley, Irtna Kadis,and Sara Straate.

~ ~ from page 2

"Chippewa," and "Neponset" in recognition of NewEngland’s tribal heritage. In 1985 Al himself wasrecognized for his pioneering work with conifers bythe naming of Prnztr parvzflora ‘A1 Fordham’.

Over the years, Al was honored for his achieve-ments by a number of amateur and professional orga-nizations. He was an honorary member of theMassachusetts Horticultural Society, which awardedhim its Jackson Dawson Medal in 1965. Other hon-ors included awards of merit from the InternationalPlant Propagators Society (1971), from the Interna-tional Lilac Society (1973), and the American Coni-fer Society (2000). He also received a professionalcitation and certificate of appreciation from theAmerican Horticultural Society.

In 1976 Al retired from his Arboretum dutiesbut continued his horticultural research as well as

writing, lecturing, and consulting. Among otherpost-retirement achievements, he published theArnoldia Manual of Propagatzon of Selected Conzferr in1977 as a separate issue of this magazine.

Al Fordham’s expertise and his insatiable curios-

ity about all aspects of plant development proved atimely fit for an institution that required a masterpropagator and a steady stream of new materialfor its living collections. Through his publishedwork, his classes, and his lectures, he taught a gen-eration of New England gardeners about plantpropagation and left a legacy to horticulturiststhroughout the world.

E8

SummerInternsThe 2001 Summer

Interns in the

Arboretum’s largestwhite oak (Quercusalba), from left to

right: John Backer,Rose Morgan, Sarah

Carter, Sara Straate,Dan March, BethanyKnorr, Emma Stark

Schiffman, Kim

Rennick, Steve

Egger, Nicole

Napoleon, MarissaFarris, WhitneyO’Hanian. Not

pictured: JeanGauthier.

Tides Foundation to Receive Plant Sale Donation

The Tides Foundation’s 9/11 Fund will receive

$5,000 from the Arnold Arboretum to support reliefefforts for the events of September 11. DirectorRobert E. Cook announced at the annual fall plantsale that proceeds from the live auction would bedonated to assist those immediately affected by thetragedy, and that a Quercus coccznea (scarlet oak) wouldbe planted on Peters Hill to commemorate the vic-tims. The Tides Foundation was chosen by Arbore-tum staff as the recipient of the Arboretum’s gift.

The Tides 9/11 Fund was established to meet

immediate and long-term needs in the wake of

September 11 and to promote a just and peacefulnational response to this cmsis. As of October 22,over $335,000 had been raised for this fund, and$215,000 in grants had been allocated to 18

organizations. In crafting their grant-makingstrategy, the Tides Foundation has worked to blend

their core values of social and economic justicewith the pressing needs created in the aftermathof the attacks.

For more information about this fund or for a

complete list of grantees, contact the Foundation at

mfoC~tides.org.

The James Arnold SocietyThe Arnold Arboretum is proud to announce the formation of the James Arnold Society,named for the benefactor whose bequest made possible the Arboretum we value so much

today. This new society will recognize those who have made life income gifts or bequestprovisions for the Arnold Arboretum. If you have included the Arboretum in your long-termcharitable plans, we would like to welcome you into this special group.

Members of the James Arnold Society will be invited to events at the Arboretum andreceive a complimentary invitation to any one of the Harvard University-sponsored financialand estate-planning seminars.

Please contact Anne McClintock or Amy Goldman of University Planned Giving at800.446.1277 or pgo@harvard.edu to discuss your current gift arrangements or to exploreways to benefit the Arboretum through a life income gift or bequest. Your inquiry will beheld in the strictest confidence.

For more information, please visit www.haa.harvard.edu/pgo.