Successional Pathways Derived from Different Vegetation Use Patterns by Lacandon Mayan Indians

Abstract

Eighty-four censuses were made at 38 “acahuals” (slash and burn old fields) in the village of Lacanhá Chansayab, Chiapas, México. The time since last cultivation of each field (the fallow period) ranged from one to twenty years. Fields were derived from two physiognomic variants of mature tropical rainforest (“monte alto” and “chaparral”). The data was subjected to ordination (DECORANA) and classification (TWINSPAN) and compared to the information on their history of usage. Two disturbance indices were also generated. The analysis allowed the description of successional pathways associated with human disturbance patterns. Acahuals and species were ordered along the first axis according to the current fallow period and disturbance intensity; the second axis corresponded to a gradient between the two original vegetation types from which acahuals derived, and the third axis corresponded to the gradient of disturbance frequency to which acahuals have been subjected. Acahual classification was based on the following groups: (1) acahuals with frequent use and short fallow, subjected to an intensive agricultural use; (2) acahuals with a prolonged use, with no previous fallow periods and derived from monte alto forests; (3) acahuals derived from chaparral forests; and (4) old acahuals (14 years to 20 years) with no previous fallow. Species classification resulted in: (1) non-persistent species; (2) disturbance-susceptible species; (3) acahual-typical species; (4) persisting under frequent and prolonged field-use species; and (5) two groups of species typical of old acahuals.

Full text

Successional Pathways Derived
from Different Vegetation Use Patterns
by Lacandon Mayan Indians
Samuel I. Levy-Tacher
J. Rogelio Aguirre Rivera
ABSTRACT. Eighty-four censuses were made at 38 “acahuals” (slash
and burn old fields) in the village of Lacanhá Chansayab, Chiapas,
México. The time since last cultivation of each field (the fallow period)
ranged from one to twenty years. Fields were derived from two
physiognomic variants of mature tropical rainforest (“monte alto” and
“chaparral”). The data was subjected to ordination (DECORANA) and
classification (TWINSPAN) and compared to the information on their
history of usage. Two disturbance indices were also generated. The anal-
Samuel I. Levy-Tacher is affiliated with El Colegio De La Frontera Sur, Depart-
amento de Ecología y Sistemática Terrestre, Carretera Panamericana y Periférico Sur
s/n, Barrio de María Auxiliadora, San Cristóbal de Las Casas, Chiapas, México C.P.
29290.
J. Rogelio Aguirre Rivera is Doctor Ingeniero Agrónomo and Profesor Invest
-
igador, del Instituto de Investigación de Zonas Desérticas, Universidad Autónoma de
San Luis Potosí, Altair 200 Fracc. del Llano, San Luis Potosí, SLP, México C.P. 78377
(E-mail: iizd@uaslp.mx).
Address correspondence to: Samuel I. Levy-Tacher (E-mail: slevy@sclc.ecosur.mx).
The authors would like to thank peasants of Lacanhá Chansayab: Manuel Castellanos
Chankin, Kin Bor, Carlos Paniagua, Arturo Paniagua y Carmelo ChanKayum, for in
-
terpreting and sharing his knowledge and experience. Thanks go to Neptalí Ramirez-
Marcial and Duncan Golicher for helpful reviews of earlier drafts.
Financial support for this study was provided by Simón Bross and Juan García
(GBA), Etnobiología para la Conservación A. C., the Comisión Nacional para el
Conocimiento y Uso de la Biodiversidad (CONABIO), Consejo Nacional de Ciencia y
Tecnología (CONACYT), Instituto Nacional de Ecología (INE), Secretaría del Medio
Ambiente y Recursos Naturales (SEMARNAT) and the Reserva de la Biosfera Montes
Azules (REBIMA).
Journal of Sustainable Agriculture, Vol. 26(1) 2005
Available online at http://www.haworthpress.com/web/JSA
 2005 by The Haworth Press, Inc. All rights reserved.
Digital Object Identifier: 10.1300/J064v26n01_06 49

ysis allowed the description of successional pathways associated with
human disturbance patterns. Acahuals and species were ordered along
the first axis according to the current fallow period and disturbance in
-
tensity; the second axis corresponded to a gradient between the two orig
-
inal vegetation types from which acahuals derived, and the third axis
corresponded to the gradient of disturbance frequency to which acahuals
have been subjected. Acahual classification was based on the following
groups: (1) acahuals with frequent use and short fallow, subjected to an
intensive agricultural use; (2) acahuals with a prolonged use, with no
previous fallow periods and derived from monte alto forests; (3) aca
-
huals derived from chaparral forests; and (4) old acahuals (14 years to
20 years) with no previous fallow. Species classification resulted in:
(1) non-persistent species; (2) disturbance-susceptible species; (3)
acahual-typical species; (4) persisting under frequent and prolonged
field-use species; and (5) two groups of species typical of old acahuals.
[Article copies available for a fee from The Haworth Document Delivery Service:
1-800-HAWORTH. E-mail address: <docdelivery@haworthpress.com> Website:
<http://www.HaworthPress.com>  2005 by The Haworth Press, Inc. All rights
reserved.]
KEYWORDS. Rain forest old fields, ordination, classification, suc-
cessional pathways, human disturbance patterns, disturbance intensity
and frequency
INTRODUCTION
The effects of human influence on the development of plant commu-
nities are often more important than natural disturbances. The suc-
cessional processes currently described are the result of such influence
(White and Pickett 1985, Shimidt 1988, Bazzaz 1996).
Slash and burn (S&B) agriculture alters rain forest succession in or
-
der to optimize economic output of the system (Watters 1971, Anony
-
mous 1980). Crop production and cattle raising in tropical areas have
drastically effected the tropical rainforest recovery process, facilitating
the proliferation of “acahuals.” Acahual is the local name used for S&B
old fields originally vegetated with mature tropical forest, in which sec
-
ondary vegetation corresponding to an intermediate successional state
has developed. These areas include colonizing species adapted to and
favored by an intensive use (Denslow 1985, Miranda 1993a, van der
Wal 1998). The reduction of the fallow periods plus frequent weeding
and burning act as selection factors tending to simplify the floristic
composition of affected areas. At the same time the complexity of vege
-
tation recovery patterns may increase over time (Connell and Slatyer
50 JOURNAL OF SUSTAINABLE AGRICULTURE

1977, Uhl et al. 1982, Purata 1986a). The replacement of temperate and
tropical rainforests by bushy savanna that persists with minimal human
intervention is one evidence of a successional stage derived from inten
-
sive use patterns (Egler 1954, Budowsky 1956).
Due to the poor information about the land’s use-history only a few
studies have evaluated the intensity or magnitude of disturbance in plant
communities (Glenn-Lewin and van der Maarel 1993). Holling (1973)
has pointed out that disturbance intensity and severity should be consid
-
ered. Denslow (1985), Kellman (1974) and Quintana et al. (1996) have
shown that the increase in disturbance frequency and intensity caused
by agricultural use leads to invasion by exotic weeds better adapted to
the new situation than local species. Local extinction of native species
can occur if this process leads to high mortality rates.
S&B agriculture has been used by the indigenous communities in the
Lacandon Tropical Rainforest of southeastern Mexico since pre-His-
panic times (Hellmuth 1977 and Marion 1991). The Lacandon Maya
constitute an ethnic group with a deep traditional knowledge of the local
flora and vegetation, together with their utilization and management
patterns (Nations and Nigh 1980, Marion 1991). Since the 70s the
Lacandon Indian communities have undergone rapid socio-economical
change as a consequence of cultural and economic globalization (Rzedowski
1978, Vázquez and Ramos 1992). These changes have caused the
progressive deterioration of the Lacandonian agriculture tradition. As
acahual use has been intensified, the variety of crops associated with
maize has declined. Reduction in maize yield has been reported, as a re-
sult of prolonged soil use along with annual fires and the use of ma-
chetes for weeding.
Given the core role that secondary and mature vegetation plays
within the Lacandonian agricultural system, as well as, the need to gen
-
erate alternatives for the sustainable use and rehabilitation of soils and
forests, it is important to define the plant associations resulting from the
various human use or disturbance patterns. Present research has aimed
to identify the multiple successional routes caused by the different agri
-
cultural use patterns at Lacanhá Chansayab, Chiapas.
Woody plant structural censuses were made in acahuals from one to
twenty years old differing in use-history. These data were subjected to
ordination (DECORANA) and classification (TWINSPAN). Results
from these multivariate analyses were compared to the known acahuals’
use-history patterns and successional routes associated to human distur
-
bance patterns were established.
Research, Reviews, Practices, Policy and Technology 51

METHODOLOGY
The Study Area
The study was conducted in the Lacanhá River Plains, at the north
-
eastern border of the Montes Azules Biosphere Reserve, Chiapas,
México (16° 46⬘ N, 91°08⬘ W, 350-400 m.a.s.l.). The climate is warm-
humid with a mean annual temperature of 25°C to 27°C, and an annual
rainfall between 180 cm to 200 cm (García 1973). Precipitation occurs
mainly (> 80%) between June and October. The rendzinic soils are de
-
rived from limestone, and support associations of tropical rainforest
(Pennington and Sarukhán 1998, Muench 1982). According to the Lac-
andon classification there are two types of mature vegetation, “monte
alto” and “chaparral.” The Lacandon also distinguishes between the
acahuals derived from them. The Monte Alto Forest corresponds to
high perennial rainforest (40 m to 60 m high). In contrast the chaparral
is a similar type of vegetation but with trees of smaller size (20 m to 30 m
high) and abundant vines and climbers. Chaparral occurs in patches of
up to 5 hectares within the Monte Alto Forest, in areas with seasonal
flooding, close to rivers and on soils with high organic matter content
(Levy and Aguirre 1999).
The use of renewable natural resources by the Lacandon at Lacanhá
Chansayab is based on S&B agriculture, which relies on the forest’s natu-
ral recovery and concomitant reestablishment of soil fertility through
long resting periods (fallow). During the fallow periods, the Lacandon
use forest resources (wildlife and plants) for self-consumption (extrac-
tion of construction and handcrafts materials, and edible, medicinal, or-
namental and leather-tanning plants), as well as, for trading (xate palm
leaves and mao fiber, Chamaedora spp. and Aechmea magdalenae,re-
spectively). They supplement this resource-use system with small-scale
cattle-raising, fishing, ecotourism and off farm labor. The “milpa” or
S&B corn and associated crops production system, represents the core
of the Lacandonian agricultural activity. Milpa agriculture may be asso
-
ciated with the production of more than 50 different plant crops over
several consecutive years, mostly for self-consumption (Nations and
Nigh 1980).
There are 57 farmers in Lacanhá Chansayab, only ten of them being
typical traditional farmers. The others can be designated atypical tradi
-
tional farmers. These groups differ in their production techniques, their
historical background concerning the use of resources and in their pro
-
duction organizational way. Typical traditional farmers, generally el
-
52 JOURNAL OF SUSTAINABLE AGRICULTURE

derly peasants, are characterized by maintaining production techniques
derived from the ancient S&B agriculture system. Atypical traditional
farmers have adopted the agricultural practices of Mayan groups immi
-
grating from the Chiapas highlands (Choles and Tzeltales), characteris
-
tic of temperate forests (Levy and Aguirre 1999, Levy 2000).
Acahual Selection
Thirty-five peasants authorized vegetation sampling within their
38 acahuals. Time since last cultivation on each acahual ranged from
one to twenty years. No older acahuals were found. A species/area curve
was produced using data derived from the preliminary censuses. A min-
imal area of 225 m
2
(15 m ⫻ 15 m square plot) was chosen for census in
the case of acahuals with a current fallow period shorter than eight
years. For acahuals with a fallow period over eight years the minimum
plot size was 400 m
2
(20 m ⫻ 20 m). Within each acahual plot parcels or
sampling units were delineated, and subdivided to facilitate plant loca-
tion. The number of plots per acahual varied between one to five de-
pending on the size and heterogeneity of its vegetation giving a total of
eighty-four. In the plots a census was carried out of the woody vegeta-
tion. Herb species associated with defined patterns of management were
also included. One important example of these species is Pteridium
aquilinum.
The structural attributes recorded included: (1) species composition,
(2) density, (3) height, (4) projected canopy diameter and (5) visual esti-
mate of relative cover for each species.
Delimitation and inventory of sampling units in two to four years old
acahuals were challenging due to the vegetation density. This compli-
cated the identification of the acahuals’ limits. In some cases, the lack of
clear boundaries between acahuals caused discrepancies between the
use-history described for them and that in the censused areas.
Acahual Use-History
The following information was obtained through interviews, in the
Mayan-Lacandon language, with the owners of the 38 acahuals in
-
cluded in the study: (1) owner’s name, (2) location, (3) current fallow
period, (4) original vegetation type (monte alto or chaparral), (5) num
-
ber of previous fallow periods, (6) length of each previous fallow
periods, (7) number of previous milpas, (8) times that the area was cul
-
tivated per year and whether it was used for summer or autumn milpa,
Research, Reviews, Practices, Policy and Technology 53

(9) number of consecutive years with milpa, (10) number of cultivated
species per production cycle, (11) burn frequency throughout the use-
history and (12) milpa’s weeding method (manual or using machetes).
Information Analysis
The data on use-history for each acahual included: (1) repetition ac
-
ronym; (2) age of the current fallow period (CF); (3) total number of
fallow periods, including the previous and the current one (TF); (4) num
-
ber of previous fallow periods (PF); (5) sum of years of all fallow peri
-
ods throughout the use history (SF); (6) average length of the fallow
period (AF); (7) sum of all milpa’s cultivation periods throughout its
use-history (SC); (8) average cultivation period (AC); (9) rate of total
length of the fallow periods divided by total duration of cultivation peri-
ods (SF/SC); (10) the rate mentioned in (9) divided by disturbance
frequency (SF/SC)/TF); (11) original vegetation type (VT); and (12) far-
mer type (FT). This database was used for the interpretation and discus-
sion of acahuals’ ordination and classification results.
Codification and Analysis of Census Data
Species data for each plot were included in a matrix and then submit-
ted to detrended correspondence analysis (DECORANA) (Hill 1979a).
Additionally, classification using TWINSPAN (Hill 1979b) was carried
out. Cover-percentage intervals chosen for data processing with TWIN-
SPAN were the following: 0% to 1%, larger than 1% to 20%; larger than
20% to 50%; larger than 50% to 80%, and > 80%. Interval width was
defined based on the frequency of species’ cover records.
RESULTS
Acahual Characteristics
Of the 38 acahuals and 84 plots studied 82% and 85%, respectively,
corresponded to sites with a fallow period shorter than ten years. The
modal fallow period is about four years (Table 1).
Of the 84 censuses, 24 correspond to 12 acahuals derived from chap
-
arral (chaparral-acahuals), and the remaining 60 were from 26 acahuals
derived from monte alto (monte alto acahuals) (Table 1).
54 JOURNAL OF SUSTAINABLE AGRICULTURE

Vegetation Use-History
To interpret acahual ordination and classification, information re-
garding use-history was essential (Appendix). This information consid-
ered acahual management data from which two indices were calculated:
the disturbance intensity index (SF/SC), calculated as the sum of years
of all fallow periods divided by the sum of years of cultivation periods
for each field, and the (SF/SC)/TF index, which weighs the disturbance
intensity according to its frequency (total number of fallow periods).
Thus, values lower or equal to one as calculated by these indices, mean
an intensive management, because the cultivation period equals or ex
-
ceeds the fallow period; and numbers greater than one are indicative of a
more persistent use, with a fallow period longer than the cultivation one.
Acahual and Species Ordination Based on Plant Composition
and Cover, and Species Importance in Acahuals, Respectively
The first axis or main gradient of acahuals ordination was correlated
with: (a) acahual’s current fallow period (r
2
= 0.6443), and (b) distur
-
bance intensity [(SF/SC)/TF] (r
2
= 0.2883). Six of the seven acahuals
Research, Reviews, Practices, Policy and Technology 55
TABLE 1. Chaparral (CH) and Monte Alto (MA) derived acahuals with corre
-
sponding current fallow period (FP) and number of censuses.
FP MA CH Frequency % Censuses %
11012.633.6
21237.9910.7
341513.21113.1
4 9 3 12 31.6 29 34.5
522410.589.5
60225.344.8
70112.622.4
82137.956.0
141012.622.4
153037.967.1
172025.344.8
201012.611.2
Total 26 12 38 100 84 100

with fallow periods of at least 14 years are located at the gradient’s up
-
per end, between DCA first axis ordenation values (OV) 287 and 434,
including a total of ten censuses or repetitions; the three remaining repe
-
titions are located from DCA first axis score 225 (Figure 1 and Appen
-
dix). Mean values of the SF/SC and (SF/SC)/TF indices for these
acahuals are 3.5 and 3.3, respectively; that is, their average fallow pe
-
riod was three and a half times longer than the period under cultivation
(Appendix). Acahuals with a fallow period from one to three years were
mainly located at the opposite end of the first gradient, between OV 0
and OV 163. Only two out of the 23 censuses having this fallow period
were placed above this range (Figure 1). Most of these acahuals have
(SF/SC)/TF and SF/SC indices lower or equal to one. Thus, several
acahuals within this group were cultivated during a period equal or lon-
ger than the resting period. The acahual’s use-intensity gradient in the
first ordination axis is evident from the progressive increase of the dis-
turbance indices values [SF/SC and (SF/SC)/TF]. In particular, the (SF/
SC)/TF index best describes the acahual use-history features (Appen-
56 JOURNAL OF SUSTAINABLE AGRICULTURE
400
350
300
250
200
150
0
0
50
100
150
200
250
300
350
400
450
500
DCA-axis 2 (Eigevalue = 0.445)
DCA-axis 1 (Eigevalue = 0.565)
100
50
FIGURE 1. Ordination diagram from detrended correspondence analysis of 84
censuses. Axis 1 is correlated with fallow period. Axis 2 is correlated with previ-
ous vegetation type ( o = 1-3 years; ⵧ
= 4-5 years; 䉭 = 6-8 years; x = 14-20
years).

dix). Eleven of the 13 censuses of acahuals with a resting period bet
-
ween 6 years and 8 years, are located towards the gradient’s center.
The species ordination along the first axis may correspond to a gradi
-
ent associated with the current fallow period, but could also be related to
a use gradient. Species ordered within the first quarter (OV 435 to 581)
(Table 2) are related to the 11 censuses from acahuals with a fallow be
-
tween 14 and 20 years. Taxa present in acahuals with fallow periods of 6
years or shorter were ordered at the lower end (Table 2). Species that are
abundant in acahuals from one to six years old, tend to be located be
-
tween VO 77 and 103 on this (Piper aduncum, P. auritum, Podach
-
aenium eminens). Also, Psychotria pubescens, Cupania rufescens, Belotia
mexicana, Bursera simaruba, Cecropia obtusifolia, Heliocarpus ap-
pendiculatus and Spondias mombin, located between OV 144 and 211,
are the most prominent species in this group.
The second axis may correspond to a gradient between the two origi-
nal vegetation types from which acahuals were derived, namely monte
alto and chaparral (Figure 2). Thus, the upper third of the gradient, be-
tween OV 258 and 362, includes only one of the 12 chaparral-acahuals.
Seven chaparral-acahuals fell in the lower-third, including a total of 15
out of the 27 censuses that comprise this gradient’s section (Figure 2).
Species ordination along the second axis also seems to correspond to
a gradient between the two mature vegetation types (monte alto and
chaparral). However, species with affinity for these vegetation types
were ordered along a large part of this gradient without a clear division
that would allow the identification of those species characteristic of one
vegetation type or the other in a specific section of the ordination axis.
The close ordination of eight species (OV 293 to 241) present only in
chaparral-acahuals is worth mentioning, three of which (Clidemia octona,
Casearia aculeata and Photinia microcarpa) were exclusive of the
JOSEGOJA, B acahual (Table 2).
The lower OV (-141 to 19) ordered 14 species, four of which are ex
-
clusive of chaparral-acahuals (Trema micrantha, Casearia sylvestris,
Ouratea lucens and Vernonia patens); two other species (Tetrapterys
macrocarpa and Oecopetalum aff. mexicanum) grow selectively in this
acahual type, since they are rarely found in monte alto-acahuals.
Apparently the third axis corresponds to a disturbance-frequency
gradient to which acahuals have been subjected; that is, the number of
previous fallow periods registered through the acahual’s use history. To
facilitate the analysis of this third gradient it was convenient to separate
acahuals into two sets based on fallow-period features: (a) acahuals
having a long fallow period, older than six years (LFA), and (b) acahuals
Research, Reviews, Practices, Policy and Technology 57

58 JOURNAL OF SUSTAINABLE AGRICULTURE
TABLE 2. Location and composition of species groups ordered by DECORANA
and classified by TWINSPAN according to their similitude of distribution in the
acahuals.
DECORANA TWINSPAN
Axis 1 Axis 2 Axis 3 Set Subset Grp. Family Species
⫺47 199 61 1 1.1 4 Polypodiaceae
Pteridium aquilinum
ssp. Feei (Schaff.) Maxon.
⫺41 245 ⫺120 2 2.1 11 Leguminosae
Leucaena leucocephala
(Lam.) DeWit.
⫺23 194 ⫺27 1 1.1 3 Ranunculaceae
Clematis dioica
L.
⫺18 189 318 1 1.1 4 Compositae
Clibadium arboreum
Donn. Smith
⫺18 233 407 1 1.1 1 Violaceae
Orthion subsessile
(Stand.) Standl. & Steyerm.
⫺11 239 70 1 1.2 6 Myrtaceae
Eugenia aeruginea
DC.
⫺9 115 286 1 1.2 7 Solanaceae
Solanum torvum
Swartz
⫺7 218 390 1 1.1 1 Leguminosae
Pithecellobium macrandrium
J.D. Smith
2 245 388 1 1.1 2 Leguminosae
Lonchocarpus
castilloi Standl.
9 241 256 1 1.1 2 Apocynaceae
Thevetia ahouai
(L.) A. DC.
9 291 ⫺15 1 1.1 3 Compositae
Pluchea odorata
(L.) Cass.
16 234 403 2 2.1 11 Boraginaceae
Cordia stellifera
I. M. Johnst.
19 242 400 1 1.1 1 Leguminosae
Senna racemosa
(P. Mill.) Irwin & Barneby
24 187 257 8 8.2 40 Zingiberaceae
Costus pulverulentus
C. B. Presl.
31 214 358 6 6.1 19 Leguminosae
Senna fruticosa
(P. Miller) I. & B.
32 66 10 3 3.1 14 Combretaceae
Bucida buceras
L.
34 182 267 1 1.2 7 Compositae
Vernonia deppeana
Lees.
40 197 325 2 2.1 11 Rubiaceae
Alibertia edulis
(L. Rich.) A. Rich. ex DC.
41 258 429 2 2.1 10 Euphorbiaceae
Croton schiedeanus
Schlecht.
49 271 442 8 8.2 38 Compositae
Verbesina chiapensis
Rob. & Greenm.
49 271 442 8 8.2 38 Malpighiaceae
Malpighia glabra
L.
50 212 136 1 1.2 7 Solanaceae
Solanum nudum
H. K. B.
55 298 ⫺26 1 1.2 6 Araliaceae
Oreopanax obtusifolius
L. O. Wms.
57 255 ⫺136 1 1.2 7 Elaeocarpaceae
Muntingia calabura
L.
77 281 ⫺6 1 1.2 7 Piperaceae
Piper aduncum
L.
81 ⫺141 48 8 8.2 40 Ulmaceae
Trema micrantha
(L.) Blume
81 ⫺141 48 8 8.2 40 Flacourtiaceae
Casearia sylvestris
Sw.
86 ⫺22 271 1 1.2 8 Bixaceae
Bixa orellana
L.
88 304 ⫺40 1 1.2 6 Leguminosae
Ormosia schippi
Pierce ex Stand. & Steyerm
89 244 352 2 2.1 10 Rubiaceae
Blepharidium mexicanum
Standl.
92 ⫺24 122 1 1.2 7 Compositae
Podachaenium eminens
(Lag.) Sch. Bip.
94 245 333 8 8.2 40 Rubiaceae
Rondeletia stachyoidea
J. D. Smith
103 52 39 3 3.1 14 Piperaceae
Piper auritum
H. B. K.

Research, Reviews, Practices, Policy and Technology 59
DECORANA TWINSPAN
Axis 1 Axis 2 Axis 3 Set Subset Grp. Family Species
109 199 230 1 1.1 4 Rubiaceae
Genipa americana
L.
110 84 303 1 1.1 3 Leguminosae
Albizia saman
(Jacq.) F. V. Muell.
116 205 54 8 8.2 40 Not identified
117 210 53 3 3.1 14 Myrtaceae
Psidium guajava
L.
119 111 167 3 3.1 14 Solanaceae
Solanum erianthum
D. Don.
122 126 265 1 1.1 3 Amaranthaceae
Chamissoa altissima
(Jacq.) H. B. K.
127 45 277 8 8.2 40 Caricaceae
Carica papaya
L.
128 260 ⫺63 8 8.2 40 Palmae
Scheelea
sp.
128 260 ⫺63 8 8.2 40 Acanthaceae
Ruellia matudae
Leonard
129 281 380 8 8.1 33 Verbenaceae
Aegiphila monstrosa
Mondenke
140 82 326 4 4.1 15 Leguminosae
Senna spectabilis
(D.C) I. & B.
144 150 136 6 6.1 20 Rubiaceae
Psychotria pubescens
Sw.
145 263 369 2 2.1 10 Verbenaceae
Vitex gaumeri
Greenm.
149 40 173 8 8.2 40 Compositae
Vernonia patens
H. B. K.
150 274 101 2 2.2 12 Anacardiaceae
Astronium graveolens
Jacq.
151 203 52 8 8.2 40 Sapindaceae
Sapindus
aff.
saponaria
L.
152 506 147 1 1.1 3 Anacardiaceae
Mangifera indica
L.
153 125 271 6 6.1 22 Boraginaceae
Cordia alliodora
(Ruiz & Pavón) Cham.
157 233 346 8 8.1 35 Apocynaceae
Aspidosperma megalocarpon
Muell. Arg.
157 354 211 1 1.2 6 Melastomataceae
Miconia argentea
(Sw.) DC.
159 230 69 8 8.2 40 Solanaceae
Lycianthes heteroclita
(Sendth.) Bitter
160 ⫺62 248 7 7.1 28 Leguminosae
Lonchocarpus guatemalensis
Benth.
163 228 252 2 2.1 9 Myrtaceae
Pimenta dioica
(L.) Merrill
170 3 116 6 6.1 21 Moraceae
Ficus maxima
P. Miller
170 179 250 2 2.1 9 Compositae
Goldmanella sarmentosa
Greenm.
170 249 126 7 7.2 29 Simaroubaceae
Picramnia aff. brachybotryosa
Donn. Sm.
172 121 92 6 6.1 20 Compositae
Critonia belizeana
B. L. Turner
172 273 275 2 2.1 11 Cucurbitaceae
Lagenaria siceraria
(Molina) Standl.
176 ⫺55 50 8 8.2 37 Ochnaceae
Ouratea lucens
(H. B. K.) Engler
176 90 133 4 4.1 15 Moraceae
Cecropia obtusifolia
Bertol.
181 228 281 7 7.2 29 Lauraceae
Nectandra coriacea
(Sw.) Griseb.
184 ⫺16 164 8 8.2 39 Moraceae
Trophis racemosa
(L.) Urban
184 118 56 7 7.1 26 Leguminosae
Pterocarpus
rohrii Vahl.
185 292 294 2 2.1 9 Bombacaceae
Pachira aquatica
Aubl.
188 40 112 4 4.1 15 Tiliaceae
Heliocarpus appendiculatus
Turcz.
193 154 242 6 6.1 22 Sapindaceae
Cupania rufescens
Triana & Planch.

60 JOURNAL OF SUSTAINABLE AGRICULTURE
TABLE 2 (continued)
DECORANA TWINSPAN
Axis 1 Axis 2 Axis 3 Set Subset Grp. Family Species
194 19 194 8 8.2 40 Icacinaceae
Oecopetalum
aff.
mexicanum
Greenm. & Thomps.
197 5 95 6 6.1 21 Urticaceae
Myriocarpa heterostachya
Donn. Sm.
197 87 46 6 6.1 22 Myrsinaceae
Ardisia paschalis
Donn. Sm.
199 395 158 2 2.1 9 Tiliaceae
Belotia mexicana
(DC.) K. Schum
211 166 203 4 4.1 15 Anacardiaceae
Spondias mombin
L.
216 235 174 2 2.2 12 Leguminosae
Inga pavoniana
Donn.
219 178 271 2 2.2 13 Burseraceae
Bursera simaruba
(L.) Sarg.
221 208 246 6 6.2 23 Combretaceae
Terminalia amazonia
(Gmel.) Exell.
228 193 170 7 7.1 26 Rubiaceae
Hamelia rovirosae
Wernham
230 201 53 6 6.1 21 Meliaceae
Swietenia macrophylla
King
231 46 64 6 6.1 19 Moraceae
Poulsenia armata
(Miq.) Standl.
233 141 135 6 6.1 21 Leguminosae
Inga punctata
Willd.
239 135 150 6 6.2 24 Annonaceae
Guatteria anomala
R. E. Fries
240 144 218 2 2.2 13 Leguminosae
Platymiscium dimorphandrum
Donn. Sm.
241 278 ⫺38 5 5.2 17 Malvaceae
Malvaviscus arboreus
Cav.
249 84 200 8 8.2 40 Annonaceae
Guatteria amplifolia
Triana & Planch
250 ⫺19 9 7 7.1 26 Not identified
250 73 47 7 7.2 29 Tiliaceae
Mortoniodendron guatemalense
Standl. & Steyerm.
255 293 110 8 8.1 33 Rubiaceae
Simira salvadorensis
(Standl.) Steyerm.
261 39 89 2 2.1 10 Lauraceae
Licaria peckii
(Johnst.) Kosterm.
261 253 109 6 6.1 21 Guttiferae
Calophyllum brasiliense
var. rekoi Standl.
264 186 317 8 8.1 36 Rutaceae
Zanthoxylum kellermanii
P. Wilson
266 80 326 8 8.1 36 Leguminosae
Zygia stevensonii
(Standl.) Record
266 166 108 7 7.2 31 Myrsinaceae
Parathesis chiapensis
Fern.
270 227 262 1 1.2 7 Bombacaceae
Ceiba pentandra
(L.) Gaertn.
271 ⫺29 128 8 8.1 34 Malpighiaceae
Tetrapterys macrocarpa
I. M. Johnston
280 153 85 7 7.1 28 Meliaceae
Trichilia breviflora
Blake & Standl.
282 227 136 8 8.1 33 Moraceae
Brosimun alicastrum
Sw.
286 228 125 5 5.2 18 Moraceae
Pseudolmedia
aff.
oxyphyllaria
J. D. Smith
289 222 131 6 6.2 25 Melastomataceae
Miconia impetiolaris
(Sw.) D. Don ex DC.
293 235 291 6 6.2 23 Violaceae
Rinorea hummelii
Sprague
297 281 163 6 6.2 23 Solanaceae
Cestrum nocturnum
L.
300 191 ⫺9 7 7.2 32 Euphorbiaceae
Alchornea latifolia
Sw.
307 170 308 5 5.2 17 Bignoniaceae
Amphitecna apiculata
A. Gentry

Research, Reviews, Practices, Policy and Technology 61
DECORANA TWINSPAN
Axis 1 Axis 2 Axis 3 Set Subset Grp. Family Species
307 256 279 8 8.1 36 Melastomataceae
Clidemia octona
(Bonpl.) L. O. Wms.
314 247 250 8 8.1 35 Meliaceae
Guarea grandifolia
DC.
322 159 227 7 7.2 30 Meliaceae
Cedrela odorata
L.
322 219 40 7 7.2 30 Burseraceae
Protium copal
(Schltdl. & Cham.) Engl.
324 302 30 5 5.2 18 Euphorbiaceae
Sebastiania longicuspis
Standl.
329 24 102 8 8.1 34 Simaroubaceae
Picramnia andicola
Tul.
331 211 31 8 8.2 37 Leguminosae
Acacia glomerosa
Benth.
334 119 98 8 8.1 33 Annonaceae
Cymbopetalum penduliflorum
(Dunal) Baill.
335 279 344 1 1.2 5 Piperaceae
Pothomorphe peltata
(L.) Miq.
337 252 174 8 8.1 36 Lauraceae
Ocotea cernua
(Nees) Mez
342 ⫺65 139 8 8.1 36 Compositae
Piptocarpha chontalensis
Baker
342 132 55 7 7.1 28 Flacourtiaceae
Pleuranthodendron lindenii
(Turcz.) Sleumer
345 173 35 6 6.2 24 Flacourtiaceae
Casearia corymbosa
H. B. K.
345 190 50 7 7.1 28 Piperaceae
Piper aequale
Vahl
347 110 ⫺45 7 7.1 28 Compositae
Eupatorium nubigenum
Benth.
349 147 115 6 6.1 22 Moraceae
Trophis mexicana
(Liebm.) Bur.
349 286 160 8 8.1 36 Rosaceae
Photinia microcarpa
Standl.
349 286 160 8 8.1 36 Flacourtiaceae
Casearia aculeata
Jacq.
351 286 374 5 5.2 17 Chrysobalanaceae
Licania
sp.
362 ⫺40 159 8 8.1 35 Hippocrateaceae
Salacia aff. impressifolia
(Miers.) A.C. Smith
363 332 139 2 2.2 12 Turneraceae
Erblichia odorata
Seem.
366 7 44 8 8.1 36 Cucurbitaceae
Cucurbita moschata
(Duch. Ex Lam) Duch. Ex Poir
368 133 48 8 8.2 37 Celastraceae
Rhacoma eucymosa
(Loes. & Pitt.) Standl.
371 278 205 8 8.1 36 Ulmaceae
Ampelocera hottlei
(Standl.) Standl.
372 110 ⫺12 7 7.2 30 Agavaceae
Dracaeca americana
Donn. Smith
374 242 294 6 6.2 23 Moraceae
Ficus yoponensis
Desv.
380 200 247 7 7.2 32 Euphorbiaceae
Sapium lateriflorum
Hemsl.
381 163 113 8 8.1 33 Meliaceae
Guarea glabra
Vahl
387 105 153 8 8.1 36 Sapotaceae
Sideroxylon
aff. salicifolium (L.) Lam.
396 145 82 7 7.1 28 Malpighiaceae
Bunchosia lanceolata
Turcz.
399 183 94 8 8.1 35 Moraceae
Clarisia biflora
ssp.
mexicana
Ruiz & Pavón
{(Liebm.) W. Burger}
400 238 76 8 8.1 34 Celastraceae
Wimmeria concolor
Schlecht. & Cham.
402 211 19 8 8.1 34 Not identified
403 236 158 8 8.1 36 Leguminosae
Acacia collinsii
Safford
407 184 72 8 8.1 34 Malvaceae
Hampea stipitata
S. Wats.

62 JOURNAL OF SUSTAINABLE AGRICULTURE
TABLE 2 (continued)
DECORANA TWINSPAN
Axis 1 Axis 2 Axis 3 Set Subset Grp. Family Species
410 127 67 8 8.1 36 Annonaceae
Malmea depressa
(Baill.) R. E. Fries
415 161 ⫺19 7 7.2 32 Rubiaceae
Psychotria miradorensis
(Oerst.) Hemsl.
416 128 179 8 8.1 34 Leguminosae
Vatairea lundellii
(Standl.) Kilip ex Record
416 194 86 8 8.1 35 Moraceae
Castilla elastica
Cerv.
417 129 117 7 7.1 27 Rubiaceae
Psychotria limonensis
Krause
418 229 305 8 8.1 36 Annonaceae
Desmopsis stenopetala
(J. D. Smith.) R. E. Fries
422 196 211 6 6.1 21 Monimiaceae
Mollinedia viridiflora
Tul.
422 241 99 8 8.1 36 Aquifoliaceae
Ilex valeri
Standl.
423 22 9 8 8.1 34 Urticaceae
Urera baccifera
(L.) Gaud.
425 203 90 7 7.2 32 Palmae
Bactris mexicana
Burret
426 141 116 7 7.1 28 Apocynaceae
Tabernaemontana amygdalifolia
Jacq.
435 178 201 8 8.1 36 Myrtaceae
Eugenia
aff. koepperi Standl.
442 325 117 6 6.1 20 Rutaceae
Citrus sinensis
(L.) Osbeck
445 126 143 5 5.1 16 Bombacaceae
Ochroma lagopus
(Cav. ex Lam) Urban
447 185 150 7 7.2 31 Leguminosae
Schizolobium parahybum
(Vell.) Blake
448 210 29 8 8.1 35 Leguminosae
Acacia mayana
Lundell
449 128 40 8 8.1 35 Monimiaceae
Siparuna andina
(Tul.) A. DC.
450 110 14 8 8.1 36 Sterculiaceae
Guazuma ulmifolia
Lam.
452 68 ⫺39 8 8.1 36 Piperaceae
Piper sanctum
(miq.) Schlecht.
454 112 134 8 8.1 34 Rubiaceae
Psychotria membachens
is Standl.
457 126 ⫺29 5 5.2 18 Lauraceae
Nectandra globosa
(Aubl.) Mez.
467 164 165 7 7.1 28 Euphorbiaceae
Tetrorchidium rotundatum
Standl.
469 173 100 8 8.1 34 Sapotaceae
Pouteria sapota
(Jacq.) H.E. Moore & Stearn.
472 175 88 8 8.1 36 Meliaceae
Trichilia pallida
Sw.
474 218 88 8 8.1 36 Guttiferae
Rheedia macrantha
Standl. & Steyerm.
485 173 13 8 8.1 36 Sapotaceae
Dipholis minutiflora
Pittier
487 228 115 1 1.2 8 Moraceae
Ficus cotinifolia
H. B. K.
493 229 255 8 8.2 38 Bignoniaceae
Parmentiera aculeata
(H. B. K.) Seem.
514 122 177 8 8.1 36 Palmae
Geonoma oxycarpa
Martius
519 237 84 8 8.1 36 Chrysobalanaceae
Licania platypus
(Hermsl.) Fritsch.
554 202 73 8 8.1 36 Lauraceae
Persea americana
Mill.
581 218 40 8 8.1 36 Not identified
581 218 70 8 8.1 36 Palmae
Astrocaryum mexicanum
Liebm. Ex Martius
581 218 70 8 8.1 36 Moraceae
Artocarpus altilis
Fosberg

with a short fallow period (SFA) of six years or less (Appendix and
Figure 3). The upper third of this axis (OV between 196 and 294)
corresponds to acahuals with a fallow period between two and 15
years. SFA acahuals show fallow periods from two (KINBORBA), three
(CKAYOMCA) and four years (JOSEGOEA, B, C, D) with two, one and
three previous fallow periods, respectively. The LFA set is composed of
three acahuals: one with eight (JOSEGOJB) and two with 15 years of
fallow (CARBORLB and OBREGOLB); the latter two lack a previous
fallow period because the milpa from which they derived was started di
-
rectly from mature vegetation (Appendix).
The opposite end of the third gradient, between OV 0 and 96, includes
acahuals from two to fourteen years. Contrary to the findings at the upper
third, SFA acahuals located at this gradient end do not present a previous
fallow in most cases (13 out of 20 censuses). LFA acahuals located at this
portion of the axis only show one previous fallow in their three censuses
(JORKINKA and CARBORJA, B) (Appendix and Figure 3).
The third species-ordination axis appears to represent a frequency
and intensity gradient of disturbance (amount of previous fallow peri
-
Research, Reviews, Practices, Policy and Technology 63
400
350
300
250
200
150
0
0
50 100 150 200 250 300 350 400 450 500
DCA-axis 2 (Eigevalue = 0.445)
DCA-axis 1 (Eigevalue = 0.565)
100
50
FIGURE 2. Ordination diagram from detrended correspondence analysis of 84
censuses. Axis 1 is correlated with fallow period and axis 2 with vegetation type
(
∆ = chaparral; ⵧ = monte alto).

ods and number of consecutive cultivation years) recorded in acahuals
use-history. Here, similarly to the third ordination axis for acahuals,
species’ ordination interpretation acahuals are divided into two sets based
on the current fallow period and frequency of previous fallow periods:
(a) acahuals with a long fallow period (longer than six years) with and
without a previous fallow period (LFA), and (b) acahuals with a short
fallow period (equal or shorter than six years), with and without a previ
-
ous fallow period (SFA) (Table 2, Figure 4).
At the gradient’s upper quarter, between OV 303 and 442, 25 species
mainly associated to six SFA with several previous fallow periods were
ordered. At the gradient’s lower quarter, between OV ⫺136 and ⫺6,
appear SFA-associated species, without a previous fallow period and
with several consecutive years of milpa, and species LFA-associated
with a previous fallow period. All these species seem to be adapted to a
management type including several consecutive years of cultivation,
64 JOURNAL OF SUSTAINABLE AGRICULTURE
350
300
250
200
150
0
0
50 100 150 200 250 300 350 400 450 500
DCA-axis 3 (Eigevalue = 0.279)
DCA-axis 1 (Ei
g
evalue = 0.565)
100
50
FIGURE 3. Ordination diagram from detrended correspondence analysis of 84
acahuals censuses along axis 1 (fallow period) and 3 (number of previous
fallows) (o = no previous fallows;
䉭 = several previous fallows; x = long fal
-
low period).

and able to persist throughout succession, since they occur both in SFA
and LFA.
The results of the third ordination axis make it easier to relate succes-
sion with disturbance frequency and intensity. Thus, species exclusive
to SFA (which are not present in mature acahuals) can be classified as
non-persistent, whereas, species present both in SFA and LFA may be
considered as persistent, regarding the successional process. In turn, the
agricultural land use-pattern including several previous fallow periods
may be considered as frequent, while the one with a cultivation persist
-
ing for several years may be denominated as prolonged. In this way,
four plant types can be recognized based on their successional duration
and the disturbance type to which they are adapted: (1) non-persisting,
frequent field-use species (Orthion subsessile); (2) non-persisting, pro
-
longed field-use species (Muntingia calabura, Leucaena leucocephala
and Pluchea odorata); (3) persisting, frequent field-use species (Croton
schiedeanus, Belotia mexicana, Cordia stellifera, Pithecellobium macran
-
drium, Lonchocarpus castilloi, Licania spp., Vitex gaumeri, Bleph
-
Research, Reviews, Practices, Policy and Technology 65
500
400
300
200
100⫺
100⫺
200⫺
0 100 200 300 400 500 600 700
DCA-axis 3 (Eigevalue = 0.445)
DCA-axis 1 (Eigevalue = 0.565)
100
0
FIGURE 4. Ordination diagram from detrended correspondence analysis of
173 species along axis 1 (fallow period) and 3 (number of previous fallows)
(䉭 = several previous fallows-associated species; o = no previous fallows-
associated species).

aridium mexicanum, Senna spectabilis, Alibertia edulis and Amphitecna
apiculata); and (4) persisting, prolonged field-use species (Eupatorium
nubigenum, Malvaviscus arboreus, Nectandra globosa, Oreopanax
obtusifolius, Psychotria miradorensis, Acacia mayana, Acacia glom
-
erosa and Sebastiania longicuspis).
Acahual Classification Based on Floristic Composition
and Cover Percentage
In respect to acahual classification, of the 84 censuses conducted in
38 acahuals, 27 groups emerged, composed from one to six censuses
each, and defined between classification levels 4 and 6 (Figure 5). The
conformation of these groups responds to the following factors (hierar
-
chically ordered): (1) current fallow period, (2) original vegetation type,
(3) relationship between fallow and cultivation periods [SF/SC and (SF/
SC)/TF indices], and (4) number of previous fallow periods recorded in
the acahual use-history (Appendix). The set of acahuals groups with a
frequent use and short fallow period (first set) includes acahuals with fal-
low periods between three and four years, derived from monte alto, with
one to three previous fallow periods. Three-year acahuals stand out by
presenting the highest number of previous fallow periods. Thus, acahuals
included in the first set have been subjected to an intensive agricultural
use, defined by a cultivation period equal or longer than the fallow period,
but with several previous fallow periods [(SF/SC)/TF < 1]. The set of
acahuals with prolonged use (second set) is comprised by acahuals with one
to six resting years, represented by 21 censuses, characterized by having
derived mainly from monte alto, lacking previous fallow periods and
having been cultivated continuously for three to eight years. The third set
includes 20 acahuals comprised by 34 censuses with a fallow period of
two to eight years; eight of the 12 chaparral-acahuals, represented by 14
censuses, were included in this set. The old-acahual set (fourth set) is con
-
stituted by 12 acahuals with 20 censuses. Here old acahuals predomi
-
nated (14 years to 20 years), derived from monte alto, without a previous
fallow period and used only by typical traditional farmers. Mean values
for SF/SC and (SF/SC)/TF indices are 4 and 3.3, respectively; these val
-
ues were the highest amongst all sets (Appendix).
Species Classification Based on Their Importance in Acahuals
Table 2 includes the classification set, subset and group to which
each species belongs, as well as, its family and scientific name; sets and
66 JOURNAL OF SUSTAINABLE AGRICULTURE

Old acahuals
(n = 20)
Chaparral
acahuals
(n = 34)
Monte alto acahuals,
prolonged use
(n = 21)
Frequent use
and short fallow
acahuals
(n=9)
(1)
(6)
(4) (4) (3)(3) (3)
(2)
(2)
(2)
(2) (2)
(1)
(1)
(3)
(3) (3) (3)
(4)
(4) (1)
(5)
(5)
(5)
(4)
(4)(4)
FIGURE 5. Dendogram derived from TWINSPAN classification showing relations among sites. The upper labels refer to
the number of acahuals in each set and subset. Values in parenthesis are the number of acahuals in each group. See
Appendix for composition of each subset.
67

subsets correspond to classification levels 3 and 4, respectively. The
classification of 174 species resulted in 40 groups, including from one
to 26 species each. The discussion of species classification will be based
on classification level 3, and comprises a total of eight sets of groups
(Figure 6).
(1) Set of non-persistent species. The first set constituted 27 species,
related to acahuals having a frequent and prolonged use, respectively
(which were in turn classified within the first and second subsets).
Particularly, from subset 1.1 and in relation to acahuals with several
previous fallow periods, it is worth mentioning indicator species such as
Orthion subsessile, Senna racemosa and Pithecellobium macrandrium;
as preferential species because they are present both in SFA and LFA,
Thevetia ahouai and Lonchocarpus castilloi (species persisting under
frequent field-use). Acahuals with several consecutive cultivation years
(prolonged use) show Pteridium aquilinum and Clibadium arboreum as
indicator species. Thus, taxa which are characteristic of acahuals having
a prolonged and frequent use-history are: Orthion subsessile, Pithe-
cellobium macrandrium and Pteridium aquilinum.
Subset 1.2 includes the species Vernonia deppeana, Podachaenium
eminens and Piper aduncum, they are abundant even in acahuals with
several previous fallow periods (first subset), as well as, in acahuals from
one to six years old (classified within the first three sets) (Appendix).
(2) Set of persistent under frequent field-use species. This second set
comprises 16 species which occur in frequently used acahuals (grouped
in the first set) and in acahuals with a prolonged use (second set), as well
as in acahuals of four, eight and fifteen years of fallow period within
subset 4.1 (Appendix). This fact is confirmed by the presence of 50% of
the total persistent species under frequent use (Belotia mexicana, Vitex
gaumeri, Croton schiedeanus, Blepharidium mexicanum, Cordia stellifera
and Alibertia edulis) within this subset. It is worth mentioning that
acahuals in subset 4.1 showed four and eight year fallow periods and
only one had a 15-year fallow period; also, that acahuals with longer fal
-
low resting period were classified in subset 4.2. Inga pavoniana stands
out by occurring at the beginning of succession and producing high
cover values in acahuals with fallow periods between 15 years and 20
years.
(3) Set of disturbance-susceptible species. The third set is defined
from the second classification level (Figure 6) and is constituted by four
species; from them, Piper auritum stands out as an indicador species,
since it grows exclusively in acahuals with one to six years of fallow,
68 JOURNAL OF SUSTAINABLE AGRICULTURE

Mature vegetation
and chaparral
acahuals
Persisting
prolonged
field-use
Mature vegetation
and monte alto
acahuals
Persisting,
frequent
and
prolonged
field-use
Acahual
types
Disturbance
suceptible
Persistent,
frequent field-use
Non-persistent
(short fallows)
Prolonged
field-use
Frequent
field-use
40
(13)
39
(1)
38
(3)
37
(4)
36
(26)
35
(7)
34
(9)
33
(5)
32
(4)
31
(2)
30
(3)
29
(3)
28
(8)
27
(1)
26
(3)
25
(1)
24
(2)
23
(4)
22
(4)
21
(6)
20
(3)
19
(2)
18
(3)
17
(3)
16
(1)
15
(4)
14
(4)
13
(2)
12
(3)
11
(4)
10
(3)
9
(4)
8
(2)
7
(7)
6
(4)
5
(1)
4
(3)
3
(5)
2
(2)
1
(3)
FIGURE 6. Dendogram derived from TWINSPAN classification showing relations between species. The upper labels
refer to the species groups. Values in parenthesis are the number of species in each group.
69

and does not prosper in acahuals subjected to heavy disturbance (first
and second set of acahuals).
(4) Set of acahual-typical species. The fourth set was also defined
from the second classification level (Figure 6), with a single group of
four species that are widely distributed with a high cover percentage
throughout the whole successional gradient. Thus, Spondias mombin,
Cecropia obtusifolia and Heliocarpus appendiculatus were present in
most acahuals sampled.
(5) Set of persisting under frequent and prolonged field-use species.
The fifth set is constituted mostly by species persisting under frequent
(Amphitecna apiculata and Licania spp.) and prolonged use (Malvaviscus
arboreus and Nectandra globosa), and is characterized by the presence
of Ochroma lagopus as an indicator species of mature acahuals (fourth
set of acahuals), being the only species present in subset 5.1, with al-
most 100% cover in most acahuals between 14 years and 20 years
(subset 4.2 of acahuals). This set grouped the last important number of per-
sisting species, and contrary to the second set (species persisting under
frequent use), species included here reached their highest cover in the
oldest acahuals (subset 4.2 of acahuals).
(6) Set of species characteristic of mature vegetation and monte alto-
acahuals. The sixth and seventh sets add 46 species with scarce in-
formation derived from acahuals’ ordination and classification. Only
Tetrorchidium rotundatum and Schizolobium parahybum can be recog-
nized as exclusive of mature vegetation, previously ordered in the first
axis. The sixth set also included Trophis racemosa and Ficus maxima,
two of the three species identified in the second species-ordination axis
as characteristic of monte alto; the third species (Piptocarpa chontalensis)
was classified within subset 8.1.
(7) The seventh set include two species persisting under prolonged
field-use (Eupatorium nubigenum and Psychotria miradorensis), as
well as, Schizolobium parahybum, already mentioned as an indicator of
natural and human disturbance (Miranda 1993b).
(8) Set of species characteristic of mature vegetation and chapar
-
ral-acahuals. This last set, the eighth, is the largest, containing 39% (68)
of all the taxa recorded. Species classified in this set are mainly those
which exclusively prosper in old acahuals, as well as, those growing in
acahuals derived from chaparral. The presence of Acacia glomerosa as
a species persisting under prolonged field-use is worth mentioning.
Subset 8.2 includes species growing in chaparral-acahuals. These spe
-
cies were previously ordered in the second axis’ last quarter (OV ⫺141
to 19). Indicator species of chaparral-acahuals were Trema micrantha,
70 JOURNAL OF SUSTAINABLE AGRICULTURE

Casearia sylvestris, Ruellia matudae, Oecopetalum aff. mexicanum and
Scheelea spp.; whereas, preferential species of this acahual type were
Ouratea lucens, Licaria peckii and Vernonia patens.
DISCUSSION
The modal fallow period (four years) is too short for persistent use
under S&B agriculture. The scarcity of acahuals having the minimum
desirable maturity for agricultural use (longer than ten years) in the
community derives from: (1) the massive arrival of Lacandon immi
-
grants from the Nahá and Metzabok communities, who used part of pre
existing acahuals for their milpas with Lacanhá Lacandons’ approval
(Marion 1991, Levy et al. 2001); (2) the current state prohibition on cut-
ting down mature vegetation (Muench 1982), and (3) the abandonment
of typical traditional Lacandon agricultural culture. Young farmers pre-
fer to grow corn monocrops alternated by annual fires and use short-fal-
low acahuals (Levy 2000, Levy et al. 2001).
The predominance of monte alto acahuals corresponds to the domi-
nance of this vegetation type in the community’s land. It is likely that
chaparral acahuals occur more frequently than expected from the rela-
tive area covered by this vegetation type. Indeed, chaparral patches,
generally smaller than five hectares, are highly appreciated by Lacandon
peasants for agricultural use because smaller trees can be easily cleared
(compared to those of monte alto), and because they form on more fer-
tile and moister soil (Nations and Nigh 1980, Levy 2000).
The finding that the length of the fallow period forms the main ordi-
nation gradient coincides with results reported by van der Wal (1998)
and Purata (1996b), who found that floristic changes depended largely
on fallow duration, in their ordination analysis of 28 and 40 acahuals,
respectively, with fallow periods ranging from 0 years to 50 years.
Most of the species ordered in the first axis have been recognized by
several authors as characteristic in advanced successional stages. Al
-
though there have been relatively few studies regarding old acahuals,
Purata (1986a, 1986b) mentions Astrocaryum mexicanum as one of the
species characteristic in advanced successional stages. In fact, this spe
-
cies was found exclusively in the 20-year-old acahual (KINBORNA)
(Appendix). Martínez (1985) recognizes the genera Rheendia and Psych
-
otria and the species Trichilia pallida and Astrocaryum mexicanum as
shade tolerant and points out that most of them are adapted to grow in the
dim light of the medium and lower strata of the primary rainforest. Fur
-
Research, Reviews, Practices, Policy and Technology 71

thermore, Miranda (1993a), Vázquez-Yanes (1979) and Quintana et al.
(1990) mention Licania platypus, Pouteria sapota, Piper sanctum and
Schizolobium parahybum as species characteristic of old acahuals in the
Lacandonian Rainforest. Particularly, Miranda (1993a) refers to Schizo
-
lobium parahybum, Cecropia obtusifolia and Muntingia calabura as
species indicative of natural or human disturbances.
The species present in acahuals whit fallow periods of six years old
have high frequency and cover in acahuals with intensive use patterns.
As found by Miranda (1993a), Vernonia deppeana, Cordia spp., Loncho
-
carpus castilloi, Trema micranta and Muntingia calabura were typical
of high levels of disturbance. According to van der Wal (1998), Vernonia
deppeana along with Croton draco, are indicator species of intensive
use under S&B agriculture. The presence of Pteridium aquilinum in
rainforest areas cleared for agriculture is widely documented for the
Chinantla area (Gliessman 1978), as well as, in others tropical regions
(Rice 1984). This fern is characteristic of disturbed areas, where its
growth is favored by frequent fires; in addition, it is distinguished by its
allelopathic capacity, leading to the formation of dense persistent patches
when the land is abandoned (Rice 1984). This phenomenon has been
described by Levy and Peña-Valdivia (1999) for Lacanhá Chansayab
lands.
Six species (Piper aduncum, P. auritum, Cecropia obtusifolia,
Heliocarpus appendiculatus, Spondias mombin and Belotia mexicana)
have been recognized by Miranda (1993a), Purata (1986 a, 1986b), and
Quintana et al. (1990) as the most abundant species in young and mature
acahuals. Moreover, Martínez (1985) includes Ochroma lagopus, Ce-
cropia obtusifolia, Siparuna andina, Acacia mayana, Piper auritum,
P. hispidum, Heliocarpus appendiculatus and Trema micrantha within
the colonizing-species group. Particularly, Quintana et al. (1990) de
-
scribe the successional process in the Lacanhá area in general terms and
in stages, starting with acahuals with 3 years to 6 years fallow, in which
the six species already mentioned are present; in those between 6 and 10
years of fallow Ochroma lagopus, Bursera simaruba and Cupania den
-
tata also occur; and in moderately mature rainforest they report the per
-
sistence of Spondias spp., Bursera simaruba and Cecropia obtusifolia.
The separation of acahuals and species based on the original vegeta
-
tion from which they derive (monte alto or chaparral) confirms Braun-
Blanquet’s statement (1979) that vegetation reflects the climate, soil na
-
ture, water and nutrients availability, as well as, human and biotic fac
-
tors to which it has been subjected.
72 JOURNAL OF SUSTAINABLE AGRICULTURE

The description of a disturbance-frequency gradient to which acahuals
have been subjected (third axis) shows the existence of acahuals com
-
posed by species tolerant and intolerant to a use-pattern that includes
several previous fallow periods. Indeed, the presence of certain species
in short or long fallow period acahuals with contrasting use-patterns
(with and without a previous fallow period) supports this assumption.
For this reason, young acahuals that have been frequently disturbed are
associated with mature acahuals. On the other hand, acahuals lacking a
previous fallow period are also linked to mature acahuals.
The characteristics described for the species subsets in the third ordi
-
nation axis are in line with those proposed by Pickett and White (1985)
to characterize disturbance according to frequency (number of events
per period of time), or duration (years per event). As regards distur-
bance frequency, these authors point out that prolonged rotations permit
the existence of more refuges or higher reinvasion opportunities for spe-
cies. Cultivated areas lacking a previous fallow period could meet this
condition, where non-persisting species under frequent use may be fa-
voured. As regards to disturbance duration, Pickett and White (1985)
suggest that prolonged field-use allows the persistence or performance
of poorly represented competitors within the plant association. This
condition is related with species persisting under prolonged use, which
represent a small, scarcely represented group within the vegetation struc-
ture.
Non-persistent and persistent under frequent field-use species may
be related to the short-life and long-life-cycle colonizing species, re-
spectively, described by Whitmore (1975). Non-persistent species in-
clude disturbance indicators, such as Muntingia calabura and Vernonia
deppeana (Miranda 1993a), associated with allelopathic effects and ex-
clusive to the first successional stage, as in the case of Piper aduncum
(Vázquez-Yanes 1979, Miranda 1993a).
Piper auritum as an indicador of disturbance-susceptible species has
captured the attention of several researchers (Anaya 1979) because of
its frequency and abundance in young acahuals, as well as, its role in the
successional process. Anaya (1979) found out that this species’ abun
-
dance in acahuals from southeastern Mexico may be due to its allelo
-
pathic capacity, which hinders the germination and growth of other
native species.
Acahual-typical species have been widely recognized by several au
-
thors as characteristic of acahuals (Gómez-Pompa 1976, Purata 1986a,
Miranda 1993a). These species are present indistinctly in acahuals re
-
gardless of differences among them as to chronology, management pat
-
Research, Reviews, Practices, Policy and Technology 73

terns and original vegetation type. In the case of Lacanhá Chansayab,
Quintana et al. (1990) stressed the dominance of Spondias mombin, Ce
-
cropia obtusifolia and Heliocarpus appendiculatus, as well as, Piper
auritum and P. aduncum in acahuals with three to six years of fallow.
The persistence of frequent and prolonged field-use species would
have allowed them to be related to those classified as primary species by
Gómez-Pompa and Vázquez-Yanes (1979) and Whitmore (1990). The
adaptation to tolerate or even be favored by disturbance gives these spe
-
cies an advantage which ensures that they repopulate during the earliest
successional stages. This phenomenon has been previously described
by del Amo and Gómez-Pompa (1979) and Janzen (1970). In this way,
species persisting under frequent or prolonged field-use are of special
interest, given that they can be used to promote acahuals’ evolution to-
wards advanced stages in relatively shorter fallow periods.
From the works by Miranda (1998), Breedlove (1973), Meave (1983)
and Durán (1999), it is possible to confirm that species included in the
sixth and seventh sets appear in the list of species considered by these
authors as distinctive of mature vegetation. That is the case of Psych-
otria pubescens, Cupania rufescens, Cordia alliodora, Calophyllum
brasiliense ssp. rekoi, Swietenia macrophylla, Poulsenia armata, Casearia
corymbosa, Trophis mexicana, Guatteria anomala and Terminalia ama-
zonia for the sixth set; and of Sapium lateriflorum, Schizolobium para-
hybum, Alchornea latifolia and Dracaena americana for the seventh
set.
Based on the works by Miranda (1998), Breedlove (1973) and Durán
(1999) the affinity of species present in old acahuals within those ma-
ture rainforest can be confirmed. Thus, species included in this set and
present in mature rainforest according to these authors are: Acacia
glomerosa, Ampelocera hottlei, Aspidosperma megalocarpon, Astrocar-
yum mexicanum, Brosimun alicastrum, Castilla elastica, Cymbopetalum
penduliflorum, Geonoma oxycarpa, Guarea glabra, Guarea grandi
-
folia, Hampea stipitata, Pouteria sapota, Licaria peckii, Licania plati
-
lipus, Malmea depresa, Ouratea lucens, Piper sanctum, Sideroxylon
aff. salicifolium, Simira salvadorensis, Siparuna andina and Vatairea
lundelli. For Martínez (1985) tolerant species are exclusive of the pri
-
mary rainforest because they are adapted to grow in the dim light of the
forest understory. Thus, from this set, genera Rheendia and Psychotria
and species Trichilia pallida and Astrocaryum mexicanum would be the
taxa recognized by this author as tolerant ones.
The difficulty in characterizing the species mentioned above as colo
-
nizing and nomadic, as defined by Martínez (1985), derives from the
74 JOURNAL OF SUSTAINABLE AGRICULTURE

imprecision in characterizing these kinds of species. By contrast, the iden
-
tification of plant associations related to management patterns and vari
-
ous ecological conditions through time and space, allowed a detailed
analysis of species to be conducted, which would have not been possi
-
ble without a more general approach. Thus, Martínez (1985) found that
colonizing species have life cycles of up to 50 years and include most
secondary vegetation taxa. The oldest acahual we studied had 20 years
of fallow period and colonizing species were distributed within the first
five sets. Nomadic species according to Martínez (1985) show a higher
correspondence with those recognized as typical of acahuals, persistent
species (fifth set), and from old acahuals or mature vegetation present
since the earliest successional stages. Nomadic species as identified by
Martínez (1985), coinciding with the results of the present work, in-
clude Spondias mombin (typical of acahuals), Ceiba pentandra and
Bursera simaruba (persistent), Calophyllum brasilense and Pseudomelia
oxyphylaria (fifth set), and Mortoniodendron guatemalense, Guarea glab-
ra, Brosimun alicastrum and Pouteria sapota (old acahuals or mature
vegetation). The definition of persistent species surpasses the natural
successional phenomenon, by associating groups of species with spe-
cific milpa management patterns.
CONCLUSIONS
• The present study found that most Lacandonian peasants are cur-
rently performing a very frequent acahual use with a modal fallow
of four years.
• The acahual’s use-history was essential to interpret the census or-
dination and classification, as well as, the successional results gen-
erated by the different Lacandonian field use patterns.
•
The (SF/SC)/TF disturbance index significantly summarizes the
features of acahual use-history by involving the fallow duration, as
well as, disturbance frequency and duration.
•
Acahuals and species were ordered along three gradients: (a) the
main axis combined time (current fallow period) and disturbance
intensity [SF/SC and (SF/SC)/TF indices]; (b) a gradient between
the two original vegetation types from which acahuals derived
(monte alto and chaparral); and (c) a third gradient related with
disturbance frequency (number of previous fallow periods).
•
Five agricultural land use-patterns by Lacandonian peasants were
identified: (a) acahuals with an intensive use, having cultivation
Research, Reviews, Practices, Policy and Technology 75

periods of the same duration or longer than the fallow, with several
previous fallow periods; (b) acahuals with a prolonged use derived
from mature vegetation (without a previous fallow), used for
growing crops just once a year, but uninterruptedly during several
years; (c) acahuals with a four-year fallow and with several previ
-
ous fallow periods; (d) acahuals with a two-to-six-year fallow
without a previous one (in management patterns “c” and “d” fal
-
low nearly triples cultivation period); (e) old acahuals (14 to 20
years) without a previous fallow. Patterns “b,” “d” and “e” corre
-
spond to the land use strategy of typical traditional peasants.
Patterns “a” and “c” are largely applied by atypical traditional pea
-
sants.
•
Disturbance frequency is identified as the disturbance factor mo
-
stly impacting the resulting secondary vegetation structure.
• Successional taxa classification brought about 40 groups, that can
be summarized into the following taxa sets based on their ecolo-
gical (spatial and temporal) affinities and their capacity to adapt
to various land use-patterns: (a) non-persisting taxa; (b) persist-
ing under frequent field-use taxa; (c) disturbance-susceptible
taxa; (d) acahual-typical taxa; (e) persisting under frequent and
prolonged field-use species; (f) species characteristic of mature
vegetation and monte alto-acahuals; and (g) species characteristic
of mature vegetation and chaparral-acahuals.
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RECEIVED: 07/15/03
REVISED: 02/04/04
ACCEPTED: 02/17/04
Research, Reviews, Practices, Policy and Technology 79

80 JOURNAL OF SUSTAINABLE AGRICULTURE
APPENDIX. Acahuals ordenation over the axis 1, and classified by subsets of
groups with main managements features.
ACRON OV SUBS CF TF PF SF AF SC AC SF/SC (SF/SC)/TF VT FT
JOSEGOEC 35 1.1 4 4 3 13 3.2 4 1 3.3 0.8 MA A
CKAYOMCB 211 1.1 3 2 1 6 3 9 4.5 0.7 0.3 CH M
JOSEGOED 76 1.2 4 4 3 13 3.2 4 1 3.3 0.8 MA A
JOSEGOEB 110 1.2 4 4 3 13 3.2 4 1 3.3 0.8 MA A
BORANABE 49 1.2 - - - - - - - - - MA T
MACASTEE 8 1.2 - - - - - - - - - MA T
BORANABD 0 1.2 - - - - - - - - - MA T
ALFKINCA 37 1.2 3 2 1 6 3 6 3 1.0 0.5 MA T
ALFKINCB 12 1.2 3 2 1 6 3 6 3 1.0 0.5 MA T
Mean 3.5 3.0 2.0 9.5 3.1 5.5 2.3 2.1 0.6 T
JOPECHGA 173 2.1 6 1 0 6 6 6 6 1.0 1.0 MA T
JOPECHGB 84 2.1 6 1 0 6 6 6 6 1.0 1.0 MA T
BORANABA 145 2.1 2 1 0 2 2 3 3 0.7 0.7 MA T
BORANABB 114 2.1 2 1 0 2 2 3 3 0.7 0.7 MA T
BORANABC 150 2.1 2 1 0 2 2 3 3 0.7 0.7 MA T
Mean 3.6 1 0 3.6 3.6 4.2 4.2 0.8 0.8 T
CARBOREA 100 2.2 4 1 0 4 4 8 8 0.5 0.5 MA T
CARBOREB 83 2.2 4 1 0 4 4 8 8 0.5 0.5 MA T
CARBOREC 91 2.2 4 1 0 4 4 8 8 0.5 0.5 MA T
MACASTEB 119 2.2 - - - - - - - - - MA T
ENPANICB 71 2.2 3 1 0 3 3 3 3 1.0 1.0 MA A
ENPANICC 55 2.2 3 1 0 3 3 3 3 1.0 1.0 MA A
ENPANICD 50 2.2 3 1 0 3 3 3 3 1.0 1.0 MA A
JKAYOMAA 113 2.2 1 1 0 1 1 3 3 0.3 0.3 MA A
JKAYOMAB 143 22 1 1 0 1 1 3 3 0.3 0.3 MA A
JKAYOMAC 162 2.2 1 1 0 1 1 3 3 0.3 0.3 MA A
MKAYOMCA 141 2.2 3 1 0 3 3 3 3 1.o 1.0 MA A
MKAYOMCB 151 2.2 3 1 0 3 3 3 3 1.0 1.0 MA A
JORPANBA 80 2.2 2 1 0 2 2 4 4 0.5 0.5 CH T
JORPANBB 122 2.2 2 1 0 2 2 4 4 0.5 0.5 CH T
ENPANICA 115 2.2 3 1 0 3 3 3 3 1.0 1.0 MA A

Research, Reviews, Practices, Policy and Technology 81
ACRON OV SUBS CF TF PF SF AF SC AC SF/SC (SF/SC)/TF VT FT
ENRPANFB 199 2.2 - - - - - - - - - CH A
Mean 2.6 1.0 0.0 2.6 2.6 4.2 4.2 0.7 0.7
KINBORHA 156 3.1 6 1 0 6 6 5 5 1.2 1.2 CH T
KINBORHB 154 3.1 6 1 0 6 6 5 5 1.2 1.2 CH T
JUNBORDA 110 3.1 4 3 2 24 8 7 2.3 3.4 1.2 MA M
ALFOCHEB 170 3.1 4 1 0 4 4 1 1 4.0 4.0 MA A
KINBORBA 163 3.1 2 3 2 6 2 6 2 1.0 0.3 CH T
KINBORBB 130 3.1 2 3 2 6 2 6 2 1.0 0.3 CH T
CHAYUKEA 185 3.1 4 3 2 19 6.3 6 2 3.2 1.1 MA M
CHAYUKEB 171 3.1 4 3 2 19 6.3 6 2 3.2 1.1 MA M
MACASTEC 105 3.1 4 2 1 10 5 6 3 1.7 0.8 MA T
MACASTED 171 3.1 4 2 1 10 5 6 3 1.7 0.8 MA T
BALTAZHA 193 3.1 6 1 0 6 6 1 1 6.0 6.0 CH A
BALTAZHB 203 3.1 6 1 0 6 6 1 1 6.0 6.0 CH A
BIILLYFA 172 3.1 5 1 0 5 5 2 2 2.5 2.5 CH M
BIILLYFB 160 3.1 5 1 0 5 5 2 2 2.5 2.5 CH M
MACASTEA 151 3.1 4 2 1 10 5 6 3 1.7 0.8 MA T
KINBORJA 223 3.1 8 1 0 8 8 5 5 1.6 1.6 MA T
JUANCHEA 155 3.1 4 3 2 24 8 6 2 4.0 1.3 MA M
ADOKINEA 186 3.1 4 1 0 5 5 2 2 2.5 2.5 CH A
ADOKINEB 142 3.1 4 1 0 5 5 2 2 2.5 2.5 CH A
PEPENUEA 180 3.1 4 1 0 4 4 1 1 4.0 4.0 CH A
PEPENUEB 219 3.1 4 1 0 4 4 1 1 4.0 4.0 CH A
Mean 4.5 1.7 0.7 9.1 5.3 4.0 2.3 2.8 2.2
VICKINEA 173 3.2 4 2 1 6 3 6 3 1 0.5 MA T
JORPAQEA 173 3.2 4 2 1 8 4 6 3 1.3 0.6 MA T
JORPAQEB 194 3.2 4 2 1 8 4 6 3 1.3 0.6 MA T
CARBORJA 186 3.2 8 2 1 12 6 7 3.5 1.7 0.9 MA T
VICKINEB 175 3.2 4 2 1 6 3 6 3 1 0.5 MA T
MARGCHFA 217 3.2 5 2 1 6 3 2 1 3.0 1.5 MA A
MARGCHFB 221 3.2 5 2 1 6 3 2 1 3.0 1.5 MA A
JUABORIB 225 3.2 - - - - - - - - - CH M
ENRPANFA 183 3.2 5 1 0 5 5 4 4 1.3 1.3 CH A
RICBOREB 190 3.2 4 3 2 9 3 3 1 3 1 MA A

82 JOURNAL OF SUSTAINABLE AGRICULTURE
APPENDIX (continued)
ACRON OV SUBS CF TF PF SF AF SC AC SF/SC (SF/SC)/TF VT FT
CARBORJB 165 3.2 8 2 1 12 6 7 3.5 1.7 0.9 MA T
ALFOCHEA 141 3.2 - - - - - - - - - MA A
JOSEGOEA 126 3.2 - - - - - - - - - MA A
Mean 5.1 2.0 1.0 7.8 4.0 4.9 2.6 1.8 0.9
BORANAEA 234 4.1 4 1 0 4 4 2 2 2.0 2.0 CH A
BORANAEB 256 4.1 4 1 0 4 4 2 2 2.0 2.0 CH A
RICBOREA 213 4.1 - - - - - - - - - MA A
OBREGOLB 225 4.1 15 1 0 15 15 3 3 5.0 5.0 MA T
OBREGOLA 231 4.1 15 1 0 15 15 3 3 5.0 5.0 MA T
JOSEGOJA 281 4.1 8 1 0 8 8 2 2 4.0 4.0 CH A
JOSEGOJB 231 4.1 8 1 0 8 8 2 2 4.0 4.0 CH A
CKAYOMCA 253 4.1 - - - - - - - - - CH M
Mean 9.0 1.0 0.0 9.0 9.0 2.3 2.3 3.7 3.7
PAVIBOMA 338 4.2 17 1 0 17 17 4 4 4.3 4.3 MA T
PAVIBOMB 346 4.2 17 1 0 17 17 4 4 4.3 4.3 MA T
CARBORLB 318 4.2 15 1 0 15 15 3 3 5.0 5.0 MA T
JUABORIA 227 4.2 7 3 2 21 7 5 1.6 4.2 1.5 CH M
JORKINKA 320 4.2 14 2 1 18 9 4 2 4.5 2.3 MA T
JORKINKB 275 4.2 14 2 1 18 9 4 2 4.5 2.3 MA T
CARBORLA 287 4.2 15 1 0 15 15 3 3 5.0 5.0 MA T
JUANCHLB 414 4.2 15 1 0 15 15 6 6 2.5 2.5 MA T
JUANCHLA 417 4.2 15 1 0 15 15 6 6 2.5 2.5 MA T
JUABORMB 365 4.2 17 1 0 17 17 4 4 4.3 4.3 MA T
JUABORMA 337 42 17 1 0 17 17 4 4 4.3 4.3 MA T
KINBORNA 434 4.2 20 2 1 22 11 10 5 2.2 1.1 MA T
Mean 15.3 1.4 0.4 17.313.7 4.8 3.7 4.0 3.3
(OV) ordenation values; (SUBS) subset; (CF) age of the current fallow period; (TF) total number of
fallow periods, including the previous and the current one; (PF) number of previous fallow periods;
(SF) sum of years of all fallow periods throughout the use-history; (AF) average length of the
fallow period; (SC) sum of all milpa’s cultivation periods throughout its use-history; (AC) average
cultivation period; (SF/SC) rate of total duration of fallow periods divided by total duration of
cultivation periods; (SF/SC/TF) the rate mentioned in (SF/SC) divided by disturbance frequency;
(VT) original vegetation type; (FT) farmer type: T, tipic; A, atipic; M, mixed.