Medicinal Flora of the Popoluca, Mexico: A Botanical Systematical Perspective

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

MARCO LEONTI, FERNANDO RAMIREZ R., OTTO STICHER, AND

MICHAEL HEINRICH1

Leonti, Marco (Departmentof Applied BioSciences, Institute of PharmaceuticalSciences,

Swiss Federal Instituteof Technology(ETH) Zurich, Winterthurerstr. 190, CH-8057 Zurich,
Switzerland),Fernando Ramirez R. (Proyecto Sierra de Santa Marta,A.C. Cuauhte6moc 10,
Centro Hist6rico, Xalapa 91000, Veracruz,Mexico), Otto Sticher (Departmentof Applied
BioSciences,Instituteof PharmaceuticalSciences, Swiss Federal Instituteof Technology(ETH)
Zurich, Winterthurerstr. 190, CH-8057 Zurich, Switzerland),and Michael Heinrich (Centre
for Pharmacognosyand Phytotherapy,The School of Pharmacy,Universityof London,29-39
BrunswickSq. London,WCINlAX, UK; e-mail:phyto@ulsop.ac.uk).MEDICINAL FLORA OFTHE
POPOLUCA,MEXICO: A BOTANICAL SYSTEMATICALPERSPECTIVE.Economic Botany 57(2):218-
230, 2003. Westudiedthe medicinalplants used by the Popoluca of the Sierrade SantaMarta
(eastern Mexico). Using Moerman'smethodof regressionanalysis we determinedwhich eth-
nomedicallyused taxa are over-representedin the Popolucanpharmacopoeia(e.g., Asteraceae)
and which are underrepresented(e.g., Orchidaceae).Moermanet al. (1999) found high cor-
relationbetweenthe holarcticpharmacopoeiasand assumedthat apartfrom the relatednessof
the northernfloras a "global pattern of human knowledge" may accountfor this finding.
Althoughthe Popoluca dwell in a habitatdominatedby a neotropicalflora but intermixedwith
importantholarctic elements, they include considerablyfewer neotropicaltaxa in theirphar-
macopoeia as one would expect if the historical transmittedknowledgewere influencingtheir
selection. Thisfinding confirmsthe theory stated by Moermanet al. However, the Popoluca
include some neotropicaltaxa in theirpharmacopoeiaand thus a moderatecorrelationexists
betweenthe Popolucanpharmacopoeiaand the neotropicalpharmacopoeiaanalysedby Moer-
man et al. We therefore conclude that apart from historically transmittedknowledgeabout
specific taxa the "globalpatternof humanknowledge"addressedby Moermanet al. is largely
based on "commonselection criteria."

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."

Wdhrendeiner ethnobotanischenFeldstudieuntersuchtenwir die Medizinalpflanzen der Po-

poluca in der Sierra de Santa Marta (ostlichesMexiko).WirwandtenMoermansMethodeder
Regressionsanalysean, um die ethnomedizinischiiberreprdsentierten (z. B. Asteraceae) und
unterreprasentierten Taxa (z. B. Orchidaceae)dieser Ethnopharmakopoe zu bestimmen.Moer-
man et al., welche eine hohe Korrelationzwischen holarktischenPharmakopoenfanden, ver-
muten,dass neben der Verwandschaftder nordlichenFlora ein "globalesMustermenschlichen
Wissens"far dieses Ergebnisverantwortlichist. Die Popoluca,welche in einemHabitatsiedeln,

1
Received 10 October2002; accepted 15 March2003.

EconomicBotany 57(2) pp. 218-230. 2003

? 2003 by The New York BotanicalGardenPress, Bronx, NY 10458-5126 U.S.A.
2003] LEONTIET AL.: MEDICINALFLORAOF THE POPOLUCA,MEXICO 219

in dem die neotropischeFlora dominiert,aber klare holarktischeEinfliusseaufweist,schliessen

bedeutendweniger neotropischeElemente in ihre Pharmakopoemit ein, als man erwarten
wurde, wenn nicht historisch tradiertesWissen ihre Selektionbeeinflussenwiirde.Dieses Er-
gebnis bestatigtdie Theorievon Moermanet al. Da jedoch in der Pharrmakopole der Popoluca
ein bestimmterTeil der neotropischenFlora vertretenist, besteht eine moderateKorrelation
zu der von Moermanet al. analysiertenneotropischenPharmakopbe.Daher schlussfolgernwir,
dass das von Moermanet al. vermutete"globale MustermenschlichenWissens"nebst histo-
risch tradiertemphytomedizinischemWissen uiberspezifische Taxa vor allem auch auf "ge-
meinsamenSelektionskriterien" beruht.
Key Words: traditionalmedicine; ethnobotany;ethnopharmacy;Isthmus of Tehuantepec;
Macro-Mayan;medicinalplant selection;Mexico; Popoluca;regressionanalysis.

Researchon medicinaland otheruseful plants
used in indigenous societies has been driven by
two complementaryinterests:The use of such
informationfor researchin the field of the nat-
ural sciences, especially with regard to 'new'
bioactive naturalproducts derived from plants
and the use of plant extracts in primaryhealth
care (Heinrichand Gibbons2001) and the inter-
est in better understandingthe anthropological
basis, if possible on a cross-culturalbasis, of the
use of these resources by humans and particu-
larly on the rationale(s)behind the selection of
these resources(Moermanet al. 1999).
One particularlyexciting developmentwhich
contributesto both lines of investigationis the
studyof botanicalsystematicaspectsof plantus-
age. Dan Moerman and his colleagues (e.g.,
1996, 1998a,b) have developed and used (par-
tially in collaborationwith several other schol-
ars) a methodwhich allows for a statisticalanal-
ysis of ethnobotanicalinformationbased on that
the numberof medically used plant taxa and the
total number of taxa in a certain region is
known.
Most of their researchfocused on the North
temperateregion of North America, but also on
three additionalnorthern(Kashmir,Korea,Chia-
pas) and one southern (Ecuador) hemispheric
ethnopharmacopoeias. Only one neotropical
lowland region (upper Napo River valley, Ec-
uador)is includedin his analyses.The main rea-
son for this is a lack of taxonomic and system-
atic researchin such regions. The results of the
comparisonby Moermanet al. (1999) show that
holarctic peoples rely on similar plant families
in their health care. The Pearsoncorrelationfac-
tors between the holarctic data sets are higher
than 0.6 while the correlation of the Ecuador
data set with the holarcticdata set is lower than
0.19 (maximumpossible correlationis one, low-
est is zero). Moermanet al. (1999) suggest that
the relatively high correlationbetween the ho-
larctic ethnopharmacopoeiasand the low corre-
lation between the holarcticpharmacopoeiasand
the neotropicalone is due to the relatednessof
the northernfloras and the possibility that the
knowledge about medicinal plants has been
passed on from prehistorictimes throughspace
and time. This implies that the peoples entering
the New Worldthroughthe BeringianStreet al-
ready shared a common knowledge about me-
dicinalplantswith the peoples migratingto other
areas and remainingon the Asiatic continent.
There is a widespreadacceptancethat North
America was first settled at least around15 000
yr B.P., but some authorspostulatea date as early
as 40,000 yr B.P. (Layrisse and Wilbert 1999).
According to these authors,three major waves
of immigrantspopulatedthe Americas.The first
wave was of Pre-Mongoloid tribes at around
40 000 yr B.P., followed by Paleo-Mongoloidsat
30 000 yr B.P., and the Diego-allele positive
Neo-Mongoloids at about9000 yr B.P. (Layrisse
and Wilbert 1999). When peoples crossed the
borderof the holarcticflorainto the realmof the
neotropicalplant kingdom they were suddenly
confrontedwith plant families that were new to
them and had to adapt themselves and their
pharmacopoeiato this new pool of plants.
We have recently completed a detailed study
of the medicinal plant use of the Popoluca in
southernVeracruz,Mexico (Leonti et al. 2001).
The Popolucainhabitthe Sierrade SantaMarta,
a range of volcanoes between the Lake of Ca-
temaco and the Gulf coast. These volcanoes
form the southernfoothills of the "Sierrade Los
Tuxtlas" mountainrange, a region particularly
well known for its biodiversity, where the ho-
larctic and neotropicalfloristic kingdoms over-
lap. The neotropicalinfluence is strongerthan
the holarctic one. According to Rzedowski
(1991) the overwhelming part of the Mexican
220 ECONOMICBOTANY [VOL. 57

flora belongs to the neotropicalplant kingdom, Postgraduadosde ChapingoCHAPA (Texcoco),

even though the pine and oak forests of the IMSS-M (InstitutoMexicano del Seguro Social,
Mexican highlandsand Chiapassharethe north- Mexico, D.E), Institutode Ecologia XAL (Xa-
ern and southernfloralinfluencesin aboutequal lapa), the Centre for Pharmacognosyand Phy-
parts. Importantvegetation zones in the Sierra totherapy,The School of Pharmacy,Univ. Lon-
de Santa Marta include the tropical montane don, and the ETH Zurich (CH). Identification
cloud forest, the tropical rain forest, and the was largely conducted at MEXU and the Cole-
semi-dry oak forest. gio de Postgraduadosen Ciencias Agricolas,
The study area of about 1350 km2 lies at al- Montecillo, Mexico, but with the help of spe-
titudes between sea level and 1720 m. Ramirez- cialists from these institutions.In this contribu-
Ramfrez (1999) published a comprehensive tion, voucher specimens are only cited if they
checklist of the flora of the region with records have not been reportedpreviously in our other
for 2400 species. Although,it was estimatedthat publications(Leonti et al. 2001, 2002).
about 3000 species of flowering plants grow in
the Sierra(Chevalierand Buckles 1995:182),we EVALUATION METHODS
consider an evaluationas feasible based on Ra- In order to determine the importance of the
mirez-Ramirez'swork. This enables an analysis medicinal plant families, we used Moerman's
of the use of plantsfor medical purposesby this (1991) method applying regression analysis
indigenous group, which is particularlyinterest- (Fig. 1). The families were then ranked ac-
ing because the unique compositionof the flora cording to their decreasing residuals (Appen-
including elements from two kingdoms. dix I), which reflects the proportion of plants
Here we reporton a regressionanalysis of the used as medicinals in a certain family. The re-
medicinal plants. In order to better understand sidual is the difference between the numberof
the culturalrationalfor the rankingof the most medicinal species predicted by the regression
salient plant families we analyse for which cat- analysis and the true, ethnographically deter-
egories of illness the respective plant families mined numberof medicinal species (Moerman
are used for, elucidatethe phytochemicalspectra et al. 1999).
of these families, and highlight ecological and Since Moermanand his colleagues used only
organolepticalcharacteristics. angiospermfamilies in his analysis, we decided
to do so as well, in order to secure the compa-
METHODS rability of the set of data. For the same reason
some plant families had to be combined: Ant-
ETHNOBOTANICAL RESEARCH
hericaceae and Amaryllidaceae into Liliaceae
The ethnobotanicalresearch was undertaken s.l.; Phyllonomaceae into Grossulariaceae;
in the municipalities(municipios) of Hueyapan Hypericaceaeinto Clusiaceae;Mimosaceae,Fa-
de Ocampo and Soteapan, southern Veracruz, baceae s.str. and Caesalpiniaceaeinto Fabaceae
from March 1999 to July 2000. Fieldwork fo- s.l.
cused on collecting information on medicinal To determinethe relationshipbetween the dif-
plant use and general ethnographicdata. Dried ferent floras the Pearson correlationcoefficient
herbariumspecimens and samples for further of the numberof species per family was calcu-
phytochemicalanalysis (for details see Leonti et lated. Similarly,the relationshipof the different
al. 2001; Leonti, Sticher,and Heinrich2002; cf. medicinal floras was calculated by the Pearson
Ankli, Sticher, and Heinrich 1999) were also correlation coefficient of the residuals (Moer-
collected. man et al. 1999).
The researchwas performedwith permitNo.
DOO. 02.-1750, obtainedfrom the InstitutoNa- RESULTS AND DISCUSSION
cional de Ecologia, of the Secretariade Medio The medicinal flora of the Popoluca consists
AmbienteRecursosNaturalesy Pesca (SEMAR- of 614 plant species, contributedby 72 infor-
NAP), Mexico. Complete sets of voucher spec- mants, with a total of 4488 use-reports(Leonti
imens (Leonti 1-599) are deposited at the Na- et al. 2001). The plantsbelong to 117 of the 174
tional Mexican HerbariumMEXU (Universidad Angiosperm families recorded for the region
Nacional Autonoma de Mexico, Mexico, D.E), (Ramirez-Ramirez1999). According to the re-
the Herbarium-Hortoriumof the Colegio de siduals of the regressionanalysis (y = 0.2716x
2003] LEONTIET AL.: MEDICINALFLORAOF THE POPOLUCA,MEXICO 221

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.

+ 0.1042) of the Popolucanethnomedicalflora 1) Ecological factors, including the importance

the top five families used by the Popoluca are
(Table 1): Asteraceae,Piperaceae,Fabaceae,s.l.,
Euphorbiaceae,and Lamiaceae. The five fami-
lies (positions 170-174, Table2) with the lowest
level of usage in the regressionanalysis are Or-
chidaceae,Poaceae, Rubiaceae,Cyperaceae,and
Moraceae.
It is of considerableethnobotanicalinterestto
understandthe culturalreasons for a taxon's us-
age or its avoidance. Reasons for a taxon's eth-
nomedicalimportanceinclude, for example:
of some of the taxa as managedhouse garden
plants. If a high percentage of medicinal
plants from a certain plant taxon (e.g., La-
miaceae) is cultivated in home gardens,
many species may well be of particulareth-
nomedical importance,and have furthereth-
nobotanicaluses (culinary,ornamental,or for
construction). Very often such species are
hard to find in the naturalhabitator are in-
troduced into the region. It is importantto
note that if the calculation of the regression

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'

Orchidaceae 174 143 100 245 134 118

Poaceae 173 144 100 255 135 51
Rubiaceae 172 82 9 250 65 109
Cyperaceae 171 142 94 254 136 102
Moraceae 170 113 8 33 21 117
Totalfamilies 174 144 100 255 136 118
' DatafromMoermanet al. 1999.

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

portantin the illness categories of gynecology,

fever and headache,urological conditions,bites
from venomous animals, and are the second
CO~~~~~~~C most importantfamily to treat gastrointestinal
and dermatologicalcomplaints.Membersof the
Fabaceae, s.l. are often rich in polyphenoles
CZO
CZ (tanninsand flavonoids)and triterpenesaponins
(Frohneand Jensen 1998). Especially the Faba-
ceae s.str. contain many toxic genera and spe-
cies such as Canavalia (lectines), Ormosia (chi-
nolizidine alkaloids), Erythrina spp. (isochino-
Z C ~ ~ a) 0 a)
line alkaloids), Crotalaria sagittalis L. (pyrroli-
CO zidine alkaloids).The seeds of Mucuna pruriens
(L.) DC., containingL-DOPAand hallucinogen-
0 COJ
ic tryptamines,are ground,toasted and used as
a coffee surrogatewhich is typical of ruralre-
gions in Guatemala and Mexico (Buckles,
Triomphe,and Sain 1998).
0 _
3
Of the 70 medicinally used Fabaceae s.str.
0~~~~~~~3 c
twelve (17%) are cultivated regularlyin home
0 a)~~~~uc gardens,ten of these are trees, having ornamen-
0 a)~~~~~ta)
O0 tc tal uses (Caesalpinia pulcherrima (L.) Sw.,
Leonti 118, Senna multijuga ssp. doylei (Britton
& Rose) H. S. Irwin & Barneby,Leonti 70) or
CZ
nutritionaluses (Tamarindus indica L., Dialium
guianense (Aubl.) Sandwith,Leonti 298). Cro-
talaria longirostrata Hook & Ar. (Leonti 479)
U * is cultivatedand eaten as a vegetablebut no sop-
a)a)~~~(U.0
0 CO O a)0~C
orific (sleep-inducing) effect, as reported by
Cl
U
Morton (1994) was mentionedby the Popoluca
informants.The frequent use of Fabaceae, s.l.
-cH CO
Z ~ CO et
for medicinal purposes is probably due to the
z
conspicuous characterof many of its species,
0 c their abundance,the diversity of highly active
U a) chemical constituentsand notableorganoleptical
Q) properties (see Leonti, Sticher, and Heinrich
2002).

Euphorbiaceae. The Euphorbiaceaeare pre-

dominatelyused in the illness categoriesof gas-
trointestinal disorders, dermatological com-
plaints, and urological ailments. Polyphenoles
(flavonoidsand condensed tannins),rubber,dif-
ferent types of alkaloids, cyanogenic glycosides
H ZE cfcc~~~~~~c1 7: -- and diterpenesare the main classes of secondary
naturalproducts of the Euphorbiaceae(Frohne
and Jensen 1998). The genus Croton, known to
0 00
contain toxic diterpene esters, is represented
. 0 ~co O 6
with five species in the Popolucan pharmaco-
~~0
U
~ 4- -~-~
( - 0 poeia.
Of the 26 medicinal species of Euphorbia-
CZ0 4
0:) u V. (A ceae, two are cultivatedfrequently,Cnidoscolus
chayamansa McVaugh (Leonti 182) being pri-
224 ECONOMICBOTANY [VOL. 57

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.

Veracruz Chiapas Kashmir NorthAmerica Korea Ecuador

Pearson correl. floras 0.94 0.74 0.68 0.51 0.83

Pearson correl. med. used families 0.81 0.62 0.57 0.54 0.36

main chemical constituents(Frohne and Jensen lack conspicuouscharacteristicswhich allow an

1998). Only two out of 35 Cyperaceaespecies easy distinctionamong variousspecies. Withthe
of this region have medicinal uses but are with- Piperaceae and the Euphorbiaceae,on the one
out importancein the pharmacopoeiaof the Po- hand, and the Rubiaceae and the Moraceae,on
poluca. Again, the lack of usage of this taxon the other hand, the differencesare not as prom-
seems to be due to the inconspicuousnatureof inent but point in the same direction.
the species. Also, Cyperaceaegenerallygrow in
humid and inaccessible habitats. INTERCULTURAL
COMPARISONWITHOTHER
REGRESSIONANALYSES
Moraceae. Very characteristicis the latex which
often contains phototoxic furanocumarinesand The Asteraceae and the Lamiaceae are gen-
toxic cardenolides (Frohne and Jensen 1998). erally ranked highly in holarctic ethnopharma-
The Moraceae have no importancein the Po- copoeias (Moermanet al. 1999). This points to
poluca's healthcare. Out of the 31 species, three the holarctic characterof the Popolucan phar-
are used medicinally. Ficus pertusa L.f. (Leonti macopoeia,while the Piperaceaereflectthe neo-
63) is sometimes grown in house yards as a me- tropicalinfluencein the pharmacopoeia.The Fa-
dicinal plant and as a fruit tree. The neglecting baceae's (s.l.) high ranking with the Popoluca,
of the Moraceae may in part be explained by is difficultto analyse (Table 1). They are ranked
their life form as climbers (Ficus sp.) which very low in North American (253rd of 255),
does not allow one to distinguish the species Chiapas (138th of 144), and Kashmir(85th of
even on the basis of theirleaves. Also manytaxa 100), but are the second most used family in
are abundantin more or less undisturbedforest Ecuador,and are ranked8th in Korea.Probably
habitats. the Popoluca's reliance on the Fabaceae, s.l. re-
flects neotropical influence. The Euphorbiaceae
INTRACULTURAL ETHNOPHARMACOLOGICALare highly ranked in Kashmir (2nd of 100), Ko-
COMPARISON rea (13th of 136) and Chiapas (21st of 144), but
Chemicalcharacteristicsseem to be an essen- surprisingly under-represented in North America
tial criterionfor plant selection. These are per- (234th of 255) and low ranked in the Ecuador
ceived by the Popoluca on the basis of the (90th of 118) set of data.
plant's taste and smell propertiesand theirphar- Generally, the Poaceae, Cyperaceae and Or-
macological effects. They are essential for se- chidaceae are very rarely selected (Table 2). The
lection and continueduse of certaintaxa as med- Rubiaceae are ranked 10th in Kashmir but are
icine. The high diversity of secondary com- relatively low rankedin the other sets of data.
pounds, taste and smell properties,but also the The Moraceaeshow a positive residualin North
distinctive plant morphology clearly separates America, Kashmirand Korea, while in Mesoa-
the Asteraceae,the Lamiaceaeand the Fabaceae, merica and South America the Moraceae are un-
s.l. organolepticallyfrom the lesser used fami- derrepresented in the medicinal floras (Table 2).
lies such as the Poaceae and the Cyperaceae.
Families with many weedy taxa or ones often FACTOR
PEARSONCORRELATION
cultivated in the home gardens are particularly Not surprisingly the ethnopharmacopoeia (e)
prominent in the pharmacopoeia. The lesser and the flora (f) of the Popoluca (southernVe-
used families, best exemplified by the Orchida- racruz) show the highest correlation (0.81, e and
ceae, exemplify problems in gatheringmaterial 0.94, f) with the one of the Tzotzil/Tzeltal from
of a plant that makes them unlikely candidates highland Chiapas (Table 4). Both ethnic groups
for medicinal plants. Both, the Poaceae and live in an area which harbors plant species from
Cyperaceae seem to be neglected because they the holarctic as well as from the neotropical
226 ECONOMICBOTANY
plant kingdoms, although the neotropicalinflu-
ence is higherin the floraof Veracruz.The most
prominentdifference is that the Popoluca dwell
in the lowland and the Tzotzil/Tzeltal in the
highlands. Both groups are part of the Macro-
Mayanlanguagestock and are culturallyrelated.
The culturalseparationmay have occurredabout
2500 years ago. The high correlationof their
pharmacopoeiasprobablyreflects both the bio-
logical relatednessof the area and the common
culturalpast of the two groups. If we compare
the relatednessof the flora of the Populucaand
the medicinal flora selected by them with other
floras/medicalfloras (Table4) an analysis using
the Pearsoncorrelationindicates the relatedness
of the Popolucan (Veracruzian)ethnopharma-
copoeia with the others. The only exception is
the data for Ecuador. The flora of Veracruz
shows a correlationof 0.83 with Ecuadorwhich
would predict a correlationof their pharmaco-
poeias of about 0.75 if the relatednessof their
flora would be the only influencingfactor.How-
ever, the correlation between the Veracruzian
and the Ecuadorianpharmacopoeiafalls at the
much lower level of 0.36.
The Veracruz(V) and the Chiapas (C) phar-
macopoeia perform similarly to the other phar-
macopoeias. Both show the highest correlations
with Kashmir (0.74 C/0.62 V), second highest
with NorthAmerica (0.73 C/0.57 V), thirdhigh-
est with Korea (0.61 C/0.54 V), and the lowest
with Ecuador (0.19 C/0.36 V). The correlation
of the Chiapasdata set to the other holarcticset
of data is always higher as comparedto Vera-
cruz. On the otherhand the Veracruzcorrelation
to the neotropical(Ecuador)set of data is con-
siderablyhigher than the one from Chiapas.
Thus, the Pearson correlation factors show
that the Veracruz flora is most related to the
nearby Chiapas flora but is more related to the
Ecuador flora than to any true holarctic flora.
Nevertheless, the Pearson correlationfactors of
the differentpharmacopoeiasshow that the Po-
poluca pharmacopoeiais generally more related
to the holarctic pharmacopoeiasbut shows a
clear neotropicalinfluence.
DICOTS COMPAREDTO MONOCOTS
The classical, but now controversial,separa-
tion of the Angiosperms into Monocots (MC)
and Dicots (DC) allows a useful analysis of still
anotheraspect. Overall, the MC seem to be less
importantas compared to the DC. Again this
[VOL. 57
may well be partlydue to the inconspicuousness
of many of its plants. The most prominentfam-
ilies of the MC are the Araceae (rank 8), the
Commelinaceae(rank10), and also the Liliaceae
(rank 13); the lattertwo are families with many
very showy species. All three families show
clear differences in their ethnopharmaceutical
uses. We documented 18 of 62 medicinal MC
species used for urological problems, with 84
(23.5%) of 357 total MC use reports,while 136
of 535 medicinalDC species have a total of 253
(6.2%) reportsin this group (DC-total number
of use reports:4029). The MC are thus more
likely to be selected for urologicalproblems.
Out of the 62 medicinalMC species, only two
(3.2%) with a total of five (1.4%)use reportsare
used to treatrespiratoryailments,while 73 spe-
cies of the 535 medicinal DC (13.6%) with 198
(5%) use reportswere recorded.
Why should a MC be more suitable to treat
urologic conditions and less useful to treat re-
spiratory ailments? The empirical medico-his-
toricalexperienceof an ethnic groupis of course
an essential basis for this, but at this stage the
specific reasonsremainspeculative.Phytochem-
ical as well as organolepticaldifferences exist.
Concerning the phytochemistry,the MC lack
ellagic acid and ellagic tannins,whereastannins
and essential oils, polyterpenesand alkaloidsare
not common, in general. Steroid saponinsoccur
in MC while the DCs have mostly triterpene-
saponinsand the mucilage of the MC lacks uro-
nic acid (Frohne and Jensen 1998). Regarding
the morphology,the MC distinguishthemselves
by not showing secondary growth, having a
moderatediameter,an un-branchedaxis, and a
watery stem. However,we arguethat the differ-
ent spectraof secondarynaturalproductsalong
with the recorded pharmacologicalproperties
are not sufficientto explain the unequaluseage
of these two groups. Such products are indeed
responsiblefor specific organolepticalproperties
of a botanical drug, a set of criteriawhich we
have previouslyshown to be an importantselec-
tion criterionfor medicinal plants with the Po-
poluca (Leonti et al. 2002) and other groups
(Heinrich1998; Brett and Heinrich1998). In the
classificatorysystem of the Popoluca and other
peoples all these factors-the organoleptical
perception of a plant, the morphological and
ecological traits,and the characteristicsin terms
of colour,as well as its smell and taste, are used
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
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smell properties,but in the lattercase on edibil- gung chemischer nerkmaleund pflanzlicherDro-
gen. 5. Auflage, WissenschaftlicheVerlagsgesells-
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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
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Larysse, M., and J. Wilbert. 1999. The Diego blood
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Leonti, M., H. Vibrans, 0. Sticher, and M. Hein-
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of Ethnopharmacology81:307-315.
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Ramirez-Ramirez, F. 1999. Flora y vegetaci6nde la
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2003] LEONTI ET AL.: MEDICINAL FLORA OF THE POPOLUCA, MEXICO 229

APPENDIX: RANKED LIST OF RESIDUALS PER FAM- APPENDIX: CONTINUED.

ILY INCLUDING THE TOTAL NUMBER OF SPECIES OF
Medic-
THE FLORA OF THE SIERRA DE SANTA MARTA AND Rank- Total inal
THE MEDICINAL PLANTS DOCUMENTED. ing Plantfamily species spp. Residual

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

APPENDIX: CONTINUED. APPENDIX: CONTINUED.

Medic- Medic-
Rank- Total inal Rank- Total inal
ing Plantfamily species spp. Residual ing Plantfamily species spp. Residual

96 Chloranthaceae 1 0 -0.3749 145 Pontederiaceae 3 0 -0.9179

97 Cunoniaceae 1 0 -0.3749 146 Thymelacaceae 3 0 -0.9179
98 Haemodoraceae 1 0 -0.3749 147 Campanulaceae 4 0 -1.1894
99 Hemandiaceae 1 0 -0.3749 148 Hippocrateaceae 4 0 -1.1894
100 Hypoxidaceae 1 0 -0.3749 149 Icacinaceae 4 0 -1.1894
101 Juncaceae 1 0 -0.3749 150 Marcgraviaceae 4 0 - 1.1894
102 Lacistemataceae 1 0 -0.3749 151 Potamogetonaceae 4 0 -1.1894
103 Lentibulariaceae 1 0 -0.3749 152 Scrophulariaceae 19 4 -1.2619
104 Molluginaceae 1 0 -0.3749 153 Dioscoreaceae 8 1 -1.2754
105 Najadaceae 1 0 -0.3749 154 Ericaceae 8 1 -1.2754
106 Nymphaeaceae 1 0 -0.3749 155 Heliconiaceae 8 1 -1.2754
107 Grossulariaceae 1 0 -0.3749 156 Iridaceae 8 1 -1.2754
108 Plumbaginaceae 1 0 -0.3749 157 Actinidiaceae 5 0 -1.4609
109 Polemoniaceae 1 0 -0.3749 158 Annonaceae 20 4 -1.5334
110 Primulaceae 1 0 -0.3749 159 Melastomataceae 46 11 -1.5924
111 Pyrolaceae 1 0 -0.3749 160 Flacourtiaceae 17 3 -1.7189
112 Rafflesiaceae 1 0 -0.3749 161 Capparaceae 10 1 -1.8184
113 Rhizophoraceae 1 0 -0.3749 162 Myrsinaceae 18 3 -1.9904
114 Ruppiaceae 1 0 -0.3749 163 Arecaceae 22 4 -2.0764
115 Saxifragaceae 1 0 -0.3749 164 Araliaceae 11 1 -2.0899
116 Typhaceae 1 0 -0.3749 165 Asclepiadaceae 19 3 -2.2619
117 Tovariaceae 1 0 -0.3749 166 Gesneriaceae 16 2 -2.4474
118 Vochysiaceae 1 0 -0.3749 167 Boraginaceae 17 2 -2.7189
119 Winteraceae 1 0 -0.3749 168 Bromeliaceae 38 6 -4.4204
120 Zygophyllaceae 1 0 -0.3749 169 Convolvulaceae 28 3 -4.7054
121 Brassicaceae 5 1 -0.4609 170 Moraceae 31 3 -5.5199
122 Lythraceae 5 1 -0.4609 171 Cyperaceae 35 2 -7.6059
123 Onagraceae 5 1 -0.4609 172 Rubiaceae 126 24 -10.3124
124 Ranunculaceae 5 1 -0.4609 173 Poaceae 93 5 -20.3529
125 Ulmaceae 5 1 -0.4609 174 Orchidaceae 99 2 -24.9819
126 Cactaceae 13 3 -0.6329
127 Burmanniaceae 2 0 -0.6464
128 Connaraceae 2 0 -0.6464
129 Cyclanthaceae 2 0 -0.6464
130 Dichapetalaceae 2 0 -0.6464
131 Gunneraceae 2 0 -0.6464
132 Hydrophyllaceae 2 0 -0.6464
133 Ochnaceae 2 0 -0.6464
134 Oleaceae 2 0 -0.6464
135 Sabiaceae 2 0 -0.6464
136 Staphyleaceae 2 0 -0.6464
137 Styracaceae 2 0 -0.6464
138 Theophrastaceae 2 0 -0.6464
139 Bignoniaceae 32 8 -0.7914
140 Aquifoliaceae 3 0 -0.9179
141 Convallariaceae 3 0 -0.9179
142 Erythroxylaceae 3 0 -0.9179
143 Gentianaceae 3 0 -0.9179
144 Juglandaceae 3 0 -0.9179