Marine Biology Commons^™

Extinction Risk And Conservation Of The World's Sharks And Rays, Nicholas K. Dulvy, Sarah L. Fowler, John A. Musick, Rachel D. Cavanagh, Peter M. Kyne, Lucy R. Harrison, John K. Carlson, Lindsay N. K. Davidson, Sonja V. Fordham, Malcolm P. Francis, Caroline M. Pollock, Colin A. Simpfendorfer, George H. Burgess, Kent E. Carpenter, Leonard J. V. Compagno, David A. Ebert, Claudine Gibson, Michelle R. Heupel, Suzanne R. Livingstone, Jonnell C. Sanciangco, John D. Stevens, Sarah Valenti, William T. White

Biological Sciences Faculty Publications

The rapid expansion of human activities threatens ocean-wide biodiversity. Numerous marine animal populations have declined, yet it remains unclear whether these trends are symptomatic of a chronic accumulation of global marine extinction risk. We present the first systematic analysis of threat for a globally distributed lineage of 1,041 chondrichthyan fishes-sharks, rays, and chimaeras. We estimate that one-quarter are threatened according to IUCN Red List criteria due to overfishing (targeted and incidental). Large-bodied, shallow-water species are at greatest risk and five out of the seven most threatened families are rays. Overall chondrichthyan extinction risk is substantially higher than for most other …

Concordance Between Phylogeographic And Biogeographic Boundaries In The Coral Triangle: Conservation Implications Based On Comparative Analyses Of Multiple Giant Clam Species, Timery S. Deboer, Maria Rio Abdon Naguit, Mark V. Erdmann, Maria Carmen A. Ablan-Lagman, Ambariyanto, Kent E. Carpenter, Abdul Hamid A. Toha, Paul H. Barber

Biological Sciences Faculty Publications

Marine habitats are in decline worldwide, precipitating a strong interest in marine conservation. The use of biogeographic data to designate ecoregions has had significant impacts on terrestrial conservation efforts. However, classification of marine environments into ecoregions has only become available in the last several years, based on biogeographic data supplemented by geomorphology, ocean currents, and water temperatures. Here we use a comparative phylogeographic approach to test for concordant phylogeographic patterns in three closely related species of Tridacna giant clams across the Coral Triangle, the most biodiverse marine region in the world and one of the most threatened. Data from a …

Effects Of Predation Upon The Long-Spined Sea Urchin Diadema Antillarum By The Spotted Spiny Lobster Panulirus Guttatus, Meredith D. Kintzing, Mark J. Butler Iv

Biological Sciences Faculty Publications

Sea urchins, important herbivores in marine ecosystems, are strongly impacted by both the direct and indirect effects of predation, and the long-spined sea urchin Diadema antillarum is no exception. Once abundant on Caribbean coral reefs, D. antillarum populations were decimated by disease in the early 1980s, and only where their natural predators have been over-fished has D. antillarum recovery been observed. Spiny lobsters (Palinuridae) are predators of sea urchins, and although there are several species of spiny lobster in the Caribbean, only the spotted spiny lobster Panulirus guttatus is restricted to coral reefs where D. antillarum dwells. We investigated the …

Developing Important Marine Mammal Area Criteria: Learning From Ecologically Or Biologically Significant Areas And Key Biodiversity Areas, Collenn M. Corrigan, Jeff A. Ardron, Mia T. Comeros-Raynal, Erich Hoyt, Giuseppe Notarbartolo Di Sciara, Kent E. Carpenter

Biological Sciences Faculty Publications

1. This paper explores how criteria to identify important marine mammal areas (IMMAs) could be developed, and nested in existing global criteria. This process would consider 134 species of marine mammals. 2. Particular attention is given to two suites of global criteria to identify areas important for the persistence of marine biodiversity: Ecologically or Biologically Significant Areas (EBSAs) developed through the Convention on Biological Diversity (CBD), and Key Biodiversity Areas (KBAs) in revision through the International Union for the Conservation of Nature (IUCN). They are seen as mutually complementary in the development of IMMAs. 3. The specificities necessary for identifying …

Oceanography, Geology, Biogeography, And Fisheries Of The Eastern Central Atlantic, Merete Tandstad, Ana Maria Caramelo, Fabio Carocci, Kent Carpenter, Jonnell C. Sanciangco

Biological Sciences Faculty Publications

[Introduction] This species identification guide focuses on the marine species occurring in the eastern Atlantic Ocean from Gibraltar in the north (36°00'N) to the middle of Namibia in the south (23°00'S) and westward to 30°00'W longitude (FAO Fishing Area 34) and to 20°00'W longitude (FAO Fishing Area 47) (Fig 1). Estuarine species, as well as species that only spend part of their life cycle in marine waters (anadromous and catadromous species) are also included.

The West African coast encompasses some of the most productive areas in the world, supporting important fishery resources and a unique diversity. Fisheries constitute an important …

The Impact Of Multiple Nursery Areas On The Population Structure Of Atlantic Menhaden, Brevooria Tyrannus, Kristen A. Anstead

OES Theses and Dissertations

Understanding the population structure and patterns of connectivity in marine fishes is essential when making predictions about a species' resiliency and persistence in an increasingly changing environment. The Atlantic Menhaden Brevoortia tyrannus is a clupeid that plays a critical role in the marine food web and supports one of the largest fisheries on the US East Coast. In addition to a decrease in overall numbers and spawning stock biomass, recruitment levels have remained low since the 1990s. Menhaden use numerous estuaries along the Atlantic coast for juvenile development before recruiting to the adult population and the contribution of each of …

Fish Species Distribution In Seagrass Habitats Of Chesapeake Bay Are Structured By Abiotic And Biotic Factors, Jason J. Schaffler, Jacques Van Montfrans, Cynthia M. Jones, Robert J. Orth

OES Faculty Publications

Seagrass habitats have long been known to serve as nursery habitats for juvenile fish by providing refuges from predation and areas of high forage abundance. However, comparatively less is known about other factors structuring fish communities that make extensive use of seagrass as nursery habitat. We examined both physical and biological factors that may structure the juvenile seagrass-associated fish communities across a synoptic-scale multiyear study in lower Chesapeake Bay. Across 3years of sampling, we collected 21,153 fish from 31 species. Silver Perch Bairdiella chrysoura made up over 86% of all individuals collected. Nine additional species made up at least 1% …

Springtime Contribution Of Dinitrogen Fixation To Primary Production Across The Mediterranean Sea, E. Rahav, B. Herut, A. Levi, Margaret Mulholland, I. Berman-Frank

OES Faculty Publications

Dinitrogen (N-2) fixation rates were measured during early spring across the different provinces of Mediterranean Sea surface waters. N-2 fixation rates, measured using N-15(2) enriched seawater, were lowest in the eastern basin and increased westward with a maximum at the Strait of Gibraltar (0.10 to 2.35 nmol NL-1 d(-1), respectively). These rates were 3-7 fold higher than N-2 fixation rates measured previously in the Mediterranean Sea during summertime and we estimated that methodological differences alone did not account for the seasonal changes we observed. Higher contribution of N-2 fixation to primary production (4-8 %) was measured in the western basin …

Distribution, Prevalence, And Genetic Analysis Of Panulirus Argus Virus 1 (Pav1) From The Caribbean Sea, Jessica Moss, Donald C. Behringer Jr., Jeffrey D. Shields, Antonio Baeza, Alfonso Aguilar-Perera, Phillippe G. Bush, Clement Dromer, Alejandro Herrera-Moreno, Lester Gittens, Thomas R. Matthews, Michael R. Mccord, Michelle T. Schärer, Lionel Reynal, Nathanial Truelove, Mark J. Butler Iv

Biological Sciences Faculty Publications

The pathogenic virus Panulirus argus virus 1 (PaV1) was first discovered in Caribbean spiny lobsters Panulirus argus from the Florida Keys (USA) in 1999 and has since been reported in Belize, Mexico, and Cuba; its distribution in the wider Caribbean is unknown. We collected tissue samples from adult spiny lobsters from 30 locations in 14 countries bordering the Caribbean Sea and used molecular diagnostics to assay for the presence of PaV1. PaV1 occurred primarily in the northern areas of the Caribbean, where its prevalence was highest. The virus was not found in lobsters from the southeastern Caribbean, and its prevalence …

Basin Isolation And Oceanographic Features Influencing Lineage Divergence In The Humbug Damselfish (Dascyllus Aruanus) In The Coral Triangle, Jeremy M. Raynal

Biological Sciences Theses & Dissertations

The Coral Triangle (CT) is a hotspot for marine species diversity as well as for intraspecific genetic diversity. Here, nuclear RAG2 and mitochondrial D-Loop genes were used to identify deep genetic divergence among Dascyllus aruanus (Linnaeus, 1758) populations across relatively short scales within the CT. Mitochondrial clades different by greater than 20 mutational steps were geographically isolated from one another across the distance between Java and the Lesser Sunda Islands, and also east and west across the Philippines. Evidence for population structure in the Sulu Sea and at the Lesser Sunda Islands is also identified. The results suggest that the …

Comparative Phylogeography Of The Emperor Snappers Lethrinus Lentjan And Lethrinus Harak (Lethrinidae: Percoidei) In The Coral Triangle, Andrew B. Hines

Biological Sciences Theses & Dissertations

Many questions remain regarding the origins of the well-known peak in marine biodiversity found in the Coral Triangle. Barriers to gene flow may promote lineage diversification and provide the potential for speciation contributing to the high biodiversity in this area. Population structure was examined in two species of Emperor Snappers (Lethrinidae), Lethrinus harak and Lethrinus lentjan. These species were selected because they share similar life-history traits and geographic distributions, but differ in their habitat preferences. Specimens were collected from within the Coral Triangle as well as other localities across the Indo-Pacific. To investigate phylogeographic patterns and structure a hypervariable …

Environmental And Physiological Influences On Productivity And Carbon Isotope Discrimination In Eelgrass (Zostera Marina L.), Meredith Leigh Mcpherson

OES Theses and Dissertations

A short-term longitudinal study was used to examine the relations of achievement motivation, general motivation, interest, and self-regulatory predictors to cognitive strategy use and academic performance. A total of 102 college undergraduates enrolled in their first college-level introductory psychology course completed an Achieving Tendency Scale of achievement motivation, the Motivated Strategies For Learning Questionnaire, and an Interest Profile in psychology during the first two weeks of the semester. At midterm they completed the Study Activity Survey of cognitive strategy use. At the end of the semester exam grades, total points earned in the course, and grade point average were obtained. …

Response Of Natural Estuarine Algal Communities To Nitrogen Additions, Christopher Schweitzer

OES Theses and Dissertations

Coastal eutrophication resulting from nutrient over-enrichment plagues coastal waterways worldwide. In addition to dissolved inorganic N, many marine microbes, including phytoplankton, have the capacity to use certain dissolved organic nitrogen (DON) compounds. However, while we know that many specific DON compounds are labile, the bulk of the DON pool in nature is uncharacterized. The bioavailability of DON to estuarine microbes currently remains unclear.

It is becoming evident that natural phytoplankton assemblages in estuarine waters are capable of using both inorganic and organic N compounds for growth, both of which are present in anthropogenic point and non-point source discharges. Methods for …

Phytoplankton In Virginia Lakes And Reservoirs, Harold G. Marshall

Virginia Journal of Science

This study involves a phytoplankton summer/autumn survey in 46 Virginia lakes and reservoirs during 2010-2012. A total of 307 taxa were identified which included several filamentous and colonial cyanabacteria in bloom concentrations. With the exception of one natural lake, the other sites sampled represent impoundments created decades ago, with the majority presently classified as meso- or eutrophic. Among the cyanobacteria were 6 known toxin producers (Anabaena circinalis, Anabaena spiroides, Aphanizomenon flos-aquae, Cylindrospermopsis raciborskii, Limnothrix redekei, and Microcystis aeruginosa). The study characterizes phytoplankton populations in these aging freshwater habitats taken from a large number …

Phylogeography Of The Christmas Tree Worm Spirobranchus Corniculatus (Annelida, Serpulidae) From The Coral Triangle, Australia, And Fiji, Abril Iñiguez, Craig J. Starger, Elena Kupriyanova, Demian Willette, Tristan Varman, Joshua A. Drew, Abdul Hamid Toha, Benedict Maralit, Kent E. Carpenter, Paul H. Barber

Biological Sciences Faculty Publications

Christmas Tree Woms is a common name given to a group of large and colourful serpulid polychaetes of the genus Spirobranchus that are obligate symbionts of hermatypic corals commonly found in tropical and subtropical regions world-wide. While current taxonomic species delimitations of these worms are based almost exclusively on the morphology of opercular structures, particularly the number and shape of calcareous opercular horns, high levels of intraspecific variation in this morphological feature have made species boundaries within the genus difficult to resolve. Molecular work to differentiate serpulid worms has long been hampered by the availability of informative markers. Recent work …

Systematic Ichthyofaunal Surveys In Urban And Non-Urban Watersheds, Eugene G. Maurakis, David V. Grimes, Amanda Schutt, Suzy Short

Virginia Journal of Science

Objectives were to model fish species richness relative to natural and anthropogenic variables in Quantico Creek, a forested undisturbed stream environment, and Cameron Run, a highly disturbed urban stream environment in the lower Piedmont-Fall Line region of the Potomac River watershed. Species richness in all stream orders (e.g. avg. range=2.5-9.65 in 1st-3rd orders) of Quantico Creek were significantly higher than those (e.g. avg. range=2.1- 7.6 in 1st -4th orders) of Cameron Run. Fish species richness in Quantico Creek watershed can be modeled by eight factors: season, stream order, elevation, river km, stream width and depth, watershed size, and percent of …

Phosphorus Cycling In The Sargasso Sea: Investigation Using The Oxygen Isotopic Composition Of Phosphate, Enzyme-Labeled Fluorescence, And Turnover Times, Karen Mclaughlin, Jill A. Sohm, Gregory A. Cutter, Michael W. Lomas, Adina Paytan

OES Faculty Publications

Dissolved inorganic phosphorus (DIP) concentrations in surface water of vast areas of the ocean are extremely low (<10 nM) and phosphorus (P) availability could limit primary productivity in these regions. We explore the use of oxygen isotopic signature of dissolved phosphate (δ¹⁸O_PO4) to investigate biogeochemical cycling of P in the Sargasso Sea, Atlantic Ocean. Additional techniques for studying P dynamics including ³³P-based DIP turnover time estimates and percent of cells expressing alkaline phosphatase (AP) activity as measured by enzyme-labeling fluorescence are also used. In surface waters, δ¹⁸O_PO4 values were lower than equilibrium by 3–6%, indicative of dissolved organic phosphorous (DOP) remineralization by extracellular enzymes. An isotope mass balance model using a variety of possible combinations of …

Using Ammonium Pore Water Profiles To Assess Stoichiometry Of Deep Remineralization Processes In Methanogenic Continental Margin Sediments, David J. Burdige, Tomoko Komada

OES Faculty Publications

In many continental margin Sediments, a deep reaction zone exists which is separated from remineralization processes near the sediment surface. Here, methane diffuses upward to a depth where it is oxidized by downwardly diffusing sulfate. However, the methane sources that drive this anaerobic oxidation of methane (AOM) in the sulfate-methane transition zone (SMT) may vary among sites. In particular, these sources can be thought of as either (i) "internal" sources from in situ methanogenesis (regardless of where it occurs in the sediment column) that are ultimately coupled to organic matter deposition and burial, or (ii) "external" sources such as hydrocarbon …

Diatom Proteomics Reveals Unique Acclimation Strategies To Mitigate Fe Limitation, Brook L. Nunn, Jessica F. Faux, Anna A. Hippman, Maria T. Maldonado, H. Rodger Harvey, David R. Goodlett, Philip W. Boyd, Robert F. Strzepek

OES Faculty Publications

Phytoplankton growth rates are limited by the supply of iron (Fe) in approximately one third of the open ocean, with major implications for carbon dioxide sequestration and carbon (C) biogeochemistry. To date, understanding how alteration of Fe supply changes phytoplankton physiology has focused on traditional metrics such as growth rate, elemental composition, and biophysical measurements such as photosynthetic competence (Fv/Fm). Researchers have subsequently employed transcriptomics to probe relationships between changes in Fe supply and phytoplankton physiology. Recently, studies have investigated longer-term (i.e. following acclimation) responses of phytoplankton to various Fe conditions. In the present study, the coastal diatom, Thalassiosira pseudonana, …

Effect Of Laser Ablation Depth In Otolith Life History Scans, Renée R. Hoover, Cynthia M. Jones

OES Faculty Publications

Life history scans of fish otoliths are bringing new insight into the structure, connectivity, and movement of fish populations. Data obtained from such scans, however, possess in-herent limitations that have not yet been fully addressed or understood. For example, several investigators have noted delays in otolith elemental uptake that do not appear to reflect habitat exposure. We hypothesized that the 3-dimensional structure of otoliths may produce sampling artifacts in the results obtained from laser ablation scans. To test this hypothesis, we sampled sagittal otoliths from juvenile Atlantic croaker Micropogonias undulatus with laser ablation inductively coupled plasma mass spectrometry to obtain …

Wintering Eiders Acquire Exceptional Se And Cd Burdens In The Bering Sea: Physiological And Oceanographic Factors, James R. Lovvorn, Merl F. Raisbeck, Lee W. Cooper, Gregory A. Cutter, Micah W. Miller, Marjorie L. Brooks, Jacqueline M. Grebmeier, Angela C. Matz, Cortney M. Schaefer

OES Faculty Publications

During late winter (March) in the Bering Sea, levels of Se in livers and Cd in kidneys of spectacled eiders Somateria fischeri were exceptionally high (up to 489 and 312 µg g⁻¹ dry mass, respectively). Comparison of organ and blood samples during late winter, early spring migration, and breeding suggests that the eiders’ high Se and Cd burdens were accumulated at sea, with highest exposure during winter. High exposure may have resulted from high metabolic demands and food intake, as well as concentrations in food. In the eiders’ remote wintering area, their bivalve prey contained comparable Se levels and …

Combined Effects Of Co2 And Light On Large And Small Isolates Of The Unicellular N2-Fixing Cyanobacterium Crocosphaera Watsonii From The Western Tropical Atlantic Ocean, Nathan S. Garcia, Fei-Xue Fu, Cynthia L. Breene, Elizabeth K. Yu, Peter W. Bernhardt, Margaret R. Mulholland, David A. Hutchins

OES Faculty Publications

We examined the combined effects of light and pCO₂ on growth, CO₂-fixation and N₂-fixation rates by strains of the unicellular marine N₂-fixing cyanobacterium Crocosphaera watsonii with small (WH0401) and large (WH0402) cells that were isolated from the western tropical Atlantic Ocean. In low-pCO₂-acclimated cultures (190ppm) of WH0401, growth, CO₂-fixation and N₂-fixation rates were significantly lower than those in cultures acclimated to higher (present-day approximate to 385ppm, or future approximate to 750ppm) pCO² treatments. Growth rates were not significantly different, however, in low-pCO₂-acclimated cultures of …

Bacterial Colonization And Extinction On Marine Aggregates: Stochastic Model Of Species Presence And Abundance, Andrew M. Kramer, M. Maille Lyons, Fred C. Dobbs, John M. Drake

OES Faculty Publications

Organic aggregates provide a favorable habitat for aquatic microbes, are efficiently filtered by shellfish, and may play a major role in the dynamics of aquatic pathogens. Quantifying this role requires understanding how pathogen abundance in the water and aggregate size interact to determine the presence and abundance of pathogen cells on individual aggregates. We build upon current understanding of the dynamics of bacteria and bacterial grazers on aggregates to develop a model for the dynamics of a bacterial pathogen species. The model accounts for the importance of stochasticity and the balance between colonization and extinction. Simulation results suggest that while …

How Do Shellfisheries Influence Genetic Connectivity In Metapopulations? A Modeling Study Examining The Role Of Lower Size Limits In Oyster Fisheries, Daphne M. Munroe, Eileen E. Hofmann, Eric N. Powell, John M. Klinck

CCPO Publications

Fisheries can potentially alter evolutionary processes such as genetic connectivity and lead to genotypic changes in stocks. Using an individual-based metapopulation genetics model, we examined the possible influence of oyster (Crassostrea virginica) fisheries on genetic connectivity. We simulated a range of realistic fishing pressures, with and without a minimum size limit (limit = 63.5 mm), over a range of fishing scenarios including single-area and stock-wide fisheries. Movement of a neutral marker gene provided an indicator of gene transfer between populations. Simulations showed that fishing may alter genetic connectivity. Increasing fishing pressure tended to decrease potential for fished populations …

Corrected Numbers For Fish On Red List, Bruce B. Collette, Beth Polidoro, Kent Carpenter

Biological Sciences Faculty Publications

(First paragraph) Kelly Swing gives inaccurate numbers for marine fish species on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species. He also mistakenly conflates the scientific process of species assessment for the Red List with the separate political process of IUCN member voting (Nature 494, 314; 2013).

Seasonal Dynamics Of Mesodinium Rubrum In Chesapeake Bay, Matthew D. Johnson, Doane K. Stoecker, Harold G. Marshall

Biological Sciences Faculty Publications

The photosynthetic ciliate Mesodinium rubrum is a common member of coastal phytoplankton communities that is well adapted to low-light, turbid ecosystems. It supports the growth of, or competes with, harmful dinoflagellate species for cryptophyte prey, as well as being a trophic link to copepods and larval fish. We have compiled data from various sources (n = 1063), on the abundance and distribution of M. rubrum in Chesapeake Bay and its tributaries. Because M. rubrum relies on obtaining organelles from cryptophyte algae to maintain rapid growth, we also enumerated cryptophyte algae in the portion of these samples that we collected …

Larval Connectivity And The International Management Of Fisheries, Andrew S. Kough, Claire B. Paris, Mark J. Butler Iv

Biological Sciences Faculty Publications

Predicting the oceanic dispersal of planktonic larvae that connect scattered marine animal populations is difficult, yet crucial for management of species whose movements transcend international boundaries. Using multi-scale biophysical modeling techniques coupled with empirical estimates of larval behavior and gamete production, we predict and empirically verify spatio-temporal patterns of larval supply and describe the Caribbean-wide pattern of larval connectivity for the Caribbean spiny lobster (Panulirus argus), an iconic coral reef species whose commercial value approaches $1 billion USD annually. Our results provide long sought information needed for international cooperation in the management of marine resources by identifying lobster larval connectivity …

Conus: First Comprehensive Conservation Red List Assessment Of A Marine Gastropod Mollusc Genus, Howard Peters, Bethan C. O'Leary, Julie P. Hawkins, Kent E. Carpenter, Callum M. Roberts

Biological Sciences Faculty Publications

Marine molluscs represent an estimated 23% of all extant marine taxa, but research into their conservation status has so far failed to reflect this importance, with minimal inclusion on the authoritative Red List of the International Union for the Conservation of Nature (IUCN). We assessed the status of all 632 valid species of the tropical marine gastropod mollusc, Conus (cone snails), using Red List standards and procedures to lay the groundwork for future decadal monitoring, one of the first fully comprehensive global assessments of a marine taxon. Three-quarters (75.6%) of species were not currently considered at risk of extinction owing …

Ontogenetic Shifts In Resource Allocation: Colour Change And Allometric Growth Of Defensive And Reproductive Structures In The Caribbean Spiny Lobster Panulirus Argus, Joshua R. Anderson, Angelo J. Spadaro, J. Antonio Baeza, Donald C. Behringer

Biological Sciences Faculty Publications

Resource allocation theory predicts a disproportionately large allocation of resources to defensive structures during early ontogeny in organisms that are subject to more intense predation at smaller than at larger body sizes. We tested this prediction on the Caribbean spiny lobster Panulirus argus, which exhibits a negative relationship between predation risk and body size with a high natural mortality of smaller individuals. Independent allometric growth analyses demonstrated that numerous defensive structures (e.g. orbital horns, segments supporting the antenna, the tail fan) display negative allometric growth throughout ontogeny. We interpret these findings as lobsters investing disproportionately more resources to defensive …

A Preliminary Examination Of An In Situ Dual Dye Approach To Measuring Light Fluxes In Lotic Systems, Elizabeth C. Minor, Elizabeth James, Jay A. Austin, Veronica Nelson, Ryan Lusk, Kenneth Mopper

Chemistry & Biochemistry Faculty Publications

Light is a critical parameter in aquatic ecosystems, affecting primary production and in situ photochemistry. However, measuring light exposure for suspended particles or dissolved components in a dynamic water column can be challenging with existing Eulerian approaches. Here, we assess the simultaneous deployment of two dyes differing in photolability (rhodamine WT and fluorescein) as a Lagrangian measure of sunlight exposure in a lotic system. Fluorescein is sensitive to light exposure; rhodamine WT is relatively photostable. We examined dye fluorescence at various pH, salinity, and temperature conditions. We also tested dye photolability as a function of pH and wavelength range. In …