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Torrey Botanical Society

The South American Species of Arracacia (Umbelliferae) and some Related Genera Author(s): Lincoln Constance Source: Bulletin of the Torrey Botanical Club, Vol. 76, No. 1 (Jan. - Feb., 1949), pp. 39-52 Published by: Torrey Botanical Society Stable URL: http://www.jstor.org/stable/2481888 . Accessed: 18/07/2013 09:21
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BULLETIN

OF

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TOIRIREY

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JANUARY, 1949

VOL. 76, No. 1, pp. 39-52

THE SOUTH AMERICAN SPECIES OF ARRACACIA AND SOME (UMBELLIFERAE) RELATED GENERA
LINCOLN CONSTANCE

In our revisionarystudy of North American Umbelliferae (1944-45), Dr. Mildred E. Mathias and I regarded Arracacia Bancroft,with twentyexfour species,as the fourthlargest genus of the familyin the continent, species, Eryngium L., ceeded only by Lomatium Raf., with seventy-eight The combined Raf., with thirty-two. and Cymoptermts with sixty-four, ranges of the species of Arracacia extendprettywell over the mountainous regionsof Mexico and Guatemala,and one reaches Costa Rica and Panama. For several years,Mr. EllsworthP. Killip, Curator of the Department of Botany of the United States National Museum,has been urging a study of this group. In addition to the two of the South Americanrepresentatives Bancroft (A. properly referredclassical species, Arracacia xanthorrhiza. DC.) and A. moschata (H.B.K.) DC., in the year 1908 Britton esculenta, describedA. andina fromBoliva and WolffdescribedA. elata, A. incisa, and Velaea peruviana,from Weberbauer's Peruvian collections.Finally, unpublishednamesto various herRose attachedno less than eightdifferent Two of Wolff'sspecies were delineatedin the absence of barium collections. maturefruit,and therehas been a good deal of confusionin applying his binomials. years since Wolff'sand Britton's concernwith Despite the lapse of forty the genus,the materialat hand is still muchtoo meagreto permitanything Many of the collectionsare betterthan a strictly"pioneer" classification. structures;othersare not readily referableto immatureor lack significant of a new any describedentitybut are scarcelyadequate for the typification with me in this one, althoughWolffwould almost certainlyhave differed view. I should be surprisedif the acquisitionof additional materialdid not necessitate a thorough reappraisal of the admittedly tentative scheme the necessityof different however, offered below.I thinkwe mustrecognize, "levels" of taxonomicwork,and be prepared to constructthe best treatin florisat hand. The personinterested mentpossiblewiththe information tic considerations will not be helped if the "specialist" refusesto prepare data at his disposal to a classification of any group until he has sufficient much investigations, make,let us say, statisticalanalyses and cytogenetical provisional of a appearance The as he mightwish to employ these tools. in need of usable basis forthe taxonomist a working treatment shouldafford
:39)

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keys and suitable names for his plants, and at the same time call attention to the utilityof furthercollectingby those in a positionto do it. With considerablemisgivings, then,but no apology, the followingprovisional key is offered to the species of Arracacia in South America, with supplementary notes on certain genera whichhave been in one way or anotherconfusedwiththem. I am greatlyindebted to those in charge of the followingherbaria for the opportunityof seeing pertinentspecimens: Chicago Natural History Museum (F) ; Gray Herbarium,Harvard University(GH) ; MissouriBotanical Garden (MO) ; New York Botanical Garden (NY) ; Uniiversity of California (UC); United States National Museum (US).
ARRACACIA Bancroft,Trans. Agr. Hort. Soc. Jamaica 1825: 3. 1825. Arracacha DC. Bibl. Univ. Sci. & Arts 40: 78. 1829. Velaea DC. Coll. Mem. 5: 61. 1829. Vellea D. Dietr.; Steud. Nom. Bot. ed. 2. 2: 746. 1841. Stout or slender, erect, herbaceous or somewhat woody, cauleseent, branching or simple, glabrous to pubescent perennials,from taproots or tubers.Leaves petiolate,once to several times ternate,pinnate, or ternatepinnate, the leafletsor ultimate divisions various. Petioles sheathing.Inof loose to somewhatcompactcompoundumbels; peduncles terfloreseenee minal and lateral, or rarelyonly terminal, occasionallysome umbels sessile. Involuerewantingor vestigial.Involucel of few shortto long,narrowbractto longerthan the fruitor wanting.Fertile rays few to nuLmerlets,shorter ous, spreading-ascending to divaricateand reflexed. Flowers white,greenishyellow,greenish,reddish-brown or maroon,or purple; petals oblanceolate to obovatewith a narrowerinflexedapex; calyx teethobsolete; stylesshort to long, erectto spreadingor reflexed, conic and conspicuthe stylopodium ous to depressedand indistinct.Carpophore2-cleft to the base or only bifid at the apex, flator terete.Fruit lanceolate or oblong to ovoid,usually narrowedat the apex, flattened laterally,glabrousor pubeseent;ribs prominent, acute to obtuse,or filiform and indistinct;oil tubessolitaryto several in the intervals, 2-several on the commissure; seed subterete in crosssection,often channeledunder the tubes,the face suleate or concave. Type species: Arracacia.xanthorrhiza. Bancroft.

Artificial Key to the Species


Leaves 1-2-terliate or -pilnliateor -terliate-piniiate; leaflets laliceolate to ovate with serrate or delntateand usually incised or lobed margins, and without a prominentcallus point; ilnvolucelevident; rays and pedicels Leaflets spinul:ose-serrate; stylopodiumn conlic; carpophore bifid about I its lelngth;fruit tapering at apex. to white; pedicels Foliage essentially glabrous; flowersgreenislh-yellow 4-6 mm. long. Mericarp ribs obtuse; oil tubes all about the same size. Feltile rays 4-8; pedicels 4-6 mm. long; mericarp ribs very thick and corky, the intervals broad and shallow; oil tubes large, 1. A. Pennellii. ?olitnryin the intervals.
unwebbed.

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Fertile rays 6-15; pedicels 5-8 mm. lonlg; mericarp ribs thin, the sharplyV-shaped; oil tubes very small and ilndistinct, ilntervals 2. A. elata. forming a cointinouslayer between seed alnd pericarp. 3. A. Wigginsii. Mericarp ribs acute; oil tubes of two sizes. Foliage squamulose; flowersusually marooll; pedicels 10-30 mm. long. 4. A. moschata. Leaflets variously serrate to incised or lobed, but lnot spillulose-serrate; stylopodiumdepressed; carpophore 2-cleft to the base (unklnownin A. peruviana); fruit blunt at apex. Rays 4-8 cm. loiig; bractlets linear, herbaceous, exceeding the reddish5. A. peruviana. brown flowers. Rays 1-4 cm. long; bractlets broad anid searious, or linear and shorter thanithe purple or greenish flowers. lanceolate to obovate. Bractlets searious or searious-marginied, Bractlets 4-8, obovate to lanceolate, searious, 5-10 mm. long, exceeding the flowers; mericarp ribs very prominentand corky; 6. A. incisa. oil tubes small, 2-3 in the intervals. narrowly searious-margined, 2-6 Bractlets 3-6, ovate-acumilnate, mm. long, shorterthan the flowers; mericarp ribs filiform;oil 7. A. equatorialis. tubes large, solitary in the intervals. Bractlets herbaceous, linear. Plants 5-12 dm. high; fruit (immature) oblong, 10 mm. long, 2-3 mm. broad, constricted below apex; oil tubes solitary in the 8. A. xanthorrhiza. intervals. Plants 3-4 dm. high; fruit ovoid, 6-7 mm. long, 4-5 mm. broad, 9. A. andina. not constricted; oil tubes 2-3 in the intervals. Leaves ternately decompound, the divisions oblong-linear to filiformwith entiremargins (in our material) and a prominentcallus point; involucel var. mdltifida. 10. A. toltucensis wanting (or vestigial); rays and pedicels webbed.

1. Arracacia Pennellii Constance,sp. nov. cauleseensglaucescensglabra, 6-36 dm. alta; Herba crassa suffruteseens 0.8-2 dm. longa, ternata deinde 1-2-pinfolia in ambituovato-triangularia, nata, foliolis ovato-lanceolatisovatisve, ad apicem acutis vel acuminatis, ad basin culneatisrotundatisve,2-5 cm. longis, 1-2 cm. latis, spinulososerratisad basin lobatis; petioli 1.5-2 dm. longi, ad basin vaginates; folia ramosa, caulina foliis basilaribus similia, vaginis inflatis; infloreseentia 6-25 cm. longis; involueellibracteolae pedunculisalternisvel terminalibus, 5-9, lineari-lanceolatae,inaequales, 2-10 mm. longae; radii fertiles 4-8 3-7 cm. longi; pedicelli fertiles2-6, 4-5 mm. graciles patenti-adscendentes albive, petalis obovatis; stylopodiumconicum, longi; fioresflavido-virides rigidum; fructusovoideus, 5 mm. stylisgracilibus; carpophorumbifidum, obtusis suberosis, pericarpo longus, 3-4 mm. latus, costis prominentibus suberoso,valleculis latibus brevibusque; vittae magnae solitariae in valleculis,4 in commissuris; semina sub valleculis canaliculata facie suleata. branching,cauleseent, glaucous herb, 6-36 dm. Coarse, suffruteseent, 0.8-2 dm. high, the foliage essentiallyglabrous; leaves triangular-ovate, the leaflets ovate-lanceolateto ovate, acute or long, ternate-1-2-pinnate, at base, the lower distinctand short-petioluacuminate,cuneateto roulnded 2-5 cm. long, 1-2 cm. broad, spinuloselate,theupper sessile and confluent, serrateand usually lobed towardthe base, the lowersurfacepaler, glabrous, and reticulate, a squamulosetuft on the upper side of the suleate rachis at the base of the larger leaflets;petioles 1.5-2 dm. long, narrowlysheathing

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at base; cauline leaves mostlywith whollysheathingand inflatedpetioles; infloreseencebranching, the peduncles arising terminally and axially membrana6-25 cm. long, squamulose at apex; involuereof 1-4 sheath-like entire ceous bracts,or wanting; involucel of 5-9 unequal, linear-lanceolate, but or trifid bractlets2-10 mm.long, the longer about equaling the flowers 3-7 shorterthan the fruit; fertilerays 4-8, slender,spreading-ascending, 4-5 mm.long, cm. long,scaberulousat apex; fertilepedicels 2-6, spreadinlg, to white,the petals obovate; styloposcaberulous; flowersgreenish-yellow dium conic, the styles slender,erect to reflexed;carpophorebifidabout 4, rigid; fruit ovoid, 5 mm.long, 3-4 mm. broad, tapering at apex, glabrous, the ribs very prominentand corky,obtuse,forminga corkycoveringover the entirepericarp; oil tubes large, solitaryin the very shallow intervals, 4 on the commissure;seed deeply channeledunder the intervals,the face suleate. Moist rockycafion,alt. 2700-2800 m., Rio San Francisco, above TYPE: 13 September1917, F. W. Pennell Bogot'a,Dept. Cundinamarea,Colomnbia, 1932 (NY: type).
Specimens examined: COLOMBIA. CUNDINAMARCA: Rio San Franicisco,above Bogota, 13-IX-17, Pennell 1932 (NY-type, GH, US); Macizo de Bogot'a, Quebrada del Rosal, Between Mutiscus NORTE DE SANTANDER: 3000 m., 29-VI-39, Cuatrecasas 5700 (US). SANTANDER: and Pamplona, 3400 m., 23-II-27, Killip & Smith 19,728 (GH, NY, US). Cordillera Oriental, paramo de Santurban, entre Cuesta Boba y el extremooeste, 3400 m., 27-VII-40, Cuatrecasas & Barriga 10,314 (US); Paramo de Romeral, 3800-4100 m., 29,30-I-27, Killip & Smith 18,541 (GH, NY, US); vicinity of Vetas, thickets along streams,3100-3200 m., 16-VI-27, Killip & Smith 17,347 (GH, MO, NY, US) ; Quebrada de Pais, northof La Baja, dense forest,ca. 3200 m., 31-1-27, Killip & Smith 18,781 (GH, MO, NY, US).

species.

This species is very similar in general aspect to the next,but its fewer dissimilarfruitsection, pedicels,and strikingly shorter rays,predominantly set it well apart. With no collectionsavailable to connectthe widely sepait seemsadvisable to regard themas distinct rate ranges of the two entities,

2. ARRACACIA ELATA Wolff,Bot. Jahrb.40: 304. 1908. cauleseent,branching,up to 4.5 m. high, the foliage Stout, clambering, the essentiallyglabrous; leaves ovate, 1-3 dm. long, ternate-1-2-pinnate, leafletslanceolate to ovate, acute or acuminate,cuneate to rounded at the theupper sessile and confluent, and short-petiolulate, base, thelowerdistinct and often incised 3-6 cm. long, 1-4 cm. broad, sharply spinulose-serrate toward the base, the lower surface paler and glabrous,stronglyreticulate, a squamulosetufton the upper side of the suleate rachis at the base of the larger leaflets; petioles 15-45 cm. long, their lower 3 broadly sheathing; cauline leaves similar,the uppermostwithpetioleswhollysheathingand inbranching,the peduncles arising axially, 10-25 cm. flated; infloreseence long, squamulose at apex; involuere wanting,or occasionally of a single unleaf sheath; involueelof 8-10 linear to lanceolate,entireor few-toothed but shorter equal bractlets3-15 mm.long, the longerexceedingthe flowers 3-6 cm. long; than the fruit;fertile rays 6-15, slender,spreading-aseending, scaberulous at apex; fertilepedicels 3-8, spreading,5-8 mm. long; flowers greenish,the petals oval; stylopodiumconic, the styles slender,recurved; carpophorebifidabout 1 its length,rigid; fruit ovoid, 5 mm. long, 3 mm.

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obtuse,with narrowsharplyV-shaped broad, glabrous,the ribs prominent, forminga continuouslayer intervals; oil tubes very small and indistinct, betweenseed and pericarp; seed deeply channeledunder the intervals,the face suleate. TYPE: "Peru: Dep. Amazonas, a Chachapoyos orientemversus; inter in 3300 m. altitudinis," in graminosis, Tambo Ventillas et Piscohuafiuma, 4423. VWeberbauer
Specimens examined: PERU. AMAZONAS: "a Chachapoyos orientem versus; inter Tambo Ventillas et Piscohuafiuma,'' 3000 m., VII-04, Weberbawer 4423 (type photos: F, GH, UC, US). HUANUCO: Tambo de Vaca, ca. 12,000 feet, 10,24-VI-23, Macbride 4456 (F, US). AYACUCHO: Choimacota Valley, 3000 m., 28-1I, 10-111-26, Weberbauer 7584 (F, GH, US).

the emendaThe last collectioncited has good, maturefruit,permitting whichwas based upon immatureones. The close tion of Wolff'sdescription, above. has been mentioned of this species to the foregoing resemblance 3. Arracacia Wigginsii Constance,sp. nov. cauleseens glabra, 2-3 m. alta; folia basalia Herba crassa suffruteseens 1-1.5 dm.longa, in ambituovato-triangularia, incognita;folia caulina mnaiora ternata deinde 1-2-pinnata, foliolis lanceolatis ovato-lanceolatisve,ad 3-5 cm. longis,0.8-2 em. apicem acuminatis,ad basin cuneatisrotundatisve, ad basin lobatis; petioli 1-2 dm. longi, ad basin latis, spinuloso-serratis anguste vaginantes; folia caulina superiora similia, petiolo ad laminamn ramosa, pedunculis terminalibus axillaribusque, vaginato; infloreseentia 10-16 cm. longis,apice squamuloso; involueellibracteolae 3-6, lineares oblanceolataeve inaequales, 2-10 cm. longae; radii fertiles 10-15 graciles 3.5-7 cm. longi; pedicelli fertiles3-8, 4-6 mm. longi; patenti-adscendentes albive, petalis obovatis; stylopodiumconicum,stylis floresflavido-virides gracilibus; carpophorum bifidum, rigidum; fructus ovoideus oblong(oovoideusve,4-8 mm. longus, 3-3.5 mm. latus, costis acutis prominentibus; inter pericarpium et vittae vel minimae obscuraeque stratum continuumi vel magnitudineintermediaeplerumque solitariae in valsemenformantes semina sub valleculis canaleculis omnibusvel aliquis, 2-4 in commissuris; liculata facie suleata. branching,cauleseent, 2-3 m. high, the foliage Coarse, suffruteseent, essentiallyglabrous; basal leaves unknown,the larger cauline leaves trithe leafletslanceolate to 1-1.5 dm. long, ternate-1-2-pinnate, angular-ovate, ovate-lanceolate, acute or acuminate,cuneate to rounded at base, the lower 3-5 em. long, the upper sessile and confluent, distinctand short-petiolulate, and oftenlobed towardthe base, the lower 0.8.2 cm.broad,spinulose-serrate surfacepaler, glabrous,and reticulate,a squamulose tufton the upper side of the suleate rachis at the base of the larger leaflets;petioles 1-2 dm. long, narrowlysheathing below; petioles of the upper cauline leaves wholly branching,the peduncles arising tersheathingand inflated;infloreseence minallyand axially, 10-16 cm.long,squamuloseat apex; involuerewanting, or of reduced leaf sheaths; involueelof 3-6 unequal, linear to oblanceolate, entire or trifidbractlets 2-10 mm. long, the longer about equaling the flowers but shorterthan the fruit; fertilerays 10-15, slender, spreadingascending,3.5-7 cm. long, scaberulous; fertilepedicels 3-8, spreading,4-6 "greenish yellowto nearlywhite," the petals mm.long,scaberulous;flowers erectto reflexed;carpophore conlie,the stylesslenider, obovate; stylopodium

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4-8 i1in. long, bifidabout 1 its length,rigid; fruit ovoid to oblong-ovoid, acute; oil tubes of two sizes: (1) very 3-3.5 mm.broad,the ribs prominent, a continouslayer betweenseed and pericarp, forming small and indistinct, size, usually solitaryin some or each of the and (2) vittae of intermediate intervals,2-4 on the commissure;seed channeled under the intervals,the face suleate. TYPE: Along Pan-American Highway 40 km. south of Cueniea,10,900 feet, Azuay Prov., Ecuador, 20 September 1944, Ira L. Wiggilns10,769 (UC 708,757: type).
Rose Specimens examined: ECUADOR.CAfAR: Vicinity of Caniar, 15-IX-18, Rose R AZUAY: 40 km. south of Cuenca, Wiggins 10,769 (UC type); "Mt. 22, 715 (NY, US). Pillshum,' 12,000 feet, Jameson 24 (GHi).

althoughless maturethan those The fruitsof the Rose & Rose collection, but no other to theirw-idth, of the type,are markedlylongerin proportion in structurehas been detected. This species is very difference significant similar to the two foregoingin general aspect and it is, again, the fruit which affordsthe principal basis for its separation. The peculiar dimorphism of the oil tubes suggestsa blend of the kinds of vittae found in A. to Pennellii and A. elata, but scarcelypermitsthese Ecuadorean specimnens be mergedwith eitherthe Colombianor the Peruvian species. 4. ARRACACIA MOSCHATA (H.B.K.) DC. Prodr. 4: 244. 1830. H.B.K. Nov. Gen. & Sp. 5: 12, pl. 430. 1821. Conitm moschatum clambering, cauleseent,branching,up to 1.5 m. high, the foliage Stout,, squamulose; leaves ovate, 1.5-3 dm. long, bipinnate or ternate-pinnate, to ovate, acute, cuneate or roulided at base, the the leafletsovate-oblong 2-5 cm. the upper sessile and confluenit, lowerdistinctand short-petiolulate, and ineised or pinnlatifid, long, 1-3 cm. broad, sharply spinulose-serrate squamulose on the larger veins on the upper surface, the lower surface paler, glabrous and reticulate,a squamulose tuft on the upper side of tne suleate rachis at the base of the larger leaflets; petioles 10-20 cm. long, usually broadly sheathing; cauline leaves with petioles wholly sheathing braliching,the pedulieles arising axially, 1.5-3 anid inflated;infloreseence dm. long,squamuloseat apex; involuereof reducedleaf sheathsor wanting; bractlets6-15 mm. involueelof 3-8 linear to lanceolate,entireor few-toothed than the fruit; rays 8-12, but shorter long,the longerexceedingthe flowers denselyscaberulous; or spreading,7-13 cm. lonig, stout,spreading-ascending fertile pedicels 1-12, spreading, 10-30 mm. long, densely scaberulous; conic, the flowersmaroon (rarely yellow), the petals oval; stylopodiuLim stylesslender,recurved; carpophorebifid about 4 its length,rigid; fruit acute; lance-ovoid, 6-8 mm.long,3-3.5 mm.broad,the ribs veryprominent, oil tubes large,usually solitaryin the intervals, about 4 on the cornrnissure, sometimessmall supplementaryones under the ribs or in the intervals; seed channeledunder the intervals,the face suleate. TYPE: "crescit in frigidis Provinciae de los Patos, prope Teindala, [Prov. Azuay, Ecuador], alt. 1400 hex.," Humboldt & Bonpland.
Specimens examined: ECUADOR. CARCHI: Wooded hills about 5 miles south of Tulen, 21,005 (GH, NY, US); between Tulca,n and Pun, 3500 2500 m., 10-VIII-23, Hitchcock? PICHINCHA: "cult. ad radicem Montes Pichincha, in., 11-VIII-35, Mexia 7580 (US). sive Andiuin Quintensium, " 111-64, Jameson (US-petals yellow); " in Andibus Ecuador-

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''Weg von1 Paso nach Ambato TUNGURAHUA: 1857-59, Sprutce5794 (NY). " 6-XI-32, am Rio Ambato, stelle steinige Uberberge,Andem um Ambato, 2850 m., haiifig, E. H[enrichs71 (NY).

and the usually maroon The long rays and pedicels, the leaf-cutting, flowersdistinguishthis species sharply from the preceding. The occurin the light of the differrence of small accessoryoil tubes is of initerest 's note that ences in vittae described in the firstthree species. Jameson this species is "cultivated" may indicate that other species than A. xarnthorrhizaeitherare or have been so employed,but I have no furthereviof A. moscha,ta. dence of the domestication 5. Arracacia peruviana (Wolff) Constance,comb.nov. Bot. Jahrb. 40: 303. 1908. Wolff, Vela,eaperuviarna branching,6-9 dm. high,squamulose to scaberulous Slender, cauleseent, the stem base clothedwith dry sheaths,froma branched tapthroughout, the leafletsovate to 2-3 dm. long, bipilnnate, root; leaves ovate-lanceolate, lanceolate,acute, cuneate at base, the lower distinctand short-petiolulate, 2-5 cm. long, 1-4 cm. broad, coarsely sessile and confluent, the terminial squamulose on the veins and marsinuatelylobed and mucronulate-serrate, gins, the lower surface paler and reticulate; petioles 1-3 dm. long, sheathing below; cauline leaves pinnate,the uppermostwithshort,whollysheathof alternate axillary peduncles, 7-15 cm. long, ing petioles; infloreseence squamulose at apex; involuere wanting, or of a single leaf sheath; involucel of 6-10 entirelinear bractlets5-9 mm. long, exceedingthe flowers but shorterthan the fruit; fertilerays 5-10, slender,spreading-ascending, 4-8 cm. long,squamuloseespeciallyat apex; fertilepedicels 2-6, spreadinlg, the reddish-brown, 5-6 mm.long,squamulose or scaberulousabove; flowers depressed,the styles slender,spreading-erect; petals obovate; stylopodium carpophoreunknown;fruitovoid, 4-6 mm. long, 3-4 mm. broad, glabrous, oil tubes large, solitary in the intervals,2 on the comthe ribs filiform; missure; seed face deeply and narrowlysuleate. TYPE: "Peru: Dep. Aneachs,prov. Cajatambo infra Ocros,in 3000-3200 2748. m. alt.," Weberbamuer
Specimens examined: PERU. ANCACHS: "'prov. Cajatambo infra Ocros," Weberbauer Carumas, 2900 m., 21-II, 6-III-25, Weberbauer 2748 (type photo: F). MOQUEGUA: Mountains northeastof Huanta, 3200 m., 1,10-11-26, WeberAYACUCHO: 7269 (F, US). baiter7513 (F).

As shownby Coulter and Rose (1900), the type species of Velaea DC. and hence Velaea must be tolucensisH.B.K.) is an Arracacia,, (Ligusti,cmm mergedunder Arracacia,.None of the specimensavailable for study bear mature fruit,so the descriptionof that organ has beelnabstracted from Wolff'soriginal description.This species is not likely to be confusedwith thosethat precede it, and it is separable fromthose that followby its conflowers.The Weberbauer colspicuous linear bractletsand reddish-brown fromHuanta has atypically short involueels,but it may be semilection, sterile. 6. ARRACACIA INCISA Wolff, Bot. Jahrb. 40: 305. 1908. Stout, cauleseent,branching,3-12 dm. high, the foliage squamulose; 1-2.5 dm. long, ternate-pilinate to ovate-lalnceolate, leaves triangular-ovate to ovate-oblong, acute, cuneate or or bipinnate,the leafletstriangular-ovate the upper sessile truneateat base, the lower distinctand short-petiolulate,

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and the larger pinnatelyincised,squamulose on the niarginsandlalonigthe veins on both surfaces,the lowersurfacepaler and reticulate,a squiamiiulose tufton the upper side of the suleate rachis at the base of the larger leaflets; petioles 8-16 cm. long, narrowlysheathingat the base, the sheathsscaberulous on the veins; cauline leaves with wholly sheathing,ineoiispicXuously the peduneles arisinig axially anid branching, inflatedpetioles; infloreseence 2-12 cm. long, squamulose at apex; involuerewantino,or of 1 terminally, or 2 sheathing bracts; involueel of 4-8, obovate to laneeolate, searious, cenltral unequal bractlets5-10 nmu.lolig, the gYreeni deenticulate-margined, but shorter portionprojectingas an acuminatepoint,exceedingthe flowers 1-4 em. long, than the fruit; fertilerays 4-8, stout, spreading-ascending, usually scaberulous at least at apex; fertilepedicels 2-6, stout,spreadiilng, the petals obo2-5 mm.long, scaberulous; flowers dark purple or greenish, vate; stylopodiumdepressed,the styles slender, divaricate; carpophore2cleftto the base, lax; fruitovoid, 5-8 mm. long, 3.5-6 mm.broad, the ribs very prominent and corky,acute; oil tubes small, 2-3 in the intervals,3-6 some accessoryones frequently under the ribs or in the on the commissure, intervals; seed scarcelychanneledunder the intervals,the face deeply sulcate. TYPE: "Peru: in deelivibusrupestribusprope Tambo, ad viami ferreanm 2650 m., Weberbauer165. interoppida Lima et Oroya," [Dept. Linma],
Specimens examined: PERU. HuANuco: Steep rocky open grassy slope, ca. 6500 feet, Huacachi, estacion near Munia,20-V,1-VI-23, Macbride 4163 (F, US); in shrub on southwesterncanyon slope, ca. 10,000 feet, Yanahuanca, 16,22-VI-22, Macbride J Featherstone LIMA: "in declivibus rupestribusprope Tambo, ad viam ferreamn 1244 (F). inter oppida Lima et Oroya,"I Weberbauer 165 (type photos: F, GH, UC); Huarochiri, Viso, sandy hillside, sun, 2800 m., 22-IV-39, Goodspeed, Storkf4H orton 11,540 (GH, UC); Prov. Huarochiri, valley of Rio Rimac near Lima-Oroya highway at km. 90 east of Lima, 4 Weberbauer 33,059 (GH, UC, US); in firmsoil of 3000 m., 15,22-III-42, Goodspeed steep southernslope, sparsely shrubby,ca. 8000 feet, Matucana, 12-IV,3-V-22, Macbride 4 Featherstone 326 (F, US); open slopes, ca. 8000 feet, Matucana, 14,18-III-23), Macbride 2949 (F, NY, US); rocky slopes, ca. 12,000 feet, Rio Blanco, 8,19-V-22, Macbride 4 Featherstone 730 (F, US). Cuzco: Cuzco, 1-IX-14, Rose 4 Rose 19,034 (US).

The plants representedby Macbride 4163 are much larger in stature and fruit than the other collectionscited, but these seem to be the only differences. Wolffapparentlywrote his original diagnosis fromhighlyunothersatisfactory, immature,and possibly sterile material; it is difficult wise to reconcilethe excellentmaterialnow available withhis very erroneous depictionof, for example,the ilnvolueels and stylopodium. As a consequence, this species and A. peruviana have been generally confused by, among others,the writer,although the involueels of the two species are The conspicuoussearious involueels settingoffthe usuentirelydifferent. ribbedfruitmake this and the blunt,prominently ally deep purple flowers, one of the most distinctive species of the group. 7. Arracacia equatorialis Constance,sp. i1yV. Herba gracilis ramosa cauleseens squamulosa, 4-8 dm. alta; folia in, 6-9 dm. longa, bipinnata, foliolis ovatis, ad ambitu ovato-triangularia, 1.5-3 cm. longis,1-2.5 cm. ad basin cuneatistruneatisve, apicem acuminatis, basi lobatis squamulosis; petioli 10-20 cm. longi, latis, mucronato-serratis ad basin vaginantes; folia caulina similia, foliolis linearibus lanceolatisve ramosa, pedunculis elongatis,petiolo ad laminamvaginante; inflorescentia

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vel solitariis,ad apicem squarnuloso;involueellibracteolae 3-6, verticillatis 2-6 mm. lolngae;radii fermembranaceae, ad margillem ovato-acuminatae, ad apicem scabriusculi, 2-4 cm. tiles 2-6, graciles, patenti-adscendentes, longi; florespurpurei,petalis obovatis; stylopodiumdepressum,stylis gra8-9 cilibus; carpophorumbipartitum,laxum; fructus ovoideo-oblongus, mm. longus, 3-4 mm. latus, costis filiformibus acutis; vittae magnae solisub vittis canaliculata facie tariae in valleculis, 2 in commissuris;semnina suleata. Slender, erect, cauleseent,branching,4-8 dm. high, the foliage squa6-9 cm. long, biternateor bipinnate, the mulose; leaves triangular-ovate, leafletsovate, acute or acuminate, cuneate to truncate at base, the lower 1.5-3 cm. the upper sessile alnd confluent, distinctand short-petiolulate, alnd the larger ilncisedor lobed, long, 1-2.5 cm. broad, mueronate-serrate with linear belneath squamuloseon the sheaths,rachises,veins, and marginis scales,squamuloseor merelyscaberulousabove, the lowersurfacepaler and a squamulose tuft on the upper side of the suleate rachis at the reticulate, only at the sheathinig base of the larger leaflets; petioles 10-20 cm. lolng, base; cauline leaves reduced upwards, with linear to lanceolate, elongate divisions,the obovate petioles wholly sheathing,little inflated; inflorescence braicehing, the peduncle arising in whorls or singly,2-12 cm. long, subsessile,squamulose at apex; involuerewanting; the terminalsometimes subequal, narrowlysearious-marenitire, involueel of 3-6 ovate-ac-uminate, gined bractlets2-6 mm. long, shorterthan the flowersand fruit; fertile 2-4 cm. long, scaberulous; fertile rays 2-6, slender,spreading-ascending, pedicels 1-3 (-5), stout, ascending, 3-5 mm. long; flowerspurple, the petals obovate; stylopodiumdepressed, the styles slender, spreadinlg-asfruit ovoid-oblong, cending; carpophore2-cleftto the base, lax, filiform; acute; oil tubes 8-9 mm. long, 3-4 mm. broad, glabrous,the ribs filifornm, seed channeledunder large, solitaryin the intervals,2 on the commissure; the tubes,the face deeply suleate. "vicinity of Las Juntas [Prov. Loja], Ecuador," 28 September TYPE: 1918, Rose, Pachano & Rose 23,215 (US 1,022,735: type).
47, B. Espinosa1305 (UC).
Specimens examined: ECUADORLOJA: Vicinity of Las Juntas, Rose et al. 23,215 (US-type, GH, NY); entre S. Pedro y Chinchas (unos 55 km. 0. Loja), 1600 m., 1-III-

Rose gave to the type collectionan herbariumname whichis preempted in the genus by A. humilis Rose, described from Guatemala. The Ecuafromthe and A. andina, differing dorean species is nearestA. xanthorrhiza formerin its fruit and from the latter in its foliage and oil tubes. The Espinosa collectionis immaturebut probably belongs here. 8. ARRACACIA XANTHORRHIZA Bancroft, Trans. Agr. Hort. Soc. Jamaica 1825: 5. 1825. ConiumArracacha Hook. Exot. Fl. pl. 152. 1825. Arracacha esculenta DC. Bibl. Univ. Sci. & Arts 40: 78. 1829. Billb. Linn. Samf. Handl. 1: 40. 1833. Bancroftiaxanthorrhiza Stout, cauleseent,branching,5-12 dm. high, glaucous, the foliage squamulose and scaberulous; leaves broadly ovate, 10-35 cm. long and broad, acuto triangular-ovate, biternateor bipinnate,the leafletsovate-lanceolate minate,cuneateto roundedat base, the lower distinctand oftenshort-petio-

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4-12 cm. long, 1.5-6.5 cm, broad, lulate, the upper sessile and confluent, and incised or lobed, squacoarsely simply or doubly mucronate-serrate mulose or scaberulous on the rachises, veins, and margins with flattened oblong or linear scales, the lower surface pale and reticulate,a squamulose tuft on the upper side of the suleate rachis at the base of the larger leaflets; petioles 0.8-4.5 dm. long, sheathingonly at base; cauline leaves reduced upwards, mostly ternate or 3-parted, with lanceolate, acuminate divisions,the lower alternate and petiolate,the upper often opposite and wholly sheathing with narrow, scarcely inflated sheaths; infloreseence the peduncles arising in whorlsor singly,3-10 cm. long, squabranching, mulose or scaberulousat apex; involuerewanting; involueel of 5-8 linear, than the flowers entire,unequal, herbaceousbractlets2-5 mm.long, shorter 1.5-4 cm. long, and fruit; fertilerays 5-12, slender,spreading-ascending, 2-4 mm. scaberulous; fertile pedicels 3-8, slender, spreading-ascenlding, long; flowerspurple or greenish,the petals oval; stylopodiumdepressed, the styles slender, ascending; carpophore 2-cleft to the base; immature fruit oblong, 10 mm. long, 2-3 mm. broad, constrictedbelow the apex, acute; oil tubes rather large, solitaryin the glabrous,the ribs prominent, intervals,2 on the commissure;seed channeled under the tubes, the face deeply suleate. TYPE: Jamaica, where cultivated from South America, probably Colombia.
Specimens examined: ECUADOR. LOJA: Viciiiity of Las Juntas, Rose et al. 23,215 COLOMBIA. ?Moritz 1803 (F. NORTE DE SANTANDER: Rose 4' Rose (US-foliage). Cordillera Oriental, region del Sarare: Hoya de rio Chitaga, a sobre La Cabuya, 16001800 m., 13-X-41, Cuatrecasas, Schultes 4 Smith 12,146 (F, GH-foliage). ANTIOQUIA: Cordillera Central, alrededores de Medellin, 1560 m., 1-V-46, W. H. Hodge 6861 (GH). Cuesta de TocotA, road from BuenaCAUCA: Popayan, Lehmann B.T.407 (GH-flowers); PUTUMAYO: velitura Cali, Western Cordillera, 1500-1900 m., XII-05, Pittier 714 (US). Valle de Sibundoy, los alrededores, ca, 2250 m., 18-II-42, R. E. Schu,ltes 3267 (GHfoliage). BOLIVIA. LA PAZ: Sorata, IX-1-58, 2697 m., Mandon 595 (GH), 22-IV-20, Sirupaya, 1800 m., 27-XII-06, Buchtien 59711 Holway 4 Holway 563 (US-foliage); ECUADOR. TUNGURAHUA: Ambato, 1918,G. Rose 38 (GH, US-photo of (US-foliage). foliage). PERU. CUZCO: Paruro, Hda. Araypallpa, 3100 m., 28-VII-37, Vargas 411 (GH); colinas del Laxaihuaman, 3600 m., XII-28, Herrera 858 (F); Ollantaytambo, Santa Ana, ea. 900 m., 29ea. 3000 m., 24-IV-15, Cook7& Gilbert 282 (US-foliage); San Miguel, Urubamba Valley, ca. 1800 m., VI-15, Cook 4' Gilbert 1583 (US-foliage); 935 (US-foliage). 26-V-15, Cook 4& Gilbert 934 (US-foliage),

South America,is cultivatedarracacha of northern This, the commonly in herbaria by only unsatisfactory material,and published derepresented are correspondingly inaccurate.I have not seen maturefruit,and scriptions it has never been described,to my knowledge; because the plants are ordifruit may not normallyripen. There apnarily propagated vegetatively, available on the cultivationof the species that pears to be no information was not incorporated by Hooker (1831) in his descriptionin the Botanical Magazine. None of the material seen is unquestionablyfromthe wild, and I findin it no clues as to the indigenous occurrenceand possible original home of the domesticated plant. Hooker's material came fromBogota, Colombia, by way of Jamaica. What appear to be the most closely related species, however,are P. equatorialis of Ecuador and P. andina. of Bolivia. It is hoped that the publication of this study may stimulate interestin

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more information discovering about this crop plant, its domestication, and its origins. 9. ARRACACIA ANDINA Britton,Bull. Torrey Club 18: 37. 1908. Stout, erect,cauleseent,branching,3-4 dm. high, the foliage squamulose; leaves ovate, 1-2.5 dm. long, 1-2-pinnate,the leafletslanceolate to ovate,usually acuminateand cuneate at base, the lower distinctand shortpetiolulate,the upper sessile and confluent, 2-8 cm. long, 1-3 cm. broad, mucronate-serrate and the larger incised or lobed, squamulose on the rachises, veins, and margins with flattenedlinear scales, the lower surface paler and reticulate,a squamulose tuft on the upper side of the suleate rachis at the base of the larger leaflets;petioles 15-35 cm. long, sheathing only at base; cauline leaves like the basal, the uppermostwith obovate, whollysheathing, inflatedpetioles; infloreseence moderately the branching, pecuncles arising in whorlsor singly,5-12 cm. long, squamulose at apex; involuerewanting; involueelof 6-8 linear entireunequal herbaceousbractlets 2-5 mm. long, shorterthan the flowersand fruit; fertile rays 5-8, slender, spreading-ascending, 2-4 cm. long, scaberulous; fertile pedicels 3-8, spreading, 2-4 mm. long; flowerspurple, the petals obovate; stylopodium depressed,the styles slender,spreading; carpophore2-cleftto the base, the divisionsfiliform; fruitovoid, 6-7 mm. long, 4-5 mm. broad, the ribs filiform, acute; oil tubes of intermediate size, 2-3 in the intervals,4-6 on the commissure;seed shallowly channeled under the larger tubes, the face deeply suleate. " Ingenio del Oro [Bolivia]," 10,000 feet, 111-86,Rutsby1776 TYPE: (NY: type).
(US);

Specimens examined: BOLIVIA. "Plantae Bolivianae," Bang 2839 (F, GH, MO, NY, Ingenio del Ovo, Rusby 776 (F, NY-TYPE, US).

The broad leaflets, different fruit,and clusteredoil tubes separate this species fromA. xanthorrhiza, and the different ilivolueel and oil tubes distinguishit fromA. equatorialis,but these appear to be its closestrelatives. To eitherthis species or to A. equatorialis may belong the three following Peruvian collectionswhich, because of their immaturity or fragmentary nature,I am unable to assign definitely to any species: Huasahuasi, Rutiz & Pavon (F); Hacienda Churiu, Paucartambo Valley, 1-27,Herrera 1391 (US); San Sebastian, Cuzco, 25-IV-25, Pennell 13,628 (F). It is possible that thereis a Peruvian entitywhich is distinctfrom eitherof these two species,but that decisionmust await bettermaterial. 10. ARRACACIA TOLUCENSIS (H.B.K.) Hemsl. var. MULTIFLORA (S. Wats.) Math. & Const. Bull. Torrey Club 68: 121. 1941. Stout, cauleseent,branching,1-3 m. high, the foliage scaberulous,the infloreseence puberulent; leaves deltoid, 2-3.5 dm. long, ternatelydecomiipound, the ultimatedivisionslinear-oblong to filiform, acute with a prominent callus point,cuneate at base, sessile,distinctor the terminalconfluent, 1-6 cm. long, 0.5-4 (-10) mm.broad, entire (in our material), scaberulous on the veins and marginsbeneath; petioles 1-4 dm. long, sheathingat base; cauline leaves like the basal, the uppermostusually opposite, greatly reduced and often simple, with obsolete sheaths; infloreseeneeeymosely branched,of several slender peduncles 3-12 cm. lolig; involuerewanting; involueel wanting,or vestigial; rays mostly 10-25, sleinder, spreading-ascending, subequal, 1.5-3 cm. long,slightly webbedat base and oftenpuberu-

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usually 2-5 mm. long, webbed at lent; pedicels short,spreading-ascending, stylopodium conic,the greenish-yellow; base and oftenpuberulent; flowers styles short,erect or spreading; carpophore2-cleftto the base, lax; fruit ovoid-oblong, 6-8 mm. long, 3-4 mm. broad, tapering at apex and base, the ribs prominent, acute; oil tubes large, solitaryin the intervals,2 on the commissure;seed channeledunder the tubes,the face deeply suleate. TYPE: "on the hills at Rio Hondo," Mexico, Pringle 3620. North American distribution: Hidalgo to Durango, south to Mexico (state) and Oaxaca, Mexico.
South American specimeiisexamined: COLOMBIA. MAGDALENA: Paramos of the Sierra Nevada de Santa Marta, about 30 miles inland from Dibulla, ca. 3850 m., VII-32, W Seifriz 4,2 (US).

in South America is a single of this enitity The basis for the recognition fragmentary, badly molded specimen,but thereseems to be no question of its identity.Dr. Seifriz's collecting clearly suggests that additional field work in the mountainsof Magdalena mightprove exceedinglyrewarding. Not Linnaea 9: 607. 1834; nomen conservandum. Tauschia Preissler, 1828. Two entities from South America have been referredto this genus, which has about twentyspecies in Mexico and the westernUnited States. The genericlines betweenTa-uschia and Arracacia have been shiftedand reof opinion clearly reveal that drawn numeroustimes,and these differences break. My recollections thereis no sharp morphological of strugglingwith this problemare still too vivid, however,to make me want to reopen the questionnow. 1. TAUSCHIA NUDICAULIS Schlecht.Linnaea 9: 608. 1834. this species,knownfromJalisco south to Vera Cruz, To my knowledge, Mexico (state), and Puebla, has been reportedfromSouth America only on the basis of Spruce 6065, "in Andibus Ecuadorensibus." According to Coulter and Rose (1900), "Spruce's specimen from South America is similar to the Mexican specimensin habit, but with a somewhatdifferent fruit section and it may yet prove a distinct species." Judging by the if any, are by no means consheet at the Gray Herbarium,the differences, that Ottoa oenanthoidesH.B.K. has a comspicuous. It may be significant from Guerreroand Oaxaca, Venezuela, being knowin parable distribution, Colombia,and Ecuador. 2. TAUSCHIA JAHNII Rose ex Pittier,Man. P1. Usual. Venez. 299. 1926 (nomen subnudum.). This name was picked up by Pittier fromRose's annotationof certain Venezuelan collections,of which I have seen the fQllowing:"La Puerta, Trujillo, 2000 m.," 16-IX-22, Jahn 1136 (US) and "Sierra Nevada de Merida, 10,000-16,000feet," XII-23, de Bellard 259 (US). Both collections are fragmentary and appear to agree in all respects,includingthe lack of to matchthesein the Gray Herbarium, any maturefruit.While attempting I discovereda third and much bettersample, but still fruitless,in Ghiesbreght687, "au bord de ruisseaux et des sources d'eau dans les montagnes, has recentlycollected the plant twice Chiapas, etc., Mexico." Steyermark in Venezuela: "LARA: betweenBuenos Aires and Paramo de las Rosas,
TAUSCHIA Schlecht.

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2285-3290 i.," 11-11-44, 55,455 (F) and 55,475 (F), likewise in an inmaturestate. Althoughall these specimensare superficially reminiscent of Apium graveolensL., they are apparently much more closely related to Arracacia eduwlis S. Wats. and Tauschia nudicaulis Schlecht.,and perhaps constitute an undescribedspecies in one of these two genera, which would be notable for its leafy stems, small umbels, entire bractlets,and small, bluntfruits(immature).' NEONELSONIA Coult. & Rose, Contr.U. S. Nat. Herb. 3: 306. 1895. 1. NEONELSONIA ACUMINATA (Benth.) Coult. & Rose ex Drude; E. & P. Nat. Pfl.38: 167. 1898. Arracacia acuminata Benth. P1. Hartw. 187. 1856. Althoughthis species is commonlyconfused with Arracacia,PennellUii, A. Wiggitnsii, and A. elata because of a close similarityin vegetativecharacters,its ellipsoid-cordate, wrinkledfruit and filiform bractletsand pedieels make it readily distinguishable. is known from acuminatac Neornelsonia Colombia to Ecuador. 2. NEONELSONIA Sp. ? To the collectionPennell 2452, "Forest, on mount,2-4 m. s. of Sibate, Cundinamarea, Colombia, 2900-3000 m.," 13,15-XII-17 (NY; F, USphotos), Rose gave an herbariumname under Arracacia, presumablyin recognitionof the laxity of the filiform pedicels. The young fruits are broadly ovoid, but too young to be assigned with any certaintyto the proper genus. The general similarityof the collection to the preceding species, however, suggeststhat it may representan undescribedspecies of Neonelsonia,distinguished fromN. acumnata by its mucronateratherthan spinulose serrations, acute ratherthan acuminate leaflets, and shorterand broader bractlets.
MYRRHIDENDRON Coult. & Rose, Bot. Gaz. 19: 466. 1894. As revisedby Coulterand Rose (1927), this genus containsfour species, one in Guatemala and Costa Rica, the second in Panama, and two othersin northern South America.The very large linear or oblong and dorsallyflattened fruitis ample to distinguishthese plants fronmi any species of Arracacia, but vegetativematerial of the two genera is commonly confused. 1. MYRRHIDENDRON GLAUCESCENS (Benth.) Coult. & Rose, Jour. Wash. Acad. Sci. 17: 214. 1927. Arracacia glaucescensBenth. P1. Hartw. 187. 1845. This is presumablythe more commonof the two species, being known from Colombia to Ecuador. The almost coriaceous leaves with a raised stipular ring at the base of the petiolules and the dissected bractletsdistinguishit fromany Arracacia knownto occur in the same area. 2. MYRRHIDENDRON PENNELLII Coult. & Rose, Jour. Wash. Acad. Sci. 17: 214. 1927.

' While filing the accumulated Umbelliferae in the Herbarium of the University of California, the writer discovered two more collections of this entity in Steyermark 's nos. 48,460 and 49,771 from Guatemala. The second of these has semi-maturefruit, and it is clear from a study of these that all the plants mentioned above under Tauschia Jahnii Rose are conspecificwith Arracacia vaginata Coult. & Rose, hithertoknown to occur only from Michoacan to Oaxaca, Mexico. Thus Arracacia vaginata, with a range extending from Michoacan to Venezuela, constitutes the eleventh species of its genus thus far reported from South America.

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Known only fromColombia,this species differs fromthe preceding in having lanceolate, acuminate, sharply serrate,rather than ovate, obtuse, incised and lobed as well as serrate leaflets,and in having the stipular ring denselyhairy.
GRAY HERBARIUM, HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS DEPARTMENT OF BOTANY, UNIVERSITY OF CALIFORNIA BERKELEY, CALIFORNIA

Literature Cited
Hooker, W. J. 1831. Arracacia esculenta. Eatable Arracacha. Bot. Mag. 58: pl. 3092. Coulter, J. M. & Rose, J. N. 1900. A synopsis of Mexican and Central American Umbelliferae. Proc. Wash. Acad. Sci. 1: 134, 136. . 1927. Revisioniof the genus Myrrh1idendron. Jour. Wash. Acad. Sci. 17: 213-215. Mathias, M. E. & Constance, L. 1944. Umbelliferae. N. Am. Fl. 28B: 90-102.

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