Search USGS⌕ Search

Geology topics

Steven A. Pothoven

Publications and source records attributed to Steven A. Pothoven.

At least 19 recordsLinked to original sources
USGS2026

The Great Lakes aquatic tissue analysis repository (GLATAR)

Empiricists and modellers use information on energy density, proximate composition, stable isotopes, fatty acids, thiamine, and bioaccumulative tracers (e.g., PCBs and mercury) to understand the state and inter-relationships of aquatic food webs. Data exist in many published and unpublished sources, but are not consolidated in an easily accessible database that would serve as a vital resource to i) provide basic estimates of these diet-derived measures of body composition, ii) understand sources of variation in the underlying data, iii) facilitate exploration and development of data proxies, and iv) assist in study design. We designed GLATAR (Great Lakes Aquatic Tissue Analysis Repository, glatar.org ) to address this need. GLATAR is an open-access, searchable database linked to a web-based toolbox to visualise and generate user-defined summaries on diet-derived ecological metrics, with a focus on taxa of importance to the Great Lakes. GLATAR currently contains over 50,000 records on energy density, chemical tracers, and proximate body composition from 67 species of fish and 72 invertebrate taxa. We hope this user-friendly interface will entice others to upload their published and unpublished data to the repository, enriching the breadth of data accessible to researchers. In this way, GLATAR will become a ‘living’ and interactive resource for empiricists and modellers working in freshwater ecosystems as they make critical decisions related to growth, production, and consumption across a diverse group of economically and ecologically important aquatic species.

Journal of Great Lakes Research↗
USGS2026

Availability of dark daytime refuge may limit mysid abundance in the Laurentian Great Lakes

The zooplankton Mysis diluviana is a major component of the Laurentian Great Lakes food web and has recently declined in abundance in both lakes Michigan and Huron. Drivers of these declines are not well understood. Here, we explore the hypothesis that recent increases in water clarity have contributed to the decline of M. diluviana (mysids) by limiting the availability of daytime dark refuge from visual predators. Using Secchi depth data from 1996 to 2021, we estimate that dark refuge has decreased substantially in lakes Michigan and Huron where mysids have declined, but dark refuge has remained more stable in lakes Ontario and Superior where mysid populations did not decline. Results from a 2021 uncrewed surface vessel hydroacoustic survey and lake-wide, net-based sampling in lakes Michigan and Huron revealed significantly more mysids in areas with dark refuge, such as Lake Michigan’s northern basin. Conversely, Lake Huron contains sparse dark refuge consistent with low mysid densities in that lake. Higher water clarity leading to increased predation may be a primary driver of mysid declines in the Great Lakes.

Laurentian Great Lakes↗
USGS2025

Drowned river mouth lakes are winter foraging habitats for the expanding Lake Michigan cisco Coregonus artedi population

Characterizing fish movements is required for understanding habitat use, energy flow, and trophic structure and can inform fisheries management. Drowned river mouth (DRM) lakes are productive inland habitats in the Laurentian Great Lakes basin used by migratory fishes. Despite recognition of their ecological connections to the Great Lakes, the value of DRM lakes as seasonal habitats is not well understood for many fishes. One such species, cisco Coregonus artedi , has recently expanded in Lake Michigan from near extirpation to higher relative abundances in the northeastern portion of the lake. Cisco are recreationally harvested in some DRM lakes during winter, but little is known about cisco movement patterns and ecology. In winter 2022 and 2023, we collected cisco from three DRM lakes along the eastern shores of Lake Michigan (Lake Charlevoix, Portage Lake, Muskegon Lake) to characterize genetics, morphometrics, and diets. We also implanted telemetry tags in 20 cisco collected in Lake Charlevoix to examine movement patterns and determine DRM lake residency (i.e., seasonal vs. year-round). We found no consistent genetic or morphometric differentiation across DRM lakes, suggesting that recolonization began from a single stock. Fish were the only diet item found in cisco guts collected during winter months. Movement patterns from Lake Charlevoix indicated strong spawning site fidelity to Grand Traverse Bay as well as non-spawning site fidelity. However, given the presence of cisco in southern DRM lakes and some site-specific differences in morphometrics, managers could benefit from further research to determine whether spawning occurs in southern Lake Michigan.

Journal of Great Lakes Research↗
USGS2025

Lake depth and light conditions alter Mysis vertical distributions

Light regulates the vertical migration of many aquatic organisms. Mysis species couple pelagic and benthic habitats in lakes by diel vertical migrations (DVM), transporting energy and nutrients through the water column and food web. Although Mysis are generally assumed to remain on the bottom during the day, some have been observed in the pelagic zone during the day, indicating incomplete benthic-pelagic coupling in some systems. The degree to which light attenuation and lake depth interact to affect occurrence of mysids within the water column during the day is understudied. We used standardized Mysis net sampling in summers 2020 and 2021 across nine north-temperate lakes to test the hypotheses that 1) Mysis remain pelagic during the day at depths with sufficiently low light levels, and 2) pelagic-caught individuals during the day are, on average, smaller than those caught at night. To test these hypotheses, we assessed light, dissolved oxygen (DO), Mysis densities, and size distribution between night and day across bathymetric depths. In deep lakes and darkly colored shallow lakes, Mysis suspended in the water column during the day where light levels decreased to their light avoidance threshold (∼10 -5 to 10 -6 lx ). Mysis suspended in the water column during the day were smaller than those collected at night. Further, Mysis were not captured when DO reached levels < 3 mg/L, regardless of light conditions. Our results suggest that benthic-pelagic coupling by Mysis is mediated through light conditions, lake morphometry, and DO conditions, and may include some degree of size-dependent behavior.

Journal of Great Lakes Research↗
USGS2024

Reconstructing half a century of coregonine recruitment reveals species-specific dynamics and synchrony across the Laurentian Great Lakes

Understanding how multiple species and populations vary in their recruitment dynamics can elucidate the processes driving recruitment across space and time. Lake Whitefish ( Coregonus clupeaformis ) and Cisco ( C. artedi ) are socioecologically important fishes across their range; however, many Laurentian Great Lakes populations have experienced declining, poor, or sporadic recruitment in recent decades. We integrated catch and age data from 38 long-term surveys across each of the Great Lakes and Lake Simcoe, resulting in a combined time series spanning 1960–2019. We estimated Lake Whitefish and Cisco year-class strength (YCS) in each lake using longitudinal mixed-effects regressions of relative cohort abundance. We subsequently quantified interspecific, spatial, and temporal synchrony in YCS using correlation and dynamic factor analyses. Lake Whitefish YCS was positively spatially synchronous on average, and YCS in all six lakes was elevated during the 1980s–1990s. In contrast, Cisco YCS was sporadic, not spatially synchronous, and highly variable around long-term, lake-specific means. YCS was not synchronous between species in any lake. Collectively, our analyses demonstrate that these species exhibit differential recruitment dynamics that may be regulated by species-specific factors. Results from this study can be leveraged in future research on the causes and consequences of cross-species, cross-basin recruitment variability.

Laurentian Great Lakes↗
USGS2024

Synthesizing professional opinion of Lake Whitefish and Cisco recruitment drivers across the Great Lakes

Disentangling the suite of ecological drivers that explain recruitment variability for Lake Whitefish Coregonus clupeaformis and Cisco C. artedi is of critical importance for their conservation, management, and stewardship in the Laurentian Great Lakes. However, recruitment is inherently variable and can be regulated by many interacting processes, the relative importance of which can vary spatially, temporally, and ontogenetically. Given this complexity, comparisons across lakes and species that identify overarching hypotheses could efficiently guide future research. Using facilitated deliberations among fishery professionals (n = 57) with expertise in Great Lakes Coregonus spp., we synthesized current knowledge regarding (1) which biophysical processes are most important for driving contemporary recruitment between species, among lakes, and across life stages and (2) mechanisms by which those drivers regulate recruitment at key life stages. Participants affirmed the hypothesis that many drivers interact in complex ways to regulate Lake Whitefish and Cisco recruitment. Large-scale climatic processes affecting early life-stage growth and survival were consistently considered important. Other drivers were only deemed influential in certain lakes, highlighting perceived context-dependent recruitment dynamics. Notably, recruitment in Lakes Superior, Michigan, and Huron was considered limited during larval and early juvenile life stages by low productivity, whereas spawning-habitat degradation and reduced metapopulation diversity were hypothesized to limit recruitment during embryonic and larval stages in Lakes Erie and Ontario. Several drivers were hypothesized to similarly impact Lake Whitefish and Cisco during early life stages, while drivers acting on post-larval life stages were typically distinct between species. The hypotheses synthesized herein can guide future research on Lake Whitefish and Cisco recruitment dynamics in the Great Lakes.

Great Lakes↗
USGS2024

Modeling nearshore total phosphorus in Lake Michigan using linked hydrodynamic and water quality models

Although the offshore water of Lake Michigan has been below the Great Lakes Water Quality Agreement (GLWQA) total phosphorus (TP) spring target concentration of 7 µg L −1 for several decades, higher TP concentrations occur in the nearshore, contributing to the resurgence of Cladophora and other nutrient related issues. The most recent update of the GLWQA specifically calls for the assessment of current nearshore conditions. Such assessment would require an intensive monitoring program supported by water quality models. Here we applied multiple versions of a phosphorus-based model linked to an unstructured-grid hydrodynamic model. We focus on the nearshore regions of Lake Michigan around the Grand and Muskegon rivers – a region with strong riverine TP influence and intensive monitoring. Results from a TP model were compared to observational data and to a previously published Phosphorus-based Nutrient–Phytoplankton–Zooplankton–Detrital–Mussel (NPZDM) model. Model results and observational data show that parts of the nearshore can be well above the target TP concentrations but, due to the dynamic nature of this region, the TP concentrations can change rapidly. The models’ skill statistics in predicting individual observations were variable, but it was able to simulate temporal and spatial trends and captured the distribution of observations in our study area. The similarity between the results of the TP and NPZDM models demonstrated the TP concentrations in this nearshore area are driven by hydrodynamics and river TP loads, which are likely the reasons for the higher observed TP concentrations. Simplicity, transparency, and ease of use of the TP model make it a useful tool for supporting nearshore assessments and estimating existing and future nearshore TP concentrations.

Michigan↗
USGS2021

Lake-wide annual status of Mysis diluviana population in Lake Michigan in 2015

Mysis diluviana is one of the most abundant zooplankton by biomass in the Laurentian Great Lakes of North America, a predator of other zooplankton and an important prey for fishes. Studies of long-term trends in Lake Michigan have shown 2005–2016 densities to be 50–80% lower than 1990s densities, but these observations have been based on annual monitoring that is either spatially or seasonally limited. We combined Lake Michigan Mysis data from three annual programs and the 2015 Cooperative Science and Monitoring Initiative to achieve broad spatial coverage during spring, summer, and fall of 2015 and broad depth coverage during spring 2016. Lake-wide, annual density and biomass were 82 (SE: 10) Mysis /m 2 and 200 (SE: 36) mg dry mass/m 2 . Density and biomass estimates were highest offshore, generally higher in the north basin, and seasonally highest in summer. Annual lake-wide averages for depths >30 m were better captured by seasonally-extensive annual programs than spatially-extensive annual programs, although spring sampling may bias annual values low. Mysis cohorts grew 0.026 mm/d (age-0) and 0.007 to 0.027 mm/d (age-1). Annual mortality was 81–98%. Reproduction was fall-spring and seasonal lake-wide estimates ranged from 0.6 to 19.1% females brooding, 13–20 embryos/brood, and 3–46 embryos/m 2 . Annual production (423 mg dry mass/m 2 /yr, SE: 31) was lower than all but one previous estimate from lakes Michigan, Huron, and Ontario. While Mysis tend to persist, low Mysis production may be a concern for prey fishes that feed on Mysis .

Lake Michigan↗
USGS2019

Evaluating consumptive and nonconsumptive predator effects on prey density using field times series data

Determining the degree to which predation affects prey abundance in natural communities constitutes a key goal of ecological research. Predators can affect prey through both consumptive effects (CEs) and nonconsumptive effects (NCEs), although the contributions of each mechanism to the density of prey populations remain largely hypothetical in most systems. Common statistical methods applied to time series data cannot elucidate the mechanisms responsible for hypothesized predator effects on prey density (e.g., differentiate CEs from NCEs), nor provide parameters for predictive models. State space models (SSMs) applied to time series data offer a way to meet these goals. Here, we employ SSMs to assess effects of an invasive predatory zooplankter, Bythotrephes longimanus, on an important prey species, Daphnia mendotae, in Lake Michigan. We fit mechanistic models in a SSM framework to seasonal time series (1994-2012) using a recently developed, maximum likelihood-based optimization method, iterated filtering, which can overcome challenges in ecological data (e.g. nonlinearities, measurement error, and irregular sampling intervals). Our results indicate that B. longimanus strongly influences D. mendotae dynamics, with mean annual peak densities of B. longimanus observed in Lake Michigan estimated to cause a 61% reduction in D. mendotae population growth rate and a 59% reduction in peak biomass density. Further, the observed B. longimanus effect is most consistent with an NCE via reduced birth rates. The SSM approach also provided estimates for key biological parameters (e.g., demographic rates) and the contribution of dynamic stochasticity and measurement error. Our study therefore provides evidence derived directly from survey data that the invasive zooplankter B. longimanus is affecting zooplankton demographics and offer parameter estimates needed to inform predictive models that explore the effect of B. longimanus under different scenarios such as climate change.

Ecology↗
USGS2019

Spatial patterns of rainbow smelt energetic condition in Lakes Huron and Erie in 2017: Evidence for Lake Huron resource limitation

Rainbow smelt ( Osmerus mordax ) is a key planktivore and prey fish in Lake Huron. Given the declining offshore productivity in the lake since the early 2000s, we described the energy content of rainbow smelt in 2017 across five different regions (North Channel, Georgian Bay, Saginaw Bay, northern main basin, southern main basin) where phytoplankton and zooplankton productivity likely varied. To increase contrast across the productivity gradient, rainbow smelt energy content was also estimated from western Lake Erie. Within the North Channel where large fish (≥90 mm, total length) were sampled most frequently, mean energy density (kJ/g wet weight) varied seasonally: 4.29 in April (month of spawning), 3.86 in June, 3.99 in July, and up to 4.35 in September. Energy density of rainbow smelt from higher productivity western Lake Erie was 37% (large fish ≥90 mm) to 60% higher (small fish <90 mm) than that of fish from Lake Huron. Within Lake Huron, energy density of rainbow smelt from North Channel was slightly higher than those from other regions; rainbow smelt from Georgian Bay generally had the lowest energy density. Across regions, including western Lake Erie, energy density increased with chlorophyll a concentration. Compared with Lake Huron studies prior to 2004, when oligotrophication had not yet accelerated, energy density of rainbow smelt in 2017 was up to 31% lower. The decline in rainbow smelt energy density is likely the result of declining primary and secondary pelagic production and increased resource limitation for planktivorous fish.

Michigan, Ohio, Ontario↗
USGS2019

Spatiotemporal variability in energetic condition of alewife and round goby in Lake Michigan

Pelagic-oriented alewife ( Alosa pseudoharengus ) and benthic-oriented round goby ( Neogobius melanostomus ) are two important prey fishes in the Laurentian Great Lakes. In 2015, we evaluated their seasonal total energy (TE) across nine Lake Michigan transects. Round goby contained at least 48% more kilojoules of TE than alewife of equal length during spring and summer. TE varied spatially for both species, but only large alewife exhibited a consistent pattern, with higher values along the eastern shoreline. Variation in TE was not explained by site-specific prey densities for either species. Round goby energy density (ED) was higher in Lake Michigan than in central Lake Erie, but comparable to other regions of the Great Lakes. Alewife ED in 2015 was similar to that in 2002–2004 in Lake Michigan, with the exception of November (small alewife ED was 21% higher) and April (large alewife ED was 30% lower). Despite oligotrophication, our study suggests that starvation of juvenile and adults has not been directly contributing to overall declining prey fish abundance, although future research should evaluate the potential for overwinter starvation.

Lake Michigan↗
USGS2018

Temperature regimes, growth, and food consumption for female and male adult walleye in Lake Huron and Lake Erie: a bioenergetics analysis

Bioenergetics modeling was used to assess the relative importance of food availability and water temperature in determining walleye ( Sander vitreus ) growth. Temperature regimes experienced by both female and male adult walleye in three basins of Lake Huron and in Lake Erie were determined by use of surgically implanted temperature loggers and acoustic telemetry. Temperatures experienced by walleye were higher in Lake Erie than in Lake Huron. Walleye from Lake Erie grew at nearly double the rate of walleye from Lake Huron, and mass at age for adult females averaged about 50% greater than that for adult males in both lakes. Food consumption rate for an average adult walleye in Lake Erie was nearly twice as high as that in Lake Huron. Interbasin and interlake variability in temperature regimes accounted for a moderate degree of variability in walleye growth. We concluded that the driver for faster growth in Lake Erie compared with Lake Huron was higher food availability in Lake Erie compared with Lake Huron. The sex difference in temperature regimes explained 15% of the sex difference in Lake Erie walleye growth.

Lake Erie, Lake Huron↗
USGS2018

Are changes in lower trophic levels limiting prey-fish biomass and production in Lake Michigan?

To improve understanding of how recent changes in lower trophic levels in Lake Michigan could be affecting prey-fish biomass and production, the Lake Michigan Committee (LMC) convened a Lower Trophic Level Task Group and provided several charges that are responded to in this report. First, we compiled a comprehensive summary of lower trophiclevel data in Lake Michigan, separating out nearshore versus offshore trends over time. Declining trends were prevalent in offshore time series for phosphorus, chlorophyll a, biomass of total crustacean zooplankton, biomass of herbivorous cladocerans, and density of Diporeia spp. In the nearshore, declining trends were evident only for biomass of cyclopoid copepods and density of Diporeia spp. Second, we hypothesized specific mechanisms by which changes in lower trophic levels could affect prey-fish biomass and production and described the degree of empirical support for each mechanism. The best-supported hypothesis was that declining invertebrate prey (especially Diporeia spp.) was responsible for declining growth of prey fish, especially over the last decade when competition for prey resources should otherwise have been lessened due to declining prey-fish densities. As a result, declining growth potentially limits the prey-fish biomass that could have been attained had growth been maintained at the levels that were achieved in the 1980s and earlier. Third, we prioritized several lower trophic-level indicators that fishery managers could use to better inform decision making. The top-ranked indicator was annual reporting of Alewife (Alosa pseudoharengus) condition. Fourth, we prioritized the key monitoring and research gaps that limit our current understanding of how lower trophic levels influence fish production. The highest-priority monitoring gap was coordinated sampling of the nearshore, which, if accomplished, would complement annual reporting on offshore sampling. The top-ranked knowledge gap was identifying bottlenecks that regulate fish recruitment, given that recent changes in zooplankton distribution and abundance could be suppressing survival of larval fish and, ultimately, the biomass and production of prey fish. We provided three specific recommendations for the LMC to consider as they seek to better incorporate lower trophiclevel changes into their management decision process: (1) implement a coordinated and standardized nearshore monitoring program, (2) encourage funding agencies to use our prioritized lists in their decision processes, and (3) foster the already improved dialogue between those researching lower trophic levels and those researching fisheries.

Miscellaneous Publication↗
USGS2016

Feeding ecology of the walleye (Percidae, Sander vitreus), a resurgent piscivore in Lake Huron (Laurentian Great Lakes) after shifts in the prey community

Recovering populations of piscivores can challenge understanding of ecosystem function due to impacts on prey and to potentially altered food webs supporting their production. Stocks of walleye (Percidae, Sander vitreus ), an apex predator in the Laurentian Great Lakes, crashed in the mid‐1900s. Management efforts led to recovery by 2009, but recovery coincided with environmental and fish community changes that also had implications for the feeding ecology of walleye. To evaluate potential changes in feeding ecology for this apex predator, we assessed diets in the main basin of Lake Huron and in Saginaw Bay, a large embayment of Lake Huron, during 2009–2011. Walleye switched their diets differently in the main basin and Saginaw Bay, with non‐native round goby (Gobiidae, Neogobius melanostomus ) and rainbow smelt (Osmeridae, Osmerus mordax ) more prevalent in diets in the main basin, and invertebrates, yellow perch (Percidae, Perca flavescens ) and gizzard shad (Clupeidae, Dorosoma cepedianum ) more prevalent in diets in the bay. Feeding strategy plots indicated that there was a high degree of individual specialisation by walleye in the bay and the main basin. Bioenergetic simulations indicated that walleye in Saginaw Bay need to consume 10%–18% more food than a walleye that spends part or all of the year in the main basin, respectively, in order to achieve the same growth rate. The differences in diets between the bay and main basin highlight the flexibility of this apex predator in the face of environmental changes, but changes in diet can alter energy pathways supporting piscivore production.

Ecology of Freshwater Fish↗
USGS2016

Differences in energy expenditures and growth dilution explain higher PCB concentrations in male summer flounder

Comparison of polychlorinated biphenyl (PCB) concentrations between the sexes of mature fish may reveal important behavioral and physiological differences between the sexes. We determined whole-fish PCB concentrations in 23 female summer flounder Paralichthys dentatus and 27 male summer flounder from New Jersey coastal waters. To investigate the potential for differences in diet or habitat utilization between the sexes, carbon and nitrogen stable isotope ratios were also determined. In 5 of the 23 female summer flounder, PCB concentrations in the somatic tissue and ovaries were determined. In addition, we used bioenergetics modeling to assess the contribution of the growth dilution effect to the observed difference in PCB concentrations between the sexes. Whole-fish PCB concentrations for females and males averaged 87 and 124 ng/g, respectively; thus males were 43% higher in PCB concentration compared with females. Carbon and nitrogen stable isotope ratios did not significantly differ between the sexes, suggesting that diet composition and habitat utilization did not vary between the sexes. Based on PCB determinations in the somatic tissue and ovaries, we predicted that PCB concentration of females would increase by 0.6%, on average, immediately after spawning due to release of eggs. Thus, the change in PCB concentration due to release of eggs did not explain the higher PCB concentrations observed in males. Bioenergetics modeling results indicated that the growth dilution effect could account for males being 19% higher in PCB concentration compared with females. Thus, the bulk of the observed difference in PCB concentrations between the sexes was not explained by growth dilution. We concluded that a higher rate of energy expenditure in males, stemming from greater activity and a greater resting metabolic rate, was most likely the primary driver for the observed difference in PCB concentrations between the sexes.

New Jersey↗
USGS2016

A shift in bloater consumption in Lake Michigan between 1993 and 2011 and its effects on Diporeia and Mysis prey

Bioenergetics modeling was used to determine individual and population consumption by Bloater Coregonus hoyi in Lake Michigan during three time periods with variable Bloater density: 1993&ndash;1996 (high), 1998&ndash;2002 (intermediate), and 2009&ndash;2011 (low). Despite declines in Bloater abundance between 1993 and 2011, our results did not show any density-dependent compensatory response in annual individual consumption, specific consumption, or proportion of maximum consumption consumed. Diporeia spp. accounted for a steadily decreasing fraction of annual consumption, and Bloater were apparently unable to eat enough Mysis diluviana or other prey to account for the loss of Diporeia in the environment. The fraction of production of both Diporeia and Mysis that was consumed by the Bloater population decreased over time so that the consumption-to-production ratio for Diporeia + Mysis was 0.74, 0.26, and 0.14 in 1993&ndash;1996, 1998&ndash;2002, and 2009&ndash;2011, respectively. Although high Bloater numbers in the 1980s to 1990s may have had an influence on populations of Diporeia, Bloater were not the main factor driving Diporeia to a nearly complete disappearance because Diporeia continued to decline when Bloater predation demands were lessening. Thus, there appears to be a decoupling in the inverse relationship between predator and prey abundance in Lake Michigan. Compared with Alewife Alosa pseudoharengus, the other dominant planktivore in the lake, Bloater have a lower specific consumption and higher gross conversion efficiency (GCE), indicating that the lake can support a higher biomass of Bloater than Alewife. However, declines in Bloater GCE since the 1970s and the absence of positive responses in consumption variables following declines in abundance suggest that productivity in Lake Michigan might not be able to support the same biomass of Bloater as in the past.

Lake Michigan↗
USGS2016

Reply to comments by Riley and Dunlop on He et al. (2015)

He et al. (2015) described piscivory patterns in the main basin of Lake Huron 1984-2010, during which there was also a pattern of stepwise declines in the abundance of dominant prey fish species. The approach of He et al. (2015) was to couple age-structured stock assessment and fish bioenergetics models to estimate prey fish consumption, and to compare these patterns with prey fish biomass from a bottom trawl survey. Riley and Dunlop (2015) were highly critical of the methods and conclusions reached by He et al. (2015). They claimed that we incorrectly interpreted the bottom trawl survey data, and did not account for uncertainty. We respond to these and other criticisms below, which we find do not undermine our findings.

Canadian Journal of Fisheries and Aquatic Sciences↗