Reserva Biolgica Allpahuayo-Mishana, Iquitos, Loreto, PERUPhoto Guide to the BURSERACEAE of Allpahuayo-Mishana

(Useful throughout the western Amazon)b y

Paul FinePhotos: Paul Fine. Produced by: P. Fine, S. Kaplan, M. Giblin, T. Wachter, R. Foster. Support from: A. Mellon Foundation and G. & B. Moore Foundation. P. Fine [paulfine@umich.edu] Assistance from Instituto de Investigaciones de la Amazona Peruana IIAP, University of Utah, and Italo Mesones. Env. & Cons. Programs, The Field Museum, Chicago, IL 60605 USA. [RRC@fmnh.org] [www.fmnh.org/plantguides/] Rapid Color Guide #176 version 1

Burseraceae is a cosmopolitan family of mostlylowland tropical trees and shrubs. In Peru, it isrepresented by six genera (five can be found in theAmazon basin and are included in this guide) andaround 45 species. At least 36 of these species havebeen collected in the Allpahuayo-Mishana Reservenear Iquitos (Loreto). Most of these species havewide distributions; therefore, this 34-species photokey will be useful for most areas throughout thewestern Amazon. For more Burseraceae picturesfrom the Neotropics, please seehttp://fm2.fieldmuseum.org/plantguides/default.asp.

Protium sagotianum

Identifying BurseraceaeThis guide is designed for identifying saplings and seedlings of Burseraceae plants, between ~1m and 2m in height.Burseraceae saplings have alternate, pinnately compound leaves and (with few exceptions) a distinctive odiferousresin. This resin is present in the twigs, the leaves, and the trunk of the tree, and when apparent, it is either a clearsticky resin (which often dries white) or a milky white latex. Burseracs almost always have a distinctive odor: theyeither smell sweet or have a strong incense or sharp turpentine odor. Even the sweet-smelling species have a trace ofthe incense odors that add a unique depth to the scent that separates them from other sweet-smelling trees, forexample, the Meliaceae. Other families have compound leaves and distinctive odors, like the two closely relatedfamilies Anacardiaceae and Rutaceae, which can have resins and/or odors. Anacards generally have a clear resin andsmell more like mangoes, and their resin almost always dries black. Rutaceae have no resin, but have a citrus-likeodor.

Many of the characters presented in this guide can also be used for much larger individuals, including adults.Keep in mind that for leaves in the canopy, many of the compound leaves become thicker, smaller, with fewerleaflets, and often exhibit somewhat less distinctive venation patterns. Also worth noting is that in very smallseedlings (less than 50cm in height), many of these species have simple leaves so although the odor and venationpatterns of the single first leaves can help you get to a species identification, the information concerning number andsize of the leaflets in this guide will be of no help.

Intrafamilial relationshipsAs seen in Figure 1, the intrafamilial relationships of the

Peruvian Burseraceae genera are not completely worked out.While morphological and molecular evidence strongly support theidea that the Canariie (Dacryodes and Trattinnickia) are eachothers closest relatives, and the Protieae (Crepidospermum,Tetragastris, and Protium) form a natural group, whether or notthese five genera form a monophyletic groups is not resolved.Recent molecular work has shown that there may be species ofProtium that are more closely related to Tetragastris andCrepidospermum than they are to other Protium species.Similarly, a recent unpublished analysis of Dacryodes andTrattinnickia found that Dacryodes cuspidata appeared to be asclosely related to Trattinnickia as it was to other Dacryodesspecies. While research continues on the classification of these

Anacardiaceae

Dacryodes

Trattinnickia

Bursera

Crepidospermum

Tetragastris

Protium

Figure 1: Phylogenetic relationships of thenamed genera in the Burseraceae

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groups, we can still recognize many natural groups in the field. This guide is focused on the species level, and whilesome of the genera do form recognizable natural groups (like Crepidospermum), most of the species are bestseparated using the combination of characters I have devised for field identification.

How to use this guideAll of the 36 species included are listed with at least 3 photos. The largest photo is the underside of a single leaflet,emphasizing my belief that the venation pattern on the underside of the leaf is ultimately the best character in tellingapart Burseraceae trees. There is a smaller picture of the entire compound leaf, with a yellow 5cm scale bar. Thenthere are one or two small pictures that focus in on the unique characteristics of the species that will help to clinchthe identification. Below each species name is a white box with the list of characters I have used to help separatethe species. The characters are 1) latex (resin), 2) odor, 3) leaf margin, 4) size and number of leaflets and 5) leaftexture.

On the compound leaf picture, with the scale bar, are one or two colored circles white, brown and red.These colors correspond with the soil types in which the species have been found in my extensive Burseraceaeinventories at Allpahuayo-Mishana (Fine et al. in press). White stands for white sand forest, brown is for brownsand or eroded river terraces, a common sandy clay soil type found in Allpahuayo-Mishana. Red is for clay soil.These soil types can be verified in the field by digging a machete several centimeters below the litter layer andchecking the consistency of the soil (sandy or clayey) and the color of the soil. Clay can be gray, red, or purple;brown sand can be reddish-orange to brown; and white sand can be gray or white, fine or coarse sand. If a species isfound in two soil types, its more common soil type is on the bottom.

Protium sagotianumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: VARIABLE AND MANYTEXTURE: SMOOTH

Secondaries prominent,symmetrical and forming loopsat margins. Long leaf apex.

Burseraceae References:Clarkson, J. J., Chase, M. W., Harley, M. M. 2002. Phylogenetic relationships in Burseraceae based on plastid

rps16 intron sequences. Kew Bulletin 57:183-193.Daly, D. C. 1987. A Taxonomic Revision of Protium (Burseraceae) in Eastern Amazonia and

the Guianas. Ph.D. thesis (City University of New York), New York, NY.Fine, P. V. A., Daly, D. C, Villa Muoz, G., Mesones, I., Cameron, K. in press. The contribution of edaphic

heterogeneity to the evolution and diversity of Burseraceae trees in the western Amazon. Evolution.Weeks, A., Daly, D. C., Simpson, B. B. 2005. The phylogenetic history and biogeography of the frankincense

and myrrh family (Burseraceae) based on nuclear and chloroplast sequence data. Molecular Phylogeneticsand Evolution 35:85-101.

underside ofleaflet, showsvenation

entirecompound

leaf

list ofcharacters,description

5 cmSoil type in whichspecies is found.

= white sand = brown sand = clay

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Asperous Leaves: DACRYODES (1) & TRATTINNICKIA

Dacryodes cuspidataLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: SMALL AND FEWTEXTURE: ASPEROUS

Protium-like petiolules,irregular leaflet margin andvenation.

Trattinnickia glazioviiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: SMALL AND MANYTEXTURE: ASPEROUS

Leaflets are papery with longapices, petiolules are longand thin.

Trattinnickia peruvianaLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: ASPEROUS

Leaflets are subcoriaceouswith short apices.

Serrated Margins: CREPIDOSPERMUM (+ 2 Protium on following page)

Crepidospermum goudotianumLATEX: CLEARODOR: SWEETMARGIN: SERRATEDLEAFLETS: SMALL AND MANYTEXTURE: SMOOTH

Like P. subserratum, but nowhite latex, and veinsterminate in the marginsteeth.

Crepidospermum pranceiLATEX: CLEARODOR: SWEETMARGIN: SERRATEDLEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

No petiolules, dense hairson leaf and rachis.

Crepidospermum rhoifoliumLATEX: CLEARODOR: SWEETMARGIN: SERRATEDLEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Can be hairy like prancei,but distinct petiolules.

Protium subserratumLATEX: WHITEODOR: NONEMARGIN: SERRATEDLEAFLETS: SMALL AND MANYTEXTURE: SMOOTH

Leaflets sessile, no distinctpetiolules. larger individualscan have entire leaflets.

Protium ferrugineumLATEX: WHITEODOR: NONEMARGIN: SERRATEDLEAFLETS: SMALL AND MANYTEXTURE: SMOOTH

Well-defined petiolules,larger individuals alwayshave entire leaflets.

Protium gallosumLATEX: WHITEODOR: NONEMARGIN: ENTIRELEAFLETS: LARGE AND FEWTEXTURE: SMOOTH

Glabrous and very shinyleaflet surface, with a longapex.

Protium amazonicumLATEX: WHITEODOR: NONEMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Dull leaf surface, midveinsoften hairy and reddish,short leaflet apex.

Protium calanenseLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: SMALL AND FEWTEXTURE: SMOOTH

Sometimes labeled P. aracouchini.Secondaries prominent. Completelyglabrous underside that reflects light(unlike similar spruceanum andkrukoffii).

Protium altsoniiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELFLTS: SMALL AND MANYTEXTURE:SMOOTH

Secondary veins are not prominent,glabrous leaflet underside reflectslight.

Protium heptaphyllum subsp. uleiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELFLTS: SMALL AND FEWTEXTURE: SMOOTH

Secondaries not prominent,glabrous underside that reflectslight. Similar altsonii always hasmany pairs of leaflets; heptaphyllumnever does.

Protium hebetatumLATEX: CLEARODOR: SWEETMARGIN: UNDULATINGLFLTS: SMALL AND MANYTEXTURE:SMOOTH

Underside of leaf can be hairy.Secondaries prominent andbrochydodromous and close, well infrom the margin.

Dacryodes peruvianaLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

No petiolular swellings, indistincttertiaries, and sweeter odorseparate from similar P.nodulosum.

Protium grandifoliumLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Very symmetrical, prominentsecondaries. Thicker leavesand stronger smelling latexthan similar P. opacum.

Protium opacumLATEX: NONE (OR CLEAR)ODOR: SWEET OR NONEMARGIN: ENTIRELEAFLETS: LARGE AND VARIABLETEXTURE: SMOOTH

Golden spots on rachis andbranchlets. Leaf very similar toP. nodulosum but no turpentineodor, and has a more paperytexture.

Protium nodulosumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE:SMOOTH

Unique yellowish green color ofleaf underside. Like P. opacumbut with oily latex that has a verystrong turpentine odor and driesto a visible white powder.

Protium decandrumLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND FEWTEXTURE:SMOOTHShort distance between petioleand basal leaflets. Prominentsecondaries on adaxial sidebut prominent midvein onleaflet underside (unlike similarpaniculatum).

Protium divaricatum subsp. divaricatumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: LARGE AND FEWTEXTURE: SMOOTH

Leaves almost alwaystrifoliolate. Fibrous corky woodin center of branchlets.

Protium paniculatumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: LARGE AND FEWTEXTURE: SMOOTHYellowish midvein is imbeddedin leaf underside, smooth tothe touch. Very stronglyturpentine-smelling resin drieswhite. Trees producecollectable resin lumps (copal).

Protium pallidumLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTHWhite nodes on petioluleand leaf rachis are unique.Prominent secondaries onadaxial surface, smooth andinconspicuous on leafunderside.

Protium glabrescensLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Large nodes where leafletsattach. Secondaries do notform loops at margin. Verysticky, strongly sweet-smellingresin.

Tetragastris panamensisLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

No petiolular swellings. Oftena reddish mark at nodes ofleaflet attachment.

Protium sagotianumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: VARIABLE AND MANYTEXTURE: SMOOTH

Secondaries prominent,symmetrical and forming loopsat margins. Long leaf apex.

Protium tenuifoliumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Secondaries prominent andsymmetrical. Leaf apexshorter than similarP. sagotianum.

Protium krukoffiiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: SMALL AND MANYTEXTURE: SMOOTH

Dull leaf surface, leaf apex oftentoothed, lanceolate leaf shape.Secondaries prominent. Fewersecondaries and intersecondariesthan similar P. spruceanum.

Protium spruceanumLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: SMALL AND MANYTEXTURE: SMOOTH

Dull leaf surface, leaf apextoothed, more ovate leaflet shape.Secondaries prominent. Moresecondaries and intersecondariesthan P. krukoffii.

Protium trifoliolatumLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: SMALL AND FEWTEXTURE: SMOOTH

Secondaries prominent andasymmetrical. Leaflet base verycuneate. Often hairy.

Protium divaricatum subsp. krukoffiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: SMALL AND FEWTEXTURE: SMOOTH

Petiolules and rachis long and thin.Prominent midvein, but not sharplike crassipetalum.

Dacryodes chimatensisLATEX: CLEARODOR: SWEETMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Coriaceous and completelyglabrous leaflets, no petiolularswellings.

Dacryodes hopkinsiiLATEX: CLEARODOR: SHARPMARGIN: ENTIRELEAFLETS: VARIABLE AND MANYTEXTURE: SMOOTH

Basal leaflets tiny withasymmetrical base. Indistincttertiary veins.

Protium crassipetalumLATEX: NONEODOR: NONE (SWEET)MARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Long, green cylindricalpetiole, leaflet midveinsvery prominent (feels likeknife edge).

Protium klugiiLATEX: NONEODOR: NONEMARGIN: ENTIRELEAFLETS: LARGE AND MANYTEXTURE: SMOOTH

Winged rachis andpetiolules. Yellow latexoften present in trunk slashin larger trees.