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How are arthopod communities structured and why are they so diverse? Answers from Mediterranean mountains using hierarchical additive partitioning

DOI:10.1007/s10531-017-1303-2
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Konstantina Zografou at University of Ioannina
Konstantina Zografou
Robert J Wilson at The National Museum of Natural Sciences
Robert J Wilson
John M. Halley at University of Ioannina
John M. Halley
Elli Tzirkalli at Open University of Cyprus
Elli Tzirkalli
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Abstract and Figures

Mountains are complex ecosystems supporting a great variety of taxa. Here, we explored the diversity patterns of arthropods in two mountains, pinpointing the spatial scale that accounts most for overall diversity variation, using an additive partitioning framework. Butterflies and Orthoptera were sampled in Rodopi (2012) and Grammos (2013) mountains. Diversity was partitioned into five hierarchical levels (mountain, elevational zone, habitat, transect and plot). We compared the estimated diversity values for each level to the respective permuted values expected by chance, for all species, as well as for species identified as “rare” or “common”. At broader spatial levels, the variation in total diversity was attributed to the beta diversity component: mountains accounted for 20.94 and 26.25% of butterfly and Orthoptera diversity, and elevational zones accounted for 28.94 and 35.87% respectively. At finer spatial scales, beta diversity was higher than expected by chance in terms of the Shannon index. The type of habitat was found to play a significant role only for rare orthopterans. Finally, common species were recognized for shaping overall species diversity. We highlight the importance of the spatial levels of elevation zone and then mountain position in conservation planning, due to the greater beta diversity recorded at this scale as compared to habitat or more finite scales. Monitoring programs might need to adapt different strategies with respect to the focal organisms, and consider patterns of common rather than rare species that found to drive the patterns of the entire community.
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ORIGINAL PAPER
How are arthopod communities structured and why are
they so diverse? Answers from Mediterranean mountains
using hierarchical additive partitioning
Konstantina Zografou
1
•Robert J. Wilson
2
•
John M. Halley
1
•Elli Tzirkalli
1
•Vassiliki Kati
1
Received: 15 May 2016 / Revised: 2 December 2016 / Accepted: 24 January 2017 /
Published online: 31 January 2017
ÓSpringer Science+Business Media Dordrecht 2017
Abstract Mountains are complex ecosystems supporting a great variety of taxa. Here, we
explored the diversity patterns of arthropods in two mountains, pinpointing the spatial scale
that accounts most for overall diversity variation, using an additive partitioning framework.
Butterflies and Orthoptera were sampled in Rodopi (2012) and Grammos (2013) moun-
tains. Diversity was partitioned into five hierarchical levels (mountain, elevational zone,
habitat, transect and plot). We compared the estimated diversity values for each level to the
respective permuted values expected by chance, for all species, as well as for species
identified as ‘‘rare’’ or ‘‘common’’. At broader spatial levels, the variation in total diversity
was attributed to the beta diversity component: mountains accounted for 20.94 and 26.25%
of butterfly and Orthoptera diversity, and elevational zones accounted for 28.94 and
35.87% respectively. At finer spatial scales, beta diversity was higher than expected by
chance in terms of the Shannon index. The type of habitat was found to play a significant
role only for rare orthopterans. Finally, common species were recognized for shaping
overall species diversity. We highlight the importance of the spatial levels of elevation
zone and then mountain position in conservation planning, due to the greater beta diversity
recorded at this scale as compared to habitat or more finite scales. Monitoring programs
might need to adapt different strategies with respect to the focal organisms, and consider
patterns of common rather than rare species that found to drive the patterns of the entire
community.
Communicated by Jan C. Habel.
Electronic supplementary material The online version of this article (doi:10.1007/s10531-017-1303-2)
contains supplementary material, which is available to authorized users.
&Konstantina Zografou
ntinazografou@yahoo.co.uk
1
Department of Biological Applications and Technology, University of Ioannina, Ioannina, Greece
2
College of Life and Environmental Sciences, University of Exeter, Exeter EX4 4PS, UK
123
Biodivers Conserv (2017) 26:1333–1351
DOI 10.1007/s10531-017-1303-2
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
... This suggests that landscape attributes were more favourable for the specialist group than the forest type. Greater beta diversity was observed at the elevation zone scale compared to habitat or ner scales in this study, supported by (Zografou et al. 2017). Furthermore, habitat specialists species were predominantly concentrated at higher altitudes and generalists dropped during severe land use changes associated with aridity and specialists were negatively impacted by temperature. ...
... One 500 m transect was placedin each forest type (Open forest: OF, Riparian forest: RF, and Dense forest: DF) in each study site. The mean distance between transects in each site was 1.5 ± 0.25 km and the mean distance among study sites was 39 ± 8.64 km, so each transect represented an independent sample(Zografou et al. 2017). ...
... An additive partitioning framework expressed by species richness and Shannon-Wiener index (H') has been used in this study to test the null hypothesis that butter y diversity is uniform in all spatial scale(Zografou et al. 2017). The pooled data from all sample months were aggregated by three spatial scale, i.e., transects (144 units), forest types(12 units), and elevation-based study sites (landscape level, 4units) in order to access the species diversity. ...
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To understand the effects of local landscape factors on functional species composition and phenology of butterflies across multiple spatial scales, a study was carried out in a tropical dry forest of the northern highlands of the Eastern Ghats of India from November 2016 to October 2017. A total of 3343 individuals of butterflies were recorded, including 88 species of butterflies under 62 genera, 18 subfamilies, and 6 families in three different forest types (open, riparian, dense). Butterfly species richness showed no significant deviations, but diversity patterns varied across transects. Beta diversity indicated differences in common species populations, likely due to uneven resource distribution in study site forests. The contribution of β transect to gamma diversity was greater than that of β elevation, except for specialists. Specialists were favoured by landscape attributes over forest type. Butterfly abundance peaks in April for open and dense forests, and May for riparian forests. Results show variation in seasonal patterns across different forest types (F = 15.92, P < 0.001). Generalists and versatilists are more prevalent in April and February, while specialists are more abundant from October to November. Relative humidity, shrub density, and temperature were the major contributors (40.2%) for richness whereas relative humidity and shrub density contributed 26.3% for abundance. The relative humidity was predominant over temperature for species richness and is a major predictor for assemblages of generalist species. Resource utilization based on elevation plays an important role for habitat specialist species and highlights the importance of the spatial levels of elevation zones in conservation planning.
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... 20°50′, lat. 40°21′; maximum elevation 2,520 m) is located in NW Greece ( Figure S1, Table S1, but see also Zografou, Wilson, Halley, Tzirkalli and Kati (2017) for detailed descriptions). Both systems share a low human population density and associated low-intensity human activities, as well as high coverage by protected areas of the Natura 2000 network. ...
... The minimum distance between nearest neighboring sites was approximately 2 km (SD ± 0.5) so that each site effectively represents an independent sampling unit. The lack of spatial autocorrelation between nearby sites was verified in terms of alpha and beta components of diversity in a previous study where we investigated diversity patterns of butterflies and Orthoptera across different spatial scales (Zografou et al., 2017). Each mountain was partitioned into four elevation zones (0-500 m, 501-1,000 m, 1,001-1,500 m, and 1,501-2,000 m) and each zone contained sites representing the three dominant habitats found in the study system (agricultural fields, grasslands, and forests), with the exception that agricultural areas were not present above 1,500 m ( Figure S1, Table S1). ...
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... Relatively large mountain ranges with diverse geological structure and climate (from Continental to Mediterranean) make south-eastern Europe an interesting region for studies of elevation gradients in species diversity. At the same time, data on butterfly diversity in this region is scattered in a multitude of faunistic papers, while few studies address the butterfly diversity pattern (Mihoci et al., 2011;Zografou et al., 2014Zografou et al., , 2017Kaltsas et al., 2018). Similarly, the effect of aspect is only rarely taken into consideration when addressing the distribution patterns of butterflies in mountains (Gutiérrez, 1997;Mihoci et al., 2011). ...
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... Evidence from insect distributions appears to contradict the distance limitation hypothesis. The spatial variation in species composition is higher among nearby bioclimatic belts along elevation gradients than among mountain systems separated by large distances (e.g., butterflies and orthopterans in Grammos and Rodopi separated by 426 km: Zografou et al. 2017; ground-dwelling beetles on mountains in north-western Patagonia separated by 40-100 km: Werenkraut and Ruggiero 2013; ants in Hengduan mountains, China: Liu et al. 2017). These studies suggest that, on a regional scale, the variation in mountain insect assemblages may be associated more with adaptation of species to local environmental conditions than by dispersal limitation imposed by distance (e.g., ants: Liu et al. 2017). ...
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