Pistia stratiotes - Chicago Botanic Garden

In August 2004, I continued myEcuadorian adventure, the firstpart reported in the last IAS

Newsletter. During the second half Ivisited areas on the Pacific andAtlantic slope. I was accompanied byGenevieve Ferry and Elisa Levy.Genevieve arrived from NancyBotanical Garden in France. I metGenevieve during a trip to Europefor Josef Bogner’s honorary Ph.D.ceremony. Genevieve has greatinterest in Araceae, but had neverbeen to a tropical rainforest. HerDirector, Romeric Pereil, wasconvinced that Genevieve and the

botanical garden would profit if shewere to join an expedition toEcuador. Elisa Levy, a botany studentat the Universidad Catolica, is anative of El Chical in Carchi Provinceand proved very useful because shewas very familiar with the area. I hadbeen warned that it might not be safebecause of Colombian guerilleraswho frequented the area, andthought having a local resident alongwho the community was familiarwith would provide us with a certaindegree of cover. She also proved to be

Continued on page 2

FIRST CLASSU.S. POSTAGE

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P.O. Box 43-1853South Miami, FL 33143 USA

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1

ContentsTable of Contents

Explorations for Araceae inEcuador, 2004: Part IIBy Tom Croat . . . . . . . . . . . . . . . . 1

Black Magic and Purple Passion:Black AroidsBy Karen Platt . . . . . . . . . . . . . . . 4

Recent Aroid Activities at theMissouri Botanical GardenBy Tom Croat . . . . . . . . . . . . . . . . 8

Pistia Stratiotes: An UbiquitousInvader By Emily Yates & KristenKordecki . . . . . . . . . . . . . . . . . . . 12

Review of Ittenbach, Stephan,Revision der afrikanischen Artender Gattung Amorphophallus(Araceae)By Dan Nicolson . . . . . . . . . . . . 15

Advertisements . . . . . . . . . . . . . 16

IAS OfficersPresident: Albert HuntingtonVice President: Dan LevinTreasurer: Patricia FrankRecording Sec.: Lester KallusCorresponding Sec: Julius BoosMembership: Paul Resslar

IAS PublicationsAROIDEANAEditor: Dr. Derek BurchScientific Editor: Dr. Tom Croat,Missouri Botanical Garden,St. Louis, Missouri

IAS NewsletterEditor: Emily Yates

IAS Website: www.aroid.orgAdmin: Scott Hyndman

A QUARTERLY PUBLICATION FOR MEMBERS OF THE INTERNATIONAL AROID SOCIETY

Explorations for Araceaein Ecuador 2004: Part IIThomas B. Croat, Missouri Botanical Garden

the IAS NewsletterVol. 27 – 3 • July 2005

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On the journey to El Chical fromQuito, most of the route consists ofarid hills with the highway adheringto long winding slopes with lots ofbus and truck traffic. We had lunchin Ibarra, the capitol of ImbaburaProvince, then continued on toTulcán, the capitol of Carchi Provincenear the Colombian border. The areabetween Tulcán and Tufino is totallydevoted to pasture due to itselevation of 3500 m. After beingstopped by a military checkpoint justbefore reaching Tufiño, we entered anarea of shrubby vegetation and begancollecting; most of the species werenew to me, but the area was too highfor Araceae. Just to the east the roadclimbed higher and we entered theParamo del Angel (Fig. 1), with vaststands of Espeletia and Puyaextending for more than 20 km. Theparamo was inhabited by llamasoccasionally right along the road, andwhen the paramo was enveloped infog, the llamas, barely visible amongthe Espeletia, gave an eerie sensation.

Beyond the paramo the road droppedsteeply to the west and wanderedthrough forested hills where Araceaesoon became conspicuous andabundant with most of the speciesbeing new. Finally we reachedMaldonado. We drove through townlooking for a hotel, but saw nothingobvious and decided to continue onto El Chical, hoping there would be ahotel there. We arrived in El Chicaljust before dark and checked into thetown’s only hotel. It was the onlyconcrete building in town andthough somewhat primitive, it wasvery comfortable after I wired acouple of receptacles so we could runthe computer and other essentialappliances. We were the only guests

in the hotel and even the owner leftthe following day on a trip to Quito,leaving us in the care of a friend wholived nearby.

El Chical lies on the Colombianborder across from a smallColombian village on the other sideof the Río San Juan with a footbridge providing its electric currentfrom the Ecuadorians. The village isnot connected by road to any otherpart of Colombia, and El Chical has agarrison of Ecuadorian army on thehill above the town so the soldiersrun through the streets in the earlymorning doing military exercises.The single road continuing from thetown across the Río Chical goes on toPeñas Blancas, which is really nothingmore than a house at the end of theroad. The house is owned by Elisa’sgrandparents and it was here thatElisa’s father Jim Levy, then a PeaceCorp worker trekking into thelowlands to work with Awa Indiancommunities, met and fell in lovewith Elisa’s mother. The Levys live inQuito part time and also in a housethey built just east of El Chical.

We spent our first day collecting inthe area of Peñas Blancas, whichcontains many Araceae. On the way,at the river bridge near El Chical, wefound a large new species ofAnthurium growing in a tree, amember of section Cardiolonchium,and one which was being pollinatedby small wasp-like bees. I managed toget good photos and collected somebees in a plastic bag for laterdetermination, but they all escapedby chewing their way out of the bag.In another area we found twoadditional new species, both alsomembers of section Cardiolonchium.One had a dark purple spathe and areddish spadix (Fig. 2) while theother had longer internodes and aflesh-colored spadix. Among thespecies previously seen in the Lita-San Lorenzo region were

Anthurium anchicayense Croat,A. andreanum Linden,A. angamarcanum Sodiro,A. dolichostachyum Sodiro, A. micheliiGuill., A. peltigerum Sodiro,Dieffenbachia tonduzii Croat &Grayum, Philodendroncurvipetiolatum Croat, P. esmeraldenseCroat (Fig. 3), P. roseocataphyllum

2

ECUADOR 2004: PART IIContinued from page 1

15

Continued on page 3

Figure 2. New species of Anthuriumsection Cardiolonchium with a dark

purple spathe & reddish spadix.

Figure 3. Philodendronesmeraldense Croat.

Review ofIttenbach, Stephan. 2003. Revision der afrikanischenArten der Gattung Amorphophallus (Araceae)Englera 25: 1-263. [Available from Botanischer Garten und Botanisches Museum,Königin-Luise-Str. 6-8, 14191 Berlin, Germany ($40)]

This is a classic revision in the tradition of the German-speaking aroidologistsSchott (1794-1865) and Engler (1844-1930), although dealing solely with theAfrican (including Madagascar) taxa (35 species and 7 subspecies). However, theauthor has worked closely with Wilbert Hetterscheid and, between them, theyare masters of this Old World splendid tropical genus. Members of the AroidSociety may remember Hetterscheid & Ittenbach’s extremely useful (standard)work “Everything you always wanted to know about Amorphophallus, but wereafraid to stick your nose into” (Aroideana 19: 7-131. 1997) with its key to all thenknown taxa of the genus, descriptions, photographs, etc.

The treatment’s frontispiece is a classic photo (taken at the same time as the onein Aroideana 1: 68. 1978) of young Josef Bogner (now an Ehrendoktor!) standingamong flowering Amorphophallus angloensis subsp. maculatus. The treatment isbased on study of about 1000 specimens and living material of most taxa. Thechapter on morphology (pp. 11-24) is well illustrated with drawings andphotographs and the chapter on distribution and habitat (pp. 26-34) gives one ageneral overview. There is an important discussion of the genus, how its partsmay have evolved leading to a classification delivered in the forms of synopses,keys, characters, and descriptions and followed by an evaluation of thesecharacters. All this careful reflection (pp. 35-65) leads to a key to African species(pp. 66-68), followed by a discussion of the taxa. The taxa are arrangedalphabetically, which means that you can move quickly from the key to the taxon.This has the disadvantage that the taxa closest to each other are not adjacentmaking it a bit harder to compare the treatments of similar taxa.

The best way to communicate the quality of this work is to report an example ofa treatment of one species and assure you that each species has a comparabletreatment, perhaps missing one or two items or adding something else, but toshow that this is meaty stuff!

Example: Amorphophallus gracilior Hutchinson 1939, which begins on p. 160and ends on p. 165, is a renaming of A. gracilis Chevalier (1920), a name whichwas an illegitimate later homonym of the same binomial published by Engler in1881. The holotype, from Benin (in Paris), was tracked down and found to beuseless because of having been consumed by herbarium beetles and thereforehe here designates a neotype collected in Nigeria in 1946. A full page photo ofthe neotype appears on p. 162, followed by a full page distribution map onp. 163 (with two adjacent dots), followed by the full page drawing on p. 164 withmany details. The text begins with the citation of the correct name, its place ofpublication, its synonyms, its typification(s), a long description, a long paragraphon observations, distribution, habitat, and ecology, and finally, citation of other (asopposed to type) material, including the herbarium where seen, with anexclamation mark if the author saw the specimen.

The work closes (after thanking various people and institutions) with (1) a classicbibliography and, for anyone who might have unannotated duplicates, (2) analphabetic list of collectors and their specimen numbers identified by a taxonnumber, and (3) an index of scientific names.

Would that all aroids were as well studied as these African Amorphophalli!

Dan H. Nicolson, Department of BotanyNational Museum of Natural HistorySmithsonian Institution, Washington, DC. 20560-0166. ❦

Kirtikar, K.R. and B. D. Basu. 2002. IndianMedicinal Plants. Delhi, Sri SatguruPublications.

Livingstone, D. Missionary travels andresearches in south Africahttp://etext.library.adelaide.edu.au/l/livingstone/david/mission/chapter32.html.

Mayo, S. J., J. Bogner, and P.C. Boyce. 1997.The Genera of Araceae. Royal BotanicGardens, Kew.

Pimentel, D., L. Lach, R. Zuniga & D.Morrison. 2000. Environmental andEconomic Costs of NonindigenousSpecies in the United States. BioScience.50(1):53-64.

Pimentel, D., R. Zuniga & D. Morrison.2005. Update on the Environmental andEconomic Costs Associated with Alien-Invasive Species in the United States.Ecological Economics. 52: 273-288.

Premkumar, V.G. & D. Shyamsundar.2005. Antidermatophytic activity ofPistia stratiotes. Indian J. Pharmacol.37:127-128.

Ramey, V. 2001. Pistia stratiotes. Universityof Florida. www.aquat1.ifas.ufl.edu/seagrant/pisstr2.html.

Rivers, L. 2002. Water Lettuce (Pistiastratiotes). University of Florida and SeaGrant. www.iisgcp.org/EXOTICSP/waterlettuce.htm

Smirnova, L. L. & N. V. Mironova. 2004.Intensification of BiologicalAfterpurification of MunicipalWastewater by Some PleistophyteSpecies. Water Resources 31 (4):470-473. ❦

rearing and release of the nativenoctuid moth Spodoptera pectinicornishas replaced the use of herbicides.Research indicates this biologicalcontrol is not as useful as originallyhypothesized (Dray Jr. et al., 2001).Chemical control can be effective, butcan be accompanied with its own set ofproblems such as bioaccumulation inbivalves, fish and other fauna in thefood chain (Colborn and Thayer,2000).

Though Pistia stratiotes can be invasive,it is also an oxygenator for fish, andhelps keep pond water clear. Thespecies provides food for manatees(although Dunn, 1934, states that thisnatural predator cannot keep Pistia incheck). Its roots can provide breedingand hiding space for small aquaticanimals.

Pistia stratiotes is reported to absorblarge amounts of heavy metals fromwater and are being used inremediation and wastewater treatmentefforts because of their ability toincrease water quality (Smirnova &Mironova, 2004). In a comparativestudy of 10 water plants, Pistia stratiotesshowed one of the highest removalrates for zinc and lead. These heavymetals were found in or adsorbed intothe roots (Awuah et al., 2001).

Pistia stratiotes has also been used as a“famine food” – plants that are notnormally considered as crops, but areconsumed in times of famine (FamineFoods Database). In India it was usedas a famine food in 1877-1878. InChina, young leaves are eaten cooked.

In traditional medicine, dried waterlettuce or tropical duckweed is burntinto an ash, then moistened with adrop of mustard oil, and applied in athin layer on chronic, enlarged tonsils,twice a day to help get rid of tonsillitis.(www.bestincosmetics.com). In the

Philippines, it is used to treatgonorrhea. Pistia stratiotes leaves areused in traditional medicine for thetreatment of ringworm infection of thescalp, syphilitic eruptions, skininfections, boils, and wounds, becauseit possesses antifungal properties(Premkumar & Shyamsundar, 2005).The oil extract of P. stratiotes is used inthe treatment of worm infestations,tuberculosis, asthma, and dysentery,and is applied externally to treat skindiseases, inflammation, piles, ulcers,syphilitic infections and burns (Kirtikar& Basu, 2000). Pistia stratiotes is listedon the checklist of medicinal plants inSoutheast Asia as effective against boilsand ezcema (www.arcbc.org).

Pistia stratiotes has a high potashcontent (Burkill, 1935). The extractionof caustic potash from wood ash for useas an emulsifier is probably one of themost ancient ethnobotanic traditions(e.g., Quiapo or Pistia stratiotes, one ofthe Araceae family; Water Hyacinth orEichhornia crassipes, of thePontederiaceae family; the Royal fern orOsmunda regalis, the fruit of the PricklyChaff flower or Achyranthes aspera ofthe Ameranthaceae family.(www.dweckdata.com). However, likeother Araceae, it is poisonous if ingestedbecause of the calcium oxalate crystals.

References:

APIRS Online, the database of aquatic,wetland and invasive plants, Centerfor Aquatic and Invasive Plants,The University of Florida, IFAShttp://plants.ifas.ufl.edu/search80/NetAns2/

Awuah, E., F. Anohene, K. Asante, H.Lubberding and H. Gijzen. 2001.Environmental conditions andpathogen removal in macrophyte-and algal-based domestic wastewatertreatment systems. Water Science &Technology 44 (6):11–18.

Bown, D. 1988. Aroids. Plants of theArum Family. Century HutchisonPress. London.

Burkill, I. H. 1935. Dictionary of theEconomic Products of the MalayPeninsula. Oxford, England: OxfordUniversity Press.

Center, T.D. 1994. Biological Control ofWeeds: Water Hyacinth and WaterLettuce, pp. 481-521. In: PestManagement in the Tropics:Biological Control – A FloridaPerspective, Rosen, D., F.D. Bennettand J.L. Capinera (eds.). InterceptLtd., Andover, UK.

Colborn, T. and K. Thayer. 2000.Aquatic Ecosystems: Harbingers ofEndocrine Disruption. EcologicalApplications, 10 (4): 949-957.

Connecticut Dept. of EnvironmentalProtection. June 27, 2005.http://dep.state.ct.us/burnatr/fishing/fishinfo/anguide_part1.pdf.

Dray Jr., F., T. D. Center & G.S.Wheeler. 2001. Lessons fromUnsuccessful Attempts to EstablishSpodoptera pectinicornis(Lepidoptera: Noctuidae), aBiological Control Agent of Waterlettuce.” Biocontrol Science &Technology. 11(3): 301-317.

Dray, Jr., T.D. Center. 2002-2004. WaterLettuce, pp. 65-78. In: BiologicalControl of Invasive Plants in theEastern United States, R. VanDriesche (ed.).

Dunn, L. H. 1934. Notes on the WaterLettuce, Pistia stratiotes Linn. as aNursery of Insect Life. Ecology 15(3): 329-331.

Famine Foods Database:www.hort.purdue.edu.

Glazier, K. 1996.Pistia stratiotes L.Ecology and Evolutionary BiologyConservatory. www.eeb.uconn.edu.

Holm, L. G., D. L. Plucknett, J. V.Pancho, and J. P. Herberger. 1977.The world’s worst weeds:distribution and biology. East-WestCenter/University Press of Hawaii.pp. 379-384

14 3

Croat & M. Mora, P. rugapetiolatumCroat (Fig. 4), P. subhastatum Engl.and P. verrucosum Mathieu ex Schott.Also present is a new species ofPhilodendron (Fig. 5) with a narrowblade similar to P. tenue K. Koch.

Known species found in the El Chicalregion that were not found in theLita-San Lorenzo region includeA. ovatifolium Engl., A. pulverulentumSodiro var. adsimile (Sodiro) Croat &Rodríguez, P. fibrosum Sodiro exCroat, P. oligospermum Engl. andStenospermation densiovulatum Engl.Many more species that occur in theEl Chical region are still unnamedand new to science. One is a long-bladed member of Anthurium sectionPolyneurium similar to Anthuriumcuspidatum Masters, as well as amember of Anthurium sect.Calomystrium with thick ovate bladesand free basal veins, sometimes evena peltate blade (Fig. 6). Another was adeeply three-lobed member ofAnthurium sect. Belolonchium similarto Anthurium herthae K. Krause fromthe Oriente of Ecuador (Fig. 7).Several new species of Philodendronwere found including one with athick, ovate blade and deciduouscataphylls (Fig. 8).

On the return to Tulcán, we madenumerous stops at elevations thatcontained Araceae, collecting manyinteresting and new species. Themajority were in flower and large setswere collected for type specimens.Between El Chical and the Paramodel Angel, ranging from 1200 to2000 m, many species of Anthuriumsection Belolonchium were found,most new to science. One speciesnear Maldonado has long internodesand stood more than two meters tall(Fig. 9). Higher up we found a largeAnthurium section Belolonchium with


Figure 4. Philodendronrugapetiolatum Croat.

Figure 5. New species of Philodendronwith a narrow blade.

Figure 6. New species of Anthuriumsection Calomystrium.

Continued on page 4

Figure 7. New deeply three-lobedAnthurium section Belolonchium.

Figure 8. New species ofPhilodendron.

Figure 9. New species of Anthuriumsection Belolonchium.

PISTIA STRATIOTESContinued from page 13

Continued on page 15

of management research and control inFlorida for at least two decades.

As a case study of invasion biology,Pistia stratiotes is intriguing. Rivers(2002) states that P. stratiotesreproduces vegetatively and by seed.Rapid vegetative reproduction allowswater lettuce to cover an entire lake,from shore to shore, with a dense matof connected rosettes in a short periodof time. This type of rapidreproduction makes Pistia aparticularly effective invader, wheredensities of rosettes can range from lessthan 100 to over 1,000 per m2 in southFlorida (APIRS Online). Seedproduction by P. stratiotes, oncethought not to occur in NorthAmerica, is now considered importantto reproduction and dispersal (Drayand Center 1989). While Pistia is notcold tolerant (Holm et al. 1977), itsleaves re-grow after moderate freezes,and it can survive for extended periodsof time on moist muck, sandbars, andbanks (Holm et al. 1977), contributingto its success as an invader.

Pistia stratiotes can spread from wholeplants or pieces of plants moved onboats or fishing equipment from aninfested to a clean body of water(Rivers, 2002). The most commonlyaccepted pathway of this species intothe United States is in ballast water inships from South America. Ships thattravel through mats of water lettuce cancarry fertile plant segments to newareas. Once established within a waterbody, P. stratiotes can be spread bywater currents and floods.

According to Ramey (2001), P. stratiotescontinues to be sold through aquariumsupply dealers and through theinternet. Rivers (2002) cites thatdumping of aquarium or ornamentalpond plants is often the means ofspread for P. stratiotes. The popularityof water lettuce as a garden plant has

also led to its spread, in a morecontrolled manner.

Although there are several other aroidsconsidered invasive in various regions,P. stratiotes is by far the most insidious.According to Pimentel et al. (2005)Pistia stratiotes, among several invasiveaquatic plant species, is altering fishand other aquatic animal species,choking waterways, altering nutrientcycles and reducing recreational use ofrivers and lakes. In a study published in2000, Pimentel et al. determinedinvasive aquatic plants species costs $10million in losses and damages and $100million in control costs in the US. In1994, Pistia stratiotes control cost wasestimated at $650,000 annually (Center,1994) in Florida, where it is currentlyunder “maintenance control.” Thisexcludes costs associated withinterference with recreationalopportunities, like swimming andfishing, and impacts associated withinsects that breed in waterways chokedby water lettuce. Dray Jr. and Center(2002) estimated expendituresincreased to over $1 million dollarsannually in Florida by 2002.

Like Water Hyacinth, Pistia also blocksirrigation canals and provides abreeding ground for mosquitoes andclogs fishery waters. Pistia stratiotes caninflict a severe impact on theenvironment and economy of infestedareas (Rivers, 2002). The dense matscreated by connected rosettes of theplant lead to the majority of problemsencountered with water lettuce. Thesemats can have a negative economiceffect by blocking waterways, thusincreasing the difficulty of navigation,boat traffic, fishing, and hinderingflood control efforts. It may become animportant constituent of a sudd – adense aggregation of a free-floatingvegetation. Pistia stratiotes even playeda role in the infamous Dr. Livingstone’sexplorations in Africa, where, whileexploring the river Shire, a northerntributary of the Zambesi, they learned

of a previous attempt by thePortuguese to ascend the river. ThePortuguese had to abandon the trip onaccount of the impenetrable duckweed(Pistia stratiotes), makingDr. Livingstone’s expedition up theriver Shire the first by Europeans(Livingstone, website).

Mats of P. stratiotes can also disruptnatural ecosystems. They can lead to alower concentration of oxygen incovered waters and sediments byblocking air-water interface and rootrespiration. They can degrade waterquality by blocking the air-waterinterface and greatly reducing oxygenlevels in the water, eliminatingunderwater animals such as fish.Extremely thick mats of P. stratiotes canprevent sunlight from reachingunderlying water, possibly eliminatingnative submerged plants. Thecumulative effect of these negativecharacteristics of the plant is a loss ofbiodiversity in invaded habitats. Animalcommunities may be altered byblocking access to the water and/oreliminating plants the animals dependon for shelter and nesting. P. stratiotesmats can also serve as a breeding placefor mosquitoes. It reproducesexponentially and reduces the oxygenlevels in a body of water by reducingthe water’s surface area (Dunn, 1934).

Mechanical controls, like the flailchopper or harvester, have beenused to control populations of waterlettuce, but this method is costly.Biological control is under investi-gation. The South American weevil,Neohydronomus affinis, can reducepopulations, but once introduced, theweevils themselves continue to spread(http://aquat1.ifas.ufl.edu/lettuce.html).The moth, Samea multiplicalis, whichattacks P. stratiotes and Salvinia spp.,has been established in Australia, but itsimpact has not been evaluated. InThailand, classical biological controlhas not been attempted, but mass

BookReview...Black Magicand PurplePassion:Black AroidsBy Karen Platt

Aroids are superb in anycolour, but it is amazing howmany of them approachblack. It is understood thatwhen we talk about “black”plants we are encompassingthe darkest colors found inhorticulture – from bronze, topurple to near black.

I have been researching darkplants for over ten years.Aroids being my favouritegroup of plants, I’ve amasseda lot of information on them,chiefly Arisaemas and Arums.I am fascinated by thespathes and spadices ofthese plants, let alone theirmagnificent foliage. I could

blades over a meter long and stubbypurplish red spadices with a broadhooding spathe (Figs. 10 & 11).

We made it back to Quito the sameday and began pressing thecollections. Chris Davidson hadarrived the previous day and wehurried to get back to the field. Ourfirst field trip with Chris was alongthe Quito-to-Santo Domingo road,departing at Chilligallo and ending inChiriboga. Two species in this regionwere a large Anthurium with longinternodes and cordate blades withconspicuously sunken tertiary veins,and a very small Anthurium with a

stipitate spadix. We arrived atTinalandia, where we spent the nightpressing plants on the front porch ofthe lodges overlooking the Río Pilatonnot far below the mouth of the RíoToachi. I finished pressing plants thefollowing day while Chris collected inthe forest around Tinalandia.

We left the next day for Ventanas andstayed in Babahoya, where we metXavier Cornejo in the morning. It wasa miserable hotel with roosterscrowing outside the window. The nextmorning we headed to HaciendaClementina, headquarters of the largebanana industry around Babahoya.We got permission to collect at CerroSamana, the ultimate goal of the trip.

Cerro Samana is an isolated hill about750 m elevation – one of the fewremaining areas at this elevation onthe western slope of the Andes. It willbe another study site for an aroidflorula, so collecting there wasintense. Species found wereAnthurium angamarcanum Sodiro,A. argyrostachyum Sodiro, A. asplundiiCroat, A. brachypodum Sodiro,A. cupulispatha Croat & J. Rodríguez(Fig. 12), A. dolichostachyum,A. guayaquilense Engl., A. ochreatum

4 13

Continued on page 5

Continued on page 5

Figures 10 and 11. New species of Anthurium section Belolonchiumwith blades over 1 m long.

ECUADOR 2004: PART II Continued from page 3 PISTIA STRATIOTESContinued from page 12


Figure 12. Anthurium cupulispathaCroat & J. Rodríguez.

Common names: laitue d’eau,pistie (French); Lechuguillade agua, lechuguita de agua,

repollo de agua (Spanish); waterlettuce, tropical duckweed, shellflower(English).

Pistia stratiotes, or water lettuce, is aninteresting, unique aroid. Like anypotential invasive species, it must bekept in check so it does not becomedestructive; however, it has valuableproperties, as a medicinal, a foodsource, and environmental remediator.

The name Pistia stratiotes comes fromthe Greek “pister” meaning hollowtrough, in the sense of a drinkingtrough (Mayo et al. 1997) or the Greek“pistos” meaning water; and “stratiotes”meaning soldier. It was classified in1943 to its own Pistiaceae family, but asa result of later research it is now in theAraceae, subfamily Aroideae, tribePistieae, and is the only species in thegenus Pistia.

Water lettuce is a perennial monocot ofthe Araceae family. It’s fairly

unmistakable; it looks like floatinglettuce (Fig. 1). Many descriptions ofthe plant abound: see Mayo et al. 1997,Bown, 1988.

There is some confusion about theorigin of water lettuce due to itsworldwide distribution, however, thereis strong evidence that it is native toSouth America (Rivers, 2002). Pistiastratiotes is now found throughout thetropics and subtropics (Glazier, 1996),and is one of the most widelydistributed hydrophytes in the tropics(Holm et al. 1977). Pistia stratiotesrequires a wet, temperate to tropicalhabitat (Rivers, 2002).

Pistia stratiotes may have beenintroduced to North America bynatural means or by humans. It wasspied as early as 1774 by WilliamBartram, in “vast quantities ... severalmiles in length, and in some places aquarter of a mile in breadth” in the St.Johns River in Florida (APIRS Online).It has been suggested that trade via St.Augustine, founded in 1565, may have

provided an early avenue forintroduction into the St. Johnswatershed (APIRS Online).

Pistia stratiotes is a “state listed,”“prohibited,” “category I,” “species ofconcern” or “noxious” plant in at leasteight states including CA, AZ, TX, LA,SC, GA, FL, HI and Puerto Rico. Whilethe abundance of water lettuce isrecorded primarily in sub-tropicalregions of the US, it has beendiscovered in northern states, includingOhio and New York (Dray Jr. andCenter, 2002). Several states imposefines for individuals who posses orparticipate in the sale or distribution ofwater lettuce, including South Carolinaand Texas. The Connecticut Dept. ofEnvironmental Protection will addPistia stratiotes to the list of prohibitedspecies in October, 2005 and will fineindividuals who possess it $100 perplant (CDEP, 2005). It occured in 68public water bodies in Florida by 1982and in 128 water bodies by 1989, buttotal abundance reduced by half overthe same time period as a result of astatewide management program(APIRS Online).

Pistia is common in dams, lagoons,lakes, and also grows in wetland rice. Itcan also be found on stagnant waterand rooting on muddy banks. It’s oftengrown in ponds and aquaria, and canbecome a serious invasive pest in thetropics. Its capable of forming vastmats that disrupt submersed plant andanimal communities and interfere withwater movement and navigation (Holmet al.1977); also serves as host for atleast two genera of mosquitoes (Holmet al. 1977). It is considered a seriousweed in Ceylon, Ghana, Indonesia, andThailand and at least present as a weedin 40 other countries (Holm et al.1977). It is listed by the NatureConservancy’s Invasive SpeciesInitiative as a pest in Australia(http://tncweeds.ucdavis.edu/global/australia/aca.html), and has been a target

Sodiro, A. obscurinervia Croat,A. palenquense Croat, A. pallidiflorumEngl., A. versicolor, Chlorospathadodsonii Madison (Fig. 13),Dieffenbachia sp., Monstera cf.adansonii Schott, Philodendronacuminatissimum Engl.,P. platypetiolatum Madison,P. purpurascens Engl., P. tenue K.Koch, P. tenuipes Engl., P. umbricolumEngl., Rhodospatha dodsonii Croat,Syngonium macrcarpum Engl. andXanthosoma sp. One species ofAnthurium is considered new,A. samanaense Croat (Fig. 14), as wellas two new species of Philodendron.One being (Fig. 15), reasonablycommon on the Pacific slope alsonear Piñas and Guasaganda. Clearlymore time will have to be spent atCerro Samana before any florula canbe prepared.

On the way back to Quito wecollected at the UniversidadPolytechnica Salesiana’s EstacíonSacha Wiwa. This is another logicalsite for an aroid florula andcollections began there with this inmind. Among the species here not yetseen on the Pacific slope in southernEcuador were Anthurium laciniosumSodiro, A. propinquum Sodiro,A. pulverulentum Sodiro, A. urbaniiSodiro, Philodendron inequilaterumLiebm. and P. rugapetiolum.

Next we headed east and into theEcuadorian Oriente. Aftercontinuously having trouble startingthe Mazda pickup for over a month itseem to be getting worse. Afterdriving around by taxi to severalneighborhoods we finally foundsome kid claiming to be a mechanic.He arrived with virtually no toolsand ended up pouring a bucket ofwater over the battery to get betterground. I had taken the battery cable


512

Continued on page 13 Continued on page 6

hardly contain myself when Ialso learned of purple and redfoliaged forms of somespecies from China. I love theelegance of Arisaemasikokianum aptly described byone visitor to my booth at theChelsea flower show in 2004as “like one of Cecil Beaton’shats in My Fair Lady.” It isprobably the most beautiful ofall Arisaemas, a truearistocrat. However, I am alsotaken by the deeply colouredspathes of A. speciosum andmany others.

I keep both of these slightlymoist in their pots out in thegarden over winter (Z7). Thisyear, for the first time in many,we had a lot of snow. Ithought I might have lostthem, as they wereunprotected. Thankfully, theycame through unharmed.Arisaema sikokianum finishedflowering in May just asA. speciosum was breakingdormancy. They both grow inlight shade. Arisaema griffithiiis another favourite, thepatterning on this species isincredible. My favoured Arumis palaestinum, for I believe itis the darkest, but this is hardto come by. As with manyaroids, the smell is notoriousbut I have learned that it oftendepends on provenance. Ifirst discovered this whiletalking to a journalist inOregon whose family hadgrown Dracunculus vulgaris,once classified as Arum, foralmost 100 years and heclaimed it had no off-puttingsmell at all. Yet it is the plant

Book ReviewContinued from page 4

Figure 13. Chlorospathadodsonii Madison.

Figure 15. New speciesof Philodendron.

Figure 14. Anthuriumsamanaense Croat.

Figure 1. Pistia stratiotes L. Croat 85014. Photo by Tom Croat.

Pistia stratiotes L. (Araceae) –Ubiquitous Aroid InvaderBy Emily D. Yates, Missouri Botanical Gardenand Kristen E. Kordecki, Chicago Botanic Garden

apart so many times that it eventuallybroke, so on the way down to Baezawe stopped at a automobile partsplace and bought a new terminal.When I removed the terminal end Idiscovered that the problem had notbeen terminal itself, but theconnection. Putting on the new endsolved the problems permanently.

We stayed at a new place in Baezaand found it well protected withabout half the rooms filled withsoldiers. It was a nifty place whereyou could drive inside and park rightnext to the room. The last time I was

in Baeza I stayed at a place madeentirely our of wood and nearlycaught it on fire with my plant dryer.Since it is no longer there, I suspectsomeone may have succeeded inburning it down.

The following day we drove to LagoAgrio, collecting along the way. Theroute is much more denuded sincethe last time I was on the road, butthere are still patches of forest witharoids. One of these areas is the roadthat goes from the main highway tothe small village of Gonzales Díaz dePineda at the gateway to the ParqueNacional Sumaco Napo Galeras. Thetown lies east of the Río Quijos andthe area between the highway and theriver was quite rich in aroids. Amongthe collections made was of a newspecies of Anthurium sectionCardiolonchium, Anthuriumamoenum Kunth, Philodendronasplundii Croat & M. L. Soares(Fig. 16), P. palacioanum Croat &Grayum, P. ruizii Schott, P. schmidtiaeCroat (Fig. 17), Rhodospatha latifoliaPoepp., Stenospermation killipii Croat(Fig. 18), Xanthosomapurpureomaculatum Croat, and anunknown Anthurium in section

ECUADOR 2004: PART II Continued from page 5

116

Figure 16. Philodendron asplundiiCroat & M.L. Soares.

Figure 17. Philodendronschmidtiae Croat.

Continued on page 7

that I receive the most phonecalls about. The conversationusually goes something likethis: ‘I have this plant thatsmells like something iswrong with the drains...’ I findthis frustrating; after all,animal lovers get used to thesmell of horses, cows andpigs. The beauty of theseplants far outshines anyoffence to the nose.

I just missed the flowering ofAmorphophallus titanum atKew by two days. I’ll regretthis for the rest of my days.No other plant is sooutrageous or flamboyant.

Having just started up mynursery again, (I have theaccolade of being the veryfirst person to run a nurseryspecializing in black plants), Ihope to be introducing manyspecies into the U.K. As moreof these plants are introducedto the West, one realises thewealth and magnitude of darkaroids and just how easymost of them are to grow.

———————————

Karen Platt is author of Black Magicand Purple Passion, the onlycomprehensive guide to 2750 darkplants with 425 stunning photos,available through Ball Publishing atwww.ballpublishing.com. Shefrequently speaks in the U.S.

You can read more about herbooks (all 14 of them) at herwebsite, www.karenplatt.co.uk,and become a member of theInternational Black Plant Society,which Karen founded in 2002 tofurther the knowledge of dark plants,at www.blackplants.co.uk.


coast. They were areas of explorationfor Padre Luis Sodiro, who mademost of his earliest trips into theregion on the back side of VolcánPichincha. Sodiro collectedfrequently in the area between Nonoand Nanegal and between Tandayapaand Mindo as well as betweenNanegal and Pacto.

The best of the ecological reservesin the Mindo area is the BellavistaCloud Forest Reserve – it isastonishingly beautiful. The roadis narrow, rough and lacking anyillumination, but suddenly I sawwhat appeared to be a broad, brightlyilluminated gate extending over theroad. It was quite startling. I foundthe manager and tried to negotiate aroom that I could afford, threateningto drive on to Quito if I couldn’t findone cheap enough. Bellavista hassome regular buildings near theroad, but most are made to looklike gigantic beehives protrudinginto the canopy. The cheapest placesare located on the top floor. The firsttwo floors have stairways, but thethird floor is reached only byclimbing up a ladder. Fortunately,one of the hotel personnel helped mestuff my plants and equipment upthrough the tiny opening thataccessed the third floor.

The room was round with bedsclustered around the periphery.The lighting was not the best, but Imanaged to get the plants describedand investigate the area the nextmorning. Although this area is rightin the center of Luis Sodiro’s oldstomping grounds, and I couldassume most of the Anthuriumspecies were already described bySodiro, I was surprised to find howmany species I did not know. TheBellavista area is laced with trailswith access to most of the region andthere are aroids literally everywhere.

Obviously, more collecting needs tobe done in the area. Among thespecies I saw in this area were:

Anthurium aristatum SodiroA. cordiforme SodiroA. corrugatum SodiroA. giganteum Sodiro,A. longicuspidatum EnglA. malacophyllum SodiroA. mindense SodiroA. ochreatum SodiroA. ovatifolium EnglA. pulverulentum SodiroA. striatipes SodiroA. tremulum SodiroA. umbraculum SodiroA. versicolor Sodiro, andPhilodendron fibrosum

Sodiro ex Croat.

I also collected along the roadbetween Bellavista and Mindo. Thefirst 10 km are quite rich and theroadsides are much more investedwith plants now than ever in the past.For the first few kilometers there aremany points of access into theBellavista property. There is a branchin the road on the right that goesback down to the Mindo Tandayapahighway. It also provides access to aseparate scientific field stationoperated by Bellavista. It is cheaper tostay there, but the facilities are betterat the tourist camp. The place hassolar hot water and lighting, so it is agood place to stay while studying.

That was the last collecting tripfor my 2004 adventure to Ecuador.Like most of the other areas I visited,it was a very rewarding experience. Iam certainly looking forward toworking on the large number ofaroids collected. With five separatearoid florulas tentatively planned forthe western slopes and two additionalaroid florulas planned for the easternslopes of the Andes, there should befairly good coverage of Araceae forthe Flora of Ecuador project. ❦


Figure 18. Stenospermationkillipii Croat.

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Questions?Please contact Scott Hyndmanat [email protected]

and traffic had passed. On the lowerside of the road, the drop off isvertical and you would need a ropeladder to get down. On the upperside of the road it is equally as steep,generally almost straight up.

At one place there was a narrow goatpath snaking up the side of the steepslope. At the top of the cliff itflattened out and the Solanaceousshrub “naranilla” was in cultivation.Aroids remained, including a largeand beautiful member of Anthuriumsection Belolonchium (Fig. 20, 21),and a Philodendron (Fig. 22), thatappeared to be neither P. verrucosumnor P. findens, yet resembled both.

We arrived in La Bonita at dark. Weleft for Santa Barbara the next dayand found the area to be much easiercollecting. This road from La Bonitato Santa Barbara was built manyyears ago and the slopes were not sosteep so that it was easier to get intothe adjacent forests. The area betweenLa Bonita and Santa Barbara was at ahigh elevation and there were fewspecies of Araceae in the region, butlots of interesting vegetation.Although there were few species ofAraceae between La Bonita and SantaBarbarba, we did find Anthuriumumbraculum, what appeared to beA. corrugatum Sodiro, a somewhatthree-lobed Anthurium sect.Belolonchium (Fig. 23), and amember of section Digitinervium.

We arrived at Santa Barbara late inthe day, assuming that we wouldspend the night there, but it provedto be a cold and barren place with noobvious sign of a hotel. Fearing thatit might not be a safe place to stay inany event, we decided to head for themain highway south of Tulcán. Wearrived in Julio Andrade after darkand it was a nasty-looking, crowded,noisy town, so we decided to move

on. We drove south to San Gabriel,searching the main plaza for a hotel,inspected a miserable example andmoved on. By now we weredesperate, assuming that we wouldhave to drive all the way to Ibarra tofind a place to press plants.Fortunately, not too far south of SanGabriel we came across a small area

of Hosterias in a special vacation areathat served Ibarra and Tulcán as aweekend resort. We pulled into aplace called the Hotel Oasis, whichproved to be very nice and we wereable to park right next to the lodging.

The following day we collected alongthe road to Pirampiro in dry,disturbed habitat finding a lot ofinteresting species that I had neverseen before. Just outside of Ibarra weran into trouble with the road beingblocked immediately in front of thetoll station and traffic was turningaround and going back to Ibarra. Iknew from past experience that itwould have been dangerous to try todrive through this crowd of ruffians.Apparently the protesters were

indigenous people demanding moremoney for something. While we weresitting there studying the map foroptions, one of the local people saidthat it was possible to drive down therailroad tracks that paralleled thehighway. This led out to the mainhighway and not only had we gottenpast the protest, but we had avoidedpaying the toll. We felt pretty goodabout that.

With Chris Davidson and GenevieveFerry gone, I set about drying plantsand began the long process of sortingout 3600 collections. During thecourse of processing the plants, Itook a little free time to do two moredays of fieldwork. I drove to ReservaMaquipucuna near Nanegal. TheReserva Maquipucuna has a florawritten and the Araceae is to someextent known. I was not able to spendenough time there to fully evaluatethe aroid flora, but a few hoursallowed me to see the aroids on thelower slopes, leading me to believethat there are species of Araceae notyet reported for the flora. Forexample, Philodendron fibrosumSodiro ex Croat was common, evennear the main station of the reserve,and this was not included in theflora. Additional trips are planned forMaquipucuna to produce a betteraroid florula.

After leaving Maquipucuna, I went tothe Reserva Bellavista, another areathat I plan to study for its aroid flora.Bellavista is located along the oldroad from Nanegalito to Mindo viaTandayapa. I remember this roadfrom trips made in the 1980’s fromQuito to Nono and on to Tandayapa.In those days Tandayapa was themain branch in the road with theroad to the left going up onto theback of Volcán Pichincha and theroad to the right going to Nanegaland Pacto. There was no connectionwith these roads to any area of the

Cardiolonchium (Fig. 19). Anotherside road to the Río Quijos andSan Raphael Falls was blocked bya huge landslide. Later in the daywe reached Lumbaquí and wentnorth on the good asphalt roadthat heads to the Colombian border.After less than 6 km we found aremnant of good forest and collectedseventeen species of Araceae, mostrelatively common to the lowlandAmazonian basin.

On August 20th, we returned to theLumbaquí road intending to make itall the way to La Bonita. At about4:00 p.m. we were stopped by a hugelandslide. Since they were alreadythere with large front-end loadermoving the deep pile of rocks andmud from the road we decided topress plants while we were waiting.Suddenly we heard a large noise andlooked up to see the side of the steepmountainside falling with the treesstill standing mostly erect. This massof rocks soil and trees hit the roadwith such force that I had to assumethat the man operating the front-endloader had to have been buried aliveor carried down into the steep valleybelow. We ran up to the site only

about 50 m ahead of us to findthat the operator had been warnedby one of his coworkers who hadbeen eyeing the forest above the cutand signaled to him to back up assoon as the soil began to loosen. Werealized we were not going anyfurther that day. By that time it wasdark and we had to drive all the wayback to Lumbaquí, where we spentthe night.

The following day we headed backnorth again. By the time we got tothe landslide area it had been cleaned

710



Figure 21. Anthurium section Belolonchium.

Figure 22. Unknown Philodendron.



Figure 23. Somewhat three-lobedAnthurium section Belolonchium.


Figure 19. Unknown Anthuriumsection Cardiolonchium.

Summers are times of greatactivity owing to the extrahelp I sometimes get from

student interns and this year isnot an exception. This summerwe have two interns, Jocelyn Tsaiand Ryan Kirkman. Jocelynapplied for a Howard Hugh’sFellowship through WashingtonUniversity, where she is anundergraduate student, towork with Anthurium sectionPorphyrochitonium. Althoughcompetition was steep, and theaward went to another student,she remained undaunted anddecided to volunteer to contributeto studies in Araceae. She is abright and enthusiastic studentwho has already masteredcharacter recognition in thiscomplicated group repletewith new species. Her goal is tosort out as many new speciesas possible that have adequatematerial to be described.These new species will bepublished in an upcomingpaper treating new species ofsection Porphyrochitonium withher as co-author.

Ryan Kirkman, a high schoolstudent from neighboring Illinois,responded to an article in theRiverfront Times featuringGarden staff. In the article, Italked about the need for moreAraceae research and indicatedthat I didn’t have enough help.

My appeal led Ryan to ask if therewas some way he could help thissummer. We have managed tosort through the backlog of boxesof plants to be confirmed tospecies by matching them withspecimens in the herbarium. Withhis help it has been a veryefficient way to get this massivejob done. He has become skilledat entering specimen data intoTropicos3, the Garden’s electronicdatabase of herbarium specimens,and with our 2005 plant sale infull swing, Ryan has been veryhelpful in organizing plant lists,helping to get plants prepared forpacking, and helping to fill plantorders. Ryan’s upcoming projectfor the summer will involveworking on a database of allAnthurium species names andtheir geographic distribution.This will eventually be placed onthe IAS webpage and will make auseful tool to better understandthe phytogeography ofAnthurium.

Emily Yates, research assistant inAraceae at the Garden, has beenwriting and editing numerousmanuscripts, turning out nearly30 in the last year. As editor of theIAS Newsletter she has organized,edited and written articles, and isinvolved with editorial work forAroideana, making the final editof not only our own papers, butthose of many others before

sending them on to AroideanaEditor Derek Burch. One majorproject Emily has worked onincludes the Araceae treatmentfor the Flora of the Río CenepaRegion in Amazonas Department,Peru. Rodolfo Vasquez, author ofthe flora, is currently editing thismanuscript. A total of 47 newspecies from that flora are alsoin press for a special issue ofthe Brazilian journal Rodriguezia,dealing exclusively with paperson Araceae. When this issuebecomes available IAS memberswill be made aware of how it canbe obtained.

We have only recently receivedmy 2004 collections fromEcuador, totaling 3600 numbers,and Emily has finished insertinglabels, choosing specimens to bemounted for MO, and aggregatingnew species we are working ondescribing. The approximately1000 collections from my 2005trip to Ecuador have not yetbeen shipped.

Another series of Araceae articleswas published in the Annals of theMissouri Botanical Garden 93(3).These papers were first presentedat the VIII International AroidConference in 1999. Other papersfrom this conference are beingpublished in the upcomingvolume of Aroideana. Annals ofthe Missouri Botanical Gardenvolume 93(4) contained

treatments of two aroid genera,a revision of Dractonium byGuanghua Zhu and Tom Croat,and a revision of Dieffenbachia ofCentral America by Tom Croat.

Recently, the revision ofChlorospatha, a long and detailedwork on this interesting genuscompleted by Lynn Hannon, wassubmitted to the Annals. Aboutfive years ago I suggested to Lynnthat she write a key to the existingspecies of Chlorospatha becauseI knew that she had a goodcollection of the genus. Later,I suggested that she go a bitfurther and prepare descriptionsof the species, then one thingled to another until we did afull revision.

It was partly due to Lynn’sefforts with Chlorospatha plusher outstanding ability to growtropical plants that she receivedthe Bette Waterbury Award foroutstanding accomplishmentsin horticulture at the IX Inter-national Aroid Conference inKuching, Sarawak last November.Lynn and I still continue tocollaborate on a variety ofpapers dealing with new speciesof Araceae, especially thoseinvolving our work in Ecuador.

Emily Colletti, who returned twoyears ago to be my greenhousemanager after a 17 year “vacation”to raise her family, has been doingan outstanding job of organizingthe greenhouse. She has beenimproving records, making new,more permanent tags for existingpots, repotting overgrown plants,and reshuffling collections toprovide better organization andimprove growth. I can always relyon her to find something for me.

Our computerized record systemhas never been in better shape.

We have recently gotten thesupport of Amparo Acebey deKrömer to work on the Araceaetreatment for the Flora ofVeracruz. Carola, as she iscommonly called, workedwith me on the treatment ofthe Araceae for the Flora ofBolivian checklist, and is veryknowledgeable about Araceae(among other epiphytic families).She and her husband, Thorsten,recently moved to Mexico, whereThorsten is working on epiphytesat the Los Tuxtlas BiologicalStation in Veracruz State ofMexico. Carola is currentlytrying to get a grant to workon Araceae of Veracruz as wellas for the Araceae of Mesoamerica(that portion of Mexico whichlies below the Isthmus ofTehuantepec).

My student Monica Carlsen,who was awarded the MonroeBirdsey Award two years ago, iscurrently in England workingat Kew on a KLARF award forAnthurium. Monica is doing amolecular study of Anthurium atthe sectional level for her Ph.D.thesis at the University ofMissouri, St. Louis. She and Iwill be presenting a poster onthe sectional classification ofAnthurium at the InternationalBotanical Congress in Vienna,Austria, in July.

Another student, Marcela Mora,last year’s awardee of the MonroeBirdsey Award, will also bepresenting a poster at the sameconference. Marcela was alsoawarded a KLARF award byKew and will be working on

Philodendron both at Kew and atthe Missouri Botanical Garden forthe next year.

Finally, our current efforts areconcentrating on completing theAraceae treatment for the Flora ofMesoamerica. Emily Yates hasalready managed to compile all ofthe species of Central Americathat were previously described,into a single document ready tobe edited for the FloraMesoamericana project.Descriptions for those species notyet described must be prepared,and keys written for genera suchas Monstera (now vastly expandedfrom those treated by MikeMadison in 1977), as well asSpathiphyllum and Xanthosoma,the three most complex genera,which remain unrevised inCentral America.

I hope this discourse about ouractivities here at MO gives yousome appreciation of what goesinto the development of aroidsystematics and how this allowsyour plants to get names. ❦

98

Recent Aroid Activitiesat the Missouri Botanical GardenThomas B. Croat, Missouri Botanical Garden

Summers are times of greatactivity owing to the extrahelp I sometimes get from

student interns and this year isnot an exception. This summerwe have two interns, Jocelyn Tsaiand Ryan Kirkman. Jocelynapplied for a Howard Hugh’sFellowship through WashingtonUniversity, where she is anundergraduate student, towork with Anthurium sectionPorphyrochitonium. Althoughcompetition was steep, and theaward went to another student,she remained undaunted anddecided to volunteer to contributeto studies in Araceae. She is abright and enthusiastic studentwho has already masteredcharacter recognition in thiscomplicated group repletewith new species. Her goal is tosort out as many new speciesas possible that have adequatematerial to be described.These new species will bepublished in an upcomingpaper treating new species ofsection Porphyrochitonium withher as co-author.

Ryan Kirkman, a high schoolstudent from neighboring Illinois,responded to an article in theRiverfront Times featuringGarden staff. In the article, Italked about the need for moreAraceae research and indicatedthat I didn’t have enough help.

My appeal led Ryan to ask if therewas some way he could help thissummer. We have managed tosort through the backlog of boxesof plants to be confirmed tospecies by matching them withspecimens in the herbarium. Withhis help it has been a veryefficient way to get this massivejob done. He has become skilledat entering specimen data intoTropicos3, the Garden’s electronicdatabase of herbarium specimens,and with our 2005 plant sale infull swing, Ryan has been veryhelpful in organizing plant lists,helping to get plants prepared forpacking, and helping to fill plantorders. Ryan’s upcoming projectfor the summer will involveworking on a database of allAnthurium species names andtheir geographic distribution.This will eventually be placed onthe IAS webpage and will make auseful tool to better understandthe phytogeography ofAnthurium.

Emily Yates, research assistant inAraceae at the Garden, has beenwriting and editing numerousmanuscripts, turning out nearly30 in the last year. As editor of theIAS Newsletter she has organized,edited and written articles, and isinvolved with editorial work forAroideana, making the final editof not only our own papers, butthose of many others before

sending them on to AroideanaEditor Derek Burch. One majorproject Emily has worked onincludes the Araceae treatmentfor the Flora of the Río CenepaRegion in Amazonas Department,Peru. Rodolfo Vasquez, author ofthe flora, is currently editing thismanuscript. A total of 47 newspecies from that flora are alsoin press for a special issue ofthe Brazilian journal Rodriguezia,dealing exclusively with paperson Araceae. When this issuebecomes available IAS memberswill be made aware of how it canbe obtained.

We have only recently receivedmy 2004 collections fromEcuador, totaling 3600 numbers,and Emily has finished insertinglabels, choosing specimens to bemounted for MO, and aggregatingnew species we are working ondescribing. The approximately1000 collections from my 2005trip to Ecuador have not yetbeen shipped.

Another series of Araceae articleswas published in the Annals of theMissouri Botanical Garden 93(3).These papers were first presentedat the VIII International AroidConference in 1999. Other papersfrom this conference are beingpublished in the upcomingvolume of Aroideana. Annals ofthe Missouri Botanical Gardenvolume 93(4) contained

treatments of two aroid genera,a revision of Dractonium byGuanghua Zhu and Tom Croat,and a revision of Dieffenbachia ofCentral America by Tom Croat.

Recently, the revision ofChlorospatha, a long and detailedwork on this interesting genuscompleted by Lynn Hannon, wassubmitted to the Annals. Aboutfive years ago I suggested to Lynnthat she write a key to the existingspecies of Chlorospatha becauseI knew that she had a goodcollection of the genus. Later,I suggested that she go a bitfurther and prepare descriptionsof the species, then one thingled to another until we did afull revision.

It was partly due to Lynn’sefforts with Chlorospatha plusher outstanding ability to growtropical plants that she receivedthe Bette Waterbury Award foroutstanding accomplishmentsin horticulture at the IX Inter-national Aroid Conference inKuching, Sarawak last November.Lynn and I still continue tocollaborate on a variety ofpapers dealing with new speciesof Araceae, especially thoseinvolving our work in Ecuador.

Emily Colletti, who returned twoyears ago to be my greenhousemanager after a 17 year “vacation”to raise her family, has been doingan outstanding job of organizingthe greenhouse. She has beenimproving records, making new,more permanent tags for existingpots, repotting overgrown plants,and reshuffling collections toprovide better organization andimprove growth. I can always relyon her to find something for me.

Our computerized record systemhas never been in better shape.

We have recently gotten thesupport of Amparo Acebey deKrömer to work on the Araceaetreatment for the Flora ofVeracruz. Carola, as she iscommonly called, workedwith me on the treatment ofthe Araceae for the Flora ofBolivian checklist, and is veryknowledgeable about Araceae(among other epiphytic families).She and her husband, Thorsten,recently moved to Mexico, whereThorsten is working on epiphytesat the Los Tuxtlas BiologicalStation in Veracruz State ofMexico. Carola is currentlytrying to get a grant to workon Araceae of Veracruz as wellas for the Araceae of Mesoamerica(that portion of Mexico whichlies below the Isthmus ofTehuantepec).

My student Monica Carlsen,who was awarded the MonroeBirdsey Award two years ago, iscurrently in England workingat Kew on a KLARF award forAnthurium. Monica is doing amolecular study of Anthurium atthe sectional level for her Ph.D.thesis at the University ofMissouri, St. Louis. She and Iwill be presenting a poster onthe sectional classification ofAnthurium at the InternationalBotanical Congress in Vienna,Austria, in July.

Another student, Marcela Mora,last year’s awardee of the MonroeBirdsey Award, will also bepresenting a poster at the sameconference. Marcela was alsoawarded a KLARF award byKew and will be working on

Philodendron both at Kew and atthe Missouri Botanical Garden forthe next year.

Finally, our current efforts areconcentrating on completing theAraceae treatment for the Flora ofMesoamerica. Emily Yates hasalready managed to compile all ofthe species of Central Americathat were previously described,into a single document ready tobe edited for the FloraMesoamericana project.Descriptions for those species notyet described must be prepared,and keys written for genera suchas Monstera (now vastly expandedfrom those treated by MikeMadison in 1977), as well asSpathiphyllum and Xanthosoma,the three most complex genera,which remain unrevised inCentral America.

I hope this discourse about ouractivities here at MO gives yousome appreciation of what goesinto the development of aroidsystematics and how this allowsyour plants to get names. ❦

98

Recent Aroid Activitiesat the Missouri Botanical GardenThomas B. Croat, Missouri Botanical Garden

and traffic had passed. On the lowerside of the road, the drop off isvertical and you would need a ropeladder to get down. On the upperside of the road it is equally as steep,generally almost straight up.

At one place there was a narrow goatpath snaking up the side of the steepslope. At the top of the cliff itflattened out and the Solanaceousshrub “naranilla” was in cultivation.Aroids remained, including a largeand beautiful member of Anthuriumsection Belolonchium (Fig. 20, 21),and a Philodendron (Fig. 22), thatappeared to be neither P. verrucosumnor P. findens, yet resembled both.

We arrived in La Bonita at dark. Weleft for Santa Barbara the next dayand found the area to be much easiercollecting. This road from La Bonitato Santa Barbara was built manyyears ago and the slopes were not sosteep so that it was easier to get intothe adjacent forests. The area betweenLa Bonita and Santa Barbara was at ahigh elevation and there were fewspecies of Araceae in the region, butlots of interesting vegetation.Although there were few species ofAraceae between La Bonita and SantaBarbarba, we did find Anthuriumumbraculum, what appeared to beA. corrugatum Sodiro, a somewhatthree-lobed Anthurium sect.Belolonchium (Fig. 23), and amember of section Digitinervium.

We arrived at Santa Barbara late inthe day, assuming that we wouldspend the night there, but it provedto be a cold and barren place with noobvious sign of a hotel. Fearing thatit might not be a safe place to stay inany event, we decided to head for themain highway south of Tulcán. Wearrived in Julio Andrade after darkand it was a nasty-looking, crowded,noisy town, so we decided to move

on. We drove south to San Gabriel,searching the main plaza for a hotel,inspected a miserable example andmoved on. By now we weredesperate, assuming that we wouldhave to drive all the way to Ibarra tofind a place to press plants.Fortunately, not too far south of SanGabriel we came across a small area

of Hosterias in a special vacation areathat served Ibarra and Tulcán as aweekend resort. We pulled into aplace called the Hotel Oasis, whichproved to be very nice and we wereable to park right next to the lodging.

The following day we collected alongthe road to Pirampiro in dry,disturbed habitat finding a lot ofinteresting species that I had neverseen before. Just outside of Ibarra weran into trouble with the road beingblocked immediately in front of thetoll station and traffic was turningaround and going back to Ibarra. Iknew from past experience that itwould have been dangerous to try todrive through this crowd of ruffians.Apparently the protesters were

indigenous people demanding moremoney for something. While we weresitting there studying the map foroptions, one of the local people saidthat it was possible to drive down therailroad tracks that paralleled thehighway. This led out to the mainhighway and not only had we gottenpast the protest, but we had avoidedpaying the toll. We felt pretty goodabout that.

With Chris Davidson and GenevieveFerry gone, I set about drying plantsand began the long process of sortingout 3600 collections. During thecourse of processing the plants, Itook a little free time to do two moredays of fieldwork. I drove to ReservaMaquipucuna near Nanegal. TheReserva Maquipucuna has a florawritten and the Araceae is to someextent known. I was not able to spendenough time there to fully evaluatethe aroid flora, but a few hoursallowed me to see the aroids on thelower slopes, leading me to believethat there are species of Araceae notyet reported for the flora. Forexample, Philodendron fibrosumSodiro ex Croat was common, evennear the main station of the reserve,and this was not included in theflora. Additional trips are planned forMaquipucuna to produce a betteraroid florula.

After leaving Maquipucuna, I went tothe Reserva Bellavista, another areathat I plan to study for its aroid flora.Bellavista is located along the oldroad from Nanegalito to Mindo viaTandayapa. I remember this roadfrom trips made in the 1980’s fromQuito to Nono and on to Tandayapa.In those days Tandayapa was themain branch in the road with theroad to the left going up onto theback of Volcán Pichincha and theroad to the right going to Nanegaland Pacto. There was no connectionwith these roads to any area of the

Cardiolonchium (Fig. 19). Anotherside road to the Río Quijos andSan Raphael Falls was blocked bya huge landslide. Later in the daywe reached Lumbaquí and wentnorth on the good asphalt roadthat heads to the Colombian border.After less than 6 km we found aremnant of good forest and collectedseventeen species of Araceae, mostrelatively common to the lowlandAmazonian basin.

On August 20th, we returned to theLumbaquí road intending to make itall the way to La Bonita. At about4:00 p.m. we were stopped by a hugelandslide. Since they were alreadythere with large front-end loadermoving the deep pile of rocks andmud from the road we decided topress plants while we were waiting.Suddenly we heard a large noise andlooked up to see the side of the steepmountainside falling with the treesstill standing mostly erect. This massof rocks soil and trees hit the roadwith such force that I had to assumethat the man operating the front-endloader had to have been buried aliveor carried down into the steep valleybelow. We ran up to the site only

about 50 m ahead of us to findthat the operator had been warnedby one of his coworkers who hadbeen eyeing the forest above the cutand signaled to him to back up assoon as the soil began to loosen. Werealized we were not going anyfurther that day. By that time it wasdark and we had to drive all the wayback to Lumbaquí, where we spentthe night.

The following day we headed backnorth again. By the time we got tothe landslide area it had been cleaned

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Figure 22. Unknown Philodendron.



Figure 23. Somewhat three-lobedAnthurium section Belolonchium.


Figure 19. Unknown Anthuriumsection Cardiolonchium.

apart so many times that it eventuallybroke, so on the way down to Baezawe stopped at a automobile partsplace and bought a new terminal.When I removed the terminal end Idiscovered that the problem had notbeen terminal itself, but theconnection. Putting on the new endsolved the problems permanently.

We stayed at a new place in Baezaand found it well protected withabout half the rooms filled withsoldiers. It was a nifty place whereyou could drive inside and park rightnext to the room. The last time I was

in Baeza I stayed at a place madeentirely our of wood and nearlycaught it on fire with my plant dryer.Since it is no longer there, I suspectsomeone may have succeeded inburning it down.

The following day we drove to LagoAgrio, collecting along the way. Theroute is much more denuded sincethe last time I was on the road, butthere are still patches of forest witharoids. One of these areas is the roadthat goes from the main highway tothe small village of Gonzales Díaz dePineda at the gateway to the ParqueNacional Sumaco Napo Galeras. Thetown lies east of the Río Quijos andthe area between the highway and theriver was quite rich in aroids. Amongthe collections made was of a newspecies of Anthurium sectionCardiolonchium, Anthuriumamoenum Kunth, Philodendronasplundii Croat & M. L. Soares(Fig. 16), P. palacioanum Croat &Grayum, P. ruizii Schott, P. schmidtiaeCroat (Fig. 17), Rhodospatha latifoliaPoepp., Stenospermation killipii Croat(Fig. 18), Xanthosomapurpureomaculatum Croat, and anunknown Anthurium in section


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Figure 16. Philodendron asplundiiCroat & M.L. Soares.

Figure 17. Philodendronschmidtiae Croat.

Continued on page 7

that I receive the most phonecalls about. The conversationusually goes something likethis: ‘I have this plant thatsmells like something iswrong with the drains...’ I findthis frustrating; after all,animal lovers get used to thesmell of horses, cows andpigs. The beauty of theseplants far outshines anyoffence to the nose.

I just missed the flowering ofAmorphophallus titanum atKew by two days. I’ll regretthis for the rest of my days.No other plant is sooutrageous or flamboyant.

Having just started up mynursery again, (I have theaccolade of being the veryfirst person to run a nurseryspecializing in black plants), Ihope to be introducing manyspecies into the U.K. As moreof these plants are introducedto the West, one realises thewealth and magnitude of darkaroids and just how easymost of them are to grow.

———————————

Karen Platt is author of Black Magicand Purple Passion, the onlycomprehensive guide to 2750 darkplants with 425 stunning photos,available through Ball Publishing atwww.ballpublishing.com. Shefrequently speaks in the U.S.

You can read more about herbooks (all 14 of them) at herwebsite, www.karenplatt.co.uk,and become a member of theInternational Black Plant Society,which Karen founded in 2002 tofurther the knowledge of dark plants,at www.blackplants.co.uk.


coast. They were areas of explorationfor Padre Luis Sodiro, who mademost of his earliest trips into theregion on the back side of VolcánPichincha. Sodiro collectedfrequently in the area between Nonoand Nanegal and between Tandayapaand Mindo as well as betweenNanegal and Pacto.

The best of the ecological reservesin the Mindo area is the BellavistaCloud Forest Reserve – it isastonishingly beautiful. The roadis narrow, rough and lacking anyillumination, but suddenly I sawwhat appeared to be a broad, brightlyilluminated gate extending over theroad. It was quite startling. I foundthe manager and tried to negotiate aroom that I could afford, threateningto drive on to Quito if I couldn’t findone cheap enough. Bellavista hassome regular buildings near theroad, but most are made to looklike gigantic beehives protrudinginto the canopy. The cheapest placesare located on the top floor. The firsttwo floors have stairways, but thethird floor is reached only byclimbing up a ladder. Fortunately,one of the hotel personnel helped mestuff my plants and equipment upthrough the tiny opening thataccessed the third floor.

The room was round with bedsclustered around the periphery.The lighting was not the best, but Imanaged to get the plants describedand investigate the area the nextmorning. Although this area is rightin the center of Luis Sodiro’s oldstomping grounds, and I couldassume most of the Anthuriumspecies were already described bySodiro, I was surprised to find howmany species I did not know. TheBellavista area is laced with trailswith access to most of the region andthere are aroids literally everywhere.

Obviously, more collecting needs tobe done in the area. Among thespecies I saw in this area were:

Anthurium aristatum SodiroA. cordiforme SodiroA. corrugatum SodiroA. giganteum Sodiro,A. longicuspidatum EnglA. malacophyllum SodiroA. mindense SodiroA. ochreatum SodiroA. ovatifolium EnglA. pulverulentum SodiroA. striatipes SodiroA. tremulum SodiroA. umbraculum SodiroA. versicolor Sodiro, andPhilodendron fibrosum

Sodiro ex Croat.

I also collected along the roadbetween Bellavista and Mindo. Thefirst 10 km are quite rich and theroadsides are much more investedwith plants now than ever in the past.For the first few kilometers there aremany points of access into theBellavista property. There is a branchin the road on the right that goesback down to the Mindo Tandayapahighway. It also provides access to aseparate scientific field stationoperated by Bellavista. It is cheaper tostay there, but the facilities are betterat the tourist camp. The place hassolar hot water and lighting, so it is agood place to stay while studying.

That was the last collecting tripfor my 2004 adventure to Ecuador.Like most of the other areas I visited,it was a very rewarding experience. Iam certainly looking forward toworking on the large number ofaroids collected. With five separatearoid florulas tentatively planned forthe western slopes and two additionalaroid florulas planned for the easternslopes of the Andes, there should befairly good coverage of Araceae forthe Flora of Ecuador project. ❦


Figure 18. Stenospermationkillipii Croat.

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Questions?Please contact Scott Hyndmanat [email protected]

Common names: laitue d’eau,pistie (French); Lechuguillade agua, lechuguita de agua,

repollo de agua (Spanish); waterlettuce, tropical duckweed, shellflower(English).

Pistia stratiotes, or water lettuce, is aninteresting, unique aroid. Like anypotential invasive species, it must bekept in check so it does not becomedestructive; however, it has valuableproperties, as a medicinal, a foodsource, and environmental remediator.

The name Pistia stratiotes comes fromthe Greek “pister” meaning hollowtrough, in the sense of a drinkingtrough (Mayo et al. 1997) or the Greek“pistos” meaning water; and “stratiotes”meaning soldier. It was classified in1943 to its own Pistiaceae family, but asa result of later research it is now in theAraceae, subfamily Aroideae, tribePistieae, and is the only species in thegenus Pistia.

Water lettuce is a perennial monocot ofthe Araceae family. It’s fairly

unmistakable; it looks like floatinglettuce (Fig. 1). Many descriptions ofthe plant abound: see Mayo et al. 1997,Bown, 1988.

There is some confusion about theorigin of water lettuce due to itsworldwide distribution, however, thereis strong evidence that it is native toSouth America (Rivers, 2002). Pistiastratiotes is now found throughout thetropics and subtropics (Glazier, 1996),and is one of the most widelydistributed hydrophytes in the tropics(Holm et al. 1977). Pistia stratiotesrequires a wet, temperate to tropicalhabitat (Rivers, 2002).

Pistia stratiotes may have beenintroduced to North America bynatural means or by humans. It wasspied as early as 1774 by WilliamBartram, in “vast quantities ... severalmiles in length, and in some places aquarter of a mile in breadth” in the St.Johns River in Florida (APIRS Online).It has been suggested that trade via St.Augustine, founded in 1565, may have

provided an early avenue forintroduction into the St. Johnswatershed (APIRS Online).

Pistia stratiotes is a “state listed,”“prohibited,” “category I,” “species ofconcern” or “noxious” plant in at leasteight states including CA, AZ, TX, LA,SC, GA, FL, HI and Puerto Rico. Whilethe abundance of water lettuce isrecorded primarily in sub-tropicalregions of the US, it has beendiscovered in northern states, includingOhio and New York (Dray Jr. andCenter, 2002). Several states imposefines for individuals who posses orparticipate in the sale or distribution ofwater lettuce, including South Carolinaand Texas. The Connecticut Dept. ofEnvironmental Protection will addPistia stratiotes to the list of prohibitedspecies in October, 2005 and will fineindividuals who possess it $100 perplant (CDEP, 2005). It occured in 68public water bodies in Florida by 1982and in 128 water bodies by 1989, buttotal abundance reduced by half overthe same time period as a result of astatewide management program(APIRS Online).

Pistia is common in dams, lagoons,lakes, and also grows in wetland rice. Itcan also be found on stagnant waterand rooting on muddy banks. It’s oftengrown in ponds and aquaria, and canbecome a serious invasive pest in thetropics. Its capable of forming vastmats that disrupt submersed plant andanimal communities and interfere withwater movement and navigation (Holmet al.1977); also serves as host for atleast two genera of mosquitoes (Holmet al. 1977). It is considered a seriousweed in Ceylon, Ghana, Indonesia, andThailand and at least present as a weedin 40 other countries (Holm et al.1977). It is listed by the NatureConservancy’s Invasive SpeciesInitiative as a pest in Australia(http://tncweeds.ucdavis.edu/global/australia/aca.html), and has been a target

Sodiro, A. obscurinervia Croat,A. palenquense Croat, A. pallidiflorumEngl., A. versicolor, Chlorospathadodsonii Madison (Fig. 13),Dieffenbachia sp., Monstera cf.adansonii Schott, Philodendronacuminatissimum Engl.,P. platypetiolatum Madison,P. purpurascens Engl., P. tenue K.Koch, P. tenuipes Engl., P. umbricolumEngl., Rhodospatha dodsonii Croat,Syngonium macrcarpum Engl. andXanthosoma sp. One species ofAnthurium is considered new,A. samanaense Croat (Fig. 14), as wellas two new species of Philodendron.One being (Fig. 15), reasonablycommon on the Pacific slope alsonear Piñas and Guasaganda. Clearlymore time will have to be spent atCerro Samana before any florula canbe prepared.

On the way back to Quito wecollected at the UniversidadPolytechnica Salesiana’s EstacíonSacha Wiwa. This is another logicalsite for an aroid florula andcollections began there with this inmind. Among the species here not yetseen on the Pacific slope in southernEcuador were Anthurium laciniosumSodiro, A. propinquum Sodiro,A. pulverulentum Sodiro, A. urbaniiSodiro, Philodendron inequilaterumLiebm. and P. rugapetiolum.

Next we headed east and into theEcuadorian Oriente. Aftercontinuously having trouble startingthe Mazda pickup for over a month itseem to be getting worse. Afterdriving around by taxi to severalneighborhoods we finally foundsome kid claiming to be a mechanic.He arrived with virtually no toolsand ended up pouring a bucket ofwater over the battery to get betterground. I had taken the battery cable


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Continued on page 13 Continued on page 6

hardly contain myself when Ialso learned of purple and redfoliaged forms of somespecies from China. I love theelegance of Arisaemasikokianum aptly described byone visitor to my booth at theChelsea flower show in 2004as “like one of Cecil Beaton’shats in My Fair Lady.” It isprobably the most beautiful ofall Arisaemas, a truearistocrat. However, I am alsotaken by the deeply colouredspathes of A. speciosum andmany others.

I keep both of these slightlymoist in their pots out in thegarden over winter (Z7). Thisyear, for the first time in many,we had a lot of snow. Ithought I might have lostthem, as they wereunprotected. Thankfully, theycame through unharmed.Arisaema sikokianum finishedflowering in May just asA. speciosum was breakingdormancy. They both grow inlight shade. Arisaema griffithiiis another favourite, thepatterning on this species isincredible. My favoured Arumis palaestinum, for I believe itis the darkest, but this is hardto come by. As with manyaroids, the smell is notoriousbut I have learned that it oftendepends on provenance. Ifirst discovered this whiletalking to a journalist inOregon whose family hadgrown Dracunculus vulgaris,once classified as Arum, foralmost 100 years and heclaimed it had no off-puttingsmell at all. Yet it is the plant


Figure 13. Chlorospathadodsonii Madison.

Figure 15. New speciesof Philodendron.

Figure 14. Anthuriumsamanaense Croat.

Figure 1. Pistia stratiotes L. Croat 85014. Photo by Tom Croat.

Pistia stratiotes L. (Araceae) –Ubiquitous Aroid InvaderBy Emily D. Yates, Missouri Botanical Gardenand Kristen E. Kordecki, Chicago Botanic Garden

of management research and control inFlorida for at least two decades.

As a case study of invasion biology,Pistia stratiotes is intriguing. Rivers(2002) states that P. stratiotesreproduces vegetatively and by seed.Rapid vegetative reproduction allowswater lettuce to cover an entire lake,from shore to shore, with a dense matof connected rosettes in a short periodof time. This type of rapidreproduction makes Pistia aparticularly effective invader, wheredensities of rosettes can range from lessthan 100 to over 1,000 per m2 in southFlorida (APIRS Online). Seedproduction by P. stratiotes, oncethought not to occur in NorthAmerica, is now considered importantto reproduction and dispersal (Drayand Center 1989). While Pistia is notcold tolerant (Holm et al. 1977), itsleaves re-grow after moderate freezes,and it can survive for extended periodsof time on moist muck, sandbars, andbanks (Holm et al. 1977), contributingto its success as an invader.

Pistia stratiotes can spread from wholeplants or pieces of plants moved onboats or fishing equipment from aninfested to a clean body of water(Rivers, 2002). The most commonlyaccepted pathway of this species intothe United States is in ballast water inships from South America. Ships thattravel through mats of water lettuce cancarry fertile plant segments to newareas. Once established within a waterbody, P. stratiotes can be spread bywater currents and floods.

According to Ramey (2001), P. stratiotescontinues to be sold through aquariumsupply dealers and through theinternet. Rivers (2002) cites thatdumping of aquarium or ornamentalpond plants is often the means ofspread for P. stratiotes. The popularityof water lettuce as a garden plant has

also led to its spread, in a morecontrolled manner.

Although there are several other aroidsconsidered invasive in various regions,P. stratiotes is by far the most insidious.According to Pimentel et al. (2005)Pistia stratiotes, among several invasiveaquatic plant species, is altering fishand other aquatic animal species,choking waterways, altering nutrientcycles and reducing recreational use ofrivers and lakes. In a study published in2000, Pimentel et al. determinedinvasive aquatic plants species costs $10million in losses and damages and $100million in control costs in the US. In1994, Pistia stratiotes control cost wasestimated at $650,000 annually (Center,1994) in Florida, where it is currentlyunder “maintenance control.” Thisexcludes costs associated withinterference with recreationalopportunities, like swimming andfishing, and impacts associated withinsects that breed in waterways chokedby water lettuce. Dray Jr. and Center(2002) estimated expendituresincreased to over $1 million dollarsannually in Florida by 2002.

Like Water Hyacinth, Pistia also blocksirrigation canals and provides abreeding ground for mosquitoes andclogs fishery waters. Pistia stratiotes caninflict a severe impact on theenvironment and economy of infestedareas (Rivers, 2002). The dense matscreated by connected rosettes of theplant lead to the majority of problemsencountered with water lettuce. Thesemats can have a negative economiceffect by blocking waterways, thusincreasing the difficulty of navigation,boat traffic, fishing, and hinderingflood control efforts. It may become animportant constituent of a sudd – adense aggregation of a free-floatingvegetation. Pistia stratiotes even playeda role in the infamous Dr. Livingstone’sexplorations in Africa, where, whileexploring the river Shire, a northerntributary of the Zambesi, they learned

of a previous attempt by thePortuguese to ascend the river. ThePortuguese had to abandon the trip onaccount of the impenetrable duckweed(Pistia stratiotes), makingDr. Livingstone’s expedition up theriver Shire the first by Europeans(Livingstone, website).

Mats of P. stratiotes can also disruptnatural ecosystems. They can lead to alower concentration of oxygen incovered waters and sediments byblocking air-water interface and rootrespiration. They can degrade waterquality by blocking the air-waterinterface and greatly reducing oxygenlevels in the water, eliminatingunderwater animals such as fish.Extremely thick mats of P. stratiotes canprevent sunlight from reachingunderlying water, possibly eliminatingnative submerged plants. Thecumulative effect of these negativecharacteristics of the plant is a loss ofbiodiversity in invaded habitats. Animalcommunities may be altered byblocking access to the water and/oreliminating plants the animals dependon for shelter and nesting. P. stratiotesmats can also serve as a breeding placefor mosquitoes. It reproducesexponentially and reduces the oxygenlevels in a body of water by reducingthe water’s surface area (Dunn, 1934).

Mechanical controls, like the flailchopper or harvester, have beenused to control populations of waterlettuce, but this method is costly.Biological control is under investi-gation. The South American weevil,Neohydronomus affinis, can reducepopulations, but once introduced, theweevils themselves continue to spread(http://aquat1.ifas.ufl.edu/lettuce.html).The moth, Samea multiplicalis, whichattacks P. stratiotes and Salvinia spp.,has been established in Australia, but itsimpact has not been evaluated. InThailand, classical biological controlhas not been attempted, but mass

BookReview...Black Magicand PurplePassion:Black AroidsBy Karen Platt

Aroids are superb in anycolour, but it is amazing howmany of them approachblack. It is understood thatwhen we talk about “black”plants we are encompassingthe darkest colors found inhorticulture – from bronze, topurple to near black.

I have been researching darkplants for over ten years.Aroids being my favouritegroup of plants, I’ve amasseda lot of information on them,chiefly Arisaemas and Arums.I am fascinated by thespathes and spadices ofthese plants, let alone theirmagnificent foliage. I could

blades over a meter long and stubbypurplish red spadices with a broadhooding spathe (Figs. 10 & 11).

We made it back to Quito the sameday and began pressing thecollections. Chris Davidson hadarrived the previous day and wehurried to get back to the field. Ourfirst field trip with Chris was alongthe Quito-to-Santo Domingo road,departing at Chilligallo and ending inChiriboga. Two species in this regionwere a large Anthurium with longinternodes and cordate blades withconspicuously sunken tertiary veins,and a very small Anthurium with a

stipitate spadix. We arrived atTinalandia, where we spent the nightpressing plants on the front porch ofthe lodges overlooking the Río Pilatonnot far below the mouth of the RíoToachi. I finished pressing plants thefollowing day while Chris collected inthe forest around Tinalandia.

We left the next day for Ventanas andstayed in Babahoya, where we metXavier Cornejo in the morning. It wasa miserable hotel with roosterscrowing outside the window. The nextmorning we headed to HaciendaClementina, headquarters of the largebanana industry around Babahoya.We got permission to collect at CerroSamana, the ultimate goal of the trip.

Cerro Samana is an isolated hill about750 m elevation – one of the fewremaining areas at this elevation onthe western slope of the Andes. It willbe another study site for an aroidflorula, so collecting there wasintense. Species found wereAnthurium angamarcanum Sodiro,A. argyrostachyum Sodiro, A. asplundiiCroat, A. brachypodum Sodiro,A. cupulispatha Croat & J. Rodríguez(Fig. 12), A. dolichostachyum,A. guayaquilense Engl., A. ochreatum

4 13

Continued on page 5

Continued on page 5

Figures 10 and 11. New species of Anthurium section Belolonchiumwith blades over 1 m long.

ECUADOR 2004: PART II Continued from page 3 PISTIA STRATIOTESContinued from page 12


Figure 12. Anthurium cupulispathaCroat & J. Rodríguez.

rearing and release of the nativenoctuid moth Spodoptera pectinicornishas replaced the use of herbicides.Research indicates this biologicalcontrol is not as useful as originallyhypothesized (Dray Jr. et al., 2001).Chemical control can be effective, butcan be accompanied with its own set ofproblems such as bioaccumulation inbivalves, fish and other fauna in thefood chain (Colborn and Thayer,2000).

Though Pistia stratiotes can be invasive,it is also an oxygenator for fish, andhelps keep pond water clear. Thespecies provides food for manatees(although Dunn, 1934, states that thisnatural predator cannot keep Pistia incheck). Its roots can provide breedingand hiding space for small aquaticanimals.

Pistia stratiotes is reported to absorblarge amounts of heavy metals fromwater and are being used inremediation and wastewater treatmentefforts because of their ability toincrease water quality (Smirnova &Mironova, 2004). In a comparativestudy of 10 water plants, Pistia stratiotesshowed one of the highest removalrates for zinc and lead. These heavymetals were found in or adsorbed intothe roots (Awuah et al., 2001).

Pistia stratiotes has also been used as a“famine food” – plants that are notnormally considered as crops, but areconsumed in times of famine (FamineFoods Database). In India it was usedas a famine food in 1877-1878. InChina, young leaves are eaten cooked.

In traditional medicine, dried waterlettuce or tropical duckweed is burntinto an ash, then moistened with adrop of mustard oil, and applied in athin layer on chronic, enlarged tonsils,twice a day to help get rid of tonsillitis.(www.bestincosmetics.com). In the

Philippines, it is used to treatgonorrhea. Pistia stratiotes leaves areused in traditional medicine for thetreatment of ringworm infection of thescalp, syphilitic eruptions, skininfections, boils, and wounds, becauseit possesses antifungal properties(Premkumar & Shyamsundar, 2005).The oil extract of P. stratiotes is used inthe treatment of worm infestations,tuberculosis, asthma, and dysentery,and is applied externally to treat skindiseases, inflammation, piles, ulcers,syphilitic infections and burns (Kirtikar& Basu, 2000). Pistia stratiotes is listedon the checklist of medicinal plants inSoutheast Asia as effective against boilsand ezcema (www.arcbc.org).

Pistia stratiotes has a high potashcontent (Burkill, 1935). The extractionof caustic potash from wood ash for useas an emulsifier is probably one of themost ancient ethnobotanic traditions(e.g., Quiapo or Pistia stratiotes, one ofthe Araceae family; Water Hyacinth orEichhornia crassipes, of thePontederiaceae family; the Royal fern orOsmunda regalis, the fruit of the PricklyChaff flower or Achyranthes aspera ofthe Ameranthaceae family.(www.dweckdata.com). However, likeother Araceae, it is poisonous if ingestedbecause of the calcium oxalate crystals.

References:

APIRS Online, the database of aquatic,wetland and invasive plants, Centerfor Aquatic and Invasive Plants,The University of Florida, IFAShttp://plants.ifas.ufl.edu/search80/NetAns2/

Awuah, E., F. Anohene, K. Asante, H.Lubberding and H. Gijzen. 2001.Environmental conditions andpathogen removal in macrophyte-and algal-based domestic wastewatertreatment systems. Water Science &Technology 44 (6):11–18.

Bown, D. 1988. Aroids. Plants of theArum Family. Century HutchisonPress. London.

Burkill, I. H. 1935. Dictionary of theEconomic Products of the MalayPeninsula. Oxford, England: OxfordUniversity Press.

Center, T.D. 1994. Biological Control ofWeeds: Water Hyacinth and WaterLettuce, pp. 481-521. In: PestManagement in the Tropics:Biological Control – A FloridaPerspective, Rosen, D., F.D. Bennettand J.L. Capinera (eds.). InterceptLtd., Andover, UK.

Colborn, T. and K. Thayer. 2000.Aquatic Ecosystems: Harbingers ofEndocrine Disruption. EcologicalApplications, 10 (4): 949-957.

Connecticut Dept. of EnvironmentalProtection. June 27, 2005.http://dep.state.ct.us/burnatr/fishing/fishinfo/anguide_part1.pdf.

Dray Jr., F., T. D. Center & G.S.Wheeler. 2001. Lessons fromUnsuccessful Attempts to EstablishSpodoptera pectinicornis(Lepidoptera: Noctuidae), aBiological Control Agent of Waterlettuce.” Biocontrol Science &Technology. 11(3): 301-317.

Dray, Jr., T.D. Center. 2002-2004. WaterLettuce, pp. 65-78. In: BiologicalControl of Invasive Plants in theEastern United States, R. VanDriesche (ed.).

Dunn, L. H. 1934. Notes on the WaterLettuce, Pistia stratiotes Linn. as aNursery of Insect Life. Ecology 15(3): 329-331.

Famine Foods Database:www.hort.purdue.edu.

Glazier, K. 1996.Pistia stratiotes L.Ecology and Evolutionary BiologyConservatory. www.eeb.uconn.edu.

Holm, L. G., D. L. Plucknett, J. V.Pancho, and J. P. Herberger. 1977.The world’s worst weeds:distribution and biology. East-WestCenter/University Press of Hawaii.pp. 379-384

14 3

Croat & M. Mora, P. rugapetiolatumCroat (Fig. 4), P. subhastatum Engl.and P. verrucosum Mathieu ex Schott.Also present is a new species ofPhilodendron (Fig. 5) with a narrowblade similar to P. tenue K. Koch.

Known species found in the El Chicalregion that were not found in theLita-San Lorenzo region includeA. ovatifolium Engl., A. pulverulentumSodiro var. adsimile (Sodiro) Croat &Rodríguez, P. fibrosum Sodiro exCroat, P. oligospermum Engl. andStenospermation densiovulatum Engl.Many more species that occur in theEl Chical region are still unnamedand new to science. One is a long-bladed member of Anthurium sectionPolyneurium similar to Anthuriumcuspidatum Masters, as well as amember of Anthurium sect.Calomystrium with thick ovate bladesand free basal veins, sometimes evena peltate blade (Fig. 6). Another was adeeply three-lobed member ofAnthurium sect. Belolonchium similarto Anthurium herthae K. Krause fromthe Oriente of Ecuador (Fig. 7).Several new species of Philodendronwere found including one with athick, ovate blade and deciduouscataphylls (Fig. 8).

On the return to Tulcán, we madenumerous stops at elevations thatcontained Araceae, collecting manyinteresting and new species. Themajority were in flower and large setswere collected for type specimens.Between El Chical and the Paramodel Angel, ranging from 1200 to2000 m, many species of Anthuriumsection Belolonchium were found,most new to science. One speciesnear Maldonado has long internodesand stood more than two meters tall(Fig. 9). Higher up we found a largeAnthurium section Belolonchium with


Figure 4. Philodendronrugapetiolatum Croat.

Figure 5. New species of Philodendronwith a narrow blade.

Figure 6. New species of Anthuriumsection Calomystrium.

Continued on page 4

Figure 7. New deeply three-lobedAnthurium section Belolonchium.

Figure 8. New species ofPhilodendron.

Figure 9. New species of Anthuriumsection Belolonchium.

PISTIA STRATIOTESContinued from page 13


a good field person and a competentplant presser.

On the journey to El Chical fromQuito, most of the route consists ofarid hills with the highway adheringto long winding slopes with lots ofbus and truck traffic. We had lunchin Ibarra, the capitol of ImbaburaProvince, then continued on toTulcán, the capitol of Carchi Provincenear the Colombian border. The areabetween Tulcán and Tufino is totallydevoted to pasture due to itselevation of 3500 m. After beingstopped by a military checkpoint justbefore reaching Tufiño, we entered anarea of shrubby vegetation and begancollecting; most of the species werenew to me, but the area was too highfor Araceae. Just to the east the roadclimbed higher and we entered theParamo del Angel (Fig. 1), with vaststands of Espeletia and Puyaextending for more than 20 km. Theparamo was inhabited by llamasoccasionally right along the road, andwhen the paramo was enveloped infog, the llamas, barely visible amongthe Espeletia, gave an eerie sensation.

Beyond the paramo the road droppedsteeply to the west and wanderedthrough forested hills where Araceaesoon became conspicuous andabundant with most of the speciesbeing new. Finally we reachedMaldonado. We drove through townlooking for a hotel, but saw nothingobvious and decided to continue onto El Chical, hoping there would be ahotel there. We arrived in El Chicaljust before dark and checked into thetown’s only hotel. It was the onlyconcrete building in town andthough somewhat primitive, it wasvery comfortable after I wired acouple of receptacles so we could runthe computer and other essentialappliances. We were the only guests

in the hotel and even the owner leftthe following day on a trip to Quito,leaving us in the care of a friend wholived nearby.

El Chical lies on the Colombianborder across from a smallColombian village on the other sideof the Río San Juan with a footbridge providing its electric currentfrom the Ecuadorians. The village isnot connected by road to any otherpart of Colombia, and El Chical has agarrison of Ecuadorian army on thehill above the town so the soldiersrun through the streets in the earlymorning doing military exercises.The single road continuing from thetown across the Río Chical goes on toPeñas Blancas, which is really nothingmore than a house at the end of theroad. The house is owned by Elisa’sgrandparents and it was here thatElisa’s father Jim Levy, then a PeaceCorp worker trekking into thelowlands to work with Awa Indiancommunities, met and fell in lovewith Elisa’s mother. The Levys live inQuito part time and also in a housethey built just east of El Chical.

We spent our first day collecting inthe area of Peñas Blancas, whichcontains many Araceae. On the way,at the river bridge near El Chical, wefound a large new species ofAnthurium growing in a tree, amember of section Cardiolonchium,and one which was being pollinatedby small wasp-like bees. I managed toget good photos and collected somebees in a plastic bag for laterdetermination, but they all escapedby chewing their way out of the bag.In another area we found twoadditional new species, both alsomembers of section Cardiolonchium.One had a dark purple spathe and areddish spadix (Fig. 2) while theother had longer internodes and aflesh-colored spadix. Among thespecies previously seen in the Lita-San Lorenzo region were

Anthurium anchicayense Croat,A. andreanum Linden,A. angamarcanum Sodiro,A. dolichostachyum Sodiro, A. micheliiGuill., A. peltigerum Sodiro,Dieffenbachia tonduzii Croat &Grayum, Philodendroncurvipetiolatum Croat, P. esmeraldenseCroat (Fig. 3), P. roseocataphyllum

2


15

Continued on page 3

Figure 2. New species of Anthuriumsection Cardiolonchium with a dark

purple spathe & reddish spadix.

Figure 3. Philodendronesmeraldense Croat.

Review ofIttenbach, Stephan. 2003. Revision der afrikanischenArten der Gattung Amorphophallus (Araceae)Englera 25: 1-263. [Available from Botanischer Garten und Botanisches Museum,Königin-Luise-Str. 6-8, 14191 Berlin, Germany ($40)]

This is a classic revision in the tradition of the German-speaking aroidologistsSchott (1794-1865) and Engler (1844-1930), although dealing solely with theAfrican (including Madagascar) taxa (35 species and 7 subspecies). However, theauthor has worked closely with Wilbert Hetterscheid and, between them, theyare masters of this Old World splendid tropical genus. Members of the AroidSociety may remember Hetterscheid & Ittenbach’s extremely useful (standard)work “Everything you always wanted to know about Amorphophallus, but wereafraid to stick your nose into” (Aroideana 19: 7-131. 1997) with its key to all thenknown taxa of the genus, descriptions, photographs, etc.

The treatment’s frontispiece is a classic photo (taken at the same time as the onein Aroideana 1: 68. 1978) of young Josef Bogner (now an Ehrendoktor!) standingamong flowering Amorphophallus angloensis subsp. maculatus. The treatment isbased on study of about 1000 specimens and living material of most taxa. Thechapter on morphology (pp. 11-24) is well illustrated with drawings andphotographs and the chapter on distribution and habitat (pp. 26-34) gives one ageneral overview. There is an important discussion of the genus, how its partsmay have evolved leading to a classification delivered in the forms of synopses,keys, characters, and descriptions and followed by an evaluation of thesecharacters. All this careful reflection (pp. 35-65) leads to a key to African species(pp. 66-68), followed by a discussion of the taxa. The taxa are arrangedalphabetically, which means that you can move quickly from the key to the taxon.This has the disadvantage that the taxa closest to each other are not adjacentmaking it a bit harder to compare the treatments of similar taxa.

The best way to communicate the quality of this work is to report an example ofa treatment of one species and assure you that each species has a comparabletreatment, perhaps missing one or two items or adding something else, but toshow that this is meaty stuff!

Example: Amorphophallus gracilior Hutchinson 1939, which begins on p. 160and ends on p. 165, is a renaming of A. gracilis Chevalier (1920), a name whichwas an illegitimate later homonym of the same binomial published by Engler in1881. The holotype, from Benin (in Paris), was tracked down and found to beuseless because of having been consumed by herbarium beetles and thereforehe here designates a neotype collected in Nigeria in 1946. A full page photo ofthe neotype appears on p. 162, followed by a full page distribution map onp. 163 (with two adjacent dots), followed by the full page drawing on p. 164 withmany details. The text begins with the citation of the correct name, its place ofpublication, its synonyms, its typification(s), a long description, a long paragraphon observations, distribution, habitat, and ecology, and finally, citation of other (asopposed to type) material, including the herbarium where seen, with anexclamation mark if the author saw the specimen.

The work closes (after thanking various people and institutions) with (1) a classicbibliography and, for anyone who might have unannotated duplicates, (2) analphabetic list of collectors and their specimen numbers identified by a taxonnumber, and (3) an index of scientific names.

Would that all aroids were as well studied as these African Amorphophalli!

Dan H. Nicolson, Department of BotanyNational Museum of Natural HistorySmithsonian Institution, Washington, DC. 20560-0166. ❦

Kirtikar, K.R. and B. D. Basu. 2002. IndianMedicinal Plants. Delhi, Sri SatguruPublications.

Livingstone, D. Missionary travels andresearches in south Africahttp://etext.library.adelaide.edu.au/l/livingstone/david/mission/chapter32.html.

Mayo, S. J., J. Bogner, and P.C. Boyce. 1997.The Genera of Araceae. Royal BotanicGardens, Kew.

Pimentel, D., L. Lach, R. Zuniga & D.Morrison. 2000. Environmental andEconomic Costs of NonindigenousSpecies in the United States. BioScience.50(1):53-64.

Pimentel, D., R. Zuniga & D. Morrison.2005. Update on the Environmental andEconomic Costs Associated with Alien-Invasive Species in the United States.Ecological Economics. 52: 273-288.

Premkumar, V.G. & D. Shyamsundar.2005. Antidermatophytic activity ofPistia stratiotes. Indian J. Pharmacol.37:127-128.

Ramey, V. 2001. Pistia stratiotes. Universityof Florida. www.aquat1.ifas.ufl.edu/seagrant/pisstr2.html.

Rivers, L. 2002. Water Lettuce (Pistiastratiotes). University of Florida and SeaGrant. www.iisgcp.org/EXOTICSP/waterlettuce.htm

Smirnova, L. L. & N. V. Mironova. 2004.Intensification of BiologicalAfterpurification of MunicipalWastewater by Some PleistophyteSpecies. Water Resources 31 (4):470-473. ❦

In August 2004, I continued myEcuadorian adventure, the firstpart reported in the last IAS

Newsletter. During the second half Ivisited areas on the Pacific andAtlantic slope. I was accompanied byGenevieve Ferry and Elisa Levy.Genevieve arrived from NancyBotanical Garden in France. I metGenevieve during a trip to Europefor Josef Bogner’s honorary Ph.D.ceremony. Genevieve has greatinterest in Araceae, but had neverbeen to a tropical rainforest. HerDirector, Romeric Pereil, wasconvinced that Genevieve and the

botanical garden would profit if shewere to join an expedition toEcuador. Elisa Levy, a botany studentat the Universidad Catolica, is anative of El Chical in Carchi Provinceand proved very useful because shewas very familiar with the area. I hadbeen warned that it might not be safebecause of Colombian guerilleraswho frequented the area, andthought having a local resident alongwho the community was familiarwith would provide us with a certaindegree of cover. She also proved to be

Continued on page 2

FIRST CLASSU.S. POSTAGE

PAIDPERMIT #1040LEESBURG, FL

34748

P.O. Box 43-1853South Miami, FL 33143 USA

www.aroid.org

1

ContentsTable of Contents

Explorations for Araceae inEcuador, 2004: Part IIBy Tom Croat . . . . . . . . . . . . . . . . 1

Black Magic and Purple Passion:Black AroidsBy Karen Platt . . . . . . . . . . . . . . . 4

Recent Aroid Activities at theMissouri Botanical GardenBy Tom Croat . . . . . . . . . . . . . . . . 8

Pistia Stratiotes: An UbiquitousInvader By Emily Yates & KristenKordecki . . . . . . . . . . . . . . . . . . . 12

Review of Ittenbach, Stephan,Revision der afrikanischen Artender Gattung Amorphophallus(Araceae)By Dan Nicolson . . . . . . . . . . . . 15

Advertisements . . . . . . . . . . . . . 16

IAS OfficersPresident: Albert HuntingtonVice President: Dan LevinTreasurer: Patricia FrankRecording Sec.: Lester KallusCorresponding Sec: Julius BoosMembership: Paul Resslar

IAS PublicationsAROIDEANAEditor: Dr. Derek BurchScientific Editor: Dr. Tom Croat,Missouri Botanical Garden,St. Louis, Missouri

IAS NewsletterEditor: Emily Yates

IAS Website: www.aroid.orgAdmin: Scott Hyndman

A QUARTERLY PUBLICATION FOR MEMBERS OF THE INTERNATIONAL AROID SOCIETY

Explorations for Araceaein Ecuador 2004: Part IIThomas B. Croat, Missouri Botanical Garden

the IAS NewsletterVol. 27 – 3 • July 2005

Figure 1. Paramo del Angel.

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