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ISSN : 2288-1115(Print)
ISSN : 2288-1123(Online)
Korean Journal of Ecology and Environment Vol.50 No.4 pp.478-482

Co-occurrence Patterns of Bird Species in the World

Young Min Kim, Sungwon Hong, Yu Seong Lee1, Ki Cheol Oh2, Gu Yeon Kim3, Gea-Jae Joo*
Department of Integrated Biological Science, College of Natural Sciences, Pusan National University, Busan 46241, Republic of Korea
1Department of Biological Sciences and Biotechnology, Hannam University, Daejeon 34430, Republic of Korea
2Nakdong River Basin Environmental Office, Changwon 30103, Republic of Korea
3Department of Science Education, College of Natural Sciences, Kyungnam University, Masan 51767, Republic of Korea
Corresponding author : +82-51-510-3344, +82-51-583-0172,
20171211 20171218 20171222


In order to identify key nations and bird species of conservation concern we described multinational collaborations as defined using network analysis linked by birds that are found in all nations in the network. We used network analysis to assess the patterns in bird occurrence for 10,422 bird inventories from 244 countries and territories. Nations that are important in multinational collaborations for bird conservation were assessed using the centrality measures, closeness and betweenness centrality. Countries important for the multinational collaboration of bird conservation were examined based on their centrality measures, which included closeness and betweenness centralities. Comparatively, the co-occurrence network was divided into four groups that reveal different biogeographical structures. A group with higher closeness centrality included countries in southern Africa and had the potential to affect species in many other countries. Birds in countries in Asia, Australia and the South Pacific that are important to the cohesiveness of the global network had a higher score of betweenness centrality. Countries that had higher numbers of bird species and more extensively distributed bird species had higher centrality scores; in these countries, birds may act as excellent indicators of trends in the co-occurrence bird network. For effective bird conservation in the world, much stronger coordination among countries is required. Bird co-occurrence patterns can provide a suitable and powerful framework for understanding the complexity of co-occurrence patterns and consequences for multinational collaborations on bird conservation.


    the Busan Green Environment Center


    Birds have long been a major focus of public attention and research interest. In particular, multinational collaboration is now a major driver in bird conservation (Donald et al., 2007). Amongst researchers and environmental groups, the consensus about bird conservation is that a comprehensive and cooperative international effort is needed to conserve viable bird populations (Williams et al., 2013). Effective bird conservation globally will require far greater coordination amongst countries (Donald et al., 2007). The Individualistic implementation of bird conservation policies may fail if previously identified interactions within the network between countries are ignored (Runge et al., 2015), also contributing to the limited success of bird conservation at the national level, conservation policies are rarely implemented in a uniform manner and research interest is often patchy due to its dependency upon the researcher.

    Although countries located along migratory routes have established networks for the protection of migratory birds, based on flyway concepts that emphasize political and governmental processes in multinational collaboration, non-migratory birds are not covered effectively in traditional flyway models. Spatial co-occurrences for both migratory and non-migratory birds, may reveal complex interactions in a network spanning many countries, based on the geographic patterns of bird co-occurrence among countries (Böhning- Gaese and Oberrath, 1999; Webb et al., 2002). The complexity of direct and indirect interactions linking birds amongst countries is so vast that their complete documentation is beyond reach. Using network analysis to understand the drivers affecting the distributional dynamics that facilitate bird conservation globally, as well as enable multinational collaboration, requires investigation using alternative approaches.

    We use an approach based on network analysis to identify potential broad-scale interactions between birds from different countries, based on the geographical patterns of bird cooccurrence. Network analysis has traditionally had a strong synergy with business models in certain industries (Proulx et al., 2005), but we believe that it could be used to identify direct and indirect relationships linking birds from countries by analyzing the structure in the network (Scott, 2012).

    Our specific objectives were to (i) propose multinational collaboration groups, defined by network analysis, that are linked by co-occurring birds, and (ii) to identify the key countries and birds of conservation concern in each network. First, we hypothesized that countries and birds with the potential to affect other countries and species will have higher closeness centrality scores. They can provide the clearest perspective of the state of the network. Second, we predicted that countries and birds that are important to the cohesiveness of the network would have a higher betweenness centrality score. Together, these indices can represent how each country can conduct bird conservation efforts independently and effectively relative to others.


    1.Data collection

    We used 10,422 bird inventories from 244 countries and territories obtained from BirdLife International (2015, The cumulative total contribution (in US Dollars) for 193 countries from 1973 to 2013 to the Environment Fund of UNEP (United Nations Environment Programme) was used to indicate conservation effort in each country including bird conservation.

    2.Network analysis

    We used network analysis to examine the patterns of bird co-occurrence and the structure and level of bird connectedness. Network analysis uses a set of procedures to identify and measure the structural properties of social systems, based on relationships among entities in the system rather than on characteristics of the entities. A network can be thought of as a set of nodes with connections or links between them. In our study, the nodes are countries and birds while the links are undirected connections between each country and species.

    Centrality indices describe the importance of specific nodes within the network (Newman, 2010). Different centrality indices measure different aspects related to the position of a node within its network. Closeness centrality expands the definition of degree by focusing on the distance between nodes. A central node is thus characterized in the networks by numerous short connections. Betweenness centrality represents a different aspect of centrality: based on the number of times a particular node is found on the shortest path between any pair of nodes in the network. Nodes that are highly central control information flow in the network and thus receive a higher score of betweenness centrality. A social network analysis and visualization software application, Gephi (which uses the ForceAtlas 2 algorithm as a type of force-directed layout algorithm, ver 0.9.2), was used to determine the position of the participants and develop network diagrams (sociograms) that indicate the relationship ties and information flow between the individual nodes. The different centralities among the groups were divided using the modularity index and were tested using a one-way ANOVA and Duncan’s post-hoc test.


    1.Bird co-occurrence

    The average number of bird species for the 244 countries was approximately 403 species. The median number of birds per country was 311 species. Countries in South America, such as Columbia (1,827 species), Peru (1,807 species), and Brazil (1,753 species), had the highest number of bird species (Fig. 1a). In Asia, Indonesia (1,615 species) also showed a high number of bird species. A total of 7,586 species occurred in more than two countries, while 2,836 species (27.2% of total bird species) were only recorded in one country. Arenariainterpres (221 countries), Hirundorustica (213 countries), Falco peregrinus (210 countries), Numeniusphaeopus (209 countries), Calidrisalba (205 countries), and Pandion haliaetus (202 countries) had the broadest distributions across the world.

    2.Bird network

    The country-bird network consisted of 10,585 nodes and 97,419 edges (average degree, or node, connectivity=18.4, Fig. 2). The average network distance between all node pairs (average path length) was 3.63 (the longest distance was 6 edges). The modularity index was 0.596 (values >0.4 suggest that the network has a strong modular structure).

    Nodes were divided into 4 groups based on their modularity. Co-occurrence networks for four groups of birds reveal different relationships with biogeographical organization (Fig. 2). Group 0 (G0) accounted for 8.68% of the network; it included 93 countries and 826 species in Europe, North Africa, the Middle East, and East Asia. Group 1 (G1) consisted of 60 countries and 4,220 species in America. G1 was the largest group in the network, with 40.44% of total nodes and edges. Group 2 (G2) accounted for 34.59% of the network, and included 48 countries and 3,613 birds in Asia, Australia and the South Pacific. Group 3 (G3) accounted for 16.29% of the network, and included 43 countries and 1,681 species in Sub-Saharan Africa.

    Fig. 1c and 1d show that several countries within middle latitudes had higher centrality scores for both closeness and betweenness centrality. Closeness centrality of countries in G3 was significantly higher than in the other groups (Post-hoc test, P<0.001) while betweenness of countries in G2 were higher than in the other groups (Post-hoc test, P<0.05). Pakistan (G0), Brazil (G1), Indonesia (G2), and Congo (G3) had the highest closeness centrality in each group. Highest betweenness centralities were recorded in Japan (G0), Brazil (G1), Indonesia (G2), and Tanzania (G3) in each group.

    Among all bird species, closeness and betweenness centralities of G0 were significantly higher than they were for the other groups (Post-hoc test, P<0.001). Porphyrio porphyria, Dendrocygnaviduata, Cisticolajuncidis, Elanuscaeruleus, and Rostratulabenghalensis had the highest scores for closeness centrality. Comparatively, D. viduata, P. porphyria, Nettaerythrophthalma, Laruscirrocephalus, and C. juncidis had the highest scores for betweenness centrality. Arenariainterpres (G0), Butoridesstriata (G1), Fregata minor (G2), and P. porphyrio (G3) had the highest values for closeness centrality for each each group. A. interpres (G0), B. striata (G1), F. minor (G2), and D. viduata (G3) had the highest values for betweenness centrality for each each group.

    The allocation to the Environment Fund of the UNEP from each country was not an appropriate measure of the importance of each country, based on centralities in the network. Many countries with high-level centrality values made small contributions (Fig. 3). This result presents a paradox and emphasizes the importance of multinational collaboration in bird conservation (Donald et al., 2007; Butchart et al., 2010).


    The present study identifies groups of birds that can be targeted for multinational collaboration, using network analysis. In addition, we identify several countries and bird species that provide important links and are crucial for multinational bird conservation in the four regional groups and globally. The geographical distribution of the groups, based on bird co-occurrence patterns, corresponds with large zoogeographic regions such as Nearctic, Neotropical, African, Palearctic, Oriental, and Australian regions. The zoogeographic region construct has already been used to highlight areas of the world that are most distinctive or represent high “value” and are, therefore, worthy of greater attention (Olson and Dinerstein, 1998; Olson et al., 2001). A large multinational group can be more appropriately conceptualized as an international bird conservation group rather than a representation of local conservation (Sodhi et al., 2011). Each group in the network may overcome a limitation of the migratory bird flyway concept, in which east-west migration is not well-recognized (Favell, 2008). However, conservation policies that are implemented across large biotic regions or hotspots often fail to discern smaller but highly distinctive areas; this can result in these areas receiving insufficient conservation attention (Olson et al., 2001). Bird conservation at the local level should also be strongly recommended, as careful attention to bird habitat may be improve the efficiency of conservation efforts.

    Countries and species with high scoring closeness and betweenness centrality may act as excellent sentinels or indicators of trends in co-occurrence bird networks, although countries in mid-latitudes had high centralities that strongly correlated with the number of species. Bird species with high score centralities had extensive distributions across the world. Generalist birds that have broad distribution with large population are often not important issues in bird conservation. However, these species could be important for identifying the current state of bird diversity and conservation effectiveness at the local level, for these species have the great role of connecting habitats and countries (Cassey et al., 2004).

    Sustained economic support for coherent global bird conservation is essential to improve the effectiveness of these responses. We believe that a bird co-occurrence network provides a suitable and powerful framework to address the complexities of co-occurrence patterns and can pave the way to improving worldwide bird conservation.


    This work was supported by the research fund of the Busan Green Environment Center (15-1-70-71).



    Maps of (a) the total number of bird species, (b) groups based on modularity from network analysis, (c) closeness centrality and (d) betweenness centrality of countries.


    The network diagram for countries and birds. Each node (dot) represents a country or a bird. Nodes of same color belong to the same group. An edge (line) represents the relationship between two individuals.


    Relationships between centralities and their contributions to the Environment Fund of UNEP, which represent bird conservation efforts in each country.



    1. (2015) datazone/country
    2. Böhning-GaeseK. OberrathR. (1999) Phylogenetic effects on morphological, life-history, behavioural and ecological traits of birds. , Evol. Ecol. Res., Vol.1 ; pp.347-364
    3. ButchartS.H. WalpoleM. CollenB. Van StrienA. ScharlemannJ.P. AlmondR.E. BaillieJ.E. BomhardB. BrownC. BrunoJ. (2010) Global biodiversity: indicators of recent declines. , Science, Vol.328 ; pp.1164-1168
    4. CasseyP. BlackburnT.M. SolD. DuncanR.P. LockwoodJ.L. (2004) Global patterns of introduction effort and establishment success in birds. , Proc. R. Soc. Lond. B Biol. Sci., Vol.271 ; pp.S405-S408
    5. DonaldP.F. SandersonF.J. BurfieldI.J. BiermanS.M. GregoryR.D. WaliczkyZ. (2007) International conservation policy delivers benefits for birds in Europe. , Science, Vol.317 ; pp.810-813
    6. FavellA. (2008) The new face of East-West migration in Europe. , J. Ethn. Migr. Stud., Vol.34 ; pp.701-716
    7. NewmanM. (2010) Networks: an introduction., Oxford University Press,
    8. OlsonD.M. DinersteinE. (1998) The Global 200: a representation approach to conserving the Earth ?(tm)s most biologically valuable ecoregions. , Conserv. Biol., Vol.12 ; pp.502-515
    9. OlsonD.M. DinersteinE. WikramanayakeE.D. BurgessN.D. PowellG.V. UnderwoodE.C. D’amicoJ.A. ItouaI. StrandH.E. MorrisonJ.C. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth A new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. , Bioscience, Vol.51 ; pp.933-938
    10. ProulxS.R. PromislowD.E. PhillipsP.C. (2005) Network thinking in ecology and evolution. , Trends Ecol. Evol., Vol.20 ; pp.345-353
    11. RungeC.A. WatsonJ.E. ButchartS.H. HansonJ.O. PossinghamH.P. FullerR.A. (2015) Protected areas and global conservation of migratory birds. , Science, Vol.350 ; pp.1255-1258
    12. ScottJ. (2012) Social network analysis., Sage,
    13. SodhiN.S. ButlerR. LauranceW.F. GibsonL. (2011) Conservation successes at micro-, meso-and macroscales. , Trends Ecol. Evol., Vol.26 ; pp.585-594
    14. WebbC.O. AckerlyD.D. McPeekM.A. DonoghueM.J. (2002) Phylogenies and community ecology. , Annu. Rev. Ecol. Syst., Vol.33 ; pp.475-505
    15. WilliamsD.R. (2002) Bird Conservation: Global evidence for the effects of interventions, Pelagic Publishing, Vol.Vol 2