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\begin{document}

\title{Edge disturbance shapes liana diversity and abundance but not liana-tree
interaction network patterns in moist semi-deciduous forests, Ghana~~~ ~}



\author[1]{Bismark Ofosu-Bamfo}%
\author[2]{Patrick Addo-Fordjour}%
\author[2]{Ebenezer Belford}%
\affil[1]{University of Energy and Natural Resources, Sunyani, Ghana}%
\affil[2]{Kwame Nkrumah University of Science and Technology}%


\vspace{-1em}



  
  \date{June 17, 2024}


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\selectlanguage{english}
\begin{abstract}
Edge disturbance can drive liana community changes and alter liana-tree
interaction networks, with ramifications for forest functioning.
Understanding edge effects on liana community structure and liana-tree
interactions is therefore essential for forest management and
conservation. We evaluated the response patterns of liana community
structure and liana-tree interaction structure to forest edge in two
moist semi-deciduous forests in Ghana (Asenanyo and Suhuma Forest
Reserves: AFR and SFR, respectively). Liana community structure and
liana-tree interactions were assessed in 24 50 \selectlanguage{ngerman}× 50 m randomly located
plots in three forest sites (edge, interior and deep-interior)
established at 0--50 m, 200 m and 400 m from edge. Edge effects
positively and negatively influenced liana diversity in forest edges of
AFR and SFR, respectively. There was a positive influence of edge
disturbance on liana abundance in both forests. We observed anti-nested
structure in all the liana-tree networks in AFR, while no nestedness was
observed in the networks in SFR. The networks in both forests were less
connected, and thus more modular and specialised than their null models.
Many liana and tree species were specialised, with specialisation
tending to be symmetrical. The plant species played different roles in
relation to modularity. Most of the species acted as peripherals
(specialists), with only a few species having structural importance to
the networks. The latter species group consisted of connectors
(generalists) and hubs (highly connected generalists). Some of the
species showed consistency in their roles across the sites, while the
roles of other species changed. Generally, liana species co-occurred
randomly on tree species in all the forest sites, except edge site in
AFR where lianas showed positive co-occurrence. Our findings deepen our
understanding of the response of liana communities and liana-tree
interactions to forest edge disturbance, which are useful for managing
forest edge.

\par\null%
\end{abstract}\selectlanguage{ngerman}%



\sloppy




\textbf{Edge disturbance shapes liana diversity and abundance but not
liana-tree interaction network patterns in moist semi-deciduous forests,
Ghana}

\textbf{Running title:}

\textbf{Edge effects on lianas and liana-tree interactions}

\textbf{Bismark Ofosu-Bamfo\textsuperscript{1}, Patrick
Addo-Fordjour\textsuperscript{2*}, E.J.D. Belford\textsuperscript{2}}

\textsuperscript{1}Department of Basic and Applied Biology, School of
Sciences,~University of Energy and Natural Resources, Sunyani, Ghana

\textsuperscript{2}Department of Theoretical and Applied Biology,
Faculty of Biosciences, College of Science, Kwame Nkrumah University of
Science and Technology, Kumasi, Ghana

*Corresponding author: paddykay77@yahoo.com;
paddofordjour.cos@knust.edu.gh

ABSTRACT

Edge disturbance can drive liana community changes and alter liana-tree
interaction networks, with ramifications for forest functioning.
Understanding edge effects on liana community structure and liana-tree
interactions is therefore essential for forest management and
conservation. We evaluated the response patterns of liana community
structure and liana-tree interaction structure to forest edge in two
moist semi-deciduous forests in Ghana (Asenanyo and Suhuma Forest
Reserves: AFR and SFR, respectively). Liana community structure and
liana-tree interactions were assessed in 24 50 \selectlanguage{ngerman}× 50 m randomly located
plots in three forest sites (edge, interior and deep-interior)
established at 0--50 m, 200 m and 400 m from edge. Edge effects
positively and negatively influenced liana diversity in forest edges of
AFR and SFR, respectively. There was a positive influence of edge
disturbance on liana abundance in both forests. We observed anti-nested
structure in all the liana-tree networks in AFR, while no nestedness was
observed in the networks in SFR. The networks in both forests were less
connected, and thus more modular and specialised than their null models.
Many liana and tree species were specialised, with specialisation
tending to be symmetrical. The plant species played different roles in
relation to modularity. Most of the species acted as peripherals
(specialists), with only a few species having structural importance to
the networks. The latter species group consisted of connectors
(generalists) and hubs (highly connected generalists). Some of the
species showed consistency in their roles across the sites, while the
roles of other species changed. Generally, liana species co-occurred
randomly on tree species in all the forest sites, except edge site in
AFR where lianas showed positive co-occurrence. Our findings deepen our
understanding of the response of liana communities and liana-tree
interactions to forest edge disturbance, which are useful for managing
forest edge..

Keywords: co-occurrence patterns, ecological networks, edge influence,
liana diversity and abundance, modularity, nestedness, specialisation

INTRODUCTION

In tropical forests, particularly in developing economies, human
interactions with forest ecosystems remain an important source of change
in their structure (FAO \& UNEP, 2011) and functioning (Pedro et al.,
2015). A common outcome of human interface with forest ecosystems is
fragmentation which results in the creation of edges (Harper et al.,
2005). Edge mediated microclimatic changes may favour
disturbance-adapted, light-loving species such as lianas (see Hawthorne,
1996; Laurance et al., 2001), but generally be disadvantageous to others
such as trees (Laurance et al., 2006). Previous studies reported that
edge effects enhanced liana diversity and abundance in some forests
(Laurance et al., 2001; Campbell et al., 2018; Ofosu-Bamfo et al.,
2019), but others did not detect changes in liana diversity in response
to edge (Mohandass et al., 2014; Ofofu-Bamfo et al., 2019). Several
properties of forest edge such as edge size, edge type, and surrounding
matrix type can mediate edge effects on plant community structure
(Martino, 2015), and be responsible for the varied responses of
community structure to edges in different forests. As liana community
assemblages respond to edge disturbance, the relationship between lianas
and trees may also be altered. Fagan et al. (1999) stated that habitat
edges can modify species interactions. Similarly, Porensky (2011)
reported that species interactions show strong responses to forest edge.
Nonetheless, there is scarcity of information on the response of
liana-tree interaction network patterns to forest edge.

The knowledge of liana-tree interactions and the factors that shape them
are key to fully understanding plant community composition and
structure. Although different patterns of liana-tree interactions have
been reported in literature, there is no consensus yet. For example,
nestedness, a network pattern in which the interactions of less
connected species form proper subsets of the interactions of more
connected species (Bascompte et al., 2003; Landi et al., 2018; Ponisio,
et al. 2019), has been used to characterise the structure of liana-tree
networks. Different patterns of nestedness are reported in literature
including nested (Sfair et al., 2010) and non-nested (Addo-Fordjour \&
Afram, 2021; Addo-Fordjour et al., 2016; Blick \& Burns, 2009; Magrach
et al., 2015; Ofosu-Bamfo et al., 2019) structures. Among the studies
that did not find nested structure in liana-tree netowrks, some reported
anti-nested structure which depicts non-random assembly (Addo-Fordjour
\& Afram, 2021; Blick \& Burns, 2009; Magrach et al., 2015), whiles
others observed non-significant nestedness that shows random assembly
(Addo-Fordjour et al., 2016; Ofosu-Bamfo et al., 2019). Ecological
networks can also be compartmentalised into modules whose members
interact more among themselves (Carstensen et al., 2016). This
phenomenon referred to as modularity, is predicted to stabilise
ecological networks (Massol et al., 2017; Thébault \& Fontaine, 2010).
For this reason, modularity analysis of liana-tree networks can have
implications for forest management and conservation. Nonetheless, only a
few studies assessed the patterns of modularity in liana-tree networks.
Sfair et al. (2015) did not find modular structure in their networks,
but Addo-Fordjour \& Afram (2021) recorded significant modular structure
in liana-tree networks.

Specialisation at the network and species levels can cause non-nested
and modular organisation of species (Addo-Fordjour \& Afram, 2021; Médoc
et al., 2017; Castledine et al., 2020). Thus, in liana-tree networks in
which coevolution leads to specialisation (Sfair et al., 2015), the
networks may tend to be non-nested and/or modular. Another important
metric used to characterise network structure is species co-occurrence,
which describes the frequency of pairs of liana species to co-occur on
the same phorophyte species (Zulqarnain et al., 2016). Like the
above-mentioned network metrics, mixed patterns of liana species
co-occurrence have been reported in literature, which include positive
co-occurrence (Addo-Fordjour et al., 2016; Zulqarnain et al., 2016),
negative co-occurrence (Blick \& Burns, 2011, 2009), and random
co-occurrence (Addo-Fordjour et al., 2016). With the mixed findings on
the structure of liana-tree interactions in literature, there is the
need for more studies to be conducted to determine the most consistent
patterns. Knowledge of co-occurrence patterns is important for
increasing our understanding of species interactions and predicting
community stability and maintenance, and ecosystem functioning, all of
which may be useful in forest conservation (Vizentin-Bugoni et al.,
2016).

The current study determined the response of liana community assemblages
and the patterns of structure of liana-tree interaction networks to edge
in two moist semi-deciduous forests in Ghana. The forest edges we
studied were surrounded by large matrices of crop farmlands, thus making
the edges much exposed. The nature and size of land matrix bordering
forest edges play a key role in determining the intensity of edge
effects on plant community structure (Arag\selectlanguage{greek}ό\selectlanguage{english}n et al., 2015). To this end,
edges bordered by wide land matrices are expected to exert stronger
effects on plant communities than edges surrounded by narrow area of
land (Addo-Fordjour \& Owusu-Boadi, 2016). Furthermore, because the
nature of the land matrix surrounding our forest edges is
physiognomically dissimilar to the forest vegetation, the microclimate
variation between the forest edge and interior may be enhanced (Arag\selectlanguage{greek}ό\selectlanguage{english}n
et al., 2015). Thus, edge effects on lianas and liana-tree interaction
patterns in the two moist semi-deciduous forests may be apparent. Edge
disturbance permits greater penetration of sunlight into forest edges,
and also increases forest edge dryness (Thier \& Wesenberg, 2016), both
of which can favour liana proliferation. On the basis of the above, we
tested the following hypotheses:

\begin{enumerate}
\tightlist
\item
  Liana diversity and abundance would be higher in edge site than
  non-edge sites.
\item
  We expected that as edge disturbance enhances liana abundance at the
  forest edge, network connectance will increase, resulting in less
  specialised, nested, and non-modular network structures in edge site,
  while the networks in the non-edge sites will be less connected, more
  specialised, non-nested, and modular.
\item
  Edge effects will cause shifts in topological roles of liana and tree
  species due to changes in the distribution and abundance of the
  species.
\item
  As sunlight and dry conditions are elevated at edge sites relative to
  the non-edge sites, competition of lianas for the resources in edge
  site may be lower. Moreover, as edge effects tend to cause tree
  mortality at forest edges (Murcia 1995), the number of available host
  species may reduce, increasing liana infestation per host. Thus, we
  expected that liana species in edge sites would show positive
  co-occurrence on host trees, while the species in non-edge sites will
  randomly co-occur on their hosts.
\end{enumerate}

The findings of our study would be useful in the management of forest
edges and conservation of edge species. Our study seeks to add valuable
information to literature, thus helping to obtain general patterns of
liana assemblages and structure of liana-tree interactions in relation
to edge effects. The findings would also contribute to the development
of a robust edge theory in view of the fact that there is dearth of
information on the role of edge disturbance in shaping the patterns of
liana-tree network structure in forests.

METHODOLOGY

Study areas

We conducted the study in two moist semi-deciduous tropical forest
ecosystems in Ghana: Asenanyo Forest Reserve (latitudes
6\textsuperscript{o}17' and 6\textsuperscript{o}36'N; longitudes
1\textsuperscript{o}50' and 2\textsuperscript{o}16'W) and Suhuma Forest
Reserve (latitudes 5\selectlanguage{ngerman}°56\selectlanguage{english}' and 6\selectlanguage{ngerman}°11\selectlanguage{english}'N; longitudes 2\selectlanguage{ngerman}°21\selectlanguage{english}' and 2\selectlanguage{ngerman}°36\selectlanguage{english}'W).

\emph{Asenanyo Forest Reserve}

The Asenanyo Forest Reserve is a production forest that was established
in the year 1940 and covers an area of 22,800 ha in the Ashanti Region
of Ghana (Wiafe, 2014). It is of the moist semi-deciduous forest
ecosystem, with the dominant tree species being \emph{Celtis
mildbraedii} , \emph{Triplochiton scleroxylon} ,
\emph{Entandrophragma}spp. and \emph{Thaumatococus} spp. (Wiafe, 2014;
Forest Services Division, 2010a). The forest has a bimodal rainy season
from April to October (maximum rainfall: May-June; minimum rainfall:
September-October) and a dry season from November to March. Annual
rainfall range is 1250--500 mm (Hall \& Swaine, 1981). Temperature in
the reserve ranges from an average high of 30.5\textsuperscript{o}C to
21\textsuperscript{o}C, with a mean annual relative humidity of about
84\%. Asenanyo Forest Reserve has about 20 admitted farms scattered
throughout the reserve, the size of each averaging approximately 5 ha
(Forest Services Division, 2010a). The reserve also has one admitted
community occupying an area of about 955.70 ha (Forest Services
Division, 2010a). Threats faced by the reserve include illegal logging,
charcoal production, illegal farming in the reserve, surface mining,
sand wining and hunting (Wiafe, 2014; Forest Services Division, 2010a).
These activities are accompanied by unauthorized development of in-roads
and clearing of forest.

\emph{Suhuma Forest Reserve}

The Suhuma Forest Reserve (latitudes 5\selectlanguage{ngerman}°56\selectlanguage{english}' and 6\selectlanguage{ngerman}°11\selectlanguage{english}'N; longitudes 2\selectlanguage{ngerman}°21\selectlanguage{english}'
and 2\selectlanguage{ngerman}°36\selectlanguage{english}'W) is also a production forest of about 36,030 ha located in
the Sefwi Wiawso Forest District (Hawthorne \& Abu-Juan, 1995). There
are 24 admitted farms in the reserve each averaging 11.5 ha (total 276
ha) and one admitted community covering an area of 389 ha (Forest
Services Division, 2010b). The reserve is exposed to active logging. Its
canopy is discontinuous due to excessive logging activity but still has
emergent trees that may reach heights of about 40 m. Most of the trees
that occur in the reserve are deciduous: (Forest Services Division,
2010b). The forest lies within the moist semi-deciduous forest zone in
Ghana, and thus its vegetation is dominated by tree species such
as\emph{C} . \emph{mildbraeddii} , \emph{Baphia nitida} ,
\emph{Nesogordonia papaverifera} , \emph{Microdesmis puberula} ,
\emph{Khaya ivoriensis} ,\emph{Daniella ogea} , and \emph{Dacryodes
klaineana} (Hall \& Swaine, 1981). The forest reserve experiences two
distinct seasons: the dry season and the rainy season. The dry season is
from December to March whereas April to October marks the rainy season.
Average annual rainfall is between 1300 and 1600 mm. Mean annual
temperature ranges between 26 and 29\textsuperscript{o}C, and relative
humidity is usually above 90 \% in the rainy season and falls to 60\%
during the dry season (Forest Services Division, 2010b).

\textbf{Sampling design and data collection}

A total of eight 50 \selectlanguage{ngerman}× 50 m plots were randomly established in each of
three forest sites namely, edge, interior and deep-interior. Each forest
site had two randomly demarcated and independent sampling areas, each of
which contained four plots. The edge site was defined as 0-50 m from the
forest edge, while interior and deep-interior sites were 200 m and 400 m
from the forest edge, respectively. Variable penetration distances of
edge have been reported in previous studies. A lot of studies revealed
that edges can extend up to 100 m from the forest edge, while other
studies also detected edge effects up to 300 m (see Ofosu-Bamfo et al.,
2019). Thus, we set our two interior sites 100 m beyond each of the
aforementioned edge penetration distance limit, resulting in 200 m and
400 m distances from the forest edge.

We surveyed and identified all lianas with diameter (at 1.30 m from the
rooting base) \selectlanguage{english}[?] 1 cm as well as trees (diameter at breast height \selectlanguage{english}[?] 10
cm) that carried lianas in the plots. The minimum inter-plot distance in
the sampling areas was 150 m. Plant species were identified by a plant
taxonomist, and through the use of herbarium specimens and
identification guides (Hawthorne \& Jongkind, 2006; Hawthorne, 1990).

\textbf{Data analysis}

Community structure

We calculated species richness and Shannon diversity index in the forest
sites. A rarefaction-extrapolation technique was used to standardise
species richness based on a constant number of individuals using iNEXT
package in R. We tested the significance of the differences in Shannon
diversity index among the forest sites using permutation tests in PAST
statistical package version 2.17c (Hammer et al., 2001).

Community abundance of lianas was compared among the forest sites by
running nested ANOVA, where sampling area was nested within forest site.
We employed aov function in the stats package in R to perform the nested
ANOVA. Using the equation of Harper et al. (2005, 2015), we calculated
magnitude of edge influence (MEI) on abundance for individual liana
species with abundance \selectlanguage{english}[?] 10 stems. The equation is given
as:\(MEI=\frac{e-i}{e+i}\), where e = species abundance in edge site, and i =
species abundance in non-edge site, which was obtained by finding the
average of the values of interior and deep-interior sites. The values of
MEI ranges from \selectlanguage{english}-1 (negative edge influence) to +1 (positive edge
influence). MEI value of zero indicates no edge influence. The strength
of MEI was determined as follows (Ofosu-Bamfo et al., 2019): 0 (no edge
influence), \selectlanguage{english}[?]0.19 (very weak), 0.20--0.39 (weak), 0.40--0.59 (moderate),
0.60--0.79 (strong), 0.80--1.0 (very strong).

Network structure of liana-tree interactions

Liana-tree network structure was quantified using the following network
metrics: nestedness, modularity, degree of specialization (H2', d'),
connectance, module connectivity and interactions (c and z values),
species co-occurrence. We used quantitative liana-tree species matrices
except in the species co-occurrence test where binary matrices were
employed. Each of matrices was made up of liana species assigned to rows
and tree species assigned to columns. We also represented the various
networks in graphs using plotweb function in the bipartite package in R.

Nestedness

Nestedness occurs when the more specialist species interact only with
subsets of the species interacting with the more generalist species
(Bascompte et al., 2003; Ponisio et al., 2019). This means that
generalists interact with one another, and specialists tend to interact
with generalists, but specialist-specialist interactions are often
absent (Bascompte et al., 2003). We calculated weighted nestedness
metric, WNODF with the networklevel function in bipartite package in R
(Dormann et al., 2020), in accordance with the nestedness equation of
Almeida-Neto and Ulrich (2010). The WNODF metric ranges from 0 (fully
non-nested) to 100 (fully nested). There are two forms of non-nested
pattern described in literature: (1) when nestedness value is consistent
with the null model expectation, and (2) when nestedness value is
significantly less than that of the null model. The aforementioned
patterns of nestedness refer to two different community assembly (random
and non-random assembly, respectively) and therefore must be
distinguished. We therefore used~anti-nestedness~to refer to the
situation where observed nestedness values were significantly lower than
those expected by chance, while we referred to networks that presented
observed nestedness values which were consistent with null model
expectation as not nested.

Degree of specialisation

The degree of specialisation was determined for the various networks and
the individual species in the networks as follows:

Using the H2' index, we quantified network specialisation of the various
forest sites. The index measures the extent to which observed
interactions deviate from the interactions that would be expected given
the marginal totals of the interactions per species (Bluthgen et al.,
2006). Generally, higher values of the H2' index indicate that the
species in the network are more selective, resulting in higher
specialisation of the network. The index ranges from 0 (no
specialisation) to 1 (complete specialisation). The H2' index was run
with H2fun function in the bipartite package.

The degree of species specialisation was determined by calculating d'
index, using dfun function in the bipartite package. This index is
defined as the deviation from a conformity expected by the overall
utilisation of potential partners (Bluthgen et al., 2007).

Network connectance

Weighted connectance was calculated to express network connnectance in
the study. It is defined as the linkage density divided by number of
species in the network (Dormann et al., 2020; van Altena et al., 2016).
The values of weighted connectance range from 0 (no interaction) to 1
(perfectly connected). Weighted connectance was run with the
networklevel function in the bipartite package.

Modularity

We measured modularity index (Q) with the DIRTLPAwb+ algorithm using
computeModules function within the bipartite package (Beckett, 2016).
Modularity measures the tendency of a network to form modules of
interacting species, which interact more with one another than with
species of other modules (Carstensen et al., 2016; Dormann et al.,
2020). The Q index ranges from 0 for networks with clustering not
different from random to 1 for networks with perfect modules. The Q
index calculation followed the equations in Newman (2006).

Test of statistical significance of the metrics

The above mentioned network metrics were tested for their statistical
significance by generating 1,000 null models and comparing them with the
observed metric values using the Patefield algorithm (Patefield, 1981)
in the bipartite package.

Module connectivity and interactions

The topological roles of liana and tree species with respect to network
modularity was assessed based on the number of links of the species. We
achieved this by calculating the weighted standardised among-module
connectivity (c) and within-module interactions (z), using species
strength of interaction (Watts et al. 2016). To obtain the corresponding
appropriate c and z thresholds for the species, we generated 100 null
models of the original networks using DIRTLPAwb+ algorithm, and 95 \%
quantiles as threshold c- and z-values. Based on the c and z values
generated, the species were grouped into four categories of topological
roles (Olesen et al., 2007) indicated below:

\begin{enumerate}
\tightlist
\item
  Peripherals: species with lower c- and z-values compared to the
  threshold values.
\item
  Network hubs: species with higher c- and z-values compared to the
  threshold values.
\item
  Connectors: made up of species with higher c-values and lower z-values
  compared to the threshold values.
\item
  Module hubs: made up of species with higher z-values and lower
  c-values compared to the threshold values.
\end{enumerate}

Species co-occurrence

Liana species co-occurrence patterns were determined with the
cooc\_null\_model function from EcoSimR package (Gotelli et al., 2015).
We used the C-score metric, which is the average number of checkerboards
for two species (Stone \& Roberts, 1990), to measure species
co-occurrence. The metric was calculated according to the equation
described by Almeida-Neto \& Ulrich (2011). To assess the patterns of
co-occurrence, 10,000 null models were generated by the quasiswap
algorithm and compared with the observed c-score values. The c-score
measures the tendency of species to not co-occur (Stone \& Roberts,
1990). Thus, the greater the c-score in relation to the null model, the
greater the tendency of the species to not co-occur (i.e., segregation),
and the smaller the c-score value in relation to the null model, the
higher the tendency of species to co-occur (i.e., aggregation).

RESULTS

\emph{Liana community structure}

There were more liana species in edge habitat than interior habitat,
which in turn had more species than deep-interior site in the Asenanyo
Forest Reserve (Table 1). Both the rarefaction and extrapolation curves
attested to this observation (Figure 1a). The rarefaction curves did not
reach asymptote, showing there could be more undetected species in the
forest sites. The species in edge, interior and deep-interior sites
belonged to 28 genera and 15 families, 26 genera and 16 families, and 24
genera and 12 families, respectively. Edge and interior sites had
similar Shannon diversity index (P = 0.691; H' = 2.93 and 2.91,
respectively), while each of them supported significantly higher Shannon
diversity index than deep-interior site (H' = 2.74) (P = 0.004 and
0.010, respectively). Species evenness (E) was similar among all the
forest sites in the Asenanyo Forest Reserve (P \textgreater{} 0.05;
edge: E = 0.48, interior: E = 0.51, deep-interior: E = 0.53).

Within the Suhuma Forest Reserve, liana species richness was comparable
among the three forest sites (Table 1). The identified liana species
belonged to 29 genera and 15 families in edge site, 27 genera and 13
families in interior site, and 27 genera and 16 family in deep-interior
site. The rarefaction and extrapolation curves of the forest sites
depicted a similar trend, with the curves showing that there could be
more undetected species in the forest sites (Figure 1b). Shannon
diversity index in edge site (H'= 2.99) did not vary from that in
interior site (H' = 3.01) (P = 0.723). However, due to significantly
higher species evenness in deep-interior site (P \textless{} 0.05;
deep-interior: E = 0.59, edge: E = 0.45, interior: E = 0.50), it
supported significantly higher Shannon diversity index (H' = 3.20) than
edge (P = 0.004) and interior (P = 0.027) sites.

The five most abundant liana species in edge and interior sites of the
Asenanyo Forest Reserve were \emph{Millettia chrysophylla}
,\emph{Salacia elegans} , \emph{Griffonia simplicifolia} , \emph{Alafia
barteri} and \emph{Motandra guineensis} (Table 1). They contributed 54.5
and 54.3 \% of total liana abundance in edge and interior sites,
respectively. The 10 most abundant species in these sites contributed
74.9 and 74.8 \% of the liana stems in edge and interior sites,
respectively. All the above mentioned species also constituted the five
most abundant species in interior site. Four of the above mentioned
species namely, \emph{M} . \emph{chrysophylla} , \emph{G}
.\emph{simplicifolia} , \emph{A} . \emph{barteri} , and \emph{S}
.\emph{elegans} , together with \emph{Strophanthus preussii} were the
five most abundant species in deep-interior site, forming 58.3 \% of the
stems in this site. The 10 most abundant species in deep-interior site
constituted 80.6 \% of the total stems in the site.

In the Suhuma Forest Reserve, \emph{M} . \emph{chrysophylla} , \emph{G}
.\emph{simplicifolia} , \emph{M} . \emph{guineensis} ,
\emph{Calycobolusafricanus} and \emph{A} . \emph{barteri} were the five
most abundant species in edge site, constituting 54.8 \% of liana stems
in the site (Table 1). The stems of the 10 most abundant species were
72.9 \% of the liana abundance in edge site. In interior site, the five
most abundant species included four of the above mentioned species in
edge site (i.e., \emph{M} . \emph{chrysophylla} , \emph{G}
.\emph{simplicifolia} , \emph{C} . \emph{africanus} , \emph{M}
.\emph{guineensis} ) and \emph{Acacia pentagona} . The abundance of
these five species was 52.3 \% of the total abundance in interior site.
With regard to the 10 most abundant species, they contributed 75.3 \% of
the liana abundance in the site. \emph{M} . \emph{chrysophylla} ,
\emph{A} .\emph{pentagona} , \emph{G} . \emph{simplicifolia} , \emph{C}
.\emph{africanus} , and \emph{A} . \emph{barteri} formed the five most
abundant species in deep-interior site, by contributing 41.3 \% of the
liana stems in deep-interior site. In the same forest site, the 10 most
abundant species contributed 65.8 \% of the total liana stems in the
site.

Liana abundance differed significantly between edge and deep-interior
sites of the Asenanyo (F = 3.84; P = 0.041) and Suhuma (F = 6.05; P =
0.010) Forest Reserves (Table 1). Nonetheless, there were no significant
differences in liana abundance among the other forest sites. In both
forest reserves, there was no significant effect of sampling site on
liana abundance (Asenanyo: F = 0.091, P = 0.964; Suhuma: F = 2.16,
0.128). MEI in the Asenanyo Forest Reserve ranged from -1 to 0.92 (Table
1). More species experienced positive MEI on their abundance than those
that had negative MEI on their abundance. \emph{Caesalpinia
cucullata}and \emph{Combretum acutum} were the only species that
experienced very strong MEI in the Asenanyo Forest Reserve.
\emph{Paullinia pinnata} was the only species with strong MEI on its
abundance. The rest of the species recorded moderate, weak, and very
weak MEI on their abundance. On the contrary, \emph{A} .
\emph{pentagona} had no MEI on its abundance. In Suhuma Forest Reserve
the MEI on liana species abundance ranged from -0.43 to 0.45 (Table 1).
Liana species abundance experienced both positive and negative MEI, with
the majority of the species experiencing positive MEI. Nevertheless,
there was no MEI on the abundance of\emph{Strophanthus sarmentosus} in
the Suhuma Forest Reserve. There was moderate MEI on the abundance of
\emph{Manniophyton fulvum} and\emph{Neuropeltis prevosteoides} , while
the MEI on the abundance of the remaining species was either weak or
very weak.

\emph{Network metrics}

We observed 179 interactions between 40 liana species and 38 tree
species in edge site of Asenanyo Forest Reserve (Appendix 1a). A total
of 123 and 119 interactions were recorded in interior (involving 34
liana species and 28 tree species; Appendix 1b) and deep-interior
(between 31 liana species and 35 tree species; Appendix 1c),
respectively. On the part of Suhuma Forest Reserve, 44 liana species
interacted with 63 tree species in edge site and produced a total of 202
interactions (Appendix 1d). In interior site, 44 liana species
interacted with 46 tree species, resulting in 173 interactions (Appendix
1e). We recorded an interaction involving 42 liana species and 46 tree
species in deep-interior site, giving rise to 175 interactions (Appendix
1f).

In the Asenanyo Forest Reserve, the observed nestedness metric was
significantly lower than the means of the null model in the three forest
sites (Table 2). Likewise, the liana-tree networks were less connected
than the null models of the three networks. However, the three networks
were more modular and specialised compared to the null networks. The
significant modularity of the networks resulted in the formation of a
number of modules in edge site (14 modules), which was more than the
number of modules in deep-interior (11 modules), which in turn, was more
than that in interior site (7 modules) (Figure 2a-c; Appendix 2). The
size of the modules varied greatly in the networks, ranging from 2-13
species in edge site, 5-13 species in interior site, and 2-12 species in
deep-interior site.

We did not observe significant differences in nestedness between the
observed and null models in the three forest sites in the Suhuma Forest
Reserve. Nevertheless, the three liana-tree networks in the forest
recorded significantly higher modularity and specialisation than
expected by chance. The networks in deep-interior forest site
(deep-interior: 14 modules) exhibited higher number of modules than the
other sites (edge site: 9 modules, interior site: 9 modules) (Figure
3a-c; Appendix 2). Furthermore, the networks showed much variation in
the size of the modules (edge: 5-19 species, interior: 6-15 species,
deep-interior: 3-11 species). Connectance of the three networks was
significantly lower than that of the null models. The specialisation
asymmetric values of the networks in the Asenanyo Forest Reserve were
close to zero, indicating weak asymmetry. The specialisation asymmetry
value of interior site network was consistent with that of the null
model; those of the other networks were significantly higher than
randomised expectations. The networks in the Suhuma Forest Reserve did
not only show weak asymmetry, but they also did not differ significantly
from that expected by chance.

\emph{Species specialisation metric}

The degree of specialisation differed considerably among the species in
the three forest sites within the Asenanyo Forest Reserve. The majority
of the liana species (edge: 61.5 \%, interior: 55.9 \%, deep-interior:
63.3 \%) had significantly higher degree of specialisation than that of
the respective null models (Appendix 3). Correspondingly, the degree of
specialisation varied widely among the species in the forest sites of
the Suhuma Forest Reserve. Most of the species in edge (58.1 \%) and
deep-interior (65.0 \%) sites were significantly more specialised than
expected by chance. The minority of the tree species in the three sites
of Asenanyo and Suhuma Forest Reserve were more specialised than
expected by chance (Appendix 4). About 43 \% of liana species in
interior site had significantly higher degree of specialisation than the
null model. In the Asenanyo Forest Reserve, the proportion of tree
species with higher specialisation than their null models (edge: 42.1
\%, interior: 42.9 \%, deep-interior: 37.1 \%) was lower than the
proportion of tree species that did not show higher specialisation than
expected by chance. A similar trend was recorded in the Suhuma Forest
Reserve (edge: 36.5 \%, interior: 39.1 \%, deep-interior: 45.7 \%).
Generally, higher proportions of lianas than tree species showed higher
specialisation than the null models.

\emph{Species topological roles in the networks}

In the Asenanyo Forest Reserve, liana species in edge site were mainly
peripherals, with the exception of four species (\emph{C}
.\emph{cucullata} , \emph{M} . \emph{chrhysophylla} , \emph{M}
.\emph{guineensis} , \emph{Morinda} \emph{morindoides} ) which acted as
connectors (Figure 4a). \emph{Millettia lutens} and \emph{Tiliacora
dielsiana} were the only module hub species of lianas in edge site.
Network hubs did not occur among lianas in edge site. The connector and
module hub species constituted 15.4 \% of liana species in this site.
The majority of the tree species also performed specialist role in edge
site, but seven of the species (\emph{C} . \emph{mildbraedii}
,\emph{Hymenostegia afzelii} , \emph{Trilepisium} sp., \emph{Baphia
nitida} , \emph{Entandrophragma utile} , \emph{Triplochiton scleroxylon}
,\emph{N} . \emph{papaverifera} ) were connectors (Figure 5a). These
generalist species formed 25 \% of the tree species. In interior site,
we had no liana connectors and network hubs, but two module hubs existed
in this site (\emph{M} . \emph{guineensis} , \emph{S} . \emph{elegans}
), making up 5.6 \% of liana species in interior site (Figure 4b). The
rest of the liana species served as peripherals in interior site. Trees
in interior site were mostly peripherals, with only one connector
(\emph{Albizia zygia} ) and one module hub (Bec) species (Figure 5b),
but no network hub trees. These generalists were 5.7 \% of the total
tree species in this site. Within deep-interior site, we recorded two
liana connectors \emph{(G} . \emph{simplicifolia} and \emph{S}
.\emph{preussii)} and one liana module hub (\emph{S} . \emph{elegans} )
(Figure 4c), which together made up 10.3 \% of the liana species in the
site. The rest of the liana species in deep-interior site were
peripherals. There were three connectors (\emph{C} . \emph{mildbraedii}
,\emph{Amphimas pterocarpoides} , \emph{Turraeanthus africanus} ) and
one module hub of tree species (\emph{Homalium dewevrei} ) in
deep-interior site, but there was no network hub species (Figure 5c).
These tree species composed of 8.6 \% of the total number of species in
deep-interior site.

We recorded \emph{G} . \emph{simplicifolia} , \emph{C} .
\emph{tarquense} ,\emph{A} . \emph{kamerunensis} and \emph{Combretum
paniculatum} as connector liana species within edge site of the Suhuma
Forest Reserve, while the majority of the liana species were peripherals
(Figure 6a).\emph{C} . \emph{africanus} was a network hub in the edge
site. The above mentioned generalists constituted 11.4 \% of the total
liana species. Five tree species acted as connectors in edge site
(\emph{C} .\emph{mildbraedii} , \emph{Celtis philippensis} ,
\emph{Entandrophragma angolense} , \emph{N} . \emph{papaverifera} ,
\emph{Trichilia prieuriana} ), while one tree species was identified as
a module hub (\emph{Calpocalyxbrevibracteatus} ) (Figure 7a). Network
hubs of tree species were not recorded in edge site. Together, the
connector and module hub species formed 9.5 \% of the total number of
tree species in edge site. In interior site of Suhuma Forest Reserve,
most of the liana species were peripherals. Generalist liana species
were module hubs (\emph{A} .\emph{pentagona} , \emph{C} .
\emph{africanus} , \emph{N} .\emph{acuminata} , \emph{M} . \emph{fulvum}
) and network hub (\emph{M} .\emph{chrysophylla} ), but with no
connector species (Figure 6b). The above mentioned generalist species
formed 12.2 \% of liana species in interior site. The majority of the
tree species in interior site of Suhuma Forest Reserve acted as
peripherals. We did not identify connector tree species in this site,
but a few module hub species occurred there (\emph{Albizia
adianthifolia} , \emph{C} .\emph{mildbraedii} , \emph{Sterculia oblonga}
) (Figure 7b). These generalist species formed 6.5 \% \% of the tree
species. In deep-interior site, lianas were mainly peripherals, except
for \emph{G} .\emph{simplicifolia} , \emph{C} . \emph{africanus} ,
\emph{Neuropeltis prevosteoides} and \emph{Alafia} sp., that acted as
connectors (Figure 6c). The above-mentioned generalists formed about 9.8
\% of the total liana species. Tree species in deep-interior site were
generally peripherals, except \emph{C} . \emph{mildbraedii} and
\emph{Ricinodendron heudelotii} (connectors), and Amp and Gut (module
hubs) which formed 8.7 \% of the tree species (Figure 7c).

\emph{Species co-occurrence of lianas}

In the Asenanyo Forest Reserve, the observed c-score of the matrix in
edge site was significantly lower than the mean of the null model (Table
2). Nevertheless, the c-scores of the observed matrices in interior and
deep-interior sites was consistent with the simulated mean c-scores. In
all the three sites in the Suhuma Forest Reserve, the observed c-score
values were not significantly different from those expected by chance.

DISCUSSION

\emph{Liana community structure}

Our study showed contrasting edge effects on species diversity in the
two moist semi-deciduous forests. In the Asenanyo Forest Reserve, edge
appeared to have enhanced diversity, while an opposite trend occurred in
the Suhuma Forest Reserve. The trend in the Asenanyo Forest Reserves is
consistent with previous studies that also recorded higher liana
diversity at forest edges in relation to forest interiors (Addo-Fordjour
\& Owusu-Baodi 2016; Laurance et al., 2001). It is important to note
that edge effect in our study was observed in edge site in relation to
only deep-interior site. Therefore, we did not observe variation in
species diversity between edge and interior sites. This trend is an
indication that edge effects on liana species diversity penetrated 200 m
into the interior of the forest. The findings of the present study also
indicated that species evenness was poorer in edge site of the Suhuma
Forest Reserve, resulting in a lower species diversity, irrespective of
its higher species richness. The forest edge appears to have influenced
species evenness by exerting differential effects on the abundance of
different species in edge site. Thus, in edge site of Suhuma Forest
Reserve, changes in species abundance mediated edge effects on liana
species diversity. The observed species diversity variation may possibly
relate with edge-related changes in variables such as microclimate, and
tree density and mortality that often characterise forest edges
(Ofosu-Bamfo et al., 2019; Wekesa et al., 2019).

Our study revealed pronounced edge effect on liana abundance at the
community level in the two forest reserves. This pattern is supported by
previous studies which showed that edge effect enhanced liana abundance
in their respective forests (Addo-Fordjour \& Owusu-Boadi, 2016;
Campbell et al., 2018; Laurance et al., 2001). Our results showed that
in both forest reserves, edge effects penetrated 200 m from the edge,
and this is in keeping with Laurance et al. (1991) who reported that
edge effects on the abundance of disturbance-adapted plants such as
lianas can penetrate 200 m into forest interior. Forest edges are often
characterised by increased levels of light and desiccation or dryness,
which can promote liana increase (Campbell et al., 2018). Given that
there was a sharp contrast between our forest edges and the surrounding
matrix, we expected the above mentioned conditions to be more pervasive
in edge site. Thus, increased levels of light and dryness at the forest
edge may be associated with the positive response of liana abundance to
edge disturbance in the two forests. At the species level, many liana
species showed diverse responses to edge disturbance in the two forests.
The values of MEI with respect to species abundance varied widely among
the liana species. A similar finding was reported in two rainforests in
Ghana (Ofosu-Bamfo et al., 2019). Our finding implies that although
optimal light and dry conditions may characterise forest edge sites,
they may not enhance the proliferation of some liana species. This may
occur when there is excessive soil desiccation causing liana mortality
(Nepstad et al., 2007). Nevertheless, some of the liana species showed
high positive MEI, indicating that they tended to prefer edge sites.

\emph{Liana-tree network structure}

We found anti-nested and modular structure in the three liana-tree
interaction networks in Asenanyo Forest Reserve. This trend has also
been reported by Addo-Fordjour et al. (2021, 2016), and to some extent
by Magrach et al. (2016) whose liana-tree networks showed anti-nested
structure (see supplementary data). Nevertheless, our study is at
variance with that of Sfair et al. (2010) which recorded nested
structure in three distinct vegetation formations in Brazil, and also
differs from the networks of Sfair et al. (2015) which did not show
modularity. In the suhuma Forest Reserve, all the three networks were
not nested but modular. Though the two nestedness patterns shown by the
networks in the Asenanyo and Suhuma Forest Reserves refer to non-nested
structure, that of the former depicts non-random assembly of species
whereas the latter indicates random assembly of species. We argue that a
clear distinction should be made between the two types of non-nestedness
in network studies so that the distribution pattern of each of them
would be fully understood. The presence of non-significant nestedness in
the Suhuma Forest Reserve may be due to differences in liana species
ability to colonise host trees and/or the use of defense strategies of
hosts to avoid lianas (Addo-Fordjour et al., 2016; Genini et al., 2012).
As a recap, a nested structure is formed when there are interactions
involving generalists and generalists, and specialists and generalists,
but no interaction of specialists and specialists (Landi et al., 2018).
Staniczenko et al. (2013) showed that for a nested quantitative network,
interactions of generalist-generalist species are strongest, followed by
those of generalist-specialist species, with no specialist-specialist
interactions (or when present with much weaker interactions). Thus, for
a nested structure to occur in a quantitative network like ours, there
should be a good number of specialist and generalist species undergoing
interactions. However, in our networks, we observed only a few
generalists of lianas and trees that interacted, but with many
specialist species interacting among themselves. This situation
increased the likelihood of specialist-specialist interactions at the
expense of generalist-generalist and generalist-specialist interactions,
resulting in absence of nested structure in the various networks. A
similar trend was observed in mycorrhizal networks (Jacquemyn et al.,
2015). The specialist-specialist interactions in our networks may
account for the non-asymmetry and weak asymmetry exhibited by the
networks. This finding shows that our networks tended to be more
symmetric in their interactions, a trend which causes non-significant
nestedness and significant modularity in ecological networks (Guimaraes
et al., 2007). Overall, the findings on liana-tree network structure
reported in the current and previous studies show that there is no
universal pattern in the structure of liana-tree interactions. The
patterns obtained may be dependent on the network complexity, and
species traits and abundance, which are known to influence the
organisation of liana-tree interactions (Sfair et al. 2018, 2010). The
existence of high modular structure in the various networks may increase
their stability and robustness by limiting diffusion of perturbations
through network (Thebault \& Fontaine, 2010). This may explain why the
patterns of network structure in edge site was consistent with those in
interior and deep-interior sites, irrespective of disturbance at edge
site. The modular structure of our networks may help conserve the
networks of species interaction, which in turn, may lead to the
conservation and maintenance of ecosystem functioning.

Though the nature of liana-tree interaction is still a subject of
debate, it tends to be antagonistic, in view of the fact that lianas are
not only structural parasites of trees (Tang et al., 2012), but also
compete intensely with trees for resources (Sfair et al., 2018). Species
of antagonistic networks often evolve high specialisation in order to
survive the antagonism of the interactions (Maliet et al., 2020). Our
results revealed strong species and network specialisation in the forest
sites, which demonstrates the existence of strong liana-host specificity
across the various networks in the two forest. Host specificity and
network specialisation have been reported to cause non-nestedness and
modularity in networks (Cordeiro et al., 2020; Dallas \& Cornelius,
2015; Wardhaugh et al., 2015; Maliet et al., 2020). Given this
information, the non-nested and modular structure observed in our
networks may be driven by the specialisation of the networks and host
specificity of the liana species. The specialisation in the liana-tree
networks may be related to co-evolution in lineages of lianas and trees
in the networks (Sfair et al., 2015). The possibility of co-evolution of
lianas and trees in our networks is supported by Ponisio et al. (2019,
2017) who showed that ecological communities that co-evolve become more
anti-nested and modular over time. Montoya et al. (2015) found out that
functional group diversity increases with modularity in complex
networks, and that functional groups form modules in communities. In
this regard, the presence of high number of modules per network in the
forest sites may reflect the existence of different liana functional
groups that interact with tree communities in the forests. Such networks
with high level of modularity may possess increased resistance to
disturbance (Olesen et al., 2007; Saunders \& Rader, 2019). Differences
in colonisation rates in fish parasites were found as a cause of
anti-nested structure in such networks (Poulin \& Guegan, 2000). In each
of the networks, different liana species showed varying degree of
specialisation, while others exhibited generalisation. This phenomenon
suggests that the rate of colonisation would differ markedly among the
species, with highly specialised species having lower rate of
colonisation, while species with low specialisation, or generalisation
exhibit higher colonisation rate. In this regard, like the parasite-fish
network (Poulin \& Guegan, 2000), the anti-nested structure in our
networks could have partly been occasioned by variation in colonisation
rates of the liana species. Generally, our study adds to the number of
studies that have demonstrated the existence of non-nestedness and
modularity in liana-tree networks (e.g., Addo-Fordjour et al., 2021;
Magrach et al., 2016).

\emph{Species role in the networks}

The finding of the current study showed that lianas and trees were
predominantly specialists (i.e., peripherals), irrespective of edge
disturbance or edge effects, indicating possible robustness of species
roles to disturbance. A similar pattern was recorded in a moist
semi-deciduous forest in Ghana (Addo-Fordjour et al., 2021). The high
specialisation of the species in the networks shows that antagonism in
the networks might have resulted in the evolution of specialisation in
both liana and tree species (Maliet et al., 2020). The role of some of
the liana and tree species was consistent in the forest sites, whiles
other species roles changed from one site to another. This phenomenon
indicates that edge effects probably caused a switch in the role of some
of the species among the forest sites, while the role of other species
remained unchanged. The change from specialist to generalist and vice
versa, and from one form of generalist to another may be related to
changes in species abundance and distribution following edge disturbance
(Addo-Fordjour et al., 2021). Some of the species identified as
structural important species (i.e., connectors, module hubs, network
hubs) in our study were also reported as species that possessed
structural importance in a moist semi-deciduous forest in Ghana
(Addo-Fordjour \& Afram, 2021). These plants which include two liana
species (\emph{G} . \emph{simplicifolia} , \emph{C} . \emph{africanus} )
and three tree species (\emph{T} . \emph{scleroxylon} , \emph{N}
.\emph{papaverifera} , \emph{C} . \emph{mildbraedii} ) may have unique
functional roles that support the functioning of the forests.

\emph{Species co-occurrence of lianas}

Generally, lianas were assembled randomly on their hosts in most of the
forest sites, suggesting that chance events rather than edge
disturbance, determined liana distribution on trees. Thus, we argue that
the liana communities might have been assembled on trees by stochastic
processes including host characteristics. Our finding is consistent with
that reported in a semi-deciduous forest in Brazil (Zulqarnain et al.,
2016). Contrary to the above, liana species in edge site of Asenanyo
Forest Reserve showed positive species co-occurrence on their hosts.
Since this network was organised into modules, the positive
co-occurrence trend could have existed within the modules. Thus, in the
modules, liana species resorted to positive or facilitative interactions
(McGarvey \& Veech, 2018), that might have arisen deterministically. At
forest edges, there is usually an elevated level of light coupled with
dry conditions, and trellis availability, all of which can work together
to enhance liana proliferation (Campbell et al., 2018). It appears that
as these resources are increased at edge, lianas tend to share rather
than compete for them, resulting in their positive co-occurrence on the
host trees. The liana species aggregation on trees could have also
arisen by facilitation, where increasing liana abundance at edge site
would cause new liana individuals to use already established stems to
climb trees (Perez-Salicrup \& Sork, 2001).

CONCLUSION

The findings of the study revealed considerable edge effects on liana
diversity and abundance in the two moist semi-deciduous forests. Despite
the enhanced diversity and abundance in edge site of each forest, the
patterns of liana-tree network structure of edge site were similar to
those in interior and deep-interior sites. All the networks in the two
forests were less connected and non-nested, but modular and specialised.
Lianas were mostly randomly distributed on host trees in all the forest
sites except edge site in the Suhuma Forest Reserve. Topologically, the
majority of liana and tree species were peripherals (i.e., specialist),
but a few species tended to be generalists, acting as connectors, module
hubs and network hubs. The role of most of the species did not change
from one site to another, even though the topological role of a few
species changed from one site to another. Overall, our study shows that
liana community structure was more susceptible to forest edge than
liana-tree network structure. The findings of our study corroborate
previous studies, and also present unique findings related to liana-tree
network structure. Our findings which enhance our understanding of
liana-tree interactions, have conservation implications relating to
stability and robustness of the networks. Finally, the findings of the
present study can potentially contribute to a robust edge theory
development.

\textbf{AKNOWLEDGEMENTS}

The authors are grateful to the Forestry Commission of Ghana for
providing access to the forest reserves.

\textbf{CONFLICT OF INTEREST}

The authors declare no conflict of interest.

\textbf{AUTHOR CONTRIBUTIONS}

\textbf{Bismark Ofosu-Bamfo:} Methodology (supporting); Investigation
(lead); Formal analysis (supporting); writing -- original draft (equal);
writing -- review and editing (supporting). \textbf{Patrick
Addo-Fordjour:} Conceptualisation (lead); Methodology (lead);
Supervision (equal); Formal analysis (lead); writing -- original draft
(equal); writing -- review and editing (lead). \textbf{Ebenezer J.D.
Belford:} Conceptualisation (supporting); Methodology (supporting);
Supervision (equal); writing -- review and editing (supporting).

\section*{DATA AVAILABILITY STATEMENT}

{\label{data-availability-statement}}

Data associated with the manuscript will be available on Dryad.

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\textbf{FIGURE TITLES}

\textbf{FIGURE 1} Individual-based rarefaction-extrapolation curves
showing liana species richness patterns in the three forest sites of the
two moist semi-deciduous forests in Ghana: (a) Asenanyo Forest Reserve,
(b) Suhuma Forest Reserve. The solid lines show the rarefaction
(interpolation) curves from the reference sample, while the dashed lines
indicate the extrapolation curves. The symbols ending the rarefaction
curves (see also legend) represent observed number of individuals for
the forest sites.

\textbf{FIGURE 2} Network modules identified by DIRTLPAwb+ in edge (a),
interior (b) and deep-interior (c) sites in Asenanyo Forest Reserve,
Ghana. The darker squares represent higher interaction frequency, while
the light squares show lower frequency of interaction. The boxes show
the modules of the networks, which are consecutively numbered. Each
module is made up of liana species on the y-axis and tree species on the
x-axis. The species constituting the modules are found in Appendix 2.

\textbf{FIGURE 3} Network modules identified by DIRTLPAwb+ in edge (a),
interior (b) and deep-interior (c) sites in Suhuma Forest Reserve,
Ghana. The darker squares represent higher interaction frequency, while
the light squares show lower frequency of interaction. The boxes show
the modules of the networks, which are consecutively numbered. Each
module is made up of liana species on the y-axis and tree species on the
x-axis. The species constituting the modules are found in Appendix 2.

\textbf{FIGURE 4} Module connectivity and interactions plots of the
networks that show liana species roles within edge (a), interior (b),
and deep-interior (c) forest sites in the Asenanyo Forest Reserve,
Ghana. The threshold values of among-module connectivity (\emph{c} ) and
within-module interaction (\emph{z} ) which were obtained from 95 \%
quantiles from 100 null models are denoted by the vertical and
horizontal lines. Species names are abbreviated to first two letters of
the genus name and at least the first letter of the specific epithet
(see appendix 3 for full species names)

\textbf{FIGURE 5} Module connectivity and interactions plots of the
networks that show liana species roles within edge (a), interior (b),
and deep-interior (c) forest sites in the Suhuma Forest Reserve, Ghana.
The threshold values of among-module connectivity (\emph{c} ) and
within-module interaction (\emph{z} ) which were obtained from 95 \%
quantiles from 100 null models are denoted by the vertical and
horizontal lines. Species names are abbreviated to first two letters of
the genus name and at least the first letter of the specific epithet
(see appendix 3 for full species names)

\textbf{FIGURE 6} Module connectivity and interactions plots of the
networks that show tree species roles within edge (a), interior (b), and
deep-interior (c) forest sites in the Asenanyo Forest Reserve, Ghana.
The threshold values of among-module connectivity (\emph{c} ) and
within-module interaction (\emph{z} ) which were obtained from 95 \%
quantiles from 100 null models are denoted by the vertical and
horizontal lines. Species names are abbreviated to first two letters of
the genus name and at least the first letter of the specific epithet
(see appendix 4 for full species names)

\textbf{FIGURE 7} Module connectivity and interactions plots of the
networks that show tree species roles within edge (a), interior (b), and
deep-interior (c) forest sites in the Suhuma Forest Reserve, Ghana. The
threshold values of among-module connectivity (\emph{c} ) and
within-module interaction (\emph{z} ) which were obtained from 95 \%
quantiles from 100 null models are denoted by the vertical and
horizontal lines. Species names are abbreviated to first two letters of
the genus name and at least the first letter of the specific epithet
(see appendix 4 for full species names)

\textbf{APPENDIX 1} Structure of the three liana-tree networks analysed
in edge, interior, and deep-interior forest sites of the Asenanyo (a, b
and c, respectively) and Suhuma (d, e and f, respectively) Forest
Reserves in Ghana. Each network shows all the observed interactions in
the forest management regime. The lower and upper bars represent liana
and tree species, respectively. In each network, the bar thickness is
proportional to the number of interactions of each species. Species
names are abbreviated to first two letters of the genus name and at
least the first letter of the specific epithet (refer to Appendices 3
and 4 for full species names).

\textbf{TABLE 1} Liana species abundance and MEI in the edge and
non-edge sites in two moist semi-deciduous forests in Ghana (ES: edge
site, IS: interior site, DIS: deep-interior site, MEI: magnitude of edge
influence)

ASENANYO SUHUMA

Liana species and families ES IS DIS MEI ES IS DIS MEI

Apocynaceae

\emph{Alafia barteri} Oliver 40 33 35 0.08 20 16 22 0.03

\emph{Alafia} sp. 29 13 12 0.40 15 9 9 0.25

\emph{Gongronema latifolium} Benth. 0 0 0 0 1 0

\emph{Landolphia dulcis} (Sabine ex G.Don) Pichon 1 0 2 1 4 1

\emph{Landolphia hirsuta} (Hua) Pichon 0 0 0 9 6 9 0.09

\emph{Landolphia owariensis} P.Beauv. 0 0 0 0 1 0

\emph{Motandra guineensis} (Thonn.) A.DC. 39 41 15 0.16 39 22 16 0.34

\emph{Oncinotis nitida} Benth. 0 0 0 0 0 2

\emph{Parquetina nigrescens} (Afzel.) Bullock 0 0 0 0 1 0

\emph{Strophanthus hirsutus} H.Hess 0 0 0 4 0 0

\emph{Strophanthus hispidus} DC. 1 0 0 0 0 0

\emph{Strophanthus preussii} Engl. \& Pax 8 21 23 -0.52 0 0 0

\emph{Strophanthus sarmentosus} DC. 0 0 0 6 12 0 0.00

Celastraceae

\emph{Hippocratea myriantha} Oliv. 0 0 0 0 0 1

\emph{Salacia debilis} (G.Don) Walp. 3 0 0 2 5 2

\emph{Salacia elegans} Welw. ex Oliv. 45 43 32 0.09 15 4 20 0.11

\emph{Salacia lateritia} N.Halle 8 1 0 0 0 0

\emph{Salacia leptoclada} Tul. 0 0 0 0 3 0

\emph{Salacia macrantha} A.C.Sm. 0 0 0 0 4 0

\emph{Salacia preussii} Loes 0 1 0 0 0 0

\emph{Salacia cerasifera} Welw. Ex Oliv. 0 0 0 0 5 0

\emph{Salacia staudtiana} Loes. ex Fritsch 0 0 0 0 1 0

\emph{Salacighia letestuana} (Pellegr.) Blakelock 0 0 0 1 0 0

\emph{Simirestis staudtii} (Loes.) N.Halle 0 0 0 0 0 2

\textbf{TABLE 1} (Continued)

ASENANYO SUHUMA

Liana species and families ES IS DIS MEI ES IS DIS MEI

Combretaceae

\emph{Combretum acutum} M.A.Lawson 0 13 0 -1.00 0 0 0

\emph{Combretum comosum} G.Don 18 3 21 0.20 0 0 3

\emph{Combretum fuscum} Planch. ex Benth. 0 0 0 0 0 1

\emph{Combretum micranthum} G.Don 1 0 0 0 0 0

\emph{Combretum mucronatum} Schumach. \& Thonn. 0 0 0 2 4 5 -0.38

\emph{Combretum oyemense} Exell 16 3 8 0.49 0 0 0

\emph{Combretum paniculatum} Vent. 6 14 0 -0.08 7 9 9 -0.13

\emph{Combretum smeathmannii} G.Don 0 0 0 9 4 7 0.24

\emph{Combretum sordidum} Exell 0 0 0 2 0 0

\emph{Combretum} sp. 0 0 0 0 1 0

\emph{Combretum tarquense} Clark 0 0 0 5 0 16 -0.23

Connaraceae

\emph{Agelaea obliqua} (P.Beauv.) Baillon 0 2 0 0 0 3

\emph{Agelaea trifolia} (Lam.) Baill. 1 1 3 1 2 0

\emph{Castanola paradoxa} (Gilg) Schellenb. 0 0 0 1 0 0

\emph{Cnestis ferruginea} Vahl ex DC. 0 0 0 0 1 0

\emph{Connarus africanus} Lam. 0 0 0 0 7 0

Convolvulaceae

\emph{Calycobolus africanus} (G. Don) Heine 19 23 7 0.12 25 25 23 0.02

\emph{Calycobolus heudelotii} (Baker ex Oliver) Heine 4 1 0 0 1 1

\emph{Neuropeltis acuminata} (P.Beauv.) Benth. 8 3 17 -0.11 16 14 12
0.10

\emph{Neuropeltis prevosteoides} Mangenot 0 0 0 4 4 16 -0.43

Dichapetalaceae

\emph{Dichapetalum dewevrei} De Wild. \& T.Durand 0 0 0 1 0 0

\emph{Dichapetalum pallidum} (Oliver) Engler 0 1 2 1 0 0

\textbf{TABLE 1} (Continued)

ASENANYO SUHUMA

Liana species and families ES IS DIS MEI ES IS DIS MEI

Dilleniaceae

\emph{Tetracera affinis} Hutch. 2 3 1 0 0 0

Euphorbiaceae

\emph{Manniophyton fulvum} Mull.Arg. 0 3 0 17 6 7 0.45

Hernandiaceae

\emph{Illigera pentaphylla} Welw. 0 0 0 2 0 1

Icacinaceae

\emph{Chlamydocarya macrocarpa} A.Chev.exHutch.\&D 0 2 0 0 0 0

Lamiaceae

\emph{Clerodendrum} sp. 0 0 0 1 0 0

\emph{Clerodendrum umbellatum} Poir 0 0 0 1 0 0

Fabaceae

\emph{Acacia kamerunensis} Gand. 1 5 2 5 7 15 -0.38

\emph{Acacia pentagona} (Schum\& Thonn.)Hook f. 10 11 9 0.00 14 25 26
-0.29

\emph{Baphia capparidifolia} Baker 0 0 0 1 0 0

\emph{Caesalpinia cucullata} Roxb. 12 0 1 0.92 0 0 0

\emph{Dalbergia hostilis} Bentham 3 3 0 3 1 0

\emph{Dalbergia oblongifolia} G.Don 0 0 1 0 0 0

\emph{Dalbergiella welwitschii} (Baker) Baker f. 6 3 5 0.20 3 11 1 -0.33

\emph{Griffonia simplicifolia} (DC.) Baill. 44 37 50 0.01 63 33 25 0.37

\emph{Leptoderris sassandrensis} Jongkind 0 0 0 4 2 0

\emph{Leptoderris cyclocarpa} Dunn 0 0 0 1 0 0

\emph{Leptoderris micrantha} Dunn 11 3 14 0.13 2 8 1 -0.38

\emph{Leptoderris miegei} Ake Assi \& Mangenot 1 0 0 2 0 8

\emph{Leucomphalos libericus} Breteler 0 0 0 1 0 0

\emph{Mezoneuron benthamianum} Baill. 0 0 0 6 0 0

\emph{Millettia chrysophylla} Dunn 84 74 67 0.09 86 84 44 0.15

\emph{Millettia lucens} (Scott-Elliot) Dunn 1 0 2 0 0 0

\textbf{TABLE 1} (Continued)

ASENANYO SUHUMA

Liana species and families ES IS DIS MEI ES IS DIS MEI

Linaceae

\emph{Hugonia planchonii} Hook.f. 0 0 0 0 1 0

\emph{Hugonia rufopilis} A.Chev. ex Hutch. \& Dalziel 0 0 0 1 0 0

Loganiaceae

\emph{Strychnos campicola} Gilg 5 8 7 -0.20 0 0 0

\emph{Strychnos longicaudata} Gilg 0 0 0 9 3 6 0.33

\emph{Strychnos malacoclados} C.H. Wright 0 0 0 0 3 5

Malpighiaceae

\emph{Acridocarpus smeathmannii} (DC.) Guill. \& Perr. 1 0 0 0 0 0

\emph{Grewia hookeriana} Exell \& Mendonca 1 4 2 0 0 0

\emph{Grewia malacocarpa} Mast. 0 0 0 7 1 7 0.27

Menispermaceae

\emph{Tiliacora dielsiana} Hutch. \& Dalziel 9 12 12 -0.14 10 5 8 0.21

\emph{Triclisia patens} Oliv. 1 3 1 0 0 1

Moraceae

\emph{Ficus} sp. 0 0 0 0 0 2

Phyllantaceae

\emph{Phyllanthus} sp. 0 0 0 0 0 1

Piperaceae

\emph{Piper guineense} Shumach. \& Thonn. 1 0 0 0 4 1

Polygonaceae

\emph{Afrobrunnichia erecta} (Asch.) Hutch. \& Dalziel 3 2 2 0 0 2

Rubiaceae

\emph{Morinda morindoides} (Baker) Milne-Redh. 10 8 0 0.43 0 0 0

\emph{Mussaenda tristigmatica} Cummins 1 0 0 0 0 0

\textbf{TABLE 1} (Continued)

Liana species ASENANYO SUHUMA

ES IS DIF MEI ES IS DIF MEI

Sapindaceae

\emph{Paullinia pinnata} Linne 2 15 3 -0.64 0 0 0

Vitaceae

\emph{Cissus adenocaulis} Steud. ex A.Rich 6 3 1 6 1 1

\emph{Cissus silvestris} Tchoume 0 1 0 0 0 0

\textbf{Total number of species 39 36 29 44 41 41}

\textbf{Abundance per 0.25 m\textsuperscript{2} plot 58 52 45 54 45 43}

\textbf{TABLE 2} Patterns of network properties of liana-tree
interactions among three forest sites of two moist semi-deciduous
forests in Ghana

ASENANYO SUHMA

Network metric Observed Null model P-value Observed Null model P-value

\textbf{Edge}

WNODF 11.63 17.82 0.001 6.47 7.51 0.108

Modularity 0.36 0.27 0.001 0.44 0.36 0.001

H2 0.27 0.14 0.001 0.24 0.14 0.001

Connectance 0.13 0.16 0.001 0.06 0.10 0.001

C-score 9.15 9.56 0.013 5.73 5.83 0.281

\textbf{Interior}

WNODF 13.53 18.60 0.001 7.98 8.82 0.233

Modularity 0.41 0.28 0.001 0.42 0.33 0.001

H2 0.32 0.16 0.001 0.23 0.14 0.001

Connectance 0.14 0.18 0.001 0.08 0.10 0.001

C-score 8.90 9.17 0.113 7.04 7.25 0.137

\textbf{Deep-nterior}

WNODF 11.96 16.36 0.008 5.27 5.87 0.229

Modularity 0.41 0.25 0.001 0.45 0.34 0.001

H2 0.33 0.15 0.001 0.24 0.11 0.001

Connectance 0.11 0.15 0.001 0.08 0.13 0.001

C-score 5.83 5.92 0.433 9.16 9.25 0.435

\textbf{APPENDIX 2} Liana and tree species forming modules in the
networks of the forest sites in the two moist semi-deciduous forests in
Ghana. Liana species are indicated in \textbf{bold} text, while tree
species are not bolded. The species are represented by codes, which are
made up of first two letters of the genus name and at least the first
letter of the specific epithet (see Table 1 for full names of the
species).\selectlanguage{english}
\begin{longtable}[]{@{}lllllll@{}}
\toprule
\begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Module \#}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Asenanyo}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Asenanyo}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Asenanyo}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Suhuma}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Suhuma}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Suhuma}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Edge}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Interior}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Deep-interior}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Edge}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Interior}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Deep-interior}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 1\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cia, Daw, Pig,} Trs\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cop, Grh, Mog, Stc, Tea, Chm,} Ala, Cep, Dis\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Agt, Mil, Nep, Afe,} Cap, Trd\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Alsp, Caa, Cap, Lao,} Ala, Amp, Buo, Cep, En.an, Ptm, Sck,
Coc\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Coa, Cop, Cot, Daw, Grs, Sas,} Cap, Dia, Disp, Fie, Ptm, Str,
Ich\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Mog, Daw, Lad,} Dia, Trp\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Module 2\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Sad,} Pia\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Agt, Alb, Daw, Grs,} Ena, Gut, Gue, Hec, Tes, Rih\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Ack, Mic,} Alz, Alb, Chp, Nep, Nel, Sck, Trp\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Lah, Mic, Mog,} Alb, Anm, Bap, Bls, Cea, Cep, Cez, Dis, Drp,
Mab, Mad, Nep, Trp, Trt\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Ack, Alb, Cnf, Hup, Les, Mic, Tid} Cab, Cog, Lot, Mip, Nav, Rav,
Sto, Stp\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Afe, Sas,} Anm, Cop\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Module 3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Mog, Nea, Trp,} Hya, Nep, Nel, Aig\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Caa, Coa, Coo, Dah, Mic, Nea, Tid,} Amp, Cem, Drg, Nep, Pia,
Stp\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Alb,} Fua, Ena\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Cls, Cos, Did, Ilp, Les, Nep, Sae, Sts, Nea,} Ant, Ban, Cap,
Cea, Cem, Fie, Hak, Heb, Zag, Aig\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Acp, Grm, Pig, Sal,} Bls, Buo, Desp, Mya, Oma, Tua\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Dah, Fisp, Maf, Tid,} Anf, Dak, Dik, Stp\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 4\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Afe, Grh, Mil, Mut,} Ena, Uac\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Sae, Sal, Sap, Stp, Cia, Trp,} Ban, Be.co, Ce.od, Ch.su, Py.an,
Tr.mo\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Acp, Cac, Coo, Lemic, Tea,} Cep, Los, Pia\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Maf, Stl, Tid,} Anp, Mip, Myl, Tei, Trm\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Alsp, Gol, Mog, Sas,} Cep, Cap, Trm, Trp, Trsp\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cof, Cop, Cot, Sts, Phsp, Cac,} Ban, Dig, Mie, Rih\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Module 5\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Cah, Coo, Cop, Grs, Lad, Sae, Mom,} Cem, Pya, Stt, Trp, Trs,
En.ut,\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Ago, Alsp, Maf,} Cea, Etu\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Pap,} Ala\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Agt, Daw, Grm, Hur, Lemie, Sal,} Alf, Bob, Fue, Hya, Rav,
Rid\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Lad, Lemic, Nea, Pan, Sae, Stm,} Ban, Cem, Dig, Tra\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Alb} Beo, Dis, Ptm\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 6\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Acp, Stp,} Pem, Rav\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Acp, Afe, Coc, Pap. Dip,} Hya, Law\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Alsp, Cah, Grs, Mog,} Bec, Ceo, Cog, Irg, Ra.vo\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Dip, Lel, Sth, Lad,} Sck, Zap\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cac, Com, Cosp, Lah, Sad,} Ala, Buc, Mua\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Caa, Lah, Nea, Cia,} Cem, Pab, Pia, Coc\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 7\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Ack, Agt, Cac,} Ban\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Lemic, Mom,} Alz, Cog, Cui\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cia,} Ank\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cia, Cop, Grs, Lec, Lemic, Nea,} Cas, Dak, Law, Mya, Pae, Str,
Stt, Sya, Rih, Enu\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cah, Lao, Nep,} Pya, Sck, Trt\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Lemic, Stm,} Fua\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 8\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Sth,} Bap\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Nea, Tid,} Enu, Maa, Sto\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Ack, Acp, Alb, Sad,} Alz, Ansp, Cap, Dia, Muc, Pia, Sto\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Agt, Caa, Cia, Dah, Maf, Stl,} Alf, Amp, Nep, Pia, Coc\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Ilp, Sae, Nep,} Cog, Ena, Hod\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 9\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Alb, Alsp, Lemie,} Ant, Ceo, Enc, Mip, Zag\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Caa, Daw, Stc,} Cop, Hya, Trm, Trs\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Com, Cos, Cot, Dah,} Tr.sc\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cos, Sam, Sts,} Bap, Bec, Muc, Myl, Pab\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Onn, Stl,} Tua\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 10\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Coc,} Irg\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Stp, Dip,} Ban\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Cah, Trp, Com, Grm,} Alz, Cab, Law, Das\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 11\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Com,} Alz\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Coc, Dao, Lad, Sae, Trp, Grh,} Cem, Elg, Nad, Pya, Stp,
Zag\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Ack, Alsp, Lemie,} Amp, Mya, Myl, Pai\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Module 12\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Caa, Stc, Acs,} Chs, Fua, Nav, Tet, Rih\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Sad, Cos, Pig,} Bap, Bls, Gut, Sck, Tes\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 13\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Dah, Lemic, Mic, Sth,} Ala, Bec, Fis, Stp, Los\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Him, Coc,} Trm\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Module 14\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Pap, Sal, Tea, Tid}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Acp, Grs, Mic}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Con, Desp\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Ant, Cap, Hya, Irg, Nep, Pesp, Pya, Trsp\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\textbf{APPENDIX 3} Standardised specialisation index (\emph{d'} ) for
liana species within liana-tree networks in three forest sites of two
moist semi-deciduous forests, Ghana. Values with an asterisk are
significantly higher than expected by chance.

Liana species ASENANYO SUHUMA

Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Acacia kamerunensis} Gand. 0.41* - 0.64* 0.30 0.45* 0.39*

\emph{Acacia pentagona} (Schum\& Thonn.) Hook f. 0.47* 0.34* 0.54* 0.55*
0.65* 0.32

\emph{Acridocarpus smeathmannii} (DC.) Guill. \& Perr. 0.66* - - - - -

\emph{Afrobrunnichia erecta} (Asch.) Hutch. \& Dalziel 0.63* 0.59* - - -
-

\emph{Agelaea obliqua} (P.Beauv.) Baillon - 0.40* - - - 0.17

\emph{Agelaea trifolia} (Lam.) Baill. 0.27 0.60* 1.00* 0.80* 0.48*

\emph{Alafia barteri} Oliver 0.18 0.28 0.29 0.54* 0.38 0.42*

\emph{Alafia} sp. 0.39* 0.30 0.30 0.43* 0.13 0.17

\emph{Caesalpinia cucullata} Roxb. 0.27 - 0.31 - 0.31 -

\emph{Calycobolus africanus} (G. Don) Heine 0.16 0.22 0.56* 0.25 0.13
0.21

\emph{Calycobolus heudelotii} (Baker ex Oliver) Heine 0.36* - 0.00 -
1.00* 0.69*

\emph{Castanola paradoxa} (Gilg) Schellenb. - - - 0.43* - -

\emph{Chlamydocarya macrocarpa} A.Chev.exHutch.\&D - 0.54* - - - -

\emph{Cissus adenocaulis} Steud. ex A.Rich 0.12 0.67* 0.58* 0.23 0.19
0.00

\emph{Cissus silvestris} Tchoume - 0.72* - - - -

\emph{Clerodendrum} sp. - - - 0.30 - -

\emph{Cnestis ferruginea} Vahl ex DC. - - - - 0.31 -

\emph{Combretum acutum} M.A.Lawson - 0.53* - - - -

\emph{Combretum comosum} G.Don 0.31 0.13 0.44* - - 0.71*

\emph{Combretum fuscum} Planch. ex Benth. - - - - - 0.41*

\emph{Combretum micranthum} G.Don 0.66* - - - - -

\emph{Combretum mucronatum} Schumach. \& Thonn. - - - 0.39* 0.03 0.42*

\emph{Combretum oyemense} Exell 0.33* 0.08 0.45* - - -

\emph{Combretum paniculatum} Vent. 0.34* 0.54* - 0.48* 0.24 0.43*

\emph{Combretum smeathmannii} G.Don - - - 0.14 0.87* 0.44*

\textbf{APPENDIX 3} (Continued)

Liana species ASENANYO SUHUMA

Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Combretum sordidum} Exell - - - 0.55* - -

\emph{Combretum} sp. - - - - 0.31 -

\emph{Combretum tarquense} Clark - - - 0.19 0.62* 0.22

\emph{Connarus africanus} Lam. - - - - 0.64* -

\emph{Dalbergia hostilis} Bentham 0.43* 0.13 - 0.25 1.00* -

\emph{Dalbergia oblongifolia} G.Don - - 0.09 - - -

\emph{Dalbergiella welwitschii} (Baker) Baker f. 0.32* 0.30 0.52* 0.35
0.13 0.49*

\emph{Dichapetalum dewevrei} De Wild. \& T.Durand - - - 0.60* - -

\emph{Dichapetalum pallidum} (Oliver) Engler - 0.34* 0.59* 0.53* - -

\emph{Ficus} sp. - - - - - 0.42*

\emph{Gongronema latifolium} Benth. - - - - 0.44* -

\emph{Grewia hookeriana} Exell \& Mendonca 0.66* 0.24 0.81* - - -

\emph{Grewia malacocarpa} Mast. - - - 0.45* 0.52* 0.56*

\emph{Griffonia simplicifolia} (DC.) Baill. 0.11 0.28 0.24 0.26 0.17
0.40*

\emph{Hippocratea myriantha} Oliv. - - - - - 0.45*

\emph{Hugonia planchonii} Hook.f. - - - - 0.63* -

\emph{Hugonia rufopilis} A.Chev. ex Hutch. \& Dalziel - - - 0.53* - -

\emph{Illigera pentaphylla} Welw. - - - 1.00* - 0.45*

\emph{Landolphia dulcis} (Sabine ex G.Don) Pichon 0.22 - 0.24 0.43* 0.00
0.49*

\emph{Landolphia hirsuta} (Hua) Pichon - - - 0.08 0.34 0.26

\emph{Landolphia owariensis} P.Beauv. - - - 0.80* 0.81* -

\emph{Leptoderris cyclocarpa} Dunn - - - 0.09 - -

\emph{Leptoderris micrantha} Dunn 0.62* 0.54* 0.54* 0.68* 0.08 0.68*

\emph{Leptoderris miegei} Ake Assi \& Mangenot 0.11 - - 0.43* - 0.47*

\emph{Leptoderris sassandrensis} Jongkind - - - 0.30 0.38 -

\emph{Leucomphalos libericus} Breteler - - - 0.53* - -

\textbf{APPENDIX 3} (Continued)

Liana species ASENANYO SUHUMA

Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Manniophyton fulvum} Mull.Arg. - 0.82* - 0.62* 0.38 0.36

\emph{Millettia chrysophylla} Dunn 0.25 0.28 0.35* 0.35 0.16 0.34

\emph{Millettia lucens} (Scott-Elliot) Dunn 0.75* - 0.59* - - -

\emph{Morinda morindoides} (Baker) Milne-Redh. 0.21 0.59* - - - -

\emph{Mezoneuron benthamianum} Baill. \emph{-} ` - - 0.26 - -

\emph{Motandra guineensis} (Thonn.) A.DC. 0.20 0.20 0.26 0.45* 0.22 0.32

\emph{Mussaenda tristigmatica} Cummins 0.66* - - - - -

\emph{Neuropeltis acuminata} (P.Beauv.) Benth. 0.26 0.45* 0.44* 0.47*
0.28 0.33

\emph{Neuropeltis prevosteoides} Mangenot - - 0.43* 0.29 0.63* 0.27

\emph{Oncinotis nitida} Benth. - - - - - 0.35

\emph{Parquetina nigrescens} (Afzel.) Bullock - - - - 0.00 -

\emph{Paullinia pinnata} Linne 0.52* 0.43* 0.85* - - -

\emph{Phyllanthus} sp. - - - - - 0.41*

\emph{Piper guineense} Shumach. \& Thonn. 0.29 - - - 0.46* 0.80*

\emph{Salacia cerasifera} Welw. Ex Oliv. - - - - 0.37 -

\emph{Salacia debilis} (G.Don) Walp. 0.92* - - 0.53* 0.33 0.48*

\emph{Salacia elegans} Welw. ex Oliv. 0.24 0.31 0.23 0.23 0.18 0.38*

\emph{Salacia lateritia} N.Halle 0.67* 0.16 - - - -

\emph{Salacia leptoclada} Tul. - - - - 0.69* -

\emph{Salacia macrantha} A.C.Sm. - - - - 0.63* -

\emph{Salacia preussii} Loes - 0.45* - - - -

\emph{Salacia staudtiana} Loes. ex Fritsch - - - - 0.81* -

\emph{Salacighia letestuana} (Pellegr.) Blakelock - - - 0.69* - -

\emph{Simirestis staudtii} (Loes.) N.Halle - - - - - 0.63*

\emph{Strophanthus hirsutus} H.Hess - - - 0.61* - -

\emph{Strophanthus hispidus} DC. 0.73* - - - - -

\textbf{APPENDIX 3} (Continued)

Liana species ASENANYO SUHUMA

Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Strophanthus preussii} Engl. \& Pax 0.37* 0.19 0.16 - - -

\emph{Strophanthus sarmentosus} DC. - - - 0.30 0.70* 0.35

\emph{Strychnos campicola} Gilg 0.51* 0.42* 0.59* - - -

\emph{Strychnos longicaudata} Gilg - - - 0.34 0.40 0.50*

\emph{Strychnos malacoclados} C.H. Wright - - - - 0.10 0.48*

\emph{Tetracera affinis} Hutch. 0.52* 0.40* 0.42* - - -

\emph{Tiliacora dielsiana} Hutch. \& Dalziel 0.38* 0.26 0.44* 0.38*
0.49* 0.49*

\emph{Triclisia patens} Oliv. 0.83* 0.53* 0.29 - - 1.00*

\textbf{APPENDIX 4} Standardised specialisation index (\emph{d'} ) for
tree species within liana-tree networks in three forest sites of two
moist semi-deciduous forests, Ghana. Values with an asterisk are
significantly higher than expected by chance.

ASENANYO SUHUMA

Tree species Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Aidia genipiflora} 0.16 - - 1.00* - -

\emph{Albizia adianthifolia} 0.39* 0.20 0.17 0.23 0.37* -

\emph{Albizia ferruginea} - - - 0.63* 0.22 -

\emph{Albizia zygia} 0.00 0.36* 0.18 0.52* - 0.29

\emph{Alstonia boonei} - - 0.13 0.49* - -

\emph{Amphimas pterocarpoides} - 0.55* - 0.30 1.00* 0.39*

\emph{Annickia polycarpa} - - - 0.42* - -

\emph{Anopyxis klaineana} - - 0.07 - - -

\emph{Anthocleista} sp. - - - 0.28 - -

\emph{Anthonotha fragrans} - - - - - 0.41*

\emph{Anthonotha macrophylla} - - - 0.00 - -

\emph{Antiaris toxicaria} 0.11 - - 0.37 - 0.15

\emph{Antrocaryon micraster} - - - - - 0.37*

\emph{Baphia nitida} 0.29 0.10 0.39* 0.34 0.15 0.28

\emph{Baphia pubescens} 0.16 - - 0.25 0.69* 0.12

\emph{Berlinia confusa} 0.12 0.33* 0.09 - 0.54* -

\emph{Berlinia occidentalis} - - - - - 0.41*

\emph{Blighia sapida} - - - 0.15 0.39* 0.27

\emph{Bombax buonopozense} - - - 0.34 - -

\emph{Buchholzia coriacea} - - - - 0.32 -

\emph{Bussea occidentalis} - - - 0.26 0.82* -

\emph{Calpocalyx} \emph{brevibracteatus} - - - 0.27 0.18 0.33

\emph{Carapa procera} - - 0.42* 0.28 0.65* 0.31

\emph{Canarium schweinfurtii} - - - 0.21 - -

\emph{Cedrela odorata} 0.33* 0.19 0.24 0.00 - -

\textbf{APPENDIX 4} (Continued)

ASENANYO SUHUMA

Tree species Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Ceiba pentandra} - 0.43* 0.65* 0.17 0.14 -

\emph{Celtis adolfi-friderici} - 0.50* - 0.24 - -

\emph{Celtis mildbraedii} 0.16 0.27 0.29 0.18 0.14 0.17

\emph{Celtis philippensis} Blanco - - - 0.12 - -

\emph{Celtis zenkeri} - - - 0.11 - -

\emph{Chrysophyllum perpulchrum} - - 0.42* - - -

\emph{Chrysophyllum subnudum} 0.48* 0.38* - - - -

\emph{Cola chlamydantha} - - - 0.34 0.46* 0.12

\emph{Cola gigantea} - 0.46* 0.09 - 0.37* 0.52*

\emph{Cola nitida} 0.14 - - - - -

\emph{Corynanthe pachyceras} - - 0.66* - 0.08 0.23

\emph{Cuviera nigrescens} - 0.34* - - - -

\emph{Dacryodes klaineana} - - - 0.80* - 0.41*

\emph{Daniela} sp. - - - - - 0.31

\emph{Desplatsia} sp. 0.73* - - - 0.39* -

\emph{Dialium aubrevillei} - - - 0.37 0.20 0.45*

\emph{Diospyros sanza-minika} - - - 0.28 - 0.41*

\emph{Diospyros gabunensis} - - - - 0.32 0.71*

\emph{Diospyros kamerunensis} - - - - - 0.60*

\emph{Discoglypremna} sp. - 0.17 - - - -

\emph{Distemonanthus} sp. - - - - 0.65* -

\emph{Drypetes gilgiana} - 0.11 - - - -

\emph{Drypetes principum} - - - 0.21 - -

\emph{Elaeis guineensis} Jacq. - - 0.25 - - -

\textbf{APPENDIX 4} (Continued)

ASENANYO SUHUMA

Tree species Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Entandrophragma angolense} 0.94* 0.22 0.38* 0.16 - 0.33

\emph{Entandrophragma cylindricum} 0.23 - - - - -

\emph{Entandrophragma utile} 0.21 0.89* 0.25 0.27 - -

\emph{Ficus exasperata} - - - 0.34 0.18 -

\emph{Ficus sur} 0.51* - - - - -

\emph{Funtumia africana} 0.09 - 0.37* - - 0.64*

\emph{Funtumia elastica} - - - 0.60* - -

\emph{Guarea thompsonii} - 0.30 - - - 0.31

\emph{Guibourtia ehie} - 0.11 - - - -

\emph{Hannoa klaineana} - - - 0.37* - -

\emph{Hevea brasiliensis} - - - 0.23 - -

\emph{Hexalobus crispiflorus} - 0.27 - - - -

\emph{Homalium dewevrei} - - - - - 0.40*

\emph{Hymenostegia afzelii} 0.22 0.26 0.77* 0.28 - 0.23

\emph{Irvingia gabonensis} 0.45* - 0.28 - - 0.32

\emph{Lannea welwitschii} - 0.59* - 0.38* - 1.00*

\emph{Lonchocarpus sericeus} 0.37* - 0.30 - - -

\emph{Lovoa trichilioides} - - - - 0.24 -

\emph{Macaranga barteri} - - - 0.00 - -

\emph{Mammea africana} - - 0.55* - - -

\emph{Margaritaria discoidea} - - - 0.64* - -

\emph{Microdesmis puberula} 0.22 - - 0.19 0.18 -

\emph{Milicia excelsa} - - - - - 0.40*

\emph{Musanga cecropioides} - - - 0.42* 0.38* -

\emph{Massularia acuminata} - - - - 0.25 -

\textbf{APPENDIX 4} (Continued)

ASENANYO SUHUMA

Tree species Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Myrianthus arboreus} - - - 0.43* 0.56* 0.54*

\emph{Myrianthus libericus} - - - 0.20 0.36 0.40*

\emph{Napoleonaea vogelii} 0.31* - - - 0.18 -

\emph{Nauclea diderrichii} - - 0.25 - - -

\emph{Nesogordonia papaverifera} 0.26 0.00 0.16 0.22 0.30 0.25

\emph{Newbouldia laevis} 0.56* - 0.00 - - -

\emph{Omphalocarpum ahia} - - - - 0.39 -

\emph{Pachystela msolo} - - - - - 0.54*

\emph{Pachystela brevipes} - - - - 0.00 -

\emph{Parinari excelsa} - - - 0.32 - -

\emph{Parkia bicolor} - - - - - 0.37*

\emph{Pentaclethra} sp. - - - - - 0.00

\emph{Petersianthus macrocarpus} 0.53* - - - - -

\emph{Piptadeniastrum africanum} 0.87* 0.19 0.45* 0.46* 0.41* 0.23

\emph{Pterygota macrocarpa} - - - 0.58* 0.29 0.20

\emph{Pycnanthus angolensis} 0.72* 0.39* 0.49* - 0.35 0.23

\emph{Rauvolfia vomitoria} 0.63* - 0.21 0.59* 0.37* -

\emph{Ricinodendron heudelotii} 0.54* 0.47* - 0.22 0.26 0.28

\emph{Rinorea} sp. - - - 0.86* - -

\emph{Scottellia klaineana} - - 0.18 0.33 1.00* 0.69*

\emph{Sterculia oblonga} - - 0.49* 0.37* 0.29 -

\emph{Sterculia rhinopetala} - - - 0.16 0.05 -

\emph{Sterculia tragacantha} 0.13 - - 0.23 - -

\emph{Strombosia pustulata} 0.18 0.22 0.23 0.79* 0.09 0.28

\emph{Synsepalum aubrevillei} - - - 0.00 - -

\textbf{APPENDIX 4} (Continued)

ASENANYO SUHUMA

Tree species Edge Interior Deep-interior Edge Interior Deep-interior

\emph{Terminalia ivorensis} - - - 0.58* - -

\emph{Terminalia superba} - 0.17 - - - 0.43*

\emph{Tetrapleura tetraptera} 0.10 - - - - -

\emph{Treculia africana} - - - - 0.07 -

\emph{Trichilia monadelpha} 0.12 0.11 0.29 0.41* 0.35 0.31

\emph{Trichilia prieuriana} 0.23 - 0.00 0.11 0.20 0.17

\emph{Trichilia tessmannii} - - - - 0.37* -

\emph{Tricalysia disco} lor - - 1.00* - - -

\emph{Trilepisium} sp. 0.18 - - 0.12 0.45* 0.00

\emph{Triplochiton scleroxylon} 0.10 - 0.22 0.55* - -

\emph{Turraeanthus africanus} - - - - 0.34 0.40*

\emph{Uapaca corbisieri} 0.90* - - - - -

\emph{Zanthoxylum gilletii} 0.13 - 0.25 0.38* - -

\emph{Zanthoxylum parvifoliolum} - - - 0.78* - -\selectlanguage{english}
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