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

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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.
... The relative abundance is defined as the percentage of each species contributed to the total number of individuals of all species (Magurran 2004). In addition, an additive partitioning framework has been used in this study to test the null hypothesis that butterfly diversity is uniform in all spatial scales i.e., transects, forest types, and elevation-based study sites (Zografou et al. 2017). ...
... This suggests that landscape attributes were more favourable for the specialist group than the forest type. Greater β diversity was observed at the elevation zone scale compared to habitat or finer scales in this study, supported by the findings of Zografou et al. (2017). Similar results were also found from studies carried out in this region (Mahata et al. 2019a(Mahata et al. , 2023Mahata and Palita 2023), which reported that the species richness in the eastern high-elevation ranges (Koraput Plateau), is higher than that of the western low elevation zone (Jeypore Plateau). ...
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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 88 species of butterflies were recorded, under 62 genera, 18 subfamilies, and six 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. Elevation-dependent resource utilization is crucial for habitat specialists, underscoring the significance of spatial elevation zones in effective conservation planning strategies.
... The change in vegetation types along elevation gradients often affects the composition of species, leading to differences in fauna between low and high elevations (e.g., Aisen et al., 2017;González-Reyes et al., 2017;García-L opez et al., 2012;Liu et al., 2018;Werenkraut & Ruggiero, 2013). Each vegetation type may be part of a unique biogeographic unit or eco-region representing a macro-habitat for insects, which is why the composition of species assemblages on summits of different mountains can be more similar to each other than that with lower vegetation belts of the same mountain (e.g., Aisen et al., 2017;Liu et al., 2018;Werenkraut & Ruggiero, 2013;Zografou et al., 2017). Changes in vegetation structure are often associated with differences in plant biomass/ production and diversity of functional attributes (Sundqvist et al., 2013), which affect the local climatic conditions and the resources available for insects. ...
... The impact of the forest-steppe transition on pre-and post-eruption beetle composition This study showed that the taxonomic composition of mountain epigaeic beetle assemblages is strongly structured along elevation gradients. The vertical division of mountains into different macro-habitats characterised by different vegetation types was more important than the geographical distance between mountains in shaping the taxonomic composition of beetle assemblages, consistent with findings in other mountain regions (Aisen et al., 2017;Escobar et al., 2005;Liu et al., 2018;Zografou et al., 2017). This pattern reflects a species sorting process (sensu Leibold et al., 2004;Liu et al., 2018), wherein environmental variation plays a more pivotal role than dispersal limitation in shaping taxonomic composition (see also Aisen et al., 2017;Werenkraut & Ruggiero, 2013). ...
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Monitoring changes in species composition near active volcanoes is important to identify environmental factors that maintain biological diversity after major disturbances. We investigated the taxonomic composition of epigaeic beetles' assemblages in five mountains in north‐western Patagonia (Argentina) in the aftermath of the 2011 Puyehue Cordón‐Caulle Volcanic Complex eruption. The associations of beetle composition with the thermal environment, vegetation structure and soil characteristics were analysed using 10 m × 10 m plots with nine pitfall traps each, established 100 m apart in altitude on each mountain in 2005, 2006 (pre‐eruption) and 2012, 2015 and 2016 (post‐eruption). We found significant differences in beetle composition between forests and high‐Andean steppes before and after the eruption. The beetle composition of forests and high‐Andean steppes differed due to higher plant cover and lower pH and herb richness in forests. Six months after the eruption, the herbs almost disappeared and forests showed high ash accumulation and lower temperature than the high‐Andean steppes. In the medium term, plant cover, temperature and soil conditions—but not ash accumulation—remained as important correlates of beetle composition. Before the eruption, forest assemblages showed higher beetle diversity and Leiodidae indicator species, but Curculionidae predominated post‐eruption. Tenebrionidae and Carabidae were high Andean steppes indicators. Post‐eruption, the beetle composition in high‐Andean steppes did not change significantly except for an increase in Tenebrionidae indicator species. We concluded that the conservation of landscape diversity and environmental gradients maintaining the differentiation between ecoregions is essential to protect epigaeic beetle composition and for species survival after volcanic disturbances.
... Butterflies were counted from 0900 to 1300 h via a transect walk method, with a constant space of 60 min for each transect. A transect of 500 m was surveyed in each sampling site (2 × 2 km 2 grids), with an average intervening distance of 12 km between grids, so that each transect represented an independent sample [36]. We counted butterflies for a six-month period during the dry season from December 15 to June 5 in the year 2019, as counting could not be undertaken during the wet season due to the cloudy skies and heavy downpours. ...
... Butterflies were counted from 0900 to 1300 h via a transect walk method, with a constant space of 60 min for each transect. A transect of 500 m was surveyed in each sampling site (2 × 2 km 2 grids), with an average intervening distance of 12 km between grids, so that each transect represented an independent sample [36]. We counted butterflies for a sixmonth period during the dry season from December 15 to June 5 in the year 2019, as counting could not be undertaken during the wet season due to the cloudy skies and heavy downpours. ...
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Understanding the factors that influence the diversity and distribution of butterfly species is crucial for prioritizing conservation. The Eastern Ghats of India is an ideal site for such a study, where butterfly diversity studies have yet to receive much attention. This study emphasized the butterfly assemblages of three prominent habitats in the region: open forests, riparian forests, and dense forests. We hypothesized that riparian forests would be the most preferred habitat for the butterflies, as they provide suitable microclimatic conditions for butterflies. The study collected samples for 35 grids of 2 × 2 km 2 for each habitat during the dry months (December-June). We considered the relative humidity, temperature, light intensity, elevation, and canopy cover to assess their influences on butterfly richness and abundance. We also considered the impact of disturbances on their distribution. We used structural equation modeling and canonical correspondence analysis to quantify the correlation and causation between the butterflies and their environment. The study recorded 1614 individual butterflies of 79 species from 57 genera and 6 families. During the study, we found that temperature was the most significant factor influencing butterfly richness. Relative humidity was also important and had a positive impact on butterfly richness. Riparian forests, where daytime temperatures are relatively low, were the most preferred microhabitat for butterflies. Open forests had greater species diversity, indicating the critical significance of an open canopy for butterflies. Though riparian forests need greater attention concerning butterfly distribution, maintaining open and dense forests are crucial for preserving butterfly diversity.
... 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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... For these reasons, Orthoptera have been used as bioindicators for conservation management [15,16] and its efficiency evaluation [17][18][19][20][21]. Greece is a hotspot for European Orthoptera fauna [22,23], hosting over one-third of all European species (35%: 378 species), with a pronounced degree of endemism (37%) and a number of red-listed species under the IUCN criteria (37%) [24,25]. However, the ecological knowledge base for Orthoptera communities is poor in Greece and is restricted to a few case studies (e.g., [15,[26][27][28]). ...
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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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