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Data analysis

All analysis was done separately for the whole Himalayan extent and separately for five
regions: Western, Central, Eastern, North-western, and Trans Himalayas [Fig. 1(a)]. We
assessed the occurrences of the passerine birds, and we furtherly looked for the
availability of the traits of species. Species with any missing trait values were removed
from the analysis, which finally 408 passerine species were selected in total.

Functional groups
A set of standard attributes compiled for all traits and grouped into eight functional
groups (Table S1).

Allocation of the bird species to functional groups


We assessed the functional requirements of each species for each trait and coded each
attribute for any of seven functional groups (Luck et al., 2012; Luck et al., 2013). We
constructed a matrix of 408 passerine birds with all selected traits (43 traits). The
occurrences of bird species were converted into the presence and absence matrix and sum
of the occurrence point of each species considered as the abundance of species in the
Himalayan region.
We calculated the FD for all traits collectively as well as for all seven functional
groups separately (see Table S1). These functional groups were directly related to the
species and demonstrated the resource utilization capacity of species and displayed their
response (e.g., morphological, feeding, reproductive) to any change in their environment
(Luck et al., 2015). The functional group diversity is more meaningful than species
diversity, and it is associated with higher long-term stability (O'Gorman et al., 2010;
Cadotte et al., 2011). We used a functional dendrogram to map the pattern of the FD
along the Himalayas. The functional dendrogram displays the relationship shared by the
species, and the FD is the total length of the functional dendrogram. This dendrogram is
essential to divide species by functional groups and guilds (Petchey & Gaston, 2007).

Estimates of Passerine diversity in relation to geographical regions of Himalayas


For taxonomic diversity, only the species richness (SR= the total number of species)
(Magurran, 2004) was considered for per grid cell as well as for each geographical
regions, (Western, Central, Eastern, North-western and Trans Himalayas) separately.
Both categories were compared among geographical strata by using analysis of variance
(ANOVA).
Estimates of FD
A species-by-trait matrix was built and to improve the comparability across the traits; the
matrix was standardized by linearly rescaling the traits from min-to-max, 0=minimum,
1=maximum. Further, the species-by-trait matrix was converted into the distance matrix
referencing Gower (1971). The four indices of FD, computed in R software using the
“FD” package (Laliberté et al., 2014; RCoreTeam, 2017) are briefly explained here
below: (1) FRic consists of the number of various functional traits that occur in a
community (Petchey & Gaston, 2002), FRic actively represents the relation with the SR.
However, it was commonly used in many studies, (2) functional evenness denotes the
aura of evenness in the abundance of species with particular traits in a community
(Villéger et al., 2008), (3) FDiv embodies the distribution of the trait value across trait
space and is constrained between 0 & 1 (Laliberte et al., 2010). (4) FDis is the pattern of
trait values within trait space measured as the distance of the centroid value (Laliberte et
al., 2010).
Community aggregated or community-weighted mean (CWM) values of
functional traits are used to compare the functional distance between communities; in
both cases, trait values are weighted according to the relative abundance of species
(Garnier et al., 2004; Májeková et al., 2016). Functional distances between all individuals
within a local community were calculated using the function “gowdis” in the R package
“FD”, which computes the Gower dissimilarity from different trait types [continuous,
ordinal, nominal, or binary (Laliberté et al., 2014)].
A detailed description of the methodology and data analysis is available in the
supplementary materials. One functional group, population status was removed from the
final analysis, because of the lack of accurate data on the population status of forty
species.

RESULTS
The SR pattern for 408 passerine birds across geographical regions of Himalayas was
high in central, followed by eastern, north-western, western, and trans-Himalayas [Fig.
1(b)]. While the FD ranked high in the eastern and low in the western Himalayas and
varied patterns are observed for each functional group.

Relationship between SR and FD


The correlation between SR and FD observed negatively correlated for the cumulative
traits. However, the correlation between SR and the components of FD varied differently.
When comparing each functional group separately for SR with FD, a positive correlation
was observed between SR and FRic, for all traits, i.e., morphological, clutch size, dietary
guild, habitat guild, nest height, and nest type, whereas FEve was found negatively
correlated with SR for clutch size, dietary guild, habitat guild, nest height, and nest types;
and showed positive correlation only for morphological traits. The positive correlation
between FDis and SR was observed for clutch size, dietary guild, habitat guild, and Nest
height, whereas we obtained a negative correlation between FDis and SR for
morphological and nest type. The negative correlation obtained between SR and FDiv for
clutch size, dietary guild, and habitat guild, whereas morphological traits, nest type, and
nest height were correlated positively (Table 3).

Patterns of SR and FD across Himalayan regions


The pattern of SR observed different from that of the FD (Fig. 5) [Table 1]. When
compared for cumulative traits and with each functional group, the following patterns
were observed for FD and SR in the study region:

High FRic for morphological traits in Central Himalayas directs the presence of a
diverse population with varying body mass and body length.
Significant variation in FD indices for the dietary guild and the lowest FD in the trans-
Himalayas seemed to be associated with the high availability of resources, as the trans-
Himalayas had less diverse faunal composition than other regions, food resources were
pivotal for trait diversity. Several other processes, including evolutionary innovations in
dietary preference and environmental constrained on the distribution of the food
resources (Kissling et al., 2012) thus defined the patterns of the FD. Our results were
concordant with those of other studies concerning habitat fragmentation effects on FD
(Magnago et al., 2014; de Frutos et al., 2015; Liu et al., 2018).

A higher FRic for flocking preference in the central Himalayas may be associated with
the higher competition for the available resources among species as well as the higher
complexity of the resources would help to support more species to provide a more stable
community.

Past studies also found an incremental pattern in land-use intensity that was
directly proportional to functional dispersion (Laliberte et al., 2010; Luck et al., 2013),
which means that more disturbed sites have higher measures of functional dispersion.
Pechony & Shindell (2010) suggested that the temperature-driven disturbance regimes
were changing at unprecedented rates globally, and the consequences of FD were not
known clearly.

Our study confirmed that the patterns of functional diversity was different from the
species richness and observed negatively correlated for functional traits. Each functional
group showed different response with species richness in the region, which might be
attributed to a unique set of traits, i.e., morphological, reproductive, and life history. It
may also be attributed that the passerine assemblages have every possible combination
for trait values that are more sensitive to ecological processes and thus maintaining the
functional diversity in the region. Which is consistent with the insurance theory (Yachi &
Loreau, 1999) that stated the species with high trait diversity were required to ensure the
functionality of the ecosystems. Although no single component of functional diversity
could explain its patterns in geographic regions of Himalayas, we found that each trait
played a specific and vital role in shaping the functional diversity patterns in each region.
Eastern Himalayas exhibited the highest functional diversity, while the western
Himalayas appeared to be more consistent with the functional evenness. However, the
diversity of species may be required to carry ecological processes in an ecosystem as
different species support different process at different times (Walker et al., 1999).

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