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FIGURE 13 in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURE 13. Phylogeny of the genus Namea modified from Rix et al. (2020; fig. 2), inferred from a partitioned Bayesian analysis of that study's 'FULL' dataset (151 taxa, 5,261 bp, 50% majority-rule consensus tree). Clades corresponding to 30 morpho-species OTUs are collapsed for the genus Namea (square brackets denote the number of specimens included in each clade), and nodes have a posterior probability of ≥ 0.98 unless otherwise stated. Seven species recorded from the D'Aguilar Range (south-eastern Queensland) are highlighted in enlarged bold text; note that specimens of an eighth species, N. gloriosa sp. nov., were not sequenced for this study.

FIGURES 22–31 in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURES 22–31. Namea gloriosa sp. nov., male holotype (QMB S10269) from Mount Glorious (south-eastern Queensland), somatic morphology: 22–23, carapace and abdomen, dorsal view; 24, cephalothorax, lateral view; 25, eyes, dorsal view; 26, mouthparts, ventral view; 27–28, cephalothorax and abdomen, ventral view; 29, leg I, prolateral view; 30, leg I tibia, retrolateral view; 31, leg I tibia, profile in standardised prolateral view, showing relative position of macrosetae (pd, prodorsal; pv, proventral; v, ventral); note the broken seta (*) at position v2. Scale bars = 3.0 mm.

FIGURES 4–12 in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURES 4–12. Live habitus images and burrows of Namea from the D'Aguilar Range (south-eastern Queensland). 4–8, Live habitus images: 4, female N. brisbanensis Raven, 1984 from Gold Creek Reserve (note left leg IV missing); 5, paratype female N. gowardae sp. nov. from Mount Glorious; 6, female N. salanitri Raven, 1984 from Mount Mee; 7, female N. dahmsi Raven, 1984 from Clear Mountain; 8, subadult juvenile N. nigritarsus sp. nov. from Mount Glorious. 9–12, Burrow images: 9, burrow of unidentified Namea (N. nigritarsus sp. nov. or N. nebo sp. nov.; not collected) from Mount Nebo, with spider at entrance; 10, burrow of N. gowardae sp. nov. from Mount Glorious, with spider at entrance; 11, burrow of N. dahmsi from Clear Mountain; 12, burrow of N. nigritarsus sp. nov. from Mount Mee. Images 4–8, 10, 11 by M. Rix; 9, 12 by J. Wilson.

FIGURES 48–54 in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURES 48–54. Namea gowardae sp. nov., female paratype (QMB S111377) from Maiala, Mount Glorious (south-eastern Queensland), somatic morphology: 48–49, carapace and abdomen, dorsal view; 50, cephalothorax, lateral view; 51, eyes, dorsal view; 52, mouthparts, ventral view; 53–54, cephalothorax and abdomen, ventral view (note that genitalia have been dissected from ventral abdominal epigastric region). Scale bars = 3.0 mm.

FIGURES 1–3. The D in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURES 1–3. The D'Aguilar Range of south-eastern Queensland: 1, elevational heat map showing upland areas surrounding Brisbane, and the relative position of the D'Aguilar Range; 2, topographic map of the D'Aguilar Range (in shaded relief), with major localities highlighted; 3, typical habitat for Namea on rainforest bank at Mount Glorious (Maiala section). Image 3 copyright Queensland Museum (used with permission).

FIGURES 68–77 in The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity

FIGURES 68–77. Namea nebo sp. nov., male holotype (QMB S65274) from Boombana, Mount Nebo (south-eastern Queensland), somatic morphology: 68–69, carapace and abdomen, dorsal view; 70, cephalothorax, lateral view; 71, eyes, dorsal view; 72, mouthparts, ventral view; 73–74, cephalothorax and abdomen, ventral view; 75, leg I, prolateral view; 76, leg I tibia, retrolateral view; 77, leg I tibia, profile in standardised prolateral view, showing relative position of macrosetae (pd, prodorsal; pv, proventral; v, ventral). Scale bars = 3.0 mm.

opencc-zeroJan 2021View details →

Ephemeral habitat supports high alpha and beta diversity during drought in a subtropical semiarid wetland

<p>Globally, the number of rivers with intermittent flow is increasing due to climate change and water abstraction for human consumption. Currently, our understanding of how hydrology in subtropical rivers with intermittent flow affects the structure and dynamics of aquatic communities is poorly understood. Here, we investigated how fish α and β-diversity patterns in intermittent channels in the lower reaches of the Okavango Delta respond to seasonal flooding and drought. Under low-flow conditions, ephemeral habitat had higher α-diversity, and this was influenced by a combination of fish aggregation and apparent transient legacy effects as habitat patches became smaller and more isolated. Investigation of changes in fish assemblage structure across different hydrological periods and habitat types showed significant species turnover when water levels fell, suggesting a strong influence from species sorting. During high-water periods, species assemblages were homogenized both at local and regional scales, suggesting a greater influence of mass effects. Our findings support hydrology as a major factor regulating diversity patterns in intermittent rivers of a major wetland in a semiarid region of subtropical Africa. We infer from these findings that maintenance of a relatively natural flow regime will be necessary for conserving aquatic ecosystem structure and function in this system.</p>

opencc-zeroJan 2021View details →

Neighboring trees regulate the root-associated pathogenic fungi on the host plant in a subtropical forest

<p>Root-associated fungi and host-specific pathogens are major determinants of species coexistence in forests. Phylogenetically related neighboring trees can strongly affect the fungal community structure of the host plant, which, in turn, will affect the ecological processes. Unfortunately, our understanding of the factors influencing fungal community composition in forests is still limited. In particular, investigation of the relationship between the phytopathogenic fungal community and neighboring trees is incomplete. In the current study, we tested the host specificity of members of the root-associated fungal community collected from seven tree species and determined the influence of neighboring trees and habitat variation on the composition of the phytopathogenic fungal community of the focal plant in a subtropical evergreen forest. Using high-throughput sequencing data with respect to the internal transcribed spacer (ITS) region, we characterized the community composition of the root-associated fungi and found significant differences with respect to fungal groups among the seven tree species. The density of conspecific neighboring trees had a significantly positive influence on the relative abundance of phytopathogens, especially host-specific pathogens, while the heterospecific neighbor density had a significant negative impact on the species richness of host-specific pathogens, as well as phytopathogens. Our work provides evidence that the root-associated phytopathogenic fungi of a host plant depends greatly on the tree neighbors of the host plant.</p>

opencc-zeroDec 2014View details →

Data from: Soil organic matter quantity and quality shape microbial community compositions of subtropical broadleaved forests

As two major forest types in the subtropics, broadleaved evergreen and broadleaved deciduous forests have long interested ecologists. However, little is known about their belowground ecosystems despite their ecological importance in driving biogeochemical cycling. Here, we used Illumina MiSeq sequencing targeting 16S rRNA gene and a microarray named GeoChip targeting functional genes to analyse microbial communities in broadleaved evergreen and deciduous forest soils of Shennongjia Mountain of Central China, a region known as 'The Oriental Botanic Garden' for its extraordinarily rich biodiversity. We observed higher plant diversity and relatively richer nutrients in the broadleaved evergreen forest than the deciduous forest. In odds to our expectation that plant communities shaped soil microbial communities, we found that soil organic matter quantity and quality, but not plant community parameters, were the best predictors of microbial communities. Actinobacteria, a copiotrophic phylum, was more abundant in the broadleaved evergreen forest, while Verrucomicrobia, an oligotrophic phylum, was more abundant in the broadleaved deciduous forest. The density of the correlation network of microbial OTUs was higher in the broadleaved deciduous forest but its modularity was smaller, reflecting lower resistance to environment changes. In addition, keystone OTUs of the broadleaved deciduous forest were mainly oligotrophic. Microbial functional genes associated with recalcitrant carbon degradation were also more abundant in the broadleaved deciduous forests, resulting in low accumulation of organic matters. Collectively, these findings revealed the important role of soil organic matter in shaping microbial taxonomic and functional traits.

opencc-zeroDec 2018View details →

Data from: Seasonal dynamics of waterbird assembly mechanisms revealed by phylogenetic and functional diversity in a subtropical wetland

Despite growing interest in phylogenetic and functional methods in ecological assembly, less attention has been paid to seasonal variation patterns of migrant species. Migrants can rapidly mediate influences of species interactions and environmental factors through seasonal movement, suggesting dynamical relative importance of different assembly mechanisms among seasons. Here we describe seasonal dynamics in phylogenetic and functional diversity of waterbirds in Mai Po Wetland, in a subtropical region with significant predictable temporal variation. Phylogenetic and functional structure of α diversity varied seasonally. Specifically, phylogenetic structures clustered in summer, while being over-dispersed in winter. However, phylogenetic structure in spring and autumn was intermediate with a transition to random. Functional structure was clustered in spring but showed over-dispersion in the other three seasons. For β diversity, summer and winter assemblages had two distinct groups, while spring and autumn assemblages were mixed. Thus, waterbird assemblages were primarily shaped by interspecific competition in winter. Random processes tended to shape assemblage in spring. Environmental factors played a more important role in summer . In addition, phylogenetic distance of probability of co-occurrence of species pairs was significantly larger in winter than in summer. These results suggest that the relative importance of assemblage mechanisms can vary seasonally in response to changing environmental conditions, suggesting that studies attempting to infer a single dominant assembly mechanism may ignore important assembly processes. Temporal shifts in assembly mechanisms may play an important role in maintaining diversity of subtropical and temperate wetlands and perhaps other dynamic systems.

opencc-zeroDec 2016View details →

Data from: Heterogeneous distributional responses to climate warming: evidence from rodents along a subtropical elevational gradient

Background: Understanding whether species' elevational range is shifting in response to directional changes in climate and whether there is a predictable pattern in that response is one of the major challenges in ecology. However, so far very little is known about the distributional responses of subtropical species to climate change, especially for small mammals. In this study, we examined the elevational range shifts at three range points (upper and lower range limits and abundance-weighted range centre) of rodents over a 30-year period (1986 to 2014-2015), in a subtropical forest of Southwest China. We also examined the influences of four ecological traits (body mass, habitat breadth, diet and daily activity pattern) on the upslope shifts in species' abundance-weighted range centres. Results: Despite the warming trend between 1986 and 2015, the eleven rodent species in analysis displayed heterogeneous dynamics at each of the three range points. Species which have larger body sizes and narrower habitat breadths, show both diurnal and nocturnal activities and more specialized dietary requirements, are more likely to exhibit upslope shifts in abundance-weighted range centres. Conclusions: Species' distributional responses can be heterogeneous even though there are directional changes in climate. Our study indicates that climate-induced alleviation of competition and lag in response may potentially drive species' range shift, which may not conform to the expectation from climate change. Difference in traits can lead to different range dynamics. Our study also illustrates the merit of multi-faceted assessment in studying elevational range shifts.

opencc-zeroDec 2014View details →

Data from: Pleistocene niche stability and lineage diversification in the subtropical spider Araneus omnicolor (Araneidae)

The influence of Quaternary climate oscillations on the diversification of the South American fauna is being increasingly explored. However, most of these studies have focused on taxa that are endemic to tropical environments, and relatively few have treated organisms restricted to subtropical biomes. Here we used an integrative phylogeographical framework to investigate the effects of these climate events on the ecological niche and genetic patterns of the subtropical orb-weaver spider Araneus omnicolor (Araneidae). We analyzed the mitochondrial (Cytochrome Oxidase I, COI) and nuclear (Internal Transcribed Subunit II, ITS2) DNA of 130 individuals throughout the species' range, and generated distribution models in three different climate scenarios [present, Last Glacial Maximum (LGM), and Last Interglacial Maximum (LIG)]. Additionally, we used an Approximate Bayesian Computation (ABC) approach to compare possible demographic scenarios and select the hypothesis that better explains the genetic patterns of A. omnicolor. We obtained high haplotype diversity but low nucleotide variation among sequences. The population structure and demographic analyses showed discrepancies between markers, suggesting male-biased dispersal in the species. The time-calibrated COI phylogenetic inference showed a recent diversification of lineages (Middle/Late Pleistocene), while the paleoclimate modeling indicated niche stability since ~120 Kya. The ABC results agreed with the niche models, supporting a panmictic population as the most likely historical scenario for the species. These results indicate that A. omnicolor experienced no niche or population reductions during the Late Pleistocene, despite the intense landscape modifications that occurred in the subtropical region, and that other factors beside LGM and LIG climate oscillations might have contributed to the demographic history of this species. This pattern may be related to the high dispersal ability and wide environmental tolerance of A. omnicolor, highlighting the need for more phylogeographical studies with invertebrates and other generalist taxa, in order to understand the effects of Quaternary climate changes on Neotropical biodiversity.

opencc-zeroJul 2019View details →

Data from: Tissue-specific carbon concentration, carbon stock, and distribution in Cunninghamia lanceolata (Lamb.) Hookplantations at various developmental stages in subtropical China

Key message Carbon (C) concentrations in Cunninghamia lanceolata (Lamb.) Hook plantations differed significantly among tissue types and were greater for aboveground than belowground tissues. Plantation C stock increased with developmental stage from young to mature to overmature, but at all stages the majority occurred as soil organic carbon (SOC) and was more influenced by belowground fine roots than by aboveground litterfall. Context Failing to account for tissue-specific variation in the C concentration can result in inaccurate forest C stock estimates. Aims We aimed to quantify the relative magnitudes of C stock for Chinese fir plantations at different developmental stages. Specifically, we focused on assessing tissue-specific C concentrations and C dynamics return of above- and belowground litterfall. Methods Carbon traits (C concentration, C flux, C stock and distribution at tree and ecosystem scales) were quantified in a chronosequence of Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) monoculture plantation stands at young (10), mature (22), and overmature (34 years old) developmental stages. Results Carbon concentrations differed significantly among tissue types, with mean values of 48.5 ± 0.1% and 42.5 ± 0.2% for above- and belowground biomass, respectively. The aboveground tissue C concentration, tree- and plantation-scale C stock, and SOC stock depended on developmental stage. Carbon return in litterfall, tree C stock, and SOC increased from the young to the overmature stage. SOC stock accounted for the majority of plantation C stock at all developmental stages (78.3, 59.6 and 55.7% in the young, mature and overmature stages, respectively) and was more highly influenced by belowground fine roots than aboveground litterfall. Carbon stocks in Chinese fir plantations were 86, 129, and 153 t ha-2 at the young, mature, and overmature stages. Conclusion Prolonging Chinese fir rotation increases C sequestration potential and should be the focus of forest management strategies. The tissue-specific C concentrations provide detailed information for more accurate biomass C stock estimates for Chinese fir plantations and other subtropical coniferous forest. They indicate that current guidelines result in an overestimation of belowground biomass C stocks. Using the standard 0.47 biomass to C conversion factor, the belowground C stock would have been overestimated by 7.6-13.0% for the Chinese fir developmental stages investigated, while tree C stock would be underestimated by 0.08-3.24%. Therefore, developing species- and tissue-specific conversion factors are required for supporting C plantation and forest C accounting strategies.

opencc-zeroDec 2016View details →

Data from: Multi-trophic guilds respond differently to changing elevation in a subtropical forest

Negative relationships between species richness and elevation are common and attributed to changes in single environmental properties associated to elevation, such as temperature and habitat area. However, research has lacked taxonomic breadth and comprehensive elevation studies that consider multiple groups from different trophic levels are rare. We thus analysed 24 groups of plants, arthropods, and microorganisms grouped into six trophic guilds (predators, detritivores, herbivores, plants, bacteria and fungi) along a relatively short elevational gradient (~600 m) in a subtropical forest in south-east China. The total species richness of all organisms was not related to elevation, nor was the richness of plants, herbivores or microorganisms. However, species richness and abundance in two major trophic guilds of arthropods changed with elevation, which was mediated by changes in elevation-associated habitat properties. Specifically, deadwood mass increased with elevation, which increased detritivore richness indirectly via detritivore abundance, thus supporting the 'more individuals hypothesis'. In contrast, lower predator richness at higher elevations was directly related to lower mean temperatures, which had no effect on abundance. Our study demonstrates that even along relatively short gradients, elevation can have strong direct and abundance-mediated effects on species richness, but with effects varying from positive to negative signs depending on local resource availability and the characteristics of groups or trophic guilds. If elevation positively influences local environmental properties that benefit a given group, richness can increase towards higher elevations. Thus, the effect of global change in mountainous regions should be evaluated within the local environmental context using multi-taxon approaches.

opencc-zeroDec 2015View details →

Data from: Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study

Using a novel combination of palaeohabitat modelling and genetic mixture analyses, we identify and assess a sea-level-driven recolonization process following the Last Glacial Maximum (LGM). Our palaeohabitat modelling reveals dramatic changes in estuarine habitat distribution along the coast of California (USA) and Baja California (Mexico). At the LGM (approx. 20 kya), when sea level was approximately 130 m lower, the palaeo-shoreline was too steep for tidal estuarine habitat formation, eliminating this habitat type from regions where it is currently most abundant, and limiting such estuaries to a northern and a southern refugium separated by 1000 km. We assess the recolonization of estuaries formed during post-LGM sea-level rise through examination of refugium-associated alleles and approximate Bayesian computation in three species of estuarine fishes. Results reveal sourcing of modern populations from both refugia, which admix in the newly formed habitat between the refuges. We infer a dramatic peak in habitat area between 15 and 10 kya with subsequent decline. Overall, this approach revealed a previously undocumented dynamic and integrated relationship between sea-level change, coastal processes and population genetics. These results extend glacial refugial dynamics to unglaciated subtropical coasts and have significant implications for biotic response to predicted sea-level rise.

opencc-zeroDec 2014View details →

Data from: ENSO and frost co-determine decade-long temporal variation in flower and seed production in a subtropical rain forest

Flower and seed production of plants can be greatly influenced by both natural climatic oscillations and local weather extremes. However, owing to the rarity of long-term monitoring studies conducted at a sufficient temporal scale to capture climatic oscillations and the unpredictability of extreme weather events, evidence that demonstrates how these two external forcings act in concert to drive plant reproduction remains scarce. In addition, considerable variation in species' phenological responses to the external climatic forcings was often observed. Phylogenetic relationships may mediate this inter-specific variation, but previous studies yielded inconsistent results when testing this hypothesis. We monitored the flower and seed production in a subtropical rainforest, Fushan, Taiwan (24°45'N, 121°35'E), for over ten years (since September 2002). In March 2005, a record low temperature (-1.3°C) occurred at Fushan and caused great frost damage to plants. We used weekly phenological records and long-term meteorological data to assess the effects of climatic fluctuations and extreme weather event on plant reproductive output. We show that the El Niño Southern Oscillation (ENSO) indices, which integrated local climatic variables at Fushan over several months, were strongly associated with flower and seed production. The 2005 spring frost also had long-lasting effects on the flower and seed production of several species. In particular, we detected phylogenetic signals in the relationships between phenological responses of flowering production and several climatic variables (maximum temperature, irradiance, and ENSO34 index). By contrast, the relationships between seed production and climatic variables, as well as phenological responses to the frost event, did not exhibit a phylogenetic signal. Synthesis: Our findings add to the growing evidence that together the natural climatic oscillation (ENSO) and the extreme weather event (frost) determined the temporal variation in flower and seed production. In addition, phylogenetically closely related species resembled each other in their flowering responses to abiotic variation in this subtropical rainforest. Improved understanding of these abiotic and biotic interactions may help predicting population- and community-level phenological responses under future climate changes.

opencc-zeroDec 2017View details →

Data from: Edge effects and beta diversity in ground and canopy beetle communities of fragmented subtropical forest

Clearing of dry forests globally creates edges between remnant forest and open anthropogenic habitats. We used flight intercept traps to evaluate how forest beetle communities are influenced by distance from such edges, together with vertical height, spatial location, and local vegetation structure, in an urbanising region (Brisbane, Australia). Species composition (but not total abundance or richness) differed greatly between ground and canopy. Species composition also varied strongly among sites at both ground and canopy levels, but almost all other significant effects occurred only at ground level, where: species richness declined from edge to interior; composition differed between positions near edges (<10 m) and interiors (> 50 m); high local canopy cover was associated with greater total abundance and richness and differing composition; and greater distances to the city centre were associated with increased total abundances and altered composition. Analyses of individual indicator species associated with this variation enabled further biological interpretations. A global literature synthesis showed that most spatially well-replicated studies of edge effects on ground-level beetles within forest fragments have likewise found that positions within tens of metres from edges with open anthropogenic habitats had increased species richness and different compositions from forest interior sites, with fewer effects on abundance. Accordingly, negative edge effects will not prevent relatively small compact fragments (if >10-20 ha) from supporting forest-like beetle communities, although indirect consequences of habitat degradation remain a threat. Retention of multiple spatially scattered forest areas will also be important in conserving forest-dependent beetles, given high levels of between-site diversity.

opencc-zeroJun 2019View details →

Data from: Disturbance history mediates climate change effects on subtropical forest biomass and dynamics

The responses of forest communities to interacting anthropogenic disturbances like climate change and logging are poorly known. Subtropical forests have been heavily modified by humans and their response to climate change is poorly understood. We investigated the nine year-change observed in a mixed conifer-hardwood Atlantic forest mosaic that included both mature and selectively-logged forest patches in subtropical South America. We used demographic monitoring data within 10 1 ha plots that were subjected to distinct management histories (plots logged until 1955, until 1987, and unlogged) to test the hypothesis that climate change affected forest structure and dynamics differentially depending on past disturbances. We determined the functional group of all species based on life-history affinities as well as many functional traits like leaf size, specific leaf area, wood density, total height, stem slenderness, and seed size data for the 66 most abundant species. Analysis of climate data revealed that minimum temperatures and rainfall have been increasing in the last few decades of the 20th century. Floristic composition differed mainly with logging history categories, with only minor change over the nine annual census intervals. Aboveground biomass increased in all plots, but increases were higher in mature unlogged forests, which showed signs of forest growth associated with increased CO2, temperature, and rainfall/treefall gap disturbance at the same time. Logged forests showed arrested succession as indicated by reduced abundances of Pioneers and biomass-accumulators like Large Seeded Pioneers and Araucaria, as well as reduced functional diversity. Management actions aimed at creating regeneration opportunities for long-lived pioneers are needed to restore community functional diversity, and ecosystem services such as increased aboveground biomass accumulation. We conclude that the effects of climate drivers on the dynamics of Brazilian mixed Atlantic forests vary with land-use legacies, and can differ importantly from the ones prevalent in better known tropical forests.

opencc-zeroDec 2011View details →

Data from: Pleistocene speciation with and without gene flow in Euphaea damselflies of subtropical and tropical East Asian islands

Climatic oscillations during the Pleistocene period could have had a profound impact on the origin of tropical species by the alternation of allopatric isolation and inter-population gene flow cycles. However, whether tropical speciation involves strictly allopatric isolation, or proceeds in the face of homogenizing gene flow, is relatively unclear. Here, we investigated geographical modes of speciation in four closely-related Euphaea damselfly species endemic to the subtropical and tropical East Asian islands using coalescent analyses of a multi-locus data set. The reconstructed phylogenies demonstrated distinct species status for each of the four species and the existence of two sister species pairs, E. formosa/E. yayeyamana and E. decorata/E. ornata. The species divergence time of the sibling Euphaea damselflies dates back to within the last one Mya of the Middle to Lower Pleistocene. The speciation between the populous E. formosa of Taiwan and the less numerous E. yayeyamana of the Yaeyama islands occurred despite significant bidirectional, asymmetric gene flow, which is strongly inconsistent with a strictly allopatric model. In contrast, speciation of the approximately equal-sized populations of E. decorata of the south-east Asian mainland and E. ornata of Hainan is inferred to have involved allopatric divergence without gene flow. Our findings suggest that differential selection of natural or sexual environments is a prominent driver of species divergence in subtropical E. formosa and E. yayeyamana; whereas for tropical E. decorata and E. ornata at lower latitudes, allopatric isolation may well be a pivotal promoter of species formation.