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Changes in tree and caterpillar communities
in secondary and primary forests along a climate
gradient in the Southern Yucatán, Mexico
August 2007
MSc. Thesis Tijl Anton Essens
Wageningen University, Chairgroup Forest Ecology and
Forest Management, the Netherlands.
Supervision
Prof. Dr. Frans Bongers, Wageningen University, the Netherlands;
Dr. Carmen Pozo de la Tijera, Ecosur, Unidad Chetumal, Mexico;
Dr. Ir. Henricus F.M. Vester, Ecosur, Unidad Chetumal
and University of Amsterdam.
Supervisors: Dr. Carmen Pozo de la Tijera (director of thesis)
Associate researcher, Insect Ecology and Systematics, Ecosur, División, El Colegio de
la Frontera Sur, Unidad Chetumal. Avenida Centenario km 5.5., Chetumal, Quintana
Roo, CP 77014, Mexico, telephone 01(983) 835 0440 ext 4301 Fax: ext 240.
Email: cpozo@ecosur‐qroo.mx
Prof. Dr. Frans Bongers
Professor, Tropical Forest Ecology, Wageningen University, Lumen no.100,
Droevendaalsesteeg 3, 6708 PB Wageningen, the Netherlands, telephone 0031 317
478029.
Email: [email protected]
Dr. Ir. Henricus F.M. Vester
Senior researcher, Tree and Forest architecture, Ecosur, División, El Colegio de la
Frontera Sur, Unidad Chetumal. Avenida Centenario km 5.5., Chetumal, Quintana
Roo, CP 77900, Mexico, telephone 01(983) 835 0440 and Invited Researcher,
University of Amsterdam, Institute of Biodiversity and Ecosystem dynamics,
Kruislaan 318, 1098 SM Amsterdam, the Netherlands.
Email: [email protected], [email protected]
Author
Tijl Anton Essens
E‐mail: [email protected]
Forest Ecology and Forest Management Chairgroup
FEM 80436, AV2007_11, Wageningen University, the Netherlands.
Index
Summary
1. Introduction……………………………………………………..….…...……1
1.1 Background………………………………………………...….………1
1.2 Vegetation of the Calakmul Reserve……………………..……..……..2
1.3 Disturbed habitats and community ecology …………………………..5
2. Hypotheses………………………………………………………..…..…..…..7
3. Materials and Methods…………….…………………………….…………..8
3.1 Vegetation sampling design………………………………………. ….8
3.2 Fauna sampling design………………………………………..….......10
3.3 Identification caterpillars……………………………………...…..….11
3.4 Statistical analyses…………………………………………..…..........12
Community composition……………………………………...……....12
Species diversity and richness……………………………..………....14
Nocturnal versus diurnal…………………………………..………….16
Equitability of caterpillar communities.………………………..…….17
Relationships between caterpillars and vegetation
structure and richness…………..……………………….………….…18
4. Results………………………………………………………………..………19
4.1 Vegetation……………………………….……………………..……..19
4.1.1 Community composition…………………...……….……...…..19
4.1.2 Diversity……………………….....……………….……...…....22
4.2 Caterpillars.…………………………………..…………….…….…...…..28
4.2.1 Community composition………………………………….….... 28
4.2.2 Diversity………………………...……………………….….......30
4.2.3 Nocturnal versus diurnal..…….……………………………...…32
4.2.4..Equitability of the caterpillar communities..…………….…… 33
4.2.5 Relationships between caterpillars and vegetation
structure and richness …………………………………………..……34
5. Discussion………………………………………………………………...….37
5.1 Changes of plant and caterpillar community structure on the gradient
of climate and forest age..…………………............................…………...37
5.2 Plant and caterpillar diversity on the gradient of climate…………….39
5.3 Does caterpillar diversity increase along a gradient of forest age?.....40
5.4 Does woody plant diversity increase along a gradient of forest age?.43
5.5 Relationships between caterpillars and vegetation
structure and richness...…….....……………………………………….... 45
6. General conclusions …………………………..……………..………..……46
Acknowledgements……………………………………………………………..…....47
References
Appendix
Changes in tree and caterpillar communities in
secondary and primary forests along a climate
gradient in the Southern Yucatán, Mexico
Tijl Anton Essens
Summary
The present study deals with the community composition and species richness of
caterpillars and woody plants in tropical primary and secondary forests along a
precipitation gradient in the Yucatán Peninsula, Mexico. This precipitation gradient
determines forest distribution that ranges from deciduous at the dry end of the
gradient, through an intermediate zone and to the humid semi-evergreen forests. To
investigate possible changes of caterpillar and woody plant communities on a gradient
of stand development, or grade of ‘forest disturbance’, we selected three forest age
classes in the ranges 5-10 years, 10-30 years and old-growth forests. We expected
significantly different community compositions in three climate zones (H1), and
significantly different community compositions in the three forest age-classes (H2). In
addition, the expected response of the two biotic communities is that they become
increasingly diverse from the dry to the humid climate zone (H3) and also from
disturbed to undisturbed habitats (H4). Moreover, the older forests were expected to
be more equitable compared to the young forests. Finally we tried to identify if there
is a relation between caterpillars with respect to the number of woody plant species of
different size classes and average canopy height of forest plots (H5).
Taking into account the species lists and species abundances, the composition of
caterpillar and vegetation communities across climate zones and forest age was tested
for significance with non-parametric ANOSIM test, analogue to the ANOVA. Results
showed that there is an overall significant dissimilarity among caterpillar communities
across climate zones (p=0.01) as expected (H1), and also in woody plant communities
among the three mature forest types of the climate zones (p=0.001). The change in
species composition of plant communities is also found along the gradient of forest
disturbance. Here, caterpillar communities among the three forest age-classes differ
significantly (p<0.01) as expected (H2), as well as plant communities (p=0.001).
Moreover, there is a tendency of an increasing dissimilarity of the caterpillar and tree
communities among the forest age-classes from the humid climate zone towards the
dry climate zone.
Sample-based rarefaction procedures were used to estimate the increase of the
caterpillar and plant species diversity with increasing sampling effort, while
individual-based rarefaction was applied to account for differences in density of
individuals in different habitats. The woody plant communities in the old growth
forests are sufficiently sampled to give a good indication of diversity across the
climate gradient. In contrast to the expected increase of species from the dry to the
wet climate zone (H3), the individual-based richness of woody plant species in the old
growth forests is highest in the medium climate zone. Individual-based richness of
woody plants in old-growth forest was indirectly tested for significance using the
Clench model. Despite the tendency that the forest in the medium climate appears
richer, this difference was not significant but is does contain the highest number of
unique species present of woody plant species. Similarly, the caterpillar communities
tend to be the most diverse in the medium climate zone and maintain the highest
number of unique species.
Furthermore, the results indicate that for woody plant species of ≥10 cm DBH, the
individual-based rarefied diversity and the Simpson index tend to increase with forest
age in all climate zones as expected (H4). The same patterns was found for woody
plant species 3≤ x 10 cm DBH in the medium climate zone, but the dry and the humid
climate zone tend to be more diverse in the forest patches of 10-30 years old. The
caterpillar communities show higher individual-based diversity and Simpson indices
in the secondary forests of 5-10 years old in the dry and the medium climate zone; in
the humid climate zone the secondary forests of 10-30 years appears most diverse.
Even so, significant differences between caterpillar diversity in forest age-classes
were only found in the dry climate zone, where the mature forest is significantly
different from the two secondary forests phases (H4). Similar to the results of woody
plant species, we found most unique caterpillar species in the medium climate zone.
Caterpillar log rank abundance rank plots did not reveal noticeable distinct
equitability for caterpillar communities on the gradient of forest age. The
hypothesized relation between caterpillars and the vegetation species richness and
structure cannot be not confirmed by ordination (CCA) (H5).
The results suggest that caterpillar and plant diversity are directly influenced by
climate and forest age causing a different species biotic compositions and abundances.
However, there is little statistical evidence showing a consistent response of
increasing or decreasing species diversity along the climate and the disturbance
gradient. One of the future challenges is to characterize meaningful groups within
caterpillar and woody plant communities in order to show effects of forest change
throughout various developmental stages upon caterpillar species composition and
diversity.
1. Introduction
1.1 Background
The South of the Mexican peninsula Yucatán holds large seasonal tropical forests.
The main protected area in this zone is the Calakmul Biosphere Reserve, located
towards the border of Guatemala (Figure 1). It extends over 723 185 ha and presents a
precipitation gradient creating pronounced differences in vegetation and
deciduousness, forming an ecocline (Vester et al. 2007). The Calakmul Reserve is part
of the Mesoamerican corridor, which connects several reserves in Belize and
Guatemala. Like many other tropical forests it contains numerous species and large
carbon stocks (Primack et al. 1988).
The Tropical Ecosystem Environment Observations by Satellites (TREES) project has
identified the Southern Yucatán Peninsular Region as a hot spot of tropical
deforestation. In the last three decades high rates of deforestation due to an increasing
population and changes in land use systems throughout the southern Yucatán, have
influenced the ecological functions and characteristics of the above mentioned
ecocline (Turner et al. 2001).
Together with deforestation by human activities, natural disturbances (i.e.; hurricanes,
fires) have substantially altered the forest structure in some areas of the Calakmul
region, leading to a mosaic of forest age-stands, including a variety of plant and
animal composition and diversity.
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El Colegio de la Frontera Sur (ECOSUR) together with Clark University, Virginia
State University and Rutger University study the vulnerability of the coupled human
environmental system in the Southern Yucatán peninsular Region (SYPR), a NASA
funded program. One of the program’s projects investigates the effects of land use
change on biodiversity (vegetation and fauna). The present study contributes to this
project comparing species richness and composition in secondary and primary forests
on a climate gradient which governs forest types from deciduous to semi-evergreen.
The contrast mature – secondary forest is taken as representative for the increasing
deforestation and the humid - dry gradient is taken as model for a possible long-term
drying trend. The biotic groups used in this study are plants and Lepidoptera,
specifically caterpillars.
1.2 Vegetation of the Calakmul Reserve
The Calakmul Reserve contains a gradient of vegetation that correlates to a north-
south rainfall gradient of 900-1500 mmyr-1. The length and intensity of the dry season
determines largely the vegetation type and deciduousness (Figure 1). Three forest
types can be recognized in the study area (Flores and Espejel 1994, Martínez and
Galindo-Leal 2002, Pérez-Salicrup 2004). Firstly, tropical low semi-deciduous dry
forest (canopy height <15 m) is mainly found in the uplands of the drier northwest
part of the study region (Lawrence et al. 2004). It presents an annual precipitation
ranging from 900-1000 mm yr-1. More than 75% of the trees loose their foliage during
the dry season between April and May. The intensity of leaf loss and its southward
extent varies annually, apparently linked to the amount of precipitation received
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during the dry season. The most common tree species for this forest are Thouinia
paucidentata, Beaucarnia pliabilis, Guaiacum sanctum, Lonchocarpus yucatanensis,
Bursera simaruba, Haematoxylum campechianum, Ceiba schotti, Pseudobombax
ellipticum and Maytenus schippi.
Figure 1. The Southern Yucatán Peninsular Region, with a precipitation gradient from the dry North to
the more humid Southern border of the Calakmul region. Circles represent the field stations in 1) low
stature semi-deciduous forest, 2) medium stature semi-evergreen and 3) high stature semi-evergreen.
The second forest type in the Calakmul Reserve is medium statured tropical semi-
evergreen forest, with canopy height between 15 and 25 m. It presents a shorter dry
season and less leaf loss (25-50% of the trees are deciduous). The annual precipitation
ranges from 1000-1200 mm yr-1. Tree species commonly found in these forests are
Brosimum alicastrum, Manilkara zapota, and Pouteria reticulata. The third forest
type is high tropical semi-evergreen (canopy height is generally >25 m). The
dominant tree species belong to the family Sapotaceae, the most common species are
Manilkara zapota, M. chicle, Pouteria sapota, P. amygdalina, P. campechiana and P.
reticulata (Galindo-Leal, 2001). This forest type presents a less characteristic dry
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318000´ N
18015´ N
19000´ N
18045´ N
18030´ N
18015´ N
90015´ W 90000´ W 89045´ W 89030´ W 89015´ W
Dry forests Humid forests Deforested areas Reserve boundaries
3
season and leaf loss is related to particular species that (partially) loose their foliage
during the dry season. The annual precipitation ranges from 1200-1400 mm yr-1. High
semi-evergreen forests are located in the east and south of the study area (especially
near the Guatemala border).
Figure 2. Impression of a vegetation mosaic in
Calakmul with agricultural crops chilli and maize
(in the front), secondary forest patches (right
middle) and old growth forests (background).
Location of this photo: Dos Lagunas Sur,
Campeche, Mexico (T. Essens).
The most important economic activities in this region are agriculture and forestry,
which have lead to a mosaic of mature forests and secondary forest and agricultural
fields (Figure 2). The traditional ”slash and burn” agriculture is a common practice
and characterized by cutting patches of mature or secondary forest followed by
burning the vegetative remains. Afterwards the patches are cultivated with crops such
as maize, beans and squash. After one year of agricultural activities comprising two
harvests, the fields are abandoned. In the period that follows, regeneration of
vegetation establishes successional communities of vegetation.
This agricultural practice leads to a mosaic of young forest stands with predominantly
pioneer species and older stands that hold both pioneers and long living species. Also
forest structure differs in the forest patches (e.g. basal area and the distribution and
4
variability of canopy heights) and related abiotic factors (e.g. light intensity, soil
moisture, humidity). Because caterpillars often consume a limited number of plant
species we can expect the changing forest composition results in different caterpillar
composition. Therefore we hypothesize that anthropogenic impact described above in
different vegetation types results in different caterpillar (Lepidoptera) composition.
1.3 Disturbed habitats and community ecology
Different theories have been developed explaining species richness on gradients of
disturbances. It is generally accepted that disturbances cause a turnover in the species
composition, principally due to differences in times for establishment and degrees of
adaptation to a more complex ecosystem, and a decline of species richness for both
flora (e.g. Eggleton et al. 1998) and invertebrate fauna (e.g. Brown 1994, Roth et al.
1994, Schowalter and Ganio 1999, Holloway 1998, Floren and Linsenmair 2001).
Frequently cited hypotheses that formulate causal relationships between disturbance
and species diversity are the Intermediate Disturbance Hypothesis (Connell 1978, Fox
and Connell 1979, Sheil and Burslem 2003) and the Dynamic Equilibrium Model
(Huston 1979, 1994). Connell (1978) hypothesizes that intermediate levels of
disturbance will maximize diversity, while other studies indicated multiple equilibria
(Hanski et al. 2002), non linearity and threshold effects (Nyström et al. 2000).
Caterpillars were collected, in order to investigate their responses to changing woody
plant communities along a forest age gradient. It has been postulated that Lepidoptera
provide a suitable group to study the effect of habitat change because they are
responsive to changes in their habitat and therefore potentially good ecological
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indicators (Eberhardt and Thomas 1991, Brown 1997, Thomas et al. 2001, Pozo
2006). Their sensitivity to changes is explained by their strong relation to their habitat
during their life cycle (Ehrlich and Raven 1965, Ehrlich 1984, Murphy and Wilcox
1986). Lepidoptera have a so-called holometabolic life, also called complete
metamorphism, which includes four life stages as an egg, larva, pupa and imago. The
dependence of the larval stages on certain host plants, and the dependence of flowers
for honey when adult (Gilbert 1980, Jennersten 1988, Rausher and Feeny 1990) are
relations typically affected by some types of habitat disturbances such as land use
change (Ehrlich et al. 1972, Murphy et al. 1990). In addition, the egg laying behaviour
of butterflies in their search for host plants seems to be directed towards optimal
conditions for offspring survival (Smiley 1978, Thompson and Pellmyr 1991, Bernays
and Chapman 1994, Santiago Lastra et al. 2006). Moreover, the distribution of many
butterfly species appears to be restricted by climatic conditions (Pollard 1979, Turner
et al. 1987, Dennis and Shreeve 1991) and so we might find strong effects of forest
perturbation in the various forest types.
6
2. Hypotheses
This research aims at contributing to our understanding of the dynamics of vegetation
and Lepidoptera communities along the climatic gradient from deciduous to semi
evergreen forests and the influence of anthropogenic disturbance on vegetation and
Lepidoptera communities. Three categories of forest age-classes were used to describe
the disturbance gradient, considering the youngest vegetation as “high disturbance”
and mature forest as “low disturbance”. The age is calculated from the moment fields
were abandoned by the farmers. The following hypotheses will contribute to respond
the questions how the structure and diversity of caterpillar and vegetation
communities changes along the gradients of forest age and climate and if caterpillar
diversity is related to vegetation diversity and structure. The response of biotic
communities to climate zones and forest age will be discussed in the light of existing
theories on these themes.
H1. Tree and caterpillar species composition are different in the climate zones.
H2. Tree and caterpillar species composition are different in the forest age-classes.
H3. Tree and caterpillar species diversity increase along the dry to wet gradient.
H4. Tree and caterpillar species diversity decrease with decreasing forest-age.
Furthermore, an exploratory analysis was done to investigate the link between
diversity and canopy height and on the other hand the diversity and composition of
caterpillar communities.
H5. There is a relation between caterpillar diversity and composition and tree
diversity vegetation structure.
7
Looking for caterpillars in the canopy
3. Materials and Methods
3.1 Vegetation sampling design
Fieldwork was done in the ejidos Conhuas, Zoh Laguna (and the bordering ejido
Nuevo Becal) and Dos Lagunas Sur that represent dry, medium and wet forests
respectively. Two sets of circular plots of 500 m2 were established. The first set is a
mixture of plots located along the climate and the disturbance gradient. For each
climate zone it comprises 12 plots in forest patches of three age groups: 4 plots in 5≤
x ≥10 year old vegetation; 4 plots in 10< x <30 year old vegetation and 4 plots in old
growth forest (in total 36 plots, Table 1). The second set is more at landscape scale
and each climate zone contains 32 mature forest plots (total 96 plots). The mature
forest plots used in the first set were randomly selected from the second set of three
times 32 plots in each old growth forest type.
For all plots, woody plants ≥10 cm DBH were identified, and diameter at breast
height (DBH) was measured as well as their total height (Figure 3). In the centre of
each plot a circular subplot of 100 m2 is located where all species 3≤ x <10 cm DBH
were identified, and their DBH and total height measured. During the field trips the
local guide and a botanist identified the plants. Samples of the plants were collected
using a botanical press for their posterior identification in the herbarium of Ecosur and
Merida (CIC).
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Table 1. The time spend on searching caterpillars and the amount of surface area covered. In the first
column, three forest types distinguished in the Calakmul area are shown. Within each of the three forest
types, the 12 circular plots of 500m2 are selected in three forest age-classes. The surface area covered
for searching caterpillars, amount to a total of 2000 m2 / forest age-class. Survey hours are calculated
for five fieldtrips (5 fieldtrips*4 plots *4 people*1 survey hour). The surface area covered for searching
caterpillars, amount to a total of 2000 m2 / forest age-class.
forest types Plots on a gradient of disturbance Surface area (total of plots)
Survey hours
80 80 80
Tropical low semi- deciduous dry forest (dry)
4 plots in mature forest 4 plots old secondary forest (10< x <30 yr) 4 plots young secondary forest (5≤ x ≥10 yr)
2000 m2 2000 m2 2000 m2
Σ 160
80 80 80
Medium tropical semi-evergreen forest (medium)
4 plots in mature forest 4 plots old secondary forest (10< x <30 yr ) 4 plots young secondary forest (5≤ x ≥10 yr )
2000 m2 2000 m2 2000 m2
Σ 160
80 80 80 High tropical semi-
evergreen forest (humid)
4 plots in mature forest 4 plots old secondary forest (10< x <30 yr ) 4 plots young secondary forest (5≤ x ≥10 yr )
2000 m2 2000 m2 2000 m2
Σ 160 Sum 36 plots of 500m2 18000 m2 Σ 480
W
Figure 3. Plot design for
m2. On the right, an imp
higher strata.
N
2 m
500 m2
100 m2
E
S
vegetation and caterpillars. On the left the two circular plots of 500 and 100
ression of a 500 m2 plot scanned for caterpillars at the forest floor and the
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3.2 Fauna sampling design
Pozo et al. (2003) found that different butterfly families in the Calakmul region occur
in different periods of the year. Therefore, it was decided that sampling of caterpillars
should cover all seasons. The sampling took place in the set of 36 plots (table 1). Two
fieldtrips were done in the wet season (July-November), two fieldtrips in the season
called ´´Nortes´´ (December-January), a period of cold northern winds and finally one
sampling in the dry season (February – June).
Furthermore, Lepidoptera are found in all strata of the forest (Schulze et al. 2001) and
we assumed that caterpillars consume leaves from shrubs as well from full grown
trees, thus, caterpillars and pupae were collected in a horizontal and a vertical
component in the thirty-six plots. Each plot was surveyed by four people for 1 hour.
Two people spent 1 hour in effective searching for caterpillars in the under storey up
to 2 meters (horizontal component) and two other people searched for 1 hour in the
higher region of the forest (vertical component, fig. 3). Professional climbing gear and
a ladder were used to reach a height >2 meters. Pilot investigations revealed that on
average 4 big trees (>20 meters in height) and about 5 smaller trees (<15 meters in
height) were scanned in old growth forest during this time.
During survey time, caterpillars were handpicked, photographed and inserted into 5
ml vials filled with 98% alcohol. Unfortunately, this is presently the only way to keep
the damage to soft-tissued insects to a minimum. The stored caterpillars were used for
visual identification based on morphological characteristics using a microscope.
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3.3 Identification of caterpillars
In Calakmul 430 species of diurnal Lepidoptera (Rhopalocera) were found (Pozo et al.
2003, Martinez et al 2005). For moths the number of species may be up to 7 times as
large (Heppner 1991). The species diversity of Lepidotera in general requires a high
level of experience for identifying species. Therefore it was necessary that larvae were
collected in the field and identified on a later moment. All individuals are deposited in
the reference collection (INE number QNR.IN.018.0497) at Ecosur, Chetumal.
Identification took place based on morphological characteristics using identification
keys and photos (e.g. Peterson 1962). The photo is an important tool for identification
as colours and shapes tend to change in alcohol. Moreover, the photos were matched
with photos of species that occur in Guanacaste, Costa Rica
(http://janzen.sas.upenn.edu/Wadults/searchcat2.lasso). For this work, all individuals
have been identified at family level, and species were separated in morpho-species.
11
Gonodonta ssp. Noctuidae
3.4 Statistical analyses
Community composition
The assumptions of homogeneity in the variance and normal distribution of the
abundance data of vegetation and caterpillars in each treatment were tested with the
Kolmogorov-Smirnov test using SPSS for Windows (2006). Deviations of normality
and of homogeneity in variance were in all cases significant. Therefore, further
analyses for community composition were done with non-parametric tests. The
ANOSIM test, an analogue to the ANOVA, was employed to test for significance in
the composition of caterpillar and vegetation communities among climate zones and
forest age using PRIMER software (Clarke and Gorley 2006). The ANOSIM results
in dissimilarity values in community composition of compared groups. Importantly,
the dissimilarity is calculated using the overlap of species lists and the species
abundances, which makes this test appropriate for comparative community
composition assessments.
Before ANOSIM was executed, transformations were applied to the abundance data
in order to limit the effect of zeros and the contrast of extreme high and low
abundance values. The abundance data of woody vegetation in the 36 plots were root
transformed. The abundance data of woody vegetation in the sample set of 96 plots in
old growth forest were log transformed. The caterpillar abundance data were 4th root
transformed. Then, Bray-Curtis resemblance matrices for woody vegetation and
caterpillars were generated (Bray and Curtis 1957) and used for differences in
community composition between the climate groups and forest age-classes. The Bray-
12
Curtis resemblance values show plot to plot combinations, and creates e.g. a 36 by 36
matrix (of which half contains values). Furthermore, the Bray-Curtis matrix expresses
resemblance values varying between 0 and 1, where 0 is no resemblance and 1 means
complete resemblance. Every value reflects the resemblance of one plot with another
considering the species list and their abundances. For the ANOSIM test, plots were
grouped based on categories of forest age and climate and the groups were tested
against each other for significance.
As mentioned, ANOSIM is an approximate analogue of the standard univariate 1- and
2-way ANOVA (analysis of variance) tests. I executed two types of tests with
ANOSIM. The first test is two-way nested, a hierarchical design which tests for the
differences between climate groups using age groups per climate as samples. This test
gives three main outcomes. The first outcome gives a global RANOSIM for the overall
difference between age groups across all climate groups, the second is a global
RANOSIM, which expresses the overall difference between climate groups using age
groups as samples. Finally, the program provides a table with pairwise differences
between climate groups. The two-way nested option was selected for the vegetation
and the caterpillar data set of 36 plots according to the hierarchical lay-out of the
sampling design, where climate is the first level and forest age-class is the second.
The second type of test executed with ANOSIM, is the one-way pairwise test which
analyses differences of vegetation and caterpillar assemblages found in the forest age-
classes for each climate zone independently. Also this test gives two types of
outcomes; the first is a global RANOSIM for the overall difference between age groups
within a climate group, the second is a table with post hoc pairwise tests showing
13
specifically the dissimilarities between the communities found in the forest age-
classes. This test was applied to vegetation and caterpillar data for the 36 plots and for
the vegetation data in the mature forest using information from the 96 plots.
The RANOSIM values can vary between -1 and 1. A value of 0 or lower means that there
are no differences or relation between sample caterpillar communities, while 1 means
that the samples of the groups are very distinct. In addition, the Bray-Curtis
resemblance matrices were used to create MDS plots (multi-dimensional scaling). The
plots show caterpillar and vegetation communities of each plot in ordinate space. The
interpretation of the MDS is in accordance with the resemblance matrix: sample sites
that are close together represent more similarity than points that are far from each
other.
Species diversity and richness
Bar graphs were drawn showing total numbers of plant and caterpillar species in each
habitat. Caterpillars and woody plant species in old growth forests were further
investigated for the number of shared and unique species.
In order to see if there are patterns or tendencies of floral and faunal diversity,
individual-based and sample-based accumulation graphs were generated.
Accumulation curves are recommended by Soberon y Llorente (1993) with the aim to
compare species lists in different environments. An advantage of its application is to
use them as predictive tools for biodiversity studies (Soberon y Llorente 1993). For
14
the calculations on expected species number, number of samples and individuals, the
software EstimateS has been used (Colwell 2004). Sample-based and individual-based
rarefaction curves were drawn for plant and caterpillar data. First, sample-based
accumulation curves were drawn to investigate how caterpillar and plant species
richness (MaoTau = expected species) changed over increasing sampling area. The
curves give the opportunity to evaluate the expected species richness for samples
simulating samples drawn at random from the pooled samples for all plots (Colwell
2004). Theoretically the asymptote of the curve is the maximum number of species in
the area. The steepness of the curve indicates whether plot to plot variation is high or
low.
Individual-based rarefaction was developed as a measure of species richness by
rarefaction to overcome sample size and species density effects, allowing the
comparison between communities where, for example, densities of animals and plants
are very different. The accumulation curves plots the number of expected species and
numbers of individuals (Gotelli and Colwell 2001) to investigate the levels of richness
between old-growth forest types and forest age-classes within a climate zone.
Individual-based accumulation curves were generated for the woody plant species in
the sample set of 96 old-growth forest plots. This procedure was repeated for
caterpillars in the different forest age-classes and for woody vegetation in the 36 plots.
The larvae of Lepidoptera have not systematically been studied in Mexico and
therefore not much is known about the study object. Moreover, the study area is
heterogeneous with many rare species (Pozo et al. 2003). In a similar situation,
Soberon y Llorente (1993) advocated that the Clench model should be used to obtain
15
asymptotes (Clench 1979). We used the Clench model on the individual-based
richness along the gradient of forest disturbance. Various comparable points were
interpolated on the Clench curves (Colwell et al. 2004) which, in turn, were used to
apply a t-test (Magurran 2004). In this way individual based rarefied diversity that
accounting for sample size and density effects can be tested indirectly. The Clench
model and t-tests were restricted to the data of caterpillars along the gradient of forest
disturbance and plants in old growth forest types to compare the biodiversity. An
interval of 50 individuals was applied on Clench curves, representing the number of
species along increasing number of individuals, to obtain the number of species that
were used to execute the t-test. For small diameter plants in the 500m2 plots, the level
on which the number of individuals (800 individuals) could be compared was much
higher for the 100m2 plots (500 individuals) for the trees with a larger diameter.
Therefore, to run the t-test for plants, 3≤ x <10 cm DBH (500m2 plots) 10 points were
interpolated, while for the plants ≥10 cm DBH (100m2 plots) 16 points were obtained.
For caterpillars 10 points were used. Though the t-test is our main interest, the curves
of the Clench model also allow to make tentative predictions about the dimensions of
plants and caterpillar communities by extrapolating the curve to obtain the asymptote.
Lande et al. (2000) advocate the use of the Simpson index as an unbiased measure of
diversity for smaller sets of non-parametric samples. The Simpson index is a
dominance measure that weighs towards abundances of the commonest species rather
than providing a measure of the species richness. It calculates the probability of two
individuals randomly drawn belonging to different species (Magurran 1988, 2004).
Accordingly, calculations were performed using the same methods.
16
Nocturnal versus diurnal
The response of Lepidoptera to forest disturbance is further explored by comparing
the relative differences of moth and butterfly populations throughout in the selected
habitats. We calculated the relative proportion of butterfly and moth larvae in relation
to the total number of caterpillars.
Equitability of the caterpillar communities
Rank order abundance plots, sometimes referred to as dominance-diversity curves,
were used as a technique to describe the equitability of caterpillar communities across
the different forest age-classes. The rank order abundance curves simply plots log
number of individuals of each species against the rank in a series of most to less
abundant. The equitability of a caterpillar assemblage can be interpreted from reading
the steepness of the line in the first 5 ranks and the length of the tail. The prototype
disturbed habitats have relatively few ranked species and the first ranked present very
high abundances. This causes a short and steep curve. Undisturbed habitats have some
abundant species, but not as numerous as disturbed habitats. In large samples from
high diverse habitats, there tends to be a long tail of rare species (between one and 5
individuals only).
17
Relationships between caterpillars and vegetation structure and richness
Multivariate analyses were executed with the intention of examining the relationship
between caterpillar communities and the structure and richness of forest plots, with
the software CANOCO (Ter Braak and Van Tongeren 1995). First, a detrended
correspondence analysis (DCA) was executed to examine which ordination technique
was most appropriate. According to the result of the DCA, the Canonical
Correspondence Analysis (CCA) was applied (Lepš and Šmilauer 2003). The
variables incorporated for forest structure are canopy height, and for diversity (1) the
number of woody plant species 3≥ x <10 cm DBH, (2) the number of woody plant
species ≥10 cm DBH, (3) the number of woody plant species ≥3 cm DBH and finally
forest age of the plot. Each of these variables will be evaluated for a possible relation
between vegetation and caterpillar assemblages.
18
4. Results
4.1 Vegetation
4.1.1 Community composition
Differences in woody plant communities in three Old-growth forest types
The data used for these results are based on the 96 plots in old growth forest types
located in the three climate zones. In total 182 species of woody plants ≥10 cm DBH
were identified (1499 individuals). Among plants 3≤ x <10 cm DBH, 148 species
were identified (1762 individuals). Together, the 96 old-growth forest plots contain
232 woody plant species ≥3 cm DBH.
The sampled vegetation in both size classes shows significant (p = 0.001) and strong
overall dissimilarities among the three old-growth forest types (Table 2). The
difference between the plant composition of the medium and the humid forest is
smallest, while the other combinations are more dissimilar.
Diameter classes 3≤ x <10 cm DBH ≥10 cm DBH
Statistics RANOSIM p RANOSIM p Overall dissimilarity 0.723 0.001 0.735 0.001
Dry / Medium 0.766 0.001 0.795 0.001
Dry / Humid 0.871 0.001 0.777 0.001
Medium / Humid 0.541 0.001 0.662 0.001
Table 2. Dissimilarity (RANOSIM) statistics and significance level for plants of two size classes in the three
mature forest types
19
The MDS plots for mature forests illustrate that the samples which belong to the same
forest type are clustered rather well. Only some samples are dissimilar from the
cluster to which they are associated. In the 3≤ x <10 cm DBH size class the cluster is
more separated (Figure 4) though not expressed clearly in the statistics (table 2).
Figure 4. The sample set of 96 plots in mature forests for trees
≥10 cm DBH (left) and plants 3≤ x <10 cm DBH (right) in MDS
based on the Bray-Curtis similarity matrix.
Differences in Plant Communities across Climate and Forest Age
The data used for these results are based on the 36 plots located in the three age-
classes along the climate gradient. In total 130 species of plants ≥10 cm DBH were
identified (818 individuals) and 132 species of plants 3≤ x <10 cm DBH (792
individuals).
Overall dissimilarity of communities among climate groups in this dataset was not
significant, for woody plants ≥10 cm DBH (RANOSIM = 0.284, p = 0.089) nor for woody
plant communities 3≤ x <10 cm DBH (RANOSIM = 0.218, p = 0.175). The largest
difference is, as expected, between the dry and the humid climate group (≥10 cm
DBH RANOSIM = 0.63, p = 0.1 and 3≤ x <10 cm DBH RANOSIM = 0.444, p = 0.1).
20
Differences in plant communities in the gradient of forest disturbance
Overall variation in woody plant communities among the three forest age-classes
across the climate zones shows significant dissimilarities. They were slightly larger in
plant communities 3≤ x <10 cm DBH (RANOSIM = 0.627, p = 0.001) than for plants ≥10
cm DBH (RANOSIM = 0.547, p = 0.001).
Considering each climate zone as an independent unit to test the variation among
forest age-classes, larger dissimilarities of plant communities ≥10 cm DBH were
found in the dry climate zone and the humid climate zone (Table 3). The two most
dissimilar plant communities in the size class 3≤ x <10 cm DBH were found in the
medium climate zone and the dry climate zone respectively.
Table 3. Dissimilarity (RANOSIM) and significance levels for woody plant communities among and
between forest age-classes in each climate zone. Old-growth forest is abbreviated with mf (mature
forest).
Moreover, the results indicate that the woody plant communities of the two secondary
forests are more similar to each other than to the mature forests. There is one
exception; the secondary humid forests in the size class 3≤ x <10 cm DBH (RANOSIM =
0.49, p = 0.029) is more dissimilar than the combination of old secondary and mature
forest (RANOSIM = 0.46, p = 0.057).
Climate zones Dry Medium Humid
Diameter class 3≤ x <10 cm ≥10 cm 3≤ x <10 cm ≥10 cm 3≤ x <10 cm ≥10 cm
Statistics RANOSIM p RANOSIM p RANOSIM P RANOSIM p RANOSIM p RANOSIM p
Overall dissimilarity 0.67 0.001 0.656 0.001 0.752 0.001 0.436 0.001 0.595 0.001 0.55 0.001
5-10 yrs /10-30 yrs 0.458 0.029 0.422 0.029 0.320 0.057 0.25 0.057 0.49 0.029 0.323 0.667
5-10 yrs/mf 0.807 0.029 0.885 0.029 0.99 0.029 0.625 0.029 0.734 0.029 0.667 0.029
10-30 yrs/mf 0.74 0.029 0.542 0.029 1 0.029 0.49 0.029 0.464 0.057 0.771 0.029
21
4.1.2 Diversity
Diversity and distribution of vegetation in old-growth forests over a climate gradient
In the sample set of 96 plots in mature forests, the number of woody plant species in
both size classes was highest in medium stature semi-evergreen forest, located in the
medium climate zone.
0
20
40
60
80
100
120
Dry Medium Wet0
20
40
60
80
100
Dry Medium Humid
Num
ber o
f pla
nt s
peci
es 3
-10
cm D
BH
Num
ber o
f pla
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peci
es >
10 c
m D
BH1
Shared species Medium and Humid
Shared species Dry and Humid
Shared species Dry and Medium
Uniques
Shared species all localities
4147
3330
24
42
83
101
7866
83
64
Figure 5. Total number of plant species ≥3 x <10 cm (left) and ≥10 cm DBH (right) in old-growth
forests of the Calakmul region.
The medium climate zone shows clearly the largest number of shared species >10 cm
DBH. For species 3≥ x <10 cm DBH the humid and the medium climate zone contain
the same number. Besides, the medium climate zone harbours most unique species
(approximately 50% of all plant species 3≥ x <10 cm DBH and 40% of plant species
≥10 cm DBH). Moreover, dry forests contained the smallest number of species
(Figure 5).
The woody plants in 96 plots in old-growth forest on the climate gradient were further
examined with sample-based and individual-based rarefaction curves (Figures 6 and
7). The sample-based curves have the advantage that they give an indication if enough
samples have been taken to characterize a community (Figure 6.1 and 6.2).
22
Number of individuals
0 200 400 600 800 1000 1200 1400
Num
ber o
f pla
nt s
peci
es >
10 c
m D
BH
0
20
40
60
80
100
120
Table 4. Species number based on sample-based (Sb) (figure 6.1 & 6.2) and individual- based (Ib)
rarefaction curves (7.1 & 7.2), and their standard deviation (SD).
Although the curves do not reach the asymptote, they have leveled to a reasonable
level to obtain an acceptable impression of which is the richer old-growth forest. The
Figures 6. Sampled-based rarefaction curves for plants 3≤ x <10 cm DBH in samples of 100 m2 (figure 6.1)
and plants ≥10 cm DBH in samples of 500 m2 (figure 6.2) in the mature forests of the Calakmul region.
Figures 7. Individual-based rarefaction curves (figure 7.1) for
plants 3≤ x <10 cm DBH and plants ≥10 cm DBH
(figure 7.2) in mature forests.
Climate zones Dry Medium Humid Statistics Sb SD Ib SD Sb SD Ib SD Sb SD Ib SD 3≤ x <10 cm 64 2.88 60 2.79 83 4.53 76 4.27 66 5.5 66 5.5 ≥10 cm 78 4.67 77 4.63 101 4.87 88 4.51 83 4.87 83 4.87
Number of samples
0 5 10 15 20 25 30 35
Num
ber
spe
0 cm
DBH
cies
3-1
of p
lant
0
20
40
60
80
100
Number of samples
0 5 10 15 20 25 30 35
Num
ber o
f pla
nt s
peci
es >
10 c
m D
BH
0
20
40
60
80
100
120
Fig. 6.1 Fig. 6.2
Number of individuals
0 100 200 300 400 500 600 700
Num
ber o
f pla
nt s
peci
es 3
-10
cm D
BH
0
20
40
60
80
100
Fig. 7.2 Fig. 7.1
High stature semi-evergreen forestMedium stature semi-evergreen forestLow stature semi-deciduous forest
Num
ber o
f pla
nt s
peci
es >
10 c
m D
BH
23
curves suggest that with increasing sample effort the medium semi-evergreen forest in
the medium climate zone is the richest and the dry one is the least rich, although the
difference between dry and humid forest is not certain given the variation.
In the individual-based rarefaction curves, the number of woody plant species that
occur in high stature semi-evergreen forest represents the number density on which
richness can be compared across the three forest types (Figure 7.1 and 7.2). The
results are similar to the sample-based rarefaction, showing that the medium stature
semi-evergreen forest seems to maintain more species than the two other forest types.
The plotted Clench models illustrate that the plant communities of the old growth
forests seem sufficiently sampled to make a meaningful comparison. The richer forest
is again the medium climate zone (Figure 8). However, the t-test executed with the
Clench values did not show significant differences between plant diversity of the
forest types and we ought to assume that the groups have equal variances.
0
20
40
60
80
100
120
0 800 1600 2400 3200 4000 4800 5600
No. individuals
No.
spe
cies
High stature semi-evergreen forestMedium stature semi-evergreen forestLow stature semi-deciduous forest
Figure 8. Accumulation curves based on the individual-based richness of woody plant species ≥10 cm
DBH, calculated according to the Clench model. The vertical line indicates the maximum number of
individuals to which points had been interpolated, meaning that the sampling effort and the number of
expected species reached until this point; beyond this line are extrapolations using the Clench model.
24
Plant species richness on the gradient of forest disturbance
Plant richness derived from sample- and individual-based rarefaction methods reflect
similar patterns, however, high values for sample based diversity are generally
suppressed compared to individual-based rarefaction (Table 5 and see Appendix 1, 2,
3, 5 for the curves). Individual-based plant richness in the smaller diameter class (3≤ x
<10 cm DBH) in the medium climate zone shows an increase of plant species along a
gradient of forest age. In the two other climate zones woody plant richness of the
young and the mature forests are equal or similar.
Climate zones Dry Medium Humid Statistics Sb SD Ib SD Sb SD Ib SD Sb SD Ib SD 3≤ x <10 cm DBH Forest 5-10 yrs 21 2.52 21 2.52 17 3.6 14 3.00 21 3.74 17 3.12 Forest 10-30 yrs 30 4.66 21 3.72 21 3.11 21 3.11 37 3.58 25 2.70 Mature forest 20 2.49 18 2.3 35 5.32 35 5.32 22 3.39 21 3.74 ≥10 cm DBH Forest 5-10 yrs 9 2.39 9 2.00 nd nd nd nd 12 3.02 12 3.02 Forest 10-30 yrs 36 5.02 12 1.84 15 2.4 15 2.4 19 2.81 5 1.46 Mature forest 31 2.94 16 2.37 49 4.78 29 2.88 38 4.16 10 1.90 ≥3 cm DBH Forest 5-10 yrs 28 3.02 28 3.02 17 3.12 17 3.12 29 4.69 29 4.69 Forest 10-30 yrs 54 5.96 28 3.50 30 3.99 25 3.50 47 3.81 30 2.73 Mature forest 39 3.84 28 3.03 68 5.72 45 4.14 49 4.32 35 3.50
Table 5. Rarefied plant species richness for different size classes in 36 plots in different age-classes
based on sample-based (Sb) and individual- based (Ib) rarefaction curves and their standard deviation
(SD). Nd means no data: EstimateS was not able to calculate richness parameters in the four young
secondary forest plots of the medium climate zone; here, only two tree species ≥10 cm DBH were
found. Note that rarefaction does not account for across climate comparison; these data are exclusively
for across forest age comparison within one climate zone!
For plants with a diameter ≥10 cm DBH in the dry and the medium climate, there is
an increase of individual-based rarefied richness with forest age. In the humid zone,
25
the sample-based richness does shows an increase in species with age, but not when
rarefied with the number of individuals (Table 5).
Individual-based plant richness considering both size classes (plants ≥3 cm DBH)
shows an increase of species with age in both the medium and the humid climate
zone. In the dry climate, plant species numbers are equal in both young and old forest
types (Appendix 4).
The results of the Simpson index shows that in particular the forests in the medium
climate zone show that the chance that two random woody plant species individuals
drawn from the population in mature forests is much higher compared to the
secondary forests (Table 6 and Appendix 6).
Table 6. Simpson estimators in forest age-classes in each climate zone. The calculation could not be
executed for plants ≥10 cm DBH in the wet climate zone, because one plot did not contain plants in this
size class.
Contrastingly, index values for woody plant species 3≤ x 10 cm DBH in the dry as in
the humid climate zone and the index values for plants ≥3 cm DBH in the dry climate
Climate zones Dry Medium Humid 3≤ x <10 cm DBH Forest 5-10 yrs 10.65 4.04 6.33 Forest 10-30 yrs 20.94 6.74 14.00 Mature forest 14.02 14.06 12.95 ≥10 cm DBH Forest 5-10 yrs 4.23 5.64 Nd Forest 10-30 yrs 19.57 5.64 5.17 Mature forest 21.53 18.78 14.23 ≥3 cm DBH Forest 5-10 yrs 14.21 4.64 8.00 Forest 10-30 yrs 27.19 8.37 8.57 Mature forest 23.49 24.42 20.58
26
zone present show higher species diversity in the old secondary forest compared to the
mature forest.
The vegetation communities in the forest mosaic
In order to compare the richness of the whole forest mosaic in each climate, species of
all forest age-classes were brought together (Figure 9). The medium forest samples
share the largest proportions of species found occurring in both dry and humid forests
and harbours in both size classes the lowest number of unique species.
Figure 9. Distribution of plant species 3≤ x <10 cm DBH and ≥10 cm DBH in 36 plots, with total species
number lumped for all forest age-classes.
0
20
40
60
80
100
Dry Medium Humid
Num
ber o
f spe
cies
plan
ts >
10 c
m D
BH
Shared species Medium and HumidShared species Dry and HumidShared species Dry and MediumUniquesShared species for all localities 0
20
40
60
80
Dry Medium Humid
Num
ber o
f spe
cies
plan
ts 3
-10
cm D
BH
27
Automeris spp. Saturniidae
4.2 Caterpillars
4.2.1 Community composition
Variation of Caterpillar Communities across Climate zones
From the 4200 individuals we identified 341 morpho-species, distributed over 5
butterfly families and 23 moth families. There is an overall significant variation
among caterpillar communities in the climate zones (global RANOSIM = 0.391, p = 0.011),
but there are no significant differences of paired climate groups, e.g. the largest
dissimilarity between communities was found in the combination of the dry and the
humid climate group (RANOSIM = 0.667, p = 0.1).
Differences between Caterpillar Communities across Forest age-classes
The caterpillar composition of three forest age-classes differ significantly across all
climate zones (RANOSIM = 0.449, P = 0.001). For each climate zone separately, significant
overall dissimilarities of caterpillar assemblages in the forest age-classes, were found
in the dry climate zone (RANOSIM = 0.495, p = 0.001), in the medium climate zone
(global RANOSIM = 0.266, p = 0.006) and in the humid climate zone (global RANOSIM =
0.128, p = 0.003) (Table 7). It shows that the dissimilarity of caterpillar assemblages
in the dry climate zone is most dissimilar and decreases along the dry to wet gradient.
For each climate zone separately, the most distinct caterpillar communities on the
gradient of forest age can be found between secondary forests (both young and old
secondary) and mature forests.
28
The caterpillar assemblages in ordinate space show that caterpillar community in the
dry climate zone is most dissimilar from the humid forests and the caterpillar
communities of the medium climate zone takes an intermediate position between
those two (Figure 10.1 and 10.2). In addition, the plots located in humid forest are
much more dispersed compared to the much more consistent cloud of samples located
in the dry forests (Figure 10.2).
Table 7. RANOSIM and significance levels for overall and pairwise differences in caterpillar communities.
Climate zones Dry Medium Humid
Statistics RANOSIM P RANOSIM P RANOSIM P
Overall dissimilarity 0.459 0.001 0.266 0.006 0.128 0.003
Forest 5-10 yrs / forest 10-30 yrs 0.51 0.029 0.135 0.171 0.011 0.373
Forest 5-10 yrs / mature forest 0.833 0.029 0.24 0.114 0.294 0.002
Forest 10-30 yrs / mature forest 0.198 0.114 0.469 0.029 0.083 0.082
Fig. 10.2
Fig. 10.1
Figure 10. Caterpillar assemblages of plots in ordinate
space (figure 9.1).On the right plots with the same
samples and equal projection as figure 9.1. The
difference consist of standardized symbols and grey
tones of sampled assemblages for the climate gradient
(figure 9.2) and forest age (figure 9.3).
Fig. 10.3
29
4.2.2 Diversity
Differences in Climate and Age class groups
Dry forestsShared species 10-30 and mfShared species 5-10 and mfShared species 5-10 and 10-30UniquesShared species all age classes
Most caterpillar species were found in the old secondary forests of the medium
climate zone. There is little difference between the absolute number of caterpillar
species among the mature forest types (Figure 11), but large differences age class 10-
30 years.
0
20
40
60
80
5_10 10_30 mf
Dry forests
Num
ber o
f spe
cies
0
20
40
60
80
5_10 10_30 mf
Medium forests
0
20
40
60
80
5_10 10_30 mf
Humid forests Figure 11. Absolute number of species, unique to an age class
within a climate group or shared with another age class in the
same climate group. Number above bars indicates total number
of species. Whiskers present the standard deviation for sampled
caterpillars in each forest age class group. Old-growth forest is
abbreviated with mf (mature forest).
Rarefied species richness of caterpillar communities does not express straightforward
patterns of how diversity changes along the forest-age gradient (Table 8 and
Appendix 5). The individual-based richness of the young secondary forest and the
mature forest are rather similar throughout climate zones. However, the old secondary
forest presents an extraordinary increase in species diversity from the dry to the humid
climate zone.
30
Table 8. Rarefied sample-based (Sb) and individual-based (Ib) species diversity. Note that rarefaction
does not account for across climate comparison; these data are exclusively for across forest age
comparison within one climate zone!
Significant differences between the caterpillar richness in the different forest age-
classes were only found in the dry climate zone; the dry young secondary forest and
old secondary forest (p=0.001) as well as the old secondary forest and the mature
forest (p<0.001).
The Simpson index shows similar patterns as found for individual based rarefaction
(Table 9 and Appendix 6). Also here, the chance that two random caterpillar species
drawn from the young or old secondary forests belong to different species is larger
than in mature forests.
Climate zones Dry Medium Humid Statistics Mean Mean Mean 5-10 yrs 6,11 11,25 8,99 10-30 yrs 2,45 7,17 16,92 mature forest 5,91 8,64 8,6
Table 9. Simpson index values in forest age-classes for each climate zone.
Climate zones Dry Medium Humid Statistics Sb SD Ib SD Sb SD Ib SD Sb SD Ib SD 5-10 yrs 54 3.80 45 3.48 63 3.66 61 3.66 48 4.52 32 3.33 10-30 yrs 25 3.98 17 2.84 73 5.79 48 4.26 51 4.02 51 4.02 mature forest 45 3.66 45 3.66 58 4.29 58 4.29 59 3.96 38 3.17
31
The caterpillar communities in the forest mosaic
The highest total caterpillar species number was found in the medium climate zone,
followed by the humid and dry climate zone respectively (Figure 12). Furthermore,
the medium climate zone shares the largest proportions of species found in both dry
and humid forests. With respect to unique species to a climate group, the humid zone
contains three species more than the medium forests.
020406080
100120140160180
Dry Medium Humid
Precipitation range (sites)
Num
ber o
f spe
cies
Figure 12. Absolute number of species, unique to a locality or shared with another. Number above bars
indicates total number of species.
Shared species Medium and HumidShared species Dry and HumidShared species Dry and MediumUniquesShared species for all localities
154 162
121
4.2.3 Nocturnal versus diurnal
The relative contribution of the occurrence and abundance of caterpillars that belong
to diurnal butterfly species showed that this group is rather small compared to the
nocturnal species throughout the sampled habitats (Table 10). The occurrence of
diurnal species is less than 10% and their abundances vary between 4 and 11%
compared to nocturnal species. Moreover, the abundance and the occurrence of
diurnal species are rather constant throughout the forest ages, which suggest that the
32
proportion of diurnal against nocturnal butterfly larvae does not change substantially
throughout the various habitats.
Climate zones Dry Medium Humid Statistics Occ (%) Ab (%) Occ (%) Ab (%) Occ (%) Ab (%) 5-10 yrs 6,1 8,8 4,7 5,9 7,1 10,7 10-30 yrs 6,0 8,6 4,0 6,1 7,8 10,7 mf 6,3 8,3 7,2 9,0 4,0 4,3
Table 10. Relative proportion of the number of diurnal species (Occ) and the relative abundances of
diurnal species (Ab) in relation to the total number and abundance of all species.
4.2.4 Equitability of the caterpillar communities
More than half of all species were found in low abundances (1 individual) in disturbed
as well as in undisturbed forests (Figure 13). There are no visible patterns in the
steepness of the curves representing on the one hand disturbed and undisturbed forests
on the other. In dry and medium forests Cisthene menea (Arctiidae) was particularly
abundant, and to a lesser extent also in humid forests. The highest ranked species in
the old growth humid forest is occupied by Chlosyne gaudealis (Nymphalidae).
Caterpillars with the higher ranks in all forest age groups belong to Pyralidae,
Elachistidae and Crambidae, of which the adults are small nocturnal species.
33
0
0,5
1
1,5
2
2,5
0 20 40
Range
Log
abun
danc
e ca
terp
illar
s
600
0,5
1
1,5
2
2,5
0 20 40 60 8
Range
Log
abun
danc
e ca
terp
illar
s
0
Dry MediumCis-men Cis-men
Pyr Pyr
0
0,5
1
1,5
2
2,5
0 20 40 60
Range
Log
abun
danc
e ca
terp
illar
s
Figure 13. Rank order log abundance plot for dry, medium and humid climate zone, each with the three
forest age-classes. The arrows exemplify the species Cisthene menea (Arctiidae) and Pyraloids in the
highest ranks.
4.2.5 Relationship between caterpillars and vegetation structure and richness
The diversity of plant species expressed in species number did not explain the
distribution of caterpillar diversity, neither did age or average height of the canopy of
the plots. With the unconstrained ordination DCA it was found that the first axis
explains 5% of the caterpillar species variability, and the first plus the second axis
reached 9.1% (Table 11). This is quite high given the species numbers registered
Humid 5-10 years10-30 yearsmature forest
Cis-men
Pyr
34
(Lepš and Šmilauer 2003), and this percentage accounts for more than 30 species of
the total number of caterpillar species registered.
1 2 3 4 Total inertia Eigenvalues 0.650 0.539 0.458 0.392 13.015 Lengths of the gradient 4.087 3.680 3.630 3.097 Cumulative variance of species data (%)
5.0 9.1 12.7 15.7
Sum of eigenvalues 13.015 Table 11. Results of the DCA analysis
The results of the CCA analysis indicate that 16.3% of the variability is explained by
all the variables (related to structure and richness of vegetation) studied, but just two
of them, number of species of all vegetation ≥3 cm DBH (Veg all) and height of the
canopy (Altura 3) were significant and contributed with 43% of the explained
variability (Figure 14, Table 12).
Table 12. Results of the CCA analysis for the environmental variables. The environmental variables at
plot level are canopy height (Altura 3), the number of woody plant species 3≤ x <10 cm DBH (Veg 3-
10), number of woody plant species ≥10 cm DBH (Veg 10), number of woody plant species ≥3 cm
DBH (Veg all) and forest age (Age).
Envir. variable F p Eigenvalue Accum. Variability explained (%) Altura 3 1.342 0.001 0.494 23.3 Veg all 1.181 0.023 0.452 43.7 Veg 10 1.136 0.105 0.415 63.2 Veg 3-10 1.071 0.251 0.441 82.3 Age 1.01 0.457 0.473 100
35
Figure 14. The species-enviromental
variables-samples triplot of CCA. Species
are represented by triangles, 1-32 circles
indicate plots and the arrows the
environmental variables.
-3 4
-46
AgeVeg 3-10
Veg .10
Veg all
Altura 3
1
2
3
4
5
6
78
9
10
11
12
13
14
15
16 17
18
19
20
21222324
25
2627
28
2930
31
32
33
34
35
36
36
5. Discussion
5.1 Changes in plant and caterpillar community structure on the gradient of
climate and forest age
The results show that the plant community structure is significantly different among
forests with different precipitation ranges. The strongest evidence is provided by the
results of the set of 96 samples which demonstrated overall and pairwise significant
differences of the three mature forest types selected. These findings in agreement with
previous observations that the medium sized semi-evergreen forest and high semi-
evergreen forest fall are floristically distinct (Martínez and Gallindo-Leal 2002, Peréz
Salicrup 2004). The same authors also argue that low semi-deciduous and medium
sized semi-evergreen forest contain similar species, and vary only in terms of
structure and abundance. Vester et al (2007) demonstrate how different structural
characteristics of low and medium sized old growth forest types are different from
high semi-evergreen forest. In disagreement with Martínez and Gallindo-Leal (2002)
and Peréz Salicrup (2004) the analyses in the work presented here, indicate that low
and medium sized old growth forest maintain also very distinct plant communities in
terms of species composition. The results show a high proportion of unique species
for each old growth forest type, moreover there is a small proportion of species that
occur in all old growth forests. The caterpillar species composition is significantly
different across the climate zones but not between pairwise combinations. The
absence of significance may be resolved with more sampling effort similarly to plant
communities.
37
We found overall significant dissimilarity among the woody plant communities of the
forest age-classes. Similarly the results show dissimilar caterpillar communities but
the separation was less compared to woody vegetation. At the same time, our data
show that dissimilarity in caterpillar communities between age-classes was strongest
in dry forests while the lowest in the humid climate zone. Vester et al. (2007) found in
the dry climate zone a much more marked reduction of butterfly diversity in the
gradient of disturbance compared to the medium climate zone. Our results and those
of Vester et al. (2007) findings can be related to the idea that changes in caterpillar
community structure along forest age and climate is characterized by an increase of
dissimilarity in caterpillar communities among forest age-classes from the humid
climate zone towards the dry climate zone. The cause for the stronger divergence of
caterpillar assemblages in dry forests along the gradient of forest age is perhaps
related to pronounced unbalanced abiotic conditions, and slower regeneration and
establishment of original plant species during succession that may affect the
caterpillar distribution. Some of the causes may be found in increased and more
variable temperatures, reduced soil water availability and susceptibility to fires,
commonly observed in secondary forests of the dry tropical zones (Nepstad 1999,
Harvey and Eastman 2005). Obviously, the described disturbances also play a role in
the recovery process of secondary forests in the humid areas, but to a lesser extent
because more frequent and long-lasting precipitation replenishes water losses leading
to the recovery of vegetation and more stable environmental conditions.
The gradual increase in overall dissimilarity between caterpillar communities of forest
age-classes from the humid climate to the dry zone, is not unambiguously supported
by the change in woody plant communities. Similar to the response of caterpillar
38
communities, there is evidence of a contrasting response of plant communities in the
dry (high dissimilarity) and the humid forest (low dissimilarity). However, the
medium climate zone accounts for the lowest dissimilarity for plant communities in
the diameter class ≥10 cm DBH but also the highest dissimilarity for plants
communities 3≤ x <10 cm DBH. Driving attributes in the succession and diversity of
tropical vegetation are variable in the gradients considered, which possibly obscure
the expected uniform response in woody plant communities. To illustrate some of
these aspects that vary along the precipitation gradient are nutrient cycling (Lawrence
2002, Santiago et al. 2005), seed and seedling growth (Ray and Brown 1995, Khurana
and Sing 2001) animal removal and dispersal of seed and seedlings (Uhl 1987), while
plot-to-plot variation could be attributed to timing of abandonment and land use
history (Chinea and Helmer 2003) and related presence of mycoryzza (Allen et al.
2003).
5.2 Plant and caterpillar diversity on the gradient of climate
It is generally suggested that the number of tree species per unit area increases with
rainfall and decreases with seasonality in tropical lowland forests (Gentry 1982, 1988;
Wright 1992, Specht and Specht 1993, Clinebell et al. 1995, Aplet et al. 1998).
Although not significant, we found a tendency which is not accordance with this
hypothesized increase i.e.; the number of woody plant species seems the highest in
medium semi-evergreen forest. Moreover, the number of unique species is also
highest in mature medium semi-evergreen forest.
Although not significant, the disturbed and undisturbed forests in the medium climate
zone maintain probably the highest caterpillar richness, in accordance with Pozo et al.
39
(2003) who found a comparable evidence for butterflies. The high caterpillar richness,
this is mainly due to the high caterpillar diversity found in the younger forest age-
classes. Pozo et al. (2003) also demonstrated that the proportion of shared butterfly
species with other climate zones was high, while unique butterfly species in each
forest type was a quarter of all species (Pozo et al. 2003). In the present study, the
proportion of unique caterpillar species was higher than the proportion of shared
species, especially in medium semi-evergreen forest (>50%). In the other climate
zones and forest age-classes the proportion of unique species was also larger than a
quarter. Although this result may be an artefact of insufficient sampling, it could
indicate that the importance of unique species for creating the differences among
Lepidoptera communities between the three distinguished climate zones may be
higher than generally was assumed. This difference can possibly be related to the
differences in selected species; butterflies can move from one forest patch to another,
whereas caterpillars are less mobile.
5.3 Does caterpillar diversity increase along a gradient of forest age?
The conventional perspective on the effect on anthropogenic disturbance is an overall
loss of biodiversity (e.g. Phillips 1997). The diversity of the caterpillar communities
of the Yucatán peninsula did not present a consistent response to disturbance.
Recently, butterfly diversity in forests patches of secondary forests were reported to
sustain higher diversity compared to mature forests in the Calakmul region (Vester et
al. 2007). This effect was attributed to the ‘source-sink relationship’ referring to the
dispersion of butterflies across forest ages in the entire forest mosaic. As mentioned,
we did not reveal clear patterns of caterpillar species diversity on the gradient of
forest age, and particularly the caterpillar diversity in the old secondary forest is
40
variable. Only the dry climate zone shows significant differences between caterpillar
communities that belong to different forest age-classes.
As we have shown already at the level of caterpillar community structure, it is clear
that habitat modification causes species composition change. Another measure of
caterpillar species composition change can be expressed by shared and unique species
distribution along forest age. Research on butterflies in the Calakmul zone showed
that about 60% of the butterfly species were shared between old-growth forest and
forest <10 years (Vester et al. 2007). In our work, the proportion of shared caterpillar
species is less than 10%, and points out the importance of unique species contributing
to the variation caterpillar diversity among habitats. In general it may be that the
number of unique species in our work is relatively higher because we also dealt with
nocturnal species, which considerably increases the chance of encountering unique
species.
The results suggest that primary forests in Calakmul do not present higher diversity
compared to secondary forests. In the light of the ecological theories explaining
diversity on a gradient of forest disturbance, this type of response could be related to
the intermediate disturbance theory (Connell 1977). Previously, it has been argued
before that decreases as well as increases of butterfly diversity in response to forest
disturbance can occur. For example, various studies showed that low disturbance
levels have a positive effects on the local butterfly diversity (Hill 1999, Lovejoy et al.
1986, Brown 1991, Wood and Gillman 1998) while other found a decrease of
butterfly and moth diversity (Bowman et al. 1990, Thomas 1991, Spitzer et al. 1993,
1997, Kremen 1994, Hamer and Hill 2000, Hill et al. 1995, 2001, Hill and Hamer
41
1998, Brown 1997, Lewis et al. 1998, Willott et al. 2000, Lewis 2000, Fermon et al.
2000, 2001).
Previous studies learn us that species (or guilds) have their own environmental
requirements and frequently lead to more sophisticated predictions instead of
considering Lepidoptera community as a whole. For example, the species composition
of Pyralidae did not change in disturbed and undisturbed sites but this family did
show different abundances (Fiedler and Schulze 2004). Similar, it has been
documented that the diversity of moths remained unchanged whereas butterfly
diversity alters at different levels of fragmentation (Daily and Ehrlich 1996). Also in
Borneo it appeared that Sphingidae do not reflect the human induced disturbance
(Schulze and Fiedler 2003). Despite a different response from nocturnal species
compared to diurnal species in various studies, our work showed that the proportion
of diurnal/nocturnal species is rather constant throughout young and old forests in the
Calakmul area. These issues make clear that it is worthwhile to opt for different
analytical measures for future habitat quality studies such as the use of indicator
families and target species, which could enhance the discriminatory power to identify
the responsiveness of Lepidoptera upon disturbance (Basset et al. 2004).
There may be also another effect that causes the rather high but variable caterpillar
diversity in the secondary forests. This effect is related is related to the abundance and
quality of host-plants in the secondary forests, which are characterized with few but
dominant plant species. To illustrate this relation, I will give two examples with the
Lepidoptera families, Hesperidae and Nymphalidae. A number of abundant species of
from the family of Hesperidae feed frequently on Cryosophila stauracantha, Sabal
japa and Chamedorea oblongata (Palmea), while Nymphalidae have often been found
42
on Croton arboreus (Euphorbiaceae), Lonchocarpus xuul (Leguminosae) and Hampea
trilobata (Malvaceae) (this is a personal observation). These plant species are very
abundant in young and disturbed forests (and dry forests). The diversity measures we
used do not account for high abundances of certain plant species as an explanatory
variable. Here, the use of caterpillar guilds related to plant growth form (Janz and
Nylin 1998, Cleary 2003) may yield clearer indicator groups. It leaves no doubt that
this aspect calls for studies to assess the above mentioned effects.
5.4 Does woody plant diversity increase along a gradient of forest age?
In general, it is likely that the disappearance of old growth forests will lead to a
decrease in organisms, particularly of those species which have restricted habitat
requirements (Thomas 1991). Our results indicate that ongoing conversion of mature
forests to secondary stands in the Calakmul region causes a steady decrease of trees
species ≥10 cm DBH in the medium climate zone. Also in the humid climate zone
there is a decrease of sample based species diversity of plant ≥10 cm DBH with
increasing forest-age.
When diversity is rarefied based on individuals, the interpolation limits the curves
causing serious loss of data, at the expense of old secondary and the primary forest.
More sampling effort is needed in the young secondary forest to express a pattern in
the humid climate zone. The rarefied curves of diversity in the dry climate zone are
still more precarious to interpret as the curves are crossing each other. The moment
the curves for two communities intersect, the ranking of the observed species richness
reverses and poses serious limitations in comparisons for species diversity (Lande et
al. 2000). Moreover, the interpolation, based on the young secondary forest species
43
diversity, leads also here to significant losses of data in the old secondary and primary
forest. With respect to plants 3≤ x <10 cm DBH in the dry forest, the problems
described above, applies here too.
It was intended to overcome problems related to smaller sampling sets using the
Simpson index. In general, the Simpson diversity index for plants and caterpillars
present similar response as the individual based rarefaction statistics. The most
apparent difference is that the Simpson index displays a stronger division between the
plant diversity in the secondary forests compared to the mature forests. The Simpson
index confirms the earlier findings of the individual based rarefaction, and supports
the hypothesized increase of plant species richness with increasing forest age,
particularly for the medium and the humid climate zone.
5.5 Caterpillar community equitability
Low levels of equitability suggest the dominance of disturbance or pollution tolerant
species (Favila and Halfter 1997, Floren and Linsenmair, 2001). We expected these
low levels of equitability in younger age-classes compared to mature forests in
Calakmul. This links up with the hypothesis that species diversity decreases with
decreasing forest-age, as equitability is a measure of how well species abundances are
distributed in the community. The rank order abundance curves did not expose
patterns of distinct curves of caterpillar communities in young forests compared to old
forests and the expected steeper of curves that represent caterpillar communities in
disturbed environments was not distinguished. Apparently, the resources in these
44
disturbed forests are sufficiently abundant and stress is sufficiently low to allow a
large diversity with a distribution of abundances similar to mature forests.
5.5 Relationships between caterpillars and vegetation structure and richness
Examination of the CCA did not show clear caterpillar species assemblages
concentrated around forest plots, nor the variables of plant diversity and forest
structure were evidently related to caterpillar distributions. It would be advisable to
repeat the exercise for each climate zone independently. Moreover, the selected
measures for plant diversity and structure are not suited to provide evidence for a
relation with caterpillar communities. The relation between plant and caterpillar
diversity may be clearer when taking into account the undergrowth, particularly herbs.
There are studies executed in south-east Asia forest plantations with high plant
species richness in the undergrowth turned out to support a significantly richer moth
family i.e.; geometrids (Intachat et al. 1997, 1999) as well as other moth (Chey et al.
1997) and butterfly faunas (Ghazoul, 2002) which is also an important family in our
study area.
45
Automeris banus Saturniidae
6. General conclusions
This work contributes to the present investigation on the effects of land use change on
biodiversity by comparing species richness and composition in secondary and primary
forests in the gradient from the dry to the humid climate zone. This work also adds to
current understanding on tropical insect biology, because knowledge on caterpillar
diversity is poor in the literature, even more if it is intended to explain the ecological
aspects involved. The medium climate zone appears to maintain a high diversity of
caterpillars and plants compared to the dry to and the humid climate, but statistical
analyses could not confirm this pattern. There is not enough evidence to support the
idea of a gradual loss of caterpillar diversity over the gradient of forest disturbance.
Instead, caterpillars communities in secondary forests can be highly diverse but the
variation between communities in forest age-classes is large and still more effort is
needed to identify the response of caterpillar communities. I believe that it is
important at this stage to interpret the tendencies rather than statistical differences
only to reflect gradual differences in the forest mosaic. Finally present and future
research on the adult stages of Lepidoptera, host plant interaction, species guilds and
the effect at different levels of the Lepidoptera community structure may enhance our
understanding.
46
Acknowledgements
This work is a product created at El Colegio de la Frontera Sur, Ecosur, Unidad
Chetumal and Wageningen University. I thank the Southern Yucatán Peninsular
Region (SYPR) project involving Clark University, the University of Virginia, El
Colegio de la Frontera Sur, and Rutger University. Its principal sponsors have been
NASA-LCLUC (Land Cover and Land Use Change) program (NAG5-6046 and
NAG5-11134, NNG06GD98G), the Center for Integrated Studies of the Human
Dimensions of Global Environmental Change, Carnegie Mellon University (NSF SBR
95-21914), and NSF-Biocomplexity (BCS-0410016). I would like to thank Hans
Vester, Frans Bongers and Carmen Pozo for reading and giving comments on my
thesis, as well as Jorge Montero and Hector Abuid Hernandez Arana for statistical
assistance. We owe our thanks to those people that have worked on the large set of
mature forest plots, Richard van Sluis en Maarten Debruyne, and specifically in the
ejido of Conhuas, Rafael Espinoza López, Jaime Haas Tzuc, Wilbert Poot Pool,
Daniel Poot Pool, William Naal Segovia, Hans van der Wal, financial aids for these
colleagues came from SEMARNAT in Plan de manejo de Balam Kin en Balam Ku. I
am also grateful to the people I have been working with both in the field and in the
laboratory, Euridice Leyequien Abarca, Angeles Islas Luna, Miguel Xijun, Emigdio
May Uc, José and Magarito and local field guides that have played an invaluable role
for their knowledge Don Magdaleno of the ejido Conhuas, Don Diego of the ejido of
Dos Lagunas, Don Nico of the ejido Nuevo Becal and Don Eliseo of the ejido Zoh
Laguna.
47
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Appendix Appendix 1 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Dry climate zone. Appendix 2 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Medium climate zone. Appendix 3 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Humid climate zone. Appendix 4 Individual-based species richness curves for caterpillars in the forest age-classes in the climate zones. Appendix 5 Rarefied diversity for plants and caterpillars in the forest age classes Appendix 6 Simpson index for plants and caterpillars in the forest age classes Appendix 7 Clench model for caterpillars in the forest age classes in each climate zone
Appendix 1 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Dry climate zone.
0 1 2 3 4 50
10
20
30
40
50
60
70
0 1 2 3 4 50
10
20
30
40
50
Num
ber o
f spe
cies
P
lant
s ≥3
cm
DB
H
0 1 2 3 4 50
10
20
30
40
Num
ber o
f spe
cies
P
lant
s ≥3
- <1
0 cm
DBH
Num
ber o
f spe
cies
P
lant
s ≥1
0 cm
DB
H
Samples Samples 5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
0 20 40 60 80 100 120 140 160 180 2000
10
20
30
40
50
60
70
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
Individuals Individuals0 20 40 60 80 100 120
0
10
20
30
40
50
Num
ber o
f spe
cies
P
lant
s ≥3
cm
DB
H
0 20 40 60 800
10
20
30
40
Num
ber o
f spe
cies
P
lant
s ≥3
- <1
0 cm
DBH
Num
ber o
f spe
cies
P
lant
s ≥1
0 cm
DB
H
Appendix 2 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Medium climate zone.
0 1 2 3 4 50
10
20
30
40
50
60
Num
ber o
f spe
cies
P
lant
s ≥3
cm
DB
H
0 1 2 3 4 50
10
20
30
40
50
60
70
80
0 1 2 3 4 50
10
20
30
40
50
10-30 yrsmf
Samples Samples Samples 5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
Individuals 0
Individuals 20 40 60 80 100 120 140
0
10
20
30
40
50
5-10 yrs 10-30 yrsmf
Individuals
0 20 40 60 80 100 120 140 160 180 2000
10
20
30
40
50
60
10-30 yrsmf
Individuals
0 50 100 150 200 250 3000
10
20
30
40
50
60
70
80
Appendix 3 Sample-based and individual-based species richness curves for plants in the forest age-classes in the Humid climate zone.
0 1 2 3 4 50
10
20
30
40
50
60
0 1 2 3 4 50
10
20
30
40
50
Num
ber o
f spe
cies
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lant
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- <1
0 cm
DBH
Num
ber o
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cies
P
lant
s ≥1
0 cm
DB
H
0 1 2 3 4 50
10
20
30
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50
Num
ber o
f spe
cies
P
lant
s ≥3
cm
DB
H
Samples Samples 5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
Samples 5-10 yrs 10-30 yrsmf
0 20 40 60 80 100 120 1400
10
20
30
40
50
Num
ber o
f spe
cies
P
lant
s ≥3
- <1
0 cm
DBH
0 20 40 60 80 100 120 140 1600
10
20
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40
50
0 50 100 150 200 250 3000
10
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60
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cies
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lant
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cm
DB
H
Num
ber o
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lant
s ≥1
0 cm
DB
H
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
5-10 yrs 10-30 yrsmf
Individuals IndividualsIndividuals
Appendix 4 Individual-based species richness curves for caterpillars in the forest age-classes in the climate zones.
0 100 200 300 400 500 600 700
Individuals
Num
ber o
f cat
erpi
llar s
peci
es
0
80
20
40
60
0 100 200 300 400 500 600 7000
20
40
60
80
0 100 200 300 400 500 600 7000
20
40
60
80
5-10 years10-30 yearsmature forest
Individuals 5-10 years10-30 yearsmature forest
Humid forestsMedium forests
5-10 years10-30 yearsmature forest
Dry forests
Num
ber o
f cat
erpi
llar s
peci
es
Individuals
Individual-based species richness curves for the age classes within each of the three types of climate. The arrows indicate the limiting curve for rarefying richness in the
other habitats. In the graph of Humid forests an example is given how rarefied richness is read from the graph.
Appendix 5 Rarefied diversity for plants and caterpillars in the forest age classes
Rarefied diversity Dry climate zone Medium climate zone Humid climate zone
Sample based
Indiv. based
Sample based
Indiv. based
Sample based
Indiv. based Plant diversity Diameter class ≥3x<10 cm DBH
No
of S
peci
es
0204060
5-10 yrs 10-30 yrs mf
0102030
5-10 yrs 10-30 yrs mf
020406080
5-10 yr s 10-30 yr s mf
0
20
40
60
5-10 yrs 10-30 yrs mf
0
20
40
60
5-10 yrs 10-30 yrs mf
010203040
5-10 y rs 10-30 yrs mf
Diameter class ≥10 cm DBH
No
of S
peci
es
0204060
5-10 yrs 10-30 yrs mf
0102030
5-10 yrs 10-30 yrs mf
020406080
10-30 yr s mf
0
20
40
60
10-30 y rs mf
0
20
40
60
5-10 yrs 10-30 yrs mf
010203040
5-10 y rs 10-30 yrs mf
All woody species ≥3 cm DBH
No
of S
peci
es
0204060
5-10 yrs 10-30 yrs mf
0102030
5-10 yrs 10-30 yrs mf
020406080
5-10 yrs 10-30 yrs mf
0
20
40
60
5-10 yrs 10-30 yrs mf0
20
40
60
5-10 yrs 10-30 yr s mf
010203040
5-10 yrs 10-30 yrs mf
Caterpillar diversity
No
of S
peci
es
0
20
40
60
5-10 yrs 10-30 yrs mf
0
20
40
60
5-10 y rs 10-30 yrs mf
0
20406080
5-10 y rs 10-30 y rs mf0
20406080
5-10 yrs 10-30 yrs mf
0
20
40
60
5-10 yrs 10-30 yrs mf
0
20
40
60
5-10 yrs 10-30 yrs mf
Appendix 6 Simpson index for plants and caterpillars in the forest age classes
Simpson Index Dry climate zone Medium climate zone Humid climate zone Plant diversity Diameter class ≥3x<10 cm DBH
No
of S
peci
es
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
5-10 year s 10-30 year s mf
Diameter class ≥10 cm DBH
No
of S
peci
es
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
10-30 year s matur e f or est
All woody species ≥3 cm DBH
No
of S
peci
es
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
5-10 year s 10-30 year s mf
05
1015202530
5-10 year s 10-30 year s mf
Caterpillar diversity
No
of S
peci
es
0
5
10
15
20
5-10 year s 10-30 year s mf
0
5
10
15
20
5-10 year s 10-30 year s mf
0
5
10
15
20
5-10 year s 10-30 year s mf
Appendix 7 Clench model for caterpillar species richness in the forest age classes in each climate zone
Dos Lagunas
0
20
40
60
80
100
0 300 600 900 1200 1500
No. individuals
No. s
peci
es
5-10 yrs10-30 yrsMf
Humid Naadzkan
0
20
40
60
80
0 300 600 900 1200 1500 1800
No. individuals
No. s
peci
es 5-10 yrs10-30 yrsMf
Dry
Nuevo Becal
0
20
40
60
80
100
0 300 600 900 1200 1500 1800
No. individuals
No.
spe
cies 5-10 yrs
10-30 yrsMf
Medium
Climate zones Dry Medium Humid Statistics 5-10 yrs 46 % 56 % 35 % 10-30 yrs 35 % 36 % 57 % mature forest 60 % 52 % 37 %
The percentage of registered species with respect to the
total number of expected species (asymptote).