Vol. 45 No. 4 Fall 1985

Page 2 The Case for Monkey-Mediated Evolution in Big-Bracted Dogwoods Richard H. Eyde

Amoldia (ISSN 0004-2633) is published quarterly m 10 Reminiscences of Collecting the Type spnng, summer, fall, and winter by the Arnold Specimens of Metasequoia glyptostroboides Arboretum of Harvard University. H. H. Hu & Cheng Subscriptions are $12.00 per year; single copies $3.50. Hsueh Chi-ju Second-class postage paid at Boston, Massachusetts.

Postmaster: Send address changes to: Amoldia The Arnold Arboretum The Arborway Jamaica Plain, MA 02130

Copyright © 1985 President and Fellows of Harvard College.

Edmund A. Schofield, Editor

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Front cover: The inflorescence and fruits of flowering dogwood (Comus flonda L.), top, and of kousa /C. kousa Hance), bottom. To find out why kousa has compound fruits and flowering dogwood does not, see page 2. Watercolor by Alice Tangenm, Smithsoman Institution. Front cover foldout. Three compound fruits of Cornus capitata Walhch, a species of dogwood from the Hima- layas. From Curtis’s Botamcal Magazme for 1852. This page: Painting by an anonymous Chinese artist of the 19 "Les a Far-Northern Garden tree from which Hsueh Chi-~u collected the type spec- Quatre Vents," imens of Metasequoia glyptostroboides H. H Hu & Francis H. Cabot Cheng m 1946. (See page 11.) Inside front cover: The Arnold Arboretum’s Plant Hardmess Zone Map. (See 32 Plant Hardiness-Zone Maps pages 32 through 34.) Inside back cover Calypso bul- Donald Wyman bosa Sallsb., an orchid of cool, mossy woods m Canada Harrison L. Flint and the Umted States. (See page 19.) Photograph by Fran- cis H. Cabot. Back cover: Bunchberry (Corns canaden- sis L.), a small-bracted dwarf comel with tight clusters 35 What Determines a Plant’s Cold Hardiness? of fruits. From The Monthly Flora, or Botamcal Maga- John W Einset zine, Volume 1 (1846). (See page 2.)

39 A Letter to Readers

41 Index to Volume 45

RESEARCH REPORT

~

The Case for Monkey-Mediated Evolution in Big-Bracted Dogwoods Richard H. Eyde

Because New World monkeys are blind to red and live only in warm regions, American big-bracted dogwoods produce only simple fruits

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Say "dogwood" to anyone from eastern North Big-bracted dogwoods, American and Asian, America and the plant that comes to mind is had a common evolutionary origin: their Comus florida, called flowering dogwood chemistry attests to that (Bate-Smith et al., because four petal-like bracts beneath each 1975), as do serological experiments (Brunner tight flower cluster make a cluster look like and Fairbrothers, 1978) and cases of cross- one big showy flower. Dogwoods with such breeding (Orton, 1969; Bean, 1970; Bond, blossoms are a minority. Most of the 50-plus 1984; Santamour and McArdle, 1985). The species in the dogwood genus-Cornus in ease with which they hybridize weighs heav- the wide Linnaean sense-bear broadly ily against the taxonomic splitting of big- branching flower clusters in which bracts are bracted dogwoods into two genera, Dendro- rudimentary or lacking. Dwarf comels-C. benthamia and Cynoxylon, a practice met canadensis and two similar species-have with often enough to need condemning. showy-bracted blossoms and tight clusters, Though most species of Comus have white, but the bracts, like the plants themselves, blue, or blue-black fruits, the fruits of showy- are small. Besides C. florida, the subgroup bracted species (dwarf cornels included) are with big-bracted clusters includes C. nuttal- red. Fruits of Old World members of the big- lii of western North America and C. disci- bracted subgroup differ from their New World flora, a mountain tree of Mexico and Central counterparts, however, in that they are com- America. (The bracts of C. disciflora are like pound. [See the cover of this issue of Amol- those of C. florida and C. nuttallii only in dia. Our flowering dogwood bears bunched the bud; they wither and fall off when buds fruits, the individuals parting readily from turn into blossoms.) There are big-bracted the bunch: in C. kousa and other Asian dogwoods in Asia, too. Of the 19 species rec- members of the subgroup, flowers can be sep- ognized in recent treatments (Poyarkova, arated, but the separability is lost as flowers 1950; Fang, 1953; Hu, 1980; Hu and Soong, change to fruits. Ripe compound fruits look 1981 only C. kousa is widely known and enough like fat strawberries that one species, grown in North America. C. capitata, is called strawberry dogwood, 4

- The evolutionary relationships among the subgroups of Cornus. Because compound fruits occur in no genus of the dogwood family but Cornus, and there only in the Old World members of one subgroup, students of dogwood evolution can tnfer with confidence that they are denved This diagram makes the comelian cherry subgroup (Cornus mas and allied species) more advanced than bractless dogwoods, yet swrtchmg these two branches of the diagram would not affect the denved status of compound fruits, and no matter which relative of Cornus is chosen as its nearest evolutionary neighbor, the conclusion remams that the change from simple fmtts to compound fruits occurred just once, as big-bracted dogwoods separated into today’s Amencan and Asian forms

and the flesh of compound fruits is soft and Indeed, the stages can be seen at different sweet, unlike the tart or bitter flesh of dog- levels in a half-ripe compound fruit. Stained wood fruits in general. [See the front-cover serial sections through a fused region show foldout.] normal epidermal cells, one layer for each of When P. F. Maevskii (1881) examined fruits the adjoining ovaries, only at the fruit’s sur- of Comus capitata microscopically, what he face. Within the fruit, the corresponding saw misled him. Seeing-he thought-free cells show signs of repeated division. They individuals, each set in a cup-like hollow, he no longer have their typical rectangular out- concluded that adjoining fruits do not really lines, and they interdigitate: cells from each unite, that they sink into an expanding fruit epidermis penetrate the other. There are, in stalk as the complex ripens. Museum tech- some places where epidermises once were, nician Stan Yankowski and I repeated Maev- nests of the thick-walled, heavily staining skii’s work with a modern microtome-the cells called sclereids. As the cluster ripens to anatomist’s precise slicer-and a modem a sphere roughly half the diameter of a golf microscope, taking pains to look at all stages ball, tissues tear and cavities develop near of growth, and we found true developmental the sclereids, hence the illusion of nonunion fusion of the ovaries. (Dogwood ovaries are that Maevskii saw. inferior; that is, petals, calyx lobes, and sta- Evolutionists from Darwin on have stud- mens are above the ovary, out of union’s way.)/ ied the selective pressures that make flowers ; 5

so diverse, but only recently have they taken Borowicz and Stephenson, in press) but not note of pressures working on the fruits of enough to cause an evolutionary change. Fur- temperate regions (Thompson and Willson, thermore, the beasts are color-blind: the red- 1979; Stiles, 1980, 1982, 1984; Stiles and ness of a fruit means nothing to them. White, 1982; Morden-Moore and Willson, 1982; Willson, 1983, Borowicz and Stephen- son, in press). Thus, evolutionists have over- looked the fact that compound-fruited dog- woods pose the following problem: any reasonable diagram connecting Cornus to a neighboring group, and subgroups of Cornus to each other, shows that ancestral dogwoods had open flower clusters, that dogwoods with bunched flowers and ordinary fruits came later and gave rise to those with compound fruits. The change to bunched flowers likely had to do with pollinating insects, but the further change to compound fruits must have been dispersal-linked. What new means of dispersal would have made a compound fruit a betterment? And why are dogwoods with such fruits found only in the Old World? Robins and other migratory birds are the principal dispersers of the flowering dogwood (Baird, 1980). They also peck at fruits of kousa dogwoods introduced to gardens. By tearing up a compound fruit, they doubtless spread some of its seed-containing stones. (As in peaches, inner cell layers of the fruit wall turn into a "stony" housing for the seed. Most dogwoods have two seeds per stone, and a compound fruit has several stones.) But birds the size of robins, which are so well fitted to the bean-sized individual fruits of New World dogwoods, can hardly have pro- vided pressure for the change to compound fruits. Grouse eat dogwood fruits, but tests with and indicate the Stamed microtome secuons of developmg Cornus sp quails pheasants gal- frul ts. linaceous does a stone more gizzard dogwood o A somewhat oblique slice through the entire fruit of Cornus capitata, harm than good (Krefting and Roe, 1949). In magnified I1 times /See the front cover foldout of this issue of Arnold~a / b The area marked with a rectangle in a, enlarged to about 70 times Asia, hornbills and fruit pigeons spread large- (Liquid-preserved specimen taken from a wild tree in Nepal ~D H Nicol son 2371~ c A section like that in b, but thinner and from a developing fruited but the birds are plants efficiently, fmtt of Cornus kousa (Liquid-preserved specimen from a cultivated dog- and Asiatic like our own, wood on the grounds of the Smithsonian Institution ) Groups of sclereids tropical, dogwoods, (arrows) have differentiated where ovaries have fused are largely temperate. Mice, raccoons, and bears disperse some dogwood stones (Martin et al., 1951; Rogers and Applegate, 1983; 6

That leaves monkeys. Monkeys disperse have emerged from Africa (Delson, 1980; seeds and stones of fleshy fruits by spitting Sohn, 1983, 1984a, 1984b). Lately, European and by voiding (Hladik and Hladik, 1967, specialists have also learned to spot the 1972; Lieberman et al., 1979). Macaques, the remains of dogwood subgroups that have dis- northernmost of monkeys, see color much as appeared from Europe. Some of the Miocene human beings do (De Valois and Jacobs, impressions once called "persimmon calyx- 1968); they eat Cornus fruits, both simple es" are now thought to be the four bracts of and compound (Uehara, 1977; Maruhashi, Comus (Gregor, 1982), and fossil fruits, when 1980), and they once ranged almost every- they are well preserved, can be identified where within the range of compound-fruited more accurately than that. Size, shape, sur- dogwoods. A critic could object that match- face features, and internal structure mark a ing the ranges of macaques and dogwoods fruit stone as belonging to the big-bracted merely shows that both of them are creatures subgroup. Then, if it is ovoid or ellipsoid, it of the deciduous forest. Taking fossils into must be from a solitary fruit like that of C. account, however, boosts the case for mon- florida. Such stones occur in Oligocene and key-mediated evolution. Macaque remains Miocene brown-coal deposits of the German are found in the Pliocene and Pleistocene of Democratic Republic (Mai and Walther, 1978).). Asia as far north as Beijing and Korea and at If, however, a stone is asymmetrical, tapered sites in Europe dating back almost to latest to the base, and faceted, it is from a com- Miocene, when macaques are thought to pound fruit. Only younger beds have yielded

The natural ranges of macaques (hatchmg) and compound-frmted dogwoods (squares). Fruit stones like those of flowering dogwood (upper inset) occur as fossils in East Germany, stones from compound-fruited dogwoods occur at younger sites m Alsace and near the Dutch-German border Compound fruited dogwoods likely came to some sites-the Ryukyus, for example-with birds or people Five thousand years of tilling made the blank spot in the Yellow River bastn The macaque range is from Zhang et al ~1981~, Fooden (1982), and Wolfheim /1983/ The dogwood range is from the Herbanum of the Arnold Arboretum, the Gray Herbanum, the U S National Herbanum, and all germane flonsnc works 7

such: a mid-Pliocene locality near Cologne References another in Alsace (Burgh, 1978, 1983), (Geis- Baird, J. W. 1980. The selection and use 5f fruit by birds sert and Gregor, 1981 and a later Pliocene m an eastern USA forest. Wilson Bulletin 92: locality at Tegelen in the Netherlands (Mai, 63-73. 1976). The Tegelen beds have also yielded Bate-Smith, E. C., I. K. Ferguson, K. Hutson, S. R. Jensen, B. J. Nielsen, and T. Swam. 1975. Phytochemical bones of a macaque (Schreuder, 1945; see interrelationships m the Comaceae. Biochemical and Van for infor- Berggren Couvering, 1979, Systemaucs and Ecology 3: 79-89. [This is an mation on the age of the Tegelen beds). important work for recognizing natural subgroups Big-bracted dogwoods formerly extended within Cornus, but one conclusion is at odds with round the Northern Hemisphere, and all had my reasoning on dogwood evolution. The authors say hedgingly that C. capitata, despite its com- ordinary fruits until monkeys came in con- pound fruits, seems chemically closest to ances- tact with them about five million years ago. tral Cornus. I read their data differently m a a meant Selection for better monkey meal review I am writing of dogwoods and their allies.] better scattering of seeds; so compound Bean, W. J. 1970. l~ees and Shrubs Hardy m the Bnush fruits replaced the simple ones, but only in Isles, 8th edition, Volume 1. London: John Mur- Eurasia. America retains the older kind ray. [Page 704 mentions the probable crossing of Cornus kousa with C. nuttalln.] because the New World blind to monkeys, Berggren, W. A., and J. A. Van Couvenng. 1979. Quater- and tied to warmer red (Terborgh, 1983) nary. In Treatise on Invertebrate Paleontology, regions, never took up foraging on dogwoods. Part A, 505-543. Boulder, Colorado, and Law- The behavioral side of this scenario is test- rence, Kansas: Geological Society of Amenca and able. China and Japan have monkeys, dog- University of Kansas Press. Bond, J. 1984. Some North American dogwoods at Wind- both kinds kind in woods of (our cultivation), sor. The Garden 109: 154-155. and primatologists to do the testing. Borowicz, V. A., and A. G. Stephenson. In press. Fruit composition and patterns of dispersal of two Cor- nus spp. Oecologia Acknowledgments Brunner, F., and D. E. Fairbrothers. 1978. A comparative For thoughts and information, I thank C. serological investigation within the Comales. Badgley, B. Beehler, A. K. Behrensmeyer, V. Serological Museum Bulletin 53: 2-5. Burgh, J. van der. 1978. The Pliocene flora of A. E. E. S. Morton, A. F. Borowicz, Delson, Fortuna-Garsdorf I. Fruits and seeds of angio- M. R. S. Richard, Leighton, J. Oppenheimer, sperms. Review of Palaeobotany and Palynology D. Ripley, B. B. Simpson, E. W. Stiles, J. N. 26: 173-211. [Stones that Burgh would recognize Thompson, M. F. Willson, and, above all, R. in 1983 as those of a compound-fruited dogwood W. Thorington. Dan Nicolson collected Cor- are here called Carpohthus natans.] . 1983. Allochthonous seed and fruit floras from nus for me D. E. capitata years ago. Boufford, the Pliocene of the lower Rhme basm. Review of S. A. Spongberg, and S.-Y. Wang eased the Palaeobotany and Palynology 40: 33-90. way to publication. Delson, E. 1980. Fossil macaques, phyletic relationships and a scenario of deployment. In The Macaques: Studies in Ecology, Behavior and Evolution, edit- ed by D. G. Lindburg, 10-30. New York: Van Nos- trand Reinhold. De Valois R. L., and G. H. Jacobs. 1968. Primate color vision. Science 162: 533-540. Fang, W.-P. 1953. Notes on Dendrobenthamia. Acta Phytotaxonomica Smica 2: 89-114; plates 11-16. [In Chinese with English summary.] Fooden, J. 1982. Ecogeographic segregation of macaque species. Pnmates 23: 574-579. 8

Geissert, F., and H.-J. Gregor. 1981. Eme neue Elsassische gen, Abteilung B, Palaobotamk 2 ~1/ [19G4]. One Phozan-Flora, die "Saugbagger-Flora" der Kies- of the figures m this earlier work-a photo of a grube von Sessenheim (Bas-Rhin). Couner For- Cornu stone from Wiesa near Kamenz-rs per- schungsmstitut Senckenberg 50: 59-71. [Dendro- suasrve enough that I added the locality to my benthamia tegehensis is listed on page 64.] map.] Gregor, H -J. 1982. Die Jungtertiaren Floren Sud- Martm, A. C., H. S. Zim, and A. L. Nelson. 1951. Amen- deutschlands ; Palaokarpologie, Phytostrarigra- can Wildlife eJ Plants. New York: McGraw-Hill. phie, Palaookologle, Palaokhmatologie. Stutt- Maruhashi T. 1980. Feeding behavior and diet of the gart : F. Enke. ["Persimmon sepals" reidentified as Japanese monkey (Macaca fuscata yakuy on dogwood bracts, pages 119-120.] Yakushima Island, Japan. Primates 21: 141-160. Hladik, C. M., and A. Hladik. 1967. Observations sur le Morden-Moore, A. L., and M. F. Wrllson. 1982. On the role des primates dans la dissemmation des ecological significance of fruit color m Prunus vegetaux de la foret gabonaise. Biologia Gabomca serotma and Rubus occidentalis: Field expen- 3: 43-58. ments. Canadian fournal of Botany 60: 1154-1160.

. et E. 1969. ornamentals. , and 1972. Disponibilites alimentaires Orton, R., Jr. Hybndizmg woody domames vitaux des primates a Ceylan. Terre et International Plant Propagators’ Society Com- Vie 26: 149-215 bmed Proceedmgs 19: 371-375. Hu, W. K. 1980. Materiae ad floram Cornacearum sini- Poyarkova, A. 1. 1950. Sektsiya Benthamia (Lindl.) nakai carum. Sichuan Daxue Xuebao (Zman Kexue) roda Cynoxylon Raf. i ee vidy. Botanicheskie [Journal of Sichuan Umversity, Natural Sciences] Matenaly Gerbanya Botamcheskogo Instituta 1980 (3): 155-174. [In Chinese.] imem Komarova Akademm Nauk SSSR 12’ T. 181-195. , and P. Soong. 1981. Cornaceae. In Flora Sichuanica (Spermatophyta), edited by Fang W. Rogers, L. L., and R. D. Applegate. 1983. Dispersal of P., Volume 1, 320-395, 468-475. Chengdu: fruit seeds by black bears. /oumal of Mammalogy Chengdu Institute of Biology. [In Chinese.] 64: 310-311. Krefting, L. W., and E. I. Roe. 1949. The role of some Santamour, F. S., Jr., and A. J. McArdle. 1985. Cultivar birds and mammals m seed germmation. Ecolog- checklists of the large-bracted dogwoods: Cornus ical Monographs 19: 269-286. flonda, C. kousa, and C. nuttalln. Journal of Lieberman, D., J B. Hall, and M. D. Swame. 1979. Seed Arbonculture 11: 29-36. dispersal by baboons in the Shai Hills, Ghana. Schreuder, A. 1945. The Tegelen fauna, with a descrip- Ecology 60: 65-75. tion of new remams of its rare components ~Lep- Maevskii, P. F. 1881. 0 stroenm ploda Benthamia fragi- tobos, Archidiskos mendionahs, Macaca, Sus fera. Izvestiya Obshchestva Lyubitelei Estestvoz- strozzy. Archives Neerlandaises de Zoologie 7: namya, Antropologii i Etnografil 37: 28-30. [Yale 153-204. Umversity’s library has this ranty. Benthamia Sohn, P. 1983. Early Man at Thrubong Cave No. 9 near fragifera is Comus capitata See the front-cover Ch’ongju, Korea Seoul Laboratory of Prehistory, foldout of this issue of Amoldia.] Museum of Yonser University. [In Korean with Mai, D. H. 1976. Dendrobenthamia tegehensis nov. English summary.] sp.-Em neues ostasiatisches Florenelement im . 1984a Early man m Korea. East Asian Tertiaryl Altquartar Europas. Abhandlungen des Staat- Quaternary Newsletter 1. 61-62. lichen Museums fur Mineralogie und Geologle . 1984b. The palaeoenvironment of Middle and zu Dresden 25: 113-123. Upper Pleistocene Korea. In The Evolution of the Asian edited R. O. , and H. Walther. 1978. Die Floren der Haselbach- East Envmonment, by Whyte, er Sene im Weisselster-Becken (Bezirk Leipzig, Volume 2, 877-893. Hong Kong: Centre of Asian DDR). Abhandlungen des Staathchen Museums Studies, University of Hong Kong. f ur Mmeralogie und Geologie zu Dresden,Volume Stiles, E. W. 1980. Patterns of fruit presentation and seed 28. [The mid-Oligocene fossils include Cynoxy- dispersal m bird-disseminated woody plants m lon (Cornus) caroln, stones like those of modem the eastern deciduous forest. Amencan Natural- C. disciflora, C. flonda, and C. nuttalhi. Mai, the ist 11G: 670-688. fruit specialist of the Mai-Walther partnership, -. 1982. Fruit flags: Two hypotheses. Amencan also mentions a suggestive similarity of C. caroln Naturahst 120: 500-509. to Miocene stones that he called simply Comus -. 1984. Fruit for all seasons. Natural History 93 sp. m his "Die Mastixioideen-Floren im Tertiar (8): 42-53. and D. W. White. der Oberlausitz," Palaontologische Abhandlun- , 1982. Additional information on temperate bird-disseminated fruits: Response 9

to Herrera’s comments. Amencan Naturahst Nakao, and T. Umesao, pages 187-232. Tokyo: 120: 823-827. Chuokoron-sha. [In Japanese.] Terborgh, J. 1983. Five New World Pnmates A Study m Willson, M. F. 1983. Plant Reproductive Ecology. New Comparatme Ecology Princeton. Prmceton Um- York: Wiley. versity Press. [Page 77 treats vision and fructivo- Wolfheim, J. H. 1983. Pnmates of the World. Seattle: ry.] University of Washmgton Press. Thompson, J. N., and M. F. Willson, 1979. Evolution of Zhang, Y. Z., S. Wang, and G. Q. Quan, 1981. On the temperate fruit/bird interactions: Phenological geographical distribution of primates m Chma. strategies. Evolution 33: 973-982. Journal of Human Evolution 10: 215-226. Uehara, S. 1977. A biogeographic study of adaptation of Japanese monkeys with regard to food habits. In Keishitsu, Shmka Reichorm [Morphology, Evo- Richard H. Eyde is curator m the Department of Botany, lution, and Primates], edited by Y. Kato, S. Smithsonian Institution, Washington, D.C. Professor Hsueh Chi-7u, the botamst who collected the type specimens of Metasequoia glyptostroboides m 1946. Photographed m October 1984 by Peter S. Ashton. Reminiscences of Collecting the Type Specimens of Metasequoia glyptostroboides H. H. Hu & Cheng Hsueh Chi-ju

After forty years a Chinese botanist has still-vivid memories of his eager and determined quest for specimens of the dawn redwood

When the Arboretum’s Director, Peter S. Ashton, visited the People’s Republic of China during the fall of 1984, he met Professor Hsueh Chi-ju, of Southwestern Forestry College, Kunming City Yunnan province. As a young man, Professor Hsueh had collected the type specimens of the recently discovered dawn redwood, Metasequoia glyptostroboides H. H. Hu ~7 Cheng. Director Ashton invited him to write a first-hand account of his collecting trips, for publication in Amoldia. To commemorate the approaching forty-fifth anniversary of the discovery of living Metasequoia trees, we are pleased to present herewith Professor Hsueh’s fascinating account of his adventures.

Forty years ago, I happened to see the speci- nospermae, since the opposite arrangement men of Metasequoia glyptostroboides that of the leaves and cone scales differed from Mr. Wang Zhang had collected at Modaoqi that of G. pensilis and other members of the [knife-grinding] village in Wanxian county, Taxodiaceae. China. The next year, following the route Since the specimen Mr. Wang collected Mr. Wang had taken, I made two trips there had no male inflorescences and since the to collect perfect specimens and to conduct cones had been picked up from the ground, further investigations. Although I am old we didn’t know how the cones grew on the now, the two trips are still fresh in my mem- branches. In addition, we had no information ory. on whether it was deciduous or evergreen, on I graduated from the Forestry Department its flowering season, or on its ecological char- of the former National Central University at acteristics and distribution. Zhongjing (Chungking) in 1945 and then Further research being necessary, Profes- worked on the gymnosperms, studying for a sor Cheng naturally advised me to collect master’s degree under the guidance of Pro- some perfect specimens and to make an fessor Cheng Wanjun. One day in 1945, Wang investigation. Since we had no funds and Zhang, who worked at the Central Forestry everybody was quite hard up, I could only go Experimental Institute, sent a cone-bearing to the place on my own, carrying a few pieces specimen collected at Modaoqi to Professor of simple baggage and specimen-clips. I left Cheng for identification. Its vernacular name Chungking city by streamboat and, after two was shui-shan (water fir), and it was some- days, arrived at Wanxian county, on the what similar to Glyptostrobus pensilis (G. northern bank of the Changjiang (Yangtze) lineatus). After making a preliminary iden- River. After crossing the river, I had to walk tification, Professor Cheng considered that 120 kilometers [72 miles] to my destination. it might belong to a new taxon of the Gym- In 1946 I made two trips from Chungking to 12

Modaoqi, in February and May, respectively, both times single-handedly.

The First Trip to Modaoqi I remember that on my first trip the boat was moored in Fengdu county for the first night. On a hill behind the county town was a tem- ple regarded in the Old China as an inferno where the "Lord of Hell" reigned. Dead souls were supposed to go there to register. So I made use of this rare opportunity to take a solitary night walk in this weird and dreadful place-evidence that I was full of vigor and curiosity in my youth. At that time there was no highway from Wanxian county to Modaoqi village. My trip was very difficult, the trails threading through the mountains being less than one foot wide. The region was inhabited by the Tu minority and had been isolated from the outside world for ages. During the war of resistance against Japan, the Hubei provincial government moved to Enshi county in its neighborhood; thenceforward its intercourse with the out- side world had somewhat increased. Since Topography of the rugged region of central Chma m this region was located on the border between which the author travelled, often by narrow mountain Sichuan and Hubei provinces, an area char- trails, m his search for Metasequoia trees, as photo- acterized difficult and hazardous graphed from a NASA satell1te m 1975 He collected by roads, the type specimens from a tree just outside Modaoqi. murder and robbery occurred frequently. It The areas are irregular white snow-covered mountam was regarded as a forbidding place and was ranges The population of trees Professor Hsueh was seldom visited travellers. lookmg for when his time ran out, and he had to return by to Wanxian, is labelled "Metasequoia Valley" The On my trip, I set out from Wanxian and approximate range of Metasequoia is the hatched area stayed at Changtanjing for the night. My fel- bounded by tnangles. low travellers were several pedlars. While we chatted around a fire at night, the innkeeper came to give us a warning: "If you go any farther you will travel along a narrow valley cut by the Modaoqi River. Travel will become more dangerous and threatened with rob- bery, which often occurs at dangerous turns of the river. Travellers from both directions are robbed by being jammed together, or ’rounded up.’ Therefore, if you see no trav- ellers coming your way for a long time, it is very likely that a robbery has occurred ahead, 13

and you had better take care. Only a few days ering where I would take my lodging. It was ago we witnessed such an incident in this February 19th, and cold. The tree was located vicinity." The innkeeper then gave a vivid at the edge of the southern end of a small and horrible description of a murder. The street. In the tmlight nothing was discerni- poor pedlars, my fellow travellers, were very ble except the withered and yellowed appear- frightened. They dared not go any farther and ance of the whole tree. My excitement returned to Wanxian the next morning. As cooled. for me, I was bent on finding that colossal "Am I to bring back just some dried tree and collecting more specimens, so I reso- branches?" I asked myself. lutely continued my trip along the route The tree was gigantic; no one could have marked out by Mr. Wang, without any fear or climbed it. As I had no specific tools, I could hesitation. only throw stones at it. When the branches Finally, at dusk on the third day, I reached fell from the tree, I found, to my great sur- my destination safely. I set out immediately prise, that there were many yellow male to search for that colossal tree despite hun- cones and some female cones on the leafless ger, thirst, and fatigue, and without consid- branches. I jumped with joy and excitement.

The city of Wanxian, lookmg downstream along the Changyang (Yangtze) River All of the expedruons to the Meta- sequoia region began m this bustlmg port situated over 900 miles from the nver’s mouth. Photograph by the Amencan members of the 1980 Smo-Amencan Botamcal Expedition (B. Bartholomew, D E. Boufford, J L Luteyn, and S A Spongberg). 14 .

The weather being cold, many plants were fagot mixed with some Podocarpus nagi. The not yet in flower. Since I was short of money, wood was said to have been cut from a nearby I returned to Chungking city three days later. mountain. I took two twigs and pressed them as specimens. This indicated that P. nagi, The Second ’Irip to Modaoqi another primeval gymnosperm, occurred in the vicintiy. The second was in of the same trip May year, This time I took measurement of the its purpose being to collect the cone-bearing Metasequoia tree. It was 37 meters [about specimens in addition to ascertaining the 122 feet] high and 7 meters [about 23 feet] in natural distribution of Metasequoia and the girth, and still grew vigorously. flora of the On to region. my way Modaoqi, To ascertain the distribution of Metase- about half a day’s walk from my destination, quoia I interviewed many local people, but I came across a peasant carrying a bundle of none of them knew. The innkeeper did tell me that a whole stretch of shui-shan trees might be found at Xiahoe, in Lichuan county, Hubei province, about 50 kilometers [30 miles] away. As I had almost exhausted my travelling allowance, and as communication was extremely inconvenient, I had to give up my attempt to extend my trip to that place. Nevertheless, the innkeeper had provided an important clue for a more thoroughgoing exploration later. All I could do was-taking the original spot as a center-to make a reconnaissance within the area I could cover in one day. In a few days I had collected more than one hundred specimens. Two things impressed me deeply. One was that I came across whole stretches of Geas- trum sp. (an earthstar fungus) mixed with small stones of a similar shape, forming a peculiar landscape. The other thing that impressed me was an incident. Not even by the day before my departure had I given up on the possibility of making a reconnais- sance. At four in the afternoon of the last day, I met a traveller coming from the south- east and asked him where the shui-shan tree could be found. He told me that it could be got near a small village about 5 kilometers [3 miles] from where we were. Upon hearing this I almost broke into a to The tree m Modaoqi from which the author collected run, intending the type specimens of Metasequoia glyptostroboides. return to the inn before dark so that I might The shrme he mentions has been dismantled but is leave for Wanxian the next day. After trotting shown m the drawmg on page 1 of this issue of Arnoldia. a met and asked Photograph by the American members of the 1980 Smo- for while, I another peasant Amencan Botamcal ExpedW on. him how far it was to the village. (I can’t be 15

sure now, but it may have been Nanpin vil- seldom met outsiders, especially "intellec- lage in Lichuan county.) "Five kilometers," tuals" such as I was. My arrival aroused their he replied. Mountain people sometimes dif- curiosity. They surrounded me, making all fer considerably in their gauge of distance. sorts of inquiries. But I was anxious to see I was wavering as to whether to go or not. the Metasequoia trees. When I was told that If I should go, it was certain that I could not there were no such trees, I was very disap- have returned to the inn before dark and that pointed. However, I did not give up hope, and the innkeeper would worry. Then, too, I had asked the villagers to accompany me to already hired a man to carry the specimens make one last reconnaissance. There was, for me; we had agreed on the next morning indeed, no Metasequoia. I did collect some as the time for departure. I could not break specimens of Tsuga chinensis, however. my word! But finally I made up my mind to I intended to return to the inn in spite of make another reconnaissance for shui-shan. the dark night. However, the friendly vil- It was getting dark when I arrived at the lagers had already made arrangements for my small village. The villagers in their isolation food and lodging, and had warned me repeat-

Male cone (left) and female cones of Metasequoia glyptostroboides. The author made his tnps to secure male as well as female cones so that the new species could be given a botamcal descmption. Photographs by LeRoy C. Johnson. 16

"Metasequoia Valley," near Xiahoe, m 1980 Note the absence of natural vegetauon around the conspicuous Metase- quoia trees. Photograph by the Amencan members of the 1980 Smo-Amemcan Botamcal Expedition

Dracvmg, by an unknown artist, ofa 420-year-old Metasequoia tree m "Metasequoia Valley," Xiahoe commune. This is the oldest Metasequoia tree known 17

edly of the frequent robberies on the way, the geomantic nature of the place. Thus, it insisting on my leaving the next day, escort- was because of feudalistic superstition that ed by some local people. Yet I could hardly the tree had survived. Its age is estimated at fall asleep, thinking that I could not cause four hundred years. them so much trouble or break my word to With the advent of well regulated highway the hired carrier. And then I thought that in communication, the poor village of the for- the depth of the night there would be no mer days changed its aspect long ago. The "bandits," since there would be no travellers Metasequoia tree, which had survived the to rob. So, at two in the morning I awoke my ravages of time and is reputed to be a "living roommates, explaining to them the reason fossil," has not only persisted, but is being for my prompt departure, and left the vil- disseminated. Now, Metasequoia trees are lagers a letter of acknowledgment. Since the "settled" in many countries of the world. It door was locked, I could only jump over the is only natural that people, when admiring wall so as not to disturb others. In the moon- this species of primeval tree, should wonder light I passed through stretches of dark pines, about its original habitat and should wish to returning to the inn before dawn. That very know how it was discovered. day I left for Wanxian.

Geomancy Spared the Type Tree Modaoqi was a very small village, to the southeast of which stood the Chiyue Moun- tains. Its altitude was 1,744 meters [about 5,755 feet]. At the time it was in Wanxian county, Sichuan province. It was so called because of its situation at the source of the river. As modao in Chinese means "knife- grinding" and suggests sinistemess, the name was changed to Moudao, which means "truth- seeking" in Chinese. At present it is under the jurisdiction of Lichuan county, Hubei province. As the local people looked upon the Meta- sequoia as a sort of divine tree, they built a shrine beside it. Among the villagers there were quite a few traditions about the Metase- quoia. As a result, the villagers considered its fruit-bearing condition to be an indication of the yield of crops, and the withering of its twigs or branches a forecast of someone’s death. It was also rumored that, some time after the founding of the Kuomin Tang gov- ernment, some foreign missionaries who The pnze. one of the many specimens of Metasequoia that Hsueh collected were passing through the village were willing glyptostroboides Professor dunng his first tnp to Modaoqi This specimen, called a syn- to the tree for a sum of The buy big money. type, is m the Herbanum of the Arnold Arboretum. villagers refused to sell, however, because of Photograph by Damd E. Boufford See next page 18i

Metasequoia was discovered in 1941 at Modaoqi in Sichuan Province near the border with Hubei Province by T. Kan (Gan Duo), of the Department of Forestry of National Central University. Kan, however, did not collect or make specimens, and it was not until 1943 that C. Wang (Wang Zhang) of the Central Bureau of Forestry made the first collections of herbarium material. The tree was initially thought to be a form of Glyptostrobus lineatus (Poiret) Drude (syn.: G. pensilis (Staunton) Koch), but W. C. Cheng (Cheng Wanjun), of National Central University in Nanjing, realized that it represented a new genus. As a result, Cheng sent one of his assistants, C. J. Hsueh (Xue Jiru), to collect more herbarium material in February and May of 1946. In the fall of 1946, H. H. Hu (Hu Xiansu), then director of the Fan Memorial Institute of Biology, Beijing, received material from W. C. Cheng. Hu recognized that the newly discovered tree belonged to the genus Metasequoia, described in 1941 from Pliocene fossils by the Japanese botanist Shigeru Miki. Miki determined that certain fossils, which for nearly 100 years had been variously assigned to either Sequoia or Taxodium actually represented a new genus, which he named Metase- quoia. At the time, he did not realize that a living species of Metasequoia was still extant in south-central China.

. -Reprmted from an article by Bruce Bartholomew, David E. Boufford, and Stephen A. Spongberg m the Journal of the Amold Arboretum, vol. 64, no. 1 (1983). Used with permission. "Les Quatre Vents;’ a Far-Northern Garden

Francis H. Cabot

Seizing opportunities presented by cool, moist summers and deep winter snows has been the key to success in this northern outpost of Zone 4

A detailed map of the plant-hardiness zones of North America shows Zone 4 sweeping east from Lake Superior to the northern-tier states of New York and New England, thence north through New Brunswick to Newfoundland. As one follows the St. Lawrence River northeast from Montreal, Zone 4 appears to peter out on the north shore of the river a few miles north of the windswept Citadel of Quebec City, which overlooks the river at a point where it suddenly changes into an ever-widening inland sea that ultimatelv seven hundred miles to the northeast, becomes the Atlantic Ocean. The view to the northeast from the heights of Quebec over the thirty-mile-long Ile d’Orleans shows the southern shore of the St. Lawrence receding into the distance as it veers slightly to the east. The view of the north shore, on the other hand, is limited by Cap Tourmente, a stark, eighteen hundred-foot-high headland that curves sharply down to the roiled gray-green salt water. At this point, just beyond the northern tip of lle d’Orleans, the granite mass of the Laurentian Mountains, clothed in the spruce and fir mantle of the boreal forest, meets the salt-water coastline for the first time. From here on, these two elements-boreal forest and salt water-are the norm until one reaches the tundra and permafrost of Labrador. They dominate the ecological and horticultural life of the region. In the following article, Francis H. Cabot, who has been gardening in a small, far-northern outpost of Hardiness Zone 4, shares the horticultural insights he has gained over the course of the past twenty years.

The departure of the greater snow geese in North Carolina to its breeding grounds on late May and their return in early October Ellesmere and Baffin islands. The seven have special significance at La Malbaie, hundred-mile flight from the Middle Atlan- Charlevoix County, Quebec, for their migra- tic coast lasts two hundred hours. No one tion flights bracket precisely La Malbaie’s knows how long the second leg of the flight, gardening season, which is seven or eight from Cap Tourmente to Baffin Island, lasts. weeks shorter than those of zones 5 and 6. Virtually the last farming area of any size For six weeks each spring, the great flock along the northern shore of the St. Lawrence, stops to feed on the sedge Scirpus amencan- La Malbaie, thanks to the tempering effect us that grows along the brackish marshes of of its fourteen-mile-wide stretch of the river, the St. Lawrence River to the south of Cap Overleaf: Part of the terraced vegetable garden at "Les Tourmente. The flock, now estimated to Quatre Vents." The beds were constructed from trees number over 250,000, thanks to its protected killed by the spruce budworm Potatoes growm the bed m the foreground. The bed m the center contams sweet makes this one in its status, only stop 2,500- peas, that at the left, red currants. All photographs are mile journey from the coasts of Delaware and by the author

21

Astilbe, delphimums, Cimicifuga racemosa, and Achillea taygetea m the perenmal border.

Allium chnstophm (center), Primula flonndae, P. alpicola var luna, Lilium martagon var album, and Meconopsis betomcrfolia m bloom m shade garden Bed No 3, mth asulbe and Aruncus dioicus m the bed m the background. 22 ,

enjoys a microclimate of Zone 4 in an area of the quaking aspen (Populus tremuloides), of some twenty-five square miles surrounded which the French Canadians call les trem- by a relatively thin band of Zone 3 that soon bles, and several kinds of Amelanchier, whose becomes Zone 2. In the best of years La Mal- burnished-orange autumn foliage comple- baie enjoys both the cool summer evenings ments the tints of the red and sugar maples. and the fog and mists of a maritime climate Acer spicatum, the ubiquitous mountain (its portion of the St. Lawrence warms to 48 maple, needs to be constantly weeded out of degrees Fahrenheit in late summer), as well woodland areas, and A. pensylvanicum as the deep early snows of the boreal forest. (moosewood) to be encouraged. If the snows arrive before the ground freezes While Betula lutea (yellow birch) can be and last throughout the winter, the horticul- found in the forests, far and away the best of turist can proudly display plants that usually the native trees is Betula papyrifera, the thrive only in the Himalayas or in Scotland. paper birch. Invariably decorative at all stages In the worst of years, when the snows don’t of its life, the paper birch enchances its sur- come until it is too late, or even when an roundings whether it is used as a lawn spec- atypical winter thaw destroys the snow cov- imen, in the garden, or as an allee. I cannot er, it is another story, and a humbler horti- decide whether its bark is more beautiful in cultural outlook prevails. Most years the early youth, when it has fawn-like spots; in results are in between and are horticulturally adolescence, when it develops coral and satisfying. peach tints; or in maturity, when it takes on a pristine whiteness. In this northern setting the paper birch seems to be whiter than it is Limitations to Northern Gardening farther south and is hard to improve upon in The principal drawback to gardening in the landscape. northern zones is the limited choice one has Many non-native species love the North of trees and shrubs, yet this limitation does also. The Lombardy poplar (Populus nigra simplify the landscaper’s task. At La Malbaie ’Italica’) achieves impressive heights at La the wisest course often has been to use native Malbaie, for example, as does the Carolina (or naturalized) species for the backbone of poplar (P. canadensis). The former’s relative- the landscape plan. On the whole, the native ly short life span and pruning needs at matu- species are a useful and appealing lot. rity give one pause, however, but, used as an The conifers are represented by the spruces accent or as a specimen, there is nothing Picea abies, P. glauca, and P. nigra; the bal- quite so effective or so appropriate for French sam fir (Abies balsamea~; the American Canada. The Carolina poplar is far less larch (Larix laricina); and the pines Pinus demanding, it would appear, and longer lived. strobus, P. resinosa, and P. banksiana. Thuja Undistinguished in its early years, the Caro- occidentalis thrives near the shores of the St. lina poplar achieves dignity and grace with Lawrence, growing to majestic proportions, age and can be a useful part of the landscape. and Juniperus communis var. depressaa The Amur maple (Acer ginnala), the ear- abounds. There are a few shoreline speci- liest maple to take on color in the autumn, mens of juniperus horizontalis, and Taxus is also useful. Hardy to Zone 2, it forms a canadensis is prevalent in the forest. graceful, medium-sized tree but requires Sorbus americana, the mountain ash, is pruning and thinning as it matures. It is everywhere, enlivening the late-summer and worth the trouble, though, for its brilliant autumn landscape with its abundant clumps scarlet foliage in September. Along one of the of red berries. There are groves and hillsides farm roads we have planted an allee of Amur 23

maple that is becoming increasingly colorful A Botanical Ramble in the Boreal Forest as the years pass. While a few of the hardier species of Malus An English friend of mine, walking through can withstand the winter temperatures of the woodlands surroundmg the gardens at La Zone 4, the diversity of blossom color avail- Malbaie, was struck by the number of great able in Zone 5 and points south is missing, botanists and plant hunters who were com- as are the flowering cherries. Apple trees are memorated in the flora. Not only the great close to their northern limits at La Malbaie. Linnaeus in the twinflower (Linnaea boreal- They grow very slowly there, but it is worth is) that carpets the woodlands, but John the wait for their crisp and flavorful fruit. Goodyer in the three species of Goodyera ’Fameuse’ is one of our favorite varieties. that abound (Goodyera oblongifolia, G. Plums do beautifully at La Malbaie and seem repens, and G. tesselata~, Sir Joseph Banks in healthier and more productive there than the stands of Pinus banksiana, and John Bar- they do in more-southern climes. Our crop tram in the serviceberry, Amelanchier bar- of ’Mirabelle’ and other small varieties of tramiana. plum is abundant. The serviceberries are edible but not par- The Zone 3 and Zone 4 climatic limita- ticularly interesting. The best of the lot are tions on trees are felt equally severely among the oval fruits of Amelanchier bartramiana, shrubs, but there are sufficient species of which are larger and more succulent than Rosa Syringa, Spiraea, Berberis, Viburnum, other varieties. It seems strange that this Philadelphus, Neillia, Lonicera, Caragana, most garden-worthy species, which is found Cornus, and the like to make do and to fur- in the Laurentians and at the higher eleva- nish the garden adequately. tions of the Appalachians, is not more widely It is when one comes to herbaceous plants used in horticulture. The white flowers are that one forgets about the lack of diversity in the largest of the genus, comparable to those trees and shrubs and begins to chortle over of Potentilla fruticosa and borne in a very the salubrious northern maritime climate. similar fashion. The leaves, as they emerge Aside from the Pacific Northwest, it has to in the spring, vary from bronze to pale green be the best spot on the continent to raise and turn to burnished orange in late Septem- perennials. While the Atlantic coast from ber. Amelanchier bartramiana is a shrub Maine north shares a comparably cool and rather than a small tree. It grows slowly, damp summer climate, it does not enjoy the attaining a maximum height of five to six heavy snow cover that is characteristic of the feet, and is compact and stoloniferous. The Laurentians. largest specimen I have seen was five feet in The sensible gardener at some point stops diameter and had a most sympathetic and fightmg his climatic limitations and sticks slightly irregular outline such as one finds in to what will do well for him in the habitats specimens of ancient English box. Why, and microclimates that he is able to create. then, is it not proffered by the trade? The joy of gardemng in a northern maritime The answer lies in the propagation records garden is that it widens his horizons consid- of the Arnold Arboretum, which show that erably. Of course, those plants that depend every effort to propagate Amelanchier bar- on heat units to live up to their promise will tramiana from seed or cuttings over the not fare so well. But then life is a series of years has failed. It can be mtroduced suc- compromises; the gardener will have to cessfully into the garden by transplanting choose between the delights of harvesting small, stoloniferous offshoots severed from sweet corn and raising exotic primulas. the main root in early spring. Princeton Nur- 24

Peomes, thahctrum, Amsonia tabemaemontana, and Aruncus dioicus hne the Goose Allee. The hedge is hawthorn. series has successfully grafted bud-wood of nuses, puffballs, and a wonderful nutty-fla- branches cut in the autumn to scions of vored, bright-orange parasitic fungus, Amelanchier canadensis, and it should be Hyphomyces lactifluorum. An experienced propagable from root cuttings, given its sto- mycologist could probably identify scores of loniferous nature. [Alfred J. Fordham descnbes other edible varieties among the hundreds the technique for doing so in Arnoldia, vol- that proliferate in the woods. We are slowly ume 28, numbers 4 and 5 (May 17, 1968), expanding the repertoire, and cautiously, too, pages 36 to 40]. I hope it will find its way into for there is little room for error. more widespread horticultural use. In July, August, and September our every spare moment is taken up with the harvest- ing and preserving of berries, from the delect- able des far to of the Boreal Forest fraises champs (so superior Gustatory Delights the fraises des bois of France), through all A botanical ramble through the boreal forest the garden varieties of strawberries, raspber- in late summer has many gustatory rewards ries, gooseberries, and currants to the low- as well. One can nibble the tiny, delicious, bush blueberries (Vaccinium angustifolium). and fragrant creeping snowberry (Chiogenes Berries, in general, have a superlative flavor [or Gaultheria] hispidula/, and the noisette and spoil one for their counterparts down (Corylus americana/, the local hazelnut. One south. can bring home baskets of mushrooms: If one hikes to the higher elevations of the chanterelles, cepes, russulas, clavarias, copri- Laurentians one finds Rubus chamaemorus, 25

the exotic, buff-colored cloudberry, so prized northern flank of the setting. To the south by the Scandinavians, along with mountain- one looks down across the bay of La Malbaie tops of the brilliant, scarlet mountain cran- to the villages on the far shore. berry (Vaccinium vitis-idaea var. minus/, the We use Pinus sylvestris, P. cembra, and P. lingonberry of Sweden and the Preisselbeer- mugo extensively, and Tsuga canadensis en of Germany. The French call them airelles (Canada hemlock) can be established in shel- rouges, and the local Indians call them ataca. tered spots. Other conifers grow well, although Whatever they are called, they make the best we have not tried a wide variety of species, of all tart preserves to accompany game, and probably because the surrounding forests are their glossy, dark-green, prostrate mats are of spruce and fir. It would be interesting to an exhilarating sight, particularly when they establish an experimental planting of coni- are interspersed with shrubby Cladonia fers to determine which of them would grow. lichens in the cracks of a lichen-covered Unfortunately, nurseries in Quebec have a rock-the perfect setting for the scarlet ber- limited number of species from which to ries. Inedible or less-edible berries abound as choose. Thuja is invaluable for hedging and well: the gray-blue of Vaccinium uliginosum shaping, as is the local Crataegus, C. foetida. var. alpinum, Empetrum nigrum, the black The native species can easily be dug from the and dark-purple crowberry, and, of course, surrounding woodlands and incorporated into the bright-red bunchberry (Cornus canaden- the landscape plan. There is ample material sis/, which sheets the ground in the right with which to create a framework and back- habitats. ground that fits in well with the natural set- ting. An entrance allee of Lombardy poplars a Cultivated Landscape at La Developing in 1926 is now in its last of Malbaie planted stages decay. What was once a dramatic feature of Our garden and house were built over the the landscape has become a spotty and past sixty years on a part of a seigniory grant- decrepit line of hangers-on. Every year their ed in 1653 by Louis XIV to Jean Bourdon, remains are cut down and removed, and Surveyor General of the Colony of New native paper birches planted between the France. The lay of the land has dictated the stumps so that in time a new and more per- development of the landscape from quite manent allee will take the old one’s place. modest beginnings to a series of gardens that A lilac hedge bordering the entrance allee now cover approximately twenty acres. The of Lombardy poplars was installed to frame cultivated landscape is still growing some- a sloping vegetable garden that runs down to what but is approaching its logical limits. a garden shed with breezeway and weather- The growth and development of the garden vane, and on to a small greenhouse. The east- make interesting study, especially because ern view is dominated by the St. Lawrence, no professional landscaping expertise has whose southern shore, some fourteen miles been brought to bear on the matter. In 1926, away, forms the horizon, and where indis- a house was built in a bare field and a small tinct villages glint in the settmg sun. perennial garden created to the west, where One’s first hesitant efforts to create a gar- the view leads over a meadow, across a den are often obliterated as one learns more stream-filled gully, to pastures, distant about garden design or about the shortcom- woodlands, and the muted cordillera of the ings of a site, or if one takes to heart the Laurentians. A steep, wooded hillside that counsel of those favored with architectural merges into sloping pastures marks the wisdom. This has been my experience. At La 26

Malbaie, aggressive winds sweeping down from the 1930s. It is only when one walks from the Laurentians (we have called the down the terrace steps towards the gardens place "Les Quatre Vents") dictated the con- that one realizes there are cross axes that struction of tall wooden windscreens. An lure one away from the tapis vert. architect, in the process of adding a guest wing to the house, installed a terrace with reflection where the had been pool garden A Tbur of the Garden and decreed a "tapis vert"-a long, narrow carpet of lawn running from the terrace to The new developments in the garden are best the edge of the stream gully. The tapis vert explored in tour sequence. An entrance in was flanked by a sunken blue garden and a the hawthorn hedge, to the right, or north, of raised white garden inspired by Vita Sack- the tapis vert, leads into a narrow perennial ville-West’s writings, centered on an oval lily allee flanked by a matching hawthorn hedge pool. A double hedge of hawthorn and bar- that finally is getting tall enough to give the berry in due course replaced the windscreens desired tunnel effect. The allee, known as on both sides of the tapis vert, and the west- Goose Allee because of frequent visitations ern view now flowed gracefully from the by the denizens of the lake at its foot, runs house to the mountains, unfettered by any parallel to the tapis vert and is about one horticultural distractions, the shallow reflec- hundred feet long. At its upper end is a seat tion pool enhancing the spectacular sunsets from which the visitor can enjoy the many of that northern clime. tall and spiky perennials planted to enhance It wasn’t until some twelve years after I the tunnel effect. The "tunnel" directs the had inherited the house and garden in La eye to a conveniently centered paper birch, Malbaie, when I had been increasingly exposed by the water’s edge, which ends the vista. to the gardens of Britain and the myriad of The visitor is still not conscious that a lake landscaping devices and plant relationships is there. they display, that the grounds’ potential as a The Goose Allee beds are terminated by horticultural tour de force became apparent. clumps of Daphne mezereum underplanted It soon became clear to me that a number of with Primula abchasica, P. vulgaris ssp. sib- gardens could be added to the basic frame- thorpii, and Scilla sibirica, a felicitous com- work, gardens that would make the whole bination of purple, mauve, and blue to start more interesting, and result in a more diverse off the season. They are followed closely by horticultural experience. There was space in Doronicum interplanted with Brunnera and, which to expand, an unlimited supply of ultimately, a succession of Aconitum, Cim- water from the stream, and a variety of dif- icifuga, Delphinium, Ligularia, Rudbeckia ferent habitats for new plants. Over a period maxima, and Thalictium providing the tall of ten years, I have filled the space and hab- accents amid Astrantia, Centaurea, Penste- itats one step at a time, without any partic- mon, Paeonia, and Trollius in variety, among ular forethought, but each step leading to the others. next. The gardens developed during this ten- A high Thuja tunnel leads off to the right year period supplement the good original and draws one across the Goose Allee, into landscape setting. Looking across the reflec- a dark-green channel to an alcove adorned tion pool and tapis vert, between the white with a statue of one of the Four Seasons. The and rose gardens and the frame of the haw- statue looks west through a rondel of Thuja, thorn and barberry hedges, toward the moun- over a millstone converted to a sundial, then tains, one is not conscious of much change down to the lake and across to a somewhat 27

battered bust of Antonia, the half-sister of and happily cross the stream over the Chinese Augustus Caesar, arising from a clump of bridge and meander around the lake, where Clematis recta and other perennials on the they catch a good view back over the lake far shore. In the background, at the edge of and up the tapis vert towards the house. the fields, a Lombardy poplar (a kind of living The visitor then crosses the top of a high obelisk), has been planted as an exclamation dam that separates the lake from the deep point. The Thuja rondel is set within a Thu- woodland ravine into which the stream falls. ja-and-hawthorn square, with Betula papyr- The banks of the dam have been planted with ifera and Acer ginnala planted in the spaces the white-flowered form of fireweed, Epilob- between the two. At this first glimpse of the mm angustifolium forma albiflorum. By lake, one’s curiosity is aroused. avoiding the tapis vert after crossing the dam Turning away from the vista through the and bearing right, along the edge of the rondel and continuing along the Thuja tun- woods, the visitor enters a new world. nel, the visitor suddenly emerges and finds The garden setting not only had fine views himself crossing the middle of a string of six of the mountain and the river, but also had rectangular lily pools, with the water cascad- a handsome grove of young spruce when the ing and stepping down the gentle slope to the site was first developed. My father could lake. The pools are flanked by Thuja planted remember jumping over them as a boy in a quincunx pattern and by occasional around the turn of the century. Thirty years matching clumps of three decorative rhu- later, they were indeed a mossy fairyland. barbs. Looking to the right, the visitor looks One of the natural tragedies of our part of up the watercourse to a dolphin at its head, French Canada has been the destruction, by the source of the water. the spruce budworm, of the spruce and bal- Crossing over the watercourse, one emerges sam fir forests that constitute 90 percent of into a longer, broader vista, again flanked by the conifer forest. It has been a nightmare to Thuja in a quincunx pattern whose size watch beloved, dark-green hills turn, first increases as one descends the slope. The vis- brown, then a ghostly gray. It has taken fif- ta slopes from the garage court down through teen years to do the killing, and millions of a rhubarb allee to the north end of the lake. acres have lost these two species. Our place A Chinese bridge (made of plywood) that was no exception, and since our spruce had crosses the stream entering the lake in the been allowed to grow to climax-forest matu- distance is the focus this time, and draws one rity, the loss is all the worse, for there is to the lake’s edge, which is packed with nat- nothing but brambles and alder scrub left to uralized perennials situated so as not to cover the Cornus canadensis, Linnaea bore- obscure any vistas. As one skirts the shore, alis, Pyrola, and other delights. Happily, the ducks and geese keeping a wary distance, regeneration has started with a vengeance, there are agreeable glimpses through branch- and the budworm cycle is terminating, but es and flowers of the bust of Antonia. it will be twenty-five years before the coun- The visitor now has two options. The tryside regains its former character. energetic visitor will bear right and walk The spruce budworm wreaked its havoc on through a ten-acre stream garden planted to the grove of spruce near the garden. Slowly native trees and hardy shrubs. but surely the trees died. As they were Since the stream garden is planted with knocked down by the wind and the light was young trees, and it requires some imagina- let in, thickets of wild raspberries and eld- tion to visualize what it will be like in the erberries ensued, and the grove became year 2000, most visitors are spared the hike impenetrable. 28

The Shade Beds combination of Lilium martagon var. album and Meconopsis betonicifolia interspersed It was at this that three shade- point large with the yellow form of Primula alpicola. garden beds the of the woodland along edge But then two specimens of Acer spicatum were made out of what had been a mixture that had shade after the of weeds and lawn. The which provided adequate scruffy beds, loss of an ancient and have been and several spruce suddenly replanted rearranged mexplicably died, and the sunlight streamed times over, now contain a profusion of shade- in. In 1985, this bed was converted into a loving plants that are at their best in early grass garden with varieties of Miscanthus, when the are "tak- August, perennial gardens Molina, Panicum, Pennisetum, and other ing a breather" before the autumn show grasses, interspersed with mauve delphin- Astilbes have been used to a begins. great iums. The Primula alpicola is still there, extent, large and small, and early and late along with Trollius ’Alabaster’, Astrantia car- bloomers. In one of the beds have been they niolica var. rubra, Allium Pri- intermixed with three of christophii, species Aruncus, mula florindae, and Primula sikkimensis. four species of Cimici f uga, and the native red The meconopsis appear to be grateful that and white baneberries (Actaea rubra and A. they were moved deeper into the woodland. pachypoda, respectively). In addition, lilies have been used liberally. The bed is divided roughly in half, and when everything works The Woodland Garden as it should one half is filled with white and The shade beds lead toward a grassy clearing astilbes which Actaea rub- pale-pink among at the end of the main perennial allee. The ra and red lilies are thrusting their stalks, far side of the clearing is one of the entrances while the other half is the Astilbe reverse, to the woodland garden and is the spot where ’Fanal’ forming a background for Actaea the first experimental species of Primula and pachypoda, Campanula persicifolia ’Alba; Soldanella were tried, with such success that and a white In and the bed is lily. June July moderation was soon thrown to the winds. a mass of green and little else, but it foliage For some reason, the spruce in this corner is worth the succession of bloom forgoing have withstood the ravages of the budworm. and for the moment of waiting glory. The earth consists of rich forest duff, rich A second bed consists of softer colors: the enough to stick one’s arm in to the elbow. mauve and astilbes rose-pink interspersed The experimental plantings grew so rapidly with a the buff tints of Lil- pale-yellow lily, that they could be divided twice, and some- ium Lilium marhan, rose-pink martagon, times three times, in the summer. In the the white form of Meconopsis betonicifolia, spring the soldanellas bloomed profusely, and the occasional blue of spikes Campanula something that doesn’t happen at points persicifolia. This bed has been changed again south. and again, principally to take out lilies that, Clearly, this wretched and impenetrable to the turned despite representations contrary, woodland held promise, and over a six-year out to be a color I have a difficult orange, period the dead trees were felled and the time with in the garden. The interplanting brambles and other trash cleaned out. It of the astilbes has also involved considerable turned out that there was an underplanting so that the succession and combi- thought, of young Betula papyrifera, Amelanchier nation of color works to advantage. canadensis, Sorbus, and Acer spicatum. These The third bed has undergone the most were thinned out and transplanted so as to changes of all. In 1984, it was a successful spread their valuable shade throughout the 29

rest of the copse. To provide the essential Planting the Ravine moisture, a two-inch plastic pipe was laid The stream and ravine curve in an arc of from the stream to the edge of the woodland, about While I was with the result that there now are four arti- seventy degrees. visiting the beautiful Glen at Wakehurst ficial streams that moisten the woodland Himalayan Place, its director, Tony Schilling, remarked floor, and a series of twenty-five overhead that he hoped someday to install a Nepalese sprinklers that are operated for an hour or across the top of the glen so that vis- two each evening. bridge itors could look down at the rhododendrons Not surprismgly, primulas love the habi- and other It wasn’t tat. We have been able over time to increase Himalayan species. long before that excellent idea manifested itself the number of species and the size of the over the ravine at La Malbaie in the form of plantings to where the little streams are now two rope from a sketch in Roy planted to great quantities of a given species bridges copied Lancaster’s Plant Hunting in Nepal. The and grouped by color variations where appro- are at about sixty feet over priate. For example, one stream is flanked bridges suspended the streambed and are just over one hundred with Primula sonchifolia and Primula rosea, feet mixed. Well over one hundred species are long. The ravine has been with grown, including a number not often seen in planted mostly eastern North America. Primula vialii flour- large-leaved species that will provide easily noticeable textural for the nervous ishes, as does Primula nutans (now called appeal cast down them. The stream Primula flaccida). Primula nutans is truly glances upon bed has been lined with Petasites perennial and has the most captivating scent. japonicus var. since there is room for it to Primula reidii var. williamsii, and Primula gigantea, and since it seems that it will reinii from Japan, grow well along with Pri- romp unlikely climb the banks. mula reptans, the smallest of the lot, which up steep five different of behaves like a ground cover, its large purple Large colonies of species laid out on the lower blossoms resting on its tiny, prostrate leaves. Rodgersia have been Primula sapphirina and Primula primulina slopes and ravine floor, and Bergenia cordi- from to also thrive, as do the petiolarids (with the folia sweeps down diagonally top exception of Primula aureata, which has yet bottom. The cirque formed by the arc has to be wintered over successfully). Of course, been planted to Rhododendron yakusiman- um the small the easy Auriculata, Candelabra, Efarinose, exclusively. When plants and Sikkimensis sections flourish, so much mature, it should be a splendid sight if there A manicata so that we have just barely been able to find is not too much shade. Gunnera places for the seedlings by diligently clearing has survived its first (and rather bad) winter new areas each summer. This work has prov- under extensive wraps, and it promises to fill a between the two en exciting because of the nearby ravine, hollow in the ravine floor which is now cleared and being planted, and bridges with its dramatic foliage. which affords an especially protected habi- Not that everything is perfect, of course. tat-one reason the petiolarids and other The hillside of Primula pulverulenta ’Bartley plants collected in the Himalayas survive. Strain’ seedlings, planted so carefully last The steep ravine slopes and deep snow cover summer on a very steep slope, succumbed to are not unlike their habitats at home. the lack of snow cover, and to the ice and What is good for Primula is good for Gen- subsequent erosion in that unfortunate, tiana, Cardamine, Glaucidium, Rodgersia, windswept spot. But the hillside of Mecon- and others of their ilk. opsis betonicifolia intermingled with Pri- 30

mula alpicola var. luna x var. alba (which great many spruce logs and have elected to we have dubbed "Ivory Tower," for want of a use some of them to convert the vegetable better name) is in fine shape, as are the plant- garden into a series of terraced beds, building ings of Heracleum on a steep and sunny spot. one bed a year. After six years, the project is (We grow some ten species of Meconopsis in finished, and the vegetable garden, now one part or another of the woodland.)/ mixed with annuals and perennials for cut- The next step will be to embellish these ting, has a new lease on life. ravine slopes with bulbs. One slope is yearn- Here again, there was no detailed plan ing for a mass of Erythronium revolutum when we started. The configuration of the intermingled with Anemone blanda ’Atro- beds was dictated by the degree to which the caerulea’ to sweep over its brow. The other land fell away on a given slope. As a conse- slope would prefer Corydalis ambigua /or quence, no two beds are the same, and each perhaps Corydalis cashmeriana or Corydalis bed’s shape conforms to the underlying con- solida var. transylvanica/. Woodland garden- tours. Some of the terraces within the beds ing is heady stuff and, in this easy habitat, as are small and some quite large, but the whole good a way as any to spend the time. provides an ample area in which to grow just A plantsman’s inspection of the woodland about everything that could be grown. In garden-or sous-bois, as it is called locally- time, as we figure out how best to use and can take longer than is necessary at this junc- combine the vegetables, annuals, and per- ture, but one of the paths leads out of the ennials within it, this garden should yield a woodland, past a gazebo where one of the lot of horticultural pleasure for all con- rope bridges terminates, and to where all cerned. Terracing is probably the only logical orange plants of whatever genus (Lilium, solution to the problems of gardening on a Primula, Trollius, Ligularia, and others) have slope. been exiled. They bloom harmoniously, There is nowhere left to walk but back without competition, against the green back- towards the house through the lilac hedge ground. (A similar corner is filled with every- enclosing the vegetable garden. There is a thing magenta.) The path joins a carriage small meadow garden here, crisscrossed and drive that skirts a snake fence with good formed into a series of more or less geometri- views of the bay and the river until it comes cally shaped panels by a series of paths. Every to the vegetable garden. autumn a hefty contingent of bulbs is plant- ed within it, and every summer new peren- nials and biennials are added to it in The Terraced Vegetable Garden hopes they will take hold and establish colonies. It In its original state, the vegetable garden con- is a long but worthwhile process. A mosaic sisted of two long beds bracketing the path of Crocus chrysanthus in sweeps of blue, from the house down to the weathervane. cream, and white appears in early May and The beds sloped both downwards and side- expands as the years go by. They are followed ways and were not particularly satisfactory. by daffodils, which in turn are underplanted The spruce trees killed by the spruce bud- with Scilla, Muscari, Puschkinia, and Chion- worms came in very useful at this juncture, odoxa, each panel with its own combination since one can salvage the value of the timber of colors. There is a moment when pissenlits and clear the land of what promises to (dandelions) seem to be the only plant in evi- become an impenetrable barrier of fallen dence, but lupines soon assert themselves trees if one cuts within three years of the and put on a grand show m June and July, trees’ death. Thus, we have had access to a along with Oriental poppies, Achillea ptar- 31

mica, Filipendula, Campanula, and Rud- could see the black tips of their wings, bisect- beckia. The trick is to extend the flowering ed the blue sky, framed for an instant in that season until the meadow is cut in early Sep- incomparable setting. tember. Later that afternoon the weather suddenly deteriorated and became threatening. I had been planting the last of the bulbs in the Return of the Snow Geese woodland garden when it began to snow Perhaps the happiest time of the garden cycle quite heavily. In a matter of minutes snow in La Malbaie is early October-the horti- covered the ground, and I could imagine what cultural evensong, when the season is over it was like to be a petiolarid primula on the and our work done. The days are brisk, the flanks of a Himal. The storm was sufficiently nights cold, and the colors of the foliage intense to cut visibility way down. All I exhilaratmg. The garden has been put to bed, could see were swirling snowflakes. Again I the bulbs planted, and notes taken on what heard snow geese, only this time there were to move the next spring to improve a given thousands of them, and I rushed to the edge planting. It is a time to cut trails, to build of the woodland to try and see them. The bridges, to make lakes, as well as a time to snow was so thick by this time that a tree sketch plans and ideas for the future. twenty feet away was barely visible and I One crystal-clear October morning in 1984, despaired of catching even a glimpse of them I had time to wander at length along the var- when, suddenly, from all sides, hundreds of ious trails and to explore rarely visited geese descended out of the clouds-on the byways that had been cleared. Along one front lawn, in the woodland garden, on the byway, down a gently sloping, moist gully, tapis vert, and in the fields beyond, which the forest had escaped any damage, and there were just becoming visible again. I wondered were still many fine ancient specimens of red if this was what heaven was like. It ended as and white pine. Partridge, woodcock, and quickly as it had begun. The snow geese, hare flushed from along the trail, the mosses appropriately wrapped in the swirling cloud and mushrooms glistened, and the whorls of of snow, and over their apparent disorienta- Cornus canadensis were deep purple. As the tion, lifted off the ground (had they ever trail came to a cul-de-sac and I was beginning touched it?) and disappeared. to explore how it could be blazed on, the What did it mean, I wondered. Did it mean unmistakable sound of the snow geese call- it was time to stop gardening for that season, ing in flight broke the stillness. I hurriedly to reflect upon the mysteries of Nature and struggled out of the thicket and back up the gather strength for the coming spring, when trail to a point where I could look up at the the snow geese would be heading north, and sky. There was only one opening, a small the gardening cycle would begin again? Was one, and it was framed by the autumn colors it a message confirming that we were fortu- of the maples, Amelanchier, and paper birch. nate in our earthly paradise, that we could Disappointed that I couldn’t see the geese do worse than to keep gardening in that through that small, baroque window of blue northern woodland until our day was done? sky, I was nonetheless struck by its beauty, a perfection worthy of Tiepolo, an intensity and combination of colors, backlighted by Francis H. Cabot, of Cold Sprmg, New York, and La Malbaie, Quebec, describes himself as a horticultural that were I at it the sun, breathtaking. gazed enthusiast. He is treasurer of the Amencan Rock Gar- for a while and was about to stop when five den Society and past chairman of the board of managers snow geese, m formation and flying so low I of the New York Botamcal Garden. Plant Hardiness-Zone Maps Donald Wyman Harrison L. Flint

Existing hardiness-zone maps are valuable, but using them requires knowing how they differ one from the other, and how "hardiness" has been defined

Plant hardiness-zone maps have been a valu- differences in lowest monthly mean temper- able aid to those interested in predicting the ature. adaptability of plants to specific climatic A few years later, the prototype of the pres- areas. Most are isotherm maps of geograph- ent Arnold Arboretum hardiness map was ical regions, based upon average annual min- prepared by Donald Wyman. It included the imum temperatures experienced at certain entire United States and was first published weather stations over some period of years. in his book Hedges, Screens and Wind- Many hardiness-zone maps are available- breaks, in 1938. This map was based on some cover small areas such as individual average annual minimum temperatures for states, while others encompass entire coun- the years 1895 to 1935, as published in the tries. Unfortunately, many do not agree in Atlas of American Agriculture, USDA, in their numbering schemes-so zone numbers 1936. A modification appeared in the second assigned to individual plant species cannot edition of Rehder’s Manual of Cultivated be used in referring to all of the existing Trees and Shrubs, in 1940. In that book, the maps. In most cases they can be related only southernmost part of the United States was to the map used in assigning them. The two not included and much of Canada was added, most widely used in this country are the in keeping with the manual’s scope. Arnold Arboretum hardiness map and the The Arnold Arboretum Hardiness Zone "Plant Hardiness Zone Map" prepared by the Map in use since 1949 mcludes the entire Agricultural Research Service, United States United States (except for Alaska and Hawaii) Department of Agriculture (USDA). and southern Canada. This was first pub- lished in Wyman’s Shrubs and Vines for American and in his Arnold Arboretum Hardiness Gardens, republished Maps books Trees for American Gardens (1951 and The original map prepared at the Arnold 1965), The Arnold Arboretum Garden Book Arboretum was published in the first edition (1954), Ground Cover Plants (1956), and The (1927) of Manual of Cultivated Trees and Saturday Morning Gardener (1962). Shrubs, by Alfred Rehder. In this map, the The Arnold Arboretum map was revised in United States and southern Canada (except 1967 and again in 1971. The 1967 version was for southern Florida) were divided into eight published in Arnoldia, while the 1971 ver- zones characterized by 5-Fahrenheit-degree sion, which is the one published on the mside front cover of this issue of Amoldia, Reprmted, with slight modifications, from Amoldia, appeared in Wyman’s Gardening Encyclo- Vol. 27, No. 6, pages 53-56 (June 30, 1967). pedia. 33

The 1967 map differed from the previous ble. Unfortunately for the casual user, these version only in that hardiness-zone lines inconsistencies are small enough to be over- were redrawn to conform to more recent looked, and in several instances, writers have weather data. The zone-numbering system erroneously applied the hardiness-zone des- was unchanged-so zone numbers applied to ignations of Rehder to the USDA map. The specific plants in Rehder’s manual and other table shows the relationship between the two publications by the Arnold Arboretum staff numbering systems. could be used with the newer map just as well as with its predecessor. Local Hardiness-Zone Maps More-detailed plant-hardiness-zone maps have been for certain states and locah- USDA Plant Hardiness-Zone Map prepared ties. The total area covered by such maps is This map (Figure 1) was issued in 1960 as still rather small. Fortunately some detailed Miscellaneous Publication No. 814 of the maps use the same zone-numbering system Agricultural Research Service, USDA. It con- as the larger, more general maps. A good tains uniform zones of 10 Fahrenheit degrees, example is a hardiness-zone map of the state and sub-zones of 5 Fahrenheit degrees. Since of Vermont (Hopp and Lautzenheiser, 1966). the Arnold Arboretum map uses zones of dif- This map uses the same zone-numbering ferent ranges (5, 10, or 15 Fahrenheit degrees), system as the USDA map, but is based upon discrepancies between the two are inevita- a larger number of weather stations in Ver-

Figure 1. The U S. Department of Agnculture’s plant hardmess zones m the contermmous Umted States and southern Canada. 34

mont, so zone lines have been drawn in more Relationship Between the Arnold Arboretum and detail than in the larger map. As more areas USDA Hardiness-Zone Numbering Systems are mapped in greater detail in this way, har- diness-zone maps will become increasingly useful.

Canadian Plant Hardiness Map The Canadian Plant Hardiness Map, released in 1967 by the Canada Department of Agri- culture, covers all but the far-northern parts of Canada. This map represented a new approach in that it was an attempt to describe hardiness zones in terms of the whole com- plex of environmental factors that contrib- ute to severity of climate, rather than in terms of a single factor such as average annual minimum temperature. To as great an extent as possible, actual observations of plant adaptability played a part in describing hardiness zones. Direct comparisons between this map and those prepared in the United States are not valid, because of the different criteria used in describing hardiness zones.

The Future

It appears that the business of preparing har- diness-zone maps and assigning plants to their proper zones is still in the experimental stage. Hopefully, we may eventually see wide adoption of a single hardiness-zone map for the United States, for North America, or Reference even for the northern hemisphere. If this is someday accomplished, the problem of Hopp, R.J., and R.E. Lautzenheiser, 1966. Extreme Wm- assigning realistic zone numbers to specific ter Temperatures m Vermont. Umversity of Ver- plants will still remain. The ability to match mont Agricultural Experiment Station Bulletin 648, 19 pages. plants with zones over wide regions will be the ultimate test of any map, and many more careful observations will to have be made Donald Wyman was horticulturist for the Arnold Arbor- before this can be done with most of our etum from 1936 to 1970, during which period he was present trees and shrubs. Meanwhile, the editor of Amoldia. His Gardemng Encyclopedia is a standard reference for and horticulturists. existing maps will continue to be useful. But gardeners Harnson L. Flint, a former staff member of the Arnold to use them most we must recog- effectively Arboretum, is professor of horticulture at Purdue Um- nize their differences and use zone published versity. Among his best-known publications is Land- references only with the right map. scape Plants for Eastern North Amenca ~Wiley, 1983). BOTANY: THE STATE OF THE ART

What Determines a Plant’s Cold Hardiness? John W Einset

Cold-tolerance depends upon a plant’s ability to keep water from leaving its cells and freezing, which severely dehydrates the cells

Banana plantations along Cape Cod, orange mits them to survive at temperatures as low groves in the Berkshire Mountains, tropical as minus 75 degrees Fahrenheit. landscaping in New England! Unlikely as What causes such wide variations in the these images seem, they are not entirely out sensitivity of plants to cold? As a conse- of the realm of possibility, especially if more quence of natural selection, plants native to can be learned about the basic mechanisms a particular hardiness zone are adapted to the that govem the tolerance of plants to cold. In temperature extremes that occur in their fact, goals less spectacular than these, yet environment. Remove them from that envi- still highly significant, are achievable in the ronment, and they may or may not survive. near future because both our understanding For example, banana plants kept at low, but of plant physiology and our ability to manip- nonfreezing, temperatures suffer an imbal- ulate plant cold hardiness have improved. ance in their metabolism that kills their cells Without question, resistance to low tem- and causes brown necrotic streaks to appear peratures is a major factor determining the on fruits. Or, hardier plants might be killed geographic distribution of plant species. So- by frost that occurs during their normal called chillmg-sensitive plants, such as the period of vegetative growth. Several cultivars tropical banana and the semitropical avoca- of rhododendrons and azaleas that are grown do, can be severely injured or even killed by successfully in Georgia and the Carolinas, long-term exposure to temperatures (50 for instance, are killed by late frosts when degrees Fahrenheit, for example) that are well they are transplanted to New England only above freezing. By contrast, chilling-resis- because their tender vegetative buds initiate tant plants, such as garden peas and potatoes, growth too early in spring. Other plants fail survive brief periods of frost but are killed to survive because late-summer frosts kill when freezing conditions continue for more vegetative shoots before they become accli- than about four hours. Cold-hardy plants, on mated to cold temperatures. the other hand, tolerate extended periods of During any given year, a species of tree or freezing, and laboratory tests indicate that shrub adapted to the north-temperate envi- cold hardmess in some of these plants per- ronment alternates between periods of cold 36

hardiness and cold sensitivity. The term ohsm slows to a standstill and, since the cells acclimation (hardening) refers to the transi- die, the entire plant dies. tion from a sensitive to a hardy condition, Apparently, species of plants that survive while deacclimation (dehardening) desig- temperatures lethal to other species do so by nates the hardy-to-sensitive transition. preventing the dehydration caused by ice for- Obviously, the seasonal timing of acclima- mation. One way in which they accomplish tion and deacclimation is of critical impor- this involves "supercooling"-the absence of tance in determining a plant’s cold hardi- ice formation even during periods of freezing ness. The magnitude and duration of the temperatures. Another way is for ice to form acclimated state are also crucial. In fact, the in the intercellular spaces but for the loss of Arnold Arboretum’s hardiness zones classify water from cells to be reduced. Often, this woody-plant species according to the mag- means of frost prevention involves osmotic nitude of the cold tolerance they exhibit in alterations in the protoplasm of hardy plants. their acclimated states. Zone 6 plants, for Halophytes (salt-tolerant plants), for exam- instance, can withstand minimum temper- ple, usually are hardier than their non-salt- atures of plus 5 degrees Fahrenheit to minus tolerant relatives because the higher osmotic 5 degrees Fahrenheit, while plants of zones concentration of their protoplasm effectively 5, 4, 3, 2, and 1 exhibit progressively greater prevents water from leaving cells and con- cold hardiness. Obviously, plants in all of tributing to extracellular ice crystals. Some these categories can tolerate some below- other hardy plants generate high internal freezing weather; it is the magnitude of their contents of organic solutes (dissolved com- tolerance that differentiates them. pounds) during acclimation. Finally, certain When a plant, regardless of its hardiness plants are cold tolerant simply because they classification, is injured by a killing frost, can recover from even the extreme dehydra- several harmful processes are involved. One tion that accompanies ice formation. Exam- of the earliest and most critical processes is ples of such species are paper birch (Betula the formation of ice crystals in the spaces papyrifera/, trembling aspen (Populus tremu- between their cells. Freezing of the water in loides/, and several willows. the intercellular spaces causes water in the In view of all these considerations, what adjacent living cells to move out of the cells practically can be done to prevent freezing into the intercellular spaces, where it, too, injury in plants? An obvious strategy is to freezes. The amount of ice in the intercel- ensure that plants are well-watered before lular spaces increases rapidly as additional periods of potential frost. By keeping their water moves out of the cells. Left unchecked, tissues turgid, or swollen, one might be able the loss of water from cells causes severe to prevent the extreme cellular dehydration dehydration. In fact, the most widely held that usually kills frosted plants. A related explanation of frost damage in plants is that treatment is to spray tender plants with death is caused directly by the advanced state water whenever temperatures are below of cellular dehydration that results when ice freezing. The rationale of this procedure is forms in tissues. According to this explana- twofold. First, it maximizes the water con- tion when the concentation of water in cells tent of living tissues and, second, the heat falls below a critical "threshold" value, pro- (known to physicists as the heat of fusion) tein molecules in the cells’ protoplasm begin given off when water on the surface of a plant to cross-link with each other, forming a sta- freezes, counteracts the effect of freezing ble but nonfunctional matrix. In this per- temperatures on water within the plant. In manently altered state of protoplasm, metab- Massachusetts, for example, cranberry grow- 37

ers routinely use water sprinklers in their in fact, are quite controversial. Researchers bogs during late-spring frosts to take advan- at the University of California, for example, tage of the heat of fusion released when water are attempting to utilize genetically engi- freezes on the surface of cranberry plants. neered strains of bacteria as frost-protection The heat released by 100 gallons of water agents. They reason that some bacterial spe- when it freezes is approximately equivalent cies called "ice-nucleating bacteria," nor- to the amount of heat produced by burning mally associated with plants, tend to sensi- one gallon of fuel oil. Obviously, the sprink- tize plants to freezing injury, since individual ler technology is an important frost-protec- bacteria act as "nucleation centers" for the tion measure. Breeding programs to intro- formation of ice crystals. Other bacterial spe- duce hardiness genes into less hardy plants cies, by contrast, are nonnucleating. Dis- may also become extremely important in the place ice-nucleating bacteria on a plant with next few years. nonnucleating ones, it is argued, and the Other ways of preventing frost injury in plant should be less prone to frost injury. plants are still in the experimental stages and Unfortunately, it is still too early to judge therefore are not yet of practical value. Some, whether theory and practice are compatible

The paper birch ~Betula papynfera), m the Arnold Arboretum (left), and the tremblmg aspen (Populus tremuloides/, m the San Francisco Mountams, Anzona, are cold tolerant simply because they can recover from the extreme dehydrauon due to the formation of ice m their mtercellular spaces Many other species are cold tolerant because they prevent dehydration of their cells m the first place Photograph of Populus tremuloides by Susan D McKelvey Both photographs from the Archmes of the Arnold Arboretum. 38

in this case. Not only that, but several of the public’s concerns about nonnucleating experts question the wisdom of introducing bacteria as frost protectants will ultimately potentially harmful bacteria into the envi- depend on the results of controlled experi- ronment. They identify several important ments designed to identify possible adverse questions. Will the engineered bacteria cause environmental effects of this practice. If undesirable plants, such as weeds, to become none are found, then significant progress will frost tolerant, too? Or, since the ice-nucleat- have been made on one strategy for manip- ing bacteria normally present on plants may ulating plant cold hardiness. As usual, have beneficial but unrecognized effects on improved scientific understanding of a phe- their host plants, might not their beneficial nomenon, such as cold hardiness, if it is effects be abolished? Lastly, is it possible that properly applied, has the potential of improv- nonnucleating bacteria could affect the ing horticultural technology. weather, a prospect that could have profound consequences? As is case whenever basic science is the John W. Einset, a staff member of the Arnold Arboretum, used to solve practical problems, resolution is associate professor of biology m Harvard University. A Letter to Readers

Amoldia has been published continuously Rated at or very near the top as a membership by the Arnold Arboretum since 1911. Next benefit, Amoldia was held to be an impor- year will mark the magazine’s seventy-fifth tant benefit by a margin of seventeen to one. anniversary. During Arnoldia’s seven and A difficult and important challenge for one-half decades of existence, its format, fre- Arnoldia in the coming year or two will be quency of publication, subject matter, and to expand its mix of subject matter without even purpose have changed dramatically- neglecting or abandoning any of the tradi- from weekly or fortnightly newsletter pub- tional subjects that have delighted readers lished only during spring, summer, and fall, over the years. To guide the staff members to monthly, then bimonthly, and finally and volunteers who produce Amoldia in that quarterly magazine covering most aspects of critical task, a questionnaire will be sent to horticulture and botany. During its early all readers, Friends and subscribers alike, ear- years the publication was called the Bulletin ly in 1986, in the hopes that they will share of Popular Information. The name was valuable insights and opinions with those of changed to Arnoldia in 1941. us who produce the magazine. In 1986, Arnoldia’s tradition of evolution The past twelve months have been a period will continue. Appropriate (but modest) of careful analysis and assessment of Amol- changes will be made: minor adjustments in dia’s role vis-a-vis the University, the Arbor- format, improvements in quality, increases etum, and the Friends. The verdict has been in content, broadening of scope and diversity, favorable in every respect. During the period etc.-all without rises in production costs. of introspection, however, Arnoldia fell sub- In fact, with respect to cost, the goal will stantially behind in its publication schedule. continue to be economy in every possible The spring and summer issues have appeared way. This goal of cost-cutting will be pursued only in the last month. With the present in part through the aid of much-appreciated issue, Arnoldia’s normal publication sched- volunteer help from Friends of the Arbore- ule resumes. Winter issues will again appear tum, and others. At the same time, a serious in January, spring issues in April, summer campaign will be launched to raise outside issues in July, and fall issues in October. funding for special issues, as well as for gen- During 1986, a variety of interesting arti- eral operating costs. cles and special issues will be published by Ten months ago, the Friends of the Arnold Arnoldia-a special issue on the Arbore- Arboretum, who receive Arnoldia as a mem- tum’s historic involvement in Chinese botany, bership benefit, were polled on their atti- as well as an issue on the vine, articles on tudes toward the Arboretum, membership camellias, enkianthus, plant collecting in benefits, and so on. From Arnoldia’s perspec- China and North Africa, propagation of tive, the response was especially gratifying. Metasequoia, and the multifarious activities 40

of the Arboretum’s staff members. In 1986 U S POSTAL SERVICE STATEMENT OF OWNERSHIP, or 1987, we hope to have articles or special MANAGEMENT, AND CIRCULATION issues on the forests of the United States, (Required by 39 U.S C 3685) Chinese food plants, plant conservation, and 1 Title of publication Arnoldia A Publication Number 0004- techniques for ameliorating the impacts of 2633. 2 Date of filing September 30, 1985 3. Frequency of issue’ Quarterly. A Number of issues published annually 4 B on air pollutants woody ornamentals and Annual subscription pnce $12.00. 4 Complete mailing address other cultivated plants. As always, we will be of known office of publication The Arnold Arboretum, The Arborway, Jamaica Plam (Boston~, Suffolk County, MA 02130 5 eager to consider manuscripts on horticul- Complete mailing address of the headquarters of general bust- tural or botanical subjects for publication. ness office of the publishers The Arnold Arboretum, The Arbor- way, Jamaica Plam (Boston), Suffolk County, MA 02130 6 Full We are grateful for the patience and enthu- names and complete mailing address of Pubhsher, Editor, and siastic support readers have accorded Amol- Managing Editor The Arnold Arboretum, The Arborway, Jamai- ca Plain ~Boston~, Suffolk County, MA 02130, Publisher, Edmund dia over the past twelve months and hope A Schofield, The Arnold Arboretum, The Arborway, Jamaica they will continue to express their support Plain ~Boston~, Suffolk County, MA 02130, Editor. 7. Owner: The Arnold Arboretum of Harvard University, The Arborway, by responding to the questionnaire next year. Jamaica Plam (Boston), Suffolk County, MA 02130. 8 Known bondholders, mortgagees, and other security holders ownmg or holding I percent or more of total amount of bonds, mortgages, or other secunties President and Fellows of Harvard College, Cambndge, MA 02138 9 For completion by nonprofit organi- zations authonzed to mail at special rates (Section 411.3, DMM only) The purpose, function, and nonprofit status of this orga- nization and the exempt status for Federal income tax purposes have not changed dunng the preceding 12 months. 10 Extent and nature of circulation A Total number of copies Average number of copies each issue dunng the precedmg 12 months 4,375. Actual number of copies of single issue pubhshed nearest to filing date 4,500 B Paid circulation. 1 Sales through dealers and carriers, street vendors, and counter sales Averagc number of copies each issue dunng precedmg 12 months’ None Actual number of copies of single issue published nearest to filing date: None 2 Mail subscription Average number of copies each issue dunng preceding 12 months 603. Actual number of copies of smgle issue pubhshed nearest to filing date 601. C Total paid circulation. Average number of copies each issue dunng preced- mg 12 months’ 603. Actual number of copies of smgle issue published nearest to filing date 601. D Free distribution by mail, carner, or other means (sample, complimentary, and other free copres~ Average number of copies each issue during preced- mg 12 months 3,015 Actual number of copies of smgle issue published nearest to filing date~ 3,023 E. Total distribution. Average number of copies each issue dunng precedmg 12 months 3,617 Actual number of copies of single issue published nearest to filing date. 3,623 F Copies not distnbuted. 1 Office use, left over, unaccounted, spoiled after printing Average num- ber of copies each issue dunng preceding 12 months 797 Actual number of copies of single issue pubhshed nearest to filing date 821 2. Return from news agents. Average number of copies each issue dunng precedmg 12 months None Actual number of copies of single issue pubhshed nearest to filing date: None G Total Average number of copies each issue dunng preceding 12 months 4,375 Actual number of copies of smgle issue pub- lished nearest to filing date. 4,500 11. I certify that the state- ments made by me above are correct and complete Edmund A Schofield, Editor Index to Volume 45

(Numbers m parentheses refer to issues, and those m boldface to illustrations )

"A Directory of Information Sources on -’Kimigayo’, (2) : 13, 13 (3): 30-32 Forestry Management," ( 1 11-14 -’Koganebana’, (2) : 13, 14 -photograph by, / 1 25 Abscisic acid, (2) : 28, 31 -’Shmofu Chmmen’, (2). 14, 15 Buxus species, (2)’ 19 -structure of, (2): 29 Aruncus species, (4) : 28 Cabot, Francis H., "’Les Quatre Vents,’ Acanthopanax szeboldzanus, (2/: 25 -dioicus, (4): 21, 24 a Far-Northern Garden," /4~. 19-31 Acchmation, (4) 36 Ashton, Peter S., photograph by, (4) 10 -photographs by, (4~: 20, 21, 24, Acer gmnala, (4). 22-23, 27 Aspen, quakmg, /4/~ 22 inside back cover -pensylvanzcum, (4/: 22 -trembling, (4) : 36, 37 Calycanthus flondus, /2~: 26 -spicatum, (4). 22, 28 Asulbe,(4/ 21 Calypso bulbosa, (4) inside back cover Achlllea ptarm1ca, (4/~ 30-31 Astilbe ’Fanal’, /4/: 28 Campanula species, (4) : 31 -taygeta, (4): 21 Astrantia species, (4): 26 -persicifolia, /4~: 28 Acomtum species, (4) : 26 -carmohco var. rubra, (4). 28 =Alba’, (4): 28 Actaea pachypoda, (4) : 28 Aucuba ~apomca, (2/: 17, 19 Canada hemlock, /4~ 25 -rubra, 14): 28 -’Meigetsu’, 12): 17 Caragana species, (4/: 23 Air pollutants, phytotoxic, ( 1 15-18 Auxm, (21: 28-33 -arborescens, /2/: 19 Airelles rouges, (4) 25 -effects of, (2/~ 30 Carex phyllocephala ’Sparkler’, /2/~ 15 Alexander, John H. III, photograph by, -structure of, (2) : 29 Carpohthus natans, (4): 7 (1): 17 Azalea, roseshell, (3): 16 "Case for Monkey-Mediated Evolution Allzum chnstophzz, (4/: 21, 28 Balsam fir, (4): 22 m Big-Bracted Dogwoods," Richard Amelanchier species, (4y 22 Benthamia fragifera, (4): front-cover H. Eyde, (4) : 2-9 -bartramzana, ~4/. 23 foldout, 8 Castille~a coccmea, /3/: 16 -canadensis, (2/: 25, (4): 24, 28 Berbens species, /4/: 23 Centaurea species, (4/~ 26 Amsoma tabemaemontana, /4/: 24 -xmentorens~s, (2/~ 26 Chang, Young June, drawmgs by, (2) : Amur maple, (4) 22-23 -~uhanae, (2/: 24 9-14 Andromeda polzfolza, (2/ 19 -thunbergm, (2) 20, 26 Chilling-resistant plants, (4): 35 Anemone blanda ’Atrocaerulea’, (4). 30 Bergema cordifoha, (4)’ 29 Chilling-sensitive plants, (4): 35 Angelo, Ray, book review by, (2). 36-38 Betula lutea, /4/: 22 Chiogenes hispidula, (4): 24 -, "Botanical Index to the Journal of -papynfera, (4/ 22, 27, 28, 36, 37 Ch~onodoxa species, (4). 30 Henry David Thoreau" (reviewed), Bioregulators, (2) : 33 Cimicifuga species, /4/ 26, 28 (3): 30-32 Birch, paper, (4) : 22, 36, 37 -racemosa, /4/: 21 -, "Thoreau as Botamst: An Appre- -yellow, (4): 22 Clematis recta, (4~ 277 ciation and a Cntique;’ (3): 13-23 Blueberry, lowbush, /4/: 24 Clethra almfoha, (2~: 20, 25 anti-ethylene compounds, (2): 32 "Books" /column/, (1/~ 28-31; (2) : 35-39, Cloudberry, (4~: 24-25 Apple’Fameuse; (4)’ 23 /3/: 30-32 Coffin, Robert L , photograph by, (3). 29 Arceuthobmm puszllum, (3/. 17 "Botamcal Index to the Journal of Hen- Cold-hardy plants, /4/: 35 Ardzsza ~aponzca ’Amanogawa’, (2/: 8, 9 ry David Thoreau," Ray Angelo "Collector’s Notebook" (column), ( 1~: =Bemyuki; (2) : 8, 9 (reviewed), (3) : 30-32 25-27 = Chmmen; (2/~ 10-11, 10 "Botany The State of the Art" (col- Comum maculatum, /1~: 28 -’Chiyoda; (2): 8, 10, 10 umn), (2) : 28-34 Comus species, 12): 25, /4~ 23 -’Hmode’, (2) 10, 11 Brumback, William E., "Raising the -canadensis, ( 1 /: 19-22, inside front -’Hi-no-Tsukasa’, (2). 11, 11 Climbmg Fem from Spores," (3) : 27-29 cover, 19, 20, 21, /4/ 3, 25, 27, back -’Hokan Nishiki’, (2y 11, 12 Brunnera species, (4): 26 cover -’Hoshiami; (2/: 12, 13 Bunchberry, ( 1 inside front cover, -relay buds of, ( 1 ~ ~ 20, 20, 21 -’Ito Fukunn’, inside front cover, 12, 18-22, (4y 25 -rhizomes of, / 1 19 13 -seedlmg of, ( 1 19 -seeds of, germmauon of, (1) 20 Bussewitz, Albert W., book review by, -capitata, (4): front-cover foldout, 3, 4 42

-frmt of, (4): 5 Emset, John W., "Chemicals That Reg- Hamamelis vmgimana, /2): 24 -disciflora, (4y 3 ulate Plants," /2/: 28-34 Hardening, (4)’36 -flonda, ( 1 ~: 19-22, 21; /4/: front cover, -"What Determmes a Plant’s Cold Hazelnut,(4):24 3,6 Hardmess?", /4y 35-38 Hemlock, Canada, /4)~ 25 -kousa, /4/: front cover, 3 Elaeagnus umbellata, (2~. 25 Heracleum specics, (4): 30 -frmt of, (4): 7 Enkianthus campanulatus, /2~: 26 Hormoroot A, ( 1 23 -mas, /4/: 4 Epilobmm angustifolium forma albi- Houttuyma cordata ’Cameleon’, (2): 16 -nuttalln, /4/: 24, 26 2, 3 florum, /4/: 27 Hsueh Chi-~u, photograph of, (4) : 10 -racemosa, (2): 24, 26 Erythromum revolutum, /4~: 30 =’Remmiscences of Collecting the -sencea, /2~: 19, 25, 26 Ethephon, /2/: 32-33 Type Specimens of Metasequoia -subgroups of, evolutionary relation- Euonymus alatus, /2~: 19, 24, 25 glyptostroboides H. H. Hu & Cheng;’ ships of, /4~: 4 = Compactus’, /2~: 24 /4): 11-18 Corydahs cashmenana, (4/: 30 Fern, climbing, /3~: 1, inside back cover Hutchmson, B. June, book review by, -sohda var. transylvamca, (4~: 30 -distmburion of m New England, /3/: (2): 35-36 Corylus amencana, (4): 24 27, 28 Hydrangea macrophylla, (2): 19-20 Cotmus coggygna, /2/: 26 -ferule pinnae of, (3): 29 -quercifoha, /2): 19, 26 Cotoneaster species, (2~: 19, 20 -frond of, /3/: inside back cover Hypemcum perforatum, (2/. 20 Crataegus foetida, /4~: 25 -spore of, /3~: 29 Hyphomyces lactifluorum, /4/. 24 Creeping snowberry, (4): 24 "Field Guide to Poisonous Plants and Ice-nucleating bactena, /4): 37-38 Crocus chrysanthus, /4/: 30 Mushrooms of North America," "If You Decide to Cut," Ernest Gould, Cynoxylon, /4/: 3 Charles Kmgsley Levy and Richard B. (1): 10 Cynsus scopanus, (2)’ 19 Pnmack (reviewed), / 1 ~ ~ 30 Ilex mtegra ’Green Shadow’, /2). 16 Cytokmn, /2~: 28-31, 33-34 Fihpendula species, (4). 31 -verticillata, /2/: 25 -structure of, (2) : 29 Fir, balsam, 14): 22 Indolebutync acid, / 1 23 2, 4-D /2, 4-dichlorophenoayacetie acid), Flmt, Harnson L., and Donald Wyman, Johnson, LeRoy, photographs by, 14): 15 (2/: 29 "Plant Hardiness-Zone Maps," (4) : -Norman K., et alm, "Garden Design Daphne mezereum, /4~. 26 32-34 History, Principles, Elements, Prac- -odora ‘Rmgmaster’, /2/: 14, 15 Forest Cutting Practices Act (Massa- tice" (reviewed), /2). 35-36 -’Zmko Nishiki, /2/: 14, 15 chusetts), (1~: 10 ~umperus commums var. depressa, /4): Deacclimation, /4~. 36 Fothergilla gardenn, /2~: 26 22 Defoliants, (2) : 31-32 Frey, Susan R., et alm, "Garden Design: -honzontahs, /4): 22 Dehardemng, /4y 36 History, Principles, Elements, Prac- Kalmia angustifoha, (2) 19 Del Tredici, Peter, photographs by, ( 1/ ~ tice" (reviewed), (2): 35-36 Kerna ~apomca, (2): 19 18, 19, 20; (3): 31 "Garden Design. History, Principles, Koller, Gary L., photograph by, /1): 26 -"The Layered Look;’ ( 1 19-22 Elements, Practice," William Lake Koten engei, (2): 7 -"What’s m a Leaf?", (2) : 2-6 Douglas et alm (reviewed), (2) : 35-36 Kousa, (4): front cover, 3 Delphmmm species, (4): 26 Gaulthena hispidula, (4): 24 Labrador tea, (3). 17 Dendrobenthamia, /4/. 3 Gentiana species, /4~: 29 Landry, Sarah B., drawmg by, (3): 1 DeSando, Cynthia, drawing by, (3) : 14 Gibberelhc acid, (2)’ 28-33 Lanner, Ronald M., "Trees of the Great Dirr, Michael A., photograph by, /2~: -effects of, (2) : 32, inside back cover Basrn: A Natural History" (reviewed), front cover -structure of, (2) 29 / 1 30-31 Distylium racemosum ’Akebono; /2y Glaucidmm species, (4) : 29 Larch, American, (4): 22 15, 16 Gleason, Herbert Wendell, photograph Lanx lancma, (4) : 22 -‘Guppy; (2)’ 15-16 by, /3~: 25 Ledum groenlandicum, /3/: 17 Dogwood, /4y 3 Goodyera oblonglfoha, /4~: 23 "’Les Quatre Vents,’ a Far-Northern Doromcum species, (4~: 26 -repens, (4): 23 Garden," Francis H. Cabot, (4): 19-31 Douglas, William Lake, et aln, "Garden -tesselata, /4/: 23 Leucothoe catesbaei, (2) : 19 Design: History, Principles, Ele- Grape ’Thompson Seedless; (2): inside -fontanesiana, (2/: 19 ments, Practice" (reviewed), (2): 35-36 back cover Levy, Charles Kmgsley, and Richard B. "Drawn from Nature: The Botanical Gray, Asa, ( 1 28-29; (3): 14, 15, 17, 18 Pnmack, "A Field Guide to Poisonous Art of Joseph Prestele and His Sons," Gould, Ernest, "Managmg a Small Plants and Mushrooms of North Charles van Ravenswaay (reviewed), Woodlot," / 1 2-10 America" (reviewed), ( 1 30-31 / 1 28-30 Gunnera mamcata, /4/: 29 Lewandowski, Rick, and Paul W. Meyer, Dwarf mistletoe, (3/ 17 Halle, Francis, / 1 ~: 19, 20, 22 "The ’Okame’ Cherry" / 1 23-24 43

Ligulana species, (4) : 26 plant communities of, (3): 16, 22 -strobus, 14): 22 Lihum marhan, ~4/: 28 Mountam ash, (4): 22 -sylvestns, (4/: 25 -martagon, (4/: 28 -cranberry, /4/. 25 Pmxter-flower, (3/ 16 -var. album, (4) : 28 Muscan, (4): 30 Pissenht, (4). 30 Lmdera benzom, ~2/~ 20 Mynca cenfera, /2/~ 19 "Plant Hardmess Zone Map," Umted Lmgonberry, (4): 25 . -pensylvamca, /2/. 23, 25 States Department of Agriculture, Lmnaea boreahs, (4) : 23, 27 Nasturtmm officmale, (31: 19 (4) : 32, 33, 33 Lmodendron tuhpifera, ~2/ 2-6 Neilha species, /4/. 23 "Plant Hardiness-Zone Maps," Donald -’Fastigiatum; flower of, (2) : front Nunes, Dawn M., drawing by, (2): 2 Wyman and Harnson L Flmt, (4/: cover "’Okame’ Cherry," Paul W. Meyer and 32-34 -flower of, ~2/: front cover, 1 Rick Lewandowski, / 1 23-24 "Plants That Ment Attention: Volume -leaves of, ~2/ 1, 2, 5 Ophiopogon japonicus ’Torafu’, (2). 16 I-Trees," edited by Janet M. Poor development of, (2): 2, 6 Oryza satma, (3) : 3 (reviewed), (2): 36-38 -stipules of, development of, (2): 6 Paeoma species, (4) : 26 Pogoma, whorled, (3): 19 Little, Christopher, photograph by, (2). Pamcum species, (4/: 28 Poor, Janet M., editor, "Plants That 35 Paper birch, (4) : 22, 36, 37 Ment Attention. Volume I-Trees" Littlefield, Susan, et alll, "Garden Penmsetum species, (4): 28 (reviewed), (2) : 36-38 Design: History, Principles, Ele- Penstemon species, (4) : 26 Poplar, Carolina, (4) 22 ments, Practice" /remewed/, ~2/: 35-36 "Persimmon calyxes," (4). 6, 8 -Lombardy, /4/. 22, 25 Lombardy poplar, (4): 25 Petasites japonicus var. gigantea, (4/. 29 Populus canadensis, (4): 22 Lomcera species, ~2/: 19; (4): 23 Philadelphus species, (4/~ 23 -nigra ‘Itahca’, (4/~ 22 Loucks, One L , photographs by, (1): 16, Phlox Alpha; (1/: 27 -tremuloides, (4/: 22, 36, 37 17 -blue, ( 1 26 Prestele, Joseph, ( 1 28-29 Lygodmm palmatum, ~3/: 1, 16, 25 -creepmg, ( 1 26 Pnmack, Richard B., and Charles -distribution of in New England, /3/: -Miss Lmgard’, ( 1 26, 27 Kmgsley Levy, "A Field Guide to Poi- 27, 28 -mountam, / 1 27 sonous Plants and Mushrooms of -fertile pinnae of, /3/: 29 = Reme du Jour’, ( 1 27 North Amenca" (reviewed), ( 1 30 -spore of, ~3/~ 39 = Rosalmde; (1). 27 Primula species, (4/. 28, 29 ’ Magnolia vmgmiana, (2): 26 -summer, / 1 /: 26, 27 -abchasica, /4/: 25 Mahoma aqmfohum, (2) : 19 Phlox amoena, ( 1 1, 26, 27 -alpicola, (4/. 28 Malus species, (4) : 23 -carolma, (1/: 25, 26, 27 -var. luna, (4): 21 -’Fameuse’, ~4/: 23 -dmancata, /1/: 26 x var. alba, (4) : 29-30 "Managing a Small Woodlot," Ernest -glabernma, (1/: 25, 27 -aureata, (4): 29 Gould, (1/: 2-9 -maculata, /1/~ 27 -flaccida, (4/: 29 Maple, Amur, ~4/: 22-23 -‘Miss Lmgard’, ( 1 26, 27 -flonndae, (4/: 21, 28 Matheny, R. Norman, photograph by, -ovata, ( 1 27 -nutans, (4). 29 ~ 1 front cover -pamculata, (1/: 26, 27 -pnmulma, (4): 29 McKelvey, Susan D., photograph by, (4) : -stolomfera, (1/. 26 -pulverulenta ’Bartley Stram; (4/~ 29 37 -subulata, ( 1 25 -reidn, (4). 29 Meconopsis species, (4) : 30 Phytoalexms, (2): 33 -reptans, 14). 29 -betomcifolla, (4/: 21, 28, 29 Phytohormones, chemical structures -rosea, (4): 29 Metasequoia glyptostroboides, (4y of, (2/ 29 -sapphirina, (4/: 29 11-18, 14, 15, 1G -horticultural applications of, (2): 29-33 -sikkimensis, (4/: 28 Meyer, Paul W , and Rick Lewandowski, -kmds of, (2): 28-29 -sonchifolia, (4/: 29 "The ’Okame’ Cherry," 1 23-24 Picea ables, (4). 22 -maln, (4/: 29 Micropropagation, (2): 30-31, 33-34 -glauca, (4) : 22 -vulgans ssp. sibthorpn, (4/: 26 Miscanthus species, (4): 28 -nigra, (4/: 22 Prunus campanulata, /1/: 22 Mistletoe, dwarf, ~3/: 17 Pinus banksiana, (4): 22, 23 2014xmcam ’Okame, (1): 23-24 Molma species, (4). 28 -cembra, (4/: 25 -mcisa, (1/~ 23 Moosewood, 14): 22 -mugo, (4): 25 Puschkmia species, (4): 30 Morris Arboretum, ( 1 23 -parm flora ’Fubuki Nishiki’, (2/: 16-17 Pyrola species, (4) 27 Moss pink, ( 1 25-26 -’Janome; 12): 17 Quaking aspen, (4)’ 22 Mount Washington, New Hampshire, -’Ogon; /2/: 17 "Raising the Climbing Fern from -resmosa, (4/: 22 Spores," William E. Brumback, (3): 44

25-27 Sp1raea species, (4) : 27, 30 Twinflower, (4): 23 "Recognizing and Treating Air Pollu- Spotted hemlock, ( 1 28 Vaccimum angustifolium, ~2/: 19, ~4): tion Damage to Familiar Cultivated Spruce budworm, (4): 27, 30 24 Plants: A Conference," ~1/~ 15-18 Staples, George, book review by, /2~: -uhgmosum var. alpmum, ~4/: 25 Research Report (occasional feature), 38-39 -mns-idaea var. mmus, /4/: 25 ~4/: 2-9 Sulawesi, Indonesia, /3/: 2-12 van Ravenswaay, Charles, "Drawn from Rhododendron species, ~2/: 19 Supercooling, /4~: 36 Nature: The Botanical Art of Joseph -bakem, /2/: 26 Syrmga species, (4): 23 Prestele and His Sons" (reviewed), / 1/: -’Janet Blarr; ~2/: 26 -tissue culture of, (2~: 30 28-30 -nudiflorum, (2/: 26 Taxus canadensis, (4~: 22 Van Valkenburgh, Michael, et alm, -prumfohum, (2/: 26 Thahctrum species, /4/: 26 "Garden Design: History, Principles, -roseum, (3) : 16 "The Book of Edible Nuts," Fredenck Elements, Practice" (reviewed), (2): -schhppenbachil, (2/: 26 Rosengarten, Jr. (remewed~, /2~: 38-39 35-36 -vasey, /2/: 26 "The Case for Monkey-Mediated Evo- Vaux-le-Vicomte, France, gardens at, -yakusimanum, (4/: 29 lution in Big-Bracted Dogwoods," (2): 35 Rhodotypos scandens, ~2/: 20, 24, back Richard H. Eyde, /4/: 2-9 Viburnum species, /2/~ 19; ~4). 23 ° cover "The Golden Waterworks: Tora~a Ritu- -dentatum, ~2/: 20 Rhus aromatica, /2/: 26 als of the Wet-Rice Landscape," -dilatatum, (2/: 25 -typhma, ~2/: 19, 26 Charles Zerner, (3) : 2-12 -lantana, (2) : 25 Rmker, Robert, photographs by, ~2/: "The Layered Look," Peter Del Tredici, -prumfolmm, ~2): 25 inside back cover, 1-14, 16 ( 1 19-22 -sieboldm, (2/: 24, 25 Rodgersia species, (4): 29 "The ’Okame’ Cherry," Paul W. Meyer Volkman, T., photographs by, (3): 2, 5, Rosa mtida, (2) : 26 and Rick Lewandowskr, / 1 23-24 6, 111 -mchuraiana, /2/: 26 Thmam, ( 1 /: 23 Weaver, Richard E., Jr., photographs by, Rose ’Fairy; /2/: 26 Thoreau, Henry D., (3) : 13-26, 30-32 ~ 1 25, 26 -’Seafoam’, (21: 26 -Journal, (3/: 13, 16, 18, 19 -, "The Appeal of Phloxes," 1/: Rosengarten, Frederick Jr., "The Book botamcal index to, (3): 30-32 25-27 of Edible Nuts" (reviewed), (2): 38-39 iliustrauons from, (3): 15, 16, 18, Wemstem, Geraldme, photographs by, Rubus chamaemorus, ~4/. 24-25 21,22 92): 20, 23, 24 Rudbeckia species, (4): 31 -quoted, /3/: 15, 17, 19, 20, 22, 23 -"Replacmg the Understory Plant- -maxima, /4/: 26 -portrait of, /3/: 14 mgs of Central Park," ~2/: 19-27 Schultes, Richard Evans, book reviews -"The Succession of Forest Trees," Wetlands Protection Act (Massachu- by, / 1 30, 30-31 (3) : 21-22 setts), (1/: 10 Scilla species, ~4/: 30 "Thoreau as Botamst: An Appreciation "What Determmes a Plant’s Cold Har- -sibmca, (4/: 26 and a Critique;’ Ray Angelo, /3/: dmess?’; John W. Emset, 14): 35-38 Scirpus amencanus, ~4/: 19 13-23 Whorled pogoma, ~3): 19 Serviceberry, (4) : 23 Thrift, / 1 26 Wise, Hope, photograph by, 1 88 Shm-shan, ~4/: 11, 14 Thu~a species, (4). 25, 26-27 Wyman, Donald, and Harnson L. Flint, Smo-Amerrcan Botamcal Expedition, -occidentahs, /4/. 22 "Plant Hardiness-Zone Maps," ~4): 1980, Amencan members of, photo- Tora~a people, /3/. front cover, inside 32-34 graphs by, (4): 13, 14, 16 front cover, back cover Yellow birch, (4): 22 Skmner, D. Henry, ( 1 23 "Trees of the Great Basm A Natural Zeatm, structure of, (2): 29 Slash Disposal Act (Massachusetts/, ~ 1/: History," Ronald M. Lanner (reviewed), Zerner, Charles, photographs by, (3): 10 /1/~ 30-31 front cover, 9, back cover Smith, Olga A., drawing by, 1 21 ’Iiembles, les, /4/: 22 -"The Golden Waterworks: Tora~a Snowberry, creeping, (4): 24 Trembling aspen, (4): 36, 37 Rituals of the Wet-Rice Landscape," Soldanella species, (4): 28 Trollms species, (4): 26 (3): 2-12 Sorbus species, ~4/. 28 -Alabaster’, (4) : 28 Zwinger, Ann Haymond, book review -amencana, /4/. 22 Tryon, Alice F., photograph by, (3) : by, ( 1 28-30 South Sulawesi, Indonesia, /3/. front inside back cover cover, inside front cover, back cover Tsuga canadensis, /4y 25