AMUR HONEYSUCKLE (LONICERA MAACKII; ASCENT, CAPRIFOLIACEAE): ITS AND DECLINE, FALL LUKEN JOHN THIERET JAMES W. O. and Department Biological Sciences of Northern Kentucky University KY Highland 41099-0400, U.S.A. Heights, ar chronology of interaction bet of arboreta, botanical gardens, ; The ;ntual naturalization. specie introduccion y eventual naturalizacion de esta planta. Se describe la biologia de esta especie, how new In an to understand non-indigenous plants occupy geo- effort graphic two processes have received the most attention: population areas, documented from herbarium 1985) and popula- spread records (Forcella as (Mack 1985). Although most plant mvasions result from accidental or in- human cultures and the historical and extant exchange systems contrib- to A when uting plant invasion. description of these systems, they operate to may become with plants that eventually escape and naturalized, be useful understanding and extent of the invasion process. for rates areal We 150-year chronology of events that eventually led to present here a shrub Lomcera maackii introduction and naturalization of the eastern Asiatic Amur North America. Herder honeysuckle, (Rupr.) (Caprifoliaceae), in Throughout most of time maackii was highly valued in gardens and this L. conservation plantings. However, the tendency of the species to naturalize woody and spread beyond points of original introduction established as a it The documented "weed" of concern eastern U.S. history of interaction in Amur The between honeysuckle and people both extensive and varied. is may made chronology given here be of value regulatory decisions are as about future plant introductions. 4S0 Sum 1995 16(3) The addition of L. mctcickii to the alien flora of North America can be traced to three historical interactions between the plant and Homo sapiens: member (1) Discover the plant, classify it, and describe it as a of the flora of eastern Asia; Introduce the plant western (2) to horticulture for attrac- its abundant showy tive foliage, flowers, and and Use fruits; the plant to (3) achieve conservation goals, e.g., soil stabilization and/or wildlife-habitat improvement. These three interactions contributed widespread to intro- duction, thus necessitating fourth and remove a final one: control or the many from plant the biotic communities that have been invaded. From the Pacific to Western Europe: The Russian Role mid In the 9th century, Russian possessions in coastal eastern Asia were 1 all north of the 55th parallel, a somewhat less than hospitable region. To extend more their holdings into favorable areas, the Russians initiated a of southward series explorations into the relatively greener pastures of Manchuria (Bretschneider This 1898). area, then "but loosely held in the feeble grasp of the Chinese government," had excellent harbors and abun- Among dant resources, including timber. was the targets the territory north Amur Amur of the River and bounded by that the and Ussuri the rivers, Sea ofJapan, and the Korean frontier. Russia eventually annexed these lands by treaty from China in 1858 and I860, thus extending domain south its to the latitude of present-day Vladivostok. One Amur of the scientific expeditions sent to explore the valley of the began at Irkutsk in April 1855, returning Irkutsk months to 9 later Accompanying (Bretschneider 1898). the expedition was a naturalist, Ri- Maack chard (1825-1886), professor in the Gymnasium of Irkutsk. Maack is remembered name today primarily in the of a genus of Fabaceae, Maackta, and in the specific epithets of several species, including one Lonkera. in Among Maack Amur the species that found along the in June was the Amur yet-to-be-described made honeysuckle; he but a single collection of — Range Amur the plant in the Bureja north of the about midway between Khabarovsk and Blagoveshchensk (Maximowwicz 1878; Ruprecht 1857). His specimens woody of plants from the trip were sent to Petersburg, St. where they provided the basis for part of the publication devoted first to plants of "Amurland" (Ruprecht Amur 1857), that area on both sides of the between 42° and 55° north and 131° and 141° (Maximowicz ca. east 1859). work In that the honeysuckle was described new as a species, Xylosteum maackii, by Ruprecht (Ruprecht Maack 1857). (1859) gave an account of Amur; journey along work his the in this the published is first illustration Amur (at least in western literature) of honeysuckle The (Fig. species 1). was soon included Maximowicz's in Pnmitiae florae Amurensis (Maximowicz Amurland, Maximowicz's 1859), the flora of written after first trip first (1853-1857) to eastern Asia. Five years after publication of the flora, the Herder (Herder (Ruprecht) species was transferred to Lonicera as L. maackti 1864). Maximowicz did not see the plant in nature before he wrote Pnmitiae However, expedition (1859-1864) he obtained florae Amurensis. in a later specimens of for St. Petersburg from five localities (Herder 1878): near it the mouth of the Amur, St. Olga Bay, and at three sites near Vladivostok. t i We Maack do not accept Bretschneider 1898) that "intro- repc^rts (e.g., Boom Petersburg Botanic Garden; duced" L. maackii to culti ration at the St. Wyman Rehder (1949b), ind (1969) even date this event as "I860." (1959), We conclude Maack did not bring seeds or living plants of L. maackii that to the Garden from eithe of his trips (1855, 1859). If he had brought back • much blooming such propagules, these would have produced plants earlier than 1883, the date recorded in Gartenflora (Regel 1884) for the first Euro- Petersburg Garden. This beginning pean flowering of the plait, at the St. Maack was some 24 of the plant's ascent in western horticulture years after s — The comes not return from eastern / species into flower in 3 to 5 last sia. 24— The from flowering of other eastern years seed (Lo "enz et 1989). al. Maack Asiatic woody plants raised from seeds that did send to St. Peters- and was reported burg Pyrus, Deutzta, Syringa) in species oiClen'atis, (e.g., the early 1860s (Bretschr eider 1898). According Thatchei plants of L. maackii were introduced to to (1922), unknown— Manchuria— Petersburg from introducer in 1880; these St. plants could well have be;n the ones that came into flower in 1883. (That Anonymous were the introduced p "opagules was maintained by {1924}.) seeds Petersburg Regel's (1884) report cf the flowering of L. maackii in St. in works 1883 was soon translated abridged, and published in horticultural England Anonyinous 1884a, 1884b; Nicholson 1888) and the in (e.g., United (Bailey 190 Davis 1899); the authors of these reports obvi- States 3; However, within decade ously had not seen living ^^xamples of the plant. a from 1884 morphological data obtainable only deiailed after the article, Germany Koehne were published (Dippel 1889; 1893), indi- plants in //i^^ German 1896 one nursery cating cultivation in that country. In at least grew maackn comm.). The National Botanic Gardens, (Cole, L. fers. Amur honeysuckle from Dublin, purchased plants of Glasnevin, Ireland, The was the French nursery Lemoiue in 1889 (Nelson, pers. comm.). plant Kew Gardens Ukraine 1898 cultivated at in 1896 (Royal 1896), in in Garden Darmstadt, Germany, (Kokhno 1986), and in the Botanical in in 1900 (Purpus 1900). The Purpus article contains the earliest photograph known of the species to us. The European introductions almost seeds or plants fcr these early and garden there had long been receiving plant marerials by Rus- collected and sian travellers in central eastern Asia. Duplicates of these collections were sent to other major European botanical gardens (Bretschneider 1898). many As botanical gardens do, the one Petersburg published annu- at St. m ally a list of seeds available; L. maackii appeared the garden's first list, its Delectus seminum, in 1887 (Hortus Botanicus Imperialis Petropolitanus 1887). Amur Within a few decades, honeysuckle was growing in botanical gar- much dens through of Europe. Seeds of the plant eventually were offered in the seed lists of various European gardens for the time in the following first years: Cambridge, 1913 (Cambridge University Botanic Garden 1913); Oslo, 1917 (Universitet Botanske Have 1917); Dublin, 1919 (Royal Botanic Gardens, Dublin, Copenhagen, 1924 1919); (Horto Universitatis Hauniensis 1924); Edinburgh, 1924 (Royal Botanic Garden, Edmburgh, Amsterdam, 1924); 1929 Qardin Botanique de I'Umversite d'Amsterdam and (Museum 1929); Pans, 1931 d'Histoire Naturelle 1931). From Western The the Europe: Pacific to English Role The earliest recorded observation of L. maackii by a European not that is Maack of but appears to be that of Robert Fortune, who the mid 1840s in collected the species in China. Where in China the specimen came from, Amoy somewhere either or in "northern China," has been matter of de- a The specimen Kew) "34" bate. (two sheets at has but scant data: "A" and Bretschneider (Fig. (1894, 1898) concluded "A" 2). that the stands for Amoy, which Fortune did indeed The specimen may visit. well have been much collected in a garden because Fortune spent time searching gardens new for plants to introduce Europe. to Anonymous Several authors (e.g.. 1929, 1934; Bean 1973; Thatcher Wilson 1922; 1929) maintained that the first introduction of L. maackii into Great Britain was in 1900 by E.H. Wilson; was one of the species he it collected China during in his first trip there for the James Veitch Nursery. We were, however, unable to reconcile this date and method of introduc- with tion the statement Bretschneider in (1898) that the Petersburg St. Garden sent to the "greater botanical institutions Europe and America, in Kew" especially to (italics ours), seeds and plants and also duplicate speci- mens from the collections received from and it central eastern Asia. Be- cause L. maackii was growing at St. Petersburg since about 1880 and was first listed in the Garden's Delectus semtnum 1887, we wondered why in Kew propagules of the plant had not been sent to before 1900. After read- ing in Truelove (1917) that L. maackii was "1894" Kew listed in in the "Hand-List and Shrubs" we of Trees finally obtained a copy of that work 485 LUKEN AND Thieret, Lonicera maackii 1896 volume the one of concern) and found that L. (date actually for 2, We maackii indeed listed there (Royal Gardens 1896). suggest that seeds is Kew from Petersburg were sent to some time before 1896 and that the St. among plant or plants from those seeds languished, unheralded, their con- geners the garden. However, what might be called the "effective" intro- in 1900 by Wilson duction of L. maackit into Britain was apparently that in Veitch which then extolled and disseminated in Britain the nursery, it for Once propaganda machine was Veitch ac- and elsewhere (Allan 1974). the many horticultural tivated in behalf, the plant received notices in lit- its The company North America. largely favorable until recently in erature, Amur meeting Royal Hor- exhibited specimens of honeysuckle at a of the Award 1907 where they received an of Merit (Floral ticultural Society in made Anonymous drawing Committee {19151 of L. maack'u 1908; see for a A award was bestowed on the from 1907 Veitch specimens.). similar the Amur Committee honeysuckle was one of the 1915 1916). plant in (Floral made such double award had been few plants which, until that time, a to European (Truelove 1917). Early mentions of the plant in continental peri- photograph with sup- include one the Belgian Tribune Hortkole, a odicals in plied by Veitch (Anonymous 1909), and one in the French Revue Horticole, through France apparently the reporting introduction of the species into United agency of Veitch (Mottet 1907). Veitch sent seeds of the plant to the 1908 (U.S.D.A. Department of Agriculture (U.S.D.A.) as early as States 1909). From and Western Europe North America the Pacific to Amur we The North American record of honeysuckle have lo- earliest Dominion Arboretum, Ottawa: plants were cated in archives of the re- is Germany comm.). 1896 from Spaeth Nurseries in (Cole, pers. ceived there in New York Botanical Garden: The U.S. record in archives of the first is Amur 1898 honeysuckle from Russia were accessioned there in of seeds comm.) (Table (Riggs, pers. 1). This U.S. record of L. maackii came about through the agency of the first Foreign Seed and Plant Introduction (S.PI.) then newly organized Section of which was mandated procure, propagate, and distrib- U.S.D.A., to of the new and 1897 the U.S.D.A. dispatched and valuable seeds plants. In ute Hansen an agricultural explorer to Russia in search of cold- Niels E. as The U.S.D.A., extended Hansen's the hardy forage plants. trip, first for from June 1897 to March 1898 (Hansen 1909; Taylor 1941). Unilaterally Hansen about accessions of forage, shrub, expanding charge, sent 9.30 his Washington, DC, between December 1897 and June 1898 and tree seeds to (U.S.D.A. 1899a, 1899b). Some of the seeds were delivered before facili- and dissemination (Fairchild 1938). were ready their storage for ties 486 SiDA 1995 16(3) Distribution of seeds received by the Section was started soon after they One were Washington. received at of the recipients of seeds was the first New York Botanical Garden: the "PIE" in the 1898 entry for L. maackii in Garden the archives indicated one of the "Plant Introduction Experi- first ments"— seed distributions— by initiated the S.RI. i.e., The and seeds plants imported by the were numbered consecu- S.P.I, tively starting in "Inventory No. 1," 1898. Hansen's collections are listed in the two inventories (U.S.D.A. 1899a, 1899b). The data number first for 246 From in Inventory are "Lonkera maackii. Russia. Received through I A N.E. Hansen, December, 1897." Prof. similar entry, dated January 1898, number Amur 391 is in this first inventory. Seeds of honeysuckle were thus among few hundred the accessions received by the first S.P.I. The geographical origin of the seeds of L. maackii sent by Hansen an is intriguing mystery. According to the inventory data (U.S.D.A. 1899a, 1899b), the sources of Hansen's collections seemed have spanned much to of Russia from Petersburg and Odessa most St. to the Pacific. Origins of of Amur the seeds, including those of honeysuckle, given broad are in terms, simply "from More often Russia." exact data are given for a few species: some came from {now "Sea Province Primorski Krai Maritime or Terri- tory], South Ussurie, Siberia" and some from "Amur." However, in spite of data indicating far eastern Russia, Hansen's 1897-1898 journey did not extend into that part of Asia. The Russian segment of his journey began and ended Petersburg at St. Omsk via Tashkent, Semipalatinsk, and (Taylor 1941). Apparently the far- when thest east he travelled was he visited Kuldja (or Kulja; also known as Gulja, Ining, and Yining), a Chinese city in western Sinkiang within 50 ca. miles from the Russian border. (A rather difficult-to-interpret map show- ing the routes of Hansen's several Asiatic trips was published in Hansen [1909]). Even though the exact western Chinese range of L. maackii uncertain, is known the species not to occur in that small portion of China by is visited Wang Hansen 1897-1898 (Hsu and The Wash- in 1988). seeds he sent to ington, then, must have come from some botanical garden, forestry sta- such establishments whenever he had the opportunity. For example, in August 1897 he was at the Petersburg Garden (U.S.D.A. 1899b); St. seeds Amur of honeysuckle certainly were available to him there from the stock maintained by Garden the exchange. for Thinking that Hansen's seeds marked simply "from Russia" might have been obtained from Petersburg, we obtained photocopy St. a of the garden's 1897— 1899 Delectus seminum (the Delectm for the year of Hansen's visit there— and 1898 were The for not available to (Hortus Botanicus us). list LuKEN AND Thieret, Loniccra maackii 487 number Imperialis Petropolitanus 1899) contains a most impressive of en- — among some 3000 them; some represented Hansen's of of the species tries seeds are in the Delectus, but most are not. Lonkera maackii is there, as it is in the 1887 Delectus. St. Petersburg, then, could have been the source of Amur come from some Hansen's honeysuckle but they could have seeds, Amur Hansen U.S.D.A. imported introduced honeysuckle, the After it through from foreign countries and released in the U.S. at least eight times it 1927 (Table Some of the introductions were from British botanical gar- 1). dens; others were collected from native habitats in Manchuria by U.S.D.A. employees. The success of this introduction effort was indicated by the fact Amur commer- from 1931 honeysuckle was available at least eight that in The nurseries throughout the U.S. (Partington 1931). history of intro- cial honeysuckle duction published by the U.S.D.A. indicates that plants of the now naturalized throughout eastern U.S. represent a mixture of genotypes of diverse origins. Beginning 1960s and culminating introductions in the in five official sponsored up to 1984, the U.S.D.A. Soil Conservation Service (S.C.S) a Amur was hoped program to develop improved cultivars of honeysuckle. It would that these cultivars further traditional goals of the S.C.S. soil stabi- : From and improvement. plants already lization/reclamation wildlife-habitat more naturalized in various parts of the U.S., genotypes were selected for abundant production, propagated vegetatively, and then cultivated in fruit production blocks various plant materials centers around the coun- seed at made The (Sharp and Belcher 1981). Seeds were available by request. try most successful of these cultivars 'Rem-Red' (Lorenz et al. 1989). is AMUR and naturalization honeysuckle: escape New World In the New World The hint we have located of the plant's escape in the earliest Morton Arboretum Chicago, which mention near archives of the its in is when brought weedy tendencies: "weed in arboretum since 1924, first in" Morton (Swink, comm.). This early hint of the plant's decline in favor at pers. has accelerated toward a the current situation there being well stated fall, by Swink and Wilhelm (1994): "It would be difficult to exaggerate the Swink remarked weedy potential of this shrub." Floyd has to us that the would arboretum "unbelievable take a spread of L. maackii in the is ... it Amur Rhamnus under worker keep honeysuckle and cathartica full-time to control." A Amur would keep honey- host of full-time workers be required to suckle "under control" in the Greater Cincinnati region (including far north- Ohio Lucy ern Kentucky) from which the plant was reported for by E. first Braun from Hamilton (1961) only County, where was "becoming abun- it dant in pastures and woodlands." (As of October 1994 specimens have been Ohio collected in 34 counties {Trisel, pers. comm.}). In Greater Cincinnati now the plant omnipresent, being by commonest is far the area's shrub, — — native or alien. Efforts in part thwarted by birds being made by are various governmental agencies to eliminate the species from woodlands and The other sites. plant's establishment has been short of phenom- little The enal. species ubiquitous, and often abundant, on open and is slopes in fencerows, pastures, thin woods, woodland prairies, borders, road rights- When of-way, railroad yards, and waste moved places. the junior author home KY) Amur into his in Alexandria (Campbell County, no in 1973, Now honeysuckle was on the property. hundreds of individuals are there. We conducted a survey of selected botanical gardens and arboreta in the eastern United States and in eastern and western Canada. Although many who and botanists arborists responded noted that L. maackii was natural- ized, the species was considered a problem weed only in the following lo- DC; National Arboretum, Morton Arboretum, calities: Butler Univer- IL; IN; Bernheim KY; Matthei sity, Forest, Botanical Gardens, MI; Beal W.J. MO; Botanic Gardens, MI; Shaw Arboretum, Morris Arboretum, PA; Core WV. Arboretum, The may Edmonton species not be winter hardy and at Amur Montreal. Non-cultivated known plants of honeysuckle are currently 24 in at least states of the eastern U.S. (Trisel and Gorchov 1994) and in Ontario (Pringle 1973).