Adi Khen

Adi Khen

*This is an algorithmically generated author record based on publicly available information.

Current Organizations / Institutes

The organizations/institutes from the scholar's latest article


Adi Khen has a total of 24 co-authors. Here are the top co-authors by number of shared publications:

  • Jennifer E. Smith

    Marine Biology Research Division, Scripps Institution of Oceanography, University of California, San Diego

    3 shared publications
  • Jennifer E. Smith

    Center for Marine Biodiversity and Conservation, Scripps Institution of Oceanography, University of California, San Diego, La Jolla, CA, United States

    2 shared publications
  • Maggie D. Johnson

    Biological and Environmental Sciences and Engineering Division, King Abdullah University of Science and Technology, Thuwal, Saudi Arabia

    2 shared publications
  • Michael D. Fox

    Biological and Environmental Sciences and Engineering Division, King Abdullah University of Science and Technology, Thuwal, Saudi Arabia

    2 shared publications
  • Amanda L. Carter

    Center for Marine Biodiversity and Conservation, Scripps Institution of Oceanography, University of California San Diego, La Jolla, USA

    1 shared publication
  • Christine A. Steinke

    Scripps Institution of Oceanography University of California San Diego La Jolla California USA

    1 shared publication

Latest Publications

  • Open Access

    Characterizing morphology of Egregia menziesii (Laminariales) in California over 2 centuries using historical and contemporary herbarium specimens

    by Adi Khen, Kai M. Moore, Siobhan A. Braybrook, Peter S. Vroom, Kathy Ann Miller, Jennifer E. Smith

    The canopy‐forming feather boa kelp Egregia menziesii exhibits remarkable morphological variability across its geographic range. Regional morphotypes of Egregia were once considered separate species, but they were not determined to be genetically distinct; instead, their morphology was thought to reflect local physical or environmental conditions. Although morphological variation in Egregia has long been observed and was previously characterized through field surveys in the early 2000s, we revisited this topic using digital morphometrics (i.e., image analysis) of 1624 macroalgal herbarium specimens from California dating back to the 19th century. We observed that the morphology of Egregia (rachis texture, lateral blade shape, and blade or pneumatocyst density) varied along a latitudinal gradient and could be predicted by seawater temperature and wave height. We also identified some region‐specific morphological changes in recent decades. Further, the monthly presence or absence of sporophylls in southern‐region specimens provided preliminary evidence into the reproductive phenology of Egregia . Herbarium collections are invaluable for studying patterns in morphology because they showcase inter‐ and intraspecific variability and establish a baseline for comparison through time. Integrating natural historical and contemporary data will be critical for understanding and predicting future trends in the context of ocean warming.

  • Open Access

    Benthic algal community dynamics on Palmyra Atoll throughout a decade with two thermal anomalies

    by Adi Khen, Maggie D. Johnson, Michael D. Fox, Jennifer E. Smith

    Coral reef algae serve many important ecological functions, from primary production to nutrient uptake and reef stabilization, but our knowledge of longer-term effects of thermal stress on algae in situ is limited. While ocean warming can facilitate proliferation of algae and potential phase shifts from coral to macroalgal-dominated states, algal responses may vary by species, genus, functional group, or type (e.g., calcareous vs. fleshy). We used 11 years of annual monitoring data (2009-2019) that spans two El Niño-associated heatwaves to examine benthic algal community dynamics on Palmyra Atoll in the central Pacific Ocean. We quantified the percent cover of algal taxa via image analysis of permanent benthic photoquadrats from two habitats on Palmyra: the deeper, wave-exposed fore reef (10 m depth) and the shallower, wave-sheltered reef terrace (5 m depth). Each habitat was characterized by distinct algal communities: predominantly calcareous taxa on the fore reef and predominantly fleshy taxa on the reef terrace. Patterns in abundance fluctuated over time and/or in response to thermal anomalies in 2009 and 2015. Fleshy algae generally increased in cover post-warming, which coincided with large declines of the calcified macroalgae, Halimeda spp. Long-term monitoring of coral reef algal communities is critical for understanding their differential responses to thermal stress and can improve projections of ecosystem functioning in the context of global change.


  • Open Access

    Inter- and intraspecific responses of coral colonies to thermal anomalies on Palmyra Atoll, central Pacific

    by Adi Khen, Michael D Fox, Maggie D Johnson, Christopher B Wall, Jennifer E Smith

    Long-term monitoring of individual coral colonies is important for understanding variability between and within species over time in the context of thermal stress. Here, we analyze an 11-year time series of permanent benthic photoquadrats taken on Palmyra Atoll, central Pacific, from 2009 to 2019 to track the growth (i.e., increase in live planar area), pigmentation or lack thereof (“discoloration”), partial or whole-colony mortality, survival, and regrowth of 314 individual coral colonies of nine focal species from two reef habitat types. During this period, thermal anomalies occurred on Palmyra in conjunction with El Niño-Southern Oscillation events in both 2009 and 2015, of which the latter heatwave was longer-lasting and more thermally-severe. We found that coral responses varied by habitat, within and among species, and/or according to the degree of accumulated thermal stress. Nearly all species, particularly Stylophora pistillata and Pocillopora damicornis, responded more negatively to the 2015 heatwave in terms of colony-specific discoloration and reduction in live planar area. While discoloration was more prominent at the shallower reef terrace compared to the fore reef for this subset of colonies, the reef terrace exhibited greater stability of community-wide coral cover. Colony fate was associated with severity of discoloration at the time of warming: one year following the 2009 heatwave, more severely discolored colonies were more likely to grow, yet following the second heatwave in 2015, colonies were more likely to experience shrinkage or mortality. However, colonies that were more severely discolored in 2009 were not necessarily more discolored in 2015, suggesting that colony-specific factors may be more influential in governing responses to thermal stress.


  • Open Access

    Standardization of in situ coral bleaching measurements highlights the variability in responses across genera, morphologies, and regions

    by Adi Khen, Christopher B. Wall, Jennifer E. Smith

    Marine heatwaves and regional coral bleaching events have become more frequent and severe across the world’s oceans over the last several decades due to global climate change. Observational studies have documented spatiotemporal variation in the responses of reef-building corals to thermal stress within and among taxa across geographic scales. Although many tools exist for predicting, detecting, and quantifying coral bleaching, it remains difficult to compare bleaching severity (e.g., percent cover of bleached surface areas) among studies and across species or regions. For this review, we compiled over 2,100 in situ coral bleaching observations representing 87 reef-building coral genera and 250 species of common morphological groups from a total of 74 peer-reviewed scientific articles, encompassing three broad geographic regions (Atlantic, Indian, and Pacific Oceans). While bleaching severity was found to vary by region, genus, and morphology, we found that both genera and morphologies responded differently to thermal stress across regions. These patterns were complicated by (i) inconsistent methods and response metrics across studies; (ii) differing ecological scales of observations (i.e., individual colony-level vs. population or community-level); and (iii) temporal variability in surveys with respect to the onset of thermal stress and the chronology of bleaching episodes. To improve cross-study comparisons, we recommend that future surveys prioritize measuring bleaching in the same individual coral colonies over time and incorporate the severity and timing of warming into their analyses. By reevaluating and standardizing the ways in which coral bleaching is quantified, researchers will be able to track responses to marine heatwaves with increased rigor, precision, and accuracy.


  • Open Access

    First known observations of brooding, development, and hatching of fertilized eggs for the North Pacific bigeye octopus, Octopus californicus

    by Adi Khen, Lillian R. McCormick, Christine A. Steinke, Greg W. Rouse, Phil J. Zerofski

    The North Pacific bigeye octopus, Octopus californicus (Berry, 1911) is a cold‐water, deep‐sea octopod. Little is known about their biology due to difficulty accessing their natural habitat and obtaining live specimens. Although they are a frequent bycatch product in commercial bottom trawl fisheries, individuals of this species have rarely been raised in captivity and their embryonic development has not yet been documented. Considering these limitations, we were fortunate to have witnessed this process leading to successful hatching in an aquarium setting. Here, we present a brief observational account of the first‐known record of brooding, development, and hatching of fertilized eggs for O. californicus. The incubation time was a maximum of 10 months at a temperature between 8–10°C and embryos hatched over a period of 2.5 months. While more detailed research is needed, this preliminary information contributes to our limited knowledge of this species and supports life history theories of prolonged embryonic development under colder temperatures.


  • Open Access

    Decadal stability of coral reef benthic communities on Palmyra Atoll, central Pacific, through two bleaching events

    by Adi Khen, Maggie D. Johnson, Michael D. Fox, Samantha M. Clements, Amanda L. Carter, Jennifer E. Smith

    The prevalence of coral bleaching due to thermal stress has been increasing on coral reefs worldwide. While many studies have documented how corals respond to warming, fewer have focused on benthic community responses over longer time periods or on the response of non-coral taxa (e.g., crustose coralline algae, macroalgae, or turf). Here, we quantify spatial and temporal changes in benthic community composition over a decade using image analysis of permanent photoquadrats on Palmyra Atoll in the central Pacific Ocean. Eighty permanent plots were photographed annually between 2009 and 2018 on both the wave-exposed fore reef (FR, 10 m depth,n = 4 sites) and the wave-sheltered reef terrace (RT, 5 m depth,n = 4 sites) habitats. The El Niño events of 2009–2010 and 2015–2016 resulted in acute thermal stress and coral bleaching was observed at both reef habitats during these events. Across 10 yr and two bleaching events, the benthic community structure on Palmyra shows evidence of long-term stability. Communities on the RT exhibited minimal change in percent cover of the dominant functional groups, while the FR had greater variability and minor declines in hard coral cover. There was also spatial variation in the trajectory of each site through time. Coral cover decreased at some sites 1 yr following both bleaching events and was replaced by different algal groups depending on the site, yet returned to pre-bleaching levels within 2 yr. Overall, our data reveal the resilience of calcifier-dominated coral reef communities on Palmyra Atoll that have persisted over the last decade despite two bleaching events, demonstrating the capacity for these reefs to recover from and/or withstand disturbances in the absence of local stressors.


  • Planktonic trophic structure in a coral reef ecosystem – Grazing versus microbial food webs and the production of mesozooplankton

    by Ryota Nakajima, Haruka Yamazaki, Levi S. Lewis, Adi Khen, Jennifer E. Smith, Nobuyuki Nakatomi, Haruko Kurihara