Author(s): Marco Leonti, Fernando Ramirez R., Otto Sticher, Michael Heinrich
Source: Economic Botany, Vol. 57, No. 2 (Summer, 2003), pp. 218-230
Published by: Springer on behalf of New York Botanical Garden Press
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MEDICINAL FLORA OF THE POPOLUCA, MEXICO: A BOTANICAL
SYSTEMATICAL PERSPECTIVE
Estudiamoslas plantas medicinales que usan los Popolucas de la Sierra de Santa Marta,
Veracruz,en el oriente de Mexico.Aplicandoel me6todo de ana'lisisde regresi6nde Moerman,
determinamoscuales taxa utilizadosetnomedicamenteestdn sobre-representados(p. ej. Aste-
raceae) y cua'lestaxa estdnsub-representados(p. ej. Orchidaceae)en lafarmacopeaPopoluca.
Moermany colaboradoresencontraronuna alta correlacion entre diversasfarmacopeas ho-
ldrticasy suponenque este hecho se debe a la semejanzade las floras borealesy a la existencia
de un "cuadro comu'nde sabidurfa humana." Los Popolucas, quienes viven en un medio
ambientedominadopor la flora neotropicalmezcladacon elementosboreales, incluyenmenos
taxa neotropicalesen su farmacopea de lo que se hubieraesperado. Este resultadoapoya la
teoria expresadopor Moermanet al. sobre la influenciade un conocimientode la etnofar-
macopea holartica que ha sido transmitidodesde tiemposprehist6ricos.Sin embargocompro-
bamos que existe una clara influencianeotropical en la farmacopea Popoluca por lo cual
concluimosque ademdsde los conocimientossobre taxa especificostransmitidoshist6ricamen-
te, el hipotetico "patr6ncomu'nde sabidurfahumana"citado por Moermanet al. se refiere
tambiena "criterioscomunesde selecci6n."
1
Received 10 October2002; accepted 15 March2003.
Researchon medicinaland otheruseful plants the relatively high correlationbetween the ho-
used in indigenous societies has been driven by larctic ethnopharmacopoeiasand the low corre-
two complementaryinterests:The use of such lation between the holarcticpharmacopoeiasand
informationfor researchin the field of the nat- the neotropicalone is due to the relatednessof
ural sciences, especially with regard to 'new' the northernfloras and the possibility that the
bioactive naturalproducts derived from plants knowledge about medicinal plants has been
and the use of plant extracts in primaryhealth passed on from prehistorictimes throughspace
care (Heinrichand Gibbons2001) and the inter- and time. This implies that the peoples entering
est in better understandingthe anthropological the New Worldthroughthe BeringianStreet al-
basis, if possible on a cross-culturalbasis, of the ready shared a common knowledge about me-
use of these resources by humans and particu- dicinalplantswith the peoples migratingto other
larly on the rationale(s)behind the selection of areas and remainingon the Asiatic continent.
these resources(Moermanet al. 1999). There is a widespreadacceptancethat North
One particularlyexciting developmentwhich America was first settled at least around15 000
contributesto both lines of investigationis the yr B.P., but some authorspostulatea date as early
studyof botanicalsystematicaspectsof plantus- as 40,000 yr B.P. (Layrisse and Wilbert 1999).
age. Dan Moerman and his colleagues (e.g., According to these authors,three major waves
1996, 1998a,b) have developed and used (par- of immigrantspopulatedthe Americas.The first
tially in collaborationwith several other schol- wave was of Pre-Mongoloid tribes at around
ars) a methodwhich allows for a statisticalanal- 40 000 yr B.P., followed by Paleo-Mongoloidsat
ysis of ethnobotanicalinformationbased on that 30 000 yr B.P., and the Diego-allele positive
the numberof medically used plant taxa and the Neo-Mongoloids at about9000 yr B.P. (Layrisse
total number of taxa in a certain region is and Wilbert 1999). When peoples crossed the
known. borderof the holarcticflorainto the realmof the
Most of their researchfocused on the North neotropicalplant kingdom they were suddenly
temperateregion of North America, but also on confrontedwith plant families that were new to
three additionalnorthern(Kashmir,Korea,Chia- them and had to adapt themselves and their
pas) and one southern (Ecuador) hemispheric pharmacopoeiato this new pool of plants.
ethnopharmacopoeias. Only one neotropical We have recently completed a detailed study
lowland region (upper Napo River valley, Ec- of the medicinal plant use of the Popoluca in
uador)is includedin his analyses.The main rea- southernVeracruz,Mexico (Leonti et al. 2001).
son for this is a lack of taxonomic and system- The Popolucainhabitthe Sierrade SantaMarta,
atic researchin such regions. The results of the a range of volcanoes between the Lake of Ca-
comparisonby Moermanet al. (1999) show that temaco and the Gulf coast. These volcanoes
holarctic peoples rely on similar plant families form the southernfoothills of the "Sierrade Los
in their health care. The Pearsoncorrelationfac- Tuxtlas" mountainrange, a region particularly
tors between the holarctic data sets are higher well known for its biodiversity, where the ho-
than 0.6 while the correlation of the Ecuador larctic and neotropicalfloristic kingdoms over-
data set with the holarcticdata set is lower than lap. The neotropicalinfluence is strongerthan
0.19 (maximumpossible correlationis one, low- the holarctic one. According to Rzedowski
est is zero). Moermanet al. (1999) suggest that (1991) the overwhelming part of the Mexican
220 ECONOMICBOTANY [VOL. 57
4'~~~~~~~~~~~~~~~~~~~~~~4
so it 2
Fig. 1. Regressionanalysis of the medicinalflora of the Popoluca.X axis, numberof species per family in
the flora;Y axis, numberof medicinalplants per family.
TABLE 1. RANKING OF THE FIVE MOST USED PLANT FAMILIES IN VERACRUZ AS COMPARED TO THE
DATASET FROM MOERMAN ET AL. (1999).
North
Family Veracruz Chiapas' Kashmir' America' Korea' Ecuador'
Asteraceae 1 1 1 1 3 45
Piperaceae 2 105 237 89 27
Fabaceae s.l. 3 138 85 253 8 2
Euphorbiaceae 4 21 2 234 13 90
Lamiaceae 5 2 4 8 4 91
Total families 174 144 100 255 136 118
' FromMoermanet al. 1999.
222 ECONOMIC BOTANY [VOL. 57
TABLE 2. RANKING OF THE FIVE LEAST USED PLANT FAMILIES FROM VERACRUZ AS COMPARED TO THE
DATASETFROMMOERMANET AL. (1999).
North
Family Veracruz Chiapas' Kashmir' America' Korea' Ecuador'
analysis would have been performedwithout species and do not occur spontaneously.The rest
the introducedspecies the Lamiaceaewould of the Asteraceae used by the Popoluca are all
rank 34th and be substitutedby the Acantha- weeds and normally gatheredin the immediate
ceae, vicinity of the community.
2) Phytochemical characteristics of the plant Asteraceae flower heads are attractive and
families relating to the presence of pharma- conspicuous. Although there are many species
cologically active secondarymetabolites, with similarlylooking yellow flower heads, and
3) Organolepticalcharacteristicsof the mem- hence are difficult to distinguish. Therefore in
bers of a family, e.g., aromatic species are folk taxonomy some species are consolidated
very prominentin the Lamiaceae (Leonti et into one taxon.
al. 2002).
Piperaceae. The Piperaceaeare predominantly
In the following the five top and bottom
used to treat dermatologicalcomplaints (Piper
rankedfamilies are discussed separately.
spp., Pothomorphe sp.) and skeleto-muscular
ToP RANKEDFAMILIES problems (Peperomia spp.) such as rheumatic
conditions. The Piperaceaeare rich in monoter-
Asteraceae. The Asteraceae are the Popoluca's
penes, sesquiterpenes,phenylpropanesand am-
most frequentlyused family, and take priorityin
ides, with a variety of significantpharmacolog-
the treatmentof illness groupsof gastrointestinal
ical effects recorded(Frohneand Jensen 1998).
disorders,dermatologicalailments,skeleto-mus-
Out of the 22 medicinal species only Piper au-
cular problems, respiratory ailments and are
ritumKunthis cultivatedregularlyin home gar-
very important(2nd most often used family) in
dens, mainly for culinary reasons to flavour
gynaecology (see Table 3). The Asteraceae are
chicken soup, pork stew and pork tamales with
phytochemicallyvery diverse and so far at least
its leaves. The main constituentof P. auritum
7000 natural compounds have been isolated
leaves is the carcinogenicsafrole and thus may
from chemical classes such as sesquiterpenelac-
constitutea health risk if consumedexcessively.
tones, diterpenes,phenols, and polyenes (Frohne
The genus Piper and the genus Peperomiaare
and Jensen 1998). This varietyof chemicalcom-
morphologicallyhomogeneousand thereforethe
pounds has been recorded to have a multitude
species are difficult to separatefrom each other.
of pharmacologicalactivities including anti-in-
In Popoluca folk taxonomy all species from the
flammatory,cytotoxic, bactericidal, fungicidal,
genus Piper except P. auritum (Acuyo) are
and appetite-inducingproperties.The pharma-
called 'Tooso' and are conceived as havingmore
cological propertiesof the compoundclasses re-
or less the same virtue.Therefore,the high rank-
flect the broad therapeuticapplicationwith the
ing accordingto the residual of the Piperaceae
Popolucaand may in partexplain the Popoluca's
seems to be largely due to the family's typical
reliance on the Asteraceae.
characteristicsin combinationwith the insepa-
Of the 57 Asteraceae species used seven
rabilityof many of the highly diversePiperaceae
(12%, Tagetes spp., Artemisia ludoviciana Nutt.,
(according to Popoluca concepts) and the Po-
Porophyllum ruderale (Jacq.) Cass., Leonti 224)
poluca's indiscriminateuse of these species.
are regularlygrown in the home gardensfor me-
dicinal purposes. They are mainly introduced Fabaceae, s.l. The Fabaceae, s.l. are most im-
2003] LEONTIET AL.: MEDICINALFLORAOF THE POPOLUCA,MEXICO 223
marily used as a vegetable. The Euphorbiaceae macopoeiathe genus Lasiacis is used for a spe-
have very inconspicuous flowers or inflores- cial dermatologicalcondition, but, as a whole,
cences but the generaare visually distincton the the Poaceae are importantin none of the illness
basis of morphologicalcharacters(herbs,shrubs, groups.Out of 93 species, five are medicinaland
trees) and all grow as weeds in the vicinity of two of these are introducedspecies cultivatedin
the villages. Taste and smell characteristicsmay house gardens: Bamboo, Bambusa sp., apart
well be importantin selecting and recognizinga from its medicinal usage is primarilyused for
species. construction,and Cymbopogonsp., rich in es-
sential oils such as citronellal and geraniol, is
Lamiaceae. The Lamiaceae are used in gynae-
used to treatgastrointestinalcomplaints.
cology and to treatgastrointestinaldisorders.Es-
While this family is abundantthroughoutthe
sential oils, tannins,bitter diterpenoids,and iri-
region and includes many weeds, the species are
doid glycosides are typical Lamiaceaeconstitu-
normallyvery inconspicuousand difficultto dis-
ents with antimicrobial,antimycotic, antiviral,
tinguish, especially in the sterile state. This
anti-inflammatory and choleretic properties
makes them unlikely candidatesto be selected
(Frohne and Jensen 1998). Of the twelve La-
as a medicine.
miaceae in the pharmacopoeiasix introduced
species, which do not grow spontaneouslyare Rubiaceae. Characteristicof this family are xan-
regularlycultivated in the home gardens.Plec- thine derivatives,anthranoides,coumarines,tri-
tranthus amboinicus (Lour.) Spreng. (Leonti terpenes, triterpenesaponines, proanthocyanidi-
253) is a medicinal plant and is often used as a nes, and indol- and emetane alkaloids (Frohne
herb to flavourmeat soups. NeitherRosmarinus and Jensen 1998). In the illness categories der-
officinalis L, nor Ocimum basilicum L., and 0. matologicalconditions and gynecology, the Ru-
micranthum Willd. (Leonti 54) are used in the biaceae are the thirdmost frequentlyused family
kitchen. and thereforeof considerableimportancein the
Popoluca pharmacopoeia. Coffea arabica L.,
LEAST USED FAMILIES coffee, is cultivatedas a cash crop in the higher
Orchidaceae. The pseudobulbs of this family regions and rarelyused as a source of medicine.
are best known for their starch and glucoman- Of the 26 medicinallyused species two are reg-
nane mucilage. The presence of a variety of al- ularly grown in home gardens. Sickingia mexi-
kaloids such as pyrrolizidineand phenylisochi- cana Bullock (Leonti272) used in gynaecology
noline alkaloids has been reported(Frohne and has become a rare species in the past years due
Jensen 1998). The Orchidaceaeare not impor- to deforestation.Recently, midwives began to
tant in any of the illness groups; only two out cultivate this tree, simultaneouslysecuringtheir
of 99 species are medicinals. The mucilage of medicine and contributingto the protectionof
Maxillaria tenuifolia Lindl. (Leonti 504) is used biodiversity. Gardenia sp. is mainly grown for
as naturalglue in the constructionof 'jaranas,' ornamentalreasons but also has medicinaluses.
(traditionallittle guitars). The Orchidaceaeare Some species of the Rubiaceae are utilised ex-
often rare,have relatively scattereddistributions tensively by the Popoluca,e.g., Hamelia patens
and often inaccessible populations,which com- Jacq. used for bleeding wounds, which is the
plicates the acquisition of plant material.Also, species with the second largest number of use
in the region of the Popoluca,the orchidsdo not reportsin the whole ethnopharmacopoeia (Leon-
grow as weeds in the secondary vegetation ti et al. 2001) but the fact that the family is rep-
zones aroundthe villages (Stepp and Moerman resented by many species, of which a over-
2001). whelming majoritygrows in primaryforest re-
mote from the Popolucavillages resultsin a rel-
Poaceae. Silicate is the main chemical charac-
low ranking. Also the Rubiaceae are
teristic of this family. Normally,low concentra- atively
(especially in the sterile state) very inconspicu-
tions of coumarine derivatives and cyanogenic
ous (Psychotriaspp.), which probablyinfluences
glycosides are common (Frohne and Jensen
the selection of species from this family.
1998). Zea mays L., maize, is the most widely
cultivatedfood plant in the region and in Mex- Cyperaceae. The Cyperaceae have a similar
ico, in general,but is used medicinallyonly very chemical spectrum as the Poaceae. Silicate,
rarely by the Popoluca. In the Popolucan phar- proanthocyanidineand essential oils are the
2003] LEONTIET AL.: MEDICINALFLORAOF THE POPOLUCA,MEXICO 225
TABLE 4. PEARSONCORRELATION.
plant kingdoms, although the neotropicalinflu- may well be partlydue to the inconspicuousness
ence is higherin the floraof Veracruz.The most of many of its plants. The most prominentfam-
prominentdifference is that the Popoluca dwell ilies of the MC are the Araceae (rank 8), the
in the lowland and the Tzotzil/Tzeltal in the Commelinaceae(rank10), and also the Liliaceae
highlands. Both groups are part of the Macro- (rank 13); the lattertwo are families with many
Mayanlanguagestock and are culturallyrelated. very showy species. All three families show
The culturalseparationmay have occurredabout clear differences in their ethnopharmaceutical
2500 years ago. The high correlationof their uses. We documented 18 of 62 medicinal MC
pharmacopoeiasprobablyreflects both the bio- species used for urological problems, with 84
logical relatednessof the area and the common (23.5%) of 357 total MC use reports,while 136
culturalpast of the two groups. If we compare of 535 medicinalDC species have a total of 253
the relatednessof the flora of the Populucaand (6.2%) reportsin this group (DC-total number
the medicinal flora selected by them with other of use reports:4029). The MC are thus more
floras/medicalfloras (Table4) an analysis using likely to be selected for urologicalproblems.
the Pearsoncorrelationindicates the relatedness Out of the 62 medicinalMC species, only two
of the Popolucan (Veracruzian)ethnopharma- (3.2%) with a total of five (1.4%)use reportsare
copoeia with the others. The only exception is used to treatrespiratoryailments,while 73 spe-
the data for Ecuador. The flora of Veracruz cies of the 535 medicinal DC (13.6%) with 198
shows a correlationof 0.83 with Ecuadorwhich (5%) use reportswere recorded.
would predict a correlationof their pharmaco- Why should a MC be more suitable to treat
poeias of about 0.75 if the relatednessof their urologic conditions and less useful to treat re-
flora would be the only influencingfactor.How- spiratory ailments? The empirical medico-his-
ever, the correlation between the Veracruzian toricalexperienceof an ethnic groupis of course
and the Ecuadorianpharmacopoeiafalls at the an essential basis for this, but at this stage the
much lower level of 0.36. specific reasonsremainspeculative.Phytochem-
The Veracruz(V) and the Chiapas (C) phar- ical as well as organolepticaldifferences exist.
macopoeia perform similarly to the other phar- Concerning the phytochemistry,the MC lack
macopoeias. Both show the highest correlations ellagic acid and ellagic tannins,whereastannins
with Kashmir (0.74 C/0.62 V), second highest and essential oils, polyterpenesand alkaloidsare
with NorthAmerica (0.73 C/0.57 V), thirdhigh- not common, in general. Steroid saponinsoccur
est with Korea (0.61 C/0.54 V), and the lowest in MC while the DCs have mostly triterpene-
with Ecuador (0.19 C/0.36 V). The correlation saponinsand the mucilage of the MC lacks uro-
of the Chiapasdata set to the other holarcticset nic acid (Frohne and Jensen 1998). Regarding
of data is always higher as comparedto Vera- the morphology,the MC distinguishthemselves
cruz. On the otherhand the Veracruzcorrelation by not showing secondary growth, having a
to the neotropical(Ecuador)set of data is con- moderatediameter,an un-branchedaxis, and a
siderablyhigher than the one from Chiapas. watery stem. However,we arguethat the differ-
Thus, the Pearson correlation factors show ent spectraof secondarynaturalproductsalong
that the Veracruz flora is most related to the with the recorded pharmacologicalproperties
nearby Chiapas flora but is more related to the are not sufficientto explain the unequaluseage
Ecuador flora than to any true holarctic flora. of these two groups. Such products are indeed
Nevertheless, the Pearson correlationfactors of responsiblefor specific organolepticalproperties
the differentpharmacopoeiasshow that the Po- of a botanical drug, a set of criteriawhich we
poluca pharmacopoeiais generally more related have previouslyshown to be an importantselec-
to the holarctic pharmacopoeiasbut shows a tion criterionfor medicinal plants with the Po-
clear neotropicalinfluence. poluca (Leonti et al. 2002) and other groups
(Heinrich1998; Brett and Heinrich1998). In the
DICOTS COMPAREDTO MONOCOTS classificatorysystem of the Popoluca and other
The classical, but now controversial,separa- peoples all these factors-the organoleptical
tion of the Angiosperms into Monocots (MC) perception of a plant, the morphological and
and Dicots (DC) allows a useful analysis of still ecological traits,and the characteristicsin terms
anotheraspect. Overall, the MC seem to be less of colour,as well as its smell and taste, are used
importantas compared to the DC. Again this in combination.In case of the preferentialtreat-
2003] LEONTIET AL.: MEDICINALFLORAOF THE POPOLUCA,MEXICO 227
ment of urological ailmentswith MC and of the the common selection criteria may be best ex-
respiratoryconditions with DC we consider the plained with the similar perceptualappreciation
organolepticalaspect to be more importantthan of human beings. We argue that the common
direct pharmacologicaleffects. knowledge is due to common selection criteria
and not to just a common knowledge sharedby
CONCLUSION differentcultures.
The fact that the Popoluca dwell in an area This researchalso has implicationsfor other
where the neotropical and the holarctic plant studies focusing on the selection of interesting
kingdoms overlap makes an ethnobotanicaland and relevant ethnomedicinalplant species for
a botanical-systematicanalysis of special inter- phytochemical and pharmacologicalinvestiga-
est. Our study provides additionalevidence for tions. Not all plants of an ethnopharmacopoeia
the concepts developed by Moermanfor the rea- fit into the emic classificationsystem. Detecting
sons for selecting certain plant taxa as medi- such outliers and subjectingthem to pharmaco-
cines. The ancestors of the Popoluca, dwelling logical studies could result in a higher hit rate
in a zone with neotropicalas well as holarctic- for promising phytochemical compounds and
flora, could choose to a certain extent from a would at the same time advanceour knowledge
similar pool of plant families as the holarctic about such elements of the indigenous ethnop-
peoples to compile theirpharmacopoeiabut also harmacopoeias and their pharmacological ef-
have a vast arrayof neotropicaltaxa to choose fects.
from. The correlationfactors show that the Po-
poluca preferand neglect plantfamilies to a sim- ACKNOWLEDGMENTS
ilar extent as holarcticpeoples do. Nonetheless, We are gratefulto all healers,midwivesand the inhabitantsof the
Popoluca sharingtheirculture,theirfriendship,and theirhospitality.
the Popoluca include neotropicalplant families We thankforthe specialistsof the MexicanNationalHerbarium(MEXU),
in their pharmacopoeia,withoutbeing forced by in particularDr. M. Sousa, A. Reyes, and FranciscoRamos,as well as
the prevailing flora to do so. The Piperaceae, Dra.HeikeVibransof the Colegiode Postgraduados,Chapingo.Prof.D.
Moerman(Michigan)gave us access to unpublisheddetails of his re-
which are the most salient difference as com- search.We are gratefulto the I.N.I. for collaborationand the SEMAR-
pared to a purely holarcticethnopharmacopoeia NAP for the permissionto collect plantmaterial.Financialsupportby
are used to treat dermatologicconditions. Der- the S.R.E. (Secretariade RelacionesExteriores,Mexico D.F) and the
S.D.C. (Swiss Agency for Developmentand Cooperation)is gratefully
matologicafflictionsare very commonin the hot acknowledged.
and humid lowlands of the tropics. Indeed,
based on the numberof taxa used and the use- LITERATURE CITED
reports, diseases of the skin form the largest Ankli, A., 0. Sticher, and M. Heinrich. 1999. Yu-
group of illnesses sufferedby the Popoluca and catec Mayan medicinal plants vs. non-medicinal
is the most importantillness category (Leonti et plants: selection and indigenous characterization.
al. 2001). We suggest that the high prevalence HumanEcology 27:557-580.
and varianceof dermalafflictions in correlation Brett, J., and M. Heinrich. 1998. Culture,perception
with the new climatic conditions led to the in- and the environment.Journalof Applied Botany
corporationof taxa new to the pharmacopoeia. 72:67-69.
Buckles, D., B. Triomphe, and G. Sain. 1998. Cover
The process that led to the incorporationof the
cropsin hillside agriculture:farmerinnovationwith
Piperaceae and other neotropicaltaxa is based Mucuna. IDRC, Ottawa, Canada, & CYMMIT,
on the cultural selection criteriaincluding taste Mexico D.E, Mexico.
and smell propertiesof plants.We recentlydem- Chevalier, J. M., and D. Buckles. 1995. A landwith-
onstratedthe importanceof such concepts in a out Gods: process theory,maldevelopmentand the
completely unrelated group the Arbereshe in Mexican Nahuas.Zed Books, London& New Jer-
northernLucania (southernItaly, Pieroni et al. sey.
2002) and there too weeds were selected based Frohne, D., and U. Jensen. 1998. Systematik des
on several characteristicssuch as their taste and Pflanzenreichs; unter besonderer Beruecksichti-
smell properties,but in the lattercase on edibil- gung chemischer nerkmaleund pflanzlicherDro-
gen. 5. Auflage, WissenschaftlicheVerlagsgesells-
ity. Thus, we suggest that the common knowl-
chaft mbH. Stuttgart,Germany.
edge as proposed by Moerman et al. (1999) Heinrich, M. 1998. Indigenousconceptsof medicinal
should be understood as a selection based on plants in Oaxaca, Mexico: Lowland Mixe plant
common organoleptic and other culturally de- classificationbased on organolepticcharacteristics.
fined criteria(Leonti et al. 2002). The origin of Journalof Applied Botany 72:75-81.
228 ECONOMIC BOTANY [VOL. 57
47 Bixaceae 1 1 0.6251
Medic-
Rank- Total inal 48 Caprifoliaceae 1 1 0.6251
ing Plant family species spp. Residual 49 Chenopodiaceae 1 1 0.6251
1 Asteraceae 116 57 25.4026 50 Cochlospermaceae 1 1 0.6251
2 Piperaceae 46 22 9.4076 51 Cuscutaceae 1 1 0.6251
3 Fabaceae,s.l. 226 70 8.5376 52 Hamamelidaceae 1 1 0.6251
4 Euphorbiaceae 70 27 7.8916 53 Myricaceae 1 1 0.6251
5 Lamiaceae 25 12 5.1091 54 Myristicaceae 1 1 0.6251
6 Acanthaceae 23 11 4.6521 55 Proteaceae 1 1 0.6251
7 Amaranthaceae 14 8 4.0956 56 Punicaceae 1 1 0.6251
8 Araceae 29 12 4.0231 57 Turneraceae 1 1 0.6251
9 Sapindaceae 19 9 3.7381 58 Valerianaceae 1 1 0.6251
10 Commelinaceae 12 7 3.6386 59 Combretaceae 5 2 0.5391
11 Anacardiaceae 9 6 3.4531 60 Passifloraceae 9 3 0.4531
12 Aristolochiaceae 9 6 3.4531 61 Vitaceae 9 3 0.4531
13 Liliaceae, s.i. 8 5 2.7246 62 Clusiaceae 13 4 0.3671
14 Rutaceae 16 7 2.5526 63 Alstroemeriaceae 2 1 0.3536
15 Oxalidaceae 5 4 2.5391 64 Cannaceae 2 1 0.3536
16 Malvaceae 35 12 2.3941 65 Cecropiaceae 2 1 0.3536
17 Sapotaceae 13 6 2.3671 66 Magnoliaceae 2 1 0.3536
18 Fagaceae 10 5 2.1816 67 Martyniaceae 2 1 0.3536
19 Tiliaceae 14 6 2.0956 68 Musaceae 2 1 0.3536
20 Crassulaceae 3 3 2.0821 69 Olacaceae 2 1 0.3536
21 Cucurbitaceae 18 7 2.0096 70 Papaveraceae 2 1 0.3536
22 Chrysobalanaceae 4 3 1.8106 71 Plantaginaceae 2 1 0.3536
23 Begoniaceae 8 4 1.7246 72 Podostemaceae 2 1 0.3536
24 Menispermaceae 8 4 1.7246 73 Apiaceae 6 2 0.2676
25 Phytolaccaceae 5 3 1.5391 74 Loganiaceae 6 2 0.2676
26 Rhamnaceae 5 3 1.5391 75 Caricaceae 3 1 0.0821
27 Malpighiaceae 13 5 1.3671 76 Ebenaceae 3 1 0.0821
28 Simaroubaceae 6 3 1.2676 77 Elaeocarpaceae 3 1 0.0821
29 Verbenaceae 32 10 1.2086 78 Portulaccaceae 3 1 0.0821
30 Burseraceae 3 2 1.0821 79 Myrtaceae 29 8 0.0231
31 Lauraceae 29 9 1.0231 80 Celastraceae 7 2 -0.0039
32 Nyctaginaceae 7 3 0.9961 81 Violaceae 7 2 -0.0039
33 Apocynaceae 26 8 0.8376 82 Caryophyllaceae 4 1 -0.1894
34 Agavaceae 4 2 0.8106 83 Clethraceae 4 1 -0.1894
35 Polygalaceae 4 2 0.8106 84 Dilleniaceae 4 1 -0.1894
36 Rosaceae 4 2 0.8106 85 Monimiaceae 4 1 -0.1894
37 Smilacaceae 4 2 0.8106 86 Theaceae 4 1 -0.1894
38 Bombacaceae 8 3 0.7246 87 Solanaceae 63 17 -0.2079
39 Loranthaceae 8 3 0.7246 88 Polygonaceae 8 2 -0.2754
40 Marantaceae 8 3 0.7246 89 Alismataceae 1 0 -0.3749
41 Zingiberaceae 8 3 0.7246 90 Balanophoraceae 1 0 -0.3749
42 Meliaceae 12 4 0.6836 91 Betulaceae 1 0 -0.3749
43 Sterculiaceae 12 4 0.6386 92 Brunnellaceae 1 0 -0.3749
44 Urticaceae 12 4 0.6386 93 Cabombaceae 1 0 -0.3749
45 Balsaminaceae 1 1 0.6251 94 Casuarinaceae 1 0 -0.3749
46 Basellaceae 1 1 0.6251 95 Ceratophyllaceae 1 0 -0.3749
230 ECONOMICBOTANY [VOL. 57
Medic- Medic-
Rank- Total inal Rank- Total inal
ing Plantfamily species spp. Residual ing Plantfamily species spp. Residual